Noncystimorph Colonial Rugose Corals of the 

Onesquethaw and Lower Cazenovia Stages 

(Lower and Middle Devonian) in 

New York and Adjacent Areas 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 
Noncystimorph Colonial Rugose Corals of the 

Onesquethaw and Lower Cazenovia Stages 

(Lower and Middle Devonian) in 

New York and A9jacent Areas 

By WILLIAM A. OLIVER, Jr. 

G E 0 L 0 G I CAL SURVEY P R 0 F E-S S I 0 N A L PAPER 8 6 9 

Paleobiology and biostratigraphy of 3 8 

species of colonial rugose coTals' from 

New York) Ontario) Ohio) Kentucky) 

Indiana) Michigan) and Quebec 

UNITED STATES GOVERNl\1ENT PRINTI'NG OFFICE, WASHINGTON 1976 
UNITED STATES DEPARTMENT OF THE INTERIOR 

THOMAS S. KLEPPE, Secretary 

GEOLOGICAL SURVEY 

V. E. McKelvey, Director 

Library of Congress Cataloging in Publication Data 

Oliver, William Albert, Jr., 1926-

Noncystimorph colonial rugose corals of the Onesquethaw and lower Cazenovia stages (Lower and Middle Devonian) in New York and adjacent areas. 

(Geological Survey professional paper ; 869) 

Bibliography: p. 

Includes index. 

Supt. of Docs. no.: I 19.16:869 

1. Rugosa. 2. Paleontology-Devonian. 3. Paleontology-Northeastern states. 4. Paleontology-Canada. I. 

Title. II. Series: United States. Geological Survey. Professional paper ; 869. 

QE778.044 563'.6 74-32012 

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

Washington, D.C. 20402 

Stock Number 024-001-02815-8 
CONTENTS 

Abstract 

Introduction ___ ------ ____________________ --------

Acknowledgments _____________ - _____ --- _________ _ 

Onesquethaw and Cazenovia Stages -----------------

Eastern North America --------------------------- Regional stratigraphy ___________________________ _ 

Appalachian Basin ---------------------------

New York and Niagara Peninsula, Ontario __ 

Central Ohio -----------------------------

Northeastern Ohio -----------------------

East-central Kentucky --------------------

Appalachians south of New York ----------

Quebec ----------------------------------

Moose River Basin, Ontario -------------------

Michigan Basin ------------------------------

Woodstock area, Ontario ------------------

Mackinac Straits, Michigan ---------------

Northwestern Ohio-southeastern Michigan __ _ 

Illinois Basin _______________________________ _ 

Falls of the Ohio, Indiana-Kentucky --------

Grand Tower area, Illinois-Missouri --------

Age of the Onesquethaw Stage --------------------

Biostratigraphy __ ----- __________________________ _ 

Paleobiogeography --------------------------------

Previous systematic work -------------------------

Coral morphology -------------------------------- Microstructure _____________________________ _ 

Carinae -------------------------------------

Page 

1 

1 

2 

2 

4 

5 

5 

5 

12 

13 

14 

15 

15 

16 

17 

17 

18 

18 

18 

19 

19 

19 

21 

23 

30 

31 

31 

31 

Intracolony and intercolony variation ---------------

Variation in Acinophyllum segregatum ---------- Terminology ____________________________ _ 

Individual variation ----------------------

Geographic and stratigraphic variation ------

Variation in other species ---------------------

Summary and conclusions ---------------------

Systematic paleontology ___ --- ___ -- ____ ---- _-------

Depositories ____ --- __ -----_------------------

Classification __ -- ____ --- _ --------------------

Classification and species list-an outline ---

Family Stauriidae ----------------------------

Genus Synaptophyllum Simpson -----------

Family Craspedophyllidae ---------------------

Subfamily Cylindrophyllinae ___ - ___ --_----

Genus Acinophyllum McLaren ---------

Genus Cylindrophyllum Simpson -------

Genus Prismatophyllum Simpson ------

Genus Asterobillingsa Oliver ----------

Subfamily Craspedophyllinae ___ - ___ -- __ - _-

Genus Eridophyllum Milne-Edwards and 

Haime ____ --- _ --------------------

Genus Grewgiphyllum Oliver ----------

Family Disphyllidae -------------------------

Genus Disphyllum? -----------------------

Family Zaphrentidae -------------------------

Genus Cyathocylindrium Oliver ------------

Genus Heliophyllum Hall -----------------

Locality data ------------------------------------ References cited ____ -- ____ ---___ ------------------

Index -------------------------------------------

ILLUSTRATIONS 

PLATE 1. 

2-4. 

5. 

6. 

7. 

8-10. 

11. 

12-14. 

15-22. 

23, 24. 

25-28. 

29. 

30. 

31-33. 

34. 

35-38. 

39, 40. 

41-43. 

44. 

[Plates follow index] 

Reconstruction of Edgecliff bioherm scene. 

Synaptophyllum arundinaceum (Billings). 

S. tabulatum (Simpson). 

S. kladion n. sp. 

S. kladion n. sp. and S. arundinaceum (Billings). 

Acinophyllum simcoense (Billings). 

A. simcoense (Billings) and A. segregatum (Simpson) . 

A. stramineum (Billings) . 

A. segregatum (Simpson). 

A. mclareni Fagerstrom. 

A. stokesi (Milne-Edwards and Haime). 

A. vermetum (Weisbord). 

Disphyllum? rectiseptatum (Rominger) . 

Cylindrophyllum elongatum Simpson. 

C. elongatum and C. sp. A. 

C. propinquum Stewart. 

C. laxtensum n. sp. 

C. deearium n. sp. 

Disphyllum? stummi (Oliver). 

III 

Page 

32 

32 

32 

33 

36 

40 

42 

43 

43 

44 

44 

46 

46 

51 

53 

53 

68 

77 

88 

96 

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108 

111 

112 

113 

117 

123 

133 

145 

153 
IV 

PLATE 45. 

46-48. 

49-52. 

53, 54. 

55, 56. 

5·7-60. 

61-64. 

65. 

66. 

67. 

68, 69. 

70-72. 

73. 

74, 75. 

76-79. 

80-86. 

87-90. 

91. 

92,9-3. 

94. 

95. 

9'6-98. 

99-102. 

103. 

104, 105. 

106. 

107. 

108. 

FIGURE 1. 

2. 

3. 

4. 

5. 

6. 

7. 

8. 

9. 

10. 

11. 

12. 

13. 

14. 

15. 

16. 

17. 

18. 

19. 

20. 

21.. 

22. 

23. 

24-37. 

CONTENTS 

D.? stummi (Oliver) and Cylindrophyllum sp. cf. C. propinquum Stewart. 

Prismatophyllum prisma Lang and Smith. 

P. ovoideum (Davis) . 

P. truncata Stewart. 

P. anna (Whitfield). 

Asterobillingsa magdisa magdisa Oliver. 

A. magdisa steorra Oliver. 

A. rugosa (Hall). 

Radiastraea arachne Stumm. 

Eridophyllum seriale Milne-Edwards and Haime. 

E. seriale Milne-Edwards and Haime, population B. 

E. subseriale Stumm. 

E. aulodokum n. sp. 

E. corniculum n. sp. 

Grewgiphyllum colligatum (Billings). 

Cyathocylindrium opulens Oliver and C. sp. cf. C. opulens. 

C. gemmatum (Hall). 

C. n. sp. A. 

Heliophyllum megaproliferum n. sp. 

H. n. sp. B. 

H. n. sp. C. 

H. coalitum (Rominger) . 

H. monticulum (Ehlers and Stumm). 

H. cassum n. sp. 

H. procellatum 'n. sp. 

H. sp. cf. H. procellatum n. sp. 

Eridophyllum conjunctum (Davis). 

E. conjunctum (Davis) and E. sp. cf. E. conjunctum (Davis). 

Page Map of study area __ _ __ ___ ____ _ _ ____ _ _ _ __ __ _ _ _ _ ___ _ _ _ _ ____ _ __ _ __ __ ___ _ _ _ ___ _ _ __ _ _ __ _ _ _ ____ _ __ 3 

Chart showing development of stage nomenclature in the New York standard sequence --------------- 4 

Correlation chart of principal stratigraphic sections, eastern North America ------------------------ 6 

West-east cross section of Onondaga Limestone in New York------------------------------------ 9 

Map of Edgecliff facies, New York and Ontario ------------------------------------------------- 10 

Correlation of Onesquethaw sequences: Ohio-Ontario-New York -------------------------------·---- 13 

Correlation of Onesquethaw sequences: OhiQo-Pennsylvania-New York ------------------------------ 14 

Map showing section lines for figures 4, 6, 7 --------------------------------------------------- 16 

Chart showing history of nomenclature for the rock sequence at the Falls of the Ohio --------------- 20 

Chart showing stratigraphic range of genera and species of colonial rugose corals described in this report ----------------------------------------------------------------------------------- 22 

Map showing distribution of studied lower and middle Onesquethaw corals ----------------------- 24 

Map showing distribution of studied upper Onesquethaw, Edgecliff assemblage corals -------------- 25 

Map showing distribution of studied upper Onesquethaw, Moorehouse assemblage corals and general localities of Seneca-age corals _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 26 

Map showing world distribution of coral genera characteristic of the Eastern Americas Realm ------ 28 

Graph of septal number versus diameter in samples of selected colonies of Acinophyllum segregatum _ 34 

Graph of intercolony septal number versus diameter in samples of Acinophyllum segregatum ------- 36 

Normal distribution of intercolony coefficients of septal number variation in Acinophyllum segregatum 40 

Transverse thin section and map of colony of Acinophyllum segregatum ------------------------ 43 

Graph showing bimodal distribution of septal number in colony Acinophyllum segregatum ----------- 44 

Graph showing intercolony V;; and range of intracolony Vn in samples of colonial rugose corals ------ 45 

Graph showing data of figure 20 rearranged by stratigraphic unit ------------------------------- 45 

Graph of n versus din Synaptophyllum spp ---------------------------------------------------- 48 

Diagram showing morphology of tabulae in Acinophyllum spp ----------------------------------- 54 

Graphs of data for: 

24. Acinophyllum simcoense __________________________________________ --- _ _ _ _ _ __ _ _ __ __ _ _ _ 58 

25. A. spp. and Disphyllum? rectiseptatum ------------------------------------------------ 59 26. A. mclareni _____________________________________________________ --- _ _ __ _ __ _ _ __ __ ___ 64 

27. A. stokesi -------------------------------------------------------------------------- 66 

28. Cylindrophyllum spp. and Disphyllum? stummi ---------------------------------------- 72 

29. Prismatophyllum spp __ _ __ _ __ _ __ ____ _ _ _ _ ___ ___ _ __ __ __ __ _ ___ _ ___ _ ___ _ _ __ _ _ _ _ _ _ __ _ __ _ _ _ 81 

30. Asterobillingsa spp. and Heliophyllum procellatum -------------------------------------- 92 

31. Eridophyllum seriale _____________________________ ---- _____ -- ____ ----- __ -- ____ ------- 101 

32. E. spp ----------------------------------------------------------------------------- 104 
FIGURES 24-37. 

TABLES 1-4. 

5. 

6. 

7. 

8. 

9-13. 

14. 

15. 

16-39. 

40-44. 

45. 

46. 

CONTENTS 

33. Grewgiphyllum· colligatum ----------------------------------------------------------- 34. Cyathocy lindrium opulens ____________________________________________________________ _ 

35. C. gemmatum ----------------------------------------------------------------------- 36. H eliophyllum phaceloid spp _______________________ ------ _ -- __ - ------------------------

37. H. massive spp --------------------------------------------------------------------

TABLES 

Geographic distribution of colonial rugose corals: 

1. Middle Onesquethaw (Bois Blanc-Schoharie) __ ----- _ -- ________ ----- _ ---- ____ ---- _ ------ _ 

2. Upper Onesquethaw, Edgecliff assemblage -----------------------------------------------

3. Upper Onesquethaw, Moorehouse assemblage -------------------------------------------- 4. Lower Cazenovia (Seneca-Delaware) _________________ --------- ___ ------ _ ---- __________ _ 

SQmmary of paleontologic data from northeastern Ohio cores ------------------------------------

Stratigraphic ranges of colonial rugose corals ______ -------------------------------------------

Acinophyllum segregatum colony data arranged by stratigraphic unit ------------------------------ Acinophyllum segregatum, summary data ________________________ ---- ____ ---- ________ ----- ___ _ 

Acinophyllum segregatum summary data: _ 

9. Edgecliff bioherm facies __________________ ------------------------_------------------

10. Edgecliff biostrome facies ----------------------------------------------------------- 11. Edgecliff fine-grained limestone facies ______ ---- ___ -- __________ -- __________ --- ________ _ 

12. Moorehouse Member (except Acinophyllum-Eridophyllum bed) -------------------------

13. Moorehouse Acinophyllum-Eridophyllum bed --------------------------------------------

Data from selected colonies of Acinophyllum segregatum arranged in order of increasing coefficient of 

septal number variation ( V n) _____ ------------ _ -----_--------------- ______ ------- _ -- _ -----

Intracolony and intercolony coefficients of variation in 24 samples of colonial rugose corals ---------

Summary data (unless otherwise noted): 

16. Synaptophyllum arundinaceum ____________ - _ -- _ ------ _________ ---- ___ ---- ___________ _ 

17. Acinophyllum simcoense __________________ - --- _ ----------------------------------- __ _ 

18. Acinophyllum stramineum and A. vermetum _ -------------------------------------------

19. Acinophyllum stokesi, Bois Blanc Formation sample -------------------------------------

20. Acinophyllum stokesi, lower Jeffersonville Limestone sample ---------------------------- 21. Cylindrophyllum elongatum __________________________________________ -- ______________ _ 

22. Cylindrophyllum propinquum ________________________________________________________ _ 

23. Cylindrophyllum laxtensum, intracolony data-------------------------------------------

24. deearium ____________________________ ----- _________________________ _ 

25. Prismatophyllum prisma ____________________________________________________________ _ 

26. Prismatophyllum ovoideum ___________________________________________ ---- __ ----- ___ _ 

27. Prismatophyllum truncata and P. anna, intracolony data --------------------------------

28. Asterobillingsa magdisa magdisa ------------------------------------------------------ 29. Asterobillingsa magdisa steorra ______________________________________________________ _ 

30. Asterobillingsa rugosa and A. affinis, intracolony data -----------------------------------

31. Eridophyllum seriale -----------------------------------------------------------------

32. Eridophyllum subseriale and E. aulodokum --------------------------------------------- 33. Eridophyllum corniculum __________________________________ ----- _____________________ _ 

34. Eridophyllum conjunctum, intracolony data __ ------------------------------------------- 35. Grewgiphyllum colligatum ________________________ ------- ____________________ ----- __ _ 

36. Disphyllum? stummi and D.? rectiseptatum --------------------------------------------- 37. Cyathocylindrium opulens ________________________________ ------- ___________ ----- ____ _ 

38. H eliophyllum monticulum __________________________________________________________ --

39. H eliophyllum procellatum, intracolony data _________________ ---~ _______________________ _ 

Distribution of colonial rugose corals in the--

40. Bois Blanc and Schoharie Formations, New York and southwestern Ontario ----------------

41. Bioherm facies of the Edgecliff Member of the Onondaga Limestone ----------------------

42. Nonbioherm facies of the Edgecliff Member of the Onondaga Limestone -------------------

43. Clarence and Nedrow Members of the Onondaga Limestone -----------------------------

44. Moorehouse Member of the Onondaga Limestone ---------------------------------------

.Location of the single colonial rugose coral known from the Seneca Member of the Onondaga Limestone -----------------~----------------------------------------------------------------- List of museum numbers of all type, illustrated, and otherwise mentioned specimens, giving the species, collection number, and locality ___________________________________________________________ _ 

v 

Page 

110 

118 

121 

125 

128 

Page 

5 

9 

11 

12 

15 

21 

37 

38 

38 

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39 

41 

48 

58 

60 

66 

66 

71 

7.4 

75 

76 

80 

84 

86 

92 

92 

95 

101 

105 

107 

108 

110 

112 

-119 

129 

132 

133 

133 

134 

135 

135 

135 

135 
NONCYSTIMORPH COLONIAL RUGOSE CORALS OF THE ONESQUETHAW AND 

LOWER CAZENOVIA STAGES (LOWER AND MIDDLE DEVONIAN) IN 

NEW YORK AND ADJACENT AREAS 

By WILLIAM A. OLIVER, JR. 

ABSTRACT 

A succession of coral faunas is recognized within the Onesquethaw and lower Cazenovia Stages in Eastern North 

America. Corals are rare in the lower Onesquethaw ( Siegenian 

and (or) Emsian age). A middle Onesquethaw (Emsian) 

coral assemblage is homogeneous and widespread and includes 

several colonial rugose corals. In New York this assemblage 

is found in the Schoharie and Bois Blanc Formations. 

The Onondaga Limestone, except for its upper member, is 

upper Onesquethaw (Eifelian) in age. The Onondaga, and 

equivalent formations in other parts of Eastern North America, 

contain a large variety of corals including many colonial rugose 

corals. The areal distribution of upper Onesquethaw corals is 

partly controlled by facies, but successions of coral assemblages 

in New York, Ohio, and Kentucky are similar and provide the 

basis for biostratigraphic subdivision of the upper Onesquethaw. Colonial rugose corals are most numerous and diverse 

in bioherms found within the Edgecliff Member of the Onondaga in New York and southwestern Ontario. They are less 

diverse and numerous in biostrome facies of the same age, but 

this facies is more widely developed. A still smaller colonial 

rugose coral assemblage in the higher Moorehouse Member of 

the Onondaga Limestone is also widespread. 

The Seneca Member of the Onondaga Limestone is in the 

lower part of the Cazenovia Stage (Eifelian). Only one 

colonial rugose coral is known from this member in New 

York, but additional forms are known from rocks of the 

same age in other areas. 

All colonial rugose coral species known from the Onesquethaw and lower Cazenovia Stages in New York and Niagara 

Peninsula of Ontario, except the cystimorphs, are described 

on the bas·is of available specimens from all of Eastern North 

America. Additional species froin Kentucky, Ohio, Michigan, 

Quebec, and Venezuela are described because of their importance in understanding the New York-Ontario genera and 

species. Genera are described and interpreted on the basis 

of all known species. Most of the genera and all of the 

species are endemic to the Eastern America~ Faunal Realm. 

Their histories in the Eastern Americas Realm and analogous 

Old World Realm genera are discussed in detail. 

A morphologic and stratigraphic succession from early 

species of Prismatophyllum to Grewgiphyllum to Eridophyllum indicates the possible origin of the Craspedophyllinae in 

Prismatophyllum (Cylindrophyllinae). The earliest known 

cylindrophyllinid, however, is Asterobillingsa, which seems 

an unlikely ancestor for the whole group. 

The nature and amount of variation, both within and between colonies, differs with the character considered, environment, and genetics. In general, number of septa is a low 

varying character within the studied colonies. Intercolony 

variation of the same character is a function of environment 

in some species. Variation tends to be greatest in situations 

of maximum coral diversity and least with low diversity. 

Variation in apparently the same ecologic situation differs 

in diffe.rent taxa. In general, intracolony variation is less in 

the Craspedophyllidae than in the Zaphrentidae. · 

Thirty-nine species in 10 genera are described and assigned to 4 families, one of which is divided into 2 subfamilies. 

The families and new or only recently described subordinate 

taxa are: Stauriidae with Synaptophyllum kladion n. sp.; 

Craspedophyllidae, Cylindrophyllinae with Cylindrophyllum 

laxtensum n. sp. C. deearium n. sp., Asterobillingsa Oliver, 

type species A. magdisa Oliver, A. magdisa rnagdisa Oliver, 

and A. magdisa steorra Oliver; Craspedophyllinae with 

Eridophyllum aulodokum n. sp., E. corniculum n. sp., and 

Grewgiphyllum Oliver (type species G. colligatum (Billings)); 

Disphyllidae with no new taxa; and Zaphrentidae with Cyathocylindrium Oliver, type species C. opulens Oliver, Heliophyllum megaproliferum n. sp., H. cassum n. sp., and H. 

procellatum n. sp. 

INTRODUCTION 

This is primarily a study of the noncystimorph 

colonial rugose corals of the Onesquethaw and lower 

Cazenovia Stages (upper Siegenian-Eifelian) in 

Eastern North America. It is based on a detailed 

analysis of the.se corals from New York and the 

Niagara Peninsula of Ontario but also includes comparisons, discussions, and some descriptions of forms 

from rocks of the s•ame age in Ohio, Kentucky, 

Indiana, Michigan, other p·arts of southwestern 

Ontario, Quebec, Virginia, and Illinois-Missouri 

1 
2 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

(fig. 1). These areas are all within the Eastern 

North American (ENA) Faunal Province of Early 

and Middle Devonian time. 

The Onesquethaw colonial rugose corals belong 

to several families: Stauriidae, Craspedophyllidae 

( Cylindrophyllinae and Graspedophyllinae), Zaphrentidae and one or more cystimorph groups. The 

Stauriidae, Craspedophyllidae. and Za.phrentidae 

are the subject of this paper; the cystimorphs have 

not been studied. Although the first emphasis is on 

corals from New York and Ontario, their congeners 

from the Columbus Limestone in Ohio and the J effersonville Limestone in Kentucky-Indiana have 

been examined and compared ; some of these are 

described and illustrated. 

The first part of this paper is a review of the 

stratigraphy of Onesquethaw- and Cazenovia-age 

rocks in the study area with discussions of detailed 

correlations both within the Province and with 

other areas, principally Western North America 

and Europe. It includes a summary of the history of 

faunal provincialis.m and its extent in time and space 

and a description of the biostratigraphic sequence. 

ACKNOWLEDGMENTS 

The principal collection on which this study is 

based was accumulated over a number of years. Collecting began while I was a temporary geologist 

with the New York State Museum and Science Service, Geological Survey. I am indebted to the late 

Winifred Goldring, former State Paleontologist, and 

J. G. Broughton, former State Geologist, for financial 

support prior to 1957, and to them and D. F. Fisher, 

State Paleontologist L. V. Rickard, Assistant State 

Paleontologist, W. L. Kreidler, Senior Scientist, J. 

F. Davis, Senior Scientist and now State Geologist, 

and B. M. Bell, Curator of Paleontology, for moral 

and logistical help as well as intellectual stimulation 

over the many years of our formal and informal association. The State Museum freely loaned specimens and exercised great patience. 

I received the full cooperation of the Geological 

Survey of Canada. D. J. McLaren and T. E. Bolton 

_arranged for long-term loans of large collections, 

and McLaren, B. V. Sanford,and A. W. Norris consulted in the field. 

Many additional institutions loaned specimens. 

These and the responsible authorities ·are: American Museum of Natural History, N. D. Newell and 

R. L. Batten; British Museum (Natural History), 

C. T. Scrutton and R. F. Wise; University of Buf- , 

falo, E. J. Buehler; University of California, Berkeley, J. H. Peck, Jr.; Harvard University, Bernhard 

Kummel; University of Iowa, H. L. Strimple; University of Michigan, E. C. Stumm, R. V. Kessling, 

and D. B. Macurda; Ohio State University, J. J. 

Burke and S.M. Bergstrom; Royal Ontario Museum, 

R. R. H. Lemon and John Monteith; and U.S. National Museum of Natural History, F. J. Collier. 

I am obliged to the following individuals for permitting me to study American type and other specimens in their institutions: Rudolf Birenheide, 

Forsch ungsinstitut Sencken berg, Frankfurt-amMain ; J iirgen Kullman, Institut fiir Geologie und 

PaHiontologie, Tiibingen; C. T. Scrutton, British 

Museum (Natural History), London; and Pierre 

Semenoff-Tian-Chansky, Institute de Paleontologie, 

Paris. These visits were supported during 1969-70 

by N.S.F. Grant No. GB 8357. 

During the course of this study, specimens were 

given to the U.S. National Museum by J. W. Wells, 

Cornell University, and by E. J. Buehler, University 

of Buffalo. These are specifically acknowledged in 

the appropriate systematic descriptions. 

Thin sections were expertly and often exquisitely 

prepared by William C. Pinckney, Jr., without whose 

help this study would not have been possible in its 

present form. 

Photographs were made by D. H. Massie (thin 

sections) and R. H. McKinney (exteriors) , except 

as noted. Film processing and printing was by H. E. 

Mochizuki and by Massie. 

The manuscript was critically reviewed by J. T. 

Dutro, Jr., U.S. Geological Survey, and by J. E. 

Sorauf, State University of New York at Binghamton. Both reviewers helped substantially by noting 

errors, inconsistencies, and unclear statements. 

ONESQUETHAW AND CAZENOVIA STAGES 

The stage nomenclature for eastern North 

America was established by Cooper and others 

(1942) because of the recognized need for a provincial standard to permit the separation of problems 

of correlation within North America from problems 

of intercontinental correlation. The scheme has been 

particularly useful for Lower and Middle Devonian 

rocks in the eastern half of the continent because 

of the provincial nature of their enclosed fossils. It 

has been less useful in western North America and 

for Upper Devonian rocks. 

The Onesquethaw Stage was defined to include 

the Esopus, Schoharie, and Onondaga Formations in 1 
ONTARIO 

ONESQUETHAW AND CAZENOVIA. STAGES 

HUDSON B AYj;; 

I 

C A \N A D A 

I 

I 

p R ECAM BIR IAN SHIELD 

( 

\ 

\ 

0 100 200 300 400 MILES 

~~1,-L'~--~~--~-L'-----' I I I I 

0 100 200 300 400 KILOMETRES 

EXPLANATION 

Onesquethaw rocks 

Structural arches and 

present boundaries of basins 

FIGURE 1.-0utline map of study area showing principal structural elements mentioned in the text. Geology from Geologic 

Map of North America, U.S. Geological Survey (1965). 

3 
4 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Albany County, N.Y., and correlative units elsewhere. The succeeding formations of the Hamilton 

Group were placed in the Cazen.ovia and Tioughnioga 

Stages. General changes in the understanding of the 

stages can be realized by comparing the correlation 

charts of Cooper and others, 1942 with those of 

Rickard, 1964, and Oliver and others, 1968 (fig. 2). 

The modifications shown by Rickard represent acceptance of both the transgressive nature of the 

Onondaga-Marcellus boundary as discussed by 

Oliver (1954, p. 645-646; 1963a, p. 12; later Oliver 

references) ·and the consequent .suggestion by Dennison (1961, p. 2, 10) that the upper limit of the 

Onesquethaw should be redefined as the top of the 

Tioga Bentonite Bed. These modifications had the 

result of placing the Seneea Member of the Onondaga Limestone in western New York in the lower 

Cazenovia Stage with time equivalent shale units 

to the east. 

Rickard subdivided the Onesquethaw Stage into 

two stages, the Sawkill and the Southwood, to emphasize the lithic and f.aunal break between the 

Schoharie Grit and equivalent formations and the 

Onondaga Limestone. Earlier, Boucot (1959, p. 737-

739) had .suggested an Esopus Stage for the Esopus 

and equivalent formations (lower Onesquethaw) 

and had informally used "Schoharie" and "Onondaga" in a stage or substage sense. The threefold 

(Boucot) or twofold (Rickard) division of the 

Onesquethaw Stage is logical, but in this report the 

units are simply termed lower, middle, and upper 

Onesquethaw. 

Changes in the correlation of the New York stages 

with the European standard are shown in figure 2. 

The position of the Lower~ Middle and Middle-Upper 

Devonian boundaries have .most recently been discussed by Oliver and others (1968, p. 1032-1035), 

and these correlations are used here. The bearing of 

the colonial rugose corals on problems of intercontinental correlation is discussed in a later section. 

Figure 3 summarizes the stratigraphic nomenclature for the area under consideration. Only those 

rock units that ~are important in the present work 

are included. A more detailed correlation chart is in 

Oliver and others, 1968 and 1969. 

EASTERN NORTH AMERICA 

Eastern North America, as here designated1 is 

that half of the present continent that was east of 

the Devoni·an Transcontinental arch (Eardley, 1951, 

p. 27-28) . During Ea,rly and Middle Devonian time, 

the arch was important in separating an eastern 

from ·a western faunal province or realm. This will 

be more fully discussed below. 

Cooper and others (1942) Rickard· ( 1964) I ...... 

--1 Oliver and others 

(1968) r 

Taghanic Tully Limestone Frasnian Taghanic Tully Limestone 

Moscow Moscow 

Formation a. Formation 

Givetian Tioughnioga a. Tioughnioga. ::J Givetian ::J 

~ Ludlowville ~ Ludlowville 

Formation <-' <.!) Formation c c Givetian 

~ ~ Skaneateles E Skaneateles 

E Formation ro Formation 

Cazenovia I 

ro Cazenovia I -?---

Marcellus Marcellus 

Eifel ian Formation r-::~ Formation 

1---? _...;;:: --?---

tfiJ"ga Bentonite Bed 

Eifel ian Southwood of Seneca Member Onondaga Limestone Eifel ian 

Onondaga Limestone 3: 

ro 1-.v .r; 

a; 

Coblenzian Onesquethaw Schoharie Grit ::J Schoharie Formation Ems·ian rr 

If) Emsian (]) 

Sawkill c Ca,rlisle _ct~nter 0 --?---

Esopus Shale Esopus Formation 

Siegenian --?---

Deer park Oriskany Sandstone Deerpark Oriskany Sandstone Siegenian 

.t IGURE ~.-uevelopment of stage nomenclature in the New York standard sequence. The terminology of Oliver and other s 

(1968) is followed in this paper. 
REGIONAL STRATIGRAPHY 5 

That part of Eastern North America from which 

colonial rugose corals are herein described or discussed is shown in figure 1. Localities and areas 

mentioned in the text are identified on this map or 

on the more specialized maps that follow. 

REGIONAL STRATIGRAPHY 

APPALACHIAN BASIN 

That part of Eastern North America east of the 

Nashville-Cincinnati-Findl·ay-Algonquin system of 

arches is termed the Appal•achian Basin (fig. 1). The· 

principal coral ass·emblages of this study are from 

the central part of the basin (New York and the 

Niagara Peninsula of Ontario), but a few corals 

from several other Appalachian areas, particularly 

central and northeastern Ohio but including eastcentral Kentucky, western Virginia, and southeastern Quebec, are also compared or described. The general stratigraphy of each area is discussed separately in the following sections to develop the regional 

picture of colonial rugose coral distribution and history. 

Oliver and others (1968, 1969, 1971) reviewed 

the Devonian of the central and southern Appalachian Basin and ·adjacent areas. The correlation· 

charts (1968, fig. 4; 1969) and Onesquethaw Stage 

isopach and lithofacies m·aps (1968, fig. 7; 1971, 

sheet 3) are summaries of much generalized data. 

NEW YORK AND NIAGARA PENINSULA, ONTARIO 

Colonial rugose corals are not known to occur in 

the lower and middle Onesquethaw Esopus and 

Carlisle Center Formations. They :are important 

elements of the fauna in the Schoharie Grit in ea~stern New York, the Bois Blanc Formation in western New York and Ontario (both middle Onesquethaw), and in the Onondaga Limestone (upper 

Onesquethaw and lower Cazenovia) throughout the 

area. No colonial rugose corals are known from the 

lower part (Cazenovian Stage) of the overlying 

H·amilton Group. 

Previous work and present understanding of the 

Schoharie Formation in eastern New York were 

summarized by Oliver (1967a, p. 1-2) and Berdan 

(1971, p. 161-162). Present facies interpret•ations 

and nomenclature were detailed by Goldring and 

Flower (1942) and Johnsen and Southard (1962). 

Colonial rugose corals are known only from the 

Rickard Hill Member of Johnsen and Southard, but 

this is equivalent to the original Schoharie Grit of 

Vanuxem (1842, p. 131) and will be referred to as 

the Schoharie Grit in this paper. 

The Schoharie Grit consists of sandy limestone or 

oalcareous sandstone containing a rich •assemblage 

of brachiopods corals, cephalopods, other mollusks, 

and trilobites. The most comprehensive listing and 

discussion of the Schoharie fauna is that of Grabau 

(1906, p. 181-192, 325-327). Grabau did not mention or list corals although these are numerically 

important at many localities. Colonies of Acinophyllum (A. stokesi) and Asterobillingsa (A. magdisa magdisa) are relatively common in the Schoharie (table 1) , but most specimens are poorly preserved and many are only casts. Cylindrophyllum 

laxtensum is very rare. Schoharie ostracodes were 

recently described by Berdan (1971). 

Oliver (1966a, 1967a, p. 2-5) reviewed previous 

work on Bois Blanc-Onondaga stratigraphy in 

southwestern Ontario. Stauffer (1915) described 

and provided lists of fossils for numerous sections 

in the Niagara Peninsula. He referred to the whole 

sequence as Onondaga but clearly recognized that 

a lower fauna was distinct from that of the Onondaga proper. Cooper (in Cooper and others, 1942, p. 

177 4) and Flower (1943) were both referring to 

beds here classed as Bois Blanc when they discussed "lower Onondaga" faunas with Schoharie 

elements. After Ehlers (1945, p. 34, 89-109) named 

the Bois Blanc Formation for rocks in the Mackinac 

TABLE !.-Geographic distribution of middle Onesquethaw (Bois Blanc-Schoharie) colonial rugose co·rals 

(Key: New York-eastern and western areas (see fig. 5): Ontario-Ningara Peninsula and central southwestern Ontario; Ohio-northeastern; 

Michigan-northern (Mackinac area) ; Ontario-north (Moose River Basin) ; Falls of the Ohio; Virginia-southwestern] 

Acinophyllum simcoense --------------------

A. stokesi -------------------------------- Cylindrophyllum laxtensum ----------------- Asterobillingsa rugosa --------------------- A. magdisa magdisa ------------------------ Grewgiphyllum colligatum ------------------ Heliophyllum procellatum ------------------

H. cassum -------------------------------- H. sp. (phaceloid) -------------------------

H. coalitum -------------------------------

New York 

E w 

X 

X 

X 

? 

X X 

? 

Ontario 

Niagara c 

X X 

X X 

X X 

X X 

X X 

X X 

X 

? 

Ohio Michigan Ontario Falls Virginia of the NE N N Ohio sw 

X 

X X X X X 

? 

? 

?cf. 

X 
6 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

2 3 4 5 6 

Louisville, Kentucky 

Lincoln County, Big Stone Gap, Ohio 

Sylvania, Ohio 

Mackinac area, 

Kentucky Virginia Columbus ~ Marblehead Michigan 

Black shale 

Black shale 

e_o~ Beechwood Limestone ~oyle LimestonE 

~£rrrnMem ber ~Vl 

~Eh·~~~~~~~~ ~:.:J .Silver Creek Limestone . 

· Memberr-,....,....~ 

? 

F. Parasp/nler Zone 

---------

<1> E. Stropheodontid c Zone .B 

Vl ---------

JfOIDIDJt 

<1> 

E 

:.::i 

~ ~--~ ~~ 

·:;: 

c C. Stromatoporoid Boyle 0 

Vl Limestone Zone ~ B. P. pnsma Zone f-~r- ? 

(i; 

A . ...., 

Aemulophyllum Zone 

Upper Silurian 

Olentangy Shale Black shale 

Black shale _,~-.............................................. ..., (Oientangy Shale I 

1/ Prout Tenmile Creek Dolomite 

Traverse 

Group 

If-.,._ Limestone I:"~ of Stewart (1938) / 

of,Stauffer (~90'7,)- '-----....:......-...:-J 1------~ 

1

/ Plum Brook Silica Shale Rogers City Shale of f S Limestone Cooper (1941 ) o tewart (1927) ~~~~~~~~~~~~~~~--------?--------~ 

<1> 

c 

.B 

Vl 

-o 

c 

ro 

(/) 

>. 

,_ ..... J 

Upper , 

~ 

~ 1-L-'M'--L-i d,.._d:-!-1 e__.___..> 

~ 1-'11-'-L..L. ......... ....L.., 

::2 

~ Lower 

Upoer Silurian 

Delaware Limestone 

Tioga Bentonite Bed~ 

H. Parasp/nler Zone 

1---------------

<1> G. c 

.B ----------....... :fl F. Brevtsp/nler Zone-----

E t--------~------

:.::i ./'~ 

~ E. <....._ Detroit River 

.o /-.... Group 

~ -----/7' (Lucas facies) 

o C-D "- u '> ------

A-B 

Upper Silurian 

Dundee Limestone 

a 

::::1 

e 

Lucas Formation ~ 

<1> 

> 

0:: 

....., 

·e 

Dundee 

Limestone 

Lucas 

Formation 

~----------~ w 1-----~ 

Amherstburg 

Formation 

0 ~mherstbuq Formation 

I I 

Bois Blanc 

Formation 

Upper Silurian 

FIGURE 3.-Correlation chart of principal 
7 

Central Southwest 

Ontario 

Black shale 

8 

Niagara Peninsula, 

Ontario 

w 

REGIONAL STRATIGRAPHY 

9 

New York 

Java Formation 

West Falls Formation 

Sonyea Formation 

Genesee Formation 

10 

Series or Stage 

E 

Cohocton 

Frasnian 

Finger Lakes 

c 

.~ 

c 

0 

> 

(I) 

0 

(ij 

a. 

a. 

::J 

7 

r.~rr~~~-----?------------------~~-----1----~r-----------~--~ Ill Tully Limestone JJJ.v--------.--+----

Hamilton Group 

L--1,...1> 

1--L--1.--L--

Taghanic 

Ludlowville Formation 

Tiougnnioga 

a. Centerfiela Limestone Member ::J 

e ~------------~ (.!) 

c 

~ 

E 

Givetian 

Cherry Valley 

f-L-L-L...L....L.....J? Marcellus Shale Limestone Member 

ro 

I Cazenovia !-----?-

""""" 

Dundee Limestone I-·...._._,_ ._?- ._.__,_ .._.__ Union Springs Shale 

1---------- 1-------------,,~-T-io_g_a_B_e_n_t_o_n-it_e_B=e::d::::::.II-__~---------------LSeneca Member ~ 

---? --- Columbu~~ (I) 

c Limestone .8 a. Moorehouse VI 

::J (I) 

e Member E 

(.!) ~ Moorehouse Member 

(ij Lucas ro 

tlD 

> Formation ro 

i:i: "0 

c +-' 0 ·e c 

Q) Clarence Member 0 

0 1--------- Amherstburg 

Formation Edgecliff Member 

Clarence. Nedrow Member 

Memberr-------------------------~ 

Edgecliff Member 

I 

Bois Blanc 

Formation 

/ 

Bois Blanc Formation ",.,-/ 

rrTI~IIIT~~~~~ t-r...,....,-,rT"".,.....,..-r-"1r-r-..--1 

Y Schoharie Grit 

'/c_ . arllsle Center Formation 

Upper 

Middle 

1/J---Es_o_p_u_s--Fo_r_m_a_t-io_n ____ ~ -- -~:;-

11 

Oriskany Sandstone 

~~a. 

~~~ M~~ 

Deerpark 

Helderberg 

~ 

ro 

.s:::. 

Q) 

::J 

r:::r 

VI 

(I) 

c 

0 

Eifel ian 

Em sian 

Siegenian 

Gedinnian 

c 

-~ c 

0 

> 

(I) 

0 

~ 

"0 

:2 

~ 

c 

-~ c 

0 

> 

(I) 

0 

(ij 

~ 

0 

....J 

1 ----------._ . ..:..:.: ne "O 

Rondout Limesto~ .._!_r--------------lf-----------.....l..-----1 

Upper Silurian Coble'skill Limesto~ Cayuga Pridoli 

stratigraphic sections discussed m text. 
8 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Straits region of Michigan, he and others (Caley, 

1947; Ehlers and Stumm, 1951b; Ehlers and others, 

1951; Stumm and others, 1956; Stauffer, 1957; and 

Sanford, 1958) correlated all or part of the New 

York-Ontario Onondaga and Bois Blanc with the 

Michigan Bois Blanc. Oliver (1960a) correlated the 

western New York zone of sm;all Amphigenia (Bois 

Blanc of later reportrs) with the Bois Blanc and 

excluded it from the Onondaga. In addition he suggested that the Bois Blanc was separated from 

overlying formations by an unconformity. These 

correlations are now generally accepted (Boucot and 

Johnson, 1968b; Sanford, 1968). 

The Bois Blanc is a medium-dark-gray, finegrained limestone. Chert is uncommon in New York 

but becomes common west of the Niagara River 

where the formation is much thicker. Within New 

York, the formation is discontinuous, clearly absent 

in many places, and has a maximum thickness 

of 4 feet (1.2 m). Only one exposure is known east 

of the Genesee Valley. 

In Ontario, the formation thickens to 15 feet ( 4.6 

m) at Port Colborne and to at least 24 feet (7.3 m) 

at Hagarsville. It is a continuous unit from not far 

west of the Niagara River to Hagarsville and on 

into the Michigan Basin. In the vicinity of Hagar·sville the lower few feet of the Bois Blanc is very 

sandy. This is the Springvale Sandstone (named. by 

Stauffer, 1913, p. 85) which contains mos.t of the 

same fossil forms as the succeeding limestone beds. 

The Bois Blanc contains a rich and varied coral 

fauna. Among the colonial rugose corals, species of 

Acinophyllum, Asterobillingsa, Cylindrophyllum, 

and Grewgiphyllum are common (table 1), including all of the species known from the Schoharie 

Grit. Other corals were lis.ted by Oliver (1968a, table 

3, p. 740). Other elements of the fauna are abundant 

brachiopods (Boucot and Johnson, 1968b) and, less 

commonly, mollusks, tabulate corals, and trilobites. 

Many identical coral and other specie's are common to both the Schoharie and Bois Blanc. The formations are considered to be time equivalents and 

of Em sian age as discussed below. 

Both the Schoharie Grit and the Bois Blanc Formation are overlain by the Edgecliff Member of the 

Onondaga Limestone. Oliver (1967a) discussed the 

contact and concluded that both lithologic and faunal 

evidence indicated a disconformity between the Bois 

Blanc and Onondaga in western New York. The 

faunal break in Ontario and in eastern New York 

between the Schoharie and Onondaga is similar, and 

it is probable that there is a time gap in these areas 

also. 

Colonial rugose corals are more abundant and 

varied in the Onondaga Limestone than in the underlying formations. This is partly due to the presence 

of more favorable conditions at the time of deposition of the Onondaga, but it also reflects the· fact 

that the Onondaga includes a succession of fauna's 

(Oliver, 1968a, p. 741-742) and probably represents 

a significantly greater amount of time. 

The Onondaga is a complex of limestone facies 

that have been described in several papers (Oliverr 

1954, 1956c, 1962, 1963a, 1966c) which should be 

consulted for detailed descriptions of lithology, local 

successions, and lists of fossils in each unit. The 

general relationships of members and facies are 

shown in figure 4. The units are briefly described below, with emphasis on those that include colonial 

rugose corals. The Onondaga Limestone is exposed 

in a narrow belt that extends some 270 miles ( 432 

km) from Buffalo to Albany, then south and southwest along the Hudson and Neversink Valleys to the 

New Jersey and Pennsylvania borders (fig. 5). 

Corals, especially colonial rugose corals, are uncommon in ~southeastern New York (south of Cab3kill), 

and the Onondaga of this area is not included in the 

following discussion. 

The Edgecliff Member is the lowest unit and is 

present with relatively constant lithology over the 

whole outcrop area from Buffalo to Catskill. In general it is a massive light-gray coarsely crystalline 

limestone characterized by the number and variety 

of included rugose and tabulate corals and brachiopods. Four coral-bearing facies are recognized (figs. 

4, 5). A biostrome (normal) facies occurs from 

Hagarsville to Catskill and contains solitary rugose 

and tabulate corals and brachiopods, but only one 

species of colonial rugose coral (Acinophyllum segregatum) is common, and only a few occur at all 

(table 2). 

Locally, in the Niagara Peninsula and both eastern 

and western New York, but rarely in the central 

area, small bioherms are found (fig. 5). These are 

distinct mound-shaped masses, ranging from 10 to 

70 or more feet (3 to 21 m) in thickness and from 

30 to 1,200 feet (9 to 365 m) in diameter (Oliver, 

1956a). The rock composing the bioherms is coarser 

grained and lighter in color than in the rest of the 

Edgecliff, although "reef cores" can be fine and· 

dark (Oliver, 1966c, p. 38). The bioherms are much 

thicker than surrounding, contemoorary beds and in 

some places project up into or through succeeding 

members. The fauna of the biohe·rm facies includes 

a variety of colonial rugose corals (table 2) in addition to solitary and tabulate corals and brachiopods. The biohermrs are interpreted as representing 

small patch reefs in a shallow-shelf sea. 
REGIONAL STRATIGRAPHY 9 

w 

z 

0 

1- (j) 

79° 

BUFFALO 

SENECA yoga Bentonite Western facies Bed 

w 

::2: MOOREHOUSE Coral facies 

A.-E. bed ...J 

<! 

<!) 

;3 CLARENCE 

z Black chert facies 

0 

z 

0 ~------ FEET METRES 

50 

100 

25 

50 ~w{ <!)Z 

<!0 

01- z(j) 

0~ ---· z_ 

0 oo....~ 

Bioherm 

76° 

Biostrome facies 

Eastern facies 

SYRACUSE 

Tioga Bentonite 

Bed 

NEDROW 

ALBANY 

MOOREHOUSE 

Limestone facies NED ROW 

Fme-grained 

limestone facies 

/1\ EDGECLIFF 

Bioherm 

FIGURE 4.-West-to-east cross section of the Onondaga Limestone in New York showing generalized distribution and relationship of members, facies, and beds discussed in the text. The A.-E. bed· is the Acinophyllum-ETidophyllum bed in the western 

Moorehouse Member. 

TABLE 2.-Geographic distributions of upper Onesquethaw, Edgeclijf assemblage, colonial rugose corals 

[Key: New York~astern, central, and western areas (see fig. 5) ; Ontario-Niagara Peninsula, Woodstock area, Formosa area; Ohi~entral, 

northeastern; Falls of the Ohio-Zones B and C; Illinois-Missouri-Grand Tower area, R. Edgecliff bioherm facies] 

Formation or unit ---------------

Location --------------------·------- E 

Synaptophyllum arundinaceum R 

S. tabulatum --------------- R 

S. kladion ------------------ ------ Acinophyllum stramineum ____ R 

A. segregatum -------------- RX 

A. mclareni ---------------- _____ _ Cylindrophyllum elongatum _ _ R 

C. propinquum ------------- RX 

C. deearium ---------------- R 

C. sp. A ------------------ _____ _ Prismatophyllum prisma __________ _ 

P. ovoideum --------------- _____ _ Asterobillingsa ma,qdisa steorra R 

A. magdisa magdisa --------- _____ _ Eridophyllum subseriale ______ RX 

Grewgiph?Jllum colligatum ________ _ 

Cyathocylindrium opulens ____ R 

C. gemmatum -------------- R 

Heliophyllum megaproliferum _ R 

H. n. sp. B ---------------- R 

H. n. sp. C ---------------- R 

H. coalitum ---------------- R 

H. monticulum -------------- R 

Edgecliff 

New York 

c 

R 

Rx 

R 

Rx 

w Niagara 

R X 

Ror X 

? 

R RX 

Rx Rx 

X 

R 

cf. 

R X 

R 

Ror X 

R 

R R? 

Ror X 

R X 

R RX 

R R 

R R 

R 

R R 

A third Edgecliff facies is found only locally in 

eastern New York (fig. 4). This is a well-bedded 

unit in the lower part of the member and is composed of medium-gray fine-grained lime,stone. It 

Amherstburg 

Ontario 

Wood- For- stock mosa 

? 

X X 

? 

? 

X 

X 

? 

? 

Columbus 

Ohio 

c NE 

X X 

? 

X X 

X X 

X 

X 

X 

B 

X 

? 

X 

X 

Jeffersonville 

Falls 

of the 

Ohio 

c 

X 

X 

Grand 

Tower 

111.-Mo. 

X 

contains a typical Onondaga assemblage of fos.sils, 

but colonial rugose corals are rare or lacking except 

in one local bed where Acinophyllum segregatum 

occurs. Presumably thi·s "dark lime-stone" facie's is 
10 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

EXPLANATION 

1x 0 BIOHERMS 

BIOSTROME 

SOUTHEASTERN. FACIES 

SENTRAL 

SOUTH WESTERN 

ONTARIO 

/,.... ______ _ // 

ADIRONDACKS 

LAKE ONTARIO "1 

n/1 

LondonO 13,14 Mt. rom group) Jl 

X ll (15-21) 

9 4 . 1 . 1

;W

8

1{)lANY/ 

1-~ / I . WESTERN AREA CENTRAL AREA EASTERN AREA/' 

(f) ,.,,.,""/ / 1 --------------------------~, __ y~¥J~N// I 

\ AP.EA/ ~ I 

0 40 80 120 MILES '"'-)(' \ 

I 11 I I 1 1 I I ,,~ // 

0 40 80 120 KILOMETRES ', 

FIGURE 5.-Distribution of Edgecliff facies in New York and 

southwestern Ontario. The dashed lines separate the "areas" 

used in the text. From west to east these are central southwestern Ontario, the Niagara Peninsula, and the western, 

central, eastern, and southeastern areas of New York. Studied bioherms are numbered from east to west; brief descriptions are given in the "Locality Data" section for those 

bioherms from which colonial corals have been described. 

Others are described by Oliver (1956b, p. 17-22). Locality 

equivalent to the normal bioherm facies to both 

the east and west (fig. 3). 

Another local facies of the Edgecliff (and Bois 

Blanc in this case) at Port Colborne, Ontario, consists of some 20 feet (6 m) of argillaceous, silty 

limestone beds with a distinct green color. This unit 

grades upward into typical Edgecliff lithology, and 

the upper part at least grades laterally into a nearby 

Edgecliff bioherm. Several colonies of coJonial rugose 

corals were collected from various positions within 

the unit as follows: 

Upper 16 ft (4.9m): 

Acinophyllum segregatum 

Grewgiphyllum colligatum 

Lower 4 ft (1.2m): 

Acinophyllum stokesi 

A. simcoense 

Asterobillingsa magdisa magdisa 

numbers are as follows: bioherm 1, locality 297; 2, 296; 3, 

248; 4, 244; 6, 81; 7, 329; 8, 285; 9, 135; 11, 185; 13 and 

14, 269; 15, 207; 16, 233; 18, 236; 20, 239; 21, 229; 22, 360; 

23, 597; 24, 438.; 25, Falkirk; 28, 348; 29, 108; 30, C3; 31, 

C5; 32, C7; 33, C9; and 34, C10. The Formosa bioherm 

area was described by Fagerstrom ( 1961). Other bioherm 

areas shown (patterned) are subsurface: Ontario, from 

Sanford (1968); New York from Kreidler (1972). 

These and other corals indicate that the lower 4 

feet (1.2 m) are Bois Blanc in age, whereas the upper 16 feet ( 4.9 m) are Edgecliff. The unit should be 

studied in more detail to determine the reasons for 

its lithologic peculiarities and to see if in any sense 

it fills the time gap between the Bois Blanc and 

Onondaga. 

A fifth Edgecliff facies, found in southeastern 

New York (fig. 5), is composed of a darker and 

finer grained limestone in which corals are uncommon and colonial rugose corals are unknown. 

The Nedrow Member overlies the Edgecliff in 

central and eastern New York and passes westward 

into the Clarence Member which overlies and interfingers with the Edgecliff in western New York. The 

typical Nedrow is an argillaceous limestone at its 

base but grades upward into a more massive, fine-
REGIONAL STRATIGRAPHY 11 

grained limestone so that its lower contact is sharp 

whereas the upper one is gradational. To the east, 

in the Helderberg area, the Nedrow is more massive 

throughout its thickness and the lower contact is 

less abrupt. Corals are common in the Nedrow, but 

are less varied than in the Edgecliff ; colonial rugose 

corals are rare, but Acinophyllum segregatum is 

known to occur. The Nedrow seems to represent 

an influx of fine clastic material, and the base of the 

unit is interpreted as being virtually a time plane 

over its area of outcrop. 

The Nedrow passes westward into the Clarence 

Member (Ozol, 1964; Oliver 1966b, c) in the vicinity 

of Geneva. The Clarence is a cherty limestone ( 45-

75 percent chert) having few corals but including 

Acinophyllum segregatum. At some loooHties the 

Clarence and Edgecliff lithologies are interbedded 

and much of the thickness irregularity of the Edgecliff in western New York and the Niagara Peninsula 

is due to the interfingering of the two members. 

The Moorehouse Member, which overlies both the 

Nedrow and Clarence Members, is the thickest member in all areas and generally contains the most diverse fauna. Brachiopods and mollusks are varied 

and numerous, but both rugose and tabulate corals 

are present or common in most facies. The Moorehouse is thinnest, finest grained, and darkest in 

central New York (central facies, fig. 4). It thickens 

toward both east and wes.t and becomes lighter and 

coarser so that in both eastern and western New 

York it is nearly a repetition of the Edgecliff but 

without the development of biostromes or bioherms. 

Corals, including colonial rugose corals, are common and varied in the eastern and western facies 

but especially so in the west (coral facies, fig. 4), 

although neither the variety nor abundance is as 

great as in the Edgecliff. Species of Synaptophyllum, 

Acinophyllum, and Eridophyllum are present in the 

Moorehouse (table 3), but only Acinophyllum segregatum is generally common or widespread. 

The Acinophyllum-Eridophyllum bed lies within 

the western coral f,acies of the Moorehouse (fig. 4). 

This is a bed of limestone and chert, so·me 20 inches 

(51 em) thick, that extends for 20 or more miles (32 

km) along the outcrop in the vicinity of oLeroy. The 

bed is composed primarily of mass·es of A. segregatum and E. seriale (population B) and few other 

fossils. 

In western and central New York and as far east 

as Cherry Valley, the Moorehouse Member is overlain by the Tioga Bentonite Bed of the Seneca. E-ast 

of Cherry Valley the Tioga has not been recognized 

but may pass into the black shale that overlies the 

Onondaga (Oliver, 1956c, p. 1466-1467). The Tioga 

represents a widespread ash fall that is known 

throughout a large part of the Appalachian Basin 

(see Oliver and others 1968, p. 1019; 1969 for summary) and as far west as Illinois (Meents and 

Swann, 1965). The Tioga is an excellent marker 

horizon and is an important aid to correlation. It is 

accepted here as marking the top of the Onesquethaw Stage. 

The Seneca Member and its Tioga Bentonite Bed 

overlie the Moorehouse in western and central New 

York and as far east as Cherry Valley. Farther east 

the Seneca is unrecognized (thin or missing) , and 

the Moorehouse is overlain by the Union Springs 

Shale Member of the Marcellus Shale. Oliver (1956c, 

p. 1465-1467) has presented evidence that the 

Seneca and Union Springs laterally interfinger and 

are time equivalent. The Seneca is overlain by the 

Union Springs which thins to the west as the Seneca 

thickens so that the Cherry Valley Limestone Member of the Marcellus, overlying the Union Springs 

in central and eastern New York may be the approximate equivalent of the uppermost beds of the 

Onondaga in western New York. 

In central New York, the Seneca Member is a succession of dark, fine-grained limestones with numerous brachiopods and corals representing only a few 

TABLE 3.-Geographic distribution of upper Onesquethaw, Moorehouse assemblage colonial rugose corals 

[Key: New York-southeastern, eastern, central and western areas (see fig. 5); Ohio-central and northeastern] 

SE 

Synaptophyllum kladion ___________________________ _ 

Acinophyllum segregatum -------------------------- X 

A. mclareni --------------------------------------- Cylindrophyllum deearium ------------------------- Prismatophyllum ovoideum _________________________ _ 

P. truncata --------------------------------------- Eridophyllum seriale _____________________________ _ 

E. aulodokum ------------------------------------- E. corniculum ------------------------------------- Cyathocylindrium opulens ---------------------------

C. n. sp. A ----------------------------------------

New York 

E 

X 

X 

? 

c 

X 

Ontado 

w .Niagara c Peninsula 

X X 

X X X 

X 

? 

? 

X X 

X 

X 

X 

Ohio Falls 

NE of the 

Ohio 

X 

X 

? 

cf. 

? 
12 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

species. Only one specimen of a colonial rugose coral 

is known (Prismatophyllum truncata) (table 4). 

Wes.t of Leroy, the member is lighter in color and 

coarser grained and is similar to the underlying 

Moorehouse, but corals are uncommon and no colonial rugose corals have been found. 

The Moorehouse ~and Seneca Members are not 

well exposed in the Niagara Peninsula. They are 

beneath Lake Erie to the east and outcrops are 

spa.rse in the western part of the area where the belt 

underlain by the Onondaga strikes away from Lake 

Erie toward Hagarsville. 

There are few or no outcrops of either the Bois 

Blanc or Onondaga northwest of H.agarsville as far 

as the Woodstock area, where the sequence is sufficiently different to w·arrant separate discussion. 

The various shale and dark limestone members of 

the Marcellus and Skaneateles Formations that succeed the Onondaga and, with the Seneca, compose 

the Cazenovia Stage in this area, contain relatively 

few corals and no known colonial rugose corals. 

TABLE 4.-Geographic distribution of lower Cazenovia (SenecaDelaware) colonial Tugose corals 

Stratigraphic unit ------------ Seneca Famine 

Location --------------------- New York Quebec 

Disphyllum? rectiseptatum 

D? stummi ------------- H eliophyllum sp. cf. H. 

proliferum Hall ------- Prismatophyllum truncata_ X 

CENTRAL OHIO 

X 

X 

"Blue" Dundee limestone 

Michigan Ohio 

X 

X 

The Onesquethaw-Cazenovia sequence in central 

Ohio (fig. 3, column 4) consists of the Columbus and 

Delaware Limestones and the Plum Brook Shale of 

Cooper (1941). In addition, Tillman (1970) has 

suggested that the lower part of the Olentangy Shale 

correlates with the Plum Brook. Colonial rugose 

corals are known only from the Columbus Limestone. Some of the Columbus species are known 

also from New York or Ontario and are discussed 

and (or) illustrated here for that reason. Both 

Eridophyllum and Prismatophyllum are more common and are represented by more species in Ohio 

than in New York. 

The Columbus Limestone crops out in a belt that 

extends from Pickaway County, some 25 miles (40 

km) south of Columbus, north through Columbus 

to Marblehead on Lake Erie. It is well exposed at 

many places and is very fossiliferous. The formation rests unconformably on rocks of Silurian age. 

The top of the Columbus is a widesp·read bone bed 

which may represent a lag concentrate and be 

mainly isochronous throughout the outcrop area 

(Wells, 1944). The Tiog'a Bentonite Bed has been 

identified at only one place in Ohio, near Sandusky 

(Oliver, 1967b; Oliver and others, 1968). It rests 

on the bone bed at the Columbus-Delaware contact 

indicating that the top of the Columbus Limestone 

correlates with the top of the Moorehouse Member 

of New York. The overlying Del a ware Limestone 

includes the Seneca equivalent, but the top of the 

Delaware is probably younger than the top of the 

Seneca at Buffalo. 

The basic biostratigraphic analysis of the Columbus Limestone is that of Stauffer (1909), reviewed 

and interpreted by Wells (1947). Stauffer proposed 

a series of Zones, A to H, based on a combination 

of lithic and biostratigraphic criteria. Zone A is a 

basal conglomerate, but Zones B through H are essentially assemblage zone's. 

Zone B is poor in megafossils and has been variously interpreted, but Ramsey (1969) listed conodonts from this unit and correlated it with the Bois 

Blanc and Schoharie of New York. 

Zone C, the coral zone, is known only in the southern half of the outcrop. It contains several Edgecliff species (table 2) and is correlated with the 

Edgecliff of New York and its equivalents elsewhere. 

The Columbus contains a succession of Eridophyllum forms with E. seriale being characteristic 

of Zone H. This specie1s is limited to the upper 

Moorehouse in New York and, with the Tioga Bentonite Bed, is the basis for correlating these units. 

At this time, the colonial rugose corals provide 

the basis for correlating only Columbus Zones C and 

H with the New York sequence, but these and other 

data indicate that Columbus Zones C through H are 

almost precisely the Edgecliff through Moorehouse 

of New York. 

Only the upper zones of the Columbus can be 

traced to northern Ohio. Zone F, with Brevispirifer 

grega1·ius, is known as far north as Parkertown, 

less than 10 miles (16 km) from Marblehead. Zone 

H extends to Marblehead on Lake Erie, with its 

fauna including Paraspirifer acuminatus and Eridophyllum seriale. Lower zones are replaced to the 

north by a brown, dolomitic facies that should be 

referred to the Detroit River Group. Zone F, in 

northern Ohio, is actually in the Detroit River facies, but its position is m'arked by the persistent B. 

gregarius. If Ramsey (1969) is correct in correlating Columbus Zone B with the Bois Blanc, then it 

is probable that the lower pa.rt of the Columbus in 

the Sandusky-Marblehead area is also of this age. 
REGIONAL STRATIGRAPHY 13 

Zone B at Columbus may represent the southernmost tongue of the Detroit River. My interpretations 

of the Columbus-Sandusky correlations are shown 

in figure 6. 

NORTHEASTERN OHIO 

The Bois Blanc Formation does not crop out in 

Ohio, although lower Columbus Zone B may represent the same time interval as discussed above. 

However, the Bois Blanc is known in the subsurface 

of northern Ohio, in the area east of the Columbus 

outcrop, and is apparently continuous from just 

east of the outcrop in Lorain County to Pennsylvania 

and New York. In Lorain and Lake Counties it is 

succeeded by a "coral biostrome" representing 

Columbus Limestone Zone C or the Edgecliff Member and then by limestones with more or less typical 

Columbus and Delaware lithologies. My knowledge 

of the subsurface paleontology is based on a study 

of the fossils extracted by University of Michigan 

students from three Lorain County cores and one 

Ashtabula County core and now preserved in the 

Museum of Paleontology, University of Michigan 

(unpub. M. S. theses: G. W. Rector, 1950; J. W. 

Parker, 1950; W. J. Truettner, Jr., 1954; and K. R. 

Newman and R. L. Woodhams, 1954). In addition, 

I was permitted to ext~act and study fossils from a 

Lake County core owned by the Diamond Alkali Co. 

All fossils listed in table 5 were studied and identified in my w~ashington laboratory during 1966 and 

1967. 

The Bojs Blanc Formation was first recognized in 

the subsurface of northern Ohio by W. J. Truettner, 

Jr., and by K. R. Newman and R. L. Woodhams 

(both unpub. M. S. theses, Univ. of Michigan 1954). 

Published notices are by Dow (1962) and Janssens 

Chillicothe Columbus Sandusky Windsor Woodstock Buffalo 

Bois Blanc 

FEET 

50 

40 

30 

20 

Lower 

Devonian 

METRES 

25 

20 

15 

10 

5 

FIGURE 6.-Correlation of Onesquethaw sequence in central Ohio with those in southwestern Ontario and western New 

York (location of sections shown in fig. 8). Sections are drawn to approximate vertical scale but to no horizontal scale. 

Time relationships are discussed in the text and indicated here by the dashed lines drawn directly from section to section. 

Capital letters are the Columbus Limestone Zones of Stauffer. Detroit River formations: Amh., Amherstburg; Lu., Lucas; 

And., Anderdon Member of Lucas. 
14 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

(1968, 1970b). Janssens (1970a, p. 21-27) summarized the information available in the Michigan 

theses and the other publications. The Bois Blanc 

of this area is certainly a southern extension of the 

Ontario unit; the subsurface Columbus and Delaware show the pass·age eastward from the outcropping formations to the Ononda~a Limestone. My 

interpretations of the stratigraphy of the northern 

Ohio subsurface sequence are shown in figure 7, 

with general location of the key sections shown in 

figure 8. Corals and other identified fossils from the 

cores are listed in table 5, and the colonial rugose 

corals are included in tables 1, 2, and 3 as appropriate. 

Silica Sandusky Lorain 

County 

Lake 

County 

EAST-CENTRAL KENTUCKY 

The Boyle Limestone crops out around the southwest, south, and southeast sides of the Cincinnati 

arch in Kentucky. Throughout its extent the Boyle 

is mostly Hamilton (early Tioughnioga) in age, but 

locally it includes older units of Onesquethaw age 

(Oliver and others, 1968, p. 1018; 1969, columns 

34-36) . One such unit, on Flax Creek, Broadhead 

quadrangle, Lincoln County, contains a coral fauna 

with mixed Columbus Limestone and Jeffersonville Limestone affinities, including Acinophyllum 

mclareni. The lithology ~f this 15-foot ( 4.6-m) unit . 

is most similar to that of the Columbus Limestone, 

Ashtabula 

County 

Erie,* 

Pa. 

Buffalo 

Plum Brook Shale of Cooper (1941) 

............ ----- ___ // ~~-------------- ...... ---------,...,. 

Dundee 

Limestone 

...:::_ _____ _ 

/ 

Marcellus Shale 

Seneca 

lioga 

Hf'n~~~;·r R0':! 

And. ~·····-_-.?, .- ..e:, ___ _ 

' ->. =-·--·-.... ... ...:::-..:::---

', ... ~ . ........_·----~- .,.l. 

-- ---- -:::~ .......-- ............. 

I • 

Lu. ~. '·, 

Am h. 

. ) .. ..,,.:,::: ·.:· 

.... L·"-·.::··.!·· 

: : ~ •,. : • I • 

I :: .... ~ • I' 

I ;I .•... :In 

METRES 

25 

20 

50 15 

40 

30 10 

20 5 

10 

0 0 

FIGURE 7.-Correlation of Onesquethaw sequences in northern Ohio sections (location of sections shown in fig. 8). Sections are 

drawn to an approximate vertical scale but to no horizontal scale. Time relationships are discussed in the text and indicated here by the dashed lines drawn directly from section to section. Capital letters are the Columbus Limestone zones 

of Stauffer. Detroit River formations: Amh., Amherstburg; Lu., Lucas; And., Anderdon Member of Lucas. Data points 

marked with the asterisk (*) are from L. V. Rickard (written commun., 1969). 
REGIONAL STRATIGRAPHY 15 

TARLE 5.-Summary of paleontologic data from northeastern 

Ohio cores 

[Ranges of key fossils in other areas indicated in parentheses : B, Bois 

Blanc; 0, Onondaga; E, Edgecliff; M, Moorehouse or equivalents] 

County 

Upper Columbus: 

Acinophyllum segregatum (0) 

Cylindrophyllum deearium (E) ___ _ 

Cyathocylindrium opulens (E) ____ _ 

Eridophyllum sp. (M) ------------ Brevispirifer gregarius (M) _____ _ 

Lower Columbus ("biostrome"): Acinophyllum segregatum (0) ____ _ 

A. mclareni (0) -----------------

A. stramineum (E) -------------- Synaptophyllum arundinaceum (E)_ 

"S" grabaui Fagerstrom (E) _____ _ 

Heliophyllum sp. cf. H. halli _____ _ Siphonophrentis sp ______________ _ 

Bois Blanc: 

Acinophyllum simcoense (B) _____ _ 

A. stokesi (B) -----..,------------- Aemulophyllum exiguum (B) _____ _ 

Homalovh1Jllum sp. (B) -----------

Scenophyllum sp. cf. S. conigerum 

(B) -------------------------- Siphonophrentis sp --------------- Amphigenia sp. (small sp.) (B) ___ _ 

Lorain Lake t~~~ia 

X 

X 

X 

? 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

? 

X 

X 

X 

X 

X 

X 

X 

X 

and it seems likely that Columbus denosition was 

originally continuous around the Cincinnati arch. 

The age of the Limestone in this i·solated patch is 

in the Edgecliff-to-Moorehouse range, with some 

suggestion that it may be Edgecliff. 

APPALACHIANS SOUTH OF NEW YORK 

Rocks of Onesquethaw age are widespread in the 

Appalachian Mountains south of New York. These 

are discussed and diagramed by Oliver and others 

(1968, p. 1018-1019 and fig. 7; 1969; 1971 pl. 

3) . A limestone facies is well developed in subsurface areas between the New York and Ohio outcrop, 

but in much of the rest of the area, including the 

outcrop belt, Onesquethaw rocks are clastjc or very 

cherty and extensive col"al faunas are not known. 

Solitary rugose corals are common in parts of the 

section and a few of them were described and illustrated by Kindle (1912, p. 66-67 pl. 2). In addition he listed rugose corals from ·several localities, 

commonly as uzaphrentis" sp. Kindle's "Onondaga" 

is now recognized as including ·rocks of Esopus?, 

Schoharie, •and Onondaga age. 

Kindle's most extensive list of corals, and the only 

one including colonial rugose corals, is that from 

Big Stone Gap, southwestern Virginia (1912, p. 50-

53) . Here a "Grey coralline limestone and interbedded black chert" was considered a coral reef in 

which corals "practically excluded other forms of 

marine life" (p. 52). Dennison (1961, p. 35) briefly 

discussed the unit as the "Onondaga Limestone 

Formation." 

Kindle's unit is part of the southwestern extremity of the Huntersville Chert in which limestone locally exceeds chert in volume. It is also the uppermost part of Miller, Harris, and Roen's (1964) 

Wildcat Valley S.andstone. Corals collected by Dennison and Miller were identified and listed by Oliver 

(1968a, table 3, p. 740) and are clearly of Schoharie 

age. Acinophyllum stokesi is the only identified colonial rugose coral but a second species of Acinophyllum is also present and fragments of a colonial 

z.aphrentid are known. 

QUEBEC 

Rocks of Onesquethaw age are widely distributed 

in New England, Quebec, and the Maritime Provinces (Boucot 1968, p. 89-93; table 6-2, EsopusOnondaga). Rugose cor,als are common, although 

mostly unstudied, but colonial rugose corals are apparently rare. 

The only known lower Onesquethaw colonial 

rugose coral is Asterobillingsa affinis (Billings) redescribed by Oliver (1964) and discussed again in 

the systematic part of this paper. A. a.fjinis is known 

from a single corallum from the upper part of the 

Grande Greve Limestone, on the Gaspe Peninsula. 

Middle Onesquethaw (Schoharie age) corals are 

known from this area (for example, Oliver, 1960b, 

p. 16-20) and are probably widespread, but no colonial rugose corals are known. 

Middle Devonian rock·s and fossil assemblages of 

the Appalachians northeast of New York have been 

reviewed by Boucot and Drapeau (1968). Onesquethaw colonial rugose corals are known from two 

areas: 

1. The Mountain House Wharf Limestone of Boucot and Drapeau crops out on the west side 

of Lake Memphremagog. Corals from this 

formation were identified by Oliver and listed 

and discussed in Boucot and Drapeau (p. 9-

10, 26-28) but, unfortunately, the one colonial rugose coral reported was not included 

in the pubHshed list. This is an Acinophyllum 

represented in the collection by two or more 

crushed and distorted coralla. The specimens 

have the internal characters of A. stramineum 

and A. segregatum, both rest-ricted to the 

Onondaga in New York. The lack of lateral 

supports suggests A. stramineum and an 

Edgecliff age, but the identification is not certain and a. Nedrow to Moorehouse age cannot be ruled out. 
16 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

ONTARIO 

MICHJGAN 

NEW YORK 

--------

PENNSYLVANIA 

INiHANA 

V lRG'lNIA 

--

N.H. 

EXPLANATION 

0 • 

Section locations 

Line of section for 

Fig11re 4 

Figure 6 

L __ _ Figure 7 

------------

0 

I 

0 

I. 

100 

100 

I 

I 

200 

I 

200 

I 

3.00 KILOMETRES 

300 MILES 

I 

FIGURE 8.-Location of sections correlated in figures 6 and 7 and the continuous section shown in figure 4. 

2. The Famine Lilnestone at St. Georges, Quebec, 

was discussed by Boucot and Drapeau (1968, 

p. 19, 22, 32-34, 37) and by Oliver (1971) 

who also described the coral fauna including 

colonial "Cylindrophyllum" stummi (here 

Disphyllum ?) and H eliophyllum sp. cf. H. proliferum Hall (not Nicholson). The Famine 

Limestone is Onondaga in age (Uyeno, Boucot, and Oliver in Boucot and Drapeau, 1968, 

p. 37). Oliver (1971, p. 194-195) cited evidence for a post-Edgecliff, Onondaga age for 

the Famine Limestone based on the corals 

and on study of the brachiopods by A. J. 

Boucot and conodonts by Gilbert Klapper 

(University of Iowa) and T. T. Uyeno (Geological Survey of Canada). According to 

Klapper (written commun., 1968), the conodonts are comparable to specimens from the 

Dundee Limestone of northwestern Ohio. 

Disphyllum? stummi is most similar to forms 

of Cazenovia and Tioughnioga age in the 

Michigan Basin. The earliest of these Disphyllum? spp. occurs in the Dundee. Taken together, these data suggest a lower Cazenovia 

(Seneca, Dundee) position for the Famine 

Limestone. 

MOOSE RIVER BASIN, ONTARIO 

The -stratigraphy of the Devonian rocks in the 

Hudson Bay Lowlands has been summarized by San-
REGIONAL STRATIGRAPHY 17 

ford, Norris, and Bostock (1968), Norris and Sanford (1969), and McGregor, Sanford, and Norris 

(1970). Formations were correlated with the Bois 

Blanc, Onondaga, and Dundee of the central southwestern Ontario sequence. Rugose corals including 

colonial forms were described from a part of this 

sequence by Cranswick and Fritz ( 1'958) . Most of 

the Cranswick and Fritz corals came from "Coral 

Rapids, west bank of the Abitibi River." Both the 

Stopping River Formation (Bois Blanc age) and 

the Kwataboahegan Formation (Edgeeliff age) crop 

out at this locality (Sanford and others, 1968, p. 

30-31), the latter being described as a eoral limestone. The Cranswick and Fritz colleetion seems to 

include some distinct Bois Blanc corals (Oliver, 

1966c, p. 33; 1968a, p. 740, table 3, col. H) and it is 

assumed that the collection include8 corals from 

both formations at Coral Rapids. 

I have reex,amined the Cranswick and Fritz collection, and the colonial rugose corals are discussed 

in the appropriate systematic discussion. Stratigraphic assignment is tentative pending study of 

new collections from known positions within the 

sequence. 

MICHIGAN BASIN 

The Michigan Basin is separated from the Appalachian Basin by the Algonquin and Findlay arches 

and from the Illinois Basin by the Kankakee arch 

(fig. 1). In the following discussions of corals, some 

colonial rugose corals are described or di:seussed 

from central southwestern Ontario, from the 

Mackinac Straits region of the northern southern 

peninsula of Michigan, and from northwestern Ohio. 

The first of these a~reas is on the arch that separates 

the Michigan and Appalachian Basins, but the formation nomenclature and lithofacies are closer to 

those of the Michigan Basin than to those of New 

York or the Niagara Peninsula. The other areas are 

well within the Michigan Basin (fig. 1). In this section, emphasis is placed on those stratigraphic units 

pertinent to the understanding of the colonial rugose 

corals, but in addition, some discus,sion is included 

of those other parts of the sequence important in 

the interpretation of time relationships. 

WOODSTOCK AREA, ONT ARlO 

The general sequence of formations in this area 

is summarized in figure 3, column 7. The Bois 

Blanc Formation is present, as it is over much of 

the basin. The weste·rly thickening trend noted in 

the Niagara Peninsula continues so that the thickness is about 50 ft (15 m) in the Woodstock a.rea 

and more than 400 ft (121 m) in central Michigan 

(Sanford, 1968, fig. 5a). Although the Bois Blanc 

includes cherty limestone and dolomite (Sanford, 

1968_, p. 979-980 and fig. 5a), the corals described 

and discussed herein are from cherty limestone fades (Sanford, 1968, fig. 5a). Throughout the Michigan Basin, the Bois Blanc is accepted as middle 

Onesquethaw, Emsian, pre-Onondaga in age (Oliver, 

1967a; Sanford, 1968, p. 980; Boucot and Johnson, 

1968b, p. 3). 

The Bois Blanc is overlain by the Detroit River 

Group, dominantly a dolomite-evaporite complex 

representing a quite different sedimentary regime 

than most of the carbonates under discussion (Sanford, 1968). However, in central southwestern Ontario, limestone units are present that contain coral 

faunas comparable to those from New York and 

Ohio. Corals occur also in southeastern Michigan 

and adjacent Ontario (Windsor area), but they are 

not discussed in this paper. 

The Detroit River Group in the Woodstock area 

consists of three distinct units. In ascending order 

these are the Amherstburg, Lucas, and "Columbus" 

limestones of Ehlers and Stumm (1951b). Sanford 

(1968) considered the "Columbus" to be only a local 

facies of the Luca~s but it contains a distinct upper 

Columbus Limestone fauna, and I use the term in 

quotation marks to emphasize this relationship. 

The Amherstburg is a light-medium-gray coarsegrained limestone like the Edgecliff, and it contains 

abundant H eliophyllum and Favosites and some (at 

least) typical Edgecliff colonial rugose corals. The 

"Formosa reef" limestone (fig. 5) northwest of 

Woodstock is eonsidered to be of the same age and 

is included in the Amherstburg by Sanford (1968, 

p. 981) and other workers. A few colonial rugose 

corals are described or discussed from both the 

Woodstock and Formosa areas (table 2) . The 

Amherstburg .is correlated with Edgecliff (Oliver, 

1960a; Fagerstrom, 1961; Sanford 1968). 

The Lucas Formation consists of brown limestones 

in the Woodstock area, but it is dolomitic and contains anhydrite over much of the Michigan Basin 

(Sanfo,rd, 1968, fig. 6a). The fauna of the Lucas is 

rather specialized, and no Lucas corals are involved 

in this study. 

The "Columbus" Limestone was first noted in the 

Woodstock area by Ehlers and Stumm (1951b) who 

listed sever'al corals, including three species of colonial rugose corals: Eridophyllum seriale, Hexagonaria (here Prismatophyllum) prisma, and Synaptophyllum sp. (which would now be considered an 

Acinophyllum). On the basis of these and a long list 
18 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

of other fossil corals and brachiopods, Ehlers and 

Stumm assigned this unit to Zone H of the Columbus Limestone. Sanford (1968, p. 985) rejected this 

assignment on the basis of subsurface tracing of 

units and correlates the Lucas and "Columbus" with 

a postulated unconformity separating the Edgecliff 

and Moorehouse Members of the Onondaga in New 

York. My work supports Ehlers and Stumm on this 

point. Eridophyllum seriale is common in the Woodstock area "Columbus" and is elsewhere known only 

from the upper Moorehouse of New York and Columbus Limestone Zone H of Ohio. Preliminary analysis of the "Columbus" brachiopods (J. T. Dutro, 

Jr., oral commun., 1971) and other corals make's 

this correlation compelling. My interpretation of the 

relationships of the Woodstock section to the New 

York and Ohio sequences is shown in figure 6. 

The Detroit River Group is overlain by the Dundee (or Delaware) Limestone. This is principally a 

medium-gray medium-fine-grained limestone in outcrop areas, similar to much of the Moorehouse Member of the Onondaga Limestone in New York and 

the Niagara Peninsula. In outcrop areas the Dundee 

is considered to be of Cazenovia age (Cooper and 

others, 1942, pp. 1754, 1756 and chart; Oliver and 

others, 1969 col. 41). As 'a lithologic unit, the Dundee 

becomes older to the south and east (figs. 6, 7). The 

Tioga Bentonite Bed occurs in the middle part of 

the Dundee (or Delaware) Limestone in the subsurface of Ontario south of Woodstock (Sanford, 

1968, p. 987). To the east this "Dundee" p~sses into 

the Moorehouse and Seneca Members of the Onondaga (fig. 6; and Sanford, 1968, p. 985-987). Sanford (1968) considered the pre- plus post-Tioga 

Dundee and the Moorehouse-Seneca to be younger 

than the Woodstock area "Columbus" (his upper 

Lucas, p. 983-985) leaving a major unconformity in 

the middle of the Onondaga Limestone (8anford, 

1968, fig. 3 and text). My interpretation is based 

on the correlation of the Woodstock area "Columbus" with Columbus Zone H and with the upper 

Moorehouse as discussed above and shown in figures 

6 and 7. 

MACKINAC STRAITS, MICHIGAN 

The Lower and Middle Devonian stratigraphy of 

the northern part of the Lower Peninsula of Michigan has been studied by Ehlers (1945) and summarized by Ehlers, Smith, and Sheldon (1959) and 

Briggs (1959). The general sequence is shown in 

column 6, figure 3. 

The Bois Blanc Formation is 300-400 ft (91-121 

m) thick and quite fossiliferous (Ehlers and others, 

1959, p. 14-15). Corals including several colonial 

rugose corals that were listed or illustrated by 

Ehlers (1945, p. 80-109) are common in the upper 

middle part. Species of Asterobillingsa, Acinophyllum, Cylindrophyllum, Prismatophyllum, and 

zaphrentids are present, but only a few of the species are the same as, or similar to, New YorkOntario species (table 1). 

The Bois Blanc Formation is succeeded by the 

Detroit River Group, the Dundee Limestone, and 

the Rogers City Limestone in that order (fig. 3, col. 

6) . Colonial rugose corals .are not known from the 

Detroit River, but they do occur in the Dundee and 

Rogers City (Ehlers, 1945, p. 114-115). Some of 

these are mentioned or discussed in following sections. 

NORTHWESTERN OHIO-SOUTHEASTERN MICHIGAN 

The general sequence of formations in this area 

is shown in figure 3, column 5. The Bois Blanc Form,ation is missing (Sanford, 1968, fig. 5a and other 

references cited below), but the Detroit River Group 

and Dundee Limestone are comparable to other 

parts of the Michigan Basin. Both contain colonial 

and other rugose corals, and some of these are mentioned in the systematic discussions. 

The Dundee Limestone is overlain by the Silica 

Shale (Stewart, 1927; Ehlers, Stumm, and Kesling, 

1951; Mitchell, 1967; Janssens, 1970b). The lower 

part of this formation (beds 1-6 of Ehlers, Stumm, 

and Kesling 1951, p. 20) has been called the "Blue" 

limestone, which term is used here to distinguish 

these beds from the shaly parts of the formation. 

The "Blue" limestone contains Prismatophyllum 

(discussed below) a:nd Eridophyllum and has previously been included in either th~e Dundee or Silica 

Formations by various workers. It is now generally 

accepted as part of the Silica and considered to be 

significantly younger than the upper Dundee. 

ILLINOIS BASIN 

The Illinois Basin lies west of the Appalachian 

Basin and south of the Michigan Basin (fig. 1). The 

Devonian stratigraphy of this area was recently 

summarized by Collinson and others (1968) and by 

Oliver and others (1968, southern part only). Corals 

from the Falls of the Ohio are important in any 

study of eastern North American Silurian or Devonian corals because of the many early studies that 

have priority over work in other areas. The only 

other part of the basin from which corals have been 

studied for this work is the Grand Tower area of 

Illinois and Missouri. 
AGE OF THE ONESQUETHAW STAGE: 19 

FALLS OF THE OHIO, INDIANA-KENTUCKY 

The sequence of rock units in the vicinity of the 

Falls of the Ohio River at Louisville, Ky., is shown 

in figure 3, column 1. The Jeffersonville Limestone is 

of Onesquethaw age. The overlying Silver Creek 

Limestone Member of the Sellersburg Limestone is 

of Cazenovia age but contains few corals and no 

known colonial rugose corals. 

The Jeffersonville Limestone at the Falls is only 

30 ft (9.1 m) thick, but it is divided into six distinct 

units on a combination of lithology and contained 

fossils. These are assemblage zones, herE! designated 

Zones A-F and named after characteristic fossils, 

some of which are widespread and important in correlation. The present subdivision is based on: (1) 

previous work by many people (fig. 9) ; (2) cooperative fie1d studies by J. T. Dutro, Jr., and myself; 

and (3) laboratory study of the corals. 

Most previous zonation schemes are inadequate 

for detailed stratigraphic study, but the present 

scheme is very close to that of Campbell (1942, 

supplemented by unpub. notes), Oliver (1960a), and 

Perkins (1963). Perkins published the rnost detailed 

description of the Falls rocks, but unfortunately his 

measurements do not agree with those of other 

workers, and the base of his column was apparently not the base of the Jeffersonville Limestone. 

Oliver (1960a) was the first to separate the Lower 

Coral Zone (Zone A) from the Upper Coral Zone 

and to recognize that it is of Bois Blanc-Schoharie 

age. Key corals from this Aemulophyllum exiguum 

Zone were listed by Oliver (1968a, p. "!40, table 3, 

col. F). 

Zone B (Prismatophyllum prisma Zone) and Zone 

C (Stromatoporoid Zone) are correlated with the 

Edgecliff and Amherstburg of New York and Ontario and Columbus Zone C. Zones D, E, and F 

(Brevispirifer gregarius Zone, Stropheodontid Zone, 

and Paraspirifer acuminatus Zone) are correlated 

with the upper Columbus (approximately Zones FH) and the Moorehouse Member. The Falls sequence 

is probably less complete than the Columbus and 

Onondaga sequences and each of the J"effersonville 

zones may be separated by unconformities. 

The indicated correlations of the Jeffersonville 

zones (fig. 3) are based on all available data and, 

except for Zone A, are conventional. The separation 

of Zone A from the rest of the formation is based 

on an analysis of the coral fauna bec-ause few other 

kinds of fossils are found in these beds. The colonial 

rugose corals support the correlations of the higher 

zones, but species common to the Onondaga and J effersonville are not numerous (tables 2-4), and other 

corals and brachiopods are of greater importance. 

Rexroad and Orr (1967, p. 65, 66) recognized two 

conodont faunules in the Jeffersonville. The lower 

faunule (Jeffersonville Zones C and D) is Onondaga 

in age. The upper faunule (upper Zone D through 

Zone F) was suggested to correlate with the Delaware and Dundee (Cazenovia age) because of the 

presence of Icriodus angustus. This correlation is 

unacceptable to Jme because the megafossils of the 

upper Jeffersonville are upper Columbus and upper 

Mooreihous·e in affinities, not Delaware-Dundee. It 

should be noted that Bultynck (1970, pl. 39) reports the /. angustus range in Belgium to be Emsian-early Eifelian. 

GRAND TOWER AREA, ILLINOIS-MISSOURI 

The Grand Tower Limestone crops out in both 

Illinois and Missouri in the area around Grand 

Tower, Ill. (fig. 1) The stratigraphy has been reviewed by Meents and Swann (1965) and by Collinson and others (1968, p. 952-954). Corals, including colonial rugose corals, are abundant and are 

generally similar to Onondaga and Jeffersonville 

forms. They have been listed by various workers 

but, with some few ~exceptions, have not been described or illustrated. One species of Prismatophyllum from the lower Grand Tow·er has been described (Fraunf4~lter, 1966) and is herein discussed 

and reassigned to a lower Jeffersonville species. 

AGE OF THE ONESQUETHAW STAGE 

Correlation of the New York standard sequence 

with the type section in western Europe has been 

discussed by Oliver and others (1968, p. 1032-1035) 

who summarized previous work on the subject. 

The lower Onesquethaw is Siegenian and (or) 

early Emsian in age according to Boucot (in Boucot 

and others, 1970, p. 50). This statement was based 

on study of relatively few brachiopods that could 

be compa.red to European forms of known position 

in the standard stage sequence. 

There is general agreement that the middle Onesquethaw (Schoharie--Bois Blanc) is Emsian, perhaps early Em~dan in age (Boucot and Johnson, 

1968; A. R. Ormiston, written commun.), and that 

most of the upper Onesquethaw (Onondaga) is 

Eifelian (Oliver, 1960a; House, 1962, p. 253, 1968, 

p. 1064; Klapper and Ziegler, 1967, p. 71). 

Edgecliff fossils are strongly provincial (corals 

and brachiopods) or as yet unknown in Europe 

(conodonts, Klapper and Ziegler, 1967, p. 70-71) 

and conceivably could be either Emsian or Eifelian 
<11 

tlD 

Cll 

en 

c 

Cll 

~ > 

(3 

c 

Cll 

2 

w 

c 

Cll 

·v; 

E 

UJ 

r::: 

'- -~ <11 > ~E 

0"0 

....J ::J 

...J 

Campbell This report (see discussion) Lyon (1860) Davis (1887) Bassler (1908) Oliver (1960a) Stumm (1965) Perkins (1963) I 

(1942) Unpublished 

notes 

~ Beechwood 3'6" Encrinital c :J Encrinal Beechwood ..... Cll Sellersburg Beechwood Beechwood .D <l.l·c (/) a.o 

~ 0.> 

::::l<ll a:; Silver Creek 15'6" Hydraulic a Silver Creek Silver Creek Silver Creek Silver Creek (/) 

Paraspmfer Spt"nft"c Spt"rtfer covered P. acum1natus F Zone 5' cultragulatus acum1natus Zone 

bed bed P acum1natus 

Sp1nfer Parasp1nfer I-- ~---- -------- ~-------- ---- Zone 

Bryozoan bed acummafus Zone Fenestrate Zone 

Stropheodontid c Nucleocrmus F bryozoanE Nucleocrmus -~ bed brachiopod Zone 6' c c Weathered clay bed 0 Cll 

> Stropheodonfa Zone 2 with bryozoans <11 "Red clay" a 

~------- bed w ;md ostracodes I-- ?---- ~ 

.-U. I i' "U 

BreviSfJirtfer Cll :2 0\ Brev1spt"nfer Spmfer c D N ~ Turbo bed :J B. greganus Zone 3' E Zone Cll 

~ greganus and Ea ..... B. gregart"us Zone I-- Sjnnfer -ro ·:;: Turbo beds t--------- r----?----- c 0 Zone 0 greganus Coral- u (/) 

D (ii 

~ Stromatoporoid white Zone Stromataporoid a. r--------- a. c ~ Zone 6' Decomposed Zone ::::> 

Cb 

chert= zone C c [/Blue clay 4-6" Ampht"pora 

"Blue clay" (/) ca Zone 

- ----?--- -?-· 

,.(' _______ -------g 

Pnsmatophyllum .D 

pnsma Zone Upper c B black Upper Coral -~ B 4'6" Coral t-----?------ c Zone 4.7' Coral 0 Coral Zone Coral > beds <11 -ro Zone a Coralline -------,_ _____ 0 

/ Aemulophyllum (ii "Ferruginous u Ac Lower Zone Zone ~ - brown A A Zone 4' 0 clay"= zone A Ab Coral I ~- ro ....J 

~ e c Lower Coral zone 4'- Zone 0 0 :J t-----?----

A a ....JU~ 

Louisville (2' exposed) Catent"pora Niagara Louisville Louisville Louisville Louisville 

beds I 

FIGURE 9.-History of nomenclature of the rock sequence at the Falls of the Ohio (Louisville, Ky., and Jeffersonville, Ind.). See text discussion. 

1.'1:) 

0 

z 

0 

z 

(') 

t-< 

00 

1-3 

~ 

~ 

0 

~ 

""d 

~ 

(') 

0 

t"4 

0 

z 

~ 

> t"4 

~ 

q 

C) 

0 

00 

t?=j 

(') 

0 

~ 

> t"4 

00 
BIOSTRATIGRAPHY 21 

in age. Edgecliff corals are of genera and species 

that are common in higher units but uncommon or 

lacking in the underlying Schoharie-Bois Blanc. To 

a lesser extent this is also true of the brachiopods. 

Therefore, it seems most logical to assign the Edgecliff to the Eifelian, at least until more definitive 

data is available. 

The Eifelian-Givetian boundary has been suggested to be as low as the lower or basal Marcellus 

(House, 1962, p. 254). Others have placed the 

boundary above the Marcellus (G. A. Cooper, on the 

basis of brachiopods, and Gilbert Klapper, on the 

basic of conodonts; both oral commun., 1966). I 

'have shown both alternatives in figure 2 and have 

nothing new to contribute on the question. The 

Seneca Member and Delaware Limestone fall into 

this questionable interval and may be either Eifelian 

or Givetian in a.ge. 

'BIOSTRATIGRAPHY 

The known geographic and biostratigraphic distribution of Onesquethaw stauriids., craspedophyllids, and zaphrentids that are described herein is 

summ·arized on tables 1 to 5. Table 6 and figure 10 

show distribution by substages or zones recognized 

on the basis of these and other fos.sils. The separation between the middle and upper Onesquethaw 

as~emblages is clear and has been noted before on 

the basis of rugose corals (Oliver, 1960a, 1968a), 

brachiopods (Boucot and Johnson, 1968b), and conodonts (Klapper and Ziegler, 1967, p. 70-71). The 

less clear sepamtion between faunas of Edgecliff 

and Moorehouse age has been suggested previously 

(Oliver, 1954, 1968a, Klapper and Ziegler, 1967, p. 

70-71) but ha1s not been as thoroughly discussed. 

Recognition of a lower Onesquethaw (Esopus) 

stage or substage was first recommended by Boucot 

(1959) after study of the brachiopod succession in 

the Hudson Valley, New York. Few corals, and only 

one colonial rugoSie coral, have been described or are 

known from rocks of this age in Eastern North 

America. (See Oliver, 1968a, p. 740, for summary). 

The earliest known craspedophyllid, Asterobillingsa 

TABLE 6.-Stratigrapkic ranges of colonial rugose corals 

Asterobillingsa a/finis ---------------------------------- Acinopkyllum simcoense -------------------------------- A. stokesi ____________________________________________ _ 

Cylindrophyllum laxtensum ---------------------------- Heliopkyllum cassum ---------------------------------- H. sp. (phaceloid other than spp. below) -----------------

Asterobillingsa rugosa --------------------------------- H eliophyllum procellatum ____ ""-- _______________________ _ 

Asterobillingsa magdisa magdisa ------------------------ Grewgiphyllum colligatum ------------------------------ Heliophyllum coalitum _______________________ :_ _________ _ 

Acinophyllum vermetum -------------------------------- Synaptophyllum arundinaceum -------------------------- 8. tabula tum _ -------- _____ ------ _____________________ _ 

Acinophyllum stramineum ------------------------------ Cylindrophyllum elongatum ___________________________ _ 

C. propinquum ----------------------------------------

C. Prismatophyllum sp. A ---------------------------------------------- prisma ______________________________ _ 

Asterobillingsa magdisa steor1·a ------------------------ Eridophyllum subseriale ---------- ________ -------------- Cyathocylindrium gemmatum --------------------------- H eliophyllum megaproliferum _________________________ _ 

H. n. sp. B ------------'------------------------------- H. n. sp. C ------------------------------------------- H. monticulum _______________________________________ _ 

Cuatkoculindrium opulens ------------------------------ Pr1'smatoph71llum ovoideum ____________________________ _ 

Sunaptovh11llum kladion _______________________________ _ Acinoph11llum segregatum ___________________ --------- __ 

A. mclareni ------------------------------------------- Cub:ndroPh'l!llum deearium _____________________________ _ 

Eridovhullum seriale _________________________________ _ F:. a.ulodokum -------- _______________________________ _ E. corn1'culu.m _______________________________________ _ r;?,rr.t.hocuUndr1um n. so. A _____________________________ _ Pt·ismntophullum truncata _____________________________ _ 

P. anna ---------------------------------------------- n{c:nh?,llum? sf.ummi ---------------------------------- D.? 1·ediseptatum ---------- ___________________________ _ 

X 

X 

X 

X 

X 

X 

? 

? 

X 

X 

? 

X 

X 

X 

X? 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

? 

cf. 

X 

X 

X 

X 

X 

X 

X 

X 

X 

? 

X 

X 

x? 

X 
l'%j 

c:1' 

t.:rj ~ d ~ ::s :a r:IJ s:lo t'l 

~r:IJ~ 

'1 'C ~ ::s ~ I 

z(l)·rn 

0 r:IJ ft 

::r'<+o: ~-~ 

~ s:~ ss-=- n·m () fD-~ 

s·~ 

zo 

(t) H> 

~~ 

0 ~~ ., 

~~ ~ 

~ ::s 

::s s:lo 

s:lor:IJ 

r:IJ'C 

0 ~ s:: ..... :;.~ 

~ 0 r:IJ H> fig 

::s -

§ 

0'"'' 

::s e. <+ 

~ ., 5·~ ~· 0 r:IJ s:lo(t) 

~ () 

::r'~ (t) ~ 

s:lo-r:IJ 

-

s· g. (t) r:IJ r:IJ () 

., s· o: 

s:lo(t) 

~· s:lo 

s- s· 

(t) :;. ~ ..... ~r:IJ 

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r:IJ ~ 

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~ s: ~ 5- r:IJ (t) 

r:IJ 

g- s· [~ 0 ~ ::s g 0 ., () § gaq 

., (t) ., 

(t) 0 ::s H> () 

cti(JQ 

r:IJ (t) 

..... ~ ::s ., 0 ~ 

g;-

(t) ::r' ., ~ 

~ < .,'< 

(t) 

~ s-(t) 0 r:IJ H>'-' 

I 

co 

0:: 

co 

3-(t) 

aQ 

:YU> 

0 ...... c Ill OQC 

:::T ~­

~ 0.: c IJ) (1Q0 () IJ) ~ 

(t) Ill 

IJ) 

n-o 

0 co ~ 0. 

~0 VJ-o 

Ill :::T ~'< (t)-

n(i 

0 IJ) 3-

3 ~ Oo. 

:::l 

......... () 

0 

0 

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Ill 

"0 

:::T 

CD 

~ 

a. 

IJ) 

:J 

s 

:;>;"" 

:::l 

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::E 

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Gedinnian 

0 

co 

~ "0 

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~ I Esopus 

::TCO"""' 

Ill < ..,., IJ) 0 co ::Jc::E 

0 =·...., -oc CT::JOQ 

~ Ill ~ :::l :::l co "0 0. () 

~ 0. 0 co -· ~ 

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< IJ) co::;;;o;-

~ ~r g 0 ~ ::E ~ 0 :::l 

S,::Jo 

Onesquethaw Cazenovia 

Schoharie 

Onondaga 

. I Moore- I Edgecl1ff house Seneca 

I I I Synaptophyllum 

1-· ------1 5. arund1~aceum I 

5. tabulatum 

1 I I I s. klad10n 

Acinophyllum 

I 

I 1-----1 A. simcoense 

1-----1 A. stokes/ 

L A. stram,ieum I 

l I A. segregatum 

~ 

a· 

c 

(1Q 

:::T 

:::l 

a· 

(1Q 

Ill 

~ 

Ill 

(1Q 

:::T 

Ill 

:::l 

()" 

:;· :;·: ~Ql::T 

g_ CD v;· 

c Ill Ill ()Vl:JQ 

L I - A. mclarehi 

I I 

C. /axtensum Cylindrophjdlum 4----

g.~~­ ~ C elongaJum I ::~:::l 

o(t)rn ~z 

CO)> 0 () . 

c ~ -o.~ IJ) -· 

0.: -· 3 (t) IJ) (t) 

0 :::l 0 ..... o ..... rn ...... 

z ::o;-3" )> :::l -· ......... 0 ~-

::E~ 

:::l 

t=j C. propm~uum I 

C.sp. A 

~-+----- C. deear/Jm 

I 

1-----+----- Prismatophyllum 

P. ovo1deum I 

---l P truncatum 

~ 1------1 A. affi171s As terobdlingsa 

I 

Siegenian 

Lower Devonian 

1--·-.J----1 A. rugosa 

A. magdisa maqdisa I . I I I A. magdisa steorra . I 

Grew~iphy/lum I 1 

G. colligatum I 

...._ __ ....o1 £ b . , Endophyllum .- . su sena1e I ~ £. seriale. 

- £. aulodokum I - £. conjunctum 

-1£. cornicu{um 

r Cy]thocylindnum 

C. gemmatum I 

1------+-----l C. opulens I 

C. n. sp. A , I 

Heliophyllum 

H. megap~oliferum· I} ~----t H. n. sp. f! .

1 

phacelo1d 

~--....o1 H. n. sp. C 

-- H. coalifu1 I} ceno1d 

·-.J /f. procellatum I I} 1-----1 H. cassu~ astraeo1d 

H. mont1c(um I I 

Em sian E·t ,. T I 1 e 1an ·-.J ·-.J I I 

Middle Devonian 

Givetian 

S~VROO ~SODilR ~VINO~OO HdROWI~SXONON 

VJ)> 

,..,., I tii3z Ill :;· c (1Q co 0 

i;OQ ~.co~-~ IJ)co -Ql()::T ~ :::l Ill 

O.:::l 

(/) 

~ 

)> 

c 

::0 

0 

)> 

rn 

() 

-< 

!:: 

z 

0 

::0 

0 

-o 

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-< 

r 

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(/) 

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0 

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-o 

I 

-< 

r 

r 

z 

)> 

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)> 

-o 

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::0 

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z 

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rn 

Frasnian 

Upper 

Devonian 

VJ·-o rn ,..... COC Ill Ill~ 

~ ::J9 

C/l 

~ c;;· 

IJ) 

66 
PALEOBIOGEOGRAPHY 23 

a/finis, is of this age. It seems safe to assume that 

early Onesquethaw rugose corals were much more 

diverse than the present record indicates and that 

the middle-upper Onesquethaw record of craspedophyllids and zaphrentid colonial corals was preceeded by a period during which the various genera 

or morphologic types were established. Corals are 

common in several facies of Helderberg age (Oliver, 

1960b, c, 1968a) but stauriids, craspedophyllids, and 

zaphrentids are lacking. Deerpark and lower Onesquethaw corals are rare. Either the precursors of 

the known craspedophyllids and zaphrentids evolved 

during this time in areas where no record has been 

preserved or recognized, or they entered Eastern 

North America during this time from another 

area. 

A middle Onesquethaw substage or zone is clearly defined by the colonial rugose corals and by other 

studied groups. All known cylindrophyllinid genera 

are present (not all are known in New York or Ontario) although species diversity is· not as great as 

in the succeeding substage. The Craspedophyllinae 

are represented by Grewgiphyllum, which may be 

an evolutionary link between the cylindrophyllinid 

Prismatophyllum and the craspedophyllinid Eridophyllum, a;s discussed in the systematic section. The 

first colonial zaphrentids also appear here and both 

phaceloid and astraeoid species of H eliophyllum are 

known. 

The geographic source areas for the middle Onesquethaw corals described or discussed in this paper 

are shown in figure 11. The middle Onesquethaw 

coral assemblage was reviewed in :somewhat greater 

detail by Oliver (1968a, p. 740-741 and table 3). It 

is dominated by endemic families and genera and is 

notable for its homog·eneity, as many species are 

represented in a variety of lithologies over a large 

area. Associated brachiopods are more widespread 

than the corals (Boucot and Johnson, 1968a). 

The upper Onesquethaw substage is characterized 

by two distinct coral assemblages that are here 

termed the Edgecliff ass·emblage and the Moorehouse assemblage. In New York the former is found 

in the Edgecliff Member and the lower Clarence 

Member. The Nedrow and upper Clarence Members 

have relatively few corals and are not characterized 

by either .assemblage. The Moorehouse Member contains the Moorehouse assemblage, especially in 

coarser facies; the fine-grained medium-gray Moorehouse of central New York lacks most of the characteristic Moorehouse colonial species (table 3). 

The Edgecliff assemblage represents the time of 

greatest diversity of rugose corals in Eastern North 

America. A favorable environment was widespread, 

and coral bioherms and bios·traomes were developed 

over a large area (figs. 5, 12). The distinct Edgecliff assemblage i:s known from the Edgecliff Member in New York and Ontario, from the Amherstburg Limestone in Ontario, from the Columbus 

Limestone, Zone C, in Ohio 1 and from the Jeffersonville Limestone, Zones B and C, in Indiana-Kentucky. 

It occurs also in th·e central Mississippi Valley, in 

both Illinois and Missouri, although it is largely 

undescribed. It is generally absent from the central 

Michigan Basin where the Detroit River facies were 

generally inimical to this assemblage of corals. 

All of the Onesquethaw colonial genera and 18 

of the 33 New York-Ontario species are present in 

the Edgecliff assemblage; 13 to 15 species are restricted to the assemblage (table 6 and fig. 10). The 

assemblage is less homogeneous on the species level 

than the preceding Bois Blanc assemblage, and many 

colonial species remain to be described from Ohio, 

Indiana-Kentucky, the Mississippi Valley, and probably other areas. 

The Moorehouse assemblage is almost as widely 

distributed as the Edgecliff one (fig. 13). It includes 

most, but not all, of the Edgecliff genera and a combination of species that are either holdovers from 

the Edgecliff, limited to the Moorehouse and equivalent units, or that range into later units. Only four 

Moorehouse colonial rugose coral species are limited 

to this assemblage, but additional forms remain to 

be studied from Ohio, Indiana-Kentucky, and other 

areas. 

The Onesquethaw Stage is succeeded by the 

Cazenovia Stage with relatively few colonial rugose 

corals in the study area. Only one species is known 

from New York and this probably occurs in the 

Moorehouse assemblage in Ohio. Additional forms 

are known to occur in the Michigan Basin and other 

areas but are not covered in this ·study. 

The generic and specific ranges given in figure 10 

are based on the presently available information but 

will certainly require some modifications as corals 

from additional parts of Eastern North America 

are studied. The craspedophyllid ranges are most 

reliable, the zaphrentid ranges least so. All are described and discussed more ·extensively in the appropriate sections on distribution, which are included in the systematic descriptions. 

PALEOBIOGEOGRAPHY 

A definitive discussion of Onesquethaw paleobiogeography must necessarily be based on all available information, not just on three families of 
24 

I 

/ 

/ 

/ 

/ 

/ 

/ 

/ 

/ 

0NTARIO 

NONCYSTIMORPH COLONIAL RUGOSE CORALS 

HUDSON BAY/) 

QUEBEC 

Coral x Rapids 1 

I 

C A \N A D A 

J-_/' 

,,J-, 

( I \ NEW 

\\ St. I \BRUNSWICK __ eorge~

1 , 

~ ,fLake ~ ' / 

) 

I 

'- '~VJ_ 

\ _ Memphremago(\ MAINE 

LAKE ~~STATESl /~yo ) ) I 

~'\ I 

'\.) NEW I VT./ 1 

YORK 'UN.H. Syracuse 

o Albany __ 

B f • o I SS indsor l{.lB u falo Helderberg 1 MA · B B ______ __Mts. ---

1.}~-I{ Erie ' lco~N. 1 \ Port Jerv1s 

Marblehead 

1

1 

ILLINOIS OHIO I PENNSYLVANIA 

\ II , :INDIANA 1 o It 

,

7 < "1_ Columrb_5J' .___ L_::-_~-M-D 

)Falls ot ' (.J J ...... · \. G ./the Ohio --._ ~ \../,., WEST ,./ ' 

\ T~~~..r-..,...-v-.Louisville \ VIRGINI~-" 

~'I KENTUCKy fiax Cree:,\__/"'-' VIRGINIA 

./ 

EXPLANATION 

~,-----._ J------ ~-f!ig,Ston_<: C!a_p ______ _ • 

)_______ T~~N:~E-L:rr""/ NORTH CAROLINA ~ 

Middle Onesquethaw coral locality 

X 

Lower Onesquethaw coral locality 

---"\__ 

goo 80° 70° 

0 100 200 300 400 MILES 

I I I 

I 

I I 

I 

I 

I I 

0 100 200 300 400 KILOMETRES 

FIGURE H.-Distribution of studied lower and middle Onesquethaw corals. 

corals, and such a discussion is beyond the scope orf 

this paper. However, a brief discussion is desirable 

as distribution in ti,me and SJpac~e of the genera being desc.ribed can be better understood in the light 

of what is known of the paleogeography. 

An Early and Middle Devonian Appalachian or 

Eastern N orlh American province has long been recognized ; in this pa:per I call it the Eastern North 
50° 

40° 

I 

/ 

/ / 

goo 

/ 

/ 

/ 

/ 

/ 

/ 

ONTARIO 

PALEOBIOGEOGRAPHY 

70° 

HUDSON BAY$; 

QUEBEC: 

I 

I 

c A IN A D A 

goo 

( 

\ 

\ 

0 100 200 300 400 MILES I I I I I I I I I I I 

0 100 200 300 400 KILOMETRES 

70° 

FIGURE 12.-Distribution of studied upper Onesquethaw, Edgecliff assemblage corals. 

25 

50° 

40° 

American (ENA) Province of the Eastern Americas Realm (fig. 14). Boucot and Johnson (1967, 

1968a) and Boucot, Johnson, and Talent (1968, 

1969) have summarized Early Devonian brachiopod 

distributions and presented paleogeographic maps 

based on the known distribution of various groups 

of fossils and rock types for as many as five Early 

Devonian time intervals, including both early and 
26 

I 

/ 

) 

( 

" 

/ 

/ 

/ 

/ 

/ 

/ 

/ 

/ 

ONTARIO 

NONCYSTIMORPH COLONIAL RUGOSE CORALS 

HUDSON BAYj) 

QUEBEC 

Coral x 

Rapids 1 

I 

c A IN A D A 

EXPLANATION 

Moorehouse age • coral locality 

Seneca age • coral locality 

0 100 200 300 400 MILES I I I I I I I I I I 

0 100 200 300 400 KILOMETRES 

FIGURE 13.-Distribution of studied upper Onesquethaw, Moorehouse assemblage corals. General localities of Seneca- (Cazenovia) age corals are also shown. 

and middle Onesquethaw. These provide ,a,n excellent fr.am,ework in which to discuss Early Devonian 

coral distribution. 

Sparssky (1967) and Dubatolov and Spa.ssky 

(1970) tabulated the distribution af Devoni~an cortal 

genera for the world and presented generalized 
PALEOBIOGEOGRAPHY 27 

maps of Early, Middle, and Late Devonian biogeography on which they clearly distinguished the 

"Appalachian" (ENA) province during the first 

two epochs. 

Hill (1957) presented the clearest statement on 

the nature and distinctness of EN A coral provincialism during Onesquethaw time. She called the 

Emsian-Givetian fauna the Heliophyllum fauna and 

noted its extension into Africa during the Middle 

Devonian. The contemporary faunas of the rest of 

the world (including western North America) were 

termed "Eurasaustralian" (Old World of Boucot 

and this paper), and Hill noted that regional differences within this Old World fauna during Emsian, Eifelian, or Givetian time were small in comparison with the differences between this fauna and 

the ENA fauna. Because of revisions in age and 

generic assignments, many of Hill's temporal distribution data are no longer useful (Hill, 1957, p. 

47-48), but the recognized faunal differences are 

sharper now than when her paper was written. 

Oliver ( 1968a, 1973, 1975) summarized the succession of rugose coral f.aunas in the Lower and Middle Devonian of Eastern North America. The 

Onesquethaw faunas were reviewed (1968a p. 740-

7 42; 1975) and lists were given ( 1968a, tables 3 

and 4; 1975, tables 2 and 3) of species of known 

stratigraphic position (their position having been 

determined partly on the basis of the families here 

described) ; many of the 1968 and 1973 data are 

updated by information tabulated in this paper. 

Early Onesquethaw paleogeography was mapped 

and discussed by Boucot and Johnson (1967, p. 49-

50, fig. 4). The earliest craspedophyllid, and the 

only known one of this age, is from the Gaspe area 

(locations A-B on the Boucot and Johnson map). 

Middle Onesquethaw paleogeography was also 

mapped and discussed by Boucot and Johnson (1967, 

p. 50-51, fig. 5, 1968a, p. 1264, fig. 3). The known 

distribution of the middle Onesquethaw coral assemblage (fig. 11; Oliver, 1975, fig. 2) is not as 

great as that of the brachiopods. Corals of this age 

are known from Maine (Oliver, 1960b, p. 16-20) 

and from the Llano region, Texas (unpub. data) 

but are few, simple, and not certainly related to 

species of the assemblage from the area of abundanc·e. No colonial corals of this age are known from 

south of Missouri-Illinois- Kentucky or from Quebec 

and New England (fig. 11). 

Upper Onesquethaw coral assemblages have approximately the same known distribution (figs. 12, 

13) as the middle Onesquethaw assemblage. This is 

the "Appalachian * * * upper Emsian to Eifelian" 

of Boucot, Johnson, and Talent (1968, p. 1249, :fig. 

4; 1969, p. 24-26, fig. 4) who indicate that brachiopods of this age are uncommon south or west of 

Illinois-Missouri or north of New York. Upper 

Onesquethaw corals, including some genera typical 

of the Edgecliff and Moorehouse assemblages are 

present in southeastern Quebec. The craspedophyllid 

Acinophyllum sp. cf. A. stramineum, associated with 

typical EN A solitary H eliophyllum? and Siphonophrentis in the Mountain House Wharf Limestone, 

Lake Memphremagog, Quebec (fig. 12), is upper 

Onesquethaw, and probably Edgecliff, in age. Colonial Heliophyllum and solitary Heliophyllum, Heterophrentis, and Siphonophrentis in the Famine 

Limestone, St. Georg·es, Quebec, are EN A forms of 

post-Edgecliff Onondaga age (see above discussion) 

and are tentatively assigned to the Seneca assemblage (table 4 and fig. 13). Colonial Disphyllum? 

in the Famine, represents the only known occurrence 

of a disphyllid within the Appalachian Basin. 

During Emsian-illiddle Devonian time, EN A 

province corals wer ~ known from parts of northern 

South Am·erica, northwestern Africa, and Spain. 

The South America.a EN A corals were from the 

Cafio Grande and Cafio del Oeste Formations in the 

Rio Cachiri area, State of Zulia, northwestern 

Venezuela. Two species were initially described by 

Weisbord ( 1926), and these were redescribed with 

two ·additional forms by Wells (1943). Acinophyllum vermetum (Weisbord) (redescribed below) 

is very close to A. stramineum (Billings) of Edgecliff age. In addition, H eliophyllum halli MilneEd wards and Haime, H eterophrentis venezuelensis 

(Weisbord), and Zonophyllum sp. were described. 

All but Zonophyllum are distinct ENA types and, 

with associated brachiopods (Liddle, 1943, p. 14-

17), indicate an Onesquethaw age. The whole fauna 

has recently been listed •and reviewed by W eisbord 

(1968, p. 217-222) who suggested again a late 

Early Devonian to Middle Devonian age range. 

Typical H eliophyllum halli suggests a Middle Devonian age, and the similarity of A. vermetum to 

Edgecliff Acinophyllum suggests an Edgecliff age. 

More definite age assignment must await study of 

more and better specimens. 

The EN A affinities of northern South American 

Devonian faunal assemblages are discussed by Harrington (1968), Krommelbein (1968), Weisbord 

(1968), and Boucot, Johnson and Talent (1968, 

1969). These authors indicate that "Appalachian" 

(ENA) elements extend as far south as centralwestern Bolivia but that the fauna was "pure" ENA 

only at the northern end of its distribution area and 
28 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

EXPLANATION 

LAND AREAS RUGOSAN GENERA 

• 

Eastern American 

Old Red Facies 

Other 
PALEOBIOGEOGRAPHY 29 

became progressively diluted with other elements 

to the south (Harrington, 1968, p. 664; Boucot and 

others, 1968, p. 1251, 1969, p. 27-28). 

Eastern North American corals have been described from Morocco by Le Maitre (1947) and 

from Mauritania by Cottreau (1941). ENA forms 

are also present in Spanish Sahara collections now 

in the U.S. National Museum. All are Middle Devonian in age, but some or most are Givetian and are 

younger than the Onesquethaw corals described 

here. 

The Moroccan corals (LeMaitre, 1947) are the 

most important because they were so well described 

and illustrated ~and because the Moroccan assemblage includes more ENA forms than any other 

known African ·assemblage. LeM·aitre illustrated 

typical H eliophyllum halli, Siphonophrentis gig antea, and Eridophyllum. I have examined her specimens (in Lille) and agree with her EN A identifica ... 

tions except that her EriaojJhyllum is probably not 

E. seriale. However, it is a typical Eridophyllum 

and the only known example of this genus from outside of Eastern North America. The three Moroccan 

species are typical Hamilton forms (Tioughnioga 

Stage) and would be considered Givetian if collected 

in North America. 

Cottreau (1941) described "Zaphrentis" sp. cf. 

"Z." gigantea and H eliophyllum halli from northern Mauritania. No sections were illustrated, but 

photographs of the H. halli exterior appear typical 

for that species and there is no reason to question 

either identification. Assodated corals and brachiopods indicate a Middle Devonian age. 

H eliophyllum halli also occurs in the Spanish 

Sahara along with Siphonophrentis sp. and H eterophrentis sp., all identified from collections sent to 

the U.S. National Museum by the Richfield Oil Co. 

FIGURE 14.--.-World distribution of coral genera characteristic 

~ of the Eastern Americas Realm. Eastern American genera 

are those known only from Eastern North America, 

northern South America, northwest Africa, and Spain. 

Old World genera are those that were widespread in the 

Old World Realm and absent in the Eastern Americas 

Realm except in. the Givetian of the Michigan Basin. (See 

text discussion.) The paleogeographic base is the bestfit map of Bullard, Everett, and Smith (1945; used by 

permission). Land areas from various sources. Possible 

migration routes (arrows) to Africa would be Emsian, 

Eifelian, or Givetian; routes to the Hudson Bay and 

Michigan Basin areas were Givetian. The Eastern Americas Realm included Eastern North America and northern 

South America. Western North America. Africa, and 

Europe were parts of the Old World Realm during Emsian and Middle Devonian time. 

and by Pan American Hispano Oil Co. The col1ections are probably of Eifelian age. Arden and Rehrig 

(1964, p. 1522) list Heterophrentis prolifica (Billings) ·and Zaphrentis phrygia (Rafinesque and 

Clifford) among other corals from Spanish Sahara 

as of Givetian age. These are EN A species of Eifelian age but identification has not been confirmed. 

In all of the African collections, EN A forms are 

associated with more numerous and varied Old 

World genera and species. Brachiopod assemblages 

are similar mixtures with EN A elements forming 

a small part of dominantly Old World assemblages 

(Boucot, Johnson, and Talent, 1969, p. 35). 

Altevogt (1968, and unpub. dissert., Munster) 

and Y. M. Cheng (unpub. dissert., Munster) have 

described typical EN A H eliophyllum and Siphonophrentis from the northern coast of Asturia, Spain. 

During a visit to Munster (1969) I was able to confirm the similarity of some of the Spanish forms to 

EN A species. I disagree, however, with some of the 

other identifications of specimens described and 

illustrated by Altevogt (1968). Scenophyllum, 

Kionelasma, and Heliophylloides, all typical ENA 

genera we·re misidentified, and his identification of 

Bethanyphyllum is uncertain. The fauna is dominated by Acanthophyllum, Disphyllum, and other 

Old World genera. 

The Middle Devonian paleobiogeography of the 

ENA coral fauna is summarized in figure 14. The 

distribution of upper Onesquethaw to Taghanic 

coral as~semblages is generalized and the boundaries 

of a reasonably "pure" Eastern Americas Realm 

are indicated. The western m'argin was the transcontinental arch of Eardley (1951) which apparently form·ed an unbroken barrier throughout Onesquethaw time. Old World genera entered Michigan 

and northern Indiana in middle or late Cazenovia 

time (Miami Bend Formation of Cooper and Phelan, 

1966; Indiana) and at various times during the 

Tioughnioga (Traverse Group, Michigan), but only 

one is known to have reached the Appalachian 

Basin (Disphyllum? stummi, Famine Limestone, 

Quebec). 

The nature of the northern or northeastern boundary is unknown. EN A corals are known from the 

southern Hudson Bay Lowland, but the Arctic 

Islands corals are Old World. Possible connections 

with Europe along the northwest side of the Old 

Red Continent see·m unlikely. 

The Old Red Continent formed the eastern side 

of the province and realm, but the size and extent 

of this land area to the south is a matter of speculation. It may well have extended farther than is 
30 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

shown in figure 14 and could have been continuous 

at times. 

Eastern North American faunas apparently migrated south to northern Venezuela and beyond, but 

in central South Americ·a they mixed with other 

forms that did not reach Eastern North America. 

EN A corals and other animals may have entered 

North Africa around the southern end of the Old 

Red Continent where they intermixed with Old 

World elements that did not reach Eastern North 

America. EN A elements moved as far east or north 

as Spain but seem to have formed progressively 

smaller percentages of the fauna as they moved 

north. More data on all assembleges concerned are 

needed before the migrations and distribution patterns can be described in any detail or understood. 

PREVIOUS SYSTEMATIC WORK 

Previous descriptions of Onesquethaw colonial 

rugose corals from New York and Ontario are those 

of Hall (1843), Billings (1859), Nicholson (1874, 

1875), Simpson (1900), Lambe (1899, 1901), 

Ehlers and Stumm (1953), Stumm (1954, 1955), 

and McLaren (1959). The later papers redescribed 

specimens collected by earlier geologists, none o.f 

whom was working within a stratigraphic framework adequate by today's standards. 

In the Niagara Peninsula, "Onondaga" or "Corniferous" corals may have been from either the Bois 

Blanc or Onondaga. In western New York the Bois 

Blanc is thin so that most "Onondaga" corals are 

Onondaga but ·some (for example, from Leroy) are 

almost certainly Bois Blanc and others may be. In 

central and eastern New York where the Bois Blanc 

is lacking, the Schoharie was recognized by the 

early geologists so there is less of a problem in making the distinction between a Lower and Middle 

Devonian source. 

Some early collectors in New York distinguished 

the "Onondaga" ( = Edgecliff) from "Corniferous" 

( =Nedrow and Moorehouse), and many distinguished the Seneca in centr1al New York. Nevertheless, most museum specimens are simply labeled 

"Onondaga" which means various things depending 

on loc-ality, collector, and date of collection. 

Insofar as it is practical to do so, the following 

descriptions are based on new collections from 

known stratigraphic positions. However, study of 

many of the rare or uncommon forms is based on ~all 

available material, and it has been necessary to try 

to determine the source of many old collections that 

contain type material or representatives of uncommon species. Procedure used in determining the 

source of important specimens from old collections 

is discussed in the ap'Propriate part of the systematic description. In general this involves one or more 

of the following: (1) knowledge of what is exposed 

at the source locality, (2) lithology of matrix or filling, ( 3) preservation characteristics, and ( 4) strar 

tigraphic position of other specimens of the taxon 

in the same area. 

Many of the New York-Ontario taxa are assigned 

to species originally described from : ( 1) the Falls 

of the Ohio, Indiana-Kentucky, (2) the Columbus 

Limestone, Ohio, and ( 3) the drift in the vicinity 

of Ann Arbor, Mich. Early collections from these 

and other areas pose the same problems of stratigraphic interpretation as do the New York and Ontario ones. 

Falls of the Ohio corals have recently been revised 

by S.tumm (1965) who briefly reviewed earlier work 

on Falls coral,s. Rominger (1876), Hall (1877, 1882, 

1883b, 1884), Davis (1887), Greene (1898-1906), 

and numerous other works dating from 1820 to 

1958 prec-eded Stumm and established approximately 900 nominal species based on type specimens from 

the Falls or its immediate vicinity. Stumm (1965) 

did much to reduce the nomenclatorial maze by 

synonymizing so·me two-thirds of the "species" and 

presenting brief descriptions of 291 species that 

he found acceptable. However, the stratigraphic 

confusion remained. The fossiliferous limestone at 

the Falls is less than 50 ft (15 m) thick but it includes rocks (and corals) of early Ludlow (Berry 

and Boucot, 1970, p. 180-181), Emsian, Eifelian, 

and Givetian age (figs. 3, 9). Stumm and some 

earlier workers sepa,mted the Silurian and Givetian 

corals with greater or lesser success, but none of the 

descriptive paleontology has been done within a 

really acceptable stratigraphic framework. In this 

report Falls specimens are assigned to stratigraphic 

units on the basis of all available information. Study 

of bed-by-bed collections made by Guy Campbell 

(later presented to the U.S. National Museum) and 

me, has indicated the probable source of many 

forms. Through the kindness of the late Prof. 

Stumm, I have a copy of the index pages to Davis' 

own copy of his 1887 Kentucky Fossil Corals (Davis, 

1887). Davis annotated his copy with information 

on source position, but this is for his "species," not 

for individual ·specimens,· and m•any indicated 

"ranges" are too great to be o.f help. 

Columbus Limestone corals were described by 

Stewart (1938), who also reviewed the sparse previ-
PREVIOUS SYSTEMATIC WORK 31 

ous work. Her descriptions were based on old collections as well as her own and some of the described species may not actually occur in Ohio or in 

the Columbus Limestone. Some, however, are well 

located in Stauffer's zone sequence (see &tl'latigraphy 

discussion), and most are clearly from the Columbus. 

Because the parts of the Columbus that include 

corals a;re all upper Onesquethaw, the problem of 

stratigraphic assignment is reduced. However, the 

Columbus does include a sequence of five or six 

zones, most of which have a characteristic asse·mblage of corals. Columbus corals referred to in this 

work are assigned to stratigraphic position on the 

basis of Stew·art's information or the position of the 

species as learned from study of my own collections. 

Rominger (1876) described many corals, including several colonial rugose corals, from "the drift 

of Ann Arbor" (Michigan). These specimens could 

have been derived from the Mackinaw Straits region, Michigan (Bois Blanc Formation), or from 

southwestern Ontario or southeastern Michigan 

(Bois Blanc, Amherstburg, or Onondaga Formations). Until the coral assemblages of these areas 

are described, the source of many of Rominger's 

specimens will rema.in speculative. Rominger taxa 

referred to in this work are tentatively assigned to 

stratigraphic positions on the basis of the known 

position of similar s.pecimens elsewhere. 

Specific problems of stratigraphic source of type 

and other specimens referred to are fully discussed 

in the systematic section. 

CORAL MORPHOLOGY 

Insofar as is practical, morphological terms are 

used as defined in the Treatise (Moore, Hill, and 

Wells, 1956, p. F245-F251). Additional terms or 

usage modifications are explained in this section or 

where used. Terminology for increase is after Hill 

(1935, p. 491-492; 1956b, p. F244-F245) and Oliver 

(1968b, p. 19-21). 

MICROSTRUCTURE 

Microstructure is consistent with each of the three 

principal families of corals studied. The three species of Synaptophyllum have the same structure, 

described under the generic heading, but it is not 

known whether this is characteristic of the family. 

The many craspedophyllid species have a common microstructure, here termed craspedophyllid 

to contrast with disphyllid structure. This structure 

is described under the family heading. Briefly, it 

consists of fine monacanths at a relatively high angle 

to the horizontal. Each carina, or part of septum between adjacent carinae, is composed of numerous 

closely spaced parallel trabeculae. 

The colonial zaphrentids have craspedophyllid 

microstructure, but in species of H eliophyllum this 

is combined with coarser monacanthine trabeculae 

that develop locally. In such forms, craspedophyllid 

structure predominates, but some carinae, or parts 

of septa between carinae, are a single, coarse monacanth. This is described in more detail in the family 

description. 

CARINAE 

Carinae are flangelike elevations on the sides of 

septa. Where present they a.re a fundamental part of 

the septa and have the same microstructure. They 

may be e;ither a thickened trabecula or be composed 

of several trabeculae, but in any case they are parallel to the trabeculae and so indicate growth direction even if the fine structure of the septum is not 

preserved. 

The orientation of carinae varies with the coral 

and particularly with the shape of the calice and the 

distal edge of the s·eptum. In the corals under study, 

carinae tend to be vertical or at a high angle in the 

outer and middle parts of dissepimentaria (pl. 84, 

fig. 3). In the outer disseptimentarium they angle 

away from the axis of corallites having a reflected 

calice platform (pl. 62, figs. 1-3) and form a broad 

fan pattern that should not be confused with or compared to the tight trabecular fans in the Phillipsastraeidae. Within the inner dissepimentarium they 

bend toward the axis and commonly are horizontal or 

at a low angle at the margin of the tabularium (pl. 

42, figs. 3-5). Carinae are best developed in, and 

commonly limited to, the dissepimentarium but are 

present and even well developed within tabularia of 

som·e species (pl. 38, figs. 8-11) where they tend to 

deflect upward and be almost vertical (pl. 38, figs. 

4, 9). 

The appearance of carinae in transverse sections 

depends to a marked extent on their orientation. At 

the inner margin of the dissepimentarium, carinae 

may be almost parallel to the section and appear distorted as a result (pl. 93, fig. 3). 

Carinae are on both sides of septa and either alternate in position or are opposite each other. Based on 

appearance in transverse section, these are termed 

zigzag and yard-arm carinae, but all gradations from 

one to the other exist. 

Zigzag carinae may quite precisely alternate in 

position (pl. 43, fig. 11) or be only slightly offset (pl. 

82, fig. 6). They may be sharp ridges on the septal 

faces (pl. 61, fig. 4) or may be little more than crenu-
32 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

lations of the septum (pl. 43, fig. 11). They may be 

strongly or weakly developed and all of these variations can occur within a single corallite and on one 

septum although there tends to be a characteristic 

pattern for each species. Zigzag carinae are the most 

common type in all of the craspedophyllid genera and 

in the zaphrentid Cyathocylindrium, although in 

most species carinae that approach the true yardarm type, occur. 

Yard-arm carinae tend to extend farther from the 

septal surface than do zigzag carinae and ideally are 

truly opposite each other (pl. 93, fig. 4). Commonly 

however, either the septum or the carina is slightly 

offset (pl. 94, fig. 9). To aid in the description of the 

various species, the term "su·byard-arm" is used for 

carinae that rise abruptly from the septal surface 

but that are slightly offset. Yard-arm carinae are 

characteristic of H eliophyllum, but they also occur in 

most of the other craspedophyllid and zaphrentid 

genera. 

It is striking and .important that the craspedophyllid and zaphrentid corals of the Eastern Americas 

Realm are characteristically carinate and, in spite of 

the noted variations, no coralla and few corallites are 

completely noncarinate. This is very much in contrast to the Old World disphyllids and phillipsastraeiids in which carinae tend to be poorly developed 

or absent. In many (most?) examples the Old World 

"carinae" are only the relief on the septal faces, 

caused by the coarse monacanths that form the s~epta. 

Some Old World cyathophyllids (Birenheide, 1963) 

have craspedophyllid-type carinae, and their distinction from the zaphrentids is difficult. (See discussion in systematic section.) 

INTRACOLONY AND INTERCOLONY 

VARIATION 

Oliver (1968b) discussed general aspects of variation within and between coral colonies (1968b, p. 

23-25) , basing most of his arguments on studies of 

some of the same colonial rugose corals as described 

herein (1968b, p. 26-31). The 1968 data have been 

modified only slightly (tables 8-13, 16-39) and not 

so as to affect the general conclusions. The following 

nomenclatural changes have been made: (1) Billingsastraea rugosa of 1968 is Asterobillingsa magdisa 

of this paper, but the sa,mple has been subdivided 

(subspecies magdisa and steorra) and a few specimens added so that the 1968 data are replaced by 

more refined data in tables 20 and 21. (2) Eridophyllum seriale n. subsp. of 1968 is E. seriale population B of this paper. (3) E. n. sp. of 1968, is E. 

corniculum of this paper. 

Variation of each of the species is described in the 

systematic part of this paper. Where practical, measurements and septal counts are analyzed and tabulated (tables 8-13, 16-39). The data in this paper 

are more ~extensive and cover more species than the 

1968 data. 

Variation in Acinophyllum segregatum is described in detail because the analysis of this species 

has provided a standard by which other species can 

be evaluated. Some biologic and stratigraphic conclusions that I consider important were based initially on this form. In a following section, results of the 

analysis of other species are tabulated and discussed 

in comparison with A. segregatum. 

VARIATION IN ACINOPHYLLUM SEGREGATUM 

Acinophyllum segregatum is more common and occurs in more lithofacies than any other Onesquethaw 

colonial coral. Its general morphology makes it ideal 

for analysis; corallites are parallel and sufficiently 

small and closely spaced that a large number of well 

oriented individuals can be studied in one transverse 

section. Data are from one or more transverse sections in 58 colonies. The analyzed colonies, collected 

by the author from the Onondaga Limestone in New 

York, include most of those considered suitable for 

study. Colonies or parts of colonies that were broken 

or distorted before or during burial and those too 

poorly preserved were rejected, as were fragments 

of colonies too small to provide data on 20 or more 

individuals. This rej-ected material, however, added 

to the distribution data and provided additional 

qualitative information on variation. 

The number of major septa (n) and diameter (d) 

have been analyzed; in each colony, 20 to 60 corallites 

were measured, and corresponding septal counts 

were made. In a g.iven thin or polished section, all 

adequately preserved individuals were measured. 

This inevitably included a few immature ·specimens 

(conical stage) which could not be so identified. 

However, study of many colonies shows that only a 

few percent of the individuals in a given transverse 

section would be immature, and the only statistic 

that is significantly affected by their inclusion is the 

observed range of diameter~ Results of the study are 

shown in tables 7 to 15 and on the scatter and bar 

diagrams (figs. 15-21). 

TERMINOLOGY 

The following characters are quantitatively 

treated in the present study: Diameter (d) , measured to the nearest 0.1 mm by occular micrometer in 

a section through the colony which intersects the 
INTRACOLONY AND INTERCOLONY VARIATION 33 

parallel corallites at right angles to their axes; septal number (n), number of major septa, counted in 

the same section as that in which corresponding diameter was measured. The septal ratio (n: d) is of 

aid in analyzing these characters. 

The following standard statistical terms (the symbols for which also appear in tables 7-39) are used; 

methods are adapted from Imbrie (1956), to which 

reference can be made for further discussions: 

Mean (x) arithmetic average of character. 

Observed range (ORx), minimum and maximum 

values of given character in given sample. 

Standard deviation (sx), measure of variation 

from sample mean in units of character being 

analyzed. 

Coefficient of variation ( V x) , staJ!dard deviation 

expressed as a percentage of the mean; a dimensionless number useful in comparing variation 

in analogous characters of differing numerical 

range. 

Correlation coefficient (r), measure of the closeness to a straight line of points plotted on a 

scaUer diagram; the coefficient may vary frbm 

one to zero, one indicating a straight-line plot 

and zero indicating lack of any correlation. 

P, the probability of the valuer being not significant-a P of 0.01 indicates a probability of significance at the 99 percent level, a P of 0.10 

indicates probability at the 90 percent level, and 

so forth; in the present work, Pis derived from 

tables in Simpson, Roe, and Lewontin (1960, p. 

426) and Fisher (1946, p. 174). 

INDIVIDUAL VARIATION 

Mean values for n and d based on 20 to 60 corallites and standard deviations and coefficients of 

variation for n and d were determined for all 58 

colonies (table 7). Data on 32 (of the 58) colonies, 

selected w.ithin stratigraphic groupings, were further analyzed to determine the n : d correlation coefficient ( r) . Of thes·e 32 colonies, 12 (see asterisks, 

table 7) were selected to illustrate extremes in n, d, 

and n: d correlation (scatter diagrams, fig. 15A-D). 

Variation between colonies (intercolony) was 

analyzed for the same factors as variation within 

colonies (intracolony), but each colony was treated 

as an individual. Intercolony mean values are thus 

mean intracolony means, and intercolony coefficients 

of variation are V's of colony means, ·and so ~erth. 

Tables 8 to 13 and the scatter diagram (fig. 16) 

show the variation in the 58 colony means. 

Intracolony variation in diameter is best indicated 

by the scatter diagrams (fig. 15A-D) and the coefficients of variation shown in table 7 (summarized in 

tables 8 to 13). Colony coefficients of diameter variation range for all 58 colonies from 5.8 to 15.9 and 

average 10.0. Intercolony size variation, based on 58 

pairs of colony means, is illustrated in figure 16; 

variation between colonies (Va=20.7, table 8) is approximately twice as great as that between individuals within colonies. 

Septal number variation in A. segregatum is 

shown on the scatter diagram (fig. 16) and tables 7 

to 13. Coefficients of intracolony variation are low 

(ranging from 1.9 to 8.7, mean value 4.4) and intercolony variation is approximately tw.ice as great 

(Vn=1l.O; table 8). 

Normal distribution of the 58 coefficients of septal 

number variation (Vn) is shown in figure 17. Some 

of the data for the 32 colonies for which the n: d 

correlation coefficient (r) was calculated are given 

.in table 14, with the colonies in the order of increasing Vn. This shows the distribution of septal number 

in the 32 colonies, and the lack of correlation between 

Vn and P. 

At the top of t3!ble 14 are two colonies in which at 

least 85 percent of the sampled corallites have the 

same number of major septa. Tw·enty-three colonies 

have at least 70 percent of their corallites in two septal number classes. At the other extreme, the bottom 

three colonies .in table 14, have strikingly high coefficients of variation and show bimodal septal number 

variation. 

From these data I have previously (1968b, p. 26-

29) suggested that septal number in this species may 

have been genetically controlled within rather narrow limits. If the "normal" colony was genetically 

programed for a septal number range of three, sexually reproduced protopolyps would display normal 

genetic variation in this character, but asexually 

reproduced polyps in each colony would be restricted 

to the narrow range of their colonies' protopolyps 

genotype. This, at least, would explain the very restricted intracolony variation accompanying the 

broad intercolony variation in septal number and 

other characters. 

Broader intracolony variation, especially if bimodal, suggests that the analyzed colony may have 

included two or more genotypes. This could have resulted from a genetic accident (somatic mutation) or 

more likely from a mechanical one such as intergrowth of two or more adjacent colonies. To test this 

possibility, colony USNM 162631 (tables 7 and 14 

and fig. 15B) was analyzed. The original sample of 

20 corallites showed bimodal distribution (table 14, 

upper line of data). Another sample of 85 corallites 
34 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

35r---------~----------~----------~---------,-----------.----------.----------..-. 

A 

30 

c:: 25 

20 

15 

2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 

d(mm) 

30r-----------~------------~------------~-----------,-------------r------------, 

8 

25 

20 X 

Colony 6 (USNM 162631) 

r =0.68; P<0.01 

15~----------_.------------~------------~----------~------------~----------~ 2.0 3.0 4.0 5.0 6.0 7.0 

d (mm) 

FIGURE 15.-Number of major septa (n) and diameter (d) in samples of selected colonies of Acinophyllum seg're.qatum. Each 

polygon encloses data from one colony The line d=0.43n-4.0 represents the intercolony mean (see fig. 16) and is included 

here as a basis for comparison of the different colony samples. A, Four Edgecliff bioherm colonies. B, The colony shown 

in figure 18 and discussed in the text as having bimodal distribution of septal number; the line n=22 separates the two 

modes. C, Four Edgecliff colonies from the biostrome and fine-grained facies. D, Three Moorehouse colonies. 
INTRACOLONY AND INTERCOLONY VARIATION 35 

35r------------r-----------.------------.------------.------------~----------~----------~ 

30 c o.9 

--"f ------------ + 4=".......... 0· ~0~ - ',o~ 

-----+ + $$t+ +* + _,., ~ 

_,.,---+ + ++ ++ ---- ti..:.... _______ t____________ --"' Coiony 9 (USNM 162618) 

~ 25 r = 0.52; P = 0.01 

20 

------og-, ---c;-~ 0 8 f:<Bcy ---- <:5 / Colony 57 (USNM 172246) ,.,::::._Q._O-G-_._--'« o§o /;/ 

---,:- • : ~'\ ,.,"" r=O.ll;P=<O.Ol «t...... ~--·= : .. : : 0 0 ~,.,., 

-~--::;::;:. - ---Colony 51 (USNM 162642) 

--"'.(xx~ - I r =0.45; P=0.05 

.(xx~ ~~ 1 \ x!~~~-~/ Colony 29 (USNM 16.2620) 

15 r = 0.32; P~ 0.20 

2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 

d (mm) 

30 ~----------~-----------.r-----------,------------.------------,------------.------------, 

25 

20 

15 

2.0 

D 

Colony 3 (NYSM 12784) 

r = 0.49; P= 0.05 --oo', 

' -- 0 8 ', -- ' ---~ 0 0 0 8 0 ~ 

(c)- o o o ___ o,., .......... 

------- ~--- --- /~ ~ x xx S('fNa:•a;.t-J-::.~-a.-------- / ',1 / // ~X* ~B~ >)..,..-.!!._L __ , ..... 

// x x~ __ >/ Colony 33 (USNM 162649) 

/ --- -~-· L lx J!.----- 1 r:-0.01; P >0.90 

/- Colony 37 (USNM 162652) 

/" r=0.45, P=0.02 

3.0 4.0 5.0 6.0 7.0 8.0 

d (mm) 

9.0 
36 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

'C:: 

,14 L-----...J....------1---..L......-----L..------J 

2.0 3.0 4.0 5.0 6.0 

d(mm) 

EXPLANATION 

X Edgecliff bioherm facies 

!5. Edgecliff nonbioherm facies 

0 Moorehouse western coral facies 

£ Edgecliff fine -grained limestone facies 

• Moorehouse A. -E. bed 

+ Nedrow 

7.0 

FIGURE 16.-Mean number of major septa (fi) and mean diameter (d) in 58 colonies of Acinophyllum segregatum from 

the Onondaga Limestone in New York. Each polygon encloses samples from a different member and facies as discussed in the text. The line d=0.43-4.0 was calculated 

from the total sample of 58 to show how the unit subsamples vary. 

(table 14, lower line of data) taken from a different 

thin section (fig. 18A) showed the same two modes 

(fig. 19). Figure 18B shows the distribution of septal numbers on the section. It seems significant that 

a line can be drawn across the section that almost 

completely separates the corallites of the two modes. 

The pattern strongly suggests that colony USNM 

162631 was formed through intergrowth of two adjacent colonies with dominant septal number ranges 

of 20 to 22 and 22 to 24. 

Intercolony variation in the correlation coefficient 

r is extreme (table 7). Scatter diagrams of selected 

colonies illustrate this (fig. 15A-C). The coefficient 

itself (r) ranges from 0.01 to 0.84, indicating almost 

no n: d correlation in the former and a very high 

probability of correlation significance in the latter. 

P values are greater than 0.50 (low correlation 

probability) in 3 colonies of 32 tested, less than 0.01 

(99 percent probability) in 11 colonies, and less than 

0.05 (95 percent probability) in all but 9 of the 32 

colonies (table 14). 

The lowest correlation is found in colony USNM 

162649 ( r = 0.01, P> 0.90; fig. 15D) in which 22 of 

26 sampled corallites have 20 major septa although 

there js considerable diameter variation (ORa= 4.6 

to 5.8 mm). Other low-correlation colonies are typified by colonies USNM 162620 ( r = 0.32, P = 0.20; 

fig. 15C) and USNM 162635 (r=0.11, P=0.60; fig. 

15A). In both colonies, average n variation is 

coupled with low d variation. 

Intercolony analysis (by colony means) shows a 

stronger correlation between diameter and septal 

number (r= 0.88; P<0.001; table 8) than does any 

one colony. There is a strong correlation in these 

variables in A. segregatum, as in other corals (see 

Oliver, 1968b, p. 29 for discussion and references), 

but the two characters vary almost independently in 

some colonies. 

The lack of n: d correlation within some colonies 

suggests that the commonly held view that septal 

insertion is a response to diameter increase--that is, 

space filling-is untenable. The two processes are 

related, however, as is shown by the high correlation 

in all samples in which genetic variation is possible. 

The correlation means that variation in distance between septa was limited. With the broad septal number and diameter variation found in most species of 

rugose corals, a significant correlation resulted. In a 

clone in which septal number variation was very 

limited, diameter variation undoubtedly was restricted but was sufficient to effectively mask any 

control. In the extreme example (theoretical) in 

which all corallites in a corallum had the same number of septa, any diameter variation would appear to 

be random and uncorrelated. It appears reasonable 

to conclude that septal number was gentically controlled in many rugose corals, and that diameter was 

loosely controlled by septal number. 

GEOGRAPHIC AND STRATIGRAPHIC VARIATION 

Most of the colonies of Acinophyllum segregatum 

in the Onondaga Limestone are found at one of two 
INTRACOLONY AND INTERCOLONY VARIATION 37 

TABLE 7.-Acinophyllum segregatum colony data arranged by stratigraphic unit 

Key to symbols used in tables 7-39; x, character; X', mean value of character; N, sample size (N1, intracolony; N2, i?tercolony~; n, number of major septa (septal number); ORx, observed range; Bx, standard deviation; Vx, coefficient of variation; r, correlatiOn coefficient; P, probability of r not being not significant; d, diameter; d 1, tabularium, diam.:!ter; dau aulos diameter. 

*USNM 

USNM 

USNM 

*USNM 

USNM 

USNM 

*NYSM 

NYSM 

USNM 

USNM 

*USNM 

NYSM 

NYSM 

USNM 

USNM 

USNM 

*USNM 

NYSM 

USNM 

*USNM 

NYSM 

NYSM 

NYSM 

*USNM 

NYSM 

USNM 

NYSM 

USNM 

*USNM 

USNM 

*USNM 

USNM 

USNM 

*NYSM 

NYSM 

NYSM 

USNM 

USNM 

USNM 

USNM 

USNM 

NYSM 

NYSM 

NYSM 

USNM 

USNM 

USNM 

USNM 

NYSM 

NYSM 

USNM 

USNM 

*USNM 

USNM 

USNM 

*USNM 

USNM 

USNM 

Corallum 

162615 

162o29 

162660 

162631 

162632 

162616 

12793 

12794 

162633 

162634 

162635 

12795 

12796 

162636 

162637 

162638 

162617 

12780 

162639 

162618 

12781 

12782 

12783 

172246 

12797 

162640 

12798 

162619 

162620 

162641 

162642 

162643 

162659 

12784 

12803 

12804 

162655 

162622 

162656 

162657 

162658 

12805 

12799 

12800 

162644 

162645 

162646 

162647 

12801 

12802 

162621 

162648 

162649 

162650 

162651 

162652 

162653 

162654 

• Colonies graphed in figure 15A.-D. 

[See text for explanation and discussion. Dimensions are in millimetres] 

N 

43 

24 

50 

20 

32 

20 

20 

20 

25 

25 

25 

30 

23 

20 

30 

23 

30 

20 

22 

28 

20 

21 

23 

30 

28 

30 

30 

30 

25 

30 

20 

23 

20 

20 

20 

30 

20 

30 

30 

23 

60 

27 

40 

25 

40 

20 

30 

30 

30 

30 

30 

30 

26 

30 

30 

30 

28 

20 

26.5 

23.5 

19.4 

22.4 

21.8 

20.1 

18.2 

20.2 

20.0 

22.8 

22.1 

24.5 

21.9 

22.2 

23.1 

19.6 

19.5 

22.1 

20.7 

27.7 

21.2 

21.3 

18.3 

21.3 

19.4 

17.9 

17.0 

16.5 

16.8 

18.0 

20.0 

17.0 

17.8 

21.9 

20.4 

18.8 

19.4 

18.8 

18.8 

18.8 

17.5 

17.5 

ORn Bn 

Edgecliff bioherm facies 

24-30 

22-26 

17-22 

20-24 

20-23 

18-21 

17-20 

18-21 

18-22 

21-25 

20-25 

22-26 

20-23 

21-24 

21-25 

17-21 

18-21 

1.2 

1.0 

1.0 

1.6 

.8 

.7 

.9 

.7 

1.1 

1.3 

1.0 

1.0 

.8 

1.0 

1.0 

1.0 

.8 

4.7 

4.3 

5.2 

7.1 

3.9 

3.6 

4.9 

3.7 

5.4 

5.6 

4.6 

4.0 

3.9 

4.4 

4.2 

5.2 

4.2 

Edgecliff biostrome facies 

19-24 

18-22 

26-29 

20-22 

20-23 

17-20 

19-23 

1.1 

1.0 

.8 

.7 

1.1 

.8 

1.0 

5.1 

4.7 

3.0 

3.3 

5.0 

4.4 

4.8 

6.8 

5.4 

5.2 

4.9 

5.2 

5.0 

3.1 

4.1 

4.6 

5.3 

5.9 

6.8 

5.1 

5.7 

5.7 

4.0 

3.9 

5.6 

5.0 

6.6 

6.0 

4.6 

4.1 

4.9 

Edgecliff fine-grained limestone facies 

18-21 

17-19 

15-18 

15-18 

16-18 

16-19 

19-21 

16-19 

15-20 

0.9 

.4 

1.0 

.7 

.7 

.6 

.4 

.8 

4.5 

2.2 

5.7 

4.1 

4.3 

3.6 

2.0 

4.8 

Nedrow member 

1.6 8.7 

3.4 

4.0 

3.1 

2.8 

3.0 

3.2 

4.0 

3.4 

3.3 

Moorehouse western coral facies 

20-24 

18-22 

18-20 

17-20 

18-20 

18-20 

17-20 

14-20 

16-19. 

1.1 

1.0 

.9 

.9 

.7 

.8 

1.0 

1.2 

.8 

5.1 

4.9 

4.8 

4.5 

4.0 

4.2 

5.5 

7.1 

4.6 

6.7 

4.4 

4.1 

4.6 

4.4 

4.2 

4.0 

3.4 

3.7 

Moorehouse Acinophyllum-Eridophyllum Bed 

19.6 

18.6 . 

19.0 

17.9 

19.1 

19.0 

19.7 

18.9 

20.2 

19.5 

19.8 

19.7 

20.3 

19.1 

19.4 

18.6 

18-20 

17-20 

17-21 

17-19 

17-20 

17-20 

18-2'2 

18-20 

18-22 

18-20 

19-20 

18-21 

19-23 

17-20 

18-21 

16-21 

0.6 

.9 

.8 

.4 

.8 

.9 

.9 

.8 

.8 

.6 

.4 

.7 

.8 

.9 

.8 

1.2 

3.0 

4.6 

4.5 

2.5 

4.2 

4.6 

4.6 

4.4 

4.0 

2.9 

1.9 

3.6 

3.8 

4.8 

4.1 

6.4 

4.7 

4.4 

4.1 

4.0 

4.5 

4.1 

4.7 

4.2 

5.0 

4.3 

5.2 

4.4 

4.8 

4.1 

4.7 

4.4 

4.8-8.2 

4.4-6.2 

4.0-5.9 

4.3-5.8 

4.0-6.2 

3.7-6.2 

2.3-3.8 

3.4-4.8 

3.1-5.3 

4.3-6.2 

4.4-6.8 

5.2-7.9 

3.4-6.8 

4.8-6.5 

4.2-7.3 

2.7-4.9 

2.8-4.7 

4.5-6.3 

3.6-5.9 

4.9-8.0 

4.4-6.7 

3.8-5.3 

3.4-4.9 

3.5-5.5 

2.9-4.1 

3.1-4.6 

2.6-3.7 

2.3-3.4 

2.7-3.4 

2.5-3.9 

3.1-4.8 

2.8-3.9 

2.8-4.0 

4.9-7.8 

3.0-5.5 

3.5-4.8 

3.2-5.3 

3.7-5.3 

3.0-4.0 

2.9-4.9 

2.5-4.7 

2.8-4.4 

3.8-5.2 

3.2-5.4 

3.1-4.8 

3.1-4.5 

3.6-5.5 

3.4-4.6 

3.4-5.3 

3.5-4.9 

3.8-5.8 

3.7-5.0 

4.6-5.8 

3.4-5.5 

4.0-5.6 

3.2-4.8 

3.8-5.3 

3.0-5.2 

Bd 

0.8 

.6 

.3 

.4 

.6 

.5 

.4 

.4 

.6 

.5 

.5 

.6 

.8 

.4 

.6 

.6 

.4 

0.4 

.5 

.7 

.6 

.5 

.4 

.5 

0.3 

.4 

.3 

.2 

.2 

.3 

.4 

.3 

0.3 

0.8 

.6 

.3 

.4 

.4 

.5 

.6 

.4 

.5 

0.3 

.4 

.4 

.5 

.4 

.3 

.5 

.4 

.4 

.3 

.3 

.5 

.4 

.4 

.4 

.6 

11.6 

10.5 

5.8 

8.0 

10.7 

10.7 

11.8 

10.0 

12.3 

9.3 

8.0 

9.3 

15.9 

7.9 

10.8 

14.1 

11.2 

7.6 

10.1 

10.0 

9.7 

10.2 

8.6 

10.6 

10.1 

9.0 

9.5 

8.8 

6.9 

10.1 

9.6 

8.7 

9.1 

11.5 

14.1 

7.4 

9.7 

9.8 

12.1 

13.9 

11.8 

12.4 

6.4 

10.3 

10.4 

11.8 

9.8 

7.5 

10.1 

9.9 

8.6 

7.9 

5.8 

10.7 

7.2 

9.1 

9.4 

13.2 

p 

0.84 <0.01 

.68 <.01 

.58 <.01 

.45 .05 

.65 <.01 

.11 .60 

.45 .05 

.66 <.01 

.31 .10 

0.56 0.01 

.52 .01 

.55 .02 

.71 <.01 

0.48 4.02 

.46 .02 

.30 .10 

.32 .20 

.45 .05 

0.51 0.05 

0.49 0.05 

.60 .01 

.29 .20 

.70 <.01 

.30 .20 

0.19 0.40 

.47 .02 

.08 .70 

'.62 <.01 

.01 >.90 

.45 .02 

.45 .02 

.45 .05 
38 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

TABLE 8.-Acinophyllum segregatum~ summary data 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 20 60 58 

n: x ------- 16.5 27.7 20.0 

OR ----- 15-18 26-29 16.5-27.7 

s ------- .4 1.6 2.2 

v ------ 1.9 8.7 4.4 11.0 

d: x ------- 2.8 6.8 4.6 

OR ----- 2.3-3.4 6.0-8.9 2.8- 6.8 

s ------- .2 .8 1.0 

v ------ 5.8 15.9 10.0 20.0 

*r ---------- .01 .84 .88 

*P ---------- >.99 <.01 <.001 

*r and P calculated for 32 of the 58 colonies. 

TABLE 9.-Acinophyllum segregatum, summary data, Edgecliff bioherm facies 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 20 50 17 

n: x 18.2 26.5 21.6 

OR ----- 17-20 24-30 18.2-26.5 

s ------- .8 1.6 2.1 

v ------ 3.6 1.7 4.6 9.9 

d: x 3.1 6.8 5.1 

OR ----- 2.3 - 3.8 4.8- 8.2 3.1- 6.8 

s ------- .4 .8 1.0 

v ------ 5.8 15.9 10.5 19.1 

TABLE 10.-Acinophyllum segregatum, summary data, Edgecliff biostrome facies 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 20 30 7 

n: x 18.3 27.7 21.8 

OR ----- 17-20 26-29 18.3-27.7 

s ------- .7 1.1 2.9 

v ------ 3.0 5.1 4.3 13.1 

d: x 4.1 6.6 5.3 

OR ----- 3.4-4.9 4.9- 2.0 4.1- 6.6 

8 ------- .4 .7 .9 

v ------ 7.6 10.6 9.5 16.3 

TABLE 11.-Acinophyll urn segregatum summary data, Edgecliff fine-grained limestone facies 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 20 30 8 

n: x 16.5 20.0 17.8 

OR ----- 15-18 19-21 16.5-20.0 

8 ------- .4 1.0 1.3 

v ------ 2.0 5.7 3.9 7.2 

d: x 2.8 4.0 3.4 

OR ----- 2.3-3.4 3.1- 4.8 2.8- 4.0 

s ------- .2 .4 .4 

v ------ 6.9 10.1 9.1 12.9 

TABLE 12.-Acinophyllum segregatum, summary data, Moo?·ehouse Member (except Acinophyllum-Eridophyllum bed) 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 20 60 9 n: x 17.5 21.9 19.3 

OR ----- 14-20 20-24 17.5-21.9 

s ------- .7 1.2 1.3 

v ------ 4.0 7.1 5.0 6.8 

d: x ------- 3.4 6.7 4.5 

OR ----- 2.5-4.7 4.9- 7.8 3.4- 6.7 s ------- .3 .8 .9 

v ------ 7.4 14.1 11.4 20.6 

TABLE 13.-Acinophyllum segregatum, summary data, Moorehouse Acinophylum-Eridophyllum bed 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 20 40 16 n: x ------- 17.9 20.3 19.3 

OR ----- 17-19 19-23 17.9-20.3 

s ------- .4 1.2 .6 

v ------ 1.9 6.4 4.0 3.2 

d: x ------- 4.0 5.2 4.5 

OR ----- 3.1-4.5 4.6- 5.8 4.0- 5.2 s ------- .3 .6 .4 

v ------ 5.8 13.2 9.3 7.8 

stratigr:ruphic levels: in the Edgecliff Member or in 

the upper part of the Moorehouse Member. These are 

plotted separately on the scatter diagram (fig. 16), 

and distinction is made between colonies from various facies or other subdivisions of the members. 

No evolutionary pattern can be discerned. The 

oldest colonies are those from the Edgecliff finegrained facies; reef and biostrome colonies are (on 

the average) ·slightly younger and the Moorehouse 

colonies are youngest. In none of the characters 

studied is there a progression from older to younger. 

Differences between stratigraphically defined 

groups of colonies are shown on the scatter diagrams 

and summarized in tables 9 to 13. These differences 

relate to the character of the lithology and associated 

fossils (that is, environment) of the stratigraphic 

unit from which each group was collected. In general, the groups with greatest variation are found in 

the light-colored coarse-crystalline facies and are 

associated with a large variety of other corals. 

Groups with least variation are found in medium- to 

dark gray fine-grained lim·estone associated with few 

other corals. 

Sufficient material for analysis has been collected 

from five stratigraphic subdivisions of the Onondaga. These and one other unit are discussed below 

to illustrate the ecological variation. 
INTRACOLONY AND INTERCOLONY VARIATION 39 

TABLE 14.-Data from selected colonies of Acinophyllum segregatum arranged in order of increasing coefficient of septal 

number variation (V n) 

Corallum 

*USNM 162649 

*USNM 162642 

*USNM 162618 

USNM 162650 

USNM 162632 

NYSM 12796 

USNM 162621 

USNM 162622 

USNM 162619 

USNM 162646 

*USNM 162617 

*USNM 16~620 

NYSM 12797 

NYSM 12800 

NYSM 12801 

NYSM 12805 

*USNM 162635 

*USNM 162615 

*USNM 162652 

*USNM 172246 

NYSM 12803 

*NYSM 12793 

NYSM 12782 

*NYSM 12784 

NYSM 12780 

USNM 162638 

USNM 162657 

USNM 162634 

NYSM 12798 

USNM 162654 

*USNM 162631 

USNM 162659 

N P Vn 

26 0.90 1.9 

20 .05 2.0 

28 .01 3.0 

30 .02 3.6 

32 .01 3.9 

23 .05 3.9 

30 .01 4.0 

30 .20 4.0 

30 .10 4.1 

30 .02 4.2 

30 .10 4.2 

25 .20 4.3 

28 .02 4.5 

25 .40 4.6 

30 .70 4.6 

27 .20 4.6 

25 .60 4.6 

43 .01 4.7 

30 .02 4.8 

30 .01 4.8 

20 .01 4.9 

20 .05 4.9 

21 .02 5.0 

21 .05 5.1 

20 .01 5.1 

23 .01 5.2 

23 .01 5.5 

25 .01 5.6 

30 .02 5.7 

20 .05 6.4 

20 .01 7.1 

85 

20 .05 8.7 

* Colonies graphed in figure 15A-D. 

Percentage of 

corallites in 1, 2, or 3 

Number of major septa ( n) per corallite septal classes 

15 16 17 18 19 20 21 22 23 :.::4 :.::5 :.::6 27 28 29 30 1 2 3 

4 22 

1 17 2 

2 7 19 

3 

1 

1 2 18 

11 13 6 

1 16 11 

1 

10 11 

2 

5 

3 

4 

5 

1 12 

3 

3 10 12 

14 10 

11 13 

9 12 

7 5 

8 15 

2 

1 

2 5 12 11 

1 6 

1 1 9 

4 10 4 2 

6 

2 

1 

1 2 5 11 

3 6 7 7 

2 

7 17 

6 10 

8 1 

3 

2 

3 1 

4 14 

9 11 

6 3 

5 7 

5 8 

4 7 

4 

5 

6 

4 

3 

3 

3 

7 

4 8 6 

3 4 12 11 

1 11 2 5 1 

4 3 1 5 

20 24 11 18 

2 3 2 7 3 3 

1 

2 

2 

1 

4 

7 

12 

3 6 16 

1 

8 10 15 7 

3 

85 100 

85 95 100 

3 57 79 89 

63 87 93 

53 75 91 

43 70 96 

60 87 93 

43 80 100 

53 90 97 

47 80 97 

43 80 90 

44 84 100 

43 75 93 

48 76 96 

50 77 87 

44 81 93 

56 72 88 

1 35 58 77 

40 77 93 

37 67 87 

45 75 90 

50 70 90 

33 57 86 

40 65 80 

35 70 90 

48 70 87 

3(} 61 87 

32 56 72 

40 77 90 

55 65 90 

35 60 65 ____ 28 52 65 

-- 35 50 65 

Bioherm and biostrome facies.-Seventeen bioherm colonies were analyzed. Variation in diameter, 

septal number, ·and correlation coefficient is almost 

coextensive with that in the speoies as a whole 

(tables 7 and 9 and fig. 16). The bioherms are composed of coral skeletons (tabulates and solitary and 

colonial rugose) in a matrix of coarse crinoidal debris and are the most favorable environment for 

corals (Oliver, 1954, 1956b, c). 

sis have be,en studied. Most of these would easily fall 

within the scatter area of the :samples in figure 16, 

but a few would fall at and below the lower end of 

this ·area. The additional material indicates even 

greater variation than is shown by tables 9 and 10. 

Colonies with small coralites are uncommon in the 

two facies but do exist. The more restricted scatter 

areas of other stratigraphic groupings all fall completely within the scatter area of the more or less 

Seven colonies were analyzed from the normal coextensive bioherm and biostrome groupings. 

Edgecliff (biostrome facies). The sample is small, 

and it is difficult to discern a significant difference 

between these and the bioherm facies (fig. 16). This 

nonbioherm facies is composed of bedded crinoidal 

debris with tabulate and solitary rugose corals 

being very common, in places forming a coral biostrome (Oliver, 1954, 1956c). Acinophyllum segregatum is not common and other colonial rugose 

corals are rare. 

Collections from both the bioherm and biostrome 

facies of the Edgecliff Member are more extensive 

than indicated. In addition to the analyzed colonies, 

numerous fragments inadequate for statistical analyFine-grained facies.-A localized facies of the lower Edgecliff Member was designated the C1 facies by 

Oliver (1956c). This is composed of medium-gray 

rather fine-grained limestone. All fossils are less 

common here than in the rest of the member, but 

colonies of Acinophyllum segregatum form small 

local coral beds in several places. Intercolony variation is much less than in the other facies of the Edgecliff Member. Limits of variation are shown on the 

scatter diagram (table 7 and fig. 16). Coefficients of 

variation are 7.2 and 12.9 (table 11), as compared 

with 9.9 and 19.1 in the bioherm group and 11.0 and 

20.7 in the whole Onondaga assemblage studied. 
40 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

16 

14 

~ 12 

z 

0 

_J 

0 10 () 

LL 

0 

0:: 8 w 

(l) 

~ 

~ 6 

4 

2 

0 

2.0 3 .0 4.0 5 .0 6.0 7 .0 8.0 9.0 

VnFIGURE 17.-Normal distribution of intercolony coefficients 

of septal number variation (Vn) in Acinophyllum segregatum. Sample size (N) is 58. 

Moorehouse coral facies.-The western coral facies of the Moorehouse Member (Oliver, 1954) as a 

whole represents a recurrence of conditions very 

much like the Edgecliff. The lithology .is somewhat 

finer and darker, but corals are locally common, 

Acinophyllum segregatum and one other colonial 

rugose coral species compose the AcinophyllumEridophyllum bed (Synaptophyllum-Cylindrophyllum bed of Oliver, 1954). Elsewhere in the Moorehouse A. segregatum is widely scattered but not uncommon; other colonial rugose corals are less common or rare. 

The sc~tter diagram (fig. 16) shows that Moorehouse A. segregaturn colonies outside of the Acinophyllurn-Eridophyllum bed are quite variable. Nine 

colonies analyzed show much of the range of bioherm 

and bioherm colonies. All but one of the colonies are 

from the western coral facies of the member. Intercolony coefficients of variation are 6.8 and 20.6 

(table 12). 

The Acinophyllum coral bed of the western Moorehouse Member represents the most restricted conditions of all. Although enclosed by the "coral facies," 

the 16 analyzed colonies show less intercolony variation than any other group in the formation ( V = 3.2 

and 7.8, table 13). The matrix of this bed consists of 

fine medium-dark limestone and dark chert. AIthough only 12 to 18 inches (30 to 45 em) in thickness, the bed extends for several miles. 

Nedrow and Clarence Members.-Fragments of 

Acinophyllum segregatum have been collected from 

these members at four localities. Three specimens 

have been studied from the western Clarence and 

three from the east-central Nedrow. Data from only 

one of these is included in the statistical analysis 

(table 7), but all specimens collected fall within the 

area of the Edgecliff fine-grained limestone group of 

figure 16. 

Two solitary rugose coral species characterize the 

Nedrow shaly limestone facies but, except for these, 

corals of all types are rare in both members. 

Summary.-Data for the five principal samples of 

A. segregatum are brought together in table 15 and 

figure 20, where the samples are presented in the 

order of decreasing numbers of coral species identified from each facies (diversity index). Table 15 

shows that for these five samples there is a general 

decrease in variation with decrease in diversity. It 

seems clear that the environmental factors that permitted high species diversity also permitted greater 

variation within the studied sample. At one extreme 

is the Edgecliff bioherm facies with high species diversity and high intercolony variation. At the other 

extreme is the Moorehouse Acinophyllum-Eridophyllum bed containing only two species of corals; although colonies are abundant, intercolony variation 

is minimal. 

Figure 20 is based on the same data but shows 

more clearly that intracolony septal number variation is not very different in the five facies, but that 

intercolony variation is high in two facies, medium 

in two, and very low in one. This seems most likely 

to reflect environmental control. 

VARIATION IN OTHER SPECIES 

Study of variation in the other species of colonial 

rugose corals was similar to that of Acinophyllum 

segregatum but limited by smaller sample sizes, the 

restriction of most species to a single stratigraphic 

unit (formation, member, or facies), and by other 

practical difficulties such as the large individual size 

of most of the zaphrentids. Summary data for many 

of the species are given in tables 16 to 39. Part of 

this data is gathered on table 15 and figures 20 and 

21. 

Analysis of other phaceloid species is by the characters and methods outlined for Acinophyllum segregatum except that aulas diameter (dau) was added 

for the Eridophyllum spp. In cerioid species, diameter was measured through the axis at right angles to 
TABLE 15.-lntracolony and intercolony coe'fficients of variation in.24 samples of colonial rugose corals 

Vn Vd or Vdaz 

Sample Species Intracolony ~ntracolony No. Inter- Inter- Nt N2 OR Mean • colony OR Mean colony 

la. Acinophyllum segregatum, Edgecliff bioherm facies __ 20-50 17 3.6- 7.1 4.6 9.9 5.8-15.9 10.5 19.1 

lb. A. segregatum, Edgecliff biostrome facies ----------- 20-30 7 3.0- 5.1 4.3 13.1 7.6-10.6 9.5 16.3 

lc. A. segregatum~ Moorehouse Member except A.-E. bed __ 20-60 9 4.0- 7.1 5.0 6.8 7.4-14.1 11.4 20.6 

ld. A. segregatum, Edgecliff fine-grained limestone facies _ 20-30 8 2.0- 5.7 3.9 7.2 6.9-10.1 9.1 12.9 

le. A. segregatum, Moorehouse, Acinophyllum- 20-40 16 1.9- 6.4 4.0 3.2 5.8-13.2 9.3 7.8 

Eridophyllum bed. 

2. A. stramineum, Edgecliff bioherm facies ------------ 15-22 8 3.3- 6.7 5.5 6.5 7.7-15.6 12.5 14.2 

3. A. simcoense, Bois Blanc Formation ----------------- 10-26 15 2.9- 6.5 4.2 7.4 6.9-15.2 11.1 16.9 

4a. A. stokesi, Bois Blanc Formation -------------------- 10-20 10 2.6- 7.6 4.6 5.6 7.3-13.6 10.1 10.1 

4b. A. stokesi, Jeffersonville Limestone, Zone A ---------- 10-20 6 2.6- 4.6 4.0 5.1 6.2-14.1 9.6 10.9 

5. C11lindrophyllum elongatum, Edgecliff bioherm facies __ 4-16 9 3.7- 7.2 5.1 7.7 3.6-20.0 12.1 12.1 

6a. C. propinquum, Edgecliff bioherm facies (includes 1 4-15 6 1.6- 6.6 3.6 8.2 9.8-22.2 15.6 21.4 

biOstrome facies colony) . 6b. C. propinquum, Columbus Limestone, Zone C _________ 3- 9 5 0- 7.3 4.5 5.5 6.7-15.9 10.8 4.1 

7. C. deearium, Edgecliff bioherm facies (includes two 3-14 9 1.5- 6.9 4.3 5.0 6.7-18.4 12.6 11.4 

biostrome facies colonies) . 8. Prismatophyllum prisma, Jeffersonville Limestone, 5-12 8 2.2- 8.4 5.7 5.6 10.8-25.3 18.7 9.6 

Zone B. 

9. P. ovoideum, Jeffersonville Limestone, Zone C ________ 7-16 7 3.6- 7.1 5.4 7.5 3.3-24.5 16.9 11.7 

10. Asterobillinnsa magdisa magdisa ( Edgecliff bioherm 15-23 6 4.9- 7.8 5.8 11.1 8.7-13.7 6.6 16.1 

and Bois Blanc) • 11. A. magdisa steorra, Edgecliff, bioherm facies --------- 10-36 15 4.4- 9.3 6.3 8.4 8.9-22.0 6.6 11.4 

12. Eridophyllum seriale, pop. B, Moorehouse 

Acinophyllum-Eridophyllum bed. 

4-14 10 3.1- 6.4 4.8 4.2 6.6-15.7 11.4 7.5 

13. E. corniculum, Moorehouse Member, western coral 5-10 11 2.4-14.8 6.7 9.9 5.3-28.9 13.9 21.7 

facies. 

14. E. subseriale, Edgecliff bioherm facies -------------- 6-14 7 3.9-11.5 6.1 5.7 5.8-18.2 12.6 13.6 

15. Grewgiphyllum colligatum, Bois Blanc and Edgecliff 

bioherm facies. 

5-20 10 3.8- 9.1 5.4 6.1 (1) C> ---

16. Cyathocylindrium opulens, Edgecliff bioherm facies ___ 4-20 13 0.0-14.4 7.5 11.9 3.6-24.5 13.7 19.6 

17. Heliophyllum monticulum, Edgecliff bioherm facies ___ 9-10 9 4.0- 7.0 5.7 15.1 10.0-18.9 13.9 15.5 

18. Synaptophryllum arundinaceum, Edgecliff bioherm 

facies. 

12-20 9 2.5- 9.3 5.3 4.8 6.6-17.4 12.5 13.7 

--- t See table 35. 

Vd or Vd•u 

Intracolony Inter- OR M colony 

-------- --- --- -------- --- --- -------- --- --- -------- --- --- -------- --- ---

-------- --- --- -------- --- --- -------- --- --- -------- --- --- -------- --- --- -------- --- ---

-------- --- --- -------- --- ---

-------- --- ---

-------- --- --- 5.6- 7.9 11.4 19.6 

4.1-13.6 14.2 19.4 

15.8-30.6 21.1 15.0 

13.4-30.7 21.1 20.5 

13.7-22.6 17.3 13.2 

10.9-19.4 14.3 18.9 

--------

3.7-11.1 7.3 21.4 

--------

Diversity 

index 

34 

26 

21 

9 

2 

--

--

--

--

--

--

' --

--

--

--

--

--

--

--

--

--

1-4 

z 

1-3 

~ 

> (J 

0 

t"'4 

0 

z 

~ 

> z 

tj 

1-4 

z 

1-3 

t_:l:j 

~ 

0 

0 

~ 

0 

z 

~ 

<: 

> 

~ 1-4 

> 

1-3 

1-4 

0 

z 

~ 

...... 
42 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

one wall; in corallites with two or more markedly 

different diameters, the shorter was recorded. Astraeoid species were analyzed for tabularium diameter (dt) ; an average corallite diameter was obtained 

by measuring the distance between the axes of adjacent corallites (axial distance, da:c), but these 

measurements cannot be related to an individual 

coral so that intracolony correlation of diameter and 

septal numbers cannot be tested in these forms. 

Each sample represented in figures 20 and 21 and 

in table 15 can be considered to show one of three 

patterns, although it is probable that there is some 

distortion due to the small size of some samples. The 

patterns are : 

1. Intercolony Vn being well above the observed 

range of intracolony Vn, including samples 1a, 

1b, and 10. 

2. Intercolony Vn being just above or at high end of 

observed range of intracolony Vn, including 

most of the samples. 

3. Intercolony Vn being low in observed range of 

intracolony Vn, including samples 1e, 8, 12, 14, 

and 18. 

Visual examination of figure 20 suggests that intracolony variation m.ay not significantly differ 

among the studied samples. The 3 patterns a.re mostly a function of the position of intercolony variation 

relative to the common intracolony range of variation, thus patterns 1 and 2 are separated by a Vn 

of 9 to 10 with only one apparent exception, and 

patterns 2 and 3 are separated by a Vn of 5 with 2 

exceptions. 

In analyzing Acinophyllum segregatum, I suggested that these patterns as observed in samples 1a 

to 1e were environmentally controlled. Figure 21 

groups the samples by stratigraphic unit and suggests that ecology is of importance in some of the 

groupings. The only two coral species in the Moorehouse Acinophyllum-Eridophyllum bed (with the 

lowest diversity index of any unit) have very low 

coefficients of intercolony variation. This fits the 

generalization that poor environments are characterized by large numbers of specimens of few species. 

On the other hand, the Edgecliff bioherm and biostrome facies, with the highest diversity indices of 

the five Onondaga facies, have representatives of all 

three patterns. Most biohermal populations show 

medium to high intercolony variation, but some are 

low, and two species (14 and 18) are very low. I suggested in my 1968 paper that these two species may 

simply have been genetically less variable (1968b, 

p. 31). 

Most of the other samples in figure 21 show medium intercolony variation and represent stratigraphic 

units in which an intermediate diversity of corals 

is found. 

SUMMARY AND CONCLUSIONS 

These data may be summarized and conclusions 

drawn as follows (Oliver, 1968b, p. 26-29) : 

1. Acinophyllum segregatum shows low intracolony 

variation, especially in septal number, and this 

narrow range of variation is found in samples 

from five different facies. This suggests that: 

a. Protopolyps were genetically programed for a 

narrow range of septal number variation, 

and this range was transmitted to each member of the colony by means of asexual 

reproduction. 

b. Broader (including bimodal) variation in some 

colonies may be due to genetic accidents or, 

more likely, mechanical accidents such as 

the intergrowth of two or more adjacent 

colonies. 

2. Comparably low intracolony variation was found 

in the other species studied. 

3. Intercolony variation in A. segregatum is related 

to gross environment; variation is greatest in 

facies with diverse coral faunas and least in 

facies wjth restricted faunas. 

4. Intercolony variation in most of the other studied 

samples is roughly comparable to that of associated A. segregatum and may have been 

environmentally controlled. 

5. Two samples (Eridophyllum subseriale and Synaptophyllum arundinaceum) show low intercolony variation although they are from the 

"best" facies (Edgecliff bioherm). These populations or species may have been genetically 

less variable. 

Intercolony variation is a product of the interaction of 

random genetic variation, ecologically induced variation, and 

astogenetic stage and is therefore analogous to variation in 

solitary animals. Such variation is commonly continuous, . but 

discontinuous variation may result from genetically induced 

di- or polymorphism, from nonuniform environmental factors, 

or from inadequate sampling. 

Intracolony variation may be very restricted because of 

genetic uniformity and the presence of relatively uniform environmental conditions. Genetic uniformity leaves room for 

phenetic differences however, and microenvironmental factors 

(situation within colony) may cause enough individual variaton to effectively mask the restrictive effects of genetic similarity. 

Variation in any character may be severly limited within 

a colony because of the lack of genetic variation. Sampling of 

colonial species for statistical analysis must take this into 

account. The analysis should be based on a) equal numbers 

of randomly selected corallites from all colonies, or b) mean 

values calculated for each colony. Sample size should refer 

to number of colonies ·not number of corallites. 
SYSTEMATIC PALEONTOLOGY 43 

0 X 

X 1 1 3 

2 1 2 0 

1 0 

0@1 

X 

X 3 

1 X 

0 x2 

0 0 

1 0 X 

X 

0 0 

0 X 0 

2 

X 

3 

4 4 X 

3 

4 3 X X X 

X 

2 

® 

2 3 4 

4 

B 

FIGURE 18.-Transverse thin section through colony 6 (USNM 162631) of Acinophyllum segregatum (X 0.7). A, Photograph. 

B, "Map" on which each numeral +20 is the number of major septa in the corresponding corallite (for example "0" 

marks corallites with 20 major septa, "3", those with 23). The line separates all corallites with 20 or 21 septa (area A) 

from those with 23 or 24 (area B) except for the three circled corallites. (See text discussion.) The small x's mark corallites in which the septa could not be counted. The bimodal distribution of septal number in this colony is shown in figure 

19. 

The general assumption that number of septa in rugose 

corals is a function of diameter may be in error. Rather, it 

seems that numbers of septa may determine diameter within 

certain limits. (Oliver, 1968b, p. 31-32). 

SYSTEMATIC PALEONTOLOGY 

~EPOSITORIES 

The following descriptions are based on specimens 

from many sources but principally on those from my 

collections made since 1950 while employed by the 

New York State Museum and the U.S. Geological 

Survey. All described material from these collections 

is now deposited in the U.S. National Museum of 

Natural History, the New York State Museum, or 

the collections of the Geological Survey of Canada. 

In addition, many specimens have been studied from 

earlier collections in these and many other institutions, and the New York-Ontario corals have been 

compared to EN A corals from still additional collections. All of these institutions .are listed below with 

the abbreviations used for referring to them in depository lists and the descriptive texts. 

Depositories of described New York-Ontario corals: 

AMNH-American Museum of Nat-sral History, New York. 

GSC-Geological Survey of Canada, Ottawa. 

NYSM-New York State Museum, Albany. 

UBuff-Dept. Geology, University of Buffalo 

(SUNY), Buffalo. 

UMMP-Univ. of Michigan, Museum of Paleontology, Ann Arbor. 

USNM-U.S. National Museum of Natural History, Washington. 

Depositories (in addition to those listed above) of 

other EN A corals serving as basis for descriptions 

or comparisons : 

2 

X 
44 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

(/) 

I.U 

1-

:::::i 

....J 

<( 

0::: 

0 

() 

25 

20 

15 

10 

5 

n=85 

20 22' ·24 

MAJOR SEPTA 

FIGURE 19.-Bimodal distribution of septal number (n) in a 

sample of 85 corallites of colony 6 (USNM 162631) of 

Acinophyllum segregatum. (See text discussion and fig. 18.) 

BMNH-British Museum (Natural History), 

London. 

FMNH-Field Museum of Natural History, 

Chicago. 

MCZ-Harvard Univ., Museum of Comparative 

Zoology, Cambridge. 

OSU-Orton Museum, Ohio State Univ., 

Columbus. 

Paris-Ecole des Mines, Paris. 

PRJ-Paleontological Research Institute, Ithaca. 

ROM-Royal Ontario Museum, Toronto. 

SlUM-Southern Illinois Univ. Museum, 

Carbondale. 

· SUI-Dept. Geology, State Univ. Iowa, Iowa 

City. 

Tiibingen-Quenstedt Collection, Paleontological Institute, Tiibingen. 

UCMP-University of California, Museum of 

Paleontology, Berkeley. 

Several specimens collected by University of Buffalo students have been donated to the USNM by 

Prof. E. J. Buehler. The only known colonial rugose 

coral from the Seneca Member of the Onondaga Formation in New York was collected by Prof. J. W. 

Wells, Cornell University, and later given to the 

USNM. 

CLASSIFICATION 

The classification of the described corals is new 

(see Oliver, 1974), although ultimately based on suggestions of many previous workers. The classification is explained, discussed, and defended in the 

appropriate sections that follow, but it is outlined 

here, in combination with a species list, for 

convenience. 

CLASSIFICATION AND SPECIES LIST -AN OUTLINE 

Family Stauriidae Milne-Edwards and Haime 

Synaptophyllum Simpson 

S. arundinaceum (Billings) 

S. tabulatum (Simpson) 

S. kladion n. sp. 

Family Craspedophyllidae Dybowski 

Subfamily Cylindrophyllinae Oliver 

Acinophyllum McLaren 

A~ simcoense (Billings) 

A. stramineum (Billings) 

A. segregatum (Simpson) 

4. mclareni Fagerstrom 

A. stokesi (Milne-Edwards and 

Haime) 

A. vermetum (Weisbord) 

Cylindrophyllum Simpson 

C. elongatum Simpson 

C. propinquum Stewart 

C. laxtensum n. sp. 

C. deearium n. sp. 

C. sp. A 

Prismatophyllum Simpson 

P. prisma Lang and Smith 

P. ovoideum (Davis) 

P. truncata Stewart 

P. anna (Whitfield) 

Asterobillingsa Oliver 

A. rugosa (Hall) 

A. magdisa magdisa Oliver 

A. magdisa steorra Oliver 

A. affinis (Billings) 

Subfamily Craspedophyllinae Dybowski 

Eridophyllum Milne-Edwards and Haime 

E. seriale Milne-Edwards and Haime 
SYSTEMATIC PALEONTOLOGY 45 

1a 

b 

d 

2 

3 

e 

4a 

b 

0:: 5 LL.J 

~ 6a 

:::> 

z b 

LL.J 

_J 7 c... 

~ 8 (/) 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

f-.---

II 

I 

0 2 

,. 

I I 

I....J 

I 

I 

I 

~ ~0 

,..., 

I....J 

I 

I 

I ,..., 

I'-' 

I 

I I....J 

4 

0 

X 

L__j 

0 

0 

I....J 

I 0 

L..J 

0 

,..., I ~ 

0 

0 

,..., I ....... 

I 

,...,.. I 

.., lo 

--, 0 

1.--. I 

1'--' 

I 

'""' 

h I 

r I 

_J 0 

I 

6 8 10 12 14 16 

Vn 

EXPLANATION 

lntercolony Vn 

Mean intracolony Vn 

Observed range of intracolony Vn 

FIGURE 20.-lntercolony coefficient of septal number variation (V;;) and observed range and mean of intracolony 

coefficients ( V,.) in samples of colonial rugose corals. See 

table 15 for key to sample numbers. 

E. subseriale Stumm 

E. aulodokum n. sp. 

E. corniculum n. sp. 

E. conjunctum (Davis) 

Grewgiphyllum Oliver 

G. colligatum (Billings) 

?Family Disphyllidae Hill 

Disphyllum Fromental 

D? rectiseptatum (Rominger) 

D? stummi (Oliver) 

Family Zaphrentidae Milne-Edwards and Haime 

Cyathocylindrium Oliver 

C. gemmatum (Hall) 

0:: 

LL.J 

co 

~ 

:::> 

z 

LL.J 

_J 

c... 

~ 

<( 

(/) 

Edgecliff bioherm and biostrome facies 

1a I [ 

1b I I ....- 0 

2 .-.I 

L..J 

5 lo 

6a I I p 

L . 7 L..J 

11 ,..., I 

-=-

14 w 

16 ~ 

..., ..... 

.._ 

17 I 

18 - I 

L..J 

Edgecliff bioherm and Bois Blanc Formation 

10 I I 0 

15 I I 

pBois Blanc Formation 

3 I 0 

4a I ,..., I 

I....J 

Jeffersonville Limestone Zones A, B, C 

4b I _j 0 

8 -.::7' 

9 I lo 

Columbus Limestone Zone C 

6b --+ - I '-'I 

Edgecliff fine-grained facies 

1d ~ . l 0 'T 

Moorehouse Member 

1c .-.I 

-= 

13 I 1... 

Moorehouse AcmophyllufTI- Endophyllum ,bed: 

le ,..., I ....... 

}.2 I 

I 1- r=-1 I I I I 

0 2 4 6 8 10 12 

Vn 

EXPLANATION 

0 lntercolony Vn 

X Mean intracolony Vn 

L___J Observed range of intracolony Vn 

I 

0 

I 

I 

14 

FIGURE 21.-Data of figure 20 rearranged by stratigraphic 

unit. 

C. opulens Oliver 

C. n. sp. A 

H eliophyllum Hall 

phaceloid species 

H. megaproliferum n. sp. 

H. n. sp. B. 

H. n. sp. C. 
46 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

cerioid species 

H. coalitum (Rominger) 

astreoid species 

H. monticulum (Ehlers and Stumm) 

H. cassum n. sp. 

H. procellatum n. sp. 

Family STAURIIDAE Milne-Edwards and Haime, 1850 

Diagnosis.-"Fasciculate and cerioid coralla with 

slender corallites; a marginarium may develop as a 

narrow peripheral stereozone; a single series of 

elongate dissepiments, discontinuous vertically may 

occur between major and minor septa." (Hill, 1956b, 

p. F296). 

Discussion.-The Stauriidae ( Columnariidae of 

many autho~s) is a family of relatively simple, colonial rugose corals that is probably polyphyletic in 

origin. At least I doubt that the Onesquethaw genus 

here described is closely related to Stauria or that 

either is related to some of the other conventionally 

included genera. 

Synaptophyllum 8impson (including Placophyllum Simpson) is the only "stauriid" known from the 

Onesquethaw Stage in Eastern North America, 

where its known range is Edgecliff to Moorehous·e. 

Depasophyllum Grabau, Hemilton to Tully in the 

same area, is morphologically similar and a likely 

descendent of Synaptophyllum. Synaptophyllum may 

have been derived from the Ordovician and Silurian 

Palaeophyllum Billings. (See Oliver, 1963b, eastern 

· North American Silurian form.) 

Genus SYNAPTOPHYLLUM Simpson 

part 1900 Synaptophyllum Simpson, p. 212-214 (type 

species only; others Adnophyllum). 

1900 Placophyllum Simpson, p. 216. 

not 1906 Synaptophyllum Simpson. Grabau and Shimer, 

p. 210 (=Acinophyllum). 

not 1909 Synaptophyllum Simpson. Grabau and Shimer, 

p. 73 ( =Acinophyllum). 

1935a Placophyllum Simpson. Lang and Smith, p. 557-

558. 

not 1935a Synaptophyllum Simpson. Lang and Smith, 

p. 561-562 (=Acinophyllum). 

not 1938 Synaptophyllum Simpson. Stewart, p. 42-43 

(=Acinophyllum). 

not 1939 Synaptophyllum Simpson. Stumm, p. 62 

(=Acinophyllum). 

1940 Placophyllum Simpson. Lang, Smith, and 

Thomas, p. 100. 

part 1940 Synaptophyllum Simpson. Lang, Smith, and 

Thomas, p. 129. 

not 1944 Synaptophyllum Simpson. Shimer and Shrock, p. 

95 (=A cinophyllum) . 

not 1948 Synaptophyllum Simpson. Stumm, p. 43-44 

(=Phacellophyllum?). 

1949 Placophyllum Simpson. Stumm, p. 30. 

not 1949 Synaptophyllum Simpson. Stumm, p. 37 (=Phacellophyllum ?) • 

not 1949 Synaptophyllum Simpson. Schouppe, p. 124-127 

( =Phacellophyllum ?) . 

not 1953 Synaptophyllum Simpson. R6zkowska, p. 16 

(=Phacellophyllum?). 

not 1955 Synaptophyllum Simpson. Stumm, card 258 [part 

= Acinophyllum; part = Phacellophyllum?, 

part = Disphyllum ?] . 

1956b Placophyllum Simpson. Hill, p. F298. 

not 1956b Synaptophyllum Simpson. Hill, p. F280 (=Acinophyllum). 

not 1958 Synaptophyllum Simpson. Cranswick and Fritz, 

p. 44 ( =Acinophyllum). 

not 1958 Synaptophyllum Simpson. Schouppe, p. 230-232 

( =P hacellopkyllum?). 

1959 Synaptophyllum Simpson. McLaren, p. 16-18. 

Type species.-By original designation, Diphyphyllum arundinaceum, Billings, 1859, p. 134-135. 

Edg"~ecliff Member of Onondaga Limestone, Niagara 

Peninsula of Ontario and New York State. 

Diagnosis.-Phaceloid or dendroid coralla with 

long, slender, cylindrical or tapering corallites connected by lateral projections of corallite walls. Septa 

noncarinate, thin, of two orders; the major may 

either be short or extend nearly to the axis, the 

minor are commonly short ·and rarely more than half 

as long as the major. The septa merge into a peripheral stereozone. There are no dissepiments. Tabulae complete, horizontal to ·arched axially·. Increase 

peripheral. (Diagnosis modified from McLaren, 

1959, p. 16.) 

Microstructure.-The septa and wall appear to be 

primarily lamellar. Septa have dark, discontinuous 

center lines probably formed by monacanthine trabeculae. Discontinuous, overlapping lam•ellae follow 

the outline of the skeleton and are continuous from 

septa to wall. In som·e specimens, faint discontinuities can be observed in the wall, midway between 

septa, and crushed corallites show a tendency to 

break along these lines of apparent weakness. 

The tabulae are extremely thin and no microstructure has been observed jn them. 

Discussion.-McLaren (1959) cleared up the confusion that has surrounded Synaptophyllum since its 

first description. Simpson (1900) named Diphyphyllum arundinaceum Billings as genotype but based his 

generic concept on Eridophyllum simcoense Billings. 

The two species are not congeneric. Lang and Smith 

(1935, p. 561-562) further confused the issue by illustrating a specimen of a third genus as Synaptophyllum arundinaceum Billings. McLaren redescribed the type material of D. arundinaceum and 

showed clearly that it is a stauriid (sensu Hill, 

1956b, p. F296). He assigned E. simcoense to a new 
SYSTEMATIC PALEONTOLOGY 47 

genus Acinophyllum which represents Simpson's and 

most American workers' concept of Synaptophyllum 

as a craspedophyllid. He reassigned Smith's material 

to Phacellophyllum. Prior to McLaren's excellent redescription of the type material of Synaptophyllum, 

all workers, using the erroneous concepts of either 

Simpson or Lang and Smith as their guides, had assumed the genus to have dissepiments. 

Increase.-Increase is lateral .in all observed examples, but separation of the offset by a skeletal tissue is effected in a relatively short growth distance, 

and protocorallite and offset are closely appressed 

through much of the offset's conical stage. This results in a very small "opening" between protocorallite and offset and .in a double wall between the two 

that persists through most of the offset's conical 

stage. These features can be observed in both longitudinal (pl. 6, fig. 4) and transverse sections (pl. 7, 

fig. 10). Transverse sections show a complete wall 

(pl. 4, figs. 7, 8) at a much smaller offset diameter 

than is characteristic of other phaceloid species 

studied ( cf. Acinophyllum spp. in pls. 14 and 22). 

This description is based on several coralla of S. 

arundinaceum and S. kladion in which increase was 

observed. The single known fragment of S. tabulatum does not preserve any offsets. 

Placophyllum Simpson has been considered unrecognizable by most workers. Until the present 

time, the only available specimens of the genus were 

two longitudinal thin sections (NYSM 300) which 

are the originals of Simpson's only illustration of the 

type species, P. tabulatum. To these have now been 

·added a transverse thin section, of similar vintage 

and workmanship as the two type slides, which bears 

the same locality number as the others and which is 

accompanied by a label in Simpson's handwriting 

identifying it as "Placophyllum tabulatum Simpson." 

The transverse section may be from the type corallum and has been added to the type collection. In 

addition I have collected a single corallum which in 

every respect agrees with the types and which unquestionably belongs to P. tabulatum. It agrees with 

S. arundinaceum in all important respects except 

size and there is no longer any question that Placophyllum is a junior synonym of Synaptophyllum as 

suggested by McLaren ( 1959, p. 17) . 

Depasophyllum Grabau (1922, 1936, not Yu, 1934; 

type species D. adnatum Grabau, 1936, basal Tioughnioga, Michigan) is simHar to Synaptophyllum in its 

general morphology, that is, short septa and strongly deflected tabulae, but differs as follows: ( 1) 

Minor septa are commonly lacking except in distal 

parts of large specimens (gerontic?) ; when present 

they tend to be contratingent. (2) Coralla are solitary and not known to produce offsets; "peripheral 

buds" of Ehlrer.s and Stumm (1949, p. 31) are young 

of the .species cemented to an adult. Only two species 

of Depasophyllum have been described, both from 

the upper Middle Devonian of eastern North America. It is possible that these are solitary descendents 

of the colonial Synaptophyllum but the differences 

are certainly great enough to warrant distinction as 

genera. 

Three species of Synaptophyllum are known from 

the Onondaga Limestone. S. arundinaceum (Billings) and S. tabulatum (Simpson) are limited to the 

Edgecliff Member (possibly to the bioherm facies) 

and are distributed from eastern New York to the 

Niagara Peninsula of Ontario. S. arundinaceum is 

also known from the Columbus Limestone (Zone C), 

near Bellepoint, Ohio, and from the Perry core, Lake 

County, Ohio. S. kladion n. sp. is known only from 

the Moorehouse Member in western New York and 

southwestern Ontario. All three species are unco·mmon. 

Synaptophyllum grabaui Fagerstrom (1961, p. 13, 

pl. 3, figs. 11-14), from the Formosa reef facies of 

the lower Detroit River Group in southwestern Ontario, is not a stauriid. It appears to be an aberrant 

form, po,ssibly related to "Tryplasma" rhopalium 

Oliver, 1960b ( = Cyathophyllum marylandicum 

Swartz, 1913). 

Silurian species that were assigned to Synaptophyllum by Grabau (1910) ·and subsequent workers 

have been discussed by Oliver, 1963b (p. G5). These 

have vertically elongate dissepiments in a single row 

and differ from species of Synaptophyllum in this 

and other respects. 

Synaptophyllum arundinaceum (Billings) 

Plates 2-4; plate 7, figures 7-11 

1859 Diphyphyllum arundinaceum Billings, p. 134-:-

135. 

not? 1874 D. arundinaceum Billings. Nicholson, p. 32-33, 

pl. 6, fig. 1 (probably Acinophyllum segregatum). 

not 1887 E. arundinaceum. Davis, pl. 112, fig. 2 (see 

Acinophyllum davisi Stumm). 

not 1901 D. arundinaceum Billings. Lambe, p. 162-163, pl. 

14, fig. 1, 1a, b (see Acinophyllum segregatum 

Simpson). 

not 1923 D. arundinaceum Billings. Clark, p. 218 (see 

Disphyllum? stummi Oliver). 

not 1935a Synaptophyllum arundinaceum (Billings). Lang 

and Smith, ( Phillipsastraeid, figs. 19, 20 p. 

561, see McLaren, 1959, p. 29, for discussion). 

not 1944 S. arundinaceum (Billings). Shimer and Shrock, 

p. 95, pl. 30, figs. 5-7 (see Acinophyllum segregatum Simpson). 
48 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

not 1945 Disphyllum [S.] cf. arundinaceum (Billings). 

Smith, p. 22, pl. 12, figs. 1, 2 (Phillipsastraeid, 

see McLaren, 1959, p. 29). 

not 1949 Macgeea (Synaptophyllitm) arundinaceum (Billings). Schouppe, pl. 12, figs. 60~ 61 (Phillipsastraeid) . 

not 1949 Synaptophyllum arundinaceum ( B i 11 i n g s ) . 

Stumm, pl. 17, figs. 19, 20 · (Phillipsastraeid, 

see McLaren, 1959, p. 29). 

not 1955 S. arundinaceum (Billings). Stumm, cards 259, 

260 (part=Phillipsastraeid; part unrecognizable). 

not 1956b S. arundinaceum (Billings). Hill, fig. 191-6 

(Phillipsastraeid, see McLaren, 1959, p. 29). 

not 1958 S. arundinaceum (Billings). Schouppe, figs. 15, 

16 (Phillipsastraeid, see McLaren, 1959, p. 29). 

1959 Synaptophyllum arundincweum (Billings). McLaren, p. 18-22, figs. 4-6; pl. 7, figs. 1-3; pl. 

8, figs. 1-5. 

not 1963 Eridophyllum a'rundinaceum. Kato, fig. 12-32 

(probably= Acinophyllum sp.). 

Occurrence of type material.-"Rama's farm, near 

Port Colborne, and in various localities in the townships of Walpole, Oneida, Cayuga, and W ainfleet, in 

the corniferous limestone" (Billings, 1859, p. 135). 

Lectotype selected by McLaren, 1959, is "from 3 

miles west of Cayuga, Ontario" ; syntypes are from 

Wainfleet township, Weiland County, and Walpole 

township, Haldimand County, Ontario (McLaren, 

1959, p. 19). All of these localities are most likely in 

the Edgecliff Member of the Onondaga Limestone. 

Diagnosis.-Phaceloid Synaptophyllum with relatively small corallite diameters and tabulae that are 

axially flat and strongly deflected downward at the 

periphery. 

External features.-Phaceloid colonies are composed of subparallel, straight or slightly flexuous, 

cylindrical corallites. The partly crushed lectotype 

corallum is 30.5 em high and 28 by 5 em wide; other 

parts of coralla on which this description is based 

range up to 19 em high and 12 by 5 em wide. M.a.ture 

corallites range in diameter from 3.5 to 8.0 mm; 

within colonies size v~ariation is more limited (table 

16); the mean diameter in 10 colonies ranges from 

4.5 to 6.9 mm (fig. 22). 

Increase is lateral at infrequent intervals; expansion is rapid in the offsets. Corallites are supported 

within the colony by tapering lateral projections of 

the corallite wall, but these are rare in the coralla 

studied. 

Corallites are irregularly rugose and marked by 

fine encircling growth striae. Longitudinal septal 

grooves and broad, rounded interseptal ridges are 

weakly to strongly developed, giving a scalloped 

cross-sectional outline to many corallites. 

22 

21 

20 

19 

/x---;c 

/ ) 

't::: 

//X 11'6-··x-·~}y/ 

I ....;xl X/~> 18 

17 

16 

15 

4.0 

j2g' I .,...., / EXPLANATION I I _........ / 

~-_.,...... r // X Synaptophyllum arundmaceum 

I // + S. tabulatum 

I / 

I / 

I / '{~// 

5.0 6.0 

6 S. klad/on 

0 Holotype or lectotype corallum 

7.0 

d(mm) 

8.0 

FIGURE 22.-Mean number of major septa (n) versus mean 

diameter (d) in Synaptophyllum spp. The polygons enclose 

samples of the different species. 

N 

n: 

d: 

TABLE 16.-Synaptophyllum arundinaceum Billings, sum- mary data 

Minimum 

---------- 12 

x 17.3 

OR ----- 15-19 

8 ------- .51 

v ------ 2.5 

x 4.5 

OR ----- 3.5- 5.9 

8 ------- .44 

v ------ 6.6 

Intracolony 

mum 

Maxi20 

20.2 

17-22 

1.76 

9.3 

6.9 

6.2- 7.9 

.90 

17.4 

Inter- Mean colony 

9 

18.9 

17.3-20.2 

.91 

5.4 4.8 

5.9 

4.5- 6.9 

.81 

12.5 13.7 

The coralla were structurally weak, and most 

specimens are broken. In some colonies the corallites 

have heen pressed close together in one direction 

(pl. 3, fig. 2) ; in others the corallites were broken 

apart and scattered before fossilization. Better preserved parts of coralla indicate that the corallites 

were subparallel to each other and separated by a 

distance approximately equal to one corallite 

diameter. 

Calices are deep and steep sided, with low septal 

ridges extending down the calioe walls and part way 

over the flat bottom. 

Internal features.-The septa are radially arranged. Major septa are amplexoid and commonly 

extend to or nearly to the axis on the surfaces of 

tabulae; between tabulae they range in length from 

one-third to three-fifths the radius and are commonly 

about one-half. The average number of major septa 

per corallite in 9 coralla ranges from 17.3 to 20.2 

(fig. 22) with limited variation in any one colony 

(table 16). Septa are thin (average 0.15 mm peripherally) and taper toward the axis. Minor septa vary 

in length but average one-half the length of the 
SYSTEMATIC PALEONTOLOGY 49 

major septa and are somewhat thinner. All .septa are 

reasonably straight .and lack carinae. 

Peripherally, all septa dilate to form the outer wall 

which ranges from 0.2 to 0.5 mm in thickness. No 

dissepiments are present. 

The tabulae are typically complete with broad, flat 

axial parts and sharply downbent peripheral parts. 

They are irregularly spaced, ranging from three to 

seven per 5 mm. 

Microstructure.-As for genus. 

Discussion.-Synaptophyllum arundinaceum was 

originally described by Billings (1859, p. 134-135), 

but internal features were first illustrated and examined by McLaren (1959). In the intervening 100 

years, the name was used for cylindrophyllinid species now referred to Acinophyllum (McLaren, 1959). 

S. arundinaceum differs from S. tabulatum (Simpson) in its smaller corallite size and in its longer 

septa and more highly arched tabulae. It differs from 

S. kladion n. sp. in the infrequency of its offsets and 

lateral connections, .in its cylindrical corallites, and 

in the regular form of the tabulae and the lesser variation in length of major septa. 

Distribution.-S. arundinaceum occurs in the 

Edgecliff Member of the Onondaga Limestone in 

N·ew York and in the Niagara Peninsula of Ontario. 

The species is widespread in the Edgecliff bioherm 

facies, having been collected from nearly one-third 

of the known bioherms in both eastern and western 

New York and in the Niagara Peninsula. None of the 

known New York specimens are from the nonbioherm facies of the Edgecliff. 

-The lectotype and syntypes and four other specimens studied are from the Niagara peninsula of 

Ontario (McLaren, 1959). They are probably all 

from the Edgecliff Member, but at least two of the 

coralla are from nonbioherm facies. 

Specimens are known from the Columbus Limestone (lower Columbus, coral zone, probably Edgecliff in age), both in the outcrop belt near Bellepoint, 

Ohio, and in the subsurface of Lake County, northeastern, Ohio. 

Material studied.-The above description is based 

primarily on 10 coralla : (a) the lectotype ( GSC 

3602) of McLaren (1959, p. 19), (b) five coralla collected by the author (USNM 162576-78 and NYSM 

12821, 22), (c) one additional corallum from the 

USNM (10719), and (d) three coralla from the Geological Survey of Canada (GSC 3422a, 31151-52). 

Counts and measurements are based on an analysis 

of seven lectotype corallites and 12 to 20 corallites in 

each of the other 9 coralla. Additional specimens providing supplemental morphologic and distribution 

data are: GSC 3431b-f, 3432 (paralectotypes, see 

McLaren, 1959, p. 19), nine specimens collected by 

the author (USNM 162579-86 and NYSM 3488), 

and GSC 22812. 

The specimens from the Columbus Limestone are 

USNM 162587-88. 

Detailed locality information for all specim·ens is 

in the locality section and tables 40 to 46. 

Synaptophyllum tabulatum (Simpson) 

Plate 5 

1900 Placophyllum tabulatum Simpson, p. 216-217, fig. 

41. 

1935a P. tabulatum Simpson. Lang and Smith, fig. 16, p. 

557. 

1949 P. tabulatum Simpson. Stumm, pl. 14, fig. 8. 

1956b P. tabulatum Simpson, Hill, fig. 203, p. 6. F299. 

1959 P. tabulatum Simpson. McLaren, pl. 10, figs. 1, 2. 

? 1961 P.? sp. Fagerstrom, p. 13, pl. 4, fig. 15. 

? 1961 Depasophyllum sp. cf. D. adnatum Fagerstrom( p. 

13, pl. 4, fig. 14. 

Occurrence of type material.-According to the 

New York State Museum label and catalog, Simpson's type corallum was collected by C. D. Walcott in 

1878 from the Onondaga Limestone at Walpole, 

Ontario. ·This is almost certainly the Edgecliff Member and may or may not be the bioherm facies. 

Diagnosis.-Synaptophyllum with large corallite 

diameters; tabulae are axially flat and smoothly bent 

downward at the periphery. 

External features.-Dendroid or possibly phacelloid coralla of unknown size and shape, the only 

corallum known other than the holotype, measures 

approximately 11 by 9 by 6 em. Mature corallites are 

straight or somewhat flexuous and range in di·ameter 

from 7.4 to 10.7 mm in the specimens available for 

study; mean diameter of 14 corallites in one corallum 

is 9.3 mm, observed range is 7.4 to 10.7 mm; the 

diameter of the holotype corallite is approximately 

9.8mm. 

Increase is presumed to be lateral at infrequent intervals, although no examples of increase have been 

obs·erved in the specimens at hand. Lateral supports 

are apparently rare but have been observed in both 

the type specimens and in the supplementary 

corallum. 

No well-preserved corallite exteriors have been 

directly observed, but weathered specimens and sections of corallites in the matrix indicate that encircling rugae and longitudinal septal grooves are present. Finer markings are not known. No calices have 

been preserved, but they may be deduced to be steep 

sided with virtually flat bottoms. 
50 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Internal features.-Septa are radially arranged. 

Major septa are amplexoid, and extend approximately two-fifths the distance to the axis on the tabulae 

surfaces, but are shorter above the tabulae. The 

number of major septa in the supplementary corallum ranges from 18 to 21 in 10 corallites and averages 19.9. Septa are thin (com·monly 0.15 mm peripherally) and taper towa:rd the axis. Minor septa are 

one-half as long as the major and somewhat thinner. 

All septa are straight and lack carinae. 

Peripherally, all septa dilate to form an outer wall 

which ranges from 0.2 to 0.35 mm in thickness. 

There are no dissepiments. 

Tabulae are typically complete with broad flat 

axial areas and gently downb{!nt peripheries. Spacing is irregular, being two to four per 5 mm in the 

specimens studied. 

Microstructure.-As for the genus. 

Discussio·n.-The above description is based on 

three thin sections of individual corallites that compose the type lot ( NYSM 300 and 1480) and on one 

additional corall urn collected in the course of this 

study. The types are from southwest Ontario; the 

supplementary cor all urn is from eastern New York. 

Placophyllum tabulatum was established by Simpson (1900, p. 216-217) by ·a generic description and 

illustration of a longitudinal s.ection, a part of which 

is preserved as two thin sections (NYSM 300). 

Simpson seems to have based his description on one 

or more large coralla, but only the illustrated thin 

sections are included in the type collection. 

The genus and species have been considered unrecognizable by most workers, and all descriptions 

have been based on Simpson. Subsequent illustrations have been cop.ies of Simpson's illustration or 

new photographs of the type slides. To the type collection ha:s now been added a transverse thin section 

of a corallite bearing the same locality number as the 

other sections and labeled Placophyllum tabulatum 

in Simpson's handwriting (NYSM 1480). This 

transverse section and the second corallum from 

eastern New York make possible a more adequate 

description of Simpson's species and its reassignment to the genus Synaptophyllum. 

S. tabulatum differs from S. arundinaceum in its 

large corallite size, short thinner septa, and broader, 

more gently flexed tabulae. Corallite size was compared in the two species, using the method for small 

samples outlined by Simpson, Roe, and Lewontin 

(1960, p. 182-183). Two tests were made: (1) Comparing the mean corallite diameter of 10 coralla of 

S. arundinaceum with that of one corallum of P. 

tabulatum gave the values t= 3.56, P<0.01. (2) Comparing the maximum diameters of the 10 coralla of 

S. arundinaceum with that of S. tabulatum gave the 

values t=4.51, P<O.Ol. According to both tests, the 

chances are less than one in a hundred that the 

single corallum was drawn from the same population as the 10 coralla. The tests suggest a significant 

difference in corallite size between S. arundinaceum 

and S. tabulatum and indicate that they may represent different species. This, plus the qualitative differenees noted, makes it seem best to recognize both 

species until more adequate collections can be 

studied. 

Placophyllum? sp. of Fagerstrom (1961, p. 13, pl. 

4, fig. 15) has flat tabulae and very short septa (compare my pl. 5, fig. 4) and may be a fragment of S. 

tabulatum. Depasophyllum sp. cf. D. adnatum of 

Fagerstrom (1961, p. 13, pl. 4, fig. 14) has downturned tabulae and amplexoid :septa and is very 

similar to typical S. tabulatum. Both forms are represented by cylindrical fragments and cannot be certainly identified. They were derived from the Formosa reef facies of the lower Detroit River Group 

and are probably Edgecliff in age. 

Distribution.-S. tabulatum is known from only 

two localities. The type thin sections are labeled Walpole, Ontario. They are probably from the Edgecliff 

Member but might be from either the bioherm or biostrome facies. The supplementary corallum is from 

the bioherm facies of the Edgecliff Member in the 

Helderberg escarpment, southwest of Albany, N.Y. 

The two localities are more than 300 miles ( 480 km) 

apart and represent almost the western and eastern 

extremities of the exposed Edgecliff Member. Apparently the species was widespread but very rare. 

The questionable specimens of Fagerstrom (see 

discussion) are from the Formosa reef limestone of 

probable Edgecliff age in central southwestern 

Ontario. 

Material.-Hoiotype, NYSM 300 (two longitudinal thin sections); syntype NYSM 12875 (one transverse thin section); supplementary corallum, USNM 

162607. Placnphyllum •sp. F!agerstrom, hypotype 

UMMP 36125A, B (two thin sections)'; Depasophyllum sp. cf. D. adnatum of Fagerstrom, hypotype 

UMMP 36121 (one fragment). 

Synaptopbyllum kladion n. sp. 

Plate 6; plate 7, figures 1-6 

Occurrence of type material.-Holotype, GSC 

31153, Moorehouse Member, Onondaga Limestone, 5 

miles ( 8 km) south-southeast of Cayuga, Ontario. 

Paratypes are from the Moorehouse Member near 

Leroy, Wilhelm, and Bellevue in western New York. 
SYSTEMATIC PALEONTOLOGY 51 

A single specimen is known from the upper part of 

the Edgecliff Member near Port Colborne, Ontario. 

Diagnosis.-Dendroid Synaptophyllum with numerous offsets and lateral connections; tabulae are 

domed or irregular, :septa are quite long in some 

corallites. 

External features.-Colonies are dendroid, composed of radiating, flexuous, ceratoid corallites. The 

largest available corallum occupies a block of limestone 19 by 17 by 8 em; the colony is broken, however, and the original was larger. 

Individual corallites tend to expand gradually and 

assume a ceratoid or subcylindrical form. Mean diameters, based on 12 to 15 individuals in 3 colonies, 

are shown in figure 22, but their significance may be 

questioned. The measurements are from random thin 

sections and do not. represent maximum dia-meters of 

individual corallites. In the following table the maximum measured diameter may be of somewhat greater value in comparing S. kladion with other species 

of the genus. 

Colony 

USNM 162589 

GSC 31153 ------- USNM 162590 ___ _ 

d(mm) ORr1(mm) n 

5.2 4.1-5.9 

5.9 5.1-6.9 

6.9 5.4-8.4 

15.5 

19.1 

18.5 

OR,. 

14-17 

18-20 

17-20 

N 

13 

15 

12 

Increase is lateral ·and offsets are common; expansion is gradual in the corallites. Corallites are supported within the colony by tapering lateral projections of the corallite wall ; these are common in parts 

of the coralla studied; ·as many a-s six lateral supports in 3 em have been observed (pl. 6, figs. 9, 10). 

Corallites are irregularly rugose and marked by 

fine encircling growth striae. Longitudinal septal 

grooves and broad, rounded interseptal ridges are 

weakly developed, giving a scalloped cross-sectional 

outline to corallites in transverse thin sections. 

The coralla were apparently not very strong. Parts 

of each of eight coralla studied are preserved with 

corallites in their original relative positions but all 

are partly broken or crushed. Additional specimens 

in the collection consist of a few to several disoriented and broken corallites. 

Calices are deep and steep sided ; low septal ridges 

extend down the sides and part way across the floors. 

Calycinal rejuvenescence was noted in two colonies. 

Internal features.-The septa are radially arranged. Major septa are amplexoid and range in 

length from one-third to two-thirds the radius; the 

length tends to be relatively less in larger sections 

and longer ·gepta may represent immature parts of 

corallites. The average number of major septa in 

three coralla ranges from 15.5 to 19.1 with limited 

variation in any one colony (table above). Apparently the full quota of septa was obtained by most corallites weB before they achieved maximum diameter. 

Septa are thin and taper toward the axis. Minor septa are one-half or less as long as the major. All septa 

are straight and lack carinae. 

Peripherally all septa dilate to form the outer wall 

which ranges from 0.15 to 0.4 mm in thickness. No 

dissepiments are present. 

The tabulae are commonly irregular in form. In 

longitudinal section they vary from flat to irregularly undulating to dome shaped having downbent peripheries as in the other species described. They are 

irregularly spaced, ranging from three to six per 5 

mm in the specimens sectioned. 

Microstructure.-As for genus. 

Discussion.-The above description is based on 

eight coralla, seven collected by myself from known 

positions iu the Moorehouse Member and one from 

the upper part of the Edgecliff Member. 

S. kladion differs from both S. arundinaceum and 

S. tabulatum in its dendroid habit, ceratoid corallites, 

and irregular tabulae; in the last two species, growth 

form is phacelloid, corallites are cylindrical, and 

tabulae are regularly arched. 

Distribution.-S. kladion is most common in the 

Moorehouse Me1mber. The eight known specimens 

are distributed as follows: six specimens are from 

the western coral facies of the Moorehouse Member 

near Leroy, Wilhelm, and Bellevue, N.Y.; one specimen is from the middle Moorehouse at Dunnville, 

Ontario. The eighth specimen is from the Edgecliff 

Member (biostrom·e facies) at Port Col borne, Ontario. One more specimen, possibly referable to this 

spec:ies, was collected by A. Murray in 1860 from 

"near W oodstrock" ( GSC 2599) . 

Material.-Holotype, GSC 31153; illustrated paratypes US.NM 162589, 90, 92, 93; unillustrated paratypes USNM 162591 and 162594, NYSM 12823, and 

GSC 31154. 

Family CRASPEDOPHYLLIDAE Dybowski, 1873 

Diagnosis.-Solitary( ?) , phaceloid, cerioid, phaceloid-cerioid, astreoid, and thamnasterioid rugose 

corals with long or short, very attenuate major septa 

marked by zigzag carinae. The septa characteristically are composed of very find monocanthine trabeculae and appear in most sections to have a dark axial· 

zone with light tissue on either side. The major septa 

are deflected to form an aulos in the Craspedophyllinae, but this structure is lacking in the Cylindrophyllinae. The dissepimentarium is formed by one to 

several rows of globose normal dissep.iments. Tabulae are variable but commonly flat or slightly arched 
52 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

axially and downbent at the margin of the 

tabulari urn. 

Included subfamilies.-Cylindrophyllinae new and 

Craspedophy llinae. 

Discussion.-The Craspedophyllidae as here recognized includes Eridophyllum ( = Craspedophyllum), 

and related form'S :in the nominate subfa.mily, and a 

group of genera previously generally classified as 

Disphyllidae, here assigned to the new Subfamily 

Cylindrophyllinae. The Cylindrophyllinae are essentially the Billingsastraeinae of Jell (1969, p. 63), 

but this name is not available as the genus Billingsastraea proves to be a Silurian coral unrelated to the 

form·s on which. our concept of Billingsastraea has 

been based. (See generic discussion.) 

The cylindrophyllinids differ from the disphyllids 

(s.s.) (essenti,ally the Disphyllinae of Jell) in having strongly attenuate septa formed of fine monocanths with well developed zigzag carinae. In contrast the disphyllids have thicker ·septa formed of 

coarse monacanths and generally lack such marked 

carinae. These characters are vari,able within genera 

and species, but it seems very significant that the.-· 

cylindrophyllinid 'structure overwhelmingly predominates in eastern North American species and 

genera, whereas the disphyllid structure predominates in other areas of the world. 

There are analagous genera in the two family 

units as outlined here: 

Eastern Americas Realm Old World Realm 

AC ctl·'t}-od.phylhluml } ·------------------------- _Disphyllum y m rop yl um Prismatophyllum _________________________ _ H exagonaria 

In general these generic "pairs" differ only in the 

characters mentioned above. The EAR Asterobillingsa ( = Billingsastraea of authors) has no 

named Old World counterpart in the Disphyllidae. 

Eridophyllum and its allies (Cra.spedophyllinae) 

differ from the Cylindrophyllinae only in the possession of an aulos and are also a distinctly ENA develop·ment. It seems likely that the Craspedophyllinae were derived from the Cylindrophyllinae in 

middle Onesquethaw time, ·as discussed below. The 

apparent close relationship of the two groups can 

best be noted by inclusion of them as subfamilies 

with the Craspedophyllidae. 

The degree of relationship between the Craspedophyllidae and the Disphyllidae is more difficult to 

analyze. Probably too little is known of the early history of either fam:ily to offer a meaningful basis for 

an opinion as to whether they are more closely related to each other than to other families. On the 

basis of my analysis of the development and history 

of the Eastern Americas biogeographic realm, and 

of the morphologic similarities and dissimilarities, I 

suggest that the Craspedophyllidae evolved independently in the EAR .at least from Siegenian time. 

I know of no pre-Siegenian North American rugose 

coral that is at all likely as an ancestor to the family. 

I doubt that either family is very closely related to 

the Phillipsastraeidae, although both have been so 

grouped by many workers in the past. 

No true disphyllids are known from EAR rocks 

older than the Cazenovia Stage. Forms indi:Jtinguisha;ble from H exagonaria are known from the Miami 

Bend Formation of Cooper and Phelan (1966) 

(Cazenovian age in Indiana) and, with Disphyllum, 

from the Traverse Group (Tioughnioga age, Michigan). These probably represent invasions from the 

north or northwest, as they are accompanied by 

other Old World genera of rugose corals although 

apparently not by other disphyllids. The only possible Old World coral known to occur east of the Nashv:ille-Cincinnati-Findlay-Algonquin ·axis is Disphyllum? stummi. 

Distribution.---'Cra.s·pedophyllids range in age 

from early Onesquethaw to Taghanic in the Eastern 

North American province. Early and Middle Devonian extensions of this provincal fauna into northern South America and northwest Africa are discussed above and under the appropriate generic 

heading. 

Family name.-Craspedophyllum Dybowski 

(1873a, p. 83; type species by monotypy, C. americanu.m Dybowski, 1837b, p. 155-159, pl. 6, figs. 1-6) 

has commonly been considered a junior synonym of 

Eridophyllum although no one has redescribed the 

type species or compared it with other described spedes of Eridophyllum. The location of the type material is not known. Under the circumstances, any 

interpretation of Craspedophyllum must be based on 

(1) Dybowski's description and illustrations and (2) 

his statement that his specimens ca.me from Columbus, Ohio. 

Dybowski's description and jllustrations are ambiguous although he clearly describes and illustrates the aulos. His figure 1 is suggestive of Grewgiphyllum colligatum in the presence of "growth 

swellings" that appear to extend in all directions 

from the corallite, but he does not show any contacts 

w.ith other corallites or a cerioid stage. Dybowski's 

figures 2 and 5 a.re longitudinal and transverse sections of an Eridophyllum, possibly E. seriale MilneEdwards and Haime. His figures 3, 4, and 6 are details and could represent most species of either 
SYSTEMATIC PALEONTOLOGY 53 

genus. All of the figures are drawings and may be 

somewhat diagrammatic. 

Dybowski makes much of the growth swellings 

but does not clearly descr:ibe the manner in which 

adjacent corallites make contact. He does describe 

the aulos clearly enough for identification of his 

specimens as Eridophyllum. Possibly he had specimens of two or more species so that his description 

is a composite. 

Dybowski's only stratigraphic or geographic information is "Columbus, Ohio." This suggests the 

Columbus Limestone in which Eridophyllum (several species) is common and Grewgiphyllum is rare or 

does not occur. (See discussion under G. colligatum.) 

The combined data :indicate that Craspedophyllum 

americanum is a Columbus Limestone Eridophyllum, 

most likely E. serial e. 

Microstructure.-The craspedophyllid wall is thin, 

thicker in the region of the lateral supports. Microscopically, it appears to be composed of an inner 

light-colored zone of minute fibers of calcite radially 

oriented in transverse section and an outer very thin 

dark zone of uncertain structure. In longitudinal section the fibers are seen to incline upward toward the 

axis at approximately the 'Same angle as the top surface of each dissepiment so that in some places the 

dissepiments seem to be structurally continuous with 

the epitheca. More generally the dissepiments, although structurally similar are separated from the 

epitheca by a discontinuity. The tabulae are commonly very thin and their microstructure is obscure. 

Septa are composed of parallel, fine, closely-spaced 

monacanthine trabeculae. Where the septal microstructure is well preserved it appears as follows. A 

dark central area, approximately 0.03 mm thick, indents the epitheca and extends the full length of the 

septum. This is apparently composed of closely 

spaced "centers of calcification." On either side of 

the dark central zone is a light-colored zone composed of m:inute fibers directed away from the central zone and inclined slightly toward the axis of the 

corallite, giving a pinnate appearance in transverse 

s,ection. Septa are separated from the wall by a dark 

line but the light-colored zone is identical to the inner 

wall in texture. Septa are thickest at their peripheral 

ends; the light-colored tissue thins toward the inner 

part of the tabularium. Carinae rorm ridges on the 

sides of the septa and show the same structure as the 

septa in transverse section. 

Subfamily CYLINDROPHYLLINAE Oliver, 197-1 

1969 Billingsastraeinae Jell, p. 63. 

1974 Cylindrophyllinae Oliver, p. 167. 

Type genus.-Cylindrophyllum Simpson. 

Diagnosis.-Craspedophyllids that lack an aulos. 

Included genera.-Cylindrophyllum Simpson, 

Prismatophyllum Simpson, Acinophyllum McLaren, 

Asterobillingsa Oliver ( =Billingsastraea of most 

authors). 

D1:scussion.-Possible relationships of the cylindrophyllinids to the disphyllids and a comparison of 

the two family-level taxa are included in the family 

discussion. 

The cylindrophyllinid genus Prismatophyllum 

probably gave rise to the Craspedophyllinae. This 

idea is developed more fully in the discussion of that 

subfamily. 

The earliest known cylindrophyllinid is Asterobillingsa ajjinis (Billings), from the Grande Greve 

Limestone, Gaspe, Quebec (Oliver, 1964), known 

from a single specimen. Except for this isolated occurrence, all four genera appeared at virtually the 

same ti•me in the middle Onesquethaw. Similarities 

suggestive of intergeneric relationships are discussed under the appropriate species headings but none 

are particularly helpful in understanding evolution 

within the subfamily. A pre-middle Onesquethaw 

record is implied. Because Asterobillingsa seems unlikely to be the primitive member of the group, a history at least as ancient as Siegenian or Gedinnian 

seems probable. 

Subfamily distribution.-Early Onesquethaw-Tioughnioga, Eastern North America; middle or upper 

Onesquethaw, Venezuela. No Taghanic or Frasnian 

form'S are known. (See summaries of distribution 

under generic descriptions.) 

Genus ACINOPHYLLUM McLaren 

part 1900 Synaptophyllum Simpson, p. 212-214. 

1906 Synaptophyllum Simpson. Grabau and Shimer, 

p. 210. 

1909 Synaptophyllum Simpson. Grabau and Shimer, 

p. 73. 

1935a Synaptophyllum Simpson. Lang and Smith, p. 

561-562. 

1938 Synaptophyllum Simpson. Stewart, p. 42-43. 

1944 Synaptophyllum Simpson. Shimer and Shrock, p. 

95. 

part 1955 Synaptophyllum Simpson. Stumm, card 258. 

1956b Synaptophyllum Simpson. Hill, p. F280 (not the 

illustrations). 

1958 Synaptophyllum Simpson. Cranswick and Fritz p.4~ , 

1959 Acinophyllum McLaren, p. 22-23. 

1967 Cylindrophyllum (Acinophyllum) McLaren, 

Pickett, p. 22. 

1974 Acinophyllum McLaren. Oliver, p. 167. 

Type species.-By original designation, Eridophyllum simcoense Billings, 1859, p. 132, text fig. 27. 
54 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

"Rama's Farm [in Port Colborne] and near the town 

of Simcoe." Southwestern Ontario; Bois Blanc 

Formation. 

Diagnosis.-Compound cylindrophyllinid corals 

with dendroid or phaceloid coralla made up of 

slender corallites often connected by lateral proj ections of corallite walls. Increase is lateral and nonparricidal. Major and minor septa with weak to 

strong, commonly zigzag carinae in peripheral zone. 

Major septa short or long but do not extend to axis ; 

radially arranged with no significant modification of 

protosepta in m·ature parts. Dissepimentarium consists of one row of globose, normal dissepiments; one 

or two additional rows of :smaller dissepiments may 

be present. Tabularium wide; tabulae complete, commonly flat or concave axially, strongly downbent 

peripherally. 

External features.-Phaceloid and dendroid colonies sm,all to large, lenticular, as much as 100 em or 

more in diameter and 30 em in height, composed of 

radiating or subparallel corallites. Mature corallites 

are cylindrical, straight to slightly flexuous with dia~meters as much as 1 em in some species. 

Increase is lateral and nonparricidal at varying 

intervals; ·expansion is rapid in new corallites, and 

the initial conical stage is short. 

Corallites are supported within the colonies of 

most species by tapering lateral projections of the 

wall; a single support may be formed by one or both 

of the adjacent individuals but the deposit of each is 

discrete and cemented to the wall or projection of the 

adjoined coral. 

Growth lines are fine and (or) coarse and individuals in some species are quite rugose. Longitudinal septal grooves are ~shallow and separated by 

broad interseptal ridges giving a scalloped appearance in transverse section. 

Calices are relatively deep ( 1;2 to 11/2 times the 

diameter) with vertical sides and flat bottom. 

Internal features.-Septa are radially arranged; 

major septa long or short, minor septa short. Zigzag 

carinae are present on peripheral parts of septa, 

commonly strongly developed as flanges. On the sides 

of the septa and in properly located longitudinal sections, these appear as low ridges directed inward and 

upward from the periphery and more steeply inclined at the periphery. 

The dissepimentarium is formed of a single peripheral layer of normal, globose dissepiments, commonly augmented by additional discontinuous layers 

of smaller dissepiments. Transverse sections give the 

impression of an inner wall at the inner boundary of 

the principal layer of dissepiments. Longitudinal sections show a peripheral row, in some forms with one 

or even two inner rows of smaller dissepiments that 

are commonly discontinuous. The inner edge of each 

dissepiment rests on the next lower dissepiment; the 

outer edge rests on the corallite wall or an adjacent 

dissepiment; there are no horseshoe or sigmoidal dissepiments. Additional dissepiments fill the lateral 

supports. 

Tabulae are thin, usually complete, commonly convex but shape varies widely in some species (fig. 23). 

Increase.-lncrease is lateral in all known species. 

Offsets project away from the protocorallite at a 

high angle and turn upward and expand to mature 

diameter in 1 em or less. Protocorallite diameter is 

not aff·ected by the offset. The lateral offsetting 

stages are similar in the various species except for 

minor details. Illustrations of offsetting stages are 

shown on plates 14, 15, 20, 22, and others. 

Microstr-uctur-e.-As for f~amily. 

Discuss-ion.-Acinophyllum was described by McLaren (1959) to include species previously assigned 

to Synaptophyllum Simpson (1900). 

McLaren cleared up many misconceptions regarding the genus Synaptophyllum. He redescribed some 

of the original material of Diphyphyllum arundinaceum Billings (type species of Synaptophyllum by 

original designation), and thereby established Synaptophyllum as a stauriid (sensu Hill, 1956b) rather 

than a disphyllid. Simpson's description of Synaptophyllum was based on "Eridophyllum simcoense 

(Billings)" (actually specimens of Acinophyllum 

segregatum) with normal dissepiments, which he 

listed as an example of his new genus. Many subsequent workers accepted Simpson's concept of the 

genus (hased on A. segregatum), and no valid revision was made before McLaren's work. 

The issue was further confused, however, by Lang 

and S.mith (1935a, p. 561-562) who illustrated· a 

specimen with horseshoe dissepiments as Synaptophyllum arundinaceum. Although Lang and Smith 

3 

4 

2 

5 

FIGURE 23.-Morphology of tabulae in Acinophyllum spp. 

Diagrammatic representation of five types described in 

text. 
SYSTEMATIC PALEONTOLOGY 55 

did not indicate the source of their specimen, McLaren (1959, p. 29) showed that it probably was from 

Upper Devonian coral collections from the Mackenzie River Valley (Canada), later described by 

Smith (1945). McLaren (1959, p. 29) redescrihed 

the Smith species as Phacellophyllum fenese. 

The Synaptophyllu.m conoepts of most workers 

since 1935 have been colored by Lang and Smith's 

work. Stumm (1939, p. 62-63) followed Simpson's 

concept of normal dissepiments, but in later work 

(Stumm 1948b, p. 43-44; 1949, p. 37; and 19·55, 

cards 258-275) followed Lang and Smith',s concept 

of horseshoe dissepiments in his generic diagnoses 

but included species with normal dissepiments in the 

genus. R6zkowska (1953, p. 16), and Schouppe 

(1949, p. 124-127, ~and 1958, p. 230-232) followed 

Lang and Smith. Hill (1956b) in the "Treatise on 

Invertebrate Paleontology," followed Simpson's concept but used Lang and 8mith'.s illustrations. 

McLaren's (1959) concept of Synaptophyllum (no 

di·ssepiments) ba.s·ed on part of Billing's type material is the only one that can be accepted. He reviewed most North American .species of H Synaptophyllum" and assigned them as follows: (1) Those 

fitting Simpson's original (but erroneous) concept 

were assigned to ·a new genus, Acinophyllum. (2) 

Those fitting the Lang and Smith conc·ept were studied in less detail, but were considered to be assignable to Phacellophyllum. (3) Only the type .species 

was assigned to Synaptophyllum. 

Planetophyllum Crickmay (1960) is similar to 

Acinophyllum :in hav,ing a single basic row of dissepiment1s. The dissepiments, however, are elongate 

rather than globose. In ~addition, Planetophyllum has 

very short noncarinate septa and irregular commonly incomplete trubulae. The genus may be a di.sphyllid 

or unrelated to both the Disphyllidae and Craspedophyllidae. 

Acinophyllum differs from Cylindrophyllum Simpson and from Disphyllum deFromental and other disphyllids in its very narrow di.ssepimentarium, basically one dissepiment thick Disphyllum and Cylindrophyllum have wide irregular dissepimentaria 

composed of m·any rows of dis:sepiments and with an 

irl"'egula.r irnner .m:a1rgin. Acinophyllum differs from 

phaceloid phillipsastraeids such a:s Peneckiella Soshkina, Thamnophyllum Penecke, and Phacellophyllum 

Giirich in lacking horsesho·e or :sigmoidal dissepiments and in many other characters. 

Acinophyllum is not Peneckiella.-McLaren 

(1959, p. 22-23) suggested that Acinophyllum might 

be a junior synonym of Peneckiella Soshkina. The 

type specie·s of the latter, Diphyphyllum minus Roemer (1855), has been variously described as having 

norm·al or horseshoe di.ssepiments. Roemer's original 

illust~ations (1855, pl. 6, fig. 12a-c) are rather crude 

sketche1s that suggest the pres:ence of a single row of 

norm.al, peripheral dissepim.ents such as in Acinophyllum. However, Fliigel (1956) redescribed Roemer's types and indica·ted the presence of a single 

row of horseshoe ditssepiments. Lang and Smith 

(1935a., p. 576-577, pl. 35, fig. 3), Schouppe (1958, 

p. 186-204, 229-230, fig. 12-14), and Pickett (1967, 

fig. 14) studied topotyp·e specimens ·and .similarly 

concluded that the species possessed hors·eshoe dissepim,ents. Strusz (1965, p. 554-556) ·summarized 

recent descriptions and described and illustrated 

various modifications of dissepiments in Peneckiella 

and its .allies. He considered horseshoe and peneckielloid dis.sepiments to be characteristic. Scrutton 

(1968, p. 271-273) redes.c.ribed the genus on the 

hasis of the holotype of P. minus and noted that "no 

true horseshoe dissep:ments can he positively identified in the holotype, although they do occur in a subsidi·ary role in the dissepimentari~ of some topotype 

spec.imens" (p. 272). Scrutton considered Acinophyllum a junior synonym of Peneckiella because of the 

"peneckielloid" dissep:iments of the former and other 

gene~al similarities (p. 272-273). 

I consider Acinophyllum to be distinct from Peneckiella in the nature of its dissepiments as well as 

in other character.s mentioned in the frumily discussion. Examination of hundreds of longitudinal sections of s.eVJeral species of Acinophyllum has produced no .suggestion of peneckielloid, sigmoidal, or 

horseshoe dissepiments. (.See 8trusz, 1965, fig. 15, 

for illustrations of these types.) Acinophyllum dissepiments .are invariably globose normal dissepiments, both in the characteristic peripheral row and 

in inner, supplemental rows. My examination of 

Peneckiella materital is more limited but I have soon 

the P. minor topotypes des:eribed by Schouppe 

(1958) and Pickett (1967) m·entioned above and 

have exceHent photographs of the holotype thin sections kindly provided by Dr. Scrutton. In ·addition I 

have examined the sections of Peneckiella minor kunthi and of Sudetia lateseptata of R6zkow.ska (1960), 

both considered Peneckiella by Scrutton (1968, p. 

273). In this material, horseshoe dissepiments are of 

oommon occurrenc-e and nonhorseshoe dissepiments 

tend to be peneckielloid or ·sigmoidal. I consider it 

clear that Peneckiella i:s a phillipsastraeid, whereas 

Acinophyllum is not. Additional family-level characters that separate the two genera are the microstructure and the presence of abundant septal cari-
56 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

/ 

nae in Acinophyllum. (See family discussion.) I have 

emphasized the dissepiments here because most Peneckiella discussions have recognized the paramount 

importance of this character in these forms. 

Acinophyllum seems to be restricted to North 

America and northern South America although the 

detailed morphology of all Old World corals referred 

to Peneckiella is not clear. Soshkina described some 

Peneckiella as having normal dissepiments (for example, P. jevlanensis Bulvanker of Soshkina 1952, p. 

103, 1954, p. 35; P. nalivkini Soshkina 1939, p. 25, 

51, 1952, p. 103; and P. minima of Soshkina 1951, p. 

106, 1952, p. 104), but her illustrations are not clear, 

and it s·eems likely that these are forms with peneckielloid rather than Acinophyllum-type dissepiments. 

Onesquethaw species.-Five species of Acinophyllum are recognized in the Bois Blanc, Onondaga, and 

equivalents :in New York, Ontario, and adjacent 

areas. One species of this age is known from V ene-

~uela. These are listed here and described in following sections. 

A. simcoense (Billings), type species, Bois Blanc 

Formation, Ontario, New York, and northern Ohio. 

A. stramineum (Billings), Edgecliff Member (bioherm facies), Onondaga Limestone, Ontario, and 

New York; Mountain House Wharf Limestone ( ?) , 

Lake Memphremagog, Quebec. 

A. segregatum (Simpson), Edgecliff, Nedrow, and 

Moorehouse Members, Onondaga Limestone, New 

York and Ontario, and Columbus Limestone, Ohio. 

This is the common Onondaga species, and the one 

on which most,-descriptions of the genus and of A. 

simcoense have· been based. 

A. mclareni Fagerstrom, Detroit River Group and 

Moorehouse Member, Onondaga Limestone, Ontario; 

Columbus Limestone, Ohio; the lower part (Onondaga age) of the Boyle Limestone (locally) in Kentucky; and the upper coral zone (Onondaga age) in 

the Jeffersonville Limestone, Falls of the Ohio, 

Kentucky. 

A. stokesi (Milne-Edwards and H.aime), Schoharie Formation, New York; Bois Blanc Formation, 

Ontario, Michigan, and northern Ohio; lower coral 

zone (Schoharie age) of the Jeffersonville Limestone, Falls of the Ohio, Kentucky; upper part 

(Schoharie age) of the Wildcat Valley Sandstone, 

Virginia; and the upper Abitibi River Limestone, 

Hudson Bay Lowlands, Ontario. 

A. vermetum Weisbord, upper? Onesquethaw age, 

northwestern Venezuela. 

Other North American "species".-No other species of Acinophyllum are known, although superficially similar forms of Givetian and Frasnian age 

have beeh commonly assigned to the genus. (See 

McLaren, 1959, p. 27-28 for summary.) The following notes are based on an examination of the type 

suites of specimens and additional specim·ens as 

noted. 

A. crassiseptatum (Ehlers and Stumm), from the 

Potter Farm Formation (Middle Devonian) in Michigan, has thick septa of coarse monacanthine trabeculae, and two or three rows of equal dissepiments, 

and is a true Disphyllum. 

A. ? rectiseptatua (Rominger) , from the Dundee 

Limestone (Cazenovia Stage) in M~chigan, has a 

narrow Acinophyllum-like dissepimentarium but the 

septa are relatively thick and are apparently composed of coar$e monacanthine trabeculae; it m·ay be 

a true Disphyllum and if so is one of the earliest 

known in eastern North America. It is described below because of its pertinence to the question of the 

relationship between the cylindrophyllinids and disphyllids and to the time of entry of the disphyllids 

into eastern North America. 

A. fasciculum (Meek), Frasnian, Devils Gate 

Limestone in Nevada, has peneckielloid and nearhorseshoe dissepiments. This and other characters 

suggest that the species is a phillipsastraeid. It is 

certainly .not Acinophyllum. 

A. accidens (Stumm), Frasnian, Martin Limestone in Arizona, is indeterminate. The holotype consists now of only the two th·in sections illustrated by 

Stumm: The original was crudely silicified, and the 

morphology of the dissepimentarium is totally 

obscured. 

A. camselli (Smith), Frasnian, Northwest Territori,es, Canada, has disphyllid septal structure and is 

a Disphyllum. . 

A. sp. D. McLaren, Frasnian, District of Mackenzie, Canada, is another Disphyllum with narrow dissepi:mentarium and closely appress.ed corallites. 

Other Frasnian "Synaptophyllum" (that is, Acinophyllum) of Smith, 1945, were reas,signed to Phacellophyllum by McLaren (1959, p. 28--30). 

Grahau (1910), Williams (1919), and Northrup 

( 1939) assigned the Silurian Syringopora multicaule 

Hall to Synaptophyllum (Acinophyllum of this pap1er) in the belief that Hall's species had a single, 

peripheral row of normal dissepiments. Reexamination of Hall's material indicates that no dissepiments 

of any kind are present and that S. multicaule is a 

species of Palaeophyllum (Oliver, 1963b, p. G5). 

Columnaria? coralliferum (Hall), from Upper Silurian rocks in New York and Quebec (Oliver, 1963c, 

p. 14), is superficially similar to Acinophyllum and 

does have a single row of peripheral dissepiments. 
SYSTEMATIC PALEONTOLOGY 57 

The dissepiments in C.? coralliferum are vertically 

elongate; in this and in other structures the species 

is quite unlike any cylindrophyllinid. It is likely that 

some of the material assigned to S. multicaule is 

congeneric o·r even conspecific with C.? coralliferum. 

Distribution.-Known Acinophyllum species are 

restricted to rocks of Onesquethaw age in eastern 

North America and Venezuela. They are not known 

from western North America or other Old World 

areas. Other corals assigned to Acinophyllum (or 

"Synaptophyllum" in pre-1959 literature) are rejected as being misassigned or indeterminate. 

Acinophyllum simcoen3e (Billings) 

Plates 8-10; plate 11, figures 1, 2 

1859 Eridophyllum simcoense Billings, p. 132, fig. 

27. 

1863 E. simcoense Billings. Billings, fig. 369 (p. 

366). 

1874 E. simcoense Billings. Nicholson, p. 34-35, pl. 

6, fig. 5. 

not · 1875 E. simcoense Billings. Nicholson, p. 228. (Silurian specimen, probably an Entelophyllum) 

not 1876 Diphyphyllum simcoense (Billings). Rominger 

(see A. segregatum). 

not 1879 E. simcoense Billings. Quenstedt, p. 511, pl. 161, 

fig. 8 (see A. segregatum) 

?not 1883b E. simcoense Billings. Hall (see A. segregatum). 

not 1887 E. simcoense Billings. Davis, pl. 112, fig. 1 

( =.Disphyllum synaptophylloides Stumm, 1965, 

by objective synonymy). 

not 1889 D. simcoense (Billings) . Sherzer (see A. 

mclareni). 

1899 D. simcoense (Billings). Lambe, p. 242-243. 

not 1900 Synaptophyllum simcoense (Billings). Simpson 

(see A. segregatum). 

1900 S. baculoideum Simpson. fig. 35-36 (p. 213). 

1901 D. simcoense Billings. Lambe, p. 161-162, pl. 13, 

fig. 6, a-b. 

not 1938 S. simcoense (Billings). Stewart (see A. 

mclareni). 

not 1944 S. simcoense (Billings). Shimer and Shrock (see 

A. segregatum) . 

not 1949 S. simcoense (Billings). Stumm, pl. 17, fig. 18 

( =Disphyllum synaptophylloides Stumm, 1965, 

by objective synonymy). 

1955 S. baculoideum (Billings). Stumm, card 261. 

part 1955 S. simcoense (Billings). Stumm, card 271-272 

(figs. after Nicholson, 187 4, only). 

?not 1958 S. simcoense (Billings). Cranswick and Fritz 

(see A. stokesi). 

1959 Acinophyllum simcoense (Billings). McLaren, p. 

24-25, pl. 8, fig. 6, pl. 9., fig. 1-2, fig. 7. 

1959 A. baculoideum (Simpson). McLaren, p. 27, pl. 

10, fig. 3-4. 

not 1967 Cylind'rophyllum (Acinophyllum) simco ense 

(Billings). Pickett, p. 56-57, fig. 15 (note correction, Pickett, p. 56; see A. segregatum) . 

Occurrence of type material.-"Rama's Farm; and 

near the town of Simcoe." (Billings, 1859, p. 132), 

southwest Ontario. "Rama's Farm" was in what is 

now Port Colborne and was the source of a large 

number of "Onondaga" corals now known to be from 

the Bois Blanc Formation, although both the Bois 

Blanc and the Onondaga crop out in the immediate 

area. "The town of Simcoe" is certainly an Onondaga 

locality. Because Billings' ilustration is clearly of 

the Boi:s Blanc species, the illustrated specimen is 

presumed to be from the Rama's Farm locality. 

Early Devonian (Emsian). 

The present whereabouts of the illustrated specimen and Billings' other syntypes is unknown, although they may be in the collections of the Geological Survey of Canada. McLaren's (1959) redescription is based on part of Lambe's (1901) material from near Woodstock, Ontario, but I agree 

that this material is conspecific with the specimen 

illustrated by Billings. To stabilize the species and 

thereby the genus, I here select as lectotype the specimen illustrated by Billings, thus affixing the name to 

the Bois Blanc species. As the lectotype is unavailable and probably lost I select as neotype a specimen 

(USGS loc. 5052-SD) from the Bois Blanc Formation at Innerkip, Ontario; it is now deposited in the 

collections of the Geological Survey of Canada and 

numbered GSC 31144 (pl. 8, figs. 1-4). 

Diagnosis.-Phaceloid Acinophyllum with abundant lateral supports and with short major septa extending les:s than half the distance to the axis. 

External features.-Phaceloid colonies as much as 

41 by 43 by 10 em or more in height, length, and 

width. Mature corallites cylindrical, more or less 

straight; they range in diameter from 4 to 10 mm in 

the collection at hand; mean diameter in 15 colonies 

ranges from 4.7 to 8.0 mm (table 17 and fig. 24). 

Increase is lateral at infrequent intervals. Corallites are supported within the colony by lateral projections that are numerous and on all sides of the 

corallites; the frequent projections (pl. 11, :fig. 1) 

give a very rugose appearance to the specimens in 

longitudinal sections and triangular and quadrangular .shapes in transverse sections. Broad, rounded, 

inters·eptal ridges with V-shaped septal grooves are 

characteristic. 

Spacing of corallites within the colony ranges 

from less than one to nearly two diameters. 

Galice deep and steep sided. 

Internal features.-Septa radially arranged. Major septa less than one-half the radius in virtually 

all individuals. Average numbers of major septa per 

corallite in 15 colonies ranges from 21 to 28 table 

17 and fig. 24). Minor septa alternate with the rna-
58 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

jor and are nearly as long. Zigzag carinae are strongly developed on the outer parts of all septa. 

The dissepimentarium is composed of one or two 

rows of globose, normal dissepiments; the inner row 

where present consists of smaller dissepiments than 

the outer row. The lateral supports are filled with 

large dissepiments and extensions of the s·epta. The 

tabula.rium is occupied by thin tabulae which are 

usually complete, variable in shape, and commonly 

convex. 

Discussion.-The above description is based on a 

collection of 23 coralla from the Bois Blanc Formation at Innerkip, Hagarsville, and Port Colborne, 

Ontario, and three coralla from Leroy, N.Y., that are 

presumed to be from the Bois Blanc Formation. 

Counts and measurements are based on samples of 

10 to 26 corallites in 15 colonies (table 17 and fig. 

24). 

A. simcoense is characterized by its short major 

septa and abundant lateral supports that give a boxlike appearance to corallites in transverse sections. 

Other species of Acinophyllum here described have 

N 

n: 

d: 

TABLE 17.-Acinophyllum simcoense (Billings), summary 

data 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

---------- 10 26 15 

x 21.4 27.5 23.8 

OR ----- 20-23 25-29 21.4-27.5 

8 ------- .7 1.7 1.8 

v ------ 2.9 6.5 4.2 7.4 

x 4.7 8.0 5.9 

OR ----- 3.7-5.2 6.6-9.9 4.7-8.0 8 ------- .36 1.05 .99 

v ------ 6.9 15.2 11.1 16.9 

28 I I I I 

X 

X 26-- - X 

X X 

0 

X I(;: 24 ~ -

x~0 0 

X 

22- EXPLANATION - X 

X .x Ontario 

0 Leroy, N.Y. 

D Neotype corallum i I I I 

5.0 20~----~------~-----~------~----~ 4.0 6.0 7.0 8.0 9.0 

J. (mm) 

FIGURE 24.-Mean number of major septa (n) versus 

mean diameter (d) in colonies of Acinophyllum simcoense. 

long major septa in all or many corallites and, except 

for A. stokesi have significantly fewer lateral 

supports. 

The early descriptions of "Eridophyllum" or "Diphyphyllum" simcoense by Billings, Nicholson, and 

Lambe, were based on Ontario .specimens, mostly 

from what is now known as the Bois Blanc Formation. Workers who based their descriptions of the 

species on m:aterial from outside Ontario were usually dealing with one of the species of Onondaga age 

so that the early concept of the species was virtually 

the pres.ent generic concept. 

Synaptophyllum baculoideum Simpson (1900, p. 

213; herein, pl. 8, figs. 5-7) was established by the 

pubHcation of drawings of two transvers·e sections 

of individual corallites. Stumm (1955, card 264) and 

McLaren (1959, pl. 10, figs. 3, 4) published photographs of the syntypes including transvers1e sections 

of two additim:al corallites, and McLaren (1959, p. 

27) provided the first description of the syntypes. 

None of Simpson's material other than the me;ntioned thin sections is known to ·exist, but several 

larg"le parts of colonies found in the New York State 

Museum are probably from the same locality (2.5 to 

3 miles [ 4 to 4.8 km] east-northeast of Leroy) and 

are conspecific with the types. S. baculoideum as interpreted from Simpson's material differs from S. 

simcoense only in its large corallite size (diameters 

6.0 to 7.4 mm). Other specimens from the Leroy 

locality are also large (including colonies 50 and 52; 

pl. 9, figs. 3-5), corallite diameter in colony 52 ranging from 6. 0 to 8. 9 mm. These specimens agree in all 

respects with Simpson's material and can be considered topotypes. At the same time they cannot be 

separated from A. simocense. Size variation in that 

species is extensive (fig. 24) and, whereas specimens 

that could be referred to S. baculoideum are toward 

the large end of the size range, they cannot be separated from the mass of colonies studied. Figure 24 

shows the size variation in specimens from Ontario 

and Leroy, N.Y. 

The type sp·ecimen of S. baculoideum and the additional Leroy specimens were cataloged as being from 

the "Onondaga Limestone," but recent work showing 

that the Bois Blanc Formation extends into New 

York (see stratigraphy section) makes this assignment questionable. The Bois Blanc Formation crops 

out in several places just north and northeast of 

Leroy, and it now seems probable that the New York 

specimens of A. simcoense are from the Bois Blanc 

Formation as are the Ontario ones. 

The synonymy presented for A. simcoense is as 

complete as is justified by the need to clear up the 
SYSTEMATIC PALEONTOLOGY 59 

distribution record. Original hypotypes have been 

studied, where possible, but ·many of the assignments are based on the published illustrations or 

description where these are clear and specimens are 

lost. "A. simcoense" was something of a catchall 

term, and the list of accepted and rejected assignments is complete only for material described from 

rocks of Onesquethaw age in eastern North America. 

Distribution.-Bois Blanc Formation (Lower Devonian, Emsian), Ontario, New York, and Ohio. The 

probable assignment of so·me New York specimens 

to the Bois Blanc was discussed in an earlier paragraph. Principal occurrence~s of described specimens 

are at Innerkip, Hagarsville, and Port Colborne, Ontario, and Leroy, N.Y. 

Material studied.-Southwest Ontario: Neotype, 

GSC 31144; hypotype of Lambe (1901) and McLaren (1959), GSC 3436; illustrated specimens (six), 

USNM 162595-98, 27961; other measured specimens 

(eight), USNM 162599-606; others, USNM 162607 

and eight more. Leroy, N.Y.: illustrated specimens 

(four), NYSM 334, 335 (holotype of Synaptophyllum baculoideum Simpson), 12811-13. Lake County, 

Ohio : two specimens from core. 

? 

? 

not 

? 

? 

? 

not 

not 

part 

part 

not 

Acinophyllum stramineum (Billings) 

Plates 12-14 

1859 Diphyphyllum stramineum Billings, p. 135. 

1874 D. stmmineum Billings, Nicholson, p. 33, pl. 5, 

fig. 6 (unrecognizable). 

1874 D. gracile McCoy, Nicholson, p. 33-34, pl. 5, fig. 

5 (unrecognizable). 

1882 D stramineum Billings, White (see (A. segregatum). 

1883b D. stramineum Billings, Hall, p. 261, pl. 9, fig. 

2 (unrecognizable) . 

1926 Diphyphyllum vermetum Weisbord, p. 5-6, pl. 1, 

figs. 6, 7, pl. 2, fig. 1. (see description of this 

form below) . 

1943 Synaptophyllum vermetum (Weisbord), Wells, 

p. 96-97, pl. 10, figs. 3, 4 (see A. vermetum). 

1945 Disphyllum [Synaptophyllum] stramineum (Billings), Smith, p. 23-24, pl. 13, figs. 1, 2, 4-12 

(par t=Phacellophyllum tructense McLaren, 

1959; figs. 3a-c are reproductions of Simpson's 

figures of S. simcoense = S. segregatum) . 

1952 Thamnophyllum stramineum (Billings), Soshkina, p. 85. 

1955 S. stramineum (Billings), Stumm, cards 273-

275, fig. after Nicholson, 1874, only. 

1959 Acinophyllum stramineum (Billings), McLaren, 

p. 25-26, pl. 9, figs. 3, 4, text fig. 8 [not pl. 

10, fig. 5, which is A. segregatum]. 

1960b T. stramineum (Billings), Spassky, p. 48-49, pl. 

15, figs. 1-3. 

Occurrence of type material.-"Common in the 

Corniferous limestone, lot 6, con. 1, Wainfl.eet." 

(Billings, 1859, p. 136.) This is 2.8 to 3 miles ( 4.5-

4.8 km) west of Port Colborne (Weiland Canal) 

and is probably identical to my locality C10. McLaren (1959) redescribed and designated as lectotype one of Billings syntypes. The lectotype (pl. 12, 

figs. 1-3) is from the bioherm facies of the Edgecliff Member. Middle Devonian (lower Eifelian). 

Diagnosis.-Acinophyllum with only rare lateral 

supports. 

External features.-The appearance of colonies of 

A. stramineum must be deduced as no coralla are 

known in which the individual corallites are in their 

original relative positions. Colonies were presumably phacelojd, but lateral supports are rare; apparently for this reason coralla are small and commonly preserved in a broken condition. 

Individual ~orallites are cylindrical except for a 

brief, initial conical stage. Mean corallite diameter 

in eight colonies ranges from 2.9 to 4.7 mm (mean 

3.6 mm) table 18 and fig. 25). 

Increas~e is lateral at infrequent intervals; expansion is rapid in new individuals and the conical stage 

is short. Except for very rare lateral supports the 

basal attachments are the only points of individual 

support. 

22r-----.-----.-----.---. 

20 

18 

16 

2.0 3.0 4.0 

d (mm) 

EXPLANATION 

0 Acmophyllum strammeum 

+ A. vermefum 

X D1sphyllum'? rect/septatum 

5.0 

FIGURE 25.-Mean number of major septa ( n) versus mean 

diameter (d) in colonies of Acinophyllum stramineum, A. 

vermetum, and Disphyllum? rectiseptatum. 
60 SYSTEMATIC PALEONTOLOGY 

TABLE 18.-Acinophyllum stramineum (Billings) and A. vermetum (Weisbord), summary data 

A. stramineum 

Intracolony 

Minimum Maximum 

N ------------- 15 22 n: x ----------- 16.9 20.8 

OR --------- 15-19 20-22 s ----------- .7 1.2 

v ---------- 3.3 6.7 

d: x ----------- 2.9 4.7 

OR --------- 2.9- 3.7 4.1- 5.4 s ----------- .3 .5 

v ---------- 7.7 15.6 r -------------- .03 .69 

p -------------- .90 <.01 

Exteriors are marked by gentle (apparently) 

widely spaced rugae in addition to the usual septal 

grooves and fine growth lines. 

Internal features.-Septa radially arranged. Ma ... 

jor septa range in length from one-fourth to 

three-fourths the corallite radius. Average number 

of major septa per corallite in 8 colonies ranges 

from 17 to 21 (fig. 25) with comparatively little 

variation in any one colony (table 18) . Minor septa 

alternate with the major and are about two-thirds 

as long. Septa are slightly amplexoid as in A. 

segregatum. 

Zigzag carinae are well developed on peripheral 

parts of septa in most corallites (pl. 14, fig. 8.) 

The dissepimentarium is composed of a single 

row of steeply inclined normal dissepiments. A few 

small dissepiments occur on the axial side of the 

single row, but these do not form a second row even 

locally. 

The tabularium is occupied by thin, usually complete, tabulae which vary in shape from convex to 

concave. The same five types described for A. segregatum (fig. 23) are found in A. stramineum (pl. 

12, fig. 4; pl. 13, figs. 7, 8). All types are found in 

many colonies ; one to five types may occur in a 

single corallite. Tabulae spacing in 8 colonies ranges 

from 5 to 16 per 5 mm to 12 to 22 per 5 mm. Average spacing is approximately 10 per 5 mm. 

The epitheca is thin, averaging 0.1 mm. Microstructure of all skeletal tissue is the same as described for the genus. 

Variation.-Intercolony and intracolony variation 

based on 15 to 22 individuals in 8 colonies is shown 

by scatter diagram (fig. 25) and in table 18. Colonies vary in mean corallite diameter and mean 

number of major septa and in the correlation coefficients of these two. Diameter-septal number correlation based on colony means, however, is very strong 

(r<0.01). 

In these and other characters, variation is much 

like that noted for A. segregatum and is discussed 

A. vermetum 

Intercolony PRI PRI Mean 21598 27637 

8 7 6 

18.1 20.7 19.8 

16.9-20.8 20-22 19-21 

1.2 .95 .75 

5.5 6.5 4.6 3.8 

3.6 4.5 4.5 

2.9- 4.7 4.1- 5.2 4.1- 5.0 

.5 .4 .3 

12.5 14.2 8.6 6.4 

.95 

<.01 

in the chapter on intercolony and intracolony variation. 

Discussion.-The above description is based on a 

large collection from the Onondaga Limestone in 

New York and Ontario. Measurements are based on 

15 to 22 individuals in each of 8 colonies. 

McLaren ( 1959) designated a lectotype and described and illustrated it, giving a firm basis for the 

present description and discussion. 

A. stramineum is very close to A. segregatum in 

most morphologic details and can be absolutely set 

off in none. Some workers have synonomized the 

two species (Rominger, 1876; Lambe, 1899, 1901; 

Smith, 1945), and others have noted their similarity. 

Stumm (1955, card 272) stated "The internal structure of S. simcoense [meaning A. segrega,tum] is 

identical with that of S. stramineum. The two 

species can be distinguished only by the connecting 

processes present on S. simcoense." 

Both species are extremely variable, and their 

separation has not been well defended because of 

limited numbers of specimens. Billings separated 

them on the presence or absence of lateral supports 

and this was followed by Stumm. Although the frequency of supports in A. segregatum varies widely, 

colonies of the A. segregatum-stramineum complex 

have either a lot of them or very few of them, and 

it is not difficult to separate colonies into groups on 

this basis. "Difficult identifications" invariably involve badly crushed specimens or small fragments. 

This morphologic difference accounts for significant 

differences in preservation. A. segregatum is commonly preserved with the individual corallites in 

their original relative positions. In contrast, A. 

stramineum corallite.s are almost invariably disoriented. The lateral processes of A. segregatum 

made the coralla rigid enough to be overturned without complete breakup, whereas A. stramineum 

coralla easily fell apart. The species are commonly 

associated in the Edgecliff bioherm facies where the 

contrast in preservation condition is marked and 
SYSTEMATIC PALEONTOLOGY 61 

provides a useful means of identification in the field. 

A. stramineum is very rare or absent from other 

members of the Onondaga although A. segregatum 

is generally common. 

In addition to the general lack of lateral supports 

and the disoriented preservation condition, the following observations can be made of A. stramineumA. segregatum differences: A. stramineum tends to 

have longer minor septa, stronger carinae, more 

closely spaced tabulae, and just a single row of 

dissepiments. 

A. stramineum differs from other described 

species of the genus ~as follows: A. stokesi, simcoense 

and mclareni have abundant or common lateral supports. In addition A. simcoense is characterized by 

short major septa and A. mclareni by long ones. 

Thamnophyllum stramineum (Billings) of Soshkina (1952) and of Spassky (1960) were based on 

Smith's (1945) erroneous generic concept and are 

not conspecific or congeneric with Billings species. 

Distribution.-Edgeclitf Member, Onondaga Limestone (Middle Devonian, lower Eifelian), New York 

and Ontario. A. stramineum is common in the bioherm facies of the Edgecliff Member in eastern and 

western New York and in the Niagara Peninsula 

of Ontario. One corallum (USNM 163380) was collected from the biostrome Edg.ecliff at Hagarsville, 

Ontario (USGS loc. 5028-SD). 

Several specimens of Acinophyllum, probably this 

species, are known from the Mountain House Wharf 

Limestone, Lake Memphra.magog, Quebec. 

Material studied.-Southwestern Ontario: Lectotype, GSC 3431; others, USNM 163381; biostrome 

specimen, USNM 163380; 1 additional specimen. 

New York: illustrated specimens ( 5) , USNM 

162608-09; NYSM 12806-08; other measured specimens (6), USNM 162610-14, NYSM 12809; others 

(19). Lake Memphramagog, Quebec: USNM 

172178-179. 

Acinopbyllum segregatum (Simpson) 

Plates 15-22; plate 11, figure 3 

? 1874 D. arundinaceum (Billings). Nicholson. 

part 1876 Diphyphyllum 8imcoen8e (Billings). Rominger, 

p. 123-124, pl. 46, figs. 2?, 3, 4. 

1879 Eridophyllum 8imcoen8e Billings. Quenstedt, p. 

511, pl. 161, fig. 8. 

? 1882 D. 8tramineum Billings. White, p. 388, pl. 48, 

fig.l. 

? 1883b E. 8imcoen8e Billings. Hall, p. 262, pl. 9, fig. 1. 

1900 Synaptophyllum 8imcoen8e (Billings). Simpson, 

fig. 33,34 (p. 213). 

1900 S. 8egregatum Simpson, fig. 37 (p. 213). 

? 1901 D. arundinaceum Billings. Lambe, p. 162-163, 

pl. 14, figs. 1, 1a, lb. 

1944 S. 8imcoen8e (Billings). Shimer and Shrock, p. 

95, pl. 30, figs. 5-7. 

part 1945 Di8phyllum [Synaptophyllum] 8tramineum (Billings). Smith, pl. 13, fig. 3a-c only (see McLaren, 1959, for description and discussion of 

Smith's other material). 

1955 S. 8egregatum (Billings). Stumm, card 270. 

part 1955 S. 8imcoen8e (Billings). Stumm, card 271-272 

(figs. after Rominger, 1876 only). 

1967 Cylindrophyllum (Acinophyllum) 8 i m co en 8 e 

(Billings). Pickett, p. 56-57, fig. 15. 

Occurrence of type material.-"Onondaga Limestone, Clarksville, Albany Co., N.Y." (Clarke and 

Ruedemann, 1903, p. 58.) Almost certainly the bioherm facies of the Edgecliff Member, Onondaga 

Limestone. Middle Devonian (Eifelian). 

Diagnosis.-Phaceloid Acinophyllum with numerous lateral supports and with both long and 

short major septa. 

External features.-Colonies small to large, lenticular, as much as 100 c.m or more in diameter and 

as much as 30 em or more in height; composed of 

radiating or subparallel corallites. Mature corallites 

range in diameter from 2.3 to 8.9 mm. In a single 

colony, size variation is limited (table 8) ; mean 

diameter in 58 colonies ranges from 2.8 to 6.8 mm 

(fig. 16). 

Increase is lateral at infrequent intervals; occasional individuals show successive offsets only 1 em 

apart, but in most individuals the offset interval is 

5 em or more. Expansion is rapid, and the conical 

stage is limited to the basal one-half em. Lateral 

supports are present in all colonies and generally 

found on all corallites but may be closely or widely 

spaced (pl. 11, fig. 3; pl. 21, fig. 6) and limited to 

one or two vertical rows on each corallite. Corallites 

are markedly rugose at irregular intervals; the lateral supports are normally an extension of the rugae. 

Spacing of corallites within a colony ranges from 

less than one to nearly two di,ameters. 

Calices are known only in sections. They are deep 

( 1 to 11;2 diameters) with steep sides from which 

the septa project only slightly. 

Internal features.-S.epta radially arranged. 

Major septa range in length from one-fourth to 

three-fourths the corallite radius, commonly less 

than one-half, but longer septa characterize an estimated 10 to 50 percent of the corallites in all sections studied. Average number of major septa per 

corallite in 58 colonies ranges from 16 to 28 (fig. 

16) with comparatively little variation in any one 

colony (table 8). Minor septa alternate with the 

major and are one-half (or less) as long. The septa 

are amplexoid, at least in some specimens; numerous 
62 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

eccentric longitudinal sections show that the major 

septa are longest on the surfaces of tabulae and 

slightly or markedly withdrawn in the sp·ace between tabulae. Septa are very thin ( 0.03-0.10 mm) 

and often slightly dilated in the peripheral region. 

Zigzag carinae are strongly or weakly developed on 

the outer parts of the septa. 

The dissepimentarium is composed of one row of 

globose, steeply inclined dissepiments. A S·econd, 

commonly discontinuous row of smaller dissepiments is locally developed in many individuals in 

many colonies (pl. 21, fig. 3). More rarely a third 

row is present. Occasional specimens lack dissepiments along part of their periphery, generally in the 

early conical stage of new individuals (pl. 20, fig. 3). 

In the cylindrical stage the number of rows of dissepiments is roughly related to size. Areas of no 

dissepiments are found only in colonies with the 

smallest diameters ; the second and third rows are 

more common in colonies of large individuals. The 

lateral supports consist of a lateral extension of 

normal septal and dissepimental tissue within the 

out-bent epitheca. 

The tabularium is occupied by usually complete, 

thin tabul3!e which vary in shape from quite convex 

to quite concave. Five basic types can be recognized 

(fig. 23) of which type 3, simple convex, is most 

common. All types are found in most colonies; one 

to five types may occur in a single longitudinal section of a single corallite. Spacing of tabulae varies 

widely within individuals as well as in the species as 

a whole. Spacing in colonies ranges from 2 to 10 per 

5 mm to 10 to 19 per 5 mm. Average spacing is 

between 5 and 10 per 5 mm. 

Microstructure.-To the family description can 

be added the following. The epitheca is thin, averaging 0.1 mm; thicker on the lateral supports. The 

septal trabecular zone (dark, axial area in transverse section) is approximately 0.03 mm thick. The 

septa average 0.1 mm thick at the periphery but thin 

toward the axis. 

Variation.-Individual and colony variation is extreme in this species and is the subject of detailed 

description and discussion in the section titled 

"Intracolony and Intercolony Variation." Morphologic characters other than size and number of septa 

are difficult to treat quantitatively, but certain observations made in the systematic description can 

be summarized here. The species as a whole is quite 

variable in most characters. Intracolony variation is 

more limited than intercolony variation in many 

features, but even individual corallites are quite 

variable in some. 

Intracolony variation is relatively low in exterior 

ornamentation (rugae and so forth), size and shape 

of dissepiments as well as number of septa and diameter, as previously discussed. All of these and colony 

size vary widely in the species as a whole. 

Variation in length of major septa and shape and 

sp·acing of tabulae is most notable within a colony 

although these vary within individuals also. 

Discussion.-The above description is based primarily on a large collection (made by the author) 

of colonies and parts of colonies from the Onondaga 

Limestone in New York. The nucleus of this collection is 58 .specim·ens, which were analyzed to determine intercolony and intracolonial variation. Supplementary material collected in Ontario and New 

York and specimens from the New York State and 

U.S. National Museums and the Geological Survey 

of Canada provided additional data on morphology 

and geographic and stratigraphic distribution of the 

species. 

This is the common Acinophyllum in the Onondaga Limestone, occurring in most facies of the 

Edgecliff, Nedrow, and Moorehouse Members 

throughout the extent of the formation. 

A. segregatum differs from other Onesquethaw 

spec:ies of the genus in these characters : major 

septa are long or short in different individuals; 

lateral connections are common but tend to be thin 

and have a narrow base. 

In contrast, A. simcoense has short septa and A. 

mclareni long septa; A. stramineum virtually lacks 

lateral connections; A. stokesi and A. simcoense 

have very broad based, thick lateral connections that 

are strikingly abundant. 

N omenclatorial problems.-Synaptophyllum segregatum Simpson was established by the publication 

of a drawing of a thin section of two connected 

corallites. The thin section from which the drawing was made is pres·erved (pl. 15, figs. 4, 6) ; it is 

the only remnant of the specimen (s) on which 

Simp•son based the species. 

Stumm (1955) published a photograph of the 

whole thin section including four additional corallites, and observed that the species was "probably 

conspecific with S. simcoense (Billings)." Because 

the accepted concept of S. simcoense at that time 

was based on the common Onondaga species (here 

assigned to S. segregatum), I am in agreement with 

this statement. 

McLaren (1959) published another, larger photograph of the holotype section and with some question placed the species in synonomy with S. stramineum ( p. 25-26) . 
SYSTEMATIC PALEONTOLOGY 63 

To most previous workers the "Onondaga" has 

included both the Bois Blanc and Onondaga of this 

report. They have tended to recognize Billings' two 

species and to differentiate between them on the 

basis of presence or absence of lateral connections. 

In consequence, the species here described as A. 

segregatum has commonly been included in A. 

simcoense. In the literature on New York most references to "S. simcoense" can be safely translated 

as A. segregatum. Thi:s is not true for reports of 

Ontario occurrences as specimens of Acinophyllum 

simcoense are as common there as are specim•ens of 

A. segregatum. 

The holotype thin section of A. segregatum shows 

the important characters of the common Onondaga 

species very well; the six corallites show both long 

and short major septa and a lateral connection. Although it would be preferable to 'base the most 

abundant known species of the genus on a large 

corallum from a known locality and stratigraphic 

position, I have no doubt of the identity of the holotype with the common species and little doubt of 

its provenance. The matrix of the holotype slide is 

characteristic of the Edgecliff bioherm facies. The 

Edgecliff outcrop belt runs through Clarksville 

(the type locality for the species) ; bioherms are 

known w:ithin a few miles of the village and may 

have occurred much closer at localities no longer exposed. 

Distribution.-Middle Devonian (Eifelian), New 

York, Ontario, and Ohio. A. segregatum is the most 

common and widely distributed colonial rugose coral 

in the Onondaga Limestone of New York and 

Ontario. It is common or abundant in all of the Edgecliff bioherms and at all outcrops of the Moorehouse Acinophyllum-Eridophyllum bed. It is of frequent occurrence, not everywhere common, in other 

facies of the Edgecliff Member and in the western 

coral facies of the Moorehouse Member. The species 

is less common in the Nedrow Member and .i:s not 

known from the Seneca Member. 

In the Niagara Peninsula of Ontario, A. segregatum is common in the Edgecliff Member (both 

facies) and was collected from probable Moorehouse 

beds in the Simcoe area. 

A., segregatum occurs in the Columbus Limestone 

in the Ohio, but its stratigraphic and geographic 

range there is yet to be worl~ed out. It may be common in the lower Columbus coral bed (Zone C) in 

the outcrop belt from northern Pickaway County to 

Delaware and has been found in the same stratigraphic position in the subsurface of Morgan 

County (southeastern Ohio). It occurs throughout 

the lower 86 feet (27m) of the Columbus-Onondaga 

in the subsurface of Lake County (northeastern 

Ohio). 

Mat erial studied.-N ew York: Holotype, NYSM 

336; illustrated and measured specimens (13), 

USNM 162615-22, NYSM 12780-84 illustrated 

(14), USNM 162623-28, NYSM 12785-92; measured (45), USNM 162629-59, 172246, NYSM 

12793-805. Ontario: Illustrated sp·ecimen:s (3), GSC 

31162-64. The distribution of this species as indicated on tables 41-44 :and 46, is based on 21 additional identified speci,mens from southwestern Ontario and more than 150 additional identified specim·ens from New York. 

Acinophyllum rnclareni Fagerstrom 

Plates 23, 24 

?1889 Diphyphyllum simcoense (Billings). Sherzer, p. 93-

95, fig. 

1938 Synaptophyllum simcoense (Billings). Stewart, p. 43, 

pl. 8, fig. 5-6. 

1961 Disphyllum sp. A Fagerstrom, p. 13-14, pl. 3, fig. 1-3. 

1961 Acinophyllum mclareni Fagerstrom, p. 14, pl. 3, fig. 

18-20. 

Occurrence of type material.-Formosa reef 

facies of the Detroit River Group, Bruce County, 

Onta.rio (Fagerstrom, 1961, p. 4, 14). Middle Devonian (Eifelian). 

Diagnosis.-Acinophyllum with uniformly long 

major septa extending approximately two-thirds the 

distanc,e to the axis. 

External features.-Colonies large, as much as 

25 c.m or more high and one to several feet in 

diameter. Corallites are stra;ight and parallel with 

relatively rare lateral supports. Mature ( cylindrical) corallites range in diameter from 4.0 to 7.6 

mm; mean diameter in 10 cor all a ranges from 4. 7 to 

6.7mm. 

Increase is lateral and apparently infrequent; offsetting interval not known. 

Corallit•e exteriors are marked by at least two 

orders of encircling rugae (growth lines) and by 

fine, longitudinal septal grooves. 

Inte1·nal features.- Septa radially arranged. 

Major septa relatively uniform in length, extend 

from 0.6 to 0.9 the distance to the axis, rarely 

shorter. Average number of major septa in 10 

coraUa ranges from 20 to 28 (fig. 26). Minor septa 

alternate with the major and are commonly onehalf or less as long. Zigzag carinae weakly to strongly developed. 

The dissepimentarium :is composed of one or two 

rows of globose dissepiments forming a continuous 

outer zone. Tabulae are commonly complete, broad-
•c: 

64 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

28,------.-------.-------.----~~------~ 

' ' _ _! __ L ~'J·~' 

26 I241--

22 I20-

Boyle Limestone, r- }J 9 

Kentucky Q ___ T __ / 

EXPLANATION ;

1 /J>'~mherstt[erg - _ Formation of 

0 Holotype corallum / /Detroit River Group 

(y/ 

_....,.,.,.,()1 

..... -- / ... tO ,,..- ~ O,/''"..,... Mo_orehouse Member of 

"""" Onondaga Limestone 

-

-

/x~ 

/ /Jeffersonville Limestone. Kentucky I X I ........ 

° Columbus Limestone, Ohio -

18~-----'~----~'~----~'------~'~----~ 3.0 4.0 5.0 6.0 7.0 8.0 

d (mm) 

FIGURE 26.-Mean number of major septa (fi) versus mean 

diameter (d) in colonies of Acinophyllum mclareni from different stratigraphic units. 

ly arched with axial s-ag and steeply descending 

periphery; four to eight per 5 mm in the specimens 

studied. 

Discussion.-The above description is ba•sed on 

specimens from five different formations and areas 

as indicated in figure 26. The species i·s rare in the 

Onondaga, occurring only in the Moorehouse Member in Ontario. The three known Moorehouse specimens are fragments (as much as 10 em) of larger 

coralla and only one is well preserved. The spec:ies 

is distinguished by its long septa and is constant in 

most characters over its area of distribution. Although available samples from the Moorehouse 

Member and the Columbus and Jeffersonville-Limestones are very small, it is probably significant that 

mean diameters are smaller in these than in specimens from the Detroit River Group and Boyle 

Limestone (fig. 26). This may relate to environm·ent 

as the former are from limestones, whereas the latter are from dolomites. The species was originally 

based on specimens from the Formosa re·ef facie•s 

of the lower Detroit Rive•r (Fagerstrom, 1961). 

This is a dolomite, and corallite diameters cited by 

Fagerstrom ( p. 14) are correspondingly large. 

A. mclareni differs from all other Onesquethaw 

species of the genus in its consistently long septa. 

A. segregatum, stramineum, and stokesi all have 

long septa in some corallites, but septa are commonly short. 

Sherzer ( 1889) described a drift specimen from 

near Ann Arbor, Mich., as Diphyphyllum simcoense 

(Billings). Major septa "extend about two-thirds of 

the distance from the outer wall to the center, rarely reaching it." It seems likely that this is a specimen of A. mclareni from the Detroit River of either 

northern M.ichigan or Ontario. 

Stewart's Synaptophyllum. simcoense agrees with 

A. mclareni in its morphology and the characteristic 

long septa are well shown in the illustrations (Stewart, 1938, pl. 8, fig. 5; herein, pl. 24, fig. 4). 

Fagerstrom's original des.cription of the species 

was based on specimens from the Formosa reef facies of the Detroit River in Bruce County, Ontario, 

some 90 miles (144 km) north of the London-Woodstock area. The present description agrees with 

Fagerstrom's in most respects. It is broadened to 

include much greater size variation than could be 

found in the type area. In addition the consistently 

long septa are emphasized as a character distinguishing this from other species with which it 

might be confused. 

Distribution.-Middle Devonian (Eifelian) ; Detroit River Group and Onondaga Lim·estone, Ontario; Columbus Limestone, Ohio; Boyle and J effersonville Limestones, Kentucky. The known Detroit 

River occurrences of A. mclareni are all :in beds 

thought to be equivalent to the Amherstburg Formation in Michigan and the Edgecliff Member in New 

York-Ontario : ( 1) The Formosa reef facies is considered lower Detroit River or Amherstburg by 

Fagerstrom (1961, p. 45; written commun., 1966). 

(2) The Gorri1e exposures (loc. C36) were convincingly shown to be lowest Detroit River or upper 

Bois Blanc by E. W. Best (unpub. PhD. thesis, 

Univ. Wisconsin, 1953, p. 87-88) ; the corals are 

clearly post-Bois Blanc (Oliver, 1960a, p. B173-

Bl74) .and the exposure is assigned to the lower 

Detroit River, Amherstburg Formation. (3) Ehlers 

and Stumm (1951b, p. 1886-1887) recognized the 

pressence of probable Amherstburg in the vicinity 

of Ingersoll, Ontario (USGS lo.c. 5051-SD) ; one 

specimen of A. mclareni was collected by the author 

from these beds. 

The Onondaga specimens of A. mclareni are from 

the Moorehouse Member at Ontario localities near 

Dunnville (loc. C18) and south of Hagarsville (loc. 

C23). This, coupled with the Detroit River information, suggests a range from Edgecliff to Moorehouse 

equivalents in Ontario. 
SYSTEMATIC PALEONTOLOGY 65 

A. mclareni is known from the lower Columbus 

Limestone (Zone C) in the outcrop belt at Bellepoint, Ohio, and from the same position :in the subsurface of Lorain County, Ohio. The species also 

occurs in the lower Boyle Limestone, in Lincoln 

County, Ky., and in the upper coral zone of the J effersonville Limestone at the Falls of the Ohio, Kentucky -Indiana. 

Throughout the area of known distribution, A. 

mclareni is present and most co·mmon in units of 

Edgecliff age, but it ranges upward into units of 

Moorehouse age in southwestern Ontario. 

Material studied.~Ontario: Detroit River-Bolotype, UMMP 36120; illustrated specdmens (two), 

USNM 162660-61; others, eight specimens from 

Gorrie. Moorehouse-illustrated (one), USNM 

162662; other measured specimens (two), USNM 

162665-66; others, one specimen. Ohio: Illustrated 

(two) OSU 17735 (hypotype of Stewart, 1938), 

UMMP 60484 ; others, three spe,cimens from Zone C 

at Bellepoint. Kentucky: Boyle-Illustrated, USNM 

162664; other measured (two), USNM 162668-69; 

others, four specimens. Jeffersonville-Illustrated, 

USNM 162663; other measured, USNM 162667. 

Acinophyllum stokesi (Milne-Edwards and Haime) 

Plates 25-28 

1851 Lithostrotion stokesi Milne-Edwards and Haime, 

p. 440, pl. 20, fig. 2. 

1860 L. stokesi Milne-Edwards and Haime. MilneEdwards, p. 429. 

1887 Eric~Jphyllum o,rundinaceum (Billings). Davis, 

pl. 112, fig. 2 (not Billings, 1859) ; objective 

synonym of Acinophyllum davisi Stumm. 

not 1897 Diphyphyllum stokesi Milne-Edwards and Haime. 

Whiteaves, p. 152, pl. 17, figs. 5-5b. 

not 1906 Columnaria stokesi Milne-Edwards and Haime. 

Whiteaves, p. 343. 

not 1915 C. (Palaeophyllum) stokesi Milne-Edwards and 

Haime. Bassler, p. 261 (synonymy of Ordovician coral) . 

not 1926 C. (P.) stokesi Milne-Edwards and Haime. Hussey, p. 151, pl. 11, fig. 15. 

not 1928 C. (P.) stokesi Milne-Edwards and Haime. 

Troedsson, p. 111, pl. 27, figs. la-d. 

1958 Synaptophyllum simcoense (Billings). Cranswick 

and Fritz, p. 45, pl. 3, figs. 1, 2. 

not 1963 Palaeophyllum stokesi Milne-Edwards and 

Haime. Nelson, p. 31, pl. 6, fig. 6. 

1965 Acinophyllum davisi Stumm, p. 42, pl. 37, figs. 

9-11. 

1968b A. davisi Stumm. Oliver, p. 26, 27, 30 (character tabulations only) . 

1973 A. stokesi (Milne-Edwards and Haime). Easton 

and Oliver, p. 915-918, pl. 1, figs. 1-6. 

Occurrence of type material.-"Carbonifere, 

Amerique du nord: Lac Wennipeg." (Milne-Edwards and Haime, 1851, p. 441). This has commonly been taken to represent an Ordovician coral 

(s·ee "nots" in synonymy) from the Lake Winnipeg 

area, Manitoba, Canada. The holotype was recently 

found by W. H. Easton and redescribed by Easton 

and Oliver (1973). It is clearly a Devonian Acinophyllum and was most likely moved from the Hudson Bay Lowlands (ENA Province) to the Lake 

Winnipeg area by glaciers. The species is known 

only from eastern North America except for this 

type occurrence. 

Diagnosis.-Phaceloid Acinophyllum in which 

corallite diameter is s.ma.Il; dissepiments are small 

except .in lateral connections which are large and 

num·erous; major septa may be long or short. 

External features.-Colonies lenticular, small to 

medium in size, composed of radiating to subparallel, cylindrical cor:allites. The largest specim·ens collected in the Bois Blanc are incomplete, one 10 em 

high and 13 em in diameter, one 23 em .in diameter. 

A corallum from the Falls of the Ohio measures 18 

em high and 28 em in maximum diameter. The largest known fr,agment from the Schoharie Grit is approximately 4 em high and 10 em in diameter. Mature corallites are straight; diameters range from 

3 to 5 mm in New York-Ontario specimens to as 

much as 6 mm in Falls of the Ohio specimens; mean 

diameters in 10 Bois Blanc colonies range from 3.3 

to 4.4 mm, 6 Falls of the Ohio colonies range from 

3.8 to 5.2 mm. Data on these and other colonies are 

shown in tables 19 and 20 and in figure 27. Spacing 

of corallites ranges from 0.5 to 1.5 times the corallite diameter. 

Increase is lateral; offsetting intervals of 1.5 to 

3.5 em have been observed. 

Corallites are gently to strongly rugose. Lateral 

connections are large and numerous and show no 

orientation. 

Calice depth approximately 0.5 to 1.0 time·s corallite diameter. 

Internal features.- Septa radially arranged. 

Major septa range in length from 0.3 to 0.9 the 

corallite radius; long and short septa .appear in different corallites in the same colony. Number of 

major septa ranges from 16 to 21 in Bois Blanc 

specimens, 18 to 23 in Falls of the Ohio specimens; 

mean septal number in 10 Bois Blanc colonies ranges 

from 17.2 to 20.1, in six Falls of the Ohio colonies 

from 18.6 to 21.2. Minor septa alternate with the 

major and extend 0.3 to 0.4 the distance to the axis. 

Carinae are abundant, commonly zigzag, rarely 

yardarm or nearly so; limited to the dissepimentarium. 
66 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

22.---,,--------~,,--------~,--~ 

20 -

IC: 

18 -

16~--~--------~~--------~'--~ 3.0 4.0 5.0 

d(mm) 

EXPLANATION 

x Bois Blanc Formation 

0 Lower Jeffersonville Limestone 

+ Schoharie Grit 

• Abitibi River Limestone 

FIGURE 27.-Mean number of major septa (n) versus mean 

diameter (d) in colonies of Acinophyllum stokesi. 

N 

n: 

d: 

TABLE 19.-Acinophyllum stokesi (Milne-Edwards and 

Haime), summary data, Bois Blanc Formation sample 

Minimum 

---------- 10 

x 17.2 

OR ----- 16-18 s ------- .47 

v ------ 2.6 

x 3.3 

OR ----- 2.7-3.6 8 ------- .24 

v 7.3 

!ntracolony 

Maximum 

20 

20.1 

19-21 

1.33 

7.7 

4.4 

3.9- 5.0 

.57 

13.6 

Inter- Mean colony 

10 

18.9 

17.2-20.1 

1.05 

4.6 5.6 

3.8 

3.3- 4.4 

.38 

10.1 10.1 

TABLE 20.-Acinophyllum stokesi (Milne-Edwm·ds and 

Haime), summary data, lower Jeffersonville Limestone 

sample 

Minimum 

N ---------- 10 n: x 18.6 

OR ----- 18-20 s ------- .50 

v ------ 2.6 

d; x 3.8 

OR ----- 3.0-4.7 8 ------- .32 

v ------ 6.2 

Tntracolony 

Maximum 

20 

21.2 

19-23 

.93 

4.6 

5.2 

4.5- 6.4 

.53 

14.1 

Inter- Mean colony 

6 

20.1 

18.6-21.2 

1.02 

4.0 5.1 

4.8 

3.8- 5.2 

.52 

9.6 10.9 

The dissepimentarium is composed of one row of 

globose dissepiments with a second incomplete row 

developed locally. The large connections are filled 

with larger, more elongate dissepiments in many 

irregular rows. 

Tabulae are complete, commonly broadly arched 

with an axial depression and sharply descending at 

the periphery. Spacing ranges from 6 to 12 per 5 

mm. 

Discussion.-The above description is based primarily on 10 coralla from the Bois Blanc, 1 from the 

Schoharie and 6 from the Jeffersonville. For purposes of n :d analysis, 10 to 20 corallites in each 

colony have been studied, but the Bois Blanc and 

Jeffersonville samples have been kept separate. Bois 

Blanc specimens have somewhat smaller corallites 

with fewer septa than do Jeffersonville specimens 

(fig. 27 and tables 19, 20), but there is considerable overlap in the observed range of these characters and specimens are otherwise very similar. 

Only one Schoharie specimen was sufficiently well 

preserved for detailed study; it is not included in 

either of the above samples, but it is well within 

the Bois Blanc range in both characters (fig. 27). 

The holotype is a fragment containing about 10 

corallites. As discussed above and more fully by 

Easton and Oliver (1973), the holotype probably 

originated in the Hudson Bay Lowl:.:o.nds in northern 

Ontario or adjacent Manitoba (Moose River Basin). 

A corallum of this ·species from the Hudson Bay 

Lowlands (the "upper Abitibi Rive·r Lim·estone") 

was studied and is included in figure 27. This is the 

specimen described and illustrated by Cranswick 

and Fritz (1958, p. 45-46, pl. 3, figs. 1, 2) as Synaptophyllum simcoense. The known presence of the 

species in the area lends support to the argument 

that the Milne-Edwards and Haime specimen was 

derived there also. Acinophyllum stokesi is one of 

several species in the Abitibi River fauna indicating a Schoharie age. Fuller discussion of this point 

will be found in the stratigraphic section. 

The species is also known from the Wildcat Valley Sandstone in southwest ·virginia. All specimens 

are crudely silicified, and they add little to the description. They fall into the Bois Blanc diameter 

range but are too poorly preserved for septal counts. 

Acinophyllum davisi was first described from the 

lower Jeffersonville Limestone (Schoharie equivalent) by Stumm (1965) who selected the specimen illustrated by Davis (1887) as holotype. No 

previous descriptions of this species from New York 

or southwestern Ontario are known; specimens 

have presumably been included in A. simcoense by 

mo·st authors. 

A. stokesi differs from other Onesquethaw species of the genus in these characters: corallites are 
SYSTEMATIC PALEONTOLOGY 67 

small; major septa are long or short :in different 

individuals; lateral connections are very common 

and tend to be thick and have a broad base. In contrast, the associated A. simcoense has large corallites and short septa; A. stramineum lacks lateral 

connections; A. mclareni has long septa; and A. 

segregatum has fewer and thinner lateral connections. 

Within the lower J effersonvilre Formation, "Disphyllum" synaptophylloides Stumm (1965, p. 42) is 

very similar to A. stokesi in size and other external 

features. "D." synaptophylloides can be recognized 

by its peripheral offsetting and by distinctly different internal morphology. This species is not 

known to occur with A. stokesi in other areas. 

Distribution.-Lower Devonian (Emsian), New 

York, Ontario, Virginia, Indiana-Kentucky, and 

northern Michigan. A. stokesi is common in the 

Schoharie, Bois Bl·anc, and lower Jeffersonville Formations, all of Schoharie age. It o·ccurs also in the 

upper Abitibi River L:i·mestone, Hudson Bay Lowlands, Ontario, and in the upper Wildcat Valley 

Sandstone, southwest Virginia, and provides evidence for the Schoharie age of these faunas (See 

stratigraphy discussion.) 

Mat erial studied.-Lower Jeffersonville Limestone, Falls of the Ohio: holotype of A. davisi 

Stumm, MCZ 7827; other measured specimens ( 5), 

USNM 163367-69 and 163378-79. Bois Blanc Formation, Ontario: here illustrated ( 5), USNM 

162670-73, GSC 31143, other measured specimens 

(7), USNM 163370-76; others (10). Schoharie Grit, 

New York: here illustrated (3), NYSM 12810, 

USNM 162674-75. Upper Abitibi River Limestone, 

Ontario: holotype, Paris Museum No. 497a (probably this formation; see discus.sion) ; hypotype (of 

Cranswick and Fritz, 1958), ROM 27064. Wildcat 

Valley Sandstone, Virginia. .Specimen l:isted by 

Kindle (1912, p. 52) as "Diphyphyllum cf. gigas," 

USNM 172165. 

Acinophyllum vermetum (Weisbord) 

Plate 29 

?1859 Diphyphyllum stTamineum Billings, p. 135 (see 

synonymy above). 

1926 Diphyphyllum vermetum Weisbord, p. 5-6, pl. 1, figs. 

6, 7, pl. 2, fig. 1. 

1943 Synaptophyllum ve'"l·metum (Weisbord). Wells, p. 96-

97, pl. 10, figs. 3, 4. 

Occurrence of type material.-N orthwestern 

Venezuela, near Columbia border, near the headwaters of the Cachiri River. Devonian, probably 

Onondaga in age. (See discussion.) 

Diagnosis.-Phaceloid Acinophyllum with rare 

lateral supports and major septa varying from long 

to short. 

External features.-Phaceloid coralla of unknown 

size. Weisbord states, "This species virtually comprises the limestone in which it is found imbedded." 

This statement suggests that colonies may be large. 

Mature corallites are straight and cylindrical; diameter ranges from 4.1 to 5.2 mm in the available 

material (table 18 and fig. 25). 

Increase is lateral. Lateral supports are present 

but unco·mmon. Broad, rounded interseptal ridges 

and septal grooves give a scalloped outline to wellpreserved corallites in transverse sections. 

Corallite spacing in corallum ranges from 0.5 to 

1.5 diameters. Galice unknown but presumably 

steepsided. 

Internal features.- Septa radially arranged. 

Major septa extend from 0.3 to 0.7 the distance to 

the axis; minor septa very short. Number of septa 

in 2 fragments of coralla ranges from 19 to 22 

(table 18 and fig. 25). Zigzag carinae are weakly 

to strongly developed on most septa. 

The diss·epimentarium is composed of one or two 

rows of normal dissepiments. In most sections the 

dissepiments are globose, but there is considerable 

variation and some are quite elongate (pl. 29, fig. 

6). 

Tabulae are variable but commonly fiat to gently 

convex. 

Discussion.-The above description is based on 

three syntypes and six topotype fragments. The 

speeies is included here because it is the only known 

Early or Middle Devonian species of Acinophyllum 

from outside of Eastern North America. With associated corals and brachiopods, it indicates that 

Venezuela wa:s part of the Eastern Americas Realm 

during this time. 

A. vermetum is very similar to A. stramineum, 

from which the available specimens differ in no 

significant way. It differs from other described species in the same way that A. stramineum does. Because of its geographic remoteness, it seems best to 

recognize A. vermetum until the scope and direction 

of ·interspecific variation within the "species" is 

known. 

Brachiopods associated with A. vermetum indicate a Schoharie or Onondaga age (J. G. Johnson, 

written commun., 1968). The similarity of A. vermetum to Onondaga rather than Schoharie species 

of Acinophyllum makes an Onondaga age more 

likely. 
68 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Distribution.-Cano Grande Formation, Rio 

Cachiri Series, on the Gafio del Norte, branch of 

the Cachiri River, northwestern Venezuela (Liddle, 

1943, p. 14-15; Wei:sbord, 1968, p. 217-222). 

Material studied.-Lectotype (here selected), 

PRI 21598; paralectotypes, PRI 21595-96; topotypes, PRI 27637-38 (here illustrated and PRI 

3332F (four unsectioned fragments). 

Genus CYLINDROPHYLLUM Simpson 

1900 Cylindrophyllum Simpson, p. 217. 

? 1910 Cylindrohelium Grabau, p. 102. 

not 1915 Cylindrophyllum Yabe and Hayasaka, p. 90 

(homonym; renamed Fletcherina by Lang, 

Smith, and Thomas, 1955). 

not 1922 Spinophyllmn Wedekind, p. 5-6. 

1935a Cylindrophyllum Simps~n. Lang and Smith, p. 

544. 

part 1938 Cylindrophyllum Simpson. Stewart, p. 43-44. 

1944 Cylindrophyllum Simpson. Shimer and Shrock, p. 

95. 

1949 Cylindrophyllum Simpson. Stumm, p. 33. 

part 1955 Cylindrophyllum Simpson. Stumm, card 136. 

1956b Cylindrophyllum Simpson. Hill, p. F280. 

? 1961 Cylindrophyllum Simpson. Fontaine, p. 98. 

part 1967 Cylindrophyllum (Cylindrophyllum) Simpson. 

Pickett, p. 21-22. 

1974 Cylindrophyllum Simpson. Oliver, p. 167. 

Type species.-Cylindrophyllum elongatum Simpson, 1900. Onondaga Limestone (probably Edgecliff 

Member, bioherm facies), ea.stern New York. 

Diagnosis.-Phaceloid cylindrophyllinids with relatively wide dissepimentarium commonly consisting 

of from three to eight rows of normal globose dissepiments. Zigzag to subyard-arm carinae are present to abundant on major and minor septa. 

Description.-Coloniai, phaceloid or less commonly dendroid corals; lateral supports between corallites may be rare or abundant. Offsetting is lateral 

or, les.s co·mmonly, peripheral. Calices deep with no 

platform. 

Septa are radially or biradially arranged. Major 

septa do not reach the axis and may be very short. 

Major and minor septa are slightly to heavily 

carinate in the dissepimentarium; carinae are commonly zigzag, rarely yard-arm. Positions of cardinal 

and (or) counter septa may be marked by short 

major septa. 

The dissepimentarium is composed of a few to 

s·everal rows of normal, globose diss,epiments. Tabulae are commonly complete; the tabularium occupies 

one-half or more of the corallite diameter. 

/ncrease.-Lateral increase is com·mon in all species studied and probably in ·all known species. Peripheral offsets are less common but are known in 

two species and may occur in others. 

Microstructure.-As for the family. Specific details are described under the individual species. 

Discussion.-Cylindrophyllum differs from phaceloid disphyllids in the presence of abundant septal 

carinae and in its relatively wide dissepimentarium. 

In contrast, Disphyllum com,monly lacks carinae and 

has coars·er trabeculae. (See family discussion). The 

evidence suggests that Cylindrophyllum is endemic 

to the Eastern Americas Realm. 

Acinophyllum differs from Cylindrophyllum in 

·having a single basic row of dissepiments ; inner 

rows, if present, are composed of smaller dissepiments. The earliest species of both genera are in 

the Bois Blanc and its equivalents, and I interpret 

the genera to represent separate lines of cylindrophyllinid development a;s distinct from each other 

as either is from cerioid or astraeoid lines. 

Cylindrohelium Grabau (1910) is generally considered to be a synonym of Cylindrophyllum 

(Stumm, 1949, p. 33; Hill, 1956b, p. F280; and 

others). The syntypes of Cylindrohelium profundum 

Grahau, the type species, are unrecognizable molds 

from Ohio and Alberta. I here select as lectotype 

the original of Grabau's plate 11, figure 5 (UMMP 

13085) from the 'Lucas Dolomite, * * * near Sylvania, Ohio." The lectotype is thus a single calice 

mold from rocks of late Onesquethaw age in northwestern Ohio. Its selection will reduce the uncertainties regarding Cylindrohelium, but will not make 

it recognizable. Cylindrohelium can best he listed as 

unrecognizable and a nomen dubium. 

Spinophyllum Wedekind (1922) is based on 

Campophyllum spongiosum Schluter (1889). Schluter described the species to include curved-conical, 

solitary horn corals whose only obvious similarity 

to Cylindrophyllum is in the possession of carinate 

septa. Wedekind apparently intended his genus to 

include colonial as well .as solitary forms and possibly for this reason Stumm (1949), Hill (1956b), 

Pickett (1967), and others have placed Spinophyllum in synonymy with Cylindrophyllum. I have 

examined the Schliiter syntypes (Bonn 17 4a, b) and 

sectioned one of them (174a). They appear to be 

conspecific with the illustrated specimen of Wedekind (1922, fig. 2; SMF 3941, 3942). S. spongiosum 

differs from :specie's of Cylindrophyllum in details 

of septal and tabulae morphology that may or may 

not be i.mportant, but it is :solitary and lacks the 

characteristic c~aspedophyllid microstructure. I 

think that these are sufficient grounds for rejecting 

synonymy, and I consider it very unlikely that the 

two taxa are even closely related. 
SYSTEMATIC PALEONTOLOGY 69 

The following Onesquethaw age species from Eastern North America are assigned to Cylindrophyllum 

and described in the following sections : 

1. Cylindrophyllum laxtensum n. sp.: Schoharie 

Grit, eastern New York, and Bois Blanc Formation, southwestern Ontario. 

2. C. elongatum Simpson; type species; Edgecliff 

bioherm facies, New York. 

3. C. propinquum Stewart; Edgecliff, bioherm and 

biostrome facies, eastern New York; Columbus Limestone, Zone C, Ohio. 

4. C. deearium n. sp.; Edgecl·iff bioherm facies, 

New York; Edgecliff biostrome facies, southwestern Onta.rio; upper Columbus Limestone, 

Moorehouse equivalent, northeastern Ohio. 

5. C. sp. A; Edgecliff bioherm facies, western New 

York. 

The following additional Onesquethaw "species" 

are synonyms, of uncertain status, or reassigned : 

1. Cylindrohelium profundum Grabau, 1910. Type 

species of Cylindrohelium (generally considered a j unioT synonym of Cylindrophyllum) ; 

Amherstburg Formation, Detroit River Group, 

Ohio (Stumm, 1955, card 151). Not recognizable. 

2. Cylindrohelium heliophylloides Grabau, 1910. 

Lucas Formation, Detroit River Group, Michigan. Probably a Cylindrophyllum but not recognizable on the species level. 

3. Cylindrophyllum confiuens Stewart, 1935. 

Columbus Limestone (Zone C), Ohio. This is 

a zaphrentid possibly conspecific with Cyathocylindrium opulens n. sp. described below. 

4. Cylindrophyllum compactum (Hall) of Stumm, 

1965. Jeffersonville Limestone, Falls of the 

Ohio, Kentucky. Stumm synonymized several 

nominal species of Hall (1882, 1884), Davis 

(1887), ·and Greene (1904). I have reexamined the types and consider that most of 

the "species" are unrecognizable but agree 

that most are or may be conspecific. Most 

have long, bilaterally arranged septa with distinct fossulae and would be zaphrentids as 

here defined. Heliophyllum gradatum Greene 

is a Cylindrophyllum and possibly a senior 

synonymn of C. propinquum Stewart, described below. Stumm also included in his 

synonymy C. propinquim and C. confiuens 

Stewart (1938) di(cussed above. / 

5. Diphyphyllum gigas Rominger, 1S76, referred 

to Cylindrophyllum by Stumm, 1955, is a 

zaphrentid and is discussed under Cyathocylindrium. 

6. Cylindrophyllum stummi Oliver, 1971, of late 

Onesquethaw or early Cazenovia age is redescribed below as Disphyllum? stummi. 

Several species of "Cylindrophyllum" have been 

described from upper Cazenovia and Tioughnioga 

Middle Devonian rocks of eastern North America. 

These are not included in this study and are listed 

here without annotations except to question the assignment of some of the species. All are from the 

Traverse Group (Givetian) of Michigan. 

1. Cylindrophyllum delicatulum Ehlers and Stumm, 1949. 

2. C.? grabaui Ehlers and Stumm, 1949. 

3. C. hindshawi Ehlers and White, 1932. 

4. C. magnum Ehlers and Stumm, 1949. 

5. C.? panicum (Winchell), 1866. 

6. C.? compactum (Ehlers and Stumm), 1949. 

Possible Middle Devonian Cylindrophyllum in 

western North America are of interest because of 

the relationship they might suggest between the 

eastern and western provinces. Only two species 

require consideration, both Givetian in age. The 

following notes are based on a restudy of the type 

specimens. 

1. Cylindrophyllum gruensis McLaren (1964, p. 9-

10, pl. 3, figs. 1a-2b), Horn Plateau Formation, Givetian, District of Mackenzie, Northwest Territories, Ganada. This has relatively 

complex septa suggestive of some of the 

ptenophyllids and a dissepimentarium with 

from 4 to 10 or more rows of small dissepiments. In thes·e respects, C. gruensis differs 

from C. spp. and may be a cyathophylloid 

rather than a craspedophyllid. 

2. Disphyllum accidens Stumm (1938), assigned to 

Cylindrophyllum by Stumm (1955, card 148) ; 

upper part of Nevada Formation (Givetian), 

Nevada. This is not a Cylindrophyllum. It 

differs in having a dendroid growth form with 

trochoid, ceratoid and cylindrical corallites attaining widely differing diameters; and in having peripherally discontinuous septa. It is a 

cyathophylloid in the broad sense but its relationship to eastern zaphrentids is unknown. 

Disphyllum was rare in Eastern North America 

during the Early and Middle Devonian. Some 

"Disphyllum" of authors are Cylindrophyllum; 

others are apparently unrelated to either Disphyllum or Cylind'rophyllum. One:squethaw species are 

briefly discuss·ed here. I have reexamined the types 

of all of the species. 
70 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

1. Cyathophyllum cohaerens Hall (1882), assigned 

to Disphyllum by Stumm (1965, p. 41), has 

weak zigzag carinae and probably belongs to 

the group discussed under Cylindrophyllum 

compactum above. 

2. Cyathophyllum septatum Hall (1882), synonymized with D. cohaerens by Stumm (1965, p. 

42) is a Silurian Entelophyllum, as is shown 

by thin sections of the holotype prepared in 

my laboratory. 

3. Disphyllum dyeri Cranswick and Fritz (1958, 

p. 38, pl. 1, figs. 8, 9) is unrecognizable. The 

published illustrations do not show the nature 

of the dissepimentarium and the holotype (and 

only) specimen has been essentially destroyed 

since illustration, apparently by attempts to 

prepare a longitudinal thin section. From remaining fragments it is not clear whether any 

dissepiments are present, but the characteristic craspedophyllid or disphyllid dissepiments are certainly lacking. 

4. Disphyllum synaptophylloides Stumm (1965, p. 

42, pl. 37, figs. 5-8) differs from craspedophyllids and disphyllids in having strongly 

concave, complete and incomplete tabulae, and 

abundant twofold, threefold, and fourfold 

peripheral offs·et; it probably belongs to 

neither family. At present the species is known 

only from the holotype in the Davis collection (MCZ 7825). According to Davis' notes, 

kindly sent to me by Prof. Stumm, the specimen is from the lower 5 feet (1.5 m) of the 

Jeffersonville Limestone at the Falls of the 

Ohio (Schoharie equivalent). The species requires more detailed study before its relationships can be established. 

5. Disphyllum sp. A of Fagerstrom (1961, p. 13-

14, pl. 3, figs. 1-3) is Acinophyllum mclareni. 

6. Disphyllum sp. B of Fagerstrom (1961, p. 14, 

pl. 3, :fig.s. 9, 10) is a Cylindrophyllum possibly conspecific with C. proprinquum Stewart. 

Only three species of Eastern North American 

post-Onesquethaw Middle Devonian age have been 

assigned to Disphyllum: 

1. Disphyllum compactum Ehlers and Stumm, 1949, 

is a weakly carinate Cylindrophyllum? possibly synonymous with C? panicum Winchell. I 

have studied the type specimens of both "species" but analysis of much larger collections is 

needed to determine the extent of variation. 

Both forms are from the Traverse Group 

(Givetian) of Michigan and both occur in the 

Potter Farm Formation. 

2. Diplophyllum callawayensis Branson, 1924 ( assigned to Disphyllum by Stumm, 1955, card 

130), from the Callaway Limestone (Givetian) in Missouri, was inadequately described 

and illustrated and has not been seen by me. 

The description and transverse section (Branson, 1924, p. 50-51, fig. 3-3) indicate that 

minor septa are either lacking or very short 

and contratingent, and it is probable that the 

species is not related to the craspedophyllids 

or disphyllids at all. 

3. Disphyllum sp. A Stumm, 1968 from the Tenmile Creek Dolomite of Stewart (1938) of 

Ohio was based on two fragments of single 

corallites. The form is indeterminate but appears to. have carinae and bilaterally arranged 

septa and may be zaphrentid. 

Two species of late Onesquethaw or early Cazenovia age are redescribed in a following s·ection as 

Disphyllum?: 

1. Diphyphyllum rectiseptatum Rominger, assigned 

to "Synaptophyllum" by Stumm, 1966, is the 

oldest probable disphyllid known from the 

Michigan Basin. 

2. Cylindrophyllum stummi Oliver is the only probable disphyllid known from the Appalachian 

Basin. 

In addition, Disphyllum? crassiseptatum (Ehlers 

and Stumm, 1949) occurs in the Potter Farm Formation (Taghanic Stage) in the Michigan Basin 

and some of the Traverse Group Cylindrophyllum? 

listed above may be Disphyllum? also. (See discussion of Disphyllum? below.) 

Three Late Devonian species have been referred 

to Cylindrophyllum. 

1. Cylindrophyllum fioydense Belanski, 1928, limestone of Shellrock stage, Iowa. Examination 

of thin sections of the holotype indicates that 

this is not a Cylindrophyllum. It lacks carinae 

and has the thickened septa and coarse monacanths of typical Disphyllum. 

2. Disphyllum nevadensc Stumm, 1939, Devils Gate 

Limestone, Nevada, reassigned to Cylindrophyllum by Stumm (1955, card 147). Carinae 

'are weakly and very locally developed in the 

holotype. The species differs from Middle Devonian Cylindrophyllum in its somewhat di-
SYSTEMATIC PALEONTOLOGY 71 

lated, spindle-shaped septa and in its very 

coarse, "European" -type trabeculae. I consider 

it unlikely that this form is a descendant of 

Cylindrophyllum s.s.; its gross characters may 

be due to convergence. 

3. Diphyphyllum tubiforme Fenton and Fenton, 

1924, Hackberry Stage, Iowa, referred to Cylindrophyllum by Stumm (1955, card 153). The 

published description and illustrations o.f this 

form are not adequate for meaningful evaluation, but Prof. J. E. Soranf states that it is a 

Disphyllum (written commun., 1971). 

Only two species have been referred to Cylindrophyllum from areas outside of North Am·erica. The 

following notes are based on the published illustrations and descriptions. 

1. Cylindrophyllum asiaticU'm Fontaine (1961, p. 

98-99, pl. 19, figs. 11-13) i:s from an unknown 

stratigraphic position within the Devonian of 

VietNam. The species has the gross characters 

of Cylindrophyllum, but its assignment to the 

genus is difficult to assess without knowledge 

of its finer structure. 

2. Cylindrophyllum planivesiculosum Chernychev 

(1941, p. 11, 53, pl. 1, figs. 4, 5) is from the 

base of the Lower Devonian, Taimyr Peninsula, USSR (Cherynchev, 1941, p. 61). This 

species has a single row of rather flattened dissepiments and may lack carinae, and it cannot 

be included in Cylindrophyllum as I understand the genus. 

Generic range.-Species of Cylindrophyllum may 

be limited to rocks of late Early and Middle Devonian age in the Eastern Americas Realm. How,ever, 

possible occurrences are noted in the late Middle 

Devonian of western North America and in the 

Devonian of Viet N am. Possible La.te Devonian occurrences have not been closely analyzed but seem 

very questionable to me. 

Cylindropbyllum elongatum Simpson 

Plates 31-33; plate 34, figures 1-7 

1900 Cylindrophyllum elongatum Simpson, p. 217-

218, fig. 42. 

1944 C. elongatum Simpson. Shimer and Shrock, p. 

95, pl. 30, fig. 8. 

1949 C. elongatum Simpson. Stumm, p. 33, pl. 15, fig. 

6. 

1955 C. elongatum Simpson. Stumm, card 139. 

1956b C. elongatum Simpson. Hill, p. F280, fig. 191, 8. 

Occurrence of type material.-"Onondaga Limestone; Clarksville, Albany County, N.Y. J. W. Hall, 

coli. 1877" (NYSM ootalog of types). Almost certainly the Edgecliff Member, probably in one of the 

bioherms that crop out within a few miles of the 

village. 

Diagnosis.-Phac.eloid coralla with cylindrical 

co:rallites (diameter 8-15 mm) offsetting laterally or 

peripherally. Major septa biradially arranged, commonly with two long counter lateral minor septa; 

major septa short; dissepimentari urn narrow containing two to five rows of globose diss·epiments. 

External features.-Phaceloid colonies are as 

much as 15 c,m or more in heJght and 10 to 12 em in 

diameter, compos,ed of cylindrical corallites virtually 

parallel to each other. Corallite spacing ranges from 

0.5 to 1.5 diameters; corallites rarely in contact except at points of origin. Observed diameters range 

fro.m 7 to 16 mm (mean diameter in nine coralla, 9.4 

to 13.4 mm) with considerable variation within each 

colony (table 21 ,and fig. 28). Increase is lateral or 

peripheral but both types of increase have not been 

ohs:erved in the sam·e colony. Of 11 colonies studied, 5 

offset laterally and 6 peripherally; no offsetting takes 

place within the holotype sections. 

Internal features.-The major septa are biradially 

arranged and extend from 0.4 to 0.8 the distance 

to the axis, commonly 0.5 to 0.6. One or two of the 

major septa may be shorter than the others ; presumably these .are the oardinal and counter septa. In 

some corallites one major septum is flanked by long 

minor septa; this is probably the counter septum. 

Major septa number from 25 to 36 (mean numbers 

in 9 co.rnll:a, 26.7 to 33.8; tabl~e 21 and fig. 28). One 

exceptional corallite has 46 m:ajo·r septa. Except a:s 

noted, minor s,epta are from one-half to two-thirds as 

long as the major septa. All septa are lightly to moderately marked by zigzag carinae. 

TABLE 21.-Cylindrophyllum elongatum Simpson, summary 

data 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 4 16 8.7 9 n: x ------- 26.7 33.8 29.2 29.5 

OR ----- 25-28 33-36 26.7-33.8 ~ ------- 1.03 2.07 2.28 

v ------ 3.7 7.2 5.1 7.7 

? 1961 C. sp. cf. C. elongatum Simpson. Fagerstrom, p. d: 

14, pl. 3, figs. 7, 8. 

x ------- OR -----

9.4 

8-11 

13.4 11.9 11.9 

12-15 9.4-13.4 

2.10 part? 1961 C sp. cf. C. propinquum Stewart. Fagerstrom, p. 

14, pl. 3, fig. 4 (not figs. 5, 6, see C. sp. A). 

s 

v -------

------ .48 1.44 

3.6 20.0 12.1 12.1 
72 

•C:: 

r 

8 

NONCYSTIMORPH COLONIAL RUGOSE CORALS 

EXPLANATION 

X Cylmdrophyllum e/ongatum 

• C laxtensum 

b. D/sphyllum? stummt 

o Samples less than 5 corallites 

d (mm) 

EXPLANATION 

Cylmdrophyllum prop/nquum 

0 Ohio 

X New York 

+ C. sp. cf C. prop/nquum, Ontario 

• C.sp. A 

• C. sp. cf C. propmquum of Fagerstrom 

I I I 

d(mm) 

The dissepimentarium is narrow, occupying from 

one-third to one-half the radius, most commonly with 

three rows of dissepiments (observed range, one to 

six rows). Dissepiments are medium to large sized 

and globose. 

Tabulae are complete or incomplete and variable 

in form. Most commonly they are complete, nearly 

flat with a broad, gentle axial depression and slightly 

down bent peripherally; others are incomplete, concave plates forming a strongly concave pattern. Intermediate types are common, but both extremes can 

he found in a single corallite or corallum. The tabularium occupies from one-half to two-thirds of the 

corallite diameter. Tabular spacing ranges from 2 to 

10 per 5 mm; the widest spacing occurs only with the 

incomplete, strongly concave tabulae. 

The production of lateral or peripheral offsets is 

as described above for the phaceloid corals as a 

group; offset intervals are not known. 

Discussion.-Simpson's original description of C. 

elongatum is brief but adequate to clearly distinguish the species in the collections. His one illustration is a somewhat diagrammatic and apparently 

composite longitudinal section. The holotype consists 

of two longitudinal thin sections, one of which 

(NYSM 246a) is clearly the basis for the upper half 

of Simpson's illustration. The second thin section 

(246b) is not well centered in the corallite but is 

apparently the basis for the lower half of the figure. 

The two sections do not fit together in any obvious 

way but are assumed to be from the same corallite or 

corallum. Simpson omitted all structures but dissepiEXPLANATION 

X New York 

0 Ontario 

35 l I X I 

c 0 

X 

X 

•c:: X 

X xx 

30 1- - 0 

I I I 

10.0 12.0 14.0 16.0 18.0 

d(mm) 

FIGURE 28.-Mean number of major septa (n) versus mean 

diameter (d) in Cylindrophyllum spp. and Disphyllum? 

stummi. Encircled symbols represent samples of less than 

five corallites. Polygons show observed ranges in species. 

A, C. elongatum, C. laxtensum, and D.? stummi. B, C. 

propinquum, C. sp. cf. C. propinquum, and C. sp. A. Straight 

lines mark indicated septal ratios. C, C. deearium. 
SYSTEMATIC PALEONTOLOGY 73 

ments, tabulae, and outer wall from his figure, although the sections show traces of ~septa with zigzag 

carinae as well. The pres~ence of both septa and carinae are mentioned in the description, but their nature has been misunderstood by some workers. Both 

types of tabulae occur in both holotype thin sections 

and are well shown in the original figure. 

The above description is based on 13 specimens including the holotyp.e. Although some of the colonies 

have been compressed, little crushing was noted and 

most of the corallites are fairly well preserved. 

Individual variation in this species is greater than 

in any of the other Onesquethaw species of Cylindrophyllum. Most notable are variation in septal length 

and tabulae shape as discussed in the description and 

in cylindrical diameters (table 21 and fig. 28). 

Most characteristic are the narrow dissepimentarium and the relatively short septa. 

Fagerstrom (1961, p. 14, pl. 3, figs. 4-10) described and illustrated four separate corallite fragments of probable Cylindrophyllum (as Disphyllum 

sp. B, C. sp. cf. C. elongatum, and C. sp. cf. C. propinquum) from the Formosa bioherm facies of the 

Detroit River Group in Ontario. The specimens are 

poorly preserved and inadequate for identification, 

but two of them are closer to C. elongatum than to 

any other described cor-al and are questionably assigned to this species as noted in my synonymy and 

list of materials studied. 

Cylindrophyllum elongatum is morphologically 

closest to C. propinquum Stewart, originally described from the Columbus· Limestone (Zone C) in 

Ohio. Study of five Ohio coralla suggests that C. propinquum differs in having longer septa that are less 

variable in length within coralla. Some New York 

Edgecliff colonies are so similar that I confidently 

assign them to C. propinquum, but I am not certain 

that much larger samples of both Edgecliff and Columbus forms won't show that there is continuous 

variation between C. propinquum and C. elongatum. 

As here defined, C. elongatum colonies all include individuals with very short septa (less than 0.5-0.6 

radius) ; commonly such individuals predominate, 

but most colonies also include individuals with longer 

septa (as long as eight-tenths radius). In contrast, 

C. propinquum colonies commonly have medium to 

long septa (0.7-0.9 radius) with minimal intracolony 

variation; corallites with short septa are very rare. 

Cylindrophyllum elongatum has decidedly longer 

septa than C. laxtensum and much shorter septa than 

C. propinquum. It differs from C. sp. A in its more 

closely spaced septa as indicated by the septal ratios 

(fig. 28). 

Distribution.-Eleven specimens that I collected 

are aU from the bioherm facies of the Edgecliff Member in New York. A twelfth specimen ·and the bolotype are NYSM specimens from "Onondaga Limestone, Clarksville, N.Y." Edgecliff bioherms are 

known from the outcrop belt both north and south of 

Clarksville (fig. 5) and others may occur even closer 

to the village. It seems most likely that the NYSM 

specimens are from the Edgecliff Member and probably from the bioherm facies, as are the other 

specimens. 

F~agerstrom's (1961) specimens indicate that C. 

elongatum may also occur in the Formosa bioherm 

facies, Amherstburg Formation, Detroit River 

Group, Ontario. 

Material.-Holotype NYSM 246 (two longitudinal 

thin s~ections); measured and (or) illustrated specimens, USNM 163382-88, 172241-42; NYSM 12814-

16; unillustrated studied specimens, USNM 163389-

90. 

Cylindropbyllum propinquum Stewart 

Plates 35-38 

1938 Cylindrophyllum propinquum Stewart, p. 46, 

pl. 8, figs 7, 8. 

1955 C. propinquum Stewart. Stumm, card 152. 

? 1961 Disphyllum sp. B, Fagerstrom, p. 14, pl. 3, 

figs. 9, 10. 

part? 1961 C. sp. cf. C. propinquum, Fagerstrom, p. 14, pl. 

3, figs. 5, 6 (see discussion under C. sp. A; 

not fig. 4 which may be C. elongatum). 

Occurrence of type material.-Columbus Limestone, Zone C, Dublin, Ohio. 

Diagnosis.-Phaceloid coralla with cylindrical 

corallites offsetting laterally. Major septa radially 

arranged but commonly with one or two short major 

septa defining the cardinal-counter plane. Major 

septa medium to long, extending seven-tenths to all 

the distance to the axis. Dissepimentarium narrow, 

commonly with three to six rows of globose 

di,ssepiments. 

External features.-Known specimens are fragments of phaceloid colonies, as much as 25 or more 

centimetres high and 15 by 40 or more centimetres 

in diameter, composed of parallel, cylindrical corallites. Corallite spacing ranges fro·m 0.1 to 1 diameters; corallites rarely in contact. Observed corallite 

diameters range from 6 to 17 mm ; mean diameter in 

five Ohio colonies ranges from 10.7 to 11.8 mm; in 

six New York colonies 7.3 to 14.4 mm (table 22 and 

fig. 28). Only lateral offsets are known in the studied 

specimens. 

Corallite exteriors are marked by broad V -shaped 

septal grooves and low, rounded interseptal ridges; 

two orders of rugae can be recognized. 
74 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

TABLE 22.-Cylindrophyllum propinquum Stewart, summary data 

Ohio sample 

Intracolony InterMinimum Maximum Mean colony 

N ----------· 3 9 5 n: x ------· 27.8 31.7 29.0 

OR -----· 27-29 29-33 8 -------· 0 2.31 1.6 

v ------· 0 7.3 4.5 5.5 

d: x -------· 10.7 11.8 11.1 

OR ----- 10.0-11.6 9.6-12.8 8 -------· .76 1.75 .46 

v ------· 6.7 15.9 10.8 4.1 

Calices, known only from sections, are flat bottomed with nearly vertical lower walls and more 

gently inclined upper walls; there is no peripheral 

platform. 

Internal features.-Most of the major septa are 

radially arranged and extend from seven-tenths to 

all the distance to the axis; corallites with shorter 

septa (six-tenths) are rare. Septa are irregularly 

bent within the tabularium. Most corallites have one 

or two major septa that are shorter than the others; 

presumably those mark the oa.rdinal-counte·r plane. 

Adjacent major septa commonly bend around one or 

both of the short septa. Minor septa are 0.5-0.7 as 

long as the major septa. 

All septa are carinate and carinae may be common 

in the tabularium as well as in the dissepimentarium. 

The pattern of carinae varies both within and between coralla. Zigzag to subyardarm carinae are 

prominent in the dissepimentarium; in some sections 

these are quite thick. At the tabularium boundary, 

thin but long yardar.m or zigzag carinae may form a 

prominent zone in transverse sections (for example, 

pl. 38, figs. 10, 11). Within the tabularium, major 

septa tend to be very irregular; zigzag carinae are 

short but distinct in many corallites but some coralla 

seem to lack these entirely. Longitudinal sections 

show the carina.e to be steeply inclined at or near the 

periphery, gerntly inclined to nearly ho,rizontal 

through mos:t of the dissepim:entarium, and turning 

to a near vertical position in the tabularium (pl. 38, 

figs. 4, 9). Vadations in carinae development can 

best be understood through reference to the plates. 

The dissepimentarium occupiers approximately half 

of the radius and consists of from three to six rows 

of globose, medium- to small-sized dissepiments. 

Tabulae are complete or incomplete, commonly 

with a broad axial depression and downbent peripherally; longer septa tend to distub the pattern and 

short, inclined incomplete tabulae are common at the 

margin of the tabularium. 

Microstructure.-See family description. 

Discussion.-This description is based on five 

New York sample Ontario 

specimen 

Intracolony Inter- USM 

Minimum Maximum Mean colony 172185 

4 15 9.8 6 5 

24.2 30.8 27.0 33.2 

22-26 30-31 31-35 

.50 1.60 2.22 1.48 

1.6 6.6 3.6 8.2 4.5 

7.3 14.4 11.1 14.1 

6-10 11-17 13.5-14.6 

1.01 2.26 2.37 .43 

9.8 22.2 15.6 21.4 3.0 

fragments of colonies from Ohio and six from New 

York. Four of the Ohio specimens are new but are 

clearly conspecific with the Stewart type material. 

The New York specimens are very similar but colo .. 

nies with both larger and smaller mean diameters 

than the Ohio forms are known and the New York 

specimens tend to have fewer septa at a given diameter (table 22 and fig. 28). An Ontario specimen 

is still larger, having even more septa, and is described :separately as C. sp. cf. C. propinquum. C. sp. 

A is also similar to C. propinquum and may be conspecific; it has a much lower septal ratio and is 

separately noted for that reason. 

Comparison of these Cylindrophyllum with long 

septa can be summarized as follows: 

n ii ii/d 

C. propinquum: 

Ohio -------------- 28-32 11-12 2.5-2.8 

New York -------- 24-21 7-14 1.9-3.3 

C. sp. cf. C. propinquum: 

Ontario 2.4 ----------- 33 14 

C. cf. C. propinquum: 

of Fagerstrom ----- 24 14 1.7 

c. sp. A -------------- 21-22 13 1.6 

These data are also plotted in figure 28B. 

Stewart ( 1938, p. 45) based her original description on "cotypes" from a single locality and to which 

she assigned only one number. It is not clear from 

her description or the type collection whether her 

"cotypes' represent diffe~rent individuals from one 

colony or from two or more colonies, but they are 

here treated as being from one colony and termed 

holotype. 

Cylindrophyllum propinquum differs from other 

described species of Cylindrophyllum in its long septa and in the arrangement of its septa within the 

tabularri urn. It is closest to C. elongatum as discussed 

under that species. 

The holotype of H eliophyllum gradatum Greene 

(1904; see discussion under genus) from the Coral 

Zone ( ?) of the Jeffersonville Limestone at the Falls 

of the Ohio is a Cylindrophyllum possibly conspecific 

with C. propinquum. 
SYSTEMATIC PALEONTOLOGY 75 

Two of the four fragments of probable Cylindrophyllum described and illustrated by Fagerstrom 

(1961, p. 14, pl. 3, figs. 4-10) ·and discussed under C. 

elongatum have long septa and may represent C. propinquum as noted in my synonymy .and list of material·s studied. (But see also discussion of C. sp. A.) 

Distribution.-The holotype and other Ohio specimens are from the Columbus Limestone, Zone C, 

within 30 miles (48 km) of Columbus, Ohio. The New 

York specimens are from the bioherm and biostrome 

facies of the Edgecliff Member, in eastern New York. 

Material.-Holotype, OSU 17779, consisting of s.ix 

thin sections assumed to be from one co,rallum. Other 

Ohio specimens, USNM 172186-189. New York 

specimens: Edgecliff bioherm, USNM 172180-183, 

NYSM 12818; Edgecliff biostrome, USNM 172184. 

Fagerstrom specimens from the Formosa bioherm 

facies and possibly this species, UMMP 36114, 36119. 

Cylindropbyllum sp. cf. C. propinquum Stewart 

Plate 45, figures 8-10 

One specimen from a bed of probable Edgecliff age 

(see distribution discussion) in Ontario is similar to 

C. propinquum in septal l-ength and arrangement, in 

having short zigz.ag carinae within the tabularium, 

and in the general arrange,ment and configuration of 

dissepiments and tabulae. The specimen differs in 

having moderately dilated septa in some parts of 

some corallites. This feature is sufficiently unusual in 

Onesquethaw craspedophyllids to warrant separate 

notice. 

The specimen is an incomplete fragment of a colony that was at least 15 em high and 15 by 20 em in 

diameter. Corallite diameters range from 13 to 15 

mm and number of major s·epta from 31 to 35 (table 

22 and fig. 28B). The corallites are large for C. propinquum but within the known range of variation in 

most characters. The dissepimentarium is relatively 

narrow and short zigzag carinae are present within 

the tabularium of several corallites. These oan be 

seen in both longitudinal and transverse sections. 

The septal dilation is such that some septa are wedge 

shaped, tapering from the periphery to the axis. Other septa are locally dilated in the inner part of the 

dissepimentarium; in some corallites the thickening 

is great enough to mask the carinae. 

Distribution.-The only known specimen is from 

beds exposed on the north shore of Lake Erie, just 

west of the mouth of the Grand River, south of 

Dunnville, Ontario. These beds are Edgecliff on the 

basis of their fossil content, lithology, and geographic position not far south of Bois Blanc outcrops. 

Material.-Illustrated specim·en, USNM 172185. 

Cylindropbyllum laxtensum n. sp. 

Plates 39, 40 

Diagnosis.-Phaceloid coralla composed of cylindrical cor.allites connected by numerous lateral supports. Cor.allite diameter1s range from 6 to 13 mm. 

Major sep:ta short, dissepimentarium narrow, tabulae complete. 

External features.-Phaceloid colonies .are as 

much as :at least 17 em .in height and diameter; corallites radiate and may be nearly horizontal in the 

lower part of the corallum but are generally erect 

and nearly parallel in the upper parts; high and 

broad lateral supports are common. Corallite spacing 

is approximately half the diam·eter. Corallites are 

more or less straight; diameters range from 6 to 13 

m.m, m·ean diameters in three coralla range from 8.0 

to 11.1 mm (table 23 and fig. 28A). Increa:se is lateral at infrequent intervals. Corallite exteriors are 

marked by prominent broad V -shaped septal grooves 

separated by rounded interseptal ridges; thes•e are 

prominent enough in well-preserved individuals to 

give a decidedly scalloped appearance in transverse 

s·ection. Gentle rugae and the lateral supports are 

additional features of the exterior. 

Calices are fl.at bottomed with steep walls and narrow peripheral platforms that gently slope toward 

the axis. 

Internal features.-Septa are radially arranged, 

commonly with no modification of the protosepta although ·short septa in so.me individuals may mark the 

cardinal-counter plane. Major septa are short, extending from 0.4 to 0.9 the distance to the axis, but 

five-tenths or les.s in most corallites. Number of 

major septa in cylindrical parts of corallites ranges 

from 21 to 29 in the studied sample; the mean number in 3 eoralla ranges from 21.3 to 26.9 (table 23 

and fig. 28A). Minor septa are 0.5 to 0.9 the length 

of the m:aj ors, commonly 0. 7 to 0.8. Zigzag carinae 

are strongly developed in some corallites, almost absent in others. Variations in septal features can best 

be judged by examination of the illustrations. 

Dissepimenta,ria are narrow, widths commonly 

four-tenths of the radius; they are composed of two 

TABLE 23.-Cylindrophyllum laxtensum n. sp., intracolony 

data 

GSC GSC USNM 3585a 31146 182813 

N ------------- 15 12 10 n: x --------- 26.9 22.1 21.3 OR -------- 25-29 21-24 21-22 8 ---------- 1.06 1.08 .48 

v --------- 3.9 4.9 2.3 d: x ---------- 11.1 8.0 8.8 OR -------- 9-13 6-10 8-10 s ---------- 1.25 1.54 .79 

v --------- 11.2 19.2 9.0 
76 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

to five rows (most commonly three) of dissepiments. 

Dissepiments are medi urn to large, usually 

globose, often elongate parallel to the carinae, 

rarely depressed. Lateral supports are filled with 

dis·sepiments. 

The tabularium occupies from 0.5 to 0.6 the diameter and is composed of usually complete, nearly 

horizontal tabulae; tabulae are broadly concave axially and gently downbent periaxially. 

Increase.-Lateral offs·ets are apparently widely 

spaced. Initial offset growth is nearly at right angles 

to the parent: full dia.m,eter is attained in 0.5 to 1 em. 

In other respects the development of offsets is as 

described for the family as a whole. 

Microstructure.-See family description. 

Discussion.-The ahov.e description is based on 9 

specimens; 2 are ca:ralla measuring 19 x 12 X 9 and 

7 X 18 X 15 em in height and long and short diameters, respectively, and including 40 to 100 or more 

corallites. The remaining specimens are smaller 

fragments including from 3 to 30 or more corallites. 

All specimens are at least partly silicified, but parts 

of the larger ones are calcitic and show the microstructure v·ery well. 

Some a:spects of individual variation within and 

between coralla is shown on table 23 and the graph 

{fig. 28A) ; variation in septal number in som.e colonies is quite restricted ·and outside the observed 

range of other colonies. Other •aspects of variation 

were qualitatively studied, but most variation is as 

great within colonies or even co·rallites as it is within the available s.ample of the species. 

Cylindrophyllum laxtensum differs fro.m both C. 

elongatum and C. propinquum in its ·short major septa, num·erous lateral supports, and nearly complete 

radial symmetry. C. elongatum and C. propinquum 

lack lateral supports .and show frequent or occasional 

modifications of the cardinal and counte~r septa. 

Distribution.-Boi·s Blanc Formation, Niagara 

Peninsula .and London area, Ontario; Schoharie Formation, Albany County, N.Y. 

M aterial.-Bois Blanc Formation, Ontario: bolotype, GSC 31146; illustrated para types, USNM 

163391, 182813, and GSC 3585a, 31147; unillustrated 

para types; USNM 163392 and GSC 31148-49. 

Schoharie Formation, N·ew York; illustrated paratype, USNM 163393; unillustrated paratype, USNM 

163394. 

Cylindrophyllum deearium n. sp. 

Plates 41-43 

Occurrence of holotype.-Onondaga Limestone, 

Edgecliff Member, bioherm facies; central New 

York, 6.5 miles (10.5 km) southwest of Oneida. 

Diagnosis.-Phaceloid co~alla with lateral or peripheral increase; corallite diameters commonly 12 

to 16 mm. Septa extend 0.6 to 0.9 the distance to 

axis; dissepim·entarium wide, commonly composed of 

five to nine rows of globos~e dissepiments; tabulae 

complete. 

External features.-Phaceloid coralla, as much as 

30 em or more in height and 15 by 25 em or more in 

di.ameter, composed of subparallel, cylindrical corallites, 10 to 20 mm in diameter. Colony means (table 

24 and fig. 28C) range from 11 to 16 mm, but are 

based on s.mall ·samples. Contacts between neighboring corallites are common, usually at expansion of 

the coarse rugae, but in places the rugae are so expanded in c .. ne direction as to form lateral supports; 

maximum observed distance between individuals is 

approximately one-half corallite diameter. Inc:rease 

is both lrateral and peripheral, but both types of offsets are not known to occur in the same colony. 

Corallites are gently to strongly rugose; longitudinal markings include broad V -.sha.ped interseptal 

grooves and low rounded septal ridges. Finer longitudinal and transvers:e striae are seen only in particul,arly well preserved areas o.f outer wall. 

Calices are relatively steep walled and flat bottom•ed ; c:alice depth equals a.pp.roximately one-half 

the corallite diameter; there is no peripheral platform. 

Internal features.-Septa are biradially arranged; 

the cardinal-oounter plane is commonly defined by 

short cardinal and counter septa, but the two defining septa are similar and specimen orientation is 

only possible in the early, conical pa.rt of a corallite 

where major septa are being inserted. 

Major septa ·extend 0.6 to 0.9 the distance to the 

axis; they rarely m·eet at the cardinal-counter plane. 

The number of mrajor septa ranges from 28 to 35 in 

cylindrical parts of corallites; the mean septal number for 8 colonies ranges from 29 to 34 (diameters 11 

to 16 mm), but samples are small and the figures are 

only approximate (table 24 and fig. 28C) . 

TABLE 24.-Cylindrophyllum deearium, n. sp., summary 

data 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 3 14 7.8 9 n: x 29.2 34.3 32.0 

OR ----- 28-31 33-35 8 ------- .50 2.12 1.64 

v ------ 1.5 6.9 4.3 5.0 

d: x 11.2 16.2 14.0 

OR ----- 9-14 13-20 8 ------- 1.00 2.99 1.59 

v ------ 6.7 18.4 12.6 11.4 
SYSTEMATIC PALEONTOLOGY 77 

Minor septa are one-half to two-thirds as long as 

the major. All septa are carinate in the dissepimentari urn ; carinae are weakly to strongly developed, 

co.mmonly zigzag. 

The dissepimentarium is composed of from 5 to 

12, commonly 7 to 9 rows of very globose dissepiments, occupying approximately one-half the radius 

of the corallite. Individual dissepiments ·are small to 

medium sized, commonly elongate parallel to the 

carinae. 

The tabularium oc.cupies approximately one-half 

the diameter. Tabulae are commonly complete, flat or 

slightly concave axially and sharply bent downward 

peripherally; inclined peri~axial tabulae connect the 

complete tabulae witih the dissepimentarium. 

Microstructure.-See family description. 

Discussion.-The above des·cription is based on 

fragments of 13 coralla; preservation is only fair 

and most of the specimens are partly compressed and 

crushed. 

In comparison with other species of the genus, 

Cylindrophyllum deearium is of large diam·eter and 

has more numerous rows of dissepiments. It is intermediate in length of septa and in this respect is 

closest to C. elongatum although intracolony variation of septal length is much greater in that species. 

Distribution.-Cylindrophyllum deearium is 

known from the bioherm facies of the Edgecliff 

Member in New York. Specimens were collected at 

four localities in central and eastern New York. Two 

specimens from the collection of the USNM and 

NYSM are labeled Leroy, N.Y.; these are probably 

from an Edgecliff bioherm also. 

Additional specimens were collected from the 

Edgecliff nonbioherm facies at Port Colborne and 

Cheapside, Ontario. The species is also known fro·m 

the lower Detroit River (Amherstburg) at Gorrie, 

Ontario. 

A single specimen is known from beds, equivalent 

to the Moorehous·e Member, in the upper part of the 

Columbus Limestone in the subsurface of northeastern Ohio (Lake County core). 

Material.-Holotype, USNM 172190; illustrated 

or measured paratypes: New York, USNM 172191-

93, 172243, 25870, NYSM 12819-20, 12871; Ontario, 

USNM 172194-95, GSC 31145, UMMP 50576. Specimen from Lake County, Ohio core: USNM 172196. 

Cylindrophyllum sp. A 

Plate 34, figures 8-13 

part? 1961 Cylindrophyllum sp. cf. C. propinquum, Fagerstrom, p. 14, pl. 3, figs. 5, 6 (not fig. 4 which 

may be C. elongatum). 

Description.-Two small corallum fragments, each 

including app,roximately six corallites, were found at 

the same locality; the specimens are very similar and 

may he fragments of one corallum. 

The cor all urn of this form is of unknown size but 

includes cylindrical corallites, 13 to 14 mm in diameter and approximately parallel to each other. 

Increase is lateral. 

The septa are radially arranged; shor ~' diametrically opposite major septa in two of four corallites thin sectioned are presumed to be the cardinal 

and counter ~septa. Major septa extend nine-tenths 

or more the distance to the axis and number 21 to 23. 

Minor septa are 0.5 to 0.6 the length of the major 

septa. Weak zigzag carinae are limited to the disseptimentarium. 

The dissepimentarium is composed of three to 

four rows of globose dissepiments and occupies approximately one-half the radius. Tabulae are incomplete and tend to be horizontal axially and irregularly inclined peripherally; spacing is two to five or 

more per 5 mm. 

Discuss.ion.-As shown by the graph (fig. 28B), C. 

sp. A is set off from other Onondaga species of Cylindrophyllum by having relatively few septa for its 

diameter. It fuTther differs from both C. elongatum 

and C. P'ropinquum in its incomplete tabulae and 

l'arger dis.sepiments. It is closes.t to C. propinquum in 

septal length and arrangement, but the low septal 

ratio is apparently distinct. 

The only previously described Cylindrophyllum 

that is clos·e to C. sp. A is one of the two s-pecimens of 

C. sp. cf. C. propinquum o.f Fagerstrom, 1961. This 

specimen is closer to C. s.p. A in septal ratio ( n = 14; 

n/d=1.71) than it is to C. sp. cf. C. propinquum but 

is closer to C. propinquum in general morphology. 

Pos,sibly C. sp. A ~hould be included in C. propinquum in spite of the noted differences. 

Distribution.-Bioherm or flank facies of the 

Edgecliff Member in the Williamsville bioherm near 

Buffalo, N.Y. (loc. 108). 

Material.-Illustrated specimens USNM 163395 

and NYSM 12817. 

part 

part 

part 

part 

Genus PRISM A TOPHYLLUM Simpson 

1900 Prismatophyllum Simpson, p. 218. 

1935a Prismatophyllum Simpson. Lang and Smith, p. 

558 (type species only). 

1938 Prismatophyllum Simpson. Stewart, p. 48-49. 

1939 Prismatophyllum Simpson. Hill, p. 229-231. 

1939 Pr-ismatophyllum Simpson. Sloss, p. 68-69. 

1940 Prismatophyllum Simpson. Stainbrook, p. 270-

273. 

1940 Prismatophyllum Simpson. Lang, Smith, and 

Thomas, p. 104. 
78 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

1944 H exagonaria Gurich. Shimer and Shrock, p. 95. 

part 1945 Prismatophyllum Simpson. Smith, p. 44-45. 

1947 Prismatophyllum Simpson. LeMaitre, p. 46-47. 

part 1948 Prismatophyllum Simpson. Wang, p. 7-8. 

1948a Hexagonaria Gurich. Stumm, p. 7-13 

part 1949 H exagonaria Gurich. Stumm, p. 33. 

part 1950 Prismatophyllum Simpson. Wang, p. 218. 

1969 Prismatophyllum Simpson. Jell, p. 63. 

1974 Prismatophyllum Simpson. Oliver, p. 167. 

Type species.-By origina.l designation, Cyathophyllum rugosum (Hall) of Milne-Edwards and 

Haime, 1851, p. 387, pl. 12, figs. 1, 1a, b (not Astraea 

rugosa Hall), renamed P. prisma by Lang and Smith, 

1935a, p. 558. Jeffersonville Limestone, Coral Zone, 

Cha.rleston Landing, Ind. (See species discussion.) 

Diagnosis.-Cerioid coralla with polygonal corallites separated by walls. Calices commonly with an 

axial pit and peripheral platform. Septa radially arranged; major commonly extend to within the tahulari urn, though in some .spereies th·ey ·are only slightly 

longer than the minor septa which are more or less 

limited to the dissepimenta.rium. The septa are thin 

with weakly to strongly developed, zigzag to subyard-arm carinae. The dissepimentarium is compos·ed of several rows of horizontal or inclined, globose diss·epiments. The tabularium is formed by complete or incomplete tabulae which in some species are 

divided into axial flat and periaxial arched series. 

Remarks.-The above diagnosis is based on a 

study of the type species a:nd the other ooste.rn North 

American species described or mentioned below. 

Prismatophyllum differs from H exagonaria, with 

which it has commonly been synonymized, in its attenuate septa com.posed of fine monacanthine trabeculae; in addition it commonly has :a relatively 

broad dissepimentarium. H exa.gonaria is typically 

disphyllid in its characters and its gros.s similarity 

to Prismatophyllum is interpreted as due to parallel 

or convergent evolution. (See also Jell, 1969, p. 68-

69.) 

The generic synonymy given here is incomplete as 

it has not been practical to ree~amine all Am•erican 

"H exagonaria" or to exhaustively study the world 

literature. Most generic descriptions have been based 

partly on Prismatophyllum and partly on Hexagonaria and some are so broad as to include virtually 

any cerioid coral. 

The type species of Prismatophyllum is P. prisma 

Lang and Smith. (See Lang, Smith, and Thomas, 

1940, p. 104, for details of its selection.) Smith 

(1945, p. 45, not Lang and Smith, 1935a, as stated by 

Smith) selected the original of Milne-Edwards and 

Haime, 1851, plate 12, figures 1a, b, as lectotype and 

illustrated thin sections for the first time. These 

authors cited "Falls of the Ohio, Charleston Landing 

* * *"as type locality, apparently not realizing that 

this was two localities. However, the lectotype in the 

de V erneuil collection, :Ecole des Mines, Paris, is 

clearly labeled Charleston Landing. This i.s the Jeffersonville Lim·estone, and almost c•ertainly the Middle Devonian part of the Coral Zone. Charleston 

Landing, Ind., is approximately 8 miles (12.8 km) 

northeast of the Falls of the Ohio, on the north side 

of the Ohio River. The following redescription of P. 

prisma is based on the lectotyp·e plus specimens from 

the Falls. 

Two species of Prismatophyllum occur in the Middle Devonian part of the Coral Zone of the J effer-

.sonville Limestone. The two can be differentiated on 

th:e has is of the configuration of the major septa 

within the tabularium in transv·ersre sections and on 

the other features described below. From the Coral 

Zone (Zone B proper), all Prismatophyllum collected 

by me have an irregular axial complex or whorl; 

from the Coral-stromatoporoid Zone (Zone C), all 

specimens have relatively straight septa. The MilneEdw:ards and Haime (1851, pl. 12, fig. 1b) illustration (drawing) of the lectotyp·e of P. prisma shows 

straight septa, but thin sections of the lectotype 

show clearly that P. prisma is the species with the 

axial complex or whorl. Stainbrook (1940) recognized that two species were present but was misled 

by the Milne-Edwards and Haime illustration. H·e 

described the species with .straight septa as Prismatophyllum prisma ~and the one with the whorl as P. 

cinctum so that now P. cinctum is a junior subjective 

synonym of P. prisma. The second species, erroneously c:aUed P. prisma by Stainbrook and subsequent 

workers, is here synonymized with P. ovoideum 

(Davis). The latter occurs in New York, southwestern Ontario, and Ohio, as well as in the Falls area. 

Prismatophyllum is very rare in New York ana 

the Niagara Peninsula of Ontario, and for this reason the genus is treated here in less detail than some 

of the other genera. Two specimens of P. ovoideum 

are apparently from the Edgecliff Member, and one 

specim·en of P. truncata Stewa.rt is from the S.eneca 

Member. These species a.re based on specimens from 

Kentucky and Ohio respectively, where coralla are 

more common. P. prisma Lang and Smith and P. 

anna (Whitfi:eld), are described for comparison with 

the New York-Ontario forms, but are not known 

from New York or Ontario. "H exagonaria" coalita 

(Rominger) from the Edgecliff Member in western 
SYSTEMATIC PALEONTOLOGY 79 

New York is a Heliophyllum. 

No Prismatophyllum is known from the postOnondaga Devonian of New York or Ontario. H exagonaria prisma Lang and Smith, listed by Oliver 

(1951, p. 712) from the Ludlowville Formation 

(Givetian) in NewYork has an aulos and is close to 

Eridophyllum arachnoideum Stumm. West of New 

York, Prismatophyllum and (or) Hexagonaria are 

common in Middle Devonian (Onondaga and Hamilton age) deposits in Michigan, Ohio, Indiana, and 

Kentucky. 

The paucity of Prismatophyllum in New York, 

even in bioherm and biostrome facies, is difficult to 

explain, especially as the rest of the New York corals 

have much in common with the more western faunas. 

This is discussed more fully in the section on paleoecology. 

H exagonaria is not known to occur in the Onesqueth·aw Stage of ·eastern North America. Hexagonaria occurs, along with A ustralophyllum and a 

variety of chonophylloid co1•als, in the Miami Bend 

Formation of Cooper and Phelan (1966), Caz.enovia 

Stage, Indiana. These Old World forms are associated with Old World brachiopods and mollusks 

(Cooper and Phelan, 1966) and apparently represent an incursion from western North America. In 

Michigan, the correlative Rogers City Limestone 

contains many of the same species (Cooper and 

Phelan, 1966) including the same Hexagonaria. In 

Michigan, the Cazenovia-Tioughnioga age Traverse 

Group includes a succession of faunas, some of which 

are dominated by Old World forms including H exagonaria. Eastern forms are also present and the details of this faunal mixing are still unclear. The 

Traverse species of "Hexagonaria" were reviewed 

by Stumm (1970). Some of these are certainly Prismatophyllum, but a detailed restudy has not yet been 

undertaken. The late Middle Devonian species from 

Iowa (Cedar Valley Limes.tone) described by Stainbrook (1940) are mostly Prismatophyllum but may 

include some Hexagonaria. These too need reexamination. 

I have not reviewed the several species of Hexagonaria that have been described from Lower and 

Middle Devonian rocks in western North America 

or thvse from the North American Frasnian. I do 

not know that any are Prismatophyllum, but close 

reexamination is needed to identify many of them, 

and this is beyond the limits of the present study. 

Generic range.-Prismatophyllum is presently 

known only from rocks of middle Onesquethaw 

(Schoharie; Em.sian) to Taghanic (Hamilton and 

Tully; Givetian) age in the Eastern North American 

Province. Possible occurrences in western North 

Ameriea and in the Upper Devonian need to be 

restudied. 

Prismatophyllum prisma Lang and Smith 

Plates 46-48 

part 1851 Cyathophyllum rugosum (Hall). Milne-Edwards 

and Haime, p. 387, pl. 12, fig. la-b (fig. 1) 

(not Astrea rugosa Hall, 1843, which is an 

Asterobillingsa). 

not 

not 

not 

not 

not 

not 

not 

not 

not 

1879 C. rugosum Edwards and Haime. Quenstedt, p. 

529, pl. 162, fig. 31a-b (=P. ovoideum). 

1887 C. rugosum (Hall). Davis, pl. 90, fig. 1, pl. 93, 

fig. 3 (seeP. ovoideum). 

1900 Prismatophyllum rugosum ( E d w a r d s and 

Haime). Simpson, 1900, p. 218 [not figs. 44, 

45] (cited as type species of Prismatophyllum 

new genus). 

1935a P. primsa Lang and Smith, p. 558-559, (not figs. 

17 and 18 which are diagramatic) (as new 

name for Prismatophyllum rugosum MilneEdwards and Haime of Simpson 1900). 

1938 P. prisma Lang and Smith. Stewart, p. 50-51, 

pl. 9, figs. 13-15 (see P. ovoideum). 

1940 P. prisma Lang and Smith. Stainbrook, p. 273-

275, pl. 37, figs. 1, 2, 4 (see P. ovoideum). 

1940 P. cinctum Stainbrook, p. 275-276, pl. 37, figs. 

3, 5, 6, pl. 36, fig. 4. 

1944 Hexagonaria prisma (Lang and Smith). Shimer 

and Shrock, p. 95, pl. 30, figs. 11, 12 (see P. 

ovoideum). 

1945 P. prisma Lang and Smith. Smith, p. 45-46, pl. 

14, figs. 1-3 (originals of figs. 2 and 3, not 

seen). 

1948a H. prisma (Lang and Smith). Stumm, p. 17-18, 

pl. 1, figs. 1, 2, pl. 8, figs. 1-7 (see P. ovoideum). 

1948a H. cincta (Stainbrook). Stumm, p. 23-24, pl. 10, 

figs. 1, 2. 

1949 H. prisma (Lang and Smith). Stumm, pl. 15, 

figs. 14, 15. 

1955 H. cincta (Stainbrook). Stumm, cards 170, 171. 

1955 H. prisma (Lang and Smith). Stumm, cards 209, 

210. 

1955 H. prisma. Carman, p. 70, fig. 3 (cf. Eridophyllum). 

1958 H. prisma (Lang and Smith. Cranswick and 

Fritz, p. 39-40, pl. 2, fig. 1 (a crushed specimen of Asterobillingsa sp.). 

1965 H. cincta (Stainbrook). Stumm, p. 43, pl. 38, 

figs. 10-12. 

1965 H. prisma (Lang and Smith). Stumm, p. 44, pl. 

38, figs. 7-9 (seeP. ovoideum). 

1966 H. mesocincta Fraunfelter, p. 347-351, figs. 3, 4. 

Occurrence of type material.-The lectotype is 

clearly labeled as from Charleston Landing, Ind. 
80 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

This is the Jeffersonville Limestone. At the Falls of 

the Ohio, 8 miles (12.8 km) down river from 

Charleston Landing, the species occurs only in the 

Middle Devonian Goral Zone (Zone B). (See distribution discussion below.) 

Diagnosis.-Prismatophyllum with medium-sized 

corallites jn which the major s·epta extend to, or 

nearly to the axis where (in transverse section) they 

appear to form an irregular network or whorl that 

in so·me individuals is almost an aulos. 

External features.-Cerioid colonies are as much 

as s·everal feet in diamete·r and 2 to 3 fe·et (0.61 to 

0.91 m) in thickness. Smaller colonies are lens 

shaped, circular in plan view, with diameter three or 

four times the maximum thickness. Corallites are 

polygonal with straight sides except in offsets. Corallite diameters range from 8 to 18 mm, mean diameters in eight coralla range from 11.5 to 15.8 mm 

(table 25 and fig. 29A). 

Calices are known only in longitudinal sections; 

shallow, depth approximately equal to tabularium 

diameter ; walls may be uniformly sloping or with 

narrow peripheral platform or slightly refl.exed. A 

low boss formed by the twisted ends of the major 

septa may have appeared in some calices. 

Internal features.-Septa radi,ally arranged, 

straight to so.mewhat irregular in dissepimentari urn. 

The m.aj or septa extend to or nearly to the axis 

where they turn to form an irregular network or a 

loose whorl. Minor ·septa are 0.5 to 0.8 the length of 

the majors. Major septa number 19 to 27 in presumably m.atur·e corallite·s; mean number in eight 

coralla ranges from 21 to 24 (table 25 and fig. 29A). 

In a few corallites a short major septum is flanked 

by long minor septa ; these may mark the counter 

position. 

Major and minor septa are commonly heavily 

carinate in the dissepim.entarium with long zigzag 

or subyard-arm carinae. Locally, dilation tissue may 

almosrt bury the carinae. Major septa are lightly 

dilated in the tabularium. A few septa are incom-

, plete at their peripheral ends, forming small Ions-

, dalioid areas. 

Corallite walls commonly thin (0.1 to 0.15 mm, 

locally ·as much as 0.4 mm), composed of a dark central zone and light-colored outer zones. Adjacent 

coralli.tes are everywhere in contact. 

The dissepimenta.ria occupy one-half or more of 

the corallite radius and consist of from four to eight 

rows of s~mall to medium-large globose dis.sepiments. 

The septal ca.rinae ar·e prominen,t features of the 

dis·sepimentaria in most longitudinal .sections. 

The tabularia occupy from one-third to one-half 

the dia.meter. In most longitudinal sections the tabulae appear to be ,s;eparated inrto a periaxial series of 

commonly complete, concave tabulae and an axial 

series of incomplete convex or concave tabulae. The 

periaxial tabulae ar·e bounded by the dissepimentarium and the axial complex; the axial tabulae are 

within the axi:al complex where they are interrupted 

by the irregul·ar septa. 

Increase.-lncrea,se is lateral or peripheral, but 

nonparicid.a.l. Seen in trans¥ers·e section, offsets commonly appea.r as a distinct bulge in the hysterocorallite wall and development is the same as in 

phaceloid craspedophyllids except that growth is 

within a confined space. One example of two conte,mporaneous offsets is known that appears to represent peripheral rather than lateral increase. Peripheral increase may be represented in one illustrated 

colony (pl. 47, fig. 4; note base of central corallite). 

Microstructure.-As for the family. Corallite 

walls are thin but show the three-layer structure 

very clearly. The light layers are distinctly fibrous 

with the fibers at right angles to the wall. Septal 

dilation within the dis,sepim·entarium is extreme in 

s-ome p.a.rts of some colonies. Locally, dilation tissue 

may completely envelope the very long carinae 

TABLE 25.-Pri8matophyllum pri8ma Lang and Smith, summary data 

Intracolony Inter- Lectotype Mini- Maxi- Mean colony BMNH mum mum R31730 

N 5 12 8 1 4 (3) n: x --- 20.7 24.0 2'2.5 22.5 

OR - 19-22 20-2:7 20.7-24.0 21 -23 

8 --- .53 1.87 1.27 1.00 

v --- 2.2 8.4 5.7 5.6 4.4 

d: x --- 11.5 15.8 13.0 12.7 

OR - 8.0-16.0 9.5-18.5 11.5-15.8 10.0-15.0 

8 1.41 3.42 1.25 2.52 

v --- 10.8 25.3 18.7 9.6 19.9 

1 Diameter measured for only three corallites. 
SYSTEMATIC PALEONTOLOGY 81 

EXPLANATION EXPLANATION 

+ Falls of the Ohio 

$ Lectotype 

$ Holotype of Hexagonar/a cmctum 

d (mm) 

26 

24 c 

tt::: 22 

20 

18 

x Falls of the Ohio ( ~ holotype) 

o New York 

Ontario 

6 Ohio 

26r---.-l-----,~,------.l------~,------~ 

...... ---------........ 

_,---- X'\ \ 24 f- - 8 

---/ I 't::: 221- ,...- X o I ,...- X I 

r. h. 6 1 '- "1!'!11 • o XI 

201-- '--- X X // -- X , _____________ ,...,., 

-

-

18~--~----~~----~~----~~------~ 8.0 10.0 12.0 14.0 16.0 

d (mm) 

EXPLANATION 

P truncafum 

X USNM 163418 

0 osu 17786 

P. anna 

Lectotype 

6 osu 3743 

.... -, /, X 1 / / / / / / / / / / / / I / 

r__~// 

16·.0 

d(mm) 

FIGURE 29.-Mean number of major septa (fi) versus mean diameter (d) in Prismatophyllum spp. A, P. prisma; dashed 

polygon shows observed range of variation in P. ov~ideum (see B). B, P. ovoideum. C, P. truncata and P. anna. 
82 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

although the dark axial planes of the carinae can be 

seen if preservation is good. 

Discussion.-The above description is based on 10 

specimens of Prismatophyllum prisma from the Falls 

of the Ohio, in addition to 2 thin sections taken from 

the holotype. 

Cyathophyllum rugosum Milne-Edwards and 

Haime was originally designated the type -spec.ies of 

Prismatophyllum Simpson (1900, p. 218) who recogniz·ed that the species described by Milne-Edwards 

and Haime was not the same as Astraea rugosa Hall 

with which Milne-Edwards and Haime had identified 

their material. The Milne-Edwards and Haime species was renamed Prismatophyllum prisma by Lang 

and Smith, 1935a, and the lectotype (the original of 

Milne-Edwards and Haime, 1851, pl. 12, figs. 1a, b) 

was described and illustrated from thin sections for 

the first time by Smith, 1945. The specimens illustrated by Simpson are not this species. 

The figures (drawings) of Milne-Edwards and 

Haime show straight septa within the tabularium although the lectotype section's illustrated by Smith 

and additionallecotype sections in the British Museum (NH) illustrated here, clearly show the axial 

complex or whorl des,cribed above. The original 

figures misled Stainbrook (1940) into describing another Falls species with straight septa as P. prisma 

and to describing the species with the axial complex 

as P. cinctum. Comparison of thin sections from the 

lectotypes of P. prisma (pl. 46, figs. 1, 2) and P. 

cincta (pl. 47, figs. 1-5) leaves no question that they 

are conspecific. The species with straight septa is 

here de,scribed as Prismato_phyllum ovoideum 

(Davis). 

H exagonaria mesocincta Fraunfelter from the 

Grand Tower Li~mestone in Illinois and Missouri falls 

well within the range of morphologic variation of P. 

prisma ~and is approximately of the sa,me age. I consider it to be a junio·r synonym of P. prisma. 

Prismatophyllum prisma is particularly characterized by its axial complex or whorl and differs from 

most other species in this respect. The older P. subcincta (Stumm, 1948a) has an even better ·and more 

distinct whorl and may have been ancestral to P. 

prisma. P. subcincta was originally described from 

the Bois Blanc Formation in Michigan but may occur 

also in the lower Coral Zone (Schoharie ~age) at the 

Falls because I consider that the specimens rillustrated by Davis (1887, pl. 91-92) a-s Cyathophyllum 

colligatum is more likely P. subcincta than "H. cincta" ( =prisma) as sta:ted by Srtumm (1965, p. 43). 

If the relationship P. subcincta ~ P. prisma is correct, a continuation of the trend toward a simpler 

axial structure could have led toP. ovoideum, which 

appears in the succeeding zone of the Jeffersonville 

Limestone. 

All other Onesquethaw species of Prismatophyllum known to me have short major septa, or lack the 

axial complexities of P. prisma. 

Distribution.-Milne-Edwards and Haime ( 1851, 

p. 387) included two Jeffersonville localities, the 

Falls of the Ohio and Charleston Landing, Ind., plus 

larCJalities in Ohio, Iowa, and Michigan in their locality list for the species. Lang and Smith (1935a, p. 

558), Lang, Smith, and Thomas (1940, p. 104) and 

S·mith (1945, p. 99) all cite "Falls of the Ohio, 

Charleston Landing, Indiana" as the type locality 

without noting that the Falls and Charleston Landing were distinct and separate localities. The lectotype is clearly labeled Charleston Landing. 

Five specimens of P. prisma collected by me in 

1957 and 1961 all cam~e from the Middle Devonian 

part of the Carral Zone (Zone B) at the Falls of the 

Ohio. Four specimens collected by Guy Campbell, and 

now in the U.S. National Mus,eum, are from the same 

zone. The type of P. cincta is from the "coral zone." 

PTismatophyllum that I collected from Jeffersonville 

Zones A, C, D and E all belong to other species. 

Therefore, it ·seem's likely that in the Jeffersonville 

Limestone, P. prisma is restricted to the early Onondaga age, Zone B. 

As noted above, I consider that P. mesocincta 

Fraunfelter is P. prisma. The only other specimen 

that I have assigned to P. prisma is one collected 

from Columbus Limestone, Zone C, in a quarry 27 

miles (43 km) west-southwest of Columbus. These 

occurrences extend the known geographic range of 

the species we,st into the Mississippi Valley and east 

into Ohio. 

H exagonaria prisma of Carman, 1955, from the 

"Columbus Limestone" in the Chillicothe test core, 

taken 45 miles (72 km) south of Columbus, has an 

i~mperfect but distinct aulos and is not this species. 

Mat erial studied.-J effersonville Limestone, Coral 

Zone (Zone B), Charleston Landing, Ind.: Lectotype, de Verneuil collection, Ecole des Mines, Paris; 

the s·ections illustrated by S.mith (1945) were not 

seen, but two thin ~sections from the lectotype were 

studied and are illustrated here (BMNH R31730 and 

BMNH R31731). Falls of the Ohio : Holotype of 

Prismatophyllum cinctum, SUI 21198; measured 

and (o.r) illustrated herein, USNM 163399-405; and 

two additional specim·ens. Columbus Limestone, 

Ohio: Co~al Zone( Zone C), locality USGS 8289-SD, 

USNM 163406. Grand Tower Limestone, Missouri: 
SYSTEMATIC PALEONTOLOGY 83 

Holotype of Hexagonaria mesocincta Fraunfelter, 

SlUM 701. 

Prismatopbyllum ovoideum (Davis) 

Plates 49-52 

1881 Cyathophyllum rugosum Milne-Edwards and 

Haime. Quenstedt, p. 529, pl. 162, fig. 3la, b. 

1887 C. rugosum. Davis, pl. 90, fig. 1, pl. 93, fig. 3. 

1887 C. ovoideum Davis, pl. 93, fig. 1. 

1938 Prismatophyllum prisma Lang and Smith. Stewart, p. 50-51, pl. 9, figs. 13-15. 

1940 P. prisma Lang and Smith. Stainbrook, p. 273-

275, pl. 37, figs. 1, 2, 4. 

1944 Hexagonaria prisma (Lang and Smith). Shimer 

and Shrock, p. 95, pl. 30, figs. 11- 12. 

1948a H. ovoidea (Davis). Stumm, p. 15-16, pl. 1, fig. 

4, pl. 6, figs. 3-5. 

1948a H. prisma (Lang and Smith). Stumm, p. 17-18, 

pl. 1, figs. 1, 2, pl. 8, figs. 1-7. 

? 1948a H. bathycalyx Stumm, p. 22, pl. 2, fig. 1, pl. 9, 

figs. 3-4. 

1965 H. ovoidea (Davis). Stumm, p. 44, pl. 38, figs. 

4-6. 

1965 H. prisma (Lang and Smith) Stumm, p. 44, pl. 

38, figs. 7-9. 

OccurTence of type material.-"Decomposed chert 

of the Middle Devonian near Louisville" (Davis 

1887, explanation, pl. 93). Probably Zone C ( Co~alStromatoporoid Zone), at or near the Falls of the 

Ohio. 

Diagnosis .-Prismatophy llum with medi um-.sized 

corallites in which the attenuate major septa extend 

nearly to the axis and are straight within the tabula,rium; zig~ag CJarinae are short and weak. 

External features.-Cerioid coralla are lens or 

dome shaped; available specimens are as much as 15 

em in diameter and 8 em in height. Coralites a.re 

polygonal with straight ·sides in top view ~except on 

offsets. Calcitic coralla tend to bl"eak betw·een individuals; separa-te corallites :are prismatic with irregular rugae and l~arger undulations along the 

l~ength of each side; septal grooves and interseptal 

ridges are also present. Cor,allite di1ameters in seven 

cor all a from the Falls of the Ohio range from 7.0 to 

16.5 mm; mean diametel"!s range from 10.1 to 13.7 

mm (table 26). Data from the holotype are not included in table 26 as this specimen is poorly preserved and has not been sectioned. Measurements 

and counts made on the surface of the holotype indicate that it falls at the smaller end of the observed 

range of variation (d=8 to 18 mm) as shown in 

figure 29. Studied specimens from New York, Ohio, 

and Ontario fall into the size range of Falls of the 

Ohio specimens or are smaller (table 26 and fig. 29). 

Increase is lateral with offsets expanding rapidly 

to mature size. The calice in silicified Falls of the 

Ohio specimens has an axial pit, nearly flat platform, 

and steeply rising outer wall; total depth is one-half 

to two-thirds the diameter. Galice, holotheca, and 

other features of the corallum exterior are not 

known from the New York or Ontario specimens. 

Internal features.-Septa are radially arranged 

and uniform in size, extending from the outer wall 

to the margin of the tabularium where the minor 

septa dilate slightly ~and terminate. The major septa 

are somewhat dilated in the tabularium; they extend 

to the axial region where most of them stop short of 

the axis itself. In a few corallites the major septa 

reach the axis and touch. Minor septa tend to be 

more irregular in length in smaller corallites, possibly an immature character. 

The number of major septa in the Falls of the 

Ohio sample and in specimens from New York, Ontario, and Ohio, are shown on table 26 and figure 29. 

Studied specimens from New York, Ontario, and 

Ohio fall within the range of intercolony variation 

of the Falls of the Ohio sample (n = 19.6-24.2). 

Most septa a.re carinate in the dissepimentari urn 

but the oarinae are short and weak and even absent 

in .som·e parts of the coralla. All carinae are zigzag. 

Where strongest, spacing of th·e carinae ranges from 

four to eight per 2.5 mm, counted on one side of a 

septum. 

In transverse sections, the corallite walls are 

straight except a~ound offs·ets where the walls tend 

to be rounded. Corallites are everywhere in contact 

but tend to break apart along wall midplanes in calcite specimens. Walls range in thickness from 0.1 to 

0.4 mm but are commonly less thon 0.15 mm. 

The tabularia !"lange from 3 to 5 mm in diameter, 

aver.aging approximately 4 mm. Tabulae are incomplete and clo·s,ely spac~ed. No inner and outer zones of 

tabellae are defined, but the inner ones tend to be 

horizontal whereas the outer ones are inclined toW3!rd the axi,g so that the overall pattern is concave. 

The dissepiments are ·small and globose, arranged 

in four o·r m'ore rows depending on size and local 

width of dissepim·enta.ri urn. The rows are steeply inclined near the axis, gently inclined to almost horizontal away from the axis, but commonly steeper 

again at the periphery. 

Microstructure.-Preservation of fine detail is 

poor in the available sp·ecimens, but the structure 

appears to be similar to that of the family. In transverse sections the midlines of septa, ca.:dnae, and 

walls are dark; tabulae, dissepiments, and surface 

layers of the septa, oarinae, and walls are light and 

more transparent. No structural detail is well preserved in either type of tissue, but there is a suggestion that the light wall and septal tissue is composed 
TABLE 26.-Prismatophyllum ovoideum (Davis), summary data 

Falls of the Ohio 1 New York Ontario 

Intracolony Inter- NYSM NYSM USNM UMMP Mini- Maxi- Mean -::o!onv 12829 12830 163414 24178 mum mum 

N ---------- 7 16 --- 7 20 15 11 9 

n: x ------- 19.6 24.2 21.2 ~1.2 22.0 20.7 20.5 21.6 

OR ----- 17-22 211-26 --- 19.6-24.2 21-23 19-22 19-~3 21-23 

8 ------- .73 1.72 --- 1.59 .60 .81 1.21 .73 

v ------ 3.6 7.1 5.4 7.5 2.7 3.9 5.9 3.4 

d: x ------- 10.1 13.7 --- 11.6 11.4 10.6 10.1 8.8 OR _____ 7.0-12.0 8.5-16.5 --- 10.1-13.7 9.0-15.0 7.0-13.5 6.0-13.0 6.7-10.5 

s ------- .44 2.77 --- 1.36 2.01 1.62 2.02 1.34 

v ------ 3.3 24.5 16.9 11.7 17.6 15.3 20.0 15.3 

1 Holotype not included. 2 d measured in only five corallites because of crushing. 

Ohio 

UMMP USNM 24176 163415 

10 7 

21.00 21.0 

19-22 210-22 

1.15 .82 

5.5 3.9 

7.2 8.9 

6.0- 8.8 8.0-10.0 

.93 .80 

13.0 9.0 

USNM 

163417 

2 10(5) 

21.3 

20-25 

1.57 

7.4 

10.9 

8.0-13.5 

2.22 

20.4 

00 

~ 

z 

c 

z 

0 

~ 

00 

~ 

;a 

0 

~ 

""0 

::c: 

0 

0 

~ 

0 

z 

...... 

> 

~ 

~ 

q 

C) 

0 

00 

tz:j 

0 

0 

~ 

> 

~ 

00 
SYSTEMATIC PALEONTOLOGY 85 

of fibers oriented at right angles to the structural 

surfaces. 

Discussion.-The ~above description is based on 

specimens from a wide geographic area. The largest 

sam:ple (seven coralla plus the holotype) is from 

Jeffersonville Limestone, Zone C, and individual 

specim~ens from N·ew York, Ontario, and Ohio add 

little to the observed r~ange of variation of this sample. The holotype has the smallest corallites of any 

Falls of the Ohio specimen but is matched in this 

rerspect by Ontario and Ohio specim·ens (fig. 29). 

Two fragments of this species in the collections of 

the New York State Museum are labeled "Onondaga 

Limestone, western New York." There is no rock 

matrix attached to either specimen but the color and 

pr·eservation suggest that th:ey were collected from 

the biohermal facies of the Edgecliff Member. No 

other specimens of the ~species ar·e known from New 

York. Four coralla were studi:ed from th~e lower Detroit River ("Amherstburg" of Ehlers and Stumm 

1951) at Beachville, Ontario. Size and s~eptal number 

data for the New York and three of the Ontario 

specimens are given in table 26 and figure 29. 

Stumm (1948a, 1965) separated his P. uprisma" 

from P. ovoideum on the basis of the greater corallite 

size and septal number of the former and on various 

other morphological features that I consider to be a 

product of individual variation. Prismatophyllum 

bathycalyx Stumm may be synonymous also. The 

holotype has very large corallites but is poorly preserved so that other feature's are difficult to analyze. 

Septa extend to or nearly to the axis and a.re arranged as in P. ovoideum. 

Distribution.-J~etfersonville Limestone, CoralStromatoporoid Zone (Zone C), Falls of the Ohio; 

Columbus Limestone, Zone C and ?H, Ohio; Onondaga Limestone, Edgecliff ( ?) bioherm facies, western New York; Detroit River Group, Amherstburg 

Formation, Beachville, Ontario. 

All specimens of P. ovoideum that were collected 

by the author are from units equivalent to th~e Edgecliff. Specimens illustrated by Stewart (1938, pl. 9, 

figs. 13-15) and Stumm (1948a, pl. 8, figs. 3, 5) from 

Columbus Limestone, Zone H (Paraspirifer Zone) 

in northern Ohio are apparently of this species. The 

precise stratigraphic position of the holotype and 

some other Falls specimens included in my synonymy 

is not known, but all specimens in my collection or in 

the Campbell collection (USNM) a,re from Zone C. 

The New York specimens are labeled only "western 

New York" but are thought to be from the Edgecliff 

Member on the basis of their color and preservation 

characters. 

M aterial.-Falls of the Ohio, Jeffersonville Formation: Holotype, MCZ 7910; measured or illustrated 

herein, USNM 163407-413 (163407-412 and three 

other studied ~specimens are from Zone C, collected 

by Guy Oampbell or by the author). W·estern New 

York, Onondaga Limestone, Edgecliff ( ?) bioherm 

facies: NYSM 12829-30 (both illustrated). Beachville, Ontario, Amherstburg Formation, Detroit River Group: USNM 163414 and UMMP 21478 (illustrated), and UMMP 24176-24177. Columbus Limestone, Zone C, Ohio: USNM 163415-417, and three 

other specimens. 

Prismatopbyllum truncata Stewart 

Plates 53, 54 

1938 Prismatophyllum truncata Stewart, p. 51-52, 

pl. 10, figs. 1, 2. 

1948a H exagonaria curta Stumm, p. 26-27, pl. 4, figs. 

2, 3, pl. 12, figs. 1-6. 

1948a H. truncata (Stewart). Stumm, p. 28-29, pl. 12, 

figs. 7-8. 

1955 H. curta Stumm. Stumm, cards 178-179. 

1955 H. truncata (Stewart). Stumm, card 223. 

1965 H. curta Stumm. Stumm, p. 43-44, pl. 38,· figs. 

1-3. 

not? 1970 H. truncata (Stewart). Brice, p. 280-281, pl. 16, 

figs. 3a-c. 

Occurrence of type material.-Stewart (1938, p. 

101) reported the holotype to be from the "Blue 

Limestone, Whitehouse Quarry," Lucas County, 

Ohio. I consider thi,s unlikely and suspect that it 

prob~bly oam~e from the Dundee Limestone or the 

highest pa·rt of the Columbus Limestone (Zone H) 

in northern Ohio. (See discus~sion.) 

Diagnosis.-Prismatophyllum with medium-sized 

corallites. Major s~epta extend just into the tabularium and minor septa are three-fourths as long to 

nearly ~equal in length; subyard-~arm carinae ·predominate. Tabulae are comple·te or incomplete, more 

or less flat in the axial region. Dissepiments tend to 

be steeply inclined and elongate on the innner side of 

the dis.sepimentari urn. 

External features.-The only known Onondaga 

specimen is a fragm.ent of a cerioid corallum that 

was broken befo,re pres~ervation ; dimensions of the 

fragment are: length, 8 em; width, 7 em; height, 11 

em. No part of the exterior surface of the colony is 

preserved. 

Longitudinal sections suggest that the calices include an axial flat area approximately one-third the 

diameter of the coral with walls sloping steeply upward to the margin. Calice depth is som·ewhat greater than the diameter of the flat axial area and there 

is no peripheral platform. 

Diameters of 15 corallites in the specimen range 
86 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

from 10 to 17 mm and average 12.5 mm (table 27, 

and fig. 29). 

Increase is lateral and frequent in the part of the 

colony preserved, as s~everal offsets can be recognized 

by their small size and low septal number. Offsets increase rapidly to mature size. 

Internal features.-Septa are radially arranged. 

The major septa extend only a short distance into the 

tabularium; the minor septa are about three-fourths 

as long and are restricted to the dissepimentarium. 

The minor septa are thinner than the major septa; 

all septa dilate slightly at their axial ends. In 15 

corallites the number of m~ajor septa varies from 22 

to 26, averaging 24.1 (table 27 and fig. 29C). All 

septa are carinate; subyard-arm carinae predominate but zigzag ones are locally common. The carinae 

are closely spaced, five or six per 2.5 m.m being 

common. 

The corallite walls a.re straight or gently curved; 

in detail they m.ay or may not zigzag depending on 

whether the septa in adjacent corallites alternate or 

correspond in position. Corallite walls range in thickness from 0.2 to 0.3 mm; the thinner walls are more 

common. 

Tabularia are comparatively large, occupying approxim·ately one-third of the diameter. Tabulae are 

complete or incomplete, axially flat or gently convex, 

bent upwards at the margin of the tabularium. 

In the peripheral one-half of the dissepimentaria, 

the diss.epiments are small, globose, and gently inclined. Toward the tabula.ri urn they become large, 

elongate, and steeply inclined. The dissepimentarium 

is composed of from five to seven rows of dissepiments. 

Microstructure.-Quality of preservation varies 

widely in the Onondaga specimen, part of which is 

silicified. Locally the calcite preservation is good 

enough to show that the microstructure is the same 

as described for the family. 

TABLE 27.-Prismatophyllum truncata Stewart and P. anna (Whitfield) intracolony data 

P. truncata P. anna 

New York Ohio Ohio 

USNM 1 0SU 1 UCMP osu 163418 17786 34218 3743 

N ---------- 15 7 15 17 

n: x 24.1 20.4 19.0 18.6 

OR ----- 22-26 18-23 18-20 18-20 8 ------- 1.1 1.7 .8 .7 

v ------ 4.6 8.4 4.4 3.8 

d: x 12.5 10.3 9.2 7.9 

OR ----- 10-17 7.0-14.5 5.0-12.0 3.5-10.0 8 ------- 1.8 2.9 2.1 2.0 

v ------ 14.1 28.0 22.8 25.1 r ----------- .94 .76 .42 .20 

p ---------- <.01 .05 .90 >.50 

1 Holotype. 

Septa and carinae have an irregular dark midplane enveloped in light, featureless tissue which is 

similar to that co.mposing the tabulae and dissepiments. The s·epta indent the outer wall and the light, 

surface tissue of the septa may be continuous with 

the wall tissue. The light wall tissue, however, is 

thicker and appears to be formed of fibers oriented 

at right angles to the surf,ac.e of the wall. Each wall 

segment was built by two adjacent corallites. The 

depo,sits of each are distinct and separated by a dark 

mid-surface1 which is more uniform in thickness 

than the presumed trabecular layer in the septa. 

Discussion.-The above description is based on the 

only known New York specimen of the species. The 

specim·en is morphologically close to the Ohio bolotype with which it has been directly compared. The 

Ohio ·specimen has smaller corallite diameters and 

fewer septa (table 27, fig. 29C) and somewhat longer 

minor septa, but both specimens show considerable 

intercolony variation in these characters. Offs.ets in 

the Ohio spec:imen show occasional discontinuous 

septa, which have not been observed in the New 

York one. 

Stewart stated that the species occurs in "hemispherical masses up to a foot or more in diameter" 

(1938, p. 51) and indicated that it occurs in the 

Columbus Limestone as well as in the "Blue" limestone (1938, p. 52). However, the only specimen in 

the Ohio State Univ·ersity type collection that can be 

related to Stewart's W'Ork is the holotype, consisting 

of two small fragments of a corallum and two thin 

sections cut from the fragments. The thin sectiOI!S 

were illustrated by Stewart (1938) and by Stumm 

(1948a, 1955). Stumm (1948a, p. 10) lists the species from the "Blue" limestone and questionably 

from the Dundee Limestone. 

P. truncata diff·ers from other ENA sp·ecies of 

Prismatophyllum in its short m.aj or septa and predominance of yardarm carinae. In comparison with 

other species that have short major septa, it is characterized by the coarse, elongate dis.s·epiments in the 

inner part of the diss·epimentaria. P. truncata is 

closest morphologically toP. anna (Whitfield) which 

is briefly redescribed below although it does not occur in New York or Ontario. 

P. truncata differs fro·m P. anna (Whitfield) in its 

larger cor.allites and greater number of septa and in 

its coarser and more variable dissepiments. Stewart 

(1938) and Stumm (1948a) .separated the species on 

the same characters. In large collections from the 

"Blue Limestone" beds of the Silica Formation of 

Stewart (1927) near Sylvania, Ohio, I have found 

only P. anna. P. truncata is the common Prismato-
SYSTEMATIC PALEONTOLOGY 87 

phyllum in Columbus Limestone Zone H, in northcentral Ohio, and in the upper Jeffersonville Limestone at the Falls of the Ohio (where Stumm described it as Hexagonaria curta). Stewart cited P. 

truncata from "Columbus limestone ( ?) : Franklin 

County, locs.? Blue Limestone: Lucas County" 

[Whitehouse quarry] (1938, p. 52) but noted the 

holotype to be from the latter locality (1938, p. 101). 

Hexagonaria truncata (Stewart) of Brice, 1970, 

from the Middle Devonian of Afghanistan, is probably not this species. It differs in the nature of its 

carinae, in having thickened septa, and in its "tendency to become phaceloid" (Brice, 1970, p. 280). 

The microstructure is not described or illustrated 

but the thickness of the septa and the bulbous appearance of the carinae suggests coarse disphyllid 

trabeculae. The Afghanistan .species is probably a 

Hexagonaria and unrelated toP. truncata Stewart. 

Distribution.-P. truncata is the only colonial rugose coral known from the 8eneca Member of the 

Onondaga Limestone. The single specimen was collected fro·m above the Chonetes lineatus Zone (Zone 

J, Oliver, 1954, p. 629-631) by Professor J. W. Wells 

of Cornell University. 

The species is common in Columbus Limestone, 

Zone H, in north-central Ohio and may occur in the 

overlying Delaware or Dundee Limestone. It also occurs in the Jeffersonville Limestone, Zones D and E 

at the Falls of the Ohio. 

Material.-Holotype, OSU 17786; New York specimen, USNM 163418. Additional specimens from 

Columbus Limestone, Zone H and Jeffersonville 

Limestone Zones D and E in the USNM. 

Prismatophyllum anna (Whitfield) 

Plates 55, 56 

1882 Stylastrea anna Whit,eld, p. 199-200. 

1891 S. anna Whitfield. Whitfield, p. 520-521, pl. 6, figs. 1-

5. 

1893 S. anna Whitfield. Whitfield, p. 420, pl. 2, figs. 1-5. 

1938 Prismatophyllum annum (Whitfield), Stewart, p. 49, 

pl. 9, figs. 11, 12. 

1948a Hexagonaria anna (Whitfield). Stumm, p. 25; pl. 5, 

fig. 3; pl. 11, figs. 1-3; pl. 13, figs. 1, 2; pl. 14, figs. 

3-6. 

1955 H: anna (Whitfield). Stumm, cards 156, 157. 

1967 H. anna (Whitfield) Stumm, p. 105-107, pl. 1, figs. 1-

21. 

1969 H. anna (Whitfield). Stumm, p. 240, pl. 10, figs. 5, 6. 

1970 H. anna (Whitfield). Stumm, p. 82, pl. 1, figs. 1, 6, 10, 

pl. 2, figs, 1, 2, 6, 7, 9, 10. 

Occurrence of type material.-The syntypes are 

from the "Upper Helderberg, Paulding Co., Ohio" 

(Whitfield, 1899, p. 146). According to Stewart 

(1938, p. 50) the specimens are probably from the 

"Blue" limestone layer (basal unit of Silica Formation of Stewart, 1927) rather than from the Columbus Limestone, as would be indicated by the term 

"Upper Helderberg." 

Remarks.-This species has not not been found in 

New York but is included for comparison with P. 

truncata Stewart. Thin sections of some of the syntyp·es are here illustrated for the first time. 

Diagnosis.-Prismatophyllum in which major and 

minor s•epta are of approxim·ately equal length and 

limited to the dissepimentarium and in which the 

dissepiments are uniformly small and globose. 

External features.-N one of the specimens studied 

are complete or show the external characters. The 

largest fragment, the lectotype, measures 12 by 7 by 

6 em, parallel ·and at right angles to the corallite 

axes. 

Longitudinal sections indicate that the calices 

have a fiat or gently arched floor formed by the uppermost tabula. The floor is approximately one-third 

the diameter of the corallite. The sides of the calice 

are steep, becoming gently inclined toward the periphery. 8eptal ridges, mark the calice walls but 

apparently not the floor. Galice depth is approximately the same as the diameter of the floor. 

Increase is lateral, as in most other species of the 

genus. Corallite diameters in two coralla, range from 

3.5 to 10.0 and from 5.0 to 12.0 mm, with mean diameters of 7.9 and 9.2 mm, respectively (table 27, 

fig. 29C). Inclusion of a few immature corallites 

affect the observed range but has no significant effect 

on the means. 

Internal features.-The septa are radially arranged. Major septa extend to or just into the tabularium; minor septa extend nearly to the tabularium 

and in some corallites and parts of corallites are only 

slightly shorter than the major septa. There is no 

significant differ·ence between major and minor septa 

in thickness, or in number, size, and spacing of 

carinae. Zigzag carinae predominate but subyardarm carinae are common ; carinae -are strongly to 

weakly developed in different parts of the same 

corallite. Number of major septa ranges from 18 to 

20 in each corallum (table 27, fig. 29C). 

The corallite walls are straight or gently curved; 

in detail they commonly zigzag as septa in adjacent 

corallites alternate in position. Corallite walls range 

in thickness from 0.1 to 0.3 mm. 

Tabularia occupy more than one-third of the diameter of .the corallites. Most tabulae are incomplete 

and gently arched. Tabularium diameters range 

from 3.5 to 5.0 mm. 
88 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Dissepim·ents are commonly small and globose, but 

locally are quite elongate. They ·are steeply inclined 

next to the tabulari urn, becoming gently inclined or 

horizontal at the periphery. The dissepimentaria are 

composed of from five to nine rows of dissepiments. 

Microstructure.-Preservation of detail i·s poor in 

the specimens studied, but the septal and wall structure seem to be the same as described for the other 

species of the family. The wall shows the dark, uniform mid-plane with fibers at right angles to it forming light-colored tis.sue on either ·side. The septal 

structure is obscure with only local suggestion of a 

dark center line. 

Discussion.-The ~above description, included here 

for comparison with H. truncata and to illustrate 

thin sections if the lectotype of H. anna, is based on 

5 Ohio speci~mens from the original Whitfield collection and 16 additional specimens from the same 

stratigraphic unit. 

Whitfield fin~t illustrated the .species in 1891 (pl. 

6, figs. 1-5) and repeated the same illustrations in 

1893 (pl. 2, fig. 1-5). In neither publication did he 

indicate whether the five figures were from one or 

more than one colony, nor did he indicate the number 

of specimens on which his species was based. In 

1899, he listed the illustrated specimen (s) as being 

located at the University of Oalifornia and referred 

to it (them) as "type," this suggesting that a single 

colony was involved (Whitfield, 1899, p. 146). Stewart based her description on two specimens labeled 

paratypes from the Whitfield collection in the Orton 

Museum, Ohio State University. Both of these were 

sectioned, ·and two sections from one specimen were 

illustrated. Stumm (1948a) redescribed the "pa.ratypes" and selected one as "lectotype" ( p. 26) in the 

belief that "Whitfield's holotype, the original of his 

plate 2, figures 1-5, is lost." Both Stewart and 

Stumm overlooked Whitfield's statement of 1899, in 

which he indicated the location of his specimens. 

Stumm (1969, p. 240, pl. 10, figs. 5, 6) briefly noted 

and illust~ated the Whitfi·eld type from thin sections 

prepared in my laborato.ry, but he cited them only as 

a paratype. 

The type lot in the Museum of Paleontology, University of California, includes three fragments of 

coralla that .are clearly the originals for Whitfield's 

figures 2, 4, and 5 and probably his figure 3. Whitfield's figure 1 represents a nearly complete corallum 

which may have been broken up to provide the sections and views shown in the other figures. (If this 

is so, some pieces are missing, .as the remaining ones 

cannot be fitted together.) As more than one colony 

may be involved, I here select specimen UCMP 

34218, a part of which was the original of Whitfield's figure 4, as lectotype. Before thin sectioning, 

this specim·en measured 12 X 8 X 7 em parallel and at 

right angles to the corallite axes. The illustrated thin 

sections are from this specimen. The "lectotype" 

selection of ~tumm (1948a, p. 26) is invalidated by 

the existence of the specimen (s) designated as 

"type" by Whitfield. 

Distribution.-According to Stewart (1938, p. 50) 

the Whitfield specimens are probably from the 

"Blue" lim·estone of the Silica Formation of Stewart 

(1927). Other studied specimens are from this same 

stratigraphic unit in northw·estern Ohio. The species 

has been reported from the upper Dundee Limestone 

of the same area and from the lower part of the 

Traverse Group in Michigan. 

.llfaterial.-Lectotype, UCMP 34218; illustrated 

paratype:s, UCMP 34216 and 34217. Illustrated by 

Stewart (1938, pl. 9, figs. 11, 12) and Stumm (1948a, 

pl. 11, figs. 1, 2), OSU 37 43-2; unillustrated specimen, OSU 3743-1. The two OSU specimens are from 

the Whitfield collection. 

I have also examined thin sections of 16 coralla 

from the "Blue Limestone" near Sylvania, Ohio 

(USGS collns. 7333, 7776, 7778, 7779-SD). 

Genus ASTEROBILLINGSA Oliver, 1974 

not 1917 Phillipsastraea (Billingsastraea) Grabau, p. 

957. 

part 1937 Billingsastraea Grabau. Stumm, p. 437-438. 

not 1937 Radiastraea Stumm, p. 439. 

part 1944 Billingsastraea Grabau. Shimer and Shrock, p. 

97. 

part 1949 Billingsastraea Grabau. Stumm, p. 35. 

not 1951 Billingsastraea Grabau. Prantl, p. 7-8. 

part 1951a Billingsastraea Grabau. Ehlers and Stumm, p. 85. 

part 1953 Billingsastraea Grabau. Ehlers and Stumm, p. 1. 

part 1955 Billingsastraea Grabau. Stumm, card 230. 

1956b Billingsastraea Grabau. Hill, p. F280. 

part 1958 Billingsastraea Grabau. Cranswick and Fritz, p. 

41-42. 

not 1958 Billingsastraea Grabau. Schouppe, p. 235-237. 

not 1962 Billingsastraea Grabau. Soshkina and Dobrolubova, p. 336. 

part 1964 Billingsastraea Grabau. Oliver, p. 2-3. 

not 1965 Billingsastraea Grabau. Strusz, p. 547. 

1967 Billingsastraea Grabau. Sorauf, p. 24-25. 

not 1967 Billingsastraea Grabau. Pickett, p. 19-20. 

part 1967 Billingsastraea Grabau. Scrutton, p. 275-276. 

1974 Asterobillingsa Oliver, p. 167-168. 

Type species.-Asterobillingsa magdisa Oliver. 

Schoharie Grit, Clarksville, N.Y. 

Diagnosis.-Astreoid to thamnasterioid cylindrophyllinid corals with calices having a central pit and 

a broad horizontal or re·flexed peripheral platform, 

com·monly with a ·raised zone around the pit. S.epta 

are radially arranged and lightly to heavily carinate 
SYSTEMATIC PALEONTOLOGY 89 

with zigzag to subyard-arm carinae. Major and minor septa extend from the periphery; minor septa 

terminate at the outer margin of the tabularium; 

major septa extend to or almost to the axis. At their 

peripheral ends the septa abut against those of adjac·ent corallites, are continuous into the next corallite, or more rarely are partly discontinuous. The 

tabularium is narrow and composed of closely 

spaced, more or less horizontal, complete and incomplete tabulae. The dissepimentarium is composed of 

gently to strongly globose dissepiments that are horizontally arranged, exc·ept next to the tabularia where 

they are inclined toward the corallite axes. 

Discussion.-Asterobillingsa was proposed to replace Billingsastraea Grabau which is unuseable in 

its conventional concept and is probably a junior 

synonym of Arachnophyllum Dana. Billingsastraea 

was established by G~abau (1917, p. 957) who merely used the word in a fossil list as "Phillipsastraea 

(Billingsastraea) verneuili E. & H." No description 

or other indication of his taxonomic concept was 

given. Clearly, the specific reference was to Phillipsastraea verneuili Milne-Edwards and Haim·e (1851, 

p. 447-448, pl. 10, fig. 5), stated to be from Wisconsin. The type .specimen of this species has been assumed to be a drift s.pecimen, originating in either 

the Onondaga or the Bois Blanc of Ontario or Michigan (Stumm, 1949, p. 35). None of the subsequent 

descriptions or diagnoses of the genus or species 

have been based on the holotype; all have been based 

on specimens of one or anoth·er of three species or 

subspecies of ·astreoid rugos·e corals from the Onondaga ·and the Bois Blanc in Ontario and New York. 

I chose to use a new generic name for these three 

species and their congeners for the following 

reasons: 

1. Inquiries and searches by interested specialists 

have failed to locate the holotype which is presumably in the de V erneuil Collection, Ecole 

des Mines, Paris. Through the kindness and 

with the cooperation of Dr. P. Semenoff-TianChansky, I also m·ade an unsuccessful search 

for the specimen (1969). Because the specimen 

cannot be found and its source is not known, 

the generic group can be stabilized only 

through a new genus with a known type 

species. 

2. Although the Milne-Edwards and Haime specim·en has not been found, Dr. Semenoff located 

another specimen in the collection that is remarkably like the original in surface appearance (the only view of the original that was 

illustrated). This specimen is labeled as being 

from an area, now in northeastern Iowa, that 

was a part of the Wisconsin Territory from 

1836 tO 1838. This s·econd specimen is clearly 

an Arachnophyllum and probably from the 

Hopkinton Dolomite (Niagaran) that outcrops 

in northeastern Iowa and southwestern Wissin. Therefore· it is possible (even probable?) 

that the holotype of Billingsastraea verneuili 

would prove to be an Arachnophyllum if found. 

3. The only known local source for a Devonian coral 

in Wisconsin, would be the small area of Devonian rocks at Milwaukee, Wis. However, no 

"BiUingsastraea" are known from this area, 

and there is no indication that this area was 

known, or that collections from this area were 

available, before about 1860 (Cleland, 1911, p. 

21). 

4. The suggestion that the holotype of Billingsastraea is a drift specimen ultimately derived 

from Michigan or Ontario does not pass critical examination. Possible sources would be the 

Mackinac Straits area of the northern Lower 

Peninsula of Michigan, and southwestern Ontario. "Billingsastraea" occurs in both areas, 

but transportation to Wisconsin would require 

.south westerly and westerly glacial movement 

across the Lake Michigan bas:in. Reference to 

the Glacial Map of North America (Flint, 

1945) and other generalized sources, suggests 

that ice move·m·ent was generally southerly, 

parallel to the Lake Michigan trough, and that 

a source east of the Lake is unlikely for any 

drift specimen found in Wisconsin. A source in 

the Silurian of southwestern Wisconsin is far 

more likely. 

Any one of these reasons is an adequate argument 

for abandoning Billingsastraea and starting fresh. 

A new start is long overdue and is logically made by 

basing a new genus on a species that will maintain, 

insofar as is practical, the common concept of "Billingsastraea." I tried to do this by basing Asterobillingsa on the species most often referred to as 

"verneuili" and s·elected a name for the genus that 

is reminisc-ent of both the first name applied to this 

genus by Vanuxem and the name under which it was 

known for so many years. 

Other names that have previously been suggested 

to be synonyms of "Billingsastraea" are Asterocycles Vanuxem, 1842, Radiastraea Stumm, 1937, 

and Keriophylloides Soshkina, 1951. 

Asterocycles Vanuxem and its type species (A. 

confiuens Vanuxem, 1842, p. 136) were described 

but not illustrated. They were probably based on a 
90 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

specimen of A. magdisa steorra (described below) 

but the holotype is lost or unrecognizable and there 

is no way to be certain of its identity. Vanuxem's 

genus and species are indeterminate and not suitable 

as a replacement for Billingsastraea. 

Radiastraea Stumm (type species R. arachne 

Stumm, 1937, p. 439-440, pl. 53, fig. 13, pl. 55, figs. 

Sa, b) is from the Emsian part of the Nevada Formation in Nevada. The holotype has been well described and illustrated and is available for study. 

Stumm originally described the form as having 

m·ajor septa "usually deflecting at their axial ends to 

form a smaH tubular ring," and without carinae 

(1937, p. 439) but later corrected his description and 

considered R. arachne ~ be a "weakly carinate 

species of Billingsastraea" (1949, p. 35). Most subsequent workers have agreed (for ex·ample, Oliver, 

1964), but Pedder (1964, p. 446-447) and Jell 

(1969, p. 63) have separated the two genera, and I 

now ·agree that they are distinct although I do not 

necessarily .accept all of Pedder's or Jell's points. 

Asterobillingsa includes a group of species that, in 

addition to the family characters and an astreoidthamnasterioid form, have incomplete tabulae that 

form a flat or gently concave or convex pattern. 

Further, specimens of A. spp. normally have raised 

(reflexed) calice rims and heavily carinate septa. In 

contrast, Radiastraea arachne (pl. 66) has incomplete tabulae, som~e of whioh are strongly arched in 

the periaxial region. (In this respect, Radiastraea 

arachne is very similar to M artinophyllum ornatum 

Jell and Pedder, 1969, p. 736-737.) Further, R. 

arachne lacks raised calice rims and is only weakly 

carinate. Family assignm~ent of Radiastraea is uncertain. It is not a dis.phyllid and may be a "par.adisphyllinid" a;s suggested by Jell (1969, p. 63). 

Keriophylloides Soshkina (type species Keriophyllum astreiforme Soshkina, 1936, p. 62-64, figs. 

71-74; 1951, p. 102-103, pl. 19, figs. la-c; Eifelian, 

Urals) was plac~ed in synonymy with "Billingsastraea" by Soshkina and Dobrolubova ( 1962, p. 336). 

However, most astreoid corals that have been assigned to Keriophylloides 'Show marked septal thickening and seem to be phillipsastraeids as suggested 

by Jell (1969, p. 63). The morphology of the type 

species is unclear. The "carinae" of Soshkina are 

not clear in her illustrations but are unlike craspedophyllid carinae and the septa appear to break into 

separate strands or to becom~e discontinuous in 

peripheral regi()llls. These characters suggest that 

Keriophylloides astreiforme is quite unrelated to the 

Craspedophyllidae. 

Prantl (1951) described two Emsian forms from 

Bohemia as Billingsastraea bohemica and B. branikensis. These have recently been restudied and 

placed in a single species of Iowaphyllum by Oliver 

and Galle (1971). 

Sohouppe ( 1958, p. 235-237) referred to Billingsastraea some Late Devonian corals previously included in Phillipsastraea but which lacked horseshoe 

dissepim,ents. These forms have a zone of septal 

thickening at the inner margin of the dissepimentari urn and display tra:becular fans. This indicates a 

relationship to the ph:illipsastraeids rather than the 

m~aspedophyllids. (See discussion by Sorauf, 1967, 

p. 24-25.) 

Strusz (1965) and Pickett (1967) followed 

Schouppe's conc·ept of the genus, basing their description primarily on Australian and European 

astreo~id corals with dilated, fusiform septa; I consider these to be probable phillipsastraeids. 

Scrutton (1967, p. 275-278) discussed Billingsastraea ·and classified it with M arisastrum R6zkowska, H aplothecia Frech, and other genera in the 

Marisastridae. H·e noted that Billingsastraea differed from the other genera in its unthickened septa 

and tentatively assigned one Devon species to Billingsastraea while noticing in this species "slight 

dilation of the septa, which are also non-carinate.'·' 

I have ex~amined Scrutton's thin sections and am of 

the opinion that these differences are significant and 

that the species is a phillipsastraeiid, possibly a 

Frechastraea. 

"Billingsastraea" as used for corals of Early and 

Middle Devonian age in eastern North America has 

included astreoid forms that I interpret as belonging 

to two groups, cylindrophyllinid.s and zaphrentids. 

Asterobillingsa is esta.blished for the cyllindrophyllinids ·and is considered to include the following 

species: 

Early Onesquethaw Stage 

A. affinis (Billings). Grande Greve Formation, Quebec. (See Oliver, 1964.) 

Middle Onesquethaw Stage 

A. magdisa magdisa Oliver. Schoharie and Bois 

Blanc FO'rm·ation, New York and Ontario; Edgecliff bioherm facies, New York. 

A. rugosa (Hall). Bois Blanc Formation ( ?) , Leroy, 

N.Y. 

Late Onesquethaw Stage 

A. magdisa steorra Olive·r. Edgecliff Member, Onondawa lim·estone, New York and Ontario. 

Tioughnioga Stage 

A. canadensis (Ehlers and Stumm). Widder Formation, Ontario. 
SYSTEMATIC PALEONTOLOGY 91 

The following species that have been assigned to 

Billingsastraea in North America are considered not 

to be Asterobillingsa and tentatively reassigned as 

indicated: 

Billingsastraea arachne (Stumm) . = Radiastraea. 

(See pl. 66, figs. 1-4.) 

B. billingsi (Calvin) ,=Heliophyllum. 

B. confiuens (Hall) ,=Heliophyllum. 

B. ingens (Davis) ,=Heliophyllum. (See Stumm 

1965 for synonymy.) 

B.longicarinata Ehlers and Stumm,=Heliophyllum. 

B. monticula Ehlers and Stumm, redescribed below 

as Heliophyllum monticulum. 

B. nevadensis Stumm,= ?cyathophyllid. 

B. pauciseptata Ehlers and Stumm,= H eliophyllum. 

B. rockportensis Ehlers and Stumm,=Heliophyllum. 

B. romingeri Ehlers and Stumm,=Heliophyllum. 

B. scheii (Loewe), possibly=Radiastraea. 

B. southworthi Ehlers and Stumm=Heliophyllum. 

B. stirps Crickmay, probably=Radiastraea. 

B. tapetiformis Crickmay, probably=Radiastraea. 

B. trichomisca Crickmay, =phillipsastraeid? 

B. verneuili Milne-Edwards, and Haime,=Arachn.ophyllum or unrecognizable. (See Asterobillingsa 

discussion.) 

B. verrilli (Meek) ,=Radiastraea. (See Pedder, 

1964.) 

B. yandelli (Rominger) ,=Heliophyllum. 

Distribution.-Asterobillingsia is known in rocks 

of early Onesquethaw to Tioughnioga age (late 

Siegenian to Givetian) in eastern North America. 

Asterobillingsa magdisa Oliver, 1974 

Plates 57-64 

? 1842 Asterocycles confluens Vanuxem, p. 136 (not 

recognizable) . 

?not 1851 Phillipsastraea verneuili Milne-Edwards and 

Haime, p. 447, pl. 10, fig. 5 (not recognizable 

but probably an Arachnophyllum; see discussio;n). 

1859 P. verneuili Milne-Edwards and Haime. Billings, 

p. 127-128, fig. 24. 

1863 P. verneuili Milne-Edwards and Haime. Billings, 

fig. 363, p. 365. 

? 1875 P. verneuili Milne-Edwards and Haime. Nicholson, p. 78 (probably this species because of 

locality). 

not 1876 P. verneuili Milne-Edwards and Haime. Rominger, p. 128, pl. 38, fig. 2 (=A. canadensis). 

not 1876 P. verneuili Milne-Edwards and Haime. deKoninck, p. 53. (Australian specimen, see discussion). 

not 1898 P. verneuili Milne-Edwards and Haime. Whiteaves, p. 365 (probably A. canadensis as stated 

by Ehlers and Stumm, 1953, p. 3) . 

1899 P. verneuili Milne-Edwards and Haime. Lambe, 

p. 250-251. ("Corniferous fm. of Ontario" 

specimens only) . 

1901 P. verneuili Milne-Edwards and Haime. Lambe, 

p. 166-167, pl. 14, fig. 4 (but excluding specimens mentioned in text other than "Comiferous fm. of Ontario") . 

part 1906 P. verneuili Milne-Edwards and Haime. Grabau 

and Shimer, p. 206, fig. 107. 

part 1909 P. verneuili Milne-Edwards and Haime. Grabau 

and Shimer, p. 69, fig. 107. 

not 1922 P. verneuili Milne-Edwards and Haime. Benson, 

p. 150, (Australian specimen, see discussion). 

part 1944 Billingsastraea verneuili Milne-Edwards and 

Haime. Shimer and Shrock, p. 97, pl. 31, fig. 

7. 

not 1949 B. verneuili Milne-Edwards and Haime. Stumm, 

p. 35, pl. 16, fig. 6. 

part 1953 B. verneuili Milne-Edwards and Haime. Ehlers 

and Stumm, p. 2-3, pl. 1, figs. 2, 3 (reillustrated herein), (not fig. 1,=A. sp.). 

part 1955 B. verneuili Milne-Edwards and Haime. Stumm, 

card 243.1 (not fig. 1, which is A. sp., and not 

card 243, which is not recognizable but probably an Arachnophyllum). 

not 1956b B. verneuili Milne-Edwards and Haime. Hill, p. 

F280, fig. 191-7. 

not 1958 B. verneuili Milne-Edwards and Haime. Cranswick and Fritz, p. 42, pl. 2, figs. 2, 4, 7 (=A. 

sp. undet.) . 

1974 Asterobillingsa magdisa Oliver, p. 168-170, figs. 

2a-e, 3a-d. 

Occurrence of holotype.-Schoharie Grit, near 

Clarksville, Albany County, N.Y. (USGS loc. 5891-

SD). 

Diagnosis.-Asterobillingsa with small to medium-sized corallites; major septa extend to or nearly 

to the ~axis where a few commonly meet; zigzag 

carinae are strongly to weakly developed on all septa 

within the dissepimentarium. 

External features.-Astreoid colonies, small to 

medium sized, disk or lens shaped, as much as 30 

em or more in diameter and 12 em or more in thickness. Gorallites are irr,egularly prismatic, radiating 

to suhpa.r,allel in outer parts of coralla. Corallite 

diameters vary, but in a single colony size variation 

is limited (tables 28, 29) ; mean diameter (dax) in 

18 colonies ranges from 7.9 to 16.9 mm (fig. 30B). 

Increase is lateral or peripheral. Offsets apparently grow rapidly as few immature individuals are 

seen in transverse sections of coralla. 

The holotheca has been observed in only one 

corall urn. This is an extremely rugose and irregular 

basal covering having concentric growth lines and 

fine radiating septal grooves (pl. 59, fig. 2). 

Calices are crater shaped with the rim of the 

"crater" corresponding to the outer margin of the 

tabularium. Inside of the rim the calice is bowl 

shaped, shallow, with a gentle boss in some corallites. 

Outside of the rim, the calice is reflexed downward, 

becoming horizontal toward the edge. 
92 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

EXPLANATION 

X Asterobillmgsa magd/sa steorra 

0 A.m. magdisa ® Edgecliff A. m. magd1SB 

• A. rugosa • He/,ophyllum procellatum 

•_A. affir11s 

20 I I I ~X I 

A 0 • ~ X • X 18 1- -

X XX 

0 X 

® xx 'C::: 16 1- X - XX X 

0 X 

® 

OX 

14 f.- -

12 I I I I I 

2 2.5 3.0 3.5 4.0 4.5 

rJt (mm) 

5 

E 4 8 x• X • ~ X .... s Xo X XX 0 .,_ 3 X X® l't:l 

0 (p 

2 

6 8 10 12 14 16 18 

(fax (mm) 

20 T I I l .X I .! 

c ~ 

.. X X 18 1-- -

X X 

X 0 

'<:::: 16 - X ® X - X X X 0 X 

"® 

xo 

14 1-- -

12 I I I I I I 

6 8 10 12 14 16 18 

(jax (mm) 

FIGURE 30.-Asterobillingsa spp. and H eliophyllum procellatum data. A, Mean number of major septa (n) versus 

mean tabularium diameter (dl). B, Mean tabularium diameter (dl) versus mean diameter (axial distance; d7z). 

C, Mean number of major septa (n) versus mean diameter 

(axial distance; ( d-;;;) . 

Internal features.-Septa radially arranged. 

Major septa extend from margin of corallite nearly 

to the axis where two or more septa may join to 

form a simple axi~al structure. Minor s·epta are limited to the dissepimentarium. Peripherally the septa 

abut against those of the adjacent corallite or in a 

TABLE 28.-Asterobillingsa magdisa magdisa Oliver, summary 

data 

Intraeolony 

Mini- Maxi- InterMean colony mum mum 

Nl --------- 15 23 19.7 6 

n: x 14.4 19.3 16.0 

OR ---- 12-16 17-23 14.4-19.3 8 ------ .8 1.5 1.77 

v ----- 4.9 7.8 5.8 11.1 

dt: x ------ 2.6 3.8 3.1 

OR ---- 2.4- 2.8 3.4 - 4.2 2.6- 3.8 8 ------ .1 .2 .50 

v ----- 5.6 7.9 6.6 16.1 r ----------- .18 .75 .89 

p ----------- .50 <.001 <.01 

das: x ------ 8.0 14.3 11.3 

OR ---- 6.4-11.0 12.2 -17.8 7.9-14.3 

8 ------ .9 1.6 2.2 

v ----- 8.7 13.7 11.4 19.6 

1 Intracolony da:c based on samples of 20 cora.llites from each colony. 

TABLE 29.-Asterobillingsa magdisa steorra Oliver, summary 

data 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

Nl --------- 10 36 18.3 15 

n: x 14.4 19.8 16.6 

OR ---- 13-16 18-22 13.4-19.8 

8 ------ .7 1.4 1.4 

v ----- 4.4 9.3 6.3 8.4 

dt: x ------ 2.9 4.3 3.7 

OR ---- 2.6- 3.2 3.9 - 5.0 2.9- 4.3 

8 ------ .1 .4 .4 

v ----- 4.1 13.6 6.6 11.4 

r ----------- .05 .86 .62 

p ----------- .90 <.001 <.01 

dn: x 8.8 16.9 12.9 

OR ---- 6.4-11.2 14.0- 21.8 8.8-16.9 

8 ------ 1.1 2.7 2.51 

v ----- 8.9 22.0 14.2 19.4 

1 da"" data from samples of 20 corallites from each of 13 colonies. 

few places appear to be continuous with those of the 

next corallite; peripherally discontinuous septa are 

rare. All septa are attenuate and uniform in thickness in the dissepimentarium; in some colonies the 

major septa thicken slightly as they enter the tabularium. The average number of inajor septa per 

corallite ranges from 14 to 20 in the studied coralla. 

(tables 28, 29 and figure 30). 

Zigzag and scattered yard-arm carinae are long 

and closely spaced in most corallites of most coralla. 

On the sides of the septa, and in p·roperly oriented 

longitudinal sections, the carinae appear as low, 

fiangelike ridges, directed inward and upward, which 

are steepest toward the periphery. 

The dissepimentaria occupy a large part of each 

corallum. In longitudinal sections the dissepiments 

are arranged horizontally in rows that are more or 

less horizontal except near the tabularia where they 

arch sharply upward, then downward to form a 

crest at the tabularium-dissepimentarium boundary. 
SYSTEMATIC PALEONTOLOGY 93 

The boundaries between adjacent corallites are not 

recognizable in longitudinal sections. In most parts 

of the coralla the dissepiments are confined to interseptal spaces, but locally the septa may be vertically 

discontinuous and the dissepiments large. 

The tabularia are narrow; the average diameter 

in 21 coralla ranges from 2.6 to 4.3 mm. Tabulae are 

incomplete, forming a shallow W -shaped pattern 

that is gently arched axially and depressed peripherally. Tabulae are closely spaced, averaging several 

per mm. 

Chronological subspecies.-Specimens of Asterobillingsa occurring in the Schoharie-Bois Blanc and 

in the Edgecliff Member are similar in most respects 

but generally differ in the morphology of their dissepimentaria. There are local lonsdaleoid areas in 

most transverse sections of the older form. These 

commonly occur where three or four corallites meet 

(pl. 60, fig. 1, and others). In longitudinal sections 

the dissepiments are generally larger in the older 

form and may be quite large where the septa are 

peripherally incomplete. The younger form has 

smaller dissepiments that are much more uniform in 

size and tend to be depressed. 

In other morphologic features, the two forms show 

considerable variation but have approximately the 

same range of variation. Figure 30 illustrates this 

for corallite and tabularium diameters and septal 

number. In both forms the major septa may or may 

not meet at the axis but are similarly arranged in 

either form. In addition, the tabularium morphology 

is quite variable, but the same variations are found 

in both forms. 

My description and analysis of A. magdisa and its 

two subspecies is based on 35 specimens. These were 

derived as follows: 

Disseviments 

"Large" - "Small" 

Schoharie ___ .;.______________ 3 

Bois Blanc ----------------- 3 

Bois Blanc ( ?) -------------- 2 

Edgecliff ( ?) ---------------- 1 12 

Edgecliff ------------------- 1 10 

Onondaga ----------------------------------- 3 

Provenance was determined by ( 1) reliable field 

identification of stratigraphic unit by collector, (2) 

reliable citation of locality where only one stratigraphic unit is exposed, and (3) associated lithology 

for eastern New York specimens where the Schoharie and the Onondaga are easily distinguished. The 

Bois Blanc and the Onondaga of western New York 

and Ontario are less easily separated so that the 

stratigraphic assignments of 15 specimens are querried although I am reasonably sure that they are 

correct. 

All known Schoharie-Bois Blanc specimens have 

"large" dissepiments. Two additional "large" types 

are probably from the Bois Blanc but one is Edgecliff and another probably Edgecliff. All of the 

"small" dissepiment types are Edgecliff, Onondaga, 

or probably On.ondaga. In the following descriptions 

and discussions all specimens are assigned to subspecies on the basis of their morphology, although it 

would be equally reasonable to assign them by stratigraphic position on the assumption of overlapping 

ranges of variation in two successive "populations." 

The two apparent survivors of the early form in the 

Edgecliff are specially designated in figure 30, and 

in all plate illustrations. 

Comparisons.-A. magdisa differs from other species of the genus in its well developed zigzag carinae, 

its arrangement of major septa at the axis, and its 

small to medium corallite size. A. magd,isa is certainly the same as Billingsastraea verneuili of many 

authors. Specimens from the "Onondaga Limestone" 

in southwestern Ontario, described as B. verneuili 

by Billings (1859), Nicholson (1875), and Lambe 

(1899, 1901), belong to A. magdisa although internal features were not described and none can be 

assigned to a subspecies. Later descriptions of "B. 

verneuili" cited in the synonymy are discussed with 

the appropriate subspecies. 

A. rugosa (Hall) differs from A. magdisa in its 

profusion of yard-arm carinae and axial complex. A. 

canadensis also has a higher percentage of yard-arm 

carinae. 

Asterobillingsa magdisa magdisa Oliver, 1974 

Plates 57-60 

part 1953 Billingsastraea verneuili (Milne-Edwards and 

Haime) Ehlers and Stumm, p. 2-3, pl. 1, 

figs. 2, 3 (not fig. 1). 

part 1955 B. verneuili (Milne-Edwards and Haime). 

Stumm, card 243.1 incl. figs. 2, 3 (not fig. 1 

and not card 243) . 

1974 Asterobillingsa magdisa magdisa Oliver, p. 170, 

figs. 2a-e. 

Occurrence of type material.-Schoharie Grit, 

Clarksville, N.Y.; USGS locality 5891-SD; collected 

1961 by the author. 

Diagnosis.-A. magdisa in which the largest dissepiments are commonly 2.0 by 0.5 mm (measured in 

longitudinal section) or larger; medium and small 

dis,sepiments are globose. 

Discussion.-A description of this subspecies and 

a comparison with A. magdisa steorra are in the 

description-discussion of A. magdisa. The subspecies 

description is based on eight specimens from or 
94 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

probably from the Schoharie and the Bois Blanc and 

two from the Onondaga. These all have large dissepiments and, with one exception, are uncrushed. 

The exception is from the "green beds" at Port Colborne, which apparently underwent more consolidation during burial than most other stratigraphic 

units under discussion. (See "Stratigraphy.") 

The specimen illustrated by Ehlers and Stumm 

(1953) and by Stumm (1955) as Billingsastraea verneuili from Port Colborne, Ontario, belongs to this 

subspecies; it is reillustrated here (pl. 58, figs. 4-6) 

because of its very large dissepiments. The specimen 

is probably from the Bois Blanc Formation. 

Distribution.-Schoharie Grit, eastern New York, 

Bois Blanc Formation, southwestern Ontario, and 

Edgecliff Member, Williamsville, N.Y. The species is 

not common anywhere, but many additional specimens, probably of this subspecies, are preserved as 

molds of the colony surface in the Schoharie Grit. 

These have the size and septal arrangement of other 

Schoharie specimens but cannot be certainly identified. Museum labels indicate ~some of these to be from 

the lower Hudson Valley and are presumably from 

the Kanouse Sandstone (Schoharie equivalent, Boucot, 1959) in the Highland Mills area. 

Material.-Schoharie Grit, eastern New York; 

Holotype, USNM 163419; paratypes, NYSM 12828 

and AMNH 641. 

Paratypes, Bois Blanc Formation, Ontario : 

USNM 163420-21, 163423, and GSC 31156. Paratypes USNM 128016, "Port Colborne, Ontario," and 

paratype NYSM 12827, "Leroy, New York" are 

probably Bois Blanc also (on the basis of lithology 

and preservation of specimens) . 

Paratype, Edgecliff Member, bioherm facies, Williamsville, N.Y.: USNM 163422. Paratype USNM 

113264, "Port Colborne, Ontario" is probably Edgecliff (based on lithology and preservation). 

Asterobillingsa magciisa steorra Oliver, 1974 

Plates 61-64 

1974 Asterobillingsa magdisa steo?"ra Oliver, p. 170, figs. 

3a-d. 

Occurrence of type material.-"Corniferous Limestone, Clarksville, Albany Co., New York." Onondaga Limestone, probably Edgecliff Member, bioherm facies. 

Diagnosis.-A. magdisa in which the largest dissepiments are relatively s.mall (maximum size in 

longitudinal section 2.0 by 0.5 em), commonly no 

larger than 1.0 by 0.4 mm; dissepiments tend to be 

depressed. 

Discussion.-A description of the morphology of 

this subspecies and a comparison with A. magdisa 

magdisa. is included in the description-discussion of 

A. magdisa. The subspecies description is based on 

25 coralla or parts of coraUa from or probably from 

the Edgecliff Member. These commonly have small 

dissepiments in thin layers that give a la.minated appearance to broken specimens; often coralla will 

split along these layers. A large percentage of specimens are partly crushed ; this is noted for some other 

bioher·m species also and may indicate either a more 

rigorous environment or gr~eater post-burial compaction than occurred in other stratigraphic units. A. 

magdisa steorra with larger dissepiments are less 

commonly crushed and A. magdisa magdisa is only 

rarely so. Possibly the forms with larger dissepiments were better engineered and more resistant to 

compression. 

DistribuUon.-Edg,eclitf Member, bioherm facies, 

eastern and western New York and Niagara Peninsula of Ontario. One specimen known from Woodstock, Ontario, is probably from the "Amherstburg" 

Limestone. Two specimens from the "Onondaga" in 

eastern New York are probably from the Edgecliff 

and possibly bioherm facies also. Eighteen of the 25 

specimens studied are fro·m the old Fogelsanger 

quarry, Williamsville, N.Y. (loc. 108), but only two 

specimens wer'e collected by me at this locality. 

Others rep~resent museum accumulations from an 

often collected locality. Apparently the species is not 

eommon. 

M aterial.-Holotype, NYSM 12824, "Corniferous 

Limestone, ClarksviHe, Albany Co., New York," 

prohably the Edgecliff Member bioherm facies. Illustrated para types, USNM 172166-68; NYSM 12826; 

and GSC 31155. Additional measured paratypes, 

USNM 26000a, 113263, 172169-76; NYSM 12825; 

other paratyp,es USNM 143121 and 172177 (five 

coralla). 

1843 

not 1844 

not 1847 

not 1850 

not 1851 

not 1876 

not 1887 

1953 

Asterobillingsa rugosa (Hall) 

Plate 65 

Astraea rugosa Hall, p. 159, pl. 32, fig. 2. 

A. rugosa Hall. Owen, p. 33 [ 407], pl. 7, fig. 

6 (possibly an Upper Devonian phillipsastraeid). 

A. rugosa Hall. Yandell and Shumard, p. 8. 

Favastraea rugosa (Hall). d'Orbigny, p. 107. 

Cyathophyllum rugosum (Hall). Milne-Edwards 

and Haime, p. 387, pl. 12, figs. 1, la-b (see 

Prismatophyllum prisma). 

C. rugosum (Hall). Rominger, p. 106, pl. 37, figs. 

1, 2 (Prismatophyllum spp.) 

C. rugosum (Hall). Davis, pl. 90; pl. 93, fig. 3 

(=Prismatophyllum spp.). 

Billingsastraea rugosa (Hall). Ehlers and 

Stumm, p. 5, pl. 2, fig. 6. 
SYSTEMATIC PALEONTOLOGY 95 

1955 B. rugosa (Hall). Stumm, cards 241, 241.1. 

part 1958 B. verneuili (Milne-Edwards and Haime). Cranswick and Fritz, p. 42-43 (not the illustrated 

specimen). 

Occurrence of holotype.-Leroy, N.Y. Probably 

the Bois Blanc Formation. (Statement based on 

lithology of specimen.) 

Diagnosis.-Asterobillingsa with medium-sized 

corallites; m·aj or septa extend to the axis where they 

irregularly meet; long subyardarm and zigzag carinae are abundant within the di:ssepimentarium. 

External features.-The holotype and only known 

New York specimen, is a partly crushed fragment of 

an astreoid or thamnasterioid colony 11 by 13 em in 

length and width and 3.5 em in height. The surface 

of the specimen is weathered so as to distort the 

original appear~ance of the colony. Sections indicate 

that calice pits were relatively deep (depth equaling 

tabulari urn diameter) having a high surrounding 

ridge and a sharply reflexed platform flattening toward the periphery. Coralite diameters ( daa;) range 

from 10 to 17 mm, average 14.3 mm (table 30, fig. 

30). 

Internal features.-Septa are radial or subbilateral. Major s·epta extend from the periphery, where 

they are commonly confluent with those of adjacent 

corallite·s (thamnasterioid), to the axis where they 

meet other septa in pairs or groups, forming irregular or distinctly bilateral patterns. Minor septa are 

limited to the dissepimentarium. All septa are 

strongly carinate within the dissepimentarium, having both zigzag and subyard-arm carinae, neither 

predominating. Some major septa thicken slightly in 

the tabularium but most septa are very attenuate 

throughout their length. 

The number of major septa in the holotype coralTABLE 30.-Asterobillingsa rugosa and A. affinis intracolony 

data 

A. rugosa A. affinis 

NYSM ROM GSC 

299 27128 3270 

Nl --------- 20 7 16 n: x 18.3 17.4 19.4 

OR ---- 17-21 15-19 18-21 8 ------ 1.1 1.4 1.1 

v ----- 6.2 8.0 5.6 

dt: x 4.1 4.5 4.3 

OR ---- 3.5- 4.5 4.0- 4.4 3.7- 4.8 

8 ------ .3 .28 .3 

v ----- 6.5 6.2 6.9 r ----------- .56 --------- .32 

p ----------- <.02 --------- .30 

dax; x ----- 14.3 14.1 12.8 

OR ---- 10.0-17.0 10.8-18.0 9.6-15.6 

s ------ 2.1 2.4 1.5 

v ----- 14.5 17.3 12.0 

1 Sample size for d"x is 20, 14, and 20 respectively in the three 

specimens. 

lum ranges from 17 to 21 in 20 corallites, average 

18.3 (table 30). 

Tabularia are narrow, occupying only slightly 

more than one-fourth the corallite diameter (dt= 

3.5-4.5 mm, average 4.1 mm). Tabulae are generally 

incomplete, nearly flat, and very closely spaced. DisS'epimentaria are correspondingly broad, composed 

of relatively small, globose dissepiments in numerous rows. 

Abitibi River Limestone specimen.-The only 

known specimen of A. rugosa other than Hall's holotype is a fragment from the Hudson Bay Lowlands 

that Cranswick and Fritz (1958) identified as Billingsastraea 1Jerneuili. This specimen is remarkably 

similar to the holotype in morphology and is les.s 

crushed, so that features of the longitudinal section 

and the contacts between corallites in transverse 

section are more easily observed. Numbers of septa 

and tabuJ.arium and corallite diamete·rs are similar to 

the holotype (table 30) . 

Discussion.-The holotype is crushed so that the 

peripheral parts of corallites are broken or distorted. The Abitibi River specimen is better preserved and shows some features more clearly than 

the holotype. 

The holotype is from Leroy, N.Y., where both Bois 

Blanc and Onondaga beds are exposed. One corner of 

the specimen is silicified; the appearance of this 

silicified area in thin section and the general color 

and preservation of the specimen as a whole is suggestive of the Bois Blanc Formation. The Abitibi 

River Limestone fauna described by Cranswick and 

Fritz (1958) c-ertainly includes Bois Blanc forms but 

may include Onondaga forms as well. The combined 

evidence avaHable suggests that both specimens are 

more likely of Bois Blanc than Onondaga age. 

Figure 30 compares A. rugosa with A. magdisa 

and H eliophyllum procellatum in size characters 

and number of septa. A. rugosa is characterized by 

the axial arrangement of major septa and by the 

presence of both yard-arm and zigzag carinae in approximately equal numbers. A. magdisa has a simpler axial arrangement of septa, commonly with only 

a few s·epta in contact; in addition, this species has 

predominantly zigzag carinae. Yard-arm carinae 

predominate in H. procellata and the axial complex 

is very involved. 

Milne-Edwards and Haime (1851) mistakenly described as Cyathophyllum rugosum (Hall) a species 

of Prismatophyllum from the Jeffersonville Limestone at the Falls of the Ohio. This was recognized 

by Simpson (1900, p. 218-220) who used the MilneEdwards and Haime (not Hall) species as the basis 
96 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

for the genus Prismatophyllum; Lang and Smith 

(1935a, p. 558-559) renamed the species P. prisma. 

Citations of Astraea or Cyathophyllum rugosum(a) 

prior to 1900 are based on specimens of either Asterobillingsa or Prismatophyllum having corallites 

in the 8 to 15 mm diameter range. In many instances, 

desC-riptions and (or) illustrations are poor and 

specimens lost or unknown. 

Material.-Holotype, NYSM 299. Specimen from 

Abitibi River Limestone, ROM 27128. 

Distribution.-Known only from Leroy, N.Y., and 

Matagami River, Ontario. Both occurrences are 

probably of Bois Blanc (Em sian) age. 

Asterobillingsa affinis (Billings) 

1870 Phillipsastraea a/finis Billings, p. 11 

1964 Billingsastraea a/finis (Billings) . Oliver, p. 3- 4, pl. 

1, figs. 1-5, pl. 2, figs. 1-4 (see for synonymy, description, and illustrations). 

Occurrence of holotype.-"Indian Cove, Gaspe, in 

the Gaspe limestone No. 8" (Billings, 1874, p. 11). 

Gaspe Li.mestone Group, Grande Greve Formation, 

Indian Cove Member of Cumming (1959, p. 4, 24). 

Lower Devonian, "Esopus Stage," Etymothyris Zone 

of Boucot (1959, p. 737-739). 

Diagnosis.-Asterobillingsa with medium-sized 

corallites; calicul·ar pits a.re deep and steep sided 

with a low, arched floor; the platform curves gently 

from the pit becoming horizontal only near the periphery (not reflexed). Major septa extend nearly to 

axi·s; zigzag carinae short and abundant. Dissepiments m·edium siz·ed and globose; tabulae incomplete, 

arched, or sagging but forming a convex pattern. 

Discussion.-The holotype and only known specimen of A. a/finis was described by Oliver (1964). 

This paper should be seen for a full synonymy and 

description with illustrations. The species is included 

here to complete the coverage of described Onesquethaw Asterobnlingsa and because of its position as 

the earliest known craspedophyllid. 

A. a/finis differs from other Asterobillingsa in its 

"normal" calice shape (lacking the reflexed, outwardly sloping platform) and in having very short 

zigzag carinae that are, however, distinct and abundant. (See also table 30 and fig. 30). 

Occurrence.-There is general agreement that the 

Indian Cove Member, Grande Greve Formation, is 

eithe'r Deerpark or early Onesquethaw in age, but 

recent opinion (see Oliver, 1964, p. 1; Boucot, 1968, 

p. 90) plaees it in the early Onesquethaw. In either 

case, A. a/finis is of Siegenian ~age, whereas other 

known Asterobillingsa are Emsian or younger in 

age. 

Mat erial.-Holotype and only known specimen, 

GSC 3270. 

Subfamily CRASPEDOPHYLLINAE Dybowski, 1873 

Type genus.-Eridophyllum Milne-Edwards and 

Haime, 1859 ( = Craspedophyllum Dybowski, 1873). 

Diagnosis.-Solitary ( ?) or compound craspedophyllids with general cylindrophyllinid characters 

except that major septa are deflected in the tabularium to form an aulos, separating the tabularium into 

axial and periaxial zones. 8epta are thin and strongly carinate, and may or may not ente.r the axial zone. 

Included genera.-Grewgiphyllum Oliver and Eridophyllum Milne-Edwards and Haime. 

Discussion.-The craspedophyllinids are morphologically close to the cylindrophyllinids and were 

probably derived from early members of that group 

within eastern North America. At this time, I recognize only two genera. Eridophyllum ( = Craspedophyllum), with a nearly perfect aulos, is limited to 

the Middle Devonian of easte~rn North America and 

Morocco. Early species are all phaceloid, but later 

ones are solitary ( ?) and cerioid as well. Grewgiphyllum ranges from the Emsian into the lower Middle 

Devonian and is restricted to Eastern North Amerioa. It is characterized by an imperfect aulos and an 

alternately cerioid and phaceloid growth form. In 

these characters it is intermediate between Eridophyllum and the cylindrophyllinid Prismatophyllum 

(middle-upper Onesquethaw) •and it seems likely 

that the morphologic sedes Prismatophyllum ~ 

Grewgiphyllum ~ Eridophyllum represents an evolutionary s1eries as well. 

Subfamily distribution.-Middle Onesquethaw 

through Taghanic (Emsian-Givetian) in eastern 

North America. One species of Eridophyllum is 

known from the Givetian of Morocco. 

Genus ERIDOPHYLLUM Milne-Edwards and Haime 

1850 Eridophyllum Milne-Edwards and Haime, p. 

lxxi. 

1851 Eridophyllum M.-E. and H. M.-E. and H., p. 

423-424. 

1857 Eridophyllum M.-E. and H. Pictet, p. 459. 

part 1859 Eridophyllum M.-E. and H. Billings, p. 130-131. 

1860 Eridophyllum M.-E. and H. Milne-Edwards, p. 

414-415. 

1873b Eridophyllum M.-E. and H. Dybowski, p. 337. 

1873b Craspedophyllum Dybowski, p. 339. 

1873a Craspedophyllum Dybowski, Dybowski, p. 160. 

part 1874 Eridophyllum M.-E. and H. Nicholson, p. 34. 

1877 Crepidophyllum Nicholson and Thompson, p. 

149-150. 

1878 Crepidophyllum Nicholson and Thompson, Nicholson, p. 51-54. 

part 1883 Eridophyllum M.-E. and H. Roemer, p. 356. 

not 1889 Craspedophyllum Dybowski. Miller, p. 180. 
SYSTEMATIC PALEONTOLOGY 97 

1889 Crepidophyllum Nicholson and Thompson. Miller, 

p. 180. 

1890 Craspedophyllum Dybowski. Sherzer, p. 59-60. 

1891 Craspedophyllum Dybowski. Sherzer, p. 290. 

1898 Craspedophyllum Dybowski. Grabau, p. 129. 

part 1901 Crepidophyllum Nicholson and Thompson. Lambe, 

p. 154. 

part 1906 Eridophyllum M.-E. and H. Grabau and Shimer, 

p. 208. 

1906 Craspedophyllum Dybowski. Grabau and Shimer, 

p. 212. 

1907 Craspedophyllum, Anderson, p. 59-66. 

1907 Eridophyllum, Anderson, p. 65-66. 

part 1909 Eridophyllum M.-E. and H. Grabau and Shimer, 

p. 71. 

1909 Craspedophyllum Dybowski. Grabau and Shimer, 

p. 75. 

not 1916 Eridophyllum, Ruffner, p. 318-319. 

1927 Eridophyllum M.-E. and H. Smith and Lang, p. 

307-308. 

1933 Eridophyllum M.-E. and H. Smith, p. 518-519. 

1935a ETidophyllum M.-E. and H. Lang and Smith, p. 

547-548. 

1938 Eridophyllum M.-E. and H. Stewart, p. 40. 

1938 Schistotoecholasma Stewart, p. 45. 

not 1940 Cmspedophyllum Dybowski. Lang, Smith, and 

Thomas, p. 41. 

1940 Crepidophyllum Nicholson and Thompson. Lang, 

Smith, and Thomas, p. 42. 

1940 Eridophyllum M.-E. and H. Lang, Smith, and 

Thomas, p. 58. 

1940 Schistotoechelasma Stewart. Lang, Smith, and 

Thomas, p. 117. 

not 1941 Eridophyllum, Hill, p. 270-271. 

not 1942b Eridophyllum, Hill, p. 186. 

1944 Eridophyllum M.-E. and H. Shimer and Shrock, 

p. 89. 

1947 Eridophyllum M.-E. and H. LeMaitre, p. 36. 

1949 ETidophyllum M.-E. and H. Stumm, p. 37-38. 

part 1950 ETidophyllum M.-E. and H. Wang, p. 223. 

1952 Eridophyllum M.-E. and H. Lecompte, p. 475. 

1954 Eridophyllum M.-E. and H. Stumm, p. 3. 

1955 Eridophyllum M.-E. and H. Stumm, card 306. 

l956b Eridophyllum M.-E. and H. Hill, p. F282. 

not 1957 Eridophyllum, Ivania (not seen, but see discussion). 

not 1960 Eridophyllum, Zheltonogova and Ivania, p. 379-

380. 

not 1962 Eridophyllum, Soshkina and Dobrolubova, p. 336. 

Type species.-By original designation, E. seriale 

Milne-Edwards and Haime, 1850, p. lxxi; erroneously renamed E. verneuilanum by Milne-Edwards and 

Haime, 1851, page 424, plate 8, figures 6, 6a. Devonian, Columbus, Ohio. Generally ·assumed to be from 

the Middle Devonian Columbus Limestone. (See following discussion.) 

Diagnosis.-Solitary or compound corals with general cylindrophyllinid characters except that the 

major septa are deflected to form a complete or nearly complete aulos separating the tabularium into 

periaxial and axial zones. Septa are carinate and 

ordinarily do not enter the axial zone. 

Description.-Solitary, phaceloid, and cerioid species have been described. "Solitary" forms often 

fo·rm s.mall colonies having from one to a few offsets. Most species al"e phaceloid and composed of 

cylindrical corallites that are or are not connected by 

lateral projedions of the corallite wall. Increase is 

lateral and nonparricidal. 

Growth lines are fine and (or) coarse and some 

species are quite rugose. Longitudinal septal grooves 

are inconspicuous. 

Calices are deep with steep walls, but the aulos 

forms an axial boss in some species. 

Major and minor septa are lightly to heavily carinate in the dissepim·entari urn, carinae are commonly 

zigz-ag but subyard-arm carinae are prominent in 

some spec-ies. Major septa do not extend to axis but 

are deflected to form an inner wall, the aulos; minor 

septa do not extend as far as the aulos. All septa are 

radially arranged except that the aulos may create a 

secondary bilateral symmetry. 

The aulos is commonly open in immature individuals, hut may be open or closed in ephebic 

individuals. The opening, when present, is most commonly (possibly always) at the position of the 

cardinal septum. Rarely, two openings are present, 

presumably cardinal and counter. 

The dissepimentarium is composed of a few to 

several rows of normal, globos.e dissepiments. The 

tabularium is divided into two zones by the auloo. 

Tabul·ae within the aulos al"e commonly complete and 

horizontal but may be incomplete and inclined. Tabulae outside of the aulos are complete or incomplete 

and ho·rizontal or inclined. 

Increase.-lncrease is lateral in all known phaceloid species; increase analagous to peripheral increase is known in one cerioid species. (See Stumm, 

1954, pl. 2, fig. 2). Lateral offsetting in phaceloid 

species is similar to that in phaceloid cylindrophyllinids. Commonly the offset aulos is open in the 

direction of the parent, although this opening does 

not necess·arily correspond to the position of the 

ephebic opening in the offset individual. The opening 

of the parental aulos (if present) may be unrelated 

to the po~ition of the offset; a secondary opening 

may form tempo·rarily at the offset position. 

Im.mature corallites tend to have relatively large 

auloi, in some s.pecimens actually larger than the 

adult aulos. Immature auloi are invariably ( ?) open 

and commonly the positions of septal insertion can 

be identified. In all examples studied, the aulos opening is in the cardinal position. 
98 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

In the earliest stages of offset development (pl. 

72, figs. 3, 4) no aulos is present and atavo- and neotissue are formed as in the cylindrophyllinids. 

Microstructure.-Craspedophyllid type. See general description. 

Discussion.-Eridophyllum was described by Milne-Edwards and Haime (1850) who named E. seriale 

·as type species. The s~am.e authors later described E. 

verneuilanum as type but this is clearly the same 

species as E. seriale and is a junior objective synonym in addition to be1ing an erroneous type designation. The holotype of E. seriale is from the "Devonian, Columbus, Ohio." This can only be the Columbus 

Limestone, but the Columbus contains at least four 

species of the genus at different stratigraphic levels. 

The growth habit illustrated by Milne-Edwards and 

Haime (1851, pl. 8, fig. 6) is characteristic of forms 

limited to the upper Columbus (Paraspirifer Zone) 

and it is assumed for present purposes that these 

forms compose E. seriale. The holotype specimen is 

thought to be in the de Verneuil collection, Ecole des 

Mines, Paris, but was not found during a recent 

visit and search (1969). 

As thus understood, E. seriale s.s. is rare in New 

York, except in the widespread AcinophyllumEridophyllum biostrome in the Mooreshouse 

Member. 

Axinura Castelnau (1843, p. 49) has been considered by some workers as a possible s·enior synonym of Eridophyllum (for example, Lang, Smith, 

and Thomas, 1940, p. 25; Hill, 1956, p. F321). Axinura is probably unrecognizable but, in any case, 

should not be ~allowed to replace the well known and 

reasonably well based Eridophyllum. 

Craspedophyllum Dybowski (1873a, b) based on 

'~C. americanum, Columbus, Ohio," is a junior synonym of Eridophyllum. Dybowski's description of the 

aulos is clear and leaves no doubt as to the generic 

affinities of his material. Specific identification of C. 

americanum is uncertain but the species is probably 

based on a speeimen of E. seriale. Craspedophyllidae 

Dybowski wa,s ·established for corals in which the 

"side faces of septa were supplied with lateral outgrowths" (carinae) or "thorny outgrowths" (translated in Sherzer, 1890, p. 60). The genus was based 

on the presence of ·an inner wall (aulos). 

Milne-Edwards and Haim·e (1850) mentioned the 

"inner mural investment" but emphasized the lateral 

connections in their first description of the genus. In 

1851 they gave more attention to the inner wall and 

clearly illustrated it but still emphasized the profusion of lateral supports. Many subsequent workers 

used the name Eridophyllum for a variety of forms 

with lateral supports; others rejected the name because late·ral supports were not an adequate basis for 

a genus. 

Under this misunderstanding of Eridophyllum, it 

is not surprising that Crepidophyllum Nicholson and 

Thompson (1877) was erected to include forms with 

an inner wall. (But as Sherzer (1890) pointed out, 

these authors misunderstood Dybowski.) The type 

species of Crepidophyllum was selected by Miller 

( 1889) and happens to be a species in which lateral 

supports are rare. This character was later used by 

some workers to separate Crepidophyllum from Eridophyllum even after it was recognized that the two 

nominal species were both characterized by the 

aulos. It is now generally accepted that the presence 

or absence of lateral supports is not of generic importance and that the two genera a.re synonym.s. 

Schistotoecholasma Stewart (1938) was separated 

from Eridophyllum because of the pres.ence of an 

open aulos rather than a clos.ed one, but this is a 

character that v·aries within many colonies as has 

been recognized by many workers. S. obliquum and 

S. typicale (both Stewart, 1938) are certainly conspecific and are probably the same as E. seriale the 

type spedes of Eridophyllum. 

The following Onesquethaw taxa are described in 

the following sections: 

1. E. subseriale Stumm; Edgecliff Member, New 

York. 

2. E. seriale Milne-Edwards and Haime; Moorehouse Member, New York. 

3. E. seriale population B; Acinophyllum-Eridophyllum bed, Moorehouse Member, western 

New York. 

4. E. aulodokum n. sp.; middle(?) Moorehouse 

Membe·r, N·ew York. 

5. E. conjunctum (Davis); Moorehouse Member 

just above A.-E. bed, western New York. 

6. E. corniculurn n. sp.; Moorehouse Me,mber above 

Acinophyllum-Eridophyllum bed, western New 

York. 

Additional Eridophyllum are known from the 

Onesquethaw-age Columbus and Jeffersonville Limestones and from various formations of Gazenovi·a 

through Taghanic age. The youngest Eridophyllum 

known to me is from the Tully Limestone in central 

N.ew York. 

Typical Eridophyllum was described from the 

Middle Devonian of Morocco by Le Maitre (1947), 

although I disagree with her assignment to E. 

seTiale. 

Other reports of Eridophyllum outside of Eastern 

North America are unacceptable to me. 
SYSTEMATIC PALEONTOLOGY 99 

From Turkey, Hiiffner (1916, p. 318-319) briefly 

described "Eridophyllum spec. (nov. spec.?)" but 

did not illustrate it. I know of no further reference 

to this occurrence and it must be discounted. 

"Eridophyllum bartrumi" Allen (1935, p. 4-6; 

Hill, 1941, p. 271-272; 1942a, p. 158) from the Middle Devonian of New Zealand and New South Wales 

has since been referred to Tipheophyllum Hill (Hill, 

1956a, p. 9). The species lacks an aulos and does not 

appear to have cvaspedo.phyllid s.eptal structure. 

"E. immersum" Hill (1942b, p. 186-187) from the 

Devonian of New South Wales has dilated septa. An 

aulos is present in some individuals but appears to 

be formed by dilation of the axial ends of septa rather than by deflection of attenuate septa as in Eridophyllum s.s. 

"E. asiaticum" Ivania (1957, p. 70-71, pl. 5, fig. 4, 

not seen; Zheltonogova and Ivania, 1960, p. 380; 

Soshkina and Dobrolubova, 1962, p. 336) from 

Eifelian deposits in Salair, USSR, has an irregula.r 

aulos or pseudoaulos which is tabular rather than 

septal in origin; the well defined innner and outer 

zones of tabulae are lacking. 

"E. crebrum" Cherepnina, 1967 (p. 172, pl. 4, figs. 

3, 4), from the Eifelian of the Gorny Altai, lacks any 

aulos. A :sharp boundary between the dissepimentarium and tabularium gives the appearance of an inner 

wall in some transverse sections. 

Distribution.-Eridophyllum is known only from 

Middle Devonian deposits in eastern North America 

and Morocco. Reported forms from Asia and Oceania 

differ significantly from the genus as clearly defined 

by numerous North Am·erioon species. In Eastern 

North America, the range is from upper Onesquethaw through Taghanic. 

Eridopbyllum seriale Milne-Edwards and Haime 

Plates 67-69 

1850 Eridophyllum se'riale Milne-Edwards and Haime, 

p. lxxi. 

1851 E. verneuilanum M.-E. and H., p. 424, pl. 8, 

figs. 6-6a. 

? 1873a Craspedophyllum americanum Dybowski, p. 155-

159, pl. 1, figs. 1-6. 

1933 E. seriale M.-E. and H. Smith, p. 519-520, pl. 1, 

fig. 12; ? figs. 13, 14. 

1938 E. seriale M.-E. and H. Stewart, p. 41-42, pl. 8, 

figs. 1-4. 

1938 Schistotoecholasma obliquus Stewart, p. 46, pl. 

9, figs. 4-6. 

1938 S. typicalis Stewart, p. 46-47, pl. 9, figs. 7-10. 

not 1947 E. seriale M.-E. and H. LeMaitre, p. 36, pl. 6, 

figs. 1-5. 

1954 E. seriale M.-E. and H. Stumm, p. 3-4, pl. 1, 

figs. 1-5. 

1956b E. seriale M.-E. and H. Hill, figs. 192-1a, b. 

not 1961 E. se'riale, Fagerstrom, p. 15 (see E. subseriale). 

part 1965 E. seriate M.-E. and H. Stumm, p. 46, pl. 41, 

figs. 2, 3, ?fig. 1. 

Occurrence of type material.-According to MilneEd wards and Haime, "Devonian, Columbus, Ohio." 

This is certainly the Columbus Limestone, Middle 

Devonian. As indicated in the following discussion, 

the type specimen was probably derived from the 

upper part of the formation, Zone H, the Paraspirifer Zone. 

Diagnosis.-Phaceloid Eridophyllum with medium-sized corallite.s (as much as 15 mm in diameter) 

that are irregularly cylindrical with very numerous 

lateral supports. The aulos is s.mall to medium sized, 

occupying as much as one-third of the corallite diam·eter; either open or closed. 

Discussion.-Eridophyllum seriale was originally 

described from the "Devonian, Columbus, Ohio." 

This is :assumed to be the Columbus Limestone. at 

Columbus since Eridophyllum is common in the Columbus and not known from other formations in the 

area. Both descriptions of Milne-Edwards and 

Haime ( 1850, 1851) are very brief, and the descriptions of most characters are' so generalized as to be 

m·eaningless except on the generic level. The growth 

form, however, is both desc·ribed and illustratedparallel corallites with numerous lateral connections , commonly lined up on the same side of each corallite. 

Further, the aulos is illustrated as being small and 

complete. 

Preliminary analysis of my own and USNM collections of Eridophyllum from the Columbus Limestone indicates the presence of as m:any as six morphologic types. Two types, both from the upper part 

of the formation (Zone H, the Paraspirifer Zone), 

have the growth form of the type specimen and are 

noted below as E. seriale and E. seriale population 

A. The form·er is widespread geographically and is 

morphologically v:ariable. The latter shows little 

variation and is known only from one bed at one 

locality. A third type is described as E. se1·iale population B; this also shows little variation and is restricted to the Acinophyllum-Eridophyllum biostrome in the Moorehouse Member, western New 

York. 

The synonymy given above is incomplete but includes the important contributions to the present 

concept of E. seriale. 

"E. seriale" w·as described from the lower part of 

the Detroit River Group by Fagerstrom (1961, p. 

15) on the basis of a fragment of one corallite. The 

fragment has a very large aulos ( 6 mm diameter) 

but is decorticated so that corallite diameter is un-
100 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

known (15 mm plus). The specimen is probably not 

identifiable but is clos.er to E. subseriale than to E. 

seriale. 

LeMaitre (1947) described a specimen of Eridophyllum from the Givetian (Hollard, 1962) of Morocco as E. seriale. The specimen is the only Eridophyllum that has been described from outside of 

eastern North America. It differs from E. seriale in 

its very small diameter ( d = 5.2 mm; n = 21.5; N = 6) 

and in its narrow dissepimentarium with only two or 

three rows of dissepiments; it probably represents a 

new species. 

The coralla that I interpret as belonging to typical 

E. seriale are found more or less randomly distributed through Zone H of the Columbus Limestone 

from Columbus to Marbl1ehead. These have the 

growth form desc·ribed and illustrated by MilneEdwards and Haime but are extre·mely variable in 

internal morphology. The aulos diameter rang·es 

from approximately 0.15 to 0.35 of the corallite diameter and may be open or closed. Aulos diameter is 

less variable within colonies than within the whole 

species, but all colonies studied have both open and 

closed auloi, although the ratio of open to closed 

varies widely. Stewart's (1938) E. seriale, Schistotoecholasma obliquus, and S. typicalis all seem to 

represent fragments of specimens of this species 

with different characters predominating. 

A 6-inch (15.4-cm) coral bed at Venice, Ohio, is 

composed of Eridophyllum colonies that are very 

consistent in morphology. These colonies fall within 

the range of variation of E. seriale and are interpreted as representing a loc,al population (population A). Ohio specimens of E. seriale, including population A, are not described here. 

A similar situation exists in New York except that 

only two specimens of typical E. seriale are known, 

both probably fro·m the Moorehouse Member. However, the Acinophyllum-Eridophyllum bed, in the 

upper Moorehouse, contains only A. segregatum and 

another relatively unvarying group of Eridophyllum, 

here interpreted and described as a second local population of E. seriale (population B). The New York 

material is described in the following paragraphs. 

I interpret E. seriale as representing a generalized 

stock persisting through much of Moorehouse time 

under somewhat varying conditions. Individuals are 

rare in New York where conditions may have been 

sufficiently different to severely limit their distribution and developm·ent. E. seriale populations A and B 

represent conse·rvative, localized developments under 

uniform conditions, probably with considerable inbreeding; as a result very little morphologic variation is present. The beds that they formed may have 

covered rather large areas, but they existed for only 

short periods of time. 

Eridophyllum seriale population A 

Diagnosis.-E. seriale with ·small aulos occupying 

0.15 to 0.25 the corallite dia.meter; open or closed, 

more commonly closed but both types occurring in all 

studied coralla. Tabul,ae within the aulos widely 

spaced, 6 to 14 per 5 mm. 

Discussion.-This form differs from typical E. 

seriale in the smaller aulos and more widely spaced 

tabulae. It differs from E. seriale population B in 

having both open and closed auloi. 

At present, this form is known only from a 6-inch 

(15-cm) bed 2 feet (0.6 m) below the top of the 

Columbus Limestone, at Venice, Ohio. The sample 

studied shows relatively little variation and is tentatively treated as representing a local population 

temporarily isolated in space. The bed is one corallum in thickness and certainly represents a very 

brief period of time. 

New York material 

Specimen 1 

Plate 67, figures 1-4 

External features.-The single specimen consists 

of 10 or more corallites not in their original relative 

positions but otherwise well preserved. The corallum 

was phaceloid, composed of relatively straight corallites. Corallite diameters range from 10 to 19 mm, 

average 14.2 mm (table 31). No lateral supports 

have been observed, but corallites are in frequent 

contact. This contact suggests that they were closely 

spaced. Increase, calice, and external markings not 

known. 

Internal features.-Major septa radially arranged 

but deflected to form an aulos; minor septa extend 

from 0.5 to 0.9 the distance to the aulos. Number of 

major septa in cylindrical parts of corallites ranges 

from 28 to 31, average 29.6, in 7 corallites (table 31 

and fig. 31). 

Septa are thin. Long zigzag carinae are common 

in all corallites studied; subyard-arm carinae are 

present in most individuals. 

The aulos diameter to total diameter ratio ranges 

from 0.25 to 0.45, ·average 0.36 (table 31 and fig. 31). 

The aulos is circular in transverse section and is 

open and closed in different corallites. Where open, 

the cardinal septum and adjacent minor septa are 

free at their axial ends or are deflected to join the 

aulos wall. Where closed, the cardinal septum seems 

to split and be deflected in both directions. 

The dissepimentarium occupies approximately 0.4 
SYSTEMATIC PALEONTOLOGY 101 

to 0.5 the corallite radius and is composed of from 

four to eight rows of globose dissepiments. 

A 

It:; 26 

I ~x---" x, 

I X I 

I X I 

~ X I \ x} \x ,,/ Population B 

24 

. 'V_$../' 

22~~------~------~------~----~--~ 

6 8 

6 8 

10 

d(mm) 

10 

d(mm) 

12 14 

12 14 

FIGURE 31.-Eridophyllum seriale data. A, Mean number of 

major septa (n) versus mean diameter (d). B, Mean aulos 

diameter ( da;,) versus mean diameter (d). Straight lines 

are selected duu I d ratios. 

The tabularium is divided into two parts by the 

aulos wall. The inner tabulae are complete or incomplete, flat or convex, from 4 to 13 per 5 mm. Incomplete tabulae may be short, arched plates. The 

outer tabulae are commonly complete, flat or concave, horizontal or inclined, from 4 to 12 per 5 mm. 

Microstructure.-As for genus and family. 

Discussion.--This specimen differs from other 

Onondaga Eridophyllum in the size and detailed 

morphology of the aulos, in the presence of long 

carinae, and in the large corallite size. It is comparable to several specimens of Eridophyllum seriale 

from the Columbus Limestone in Ohio, including the 

specimen described and illustrated by Stewart 

( 1938) as E. seriale. 

Mat erial and distribution.-Illustrated specimen, 

USNM 163424, Moorehouse Member of Onondaga. 

Limestone, locality 497, western New York. 

Specimen 2 

Plate 67, figure 5-7 

External features.-A fragment is 13 em high and 

9 by 6 em in diameter; it contains 10 or more cylindrical corallites. Spacing between corallites is less 

than one diameter, commonly they are in contact for 

part of their length. Mature corallites 11 to 15 mm in 

diameter; average diameter 13.2 mm (table 31 and 

fig. 31). 

Increase is lateral and frequent in the small fragment available. Lateral supports are present. 

Corallites are coarsely rugose; fine growth lines 

and septal grooves are also present. 

Calice, known from thin sections, is moderately 

deep (two-thirds corallite diam-eter) with steep sides 

and a very narrow peripheral platform. A floor, corresponding to the outer tabularium, is flat; a cylinTABLE 31.-Eridophyllum seriale Milne-Edwards and Haime, summary data 

Population B Specimen 1 Specimen 2 

Intracolony Intercolony lntracolony Intracolony Minimum Maximum Mean 

N --------------------- 4 14 9.0 10 7 6 

n: x --------------- 22.4 25.2 24.0 29.6 26.0 

OR ------------- 20-24 24-26 22.4-25.5 28-31 25-27 

8 --------------- .76 1.58 1.01 1.27 .63 

v -------------- 3.1 6.4 4.8 4.2 4.3 2.4 

d: x -------------- 8.4 10.4 9.5 14.2 13.2 

d"u: x -------------- 7.2 - 9.7 8.4-12.3 8.4-10.4 10.0-18.8 11.1 -14.8 

OR ------------- .63 1.56 .72 2.70 1.61 

s --------------- 6.6 15.7 11.4 7.5 19.1 12.2 

v -------------- 1.8 2.7 2.3 5.1 2.6 

OR ------------- 1.4 - 2.6 1.8- 3.1 1.8- 2.7 4.6- 5.8 1.8 - 2.9 

s --------------- .40 .72 .35 .39 .40 

v -------------- 15.8 30.6 21.1 15.0 7.7 15.6 

r/d: x --------------- 2.3 2.8 2.5 2.1 2.0 

OR ------------- --------- --------- 2.3- 2.8 --------- ---------- dau/d: x --------------- .17 .30 .24 .36 .20 

OR ------------- --------- --------- .17- .30 --------- ----------
102 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

drie:al axial pit, co-rresponds to the inner tabularium 

and also has a flat bottom. 

Internal features.-Septa radially arranged; 

major septa extend eight-tenths the distance to the 

axis, where they are sharply deflected to form a 

small, open aulos. Minor septa extend from 0.4 to 

0.6 the distance to the aulos; two minor septa adjacent to cardinal ( ?) septum may be longer than the 

other minor septa. Number of major septa in six 

corallites ranges from 25 to 27, average 26 (table 31 

and fig. 31). Septa thin to slightly dilated; zigzag 

and subyard-arm carinae very common. 

The aulos is small, averaging only two-tenths the 

total diameter in six corallites (table 31). The aulos 

is open, presumably in the direction of the cardinal 

septum; the opening is occupied by only one, two, or 

three septa depending on whether or not the cardinal 

and adjacent minor s-epta are deflected to help form 

the aulos wall. 

The dissepimentarium occupies approximately sixtenths of the radius and consists of seven to nine 

rows of globose dissepiments. 

The tabularium is divided into two parts. Within 

the aulos, tabulae are complete, flat, or concave and 

are spaced four to five per 5 mm; at the aulos opening the tabulae bend downwards and rest on the next 

lower tabula. Outside the aulos, tabulae are concave 

or inclined, complete or inco-mplete; spacing five to 

eight per 5 mm. 

Microstructure.-As for the genus and family, but 

specimen is not well preserved, and the two types of 

tissue are indistinct. 

Discussion.-The single specimen was found in the 

USNM collections labeled "Albany Co., New York." 

The stratigraphic position is not recorded but the 

adhering rock is suggestive of the middle Moorehouse of this area. 

The specimen is characterized by its very small 

open aulos and by its well-developed carinae. In these 

and other known characters the New York specimen 

is similar to the specimens described and illustrated 

as Schistotoecholasma obliquus and S. typicalis by 

Stewart (1938). 

Material and distribution.-Illustrated specimen, 

USNM 10721, from the Onondaga Limestone, Albany County, N.Y.; horizon and exact locality unknown, but most likely the middle part of the Moorehouse Member. 

Eridophyllum seriate population B 

Plates 68, 69 

Occurrence of typical material.-AcinophyllumEridophyllum bed, western coral facies, Moorehouse 

Member, Onondaga Limestone, Leroy and Stafford, 

N.Y. 

Diagnosis.-E. seriale with a relatively small, 

closed aulos and dissepimentaria with alternate 

zones of small and large dissepiments. Interpreted as 

a population developed in a restricted area over a 

short period of time. 

External features.-Phaceloid coralla as much as 

20 or more centimetres in height and probably several times that in diameter, composed of subparallel 

corallites. Mature corallites irregularly cylindrical, 

8 to 13 mm in diameter (colony means ranges from 

8.4-10.4 mm, average 9.5 mm; table 31 and fig. 31). 

Increase is lateral at infrequent intervals. Expansion is rapid and the conical stage is limited to the 

proximal 1.0 to 1.5 em. Lateral supports are numerous; they tend to be closely spaced and oriented in 

one direction in many coralla. Corallites are irregularly rugose and the lateral supports are extensions 

of the coarser rugae. 

Spacing of corallites within a colony varies, commonly from one-half to one diameter. 

Calices not known but apparently rather broad, 

depth approximately one-half diameter. 

Internal features.-Septa radially arranged. 

Major septa extend into the tabularium where they 

are deflected to form the aulos, which is closed in 

transverse section. Average number of major septa 

in mature corallites of 10 coralla ranges from 22.4 to 

25.2 (intercolony average 24.0; table 31 and fig. 31) ; 

mature corallites have as few as 20 and as many as 

28 major septa. Minor septa extend 0.5 to 0.7 the 

distance to the aulos. 

Primary septa are not recognizable in sections 

through the cylindrical parts of most corallites. A 

short major septum, commonly "leaning" on an 

adjacent septum, may mark the cardinal position in 

some individuals. The deflection of the major septa 

at. the aulos tends to be away from the cardinalcounter plane and marks the general position of 

these septa in some corallites but doesn't differentiate between them; in such corallites, the cardinal 

and (or) counter septa seem to split and deflect in 

both directions. 

The aulos is circular in transverse section and 

almost invariably closed except in the presence of 

"worm" tubes. (See discussion.) The open aulos 

in some specimens is a pathologic condition. 

The aulos is extremely irregular in any individual 

corallite so that shape and diameter commonly vary 

in successive serial transverse sections a few millimetres apart. The figures for aulos diameter given in 

table 31 and figure 31 are of limited value but serve 
SYSTEMATIC PALEONTOLOGY 103 

to show the approximate average value and range of 

variation. "Colony averages" of 0.17 to 0.30 of corallite diameter indicate a relatively small aulos in comparison with other described species. 

Zigzag carinae are present in all corallites but 

may be strongly or weakly developed. They are commonly limited to the disS'epimentarium. 

8epta are thin but slightly thicker peripherally. 

The dissepimentarium is composed of four or more 

rows of globose, steeply inclined dissepiments. Both 

large and small dissepim'ents may occur in a corallite, commonly in alternating growth zones. 

The tabularium is divided into two zones by the 

aulos. Tabulae within the aulos are flat and commonly horizontal, two to six per 5 mm. Tabulae outside the aulos are flat or concave, 6 to 11 per 5 mm. 

Tabulae and diss·epiments are thin. 

Increase.-Lateral offsets are formed as described 

for the genus. The offset aulos tends to open in the 

direction of the parent but the parental aulos is 

closed throughout the offsetting stage. 

Microstructure.-As for family and genus. 

Variation.-lntracolony and intercolony variation 

in selected characters is shown in table 31 and figure 31. Samples are small because of the difficulty of 

finding undisturbed coralla, but semiquantitative 

examination of many additional specimens shows 

that the indicated observed ranges encompass most 

of the specimens collected. 

Discussion.-Eridophyllum seriale population B 

and Acinophyllum segregatum compose the Acinophyllum-Eridophyllum biostrome within the western 

coral facies of the Moorehouse Member. The biostrome is 1 to 2ft (0.3 to 0.61 m) thick and extends 

for 20 or more miles (32 or more km); the two coral 

species are approximately equally abundant. Comparable specim·ens of E. seriale are known from two 

other localities in the western Moorehouse; both are 

in the upper part of the member and may represent 

the horizon of the Acinophyllum-Eridophyllum bed 

and parts of the same population. Comparable forms 

are not known from central or eastern New York or 

from Ontario. 

Within the Acinophyllum-Eridophyllum bed, many 

coraUa are in growth positions but are so closely set 

that a continuous bed is formed. The bed is partly 

chertified and known only in vertical quarry walls 

and in one deeply weathered near-surface exposure. 

For these reasons, coralla boundaries cannot be recognized and samples consist of blocks of varying size 

removed from the bed at inte~rvals of no less than 

10 ft (3 m). Studied "colonies" may be from one 

corallum or from two or more intergrown coralla. 

"Worm" tubes are commonly found within corallites of this species (pl. 69, figs. 6-9). The "worm" 

apparently lived within the tentacular ring of the 

coral and grew upwards with the coral. The "worm" 

tubes are long, straight, and hollow and are attached on their outer sides to dissepiments, tabulae, 

and septa within the dissepimentarium or the outer 

tabulari urn of the coral. 

Corallite structures are irregular adjacent to the 

tubes and the aulos is commonly open toward the 

tube (pl. 69, fig. 9). The opening, however, is a 

pathologic condition resulting from the presence of 

the "worm." "Worm" tube microstructure consists 

of a thin, dark inner layer and a thicker and lighter~ 

colored outer layer. By analogy with the structure of 

dissepiments and tabulae, it is clear that the tube 

was deposited by the coral to protect itself from the 

"worm" parasite. Somewhat similar tubes have been 

described in horn corals (Clarke, 1908, 1921) and in 

favositid corals (Sokolov, 1948). 

Distribution.-Abundant in the AcinophyllumEridophyllum bed (rare elsewhere) in the western 

coral facies, Moorehouse Member, Onondaga Limestone, western New York; Middle Devonian, 

Eifelian. 

Material studied.-lllustrated or measured specimens, USNM 163426-163435 and NYSM 12831. Additional studied specimens, USNM 5308C, D, 

5308aB, 5311A, B, and 5334D; NYSM 3488-3491. 

Eridopbyllum subseriale Stumm 

Plates 70-72 

1954 Eridophyllum subseriale Stumm, p. 5, pl. 1, 

figs. 6, 7. 

? 1961 E. seriale, Fagerstrom, p. 15, pl. 3, figs. 15, 16. 

Occur-rence of type material.-"Lower Onondaga 

Limestone; Caledonia, New York" (Stumm, 1954, 

p. 10). This is certainly the Edgecliff Member. Middle Devonian (Eifelian). 

Diagnosis.-Phaceloid Eridophyllum with large, 

open aulos, and relatively few lateral supports. 

External features.-Colonies as much as 25 or 

more centimetres in diameter, 15 em or more in 

height; composed of radiating or subparallel corallites. Corallites closely spaced, one-half to one diameter apart, rarely in contact, with few lateral supports. Mature corallites range in diameter from 8 to 

16 mm; average diameter in six reasonably well preserved coralla ranges from 9.3 to 13.5 mm, average 

10.9 mm (table 32 and fig. 32). 

Increase is lateral at infrequent intervals. Expansion is rapid and the conical stage is limited to the 

basal 1 to 1.5 em. 
104 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Corallites are relatively smooth with subdued 

rugae: septal grooves very lightly marked. 

Calices not known; the morphology of the dissepiments and carinae suggest that they were broad and 

shallow. 

Internal features.-Septa radially arranged; 

major septa extend more than half the distance to 

the axis, where they are sharply deflected to form the 

aulos; minor septa variable, extend 0.2 to 0.7 the distance to the aulos. Number of major septa in cylindrical parts of corallites ranges from 22 to 30, average number in six coralla ranges from 23.8 to 27.7, 

average, 25.8 (table 32 and fig. 32). 

28r--.------,------~-~,---/~~-~~~J------~~--~ ~;;..'"' 

// VJ \ 

't::: 

........ \ 

/'" /XJ ', 

(o /I ' I/ \ 

E. aulodokum --jt7 I \, 

/1 I o .,.r:;; 

/ I q /...-

/ I I _,.. 

/ to/ --- / \ ;--- £. subsenale 

A 

26 r24-

/X X I lx I 

22- I >0< X I 

( I 

/ X I £. corn/culum 

2o--lx I 

X I 

\ I 

18 '~1 6 

I I 

8 10 

} (mm) 

I 

12 

'E 

s 4 

2 

6 8 10 12 

d (mm) 

EXPLANATION 

X £ndopf?yllu,m cormculum 

0 £. subsenale 

+ £. aulodokum 

-

-

-

-

I 

14 

14 

FIGURE 32.-Eridophyllum spp. data. A, Mean number of 

major septa (n) versus mean diameter (d). B, Mean aulos 

diameter (dav.) versus mean diameter (d). Straight lines 

are selected dau I d ratios. 

Septa are thin. Zigzag carinae weakly to strongly 

developed; present in all corallites but variably developed within corallites and coralla. 

The aulos is large, averaging from 0.32 to 0.49 of 

total diameter in seven colonies analyzed (table 32 

and fig. 32B). Within colonies, the aulos is rarely less 

than three-tenths, but m·ay exceed five-tenths of total 

di·ameter; there is some variation along the length 

of individual corallites. The aulos is commonly bulb 

shaped in transverse section, open at the position of 

the cardinal septum ; co•mmonly this opening is occupied by the c~ardinal septum and by the two adjacent 

minor septa which may be longer than the other 

minor septa (pl. 71, fig. 3) ; in some transverse sections, the cardinal septum curves to one side and may 

join the aulos leaving a single minor septum to occuPY the opening in the aulos. The counter septum is 

flanked by long minor septa in some corallites (pl. 

71, fig. 3). The dissepimentarium occupies 0.25 to 

0.40 of the corallite radius and is composed of from 

two to six, commonly three or four rows of globose, 

rarely elongate, dissepiments. The lateral supports 

consist of a lateral extension of normal septal and 

dissepimental tissue within the outbent epitheca. 

The tabularium is divided into two parts. Within 

the aulos, tabulae are complete or incomplete, flat or 

slightly arched; they terminate abruptly against the 

aulos wall; at the opening in the aulos they bend 

down and rest on the next lower tabula (pl. 71, fig. 

4) ; spacing within the aulos ranges from 2 to 11 per 

5 mm. Between the aulos and the disse·pimentarium, 

tabulae are complete or incomplete, either flat or 

concave, and horizontal or steeply inclined; at the 

aulos opening the outer tabulae terminate against 

the down bent inner tabulae; spacing of outer tabulae 

ranges from 3 to 11 per 5 mm. There is considerable 

variation in shape, orientation, and spacing of both 

inner and outer tabulae within any individual. 

Microstructure.-As for the genus and family. 

Increase.-Offsetting is lateral as in all species of 

the genus; commonly only one offset is formed, but 

one example of "twin" offsets is known. In transverse section the aulos in the hystero-corallite breaks 

at the position of the offset, commonly not the position of the cardinal break. The septa in the offset are 

short; the offset aulos may form after separation 

from the hystero-corallite, or may be partly formed 

before separation. 

One corallite is known that was apparently formed 

by two polyps that failed to separate. For a length of 

2 em, the corallite is elongate in cross section (15 by 

19 mm), with an elongate aulos ( 4.6 by 9.5 mm), two 

openings in the a ulos ( ? two cardinal septa) , and 
SYSTEMATIC PALE ONTOLOGY 105 

TABLE 32.-Eridophyllum subseriale Stumm and E. aulodokum n. sp., summary data 

E. subseriate E. 

aulodokum 

Intracolony 

Minimum Maximum Mean Intercolony Intra colony 

N ------------------------ 6 14 8.7 7 10 

n: x ----------------- 23.8 27.7 25.8 25.6 

OR --------------- 22-25 26-30 23.8_:_27.7 23-29 8 ----------------- .98 2.86 1.46 1.7 

v ---------------- 3.9 11.5 6.1 5.7 6.7 

d: x ----------------- 9.3 13.5 10.9 9.2 

OR --------------- 8.2-10.5 12.5-16.0 9.3-13.5 7.9-12.0 8 ----------------- 1.0 2.1 1.5 1.3 

v ---------------- 5.8 18.2 12.6 13.6 14.1 

d·": x ----------------- 3.2 4.9 4.0 4.9 

OR --------------- 2.5- 3.7 4.1- 6.0 3.2-4.9 3.7-6.3 8 ----------------- .4 .9 .5 .8 

v ---------------- 13.7 22.6 17.3 13.2 16.3 

n/d: x ----------------- 1.9 2.8 2.4 2.8 

OR --------------- --------- ---------- 1.9-2.8 --------- da"/d: x ----------------- .32 .43 .37 .53 

OR --------------- --------- ---------- .32-.43 ---------

nearly twice the normal complement of major septa 

( 42; average of 11 round corallites, 25) (pl. 71, figs. 

6-8). This might represent an example of incomplete 

fission ('axial increase) or of parent and lateral offset that failed to completely separate. 

Discussion.-The above description is based on 

more than 30 specimens but most of these are fragments containing only a few corallites. Figures in 

table 32 are based on 6 to 14 corallites in seven coralla including the holotype. 

Eridophyllum subseriale is the only species of the 

genus known to occur in the Edgecliff Member. It is 

most common in eastern New York where specimens 

have been collected from nearly half of the known 

bioherm localities. It is known from the biostrome 

facies of the Edgecliff at two localities. The holotype 

specimen is from western New York (Caledonia), 

presumably from the Edgecliff Member. A number of 

overturned and crushed coralla were found in the top 

bed of the Edgecliff at a locality near Syracuse. None 

of these specimens are suitable for quantitative 

study but they indicate that the species was at least 

locally common in the nonbioherm facies of the 

Edgecliff. It is not known with certainty that the 

species occurs elsewhere. 

Fagerstrom described as E. seriale a single fragment of a cylindrical corallite of Eridophyllum from 

the Formosa bioherm facies of the Detroit River 

Group in southwestern Ontari~. Restudy of the specimen indicates that it is morphologically closer to E. 

subseriale in its very large aulos (6 mm in diameter), relatively large dissepiments, and widely 

spaced tabulae. Identification of the fragment is not 

possible, but it is tentatively placed here pending the 

discovery of more adequate material. 

E. subseriale is characterized by its large aulos, 

opening at the position of the cardinal septum, and 

differs from most other species of the genus in this 

character. Eridophyllum aulodokum, in the Moorehouse Member, has an even larger aulos and fewer 

and weaker carinae. Some specimens of E. seriale 

from the Columbus Limestone and specimen 1 from 

the Moorehouse Member have auloi that average 

only slightly smaller than those of E. subseriale, but 

these may be open or closed in the same corallum. 

Examination of numerous coralla has convinced me 

that the Edgecliff ·and Columbus-Moorehouse forms 

are distinct. 

Distribution.-Known only from the Edgecliff 

Member of the Onondaga in New York. Most specimens are from the bioherm facies in the east-central 

and eastern areas; a single specimen is known .from 

the biostrome facies near Schoharie (eastern area) 

and 20 or more coralla are known from the top bed 

of the Edgecliff near Syracuse (central area). The 

holotype was collected from the "Lower Onondaga 

Limestone, Oaledonia, N.Y." (Stumm, 1954, p. 10) ; 

this is almost c-ertainly the Edgecliff Member, and is 

the only specimen known from the western area. 

Material studied.-Holotype, UMMP 5272; illustrated or measured specimens USNM 163454-457, 

NYSM 12832-34; 11 additional sectioned specimens, 

U.SNM 163458a-d, 163459-462, NYSM 3492-94 ; 

nine more specimens provided additional distribution data. 

Questionable specimen from the Detroit River 

Group, UMMP 36122 (hypotype of Fagerstrom, 

1961). 
106 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Eridophyllum aulodokum n. sp. 

Plate 73 

Diagnosis.-Phaceloid with few or no lateral supports; very large aulos averages more than half the 

total diameter ; carinae very weak; tabulae closely 

spaced. 

External features.-A single specimen is in a 

block of limestone approximately 40 by 20 by 14 em; 

the cor,allites are not in their original relative positions, but the corallum evidently was somewhat larger than the block. The corallum was apparently 

phaceloid, with straight to somewhat curved cylindrical corallites. Diameters of cylindrical parts of 

corallites range from 8 to 12 mm (table 32). Increase is lateral; conical stage is short. Lateral supports are rare. 

Corallite exteriors are relatively smooth; septal 

grooves are very fine. Very fine growth lines appear 

on well-preserved exteriors; coarser rugae are virtually absent. Calice not known. 

Internal features.-Major septa radially arranged 

but deflected to form a large aulos; minor septa extend 0.3 to 0.6 the distance to the aulos wall. Number 

of major septa in cyclindrical parts of corallites 

ranges from 23 to 29, average number in 10 corallites is 25.6 (table 32 and fig. 32). 

Septa are thin. Zigzag carinae very weakly developed to absent. 

The aulos is large, ranging from 0.46 to 0.71 of 

total diameter in 10 corallites, averaging 0.53 (fig. 

32B). The aulos is bulb shaped in transverse section, open at the position of the cardinal septum; the 

opening is occupied by 0, 1, 2, or 3 septa, as any or 

all of the cardinal and the two adjacent minor septa 

may be deflected to help form the "neck" of the 

aulos. The minor septa that are adjacent to the 

cardinal and counter septa are commonly longer than 

other minor septa. 

The dissepimentarium ·occupies approximately 

one-fourth of the corallite radius and is composed of 

from two to four rows of globose dissepiments. 

The tabularium occupies approximately threefourths the corallite diameter and is divided into 

two zones. Tabulae within the aulos are complete, 

flat or slightly concave; spacing ranges from 5 to 12 

per 5 mm. Between the a ulos and the dissepimentarium, tabulae are complete, and commonly concave and 

inclined away from the aulos; spacing averages six 

per 5 mm with little variation in the small sample 

studied. 

Microstructure.-As for genus and family. 

Increase.-As for genus. The position of the offset 

appears unrelated to the aulos opening or the cardinal position of the parent. 

Discussion.-Eridophyllum aulodokum is known 

from a single block of limestone that was knocked 

loose during normal quarry operations but was set 

aside by a quarry employee and given to me while I 

was collecting in the quarry (loc. 112, western area). 

The stratigraphic position of the specimen is not 

known; the quarry wall exposes all but the lower few 

feet of the Moorehouse Member of the Onondaga 

Limestone and locally the basal few feet of the overlying Seneca Member. Therefore, the specimen is 

probably from the Moorehouse but possibly from the 

Seneca. A small fragment of an Eridophyllum from 

the Moorehouse Member in eastern New York is tentatively assigned to the same species. 

E. aulodokum is unique in the possession of an 

aulos occupying more than half the corallite diameter. It differs from E. subseriale in its larger aulos 

(fig. 32), complete tabulae, and weak or absent carinae. E. aulodokum may have been derived from E. 

subseriale but, in the absence of more material, this 

is pure speculation. 

E. seriale from the Moorehouse Member is larger 

with a relatively smaller aulos and with well developed, long carinae. 

Mat erial and distribution.-Holotype, USNM 

163436, locality 112, western area, Moorehouse ( ?) 

or Seneca ( ?) Member. A second specimen including 

fragments of four corallites is tentatively assigned 

to this species; USNM 163437, locality 246, eastern 

area, upper Moorehouse Member. 

Eridophyllum corniculum n. sp. 

Plates 74, 75 

Occu1-rence of type material.-Moorehouse Member of Onondaga Limestone, Harris Hill and Pembroke Station, New York. 

Diagnosis.-Eridophyllum with an auloporoid 

growth habit; corallites commonly short and cone 

shaped. Aulos variable, tending to be medium sized 

and open at position of cardinal septum. 

External features.-Colonies small, varying from 

a few individuals to several hundred; the largest 

known coral] urn is 37 by 20 em in diameter and 6 em 

in height. Most known coralla are formed of short, 

curved cone-shaped, reptant corallites, attached at 

points of lateral budding (pl. 74, fig. 5). Lateral supports are present and help to form a fairly rigid network by attaching the corallite to another corallite 

or to the substrate. Some coralla are only one corallite thick; each corallite rests on the substrate and 

curves upward so that the upper surface of the 
SYSTEMATIC PALEONTOLOGY 107 

colony shows upward directed, uniformly spaced 

calices. Each corallite gives rise to from one to three 

or more offsets, commonly on the underside of the 

parent, just above the substrate position. 

Individual corallites commonly range in length 

from 1 to 3 em, but one corallum is known in which 

individuals lengthened to 6 or more centimetres. Diameters of mature corallites range from 5 to 13 mm 

(colony averages 5.6 to 11.1 mm; table 33 and fig. 

32). 

TABLE 33.-Eridophyllum corniculum n. sp., summary data 

Intn.colonx 

Mini- Maxi- Mean Intermum mum colony 

N ---------- 5 10 6.7 11 

n: x --- 18.2 26.7 21.7 

OR -- 17-19 24-30 18.2-26.7 

8 ---- .55 1.94 2.15 

v --- 2.4 14.8 6.7 9.9 

d: x 5.6 11.1 6.9 

OR -- 5.3-6.0 8.5-13.2 5.6-11.1 

8 ---- .30 1.91 1.49 

v --- 5.3 28.9 13.9 21.7 

dau: x 1.4 2.9 2.0 

OR -- 1.1-2.0 1.9-3.6 1.4-2.9 

8 ---- .23 .64 .40 

v --- 13.4 30.7 21.1 20.5 

r/d: x 2.4 3.9 3.3 

OR -- --------- --------- 2.4-3.9 

da;./d: x ---- .21 .37 .29 

OR -- --------- --------- .21-.37 

Corallites are strongly rugose, especially at positions of lateral supports. Shallow septal grooves and 

low, broad interseptal ridges are present. 

Spacing of corallites ranges from zero to approximately one diameter. 

Calices are broad and shallow with an axial pit, 

corresponding to the tabularium, a rather steep inner wall, and a gently sloping to nearly flat peripheral platform. 

Internal features.-Septa radially arranged. Major septa extend 0.6 to 0.8 the distance to the axis, 

where they are sharply deflected to form an aulos. 

Number of major septa ranges from 17 to 30 in 

corallites of 5 to 13 mm di31meter; average number 

of major septa in 11 colonies ranges from 18.2 to 

26.7 (table 33 and fig. 32). The cardinal septum is 

commonly straight with a break in the ·aulos at this 

point. Minor septa a.re short, extending from 0.1 to 

0.5 the distance to the aulos. Septa are thin; zigzag 

carinae are strongly or weakly developed within the 

dissepimentarium; subyard-arm carinae are present 

in some sections. 

The dissepiment•arium is composed of two to six, 

commonly two to three, rows of globose dissepiments. The lateral supports are an outpocketing of 

the outer wall filled with dissepiments. The tabulariurn consists of outer and inner zones of more or less 

horizontal, complete tabulae, separated by the aulos. 

Aulos di,ameter ranges from 0.2 to 0.4 of the total 

diameter; tabularium diameter is as much as sixtenths of total diameter. Tabulae within the aulos are 

flat, either horizontal or slightly inclined; three to 

nine per 5 mm. Tabulae outside of the aulos are horizontal and flat, or sagging or inclined toward the 

axis ; commonly seven to nine per 5 mm. 

Microstructure.-All known specimens of E. corniculum are at least partly silicified and microstructure is not well preserved. Insofar as it can be deterInined, septal and wall structure are as described for 

the genus. 

Increase.-Offsetting is lateral; commonly a single 

set of from one to three offsets is present a short distance below the distal margin of the calice on the 

convex side of the corallite. The offset is commonly 

1ocated at the position of the opening in the parental 

aulos; the offset aulos forms early and opens toward 

the parent. 

Discussion.-The above description is bas·ed on a 

eollection of 25 complete and incomplete coralla from 

Harris Hill, N.Y. (loc. 112) and seven fragmentary 

eoralla from Pembroke Station, N.Y. (loc. 393) ; one 

of the latter is a silicified fragment, naturally etched 

:from the rock and showing the auloporoid habit to 

good advantage (pl. 74, fig. 5). Fourteen coralla, including two from Pembroke Station were thin sectioned, and all measurements and counts are based 

on from 5 to 10 corallites in 10 of the coralla. The 

Harris Hill coralla were found at a single horizon 

over an area 10 by 40 feet (3 by 12 m) and may 

:represent a single living population. The Pembroke 

Station fragments, with the exception of the naturally etched one, are possibly fragments of a single 

corall urn. In any case the type lot consists of specimens drawn from two small areas and variation 

within the species is likely to be significantly greater 

than that indicated by this description. 

E. corniculum has typical craspedophyllid internal 

structures and the characteristic Eridophyllum type 

of aulos. It differs from all other known Eridophyllum in its auloporoid (reptant) growth habit although the lateral offsetting is characteristic of the 

genus. Disphyllum cat'enatum Smith 1945 (p. 21-22, 

pl. 11, figs. 9-12) has a nearly identical growth habit 

and may be profitably compared in this respect. 

Smith noted that this mode of colony formation is 

unusual in rugose corals but "that the type specimens of D. catenatum may be merely the beginnings 

of a phaceloid colony and that possibly this mode of 

colony building is less uncommon than it appears to 
108 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

be." Eridophyllum seriale and E. aulodokum have 

not been found associated with E:- corniculum although all three species occur in the Moorehouse 

Member in western New York. E. seriale and E. 

aulodokum differ in the size and nature of the aulos 

as well as in growth form so it is unlikely that they 

represent different growth forms of the same 

species. 

In its general form, E. corniculum is distinct from 

all other known Onesquethawan species. 

Distribution.-Moorehouse Member, Onondaga 

Limestone (Middle Devonian), western New York. 

The Harris Hill "population" was taken from a level 

approximately 10 ft (3 m) below the Tioga Bentonite Bed and 20 ft ( 6.1 m) above the level of the 

Acinophyllum-Eridophyllum bed. The Pembroke 

Station specimens were taken from an isolated 16-ft 

( 4.85-m) exposure of the middle or upper Moorehouse and are probably older than the Harris Hill 

population. 

Material studied.-Holotype, USNM 163438; illustrated or measured paratyp.es, USNM 163439-450, 

and NYSM 12835; unillustrated paratypes, USNM 

163451-452 (12 specimens), 163453 (2 specimens), 

and NYSM 12836-41 (6 specimens). 

Eridopbyllum conjunctum (Davis) 

Plates 107, 108 

1887 Diphyphyllum conjunctum Davis, pl. 116, figs. 

1-3. 

1887 D. coalescens Davis, pl. 117, fig. 1. 

1900 D. adjunctum Greene, p. 27, pl. 12, fig. 1. 

1965 Eridophyllum conjunctum (Davis). Stumm, 

cards 307, 322. 

1966 E. coalescens (Davis). Stumm, card 317. 

? 1965 H exagonaria prisma. Carman, p. 70, fig. 3. 

1966 E. conjunctum (Davis). Stumm, p. 45-46, pl. 42, 

figs. 5-7. 

Occurrence of type material.-"J effersonville 

Limestone, above coral zone" (Stumm, 1965, p. 46), 

Falls of the Ohio. 

Diagnosis.-Cerioid Eridophyllum with corallites 

12 to 18 mm in diameter. Aulos diameter, 2 to 4 mm, 

complete or open at cardinal position. Major septa 

number 23 to 25 in two Falls specimens. Carinae consistently present but variously developed. 

Description.-A single specimen of a cerioid craspedophyllid with well-developed Eridophyllum-type 

aulos was found in the upper part of the Moorehouse 

Member in western New York. The specimen is lens 

shaped, 27 em in diameter and 8 em high. Corallite 

and aulos diameter and number of major septa are 

given in table 34, where they can be compared with 

corresponding data from the Falls of the Ohio specimens. There are no obvious differences between the 

Moorehouse specimens and those from the Falls that 

cannot readily be accounted for by even less intercolony variation than is common in other craspedophyllids. 

Discussion.-! agree with Stumm (1965) in synonymizing the three Falls species of Davis and 

Greene: thus making E. conjunctum the only described cerioid Eridophyllum from rocks of Onesquethaw age. Another species, E. arachnoideum Stumm 

(1954, p. 6), is also cerioid but has a much smaller 

aulos and differs in other ways also. This species is 

known from beds of Centerfield and younger age in 

the Hamilton Group of New York. 

Carman ( 1955, p. 70, fig. 3) described and illustrated as Hexagonaria prisma a cerioid craspedophyllid with a weak aulos from an unknown position 

within the Columbus Limestone in a core taken at 

Chillicothe, Ohio, south of Columbus. The specimen 

was crushed before burial and is known only from a 

small fragment, but it may be conspecific with E. 

conjunctum (table 34) . 

TABLE 34.-Eridophyllum conjunctum (Davis) intracolony 

data 

New Falls of Ohio Ohio York 

USNM MCZ MCZ osu 

172245 7859 7877 21336 

N ---------- 10 2 4 5 

n: x ------ 26.1 24.0 24.0 23.0 

OR ---- 23-30 23-25 23-25 22-25 

d: x ------ 15.5 16.0 13.0 12.2 

OR ---- 13-18 13-18 8-18 10-14 

da": x ------ 2.8 2.8 3.6 -------- OR ---- 2.6-3.5 2.2-3.4 3.2-4.0 1.6-2.0 

Distribution.-Moorehouse Member, Onondaga 

Limestone, 18 inches ( 46 em) above (AcinophyllumEridophyllum bed, at locality near Stafford, western 

New York (collected by A. Ferrari, Bologna). 

Jeffersonville Limestone, Falls of the Ohio. "Above 

coral zone" (Stumm, 1965, p. 46); probably ZoneD, 

E,orF. 

Material studied.-Lectotype of D. conjunctum 

Davis, MCZ 7859; holotype of D. coalescens Davis, 

MCZ 7877; holotype of D. adjunctum Greene, 

AMNH 23501. Additional specimens in the Davis 

collection were studied but not sectioned. 

Moorehouse specimen, USNM 172245. 

Specimen illustrated by Carman from the Columbus Limestone, OSU 21336. 

Genus GREWGIPHYLLUM Oliver, 1974 

1974 Grewigphyllum Oliver, p. 170. 

Type species.-H eliophyllum colligatum Billings, 

1859, p. 126-127. First illustr·ated by Nicholson, 
SYSTEMATIC PALEONTOLOGY 109 

187 4, p. 28, pl. 5, figs. 3, 3a. "Rama's Farm, near 

Port Colborne [Ontario]" (Billings, 1859, p. 127). 

Presumably the Bois Blanc Formation, Emsian. (See 

discussion.) 

Diagnosis.-Craspedophyllinids with an imperfect 

and irregular aulos formed by deflected major septa, 

some of which continue into the aulos and are irregularly bent and twisted. Colonial, with corallum composed of straight corallites that are at alternate 

levels expanded to form a cerioid stage and contracted to form a phaceloid stage. Increase is lateral. 

Description.-Only one species is known; see description of G. colligatum. 

Discussion.-Grewgiphyllum colligatum (Billings) has generally been assigned to Eridophyllum 

because of the presence of a distinct aulos. However, 

it differs f.rom that genus in its imperfect aulos 

(possibly primitive) and in its distinctive growth 

form, not otherwise known in the Craspedophyllidae. 

It could be considered an early (primitive) Eridophyllum, but its distinct morphology and stratigraphic position can be emphasized by giving it a 

new generic name. It is the earliest known craspedophyllinid and may be the necessary evolutionary intermediate between the cylindrophyllinid Prismatophyllum and ETidophyllum as discussed elsewhere 

in this paper. 

Smith and Lang (1927, p. 308) and Lang, Smith, 

and Thomas (1940, p. 41) synonymized Craspedophyllum americanum Dybowski ( 1873, type species 

of Craspedophyllum Dybowski, 1873) with G. colligatum. If correct this would make Craspedophyllum 

a senior synonym of Grewgiphyllum. I do not accept 

this for the following reasons: (1) Dybowski's aulos 

description is of an ETidophyllum; his figure 1 could 

be either an E1·idophyllum or a Grewgiphyllum, but 

his figures 2 and 5 are of a good Eridophyllum, (2) 

Stewart (1938, p. 40) reported two specimens of "E. 

colligatum" from Columbus, Ohio (type locality of 

Dybowski's species), as drjft specimens; one of these 

is an Arachnophyllum; the other is correctly identified as G. colliyatum but is similar in preservation to 

Ontario Bois Blanc specimens and may not be from 

the Columbus at all. I know of no other reports of 

G. colligatum in the Columbus Limestone and on this 

basis it seems far more likely that C. americanum 

was based on one of the common Columbus Eridophyllum rather than on a (at least rare) Grewgiphyllum. 

Distribution.-Only one species is known. This is 

limited to rocks of middle and early late Onesquethaw age (Bois Blanc-lower Edgecliff) in eastern 

North America. 

Grewgipbyllum colligatum (Billings) 

Plates 76-79 

1859 Heliophyllum colligatum Billings, p. 126-127. 

1874 H. colligatum Billings. Nicholson, p. 28, pl. 5, 

figs. 3, 3a. 

1876 Diphyllum colligatum (Billings). Rominger, p. 

127' pl. 38, fig. 3. 

not 1887 Cyathophyllum colligatum Davis, pls. 91, 92 and 

pl. explanations. 

1901 Crepidophyllum colligatum (Billings). Lambe, p. 

155-156, pl. 13, figs. 2a, b. 

1906 Eridophyllum colligatum (Billings). Grabau and 

Shimer, p. 209. 

1909 E. colligatum (Billings). Grabau and Shimer, p. 

72-73. 

1933 E. colligatum (Billings). Smith, p. 520. 

part 1938 E. colligatum (Billings). Stewart, p. 40, pl. 7, fig. 

7, (not fig. 8 which is an Arachnophyllum sp.). 

part 1944 E. colligatum (Billings). Shimer and Shrock, p. 

91, pl. 27, figs. 6-9 (not fig. 10 which is an 

Arachnophyllum sp.). 

not 1949 E. colligatum (Billings). Stumm, pl. 98, figs. 7-

11 (fig. 7 is Arachnophyllum sp., figs. 8-11 

are Eridophyllum spp.). 

1955 E. colligatum (Billings). Stumm, cards 320, 321. 

1974 Grewgiphyllum colligatum (Billings). Oliver, 

figs. 4a-b. 

OccurTertce of type material.-The neotype is from 

the Bois Blanc Formation at Hagarsville, Ontario. 

N eotype.-Billings cited "Rama's Farm, near Port 

Colborne" (Billings, 1859, p. 127) as the type locality. Billings specimen is apparently lost and as it was 

not illustrated would not be recognizable unless it 

was clearly labeled in Billing's hand. Nicholson, 

1874, was first illustrator but his specimens apparently were destroyed by fire. The specimen on which 

Lambe, 1901, apparently based his description and 

illustrations (diagrammatic) is available (GSC 

3590) but is poorly preserved and is not from the 

type area. Both the Bois Blanc and the lower Onondaga crop out at and near Port Colborne and both 

formations are represented in collections from 

"Rama's Farm." During this study, specimens of G. 

colligatton were found in the Edgecliff Member at 

Port Colborne and in the Bois Blanc Formation 

farther west at Hag·arsville. I here select as neotype, 

specimen GSC 31158 (pl. 76, figs. 1-5), collected by 

J. DeCew, from the Bois Blanc Formation near 

Hagarsville, Ontario. (Oliver, 1974, p. 172, erroneously cited this as lectotype.) 

External features.-Semimassive coralla are composed of straight, parallel, or radiating corallites 

that are alternately free (round in transverse section) and in lateral contact with neighboring corallites (polygonal in section). The expansions are at 

the same level throughout so that the covallum alternately appears to be cerioid and phaceloid. Coralla 
110 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

may be as large as 35 c.m in diameter and more than 

15 em high. Four specimens from the Edgecliff bioherm facies show considerable breakage along surfac.es passing through the dissepimentaria; unprepared specimens give the impression of a halysitoid 

growth habit. S.pecimens from the Bois Blanc Formation don't show this effect. This fact suggests a 

difference in living or preservation environment effecting a structurally weak corallum. 

Mature corallites range from 9 to 19 mm in diameter although muc.h of this variation results from the 

alternate expansion and contraction. (See table 35 

and fig. 33.) 

Increase is lateral and ·apparently infrequent in 

mature coralla. Lateral expansions are characteristic ; these tend to be concave on the underside, 

nearly flat on the upper. Each corallite is expanded 

uniformly in all directions. Expansion interv·al 

ranges from 7 to 14 per 5 em in the examples studied 

but is commonly 7 or 8 per 5 em. 

•c 

28.--.-----.-----.-----.-----.-,~~-.-,~ 

A 

26- 0 

0 

0 

DOD 

22 I I 

10 12 

10 12 

0 

0 

CJD 

I 

14 

0 =dp 

. 14. 

0 

-

-

I I I 

16 18 20 

0 =dc(mm) 

16 18 20 

o =dc(mm) 

FIGURE 33.-G-rewgiphyllum colligatum data. Mean diameters are plotted for both cerioid stage (de) and phaceloid stage ( dp) but not for all coralla. A, Mean number of 

major septa (n) versus mean diameter (dp and de). B, 

Mean aulos diameters (d"'ii'U) versus mean diameter (dp 

and de) • Straight lines are selected do:;. I d ratios. 

Corallites are gently rugose between expansions; 

septal grooves are fine and inconspicuous. 

Calice inferred to be shallow with an axial boss 

corresponding to the aulos; a peripheral platform is 

present and especially broad during stages of 

expansion. 

TABLE 35.-Grewgiphyllum colligatum (Billings), summary 

data 

n: 

dp: 

de: 

d"v: 

[dp, phaceloiJ diameter; de, cerioid diameter] 

Minimum 

N ----- 5 

x ------ 23.1 

OR ---- 21-26 

s ------ .95 

v ----- 3.8 

N ----- 3 

x ------ 10.4 

OR ---- 8.7-12.6 

8 ------ 1.41 

v ----- 11.0 

N ----- 3 

x ------ 11.5 

OR ---- 9.3-14.5 

8 ------ .89 

v ----- 5.3 

N ----- 5 

x ------ 3.1 

OR ---- 2.6- 3.8 

8 ------ .38 

v ----- 10.9 

Intracolony 

Maximum 

20 

27.8 

26-29 

2.35 

9.1 

6 

18.8 

16.2-22.0 

2.11 

18.3 

12 

16.7 

16.0-17.7 

2.08 

17.1 

18 

5.2 

4.0- 6.3 

.83 

19.4 

ToterMean ~olony 

10 

25.1 

23.1-27.8 

1.53 

5.4 6.1 

7 

13.0 

10.4-18.8 

2.78 

14.3 21.3 

5 

13.7 

11.5-16.7 

2.07 

10.7 15.2 

8 

3.7 

3.1- 5.2 

.70 

14.3 18.9 

Internal .features.-Septa radially arranged. Major septa extend to within 1.5 to 2.6 mm of axis, 

where they are irregularly deflected to form an irregular aulos; several major septa extend into ·aulos 

after deflection to form an erratic pattern. Average 

number of major septa in 11 coralla ranges from 23 

to 28, average 25 (table 35 ·and fig. 33) ; individual 

corallites hav·e from 21 to 29 major septa. Minor 

septa are long, extending 0.6 to 1.0 the distance to 

the aulos, commonly 0.7 to 0.8. All septa are very 

thin. 

Zigzag carinae are abundant in the diss.epimentarium and commonly are found in both the inner 

and outer tabularium as well. 

The aulos is irregular and variable although commonly closed. Mean diameter in 11 coralla range 

from 2.9 to fi.2 mm, approximately one-fourth the 

mean cerioid diameter (fig. 33B). 

The dissepimentari urn is composed of three to 

eight rows of dissepiments; these •are commonly 

large although zones of small dissepiments may alternate with zones of larg.e ones. The periodic expansions of the outer wall are filled with dissepiments 

and the diameter of the tabularium is unaffected. 
SYSTEMATIC PALEONTOLOGY 111 

The tabulari urn is in two parts. Outside of the 

aulos, tabulae are commonly complete, concave and 

inclined away fro.m the aulos w·all; spacing ranges 

from 4 to 17 per 5 mm. Within the aulos, tabulae are 

ir-regular because of the erratic septa, but they tend 

to be complete and horizontal where uninterrupted; 

spacing ranges from 6 to 15 per 5 mm. 

Microstructure.-As for genus and family. 

Discussion.-The above description is based on 

specimens from the Bois Blanc Formation in Ontario and from the lower few feet of the Edgecliff 

Member of the Onondag·a Limestone in Ontario and 

western N·ew York. This is one of two colonial rugosans known to occur in both formations. 

The .species is distinguished by its imperfect aulos 

and erratic septa and by its growth form. In the 

field, the species is most easily confused with .species 

of Prisntatophyllum of com-parable size, but these 

are truely cerioid and lack the •aulos. 

Distribution.-Common in the Bois Blanc Formation and in the lower few feet of the Edgecliff Member of the Onondaga in the Niagara Peninsula of 

Ontario. Four specimens are from the lower Edgecliff bioherm facies near Buffalo, N.Y. Three specimens from Michigan ("Drift, Ann Arbor") are 

probably from the Bois Blanc Formation in the 

Mackinac area. 

Two specimens reported from the Columbus 

Limestone by Stewart (1938, p. 40) are suspect. One 

(her fig. 8) is an Arachnophyllum; the other is a 

drift specimen, possibly from the Bois Blanc Formation in Ontario. 

Material.-Neotype: GSC 31158. Illustrated or 

measured spec i men s : USNM 163463-466, 

172161-162, 113554, 113556; NYSM 12842; GSC 

31157. Other studied s-pecimens: USNM 172163, 

172164a-c ; 113552 ; pi us three specimens from 

"Drift, Ann Arbor, Michigan" in USNM. 

Family DISPHYLLIDAE Hill, 1939 

Diagnosis.-Solitary ( ?) and colonial rugose corals 

with thin or thickened major septa composed of 

coarse monacanthine trabeculae. The dissepimentari urn is formed by one to sev·eral row·s of norm·al 

globo,se to vertically elongate dissepiments. Tabulae 

are variable but com.monly incomplete with central, 

flat, horizontal tabulae and periaxial tabulae inclined toward the axis. 

Discussion.-The above is not intended as a complete diagnosis, and no description is offered of the 

family or its assigned genera. My interpretation of 

the family i'S conventional except for excluding the 

genera here grouped as the Cylindrophyllinae and 

those commonly known as phillipsastraeids (Phillipastraeinae of Jell, 1969); it is essentially the 

Disphyllinae of Jell (1969) but excluding Acinophyllum, which is clearly a craspedophyllid as discussed 

above. 

A comparison of the Cylindrophyllinae and Disphyllidae and annotated lists of species p·reviously 

assigned to the disphyllid genera Disphyllum and 

Hexagonaria are included in the discussions of the 

Craspedophyllidae and the genera Cylindrophyllum 

and Pristnatophyllum. Except as noted below, all 

8uch previous assignments are considered erroneous. 

Disphyllids occur in the Michigan Basin in rocks 

of middle Cazenovia to Tioughnioga age (Givetian). 

Thes-e have been interpreted as being part of an 

invasion of Old World forms from the northwest 

(Oliver, 1973, 1975). Both Hexagonaria and 

Disphyllurn are represented and associated with 

species of typical cylindrophyllinids. Further discussion of these form·s is beyond the scope of this paper. 

Two older species of possible Disphyllum are of 

immediate interest because they are of the same age 

(or only slightly younger) as some of the upper 

Onondaga Limestone species. One of these is from 

the Dundee Limestone (lower Cazenovia, probably 

l~ifelian, and approximately equivalent in age to the 

Seneca Member of the Onondaga Limestone) in the 

Michigan Basin. The other is from the Famine 

Limestone (upper Onesquethaw or Cazenovia; 

I~ifelian) in Quebec. Both are redescribed below for 

comparison with species of Acinophyllum and Cylindrophyllum and because of their importance in 

paleobiogeographic considerations. If properly assigned, they indicate that the incursion of Old World 

g-enera occurred earlier than previously thought 

I ikely. The Quebec species would be the only known 

disphyllid in the Eastern Americas Realm except in 

the Michigan Basin. 

Distribution in Eastern North America.-The age 

of the Quebec Disphyllum? is in doubt (see discussion of F.amine Limestone under "Stratigraphy") 

but the evidence suggests that it is post-Edgecliff 

Onesquethaw or Caz,enovia. The Michigan Disphyllum? is of Cazenovia age (Dundee). Other Disphyllum and Disphyllum? are from the Traverse Group 

(Givetian; upper Cazenovia and Tioughnioga) in 

:Michigan. 

H exagonaria s.s. first appears in the Miami Bend 

Formation of Cooper and Phelan (1966) (middle 

Cazenovia; Givetian) in northern Indiana. The 

Traverse Group contains both H exagonaria and 

Prismatophyllum. 
112 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

?Genus DISPHYLLUM Fromental 

The two species described below plus Disphyllum? 

crassiseptatum (Ehlers and Stumm) 1949 and possibly some additional Cylindrophyllum? spp. from 

the Michigan Basin Traverse Group make a group of 

Cylindrophyllum- or Disphyllum-like corals with 

thickened septa composed of coarse monacanthine 

trabeculae. These are quite strikingly different from 

the Onesquethaw Cylindrophyllum but may have 

evolved from Cylindrophyllum through thickening 

of the trabeculae. As noted above, they more likely 

are true disphyllids representing stocks that entered 

the Eastern Americas Realm from the northwest by 

way of the Michigan Basin. 

Dispbyllum? rectiseptatum (Rominger) 

Plate 30 

1876 Diphyphyllum rectiseptatum Rominger, p. 124. 

1955 Synaptophyllum rectiseptatum (Rominger). Stumm, 

card 269, three figures (of two syntypes) . 

1959 Acinophyllum? 'rectiseptatum (Rominger). McLaren, p. 

28 (synonymy only) . 

Occurrence of type material.-"Dundee Limestone, Quarry of the Michigan Limestone and Chemioal Co., Rogers City, Michigan" (Stumm, 1955, card 

269) , Middle Devonian, upper Eifelian or lower 

Givetian. 

Diagnosis.-Disphyllum? with very short septa, 

narrow dissepimentarium, numerous and vertically 

elongate lateral connections, and both complete and 

incomplete tabulae. 

External features.-Phaceloid coralla of unknown 

size. The largest fragment available to me is 6 em 

high and 3 by 6 em in diameter and includes 50 or 

more corallites. Corallite diameter ranges from 

approximately 3.5 to 6.0 mm; measurements from 

two colonies are recorded in table 36. 

Increase is lateral. Lateral supports are common 

and tend to be vertically elongate; in some sections 

the colonies appear cateniform because of the numhers of connections (pl. 30, figs. 1, 4). Corallites are 

closely spaced, commonly less than one diameter 

apart. 

Internal features .-Septa are radially arranged 

and very short; commonly the major septa barely 

enter the tabularium; minor septa are nearly as long. 

Major septa number 15 to 19 in cylindrical parts of 

corallites in two measured coralla (table 36; fig. 25). 

Zigzag carinae are present but less sharply defined 

than in species of Acinophyllum or Cylindrophyllum 

because of the general thickening of the septa. 

Carinae are well defined on attenuate septa but subdued or apparently lacking on dilated septa. 

One, rarely two, rows of normal globose dissepiments are regularly present and give the appearance 

of an inner wall in most transverse sections. Dissepiments are lacking in early parts of some coralla 

(pl. 30, fig. 3). 

Tabulae are complete or incomplete and vary from 

horizontal to strongly convex or less commonly concave. 

All structures are relatively thicker than in the 

described species of the Craspedophyllidae. 

Septal microstructure is uncertain because of 

mediocre preservation. There is a suggestion of disphyllid structure (coarse monacanths) in some 

longitudinal sections (pl. 30, fig. 2) and in the thickness of the septa (pl. 30, fig. 6). 

Discussion.-D. ? rectiseptatum was originally described but not illustrated by Rominger (1876). The 

first illus,trations were published by Stumm (1955), 

who identified three syntype specimens in the 

Rominger collections. I here select the original of 

Stumm's thin section illustrations (UMMP 14378) 

as lectotype. The above description is based on the 

three Rominger specimens only and is included here 

to complement the description and illustration of 

other colonial rugose corals. 

D. ? rectiseptatum is anal ago us to species of Acinophyllum in its narrow dissepimentarium consisting 

TABLE 36.-Disphyllum? stummi (Oliver) and D? rectiseptatum (Rominger), summary data 

12' z:ca£iacutatum 

USNM USNM USNM Mean Inter- UMMP UMMP 163237 163397 163288 colony 14378 14380 

N --------------------- 6 6 3 8 --------- 4 20 13 n: x ------------------ 24.3 23.8 24.7 22.6 --------- 23.8 15.9 17.8 

OR ---------------- 24-25 22-27 24-25 22-24 --------- 22.6-24.7 15-17 16-19 8 ------------------ .5 1.7 .6 .7 --------- .9 .4 .8 

v ----------------- 2.1 7.2 2.4 3.3 3.8 3.8 2.8 4.5 

d: x ------------------ 8.6 7.3 8.8 9.2 --------- 8.5 4.2 5.1 

OR ---------------- 7.2-10.5 6.6-8.5 7.4-9.7 8.5-9.8 --------- 7.3-9.2 2.9-5.2 4.4-6.0 8 ------------------ 1.14 .81 1.25 .42 --------- .82 .55 .44 

v ----------------- 13.3 11.0 14.2 4.5 10.8 9.7 13.1 8.6 
SYSTEMATIC PALEONTOLOGY 113 

of one basic row of dissepiments. It differs in its very 

short, thickened septa and in the probable presence 

of disphyllid microstructure. 

Distribution.-Dundee Limestone, early Cazenovia 

age. The syntypes are from the quarry of the Michigan Limestone and Chemical Co., Rogers City, 

Mich. (Crawford's quarry of Rominger), but 

Rominger reported the species from other Dundee 

localities as well. 

M aterial.-Lectotype, here selected, UMMP 

14378; illustrated paralectotypes, UMMP 14379, 

14380. 

Dispbyllum 1 slummi (Oliver) 

Plate 44; plate 45, figs. 1-7. 

1971 Cylindrophyllum stummi Oliver, p. 195-196, pl. 1 figs. 

1-6, pl. 2, figs. 4, 5. 

Occurrence of type material.-Famine Limestone, 

St. George, Quebec. Middle Devonian. 

Diagnosis.-Phaceloid coralla with cylindrical 

corallites commonly in lateral contact with each 

other so as to form short "chains." Septa short, 

wedge shaped, radi,ally arranged; minor septa nearly 

equal to major septa in length; septal carinae present or very common. Dissepimentarium narrow with 

one to four rows of normal, globose dissepiments. 

Tabulae complete or incomplete, variable in form. 

ExteTnal features.-Phaceloid (partly cateniform) colonies are as much as 17 em or more in 

height and 30 em or more in diameter, composed of 

cylindrical corallites 6 to 11 mm in diameter; mean 

diameter in four coralla ranges from 7.3 to 9.2 mm 

(table 36 and fig. 28). Corallites are virtually parallel in astogenetically advanced parts of corallum, 

radiating in early stages. Corallite spacing was apparently less than one diameter and many corallites 

grew in lateral contact with one or two neighboring 

corallites to form short "chains" (pl. 44, fig. 3). 

Average or maximum distance between corallites is 

unknown, as all specimens are from a structurally 

disturbed area and some crushing and distortion is 

evident in parts of most coralla. Offsetting is lateral. 

Internal features.-The major septa are radially 

arranged and very short, commonly extending from 

0.2 to 0.5 the distance to the axis; minor septa are 

nearly as long. Major septa number 19 to 27 (average 22.6 to 24.7 in four coralla) in cylindrical parts 

of studied corallites (table 36, and fig. 28) . Septa 

are wedge shaped, somewhat thickened peripherally. 

Microstructure poorly preserved except locally 

where septa seem to be composed of coarse and 

medium-sized trabeculae (pl. 44, fig. 1). Zigzag 

carinae are inconspicuous in most transverse sections but are more prominent in parts of most longitudinal sections. Locally, septa expand at their axial 

ends; in such places the coarse trabecular structure 

of the septa is particularly clear (pl. 44, figs. 3, 5). 

The dissepimentarium is narrow, commonly occupying less than three-tenths the corallite radius. 

Dissepiments are normal and commonly globose; 

arranged in one to four rows. Inner dissepiments 

where present are as large or larger than peripheral 

ones. Size and shape variation is considerable as can 

be seen from the illustrations. 

The tabularium is wide and composed of complete 

tabulae supplemented by incomplete peripheral tabulae. Tabula shape varies from nearly flat to irregularly but gently convex or concave. 

Discussion.-Disphyllum? stummi is characterized by its very short, relatively thick septa. It is 

analagous to species of Cylindrophyllum described 

here but these have longer, very attenuate septa and 

more prominent septal carinae. In transverse section, C. stummi is similar to Disphyllum? crassiseptatum (Ehlers and Stumm) (1949, p. 28-29, pl. 

6, figs. 1-6), but D.? cra.ssiseptatum differs in having very coarse septal trabeculae and a peripheral 

row of dissepiments that are larger than any inner 

dissepiments. D.? rectiseptatum (Rominger) is 

much smaller in diameter and has fewer rows of 

dissepiments. 

Distribution.-D. ? stummi is known only from the 

Famine Limestone, St. George, Quebec, and is of 

Onesquethaw or early Cazenovia age. (See discussion 

in chapter on stratigraphy.) 

Material.-Holotype, USNM 163236; paratypes 

il1ustrated by Oliver (1971), GSC 24639 and USNM 

1H3237; additional paratypes illustrated here, GSC 

2L1797-98 and USNM 163237, 163398; unillustrated 

paratypes, GSC 24640 and two others, USNM 

1()3396-97, and additional topotype fragments in 

both collections. 

Family ZAPHRENTIDAE Milne-Edwards and Haime, 1850 

Tentatively, I am accepting the commonly assumed relationship between Zaphrentis and H eliophyllum but separating both from Cyathophyllum 

and the Cyathophyllidae. I consider the Zaphrentidae to consist of predominantly solitary forms but 

recognize that H eliophyllum, at least, includes small 

colonies within its type species (H. halli MilneEdwards and Haime, 1850). I am also including 

Eastern North American colonial cyathophylloid 

species in H eliophyllum and in the genus Cyatho-
114 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

cylindrium Oliver but with some reservations, as 

noted in the following paragraphs. 

There are striking similarities between Cyathophyllum and some forms that I assign to the Cylindrophyllinae or to the Zaphrentidae. Indeed, my 

present interpretation of relationships has required 

shifting of several species from one family to another and some of both families have, at one time or 

another, been assigned to Cyathophyllum or the 

Cyathophyllidae. 

Hill (1956b, p. F278), Birenheide (1963, p. 368, 

404), and Jell and Hill (1969, p. 3) included 

Zaphrentis, Heliophyllum, and Cyathophyllum in the 

same family (Zaphrentidae or Cyathophyllidae). 

The type (C. dianthus Goldfuss) and other species of 

Cyathophyllum from the Eifel region, Germany, 

have been redescribed by Birenheide ( 1963). My 

interpretation of Cyathophyllum used in the following comparisons and analysis are based on (1) Birenheide's published work, (2) an examination of 

Birenheide's material in the Seckenberg Museum, 

Frankfurt-am-Main, and ( 3) preparation and study 

in my laboratory of specimens of colonial Cyathophyllum from the Eifel area. The Middle Devonian 

Eifel species form a distinct group and Birenheide's 

study of them gives a good basis for understanding 

the genus, if not necessarily the Family Cyathophyllidae. 

Cyathophyllum can be diagnosed as follows: Solitary or colonial corals with numerous biradially 

arranged septa, long or variously withdrawn from 

the axis, smooth to strongly marked with zigzag 

carinae; cardinal and counter septa commonly 

shorter than other major septa. Tabulae arched, flat, 

or sagging axially, complete or incomplete and tending to be more complete in species with withdrawn 

septa. Dissepimentarium wide, composed of small to 

medium-sized, globose dissepiments. 

Birenheide (1963) separated the Eifel species of 

Cyathophyllum into two subgenera. C. ( Cyathophyllum) has major septa not extending to the axis and 

more or less complete tabulae; C. (Peripaedium) 

has major septa extending to the axis and incomplete 

tabulae. Comparable morphologic types are represented in the Onesquethaw Stage in Cyathocylindrium. 

The type species of both Zaphrentis (Z. phrygia 

Rafinesque and Clifford, 1820) and Heliophyllum 

(H. halli Milne-Edwards and Haime, 1850) are 

basically solitary although H. halli certainly includes 

small colonies. Stumm (1949, p. 17) suggested that 

Heliophyllum (with wide dissepimentarium) 

evolved from Zaphrentis (no dissepiments) and 

assigned intermediate forms to his genus H eliophylloides (narrow dissepimentarium). Schindewolf 

(1938, p. 445) and Smith (1942, p. 341) had already 

suggested the derivation of H eliophyllum from 

Zaphrentis but they were interpreting Zaphrentis 

as having a narrow dissepimentarium ( =Heliophylloides of Stumm). Lecompte (1952, p. 475) and Hill 

(1956b, p. F278) both accepted this derivation while 

interpreting Zaphrentis as having a..t least some dissepiments. Oliver (1960b, p. 17-18) and Stumm 

(1965, p. 34-35) have since redescribed Zaphrentis 

phrygia on the basis of topotype collections. Oliver 

described Zaphrentis as having a very narrow dissepimentarium, possibly limited to the ephebic stage, 

whereas Stumm again described the genus as lacking 

dissepiments. Stumm selected a neotype but the 

specimen was not sectioned. 

Both Zaphrentis and H eliophyllum are characterized by a bilateral (subpinnate) septal arrangement 

and by the presence of strong yard-arm septal 

carinae. Whether or not Zaphrentis has a dissepimentarium, there seem to be gradations from very 

broad to very narrow dissepimentaria within the 

Zaphrentis-H eliophyllum group. In addition, Zaphrentis and solitary (and some colonial) Heliophyllum have strongly dilated dilated septa in early ontogenetic stages that may completely fill the lumen; 

septa become attenuate in later stages with the 

attenuation appearing at the periphery and migrating toward the axis. Commonly the dilation emphasizes the bilaterality of the corals, as in any given 

section dilation may be stronger in two of the quadrants than in the others. The ontogenies of Zaphrentis and most species of Heliophyllum are unknown. 

Within Eastern North America, H eliophyllum 

ranges from the middle Onesquethaw (Bois Blanc 

and equivalent; Emsian) to the Taghanic Stage 

(late Givetian). The range of Zaphrentis is not 

known but the genus may be restricted to the upper 

Onesquethaw (early Eifelian). This throws doubt 

on the evolutionary sequence Zaphrentis~H eliophyllum as postulated, but doesn't affect the morphological similarity. In spite of confusion and lack 

of information on many aspects of the genera, a 

close relationship between Zaphrentis and H eliophyllum still seems likely and at present it seems best 

to assign them to the same family (Zaphrentidae). 

Cyathophyllum appeared earlier than H eliophyllum (Jell and Hill, 1969, p. 3-4) so, if one accepts 

the Zaphrentis~H eliophyllum evolution, a relation-
SYSTEMATIC PALEONTOLOGY 115 

ship between Cyathophyllum and H eliophyllum is 

unlikely since all similarities are with the mature 

stage of H eliophyllum. 

Cyathophyllum and H eliophyllum are similar in 

adult morphology and in microstructure. H eliophyllum is more bilateral in septal arrangement and is 

characterized by long yard-arm carinae and by extreme septal dilation in early stages. Cyathophyllum 

is less bilateral, has short zigzag carinae and less 

globose dissepiments and apparently lacks the septal 

dilation. These differences, coupled with the available distribution data, suggest that the genera are 

not closely enough related to justify placing them in 

the same family. If the postulated relationship of 

H eliophyllum to Zaphrentis is accepted, the Cyathophyllum-H eliophyllum relationship is even less tenable. I would speculate that Cyathophyllum evolved 

from a Silurian coral with a wide dissepimentarium 

(such as Entelophyllum), whereas Heliophyllum 

evolved in eastern North America from either a 

zaphrentoid ancestor or a lykophyllid, possibly 

Phaulactis, with which H eliophyllum has much in 

common. 

If Cyathophyllum and Heliophyllum are unrelated, 

then what are the affinities of the numerous large, 

heavily carinate, bilateral colonial corals in the 

Onesquethaw Stage of eastern North America? With 

some hesitation I divide them into two genera mainly 

on the basis of their carinae. Some have extremely 

long, H eliophyllum-like carinae and may (or may 

not) show septal dilation. These are assigned to 

H eliophyllum although the lack of dilation in some 

forms makes this questionable. The second group, 

composed of forms with short zigzag carinae are 

included in Cyathocylindrium. These show no septal 

dilation. 

Specimens in both groups consistently differ from 

Cyathophyllum in having a small but well-developed 

fossula in the calice, in having more bilateral septal 

arrangement, and in having more globose and generally larger dissepiments. Specimens and species 

show considerable variation in these characters and 

on an individual basis the differences are often 

slight. However, the Onesquethaw complex of 

cyathophylloid individuals and species are clearly 

separable from Eifel Cyathophyllum on these characters, and I take this to indicate a long period of 

separate development and probably an independent 

origin, as suggested above. 

The separation of Cyathocylindrium from the 

Craspedophyllidae is also a problem. Phaceloid colonies of both are similar in many morphologic details. Craspedophyllids tend to be smaller, to have a 

more radial septal arrangement, and to have a characteristic mature diameter. Cyathocylindrium tend 

to be larger with greater size irregularities within 

colonies and to be more bilateral in septal arrangement. Septal microstructure and the shape and arrangement of tabulae and dissepiments are very 

shnilar. I consider the similarities due to convergence because the group can best be interpreted as 

having independent origins and because most craspedophyllids are very different from most Cyathocylindrium. It is possible that Cyathocylindrium 

represents a craspedophyllid "end point" of convergence with H eliophyllum, but at present an evotionary relationship with H eliophyllum seems 

more likely. 

In considering possible relationships of the 

Onesquethaw colonial corals, I have studied typical 

H eliophyllum, both solitary and colonial, in some 

detail. However, I have not considered other solitary 

genera that have been assigned to the same family 

as Heliophyllum by other worke·rs, and I am not 

ready to define or further discuss the family at this 

time. 

Ontogeny and increase.-Increase in the H eliophyllum and Cyathocylindrium species described 

below follows a common pattern for each growth 

form with only minor deviations in individual 

species. The following descriptions are made to avoid 

repetition. The terminology used is that of Hill 

(1H35, p. 490-495). In the studied zaphrentids, 

phaceloid and dendroid forms display either peripheral or lateral increase; cerioid and astreoid forms 

increase peripherally. In peripheral increase several 

offsets (new corallites) are formed on the peripheral platform of the protocorallite (parent). In all 

species here described, the offsets crowd together in 

suc.h a way that the parent could not have survived 

the reproductive stage (parricidal increase). In lateral increase a single (rarely two) offset projects 

laterally from the margin of the calice (nonparricidal increase) . 

Peripheral increase.-ln no example is the founder 

or initial corallite of the colony known. Presumably 

this individual was produced sexually, and its ontogeny may have differed significantly from that of 

subsequent, asexually produced individuals. 

In the species studied, from three to eight offsets 

I 

were formed on the horizontal or slightly inclined 

, calicular platform of the protocorallite. Earliest in-
116 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

dication of the reproductive phase is the formation 

of a partial or complete wall connecting the inner 

edges of the protosepta at the inner margin of the 

calicular platform (pl. 85, fig. 9; pl. 89, fig. 1). This 

wall was built by the offset polyp as is clearly indicated by the "dark line" on the side toward the 

parental axis (pl. 89, fig. 4). A few new short thin 

septa may have formed on this wall (pl. 89, fig. 1) 

or these may he the axial ends of parental septa. 

Subsequently the wall became curved to form the 

axial sides of the offsets (pl. 85, fig. 9). 

At their earliest recognizable stage, offsets had 

significant diameters (6-10 m,m) and numerous 

septa. New septa appeared on the new wall, and the 

parental septa were modified along the old (parental) wall. Commonly, there is no indication of the 

neanic or early brephic ontogenetic stages characteristic of solitary corals. Soon after their formation, 

offsets were in lateral contact with each other, in 

some coralla being sufficiently crowded to fill the 

parental calice and appear triangular in cross section (pl. 89, fig. 2). At diameters of 9 or 10 mm, 

offsets commonly had complete, encircling walls and 

were spread out sufficiently to be free on all sides. 

At this stage they were miniature adults except that 

septa were still being introduced (the insertion positions can be recognized in many individuals (pl. 85, 

fig. 9). Major and minor septa were inserted 

throughout the conical stage of each corallite; the 

insertion frequency decreased as cylindrical growth 

was assumed. 

Peripheral increase in these species is similar to 

that described and illustrated by Smith and Tremberth (1927) for Entelophyllum articulatum. 

Lateral increase.-This description should be 

compared with that of the craspedophyllids which 

commonly reproduced in a similar way although individuals are much smaller. In no colony studied was 

the founder corallite recognized. Its morphology may 

be somewhat different from that of the subsequent 

asexually reproduced offsets. 

In lateral increase, one (rarely two) offset was 

formed at one time. In serial sections taken at right 

angles to the axis of the protocorallite, the earliest 

indication of budding is a bulge in the corallite wall; 

the protosepta extend into the bulge, and there is no 

indication of neotissue. In succeeding sections, offset septa were first differentiated on the side away 

from the protocorallite. As an incomplete wall gradually developed between offset and parent, new septa 

were differentiated on the side toward the parent. 

The wall remained incomplete until the offset was 

virtually free of the parent. Many septa are present 

at the earliest recognizable stage of the offset and 

characters are virtually adult except in size and 

number. 

Microstructure.-The fine structure of septa and 

walls in many of the zaphrentid species described below is similar to that of the craspedophyllids. This 

is particularly so where septa and walls are thin. 

Microscopically, the wall appears to be composed of 

an inner, light-colored zone of minute fibers of calcite radially oriented in transverse section and an 

outer, very thin, dark zone of uncertain structure. In 

longitudinal section the fibers are seen to incline upward toward the axis, at approximately the same 

angle as the top surface of each dissepiment, so that 

in some places the dissepiments seem to be structurally continuous with the epitheca. More generally, 

the dissepiments, although structurally similar, are 

separated from the epitheca by a discontinuity. The 

tabulae are commonly very thin and their microstructure is obscure. 

Septa are composed of parallel, fine, closely spaced 

monacanthine trabeculae. Where the septal microstructure is well preserved it appears as follows. A 

dark central area, approximately 0.03 mm thick, 

indents the epitheca and extends the full length of 

the septum. This is apparently composed of closely 

spaced "centers of calcification." On either side of 

the dark central zone is ·a light-colored zone composed of minute fibers directed away from the central zone and inclined slightly toward the axis of 

the corallite, giving a pinnate appearance in transverse section. Septa are separated from the wall by 

a dark line, but the light-color.ed zone is identical to 

the inner wall in texture. Septa are thickest at their 

peripheral ends; the light-colored tissue thins toward 

the inner part of the tabularium. Carinae form 

ridges on the sides of the septa and show the same 

structure as the septa in transverse section. Carinae 

and parts of septa between carinae are composed of 

several parallel monacanths. 

The size of the monacanths varies in some of the 

zaphrentids, so that a single carina or part of a septum between carinae may be form.ed by as few as 

two or three monacanths. In some H eliophyllum with 

dilated septa, carinae may be thickened, triangular 

or diamond shaped in cross section and formed by a 

single coarse monacanth. This is especially characteristic of H eliophyllum halli, less so of the colonial 

species. But in all of these, very .fine monacanths ~are 

the most common structural elem,ent. If medium or 

coa~se monacanths are present in an individual 
SYSTEMATIC PALE~ONTOLOGY 117 

there ,is a complete gradation ,from very fine to medium or very fine to coarse. 

Distribution.-Colonial H eliophyllum range from 

middle Onesquethaw (Emsian) to Taghanic (late 

Givetian) in ea·stern North America. Cyathocylindrium may be restricted to the middle and late 

Onesquethaw (Emsian-Eifelian). Possible occurrences elsewhere have not been studied but are apparently uncommon. Solitary H eliophyllum are 

known from North Africa, Spain, and Venezuela 

and many Old World areas as w·ell. 

Genus CYATHOCYLINDRIUM Oliver, 1974 

1974 CyathocylindTium Oliver, p. 172. 

Type species.-Cyathocylindrium opulens Oliver, 

197 4, Onondaga Limestone, Edgecliff 1bioherm facies, 

New York. 

Diagnosis.-Dendroid or phaceloid rugose corals. 

Septa num·erous, biradially or bilaterally arranged, 

long or variously withdrawn from the axis; attenuate and weakly to strongly carinate (zigzag) ; 

cardinal and counter septa com·monly shorter than 

other major septa. Wide dissepimentarium is composed of many rows of very globose dissepiments. 

Tabulae arched, complete or .incomplete, tending to 

be ·more complete in for·ms with ·shorter major septa. 

Description.-Colonial, phaceloid, or dendroid 

colonies composed of cylindrical or ceratoid individuals; corallites tend to be rugose and may or may 

not be in lateral contact at points of maximum expansion. Increase .is lateral or peripheral. Calices are 

moderately deep with broad peripheral platforms. 

Septa are biradially or bilaterally arranged although this is not obvious in species with short 

septa. Cardinal and counter septa are short; other 

major septa tend to become longer away from the 

cardinal-counter plane; the counter septum may or 

may not be flanked by long minor septa. 

Major septa may reach the axis and meet ~along 

the cardinal-counter plane or may be withdrawn 

from the axis. All septa are attenuate and carinate. 

Oarinae .are zigzag to subyard-arm but the former 

predominate; they may be weakly to strongly developed but commonly are relatively short. 

Dissepimentaria are wide, compos·ed of many rows 

of very globose dissepiments. Ta;bulae are complete, 

arched but axially depressed or concave in species 

witht major s·epta that do not re,ach the axis.; incomplete in species with very long major septa. 

Microstructure.-As for the family and the Craspedophyllidae. 

Discussion.-This genus is established for •species, 

formerly referred to Cylindrophyllum or H eliophyllu?n, but t.hat differ from species of the former in 

their larger size and decidedly more pronounced 

bilateral septal arrangement. In both of these characters, Cyathocylindrium is close to Heliophyllum. 

However, the latter differs in having predominant 

long, yard-arm carinae and dilated early-stage septa. 

Cyathocylindrium is most similar to Cyathophyllum 

from which it diff.ers primarily in its more globose 

dissepiments and more variable carinae. Possible 

relationships and reasons .for placing it in the 

Zaphrentidae are discussed under the fam·ily heading. 

Three Onesquethaw species are here referred to 

Cyathocy lindrium. 

1. C. opulens Oliver; Edgecliff Member, bioherm 

facies, Ni,agara Peninsula, Ontario, and ~eastern and western New York; non bioherm 

facies, Niagara Peninsula; Jeffersonville Limestone, Zone B ( ?) , Falls of the Ohio; upper ( ?) 

Columbus Lim·estone, northeastern Ohio. 

2. C. gemmatum (Hall) ; Edgecliff mem1ber, bioherm facies, Niagara Peninsula, Ontario, eastern and western N.ew York. 

3. C. n. sp. A; Moorehouse Member, western New 

York. 

Cyathocylindrium opulens Oliver, 1974 

Plates 80-86 

? 1874 Heliophyllum pTolije?'Um Nicholson, p. 59. (not 

Cyathophyllum pTolijeTum Roemer, 1855, p. 

29, or C. proliferum Dybowski, 1874, p. 445-

447. 

? 1874 H. p?·olijeTum Nicholson, p. 27-28. 

not 1877 H. prolife<t·urm Hall, pl. 26, figs. 1, 2, 5 (junior 

homonym of Nicholson's name). 

? 1882 H. late1·icrescens Hall, 49-50. 

? 1883a H. late1·icTescens Hall. Hall, p. 314, pl. 27, fig. 1. 

not 1938 H. proliferum Nicholson. Stewart, p. 38-39, pl. 7, 

fig. 6. 

pad? 1965 H. latericTescens Hall. Stumm, p. 37, ?pl. 29, fig. 

8 (not? fig. 7 or pl. 30, fig. 13. 

1974 CyathocylindTium opulens Oliver, p. 172-174, 

figs. 5a-e. 

Occu-rrence of holotype.-Onondaga Limestone, 

Edgecliff Member, bioherm facies, Thompson's Lake 

bioherm (loc. 81), 15 miles (24 km) west o.f Albany, 

N.Y. 

Diagnosis.-Phaceloid and dendroid Cyathocylindrium in which the septal arrangement is strongly 

bilateral; major septa extend to the cardinal-counter 

plane which is defined by short cardinal .and counter 

septa. Tabulae are incomplete. Col}allites are only 

rarely in contact. 
118 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

External features.-Coralla are phaceloid or dendroid and composed of subparallel or radiating corallites which are -rarely in contact except at their lower 35 

ends; there are no lateral supports. Colony heights 

and dia,meters in ~excess of 60 em are not uncommon. 

Two silicified parts of colonies (pls. 80 and 81) 

measure 13 by 15 by 18 and 18 by 13 by 12 em in c:: 30 

height and diam,eters; these include 35 and 20 or 

more corallites respectively. In both of these specimens, corallites are in living position relative to each 

other, but both rare incomplete and may be parts of 25 

much larger colonies. Other specimens studied are 

fragm·ents of eoralla eonsisting of a few to several 

dozen corallites. Colonies are commonly preserved 

with corallites in parallel arrangement hut pressed 

together and broken at their points orf origin. 

Corallites are trochoid or eeratoid becoming cylindric,al and attaining diameters of from 10 to 32 mm 

(table 37 and fig. 34) . Corallites are coarsely and 

more or less regularly rugose at intervals of a few to 

10 mm, averaging 6 to 8 mm. Finer encircling 

growth lines and longitudinal septal grooves are also 

present. 

Increase is late~al and peripheral. Of 33 colonies 

in which increase was observed, 24 offset laterally, 

8 offset peripherally and 1 was dominantly lateral 

21 

10 15 20 25 

d(mm) 

EXPLANATION 

N 

0 USNM 172198 17 

+ NYSM 12845 6 

D USNM 172207 6 

- NYSM 12847 9 

X USNM 172204 20 

• USNM 172201 7 

30 

Mode of 

increase 

L 

L 

L 

L 

p 

L 

with a single peripheral offset. Mode of increase was 2.2 r--~----,-------.-------.---, 

not determined in s~everal additional specimens. 1 1 1 I 

EXPLANATION 

x Lateral increase 

o Peripheral increase 

®©N<4 

I I 

o© ® I 

35 t- A X - 0 

~ X 

0 ® 0 ® 

® - •c:: 30- X 

X ® 

0 X X 

® X - 25 1-

I I I I 

15 20 25 30 

d (mm) 

FIGURE 34.-Cyathocylin&rium opulens data. A, Mean number of major septa (n) versus mean diameter (d). Data is 

from 21 coralla but sample size (N) is very small in several (see text discussion). B, Number of major septa (n) 

versus diameter (d) in six selected colonies. Sample size 

(N) and mode of increase (P, peripheral; L, lateral) are 

given. C, Mean septal ratio (n/d) versus mean diameter (d) 

showing decrease in ratio with increasing diameter. 

2.0-

1.8 f1'1::> 1.6 -

I~ 

1.4 f1.2 -

1.0 f12 

X 

c 

0 

0 

I 

15 

EXPLANATION 

x Lateral increase 

o Peripheral increase 

@ ® N < 4 

0 

@< 0@ 

X ® 0 

X 

X X® 

I I J 

20 25 30 

d (mm) 

-

-

-

-

-

-

Commonly only one lrateral offset is formed at a 

time in this s'pecies although examples of two contemporaneous offsets are known. Peripheral offsets 

commonly number three to six. Initial offset diam-
SYSTEMATIC PALEONTOLOGY 119 

TABLE 37.-Cyathocylindrium opulens, n. sp., summary data 

Intracolony Inter- Mini- Maxi- Mean colony mum mum 

N ---------- 4 20 7.5 13 n: x ------- 25.3 35.8 30.5 

OR ----- 23-27 32-41 ---------- s ------- .00 4.58 3.63 

v ------ .0 14.4 7.5 11.9 

d: x ------- 14.0 26.7 22.5 

OR ----- 11-17 19-32 ---------- s ------- .96 5.77 4.40 

v ------ 3.6 24.5 13.7 19.6 

eters range from 8 to 15 mm; offsets expand to 

mature diameters in 20 to 60 mm. Offsetting intervals of 5 to 8 em were observed although they are 

commonly .much greater than this. 

Calices ar·e moderately deep (1average one-half 

corallite diameter), having gently sloping walls and 

broad peripheral platform.s that commonly .slope .inward hut may ·be horizontal or reflexed in .some individuals. A cardinal fossula is formed by ,a short 

cardinal septum. The calice floor is flat. 

Internal features.-Septa are bilaterally arranged. 

The cardinal and counter septa ar.e commonly 

shorter than the adjacent major septa and one of 

these may be shorter than the other. In some sections 

the counter septum is flanked by long minor .septa 

aMJhough other sections of the same corallite may not 

show this feature. Com·monly the cardinal and 

counter septa are of approximately ·equal length and 

not individualized by the arrangement of other 

s·epta; a cardinal-counter plane is clearly defined, 

however. Adjacent major septa turn toward the 

symmetry axis at their axial ends and are longer 

toward the 90° positions, so that the ·major septa 

tend to meet along ~the cardinal-counter plane but 

not at a point. Alar septa can be recognized only in 

some early (conical) stage·s where new septa are 

being .introduced. 

The length of the minor septa varies with position 

but is commonly one-half to two-thirds the length 

of the adjac·ent major septa. Major and minor septa 

are slightly thickened in the inner dissepi.mentari urn 

and the outer tabularium. 

Observed numhers of major .septa are shown in 

table 37 and on the graphs (fig. 34). The value of 

colony m·eans is limited ·by s.mall sam.ple size and by 

the difficulty of elim'inating the effects of ontogeny. 

S.ections through cylindrical parts of corallites range 

in diameter from 10 to 32 m.m ; colony means range 

from 14 to 27 mm. Numbers of major septa range 

from 23 to 41; colony .means range from 25 to 36. 

Carinae abound; they are predominantly zigzag 

but are locally yard-arm or nearly so. Length and 

spacing of carinae varies and can best be judged by 

reference to the illustrations. Carinae are limited to 

the dissepimentarium and are peripherally steep, 

flattening at their axi,al ends. 

The dissepimentari urn consists of from 7 to 12 or 

more rows of globose dissepiments ranging in height 

Ifrom 0.5 to 2.5 or more millimeters. Some dissepiments are elongate parallel to the carinae; near the 

inner margin of the dissepimentarium they tend to 

bE! less globose and may be elongate in the other 

direction. 

The tabularium is clearly separated from the dissepimentarium. Tabulae are incomplete, short, 

arched plates, forming an arched or double-arched 

pattern. 

Microstructure.-See under family description. 

Individual variation.-Both intercolony and intracolony vadation are extensive in C. opulens as in the 

other species described. Variability of many structures is discussed in the above description. Figure 

34 provides additional data !but must be interpreted 

carefully. Figure 34A shows observed intercolony 

size and septal number variation, but the samples on 

which it is based are small. Tw.enty-one colonies are 

represented by from 1 to 20 corallites. Because of the 

large corallite size it was impractical to collect or 

sHction adequate numbers of individuals in all colonies. N.evertheless, it seems better to present only 

one value per colony on this graph rather than to 

show a much larger number of corallites, but with 

most from only a few coralla. T~able 37 summarizes 

data from 13 colonies (N =4-20). 

Figure 34B shows the data for 6 to 20 corallites in 

each of .six colonies. Colonies were selected to show 

rnaximum size and septal number differences between colonies. .Comparison of figures 34A and B 

will show how other coralla tend to occupy the 

middle area. 

Septal ratio variation is similar to that noted in 

other species in which there is adequate data. Figure 

34C illustrates the decrease of septal ratio with increasing diameter. The negative correlation is significant at the 0.001 level (P<0.001). 

An example of discontinuous variation in C. 

opulens is found in the presence of two distinct 

n1odes of colony development. Of 33 specimens in 

which offsetting was determined, 25 offset laterally 

and 8 offset peripherally .In one of the lateral coralla, 

a single peripheral offset was formed at the same 

time as a lateral offset (pl. 84, fig. 6) ; the parent 

corallite was itself offset laterally. The single peripheral offset attained a length of only a few millimeters and a diameter of 10 mm. This is the only ob-
120 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

served example of the occurrence of both types of increase within a single colony in the Onesquethaw 

fauna. In none of the coralla seemingly characterized 

by peripheral offsetting were fewer than three offsets noted for any one reproductive phase. 

In respect to mode of offsetting, C. opulens is 

dimorphic, although on a colony rather than individual level. Such polymorphism is commonly g·enetically controlled and the 25 :8 ratio suggests that 

mode of offsetting may have been a case of simple 

Mendelian inheritance with lateral offsetting dominant over peripheral offsetting and the forms appearing in a 3 :1 ratio. 

Discussion.-The preceding description is based 

on 39 coralla or parts of coralla; other fragments 

provided additional distribution data but have not 

been otherwise used; all are from the Edgecliff 

Member in New York and southwestern Ontario. 

In the field, C. opulens can be confused with only 

one other Onondaga species, C. gemmatum. It differs 

from C. gemmatum in its long septa, strong bilateral 

septal arrangement, incomplete tabulae, and infrequent contacts between corallites. From all other 

Onesquethaw species it differs markedly in corallite 

size or corallum form. 

Cyathocylindrium opulens is probably the same as 

Heliophyllum. proliferum Nicholson, 1874 (not H. 

proliferum Hall, 1877), but this form was never 

illustrated, and no type specimens are known. 

Nicholson's species is "extraordinarly abundant in 

one bed of the Corniferous Limestone of Ridgeway" 

(Nicholson, 1874, p. 28). This locality is some 10 

miles (16 km) east of Port Colborne, Ontario, and 

might be within either the Bois Blanc or lower 

Onondaga. "Extraordinarily abundant," however, 

can only apply to the Edgecliff bioherm facies and 

probably to C. opulens. 

H eliophyllum proliferum Hall ( 1877, not Nicholson), from the Hamilton Group in New York, differs from C. opulens in initial offset growth nearly · 

at right angles to the parent corallite and in abundant and prominent yard-arm carinae. 

H eliophyllum latericrescens Hall, 1882, from the 

Jeffersonville Limestone at the Falls of the Ohio, is a 

Cyathocylindrium similar to C. opulens. Stumm, 

1965 (p. 37) placed several Falls of the Ohio 

"species" of Greene (1898-1906) in synonymy with 

Hall's species but all of these are represented by 

fragmental, poorly preserved, and unrecognizable 

holotypes. Hall's holotype is not well preserved but 

has between 45 and 50 major septa and is probably 

not conspecific with C. opulens. C. opulens does occur 

at the Falls as discussed under distribution below, 

but no previously described Falls "species" can be 

so identified at present. 

Heliophyllum proliferum Nicholson of Stewart 

(1938) is a Hamilton form, possibly closer to H. 

proliferum Hall (not Nicholson) than to any Onesquethaw species. 

Cyb:ndrophyllum confiuens Stewart (1938), from 

Columbus Limestone, Zone C, in Ohio is poorly 

known. Stewart's "cotypes" include two or more 

species, one of which is similar to Cyathocylindrium 

opulens in having long septa and incomplete tabulae. 

But this form has a relatively small diameter (approximately 16 mm) with more septa (n=37, 38) 

than are known in the larger C. opulens. 

"H exagonaria" martinsoni Cranswick and Fritz 

differs from C. opulens in its somewhat larger corallite size and in its semicerioid habit at some growth 

stages. 

Distribution.-Cyathocylindrium opulens is the 

most common colonial zaphrentid known from the 

Onondaga. It is common in the bioherm facies of the 

Edgecliff Member, and specimens were collected 

from every known bioherm area from Port Colborne, 

Ontario, to Albany in eastern New York. 

Two NYSM specimens from "Leroy, New York" 

(pis. 80 and 81) are probably from an Edgecliff bioherm also. 

The species also occurs in the nonbioherm Edgecliff in the Niagara Peninsula (two specimens 

known). 

Two specimens were collected from the Jeffersonville Limestone at the Falls of the Ohio. Both were 

from the base of Zone B or the top of Zone A. 

One specimen from the upper part of the Columbus Limestone in the subsurface of northeastern 

Ohio is tentatively assigned to this species. 

llfaterial.-Edgecliff Member, New York and Ontario: Holotype, USNM 172197; illustrated or measured paratypes, USNM, 172198-208, NYSM 12843-

48, GSC 31159; additional sectioned paratypes, 

USNM 172211, 182846-53; NYSM 12849-50; GSC 

31160. Identified specimens: Jeffersonville Limestone, Zone A or B, USNM 172209-210. Upper 

Columbus Limestone, Ashtabula County, Ohio, 

UMMP 60485. 

Cyathocylindrium gemmatum (Hall) 

Plates 87-90 

1882 Heliophyllum gemmatum Hall, p. 49. 

1883a H. gemmatum Hall. Hall, p. 310, pl. 26, fig. 12. 

1884 H. gemma.tum Hall. Hall, p. 453. 

? 1887 Cyathophyllum oedipus Davis, pl. 84, fig. 1 

(probably not pl. 83, figs. 6-9, which are unrecognizable) . 
SYSTEMATIC PALEONTOLOGY 121 

part 1965 Cylindrophyllum compactum (Hall), Stumm p. 

42 (not H eliophyllum com pactum Hall, or most 

of Stumm's synonymy or any of the illustrations; see discussion) . 

Holotype.-The original of Hall's only illustration, 

FMNH 35872WM. Jeffersonville Limestone (position unknown) , Falls of the Ohio. 

Diagnosis.-Cyathocylindrium in which the septal 

arrang,ement is bilateral to biradial but the septa 

extend only 0.8 to 0.9 the distance to the axial plane, 

leaving an elliptical open space around the axis. The 

septa are thin; tabulae are complete and incomplete. 

Corallites are in lateral contact at several points, 

usually at expanded rugae. 

External features.-N o complete colonies are 

known; most available fragments are small, as much 

as 30 em high and 21 em in diameter and include 25 

to 100 or more corallites. The largest known specimen (pl. 87, figs. 1, 2) was found in a limestone 

block 16 em high and 95 by 75 em in diameter; the 

specimen is incomplete but apparently not crushed 

or compressed; corallites as much as 50 em in length 

are included. ·Colony form is phaceloid in the larger 

specimens. Individual corallites ar·e trochoid, become 

ceratoid or cylindrical, and reach diameters of 30 

mm or more (fig. 35). 

Corallites are strongly and more or less regularly 

rugose. There are no lateral supports, but individuals are closely spaced and the coarse rugae of 

adjacent coralJi.tes are in frequent contact. The restoration shown on plate 1 displays these characters 

very well. 

Increase is peripheral and parricidal, three to five 

offsets commonly being formed at the same time. 

Initial offset diameters orf less than 10 mm have been 

observed; offsets expand to their mature diameters 

in 40 to 60 mm. Offsetting intervals have not been 

observed. 

External markings consist of fine longitudinal 

septal grooves in addition to encircling rugae. 

Calices are moderately deep (average one-half the 

diameter) with gently inclined sides and a peripheral platform that is nearly flat or gently inclined 

toward the axis. A c·ardinal fossula is formed by a 

short cardinal septum. 

Internal features.-Septa are bilateral or biradial, 

but the major septa extend only from 0.8 to 0.9 the 

distance to the axis, leaving a central, elongate open 

area. The cardinal-counter plane is commonly well 

defined but primary septa are not individually recognizable except in some of the immature sections. 

Numbers of major septa relative to corallite diameters are shown in figure 35 for a sample of the 

available specimens. No statistics other than means 

have been calculated because of the small sample size 

and the impracticality of eliminating data from early 

ontogenetic stages. Diameters measured on cylindrical parts of corallites range from 12 to 28 mm; the 

corresponding range in numbers of major septa is 

22 to 44. Minor septa are from two-thirds to threefourths the length of adjacent major septa. 

Carinae are abundant in most corallites; they are 

mostly zigzag but are locally subyard-arm in appearance. Length and spacing of carinae can best be 

judged by reference to the illustrations. Carinae are 

limited to the dissepimentarium; they are steeply 

inclined peripherally, more gently so within the dissepimentarium, steeper at their axial ends. 

The dissepimentarium consists of from 5 to 10 or 

more rows of normal, globose dissepiments. Height 

of the spaces between dissepiments ranges from 0.6 

to 2.0 mm but is commonly ±1.0 mm. Some dissepimtental spaces are elongate parallel to the carinae. 

Tabulae are complete or nearly so and commonly 

horizontal in the axial par.t of the tabularium, but in45 

EX PLANATION 0 

X N =6-10 

0 N =3 0 

0 N =1 

40 

6.. Holotype ( N ::;;::.1) 

0 

' 35 

(~ 

0 ~ ' rh -o IL c 

Cll 

lc:: 

25 

20L-------~--------~--------~------~ 

10 15 20 25 30 

d and ORd (mm) 

FIGURE 35.-Cyathocylindrium gemmatum da'ta showing mean 

and observed range of number of major septa (n and ORn) 

versus mean and observed range of diameter ( d and ORd). 

Mean shown by symbols; observed ranges shown by vertical and horizontal lines through symbol. Sample size (N) 

shown by symbols. 
122 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

complete and arched peripherally, forming an overall concave pattern. 

Microstructure.-See family description. 

Discussion.-This description is based on 16 coralia or parts of coralla from the Edgecliff in New 

York and Ontario. 

In the field, C. gemmatum can be confused with 

C. opulens, which differs in having a more obviously 

bilateral septal arrangement, incomplete tabulae, 

and less frequent contacts between corallites. From 

all other Onondaga species, C. gemmatum differs 

markedly in corallite size and corallum form. 

I consider this Onondaga form to be conspecific 

with the single specimen described by Hall (1882, 

1883, 1884) as Heliophyllum gemmatum. Unfortunately, the species has not been found in any more 

recent Falls of the Ohio collections and the exact 

stratigraphic position of the holotype is not known. 

Stumm (1965, p. 42) redescribed H eliophyllum 

compactum Hall as Cylindrophyllum compactum and 

included H. gemmatum in the synonymy of that 

species along with five other Falls species of Hall, 

Davis, and Greene. H. compactum Hall and H. pocillatum Hall are based on silicified fragments of single 

corallites that may or m·ay not have been parts of 

colonies; I consider both species to be unrecognizable. H. fecundum Hall is based on a silicified cylindrical fragment having four peripheral offsets; this 

may be a Cyathocylindrium but is present unrecognizable on the species level. Cyathophyllu1n oedipus 

Davis may be a junior synonym of C. gemmatum but 

more detailed study is required to determine this. 

Cyathophyllum coralliferum Davis. and Heliophyllum 

gradatum Greene are probably Cylindrophyllum and 

are discussed under that genus. Of the species in 

Stumm's synonymy, H eliophyllum gemmatum Hall 

is the senior recognizable form. 

Distribution.-Edgecliff Member, New York and 

the Niagara Peninsula of Ontario. Thirteen of the 

16 specimens on which this description is based were 

collected from the Edgecliff bioherm facies at Williamsville, N.Y. (Fogelsanger quarry). Single coralia were collected from two bioherms in eastern 

New York and from the Edgecliff "green beds" at 

Port Colborne, Ontario. The holotype fragment is 

from the Jeffersonville Limestone at the Falls of the 

Ohio, stratigraphic position not known. Cyathophyllum oedipus Davis, which may be conspecific, is probably from the Middle Devonian coral zone (Zone B). 

Material.-Holotype: FMNH 35872WM. Illustrated or measured specimens : USNM 172212-217 

and NYSM 12860-63. Other specimens: USNM 

172218 and 172219 a-e. 

Cyathocylindrium n. sp. A 

Plate 91 

Diagnosis.-Phaceloid Cyathocylindrium with 

strongly rugose, ceratoid-cylindrical corallites that 

are commonly in lateral contact with other corallites 

at :positions orf maximum ·expansion; ·maximum corallite diameters as much as 50 mm. Septa long, 

bilaterally arranged; tabulae are complete and incomplete. 

External features.-Dimensions in centimeters 

and approximate number of corallites in the two 

specimens on which this description is based are 

tabulated 1below. As explained under remarks, they 

may be parts of the same colony. 

USNM 172220 ------

NYSM 12864 -------

Specimen.-

Height Length 

35 50 

25 15 

Width CQrallites 

10 50+ 

10 15+ 

Colony form is phaceloid, with large cylindrical 

corallites parallel or •slightly diverging; ceratoidcylindrical corallites range from 30 to 50 m·m or 

more in diameter. Ther·e are no lateral supports, but 

individuals are markedly rugose and in lateral contaot at points of maximum expansion. 

Increase is peripheral ( parric.idal) ; five and seven 

offsets were form·ed in the studied examples. New 

individuals expand and diverge rapidly, after which 

growth is approximately cylindrical and parallel. An 

offsetting interval of 12 em was observed in one 

specimen. 

Corallites are strongly and regularly rugose at 

intervals of from 5 to 15 m·m; finer encircling rugae 

and longitudinal septal grooves are also present. 

Calices as inferred from internal ·structure have 

moderately steep inner walls ·rising to an almost 

horizontal peripheral platform. Calices are relatively 

shallow and flat bottomed; cardinal fossula not 

prominent. 

Internal features.-Major septa are bilaterally 

arranged. The cardinal and counter septa are shortest; in some specimens one of these, presumably the 

counter, is flanked by extra-long minor septa; adj·acent major septa become longer toward the sides 

and extend to the cardinal-counter plane. Minor 

septa are approximately two-thirds as long as the 

adjacent major septa except as noted. Alar septa 

can be recognized only in early stages where new 

septa are being introduced. All septa are somewhat 

thickened in the inner part of the dissepimentarium. 

Number of major septa in five corallites ranges 

from 30 to 38; diameters range from 25 ·to 35 m·m. 

The measured specimens have larger individual 
SYSTEMATIC PALEONTOLOGY 123 

diameters and fewer major septa at a given diameter ( n/ d = 1.1) than the other species of Cyathocylindrium or H eliophyllum with which this species 

is most comparable. Cylindrical stage diameters 

range from 25 to 40 mm or more. 

Carinae are long or short, strongly or weakly 

developed, co-mmonly zigzag but subyard-arm occasionally, all within the same thin section. Carinae 

are limited to the dissepimentarium and are steeply 

inclined peripherally, gently inclined or nearly horizontal as they approach their inner limit. 

The dissepimentarium consists of many rows of 

normal dissepiments. These are mostly globose but 

the three or four inner rows are elongate and steeply 

inclined. The globose dissepiments are commonly 

medium size but may be as much as 2.5 mm in 

height. 

Tabulae are closely spaced, complete or incomplete, individually convex or concave but forming a 

horizontal or concave pattern. 

M icrostructure.-See family description. 

Discussion.-The preceding description is based 

on two specimens that were collected in different 

years from the same locality in the Moorehouse 

Member, western New York, and may be parts of 

the same colony. The specimens are of particular 

interest because they are the only large colonial 

corals known from the Moorehouse in New York 

or Ontario. 

C. n. sp. A differs from other Cyathocylindrium in 

the combination of characters noted. It has longer 

S'epta and larger size than C. gemmatum and has 

complete tabulae, unlike C. opulens. C. n. sp. A could 

have been derived from C. gemmatum by a lengthening of the major septa, an increase in diameter, and 

a decrease in distance between corallites. 

Distribution.-Known only from one locality in 

the Moorehouse Member of the Onondaga Limestone, 

western New York. 

Material.-IIlustrated specimens: USNM 172220 

and NYSM 12864. 

Genus HELIOPHYLLUM Hall 

1846 Cyathophyllum (Heliophyllum) Hall, in Dana, p. 183. 

1848 C. (Heliophyllum) Hall, in Dana, p. 356. 

1850 Heliophyllum Hall, Milne-Edwards and Haime, p. lxix. 

1940 Heliophyllum Hall, Lang, Smith, and Thomas, p. 66. 

1956b Heliophyllum Hall, Hill, p. F278. 

Type species.-Strombodes helianthoides? Hall, 

1843, (not Goldfuss, 1826) p. 209, fig. 87-3, and no. 

48, fig. 3, p. 44 of tables; renamed H eliophyllum 

halli by Milne-Edwards . and Haime, 1850, p. lxix. 

Ludlowville and Moscow Formations, Tioughnioga 

Stage, central and western New York. 

Diagnosis.-Solita.ry and colonial rugose corals, 

widely varying in shape and type of colony, characteristically having a moderately deep calice with 

broad axial pit, a steeply inclined lower calice wall, 

a broad horizontal or gently inclined peripheral platform, and a shallow fossula. 

'Major septa tend to be long and are bilaterally 

arranged about the cardinal-counter plane; mature 

s,epta may be attenuate, but septa are moderately 

to heavily dilated at least in early ontogenetic 

stages. Carinae are abundant and prominent, commonly long yard-arm but short zigzag carinae may 

also occur; they are commonly, but not necessarily, 

lin1ited to the dissepimentarium. 

Dissepiments are globose and in numerous rows. 

Tabulae are incomplete, commonly convex but forming a generally concave pattern. 

.Discussion.-The genus badly needs revision a~ 

its limits are poorly understood at present, even 

though some "populations" of H. halli have been 

admirably described (Wells, 1937). Of Onesquethaw 

colonial corals, I am including phaceloid, ceriod, 

and astreoid species characterized by long yard-arm 

carinae and, in some examples, by septal dilation. 

.Distribution.-H eliophyllum halli is common in 

upper Onesquethaw to Taghanic formations, and a 

Heliophyllum that may be H. halli is present but not 

cmnmon in the middle Onesquethaw Bois Blanc 

Formation. However, small colonies of H. halli are 

known only from the upper part of this range 

(Tioughnioga and Taghanic) and none are described 

in this paper. 

Colonial H eliophyllum described here are widespread in both middle and upper Onesquethaw formations, but most species are represented by few 

specimens. The following species are described from 

New York and Ontario : 

Phaceloid: 

1. H. megaproliferum n. sp., Edgecliff bioherm 

facies, New York and Niagara Peninsula, 

Ontario. 

2. H. n. sp. B, Edgecliff bioherm facies, eastern 

New York. 

3. H. n. sp. C, Edgecliff bioherm facies, eastern 

New York. 

Cerioid: 

·1. H. coalitum (Rominger), Edgecliff bioherm 

facies, New York; ?Ontario; Bois Formation ( ?) , Ontario and Michigan. 
124 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Astreoid: 

5. H. monticulum (Ehlers and Stumm), Edgecliff bioherm facies, New York and Niagara 

Peninsula, Ontario. 

6. H. cassum n. sp., Bois Blanc Formation, 

Niagara Peninsula and Woodstock area, 

Ontario. 

7. H. procellatum, n. sp., Bois Blanc Formation ( ?) , Falkirk, N.Y. 

The phaceloid species are types that have previously 

been referred to Cylindrophyllum or H eliophyllum, 

depending on source and specialist. The single 

cerioid species and similar forms were referred to 

Hexagonaria by Stumm (1948, 1955, 1965) because 

of the growth form with well-developed intercoralIite walls. The astreoid species have been assigned 

to Billingsastraea by recent workers solely on the 

basis of growth form. Differences betwee'n these 

species and analagous craspedophyllid forms are 

discussed in the description of families and genera 

and in some of the species comparisons. 

Phaceloid Species 

Heliophyllum megaproliferum n. sp. 

Plates 92, 93 

Occurrence of type mateTial.-Edgecliff Member, 

bioherm facies, Niagara Peninsula, Ontario, and 

eastern and western New York. 

Diagnosis.-Phaceloid coralla composed of large 

(25-45-mm diameter) ceratoid to cylindrical corallites that increase laterally. Septa are long and thin 

or dilated and have abundant long yardarm carinae. 

External features.-The holotype is a fragment 18 

em high and 11 by 32 em in diameter and includes 

17 corallites; the size of the complete colony is not 

known. Three other specimens are smaller than 

the holotype. The description is based on these four 

partial coralla. 

Individual corallites are ceratoid becoming cylindrical; m·easured cylindrical stage diameters 

range from 26 to 42 mm (fig. 36). Increase is 

lateral; after initial growth away from parent, 

corallites tend to become parallel;-- no lateral sunports are present and corallites are only incidentally 

in lateral contact with each other. 

Epitheca is marked by shallow septal grooves and 

broad, rounded interseptal ridges; corallites are 

regularly rugose. 

No calices have been observed, but the arrangement of structures in longitudinal seetions suggests. 

the following: diameter at b~se from one-third to 

two-fifths that of the corallite, depth 10 to 15 mm; 

lower sides nearly vertical but curving away to form 

a flat or gently inclined peripheral platform; low 

septal ridges may be present on floor and walls and 

a shallow cardinal fossula is probable. 

Internal features.-Major septa reach or nearly 

reach the axis and are biradially arranged. The 

cardinal-counter p~ane is defined by short cardinal 

and counter septa; adjacent major septa are arranged symmetrically about the plane; in some 

corallites, the presum~ed counter septum is flanked 

by extra-long minor septa. Number of major septa 

ranges from 26 to 41 (fig. 36). Minor septa are 

two-thirds as long as the major. All septa are thin 

but may thicken slightly in the inner part of the 

dissepimentarium. Carinae are well developed, commonly long, and of the yard-arm type; they are 

limited to the dissepimentarium and vary in inclination from nearly vertical at the periphery to nearly 

horizontal at their inner ends. 

The dissepimentarium is wide, occupying from 

three-fifths to two-thirds of the radius, and consisting of 12 o·r more rows of relatively small dissepiments. Dissepiments ·are globose and horizontal near 

the periphery, vertical and elongate near the tabularium, with intermediate shapes and orientations 

in between. 

Tabulae are incomplete; flat or gently curved 

plates form a flat or slightly convex pattern. Tabularia are sharply sep~rated from dissepimentaria 

and occupy from one-third to two-fifths the total 

diameter. 

Microstructure.-The specimens assigned to this 

snecies are silicified and microstructure is obscured. 

Dark and light layers can be recognized in some 

corallites, and the structure is apparently as described for the genus. 

Discussion.-The above description is based on 

four partial coralla, all collected from the Edgecliff 

bioherm facies. The two Ontario specimens are from 

the same locality and might have been part of the 

same colony although this seems unlikely. The other 

specimens are from Williamsville and Coxsackie, 

N.Y. Each of the specimens shows considerable 

intracolony variation, but they are very 'Similar to 

each other. The three western specimens are partly 

silicified but not in such a way that they can be 

etched from the enclosing rock. 

Within the Onondaga fauna, H. megaproliferum 

is most similar to Cyathocylindrium opulens and H. 

sp. B. C. opulens differs in its smaller corallite size, 

and in its irregular septa having predominantly zigzag carinae. H. sp. B differ'S in having peripheral offsets, large coarse dissepiments and markedly dilated 

septa. 

Distribution.-Known only from the bioherm 

facies of the Edgecliff Member in areas adjacent to 
SYSTEMATIC PALEONTOLOGY 125 

50 

45 

40 

35 

30 

25 

EXPLANATION 

Heliophyllum megaproliferum 

0 GSC 31161 

0 

15 

0 USNM 172221 

0 USNM 172222 

!::::. USNM 172223 

20 

H. n. sp. B 

0 USNM 172225 

0 USNM 172224 

& NY SM 12865 

H. n. sp C 

X USNM 172226 

• NYSM 12866 

e USNM 172227 

25 30 35 40 

d(mm) 

/e\ 

I \ 

I \ ~ I 

\._;_) 

----o----------- r6- t'&\ 0 o_·) ' .._,/ --- ------------------

45 50 55 

FIGURE 36.-Heliophyllum n. sp. B. H. megaproliferum, and· H. n. sp. C data showing number of major septa (n) versus 

diameter (d) in small samples of several colonies. Polygons enclose data for H. n. sp. C and for each corallum in the 

other species. The three unenclosed symbols in the lower left are offsets in the H. megaprolife·rum coralla indicated. Other 

data are from cylindr~cal (mature) stage. 

the Niagara Rhr·er in both Ontario ·and New York 

and in eastern New York. 

Material.-Holotype, GSC 31161; paratypes, 

USNM 172221-223 (specimen 172221 was collected 

at the type locality and may be a part of the bolotype). 

Heliophyllum n. sp. 8 

Plate 94 

Diagnosis.-Phaceloid coralla increasing peripherally, composed of large (as much as 55-mm 

diameter) cylindrical corallites. Septa a.re attenuate 

in the dissepimentarium and have short, zigzag 

c-arinae but are dilated in the tabularium. 

External features.-Three fragments of phaceloid coralla include two to five large corallites and 

appear to represent a distinct new species. Because 

of the inadequacy of the material, the species is 

described informally. Individual cor·allites are cylindrical, having diameters ranging as much a.s 55 

m1m (fig. 36); maximum observed length for a 

single corallite is 13 em. Increase is peripheral with 

five offsets in one observed example. Offsets expand 

within a few centimeters to their full diameters. 

There are no connections between corallites, except 

at poi~ts of origin. External markings are not 

known. 

Calices are inferred from orientation of dissepiments and carinae to be moderately deep (approxi-

·mately one-half corallite diameter) and have steep 

inner· walls and a broad peripheral platform. 

Internal features.-The septa are bilaterally a·rranged. Cardinal and counter septa are relatively 

short and the counter septum is flanked by extralong minor septa. Other major septa are mainly 

radial in arrangement; commonly they meander 

slightly in the tabularium and extend to or nearly 

to the axis, where some of them meet. Observed num-
126 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

bers of major septa range from 37 to 42 (fig. 36) . 

Minor septa, except the counter laterals, are twofifths to two-thirds the length of the majors. 

Septa are attenuate in the dissepimentarium but 

thicken in the tabularium by a factor of four or 

more. Abundant zigzag carinae are short; long 

yard-arm carinae are also present; considerable 

variation occurs within a single cross section of a 

corallite. Carinae are steep peripherally, gently inclined at their inner ends. 

The dissepimentari urn is formed by 8 to 15 rows 

o.f medium to larg,e dissepiments and occupies approximately two-thirds of the corallite radius. Outer 

rows ,are composed of large dissepiments that are 

nearly circular (2 mm hiwh) in longitudinal section; 

inner dissepiments are smaller and vertically elongate. 

The tabularium occupies one-third of the diameter 

of the corallite. Tabulae are incomplete; tabulae 

pattern in longitudinal sections is obscured by the 

axial ends of the major septa. 

Microstructure.-Septa are composed of a dark, 

granular-appearing mid-layer and light brown surface layers of fibers arranged perpendicular to the 

plane of the septum. Structure is particularly well 

preserved in dilated axial parts of the septa. 

Discussion.-The above description is based on 

three fragments of coralla, each including two to 

five large corallites. The specimens are worn and 

partly crushed but otherwise well preserved. The 

specimens vary in detail but are united by their 

large corallite size, septal dilation, and large dissepiments. In these characters, H. sp. B differs 

from all other phaceloid corals known from the 

Onondaga within the study area. 

H. n. sp. B is most similar to H. n. sp. C. The 

two species differ however in the form of their 

carinae, dissepiments, and calice: 

Carinae _______ _ 

Dissepiments __ _ 

Calice ---------

H. n. SJJ. B 

short, zigzag 

large, round 

broad platform 

H. n. sp. C 

long, yard· arm. 

medium large. 

no platform. 

It differs from the other H eliophyllum spp. in 

having predominantly zigzag carinae; long yardarm, H eliophyllum-type carinae are present but less 

common. 

Distribution.-Known only from the bioherm 

facies of the Edgecliff Member at one locality near 

Schoharie, in eastern New York. 

Material.-Illustrated specimens, USNM 172224, 

172225 and NYSM 12865. 

Heliophyllurn n. sp. C 

Plate 95 

Diagnosis.-Phaceloid coralla, increasing peripherally, composed of large (as much as 50-mm 

diameter) cylindrical corallites. Septa are dilated 

in the tabularium; long yardarm carinae are common in the dissepimentarium. 

Description.-Three coralla fragments are composed of two to eight ceratoid-cylindrical individuals 

and appear to represent a distinct new species. 

Corallite diameters average 50 mm; corallites have 

43 to 54 major septa (fig. 36). Increase is peripheral 

with eight offsets in one observed example. There 

are no connections between corallites except at 

points of origin. Calices are inferred to be moderately deep (approximately three-fourths corallite 

diameter) and have walls sloping from the corallite 

margin to the floor of the calice; peripheral platform absent or very narrow. 

Septa are bilaterally arranged. Cardinal and 

counter septa are relatively short and the counter 

septum is flanked by long minor septa. Other major 

septa extend to or nearly to the axis. Septa are 

attenuate in the dissepimentarium but thicken in 

the tabularium by a factor of three or four. Yardarm carinae are long and abundant; zigzag carinae 

are present but are estimated to represent less than 

10 percent of the carinae in any one transverse section. Carinae slope gently from the inner to outer 

margins of the dissepimentarium. This slope suggests a broad V -shaped calice profile. 

The dissepimentarium is composed of 8 to 15 or 

more rows of medi urn to large dissepiments and 

occupies approximately one-half of the corallite 

radius. Outer dissepiments are globose; inner ones 

are vertically elongate. 

Tabulae are incomplete, and the pattern is obscure. 

Microstructure.-As for genus. 

D'iscussion.-This description is based on three 

fragments of coralla from a single locality. The 

species is distinguished from all other colonial corals 

in the Onesquethaw by its combination of characters, 

but it is most similar to Heliophyllum n. sp. B, from 

which it differs in having long yard-arm instead of 

short zigzag carinae, in smaller and less regularly 

globose dissepiments, and in lacking a calicular platform. 

Distribution.-Known only from the Edgecliff 

bioherm facies at one locality in eastern New York. 

Material.-VSNM 172226-227, NYSM 12866. 
SYSTEMATIC PALgONTOLOGY 127 

Cerioid Species 

Heliophyllum coalilum (Rominger) 

Plates 96-98 

1876 Cyathophyllum coalitum Rominger, p. 108-109, 

pl. 38, fig. 4. 

not 1881 C. coalitum Rominger. Quenstedt, pl. 161, fig. 3 

(probably Heliophyllum yandelli). 

part? 1883b Heliophyllum coalitum (Rominger). Hall, p. 259-

260, pl. 7, fig. 3 (not figure 2). 

1887 C. coalitum Rominger. Holmes, p. 16, pl. 10, figs. 

4a-e. 

? 1901 Cyathophyllum coalitum Rominger. Lambe, p. 

152. 

1948a Hexagonaria coalita (Rominger). Stumm, p. 20-

21, pl. 1, fig. 3; pl. 9, figs. 5, 6. 

? 1948a H. bathycalyx ponderosa Stumm, p. 23. pl. 3, 

fig. 1. 

1955 H. coalita (Rominger). Stumm, cards 172-173. 

? 1955 H. bathycalyx ponderosa Stumm. Stumm, card 

208. 

? 1965 H. ponderosa Stumm. Stumm, p. 44, pl. 39, fig. 1. 

Occurrence of type material.-Ann Arbor, Mich.; 

"Found frequently in silicified condition in the drift, 

connected with fossils of the corniferous limestone." 

(Rominger, 1876, p. 109). Presumably derived 

from (1) Amherstburg, Onondaga, or Bois Blanc of 

southwestern Ontario, or (2) Bois Blanc of Mackinac Straits region, Michigan. (See discussion.) 

Diagnosis.-Cerioid or subcerioid H eliophyllum 

composed of large corallites separated by very thin 

walls; calices with axial boss and broad peripheral 

platform. Septa attenuate; major extend to axis 

where they bend and twist slightly without forming 

an axial structure. 

External features.-The coralla are large, as 

much as at least 30 by 45 em in diameter and 15 em 

high. Mature corallite diameters range from 23 to 

41 mm; in one large corallum 10 corallites range 

from 25 to 32 mm (average 29.9 mm; fig. 37). 

Calicular pits are broad and shallow with a low 

boss, formed by the ·axial ends of the septa, covering most of the ·base; the peripheral platform is 

horizontal and wide. 

Increase is peripheral, four or more offsets being 

produced at one time. 

Internal features.-The septa are bilaterally arranged although this is obscured in many corallites 

by slight twisting of septa at the axis. Major septa 

extend to or nearly to the axis and tend to bend or 

twist at their axial ends without forming an axial 

structure (except in one aberrant colony in which 

an aulos is formed ; see pl. 98 and discussion) . 

Cardinal and counter septa are shorter than other 

major septa and (in many corallites) adiacent 

m~ajor ·septa tend to bend toward the cardinalcounter plane so that they meet along a line or form 

a small elliptical area with their ends. Minor septa 

, extended to the edge of the tabularium without entering it. All septa are uniformly thin in the dissepimentarium; the major septa are slightly dilated 

within the tabularium. The number of major septa 

per corallite ranges from 31 to 49 in the few specimens analyzed (fig. 37) ; in one large corallum the 

number of major septa in 10 corallites ranges from 

34 to 40 and averages 38.1. 

All septa are carinate in the dissepimentarium. 

Long yard-arm carinae predominate but zigzag 

carinae are not uncommon. Carinae are closely 

spaced, ranging from 6 to 11 per 5 mm, averaging 

approximately 9 per 5 mm, in transverse sections. 

Short zigzag carinae are present in the tabularium. 

The corallite walls are straight or curved, forming an irregular polygonal pattern in transverse 

sections. Individual corallites are not everywhere in 

contact so that open spaces between individuals exist 

locally. At these places, the wall between two corallites splits into its two parts and is curved as in a 

phaceloid coral. Wails between adjacent corallite 

range in thickness from 0.15 to 0.50 mm. 

The tabularia are wide, occupying approximately 

two-fifths the diam~eter of e-ach corallite. T~abulae are 

incomplete, forming a pattern that is axially arched 

and peripherally concave. Spacing of tabulae varies. 

The dissep:imentaria are formed by sm,all to 

medium-sized globose dissepiments arranged in rows 

that are steeply inclined near the tabularium, almost 

horizontal in peripheral parts of corallites. 

Discussion.-The above description is based on 

five fragments of coralla from the Onondaga Limestone in New York which have been compared with 

the Michigan type specimens. The largest of these 

measures approximately 30 by 4·5 by 15 em and 

includes several dozen corallites. Other fragments 

are smaller including, 4 to 20 corallites. None of the 

fragments preserves the original form of the colony 

or any part of the holothec·a and all are broken or 

crushed internally, apparently because of very thin 

septa and walls. 

The New York specimens are similar to the bolotype and paratypes from Michigan. Most characteristic seems to be the large corallite size, the occasional curved walls, and the attenuate structures. 

In these characters, H. coalitum differs from other 

dE~scribed species of H eliophyllum or Cyathocylindrium. In the field, H. coalitum is most easily confused with H. 1nonticulum because of similar corallite size and delicate structure. The latter species 

differs mainly in the absence of a wall between corallites and in the arrangement of dissepiments. 

Variation in Heliophyllum coalitum is consider-
128 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

50~~------~~--------~~------~~------~~------~l--------~l------.-l------. 

45-

40 I-

•t:: 

EXPLANATION 

X Heltophyllum monftculum 

0 H. coaltfum 

0 H. cassum 

I .1 

---------.-~----------- 1 I -------}--i 

I I 

I I 

I I 

I I 

I I 

I 

I 

I 

-------H~------~--------------

-

-

-

-

-

! 'I 

-----=+--~--=--- -

1 I 

I I I I I I 

5 10 

d' (mm) 

15 20 25 

d (mm) 

30 35 

dax (mm) 

40 45 

FIGURE 37.-Massive Heliophyllum spp. data. Mean and observed range of number of major septa (n and ORn). versus 

mean and observed range of diameter (d and ORd). In the two astreoid species, diameter is measured as axial distance 

(d(ij,) and n is also plotted against mean tabularium diameter (d1 ). 

able, particularly in the size and shape of corallites 

(straight or curved walls) and in the axial arrangement of major septa. The small sample and the 

crushing of most specimens makes analysis difficult. 

One ·specimen consisting of parts of three or four 

corallites is par_ticularly interesting in having a welldeveloped aulos, formed as in Eridophyllum by the 

de·flection of the major septa (pl. 98). The auloi are 

large, 4 to 6 mm in diameter and apparently continuous, as one aulos has a length of 2 em or more 

with complete, approxi·mately horizontal tabulae inside the aulos. 

Distribution.-Rominger's types were drift specimens collected near Ann Arbor, Mich. According to 

Stumm (1948a, p. 21), they were probably derived 

from the Onondaga Limestone in southwestern 

Ontario, ·but it seems ·equally possible that they could 

have come from the Bois Blanc Formation of the 

same area or from the Mackinac Straits region. 

The New York specimens described in this paper 

were collected from the bioherm facies of the Edgecliff Member. Three of the specimens are from near 

Leroy, in western New York; two specimens are 

from near Cox'sackie, in the Hudson Valley. The 

species was apparently widespread, although rare, 

and in N·ew York may have been restricted to the 

bioherm facies of the Edgecliff Member. 

One specimen in the collections of the Geological 

Survey of Canada, is from "N. Oxford township, 

Oxford County, Ontario." This is the Woodstock 

area and the specimen is most likely from the Bois 

Blanc Formation but possibly from the Amherst-
SYSTEMATIC PALEONTOLOGY 129 

burg. The specimen is silicified; this and the general 

appearance of the specimen also suggests a Bois 

Blanc .source. 

Hall (1883b) illustrated a cerioid coral from the 

Jeffersonville Limestone as this species. This may 

be the form later described by Stumm (1948a) as 

H exagonaria ponderosa and both may be conspecific 

with H eliophyllum coalitum. The holotype of H. 

ponderosa is proba·bly indeterminate, ·and I have not 

seen Hall's specimen; none of the corals illustrated 

by Davis or ·Greene in their Falls monographs is 

at all similar, so the presence of this form at the 

Falls must remain conjectural. 

M aterial.-Michigan : holotype, UMMP 8579 ; 

Stum·m hypotype, UMMP 24195. New York: specimens herein illustrated, U8NM 172228-231 and 

NYSM 12867 (part of USMN 172229). Additional 

specimen, NYSM 3495. Ontario: here illustrated, 

GSC 3586. 

Astreoid Species 

Heliophyllum monticulum (Ehlers and Stumm) 

Plates 99-102 

? 1899 Phillipsastraea billingsi Calvin, Lambe, p. 249 

(not Calvin, 1893). 

? 1901 P. billingsi Calvin, Lambe, p. 169. 

1953 Billingsastraea monticula Ehlers and Stumm, p. 

4, pl. 2, figs. 1-3. 

1955 B. monticula Ehlers and Stumm, o Stumm, card 

236.4. 

? 1958 B. billingsi (Calvin), Cranswick and Fritz, p. 

43, pl. 2, fig. 3. 

Occurrence of type material. -Bioherm facies of 

Edgecliff Member, Onondaga Limestone, Williamsville, N.Y. 

Diagnosis.-Astreoid H eliophyllum with large 

corallites, prominent yard-arm carinae, and long 

m·aj or septa which are irregular in the axial region. 

External features.-The astreoid coralla are very 

large, as much as 60 em or more in diameter and 

15 em or more in height. Corallites are irregularly 

pdsmatic with indefinite boundaries. Diameters 

range from 17 to 44 mm but variation is more 

limited within individual colonies (table 38 and 

fig. 37) ; mean diameter in eight colonies ranges 

from 22 to 36 ·mm. 

Increase is peripheral but offsets expand rapidly 

and few can be recognized in any one colony. 

Calices ar·e crater shaped with the rounded rim 

of the crater projecting 4 to 5 mm above the marginal level. Inside the rim the calicular pits are bowl 

shaped, approximately 1 to 2 mm deep and 1> to 12 

·mm in diameter. No boss is pres·ent, but the pattern 

of the major septa is apparent. Outside of the rim, 

the calicular platform is reflexed downward. becoming horizontal toward the edge of the corallite. 

TABLE 38.-Heliophyllum monticulum (Ehlers and Stumm), 

summary data 

Nl --=------ n: x ____ _ OR ----

s ------

v ----- d"'<: x ------ OR ___ _ 

8 ------

v ----- dt: x ------ OR ___ _ 

8 ------

v -----

Intracolony InterMinimum Maximum Mean colony 

9 

21.6 

20-23 

1.07 

4.0 

22.1 

17-29 

2.75 

10.0 

5.6 

5.0-5.9 

.27 

3.7 

10 

3~.4 

31-36 

2.07 

7.0 5.7 

36.4 

28-44 

5.96 

18.9 13.9 

10.5 

9.0-11.5 

1.08 

11.1 7.3 

9 

27.3 

4.11 

15.1 

29.1 

4.51 

15.5 

8.2 

1.74 

21.4 

1 Axial distance (da"') based on 15 or 20 measurements in each of 

8 coralla; tabulalium diameter (d1 ) was measured in 4 to 10 corallites 

in each of 7 c01·alla. 

Internal features.-The septa are bilaterally arranged; the cardinal and (or) counter septa are 

commonly shorter than the other major septa which 

are longer away from the cardinal-counter plane. 

Major septa extend to or nearly to the axis but 

rarely meet; axial ends tend to form a linear or 

ellipsoidal pattern along the cardinal-counter plane. 

M:inor septa on either side of the counter septum 

m·ay be slightly longer than other minor septa but 

commonly are undifferentiated. In some specimens 

maJor septa are irregularly deflected to form a weak 

whorl (as described by Ehlers and .Stumm, 1953, p. 

4) ; in the holotype this is accentuated by crushing. 

The minor septa are limited to the dissepimentari urn, are thinner, and have shorter c·arinae than 

the major septa. Peripherally the septa commonly 

continue into the next corallite but may abut against 

other septa. Septa are attenuate, but m·ay dilate 

slightly in the inner dis·sepimentarium or tabulari urn. Number of major septa ranges from 20 to 

3(;; the average number per corallite in 9 coralla 

ranges from 21.6 to 33.4 (average 27.3) (table 38 

and fig. 37). 

Yard-arm and fewer zigzag carinae are strongly 

developed along all septa in the dissepimentari urn. 

The average number of carinae in the 5 ·mm closest to 

the tabularium ranges from 7.5 to 12.4, averaging 

9 to 10. On the sides of the septa, in longitudinal 

sections, the carinae appear as abruptly projecting 

ridges directed upward and inward which become 

les·s steep toward the axis. Very short zigzag carinae 

are commonly ·present within the tabularium. 

The dissepimentaria occupy a large part of each 

corallum. Dissepiments are small and inflated, 

steeply inclined adjacent to the tabularia, and gently 

inclined to horizontal near corallite boundaries. 

Tabularia range in diameter from 5.0 to 12.0 mm 

but show less variation within coralla; in seven well-
130 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

preserved colonies, mean tabularium diameter 

ranges from 5.6 to 10.5 mm and averages 8.2 mm 

(table 38 and fig. 37). The tabularia are composed 

of closely spaced, short, arched, incomplete tabulae 

that form a horizontal pattern of successive calice 

floors. 

Septa, dissepiments, and tabulae commonly are 

very thin and apparently were delicate. Most coralla 

colla:psed slightly during burial and now exhibit 

broken septa. A few specimens were obviously distorted, but in most specimens, horizontal dimensional changes appear to have been slight. 

Microstructure.-Same as for the family. Some 

carinae are dilated and formed of a single, coarse 

monacanth, but craspedophyllid microstructure predominates. 

Discussion.-The ~above description is based on ~5 

colonies or fragments of colonies from the Onondaga 

Limestone in New York. Only nine of these were 

large enough and sufficiently well preserved for the 

measurem·ents and counts indicated in the text 

(ta:ble 38 and fig. 37). 

This and the following species have been referred 

to Billingsastraea ( = Asterobillingsa) .by Stumm 

and other recent workers in eastern North America. 

~he species differ from that genus in their large 

corallites with marked bilateral symmetry, their 

long yardarm carinae, and their septal dilation. 

Large, astreoid H eliophyllum differ from Taimyrophyllum Chernychev (or Eddastraea Hill) in the 

morphology of both carinae ~and dissepiments; the 

latter have zigzag carinae or no carinae and depressed dissepiments. (See Pedder, 1964, for description and discussion of Taimyrophyllum.) 

Large astreoid corals, with corallite diameter in 

the range of 2-4 em are not uncommon in the E-N A 

Onesquethaw Stage. Early workers referred to these 

as Phillipsastraea gigas (a Silurian Arachnophyllum) or P. billingsi (Late Givetian, Iowa), but none 

was illustrated until 1953 when Ehlers and Stumm 

described the Edgecliff species as BUlingsastraea 

monticulum. 

A species distinct from H. monticulum is present 

in the Bois Blanc Formation, Ontario. Material described by Billings (1859) and Nicholson (1875) 

is probably from the Bois Blanc and presumably not 

H. monticulum . .Specimens of Lambe (1899, 1901) 

may be either the Bois Blanc or Edgecliff species, 

but Lambe describes septa "reaching the centre 

and becoming somewhat twisted," a character of 

H. monticulum rather than the Bois Blanc form. 

The longitudinal section from northern Ontario, 

illustrated by Cranswick and Fritz (1958, p. 43, pl. 

2, fig. 3) as Billingsastraea billingsi (Calvin), shows 

morphologic characters closer to those of H. monticulum than to H. billingsi. Cranswick and Fritz 

partly based their identification on specimens of 

"B. billingsi" from the "Onondaga" limestone in 

Ontario (Cranswick and Fritz, 1958, p. 44) although 

B. billingsi was described from the Cedar Valley 

Limestone in Iowa (Ehlers and Stumm, 1953, p. 

10). Pending redescription o.f the Cranswick and 

Fritz material, their specimens are tentatively 

placed in H. monticulum. 

H elioph1Jrllum yandelli and a second form from the 

J effersonviUe Limestone at the Falls of the Ohio 

consistently differ from H. monticulum in having 

long, prominent minor septa on either side of the 

counter ( ?) septum. H.? billingsi differs in that the 

·major septa tend to meet at the axis and the 

carinae are dominantly zigzag (.possibly this should 

be referred to Cyathocylindrium). The septa also 

meet at the axis in H. ingens, but this form has distinct H eliophyllum-type carinae. 

Distribution.-H eliophyllum monticulum is known 

only from the Edgecliff Member of the Onondaga 

Limestone and is probably restricted to the bioherm 

facies. Of the 33 studied ,colonies or fragments, 29, 

including the holotype, are from a single Edgecliff 

bioherm near Williamsville, N.Y. (loc. 108). Half 

of these were colle.cted during the present study, the 

rest were borrowed from collections in the Geology 

Department, University of Buffalo, and in the 

USNM. 

The largest and best preserved colony (pl. 101, 

figs. 1-3) was collected from the Edgecliff bioher·m 

north of Thompson's Lake, New York (loc. 81). 

Three specimens borrowed .from the New York 

State Museum are labeled Thompsons Lake (NYSM 

3497), Falkirk (NYS·M 12868), and Falkirk (NYSM 

12869) . Thes·e are almost certainly from the Edgecliff Member and probably from the bioherm facies. 

A single specimen is known from the Edgecliff 

bioherm facies near Port Colborne, Ontario. 

Maten'al.-Holotvpe: UMMP 28519 (fragment is 

USNM 128015). Illustrated or measured specimens: 

USNM 172232-236, 136711, 136692; NYSM 12868-

70. Other thin-sectioned specimens: USNM 9003a; 

NYSM 3498-3500; and UBuff (five specimens) . 

Other studied specimens: USNM 172237 a-d, 172238, 

113262, 182908, and UBuff (six specimens). 

Heliopbyllum cassum n. sp. 

Plate 103 

? 1859 Phillipsastrea gigas Owen, Billings, p. 128. 

? 1875 P. gigas Owen, Nicholson, p. 77-78. 

Occurrence of type material.-Bois Blanc Formation (Emsi8.n), Hagersville and Innerkip, Ontario. 
SYSTEMATIC PALEONTOLOGY 131 

Diagnosis.-Astreoid H eliophyllum with moderately large corallites and relatively short major 

septa. Septa are locally discontinuous with development of lonsdaleoid dissepimentarium; tabulae are 

compl·ete or incomplete. 

External features.-Two known specimens measure 11 by 10 by 3 and 15 by 10 by 12 em in length, 

width and height; both are fragments of larger 

coralla. No part of the original surface of either 

colony is preserved, but parts of the holotheca are 

preserved on both. Corallite diameters (da) range 

from 13 to 27 m·m and average 22.5 and 19.7 mm in 

the two specimens (fig. 37). 

Increase is peripheral and offs·ets are very numerous in the holotype which was a young, ·expanding 

corallum. In both specimens, increase was primarily 

on the outside half of the outside corallites of the 

colony. 

Calices interpreted from longitudinal sections are 

shallow and have moderately steep walls, a rim, 

and a slightly reflexed platform; the calice floor 

is approximately horizontal. 

Internal features.-The septa are bilaterally arranged although this is obscured in many corallites 

by irregularities. The major septa extend approximately halfway through the tabularium and are 

variously arranged within the tabularium. Bilateralism ·may be ·marked by relatively shorter cardinal 

and counter septa, by adjacent septa that bend 

toward the cardinal-counter plane, and by longer 

than normal minor septa on either side of the 

cardinal septum. In some corallites the septa irregularly bend within the tabularium and no plane of 

bilateral symmetry can be discerned. 

Number of m·ajor septa ranges from 18 to 22 and 

averages 19.9 and 19.8 in the available colonies 

(fig. 37). Minor septa are limited to the dissepimentarium. 

All septa are heavily carinate, commonly with 

long yard-arm carinae, but subyard-arm and zigzag carinae are also present. The septa are co·mmonly continuous from one corallite into an adjacent one, but spaces where three or four corallites 

join may be without septa and occupied by large dissepiments. Such spaces are characteristic of all 

transverse and longitudinal sections. 

The dissepimentarium is broad, composed of small 

to medium-sized globose dissepiments; in spaces 

lacking septa the dissepim·ents are large; spaces 

between dissepiments are as much as 1.5 mm high 

and 6 mm or ·more in diameter. 

The tabul'arium diameter ranges from 5 to 8 mm 

and averages 6.4 and 7.0 in the two coralla. Tabulae 

are complete or incomplete, flat or gently concave 

axially, and strongly downbent peripherally. ~ 

Microstructure.-Not well preserved but apparently as for family. No significant dilation was 

noted in either specimen. 

Discussion.-The above description is based on 

only two specimens, but they seem distinct enough to 

warrant description as a new species. They differ 

from all known astreoid H eliophyllum in their short 

rnajor septa that are lonsdaleoid in peripheral 

"corners." 

Large astreoid corals described by Billings ( 1859) 

and Nicholson (1875) have not been examined but 

were from the Bois Blanc Formation and may have 

been this species. 

Distribution.-Known only from the Bois Blanc 

Formation at Hagarsville (loc. C19) and Innerkip 

(loc. C28) , Ontario. 

Material.-Holotype, GSC 31165 (and US·NM 

172244) ; paratype, UMMP 31601. 

Heliophyllum procellatum n. sp. 

Plates 104, 105 

Occurrence of type material.-The holotype and 

paratype are from Falkirk, N.Y., and probably the 

Bois Blanc Formation. (Statement based on occurr·ence of similar forms in Bois Blanc of Michigan; see discussion.) 

Diagnosis.-H eliophyllum with medium-sized 

corallites in which long yard-arm carinae predominate. The major septa for·m an anastomosing network 

in the tabularium. 

External features.-Astreoid or thamnasterioid 

corals; two New York specimens studied are 19 by 

13 by 2 em and 13 by 13 by 3 em. Corallites are 

irregularly bounded, radiating to subparallel. Corallite diameters (dax) range from 13 to 23 mm and 

average 17.9 and 18.2 mm in the two coralla studied 

(table 39 and fig. 30). 

Calices are crater shaped with the inner slope of 

the crater corresponding to the outer margin of the 

tabulari urn. Inside the rim the calicular pit is shallow, 1 to 2 mm deep, and has ·a broad gentle boss. 

The calicular platform is deflected sharply downward so that the rim of the crater projects 2 to 

4 mm above the margins of the corallites. 

Internal features.-The septa are radially arranged. The major septa extend to the axial region 

where they are irregularly deflected in all directions to form an anastomosing structure one-half 

to two-thirds the diameter of the tabularium. The 

num·ber of major septa ranges from 18 to 21 (average 19.6), and 17 to 20 (average 18.6) in the two 

coralla (table 39 and fig. 30). 
132 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

TABLE 39.-Heliophyllum procellatum, intracolony data 

1V ----------------

n: x ----------- OR --------- 8 -----------

v ----------- d1: x ----------- OR ---------

8 -----------

v ----------- r ----------- /:.. ----------- X -----------

New York 

NYSM .NYSM 12872 12873 

20 

19.6 

18-21 

1.1 

5.4 

3.9 

3.5-4.6 

.3 

8.1 

.27 

.30 

17.9+ 

20 

18.6 

17-20 

.9 

4.7 

4.0 

3.5-5.0 

.4 

9.2 

.44 

<.05 

18.2+ 

Michigan 

(1) 

23.7 

21.3-27.0 

2.23 

9.4 

6.8 

5.7-7.5 

.72 

10.5 

1 I.ntercolony variation based on three to six corallites in five crushed co rail a. 

Minor septa are limited to the dissepimentarium 

and are slightly thinner than the major ones. All 

septa are att·enuate and relatively uniform in thickness throughout their lengths. Peripherally some 

septa are continuous into the next corallite, but 

others abut against other septa. 

Yard-arm and zigzag carinae are strongly dev·eloped; the former appear to predominate, but 

many of the yard-arm carinae are zigzag with only 

slight offset. The carinae are closely spaced, averaging eight per 2.5 mm at the inner margin of the 

dissepimentarium. Within the tabularium major 

septa have short zigzag carinae irregularly distributed. 

The dissepimentaria are composed of small, globose dissepiments, steeply inclined toward the coralite axes near the tabularium, irregularly inclined 

to horizontal elsewhere. The dissepiments are confined to interseptal spaces. The boundaries between 

adjacent corallites cannot be recognized in longitudinal sections. 

The tabularia range in diameter from 3.5 to 4.6 

mm (average 3.9) and from 3.5 to 5.0 mm (average 4.0) in the two coralla (table 39 and fig. 30). 

Tabulae are incomplete and dissepimentlike and are 

almost lost in the anastomosing axial ends of the 

major septa. 

Discussion.-The above description is based on 

two incomplete specimens from the Bois Blanc Formation ( ?) at Falkirk, N.Y. Eight thin sections 

la:beled Falkirk, from the NYSM are of this species 

and may have been cut from the same two specimens. 

H. procellatwm is distinguished by the anastomosing arrangement of major septa in the tabularia, by 

the predominance of yard-arm carinae in the dissepimentarium, and by the presence of zigzag 

carinae in the tabularium. In these characters it is 

so different from other Onesquethaw sp·ecies of the 

genus that it seems worth naming in spite of the 

small sample and limited information on provenance. 

The two knowp specimens of H. procellatum were 

found in the NY8M labeled "Falkirk, New York." 

The labels indicate that they had been selected by 

G. B. Simpson as types of a new species which was 

to have been described in a monograph on the corals 

of New York. This was never completed, and no 

written notes or description are known. Both the 

Bois Blanc Formation and the Edgecliff Member 

crop out at Falkirk (near Akron) ; there h~ no 

matrix adhering to either specimen and no direct 

way of determining the source bed. 

As discussed below, similar corals, possibly conspecific, occur in the Bois Blanc Formation of 

Michigan. This limited evidence suggests that a 

Schoharie age is more likely than an Onondaga one 

(see discussion in section on "Stratigraphy") and 

that H. procellatum in New York may be from the 

Bois Blanc rather than the Onondaga. 

Material.-Holotype, NYSM 12872; illustrated 

para type, NYS.M 12873; eight additional thin sections labeled Falkirk, NYSM 12874 (probably from 

one or both of the above .specim·ens). 

Heliophyllum sp. cf. H. procellatum n. sp. 

Plate 106 

1876 Phillip8astraea gigas, Rominger (not Owen, 1844) p. 

129-130, pl. 37, fig. 3. 

1945 Billingsastraea sp., Ehlers, pl. 13, figs. 1, 2. 

Ten specimens from the Bois Blanc Formation of 

Michigan are similar enough to H. procellatum to 

warrant notice and comparison. The specimens are 

all partly crushed so that corallite diameters and 

features of the corallite peripheries cannot be accurately determined. 

The Michigan specimens are strikingly similar to 

H. procellatum in the morphology of their carinae 

and in the axial arrangement o.f major septa. 

Carinae are long, closely spaced (six to nine per 

2.5 mm) and predominantly yard-arm or only 

slightly offset. Major septa extend to the axis where, 

in most corallites, they irregularly bend and twist 

as in H. procellatum; short, zigzag carinae are 

present in the tabularium. 

The Michigan specimens have a much larger tabularium diameter and correspondingly more septa 

than H. procellatum. Limited data from five better 

preserved coralla are given in table 39. Tabularium 

diameter in other specimens is similar. Total 

diameter ( dax) in the Michigan· specimens has not 

been measured because of the crushing, but it appears to be not significantly greater than in H. 

procellatum. 
SYSTEMATIC PALEONTOLOGY 133 

At present, too few specimens are known from 

New York or Michigan to determine the limits of 

individual variation. The Michigan specimens are so 

similar to the New York ones in the morphology of 

septa and carinae that some relationship seems 

probable. Size and septal number differences ·may 

or may not be significant. 

Distribution.-Bois Blanc Formation, Mackinac 

Straits region, Emmet County, Mich. 

MateriaL-Illustrated specimen, UMMP 18804; 

other studied specimens, USNM 172239, 172240, 

and UMMP 23602 (figured by Ehlers, 1945, pl. 13, 

fig. 1, as Billingsastraea sp.), 23603 (figured by 

Ehlers, 1945, pl. 13, fig. 2 as B. sp.), 8574 (figured 

by Rominger, 1876, pl. 37, fig. 3, as Phillipsastraea 

gigas),22953,22954,50630a,50749. 

LOCALITY DATA 

New localities in New York and southwestern 

Ontario are described in sufficient detail so that anyone with the proper map should be able to find the 

locality and identify the stratigraphic units present. 

Each locality represents one to many collections. 

Data on the position of collections at the localities 

is not giv·en because no pattern or significance to 

position within members or facies has been recognized. The collection data is preserved in the records 

of the U.S. Geological Survey (Silurian-Devoni~n 

catalog and field notebooks). Assignment of all specimens and species to members, facies, and beds discussed in the stratigraphy section is indicated in the 

systematic descriptions and :in tables 40-46. 

Old localities in N·ew York and southwest Ontario 

a1~e identified in as much detail as is possible, but 

many localities can only be assigned to a general 

area. 

Localities of comparative material from other 

areas are similarly described if new. Data on locality 

of type and previously described specimens can be 

obtained in the original publication. 

TABLE 40.-Distribution of colonial rugose corals in the Bois Blanc and Schoharie Formations, New York and southwestern Onta'rio. 

[Localities are listed in geographic order from west to east. Numbered localities are. "new" (see locality list); letters refer to "odd" localities as follows: A, Woodstock and Oxford; B, Hagarsville and Walpole; C. Port Colborne; D, Falkirk; E. Leroy; F, Schoharie and Schoharie County; G, in Berne 15-min quadrangle; H, in Albany 15-min quadrangle. G and H consist only of external molds but are identified with reasonable certainty. Num- ber of coral!a known from each locality is indicated]. 

Bois Blanc Formation Schoharie Grit 

Localities ------ _________________ Central Western southwest Niagara Peninsula, Ontario New York Eastern New York Ontario 

A C28 C19 C20 C22 B C17 ClO c D E F 79 G 505 73 H 

Acinophyllum simcoense ,., 1 4 1 6 1 1 2 3 

A. stokesi ---------------- 2 1 1 13 4 1 1 2 (4) 1 (5) 

Cylindrophyllum laxtensum_ 2 4 2 1 2 

Asterobillingsa rugosa ---- 1 

A. ma.qdisa magdisa ------ 1 1 1 1 1 2 1 

Grewgiphullum coll1gatum _ 2 2 4 

H eliophyllum p1·ocellatum __ 2 

H. cassum --------------- 1 1 

TABLE 41.-Distribution of colonial rugose corals in the biohBrrn facies of the Edgecliff Member of the Onondaga Limestone 

[Localities are listed in geographic order from west to east. Numbered localities are "new" (see locality list); letters refer to "old" localities as follows: 

A, Port Colborne; B, Falkirk; C, Leroy; D, Thompsons Lake. Albany County; E, Clarksville. Number of coralla known from each locality is indicated]. 

Niagan Peninsula, Western 

Localities --------------____ 0-'--n.c.:._t.c.:._a_ri_o___ New York 

Synaptophyllum 

arundinaceum. 

S. tabulatttm ---------- Acinophyllum stramin- eum. .4. segregatum --------- Cylindrophyllum 

elogatttm. 

C. propinquum --------

C. deearium ------------ C. sp. A --------------- Asterohillingsa magdisa steorra. A. mr-gdi11a magdisa 

E,.idophyllwm subseriale_ 

Gre•naiphullttm 

colligrrtum .. Cuathocylindrium opulens 

C .. oemm.ntum ---------- H elior.hullum megaproliferum. 

1-7. n. so. B ------------ H. n. sn. C ----------- H. coalitnm ----------- H. monticulum --------

C10 A Crl C7 C5 C:l lOR 348 B 438 

2 

2 

1 

3 

2 

23 

34 

1 

18 

1 

4 

8 

13 

30 2 

2 

3 

10 

3 

Cent':"al Eastern New York 

New York ----:-----:-:-::-,..-:-:--:-::-.,---::-::::---::-::::---::-::-:::--::-=-::---=---:-:-~-=:-::-:-::--::-:-:-=-::-:: C 597 :HJO 207 233 236 239 229 269 185 135 285 329 D 81 244 E 248 296 297 

2 

2 

2 

2 

1 

13 

--

1 

2 10 

1 

1 

2 

2 

7 

1 

2 

--

1 

4 

2 

4 

--

1 

2 

5 

3 

2 

1 

3 

2 

2 

3 

2 

2 

1 

2 

3 

2 

7 

3 
TABLE 42.-Distribution of colonial rugose corals in the nonbioherm facies of the Edgecliff Membe'r of the Onondaga Limestone 

[Localities are listed in geographic order from west to east. Numbered localities are "new" (see locality list) ; letters refer to "old" localities as follows: A, Formosa area (see discussion 

of stratigraphy in central-southwestern Ontario); B, Woodstock; C, Walpole; D, Cheapside; E, Cayuga. Number of coralla known from each locality is indicated. The asterisk marks 

collections from the Edgeclitf dark-limestone facies] 

Localities ------------------

Synaptophyllum 

arundinaceum. 

S. tabulatum ------------- 8. kladion ---------------- Acinophyllum stramineum __ 

A. se_gregatum 

A. mclareni --------------- Cylindrophyllum propinquum 

C. deearium ------------- Prismatophyllum O'll".;deum _ Asterobillingsa magdisa steorra. Eridophyllum subseriale ___ _ Grewgiphyllum colligatum __ Cyathocylindrium opulens __ 

<::S\ralO~~~;~- Niagara Peninsula, Ontario N':!s~~~k Central New York Eastern New York 

A C36 C27 B C22 C C23 D E C45 C11 C10 C5 339 109 114 119 104 101 56 3 7 598 54 215 206 198* 188 188* 191 * 337* 336* 164 154 135 90 286 82 77 292 

9 

4 

1 

6 3 

2 

2 

cf. 

5 

2 2 2 2 11 3 4 2 3 2 

12 

........ 

00 

~ 

z 

0 

z 

0 

-< 

UJ. 

~ 

~ 0 

~ 

1-d 

::q 

0 

0 

~ 

0 

z 

1-1 

> 

~ 

~ 

c:::: 

Q 

0 

UJ. 

t_%j 

0 

0 

~ 

> 

~ 

UJ. 
SYSTEMATIC PALEONTOLOGY 

TABLE 43.-Distribution of colon'ial rugose corals in the Clar- ence and Nedrow Membe·rs of the Onondaga Limestone 

[Only four specimens of one species (one from each of four localities) were studied. The localities are listed :in geographic order from west to 

eastJ 

Clat·ence Member Nedrow Member West•~rn New York Eastern New York 

Localities 114 421 119 132 

Acinophyllum segregatum __ 1 1 1 1 

TABLE 44.-Distribution of colonial rugose corals in the Moorehouse Member of the Onondaga Limestone 

135 

[Localities are listed in geographic order from west to east. Numbered lo::alitie:> are "new" (see locality list); letters refer to "old" localities as follows: A, \Voodstock; B, Port Dover; C, Naticoke; D, Albany County. The number of c01·alla known from each locality is indicated. The atserisk marks collectwns from tne Acinophyllum-bridophyllum bedJ 

Southwest Ontario Central Eastern SouthWestern New York east Central Peninsula 

Niagara New York New York New York 

Localities ------------------------------ A B c CIS 110 111 112 497 373 393 118" 120 120* 493* io2 336 86 D 246 301 324 

Synaptophyllum kladion -------- 1 1 1 2 3 

Acinophyllum segregatum ------ 1 1 3 9 3 3 4 14 9 10 4 1 1 1 2 1 

A. mclareni ------------------- 1 

Eridophyllum seriale ----------- 1 1 

E. seriale population B --------- 1 1 17 7 2 

E. aulodokum ----------------- 1 1 

E. conjunctum ---------------- 1 

E. corniculum ----------------- 28 6 

Cyathocylindrium n. sp. A ------ 2 

TABLE 45.-Location of the single colonial rugose coral known 

f'rom the Seneca Member of the Onondaga Limestone 

Prismatophyllum truncata 

TABLE 46.-List of museum numbers of all type, illustrated, 

and otherwise mentioned specimens, giving the species, collection number, and locality 

[Museum numbers are in numerical order within the following sequence of institutions 1. U.S. N·ational Museum of Natural History. 2. New 

York State Museum. 3. Geological Survey of Canada. 4. American 

Museum of Natural History. 5. British Museum (Natural History). 6. Harvard Univetsity Museum of Comparative Zoology, 7. University of 

Iowa, Department of Geology. 8. University of Michigan, Museum of 

Paleontology. 9. Orton Museum, Ohio State University. 10. Royal Ontario 

Museum. Species are keyed as followa: 1. Synaptophyllum arundinaceum, 

2. S. tabulatum, 3. S. kladion. 4. Acinophyllum simcoense, 5. A. stra- mineum, 6. A. segregatum, 7. A. mclareni, 8. A. stokesi, 9. Cylindrophyllum 

elongatum., 10. C. propinquum. 11 C. laxtensum, 12. C. deearium. 13. 

Prismatophyllum ovoideum. 14. P. truncata. 16. Asterobillingsa magdisa steorra, 17. A. magdisa magdisa, 18. A. runosa. 19. Eridophyllum seriate, 20. E. seriate population B, 21. E. corniculum, 22. E. subseriale. 23. Grewgiphyllum colligatum. 24. Cyathocylindrium opulens, 25. C. gemmatum, 26. C. n. sp. A., 27. Heliophyllum megaproliferum, 28. H. n. sp. B, 29. 

H. n. sp. C, 30. H. coalitum, 31. H. monticulum, 32. H. cassum. 33. Cylindrophyllum sp. A. 34. Disphyllum? stummi. 35. Prismatophyllum 

prisma. 36 Eridophyllum aulodokum. 37 H eliophyllum procellatum. 38. Eridophyllum conjunctum) 

Museum Species number number 

Collection 

number Locality 

U.S. National Museum of Natural History 

Washington, D.C. 

10719 -----1--------------------- 10721 ____ 19 ____________________ _ 

25870 ____ 12 ____________________ _ 

26000a ___ 16 ____________________ _ 27961 _____ 4 ____________________ _ 

113262 ____ 31 ____________________ _ 113263 ____ 16 ____________________ _ 

113264 ____ 17 ____________________ _ 113552 ____ 23 ____________________ _ 

113554 ____ 23 ____________________ _ 

113556 ____ 23 ____________________ _ 

Williamsville. 

Albany County. 

Leroy. 

Schoharie County. 

Hagarsville, Ontario. 

Williamsville. 

Do. 

Port Colborne, Ontario. 

Manitoulin Island 

(Ontario) drift. 

Hagarsville, Ontario. 

Do. 

Central New Yo!·k, locality 46 

1 

TABLE 46.-List of museu.m numbers of all type, illustrated, 

and otherwise mentioned specimens, giving the species, collection number, and locality-Continued 

Museum Species number number 

Collection 

number Locality 

128015 

128016 

136692 

136711 

143121 

162576 

162577 

162578 

162579 

162580 

162581 

162582 

162583 

162584 

162585 

162586 

162587 

162588 

162589 

162590 

162591 

162592 

162593 

162594 

162595 

162596 

162597 

U.S. National Museum of Natural History 

Washington, D.C.~Continued 

____ 31 _____________________ 108 (part of UMMP 

____ 17 ____________________ _ ____ 31 ____________________ _ 

____ 31 ____________________ _ 

____ 16 ____________________ _ 

_____ 1 236n _____ 1 5318-SD 

_____ 1 5312-SD 

_____ 1 81-(1)-5a _____ 1 1815r 

_____ 1 5358b-SD _____ 1 236 

_____ 1 348-3 

_____ 1 USNM 9003 _____ 1 7525-SD 

_____ 1 7525-SD 

_____ 1 7807-SD _____ 1 Core spec. _____ 3 5338-SD 

_____ 3 5310-SD 

_____ 3 5309b-SD _____ 3 5310-SD _____ 3 373 

_____ 3 7524-SD 

____ _4 5052-SD 

_____ 4 5052-SD 

____ _4 5052-SD 

28519). 

Port Colborne, Ontario. 

Williamsville. 

Do. 

Do. 

236 

438 

438 

81 

185 

207 

236 

348 

108 

C10 

C10 

Ohio. 

Do. 

373 

120 

120 

120 

373 

Cll 

C28 

C28 

C28 
136 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

TABLE 46.-List of museum numbers of all type, illustrated, TABLE 46.-List of museum numbers of all type, illustrated, and otherwise mentioned specimens, giving the species, col- and otherwise mentioned specimens, giving the species, col- lection number, and locality-Continued lection number, and locality-Continued 

Museum Species Collection Locality MwJeum Species Collection number number number Locality ruumber number number 

U.S. National Museum of Natural History U.S. National Museum of Natural History Washington, D.C.-----Continued Washington, D.C.-Continued 

162598 _____ 4 5052-SD C28 162668 _____ 7 7101-SD Ky 162599 _____ 4 5052-SD C28 162669 _____ 7 7101-SD Ky 162600 _____ 4 5024-SD C19 162670 -----8 7534-SD C10 162601 ____ _4 5014-SD C10 162671 _____ 8 7516-SD C22 162602 _____ 4 5052-SD C28 162672 -----8 7516-SD C22 162603 ____ _4 7509-SD C22 162673 _____ 8 7511-SD C22 162604 _____ 4 7505-SD C22 162674 _____ 8 5887a-SD 505 162605- _____ 4 7518-SD C19 162675 _____ 8 5895-SD 79 162606 _____ 4 7515-SD C22 163367 -----8 4721-SD Falls 162607 _____ 2 244 244 163368 _____ 8 4725-SD Do. 162608 _____ 5 108-17b 108 163369 _____ 8 4721-SD Do. 162609 _____ 5 135 135 163370 _____ 8 7515-SD C22 162610 -----5 108-17b-e 108 163371 _____ 8 7503-SD C20 162611 _____ 5 108-15b 108 163372 _____ 8 7506-SD C22 162612 -----5 108-15b 108 163373 _____ 8 7517-SD C22 162613 _____ 5 5351a-SD 229 163374 _____ 8 5028-SD C22 162614 -----5 108-17b-e 108 163375 _____ 8 5028-SD C22 162615 _____ 6 5353-SD 269 163376 _____ 8 7508-SD C22 162616 _____ 6 5362d-SD 329 163377 _____ 8 5895-SD 79 162617 _____ 6 5353-SD 269 163378 _____ 8 4721-SD Falls 162018 _____ 6 5357b-SD 188 163379 _____ 8 4721-SD Do. 162619 -----6 5354c-SD 198 163380 _____ 5 5028-SD C22 162620 _____ 6 5354c-SD 198 163381 _____ 5 5013-SD C9 162621 _____ 6 5308a-SD 118 163382 _____ 9 108-11 108 162622 _____ 6 5309a-SD 120 163383 _____ 9 108-8f-a 108 162623 _____ 6 5350b-SD 233 163384 _____ 9 5350e-SD 233 162624 _____ 6 233-3 233 163385 _____ 9 5362c-SD 329 162625 _____ 6 269-r2N 269 163386 _____ 9 5362c-SD 329 162626 _____ R 329-1 329 163387 _____ 9 108-8f 108 162627 _____ 6 135 135 163388 _____ 9 296-1c 296 162628 _____ 6 108-171 108 163389 _____ 9 207 207 162629 _____ 6 5352-SD 269 163390 _____ 9 135 135 16211~0 _____ 6 5350c-SD 233 163391 ____ 11 5052-SD C28 162631 _____ 6 5319-SD 438 163392 ____ 11 7518-SD C19 1626~2 _____ G 5351a-SD 229 163393 ____ 11 5887a-SD 505 1626~3 _____ 6 5312-SD 438 163394 ____ 11 5887a-SD 505 1626~4 _____ fi 5364-SD 81 163395 ____ 33 108-19d 108 1621)~5 _____ h 5314-SD 438 163396 ____ 34 5828-SD Quebec 162636 _____ 6 5313-SD 438 163397 ____ 34 5828-SD Do. 1626~7 _____ 6 5330-SD 108 163~98 ____ 34 5828-SD Do. 1626~8 _____ 6 296-1b 296 16~~99 ----~f) 4724-SD Falls 

162639 _____ 6 5357a-SD 188 16~400 ____ 35 5933-SD Do. 1621140 _____ 6 5354c-SD 198 16~401. !ll'\ 5934-SD Do. 

162641 _____ 6 5355-SD 188 163402 ____ 35 USNM Campbell 368B Do. 

162642 _____ 6 5354a-SD 198 163403 ____ 35 USNM Campbell 368B Do. 

162643 _____ 6 5355-SD 188 16~404 ____ 35 USNM Campbell 368B Do. 

162644 _____ 6 4673-SD 493 16~405 ____ 35 USNM Can,nbell 368B Do. 

162645 _____ 6 4673-SD 493 16~406 ____ 35 8289-SD Ohio 

1621146 -----6 5308a-SD 118 163407 ____ 13 4728-SD Falls 

162647 ______ 6 531la-SD 120 16~408 ____ 13 4728-SD Do. 

162648 _____ 6 5308b-SD 118 16~409 ____ 13 4728-SD Do. 

162649 _____ 6 5308b-SD 118 16~410 ____ 1~ 4728-SD Do. 

1621150 _____ 6 5311b-SD 120 16~411 ____ 13 USNM Campbell 368C Falls 

162651 _____ 6 5308b-SD 118 163412 ____ 1~ USNM Campbell 368C Do. 

162652 _____ 6 5311a-SD 120 16~413 ____ 13 USNM 1558 Do. 1r.?.R!'l~ _____ 6 5308a-SD 118 163414 ____ 13 5051-SD C27 

162654 _____ 6 5311a-SD 120 16~415 ____ 13 7808-SD Ohio 

162655 _____ 6 5310-SD 120 16~416 ____ 1~ 8289-SD Do. 

162656 _____ 6 5338-SD 373 16~417 ____ 13 7809-SD Do. 

162fl57 _____ 6 5329-SD 112 16~418 ____ 14 Cornell U. 202 46 

162fl58 _____ 6 5361-SD 336 16~419 ____ 17 5891-SD 73 162~!)9 _____ 6 421-N 421 16~420 ____ 17 5027-SD C20 

162Rfl0 _____ 7 5051-SD C27 16~421 ____ 17 7530-SD C10 

1R2flfl1 _____ 7 5063-SD C36 16~422 ____ 1'7 108-S 108 

1R2RR2 _____ 7 5021-SD C18 1~'U9~ 1f7 USNM (no number) Hagarsville 1R2nn~ _____ 7 4728-SD Falls 16~424 ____ 19 53~4-SD 497 

162nh4 _____ 7 7101-SD Ky 1R~425 ____ 19 7786-SD Ohio lR2~n5 _____ 7 5030-SD C23 11)~4.2() ____ 20 5~os:~a-SD 118 1~?.R~f) _____ 7 5030-SD C23 ]RQ4.?,7 ____ 20 5~08b-~D 118 

162667 _____ 7 4724-SD Falls 163428 ____ 20 5308a-SD 118 
SYSTEMATIC PALEONTOLOGY 137 

TABLE 46.-List of museum numbers of all type, illustrated, 

and otherwise mentioned specimens, giving the species, collection number, and locality-Continued 

Mu»eum Species 

n:umber number 

Collection 

number Locality 

U.S. National Museum of Natural History 

Washington, D.C.-Continued 

163429 ----20 5311a-SD 120 

163430 ____ 20 5308a-SD 118 

163431 ____ 20 5308a-SD 118 

163432 ____ 20 5308a-SD 118 

163433 ____ 20 5308a-SD 118 

163434 ____ 20 5308a-SD 118 163435 _____ 20 4673-SD 493 

163436 -----36 112 112 

163437 _____ 36 246 246 

163438 _____ 21 5328-SD 112 

163439 _____ 21 5328-SD 112 

163440 _____ 21 5328-SD 112 

163441 _____ 21 5328-SD 112 

163442 _____ 21 5328-SD 112 163443 _____ 21 5328-SD 112 

163444 _____ 21 5327b-SD 112 

163445 _____ 21 5328-SD 112 

163446 _____ 21 5325-SD 393 

163447 _____ 21 5325-SD 393 

163448 _____ 21 5328a-SD 112 

163449 _____ 21 393 393 

163450 _____ 21 5328a-SD 112 

163451 _____ 21 5328-SD 112 6 specimens 163452 _____ 21 5328a-SD 112 5 specimens 163453 _____ 21 5325-SD 393 2 specimens 163454 _____ 22 5362a-SD 329 

163455 _____ 22 5349-SD 360 

163456 _____ 22 5362b-SD 329 163457 _____ 22 5349-SD 360 

163458 _____ 22 7541-SD 7 4 specimens 163459 _____ 22 5363-SD 90 

163460 _____ 22 5362a-SD 329 

163461 _____ 22 239 239 2 specimens 163462 _____ 22 297-3 297 

163463 _____ 23 7518-SD C19 

163464 _____ 23 7518-SD C19 

163465 _____ 23 7529-SD C10 

163466 _____ 23 7527-SD C10 

172161 _____ 23 5331-SD 108 

172162 _____ 23 USNM 9003 108 

172163 _____ 23 USNM 900~ 108 

172164 _____ 23 7529-SD C10 3 specimen~ 

172165 ______ 8 59b of Kindle Virginia 

172166 _____ 16 Buffalo Univ. 108 

172167 _____ 16 108-16, 151 108 

172168 _____ 16 108 108 

172169 _____ 16 Ruffalo TTniv. Akron 

172170 _____ 16 Buffalo TTniv. 108 

172171 _____ 16 Ruffalo TTniv. 108 

172172 _____ 16 Buffalo TTniv. 108 

172173 _____ 16 Buffalo Univ. 108 

172174 _____ 16 Buffalo TTniv. 108 

172175 _____ 16 81(1)-3 81 

172176 _____ 16 Ruffa]o T.Tniv. 108 

172177 _____ 16 USNM 9003 108 5 specimens 

172178 ______ 5 TTRNM Roucot 1 Quebec 

172179 ______ 5 USNM Boucot 1 Do. 1721~0 _____ 10 296 296 

172181 _____ 10 5~67-~D 296 

172182 _____ to 5R52-SD 269 

172183 _____ 10 5R58!1-~D 207 

172184 _____ 10 5~57h-~D 188 

172185 _____ 10 7520-RD C45 

172186 _____ 10 7~09-SD Ohio 

172187 _____ 10 7~08-~D Do. 1721~8 _____ 10 8289-SD Do. 

172189 _____ 10 8289-RD Do. 

172190 _____ 12 5~49-SD 360 

172191 _____ ]2 81 81 

172192 _____ 12 81-(1)-2 81 

TABLE 46.-List of museum numbers of all type, illustrated, 

and otherwise mentioned specimens, giving the species, collection number, and locality-Continued 

Muneum Species 

number number 

Collection 

niUmber Locality 

U.S. National Museum of Natural History 

Washington, D.C.-Continued 

172193 _____ 12 244 

172194 _____ 12 5063-SD 

172195 _____ 12 7529-SD 

172196 _____ 12 Perry 202 

172197 _____ 24 81 

172198 _____ 24 5350e-SD 

172199 _____ 24 5029-SD 

172200 _____ 24 5010-SD 

1'12201 _____ 24 5350d-SD 

172202 _____ 24 5362b-SD 

172203 _____ 24 5350a-SD 

172204 _____ 24 329 

172205 _____ 24 5030-SD 

172206 _____ 24 5350e-SD 

172207 _____ 24 5330-SD 

172208 _____ 24 5038-SD 

172209 _____ 24 4721-SD 

172210 _____ 24 4721-SD 

172211 _____ 24 5317a-SD 

172212 _____ 25 USNM 9003 

172213 _____ 25 5330-SD 

172214 _____ 25 108-51 

172215 _____ 25 108-51 

172216 _____ 25 81 

172217 _____ 25 5014-SD 

172218 _____ 25 233-3 

172219 _____ 25 Various 

172220 _____ 26 5338-SD 

172221 _____ 27 5004-SD 

1'72222 _____ 27 5330-SD 

172223 _____ 27 5621-SD 

172224 _____ 28 5362a-SD 

1'/2225 _____ 28 329-2 

172226 _____ 29 5625-SD 

172227 _____ 29 297-1 

172228 _____ 30 7149-SD 

172229 _____ 30 5316b-SD 

172230 _____ 30 5314-SD 

1'/2231 _____ 30 296-53 

172232 _____ 31 5331-SD 

172233 _____ 31 5364-SD 

172234 _____ 31 5331-SD 

172235 _____ 31 7528-SD 

1'12236 _____ 31 5331-SD 

1rl2237 _____ 31 5330-SD 

172238 _____ 31 5331-SD 

172239 _____ 37 7662-SD 

172240 _____ 37 7662-SD 

172241 ______ 9 207-5 

172242 ______ 9 108-8f-a 

1'/2243 _____ 12 207 

172244 _____ 32 7518-SD 

172245 _____ 38 8773-SD 

172246 ______ 6 154-lb-2 

182813 _____ 11 ------------------ 182~46 _____ 24 5R50b-SD 

182847 _____ 24 5367-SD 

182848 _____ 24 5358d-SD 

182849 _____ 24 248 

182850 _____ 24 5~50b-SD 

182851 _____ 24 5330-SD 

182852 _____ 24 5R58a-SD 

182853 _____ 24 7525-SD 

182908 _____ 31 USNM (no number) 

Ohio 

244 

C36 

C10 

81 

233 

C22 

C7 

233 

329 

233 

329 

C23 

233 

108 

C25 

Falls 

Do. 

438 

108 

108 

108 

108 

81 

C10 

233 

108 5 specimens 

373 

C3 

108 

248 

329 

329 

297 

297 

296 

438 

438 

296 

108 

81 

108 

C10 

108 

108 4 specimens 

108 

Michigan 

Do. 

207 

108 

207 

C19 

118 

154 

Woodstock 

233 

296 

207 

248 

233 

108 

207 

C10 

Hagarsville, Ontario. 

New York State Museum 

Albany, N.Y. 

246 _________ 9 ------------------ Clarksville. 

299 ________ 18 ------------------ Leroy. 
138 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

TABLE 46.-List of museum numbers of all type, illustrated, 

and. otherwise mentioned .specimens, giving the species, collectwn number, and locahty-Continued 

Museum Species number number Collection 

n.umber Locality 

New York State Museum 

Albany, N.Y.-Continued 

300 ---------2 ------------------ 334-335 ____ _4 ------------------

336 _________ 6 ------------------

337 ---------6 ------------------ 338 _________ 6 ------------------ 12780 -------6 164-lb 

12781 -------6 188-lb-2 

12782 -------6 286-1 

12783 _______ 6 90-1 

12784 _______ 6 112-71 

12785 -------6 NYSM E828B 

12786 _______ 6 108-17k 

12787 _______ 6 269-r2N 

12788 _______ 6 120-5 

12789 _______ 6 236 

12790 _______ 6 336-1a1 

12791 -------6 337-1-A 

12792 _______ 6 206-1b 

12793 -------6 233-5 

12794 _______ 6 207 

12795 _______ 6 5330-SD 

12796 _______ 6 5330-SD 

12797 _______ € 336-1a1 

12798 _______ 6 5354b-SD 

12799 _______ 6 118-cs 

12~00 -------6 118-cs 

12801 _______ 6 5308b-SD 

12802 -------6 5311a-SD 

12803 _______ 6 373 

12804 -------6 118 

12805 _______ 6 102e-3 

12806 _______ 5 108-8f-a 

12807 _______ 5 108-8e-c 

12808 _______ 5 108-16b 

12809 _______ 5 NYSM 1453 

12810 _______ 8 NYSM 1415 

121311 ______ _4 NY~M: 1 M12 

12812 ______ _4 NYSM E828a 

12813 ______ _4 NYSM 1452 

12814 _______ 9 NYSM 1466 

12815 _______ 9 296-2 

12816 _______ 9 207-SW 

12817 ______ 33 108-19d 

12818 ______ 10 1R5 

12819 ______ 12 296-2 

12820 ______ 12 81 

12821 _______ 1 108-17k 

12822 _______ 1 81NW 12~23 _______ 3 J 10-1 

12824 ______ 16 NYSM 1466 

12825 ______ 16 NYSM 608 

12826 ______ 16 2fl9-r1 12~27 ______ 17 NV~M 1452 

12828 ______ 17 NYSM G4033 12~29 ______ 13 NYSM F-691 128~0 ______ 1~ NYSM F.691 

128~1 ______ 20 120-5 128~2 ______ 22 297 128~3 ______ 22 2~5 

12834 ______ 22 329-1 

12835 ______ 21 5~2~-SD 

12836 ______ 21 5~28-SD 

12R37 ______ 21 5~2~-SD 

128~8 ______ 21 5~2~-sn 

12839 ______ 21 5~?.Ra-SD 

12~40 ______ 21 5~2~:::J-SD 

12841 ______ 21 5325-SD 

12042 ______ 23 J 0~-9 

12843 ______ 24 n350e-SD 

12844 ______ 24 NYSM (no number) 

Walpole, Ontario. 

Leroy. 

Clarksville. 

Leroy. 

Do. 

164 

188 

286 

90 

112 

Leroy 

108 

269 

120 

236 

336 

337 

206 

233 

207 

108 

108 

336 

198 

118 

118 

118 

120 

373 

118 

102 

108 

108 

108 

Falkirk. 

Clarksville. 

Leroy. 

Do. 

Do. 

Clarksville. 

296 

207 

108 

135 

296 

81 

108 

81 

JlO 

Clarksville. 

Schoharie County. 

269 

Leroy. 

Schoharie. 

Western New York. 

Western New York. 

120 

297 

285 

329 

112 

112 

112 

112 

112 

112 

112 

108 

233 

Leroy. 

TABLE 46.-List of museum numbers of all type, illustrated, 

and othe1·wise mentioned specimens, giving the species, collection number, and locality-Continued 

Mu13eum Species number number Collection 

number Locality 

New York State Museum 

Albany, N.Y.-Continued 

12845 ______ 24 

12846 ______ 24 

12847 ______ 24 

12848 ______ 24 

12849 ______ 24 

12850 ______ 24 

12851 ______ 24 

12852 ______ 24 

12853 ______ 24 

12854 ______ 24 

12855 ______ 24 

12856 ______ 24 

12857 ______ 24 

12858 ______ 24 

12859 ______ 24 

12860 ______ 25 

12861 ______ 25 

12862 ______ 25 

12863 ______ 25 

12864 ______ 26 

12865 ______ 28 

12866 ______ 29 

12867 ______ 30 

NYSM (no number) 

5362b-SD 

108-SW 

135 

135 

236 

248 

108-51 

296-2 

244 

135 

296-2 

108-171 

269r2N 

329a 

108-51 

108-SW 

108-SW 

108-51 

373 

329-1 

5625-SD 

5316b-SD 

12868 ______ 31 NYSM 1453 

12869 ______ 31 NYSM 1462 

12870 ______ 31 108-12e 

12871 ______ 12 NYSM 1452 

12872 ______ 37 NYSM 1462 

12873 ______ 37 NYSM 1462 

12874 ______ 37 NYSM 1462 

12875 _______ 2 NYSM 1480 

Do. 

329 

108 

135 

135 

236 

248 

108 

296 

244 

135 

296 

108 

269 

329 

108 

108 

108 

108 

373 

329 

297 

438 (part of USNM 

172229). 

Falkirk. 

Do. 

108 

Leroy. 

Falkirk. 

Do. 

Do. 

Walpole, Ontario. 

Geological Survey of Canada 

Ottawa, Ontario 

3422a _______ 1 

3431, a ______ 5 

3431b-f _____ 1 ------------------ 3432a-d _____ 1 

3436 _______ _4 

3585a ______ 11 

3586 _______ 30 

3602 ________ 1 ------------------ 31143 _______ 8 GSC 2596 

31144 _______ 4 5052-SD 

31145 ______ 12 5014-SD 

31146 ______ 11 5026-SD 

31147 ______ 11 GSC 23676 

31148 ______ 11 GSC 2599 

31149 ______ 11 GSC 2599 

31151 _______ 1 GSC 22795 

31152 _______ 1 GSC 2645 

31153 _______ 3 5021-SD 

31154 _______ 3 GSC 2599 

31155 ______ 16 GSC 3877 

31156 ______ 17 GSC 23676 

31157 ______ 23 GSC 2599 

31158 ______ 23 GSC 2617 

31159 ______ 24 5010-SD 

31160 ______ 24 7525-SD 

31161 ______ 27 5004-SD 

31162 _______ 6 G~C 2~521 

31163 _______ 6 GSC 22978 

Cayuga, Ontario. 

Wainfleet Township, 

Ontario. 

Do. 

Walpole Township and 

Wainfleet Township, 

Ontario. 

Woodstock, Ontario. 

Walpole Township, 

Ontario. 

North Oxford Township, Ontario. 

Cayuga, Ontario. 

Oxford, Ontario. 

C28 

C10 

C19 

C28 

Woodstock, Ontario. 

Do. 

Port Maitland, Ontario. 

?W ood'stock, Ontario. 

C18 

Woodstock, Ontario. 

Do. 

C28 

Woodstock, Ontario. 

Walpole Township, 

Ontario. 

C7 

C10 

C3 

Port Dover, Ontario. 

Naticoke, Ontario. 
SYSTEMATIC PALEONTOLOGY 139 

TABLE 46.-List of museum numbers of all type, illustrated, 

and otherwise mentioned specimens, giving the species, collection number, and locality-Continued 

Museum Species 

number number 

Collection 

number Locality 

Geological Survey of Canada 

Ottawa, Ontario-Continued 

31164 _______ 6 GSC 23657 

31165 ______ 32 7518-SD Chepstow, Ontario. 

C19 (part of USNM 

172244). 

American Museum of Natural History 

New York, N.Y. 

641 ________ 17 ------------------ 23501 ______ 38 ------------------

Schoharie, N.Y. 

Falls. 

British Museum (Natural History) 

London, England 

31730-31 ___ 35 ------------------ Falls,, 

Harvard University Museum of Comparative Zoology 

Cambridge, Mass. 

7827 ________ 8 

7859 _______ 38 

7877 _______ 38 

Falls. 

Do. 

Do. 

Department of Geology, University of Iowa 

Iowa City, Iowa 

21198 ______ 35 ------------------ Falls. 

University of Michigan, Museum of Paleontology 

Ann Arbor, Mich. 

8579 _______ 30 

18804 ______ 37 

24178 ______ 13 

24195 ______ 30 

28519 ______ 31 

31601 ______ 32 

36120 _______ 7 

50576 ______ 12 

60484 -------7 60485 ______ 24 

Drift, Ann Arbor, 

Mich. 

Mackinac County, 

Mich. 

C27 

Drift, Ann Arbor, 

Mich. 

108 

C28 

Formosa area, Ontario. 

Cheapside, Ontario. 

Loraine County, Ohio. 

Ashtabula County, 

Ohio. 

Orton Museum, Ohio State University 

Columbus, Ohio 

17735 _______ 7 

17779 ______ 10 

21336 ______ 38 

27064 _______ 8 

27128 ______ 18 

Bellepoint. Ohio. 

Dublin, Ohio. 

Chillacothe, Ohio. 

Royal Ontario Museum 

Toronto, Canada 

Northern Ontario. 

Do. 

NEW YORK LOCALITIES 

Base maps for fieldwork in New York were the 7 1h-minute 

(1: 24,000) topographic quadrangle maps published by the 

U.S. Geological Survey. New localities are those studied and 

collected by the author; old localities are those known from 

museum labels or catalogs. Where an old locality can definite.,. 

ly be :identified with a new locality, the new locality number 

is used in parentheses. 

Western, central, eastern and southeastern areas are shown 

in figure 5. 

NEW LOCALITIES 

3. Syracuse West quad., central area. Split Rock quarry 

near Highway Dept. Shops on southwest side of Syracuse. 

Onondaga Limestone: 

Feet 

Nedrow Member (top not exposed) --------- 10 

Edgecliff Member (type locality) ----------- 8 

Unconformity. 

Helderberg Group. 

7. South Onondaga quad., central area. Abandoned quarry on 

Onondaga Indian reservation, 1 mile (1.6 km) south of 

Nedrow. 

Onondaga Limestone: 

Feet 

Seneca Member (top not exposed) ---------- 15 

Tioga Bentonite Bed ---------------------- 1h 

Moorehouse Member ----------------------- 19 

Nedrow Member -------------------------- 13 

Edgecliff Member (base not exposed) ------- 10% 

4(). Romulus quad., central area. Warren Bros. quarry, 1 mile 

(1.6 km) west of Canoga. 

Onondaga Limestone: 

Feet 

Marcellus Shale (black shale). 

Seneca Member --------------------------- 21 

Tioga Bentonite Bed ----------------------- 1h 

Moorehouse Member (only top few inches exposed). 

5·1. Munnsville quad., central area. Munnsville Limestone 

Corp. quarry, 1 mile (1.6 km) southeast of Munnsville. 

Onondaga Limestone: 

Feet 

Edgecliff Member (top not exposed) -------- 22 

Unconformity. 

Helderberg Group. 

56. Phelps quad., central area. General Crushed Stone Co. 

quarry at Oaks Corners, northwest of Geneva. 

Onondaga Limestone: 

Feet 

Moorehouse Member _____ .:_ ____________ ;----- 36 

Clarence Member ------------------------- 16 

Edgecliff Member -------------------- ____ 8-10 

Unconformity with 1ft+ relief. 

Silurian rocks. 

73. Clarksville quad., eastern area., Route 32 roadcut 2 miles 

(3.2 km) southeast of Clarksville and ravine parallel 

to and west of road. 

Onondaga Limestone and Schoharie Formation. Section in 

roadcut (Oliver, 1963a, p. 29): 

Feet 

Moorehouse Member (top not exposed) 49 

Nedrow Member (base not exposed) ________ 12 

Edgecliff Member and the Schoharie are exposed in ravine to west. 

'77. Clarksville quad., eastern area. Outcrop along Indian 

Ledge Road, 1.3 miles (2.1 km) south of the north 

edge of the map. Edgecliff Member rests on 2 ft + of 

the Schoharie Formation. (base not exposed). Higher 

Onondaga members are discontinuously exposed to 

south. 

79. Altamont quad., eastern area. Mine lot Creek in Thacher 

State Park, 0.6 of a mile (1 km) south of Route 157. 
140 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

Schoharie Formation crops out below the poorly exposed Edgecliff Member. 

81. Altamont quad., eastern area. Hill and roadcut on south 

side of east-west road north of· Thompsons Lake, 0.2 of 

a mile ( 0.3 km) east of Route 157 (Thompsons Lake 

reef of Oliver, 1956b, p. 18; bioherm 6 in fig. 5). Roadcut .shows Edgecliff bioherm facies resting on 4% ft 

of bedded, biostromal Edgecliff that is underlain by 

2 ft of Schoharie Formation. 

82. Altamont quad., eastern area. Outcrop on north-south 

private road, half a mile (0.8 km) east of Knox. Tenfoot exposure of :lower Edgecliff Member. 

86. Gallupville quad., eastern area. Bed of Fox Creek in Berne. 

Fifty-foot exposure of upper Onondaga; collection only 

from middle Moorehouse Member. 

90. Schoharie quad., eastern area. Glen on southwest side of 

Schoharie Hill, south of light-duty road at 1,080-ft 

contour. Lower Edgecliff rests on poorly exposed Schoharie Formation. 

101. Victor quad., central area. Great Brook south of Victor 

(0.2 of a mile (0.3 km) south of east-west Dryer 

Rd.) Small outcrop of lower Edgecliff Member. 

102. Honeoye Falls quad., central area. In Honeoye Falls 

village, just above dam. 

Onondaga Limestone: 

Feet 

Clarence Member (top not exposed) -------- 16 

Edgecliff Member ------------------------- 2 

Unconformity. 

Silurian. 

Additional outcrops in Honeoye Creek southeast of village on 

both sides of ~oute 65 and extending to North Bloomfield expose 40ft or more of Moorehouse Member. 

104. Honeoye Falls quad., central area. Railroad cut south 

of Honeoye Creek, 0.9 of a mile (1.45 km) northwest 

of Honeoye Falls. 

Onondaga Limestone: 

Feet 

Clarence Member (top not exposed) --------- 9 

Edgecliff Member ------------------------- 3 

Unconformity. 

Silurian. 

108. Buffalo NE quad., western area. Complex of quarries 

north of Route 5 between Williamsville and Snyder; 

outcrops now largely destroyed or inaccessible due to 

construction of Y oungmann Expressway interchange. 

Bioherm 29 in figure 5. This is the old Fogelsonger 

quarry (name variously spelled, sometimes with initial "V") of the literature. The Edgecliff bioherm facies was well exposed here; one or more bioherms 

rested on the Akron Dolomite with bedded Edgecliff 

and Clarence dipping away in all directions. Numerous 

collections from different sites and subfacies were examined, but no faunal differentiation was found except between bioherm and off-bioherm facies. The 140 

identifiable coralla of 12 species of colonial rugose 

corals, noted on table 41, represent the largest collections and diversity of any locality in the study area. 

109. Clarence quad., western area. Creek-side exposure in 

Clarence. Probably "Clarence Hollow" of early reports. Edgecliff Member (3% ft) between poorly exposed Clarence Member and Bois Blanc ( ?) Formation. 

110. Lancaster quad., western area. Federal Crushed Stone 

Co. quarry at Bellevue. 

Onondaga Limestone: 

Seneca Member (top not exposed) ----------

Tioga Bentonite Bed ----------------------

Moorehouse Member (western coral facies) --

Clarence Member (top only) . 

Feet 

6 

'% 

52 

111. Lancaster quad., western area. Ellicott Creek at Browmansville. Five-foot exposure of upper Moorehouse 

Member. 

112. Lancaster quad., western area. Active quarry of Buffalo 

Crushed Stone Co., north of Thruway near Harris 

Hill. Forty-five-foot exposure of Moorehouse Member 

(western coral facies) . 

114. Clarence quad., western area. Quarry north of Route 

5, 3 miles ( 4.8 km) east of Clarence. 

Onondaga Limestone: 

Feet 

Clarence Member (top not exposed) -------- 20 

Edgecliff Member ------------------------- 4 

Unconformity. 

Silurian. 

118. Stafford quad., western area. Active quarry of Genesee 

Stone Products Corp., 1 mile (1.6 km) west of Stafford. Forty-two-foot west face of quarry is entirely 

within Moorehouse Member. Acinophyllum-Eridophyllum bed is 25 ft below the top of the face. Tioga 

Bentonite Bed exposed at top of east quarry face so 

measured section extends virtually to top of Moorehouse. 

119. Leroy quad., western area. Quarry of General Crushed 

Stone Co., 1.4 miles (2.2 km) northwest of Limerock. 

Onondaga Limestone: 

Feet 

Clarence Member (top not exposed) -------- 21 

Edgecliff Member ------------------------- 5 

Unconformity ( ? ) . 

Bois Blanc Formation --------------------- 2 

Unconformity. 

Silurian. 

120. Leroy quad., western area. Large quarry north of railroad, 1.5 miles (2.4 km) east of Leroy. 

Moorehouse Member of Onondaga Limestone: 

Limestone -------------------------------- Bed with Acinophyllum and favositids ------

Limestone with chert ----------------------

Acinophyllum-Eridophyllum bed ------------

Limestone with chert ---------------------

132. Cobleskill quad., eastern area. Quarry on 

boundary of Cobleskill. 

Feet 

2 

1 

4% 

1'% 

22'% 

northeast 

Onondaga Limestone (see Oliver, 1956c, p. 1454, for section 

and discussion) : Feet 

Nedrow Member (top not exposed) --------- 13 

Edgecliff Member ------------------------ 30 

Unconformity? 

Schoharie Formation ---------------------- 2'% 

Carlisle Center Formation. 

135. Cobleskill quad., eastern area. Outcrop 0.15 of a mile 

(0.24 km) southwest of farmhouse on Route 7 (at 

intersection) 0.7 of a mile (1.12 km) west of East 
SYSTEMATIC PALEONTOLOGY 141 

Cobleskill. Small Edgecliff bioherm (East Cobleskill 

reef of Oliver, 1956b, p. 19; bioherm 9 in fig. 5). Bioherm facies rests on, and passes west into normal 

bedded Edgecliff. 

154. Sharon Springs quad., eastern area. Roadcuts on Route 

20, 1 mile (1.6 km) east-southeast of Sharon Center. 

Onondaga Limestone: exposures above the highway 

are in the Moorehouse Member; small exposures between the new and old roads are Nedrow ( ? ) or lower 

Moorehouse; 14 ft of Edgecliff is exposed below the 

old road. 

164. Sprout Brook quad., eastern area. Field west of School 

No. 2 road and south of Creek, 1.45 miles (2.4 km) 

west of Leesville. Extensive field outcrop of Edgecliff 

Member. 

185. Sprout Brook quad., eastern area. Small hill, just east 

of north-south road between railroad and creek on 

north edge of Cherry Valley. Edgecliff bioherm forms 

hill; small quarry 100 ft east of reef is in normal 

Edgecliff; Nedrow crops out in railroad cut just south 

of quarry. This is the Cherry Valley reef of Oliver, 

1956b, p. 19; bioherm 11 in figure 5. 

188. Sprout Brook quad., eastern area. Field exposures by 

W. Judd Falls road, 1.2 miles (2.1 km) north of 

Route 20 at west edge of map. Edgecliff Member of 

Onondaga Limestone: 

Feet 

Edgecliff, normal (coar,se, coral:line) facies __ 10 

Edgecliff, dark-limestone facies ------------- 6% 

Unconformity? 

Schoharie Formation ( ?) (few inches). 

Carlisle Center Formation. 

191. Sprout Brook quad., eastern area. Outcrops by west 

Judds Falls road, 1.5 miles (2.4 km) north of Route 

20 at west edge of map. Fifteen ft Edgecliff; lower 

3 ft are dark-limestone facies. Top and bottom of 

member not exposed. 

198. East Springfield quad., eastern area. Ravine at east 

edge of map, 2. 7 miles ( 4.5 km) north of old Route 

20 in Cherry Valley. 

Incomplete exposure of Edgecliff Member: 

Feet 

Edgecliff normal (coarse, coralline) facies) 9 

Edgecliff dark-limestone facies ------------- 6 

206. East Springfield quad., eastern area. Northwest-facing 

hillslope, 1.2 miles (2.1 km) east-northeast of Summit 

Lake and 0.7 of a mile (1.12 km) south of north edge 

of map at 1,600-1,620-ft contour. Edgecliff Member 

exposed in extensive ledges; section incomplete with 

scattered exposures of Carlisle Center Formation below and higher members of Onondaga above. 

207. East Springfield quad., eastern area. Mount Tom in 

northwest one-ninth of map. Edgecliff bioherm ( 700 X 

500X70 ft) forms upper part of hill, rests on 6-10 

ft of bedded Edgecliff. This is the thickest of the 

known, exposed bioherms. It is the Mount Tom reef 

of Oliver, 1956b, p. 20-21; bioherm 15 in figure 5. 

215. Jordanville quad., eastern area. Field outcrops along 

Herkimer-Otsego county boundary at corner of Otsego 

County. Extensive ledges of Elgecliff Member in its 

normal facies. 

229. Jordanville quad., eastern area. Hillocks 0.2 of a mile 

(0.32 km) north of Chyle Road just west of northsouth road that crosses Merry Hill. Edgecliff Member: 

six to eight biohermal mounds may represent the dissected remnants of a large bioherm. or several P~all 

ones; they rest on bedded Edgecliff that crops out 

along the south side of the exposure. This is the Mount 

Tom No. 7 reef of Oliver, 1956b, p. 22; bioherm 21 

in figure 5. 

233. East Springfield and Richfield Springs quads., eastern 

area. Broad, low hill 0.5 of a mile (0.8 km) westsouthwest of Mount Tom. Edgecliff bioherm rests on 

bedded Edgecliff; described as Mount Tom No. 2 reef 

by Oliver, 1956b, p. 21; bioherm 16 in figure 5. Bioherm dimensions of 1,200X900X40 ft makes this the 

largest of the known, exposed bioherms. 

23ti. Richfield Springs quad., eastern area. Hillside at southeast corner of woods, 0.4 of a mile (0.64 km) north of 

Marshall cemetery and 1.4 miles (2.2 km) northnortheast of Warren. Edgecliff bioherm; described as 

Mount Tom No. 3 reef by Oliver, 1956b, p. 21; b!oherm 18 in figure 5. 

23H. Richfield Springs quad., eastern area. Low hills just 

west of road, 1.5 miles (2.4 km) due north of Warren. Edgecliff' bioherm; described as Mount Tom No. 

4 reef by Oliver, 1956b, p. 22; bioherm 20 in figure 5. 

244. Clarksville quad., eastern area. Hill and cliff face 

around house located directly above Route 157, 1.6 

miles (2.56 km) north of Route 85 intersection and 

1.3 miles (2.1 km) northwest of New Salem. Edgec:liff bioherm very well exposed; grades northward 

into normal Edgecliff. This is the New Salem reef of 

Oliver, 1956b, p. 18; bioherm 4 in figure 5. 

246. Ravena quad., eastern area. Small quarries on either 

side of dead-end road and 0.3 of a mile (0.48 km) 

southwest of Aquetuck School (south of Route 143 

near west side of map). Twenty-five ft of upper 

Moorehouse Member. 

248. Ravena quad., eastern area. Hill, 400 ft west of northsouth road at point 0.25 of a mile (0.4 km) south of 

BM 390 (in northwest corner of map). Edgecliff bioherm; described as Northern Coxsackie reef by Oliver, 

1956b, p. 18; bioherm 3 in figure 5. 

269. Jordanville quad., eastern area. Hills at Deck triangulation station, 1.1 miles (1.75 km) southwest of Deck. 

Complex of small Edgecliff bioherms or disssected 

remnants of one or two larger ones. Described as Deck 

Reef Group by Oliver, 1956b, p. 20; bioherms 13, 14 

in figure 5. 

285. Schoharie quad., eastern area. Northern extremity of 

Schoharie Hill, 2.4 miles (3.8 km) northwest of 

Schoharie. Edgecliff bioherm; described as Schoharie 

Hill reef by Oliver, 1956b, p. 19; bioherm 8 in figure 

5. 

286. Gallupville quad., eastern area. Hillslope 0.8 of a mile 

(1.3 km) south of Gallupville; between quarry 

(Esopus Formation) ·and west Berne road. Normal 

Edgecliff Member in ledges. 

2H2. Clarksville quad., eastern area. Roadcuts 0.5 of a mile 

(0.8 km) south of Callanans Corners. Lower Edgecliff ( 8 ft) rests on Schoharie Formation ( 3 ft) . 

2B6. Ravena quad., eastern area. Hills on both sides of northsouth road, 1.6 miles (2.56 km) west-southwest of 

Hannacroix and 0.3 of a mile (0.48 km) northwest 

of Lish Homestead Cemetery. Edgecliff bioherm facies; 

described as Albrights reef by Oliver, 1956b, p. 17-

18; bioherm 2 in figure 5. 
142 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

297. Ravena quad., eastern area. Hill on east side of Limekiln Road, just north of Hass Hill Road junction; 0.7 

of a mile (1.1 km) west of Roberts Hill. Edgecliff 

bioherm facies; described as Roberts Hill reef by 

Oliver, 1956b, p. 17; bioherm 1 in figure 5. 

301. Leeds quad., southeastern area. Gorge of Catskill Creek 

at millpond in Leeds. 

Onondaga Limestone: 

Feet 

Moorehouse Member (top not exposed) _____ 37 

Nedrow Member -------------------------- 43 

Edgecliff Member (southeastern facies) ____ 36 

Schoharie Formation. . 

Beds are nearly vertical, older to west in the area of the 

measured section. Described by Oliver, 1962, p. A16-A17. 

324. Port Jervis South quad., southeastern area. Tristates 

Point between Laural Grove Cemetery and Delaware 

River. Moorehouse Member (estimated 25 ft exposed). 

Locality briefly described by Oliver, 1962, p. A-20. 

329. Schoharie quad., eastern area. Terrace Mountain, 2.2 

miles (3.52 km) north-northwest of Schoharie. Edgecliff bioherm forms the northwestern high point of 

hill. Described as Terrace Mountain reef by Oliver, 

1956b, p. 19; bioherm 7 in figure 5. 

336. Sprout Brook quad., eastern area. Roadcuts on "new" 

Route 20 extending from just east of Judds Falls to 

top of hill. 

Onondaga Limestone (diagramed and discussed by Oliver, 

1963a, p. 26-27): 

Union Springs Shale Member of Marcellus: 

Feet 

Seneca Member ---------------------------

Tioga Bentonite ·Bed ---------------------

Moorehouse Member (upper third not exposed) 

Nedrow Member --------------------------

Edgecliff Member, normal facies ------------

Edgecliff Member, dark-limestone facies ____ _ 

Unconformity. 

Carlisle Center Formation. 

6% 

% 

78 

12 

17 

6 

337. Sprout Brook quad., eastern area. Roadcuts on "new" 

Route 20 just west of Judds Falls. Onondaga Limestone in section similar to that of ioc. 336; top part 

incomplete. 

339. Buffalo NE quad., western area. Inactive quarry, 0.4 

of a mile (0.64 km) east of Bennett High School, 

Buffalo (Bennett quarry in early reports). 

Onondaga Limestone: 

Feet 

Clarence Member (top not exposed) __ 20 (estimated) 

Edgecliff Member ------------------------- 1-9 

Unconformity (with 2-3 ft relief) . 

Akron Dolomite (Silurian). 

Bois Blanc is apparently absent. 

348. Lancaster quad., western area. Old quarry east of 

Youngs Road in Buffalo Country Club (Young's 

quarry of early reports); just northeast of Williamsville. Edgecliff bioherm facies; bioherm was largely 

quarried away but flanking and margin beds are well 

exposed. Bioherm 28 in figure 5. 

360. Oneida quad., central area. Limestone Creek bed, 1.45 

miles (2.4 km) north of south edge of map; 0.1 of a 

mile (0.16 km) east of light-duty road. Boulders of 

bioherm facies Edgecliff; not found in place but these 

indicate a nearby bioherm, one of the two known in the 

central area. Bioherm 22 in figure 5. 

373. Lancaster quad., western area. Lehigh Properti~ Corp. 

test opening for possible quarry development; 0.25 of 

a mile (0.4 km) west of Gunnville Road at point 0.5 

of a mile (0.8 km) north of Wilhelm (intersection of 

Gunnville Road and Route 33). Small exposure of 

Moorehouse Member (western coral facies) of Onondaga Limestone. 

393. Akron quad., western area. Abandoned quarry on Lake 

Road, 0.35 of a mile (0.49 km) south of Pembroke 

Station. Moorehouse Member ( 14 ft; western coral 

facies). 

421. Oakfield quad., western area. Small quarry on U.S. 

Gypsum Co. property, 0.8 of a mile (1.4 km) west of 

Pearl Road and 0.6 of a mile (1.1 km) north of 

Town Line Road; 1.2 miles (1.9 km) southwest of 

Oakfield. 

Onondaga Limestone: 

Feet 

Clarence Member (lower part only) ________ 10 

Edgecliff Member ------------------------ 5% 

Unconformity. 

Silurian. 

438. Byron quad., western area. Old Empile Limestone Co. 

quarry, 0.4 of a mile (0.64 km) north of Walcott 

Road-Keeney Road junction in southeast part of map. 

Edgecliff bioherm facies with flanking beds and partly equivalent Clarence Member well exposed in im493. 

497. 

mediate area. Bioherm 24 in figure 5. 

Leroy quad., western area. Small quarry just south of 

railroad and 0.3 of a mile (0.48 km) northwest of 

St. Anthony Church, Limerock. Moorehouse Member 

( 8 ft; western coral facies) of Onondaga Limestone. 

Acinophyllum-Eridophyllum bed at ground surface so 

exposure is principally of section just below this unit. 

Lancaster quad., western area. Quarry of Lancaster 

Stone Products Corp. Just east of Barton Road and 

north of Thruway; 1.2 miles (1.9 km) southeast of 

Harris Hill. Moorehouse Member ( 18 ft; western coral 

facies). 

505. Clarksville quad., eastern area. Mouth of Gorge of 

Onesquethaw Creek at Clarksville. 

Edgecliff Member (lower part only). 

Feet 

Schoharie Formation ---------------------- 2 

Carlisle Center Formation. 

597. Jamesville quad., central area. Wooded hill on west side 

of Taylor Road at point where road diverges from 

Lafayette-Pompey town line. Edgecliff bioherm; only 

exposed one in central area. Bioherm 23 in figure 5. 

598. Jamesville quad., central area. Quarry of Solvay Process 

Co., north and east of Jamesville. 

Onondaga Limestone: 

Feet 

Seneca Member (top not seen) ------------- 18 

Tioga Bentonite Bed ---------------------- %. 

Moorehouse Member ----------------------- 32 

Nedrow Member -------------------------- 12 

Edgecliff Member ------------------------- 20 

Unconformity. 

Oriskany ( ?) Sandstone. 
SYSTEMATIC PALEONTOLOGY 143 

See Oliver, 1963a, p. 18-21 for discussion of comparable section just southeast of Jamesville. 

OLD LOCALITIES 

Akron. Akron and Walcottsville quads., western area. Several natural outcrops and old quarries in and near the city 

expose the Bois Blanc Formation and the Edgecliff and Clarence Members of the Onondaga. There is evidence from old 

collections and the field that an Edgecliff bioherm existed near 

Akron; possibly this was located in one of the two large 

quarries on the east side of Akron and was quarried away. 

Albany County. Eastern area. The Schoharie Formation and 

all members of the Onondaga crop out in southwest Albany 

County. 

Clarksville. Clarksville quad., eastern area. Onondaga Limestone and Schoharie Formation crop out in or near the village. 

Collections of apparent bioherm corals labeled Clarksville may 

have come from the New Salem bioherm (loc. 244; bioherm 4 

in fig. 5) or an unknown bioherm. 

Falkirk. Akron quad., western area. Located in what is 

now eastern Akron. The falls of Murder Creek and a quarry 

just north of the falls expose the lower Onondaga (Edgecliff 

and Clarence Members). "Reefy" beds in the falls section suggest that an Edgecliff bioherm may be or may have been in 

the vicinity. 

Leroy. Leroy quad., western area. The Bois Blanc Formation and all members of the Onondaga crop out in or near 

Leroy. Locs. 119, 120, and 438 could have been referred to as 

"Leroy." Collections of apparent Bois Blanc and biohermfacies Edgecliff corals from "NE of Leroy" must have come 

from loc. 120 or nearby. 

Schoharie. Schoharie quad., eastern area. The Schoharie 

Formation and. all members of the Onondaga crop out in or 

near Schoharie. 

Schoharie County. Eastern area. Numerous outcrops of 

Schoharie Formation and all members of the Onondaga Limestone occur in the county. 

Thompsons Lake. Altamont quad., eastern area. Probably 

Edgecliff bioherm: see 1loc. 81. 

Western New York. Locality citation almost meaningless. 

Two specimens of Prismatophyllum ovoideum are the only 

known representatives of the species from the State. 

Williamsville. Buffalo NE and Lancaster quads., western 

area. Collections 1labeled "Williamsville" most likely came from 

loc. 108 but other exposures in the village and nearby (e.g., 

loc. 348) could be sources also. 

SOUTHWESTERN ONTARIO 

Base maps for fieldwork in Ontario were the 1: 50,000-scale 

topographic maps available from the Canadian Department 

of Mines and Technical Surveys. All references here are to 

the editions available in 1958. The Niagara Peninsula here 

refers to the area between the Niagara River and the vicinity 

of Hagarsville, Ontario; as noted in the stratigraphy discussion, Onondaga and Bois Blanc facies in the Niagara Peninsula are similar to those in western New York and this is 

considered as part of the principal study area. Collections 

from central southwestern Ontario (the London-Woodstock 

area and north) were mostly used for comparative purposes 

only, but some are so similar to those in the Niagara Peninsula 

and New York that they were included in the principal study. 

NIAGARA PENINSULA 

NEW LOCALITIES 

C3. Fort Erie map sheet. Windmill Point Station; two quarries just north of Route 3C (near west edge of map) 

(lot 12, Bertie T.). Edgecliff bioherm complex with 

bioherm and flank facies and Clarence Member exposed 

in various parts of quarries. Bioherm 30 in figure 5. 

These are the Buel quarries in Stauffer, 1915, p. 22-24. 

C5. Weiland East map sheet. Quarry on west side of road, 

3.2 miles (5.1 km) northeast of Ridgeway (lot 4, con. 

VIII, Bertie T.). Edgecliff bioherm in north face of 

quarry; Clarence Member overlies normal Edgecliff 

Member on east and west faces. Bioherm 31 in figure 

5. This is the Baxter quarry in Stauffer, 1915, p. 20-

22. 

C'"/. Welland East map sheet. Quarry at Shisler Point on Lake 

Erie. Edgecliff bioherm. This is the Empire Limestone 

Co. quarry of Stauffer, 1915, p. 25-26; bioherm 32 in 

figure 5. 

CH. Wel·land West map sheet. Quarry of Canada Portland 

Cement Co., Ltd., on west side of Port Colborne (lot 

33, con. I, Humberstone T.). Edgecliff bioherm facies. 

Bioherm 33 in figure 5. This is probably the quarry 

discussed by Stauffer, 1915, p. 31-35. 

ClO. Welland West map sheet. Quarry on west side of road 

that extends north from Camelot Beach ( Reebs Bay) 

on Lake Erie (lot 6, con. I, Wainfleet T.) . Studied 

exposures are between Canadian National Railroad and 

quarry spur line track and are just .south of spur. An 

Edgecliff bioherm is well exposed south of the spur 

and in the highest part of the east wall just north 

of the spur (bioherm 34 in fig. 5). North of the spur 

and extending to the Canadian National Railroad is 

the anomalous exposure discussed in the stratigraphy 

section. A sequence of argillaceous beds some 20 ft 

thick (beds a-c) dips gently south and grade upward 

into typical Edgecliff bioherm facies lithology. Included 

fossils clearly indicate that the lowest part of the 

anomalous facies sequence is Bois Blanc in age ( approx 4 ft); the upper part (16 ft; beds b and c) contains typical Edgecliff corals. 

Composite generalized section follows: 

e. Beds of bioherm facies. \ 

d. 6 ft of gray limestone with Gnewgiphyllum colligatum 

and H eliophyllurit monticulum. 

c. 7-11 ft of green Cystiphylloides biostrome (thickness 

varies). 

b. 6 ft of green argillaceous beds with Acinophyllum 

segregatum and G. colligatum. 

a. 4 ft of green argillaceous beds with A. simcoense, 

A. stokesi, Asterobillingsa magdisa magdisa, and 

Aemulophyllum exiguum. 

Beds c and d may pass laterally (south) into the bioherm 

facies, but the gap in the quarry wall just north of the bioherm makes this uncertain. Compare section C11. 

Cll. Welland West map sheet. R. E. Law quarry, on north 

side of Route 3, 1.1 miles (1.75 km) west of County 

line (lot 5, con. II, Wainfleet T.). 

Composite section: 

e. 9ft of gray limestone, upper half very cherty. 

d. 7 ft of green argillaceous beds. 

c. 8 ft, 6 in. of Bois Blanc facies. 
144 NONCYSTIMORPH COLONIAL RUGOSE CORALS 

b. 4 in. of green argillaceous beds. 

a .. 2 ft of Bois Blanc facies with scattered blobs and 

grains of sand in basal 4 in (Springvale Sandstone) 

Unconfprmity. 

Silurian 

Unit e is clearly. Onondaga and probably Edgecliff; units a, 

b, and c are typical Bois Blanc in lithology and fossils with 

beds of the green "anomalous" facies noted at loc. ClO; unit 

d is of uncertain age as it has not been studied in detail. 

This section is less than a mile from loc. ClO with which it 

should be compared. 

C17. Dunnville West map sheet. Quarry 300 yd (273 m) 

north of Route 3, 3.3 miles ( 5.3 km) west-southwest 

of Grand River at Cayuga (lot 45, con. I, North 

Cayuga T.) . Some 5 ft of Bois Blanc rests unconformably on Si·lurian rocks This is discussed in 

Stauffer, 1915, p. 56-58. 

Cl8. Dunville West map sheet. Quarry 0.6 of a mile (1.1 km) 

south of Mount Olivet, 150 yd (136 m) north of Old 

Indian Line (road) intersection with road running 

west-southwest (con. IV, South Cayuga T.) Moorehouse Member (15-20 ft). 

Cl9. Simcoe East map sheet. Large quarry of Haldimand 

Quarries and Construction, Ltd., north of Michigan 

Central Railroad on east side of Hagarsville (con. I, 

Oneida T.). Bois Blanc Formation (15 ft) with 

Springvale Sandstone in lower part. This is the R. 

Hamilton quarry of Stauffer, 1915~ p. 68-70. 

C20. Simcoe East map sheet. Large quarry of Canada Crushed 

Stone Co. on west side of Hagarsville just north of 

Michigan Central Railroad (lot 13, con XIII, Walpole T.). Bois Blanc Formation (25 ft); Springvale 

Sandstone Member of Edgeville was seen in floor of 

quarry in 1958; by comparison with section at nearby 

:loc. C22, the section extends almost to the ton of the 

formation. This is the J. C. Ingles' quarry of Stauffer, 

1915, p. 71-73. 

C22. Simcoe East map sheet. Large quarry of Hagarsville 

Quarries, Ltd., west of Hagarsville and just south of 

Michigan Central Railroad (lot 12, con. XIII, Walpole 

T.). 

Bois Blanc and Onondaga section: 

Feet 

Edgeclic Member (top not exposed) --------- 14 

Bois Blanc cherty limestone facies __________ 16 

Bois Blanc (Springvale Sandstone) base not 

exposed) ------------------------------- 4 

There is no evidence here of an unconformity between the 

Edgecliff and Bois Blanc. 

C23. Simcoe East map sheet. Falls of Sandusk Creek, east 

of road 0.2 of a mile ( 0.32 km) south of Sandusk 

(lot 13, con IV, Walnole T.). Fifteen-foot section of 

Edgecliff Member. This is the Haggerty Falls of 

Stauffer, 1915, p. 51-53. 

C45. Dunnville East map .sheet. Lake Erie shore at end of 

road running south from Dunville (lot 19. con. V, 

Dunn T.). Fifteen-foot exposure interpreted as Edgecliff Member; neither top nor base of unit is exposed. 

OLD LOCALITIES 

Cayuga. Dunnville West map sheet. "3 mi [ 4.8 km] W of 

Cayuga" would be near my loc C17 ( GSC 3602). CSC 3422a 

is simply labeled "Cayuga, Ont." Both are thought to be 

from the Edgecliff Member. 

Cheapside. Dunnville West map sheet. UMMP 50576 is 

from "bed 1 of Stauffer" (1915, p. 49); this is in Dry Creek, 

0.75 mile (1.2 km) north of Cheapside. Stauffer',s unit is 

probably Edgecliff. 

Hagarsville. Simcoe East map sheet. Several large quarries 

(loc. C19, C20, C22) and natural outcrops expose the lower 

part of the Onondaga and the Bois Blanc Formation. USNM 

27961, 113554, 113556, 163423, 182908 are labeled "Hagarsville, Ont." only. 

Naticoke. Simcoe East map sheet. GSC loc. 22978 is "Naticoke quarry, Ont." This is a small quarry in the Naticoke 

Creek valley just south of the road. Moorehouse Member of 

Onondaga. 

Port Col borne. W elland East and W elland West map sheets. 

Lake Erie terminus of W elland Canal on boundary of two 

map areas. This is the general locality of many of the early 

collections that are generally referred to "Port Colborne" 

only or to "Rama's (Ramey's) Farm" (see below as separate 

locality) . Bois Blanc Formation and Edgecliff Member of 

Onondaga Limestone USNM 113264 and 128016 are from 

"Port Colborne." 

Port Dover. Simcoe East map sheet. GSC loc. 23521 is 

"Lake Erie shore, 3.5 miles [5.6 km] east of Port Dover." 

Moorehouse Member of Onondaga Limestone. 

Port Maitland. Dunnville East map sheet. GSC loc. 22795 

is "1%, mi [2.8 km] W of Port Maitland, Lake Erie shore." 

This is close to my loc. C45 and is in approximately the 

same stratigraphic position; Edgecliff Member of Onondaga 

Limestone. 

Ramey's Farm, Port Col borne. (See Port Col borne as separate entry.) Stauffer (1915, p. 27-28) notes the H .S. Ramey 

farm. lot 27, con. II, Humberstone Twp. as having been on 

the W elland canal: "There is no section expo.sed at that place, 

but over t.he fields and along the canal much weathered out 

material may be found." Stauffer's list of fossils and earlier 

reports make it clear that both Bois Blanc and Edgecliff were 

the source. The site of "Ramey's Farm" is now entirely urbanized but many excellent localities are near Port Colborne 

(for example locs. ClO, Cll). 

Wainfleet. Welland West maP sheet. Townshin just west 

of Port Col borne; Boi,s Blanc Formation and Edgecliff and 

Clarence Members of Onondaga Limestone are exposed (for 

example locs. ClO, C11). GSC loc. 3431 is "lot 6, con. I, 

Wainfleet." This is the position of loc. ClO and the GSC 

specimens may have come from the same auarrv. GSC 3432 

is from "Walpole Twp. [see following] and Wainfleet Twp." 

Walpole. Simcoe East and Dunnville West map sheets. 

Township including the south half of Hagarsvme and a large 

area south of Hagarsville. The Bois Blanc and all members 

of the Onondaga are exposed (for example locs. C20, C22, 

C2~. Cheapside). NYSM 300 and 1480 and GSC 3432 are 

"Wanole" with no further data. GSC 3585 is "lot 3, con. 

XIV, Walpole Twp."; this is west of Springvale and either 

Bois Blanc or Edgecliff. GSC 2617 is "lot 5. con XIV and lot 

3, con. XV, WaJnole Twp.", also west of Springvale and either 

Bois Blanc or Edgecliff. 

CENTRAL SOUTHWESTERN ONTARIO 

NEW LOCALITIES 

C27. Woodstock West map sheet. Quarry of Gypsum, Lime 

and Alabastine, Canada, Ltd., 1.5 miles (2.4 km) 

southwest of Beachville (lot 18, con. III, North Oxford T.). 
REFERENCES CITED 145 

Detroit River Group: 

Feet 

Lucas Formation (top not exposed) ________ 62 

Amherstburg Formation (base not exposed) __ 16 

C28. Woodstock East map sheet. Inactive quarry on South 

side of Innerkip. Lower part of Bois Blanc Formation 

(12ft) rests on Silurian rocks. 

C36. Wingham East map sheet. Small quarry 1.6 miles 

(2.56 km) east of Gorrie, on east side of Maitland 

River (lot 11, con. VIII, Howick T.). Amherstburg 

Formation. 

OLD LOCALITIES 

Chepstow. Walkerton West map sheet. This is the area of 

the "Foremosa reef" facies of the Amherstburg Formation 

GSC loc. 23657 is "dump on East bank of Teeswater River, 

due N of church at Chepstow." 

Formosa area. Area of "Formosa reef" facies of Amhertsburg Formation. UMMP 36120 is from this unit and area 

(Fagerstrom, 1961). 

Manitoulin Island is in Lake Huron, just east of the Northern Peninsula of Michigan. 

Oxford. The name of three townships and a county. They 

include Ingersoll, Beach ville, Woodstock, and Inner kip. (See 

locs. C27 and C28.) GSC 3586 is from "North Oxford Twp."; 

GSC 2596 is "Oxford, Ontario." Bois Blanc and Amherstburg 

Formations crop out in the area. 

Woodstock. Woodstock West and East map sheets. GSC 

loc. 2599, 3436, 3437, and 3877 are all "Woodstock" or "near 

Woodstock." GSC 2645 is "unknown but with fossils from 

Woodstock and Kilsworth, Ontario." Bois Blanc and Amherstburg Formations crop out in the area. 

OTHER AREAS 

Specimens from these localities are cited or illustrated as 

type or previously ihlustrated specimens of species occurring 

also in New York or .southwestern Ontario or as new comparative material. Site data is generalized and, where feasible, 

reference is made to locality information in original descriptions. 

FALLS OF THE OHIO 

Ohio River at Louisville, Ky., and Jeffersonville, Ind. The 

north side of the river is the State boundary so that outcrops 

in the bed of the river and on the south side are in Kentucky, 

whereas the bluffs on the north side are in Indiana. The 

stratigraphic sequence is discussed in the stratigraphy section, 

and species and specimens are assigned to formation and zone 

in the appropriate systematic description. 

EAST-CENTRAL KENTUCKY 

USGS loc. 7101-SD; Broadhead quad.; just north of Route 

150, west of south tributary of Flax Creek on hillslope at 

approximate elevation of 950 ft, Boyle Limestone; see stratigraphy section for further discussion of age. 

MICHIGAN 

USGS loc. 7662-SD is "2 mi [3.2 km] W of Mackinaw 

City, NW14, NW1,4, sec. 14, T 39N, R 9W; field of Pierce 

and Son Dairy Farm; Bois Blanc Formation." Collection presented to the U.S. Geological Survey by the late Prof. E.C. 

Stumm, University of Michigan. 

"Drift, Ann Arbor" includes material from several stratigraphic units; specimens cited here may be from the Bois 

Blanc Formation in Michigan or from the Bois Blanc or 

Amherstburg Formations in Ontario. 

"Mackinac County" is on the north side of the Mackinac 

Straits and is most likely Bois Blanc Formation. 

OHIO 

USGS LOCALITIES 

7786-SD. Kelly Island quad. Marblehead quarry. Columbus 

Limestone, Paraspirifer Zone (Zone H). 

7807-SD. Shawnee Hills quad. Union Stone Co. quarry, 2.4 

miles (3.84) southwest of Bellepoint. Columbus Limestone 

Coral Zone (Zone C) . 

7808-SD. Same as 7807. 

7809-SD. Same as 7807. 

8289-SD. Clarksburg quad. Hal-Mar Stone quarry, 1 mile 

(1.6 km) east-southeast of Crownover Mill; 0.2 of a mile 

{0.32 km) north of road. Columbus Limestone, Coral Zone 

(Zone C). 

USNM 162588. Core from well in lot 18, tract 4, Perry T., 

Lake County. Specimen from 1430.5 ft. Columbus Limestone, Coral Zone (Zone C). 

OSU LOCALITIES 

17'735. Bellepont, quarry opposite mouth of Mill Creek. Columbus Limestone, Zone C (Stewart~ 1938, p. 11, 99). 

17779. Dublin, quarrr at south edge of town. Columbus Limestone, Zone C (Stewart, 1938, p. 10, 99). 

21336. Well, 3 miles (4.8 km) southwest of Chillicothe (see 

Carman, 1955 for details of core). 

UMMP LOCALITIES 

60484. International Salt Co. well 1 Nagel, Avon T., sec. 28, 

Loraine County (see Janssens, 1970a, p. 25). Specimen from 

from 1,097 ft. Columbus Limestone, Coral Zone (Zone C). 

60485. International Salt Co., Ohio well 1; 2 mile.s (3.2 km) 

southwest of Ashtabula, Ashtabula County. Specimen from 

1,613 ft. Columbus Limestone, upper part. 

MOOSE RIVER BASIN, ONTARIO 

Specimens illustrated or noted from this area in the Hudson 

Bay Lowlands, were described by Cranswick and Fritz (1958) 

and are deposited in the Royal Ontario Museum, Toronto. 

QUEBEC 

Studied specimens are from three areas: St. Georges, Famine 

Limestone; see Oliver, 1971. Lake Memphramagog, Mountain 

House Wharf Limestone; see stratigraphy discussion and 

Boucot and Drapeau, 1968. Gaspe, Grande Greve Limestone; 

see Oliver, 1964. 

VIRGINIA 

Los. 59b of Kindle (field notes 9/15/10) is "Big Stone 

Gap, Va., near old Woolen 'Mills." See Kindle, 1912, p. 51-52. 

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Weisbord, N. E., 1926, Venezuelan Devonian fossils: Bulls. 

Am. Paleontology, v. 11, no. 46, 52 p. 

--- 1968, The Devonian System in western Venezuela, in 

Oswald, D. H., ed., Internat. Svrnposium on the Devonian 

System, Calgary, Alberta, 1967 [Proc.l: Cale:arv, Alberta Soc. P€troleum Geologists, v. 2, p. 215-226 [1967]. 

Wells, J. W., 1937, Individual variation in the rugose coral 

species H eliophyllum halli E & H: Palaeontographica 

Americana, v. 2, no. 6, 22 p., 1 pl., 31 figs. 

--- 1943, The Rio Cachiri section in the Sierra de Perija, 

Venezuela, pt. 2b, Anthozoa: Bulls. Am. Paleontology, v. 

27,no. 108,p. 95-100. \ 

--- 1944, New tabulate corals from the Pennsylvanian of 

Texas: Jour. Paleontology, v. 18, no. 3, lp. 259-262, pls. 

40, 41. 

--- 194 7, Provisional paleoecological analysis of the Devonian rocks of the Columbus Region [Ohio]: Ohio Jour. 

Sci., v. 47, no. 3, p. 119-126. 

White, C. A., 1882; Fossils of the Indiana rocks: Indiana 

Dept. Geol. Nat. History Ann. Rept. 11, p. 347-401. 

Whiteaves, J. F., 1897, The fossils of the Galena-Trenton and 

Black Riv€r Formations of Lake Winnipeg and its vicinity: Canada Geol. Survey, Palaeozoic Fossils v. 3, no. 3, 

p. 129-242. 

--- 1898, On some additional or imperfectly understood 

fossils from the Hamilton formation of Ontario, with a 

revised list of the species therefrom: Canada Geol. Survey, 

Contr. Canada Paleontology 1, p. 361-418. 

--- 1906, Paleozoic fossils-Appendix-Errata et corrigenda: Canada Geol. Survey, Palaeozoic Fossils, v. 3, p. 

341-345. 

Whitfield, R. P., 1882, Palaeontology-Part 3 in Geology of 

Wisconsin, survey of 1873-1879: Wisconsin Geol. Survey, v. 4, p. 163-363. 

--- 1891, Contributions to invertebrate paleontology [of 

Ohio]: New York Acad. Sci., Ann. 5, p. 505-620 (1891); 

or Ohio Geol. Survey, Rept. 7, p. 407-494 (1893). 

--- 1899, List of fossils, types and figured specimens used 

in the paleontological work of R. P. Whitfield, showing 

where they are probably to be found at the present time: 

New York Acad. Sci. Annals 12, p. 139-186. 

Williams, M. Y., 1919, The Silurian geology and faunas of 

Ontario Peninsula and Manitoulin and adjacent islands: 

Canada Geol. Survey Mem. 111, 195 p., 34 pls. 

Winchell, Alexander, 1866, The Grand Traverse region-a report on the geological and industrial resources of the 

counties of Antrim, Grand Traverse, Benzie, and Leelanaw in the Lower Peninsula of Michigan: Ann Arbor, 

97 p. 

Yabe, H. and Hayasaka, I., 1915, Paleozoic corals from 

Japan, Korea and China: Tokyo, Geol. Soc. Japan Jour., 

XXII, p. 55-70, 79-92, 93-109, 127-142. 

Yandell, L. P., and Shumard, B. F., 1847, Contributions to 

the Geology of Kentucky: Louisville, 36 p. 

Zheltonogova, V. A., and Ivania, V. A., 1960, ed., [Subclass 

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D27-D57. 
Page 

A 

Acanthophyllum ---------------------- 28 

Acinophyllum --------- 5, 8, 9, 11, 15, 17, 18, 27, 

40, 41, 44, 46, 47, 49, 52, 59, 

57, 58, 59, 61, 62, 63, 65, 98, 99, 

100, 102, 103, 108, 111, 112, 141 

baculoideum ---------------------- 57 

camselli -------------------------- 56 

crassiseptatum ------------------- 56 

davisi ------------------------- 65, 66, 67 

fasciculum ------------------------ 56 

mclareni ---------- 9, 11, 13, 15, 21, 44, 56, 

57, 61, 69, 67, 70, 134, 135; 

pls. 23,24 

accidens -------------------------- 56 

rectiseptatum --------------------56, 112 

segregatum ------ 8, 9, 10, 11, 14, 15, 21, 32, 

33, 36, 37, 38, 39, 40, 41, 42, 43, 

44, 47, 54, 56, 57, 60, 61, 62, 

63, 64, 67, 100, 103, 133, 134, 

135, 144; pls. 11, 15-22 

variation --------------------- 92 

segregatum-stramineum ----------- 60 

simcoense -------- 5, 10, 15, 2.1, 41, 44, 56, 

5~61,6~6~66,67,13~ 

135, 144; pls. 8-11 

stokesi --------- 5, 10, 15, 21, 41, 44, 56, 58, 

61, 62, 64, 65, 133, 135, 144; 

pls. 25-28 

atramineum -------- 5, 15, 21, 27, 41, 44, 56, 

59, 62, 64, 67, 133, 134, 135; 

pis. 12-14 

vermetum --------- 21, 27, 44, 56, 57; pl. 29 

(Acinophyllum), Cylindrophyllum ------ 53 

simcoense, Cylindrophyllum ------- 57, 61 

acuminatus, Paraspirifer -------------- 12, 19 

adjunctum, Diphyphyllum ------------- 108 

adnatum, Depasophyllum -----------47,49, 50 

Aemulophyllum exiguum ----------- 15, 19, 144 

affinis, Asterobillingsa _ 15, 21, 23, 44, 53, 90, 96 

Billingsastraea ------------------- 96 

Phillipsastraea -------------------- 96 

americanum, Craspedophyllum -----52, 53, 9·8, 

99, 109 

Amherstburg Limestone --------------- 17,29 

Amphigenia -------------------------- 8 

sp ------------------------------- 15 

angustus, Icriodus -------------------- 19 

anna, Hexagonaria -------------------- 87, 88 

Prismatophyllum __ 21, 44, 86, 87; pls. 55, 56 

Stylastrea ------------------------ 87 

annum, Prismatophyllum -------------- 87 

Appalachian Basin --------------- 5, 11, 17, 18 

Appalachians south of New York ------ 15 

arachne, Billingsastraea --------------- 91 

Radiastraea ---------------------- 90 

arachnoideum, Eridophyllum ---------- 79, 108 

Arachnophyllum --------- 89, 91, 109, 111, 130\ 

articulatum, Entelophyllum ----------- 116 

arundinaceum, Diphyphyllum ---- 46, 47, 54, 61 

Diaphyllum ----------------------- 48 

Eridophyllum ------------------ 47, 48, 65 

Macgeea (Synaptophyllum) ------- · 48 

Synaptophyllum ___ 9, 15, 21, 41, 44, 46, 47, 

50, 51, 54, 133, 134, 135; pls. 2-4, 7 

INDE2( 

[Italic numbers indicate major references and descriptions) 

Page 

asiaticum, Cyclindrophyllum ---------- 71 

Eridophyllum ---------··----------- 99 

Aaterobillingsa --------------- 5, 8, 18, 44, 52, 

53, 80, 88, 93, 95, 96, 130 

affinis ------------ 15, 21, 23, 44, 53, 90, 96 

canadensis --------------------- 90, 91,93 

'confiuens -------------·------------ 89 

magdisa ------------- H2, 88, 91, 93, 94, 95, 

pis. 57-64 

magdisa ----------------- 5, 9, 10, 21, 

32,41,44,9~99,9~ 13~ 

135, 144; pis. 57-60 

steorra ----------- 9, 21, 32, 41, 44, 90, 

93, 94, 133, 134, 135; pls. 61-64 

rugosa ------------------- 5, 21, 44, 90,94, 

133, 135; pl. 65 

spp ------------------------------ 92 Asterocycles _ _ _ _ __ _ _ _ _ __ _ _ _ ___ _ __ ___ _ _ 89 

confiuens ------------··------------ 89,91 Astraea _ _ _ _ _ _ _ __ _ __ _ __ _ __ _ __ _ _ _ __ ___ _ _ 96 

rugosa --------------------- 78, 80, 82,94 

astreiforme, Keriophyllum ------------- 90 

Astreoid species ----------------------- 129 

aulodokum, Eridophyllum _ .. _ 11, 21, 45, 98, 105, 

106, 135, pl. 73 

Australophyllum _________ -·- _ __ _ __ _ ____ 80 

Axinura _________________ -·-- _ __ _ __ ___ _ 98 

B 

baculoideum, Acinophyllum ------------ 57 

Synaptophyllum ----------------57, 58, 59 

bartrumi, Eridophyllum ___ .. ____________ 99 

bathycalyx, Hexagonaria -------------- 83 

ponderosa, Hexagonarict ----------- 127 

Bethamphyllum ----------------------- 29 

Billingsastraea --------------52, 53, 88, 89, 90, 

91, 124, 130 

affinis ---------------------------- 96 

arachne -------------------------- 91 

billingsi ------------------------- 91, 129 bohemica _ __ _ _ _ _ __ _ ____ _ __ __ _ _ __ _ _ 90 

branikensis _ _ _ _ __ _ __ _ _ _ _ __ _ __ ___ _ _ 90 

confiuens ------------------------- 91 

ingens ---------------------------- 91 lonuicarinata ___ ___ __ _ _ _ __ _ __ _ __ _ _ 91 

monticula -----------··------------ 91, 129 

monticulum ---------------------- 130 nevadensis _ __ _ __ _ _ _ _ _ _ _ __ ___ _ _ _ _ __ 91 

pauciseptata ---------------------- 91 

rockportensis --------------------- 91 

romingeri __________ -·- __ _ __ ___ _ _ __ 91 

rugosa ------------------------- 32, 94, 95 

scheii ---------------··------------- 91 

southworthi ---------··------------- 91 

stirps --------------··------------- 91 tapetiformis ________ .. _ _ _ __ __ _ ___ _ _ 91 

trichomisca ----------------------- 91 

verneuili -----------·----- 89, 91, 93, 94, 95 verrilli _ __ _ __ _ __ _ __ _ __ _ __ _ __ ___ _ __ 91 

yandelli _ __ _ __ _ __ _ _ _ _ _ _ _ _ _ __ _ __ _ __ 91 

sp ------------------------------ 132, 133 

(Billingsastraea), Phillipsttstraea ------ 88 

verneuili, Phillipsastrctea ---------- 89 

Billingsastraeidae ---------------------- 53 

Page 

billingsi, Billingsastraea -------------- 91, 129 

Heliophyllum --------------------- 130 

Phillipsastraea ------------------ 129, 130 

Biostratigraphy ----------------------- 21 

bohcmica, Billingsastraea -------------- 90 

Bois Blanc Formation ------------- 5, 8, 17, 18 

Boyle Limestone ---------------------- 14 

branikensis, Billingsastraea ------------ 90 

Brevispirifer gregarius ---------------- 12, 15 

Zone ----------------------------- 19 

c 

callawayensis, Diplophyllum ------------ 70 

CampophyUum spongiosum ------------- 68 

camselli, Acinophyllum ---------------- 56 

canadensis, Asterobillingsa --------- 90, 91, 93 

Carinae ------------------------------ 81 

Carlisle Center Formation ------------- 5 

cassum, Heliophyllum ----------- 5, 21, 46,124, 

190, 133, 135; pl. 103 

catenatum, Disphyllum ---------------- 107 

Cazenovia and Onesquethaw stages ---- 2 

Central Ohio stratigraphy ------------- 1B 

Cerioid species ----------------------- 127 

Cherry Valley Limestone Member ------ 11 

Chonetes lineatus Zone ---------------- 87 

cincta, Hexagonaria ---------------- 80, 81, 82 

Prismatophyllum ------------------ 82 

cinctum, Prismatophyllum --------- 78, 80, 82 

Clarence Member --------------- 10, 11, 23,40 

Classification of described corals ------ .u 

coalescens, Diphyphyllum -------------- 108 

Eridophyllum --------------------- 108 

coalita, Hexagonaria ------------------ 78, 127 

coalitum, Cyathophyllum -------------- 127 

Heliophyllum -------- 5, 9, 21, 46, 123, 127, 

129, 133, 135; pls. 96-98 

cohaerens, Cyathophyllum ------------- 69, 70 

Disphyllum ----------------------- 70 

colligatum, Crepidophyllum ------------ 109 

Cyathophyllum ------------------- 82, 109 

Diphyllum ------------------------ 109 

Eridophyllum --------------------- 109 

Grewgiphyllum ------ 5, 9, 10, 21,41 45, 52, 

53, 109, 133, 134, 

135, 144; pis. 76-79 

Heliophyllum ------------------- 108, 109 

Columbus Limestone ------------- 2, 12, 17, 31 

Columnaria coralliferum --------------- 56, 57 

stokesi ---------------------------- 65 

(Palaeophyllum) stokesi ----------- 65 

Columnariidae _____________ ----- _ ----- 46 

compactum, Cylindrophyllum __ 69, 70, 121, 122 

Disphyllum ----------------------- 70 

Heliophyllum ------------------- 121, 122 

confiuens, Asterobillingsa -------------- 89 

Asterocycles ----------------------- 89, 91 

Billingsastraea -------------------- 91 

Cylindrophyllum ----------------- 69, l20 

conigerum, Scenophyllum -------------- 15 

conjunctum, Diphyphyllum ------------ 108 

Eridophyllum _ 45, 98, 108, 135; pis. 107, 108 

Coral morphology --------------------- 31 

coralliferum, Columnaria --------------- 56, 57 

Cyathophyllum -------------------- 122 

153 
154 

Page 

corniculum, Eridophyllum ____ 11, 21, 32, 41, 45, 

98, 106, 135; pls. 74, 75 

Craspedophyllidae ------2,44,51,90,111,117 

Craspedophyllinae ------------ 2, 23, 44, 52,96 

Craspedophyllum ----------- 52, 96, 97, 98, 109 

americanum ------------52, 53, 98, 99, 109 

crassiseptatum, Acinophyllum ---------- 56 

Disphyllum ------------------ 70, 112, 113 

crebrum, Eridophyllum ---------------- 99 

Crepidophyllum -------------------- 96, 97, 98 

colligatum ------------------------ 109 

curta, Hexagonaria -------------------- 85, 87 

Cyathocylindrium ---------- 32, 45, 69, 114, 115, 

11~ 120,121,122, 12~ 127,130 

gemmatum ------------- 9, 21, 45, 117, 120, 

123, 133, 135; pls. 87-90 

opulens ____________ 9, 11, 15, 21, 41, 45, 69, 

117, 122, 123, 124, 133, 

134, 135; pls. 80-86 

n. sp. A ----- 11, 21, 45, 117, 122, 135; pl. 91 

Cyathophyllidae ----------------------- 113 

Cyathophyllum -------------- 113, 114, 115, 123 

coalitum _ __ ___ _ __ _ __ ___ _ _ __ __ ___ __ 127 

cohaerens ------------------------- 69,70 

colligatum ----------------------- 82, 109 

coralliferum ---------------------- 122 

marylandicum -------------------- 47 

oedipus ------------------------- 120, 122 ovoideum _ __ ___ _ __ _ _ _ ____ __ _ __ ___ _ 83 

proliferum _ __ _ __ _ __ _ ____ _ __ _ __ ___ _ 117 

rugosum ------------------- 78, 79, 80, 82, 

83, 94, 95, 96 

septa tum ___ _ __ _ _ _ _ ____ ___ _ _ _ ___ _ _ 70 

(Heliophyllum) -------------------- 123 

Cylindrohelium ------------------------ 68, 69 

heliophylloides -------------------- 69 

profundum ------------------------ 68, 69 

Cylindrophyllinae ---------------- 2, 44, 52, 53 

Cylindrophyllum ----------------- 8, 18, 40, 44, 

52,5~55,68,73,7~75,77, 

111,112,113,117,124 

asiaticum ------------------------- 71 

compactum --------------- 69, 70, 121, 122 

confluens ------------------------ 69, 120 

deearium ---------------- 9, 11, 15, 21, 41, 

44, 69, 72, 76, 133, 

134, 135; pls. 41-43 

delicatulum __________ ---- ___ _ __ _ __ 69 

elongatum ------------ 9, 21, 41, 44, 68, 69, 

71, 74, 75, 76, 77, 133, 

135; pis. 31-34 

floydense ------------------------- 70 

grabaui -------------------------- 69 gruensis ___ _ __ _ __ ___ _ _ ____ __ _ __ _ __ 69 

hindshawi ------------------------ 69 

laxtensum ------------ 5, 21, 44, 69, 72, 73, 

75, 133, 135; pis. 39, 40 

magnum -------------------------- 69 

panicum -------------------------- 69, 70 

planivesiculosum ------------------ 71 

propinquum ---------- 9, 21, 41, 44, 69, 70, 

71, 72, 73, 76, 77, 133, 134, 

135; pis. 35-38, 45 

stummi ----------------------- 16, 70, 113 

(Acinophyllum) ------------------- 53 

simcoense ---------------------57, 61 

sp. A ---------------- 9, 21, 44, 69, 74, 77, 

133, 135; pl. 34 

spp ------------------------------ 72, 112 

Cystiphylloides ------------------------ 144 

D 

davisi, Acinophyllum ------------------ 65 

deearium, Cylindrophyllum ____ 9, 11, 15, 21, 41, 

44, 69, 72, 76, 133, 134, 

135; pis. 41-43 

Delaware Limestone ------------------- 12, 18 

delicatulum, Cylindrophyllum ---------- 69 

Depasophyllum ------------------------ 46,47 

adnatum ----------------------- 47, 49, 50 

INDEX 

Page 

Depositories _______________________ ---- 43 

Detroit River Group ------------------ 17, 18 

Diphyphyllum adjunctum -------------- 1108 

arundinaceum --------------46,47, 54, 61 

coalescens ------------------------- 108 

conjunctum ----------------------- 108 

gigas ----------------------------- 67, 69 gracile _ __ _ __ _ _ _ _ __ __ _ __ _ __ _ __ _ ___ 59 

minus ---------------------------- 55 

rectiseptatum -------------------- 70, 112 

simcoense ------------------57, 58, 61, 64 

stokesi --------------------------- 65 

stramineum --------------------59, 61, 67 tubiforme _ _ _ ___ _ __ _ _ _ __ _ __ __ _ _ __ _ _ 71 

vermetum -------------------------59, 67 

Diplophyllum callawayensis ------------ 70 

Disphyllidae -----------------------45,52, 111 

Disphyllinae -------------------------- 52 

Disphyllum ------------------ 16, 27, 28, 45, 46, 

arundinaceum 

52, 55, 56, 63, 68, 

69, 70, 71, 111, 112 

48 

catenatum ------------------------ 107 cohaerens _ __ _ __ ___ _ _ _ _ _ _ _ __ _ __ ____ 70 

compactum ----------------------- 70 

crassiseptatum --------------- 70, 112, 113 dyeri _ _ _ ___ _ __ _ _ _ _ __ _ __ _ __ ___ _ ____ 710 

nevadense _ _ __ __ ___ _ _ _ __ __ _ ___ _ _ _ __ 70 

occidens _ __ __ _ __ _ __ _ __ _ __ __ _ ___ _ __ 69 

rectiseptatum ------- 12, 21, 45, 112,· pl. 30 

stummi -------------- 12, 16, 21, 29, 45, 52, 

69, 72, 113, 135; pls. 44, 45 

synaptophylloides --------------- 57, 67, 70 

(Synaptophyllum) stramineum _____ 59, 61 

sp. A ----------------------------- 63, 70 

sp. B ----------------------------- 70, 73 

Dundee Limestone --------------------- 16, 18 

dyeri, Disphyllum --------------------- 70 

E 

Eastern North America ---------------- 4 

Eddastraea --------------------------- 130 

Edgecliff bioherm facies --------------- 37,41 

Edgecliff biostrome facies ------------- 37,41 

Edgecliff Member of Onondaga 

Limestone ----------- 8, 11, 19, 23 

elongatum, Cylindrophyllum _____ 9, 21, 41, 44, 

68, 69, 71, 74, 75, 76, 77, 

133, 135; pis. 31-34 

Entelophyllum ----------------------- 70, 115 articulatum _ __ _ __ _ _ _ _ _ _ __ _ ____ _ ___ 116 

Eridophyllum --------- 11, 12, 18, 21, 23, 29, 40, 

41, 44, 52, 53, 58, 63, 96, 

!tOO, 101, 102, 103, 105, 106, 

107, 108, 109, 128, 141 

arachnoideum -------------------- 79, 108 

arundinaceum ------------------47,48, 65 

asiaticum _ __ _ _ _ _ __ _ ____ ___ _ __ _ ___ _ 99 

aulodokum ------------------11,21,45,98, 

105, 106, 135; pl. 73 

bartrumi ------------------------- 99 coalescens _ __ _ __ ___ _ __ _ ____ _ __ _ _ _ _ _ 108 

colligatum _ __ _ __ _ __ __ _ _ __ ___ _ ___ __ 109 

conjunctum ___ 45, 98, 108, 135; pis. 1107, 108 

corniculum -------------- 11, 21, 32, 41, 45, 

98, 106, 135; pis. 74, 75 

crebrum -------------------------- 99 immersum ___ ___ _ _ _ _ ____ _ __ ___ _ __ _ 99 

seriale ------------ 11, 12, 17, 18, 21, 29, 32, 

44, 52, 53, 97. 98, 99, 101, 

102, 105, 108, 135; pis. 67-69 

population A ----------------- 100 

population B ____ 11, 32, 41, 100, 102, 

135; pis. 68, 69 

simcoense --------------- 46, 53, 54, 57, 61 

subseriale ------------ 9, 21, 41, 44, 45, 100, 

108, 133, 134, 135; pis. 70-72 

verneuilanum ------------------ 97, 98,99 

sp ------------------- 9, 11, 12, 15, 40, 104 

Page 

Esopus Formation -------------------- 3, 6 

Etymothyris Zone --------------------- 96 

exiguum, Aemulophyllum ---------- 15, 19, 144 

F 

Falls of the Ohio, Indiana-Kentucky ___ 19, 146 

Famine Limestone -------------------- 16, 27 

fasciculum, Acinophyllum ------------- 56 

Favastraea rugosa --------------------- 94 

Favosites _ _ __ __ _ _ ___ _ __ _ _ _ _ __ _ _ _ ___ ___ 17 

fecundum, Heliophyllum --------------- 122 

fencse, Phacellophyllum --------------- 66 

fioydense, Cylindrophyllum ------------ 70 

Frechastraea -------------------------- 90 

G 

gemmatum, Cyathocylindrium ___ 9, 21, 45, 117, 

120, 123, 133, 135; pis. 87-90 

Heliophyllum ------------------- 120, 122 

Geoaraphic and stratigraphic variation _ 96 

gigantea, Siphonophrentis -------------- 29 

Zaphrentis ------------------------ 29 

gigas, Diphyphyllum ------------------ 67, 69 

Phillipsastraea -------------- 130, 132, 133 

grabaui, Cylindrophyllum ------------- 69 

Synaptophyllum ------------------- 15,47 

gracile, Diphyphyllum ----------------- 69 

gradatum, Heliophyllum ---------- 69, 74, 122 

Grand Tower area, Illinois-Missouri ---- 19 

Grand Tower Limestone --------------- 19 

gregarius, Brevispirifer --------------- 12, 16 

Grewgiphyllum -------- 8, 21, 23, 45, 53, 96, 108 

colligatum ------------------- 5, 9, 10, 21, 

41, 45, 52, 63, 109, 133, 

134, 135, 144; pis. 76-79 

gruensis, Cylindrophyllum ------------- 69 

H 

halli, Heliophyllum _ 15, 27, 28, 113, 114, 116, 123 

Hamilton Group ----------------------- 4 

Haplothecia --------------------------- 90 

helianthoides, Strombodes -------------- 123 

Heliophylloides ----------------------- 29, 114 

heliophylloides, Cylindrohelium --------- 69 

Heliophyllum ------------ 17, 21, 23, 27, 29, 31, 

32, 46, 78, 91, 113, 

114, 115, 116, 117. 129, 

126, 127, 129, 130, 131 

billingsi -------------------------- 130 

cassum --------------------- 5, 21, 46, 124 

180, 133, 135; pl. 103 

coalitum ----------------- 6, 9, 21, 46, 123, 

127, 129, 133, 135; pis. 96-98 

colligatum ---------------------- 108, 109 

compactum ---------------------- 121, 122 

fecundum ------------------------- 122 

gemmatum ---------------------- 120, 122 

gradatum --------------------- 69, 74, 122 

halli ---------- 15, 27, 29, 113, 114, 116, 123 

ingens _ _ _ _ __ _ _ _ _ _ _ __ _ __ _ __ ___ _ ___ _ 130 

latericrescens ------------------- 117, 120 

megaproliferum ------------ 9, 21, 45, 123, 

124, 133, 135; pis. 92, 93 

monticulum ---------- 9, 21, 41, 46, 91, 124, 

127, 129, 133, 135; pis. 99-102 

pocillatum _ __ _ __ _ __ _ _ _ _ ___ __ ___ _ __ 122 

procellata _ __ ___ _ __ _ __ _ _ _ _ __ _ __ __ __ 96 

procellatum --------- 5, 21, 46, 92, 95, 124, 

131, 133, 135; pis. 104-106 

proliferum ---------------- 12, 16, 117, 120 yandelli ___ _ __ _ __ _ _ ___ __ __ __ __ _ __ _ 1310 

n. sp. B --------------- 9, 21, 45, 123, 124, 

125, 133, 136; pl. 94 

n. sp. C ----------- 9, 21, 45, 123, 125, 126, 

133, 135; pl. 95 

sp ------------------------- 5, 21, 126, 128 

(Heliophyllum), Cyathophyllum -------- 123 
Page 

Heterophrentis ------------------------ 27, 28 

prolifica -------------------------- 29 venezuelensis _ __ __ _ _ __ _ _ ___ _ __ ___ __ 27 

sp -------------------------------- 29 

Hexagonaria ---------------52, 78, 80, 111, 124 

anna ----------------------------- 87, 88 

bathycalyx ------------------------ 83 ponderosa _ __ ___ _ __ _ __ __ __ __ _ _ 127 

cincta ------------------------- 80, 81, 82 

coalita --------------------------- 78, 127 

curta ----------------------------- 85, 87 martinsoni ___ _ __ _ _ __ _ __ _ __ ___ _ __ __ 120 

mesocincta ----------------- 79, 81, 82, 83 

ovoidca --------------------------- 83 

ponderosa ----------------------- 127, 129 

prisma ---------- 17, 79, 80, 81, 82, 83, 108 

truncata ----------------------- 85, 87, 88 

Cylindrophyllum ------------------ 69 

Homalophyllum sp -------------------- 15 

Icriodus angustus --------------------- 19 

Illinois Basin stratigraphy ------------- 18 

immersum, Eridophyllum -------------- 99 

ingens, Billingsastraea ---------------- 91 

Heliophyllum --------------------- 130 

Intercolony and intracolony variation __ 92 

lowaphyllum -------------------------- 90 

J 

Jeffersonville Limestone ------------- 2, 14, 19 

jevlanensis, Peneckiella ---------------- 56 

K 

Kentucky, east-central, stratigraphy __ 14, 145 

Keriophylloides ------------------------ 89, 90 

Kcriophyllum astreiforme -------------- 90 

Kionelasma --------------------------- 29 

kladion, Syanptophyllum ------ 9, 11, 21, 44, 47, 

49, 50, 134, 135; pis. 6, 7 

kunthi, Peneckiella minor -------------- 55 

Kwataboahegan Formation ------------ 17 

L 

latericrescens, Heliophyllum ---------- 117, 120 

latcseptata, Sudetia -------------------- 55 

laxtensum, Cylindrophyllum __ 5, 21, 44, 69, 72, 

73, 75, 133, 135; pis. 39, 40 

Lithostrotion stokesi ------------------ 65 

Locality data ------------------------- 199 

longicarinata, Billingsastraea ---------- 91 

Lucas Formation ---------------------- 1'r 

Lucas Limestone ---------------------- 17 

M 

Macgeea (Synaptophyllum) 

arundinaceum ------------- 48 

Mackinac Straits, Michigan, 

stratigraphy -------------- 18 

mclareni, Acinophyllum ______ 9, 11, 13, 15, 21, 

44,5~5~6~6~ 67, 

134, 135; pis. 23, 24 

magdisa, Asterobillingsa -------- 32, 88, 91, 93, 

94, 95; pis. 57-64 

Asterobillingsa magdisa ___ 5, 9, 10, 21, 32, 

41, 44, 90, 99, 94, 133, 

135, 144; pis. 57-60 

magdisa Asterobillingsa ____ 5, 9, 10, 21, 32, 

41, 44, 90, 99, 94, 133, 

135, 144; pis. 57-60 

steorra, Asterobillingsa ------ 9, 21, 32, 41, 

44, 90, 93, 94. 133, 134, 

135; pis. 61-64 

magnum, Cylindrophyllum ------------- 69 

Marcellus Shale ----------------------- 11, 21 

Marisastridae ------------------------- 90 

INDEX 

Page 

Marisastrum --------------··----------- 90 

Martinophyllum ornatum --------------- 90 

martinsoni, Hexagonaria --------------- 120 

marylandicum, Cyathophyllum --------- 47 

megaproliferum, Heliophyllwm __ 9, 21, 45, 123, 

124, 133, 135; pis. 92, 93 

mesocincta, Hexagonaria -------- 79, 81, 82, 83 

Prismatophyllum ------------------- 82 

Michigan, southeastern, stratigraphy -- 18, 145 

Michigan Basin ----------------------- 18, 23 

stratigraphy ----------------------- 17 Microstructure __________ --·--- ____ -- _ -- 91 

minima, Peneckiella ------------------- 56 

minor kunthi, Peneckiella --·------------ 55 

minus, Diphyphyllum ------·------------ 55 

monticula, Billingsastraea ------------- 91, 129 

monticulum, Billingsastraea ------------ 130 

Heliophyllum ____ 9, 21, 41, 46, 91, 124, 127, 

129, 133, 135, 144; pis. 99-102 

Moorehouse Member ---------------- 11, 18,41 

Moorehouse western coral facies ------- 37, 39 

Morphology, coral --------------------- 91 Mountain House Wharf Limestone ____ 15, 27 

multicaule, Syringopora --------------- 56, 57 

N 

nalivkini, Peneckiella ------------------ 56 

Nedrow Member ---------------- 10, 23, 37, 40 

nevadense, Disphyllum ----------------- 70 

nevadensis, Billingsastraea ------------ 91 

New localities ------------------------- 199 

New York and Niagara Peninsula, 

Ontario ------------------ 5 

New York localities ------------------- 199 

New York material, specimen 1 ---- 100; pl. 67 

specimen 2 ------------------- 101; pl. 67 

Niagara Peninsula localities ----------- 149 

0 

obliquum, Schistotoecholasmrt ---------- 98 

obliquus, Schistotoecholasma ______ 99, 100, 102 

occidens, Acinophyllum ---------------- 56 

Disphyllum ----------------------- 69 

oedipus, Cyathophyllum ------------- 120, 122 

Ohio, central, stratigraphy------------- 12 

northeastern, stratigraphy --------- 19 

northwestern, stratigraphy -------- 18 Old localities ___ _ __ _ __ _ __ _ __ _ __ _ _ _ _ _ _ _ 149 

Olentangy Shale ---------------------- 12 Onesquethaw and Cazenovia Stages ____ !2 

Onesquethaw Stage, age -------------- 19 

Onondoga Formation ------------------ 9 

Onondoga Limestone ------------------- 4, 18 

Ontario, central southwestern --------- 144 

southwestern localities ------------- 149 

opulens, Cyathocylindrium ··--- 9, 11, 15, 21, 41, 

45, 69, 117, 122, 123, 124, 

133, 134, 135; pis. 80-86 

ornatum, Martinophyllum -------------- 90 

ovoidea, Hexagonaria ----·------------- 83 

ovoideum, Cyathophyllum --·------------ 83 

Prismatophyllum -----·- 9, 11, 21, 41, 44, 78, 

82, 89, 134, 135; pis. 49-52 

p 

Paleobiogeography --------------------- !29 

Palaeophyllum ------------------------- 46, 56 

stokesi --------------·------------- 65 (Palaeophyllum) stokesi, Columnaria ___ 65 

panicum, Cylindrophyllum ------------- 69, 70 Paraspirifer __________________________ 85, 98 

acuminatus ----------------------- 12, 19 

Zone ----------------------------- 99 

pauciseptata, Billingsastraea ---------- 91 

Peneckiella --------------------------- 55, 56 

jevlanensis ----------------------- 56 

minima --------------------------- 56 

minor lcunthi --------------------- 55 nalivkini ___ _ _ ____ ___ _ _ ___ _ __ _ __ _ _ 56 

155 

Page 

Phaceloid species ---------------------- 124 

Phacellophyllum ---------------- 46, 47, 55, 56 

fenese ---------------------------- 55 

Phaulactis ---------------------------- 115 

Phillipsastraea ------------------------ 90 

affinis ---------------------------- 96 

billingsi ------------------------ 129, 13•0 

gigas ----------------------- 130, 132, 133 

verneuili -------------------------- 89,91 

(Billingsastraea) ------------------ 88 

verneuili ------------------------- 89 

Phillipsastraeidae --------------------- 31, 52 

phrygia, Zaphrentis ------------------ 29, 114 

Placophyllum ----------------------- 46, 47, 50 

tabulatum ---------------------- 47, 49,50 

sp -------------------------------- 50 

Planetophyllum ----------------------- 55 

planivesiculosum, Cylindrophyllum ----- 71 

Plum Brook Shale --------------------- 12 

pocillatum, Heliophyllum -------------- 122 

ponderosa, Hexagonaria ------------- 127, 129 

Hexagonaria bathycalyx ----------- 127 

prisma, Hexagonaria __ 17, 79, 80, 81, 82, 83, 108 

Prismatophyllum ------ 9, 17, 21, 41, 44, 78, 

79, 83, 85, 96, 135; pis. 46-48 

Prismatophyllum ------------ 12, 17, 18, 19, 21, 

23,44,52,53, 7~80,82, 

83,85,86,87,95,96, 109,111 

anna ------------- 21, 44, 86, 87; pis. 55,56 

annum --------------------------- 87 

cincta ---------------------------- 82 

cinctum ----------------------- 78, 79, 82 mesocincta _ __ _ _ _ ___ _ _ _ _ _ _ __ _ _____ _ 82 

ovoideum --------- 9, 11, 21, 41, 44, 78, 82, 

89, 134, 135; pis. 49-52 

prisma ------------ 9, 17, 21, 41, 44, 78, 79, 

83, 85, 96, 135; pis. 46-48 

Zone ------------------------- 19 

rugosum -------------------------- 79 

subcincta ------------------------- 82 

truncata --------------- 11, 12, 21, 44, 78, 

85, 135; pis. 53, 54 

procellata, Heliophyllum --------------- 95 

procellatum, Heliophyllum ---- 5, 21, 46, 92, 95, 

124, 181, 133, 135; p)s. 104-1106 

profundum, Cylindrohelium ------------ 68, 69 

proliferum, Cyathophyllum ------------ 117 Heliophyllum _______ : _____ 12, 16, 117, 120 

prolifica, Heterophrentis --------------- 29 

propinquum, Cylindrophyllum ---- 9, 21, 41, 44, 

69, 70, 71, 72, 78, 76, 77, 

133, 134, 135; pis. 35-38, 45 

Q 

Quebec stratigraphy 16 

R 

Radiastraea --------------------- 88, 89, 90,91 arachne _ __ ___ _ __ _ __ _ _ ____ _ _ _ ___ _ __ 90 

rectiseptatum, Acinophyllum ---------56, 112 

Diphyphyllum -------------------- 70, 112 

Disphyllum ---------- 12, 21, 45, 112,· pl. 30 

Synaptophyllum ------------------- 112 

Regional stratigraphy ----------------- 6 

rockportensis, Billingsastraea ---------- 91 

Rogers City Limestone ---------------- 18 

romingeri. Billingsastraea ------------- 91 

rugosa, Asterobillingsa ------- 5, 21, 44, 90, 94, 

95, 133, 135; pl. 65 

Astraea -------------------- 78, 80, 82,94 

Billingsastraea ----------------- 32, 94, 95 

Favastraea ----------------------- 94 

rugosum, Cyathophyllum -------- 78, 79, 80, 82, 

83, 94, 95, 96 

Prismatophyllum ------------------ 79 

s 

Scenophyllum 

conigcrum ------------------------

29 

15 
156 

Page 

scheii, Billingsastraea ----------------- 91 

Schistotoecholasma -------------------- 97,98 

obliquum -------------------------- 98 

obliquus --------------------- 99, 100, 102 typicale _ __ _ __ _ __ ___ _ __ ___ _ _ _ ___ _ __ 98 

typicalis --------------------- 99, 100, 102 

Schoharie Formation ------------------ 3 

Schoharie Grit ------------------------ 4, 8 

segregatum, Acinophyllum _____ 8, 9, 10, 11, 14, 

15, 21, 32, 33, 36, 37, 

38, 39, 40, 41, 42, 43, 

44, 47, 54, 56, 57, 60, 

61, 62, 63, 64, 67. 1100, 

103, 133, 134, 135, 144; 

pls. 11, 15-22 

Synaptophyllum ------------------- 61,62 

segregatum-stramineum, Acinophyllum__ 60 

Sellersburg Limeston<! ----------------- 19 

Seneca Member of Onondaga 

Limestone -------------- 4, 11, 18 

septatum, CyathophyUum -------------- 70 

seriale, Eridophyllum ----- 11, 12, 17, 18, 21, 29, 

32, 44, 52, 53, 97, 98, 

99, 101, 102, 105, 108, 

135; pls. 67-69 

Silica Formation ---------------------- 18 

Silver Creek Limestone Member ------- 19 

simcoense, Acinophyllum ------ 5, 10, 15, 21, 41, 

44,5~5~ 61,62,63, 6~ 

67, 133, 135, 144; pls. 8-11 

Cylindrophyllum (Acinophyllum) ___ 57, 61 

Diphyphyllum --------------- 57, 58, 61, 64 

Eridophyllum ------------46, 53, 54, 57,61 

Synaptophyllum ------------57,60, 61, 62, 

63, 64,65, 66 

Siphonophrentis ---------------------- 27, 29 gigantea _ _ _ _ __ _ __ ___ _ __ _ __ _ __ ____ _ 29 

sp -------------------------------- 15, 29 

Skaneateles Formation ---------------- 12 

southworthi, BiUingsastraea ----------- 91 

Spinophyllum ------------------------- 68, 69 

spongiosum ----------------------- 68 

sponuiosum, Campophyllum ------------ 68 

Spinophyllum --------------------- 68 

Springvale Sandstone ----------------- 8 Stauria _ _ __ __ _ __ _ _ _ ___ _ __ _ __ ___ ___ _ ___ 46 

Stauriidae -------------------------- 2, 44,46 

stirps, Pillingsastraea ----------------- 91 

steorra, Asterobillingsa magdisa __ 9, 21, 32, 41, 

44, 90, 93, 94, 133, 134, 

135; pls. 61-64 

stckesi, Acinophyllum ----- 5, 10, 15, 21, 41, 44, 

56, 58, 61, 62, 64, 65, 

133, 135, 144; pls. 25-28 

Columnaria ----------------------- 65 

(Palaeophyllum) -------------- 65 

Diphyphyllum -------------------- 65 

INDEX 

Page 

stokesi, Acinophyllum-Continued 

Lithostrotion --------------------- 65 

Paleophyllum --------------------- 65 

Stopping River Formation ------------ 17 

stramineum, Acinophyllum ____ 9, 15, 21, 27, 41, 

44, 56, 59, 62, 64, 67 .. 133, 

134, 135; pls. 12-14 

Diphyphyllum -----------------59, 61, 67 

Disphyllum (Synaptophyllum) ----- 59, 61 

Synaptophyllum ---------------- 59, 60, 62 

Thamnophyllum -------------------59, 61 

Stratigraphic and geographic 

variation ----------------- 96 

Stratigraphy, regional ---------------- 5 

Stromatoporoid Zone ----------------- 19 

Strombodes helianthoides -------------- 123 

Stropheodontid Zone ------------------ 19 

stummi, Cylindrophyllum --------- 16, 70, 113 

Disphyllum --------- 12, 16, 21, 29, 45, 52, 

69, 72, 119, 135; pls. 44, 45 

Stylastrea anna ----------------------- 87 

subcincta, Prismatophyllum ----------- 82 

subseriale, Eridophyllum ------ 9, 21, 41, 44, 45, 

100, 109, 133, 134, 135; 

pls. 70-72 

Sudetia lateseptata -------------------- 55 

Summary and conclusions ------------- 42 

synaptophylloides, Disphyllum -------57,67, 70 

Synaptophyllum ------- 11, 31, 410, 44, 46, 48, 49, 

50, 51, 53, 54, 55, 56, 57. 70 

arundinaceum ----- 9, 15, 21, 41, 44, 46, 47, 

50, 51, 54, 133,134, 135; 

pls. 2-4, 7 

baculoideum --------------------57, 58, 59 

grabaui -------------------------- 15,47 

kladion ------------ 9, 11, 21, 44, 47, 49, 50, 

134, 135; pls. 6, 7 

rectiseptatum --------------------- 112 

segregatum ----------------------- 61, 62 

simcoense ------57, 60, 61, 62, 63, 64, 65, 66 

stramineum --------------------59, 60, 62 

tabulatum ------------ 9, 21, 44, 47, 49, 51, 

133, 134, 135; pl. 5 

vermetum -------------------------59, 67 

sp -------------------------------- 17 

( Synaptophyllum) arundinaceum, 

Macgeea ------------------ 48 

stramineum, Disphyllum -----------59, 61 

Syringopora multicaule ----------------56,57 

Systematic paleontology --------------- 49 

Systematic work, previous ------------- 90 

T 

tabulatum, Placophyllum ---------------47, 50 

Synaptophyllum ------- 9, 2:!., 44, 47, 49, 51, 

133, 134, 135; pl. 5 

Page 

Taimyrophyllum ---------------------- 130 

tapeti!ormis, Billingsastraea ---------- 91 

Terminology -------------------------- 92 

Thamnophyllum ----------------------- 55 

stramineum -----------------------59, 61 

Tioga Bentonite Bed ---------------- 4, 11, 18 

Tipheophyllum ------------------------ 99 

trichomisca, Billingsastraea ----------- 91 

truncata, Hexagonaria -··----------- 85, 87,88 

Prismatophyllum ----- 11, 12, 21, 44, 78, 85, 

135; pls. 53, 54 

Tryplasma rhopalium ------------------ 47 

tubiforme, Diphyphyllum -------------- 71 

typicale, Schistotoecholasma ----------- 98 

typicalis, Schistotoecholasma _____ 99, 1QO, 102 

u 

Union Springs Shale Member 11 

v 

Variation, Acinophyllum segreoatum __ 92 

individual ------------------------- 9!2 

in other species -----------· ------- 40 

venezuelensis, Heterophrentis ---------- 27 

vermetum, Acinophyllum -------- 21, 27, 44, 56, 

57; pl. 29 

Diphyphyllum --------------------- 59, 67 

Synaptophyllum -------------------59, 67 

verneuilanum, Eridophyllum --------- 97, 98, 99 

verneuili, Billingsastraea ----- 89, 91, 93, 94, 95 

Phillipsastraea -------------------- 89,91 

(Billingsastraea) -------------- 89 

verrilli, Billingsastraea ---------------- 91 

w 

Wildcat Valley Sandstone -------------- 15 

Woodstock area, Ontario, stratigraphy __ 17 

y 

yandelli, Billingsastraea --------------- 91 

Heliophyllum --------------------- 130 

z 

Zaphrentidae ----------------------- 2, 45, 119 

Zaphrentis ------------------ 28, 113, 114, 115 

gigantea -------------------------- 29 

phrygia -------------------------- 29, 114 

sp -------------------------------- 15 

Zonophyllum -------------------------- 27 

sp -------------------------------- 27 

*U.S.GOVERNMENT PRINTING OFFICE: 1976 0-211-317/122 
PLATES 1-108 

All figures on plates 2-108 are unretouched photographs of thin sections unless otherwise 

indicated. 

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

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

Reconstruction of Edgecliff bioherm scene based on fossils collected from bioherm near Buffalo, N.Y. (Old Fogelsanger quarry, Ioc. 108). Skeletalform 

is accurately shown as molds were prepared from actual specimens. Soft parts (polyps) are based on living forms in analogous environments and are, at best, 

reasonable guesses. A key to the colonial rugose corals represented is given below. See plate 1B for detail views and key to other invertebrates. 

Models and diorama are by George Marchand in the Hall of Invertebrate Paleontology, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. Photographed by Victor Krantz, Smithsonian Institution. 

Key to colonial rugose corals: 1, Cyatkocylindrium gemnwtum (Hall); 2, Heliopkyllum monticulum (Ehlers and Stumm); 3, P1·ismatopkyllum ovoidewrn 

(Davis); 4, Acinopkyllum segregatum (Simpson); 5, Asterobillingsa magdisa. Oliver. Scale: diameter of C. gemmatunn corallite (1 in key) is approximately 1 inch (2% em). 
GEOLOGICAL SURVEY 

I 

\: 

' 

RECONSTRUCTION OF EDGECLIFF BIOHERM SCENE 
PLATE lB 

Details of Edgecliff bioherm reconstruction .shown in plate 1A. Models based on specmens from locality 108. A key to the colonial rugose •corals and other fossils represented 

is given below. Photographs by Victor Krantz, Smithsonian Institution. 

Key 

COLONIAL RUGOSE CORALS: 1, Cyathocylindrium gemmatum (Hall); 2, Heliophyllum mont·iculum (Ehlers and Stumm); 3, Prismatophyllum ovoideum (Davis); 4, 

Acinophyllum segregatum (Simpson); 5, Aste1·obillingsa magdisa Oliver. 

OTHER RUGOSE CORALS: a, Heliophyllum halli Milne-Edwards and Haime; b, 

Siphonoph1·entis elongata (Rafinesque and Clifford); c, Zaphrentis? sp. 

TABULATE CORALS: d, Alveolites mordax Davis; e, Emmonsia epidermata (Rominger); f, T1·achypora sp.; d and fare not known to occur in the Edgecliff Member. 

BRYOZOAN: g, Fenestellate. 

GASTROPOD: h, PlatyceTas (Platyostoma) lineata Conrad. 

NAUTILOID CEPHALOPODS: i, "Michelinoceras" sp.; j, Goldringia sp. 

TRILOBITES: k, Terataspis grandis (Hall); 1, Calymene ptatys Green; m, Phacops 

cristata Hall; the trilobite in the center foreground of plate 1A is Anchiopella anchiops 

(Green). All four species are characteristic of the Schoharie and Bois Blanc Formations 

and are not certainly known from the Edgecliff Member. 

BLASTOID: n, Br·achyschisma corrugatum (Reimann). 

CRINOIDS: o, Dolatoerinus sp.; p, Arachnocrinus sp. 

(Identifications made or confirmed by E. L. Yochelson, gastropod; Mackenzie Gordon, Jr., 

Goldringia; M. E. Taylor, trilobites; D. B. Macurda, echinoderms; T. L. Chase, University 

of Michigan, kindly sent me a list prepared by I. G. Reimann, of the Marchand molds, 

which· confirmed most of the identifications and added the Michelinoceras and 

A 1'achnoc?·inw3.) 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 69 PLATE lB 

DETAILS OF EDGECLIFF BIOHERM RECONSTRUCTION 
PLATE 2 

FIGURES 1-8. Synaptophyllum arundinaceum (Billings) (p. 47). 

Edgecliff Member, Onondaga Limestone, southwestern Ontario. 

1-5. Lectotype, GSC 3602. Locality 3 miles ( 4.8 km) west of Cayuga. 

1, 3. Longitudinal and transverse thin sections of two corallites 

(X 4). 

2, 4. Longitudinal and transverse thin sections of another 

corallite ( X 4) . 

5. Exterior ( X 1) . 

6. GSC 31151. Longitudinal thin section (X 5); see also pl. 4, fig. 

10. Loc. GSC 22795. 

7. GSC 31152. Transverse thin section (X 2%). Loc. GSC 2645. 

8. GSC 3422a. Exterior of fragment (X 1). Cayuga, Ontario. 
GEOLOGICAL SURVEY 

2 

5 

6 8 

SYNAPTOPHYLLUM A R UNDINACEUM (BILLINGS) 
PLATE 3 

FIGURES 1-5. Synaptophyllum arundinaceum (Billings) (p. 47). 

USNM 10719, Edgecliff bioherm facies; loc. 108. See also plate 4, 

figure 9. 

1. Detail (X 50) of one corallite in upper right part of fig. 2. 

2. Transverse thin section ( X 2%). 

3-5. Longitudinal thin sections (X 5). 
GEOLOGICAL SURVEY 

3 

4 

2 5 

S 1'XA P T OPH Y L1--UM A R UNDINACEUJv! (BILLINGS ) 
PLATE 4 

FIGURES 1-10. Synaptophyllum arundinaceum (Billings) (p. 47). 

Edgecliff bioherm facies. 

1. NYSM 12822. Longitudinal thin section (X 5). Loc. 81. 

2-5. NYSM 1282,1. Transverse thin sections of three corallites (X 

5) and part of corallite in fig. 2 (X 50). Loc. 108. 

6. USNM 162577. Transverse thin section (X 5). Loc. 438. See 

also pl. 7, figs. 7-9. 

7, 8. USNM 162576. Serial transverse sections of laterally offsetting 

corallite ( X 5) . Loc. 236. 

9. USNM 10719. Transverse thin section (X 5). These three 

corallites are top center in pl. 3, fig. 2. 

10. GSC 31151. Transverse thin section (X 2%). Same corallum 

as pl. 2, fig. 6. 
GEOLOGICAL SURVEY 

9 10 

SL\lA PTOPHYLL UM ARUNDINACEUM (BILLINGS) 
PLATE 5 

FIGURES 1-8. Synaptophyllum tabulatum (Simpson). (p. 49). 

Edgecliff Member, Onondaga Limestone. 

1-3. Holotype, NYSM 300 (and 12875, see text). Transverse and 

two longitudinal sections presumed to be from the same corallite 

or corallum ( X 5). Walpole, Ontario. 

4-8. USNM 162607. Loc. 244. 

4-7. Longitudinal and transverse thin sections of different 

corallites ( X 5) . 

8. Part of corallite shown in fig. 7 (X 50). 
PROFESSIONAL PAPER 869 PLATE 5 

8 7 

SYNAPTOPJIYLL UM TABULATUM (SIMPSON) 
PLATE 6 

FIGURES 1-1'0. Synaptophyllum kladion n. sp. (p. 50). 

Moorehouse Member, New York and Ontario. 

1-6. Holotype, GSC 31153. Loc. C18. 

1, 6. Longitudinal thin sections (X 2%). 

2-5. Transverse and Longtudinal thin sections ( X 5). 

7. Paratype, USNM 16259·2. Silicified and weathered specimen 

showing characteristic growth form (X 1). Loc. 120. 

8-10. Two paratypes, USNM 162593. Silicified fragments showing 

shape and offset frequency ( X 1 '"h) . Loc. 373. 
5 7 

SYNAPTOPHYLLUM KLADION T P. 

PROFESSIONAL PAPER 869 PLATE 6 

9 10 
PLATE 7 

FIGURES 1-6. Synaptophyllum kladion n. sp. (p. 50). 

Moorehouse Member, New York. 

1-4. Paratype, USNM 1625·89. Loc. 373. 

1, 3, 4. Transverse thin sections (X 2% and X 5). 

2. Part of corallite in fig. 4 ( X 50). 

5, 6. Paratype, USNM 162590. Transverse and longitudinal thin sections (X 5). Loc. 120. 

7-11. Synaptophyllum arundinaceum (Billings) (p. 47). 

7-9. Paratype, USNM 162577. Three corallites with lateral offsets 

( X 5). See also pl. 4, fig. 6. 

10, 11. USNM 162587. Transverse and semilongitudinal thin sections 

of corallites with lateral offsets (X 5). Columbus Limestone, 

USGS loc. 7807-SD. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 7 

2 

3 

6 

7 9 10 11 

S1"XAPTOPHYLLUM KLADION N . SP. A D S. ARUNDINACEUM (BILLINGS) 
PLATE 8 

FIGURES 1-7. Acinophyllum simcoense (Billings) (p. 57). 

Bois Blanc Formation. 

1-4. Neotype, GSC 31144. Loc. C28. 

1. Transverse thin section ( X 2,% ) . 

2. Detail ( X 100) of low-central coraHite in fig. 3. 

3. Part of fig. 1 (X 5). 

4. Longtudinal thin section ( X 5) . 

5-7. Holotype of Synaptophyllum bacu.loideum Simpson 

NYSM 334, 335. Transverse thin sections of three corallites. Leroy, N.Y. 
GEOLOGICAL SURVEY 

3 

4 5 

ACINOPHYLLUM JvJ COE~SE (BILLI G ) 
PLATE 9 

FIGURES 1-5. Acinophyllum simcoense (Billings) (p. 57). 

Bois Blanc Formation. 

1, 2. USNM 27961. Transverse and longitudinal thin sections (X 

2%, X 1) . Hagarsville, Ontario. 

3-5. NYSM 128111. Transverse and two longitudinal thin sections 

(X 2%). Leroy, N.Y. 
GEOLOGICAL SURVEY 

4 5 

ACINOPHYLLU}}! SIMCOENSE (BILL! GS) 
PLATE 10 

FIGURES 1-7. Acinophyllum simcoense (Billings) (p. 57). 

Bois Blanc Formation. 

1. USNM 162596. Transverse thin section (X 10). Loc. C28. 

2, 5. USNM 162595. Longtudinal (X 5) and transverse (X 10) 

thin sections. Loc. C28. 

3. NYSM 12813. Transverse thin section (X 5). Leroy, N.Y. 

4, 6. USNM 162597. Transverse (X 2%) and longitudinal (X 10) 

thin sections. Loc. C28. 

7. USNM 162598. Transverse thin section (X 2%). Loc. C28,. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 10 

3 

4 

7 6 

ACINOPHYLLUlvf SIMCOENSE (BILL! GS) 
PLA!TE 11 

FIGURES 1, 2. Acinophyllum simcoense (Billings) (p. 57). 

Bois Blanc Formation. NYSM 12812. Lateral and distal views of silicified and etched corallum (X 1). Leroy, N.Y. 

3. Acinophyllum segregatum (Simpson) (p. 61). 

Onondaga Limestone. NYSM 12785. Lateral view of silicified and etched 

corallum (X 1). Leroy, N.Y. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 11 

3 

A CINOPIIYLL U,V SJ:'vlCOENSE (BILLINGS) A D A . SEGREGA TUM (SIMPSON) 
PLATE 12 

FIGURES 1-6. Acinophyllum stramineum (Billings) (p. 59). 

Edgecliff Member. 

1-3. Lectotype, GSC 343,1. Wainfl.eet, Ontario. Photographs provided 

by the G.S.C. 

1, 2. Transverse and longitudinal thin sections (X 4). 

3. Surface of lectotype fragment (X 1). 

4. NYSM 12806. Longitudinal thin section ( X 5). Loc. 108. See 

also pl. 13, fig. 1. 

5'. USNM 162608. Transverse thin section ( X 21h). Loc. 108. See 

also pl. 13, figs. 6, 7, and pl. 14, figs. 8-10. 

6. NYSM 12807. Thin section through broken corallum (X 21h). 

Loc. 1108. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 12 

4 

5 6 

ACINOPHYLLUM STRAMINEUM (BILLINGS) 
PLATE 13 

FIGURES. 1-8. Acinophyllum stramineum (Billings) (p. 59). 

Edgecliff bioherm facies, New York. 

1. NYSM 12806. Transverse section of two corallites (X 10). 

See also pl. 12, fig. 4. 

2-5. USNM 162609. Transverse thin sections of several corallites 

(X 10). Loc. 135. See also pl. 14, figs. 1-7 

6, 7. USNM 162608. See also pl. 12, fig. 5, and pl. 14, figs. 8-10. 

6. Detail of section shown in fig. 7 (X 50). 

7. Longitudinal thin section ( X 10). 

8. NYSM 12808. Longitudinal thin section (X 10). Loc. 108. 
GEOLOGICAL SURVEY 

6 

2 

7 

ACINOPHYLLUM STRAMINEUM (BILLINGS) 

PROFESSIONAL PAPER 869 PLATE 13 

3 

4 

8 
PLATE 14 

FIGURES 1-10. Acinophyllum stramineum (Billings) (p. 59). 

Edgecliff bioherm facies, New York. Loc. 108. 

1-7. USNM 162609. Serial transverse sections of lateral offsetting 

( X 10) . Average distance between successive sections < 1 mm 

(range, 0.8-1.4 mm); total distance from lowest to highest 

section, 5.8 mm. See also pl. 13, figs. 2-5. 

8-1'0. USNM 16,2608. Bee also pl. 12, fig. 5, and pl. 13, figs. 6, 7. 

8. Detail of left upper part of corallite in fig. 10 (X 100). 

9, 10. Transverse thin sections of two corallites (X 10). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 14 

8 

9 10 

7 

ACINOPHYLL UM STRAMINEUM (BILL! GS) 
PLATE 15 

FrGURES 1-6. Acinophyllum segregatum Simpson (p. 61). 

Onondaga Limestone, New York. 

1-3. USNM 162615. Edgecliff bioherm facies. Loc. 269. See also pl. 

16, fig. 1 

1. Transverse thin section of two corallites and lateral support 

(X 5,). 

2. Longitudinal thin section of parent and lateral offset (X 5). 

3. Longitudinal thin section of another corallite ( X 5) . 

4-6. Holotype, NYSM 336. Successive enlargements of single thin 

section (X 2¥2, X 5·, X 20). Clarksville, N.Y. 
PROFESSIONAL PAPER 869 PLATE 15 

6 3 

A CINOPHYLL [AI SEGRECATUM (SIMPSON) 
PLATE 16 

FIGURES 1-4. Acinophyllum segregatum Simpson (p. 61). 

Four transverse thin sections to show intercolony variation in coramte 

diameter and spacing (X 21h). Edgecliff Member, New York. 

1. USNM 162·615. Bioherm facies. Loc. 269. See also pl. 15, figs. 1-3. 

2. NYSM 12780. Biostrome facies. Loc 164·. 

3. USNM 162617. Bioherm fades. Loc. 269. 

4. USNM 162620. Fine-grained facies. Loc. 198. 
GEOLOGICAL SURVEY 

2 

3 4 

ACINOPHYLLUM EGREGATUM (SIMPSO ) 
PLATE 17 

FIGURES 1-5. Acinophyllum segregatum Simpson (p. 61). 

Transverse sections through five coralla (X 2.%). Moorehouse Member 

and equivalent part of Detroit River Group. 

1. GSC 31164. Detroit River Group, Chepstow, Ontario. Loc. GSC 

236·57. 

2. NYSM 12784. Moorehouse Member, Loc. 112 .. 

3. GSC 3tl162. Moorehouse Member, Port Dover, Ontario. Loc. GSC 

23·52.1. 

4. GSC 31163. Moorehouse Member, Naticoke, Ontario. Loc. GSC 

2.2978. 

5. USNM 162621. Moorehouse Member, Acinophyllum-Eridophyllum 

bed. Loc. 118. 
2 

3 

4 5 

ACINOPHYLLUM SEGREGATUM (SIMPSON) 
PLATE 18 

FIGURES 1-6. Acinophyllum segregatum Simpson (p. 61). 

Edgecliff Member, New York. 

1-3. USNM 162619. Transverse and two longitudinal thin sections; 

note lateral connections in figs. 1 and 2 ( X 5) . Fine-grained 

facies. Loc. 19'8. 

4. NYSM 12782. Transverse section of two corallites and lateral 

connection ( X 5) . Biostrome facies. Loc. 286. 

5, 6. NYSM 12786. Longitudinal thin section ( X 10) and detail 

(X 100). Bioherm facies. Loc. 108. 
6 

ACINOPHYLLUM SEGREGATUM (SIMPSO 

3 
PLATE 19 

FIGURES 1-5. Acinophyllum segregatum Simpson (p. 61). 

Edgecliff Member, New York. 

1, 2. USNM 162618. Transverse thin section; detail (X 100) and 

corallite (X 10) ; d~tail is from upper margin as corallite is 

oriented in fig. 2. Loc. 188. 

3-5. NYSM 12787. Bioherm facies. Loc. 269. 

3. Transverse thin section of two corallites and lateral connection (X 5). 

4, 5. Longitudinal thin section; detail (X 50) and corallite 

(X 10). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 19 

4 

2 

5 

3 

ACINOPHYLLUM SEGRECATUM (SIMPSON) 
PLATE 20 

FIGURES 1-11. Acinophyllum segregatum Simpson (p. 61). 

Onondaga Limestone, New York. 

1, 2. NYSM 337, 338. Illustrated thin sections of Synaptophyllum 

simcoense of Simpson (1900) not Billings (X 2). Leroy, N.Y. 

3. NYSM 12790. Longitudinal thin section of parent and offset (X 

5). Edgecliff fine-grained facies. Loc. 337. 

4. NYSM 12789. Transverse thin section of lateral connection (X 

10). Edgecliff bioherm facies. Loc. 236. 

5, 6. NYSM 12783. Transverse thin sections showing three corallites 

with lateral connections ( X 5) . Edgecliff biostrome facies. Loc. 90'. 

7. NYSM 12788. Longitudinal thin section (X 5). Moorehouse Member, Acinophyllum-Eridophyllum bed. Loc. 120. 

8. NYSM 12792. Longitudinal thin section of lateral connections 

meeting midway between corallites (X 10). Edgecliff biostrome 

faces. Loc. 206. 

9. NYSM 12791. Transverse section of three corallites with lateral 

connections ( X 5) . Edgecliff fine-grained facies. Loc. 337. 

10. USNM 162616. Longitudinal thin section of offsetting ( X 5). 

Edgecliff bioherm facies. Loc. 329. 

11. USNM 162623. Longitudinal thin section of corallite and offset 

( X 5) . Edgecliff bioherm facies. Loc. 233. 
GEOLOGICAL SURVEY 

3 

4 

8 

5 

7 

6 

9 10 11 

ACINOPHYLLUM SEGREGATUM (SIMPSON) 
PLATE 21 

FIGURES 1-6. Acinophyllum segregatum Simpson (p. 61). 

Edgecliff Member, New York~ 

1. USNM 162626. Longitudinal thin section (X 10). Bioherm 

facies. Loc. 329. 

2. NYSM 12781. Longitudinal thin section of two corallites showing coarse rugae and lateral connections ( X 5) . Biostrome facies. Loc. 188. 

3. USNM 162628. Longitudinal thin section (X 10). Bioherm facies. Loc. 108. 

4. USNM 162625. Transverse thin section through calice (X 10). 

Bioherm facies. Loc. 269. 

5. USNM 162624. Transverse thin section (X 10). Bioherm facies. Loc. 233. 

6. USNM 162627. Longitudinal thin section of two corallites with 

lateral connections (X 5). Bioherm facies. Loc. 135. 
PROFESSIONAL PAPER 869 PLATE 21 

2 

4 5 

3 6 

ACINOPHYLLUM EGREGATUM (SIMP 0 ) 
PLATE 22 

FIGURES 1-8. Acinophyllum segregatum Simpson (p. 61). 

USNM 162622. Serial transverse thin sections through offsetting 

corallite. 

Total distance from lowest to highest section in sequence, approximately 8 mm; sections 1-3 represent 1.7 mm; 4-8 represent 3.4 

mm; sections 3 and 4 are separated by approximately 2 mm; all 

measurements perpendicular to axis of parent. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 22 

1 8 

2 7 

3 6 

4 5 

A CJNOPHYLL UM SEGREGA TUM SIMPSON 
PLATE 23 

FIGURES 1-8. Acinophyllum mclareni Fagerstrom (p. 63). 

Ontario. 

1-3. USNM 162662. Moorehouse Member. Loc. C18. 

1. Transverse thin section ( X 2% ) . 

2. Detail of corallite nearest to center of fig. 1 ( X 50) . 

3. Longitudinal thin section (X 5). 

4-7. Holotype, UMMP 36120. Formosa reef facie-s of Detroit River 

Group, Formosa, Ontario. 

4, 5. Transverse thin section ( X 5, X 2%). 

6, 7. Longitudinal thin sections of two corallites ( X 5) . 

8. USNM 162661. Transverse thin section of two corallites with 

lateral offsets ( X 5). Loc. C36. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 23 

3 

2 

6 7 8 

ACINOPHYLL Ulvf JI!ICLARENI FAGERSTROM 
PLATE 24 

FIGURES 1-7. Acinophyllum mclareni Fagerstrom (p. 63). 

1, 2. USNM 162664. Transverse (X 2%) and longitudinal (X 5) thin 

sections. Boyle Limestone, Flax Creek, Ky. 

3. USNM 162663. Transverse thin section (X 2%). Jeffersonville 

Limestone, Zone C, Falls of the Ohio, Ky. 

4. OSU 17735. Illustrated specimen of Synaptophyllum simcoense 

Stewart (1938) not Billings. Transverse view (X 5) of thin 

section illustrated by Stewart. Columbus Limestone, Ohio. 

5. USNM 162660. Transverse thin section (X 2%). Amherstberg 

Formation, Beachville, Ontario. 

6, 7. UMMP 60484. Transverse thin section (X 2%), and two corallites from same (X 10). Upper part of Columbus Limestone, 

Loraine County, Ohio. Core from Nagel No. 1 at 1,097 ft. (See 

Janssens, 1970, pl. 21, for general discussion.) 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 24 

5 7 

ACJNOPH YLLUM MCLAREN! FAGERSTROM 
FIGURE 1-8. 

PLATE 25 

Acinophyllum stokesi (Milne-Edwards and Haime) (p. 65). 

1-5. Holotype of A. davisi Stumm, MCZ 7827. Jeffersonville Limestone, Zone A (lower coral zone, Schoharie equivalent, Falls of 

the Ohio). 

1-3. Transverse thin section (X 2%) and parts of same section 

(X 5). 

4. Longitudinal thin section ( X 5) . 

5. Exterior ( X 1) . 

6-8. Holotype, original of Milne-Edwards and Haime, 1851, pl. 20, 

fig. 2. Paris Museum No. 497a. 

6. Exterior ( X 1) . 

7, 8. Tangential and transverse thin sections ( X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 2.'5 

3 7 5 

A CINOPHYLL UM STOKES! (MIL E-EDW ARDS A D HAIME) 
PLATE 26 

FIGURES 1-8. Acinophyllum stokesi (Milne-Edwards and Haime) (p. 65). 

Schoharie Formation New York. 

1-6. NYSM 12810. Clarksville, N.Y. 

1, 2. Longitudinal and transverse thin sections (X 2%). 

3-6. Individual corallites of sections shown in figs. 1 and 2 

(X 10). 

7. USNM 162675. Transverse thin section (X 5). Loc. 79. 

8. USNM 162674. Transverse thin section of two corallites (X 

10). Loc. 505. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 26 

2 

8 5 6 

ACINOPHYLL UM STOKES! (MILNE-EDWARDS A D HAIME) 
PLATE 27 

FIGURES 1-6. Acinophyllum stokesi (Milne-Edwards and Haime) (p. 65). 

Bois Blanc Formation, Ontario. 

1-5. USNM 162670. Loc. C10. 

1, 4. Two parts of one transverse thin section (X 5). 

2, 5. Two parts of one longitudinal thin section (X 5). 

3. Detail of corallite from another part of same transverse 

thin section ( X 50) . 

6. GSC 31143. Basal view of partly etched corallum (X 1). Oxford, Ontario. Loc. GSC 2596. 
C80LOG!CAL SlJR\'EY 

5 

PROFESSIONAL PAPER 869 PLATE 27 

2 

1 

6 

ACINOPHYLLUM STOKES! (MILNE-EDWARDS AND IIAIME) 
PLATE 28 

FIGURES 1-5. Acinophyllum stokesi (Milne-Edwards and Haime) (p. 65). 

Bois Blanc Formation, Ontario. Loc. C22. 

1, 2. USNM 162671. Longitudinal (X 5) and transverse (X 2%) 

thin sections. 

3, 4. USNM 162673. Longitudinal and transverse thin sections (X 

2%). 

5. USNM 162672. Transverse thin section (X 2%). 
3 

4 5 

ACINOPHYLLUM STOKES! (MIL E-EDWARDS AND HAIME) 
PLATE 29 

FIGURES 1-10. Acinophyllum vermetum (Weisbord) (p. 57). 

Caiio Grande Formation, northern Venezuela. 

1-4. Lectotype, PRI 21598. 

1, 2. Transverse thin section (X 2%) and two of the corallites (X 10). 

3, 4. Longitudinal thin sections of two corallites (X 10). 

5-7. PRI 27637. Transverse (X 2%) and two longitudinal (X 10) 

thin sections. 

8. PRI 27638. Transverse thin section (X 2%). 

9, 10. Paralectotype, PRI 21595. Transverse and longitudinal thin 

sections ( X 5) . 
8 10 

ACINOPHYLLUM VERMET[ M (WEISBORD) 

PROFESSIONAL PAPER 869 PLATE 29 

7 
PLATE 30 

FIGURES 1-7. Disphyllum? rectiseptatum (Rominger) (p. 112). 

Dundee Limestone, Michigan. 

1-3. Syntype, UMMP 14378. 

1. T,ransverse thin section (X 2.%). 

2, 3. Longitudinal thin section, detail ( X 25) and general (X 

10). 

4-6. UMMP 14380. Transverse thin section ( X 2%) ; longitudinal 

and transverse thin sections ( X 10). Corallite in fig. 6 is in 

lower right quarter of fig. 4. 

7. Syntype, UMMP 14379. Fragment (X 1). 
GEOLOGICAL Sl ' RYEY PIWFES ' IONAL PAI'I~R 869 PLATE 30 

DISPHYLL Uivi? RECTISEPTA T UM (ROMI GER) 
PLATE 31 

FIGURES 1-9. Cylindrophyllum elongatum Simpson (p. 71). 

Edgecliff Member, New York. 

1, 2. Holotype, NYSM 246. Two longitudinal thin sections (X 2%) 

that together are the original of Simpson's single drawing. 

Clarksville, N.Y. 

3-9. USNM 163382. Loc. 108. See also pl. 32, figs. 4, 5. 

3, 9. Longitudinal thin section (X 2%) and detail (X 25). 

4-8. Serial transverse thin sections through one corallite, showing peripheral increase. Total distance (parallel to corallite axis) from lowest to highest section is 7.0 mm. Sections in figs. 5 and 6 are separated by approximately 3 

mm; other intervals are 1.2-1.4 mm. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 31 

9 

6 7 8 

CYLINDROPHYLLUM ELONGATUM SIMPSO 
PLATE 32 

FIGURES 1-9. Cylindrophyllum elongatum Simpson (p. 71). 

Edgecliff bioherm facies, New York. 

1-3. USNM 163384. Transverse (X 5) and two longitudinal (X 

272 ) thin sections. Loc. 233. 

4, 5. USNM 163382. Details of corallites illustrated in pl. 31, fig. 

6 (X 50) and fig. 8 (X 10) respectively. 

6-9. USNM 163383. Three transverse (X 5) and a longitudinal 

( X 2%) thin sections. The corallite in fig. 6 shows a .single 

peripheral offset. Figs. 8 and 9 are of the same corallite. Loc. 

108. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 69 PLATE 32 

5 9 8 

CYLI DROPHYLL UJVI ELO GATUM IMPSON 
PLATE 33 

FIGURES 1-11. Cylindrophyllum elongatum Simpson (p. 71). 

Edgecliff bioherm facies, New York. 

1-5. USNM 163388. Loc. 296. 

1-4. Serial transverse thin sections (X 2%) showing stages 

in development of peripheral offsets. The lowest and 

highest sections shown are separated by 4.0 mm (measured parallel to corallite axis); distance between adjacent sections ranges from 1.0 to 1.8 mm. 

5. Longitudinal thin section of another corallite (X 2%). 

6, 7. USNM 172241. Longitudinal thin section of perpheral offset 

(fig. 7, X 2% ) and detail of basal plate ( X 50) . 

8-11. USNM 163387. Loc. 108. 

8-10. Serial transverse .sections showing stages in peripheral 

offsetting. Distance between lowest and highest sections 

is approximately 12 mm (measured parallel to corallite axis). Section in fig. 9 is approximately midway 

between other two. 

11. Detail of section in fig. 8 ( X 25). 
GEOLOGICAL SURVEY 

4 11 

CYLINDROPHYLLUM ELO~ GATUM SIMPSON 

PROFESSIONAL PAPER 869 PLATE 33 
PLATE 34 

FIGURES 1-7. Cylindrophyllum elongatum Simpson (p. 71). 

Edgecliff Member, New York. 

1-4. NYSM 12814. Clarksville. 

1, 3, 4. Transverse thin sections of three corallites (X 5). 

2. Detail of section in fig. 1 (X 50). 

5, 6. USNM 163386. Longitudinal (X 2%) and transverse (X 

5) thin sections. Bioherm facies. Loc. 329. 

7. USNM 172242. Specimen broken to show surface (with 

growth lines) of base of two peripheral offsets (X 2%). 

Bioherm facies. Loc. 108. 

8-13. Cylindrophyllum sp. A (p. 77). 

Edgecliff bioherm facies, New York. Loc. 108. 

8, 9. NYSM 12817. Transverse and longitudinal thin sections 

of same corallite ( X 2% ) . 

10-13. USNM 163395. 

10, 11. Longitudinal and transverse thin sections of one 

corallite ( X 2% ) . 

12, 13. Transverse thin section of another corallite (X 2%) 

and detail ( X 50) . 
GEOLOGICAL SURVEY 

13 12 11 

CYLit.DROPHYLL UM ELO GATUMSIMPSO A D C. SP. A 
PLATE 35 

FIGURES 1-10. Cylindrophyllum propinquum Stewart (p. 73). 

Columbus Limestone, Zone C, Ohio. 

1-6. Holotype, OSU 17779. Various transverse and longitudinal 

thin sections (all X 2%). Dublin quarry. 

7, 8. USNM 172188. Transverse thin section (X 2%) and one 

corallite (X 5). USGS loc. 8289-SD. 

9, 10. USNM 172187. Longitudinal and transverse thin sections of 

different corallites (X 5). USGS loc. 7808-SD. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 35 

9 10 

CYLINDROPHYLLUJJ PROPINQUU/vl STEWART 
PLATE 36 

FIGURES 1-7. Cylindropkyllum propinquum Stewart (p. 73). 

USNM 172182. Edgecliff bioherm facies, New York. Loc. 269. 

1. Transverse thin section ( X 2% ) . 

2, 4, 5. Serial transverse thin sections ( X 2% ) showing formation 

of lateral off.set. Distance between lowest and highest section 

is 6.6 mm (measured parallel to parental axis). Intermediate 

section is 4.4 mm above lowest section. 

3. Detail of upper right corallite in fig. 2 (X 50). 

6, 7. l.~ongitudinal thin sections of two corallites ( X 2% ) . 
CYLINDROPHYLLUM PROPINQUUM STEWART 
PLATE 37 

FIGURES 1-9. Cylindrophyllum propinquum Stewart (p. 73). 

Edgecliff biohenn facies, New York. 

1-3. USNM 172180. Two parts of one thin section through colony 

( X 2% ) and two corallites from fig. 1 ( X 5) . Loc. 296. 

4, 5. USNM 172181. Two transverse thin sections (X 2%) through 

same corallite. Loc. 296. 

6-9. NYSM 12818. Loc. 135. 

6, 7. Transverse thin sections through one corallite showing 

two stages of same lateral offset (X 2%). 

8, 9. Longitudinal thin sections of two corallites (X 2%). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 37 

4 5 8 9 

CYLINDROPHYLLUM PROPINQUUM STEWART 
PLATE 38 

FIGURES 1-12. Cylindrophyllum propinquum Stewart (p. 73). 

USNM 172184. Edgecliff biostrome facies, New York. Loc. 188. 

1-3. Wide ... spaced aerial sections of corallite with offset in ad-· 

vanced and very early stages and below same offset (X 2%). 

Total distance between lower and higher sections estimated 

at 15 mm. 

4, 5. Longitudinal thin sections of two other corallites (X 2%). 

6, 7. Transverse and longitudinal thin sections of another corallite (X 2%). 

8. Transverse thin section of another corallite (X 2%). 

9, 10. Longitudinal and transverse thin sections of another corallite (X 2%). 

11. Detail of central part of corallite in fig. 10 (X 10). 

12. Exterior of corallite and offset ( X 1% ) . 

Note carinae in tabularium of several of the corallites. 
9 

10 

12 

3 

2 

6 

5 

7 

11 

CYLINDROPHYLLUM PROPINQUUNI STEWART 

PROFESSIONAL PAPER 869 PLATE 3 

8 
PLATE 39 

FIGURES 1-8. Cylindrophyllum laxtensum n. sp. (p. 75). 

Bois Blanc and Schoharie Formations, New York and Ontario. 

1-4. Holotype, GSC 31146. Bois Blanc Formation. Loc. C19. 

1, 2. Longitudinal thin section; three coral lites (X 2% ) , the 

same and additional corallites ( X 1) . 

3, 4. Transverse thin section; fig. 4, several corallites (X 2lh) 

and fig. 3, detail of lower-middle corallite in fig. 4 (X 

25). 

5. Paratype, USNM 163391. Transverse thin section (X 2lh). 

Bois Blanc Formation. Loc. C28. 

6, 7. Paratype, GSC 31147. Longitudinal thin sections of two corallites (X 2%). Bois Blanc Formation. Innerkip, Ontario, GSC 

loc. 23676. 

8. Paratype, USNM 163393. Longitudinal thin section (X 5). 

Schoharie Grit. Loc. 505. 
GEOLOGICAL SURVEY 

4 

CYLINDROPHYLLUM LAXTENSUM N. SP. 

PROFESSIONAL PAPER 869 PLATE 39 
PLATE 40 

FIGURES 1-6. Cylindrophyllum laxtensum n. sp. (p. 75). 

Bois Blanc Formation, Ontario. 

1-2. Paratype, USNM 182813. Transverse thin section: fig. 2, (X 

2%) ; fig. 1, detail of parts of two corallites in center left of 

fig. 2 (X 50) ; horizontal structure in fig. 1 is wall between 

two lateral connections; other structures are parts of four 

septa (with dark center lines) and two dissepiments. Woodstock Ontario. 

3-6. Paratype, GSC 3585a. Walpole, Ontario. 

3. Exterior of corallites ( X 1) . 

4. Transverse thin section ( X 1) . 

5, 6. Longitudinal thin sections (X 2%). Section in fig. 5 

shows two lateral connections and part of adjacent corallite. 
Gb:OLOGLCAL SURVEY 

2 6 

CYLINDROPHYLLU!v! LAXTENSUM . SP. 
PLATE 41 

FIGURES 1-5. Cylindrophyllum deearium n. sp. (p. 76). 

Edgecliff bioherm facies, New York. 

1-3. Holotype, USNM 172100. Transverse and longitudinal thin 

sections ( X 2%). Loc. 360. 

4, 5. Paratype, NYSM 12819. Longitudinal and transverse thin 

sections ( X 2%) . Loc. 296. 
GEOLOGICAL SURVEY 

4 

5 

CYL INDROPHYLL UM DEEARIUM N. SP. 

PROFESSIONAL PAPER 869 PLATE 41 
PLATE 42 

FIGURES 1-7. Cylindrophyllurn deearium n. sp. (p. 76). 

Edgecliff Member, New York. 

1-4. Paratype, NYSM 12820. Bioherm facies. Loc. 81. 

1, 2. Transverse and longitudinal thin sections (X 2%). 

3. Longitudinal thin section of another corallite ( X 5). 

4. Detail of section in fig. 3 ( X 25). 

5. USNM 172243. Longitudinal thin section of fragment only 

tentatively identified as this species (X 2%) ; shows peripheral 

offsets. Bioherm facies. Loc. 207. 

6. Paratype, NYSM 12871. Exteriors of several corallites and 

offset (X 1). Leroy, N.Y. 

7. Paratype, USNM 172191. Longitudinal thin section including 

peripheral offsets (X 2%). Bioherm facies. Loc. 81. 
GIWLOGICAL S RVI~Y 

6 

CYLINDROPHYLLUAI DEEARJ[ M N. SP. 

7 
PLATE 43 

FrGURES 1-11. Cylindrophyllum deearium n. sp. (p. 76). 

Edgecliff Member, New York and Ontario. 

1, 2. Paratype, USNM 25870. Longitudinal and transverse thin sections (X 2%). Leroy, N.Y. 

3-5. Paratype, UMMP 50576. Longitudinal and transverse thin sections ( X 2% ) . Cheapside, Ontario. 

6-8. Paratype, USNM 172192. Longitudinal and transverse thin 

sections ( X 2%). Bioherm facies. Loc. 81. 

9-11. Paratype, USNM 172193. Bioherm facies. Loc. 244. 

9-10. Transverse and longitudinal thin sections ( X 2%). 

11. Detail of lower end of transverse section as oriented in 

fig. 9 ( X 50) . 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 43 

7 11 

CYLINDROPHYLLUM DEEARIUM N. SP. 
PI.~ATE 44 

FIGURES 1-8. Disphyllum? stummi (Oliver) (p. 113). 

Famine Limestone, St. George, Quebec. 

1-5. Paratype, GSC 24639. 

1, 2. Detail (X 25) and general view (X 5) of longitudinal 

thin section. 

3-5. Detail (X 50), general view (X 5), and second detail 

( X 50) from same transverse thin section. 

6-8. Holotype, USNM 163236. 

6, 7. Transverse and longitudinal thin sections (X 5). 

8. Transverse thin section ( X 2%) ; corallite in fig. 6 is 

in low left center. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 44 

4 

3 

5 8 

DISPHYLL UM ? STUMM! (OLIVER) 
PLATE 45 

FIGURES 1-7. Disphyllum? stummi (Oliver) (p. 113). 

Famine Limestone, St. George, Quebec. 

1-4. Paratype, GSC 24797. Serial transverse thin sections showing 

stages of lateral increase ( X 5). 

5. Paratype, GSC 24798. Longitudinal thin section of corallite and 

lateral offset ( X 5) . 

6. Paratype, USNM 163398. Longitudinal thin section (X 5). 

7. Paratype, USNM 163237. Transverse thin section of corallite 

and lateral offset ( X 5). 

8-10. Cylindrophyllum sp. cf. C. propinquum Stewart 

Edgecliff Member ( ?) , Ontario. USNM 172:185. Loc. C45. 

8. Transverse thin section of several corallites ( X 2% ) . 

9, 10. Longitudinal and transverse thin sections (X 5). Fig. 10 

corallite is right center in fig. 8. 
9 10 

DISPHYLLUM? STUMM! (OLIVER) AND CYLINDROPHYLLUM SP. CF. C. PROPINQUUM STEWART 
PLATE 46 

FIGURES 1-7. Prismatophyllum prisma Lang and Smith (p. 79). 

Jeffersonville Limestone, Zone B. 

1, 2. Holotype, BMNH R31730, 31. Longitudinal and transverse thin 

sections ( X 2 '-h). Charleston Landing, Ind. 

3-5. USNM 163401. Falls of the Ohio, Kentucky-Indiana. 

3, 4. Longitudinal thin section ( X 5, X 2 '-h). 

5. Transverse thin section ( X 2 '-h ) . 

6, 7. USNM 163400. Falls of the Ohio, Kentucky-Indiana. Longitudinal and transverse thin sections ( X 2 '-h) . 
PROFESSIONAL PAPER 869 PLATE 46 

7 5 

PRISNIATOPHYLLU!vf PRIS!VIA LA G A D SMITH 
PLATE 47 

FIGURlES 1-5. Prismatophyllum prisma Lang and Smith (p. 79). 

SUI 21198. Holotype of P. cinctum Stainbrook. Falls of the Ohio. 

1-3. Transverse section (X 10, X 2lh, X 25). 

4, 5. Longitudinal section ( X 2lh, X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 47 

3 5 

PRISM A TOPHYLL UM PRISM A LANG AND SMITH 
PLATE 48 

FIGURES 1-4. Prismatophyllum prisma Lang and Smith (p. 79). 

USNM 163405. Jeffersonville Limestone, Zone B, Falls of the Ohio, 

Ky.-Ind. 

1, 3, 4. Transverse thin section ( X 5, X 2%, X 25). 

2. Longitudinal thin section ( X 5). 
PROFESSIONAL PAPER 869 PLATE 4 

2 

1 

3 4 

PRISMATOPHYLLUM PRISMA LANG AND SMITH 
PLATE 49 

FIGURES. 1-5. Prismatophyllum ovoideum (Davis) (p. 83). 

Jeffersonville Limestone, Falls of the Ohio, Ky.-Ind. 

1. USNM 163413. Longitudinal thin section (X 5). 

2. Holotype, MCZ 7910. Surface of silicified corallum (X 2lh). 

3-5. USNM 163407. Zone C. 

3. Longitudinal thin section ( X 5). 

4, 5. Transverse thin section ( X 2lh) and part of same ( X 

10). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 49 

5 

4 

PRISMATOPHYLLUM OFOIDEU1VI (DAVIS) 
PLATE 50 

FIGURES 1-5. Prismatophyllum ovoideum (Davis) (p. 83). 

USNM 163408. Jeffersonville Limestone, Zone C, Falls of the Ohio, 

Ky.-Ind. 

1, 2, 5. Transverse thin section ( X 2%), one corallite from another 

part of same ( X 5) , and detail of wall and septa of same 

corallite (X 25). 

3, 4. Longitudinal thin section (X 2%, X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 50 

1 

3 5 

PRISMATOPHYLLUM OVOIDEUM (DAVIS) 
PLATE 51 

FIGURES 1-5. Prismatophyllum ovoideum (Davis) (p. 83). 

NYSM 12829. Onondaga Limestone, western New York. 

1, 4, 5. Transverse thin section ( X 2%) , two corallites ( X 5), and 

detail from margin of smaller corallite ( X 5()) . 

2, 3. Longitudinal thin sections (X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 51 

4 5 

PRISMATOPHYLLUM OVOIDEUM (DAVIS) 
PLATE 52 

FIGURES 1-7. Prismatophyllum ovoideum (Davis) (p. 83). 

Edgecliff Member and equivalents, Ontario and New York. 

1, 2, 7. USNM 163414. Transverse thin section (X 2%, X 5) and 

longitudinal thin section (X 5). Amherstburg Formation. 

Loc. C27. 

3, 6. UMMP 24178. Transverse and longitudinal thin sections ( X 

2%). Amherstburg Formation. Beach ville, Ontario. 

4, 5. NYSM 12830. Transverse thin section (X 2%, X 5). Onondaga Limestone, western New York. 
GEOLOGICAL SURVEY 

5 

2 

6 

PRISMATOPHYLL UN! 0 VOIDEUM (DAVIS) 

PROFESSIONAL PAPER 869 PLATE 52 

7 
PLATE 53 

FIGURES 1-6. Prismatophyllum truncata Stewart (p. 85). 

Holotype, OSU 17786. Upper Columbus or Dundee Limestone, Ohio. 

1-4. Transverse thin section ( X 2%) and details of three parts of 

same section ( X 5) . 

5, 6. Longitudinal thin sections ( X 2%, X 5). 
GEOLOGICAL S RVEY 

2 

3 

5 

6 

PRISMATOPHYLLUM TRUNCATA STEWART 
PLATE 54 

FIGURES 1-4. Prismatophyllum truncata Stewart (p. 85). 

USNM 163418. Seneca Member, New York. Loc 46. 

1, 2. Transverse thin section (X 2%) and three corallites in same 

(X 5). 

3. Longitudinal thin section ( X 2%) . 

4. Detail of wall and septa of two small corallites shown in fig. 2 

(X 50). 
GEOLOGICAL SURVEY 

2 

4 

PRISMATOPHYLLUM TRUNCATA STEWART 
PLATE 55 

FIGURES 1-7. Prismatophyllum anna (Whitfield) (p. 87). 

Lectotype, UCMP 34218. Dundee or "Blue" Limestone, Paulding County, 

Ohio. See also plate 56, figure 7. 

1, 2. Two views of lectotype fragment (X 1). Whitfield's fig. 4 is 

from the face in fig. 2. 

3, 5, 7. Transverse thin section (X 5, X 5, X 2%). 

4. Detail of wall and septa in same transverse thin section (X 50). 

6. Longitudinal thin section ( X 5) . 
GEOLOGICAL SURVEY 

6 

4 

7 

PRISMATOPI!YLLUM ANNA (WHITFIELD) 

PROFESSIONAL PAPER 869 PLATE 55 

3 
PLATE 56 

FIGURES 1-7. Prismatophyllum anna (Whitfield) (p. 87). 

"Blue" ( ?) Limestone, Ohio. 

1-3, 6. Paralectotype, OSU 3743-2. 

1, 2. Transverse thin section ( X 2% ) and part of same ( X 5) . 

3. Part of longitudinal thin section shown in fig. 6 (X 5). 

6. Longitudinal thin section ( X 2%) . 

4. Paralectotype, UCMP 342~16. Transverse thin section (X 5). 

5. Paralectotype, UCMP 342117. The original of Whitfield's fig. 2. 

7. Lectotype, UCMP 342,18. Part of section Hlustrated in pl. 55, 

fig. 7 (X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 56 

2 

4 

6 7 

PRISMATOPHYLLUM ANNA (WHITFIELD) 
PLATE 57 

FIGURES 1-5. Asterobillingsa magdisa magdisa Oliver (p. 93). 

Schoharie Formation, New York. 

1, 2, 5. Holotype, USNM 163419. Loc. 73. 

1, 2. Transverse thin section (X 2%) and two corallites of 

same (X 5). 

5. Longitudinal thin section ( X 5) . 

3, 4. Paratype, NYSM 12828. Transverse and longitudinal thin sections. Schoharie County. 
GEOLOGICAL STJRVEY PROFESSIONAL PAPER 869 PLATE 57 

.r ~~ 

3 5 

ASTEROBILLINGSA MAGDISA MAGDISA OLIVER 
PLATE 58 

FIGURES 1-6. Asterobillingsa magdisa magdisa Oliver (p. 93). 

1-3. Paratype AMNH 641. Transverse (X 2%, X 5) and longitudinal ( X 2% ) thin sections. Schoharie Grit, Schoharie, N.Y. 

4-6. Paratype, USNM 128016. Two longitudinal and one transverse 

thin sections ( X 2%) . Bois Blanc Formation, Port Col borne, 

Ontario. This is original of Ehlers and Stumm, 19·53, pl. 1, figs. 

2-3, as Billingsastraea verneuili. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 58 

ASTEROBILLINGSA MAGDISA MAGDISA OLIVER 
PLATE 59 

FIGURES 1-5. Asterobillingsa magdisa magdisa Oliver (p. 93). 

Bois Blanc Formation, Ontario. 

1. Para type, U SNM 113264. Transverse thin section ( X 2%) . 

Port Colborne. 

2. Paratype, USNM 163423. Base of silicified corallum, to show 

holotheca .(X 1). Hagarsville. 

3-5. Paratype, USNM 163420. Transverse thin section (X 5) and 

two parts of one longitudinal thin section (X 5). Loc. C20. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 59 

ASTEROBILLINGSA MAGDISA MAGDISA OLIVER 
PLATE 60 

FIGURES 1-4. Asterobillingsa magdisa magdisa Oliver (p. 93). 

Paratype, USNM 163422. Edgeciff bioherm facies. Loc. 108. 

1-3. Transverse thin section (X 2%) and parts ( X 5, X 10). 

4. Longitudinal thin section ( X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 60 

2 

4 

3 

ASTEROBILLINGSA MAGDISA MAGDISA OLIVER 
PLATE 61 

FIGURES 1-4. Asterobillingsa magdisa steorra Oliver (p. 94). 

Holotype, NYSM 12824. Onondaga Limestone, Clarksville, N.Y. 

Four photographs of one transverse thin section to show morphology 

of septa and carinae (X 2%, X 5, X 10, X 50). See also pl. 62. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 61 

2 

3 4 

ASTEROBILLINGSA MAGDISA STEORRA OLIVER 
PLATE 62 

FIGURES 1-4. Asterobillingsa magdisa steorra Oliver (p. 94). 

Holotype, NYSM 12824. Onondaga Limestone, Clarksville, N.Y. Four 

photographs of one longitudinal thin section to show general morphology and details of carinae and trabeculae (X 5, X 25, X 25, X 

50) . See also pl. 61. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 62 

2 

3 4 

ASTEROBILLINGSA MAGDISA STEORRA OLIVER 
, 

PLATE 63 

FIGURES 1-5. Asterobillingsa magdisa steorra Oliver (p. 94). 

1-3. Paratype, GSC 31155. Woodstock, Ontario. 

1, 2. Transverse thin section (X 2%, X 5). 

3. Longitudinal thin section ( X 5) . 

4. Paratype, USNM 172167. Longitudinal thin section (X 5). Edgecliff bioherm facies. Loc. 108. 

5. Paratype, USNM 172168. Longitudinal thin section (X 5). Edgecliff bioherm facies. Loc. 108. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 63 

3 5 

ASTEROBILLINGSA MAGDISA STEORRA OLIVER 
PLATE 64 

FIGURES 1-4. Asterobillingsa magdisa steorra Oliver (p. 94). 

Edgecliff bioherm facies, New York. 

1, 2. Paratype, USNM 172166. Two corallites from same transverse 

section (X 5). Loc. 108. 

3, 4. Paratype, NYSM 12826. Two corallites from same transverse 

section ( X 5) . Loc. 269. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 64 

2 

3 4 

ASTEROBILLINGSA MAGDISA STEORRA OLIVER 
PLATE 65 

FIGURES 1-5. Asterobillingsa rugosa (Hall) (p. 94). 

Holotype, NYSM 299. Leroy, N.Y. 

1, 2. Longitudinal thin section (X 10, X 5). 

3-5. Transverse thin section (X 5, X 5, X 2lh). 
GEOLOGICAL SURVEY 

2 

ASTEROBILL!NGSA RUGOSA (HALL) 
PLATE 66 

FIGURES 1-4. Radiastraea arachne Stumm ( p. 90') . 

Holotype, USNM 94458. Lower part of Nevada Formation, Nevada. 

Illustrated here for comparison with Asterobillingsa spp. 

1-3. Transverse thin .section (X 2%, X 5, and X 10). 

4. Longitudinal thin section ( X 10). 
3 

2 4 

RAIJIASTRAEA ARACII~VE ST MM 
PLATE 67 

FIGURES 1-10. Eridophyllum seriale Milne-Edwards and Haime (p. 99). 

New York and Ohio. 

1-4. USNM 163424. Moorehouse Member. Loc. 497. 

1, 2. Two transverse thin sections (X 1, X 2%). 

3, 4. Two longitudinal thin sections (X 2%). 

5-7. USNM 10721. Onondaga Limestone, Albany, N.Y. 

5, 7. T:r;ansverse thin section ( X 1) and one corallite from 

same ( X 2% ) . 

6. Longitudinal thin section ( X 2% ) . 

8-10. USNM 163425. Columbus Limestone, Zone H, Marblehead, 

Ohio (USGS loc. 7786-SD). 

8, 9. Transverse thin section, one corallite (X 5) and section 

(X 1). 

10. Longitudinal thin section (X 2%). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 67 

8 9 10 

ERIDOPHYLLU~Vf Sr-',RIALE MILNE-EDWARD ND HAIME 
PLATE 68 

FIGURES 1-10. Eridophyllum seriale Milne-Edwards and Haime,. population B (p. 99). 

Moorehouse Member, Acinophyllum-Eridophyllum bed, New York. 

1-7. USNM 163426. Loc. 118. 

1, 4. Transverse thin section (X 1) and corallite with 

offsets (X 2%). 

2, 3, 5. Corallites from second transverse thin section (X 2%, 

X5,X2%). 

6. Detail of wall and septa of corallite illustrated in fig. 

3 (X 50). 

7. Longitudinal thin section of corallite with lateral offset (X 2%). 

8. NYSM 12831. Longitudinal thin section showing lateral support (X 2%). Loc. 120. 

9, 10. USNM 163428. Transverse (X 5) and longitudinal (X 2%) 

thin sections. Note parasite tubes in fig. 9 and the resulting 

interruptions of the aulos. Loc. 118. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 68 

6 7 10 

ERIDOPHYLLUM SERIALE MILNE-EDWARDS AND HAIME, POPULATION B 
PLATE 69 

FIGURES 1-10. Eridophyllum seriale Milne-Edwards and Haime,, population B (p. 99). 

Moorehouse Member, Acinophyllum-Eridophyllum bed, New York. 

1-5. USNM 163427. Loc. 118. 

1-4. Transverse thin section (X 1), two corallites from 

same (X 5), and detail of corallite in fig. 3 (X 25). 

5. Longitudinal thin section ( X 5) . 

6-10. USNM 163429. Loc. 120. 

6, 7. Longitudinal thin sections including parasite tubes (X 

5). 

8-10. Three corallites from one transverse section (X 5). 

Parasite tubes in first two have caused disruption of 

the aulos; the third is "normal." 
GEOLOGICAL SURVEY 

4 7 

8 10 

ERIDOPHYLLUM SERIALE MILNE-EDWARDS AND HAIME, POPULATION B 
PLATE 70 

FIGURES 1-11. Eridophyllum subseriale Stumm (p. 103). 

Edgecliff Member and equivalents,, New York and Ontario. 

1, 2. USNM 163456. Longitudinal thin sections of two corallites 

(X 2%). Edgecliff bioherm facies. Loc. 329. See also pl. 71, 

figs. 6-8 and pl. 72, figs. 1-7. 

3, 4. UMMP 36122. Specimens illustrated as E. seriale by Fagerstrom, 1961 (pl. 3, figs. 15, 16) and here questionably assigned to E. subseriale. Longitudinal and transverse thin 

sections ( X 2% ) . Forma sa reef facies, Detroit River Group, 

Ontario. 

5, 6. Holotype, UMMP 5272. Longitudinal and transverse thin 

sections (X 2%). Edgecliff Member, Caledonia, N.Y. See 

also pl. 72, fig. 8. 

7-11. NYSM 12833. Edgecliff bioherm facies. Loc. 285. 

7-10. Four corallites from one transverse thin section (X 5). 

11. Detail of corallite in fig. 7 (X 25). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 70 

9 

10 11 

ERIDOPHYLLUM SUBSERIALE STUMM 
PLATE 71 

FIGURES 1-8. Eridophyllum subseriale Stumm (p. 103). 

Edgecliff bioherm facies, New York. 

1, 3, 4. USNM 163455. Loc. 360. 

1. Transverse thin section (X 1). 

3. Longitudinal thin section ( X 2% ) . 

4. One corallite from a second transverse thin section ( X 5) . 

2. USNM 163457. Longitudinal thin section (X 2%). Loc. 360. 

5. NYSM 12834. Detail of transverse thin section not otherwise 

illustrated (X 50). Loc. 329. 

6-8. USNM 163456. Serial transverse thin sections of corallite 

that apparently started to increase axially (X 2%). Lower 

and upper sections are approximately 2 ern apart (measured 

parallel to axis). See text discussion and illustrations of 

"normal corallites" in same corallurn on pl. 70, figs. 1, 2 

and pl. 72, figs. 1-7. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 71 

2 

5 

8 

ERIDOPHYLLUM SLrBSERJALE STUMM 
PLATE 72 

FIGURES 1-8. Eridophyllum subseriale Stumm ( p. 103). 

Edgecliff Member, New York. 

1-7. USNM 163456. Bioherm facie.s. See also pl. 70, figs. 1, 2 and 

pl. 71, figs. 6-8. Serial transverse thin sections (X 2%) from 

highest to lowest showing development and separation of several offsets. Spacing of sections ranges from 3 to 5 mm; total 

separation of upper (fig. 1) and lower (fig. 7) sections approximately 25 mm. Individual corallites are identified in 

tracing below. 

8. Holotype, UMMP 5272. Exterior (X 1). See also pl. 70, figs. 

6, 6. 
4 8 

ERIDOPHYLLUM SL BSERIALE ST MM 
PLATE 73 

FIGURES 1-9. Eridophyllum aulodokum n. sp. (p. 106). 

Holotype, USNM 163436. Moorehouse ( ?) or Seneca ( ?) Member. Loc. 

112. 

1, 2, 5-7. Various corrallites from one transverse thin section (fig. 

1, X 5; others, X 21h). 

3. Corallite and offset from another transverse thin section 

(X 2.1h). 

4. Detail of left corallite in fig. 1 ( X 50). 

8, 9. Longitudinal thin section (X 10, X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 73 

2 

8 9 

ERIDOPHYLLUlvf AULO[)OKUlvl N. SP. 
PLATE 74 

FIGURES 1-8. Eridophyllum corniculum n. sp. (p. 106). 

Moorehouse Member, western New York. 

1-4. Paratype, USNM 163448. Transverse thin ,section (X 1) and 

three corallites from same ( X 5) • Loc. 112. 

5. Paratype, USNM 163449. View of reptant colony from above 

(X 1). Loc. 393. 

6-8. Holotype, USNM 163438. Loc. 112. 

6, 8. One corallite ( X 5) and general view of same transverse 

thin section ( X 1) . 

7. Longitudinal thin section ( X 2% ) . 
7 8 

ERIDOPHYLLUJ;J CORNICULUM N. SP. 
PLATE 75 

FIGURES 1-11. Eridophyllum corniculum n. sp. (p. 106). 

Moorehouse Member, western New York. 

1, 2. Paratype, USNM 163450. Transverse thin sections of corallites 

with lateral offsets (X 5). Loc. 112. 

3-6. Paratype, USNM 163441. Loc. 112. 

3-5. Transverse thin sections of corallites (X 5). 

6. Transverse thin section of immature part of another 

corallite (X 10). 

7. Paratype, USNM 163446. Transverse thin sections of one 

corallite ( X 5) . Loc. 393. 

8. Paratype, NYSM 12835. Longitudinal thin section of corallite 

and offset (X 2%). Loc. 112. 

9, 10. Paratype, USNM 163444. Longitudinal thin sections of two 

corallites ( X 5) . Loc. 112. 

11. Paratype, USNM 163442. Transverse thin section (X 2%). 

Loc. 112. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 75 

ERIDOPHYLLUJI CORNICULUM N. SP. 
PLATE 76 

FIGURES 1-5. Grewgiphyllum colligatum (Billings) (p. 109). 

Neotype, GSC 31158. Bois Blanc Formation, Walpole, Ontario. 

1. Part of weathered top of corallum ( X 1), showing lower sides 

of corallite expansions to the cerioid stages. 

2. Part of weathered base ( X 1) , showing cylindrical stages and 

expansions. 

3, 4. Transverse thin section (X 1) and part of same (X 2%). 

5. Longitudinal thin section ( X 2% ) . 
GEOLOGICAL SURVEY 

3 

5 

2 

4 

GREH'GIPHYLLCJJ COLLJGATU."vl (BILLI G l 

PROFESSIONAL PAPER 69 PLATE 76 
PLATE 77 

FIGURES 1-5. Grewgiphyllum colligatum (Billings) (p. 109). 

Bois Blanc Formation, Ontario. 

1-3. USNM 163464. Transverse thin section, part (X 2%) and 

whole (X 1); and longitudinal thin section (X 2%). Loc. C19. 

4, 5. USNM 163463. Transverse and longitudinal thin sections (X 

2%, X 1). Loc. C19. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 77 

5 

GREWGIP!IYLLUAI COLLIGA. TUJI (BILLI GSl 
PLATE 78 

FIGURES 1-6. Grewgiphyllum colligatum (Billings) (p. 109). 

Ontario. 

1, 2. USNM 163465. Longitudinal thin section, part (X 2%) and 

whole (X 1). Green beds~ Edgecliff ( ?) age. Loc. C10. 

3. USNM 163466. Transverse thin section (X 1). Green beds, 

Edgecliff (?) age. Loc. C10. 

4-6. GSC 31157. Parts of two transverse and one longitudinal thin 

sections ( X 2% ) . Bois Blanc ( ? ) Formation, Woodstock. 
GEOLOGICAL S RVEY PROFESSIONAL PAPER 869 PLATE 78 

2 

3 

6 

GREWGJPHYLL U}vf COLLIGATUM (BILLINGS) 
PLATE 79 

FIGURES 1-7. Grewgiphyllum colligatum Billings (p. 109). 

1-5. USNM 172162. Edgecliff bioherm facies. Loc. 108. 

1, 2. Two longitudinal thin sections (X 2%). 

3. Transverse thin section ( X 1) . 

4. Part of same section ( X 2%) . 

5. Detail of corallite that is right-middle in fig. 4 (X 50). 

6, 7. USNM 113556. Parallel transverse thin sections, of same corallites, taken approximately 5 mm apart (X 1). Bois Blanc (?) 

Formation, Hagarsville, Ontario. 
3 4 

6 

5 

GREWGIPHYLLUM COLLIGATUM BILLI GS 
PLATE 80 

Cyathocylindrium opulens Oliver (p. 117). 

Paratype, NYSM 12844. 

Growth fonn and exwrior of silicified specimen of fonn with lateral offsets (X 1). 

Leroy, N.Y. Probably Edgecliff Member. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 80 

1 

CYA THOCYLINDRIUM OPULEN:::> OLIVER 
PLATE 81 

FIGURES 1, 2. Cyathocylindrium opulens Oliver (p. 117). 

Paratype NYSM 12845. 

Top and lateral views of part of silicified corallum of form with peripheral offsets (X 1). Leroy, N.Y. Probably Edgecliff Member. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 81 

CYA THOCYL INDRI UM OPULENS OLIVER 
PLATE 82 

FIGURES 1-9. Cyathocylindrium opulens Oliver (p. 117). 

Holotype, USNM 172197. Edgecliff bioherm facies. Loc. 81. 

1, 2. Transverse and longitudinal thin sections of one corallite (X 

2%). 

3-6. Transverse thin section of another corallite ( X 2%) and details (X 50, figs. 4 and 5; X 10, fig. 6). Septum in fig. 4 is 

second from right in fig. 6. Area of fig. 6 is upper left in fig. 3. 

Area of fig. 4 is just left of the axis in fig. 3 and reversed. 

7-9. Longitudinal and transverse thin sections of additional corallites (X 2%). 
GEOLOGICAL SURVEY 

1 

7 9 

CYATHOCYLINDRIUM OPULENS OLIVER 
PLATE 83 

[All figures X 2% except fig. 1, X 1] 

FIGURES 1-6. Cyathocylindrium opulens Oliver (p. 117). 

Edgecliff Member, New York and Ontario. 

1, 2. Paratype, USNM 172198. Two transverse thin sections. Bioherm 

facies. Loc. 233. 

3, 4. Paratype, GSC 31159. Transverse and longitudinal thin sections 

of same corallite. Bioherm facies. Loc. C7. · 

5, 6. Paratype, USNM 172199. Longitudinal and transverse thin 

se(!tions of one corallite. Nonbioherm facies. Loc. C22. 
GEOLOGICAL SURVKt PROFESSIONAL PAPER 869 PLATE 83 

5 

CYATHOCYLINDRIUM OPULENS OLIVER 
PLATE 84 

FIGURES 1-6. Cyathocylindrium opulens Oliver (p. 117). 

Edgecliff bioherm facies, New York and Ontario. 

1-4. Paratype, NYSM 12843. Loc. 233. 

1, 2. Transverse thin section ( X 2 'h) and detail ( X 50) . 

3, 4. Longitudinal thin section of same corallite ( X 2 'h) and 

detail (X 25). 

5, 6. Paratype, USNM 172200. Longitudinal and transverse thin 

sections of different corallites. Fig. 6 shows single peripheral 

offset in colony that otherwise has only lateral offsets. This is 

the only example known in this species of the occurrence of both 

lateral and peripheral offsets in the same colony. 
GEOLOGICAL SURVEY 

CYATHOCYLINDRIUM OPULENS OLIVER 
PLATE 85 

[All figures X 2%] 

FIGURES 1-9. Cyathocylindrium opulens Oliver (p. 117). 

Edgecliff bioherm facies, New York. 

1-4. Paratype, USNM 172.201. Three transverse and one longitudinal 

thin sections of four different corallites. Loc. 233. 

5, 6. Paratype, USNM 172204. Transverse and longitudinal thin 

sections of two corallites. Loc. 329. 

7. Paratype, USNM 172203. Longitudinal thin section. Loc. 233. 

8. Paratype, NYSM 12846. Transverse thin section of two corallites. Loc. 329. 

9. Paratype, USNM 172202. Transverse thin section of corallite 

~howing two generations of peripheral offsets. Loc. 329. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 85 

4 7 

3 

9 

CYATHOCYLINDRIUM OPULENS OLIVER 
PLATE 86 

[All figures X 2%] 

FIGURES 1-4. Cyathocylindrium opulens Oliver 

Paratype, USNM 172:205. Edgecliff nonbiostrome facies. Loc. C23. 

1, 2, 4. Two transverse and one longitudinal thin s,ection of one corallite. 

3. Transverse thin section of another coral1ite. 

5-7. Cyathocylindrium sp. cf. C. opulens Oliver. 

UMMP 60485. Transverse and longitudinal thin sections. Fig. 6 shows 

peripheral offsets in advanced stage. Upp,er Columbus Limestone, 

Ashtabula County core, Ohio. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 86 

6 

CYATHOCYLINDRIUM OPULENS OLIVER AND C. SP. CF. C. OPULENS 
PLATE 87 

FIGURES 1, 2. Cyathocylindrium gemrnatum (Hall) (p. 120). 

USNM 172213. Polished base and weathered upper surface of same part of rock slab approximately 5 em thick (X 1h). Shows profuse 

growth with peripheral increase near center of corallum. Edgecliff bioherm facies. Loc. 108. See also pl. 88, figs. 4-6. 

PLATE 88 

FIGURES 1-8. Cyathocylindrium gemmatum (Hall) (p. 120). 

Edgecliff Member, New York and Ontario. 

1, 2. USNM 172217. Transverse (X 1) and longitudinal (X 2%) 

thin sections. Edgecliff "'green beds." Loc. C10. 

3. USNM 172215. Longitudinal thin section ( X 2%). Bioherm 

facies. Loc. 108. See also pl. 90, figs. 4-7. 

4-6. USNM 1722,13. Bioherm facies. Loc. 108. See also pl. 87, figs. 

1, 2. 

4. Transverse thin section ( X 2%). 

5, 6. Longitudinal thin section of another corallite. Detail ( X 

50) of wall from left-center area of section in fig. 6 

(X 2%). 

7, 8. USNM 172212. Transverse and longitudinal thin sections of one 

corallite (X 2%). Bioherm facies. Loc. 108. 
GEOLOGICAL S RVEY 

7 

8 

4 

CYA THO (' YLJ.'\'DRIU:tf GE.T'Jiv!A T UM (HALL ) 
PLATE 89 

FIGURES 1-6. Cyathocylindrium gemmatum (Hall) (p. 120). 

Edgecliff bioherm facies. Loc. 108. 

1-5. USNM 172214. 

1-3. Serial transverse thin sections of axial increase (X 2%). 

Distance between upper and lower section, estimated at 

1 em. 

4. Detail of section in fig. 3 ( X 10) . 

5. Detail of section in fig. 2 (X 50). 

6. NYSM 12860. Transverse thin section (X 2%). 
GEOLOG ICAL SURVEY 

2 

5 6 

CY.iTIIOCYJ.,JX!JRf{ 'JI Gf_,'J!JIATCJJ (HALL) 
PLATE 90 

FIGURES 1-7. Cyathocylindrium gemmatum (Hall) (p. 120). 

1-3. Holotype, FMNH 35872WM. Transverse thin section ( X 2%) 

and two views of specimen ( X 1% ) . Jeffersonville Limestone, 

Falls of the Ohio . 

.'-7. USNM 172215. Edgecliff bioherm facies. Loc. 108. See also pl. 

88, fig. 3. 

4, 5. Transverse thin section ( X 5, X 2%) . 

6, 7. Longitudinal thin section of partial corallite in fig. 5. 

6. Part of section shown on pl. 88, fig. 3 ( X 5) . 

7. Detail of same section ( X 50). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 90 

5 7 

CYATHOCYL!NfJRJC.V GE)r!MATL .l\!1 (HALL) 
PLATE 91 

FIGURES 1-7. Cyathocylindrium n. sp. A (p. 122). 

Moorehouse Member. Loc. 373. 

1-5. USNM 172220. 

1, 2. Transverse and longitudinal thin sections of different 

corallites ( X 2% ) . 

3-5. Serial transverse thin sections showing axial increase. 

Upper section (fig. 3) is approximately 17 mm above 

lower section. Middle section (fig. 4) is 5 to 6 mm above 

lower section. Corallite to left in fig. 3 is enlarged in 

fig. 1. 

6, 7. NYSM 12864. Transverse and longitudinal thin sections (X 

2%). 
GEOLOGICAL SURVEY 

6 7 

CYA TJIOC1'LINDRJUJI . 'P. A 
PLATE 92 

FIGURES 1-5. Heliophyllum megaproliferum n. sp. (p. 124). 

Edgecliff bioherm facies. Loc. C3. 

1, 3, 5. Paratype USNM 172221. 

1, 3. Transverse thin section: detail (X 10) and general 

( X 2% ) . Detail is of inner dissepimentarium, above 

center in fig. 3. 

5. Longitudinal thin section of same corallite (X 2%). 

2, 4. Holotype, GSC 31161. Transverse and logitudinal thin sections (X 1). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 92 

HELIOPHYLLUi\1 J!EGAPROLIFERUAJ N. SP. 
PLATE 93 

FIGURES 1-6. H eliophyllum megaproliferum n. sp. (p. 124). 

Edgecliff bioherm facies, New York. 

1-4. Paratype, USNM 172223. Transverse thin section (X 2%, 

X 1) and details (X 10, X 50). Fig. 3 is right center part of 

corallite in fig. 1 (reversed). Loc. 248. 

5, 6. Paratype, USNM 172222. Longitudinal and transverse thin 

sections (X 2%). Fig. 5 is of partial corallite (parent) in fig. 

6. Fig. 6 shows lateral offset. 
GEOLOGICAL SURVEY PROFESSIO AL PAPER 869 PLATE 93 

2 

4 

5 6 

HELIOPJIYLLCJI .HEGAPROLIFEReJI N. SP. 
PLATE 94 

FIGURES 1-9. Heliophyllum n. sp. B (p. 125). 

Edgecliff bioherm facies. Loc. 329. 

1, 2, 8, 9. USNM 172225. 

1, 2. Transverse and longitudinal thin sections of one coralUte (X 1). 

8, 9. Details of septal morphology in section shown in 

fig. 2 (X 10). 

3-5. USNM 172224. Transverse and longitudinal thin sections 

of different corallites ( X 1) and detail of transverse section (X 50). 

6, 7. NYSM 12865. Longitudinal thin section (X 1) and detail of same (X 5). 
GEOLOGICAL SURVEY 

6 

2 

8 9 

HELJOPIIYLLUM .SP.B 
PLATE 95 

FIGURES 1-6. Heliophyllum n. sp. C (p. 126). 

USNM 172227. Edgecliff bioherm facies. Loc. 297. 

1, 2. Transv.erse thin section ( X 1) and part of same ( X 5) . 

3, 4. Details of septal morphology in same section (X 10, X 25). 

5, 6. Longitudinal thin section of another corallite (X 1) and detail of same (X 5). 
69 PLATE 95 

IIELIOPH YLLUM SP.C 
PLATE 96 

FIGURES 1-4. Heliophyllum coalitum (Rominger) (p. 127). 

1. Holotype, UMMP 8579. Top surface of fragment (X 1). Drift, 

Ann Arbor, Mich. 

2, 3. USNM 172229. Transverse thin section: one corallite (X 2%) 

and several ( X 1) . Edgecliff bioherm facies. Loc. 438. 

4. USNM 172228. Transverse thin section (X 1). Edgecliff bioherm facies. Loc. 296. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 96 

HELIOPHYLLe.~I C'OA.LITCM (ROMI GER) 
PLATE 97 

FIGURES 1-7. Heliophyllum coalitum (Rominger) (p. 127). 

1-6. USNM 172230. Edgecliff bioherm facies. Loc. 438. 

1, 5, 6. Transverse thin section (X 2%, X 1) and detail of wall 

and septa (X 10). 

2, 3. Longitudinal thin sections (X 2%). 

4. Second transverse thin section ( X 1) . 

7. GSC 3586. Weathered .surface of silicified colony. North Oxford 

Township, Ontario. 
7 6 

HELIOPHYLL UAJ CO lLTTUJI (ROM INGER) 
PLATE 98 

FIGURES 1-5. Heliophyllum coalitum (Rominger) (p. 127). 

USNM 172231. Edgecliff bioherm facies. Loc. 296. Morphology of aulos 

known only in this one corallum. 

1-3. Serial transverse sections of lateral offset with aulos (descending order; sections in figs. 2 and 3 were taken 9 mm and 15 

mm below that in fig. 1) (X 2%). 

4. Detail of aulos in fig. 1 ( X 10). 

5. Detail of earliest stage of offset and aulos in section seen in 

fig. 3 (X 5). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 98 

4 

HELIOPHYLL UM COALITUM (ROMI GER) 
PLATE 99 

FIGURES 1-5. Heliophyllum monticulum (Ehlers and Stumm) (p. 129). 

Edgecliff bioherm facies. Loc. 108 (Williamsville), New York. 

1-3. Holotype, UMMP 28519 and USNM 128015. Transverse and 

longitudinal thin sections (X 2%) and upper surface of corallum (X 1). 

4, 5. USNM 172234. Longitudinal and transverse thin sections (X 

2%, X 1). 
GEOLOGICAL SURVEY PROFES IONAL PAPER 869 PLATE 99 

5 

HELIOPHYLLUM MONTICULU!VI (EHLERS A D STUMM) 
PLATE 100 

FIGURES 1-5. Heliophyllum monticulum (Ehlers and Stumm) (p. 129). 

USNM 172232. Edgecliff bioherm facies. Loc. 108. 

1-3. Transverse thin section: general view ( X 2% ) ; detail of inner 

ends of septa of same corallite ( X 50) and its axial area ( X 

10). The three figures are similarly oriented. 

4, 5. Longitudinal thin section ( X 5) and detail ( X 50) . 
PROFESSIONAL PAPER 8~9 PLATE 100 

3 5 

HELIOPHYLLUM MONTICULUJ\!I (EHLERS AND TUMM) 
PLATE 101 

FIGURES 1-4. Heliophyllum monticulum (Ehlers and Stumm) (p. 129). 

Edgecliff bioherm facies, New York 

1-3. USNM 172233. Loc. 81. 

1, 2. Transverse and longitudinal thin sections ( X 1) . 

3. Detail of parts of two corallites in center of fig. 1 (X 5). 

4. USNM 136711. Upper surface of corallum (X 1). Loc. 108. 

See also pl. 102, figs. 1, 2. 
GEOLOGICAL SURVEY 

4 

HELIOPHYLLUM MONTICULUNI (EHLERS AND TUMM) 
PLATE 102 

[All figures X 2%] 

FIGURES 1-4. Heliophyllum monticulum (Ehlers and Stumm) (p. 129). 

Edgecliff bioherm facies, New York and Ontario. 

1, 2. USNM 136711. Transverse and longitudinal thin sections. Loc. 

108. See also pl. 101, fig. 4. 

3. USNM 172235. Transverse thin section. Loc. C10. 

4. USNM 136692. Transverse thin section. Loc. 108. 
3 

2 

4 

IIELIOPHYLLUJ;J MOr-. TICULUM (EHLER A D STUMM) 
PLATE 103 

[All figures X 2%] 

FIGURES 1-5. Heliophyllum cassum n. sp. (p. 130). 

Bois Blanc Formation, Ontario. 

1, 3, 5. Holotype, USNM 172244. Loc. C19. 

1, 5. Different parts of one transverse thin .section. Fig. 5 

shows several offsets. 

3. Longitudinal thin section. 

2, 4. Paratype, UMMP 31601. Transverse and longitudinal thin 

sections. Innerkip. 
GEOLOGICAL SURVEY 

2 

5 

4 

HELIOPHYLLUM C 1SSUM N. P. 
PLATE 104 

FIGURES 1-4. Heliophyllum procellatum n. sp. (p. 131). 

Holotype, NYSM 12872. Bois Blanc ( ?) Formation, Falkirk, N.Y. 

1. T·ransverse thin section ( X 2% ) . 

2. Central corallite in fig. 1 ( X 5) . 

3. Longitudinal thin section ( X 5) . 

4. Top surface of corallum ( X 1) . 
GEOLOGICAL SURVEY 

. ; ,'-':,;r . .fl... ~~...,.""~ 

4 

HELIOPHYLLUM PROCELLATUJI N. SP. 
PLATE 105 

FIGURES 1-5. Heliophyllum procellatum n. sp. (p. 131). 

Bois Blanc ( ?) Formation, Falkirk, N.Y. 

1, 2. NYSM 12874; thin sections of either NYSM 12872 or 12873. 

Transverse thin sections ( X 5) . 

3-5. Paratype, NYSM 12873. 

3. Transverse thin section of corallite in lower left of fig. 5 

(X 5). 

4. Longitudinal thin section ( X 5) . 

5. Transverse thin section ( X 2% ) . 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 105 

3 

5 

HELIOPHYLLUM PROCELLATUM . SP. 
PLATE 106 

FIGURES 1-4. Heliophyllum sp. cf. H. procellatum n. sp. (p. 131). 

UMMP 18804. Bois Blanc Formation, Mackinac County, Michigan. 

1, 2. Transverse thin section (X 5, X 2%). 

3. Another transverse thin section ( X 2%) . 

4. Longitudinal thin section (X 4). 
HELIOPHYLLUM SP. CF. 11. PROCELLATUlVI . SP. 
PLATE 107 

FIGURES 1-6. Eridophyllum conjunctum (Davis) (p. 108). 

USNM 172245. Moorehouse Member, New York. Loc. 118. 

1. Transverse thin section ( X 1) . Upper right-center corallite 

with upward projecting offset; same corallite as in figs. 2 

and 3. 

2, 3. Upper and .lower transverse thin section ( X 2% ) , showing 

early and middle stages of lateral offset to left of large corallite and another lateral offset from large left corallite that is 

only partly shown. Two sections are parallel and 4 mm apart. 

Section in fig. 1 is parallel and approximately 8 mm higher 

than section in fig. 2. 

4. Detail of offsets in fig. 3 ( X 10). 

5. Longitudinal thin section (X 2%). 

6. Detail of aulos in offset shown in figs. 1-3 but in additional 

transverse section taken between sections shown in figs. 1 and 

2(X10). 
GEOLOGICAL SURVEY 

2 

3 

4 

5 6 

ERIDOPHYLLUM CONJUNCTUM (DAVIS) 
PLATE 108 

FIGURES 1-3. E'tidophyllum conjunctum (Davis) (p. 108). 

Jeffersonville Limestone (above coral zone), Falls of the Ohio (X 2%). 

1. Lectotype of Diphyphyllum conjunctum Davis, MCZ 7859. 

Transverse thin section. 

2, 3. Holotype of Diphyphyllum coalescens Davis, MCZ 7877. Transverse and longitudinal thin sections. 

4-7. Eridophyllum sp. cf. E. conjunctum (Davis) 

Columbus Limestone, Chillicothe, Ohio core, OSU 21336. 

4. Longitudinal thin section ( X 2% ) . 

6-7. Transverse thin sections (X 5, X 5, X 2%). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 869 PLATE 108 

6 7 

ERIDOPHYLL UlVI CONJUNCTU11:1 (DAVIS) A D E. SP. CF. E. CONJUNCTUM (DAVIS)