Shorter Contributions to 

Paleontology and Stratigraphy, 1992 

U.S. GEOLOGICAL SURVEY BULLETIN 2024 

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Shorter Contributions to 

Paleontology and Stratigraphy, 1992 

Edited by WILLIAM j. SANDO 

This volume is published as chapters A and B. 

These chapters are not available separately. 

Chapter titles are listed in the volume table of 

contents 

U.S. GEOLOGICAL SURVEY BULLETIN 2024 
U.S. DEPARTMENT OF THE INTERIOR 

BRUCE BABBITI, Secretary 

U.S. GEOLOGICAL SURVEY 

Dallas L. Peck, Director 

Any use of trade, product, or firm names in this publication is for 

descriptive purposes only and does not imply endorsement by the 

U.S. Government 

UNITED STATES GOVERNMENT PRINTING OFFICE: 1993 

For sale by 

Book and Open-File Report Sales 

U.S. Geological Survey 

Federal Center, Box 25286 

Denver, CO 80225 

Library of Congress Cataloging in Publication Data 

ISSN 1062-4295 
CONTENTS 

[Letters designate the chapters] 

(A) Upper Cretaceous Heteromorph Ammonites from the Baculites compressus 

Zone of the Pierre Shale in North-Central Colorado, by William A. Cobban, 

W. James Kennedy, and Glenn R. Scott 

(B) The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America, 

by William A. Oliver, Jr. 

Volume Contents Ill 

Chapter A 

Upper Cretaceous Heteromorph Ammonites from 

the Baculites compressus Zone of the Pierre Shale 

in North-Central Colorado 

By WILLIAM A. COBBAN, W. JAMES KENNEDY, and 

GLENN R. SCOTT 

Descriptions and illustrations of an unusual ammonite 

assemblage of both Western Interior and Gulf Coast aspects 

U.S. GEOLOGICAL SURVEY BULLETIN 2024 

SHORTER CONTRIBUTIONS TO PALEONTOLOGY AND STRATIGRAPHY, 1992 

CONTENTS 

Abstract Al 

Introduction Al 

Acknowledgments A3 

Fossil Localities A3 

Systematic Paleontology A3 

Suborder Ancyloceratina Wiedmann, 1966 A3 

Superfamily Turrilitaceae Gill, 1871 A3 

Family Nostoceratidae Hyatt, 1894 A3 

Genus Axonoceras Stephenson, 1941 A3 

Axonoceras compressum Stephenson, 1941 A3 

Genus Anaklinoceras Stephenson, 1941 AS 

Anaklinoceras rejlexum Stephenson, 1941 AS 

Anaklinoceras gordiale n. sp. A6 

Family Dip1omoceratidae Spath, 1926 A6 

Subfamily Diplomoceratinae Spath, 1926 A6 

Genus Solenoceras Conrad, 1860 A6 

Solenoceras reesidei Stephenson, 1941 A6 

Family Baculitidae Gill, 1871 A7 

Genus Baculites Lamarck, 1799 A 7 

Baculites compressus Say, 1820 A7 

Baculites undatus Stephenson, 1941 A7 

Superfamily Scaphitaceae Gill, 1871 AS 

Family Scaphitidae Gill, 1871 AS 

Subfamily Scaphitinae Gill, 1871 AS 

Genus Jeletzkytes Riccardi, 1983 AS 

Jeletzkytes nodosus (Owen, 1852) AS 

Genus Hoploscaphites Nowak, 1911 A9 

Hoploscaphites cf. H. landesi Riccardi, 1983 A9 

References Cited A9 

PLATES 

[Plates follow references cited] 

1. Axonoceras compressum Stephenson, Anaklinoceras gordiale n. sp., and 

Anaklinoceras rejlexum Stephenson. 

2. Baculites compressus Say. 

3. Baculites compressus Say, Jeletzkytes nodosus (Owen), and Hoploscaphites cf. H. 

landesi Riccardi. 

Contents Ill 
IV Contents 

FIGURES 

1. Chart showing Campanian and Maastrichtian ammonoid zones in the Western 

Interior and the Campanian stratigraphic sequence in the Kremmling area, Grand 

County, Colo. A2 

2. Map of part of Grand County in Middle Park, Colo., showing localities where 

fossils were collected A4 

3-6. Drawings showing: 

3. Restoration of Axonoceras compressum Stephenson A4 

4. Suture of Axonoceras compressum Stephenson A4 

5. Restoration of Anaklinoceras reflexum Stephenson AS 

6. Restoration of Anaklinoceras gordiale n. sp. A6 

TABLE 

1. Localities in Middle Park in Grand County, north-central Colorado, from which 

heteromorph ammonites discussed in this report were collected AS 
Upper Cretaceous Heteromorph Ammonites from 

the Baculites compressus Zone of the Pierre Shale 

North-Central Colorado 

. 1n 

By William A. Cobban, 1 W. James Kennedy/ and Glenn R. Scote 

Abstract 

The Baculites compressus zone of the Pierre Shale in 

Grand County, Colo., contains the following assemblage 

of heteromorph ammonites: Axonoceras compressum 

Stephenson, Anaklinoceras reflexum Stephenson, A. gordiale n. sp., So/enoceras reesidei Stephenson, Baculites 

compressus Say, B. undatus Stephenson, }eletzkytes 

nodosus (Owen), and Hoploscaphites cf. H. landesi Riccardi. The first two species were described from the 

Neylandville Marl in northeastern Texas. Anaklinoceras 

reflexum is also known from the Mount Laurel Sand in 

New Jersey as well as from the Mishash Formation in 

Israel. }eletzkytes nodosus, which also occurs in the 

younger Baculites cuneatus, B. reesidei, and B. jenseni 

zones of the Western Interior, has been found in northeastern Texas, Tennessee, Arkansas, and New jersey. In 

the well-documented Upper Cretaceous sequence in 

Poland,}. nodosus is restricted to the upper Campanian; 

thus, the presence of}. nodosus confirms a late Campanian age for the B. compressus zone in Colorado. 

INTRODUCTION 

The Baculites compressus zone of the Pierre Shale 

(fig. 1) in the Western Interior of the United States usually 

contains a rather limited ammonite fauna of Baculites 

compressus Say (1820), Jeletzkytes nodosus (Owen, 1852), 

and Placenticeras meeki Bohm (1898). A more varied 

ammonite fauna occurs in two areas in Middle Park plateau 

in Grand County, north-central Colorado (fig. 2), where 

uncoiled ammonites (heteromorphs) occur. The fauna of 

these two areas is of much interest in that it represents the 

only occurrences in the Western Interior of several species 

Manuscript approved for publication February 26, 1992. 1 U.S. Geological Survey, Denver, CO 80225. 2 Geological Collections, University Museum, Parks Road, University of Oxford, Oxford OXl 3PW, U.K. 3 60 Estes Street, Lakewood, CO 80226. 

described from northeastern Texas. Two of the species are 

also known from areas as far away as New Jersey and 

Israel. 

The total ammonite fauna from the Baculites compressus zone of Middle Park consists of Placenticeras 

meeki Bohm (1898) and P. intercalare Meek and Hayden 

(1860) and the heteromorphs A.xonoceras compressum 

Stephenson, 1941, Anaklinoceras reflexum Stephenson, 

1941, A. gordiale n. sp., Solenoceras reesidei Stephenson, 

1941, Baculites compressus Say, 1820, B. undatus 

Stephenson, 1941, Jeletzkytes nodosus (Owen, 1852), and 

Hoploscaphites cf. H. landesi Riccardi, 1983. The fauna is 

assigned a late Campanian age mainly because Jeletzkytes 

nodosus, described as the new species Acanthoscaphites 

praequadrispinosus by Blaszkiewicz (1980), occurs in the 

upper Campanian zone of Nostoceras pozaryskii in Poland 

but is unknown in the succeeding lower Maastrichtian 

Belemnella lanceolata zone. Nostoceras pozaryskii Blaszkiewicz (1980), a synonym of N. hyatti Stephenson (1941), 

occurs in Poland in the upper part of the range of Belemnitella langei, a guide to one of the upper Campanian 

belemnite zones of northwestern Europe (Jeletzky, 1951 ~ 

Schulz, 1978; Ernst and others, 1979~ Blaszkiewicz, 1980; 

Schulz and others, 1984~ Robaszynski and Christensen, 

1989). 

Jeletzkytes nodosus begins in the zone of Baculites 

compressus in the Western Interior and ranges on up 

through the overlying zones of B. cuneatus, B. reesidei, and 

B. jenseni (fig. 1). A few poorly preserved fragments of a 

nostoceratid ammonite from the B. jenseni zone in the upper 

part of the Pierre Shale in south-central Colorado seem 

assignable to Nostoceras hyatti. If correctly identified, 

these fragments indicate a later Campanian age than that of 

B. compressus and suggest that the top of the Campanian 

lies at the top of the B. jenseni zone. The CampanianMaastrichtian boundary has been drawn even a little higher 

at the base of the Baculites baculus zone by Jeletzky (in 

Cobban and Reeside, 1952, p. 1026-1028) and more 

recently by Hancock and Kauffman (1989, p. 23). 

Upper Cretaceous Heteromorph Ammonites from the Baculites compressus Zone of the Pierre Shale, Colorado A 1 
t 

C1j 

c.. 

c 

C1j 

·.;::; 

..c 

u 

·;:: 

+-' 

en 

C1j 

C1j 

~ 

CRETACEOUS 

STAGE 

Middle 

Lower 

t-?--+----

AMMONOID 

ZONE 

Jeletzkytes nebrascensis 

Hoploscaphites nicolleti 

Hoploscaphites aff. nicolleti 

Baculites clinolobatus 

Baculites grandis 

Baculites baculus 

Bacu/ites eliasi 

Baculites jenseni 

Baculites reesidei 

Baculites cuneatus 

FORMATION MEMBER 

Gunsight Pass Member 

Upper •• ,:, ; .•• ~ac~~;;./co+~+s:;y;:·····u:-·· .•..• ·x;· :. ;;;? .:(•) / s~~ ~~--•• ,:;". ••• )\ 

c 

C1j 

c 

C1j 

c.. 

E 

C1j 

u 

Middle 

Lower 

Didymoceras cheyennense 

Exiteloceras jenneyi 

Didymoceras stevensoni 

Didymoceras nebrascense 

Baculites scotti 

Baculites reduncus 

Baculites gregoryensis 

Baculites perplexus 

Baculites sp. (smooth) 

Baculites asperiformis 

Baculites mclearni 

Baculites obtusus 

Baculites sp. (weak flank ribs) 

Baculites sp. (smooth) 

Scaphites hippocrepis Ill 

Scaphites hippocrepis II 

Scaphites hippocrepis I 

Scaphites leei Ill 

t 

Cll 

Q. 

c: 

0 

·.;::::; 

Cll 

E 

0 

LL 

~ 

Cll 

..0 

0 

z 

Carter Sandstone Member 

Shale 

Hygiene Sandstone Member 

Shale 

Muddy Buttes and 

Kremmling Sandstone Members 

Shale 

Sharon Springs Member 

Shale 

Figure 1. Campanian and Maastrichtian ammonoid zones in the Western Interior of the United States and 

the Campanian stratigraphic sequence in the Kremmling area, Grand County, Colo. Near Kremmling, the 

Baculites compressus zone (stippled) contains an unusually varied ammonite fauna, which is described in 

this report. 

A2 Shorter Contributions to Paleontology and Stratigraphy, 1992 
Acknowledgments 

R.E. Burkholder, retired from the U.S. Geological 

Survey (USGS), photographed many of the specimens. 

John R. Stacy, formerly of the USGS, prepared the four 

drawings. Kennedy acknowledges the financial support of 

the Natural Environment Research Council (U.K.), Royal 

Society, and Astor Fund (Oxford) and the technical support 

of the staff of the Geological Collections, University 

Museum, Oxford, and Department of Earth Sciences, 

Oxford. 

FOSSIL LOCALITIES 

The ammonites from Middle Park came from grayand brown-weathering, sandy, calcareous concretions in the 

Pierre Shale in the Kremmling and Granby areas. The 

stratigraphic position of the fossiliferous concretions in the 

Kremmling area is shown in columnar sections (lzett and 

others, 1971, pl. 1, level of D 1351; Izett and Barclay, 

1973, Baculites compressus zone). The collections are from 

a little below the middle of the unnamed shale member of 

the Pierre Shale about 116 m above the top of the Carter 

Sandstone Member (lzett and others, 1971 , p. A 15). The 

ammonites are associated with an abundant bivalve fauna 

that consists of the genera Perrisonota, Nucula, Nemodon, 

Breviarca, Phelopteria, Inoceramus (Endocostea), Ostrea, 

Pecten (Camptonectes), Anomia, Cymella, Crassatella, 

Lucina, Tenea, and Protocardia. 

Localities where ammonite specimens discussed in 

this report were collected are shown in figure 2 and listed in 

table 1. Most of the localities are shown on geologic maps 

of the Kremmling quadrangle (lzett and Barclay, 1973) and 

the Trail Mountain quadrangle (lzett, 1974). 

All specimens described in this report are kept in the 

National Museum of Natural History, Washington, D.C. , 

where they have USNM catalogue numbers. Casts of a few 

of the specimens are at the U.S. Geological Survey, Federal 

Center, Denver, Colo. 

SYSTEMATIC PALEONTOLOGY 

Suborder ANCYLOCERATINA Wiedmann, 1966 

Superfamily TURRILITACEAE Gill, 1871 

Family NOSTOCERATIDAE Hyatt, 1894 

Genus AXONOCERAS Stephenson, 1941 

Type species. -Axonoceras compressum Stephenson, 

1941, p. 422, pl. 89, figs. 1-5, by original designation. 

Axonoceras compressum Stephenson, 1941 

Plate 1, figures 1-17, 50, 51; text-figures 3, 4 

1941. Axonoceras compressum Stephenson, p. 422, pl. 89, 

figs. 1-5. 

1941. Axonoceras pingue Stephenson, p. 423, pl. 89, figs. 

6-8. 

1941. Axonoceras multicostatum Stephenson, p. 423, pl. 89, 

figs. 9-11. 

1941. Axonoceras multicostatum rotundum Stephenson, p. 

424, pl. 89, figs. 12-14. 

Types.- Holotype is USNM 77290, the original of 

Stephenson (1941, p. 422, pl. 89, fig. 1) from the Neylandville Marl on the Corsicana Road at USGS Mesozoic 

locality 17361, 4.1 km north of Corbet, Navarro County, 

Tex. 

Description. -Planispiral, small, as much as 30 mm 

in diameter (text-fig. 3). Coiling varies markedly from 

regular tightly criocone (pl. 1, figs. 1-9) that have whorls 

barely separated to regular open criocone (pl. 1, fig. 12), to 

irregularly coiled (pl. 1, fig. 10). Most specimens consist of 

three whorls. 

The protoconch is succeeded by half a whorl or more 

on which ornament is weak and irregular (pl. 1, fig. 14; textfig. 3); the most conspicuous ornament consists of marked 

constrictions and associated collar ribs spaced three per half 

whorl. Succeeding whorls range from compressed to 

depressed in intercostal section. Ornament is of numerous 

narrow, prorsiradiate, slightly flexuous, mainly single ribs 

that are weak at the umbilical shoulder, but strengthen 

across the flanks, and pass straight across the venter. 

Ribbing generally coarsens throughout ontogeny, but there 

is wide variation in both strength and number of ribs per 

whorl. Small ventral tubercles are present on most ribs. In 

finely ribbed individuals, ventral tubercles are minute, 

coarsen as ribs coarsen, and ultimately develop into small 

clavi. Nontuberculate ribs, where present, may be weaker 

than the tuberculate ones. Constrictions with associated 

flared collar ribs may persist or they may be lost at any early 

stage in development. 

Most of the specimens from Middle Park are about 20 

mm in diameter. Smaller specimens seem to be juveniles. A 

single adult body chamber (pl. 1 , figs. 50, 51) that has a 

diameter of about 32 mm may belong to this species. 

Ornament is fairly coarse like the form described by 

Stephenson (1941, p. 423, pl. 89, figs. 6-8) as the new 

species A. pingue. The Middle Park specimen has pronounced ventral clavi on some ribs separated by one or two 

nontuberculate ribs. Pairs of ribs loop between the clavi 

across the venter. Occasional specimens from Middle Park 

have loose, irregular coiling (pl. 1, fig. 10). 

The very simple suture (text-fig. 4) has a broad bifid 

saddle that separates the external (E) and lateral (L) lobes, 

and it has broad lateral and umbilical (U) lobes. 

Discussion. -Stephenson (1941) described Axonoceras compressum, A. pingue, A. multicostatum, and A. 

multicostatum rotundum from a single locality in Navarro 

County, Tex. Each of these forms is represented in the 

collection from USGS Mesozoic locality D1351 in the 

Kremmling area, and there are transition forms between 

them. We believe only a single variable species is present, 

Upper Cretaceous Heteromorph Ammonites from the Baculites compressus Zone of the Pierre Shale, Colorado A3 
R. 81 W. R. 80 W. R. 79 W. R. 78 W. R. 77W. R. 76W. 

08092r X-05991 (D"" 40°15' 

08093 ~ X......-- 05416 

T. 3 N. 08060 X-01351 

x'05862 Trail Mountain 

quadrangle 17 1/2') 

08074-X 

T. 2 N. Kremmling i d 1 e p a r k quadrangle 115') 1 

Granby 

Colorado Granby 

T. 1 N. quadrangle 17 1/2') 

40°00' 

1 06°30 ' 106°15' 1 06°00' 

0 10 MILES 

COLORADO 

0 10 KILOMETERS 

Acea offigme~ 

Figure 2. Part of Grand County in Middle Park, Colo., showing localities where fossils were collected. 

Figure 3. Restoration of Axonoceras compress urn Stephenson, x 4, based on many specimens. Stipple board drawing by john R. Stacy. 

A4 Shorter Contributions to Paleontology and Stratigraphy, 1992 

u L 

2mm 

Figure 4. Most of the suture of Axonoceras compressum 

Stephenson, from USGS Mesozoic locality 05991. E is the 

external lobe, L is the lateral lobe, and U is the umbilical 

lobe. 

for which, as first revising authors, we select the name 

Axonoceras compressum. 

Axonoceras compressum differs from A. angolanum 

Haas (1943, p. 8, figs. 3, 10-13) in its more closely coiled 

shell and more rectiradiate ribbing. Axonoceras multicostatum ellipticum Collignon (1971, p. 13, pl. 644, fig. 2384), 

from the supposed lower Maastrichtian of Madagascar, has 

an elliptical whorl section, has more intercalated ribs than 

A. compressum, and has every sixth to ninth rib strengthened. Axonoceras sohli Cobban (1974a, p. 85, figs. ll-p, 

4) has flattened flanks and venter with tubercles on every rib 

of the last whorl of the phragmocone. 

Occurrence. - Baculites compressus zone of Pierre 

Shale at USGS Mesozoic localities D1351, D5416, D5991, 
Table 1. Localities in Middle Park in Grand County, northcentral Colorado, from which heteromorph ammonites 

discussed in this report were collected 

U.S. Geological 

Survey Mesozoic 

locality 

Collector, year of collection, 

and description of locality 

Kremmling quadrangle: 

01349 .............. G.R. Scott, 1957. SE1/4NE1/4 sec. 

18, T. 3 N., R. 80 W. 

01351 .............. G.R. Scott and W.A. Cobban, 1957; 

G.A. Izett, 1967. SWl/4 sec. 17, 

T. 3 N., R. 80 W. 

D5416 .............. G.A. Izett, 1966. SE1/4NE1/4 sec. 

18, T. 3 N., R. 80 W. 

D5862 .............. G.A. Izett, 1967. NWl/4 sec. 21, T. 

3 N., R. 80 W. 

05991 .............. G.A. Izett and W.A. Cobban, 1967. 

Center of Nl/2 sec. 12, T. 3 N., R. 

81 w. 

08092 .............. G.A. Izett and W.A. Cobban, 1971. 

SE1/4SE1/4 sec. 12, T. 3 N., R. 81 

w. 

08093 .............. G.A. Izett and W.A. Cobban, 1971. 

NE1/4NE1/4 sec. 13, T. 3 N., R. 

81 w. 

08060 .............. G.A. Izett and W.A. Cobban, 1971. 

NW1/4SE1/4 sec. 12, T. 3 N., R. 

81 w. 

Granby quadrangle: 

08086 ......... . .... G.A. Izett, 1971. Sec. 34, T. 2 N., 

Trail Mountain 

quadrangle: 

R. 77W. 

08074 .............. G.A. Izett, 1971. SW1/4SW114 sec. 

2, T. 2 N., R. 77 W. 

D8074, D8086, D8092, and D8093 in Grand County, Colo. 

(fig. 2); Neylandville Marl, 4.1 km north of Corbet in 

Navarro County, Tex. 

Genus ANAKLINOCERAS Stephenson, 1941 

Type species.-Anaklinoceras reflexum Stephenson 

(1941, p. 414, pl. 83, figs. 1-5), by original designation. 

Anaklinoceras reflexum Stephenson, 1941 

Plate 1, figures 23-49; text-figure 5 

1941. Anaklinoceras rejlexum Stephenson, p. 414, pl. 83, 

figs. 1-5. 

1960. Anaklinoceras rejlexum Stephenson. Easton, text fig. 

11.26-6. 

1986. Anaklinoceras reflex urn Stephenson. Lewy, p. 1' fig. 

1B-E. 

Types.- Holotype is Texas Memorial Museum (Austin) 17300, from the Neylandville Marl on the Corsicana 

Road 4.1 km north of Corbett, Navarro County, Tex. 

Description. -Anaklinoceras reflexum is dimorphic. 

Macroconchs from Middle Park are 33 to 35 mm long; 

microconchs are 15 to 22 mm long. The shell has an initial 

whorl or two coiled in an open helix that is smooth except 

for occasional constrictions (text-fig. 5). This loosely coiled 

Figure 5. Restoration of the macroconch of Anaklinoceras reflexum Stephenson, x 3, based on many specimens. Stipple board drawing by john R. Stacy. 

stage is followed by three or four whorls that form a tightly 

coiled helix ornamented by uniform rursiradiate ribs. Three 

to five narrow, deep constrictions per whorl are present on 

the helix. The body chamber leaves the helix and wraps 

around it in a plane. 

Macroconchs have a helix of four or five whorls that 

form an apical angle of about 40° (pl. 1, figs. 29-49). Ribs 

on the larger two or three whorls have a row of small, 

pointed tubercles on the lower part of the flank and a row of 

similar tubercles at the base. The body chamber, which 

begins at the large end of the last whorl of the helix, turns 

upward abruptly and lies against one side of the helix all the 

way to just beyond its initial loosely coiled end, where the 

body chamber bends over to form an inverted U above the 

helix (text-fig. 5). Ribbing coarsens and becomes irregular 

on the body chamber, where strong ribs may be separated 

by one or two weaker secondary ribs. Stronger ribs bear 

umbilical bullae and prominent nodate ventral tubercles. 

Secondary ribs may or may not support small, inconspicuous, bullate tubercles. Ribs are rectiradiate at the beginning 

of the body chamber, prorsiradiate on the straight part, and 

rursiradiate on the final curved part. 

Microconchs have more slender whorls than macroconchs, and the older part of the body chamber, although 

straightened, is not pressed against the helix (pl. 1, figs. 24, 

Upper Cretaceous Heteromorph Ammonites from the Baculites compressus Zone of the Pierre Shale, Colorado A5 
/ 

Figure 6. Restoration of Anaklinoceras gordiafe n. sp., x 3. Stipple 

board drawing by john R. Stacy. Lines show axes of coiling. 

25). In addition, the helix consists of only three or four 

whorls in contact, whereas macroconchs have four or five. 

Small, inconspicuous tubercles may be present only on the 

last whorl of the helix. The body chamber may not begin 

until part way up the straight shaft that parallels the axis of 

the helix (pl. 1, figs. 24, 25). Ribbing coarsens on the body 

chamber and becomes a little irregular like that on macroconchs. The poorly exposed sutures show a rather simple 

pattern of little divided lobes and saddles; lobes are bifid 

and about half as wide as the saddles. 

Occurrence.-Baculites compressus zone of Pierre 

Shale at USGS Mesozoic localities D1351 and D5862 in 

Grand County, Colo. (fig. 2); Neylandville Marl of northeastern Texas; Mount Laurel Sand of New Jersey; Mishash 

Formation, middle phosphorite unit, Oron phosphate field, 

Israel. 

Anaklinoceras gordiale n. sp. 

Plate 1, figs. 18- 22; text-figure 6 

1976. Axonoceras sp. Kennedy and Cobban, p. 42, pl. 11, 

fig. 6. 

Derivation of name.-Gordios (Greek) of knot fame. 

Types.- Holotype is USNM 449783, paratype is 

USNM 449784, from USGS Mesozoic locality D1351, 

Baculites compressus zone of Pierre Shale. 

Description. -A small species about 20 mm in diameter that consists of a small helix surrounded by a planispiral 

coil set at an angle to it (text-fig. 6). Smallest whorl 

preserved is a loose planispire set at a low angle to the helix. 

Ornament on this early whorl consists of rather widely 

spaced, rounded, rectiradiate ribs and about five constrictions. The next two whorls are in contact and form the main 

part of the helix; they have rursiradiate ribbing and three 

constrictions per whorl. Remainder of the shell consists of 

A6 Shorter Contributions to Paleontology and Stratigraphy, 1992 

a planispire set at angles of 40° to 50° to the helix. Cross 

section of the planispire is ovate; height is a little more than 

the width; flanks are broadly rounded, and dorsum and 

venter are more narrowly rounded. Ornament consists of 

rursiradiate ribs of slightly irregular strengths. Stronger ribs 

have small, inconspicuous, bullate ventral tubercles, but no 

umbilical bullae. The base of the body chamber is well out 

on the planispire. 

The holotype (pl. 1, figs. 18-20) is a nearly complete 

shell 20.7 mm in diameter. Its last half whorl is the body 

chamber, which is 6 mm high and 6 mm wide at the 

aperture. The last complete whorl has 64 ribs that cross the 

venter. 

Occurrence.-This species has been found in the 

Baculites compressus zone of the Pierre Shale only at USGS 

Mesozoic localities D1351, D5862, and D5991 in Grand 

County, Colo. (fig. 2). 

Family DIPLOMOCERATIDAE Spath, 1926 

Subfamily DIPLOMOCERATINAE Spath, 1926 

Genus SOLENOCERAS Conrad, 1860 

Type species. -Hamites annulifer Morton, 1842, p. 

213, by original designation by Conrad, 1860, p. 284. 

Solenoceras reesidei Stephenson, 1941 

1941. Solenoceras reesidei Stephenson, p. 401, pl. 77, figs. 

1-3. 

Types.- Holotype is USNM 77238, the original of 

Stephenson, 1941, p. 401, pl. 77, figs. 1-3; there are four 

paratypes, all given the number USNM 77239 and all from 

the Neylandville Marl 4.1 km north of Corbet, Navarro 

County, Tex. 

Diagnosis.-A small species characterized by its 

densely ribbed phragmocone. 
Remarks. -A specimen from USGS Mesozoic locality D1351 of Solenoceras reesidei was recorded from the 

Baculites compressus zone near Kremmling, Colo. (lzett 

and others, 1971, p. A15). 

Family BACULITIDAE Gill, 1871 

Genus BACULITES Lamarck, 1799 

Type species.-Baculites vertebra/is Lamarck, 1801, 

p. 103, by subsequent designation of Meek, 1876, p. 391. 

Baculites compressus Say, 1820 

Plate 2. figures 1-12; plate 3, figure 1 

not 

1820. Baculites compressa Say, p. 41. 

1833. Baculites compressa Say. Morton, p. 291. 

1834. Baculites compressus Say. Morton, p. 43; not pl. 

9, fig. 1. 

1852. Baculites compressus Say?. Owen, pl. 7, fig. 6. 

1856. Baculites compressus Say. Hall and Meek, p. 400, 

pl. 5, fig. 2a--c; pl. 6, figs. 8, 9. 

1876. Baculites compressus Say. Meek, p. 400, text figs. 

55, 56; pl. 20, fig. 3. 

1891. Baculites compressus Say. Brown, p. 159, text 

figs. 1-6. 

1892. Baculites compressus Say. Brown, p. 136, pl. 9, 

figs. 1-11. 

1892. Baculites compressus Say. Whitfield, p. 277, pl. 

46, figs. 1, 2. 

1894. Baculites compressus Say. Hyatt, pl. 3, figs. 

13-18; pl. 4, fig. 1. 

1896. Baculites compressus Say. Gilbert, pl. 62. 

1910. Baculites compressus Say. Grabau and Shimer, p. 

181, figs. 1435, 1436. 

1917. Baculites compressus Say. Dowling, p. 31, 47, pl. 

30, fig. 1. 

1921. Baculites compressus Say. Grabau, text fig. 

1927. 

1931. 

1933. 

1940. 

1965. 

1973. 

1975. 

1975. 

1975. 

1977. 

1982. 

1698a--c. 

Baculites compressus Say. Reeside, p. 10, pl. 9, 

figs. 1-5 [=B. reesidei Elias]. 

Baculites compressus Say. Warren, pl. 2, figs. 

1, 2. 

Baculites compressus Say. Elias, p. 299, 300, pl. 

32, fig. 3 only. 

Baculites compressus Say. Landes, p. 172. 

Baculites compressus Say. Scott and Cobban, 

unnumbered figure. 

Baculites compressus Say. Gill and Cobban, p. 10, 

text fig. 2n. 

Baculites compressus Say. Scott and Cobban, 

unnumbered figure. 

Baculites compressus Say. Hirsch, text fig. 5b. 

Baculites [compressus Say]. Nelson, pl. 65, figs. 

3, 4. 

Baculites compressus Say. Kauffman, pl. 27, figs. 

1, 2. 

Baculites compressus Say. Case, text figs. 12.73, 

12.74. 

1986a. Baculites compressus Say. Scott and Cobban, 

unnumbered figure. 

1986b. Baculites compressus Say. Scott and Cobban, 

unnumbered figure. 

Types. -Morton's types were from the Pierre Shale 

near or at the Great Bend of the Missouri River below Fort 

Pierre, S. Dak. 

Diagnosis.-A moderately large species that has a 

compressed whorl section, smooth flanks in most growth 

stages, a smooth or weakly ribbed venter, and a complex 

suture characterized by the terminal branches of the lateral 

lobe being almost pinched off. 

Description.- Specimens from Middle Park have 

whorl heights of as much as 55 mm. The largest collection, 

from USGS Mesozoic locality Dl351, consists of 63 

fragments that have whorl heights from 5 to 53 mm. Angles 

of taper are mostly 9° for specimens of less than 10 mm in 

diameter, 8° for most specimens from 10 to 19.9 mm, 7° for 

most specimens from 20 to 29.9 mm, and 3° to 5° for 

specimens more than 30 mm. Specimens of less than 30 mm 

in whorl height have smooth flanks and smooth to nearly 

smooth venters (pl. 2, figs. 1-3, 7-9). Larger specimens 

may have weak crescentic flank ribs (pl. 2, figs. 4-6) and 

weak ventral ribs (pl. 2, figs. 6, 12). 

Discussion.-As Gill and Cobban (1973, p. 10) 

noted, the constriction of the terminal branches of the lateral 

lobe of the suture readily separates this species from the 

older Baculites gregoryensis Cobban (1951, p. 820, pl. 

118, figs. 1-5; text figs. 3-13) and B. scotti Cobban (1958, 

p. 660, pl. 90, figs. 1-9; text fig. 1a--e, h). Succeeding 

Baculites cuneatus has a trigonal whorl section, marked 

ventral ribbing, and well-developed flank ribs. 

Occurrence.-Abundant in the Baculites compressus 

zone of the Pierre Shale in Grand County, Colo. , at USGS 

Mesozoic localities D1349, D1351, D5416, D5862, 

D5991, D8092, D8093, D8060, D8086, and D8074 (fig. 

2). 

Baculites undatus Stephenson, 1941 

1941. Baculites undatus Stephenson, p. 405, pl. 79, figs. 

5-10. 

1973. Baculites undatus Stephenson. Cobban, p. 459, figs. 

2-5. 

1974b. Baculites undatus Stephenson. Cobban, p. 5, fig. 3. 

Types.- Holotype is USNM 77245, the original of 

Stephenson (1941 , pl. 79, figs. 5-7), from the N acatoch 

Sand near Chatfield, Navarro County, Tex. Paratypes are 

USNM 77246 and 77247 from the Nacatoch Sand in the 

Chatfield area. 

Diagnosis.-A moderately sized species that has a 

stout, subelliptical cross section; an ornament of strong, 

arcuate, nodelike ribs; and a fairly simple suture characterized by its rectangular, bifid lateral lobe. 

Description.-Two specimens from the Baculites 

compressus zone of the Pierre Shale in the Kremmling area, 

Colo. , were described and illustrated by Cobban (1973, p. 

461, figs. 3k, 1, 4a, 5h-j). These specimens (USNM 

182432 and 182433) have whorl heights of 10 and 20 mm, 

Upper Cretaceous Heteromorph Ammonites from the Baculites compressus Zone of the Pierre Shale, Colorado A7 
stout elliptical sections, and strong crescentic flank ribs. 

The larger and longer specimen has an angle of taper of 5°. 

Occurrence. -USGS Mesozoic locality D8060 near 

Kremmling, Grand County, Colo. The species is known 

from the zones of Didymoceras cheyennense, Baculites 

compressus, B. cuneatus, and B. reesidei in the Pierre 

Shale. 

Superfamily SCAPHIT ACEAE Gill, 1871 

Family SCAPHITIDAE Gill, 1871 

Subfamily SCAPHITINAE Gill, 1871 

Genus JELETZKYTES Riccardi, 1983 

Type species. -Scaphites nodosus Owen (1852, p. 

581 , pl. 8, fig. 4), by original designation of Riccardi 

(1983, p. 14). 

Jeletzkytes nodosus (Owen, 1852) 

Plate 3. figures 2-5, 7-11 

? 

? 

? 

? 

not 

? 

1852. 

1880. 

1892. 

1896. 

1899. 

1905. 

1907. 

1911. 

1916. 

1917. 

1926. 

1933. 

1940. 

1941. 

1970. 

1975. 

1975. 

1980. 

1982. 

Scaphites (Ammonites?) nodosus (N.S.) Owen, p. 

581, pl. 8, fig. 4. 

Scaphites nodosus Owen. Whitfield, p. 441, pl. 

13, fig. 12. 

Scaphites nodosus (Owen). Whitfield, p. 261, pl. 

44, figs. 13, 14. 

Scaphites nodosus Owen. Gilbert, pl. 65, fig. 2. 

Scaphites nodosus (Owen). Logan, p. 209, pl. 22, 

fig. 2; pl. 23, figs. 1-4, 6-12. 

Scaphites nodosus (Owen). Smith, p. 638, fig. 3. 

Scaphites nodosus Owen?. Weller, p. 824, pl. 107, 

figs. 1, 2. 

Scaphites nodosus Owen. Haug, pl. 118 

[=Jeletzkytes quadrangularis (Meek and Hayden)]. 

Scaphites nodosus Owen. Diener, p. 565, text fig. 

6. 

Scaphites nodosus Owen. Dowling, p. 32, pl. 32, 

fig. 3. 

Scaphites reesidei Wade, p. 198, pl. 41, figs. 3-7. 

Acanthoscaphites nodosus (Owen). Elias, p. 320, 

pl. 38, figs. 1-3. 

Acanthoscaphites nodosus (Owen). Landes, p. 

177-178. 

Scaphites rugosus Stephenson, p. 425, pl. 89, figs. 

15-18. 

Scaphites nodosus Owen. Jeletzky, pl. 27, fig. 7. 

Hoploscaphites nodosus (Owen). Hirsch, text fig. 

9b. 

Scaphites [Jeletzkytes nodosus]. Nelson, pl. 64, 

figs. 1, 2 only. 

Acanthoscaphites praequadrispinosus Blaszkiewicz, 

p. 38, pl. 19, figs. 2, 3, 6-8; pl. 20, figs. 1-3, 

6-8; pl. 21, figs. 1-6. 

Acanthoscaphites nodosus (Owen). Case, fig. 12 

(29). 

1983. Jeletzkytes nodosus (Owen, 1852). Riccardi, p. 15, 

pl. 2, figs. 1-8; text figs. 5, 6, 7a. 

Type.-Holotype, by monotype, no. 6381 in the 

collections of the Field Museum of Natural History, ChiA8 Shorter Contributions to Paleontology and Stratigraphy, 1992 

cago, is from the Pierre Shale of the Sage Creek area east of 

the Black Hills in South Dakota. The type, which was 

refigured by Riccardi (1983, pl. 2, figs. 1-3), probably 

came from the Baculites compressus zone. 

Description.-The species is dimorphic. Macroconchs in the collection from Middle Park are as long as 90 

mm, whereas microconchs are 65 mm or less. The phragmocone of macroconchs is very involute and has a deep 

umbilicus (pl. 3, figs. 3, 11). Whorl section ranges from 

compressed ovate to depressed reniform. Coarse distant 

primary ribs arise at the umbilical seam and strengthen 

across the wall and shoulder; they are prorsiradiate on the 

inner to middle flank, flex back over the outer flank, and 

trend nearly straight across the venter (pl. 3, figs. 2, 3). 

Ribs increase by bifurcation and intercalation both high and 

low on the flank. Two rows of tubercles are present. A 

ventrolateral row of conical tubercles arises on some or all 

of the primary ribs when the shell attains a diameter of 

about 10 mm. One to four nontuberculate ribs are intercalated. Pairs of ribs or groups of three ribs loop between the 

ventrolateral tubercles and cross over the venter. Umbilicolateral tubercles are generally present well out from the 

shoulder; they are separated by as many as six nontuberculate ribs. The tubercles give rise to groups of two to four 

ribs that loop or zigzag between the umbilicolateral and 

ventrolateral rows. Macroconch body chambers have a 

straight to slightly convex umbilical wall that occludes part 

of the umbilicus of the phragmocone. The whorl section is 

generally depressed; inner flanks are broadly rounded, outer 

flanks are flattened and convergent, and the venter is very 

broadly rounded. Four or five strong umbilicolateral bullae 

are present on the shaft but disappear on the final hook. 

Corresponding to these bullae are rather coarse ventrolateral 

tubercles, which persist in a much weakened form to the 

final sector of the hook (pl. 3, fig. 11). Narrow ribs cover 

the surface between tubercles, and groups of ribs that 

increase by branching and intercalation link umbilical and 

ventral tubercles. Ribs crowd and weaken toward the adult 

aperture. 

Microconchs have essentially the same ornament on 

the phragmocone as do the macroconchs (pl. 3, fig. 7). The 

body chamber is, in contrast, much less massive and does 

not occlude the umbilicus of the phragmocone. Inner flanks 

are broadly rounded, outer flanks are flattened and convergent, ventrolateral shoulders are broadly rounded, and the 

venter is slightly arched. Six or seven strong umbilicolateral 

bullae are on the shaft but weaken markedly on the final 

hook. Strong, conical ventrolateral tubercles are more 

numerous than the umbilicolateral tubercles; they may be 

restricted to the shaft only or extend onto the final hook, 

where they weaken and generally do not extend to the 

aperture. Delicate ribs arise at the umbilical seam and 

become slightly flexuous and prorsiradiate on the flanks 

where they link in groups of two to four at the tubercles and 

loop or zigzag between them. As many as four nontuber-
culate ribs lie between the tuberculate groups and increase 

by branching and intercalating. Ribs are stronger on the 

venter of the shaft than on the flank; they are slightly 

convex and loop in groups between the tubercles as well as 

intercalate between the tuberculate groups. Ribbing generally weakens and crowds towards the adult aperture. 

Discussion. -Scaphites rugosus Stephenson (1941, 

p. 425, pl. 89, figs. 15-18), from the Nacatoch Sand of 

Navarro County, Tex., is based on a microconch of 

Jeletzkytes nodosus. A further synonym is Scaphites reesidei Wade (1926, p. 198, pl. 41, figs. 3-7), of which 

Scaphites reesidei Collignon (1969, p. 51, pl. 533, figs. 

2098, 2099) is a homonym. Acanthoscaphites praequadrispinosus Blaszkiewicz (1980, p. 38, pl. 19, figs. 2, 3, 6-8; 

pl. 20, figs. 1-3, 6-8; pl. 21, figs. 1-6) is also a synonym 

on the basis of a study of casts of the type material and 

topotypes from the upper Campanian of Poland. 

Occurrence. - Baculites compressus zone of Pierre 

Shale at USGS Mesozoic localities D5991, 05862, D1349, 

D1351, D5416, D8060, D8074, D8086, and D8093 in 

Grand County, Colo. (fig. 2); Baculites compressus, B. 

cuneatus, B. reesidei, and B. jenseni zones in the Western 

Interior; Nacatoch Sand near Chatfield, Navarro County, 

Tex.; Saratoga Chalk, Arkansas; Coon Creek Tongue of 

Ripley Formation, Tennessee; basal Navesink Formation, 

New Jersey; Nostoceras pozaryskii zone, Vistula River 

valley, Poland. 

Genus HOPLOSCAPHITES Nowak, 1911 

Type species. - Scaphites constrictus J. Sowerby, 

1818. 

Hoploscaphites cf. H. landesi Riccardi, 1983 

Plate 3, figure 6 

Compare: 

1983. Hoploscaphites landesi Riccardi, p. 10, pl. 1, figs. 

12-22. 

Description.-A body chamber (USNM 449800), 74 

mm in length, represents a compressed, densely ribbed 

species that resembles H. landesi Riccardi from the Bearpaw Formation of Saskatchewan. The older two-thirds of 

the specimen has flattened flanks and a narrow, flattened 

venter bounded by large ventrolateral clavi. The younger 

third has flattened flanks and a rounded venter that has two 

small nodate ventrolateral tubercles on its older part. The 

preserved part of the aperture is constricted. Ribbing on the 

specimen is flexuous, dense, and prorsiradiate. About 102 

equally spaced ribs cross the venter with slight forward 

bending. A few weak umbilicolateral bullae are present on 

the older half of the body chamber. The concave umbilical 

wall suggests that the specimen is a microconch. 

Discussion.-The specimen resembles H. landesi in 

its compressed form, dense flexuous ribbing, and weak 

umbilicolateral bullae. Differences are mainly the larger 

size of the Middle Park specimen and its larger ventrolateral 

clavi. Riccardi (1983, p. 10) gave lengths of 38 to 48 mm 

for the types. 

Occurrence. -The body chamber described in this 

report is from the Baculites compressus zone of the Pierre 

Shale at USGS Mesozoic locality D 1351, Grand County, 

Colo. (fig. 2). Riccardi (1983, p. 10) stated that the types of 

H. landesi are from the "?Demaine Sandstone, Bearpaw 

Formation" in Saskatchewan. Caldwell (1968, fig. 16) 

showed the Demaine Member of the Bearpaw Formation as 

lying high in the "(?) zone of Didymoceras cheyennense" 

and extending into the lower part of the zone of Baculites 

compressus. 

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p., 45 pis. 

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354, 103 p., 26 pis. 

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Campanian-lower Maastrichtian chalks of the Mons basin, 

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in the Harmignies and Ciply areas: Geologie en Mijnbouw, v. 

68, p. 391--408. 

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Schulz, M.G., Ernst, Gundolf, Ernst, Hartmut, and Schmid, 

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Upper Cretaceous Heteromorph Ammonites from the Baculites compressus Zone of the Pierre Shale, Colorado A 11 

PLATES 1-3 

All specimens are kept at the U.S. National Museum of Natural History (USNM), Washington, 

D.C. Locality numbers refer to U.S. Geological Survey Mesozoic localities in Grand County, Colo. 

(fig. 2, table 1). Contact photographs of the plates in this report are available, at cost, from the U.S. 

Geological Survey Photographic Library, Federal Center, Denver, CO 80225. 
PLATE 1 

[All figures natural size] 

Figures 1-17, 50, 51. Axonoceras compressum Stephenson (p. A3). 

1-3. Hypotype USNM 449773, from locality D1351. 

4, 5. Hypotype USNM 449774, from locality D1351. 

6, 7. Hypotype USNM 449775, from locality Dl351. 

8, 9. Hypotype USNM 449776, from locality Dl351. 

10, 11. Hypotype USNM 449777, from locality D1351. 

12, 13. Hypotype USNM 449778, from locality D1351. 

14, 15. Hypotype USNM 449779, from locality D1351. 

16, 17. Hypotype USNM 449780, from locality D1351. 

50, 51. Hypotype USNM 449781, from locality D5991. 

18-22. Anaklinoceras gordiale n. sp. (p. A6). 

18-20. Holotype USNM 449783, from locality D 1351. 

21, 22. Paratype USNM 449784, from locality D1351. 

23-49. Anaklinoceras reflexum Stephenson (p. AS). 

23-26. Hypotype USNM 449785, from locality D1351. 

27, 28. Hypotype USNM 449786, from locality D1351. 

29-32. Hypotype USNM 449787, from locality D1351. 

33-36. Hypotype USNM 449788, from locality D1351. 

37-40. Hypotype USNM 449789, from locality D1351. 

41, 42. Hypotype USNM 449790, from locality D1351. 

43, 44. Hypotype USNM 449791, from locality D1351. 

45-47. Hypotype USNM 449792. Specimen from the Kremmling area donated by Wayne L. Smiglewski, 

Denver, Colo. 

48, 49. Hypotype USNM 449793, from locality D1351. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024--A PLATE 1 

2 3 

4 5 6 7 8 9 10 11 

~ $i 

\, 

12 13 14 15 16 17 18 19 20 

21 

22 23 24 25 26 

27 28 29 30 

31 32 33 34 

35 36 

37 

38 39 40 

41 42 43 44 

45 46 47 48 49 50 51 

AXONOCERAS AND ANAKLINOCERAS 
PLATE 2 

[All figures natural size] 

Figures 1-12. Baculites compressus Say (p. A7). 

1-3. Hypotype USNM 449794, from locality 01351. 

4-6. Hypotype USNM 449795, from locality 01349. For end view, see pl. 3, fig. 1. 

7-9. Hypotype USNM 449796, from locality 01351. 

10-12. Hypotype USNM 449797, from locality 01351. 
U.S. GEOLOOlCAL SURVEY BULLETIN 2024--A PLATE 2 

6 

4 

2 

3 

7 8 9 10 11 12 

BACULITES COMPRESSUS 
PLATE 3 

[All figures natural size] 

Figure 1. Baculites compressus Say (p. A 7). 

Hypotype USNM 449795, from locality D1349. For other views, see pl. 2, figs. 4-6. 

2-5, 7-11. Jeletzkytes nodosus (Owen) (p. A8). 

2, 3. Hypotype USNM 449798, from locality D1349. 

4, 5. Hypotype USNM 449799, from locality D1349. 

7-9. Hypotype USNM 449801, from locality D1349. 

10, 11. Hypotype USNM 449802, from locality D1351. 

6. Hoploscaphites cf. H. landesi Riccardi (p. A9). 

Figured specimen USNM 449800, from locality D 1351. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024--A PLATE 3 

2 

3 

9 10 

BACULITES, JELETZKYTES, AND HOPLOSCAPHITES 

Chapter B 

The Siphonophrentidae (Rugose Corals, 

Devonian) of Eastern North America 

By WILLIAM A. OLIVER, jR. 

A systematic description and analysis of the genera and species 

of a family of simple rugose corals that was a characteristic part 

of Early and Middle Devonian faunas within the Eastern 

Americas Biogeographic Realm 

U.S. GEOLOGICAL SURVEY BULLETIN 2024 

SHORTER CONTRIBUTIONS TO PALEONTOLOGY AND STRATIGRAPHY, 1992 

CONTENTS 

Abstract Bl 

Introduction Bl 

Acknowledgments Bl 

Biogeography B2 

Stratigraphic Setting B2 

Coral Morphology B4 

Systematic Paleontology BS 

Depositories BS 

Family Heterophrentidae Kullmann, 1965 BS 

Genus Heterophrentis Billings, 1875 BS 

Heterophrentis spatiosa (Billings), 1858 BS 

Family Unknown B6 

Genus Triplophyllum Simpson, 1900 B6 

Triplophyllum terebratum (Hall), 1883 B7 

Family Siphonophrentidae Merriam, 1974 B7 

Genus Briantelasma Oliver, 1960 BS 

Briantelasma boucoti n. sp. BS 

Genus Pseudoblothrophyllum Oliver, 1960 BS 

Genus Compressiphyllum Stumm, 1949 B9 

Compressiphyllum davisana (Miller), 1889 B9 

Genus Metaxyphrentis n. gen. B9 

Metaxyphrentis prolifica (Billings), 1858 BlO 

Genus Enallophrentis n. gen. B12 

Enallophrentis simplex (Hall), 1843 B12 

Enallophrentis broweri n. sp. B16 

Genus Breviphrentis Stumm, 1949 B18 

Breviphrentis invaginata (Stumm), 1937 B19 

Breviphrentis halli (Milne-Edwards and Haime), 1851 B20 

Breviphrentis cista n. sp. B21 

Breviphrentis pumilla n. sp. B24 

Genus Siphonophrentis O'Connell, 1914 B25 

Siphonophrentis elongata (Rafinesque and Clifford), 1820 B26 

Locality Data B28 

Onondaga Limestone Localities B28 

Hamilton Group Localities B28 

Skaneateles Formation, Delphi Station Member, Delphi Coral Bed B28 

Ludlowville Formation, Centerfield Member B29 

Ludlowville Formation, Otisco Shale Member, Staghom Point Coral 

Bed B29 

Moscow Formation, Windom Shale Member, Fall Brook Coral Bed B29 

References Cited B29 

Contents Ill 
IV Contents 

PLATES 

[Plates follow references cited] 

1. Heterophrentis spatiosa (Billings), Triplophyllum terebratum (Hall), Metaxyphrentis prolifzca (Billings), and Kionelasma? sp. cf. Heterophrentis spatiosa 

(Billings). 

2. Heterophrentis spatiosa (Billings) and Metaxyphrentis prolifzca (Billings). 

3. Metaxyphrentis prolifzca (Billings). 

4. Compressiphyllum davisana (Miller). 

5-9. Enallophrentis simplex (Hall). 

10. Enallophrentis simplex (Hall) and E. broweri n. sp. 

11, 12. Enallophrentis broweri n. sp. 

13. Breviphrentis invaginata (Stumm). 

14. Breviphrentis halli (Milne-Edwards and Haime) and B. cista n. sp. 

15, 16. Breviphrentis halli (Milne-Edwards and Haime). 

17, 18. Breviphrentis cista n. sp. 

19. Breviphrentis pumilla n. sp. 

20-22. Siphonophrentis elongata (Rafinesque and Clifford). 

FIGURES 

1. Chart showing Devonian lithologic units and approximate stage boundaries in 

New York B3 

2-10. Graphs of morphologic data for: 

2. Metaxyphrentis prolifzca Btl 

3, 4. Enallophrentis simplex B14 

5. Enallophrentis broweri B17 

6. Breviphrentis invaginata B20 

7. Breviphrentis halli B22 

8. Breviphrentis cista B23 

9. Breviphrentis pumilla B24 

10. Siphonophrentis elongata B27 
The Siphonophrentidae (Rugose Corals, Devonian) 

of Eastern North America 

By William A. Oliver, Jr. 1 

Abstract 

Rugose corals belonging to the Family Siphonophrentidae Merriam, 1974, are morphologically simple 

and extremely variable. They are widely distributed in 

Lower and Middle Devonian strata in areas that were 

formerly parts of the Devonian Eastern Americas Biogeographic Realm. The family, its constituent genera, and all 

type species are reviewed or redescribed. In addition, all 

previously named species and some additional species 

from New York are described. 

Two new genera are Metaxyphrentis, type species M. 

prolifica (Billings), 1858, and Enallophrentis, type species 

E. simplex (Hall), 1843. New species are Briantelasma 

boucoti (Lochkovian) and Enallophrentis broweri, Breviphrentis cista, and B. pumilla (all Givetian). Heterophrentis 

Billings, 1874 (and Heterophrentidae Kullmann, 1965), and 

Triplophyllum Simpson, 1900, are restricted to the type 

specimens of their type species because they are unrecognizable. 

INTRODUCTION 

The Siphonophrentidae are an important eastern 

North American family of Early and Middle Devonian 

rugose corals. Well over 100 nominal species have been 

described in a few genera, mostly from Middle Devonian 

strata. Siphonophrentids occur also in western North America and other parts of the world but are most common, and 

apparently originated, in the Eastern Americas Realm 

(EAR). This Devonian biogeographic area included eastern 

North America and northern South America. 

Siphonophrentids are morphologically simple and, 

for this reason, are a taxonomically difficult group. They 

are solitary hom-shaped corals consisting of wall, septa, 

and tabulae; there are no dissepiments or other features that 

tend to add character and make coral species recognizable. 

Individual variation is extensive. This variability, combined 

with the simple morphology and small number of characManuscript approved for publication February 26, 1992. 1 U.S. Geological Survey, U.S. National Museum of Natural 

History, Washington, DC 20560. 

ters, makes discrimination of taxa questionable. If all the 

siphonophrentids from eastern North America were thrown 

into one large container, I doubt that any consistent grouping could be made; boundaries between species and even 

genera would be fuzzy at best. 

Siphonophrentid microstructure is also simple, being 

originally trabeculate although commonly preserved as 

fibro-lamellar. It is consistent within the family, but it is a 

common kind of structure that is also found in many other 

groups of rugose corals. 

My approach to the species problem is through the 

"population." This, for my purpose, is the assemblage of 

individuals in a limited stratigraphic interval and geographic 

area that one might recognize as representing a species if no 

other information were available. Populations from different intervals and (or) areas that seem to be sufficiently alike 

are combined into species. In each species, I use a sample 

of the topotype population plus the type specimen(s) as the 

basis for my species concept; then I expand upon this 

through analysis of additional population samples if available. It has not been practical to describe all populations, 

but I have examined enough to at least roughly determine 

variation within the New York sequence. Material from 

places outside New York has been used to settle questions 

of priority and to describe taxa originally based on non-New 

York specimens. 

My original intention was to describe all the upper 

Middle Devonian (Givetian; Hamilton Group and Tully 

Limestone) species from New York. To do this within a 

reasonable period of time proved impractical because of the 

large number of faunas and the amount of individual 

variation. The study remains centered in New York, but it 

is not monographic in scope. I describe the topotype 

populations of the type species of the Middle Devonian 

genera, and all other species previously described from 

New York. Many populations, and probably some species, 

remain to be analyzed. 

Acknowledgments 

Thin sections were prepared by William C. Pinckney, 

Jr., and Keith Moore; most of the photography is by 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 81 
Mr. Moore, although the photographs on plate 1 are by 

N.W. Shupe and R.H. McKinney. All work was done in 

laboratories of the U.S. Geological Survey (USGS), Washington, D.C. 

I am indebted to the following individuals and institutions for the loan of important specimens: T.E. Bolton, 

Geological Survey of Canada, Ottawa; G .R. Gunnell, 

University of Michigan, Museum of Paleontology, Ann 

Arbor; Melvin Hinkley, American Museum of Natural 

History, New York; Ed Landing, New York State Museum, 

Albany; Abel Prieur, Ecole des Mines Collection, Universite Claude Bernard, Lyon; and Pierre Semenoff-TianChansky, Museum National d'Histoire Naturelle, Paris. Dr. 

Semenoff and Mme. Daniele Decrouez, Museum d'Histoire 

Naturelle de Geneve, answered questions about additional 

specimens. As always, the personnel of the U.S. National 

Museum of Natural History, Washington, D.C., especially 

Jann Thompson, have been very cooperative and helpful. 

Drafts of the paper were critically reviewed by J. T. 

Dutro, Jr., and W.J. Sando, both USGS, Washington, 

D.C., and by J.E. Sorauf, State University of New York, 

Binghamton, N.Y. I am particularly indebted to Professor 

Sorauf for many discussions of coral microstructure and its 

interpretation. 

BIOGEOGRAPHY 

The Early and Middle Devonian world was remarkably provincial, and the corals were among the most 

provincial of marine animals. Three marine biogeographic 

realms are recognized: the Eastern Americas Realm (EAR), 

the Old World Realm (OWR), and the Malvinokaffric 

Realm. The EAR, discussed below, and the much larger 

OWR had extensive coral faunas during most of Early and 

Middle Devonian time; the Malvinokaffric Realm was coral 

poor, presumably because of high latitudes. Pertinent 

descriptions and discussions of Devonian rugose coral 

biogeography are in Oliver (1977, 1980, 1990) and Oliver 

and Pedder (1979, 1989). 

The Eastern Americas Realm included the North 

American plate east of the Transcontinental Arch and south 

of the central Canadian Shield, plus northern South America (principal coral faunas are in Venezuela and Colombia). 

During the Pragian and early Emsian, the EAR extended 

west to include the Great Basin (Nevada and surrounding 

areas), although at other times this was part of the OWR. 

Four EAR provinces are recognized: ( 1) the Appohimchi 

Province included all of eastern North America during the 

Early Devonian but was essentially limited to the 

Appalachian-Ouachita belt during the Middle Devonian; 

(2) during the Middle Devonian, the Michigan Basin 

Province included eastern North American areas north of 

the Ozark and Nashville domes and west of the Cincinnati, 

Findlay, and Algonquin arches; (3) the Great Basin Prov82 Shorter Contributions to Paleontology and Stratigraphy, 1992 

ince was part of the EAR during Pragian and early Emsian 

times; and (4) the Venezuelan-Colombian Province is recognized during late Emsian or early Eifelian to early 

Givetian time. The most recent description of the EAR 

provinces is in Oliver and Pedder (1989). 

The Siphonophrentidae are most common and diverse 

in eastern North America, and the earliest known family 

members are from this area. It seems likely that the family 

and most of the genera were initially endemic to the EAR. 

STRATIGRAPHIC SETTING 

In eastern North America, siphonophrentids range 

through most of the Lower and Middle Devonian. Lochkovian (Helderbergian) and Pragian (Deerparkian) corals are 

best known from New York and eastern Quebec, respectively, although Pragian coral faunas are poorly developed 

almost everywhere. Ems ian and Middle Devonian coral 

faunas are widespread, and siphonophrentids have been 

described from many areas, but principally New York, 

southwestern Ontario, Michigan, and the region around the 

Falls of the Ohio River in Kentucky and Indiana. The taxa 

described or discussed in this paper are from all of these 

areas except Michigan, with some added from Nevada. 

Most of the corals are from New York, and the stratigraphic 

terminology used in this paper is principally from this area. 

Figure 1 is a simplified version of the New York sequence 

showing major formations, important coral-bearing members and beds, and other units that are referred to in the text. 

The positions of the internationally recognized stage 

boundaries in eastern North America were reviewed by 

Oliver and Rickard (in Kirchgasser and others, 1985) and 

are shown in figure 1. The positions of all the boundaries, 

with the exception of the Middle-Upper Devonian series 

boundary, are approximate because of the EAR provincialism, but this uncertainty has little effect on the stage 

assignment of the corals because so few are derived from 

the questionable intervals. However, the position of the 

Emsian-Eifelian boundary does affect the age of the corals 

in the Edgecliff Member of the Onondaga Limestone. 

Briefly, the upper half of the Nedrow Member of the 

Onondaga is well dated by both conodonts and ammonoids 

as Eifelian, but not earliest Eifelian. The Bois Blanc 

Formation is dated as (early?) Emsian by conodonts and 

brachiopods. The Edgecliff and lower half of the Nedrow 

are in between, and I (Oliver, 1989) have argued that the 

Figure 1. Generalized Devonian succession in New York ....,. 

showing lithologic units mentioned in text and approximate stage boundaries. The standard conodont zones 

recognized in New York are from Kirchgasser and 

others (1985), with minor updating; the asterisk (*) 

signifies the presence of the Lowermost asymmetricus 

Zone and probably the upper part of the Upper disparalis Zone in the Lodi Limestone. Diagonal lines mark 

missing parts of the succession. 
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recognized 

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Lodi Limestone 

~-n·e--s-eo __ s_ha_l_e ________________ __ 

~ \\\\\\\ ~ )~~d xxxxxxxxxxxxxxxxxxxxx Tully Limestone 

Windom Shale Member 

xxxxxx Fall Brook coral bed 

Windom 

Shale 

Member 

Deep Run Shale Member Portland Point Limestone Member 

\ Tichenor Limestone Member 

Jaycox Shale Member King Owasco Member 

Ferry Spafford Member 

Member Ivy Point Member xxx Coral bed 

Wanakah Shale Member 

xxxx Joshua coral bed 

Ledyard Shale Member Otisco Shale Member 

xxxxxxxxx Stag horn Point coral bed 

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

~-Q) 

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Tully 

Limestone 

c: 

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Centerfield Member Stone Mill Limestone Member c: ~--------------------------------------------------------------------------------4---~ ~ 

Levanna Shale Member 

Butternut Shale Member 

Pompey Member 

----xxx Delphi coral bed 

Delphi Station Member 

r---------------------------------------------------------- - - - Stafford Limestone Member Mottville Member xxx Case Hill coral bed 

Pecksport Member 

Cardiff Solsville Member Solsville coral bed xxx Oatka Creek Shale Member Shale Member 

Bridgewater Member 

Chittenango Shale Member 

~-- _ _ _ Cherry Valley Limestone Member Stony Hollow Member 

Seneca Member ~----------------''-U-n_i....:on--S-pr-in_g_s_S_h_a_le_M __ e_m_b_e..:..r ----------1 

Clarence Member I 

\\\\\\\\\\\\ 

Bois Blanc Formation 

I 

Akron Dolomite 

Moorehouse Member - - _________________________ _, Nedrow Member 

Edgecliff Member 

Oriskany Sandstone 

Cobleskill Limestone ----- - - - - - - - ·---------'--------------------1 

.E en CO ~c::::c 

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varcus 

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co status 

co status 

serotinus 

delta 

woschmidti 
Edgecliff is most likely Eifelian because of faunal relations 

in Pennsylvania; it is so treated in this paper. 

Southwestern Ontario corals referred to in the text are 

from the Niagara Peninsula, in the vicinity of Port Col borne 

(29 km west of Buffalo, N.Y.). The stratigraphic section is 

essentially the same as that in western New York (fig. 1; 

Oliver, 1976, p. 5-12), except that the Hamilton Group is 

inaccessible beneath Lake Erie. Because early workers in 

the area did not separate the Bois Blanc Formation and 

Onondaga Limestone, the provenance of many type and 

illustrated specimens is in doubt. 

The Falls of the Ohio River are in Louisville, Ky., 

and Jeffersonville, Ind., but numerous outcrops, both north 

and south of the river, have been additional sources of 

described corals. The Falls stratigraphy was reviewed by 

Oliver (1976, p. 19-20). The Jeffersonville Limestone is 

only 9 m thick but can be divided into six assemblage 

zones. The lowest zone is Emsian; the others are Eifelian in 

age. The "coral zone," cited by Stumm (1965) and others, 

includes the lower two or three zones and is both Emsian 

and Eifelian in age. The Jeffersonville is overlain by the 

Sellersburg Limestone, of which the Beechwood Limestone 

Member is an important source of corals. This unit is 

Givetian in age and is the approximate equivalent of the 

Centerfield Member of the Ludlowville Formation in New 

York. 

The coral biostratigraphy of the Lower and lower 

Middle Devonian in Nevada was revised and summarized 

by Johnson and Oliver (1977) and was based on the work of 

Merriam (1972, 1974a,b). Siphonophrentid corals there are 

of Pragian and Emsian age; those described in this paper are 

Emsian, from Coral Zone D2 of Merriam (1974b), which is 

within the Eurekaspirifer pinyonensis Zone. 

CORAL MORPHOLOGY 

Insofar as is practical, descriptive terminology is 

based on Hill (1981, Glossary, p. F32-F36), but a few 

terms are defined or discussed in this section because they 

were omitted by Hill or because of their particular application or importance to the siphonophrentids. 

Siphonofossula. A siphonofossula is a cardinal fossula formed by "the down-bending of the successive tabulae 

to form a series of invaginated funnels, . . . a peculiar type 

[of fossula] which may or may not be accompanied by an 

abortion of the cardinal septum" (O'Connell, 1914, p. 187). 

Cardinal fossulae are commonly formed by a short cardinal 

septum; adjacent major septa may be deflected or modified 

to further define the fossula, and some tabular downbending 

is common. O'Connell introduced the term to describe the 

extreme downbending found in a few corals such as 

Siphonophrentis. In the siphonophrentids, the cardinal fossula is formed by a combination of a short cardinal septum, 

modification of adjacent major septa, and tabular down84 Shorter Contributions to Paleontology and Stratigraphy, 1992 

bending. The downbending is most extreme in Siphonophrentis but is present in all family members and is 

sufficient in most to be called a siphonofossula. The 

important point is that there is a continuum from greatest to 

least, and no boundary between genera can be recognized 

on this character. 

Protosepta. Four protosepta are recognized: cardinal 

(C), counter (K), and two alar septa. In the systematic part, 

these are generally described from transverse sections and 

used to define the bilateral symmetry, which commonly is 

best developed in the cardinal quadrants. 

Septotheca. The siphonophrentid outer wall includes 

two layers, an outer epitheca and an inner theca formed by 

the thickened and joined peripheral ends of the septa. The 

epitheca is very thin and may or may not be preserved. The 

inner layer, of septal origin, is thicker and is commonly 

well preserved and prominent; it is here referred to as a 

septotheca. This term has been most commonly used for a 

similar wall structure in scleractinian corals but was 

extended to rugosans by Kato (1963, p. 611-613) and is 

used here even though the scleractinian wall may have been 

formed differently. 

Ontogenetic stages. The terms brephic, neanic, 

ephebic, and gerontic are used more or less conventionally. 

The brephic stage is the earliest recognizable in the skeleton; it extends from the first deposition of skeletal matter 

through the insertion of the protosepta, but it has not been 

recognized in any of the described species. The neanic stage 

represents adolescence, between the brephic stage and the 

ephebic stage with the fully developed adult skeletal characters. The gerontic stage represents senescence, but may or 

may not be present in a given corallum. Only the brephic 

stage is clearly defined. In cylindrical corals it can be 

argued that conical, postbrephic growth is neanic and 

cylindrical growth is ephebic, but in conical corals this 

criterion cannot be used. In descriptions of the various 

siphonophrentid species, it is convenient to use the ontogenetic stage names for successive morphologies, but the 

stages are separately defined for each species, and I do not 

mean to imply that the stages in the different corals are 

necessarily analogous or represent comparable ages or 

maturity. 

The differences between apparent ontogeny as seen in 

serial transverse sections and the true ontogeny experienced 

by the growing individual are significant. The youngest part 

of the skeleton included in a transverse section tends to be 

in the center, and the oldest part is toward the outside; 

irregularities, such as the fossula in siphonophrentids, 

violate the generality, but the point is that the skeletal units 

in any section are of varied ages and can belong to different 

growth stages. In addition, structures formed during the 

later growth of any individual would have been somewhat 

changed if that individual had lived longer. These ontogenetic factors must be considered in the analysis of variation 

in any population of rugose corals. 
SYSTEMATIC PALEONTOLOGY 

Depositories 

The following descriptions are based principally on 

collections I have made while employed by the U.S. 

Geological Survey. Described specimens from these collections are now deposited in the U.S. National Museum of 

Natural History. Specimens from older collections are from 

several additional sources. Abbreviations used in the text 

are as follows: 

AMNH -American Museum of Natural History, New 

York, N.Y. 

EM -Ecole des Mines (Paris), collection de Verneuil; now at Universite Claude Bernard, 

Lyon, France. 

GSC -Geological Survey of Canada, Ottawa. 

NYSM -New York State Museum, Albany. 

UMMP -University of Michigan, Museum of Paleontology, Ann Arbor. 

USGS -U.S. Geological Survey, Washington, D.C. 

USNM -U.S. National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 

Family HETEROPHRENTIDAE Kullmann, 1965 

part 1965. Heterophrentinae Kullmann, p. 140. 

not 1978. Heterophrentinae Kullmann, Birenheide, p. 68. 

not 1989. Heterophrentidae Kullmann, Oliver, p. 5. 

Discussion.-The genus Heterophrentis is here 

restricted to two syntype specimens of its type species (see 

following). Under the rules of zoological nomenclature, the 

family name, Heterophrentidae, which is ultimately based 

on the same specimens, is also so restricted. The family 

concept of the original author of the name, and of all 

subsequent users, is preserved in the name Siphonophrentidae Merriam, 1974a (see diagnosis and discussion). 

Genus HETEROPHRENTIS Billings, 1875 

part 1875. Heterophrentis Billings, p. 235. 

part 1889. Heterophrentis Billings, 1875. Miller, p. 193 

(designated H. spatiosa as type). 

part 1914. Heterophrentis Billings. O'Connell, 1914, p. 183 

(her suggestion that H. proli.fica was the type 

species is incorrect). 

part 1940. Heterophrentis Billings. Lang, Smith, and Thomas, 

p. 68. 

part 1981. Heterophrentis Billings. Hill, p. Fl47. 

not Heterophrentis of most authors. 

Type species. -Zaphrentis spatiosa Billings, 1858a, 

by subsequent designation of Miller, 1889, p. 193. See 

below. 

Diagnosis or description. -As for species, below. 

Discussion.-The genus and type species are unrecognizable (see species description that follows), and the use 

of the names should be restricted to the two original 

syntypes. The above synonymy includes references that are 

of particular help in understanding the nomenclatural situation, but no one has previously illustrated or accurately 

described either of the two specimens on which the genus is 

ultimately based. The Heterophrentis concept of most 

authors is based on Zaphrentis simplex Hall, 1843, as 

illustrated by Hall, 1877, pl. 21 , particularly in figures 5, 8, 

9, and 10. This species is redescribed below and assigned to 

Enallophrentis n. gen. in the Family Siphonophrentidae 

Merriam, 1974a. 

Heterophrentis spatiosa (Billings), 1858 

Plate 1, figures 1-4; plate 2, figures 1, 2 

1858a. Zaphrentis spatiosa Billings, p. 178. 

1858b. Z. spatiosa Billings. Billings, p. 430. 

1859. Z. spatiosa Billings. Billings, p. 123. 

1874. Z. spatiosa Billings. Nicholson, p. 24. 

1875. Heterophrentis spatiosa (Billings). Billings, p. 

235-236. 

1889. H. spatiosa (Billings). Miller, p. 193. 

part 1901. Streptelasma prolifica (Billings). Lambe, p. 

115-117. 

1958. H. spatiosa (Billings). Sutherland, p. 45. 

1960. Z. spatiosa Billings. Bolton, p. 60. 

Type specimens. -Lectotype, here selected, GSC 

3451, one of the two syntypes of Billings (Bolton, 1960, p. 

60), pl. 1, figs. 1, 2; pl. 2, fig. 2. Paralectotype, GSC 

3451a, syntype of Billings (Bolton, 1960, p. 60), pl. 1, 

figs. 3, 4; pl. 2, fig. 1. Bois Blanc Formation or lower part 

of Onondaga Limestone (Edgecliff Member), Rama's 

Farm, Port Colborne, Ontario. 

Original description.-Billings' original description 

follows: 

Corallum short, turbinate, moderately curved and 

very broadly expanding. At the margin of the cup about 

ninety radiating septa alternately a little unequal and with 

their edges broadly rounded as in Z. prolifica. Length 

measured on the side of the greater curvature, about 

three inches [7.6 em], width of cup two inches and a half 

[6.4 em]. Septal fossette unknown. 

This species is closely related to Z. prolifica, and 

may perhaps be united with it when its characters become 

more fully known. 

Formation and locality.-Devonian, Onondaga and 

Corniferous limestones, Rama's Farm, near Port Colborne 

Canada West. [Billings, 1858a, p. 178.] 

Description of type specimens. -Both specimens are 

silicified and have silicified matrix filling the lower part of 

the calice; they are very similar and probably conspecific. 

Coralla are smoothly curved, turbinate to trochoid; the 

initial expansion angle of 60 degrees decreases to 40 

degrees (midway) and to 25 degrees at the upper edge. 

Maximum diameter, 65 mm; length on convex side, 7 5 and 

70 mm. Cardinal septum irregularly located relative to 

curvature. Calice broad and shallow, depth is less than half 

the diameter; a pointed axial boss projects some 4 mm 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 85 
above the calice floor. Lower cal ice not exposed; no fossula 

evident on upper half of calice wall. 

Major septa number 47(?) in calice. In the middle 

calice, distal edges of septa are sharp becoming rounded 

upward; at the calice margin, they are flattened parallel to 

the calice wall (spatulate); rounded and flattened distal 

edges and lateral surfaces of thicker parts of septa are 

markedly dimpled, suggesting the presence of multiserial 

small trabeculae. 

Internal features are poorly preserved because of 

complete silicification; description based on transverse thin 

section and resulting polished faces of lectotype (the section 

is through the lowest part of calice and boss and is tilted so 

that the cardinal septum and right quadrant, looking down, 

are higher in the calice than the rest of the section) and 

longitudinal thin section (in plane of curvature) of paralectotype. Septa are bilateral in arrangement; cardinal septum 

short with major septa on either side subparallel, becoming 

longer and radial toward alar positions; major septa in 

counter quadrants are thickened and irregularly meet, with 

many or most extending to the axis. The apparent contrast 

in septal length and thickening is partly due to the poor 

orientation of the section, but the arrangement indicates the 

presence of a fossula in the lower calice. Tabulae incomplete, but form irregular tent-shaped pattern. No dissepiments. Morphological features in the thin and polished 

sections are discernible where structures are discrete or 

separated by matrix or clear silica; in other areas, features 

tend to merge and be indistinct; microstructure not preserved. 

Discussion. -H. spatiosa was selected as the type 

species of Heterophrentis by Miller (1889). However, 

concepts of the genus have generally been based on 

Zaphrentis prolifica Billings, 1858a, which was one of 

several species assigned to Heterophrentis by Billings 

(1875) in his original description of the genus, or, more 

commonly, on Zaphrentis simplex Hall, 1843, which was 

well illustrated by Hall (1877, pl. 21), and was referred to 

Heterophrentis by Stewart (1938), Stumm (1942 and later), 

and most subsequent workers. H. spatiosa is not congeneric 

with either of these and does not match any concept of the 

genus in current use. 

The discernible characters in the H. spatiosa syntypes 

that are likely to be important in determining relationships 

are the boss and small fossula in the lower calice, the 

distally thickened and spatulate septa with multiserial small 

trabeculae, and the tent-shaped tabulae. These are suggestive of various nominal species, many now assigned to 

Kionelasma Simpson, 1900 (for example, pl. 1, figs. 

16-18), although typical Kionelasma has a more strongly 

developed axial structure and boss and may lack multiserial 

trabeculae (not described in that genus). The size of the 

axial structure is unlikely to be important, and the trabeculae 

may not have been recognized if developed only at the 

calice margins of mature individuals. However, given the 

86 Shorter Contributions to Paleontology and Stratigraphy, 1992 

limited morphologic data in the syntypes, other specimens 

would have to be very similar in those features that are 

known, including the wide angle of expansion, before they 

could be considered synonymous. At present, I am not 

aware of any form that fulfills these requirements, and it 

seems best to consider both Heterophrentis and its type 

species, H. spatiosa, to be unrecognizable except in the two 

type specimens. 

Family Unknown 

Genus TRIPLOPHYLLUM Simpson, 1900 

part 1900. Triplophyllum Simpson, p. 209. 

part 1914. Triplophyllum Simpson. O'Connell, p. 184, 190. 

part 1938. Triplophyllum Simpson. Stewart, p. 27. 

1940. Triplophyllum Simpson. Lang, Smith, and Thomas, 

p. 135. 

part 1944. Triplophyllum Simpson. Easton, p. 38. 

part 1944. Triplophyllum Simpson. Shimer and Shrock, p. 93. 

not? 1962. Triplophyllum Simpson. Ivanovsky, p. 122-123. 

not? 1963. Triplophyllum Simpson. Ivanovsky, p. 55. 

1981. Triplophyllum Simpson. Hill, p. F160. 

Type species. -Zaphrentis terebrata Hall, 1883, p. 

316, pl. 23, fig. 5, by original designation. Devonian, Falls 

of the Ohio. 

Diagnosis or description. -As for type species, 

below. 

Discussion. -Simpson (1900) designated a Devonian 

coral as type, but illustrated and seems to have mainly based 

his description on a Carboniferous species. In the only 

serious study of the genus and its type species, Easton 

(1944, p. 38-39) separated the Carboniferous species as 

Triplophyllites. His description of Triplophyllum terebratum was based on photographs of the holotype plus four 

specimens from the type locality that he considered to be 

conspecific. Easton convincingly showed that the Devonian 

and Carboniferous corals were generically different, but he 

did not provide a satisfactory basis for understanding 

Triplophyllum. 

Simpson (1900) separated Triplophyllum from 

"Zaphrentis" on two characters: (1) the presence of two alar 

fossulae in addition to the cardinal fossula and (2) microstructure. Alar fossulae are variably developed in many 

"Heterophrentis," and most workers have not considered 

this an adequate basis for generic discrimination, although 

Stewart (1938) and Ivanovsky (1962, 1963) both assigned 

species (other than the type) to Triplophyllum on this basis. 

As Easton noted, the alar fossulae are weak or absent in T. 

terebratum in any case. 

Simpson's (1900) description of the microstructure is 

unclear, but it was apparently based on Carboniferous 

specimens; the T. terebratum holotype is silicified and fine 

structure is not preserved. Easton's "topotypes" are also 

poorly preserved internally, and he did not describe microstructure. 

Stumm (1949, p. 12) placed Triplophyllum in synonymy with Heterophrentis Billings, and most subsequent 
workers have accepted this. With the realization that Heterophrentis cannot be used in the way that most authors 

have intended, it becomes important to determine if Triplophyllum is a suitable replacement name. It is not, because 

the type species and genus are unrecognizable (see species 

description), and the use of the name should be restricted to 

the holotype. 

Triplophyllum terebratum (Hall), 1883 

Plate 1 , figures 5-7 

1883. Zaphrentis terebrata Hall, p. 316, pl. 23, fig. 5. 

1890. Z. terebrata. Lesley, p. 1278, text-fig. 

part? 1944. Triplophyllum terebratum (Hall). Easton, p. 38, pl. 

17, figs. 17, 18 (? fig. 16 and pl. 11, figs. 1, 2). 

part? 1949. Heterophrentis terebrata (Hall). Stumm, p. 11, pl. 

5, fig. 12 (? fig. 13). 

part? 1965. H. terebrata (Hall). Stumm, p. 22, pl. 9, figs. 3, 4 

(?figs. 1, 2 and synonymy). 

1981. Triplophyllum terebratum (Hall). Hill, p. F160, 

fig. 89:2. 

Type specimen.-Holotype (the only specimen of 

Hall, 1883), NYSM 341, pl. 1, figs. 5-7; Jeffersonville 

Limestone, Falls of the Ohio at Louisville, Ky. The 

specimen could be from either the Emsian or Eifelian part of 

the formation. 

Description.-The holotype is ceratoid, 22 mm in 

maximum diameter, and nearly 60 mm long; the exterior is 

worn but there is a suggestion of weak rugae. The calice 

margin is broken, and so original depth and shape are 

unknown, but it appears to have been deep with a narrow 

axial pit. The cardinal fossula is deep, parallel sided, and 

occupied by a thin, distally short cardinal septum; flanking 

major septa are pinnately arranged and radially short, 

becoming longer away from the fossula; septa in the counter 

quadrants are radially arranged. Major septa number 48; in 

addition to the four protosepta, there are 8 majors in each 

cardinal quadrant and 14 in each counter quadrant. Most 

major septa extend 0.7 to 0.8 the radius and are bent 

counterclockwise (viewed distally) at their axial ends. 

There are no alar fossulae of special note. Minor septa are 

inconspicuous, possibly contratingent. 

The tabulae that can be seen are concave in the axial 

area; their morphology away from the axis is not known. No 

dissepiments. 

The above description is based on examination of the 

uncut holotype. The specimen was subsequently cut transversely, but internal preservation is so poor that very little 

can be added to the description. The cut was made approximately 5 mm below the bottom of the calice at a diameter 

of 13 mm; here the cardinal septum is long and prominent, 

but other septa are irregularly broken and cannot even be 

counted. The interior is mostly empty. 

Discussion. -Simpson (1900) emphasized the alar 

fossulae in his generic description, but Hall's (1883) original description did not mention them, and neither did any 

subsequent description of the holotype; Easton (1944, p. 

38) stated, "alar fossulae either absent or doubtfully identified." Easton's species description differs from my 

description of the holotype in noting the presence of 

dissepiments and of minor septa in the "topotypes." I have 

not examined these, but they may not be conspecific with 

the holotype. 

Any understanding of Triplophyllum must ultimately 

be based on the single specimen. The specimen does not 

preserve the characters needed to characterize the species or 

genus, and clearly the generic name is not a suitable 

replacement for Heterophrentis. 

Family SIPHONOPHRENTIDAE Merriam, 1974 

part 1965. Heterophrentinae Kullmann, p. 140. 

1972. Siphonophrentinae Merriam, p. 28, nomen nudum. 

1974a. Siphonophrentinae Merriam, p. 23 (dated 1973, 

pub. January 1974). 

1974b. Siphonophrentinae Merriam, p. 41 (pub. October 

1974). 

1974. Siphonophrentinae Birenheide, p. 253 (pub. 

November 4, 1974). 

1978. Heterophrentinae Kullmann. Birenheide, p. 68. 

1989. Heterophrentidae Kullmann. Oliver, p. 5. 

Diagnosis.-Trochoid to cylindrical solitary Rugosa 

with moderately to very deep calice and a distinct cardinal 

fossula, commonly formed by a distally short cardinal 

septum and downbent tabulae. Septal arrangement bilateral; 

major septa long or short and slightly to greatly amplexoid, 

with a pinnate arrangement in the cardinal quadrants. 

Arrangement tends to be radial in the counter quadrants and 

to become radial in the cardinal quadrants of larger forms 

and those in which the septa are withdrawn from the axis. 

The wall consists of a septotheca formed by the dilation of 

both major and minor septa and a thin epitheca. Tabulae 

tend to be complete, arched with an axial depression; this 

M -shape in longitudinal section may be strongly marked or 

only gently developed. No dissepiments or axial structure. 

In early ontogenetic stages, septa are commonly 

dilated and may fill the lumen. Such dilation persists to 

maturity in some forms. Attenuation takes place with 

growth in most family members and tends to be uniform in 

all quadrants, although it is irregular in some species. 

The microstructure is characteristic of the family. 

Microstructure. -The preserved microstructure of 

the septa is consistent with an original composition of fine 

trabeculae, uniserially arranged in the axial plane of the 

septum. Septal thickening resulted from the lateral growth 

of the pinnate trabecular fibers, and the septotheca was 

formed by such lateral growth. Recrystallization has 

changed the microstructure to a secondary, but characteristic, lamellar or fibro-lamellar appearance (in the terminology ofKato, 1963). In transverse thin sections, septa appear 

to be composed of two parts: a featureless core zone of 

irregular width is clearly separated from an outer zone by 

dark lines. The core zone may terminate axially in a 

rounded end or fade into the stereome of the outer zone of 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 87 
the septa or of the axis where present. Peripherally, the core 

zone thins and passes into the wall. The outer zone is most 

commonly lamellar, with the layers pinnately arranged and 

angled toward the wall. Exceptionally, the axial dark line, 

which is characteristic of trabecular septa, is discontinuously present in the septum (with or without the core zone), 

and pinnate fibers, angled toward the axis of the coral, can 

be seen in the septum on either side of the axial plane. 

In longitudinal sections, the septa are commonly 

featureless, but there are local areas in which growth 

lamellae can be seen (these incline toward the axis, as 

would be expected) that presumably mark the shape of the 

upper edge of the septa at earlier stages. The lamellae are 

composed of fine fibers (perpendicular to the lamellae), and 

there are occasional longer structures that cross several 

lamellae and presumably are remnants of trabeculae. 

The typical microstructure is shown in many of the 

illustrations, but particularly well in pl. 2, figs. 4, 5, 8; pl. 

9, fig. 9; pl. 12, fig. 6; and pl. 18, fig. 8. Both the original 

and recrystallized structures are similar to those described 

by Hill (in Hill and Butler, 1936, p. 522-524, text-figs. 

6-10) in some Silurian corals. The structures are characteristic of the siphonophrentids, but they are not limited to that 

group. 

Discussion.-Kullmann (1965), Merriam (1972, 

1974a,b), and Birenheide (1974) independently recognized 

the basic similarity of "Heterophrentis," Breviphrentis, and 

Siphonophrentis, but all three erected subfamilies implying 

other relationships that I consider unlikely. I consider the 

genera to represent a distinct family (see Oliver, 1989, for 

additional discussion). With the revision of Heterophrentis, 

Kullmann' s Heterophrentinae becomes unusable as a name 

for the family being discussed, leaving Siphonophrentinae 

as the senior available name. Both Birenheide (1974, 1978) 

and Oliver (1989) overlooked Merriam's introduction of the 

name in 1972 and formal descriptions in 1974 (1974a,b), 

both of which have priority over Birenheide (1974). 

Included genera. -Breviphrentis Stumm, Briantelasma Oliver, Compressiphyllum Stumm, Enallophrentis 

n. gen., Metaxyphrentis n. gen., Pseudoblothrophyllum 

Oliver, and Siphonophrentis O'Connell. 

Genus BRIANTELASMA Oliver, 1960 

1960a. Briantelasma Oliver, p. 89. 

1960b. Briantelasma Oliver. Oliver, p. 6-7. 

not 1963. Briantelasma Oliver. Oliver, p. 26-27 (see Oliver, 

1989, p. C6). 

part 1973. Briantelasma Oliver. Scrutton, p. 247-248 (see 

Metaxyphrentis discussion). 

1981. Briantelasma Oliver. Hill, p. Fl54. 

part 1989. Briantelasma Oliver. Oliver, p. C5-C6 (see 

Metaxyphrentis discussion). 

Type species.-Briantelasma americanum Oliver, 

1960a, p. 89-91, pl. 14, figs. 1-14. Biohermal facies of 

Coeymans Limestone, Helderberg Group (Lochkovian), 

Madison and Oneida Counties, N.Y. 

88 Shorter Contributions to Paleontology and Stratigraphy, 1992 

Diagnosis. -Simple, trochoid to ceratoid corals with 

major septa extending to or nearly to the axis; minor septa 

one-half or less this length. Dilated septa and thickened 

tabulae entirely fill the lumen except for small, irregularly 

distributed spaces in later growth stages. 

Discussion.-My thoughts on the definition and limits of this genus have continued to evolve as I have become 

acquainted with more siphonophrentids. I here modify my 

1989 analysis by reassigning the early Eifelian B. oliveri 

Scrutton (1973, p. 248-251; Venezuela) and B. mo()rense 

Oliver (1989, p. C6-C7; Pennsylvania) to Metaxyphrentis 

n. gen. See further discussion under that genus. 

Reports of Briantelasma from areas other than eastern 

North America are erroneous or unconvincing. Briantelasma? sp. Yu and Liao, 1978, and B. breviseptata Guo, 

1989, are not determinable from the published illustrations; 

B. sibiricum Ivanovsky, 1988, is not a Briantelasma. 

Included species. -B. americanum and B. knoxboroense, both Oliver, 1960a, Lochkovian, New York; and 

B. boucoti n. sp. (see below), Lochkovian, Maine. 

Distribution. -Lochkovian, eastern North America 

(Pennsylvania, New Jersey, New York, and Maine). 

Briantelasma boucoti n. sp. 

part 1960b. Briantelasma mainense Oliver. Oliver, p. 7-8, pl. 

1, figs. 1, 2, 8, 9 (not figs. 3-7). 

Type specimens.-Holotype, USNM 137812, the 

original of Oliver, 1960b, pl. 1, fig. 2; illustrated paratypes 

(from Oliver, 1960b, pl. 1): USNM 137811 (fig. 1), 

137816 (fig. 8), 137817 (fig. 9); unillustrated paratypes, 

USNM 137818d, g, m. All specimens from USGS colin. 

3499-SD from the Beck Pond Limestone (Lochkovian), 

Maine; see Oliver, 1960b, p. 3, for locality data. 

Discussion. -Pedder (1983, p. 342) noted, and I 

agree (Oliver, 1989, p. C6), that the holotype of Briantelasma mainense Oliver, 1960b, is a ptenophyllid. Examination of the paratypes indicates that the described lot 

includes typical Briantelasma, additional ptenophyllid 

specimens, and specimens represented by transverse thin 

sections that require additional preparation for satisfactory 

identification. I am basing B. boucoti n. sp. on the B. 

mainense paratypes that can be identified as Briantelasma 

with reasonable certainty (listed above). The 1960b description (p. 7-9) is an accurate description of B. boucoti except 

that there is less open space in late developmental stages of 

B. boucoti than was originally indicated for "B. mainense. " 

Genus PSEUDOBLOTHROPHYUUM Oliver, 1960 

1960a. Pseudoblothrophyllum Oliver, p. 91-92. 

1981. Pseudoblothrophyllum Oliver. Hill, p. Fl73-Fl75. 

Type species.-Pseudoblothrophyllum helderbergium Oliver, 1960a, p. 92-93, pl. 15, figs. 1-12; pl. 16, 

figs. 1-8; and pl. 17, figs. 16, 17. Biohermal facies of 

Coeymans Limestone, Helderberg Group (Lochkovian), in 

Onondaga, Madison, and Oneida Counties, N.Y. 
Diagnosis. -Simple, large, cylindrical corals with 

major septa extending nearly to the axis; minor septa 

one-third to two-thirds as long, limited to a wide marginanum; septal dilation entirely fills marginarium, which is 

very wide in ephebic stage, narrower in neanic stage. 

Cardinal fossula limited to central tabulate zone, formed by 

short cardinal septum. Tabulae complete, axially concave, 

periaxially arched, strongly bent downward at the outer 

edge of tabularium. Exterior coarsely rugose. (Modified 

from Oliver, 1960a, p. 91.). 

Discussion. -Pseudoblothrophyllum is known only 

from the type area. P.? elegans Yu and Liao, 1978, lacks 

the thick marginarium and has very irregular tabulae; in my 

opinion, it is not Pseudoblothrophyllum. 

Included species.-P. helderbergium, P. munnsvillium, and P. arrectum, all Oliver, 1960a, Lochkovian, New 

York. 

Distribution.-Lochkovian, eastern North America; 

known from Pennsylvania, New Jersey(?), and New York. 

Genus COMPRESSIPHYLLUM Stumm, 1949 

1949. Compressiphyllum Stumm, p. 13. 

1956. Compressiphyllum Stumm. Hill, p. F270. 

1958. Heterophrentis (Compressiphyllum) Stumm. Oliver, p. 

825-826. 

1965. Compressiphyllum Stumm. Stumm, p. 22 (type data). 

1976. Compressiphyllum Stumm. Ivanovsky, p. 46. 

1981. Compressiphyllum Stumm. Hill, p. F164-F165. 

Type species. - Zaphrentis compressa Rominger, 

1876, p. 151-152, pl. 53, lower tier, upper row, second 

specimen from right; not Z. compressa Milne-Edwards, 

1860. Rominger's species was renamed Z. davisana by 

Miller (1889, p. 209). Jeffersonville Limestone, Falls of the 

Ohio; probably the lower part of the coral zone, Emsian. 

Description. -Simple trochoid corals strongly compressed parallel to the cardinal-counter plane so that the 

mature cardinal-counter diameter is two or three times the 

alar diameter. Major septa attenuate and markedly amplexoid, extend to or nearly to the axis on and just above 

tabulae, withdrawn higher; minor septa very short. Septa 

dilate at periphery to form a wall (septotheca) 2-3 mm 

thick. Calice deep with well-developed siphonofossula at 

cardinal end around a distally short cardinal septum. Transverse thin sections show a long cardinal septum bounded by 

one or more U -shaped tabular intercepts. The counter 

septum can be long or short. and can be flanked by long 

minor septa. Tabulae mostly complete, broadly arched, 

downbent peripherally. 

Discussion. -Compressiphyllum davisana is a typical siphonophrentid with a compressed growth form that is 

so extreme as to deserve recognition on the generic level. 

Only this one species is known. 

Compressiphyllum davisana (Miller), 1889 

Plate 4 

1876. Zaphrmtis compressa Rominger, 1876, p. 151-152, pl. 

53, lower tier, upper row, second specimen from right. 

Not Milne-Edwards, 1860, p. 342. 

1883. Z. compressa Rominger. Hall, p. 295-296, pl. 21, figs. 

4-5; pl. 22, fig. 5. 

1884. Z. compressa Rominger. Hall, pl. 30, figs. 1-3. 

1887. Z. compressa Rominger. Davis, pl. 134, fig. 3; pl. 138, 

fig. 4. 

1889. Z. davisana Miller, p. 209. 

1932. Z. davisana Miller. Werner, p. 114-115. 

1949. Compressiphyllum davisana (Miller). Stumm, p. 13, pl. 

5, figs. 20-21. 

1956. C. davisanum (Miller). Hill, p. F270, fig. 183:3a-b. 

1958. Heterophrentis (Compressiphyllum) davisana (Miller). 

Oliver, p. 826, pl. 106, figs. 1-9. 

1965. C. davisana (Miller). Stumm, p. 22, pl. 12, figs. 

30-33. 

1981. C. davisanum (Miller). Hill, p. F164-F165, fig. 

93:4a--c. 

Holotype.- UMMP 8620, the only specimen illustrated by Rominger (1876); this was reillustrated by Stumm 

(1965, pl. 12, figs. 30, 33) and by Hill (1981, fig. 93:4c). 

The specimen has not been sectioned, but the species is so 

distinctive that this has not been considered necessary. 

Locality and horizon.-The species is known only 

from the Falls of the Ohio River and vicinity, near Louisville, Ky. Both Davis (his own annotations in his copy of 

his 1887 monograph) and Stumm (1965, p. 22) indicated 

that the species is limited to the coral zone. In several 

collecting trips to the Falls, I have found it only in the lower 

(Emsian) part of the coral zone, the Aemulophyllum Zone of 

Oliver (1976, fig. 9, p. 20); it seems likely that all the 

known specimens are from this lower zone. 

Description.-See under genus and in Oliver, 1958 

(p. 826). The species is outside of the geographic and 

stratigraphic area of this study and is included only to make 

a more complete review of the family. The most detailed 

description is in Oliver, 1958. For this study, I have 

reviewed earlier work and some additional preparations and 

have little to add except some new illustrations, which show 

the striking differences between longitudinal sections cut 

along the short diameter (pl. 4, figs. 10-12) and those cut 

on the long diameter (pl. 4, figs. 2, 3, 7). Previously 

illustrated longitudinal sections (Oliver, 1958; Stumm, 

1965) have been short-diameter sections, which are more 

difficult to interpret because they are more likely to intersect 

the septa. For illustrations of exteriors, see Hall (1884), 

Davis (1887), and Stumm (1949, 1965); for serial transverse thin sections, see Oliver (1958). 

Material. -Seven illustrated specimens: USNM 

453996, colln. USGS 4723-SD, Aemulophyllum exiguum 

Zone (lower coral zone, Emsian); USNM 453997-454002, 

coral zone (probably lower part and Emsian). Jeffersonville 

Limestone, Falls of the Ohio, near Louisville, Ky. 

Genus METAXYPHRENTIS n. gen. 

part Heterophrentis of many authors (not Billings, 

1875). 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 89 
part? 1973. Briantelasma Oliver. Scrutton, p. 247-251. 

part 1989. Briantelasma Oliver. Oliver, p. C5-C7. 

Type species. - Zaphrentis prolifica Billings, 1858a. 

Probably Edgecliff Member of the Onondaga Limestone 

(early Eifelian), near Port Colborne, Ontario, Canada. 

Diagnosis.-Small to medium-sized, trochoid to ceratoid horn corals with a moderately deep, broadly U-shaped 

calice and a well-marked cardinal fossula; an elongate boss 

of septal origin may be present. Bilaterally arranged major 

septa extend to axial area and are commonly dilated so as to 

completely fill the lumen (with or without additional sclerenchyme), except in late stages of growth. Septal microstructure is as described for the family. 

Discussion. -Metaxyphrentis is established for those 

"Heterophrentis" in which complete septal dilation or 

sclerenchyme infilling persists into the ephebic stage but is 

followed by some attenuation if only in the sub-calice or 

calice. In this respect, the genus is morphologically intermediate between Briantelasma and Enallophrentis and may 

represent a developmental grade. Briantelasma Oliver 

(1960a) differs in being almost completely filled with 

sclerenchyme; Enallophrentis n. gen. is filled only in early 

stages. The new genera Metaxyphrentis and Enallophrentis 

are based on the two species that have most often served to 

define "Heterophrentis." As explained above, the type 

specimens on which Heterophrentis must ultimately be 

based are not related to either of the new genera. 

At this time, only three species are assigned to 

Metaxyphrentis: Z. prolifica Billings (1858a), Briantelasma 

moorense Oliver (1989), and B. oliveri Scrutton, 1973, 

although the last is assigned with some question. Most of 

the EAR "Heterophrentis" type specimens (there are over 

75 nominal species) have not been sectioned and cannot be 

generically identified at this time (but see Enallophrentis 

discussion). 

Distribution. -Early Eifelian, Eastern Americas 

Realm. 

Metaxyphrentis prolifica (Billings), 1858 

Plate 1, figures 8-15; plate 2, figures 3-10; plate 3 

part 1858a. Zaphrentis prolifica Billings, p. 176--177. 

part 1858b. Z. prolifica Billings. Billings, p. 429-430. 

part 1859. Z. prolifica Billings. Billings, p. 121-123, textfigs. 22-23. 

not? 1861. Z. prolifica Billings. Chapman, p. 510-511, textfig. 80. 

1863. Z. prolifica Billings. Billings, text-figs. 360a,b, p. 

365. 

not? 1863. Z. prolifica Billings. Chapman, text-fig. 230, p. 

442. 

not? 1864. Z. prolifica Billings. Chapman, text-figs. 80, 230, 

p. 104, 193. 

part 1874. Z. prolifica Billings. Nicholson, p. 23-24 (? pl. 3, 

figs. 2, 2a). 

part 1875. Heterophrentis prolifica (Billings). Billings, p. 

236--237. 

81 0 Shorter Contributions to Paleontology and Stratigraphy, 1992 

? 1875. Z. prolifica Billings. Nicholson, p. 237-238. 

part? 1876. Z. prolifica Billings. Rominger, p. 147, pl. 53, 

upper tier. 

part? 1886. Z. prolifica Billings. Winchell, p. 206, fig. 117 

(? p. 204, figs. 112-113). 

not? 1887. Z. prolifica Billings. Davis, pl. 135, fig. 1; pl. 

138, fig. 9. 

part? 1890. Z. prolifica Billings. Lesley, p. 1275, text-figs. p. 

1276 (? p. 1275 text-figs.). 

? 1901. Streptelasma prolificum (Billings). Lambe, p. 

115-117, pl. 8, figs. 1, 1a. 

not 1909. H. prolifica (Billings). Brown, p. 71-75, text-figs. 

20a-f. 

1909. Z. prolifica Billings. Grabau and Shimer, p. 57, 

figs. 81a,b. 

part 1914. H. prolifica (Billings). O'Connell, p. 183-184. 

not? 1927. Z. prolifica Billings. Stewart, p. 16--17, pl. 1, fig. 

1. 

not? 1938. H. prolifica (Billings). Stewart, p. 22-23, pl. 2, 

figs. 3-4. 

not? 1939. Z. prolifica Billings. Willard and others, pl. 15, 

figs. 6--7. 

not? 1942. H. prolifica (Billings). Stumm, pl. 82, fig. 4. 

part 1944. H. prolifica (Billings). Shimer and Shrock, p. 91, 

pl. 28, figs. 7-8 (?figs. 9-11). 

part? 1949. H. prolifica (Billings). Stumm, pl. 5, ? figs. 8, 11 

(not figs. 9-10). 

not? 1950. H. prolifica (Billings). Wang, pl. 6, fig. 32; pl. 9, 

fig. 73. 

part 1956. H. prolifica (Billings). Hill, p. F270 (not fig. 

183:2a-b). 

? 1956. Z. prolifica Billings. Ma, p. 41, pl. 62, figs. 20, 

21 (indet.). 

1960. Z. prolifica Billings. Bolton, p. 59 (GSC type 

cat.). 

not 1981. H. prolifica (Billings). Hill, p. F147. 

Lectotype. -GSC 3449h, the original of Billings, 

1859, text-figs. 22-23, and 1863, text-figs. 360a, b; herein 

pl. 1, figs. 9-12; pl. 2, figs. 3-5. Billings' original 

descriptions (1858a,b, 1859) were based on numerous 

specimens that he subsequently separated. In 1875 (p. 236) 

he stated, "I now propose to confine it [the name Heterophrentis prolifica] to the group typified by the specimen 

figured with the original description [ 1859] and in the 

Geology of Canada [1863], page 365." I accept this as the 

designation of a lectotype, because, in effect, Billings 

selected one of his original specimens to typify the species. 

The specimen cannot be a holotype (Bolton, 1960, p. 58) 

because it was neither unique nor designated in the original 

description. Two specimens from the syntype lot are conspecific with the lectotype and are accepted here as paralectotypes: GSC 3449i, 3449k. The other original syntypes 

either belong to different species or are undetermined. 

Type locality. -Rama's farm, Port Colborne, 

Ontario. This locality is the source of many 19th century 

collections that include material from both the Bois Blanc 

Formation (Emsian) and the Edgecliff Member of the 

Onondaga Limestone (early Eifelian). The type specimens 
55 I I I I I I I 

5 

50 - X X -

X 

X 45 - -

--.. D X X X 

,=.. X <( 40 - I- 4 - a_ 

LLI 

CJ) 

a: 

0 

'""') 

<( 35 1- - ~ 

u.. 

0 

a: 

LLI 

CD 

~ 30 1- - ::::> 2 

z 

25 r- -

20 1- -

15 ~----~~----~I ______ IL_ ____ L_I ____ ~I ____ ~IL_ ____ L_I __ ~ 

5 1 0 15 20 25 30 35 40 45 

DIAMETER (d), IN MILLIMETERS 

Figure 2. Number of major septa (n) plotted against diameter (d) for Metaxyphrentis 

prolifica (Billings). Data are from 25 sections of 14 individual corals from the 

Edgecliff Member of the Onondaga Limestone in New York and 1 section of the 

lectotype (from Port Colborne, Ontario). Serial plots from individual corals are 

marked by numbers (for example, data from six sections of one coral are marked 

4); the X's represent corals with only one section; the lectotype is marked by the 

square; symbols falling on the same point are placed immediately above and below 

the point. 

could be from either unit, but the species is otherwise 

known only from the Edgecliff, so this seems to be the more 

likely source. 

Diagnosis. -Metaxyphrentis in which a low, elliptical calicinal boss is formed by the upward extension of the 

cardinal (C) or counter (K) septum; this boss and a deep 

keyhole cardinal fossula mark the C-K plane in the calice. 

Below the calice, the lumen is commonly filled with dilated 

septa with or without additional axial sclerenchyme. 

External features. -Trochoid horn corals as much as 

4.5 and 10 em in maximum diameter and length. Most 

coralla are smoothly curved with the cardinal septum more 

or less on the convex side although the cardinal position 

varies as much as 90 degrees from the plane of curvature. 

Exteriors commonly worn but marked by shallow septal 

grooves and gentle rugae. Calice broadly U-shaped with 

bottom flat except for septal ridges, boss, and fossula; 

calice depth approximately equals diameter at distal margin. 

Cardinal fossula deep, formed by downbent tabulae and low 

cardinal septum; extends two-thirds the distance to the axis 

and is wider in its axial half, giving a keyhole shape. The 

boss is low, elongate in the C-K plane, and formed by the 

upward extension of the axial part of either the C or K 

septum; additional major septa irregularly impinge on the 

sides of the boss. 

Internal features.- Major septa extend to or nearly to 

the axis, dilated and in lateral contact with adjacent septa 

except in late stages; they number 40 to 51 in transverse 

sections 17 to 40 mm in diameter (text-fig. 2). Septa are 

bilaterally arranged although, in sections below the bottom 

of the fossula, it is difficult to differentiate between C and 

K on the basis of septal arrangement alone. In transverse 

sections through the fossula, C is of medium length; 

adjacent major septa are short, deflected away from, then 

curving toward C; successive septa are similarly curved but 

longer, and one pair may meet in a U-s'hape at the axial end 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 811 
of the fossula. More distant major septa tend to be straighter 

but to maintain the bilateral arrangement by making a 

smaller angle with the C-K plane than with a plane 

perpendicular to it. The cut edges of downbent tabulae can 

also form U-shaped lines around the inner end of the 

fossula. 

Most of the major septa are more or less straight for 

two-thirds or more of the distance to the axis; commonly 

several extend to the axis but may be bent and irregular; 

others terminate at the two-thirds point or fade into axial 

sclerenchyme (for example, pl. 3, figs, 2, 3, 8). The axial 

area may be filled by the irregular septa or by a combination 

of septa and other sclerenchyme that seems to originate 

through thickening of tabulae. In some sections, a very 

thick septal segment is present in the position of the 

calicinal boss (pl. 3, fig. 12), but this feature is not 

vertically continuous, and there is no columella. 

Minor septa are consistently present and extend from 

the margin approximately 0.3 of the radius. 

Tabulae tend to be complete, broad and flat, and 

strongly downbent peripherally with upturned margins; 

commonly thickened by sclerenchyme. In many longitudinal sections, thickened septa and tabulae fill the lumen to or 

nearly to the bottom of the calice; in these, the tabulae may 

not be distinguishable. In the upper part of the corallum, the 

major septa may be thinner and withdrawn from the axis, 

and the tabulae may be separated by open spaces, giving a 

more typical "Heterophrentis" appearance. 

Individual variation is most marked in the nature and 

amount of septal and tabular thickening (or sclerenchyme 

infilling). Small open spaces can occur at any place in a 

corallum, and the regular development of interseptal space 

can begin at some distance below the calice, for example in 

USNM 454009 (pl. 3, fig. 1); however, in this individual, 

the overall impression is of gradual decrease of thickening 

with growth. Variation in length of major septa is hard to 

analyze because of the difficulty in seeing the nature of the 

septal ends in the axial sterome. 

Microstructure is as described for the family. 

Discussion.-The Billings (1875) lectotype and paralectotype specimens are from Port Colborne, Ontario, 

some 30 km west of Buffalo, N.Y., and may be from either 

the Bois Blanc Formation or the Edgecliff Member of the 

Onondaga Limestone. The above description is based on the 

Billings types plus a sample of 14 specimens from the 

Edgecliff Member in New York that were collected by the 

author. No similar specimens have been recognized in 

collections from the Bois Blanc Formation. The Billings 

specimens are very similar to the Edgecliff sample and are 

thought to be conspecific; it seems most likely that they also 

came from the Edgecliff. 

The Billings specimens are silicified and free of 

matrix. The lectotype and one of the paralectotypes have 

been sectioned transversely; the other paralectotype is 

broken and shows similar transverse structure. None of the 

812 Shorter Contributions to Paleontology and Stratigraphy, 1992 

three has been cut longitudinally, but all show the lower 

part of the calice. In contrast, the New York specimens are 

known primarily from thin sections, and calicinal details, 

including the boss and fossula, are from sections taken just 

above and below the calice floor. 

Occurrence. -Known only from the Edgecliff Member of the Onondaga Limestone in New York and from Port 

Colborne, Ontario (Billings' types, probably Edgecliff). 

Material. -Lectotype, GSC 3449h; paralectotypes, 

GSC 3449i and 3449k; all from Port Colborne, Ontario. 

Edgecliff sample (text-fig. 2), 14 specimens (locality no. in 

parentheses); illustrated: USNM 454003 (108), 454004 

(286), 454005 (338), 454006 (137), 454007 (124), 

454008-454009 (154); unillustrated (measured for text-fig. 

2): 454010 (154), 454011-454013 (164), 454014 (338), 

454015-454016 (206). Edgecliff Member of the Onondaga 

Limestone, at indicated localities in New York. 

Genus ENALLOPHRENTIS n. gen. 

part Heterophrentis of many authors (not Billings, 

1875). 

Type species.-Strombodes simplex Hall, 1843. 

Moscow Formation, Hamilton Group; Leicester, N.Y. 

Diagnosis. -Small to medium-sized, trochoid to ceratoid horn corals with a moderately deep, broadly U-shaped 

calice and a well-defined cardinal fossula. Major septa 

extend to the axial area and are bilaterally arranged. Septa 

and tabulae are partly or completely dilated in early stages. 

Microstructure is as described for the family. 

Discussion.-Enallophrentis is established for those 

"Heterophrentis" in which complete infilling of the lumen 

is limited to the early stages. In both Briantelasma and 

Metaxyphrentis, sclerenchyme fills the lumen to a much 

later stage. The genus is known from the Oriskany Sandstone (Pragian) and from parts of the Hamilton Group 

(Givetian). Many of the nominal species of "Heterophrentis" from Middle Devonian rocks in eastern North 

America seem to belong to Enallophrentis, but most have 

not been sectioned, and studies of individual variation are 

needed in almost every case before species-level taxonomy 

can be considered. 

Distribution.-Pragian to Givetian, eastern North 

America. 

Enallophrentis simplex (Hall), 1843 

Plates 5-9; plate 10, figures 1-7 

1843. 

1847. 

? 1852. 

part 1877. 

1877. 

1890. 

Strombodes simplex Hall, p. 209-210, fig. 87-6, 

pl. 48, fig. 6. 

S. simplex. Owen, text-fig. 6, p. 69 (after Hall, 

1843). 

S. simplex. Giebel, p. 34. 

Zaphrentis ampla Hall, pl. 21, tigs. 2-3 (? fig. 1, 

not fig. 4). Not Z. ampla Ludwig, 1865, p. 165, 

pl. 38, figs. 1a,b. 

Z. simplex Hall. Hall, pl. 21, figs. 5-11. 

Z. simplex Hall. Lesley, p. 1276, text-figs. (after 

Hall). 
1899a. Z. simplex Hall. Grabau, p. 123, fig. 4. 

1899b. Z. simplex Hall. Grabau, p. 288. 

1906. Z. simplex Hall. Grabau and Shimer, p. 194. 

1909. Z. simplex Hall. Grabau and Shimer, p. 57, fig. 

82. 

? 1912. 

? 1927. 

not? 1938. 

not? 1939. 

Z. cf. simplex Hall. Kindle, p. 66, pl. 2, fig. 

(unrecognizable). 

Z. simplex Hall. Stewart, p. 17-18, pl. 1, fig. 2. 

Heterophrentis simplex (Hall). Stewart, p. 23, pl. 

2, figs. 5-7. 

Z. simplex Hall. Willard and others, pl. 15, figs. 

8-9. 

not? 1942. H. simplex (Hall). Stumm, pl. 82, figs. 5-6. 

1944. H. simplex (Hall). Shimer and Shrock, p. 91, pl. 

28, fig. 12. 

not? 1956. H. simplex (Hall). Ma, p. 67 (from Stewart, 1938). 

part? 1965. H. simplex (Hall). Stumm, p. 21, pl. 11, fig. 19; 

pl. 14, figs. 5-6. 

part 1973. H. (Heterophrentis) simplex (Hall). Scrutton, p. 

254-255, pl. 5, fig. 3 only. 

Holotype. -NYSM 360, the original and only illustrated specimen of Hall, 1843, text-fig. 87--6, p. 209; herein 

pl. 5, figs. 1-4; pl. 6, figs. 1-2. The specimen was 

reillustrated by Hall, 1877 (pl. 21, fig. 8), who stated, 

"This is the original specimen of the species." (ex pl. pl. 

21). 

Type locality.-"Moscow" (now Leicester), Leicester quadrangle, Livingston County, N.Y. This is most likely 

the outcrop of the Moscow Formation on the east side of 

Little Beards Creek, 1.3 km north-northeast of the village 

center. The specimen is probably from the Fall Brook coral 

bed (Baird and Brett, 1983), Windom Shale Member, 

Moscow Formation, Hamilton Group (Givetian). 

Diagnosis. -Small to medium-sized, trochoid to ceratoid hom corals having a moderately deep, broadly 

U-shaped calice and a well-marked cardinal fossula. Septa 

bilaterally arranged, more radial in the counter quadrants. 

Major septa vary in length, commonly extend from 0.7 to 

1.0 the distance to the axis without forming an axial 

structure; minor septa are 0.3 as long. Tabulae complete or 

incomplete, broadly M-shaped; no dissepiments. Septa and 

tabulae attenuate in mature stage (diameter > 15 mm), but 

may be partly or completely dilated in early stages. Microstructure as described for the family. 

Discussion.-Hall (1843, p. 210) gave only "Moscow" as the locality of the single specimen illustrated. This 

is now Leicester, Livingston County, in the Genesee Valley 

of western New York. Hall (1877, ex pl. pl. 21) cited 

"Moscow" for two of the six illustrated specimens; a third 

(the holotype) is also from "Moscow" (Hall, 1843). Hall 

(1877) gave no locality for the other three, but these were 

cited as "Genesee Valley, N.Y." by Whitfield ( 1900, p. 

194-195, catalog of the AMNH) and may be from the same 

locality. A suite of 21 specimens from the Little Beards 

Creek locality, which are probably all from the Fall Brook 

coral bed, is described below as the topotype population. 

The holotype and the other 1877 specimens are conspecific 

with this suite and may all be from the Fall Brook coral bed 

on Little Beards Creek, but there is no apparent way to 

demonstrate this. In any case, the 1843 and 1877 specimens 

plus the topotype population are the central and principal 

basis for my concept of the species. 

Zaphrentis ampla Hall, 1877, is a junior synonym of 

E. simplex. The lectotype, here selected, is NYSM 352, the 

original of Hall, 1877, pl. 21, figs. 2-3 (herein pl. 5, figs. 

7-9; pl. 6, fig. 3). The lectotype is from York, N.Y., and 

is most likely from the Centerfield Member of the Ludlowville Formation (see Centerfield population below). Hall 

(1877) illustrated three specimens as Z. ampla; the only 

apparent difference between Z. ampla and E. simplex is that 

the former has a somewhat wider apical angle, but there is 

considerable variation in this character, and the ampla 

lectotype is easily accommodated in E. simplex. Hall's 

other two specimens are questioned. NYSM 351 (the 

original of Hall's pl. 21, fig. 1, Darien, N.Y.) appears to 

have a streptelasmatoid axial structure in a thin section 

taken just below the calice (pl. 6, fig. 4) but is like E. 

simplex in lower sections and in the calice; probably it is a 

minor variant of E. simplex. AMNH 3050 (the original of 

Hall's fig. 4) is tentatively assigned to Metaxyphrentis sp. 

cf. M. prolifica. It is apparently from the Onondaga 

Limestone at Clarksville, N.Y. (AMNH catalog, Whitfield, 

1900, p. 194-195; and preservation characteristics), but it is 

not well enough preserved for certain identification. 

Topotype population (plate 5, figures 1--6, 10-19; 

plates 6-8).-Twenty-one specimens from the Little Beards 

Creek locality and from, or probably from, the Fall Brook 

coral bed are considered as topotypes. 

External features: Twenty-one topotype specimens 

are trochoid to nearly ceratoid. Preserved length ranges 

from 20 to 100 mm, and diameters at the calice margin 

(incomplete) are as large as 38 mm. Coralla are smoothly or 

somewhat irregularly curved. Exteriors are rugose; septal 

ridges and interseptal grooves appear only on worn specimens. Calice broadly U-shaped; if restored, depth would be 

approximately half the diameter; septal ridges on calice 

floor extend to, or nearly to, the axis; there is no axial boss. 

Cardinal fossula, formed by downbent tabulae and short 

cardinal septum, is prominent on calice floor but hardly 

recognizable on wall; position of fossula relative to plane of 

curvature is irregular but it is commonly not in plane of 

curvature. 

Internal features: Major septa are attenuate in later 

stages, extend to, or nearly to the axis, but are commonly 

somewhat withdrawn and amplexoid; they number 33 to 53 

in transverse sections 20 to 38 mm in diameter (text-fig. 

3A); minor septa approximately 0.3 as long. In early stages 

(diameter less than 20 mm), septa extend to the axis and are 

somewhat dilated; at diameters less than 15 mm, dilated 

septa fill, or nearly fill, the lumen; major septa number 23 

to 40 in sections 9 to 17 mm in diameter (text-fig. 3A). 

Septal arrangement bilateral; the cardinal septum is comThe Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 813 
55 

A 14 

50 

10 

11 X 

45 X 

D 12 

-.. X 

~ 

<( 40 14 X 3 8 f- 1 a... 13 w 

(/) 

a: 

0 10 11 --, 

<( 8 35 X ~ 9 12 LL 

0 13 

a: 

w X Ill 

~ 30 2 5 4 6 

X ::J 

z 

25 

20 

10 15 20 25 30 35 40 

DIAMETER (d), IN MILLIMETERS 

Figure 3. Number of major septa (n) plotted against diameter (d) for 

Enallophrentis simplex (Hall). A, Fall Brook coral bed population based 

on 39 sections of 20 individuals plus 4 specimens (5 sections) illustrated 

by Hall from Moscow or the Genesee Valley (see "Discussion" in text). 8, 

Centerfield population based on 53 sections of 30 individuals plus 2 Hall 

specimens (1 section each), which may be from the Centerfield Member 

of the Ludlowville Formation (see text). See figure 2 for meaning of 

symbols; in figure 3A, 1-13 represent specimens with 2 or 3 sections, and 

14 is the holotype; in 38, 1-20 represent specimens with 2 or 3 sections; 

in both graphs, individuals with one section are represented by an X, and 

the Hall specimens other than the holotype, by a square. The boundary 

lines are identical in the two graphs and are included to facilitate 

comparison of the populations. 

manly shorter than other major septa, but length varies with 

growth stage and position of section; adjacent major septa 

bend toward the cardinal septum at their axial ends. The 

counter septum is commonly long and marked by longer 

counter-lateral minors (for example, pl. 7, fig. 12). There is 

a tendency for the other major septa to be pinnately 

arranged in the cardinal quadrants, radial in the counter 

quadrants; other minor septa tend to lean in the counter 

direction but are not contratingent. The sides of the septa 

are smooth except in some large specimens (28 to 38 mm in 

diameter) where "bumps" developed in the latest (gerontic?) 

stage (pl. 7, figs. 1-2). 

Longitudinal sections in mature parts of coralla show 

tabulae to be commonly thin and closely spaced, complete 

814 Shorter Contributions to Paleontology and Stratigraphy, 1992 

or incomplete, and strongly arched with a flat or depressed 

center; peripheral parts flat or slightly upturned; axial ends 

of septa form ridges of varying heights on the tabulae. 

Earlier stages show thicker tabulae and thicker and more 

complete septa. 

Microstructure in both transverse and longitudinal 

sections is as described for the family. 

Variation in the topotype population is principally in 

length of septa and degree of dilation at given diameters. 

The brephic stage is not known because septa cannot be 

clearly distinguished in sections smaller than about 10 mm, 

and juveniles have not been found or recognized. Between 

diameters of 10 and 20 mm, most of the attenuation takes 

place, but it can begin at smaller diameters and be essen-
10 15 20 25 30 35 40 45 50 55 

DIAMETER (d), IN MILLIMETERS 

Figure 3.-Continued. 

tially complete at 14 mm or less. If the neanic stage is 

defined by heavy dilation and the ephebic stage by attenuation, then the neanic stage ended, and the ephebic began, 

at diameters of 15 to 20 mm. If the "bumps" on the sides of 

septa mark a gerontic stage, it began at diameters of some 

30 mm (text-fig. 4). 

Centerfield population (plate 5, figures 7-9; plate 9; 

plate 10, figures 1-7). -Enallophrentis simplex is most 

common in the Centerfield Member of the Ludlowville 

Formation. A sample of 30 specimens from collections 

made at several localities in the Genesee Valley and in areas 

just east and west of there was analyzed to broaden the 

species concept. As noted above, the lectotype of Hall's 

Zaphrentis amp/a is an E. simplex from York, in the 

Genesee Valley, and probably is from the Centerfield 

Member. The paralectotype of Z. amp/a, NYSM 351, is 

from Darien, N.Y., and possibly also from the Centerfield, 

but the Darien section includes most Ludlowville Formation 

members and the lower part of the Moscow Formation, so 

the provenance is uncertain. 

In general, Centerfield specimens are larger (text-fig. 

3B) and dilation persists to a larger diameter (text-fig. 4) 

than in the topotype population, but the fundamental characters are the same. Septal counts and diameters measured 

from thin sections are compared in the following table and 

in text-figures 3 and 4 (n=no. of major septa; d=diameter 

in millimeters; neanic, ephebic, and gerontic are defined for 

this purpose in the description of the topotype population): 

nat d=8-19 mm ................... . 

nat d=20-38 mm .................. . 

nat d=52 mm ..................... . 

neanic d, observed range ........... . 

ephebic d, observed range ........... . 

gerontic d, observed range .......... . 

Topotype 

23-40 

33-50 

none 

9-20 

14-28 

28-39 

Centerfield 

22-40 

32-52 

60 

5-23 

13-28 

37-52 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 815 
w 

~ 

~ 3 (/) 

z 4 0 

~ 5 _J 

0 

10 

-------- Gerontic 

} Ephebk 

20 30 40 50 60 70 

DIAMETER {d), IN MILLIMETERS 

Figure 4. Dilation stage plotted against diameter (d) for 

Enallophrentis simplex (Hall). An arbitrary scale, from 

attenuate (1) to fully dilated (7), was established with 

which to compare each section. The range of diameters 

for each dilation stage is plotted for both population 

samples (Fall Brook above, Centerfield below); stage 2 

was not observed in the Centerfield population. Ontogenetic stages, defined by persistence of dilation, are 

labeled in accordance with my text definitions. The dilation scale standards are illustrated on plates 6 and 7 as 

follows: stage 1, pl. 7, fig. 1; stage 2, pl. 6, fig. 8; stage 3, 

pl. 7, fig. 12; stage 4, pl. 6, fig. 10; stage 5, pl. 7, fig. 7; 

stage 6, pl. 6, fig. 12; stage 7, pl. 7, fig. 8. 

These data show that there is no significant difference in 

number of septa at any diameter but that the largest single 

specimen is from the Centerfield Member. Growth stages 

based on dilation and attenuation show more overlap in size 

range in the Centerfield population. 

Material. -(1) From, or probably from, the Fall 

Brook coral bed, Windom Member, Moscow Formation, 

Leicester, N.Y. (see discussion and locality data): Bolotype, NYSM 360, the original of Hall, 1843, text-fig. 87-6, 

p. 209 (also Hall, 1877, pl. 21, fig. 8). Illustrated specimens of Hall, 1877, pl. 21 (Hall fig. in parentheses): 

NYSM 6266 (5), 6267 (11), AMNH 39407 (6, 7), 39408 

(9), 39409 (10). Topotype sample (text-fig. 3A), illustrated: 

USNM 454017-454019, coHn. USGS 5700-SD; 454020-

454023, 5701-SD; 454024-454030, 8770-SD; 454031, 

11476-SD; unillustrated: USNM 454032, USGS 5700-SD; 

454033-454034, 5701-SD; 454035, 8770-SD; 454036-

454037, 11190-SD. 

(2) From the Centerfield Member, Ludlowville Formation, western New York (see discussion and locality 

data): NYSM 352, lectotype, here selected, of Zaphrentis 

amp/a Hall, 1877, pl. 21, figs. 2, 3 (York). NYSM 351, 

paralectotype of Z. amp/a Hall, 1877, pl. 21, fig. 1 

(Darien), may or may not be from the Centerfield but is 

listed here. Centerfield population sample (text-fig. 3B), 

illustrated: USNM 454038-454041, colin. USGS 

5686-SD; 454042-454043, 5692-SD; 454044, 4678-SD; 

unillustrated: 454045-454051, USGS 4678-SD; 454052, 

5324a-SD; 454053-454060, 5686-SD; 454061-454062, 

816 Shorter Contributions to Paleontology and Stratigraphy, 1992 

5692-SD; 454063, 5705-SD; 454064, 5714-SD; 454065, 

5716-SD; 454066, 11197-SD; 454067, 11199-SD. 

Enallophrentis broweri n. sp. 

Plate 10, figures 8-10; plates 11, 12 

Type population.-The following description is 

based primarily on a sample of 34 specimens from the 

Delphi coral bed (Oliver, 1951, p. 719-720), Delphi 

Station Member, Skaneateles Formation, Hamilton Group, 

at Pompey Center, N.Y. The Pompey Center population 

includes specimens of a wide size range that are interpreted 

as representing neanic, ephebic, and gerontic individuals 

although there are no sharp morphologic boundaries 

between stages, and no age or growth classes have been 

specifically identified. 

Diagnosis. -Small to medium-sized, trochoid horn 

corals with a moderately deep, broadly U-shaped calice and 

a weak to well-marked cardinal fossula. Septa weakly 

bilateral in early stages; bilaterality becomes more marked 

in later growth stages, especially in cardinal quadrants. 

Major septa thick, extend 0. 8 to 1.0 the distance to the axis; 

minor septa extend 0.2 the distance. A counterclockwise 

deflection of the septa is very marked in early stages, less so 

in later stages, but there is no axial structure. Tabulae 

complete or incomplete, broadly arched, commonly with 

flat axial area. Septa and tabulae very thick in early stages, 

gradually thinning later so that mature part of large individuals can be quite open. Microstructure as described for the 

family. 

External features.- Mostly trochoid, rarely ceratoid 

or irregular. Size range in studied population: diameter at 

calice margin, 25 to 50 mm; length along convex side, 40 to 

100 mm. Most coralla are smoothly curved, rarely somewhat irregular; the cardinal septum tends to be on the 

convex side, but the C-K plane and the plane of curvature 

do not coincide. Exteriors are gently rugose; there are 

clearly incised septal grooves and flat-topped interseptal 

ridges where the surface is well preserved. Calice broadly 

U-shaped; depth approximately 0.7 the diameter. Septal 

ridges on calice floor extend to, or nearly to, axis. A 

siphonofossula, formed by a distally short cardinal septum 

and downbent tabulae, is on calice floor and wall. 

Internal features. -Ontogenetic changes in this species are so marked that it is difficult to generalize. Instead, 

I use the late ontogenetic characters of individual corals to 

arbitrarily define stages. Ephebic individuals have long, 

moderately thickened major septa, with considerable 

interseptal and intertabular space and a clearly defined 

cardinal fossula. The smaller, more youthful, neanic individuals have thicker septa and tabulae and little open space 

within the skeleton; the fossula is weak. Postephebic 

individuals have attenuate, withdrawn septa, a siphonofossula, and thin tabulae in their latest stage; these are termed 

gerontic although there is no evidence that they were 

senescent. There are gradations between the stages, espe-
60 I I I I I I 

Gerontic24 55 r- -

23 25 

21 

50 - X 12 -

21 14 11 22 

13 23 

19 

20 10 17 

45 - 12 25 9 16 - --E. 15 

X 7 22 24 

14 

<( 

fa... 

L.U 

(/) 

a:: 40 11 -

0 X 3 X 

6 

15 5 X 8 

l~ ~ 17 20 - 4 -Ephebic -

X 

16 1 9 13 15 

7 4 

"'"") 

<( 

~ 

LL 

0 35 a:: 

L.U 

co 

~ 

::J 

z 

30 - 8 3 2 -

18 ~ 

X 

5 

25 - 6 -

1 

-Neanic 

20 - -

15 I I I I I I I 

5 10 15 20 25 30 35 40 45 

DIAMETER (d), IN MILLIMETERS 

Figure 5. Number of major septa (n) plotted against diameter (d) for Enallophrentis 

broweri n. sp. Data are from 60 sections in a sample of 34 individuals from the 

Delphi coral bed, Pompey Center, N.Y. See figure 2 for meaning of symbols. Boxes 

mark the observed ranges of septal number and diameter for the ontogenetic 

stages as discussed in the text. 

cially within individual corals, but, in the population, the 

differences are striking enough that the possibility of there 

being three or four species was considered. 

The differences between apparent ontogeny as seen in 

serial transverse sections and the true ontogeny experienced 

by the growing individual are particularly significant in this 

species (see discussion in section on coral morphology). 

The most obvious ontogenetic changes in E. broweri 

involve thickening of septa and tabulae so that dilation tends 

to persist to a larger diameter in corals that lived longer. 

In all growth stages, the outer wall is a septotheca 

formed by dilation of both major and minor septa; the 

thickness ranges from one to several millimeters both 

ontogenetically and among individuals of the same size and 

growth stage. Minor septa are short, consistently about 0. 2 

the radius. The earliest (brephic) stage is not known. 

The stages are described in growth order. Ephebic 

individuals are the most common (50 percent of sample), 

and this is the stage that I would expect to be found most 

frequently at other localities. Text-figure 5 shows the range 

of number of septa at given diameters in the studied 

population. 

Neanic stage: Ten individuals with calice-rim diameter and length up to approximately 28 and 40 mm, 

respectively, represent the neanic stage. In transverse sections just below the calice, major septa extend to or nearly 

to the axis and number 28 to 40 at diameters of 10 to 18 mm 

(text-fig. 5). The septa are dilated so that they fill or nearly 

fill the lumen and are strongly deflected to the right, 

forming a counterclockwise whorl. The approximate position of the cardinal septum and the C-K plane may be 

indicated by a short septum and (or) by long minors 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 817 
flanking the counter septum, but the fossula is not marked 

although it can be seen on the calice wall and on the calice 

floor of some individuals. Minor septa short, approximately 

0. 2 the radius. Tabulae are obscured by the dilated septa 

except near the axis, where parts of thickened tabulae can 

be seen in some specimens. 

Ephebic stage: Seventeen individuals with calice-rim 

diameter and length up to approximately 40 and 80 mm, 

respectively, compose the ephebic group. Major septa 

extend at least 0.8 the distance to the axis and are moderately dilated; in sections just below the calice (d= 15-30 

mm), they number 35 to 50 (text-fig. 5). Minor septa are 

short, commonly 0.2 the radius. The C-K plane is marked 

by a short cardinal septum in a narrow fossula, and by a 

counter septum flanked by relatively long minors. The 

cardinal fossula is bounded on each side by one or two 

major septa that curve around it; downbending of the 

tabulae is indicated in some sections. Except near the C 

position, the septa are more or less radial, although the alar 

positions are commonly marked by deviations. Tabulae are 

thick and widely spaced; they are commonly flat axially and 

downbent peripherally although this shape tends to be 

obscured by the thickened septa. Sections through the early 

(neanic) stage of ephebic individuals are characterized by 

thick septa that nearly fill the lumen; the major septa are 

strongly deflected to the right as described above. 

Gerontic stage: Seven large individuals with calicerim diameter and length up to 50 and 100 mm, respectively, 

are considered gerontic. Major septa are attenuate and 

withdrawn, and the upper tabulae are complete, thin, and 

downbent peripherally with a broad flat axial area. In 

sections 25 to 45 mm in diameter, taken at or just above the 

calice floor, there are 49 to 56 major septa (text-fig. 5); in 

these sections, the cardinal fossula is formed by down bent 

tabulae in addition to the incurving of the adjacent major 

septa. Sections through the earlier stages of gerontic individuals exhibit neanic and ephebic characters as would be 

expected, except that the earlier stages tended to persist 

longer, or to larger diameters. 

Discussion. -E. broweri is characterized by its thick 

septa and tabulae and by the neanic whorl. It differs from E. 

simplex in these characters plus the axially flat tabulae and 

weak fossula. 

Occurrence. -Known only from the Delphi coral 

bed(s) in the upper part of the Delphi Station Member, 

Skaneateles Formation (early Givetian), Hamilton Group, 

New York. See discussion in locality data. 

Material.-The description is based on a sample of 

34 specimens from the Delphi coral bed at Pompey Center, 

N.Y. (USGS 9745-SD): Holotype, USNM 454068; illustrated paratypes, USNM 454069-454078; unillustrated 

(measured, text-fig. 5) paratypes, USNM 454079-454101. 

Additional illustrated paratype, USNM 454102, coHn. 

USGS 9132-SD, Delphi coral bed, De Ruyter quadrangle, 

New York. 

818 Shorter Contributions to Paleontology and Stratigraphy, 1992 

Genus BREVIPHRENTIS Stumm, 1949 

1949. Breviphrentis Stumm, p. 13. 

1956. Breviphrentis Stumm. Hill, p. F271. 

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

not 1961. Breviphrentis Stumm. Fontaine, p. 90. 

part 1965. Siphonophrentis O'Connell. Stumm, p. 23-25. 

1974b. Siphonophrentis (Breviphrentis) Stumm. Merriam, 

p. 42. 

? 1978. Breviphrentis Stumm. Birenheide, p. 69. 

1981. Breviphrentis Stumm. Hill, p. F166. 

Type species. -Amplexus invaginatus Stumm, 1937, 

p. 427, pl. 53, fig. 2; pl. 54, figs. 2a-c; Nevada Formation, 

Eurekaspirifer pinyonensis Zone (Emsian), Nevada. 

Diagnosis. - Ceratoid to cylindrical siphonophrentids 

in which septa are withdrawn from the axis from an early 

stage so that late neanic and ephebic septa extend half to 

two-thirds the distance to the axis and are markedly amplexoid. Tabulae axially flat to strongly depressed and peripherally deflected downward; some have a marginal trough 

and tabellae. Siphonofossula variably developed. No dissepiments or axial structure. 

Discussion. -Stumm (1949) established Breviphrentis more or less as an intermediate between "Heterophrentis" and Siphonophrentis; he considered the fossula of 

Breviphrentis to be formed by a short cardinal septum 

without downbending of the tabulae and noted this feature 

as a principal difference from Siphonophrentis. Oliver 

(1960a) thought that both forms had downbent tabulae and 

that the only difference was in degree. Stumm (in discussions) accepted this, and both Oliver (1960a) and Stumm 

(1965) synonymized the two genera. Merriam (1974b) 

considered them different enough to recognize as subgenera. Birenheide (1978) noted the question but used both 

names. Hill (1981) rightly stated that both type species 

needed to be redescribed before the generic or family names 

could be properly used. 

Since the early 1970's, I have used both Breviphrentis and Siphonophrentis in biogeographic works, initially for pragmatic reasons-they were useful for separating morphologic end members. They continue to be useful, 

and in this restudy of the family I have become satisfied that 

they are reasonably distinct and that the EAR species can be 

confidently assigned. They are as likely to be valid genera 

as can be expected in families with so few morphologic 

characters. However, the identifications used biogeographically are not in all cases the same as those given here, and 

some revisions of the generic distributions given in earlier 

papers are needed. 

Breviphyllum Stumm, 1949, type species, Amplexus 

lonensis Stumm, 1937 (p. 428, pl. 53, fig. 4; pl. 54, figs. 

4a-b), differs from Breviphrentis in having well-developed, 

vertically elongate dissepiments in the mature stage (Hill, 

1981, fig. 95:3a-b). The type species was inadequately 

illustrated initially and, as a result, misinterpreted by Oliver 

(in correspondence) and others (for example, Merriam, 

1974b, p. 41) who have synonymized it with Breviphrentis. 
Included species. - Zaphrentis roemeri MilneEd wards and Haime, 1851 (Lochkovian, New York); 

Zaphrentis halli Milne-Edwards and Haime, 1851 (Givetian, New York); Amplexus invaginatus Stumm, 1937 (Emsian, Nevada); Siphonophrentis variabilis Oliver, 1960a 

(Lochkovian, New York); Siphonophrentis sp. aff. S. halli 

(Milne-Edwards and Haime) Oliver, 1987 (Givetian, Illinois); Breviphrentis cista n. sp. (Givetian, New York); and 

B. pumilla n. sp. (Givetian, New York). 

Distribution.-Lochkovian to Givetian, Eastern 

Americas Realm, including all four provinces. 

Breviphrentis invaginata (Stumm), 1937 

Plate 13 

1937. Amplexus invaginatus Stumm, p. 427, pl. 53, fig. 

2; pl. 54, figs. 2a-c. 

1937. Heterophrentis nevadensis Stumm, p. 426-427, pl. 

53, fig. 1; pl. 54, figs. la-c. 

not? 1937. Amplexus nevadensis Stumm, p. 427-428, pl. 53, 

fig. 3 (not pl. 54, figs. 3a-c). 

not 1940. Amplexus invaginatus Stumm. Merriam, pl. 16, 

figs. 3-4. 

1949. Breviphrentis invaginatus (Stumm). Stumm, p. 

13-14, pl. 5, figs. 22-24. 

1956. Breviphrentis invaginata (Stumm). Hill, p. F271, 

fig. 183:6a-b. 

not? 1956. Amplexus invaginatus Stumm. Warren and Stelck, 

pl. 4, fig. 4. 

part 1974b. Siphonophrentis (Breviphrentis) invaginatus 

(Stumm). Merriam, p. 42-43, pl. 15, figs. 3-4, 9, 

11 (not figs. 1-2, 5-8, 10); pl. 16, figs. 3-8, 

15-16 (not figs. 1-2, 9-13); not pl. 17, figs. 

12-13. 

1981. Breviphrentis invaginata (Stumm). Hill, p. Fl66, 

fig. 95: la-b. 

Type specimens.-Lectotype, here selected, USNM 

94443, the original of Stumm (1937) pl. 53, fig. 2, herein 

pl. 13, figs. 1, 2. Paralectotype, USNM 454103 (formerly 

also 94443), original of Stumm (1937), pl. 54, fig. 2a, 

herein pl. 13, fig. 3. Stumm (1937, plate explanations) 

labeled both an exterior (pl. 53, fig. 2) and a transverse thin 

section (pl. 54, fig. 2a) as holotype, but they are not the 

same specimen. I here select the specimen, rather than the 

single thin section, as lectotype because sections cut from 

the specimen provide a suitable basis for the species; such 

sections were prepared earlier and have already been 

illustrated in the Treatise (Hill, 1981 , fig. 9 5: 1 a-b). 

Stumm's paratype, USNM 94443a (renumbered 454104, 

thin section), is the original of Stumm (1937) pl. 54, fig. 

2b; the thin section original of fig. 2c may be from the same 

specimen but is apparently lost; the specimen fragment 

labeled USNM 94443a (unillustrated Stumm paratype) is 

not the one from which either thin section was cut and is 

probably not B. invaginata. 

Heterophrentis nevadensis Stumm, 1937, is a subjective synonym of B. invaginata. The original description of 

H. nevadensis immediately preceded that of Amplexus 

invaginatus in Stumm, 1937. Under the rules of the 

International Commission on Zoological Nomenclature 

(1985 ed.), the first reviser determines which name to use in 

this situation. Recommendation 24A states that the reviser 

should select the name "that will best serve stability" and 

that appearing first in the same publication is a secondary 

criterion. In this case, stability is served by maintaining B. 

invaginata as the name of the type species and declaring H. 

nevadensis a junior subjective synonym. Merriam (1974b) 

placed H. nevadensis in synonymy with B. invaginatus with 

a query but did not discuss the point except to comment that 

some B. invaginatus resemble Stumm's H. nevadensis. 

Diagnosis. -Small, ceratoid to cylindrical Breviphrentis with siphonofossula and relatively long major and 

minor septa. Tabulae closely spaced, axially flat to irregular, strongly down bent at periphery. 

Description. -Ceratoid to cylindrical, tending to 

cylindrical when fully developed; mature diameters 18 to 22 

mm; maximum measured length is 5.5 em (in the holotype, 

broken at both ends). Calice not known but longitudinal 

sections indicate that it is probably broadly shallow with a 

flat bottom, with or without septal traces on the upper 

surface of the tabula forming its floor. 

In sections 18 to 22 mm in diameter, major septa 

number 37 to 46 (text-fig. 6) and extend 0.5 to 0.8 the 

radius; a few septa may be longer and extend nearly to the 

axis (possibly only on the upper surfaces of tabulae). Minor 

septa relatively long, commonly 0.4 the radius. Septa 

attenuate to moderately thickened, but dilate at periphery to 

form a septotheca. Cardinal septum short; adjacent major 

septa may bend toward it or be straight; counter septum 

commonly flanked by long minors. Other major septa are 

radially arranged but may tum at their inner ends; those that 

extend into the axial area tend to be quite irregular. 

In sections through early growth stages (one specimen, sectioned at 7.5 and 14 mm), major septa are 

relatively longer and thicker, but there is no indication of 

complete infilling of the lumen. 

In longitudinal sections, tabulae are thin and closely 

spaced. They tend to be complete, axially flat, and strongly 

downbent peripherally. Axial ends of the longer amplexoid 

septa commonly project a short distance upward from the 

tabular surfaces. 

Microstructure is as described for the family. 

Discussion. -Merriam's description of B. invaginatus (1974b, p. 42-43) was based on 16 specimens in 

addition to the Stumm types and his own 1940 illustrated 

specimen. All of Merriam's illustrated thin sections (1974b) 

and acetate peels (1940) are in the collections of the USNM, 

but remnants of the specimens from which they were cut 

have been found for only three of them (marked * in 

Material list). I consider Merriam's specimens to belong to 

three species: eight are B. invaginata, seven (including the 

1940 specimen) are an unnamed species (not a siphonophrentid) characterized by coarse monacanthine trabeculae 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 819 
50 

X X 

45 

X X X 

X X 

X 

40 

....... 

~ X <( 

fa... 

UJ 

(/) 35 a: 

0 

"""') 

<( 

~ 

u.. 

0 Cylindrical a: 

UJ 30 IJl 

~ 

::> 

z 

Conical 

25 

20 

15 ~------~---------L--------~------~ 

5 10 15 20 25 

DIAMETER (d), IN MILLIMETERS 

Figure 6. Number of major septa (n) plotted against diameter (d) for Breviphrentis invaginata (Stumm). Data are 

from 10 individuals from the Eurekaspirifer pinyonensis 

Zone, Nevada Formation, Emsian, Nevada. Included are 

the five Stumm (1937) specimens and five of the eight 

Merriam (1974b) specimens listed in the "Material" section. The two smallest thin sections are from the conical 

part of one specimen; the others are from cylindrical 

parts, and each X represents one individual. 

and rejuvenescence (USNM 159296, 159299, 159301, 

159303, 159309, 159310, 163269), and one is Breviphyllum lonense (USNM 159298); in addition, one is indeterminate (USNM 159308). Both Stumm (1937) and Merriam 

(1974b) emphasized rejuvenescence in their descriptions, 

but the holotype and other specimens that I accept as B. 

invaginata have simple rugae and siphonophrentid microstructure, whereas those specimens with true rejuvenescence (including abandoned calice rims at more or less 

consistent intervals) have a quite different microstructure in 

which coarse monacanthine trabeculae are prominent where 

septa are well preserved. This form is best illustrated in 

820 Shorter Contributions to Paleontology and Stratigraphy, 1992 

Merriam's (1974b) pl. 16, figs. 10-13. The Stumm (1937) 

unillustrated "paratype" (specimen USNM 94443a) probably belongs to this species also. 

My description is based on the Stumm types of B. 

invaginata and H. nevadensis and on the eight Merriam 

specimens. 

Material.-From Stumm, 1937: Lectotype, USNM 

94443 (original of pl. 53, fig. 2; two thin sections were 

prepared later and illustrated by Hill, 1981, fig. 95:1a-b). 

Paralectotype, USNM 454103 (originally numbered 94443; 

pl. 54, fig. 2a). "Paratype," thin section USNM 454104 

(originally numbered 94443a; pl. 54, fig. 2b). 

From Merriam, 1974b (*=specimen remnants in 

addition to thin sections; plate and figure numbers in 

parentheses): USNM 159297* (15, 3-4), 159300 (15, 9), 

159302 (15, 11), 159304 (16, 3), 159305* (16, 4), 159306 

(16, 5-7), 159307 (16, 8), 159311 * (16 ' 15-16). 

Types of Heterophrentis nevadensis Stumm, 1937: 

holotype, USNM 94442 (pl. 53, fig. 1; pl. 54, fig. 1a; a 

longitudinal section prepared later is illustrated here for the 

first time); paratype(s), USNM 94442a (pl. 54, fig. 1b--c; 

remnant and two thin sections may represent two specimens). 

All of the above specimens are from the Nevada 

Formation, Eurekaspirifer pinyonensis Zone, Emsian, 

Nevada. 

Breviphrentis halli (Milne-Edwards and Haime), 1851 

Plate 14, figures 1-6; plates 15, 16 

1851. Zaphrentis halli M.-E. and H., p. 341. 

1860. Z. halli M.-E. and H. Milne-Edwards, p. 347. 

1877. Z. halli M.-E. and H. Hall, pl. 20, figs. 1, 3, 5, 

8. Not figs. 4, 6, and 7, which are B. cista 

(described below); not figs. 2 and 9, which are 

undetermined but not Breviphrentis. 

? 1904. Z. halli M.-E. and H. Greene, p. 179, pl. 53, figs. 

3-4. 

1951. Siphonophrentis. Oliver, pl. 2, fig. 1. 

? 1965. Siphonophrentis halli (M.-E. and H.). Stumm, p. 

24, pl. 12, fig. 8; pl. 14, figs. 1-4, 7-10. See 

Stumm synonymy for other possible forms from 

the Falls of the Ohio area. 

? 1987. S. sp. aff. S. halli (M.-E. and H.). Oliver, p. 3, 

pl. 1 , figs. 5-7. 

Type specimens.-Holotype, Paris EM 15173, not 

previously illustrated; herein pl. 14, figs 1-3; pl. 15, figs. 

1-2. Illustrated specimens of Hall, 1877, pl. 20 (fig. no. in 

parentheses): AMNH 39405 (1), 4942 (5), 39404 (8); 

NYSM 6262 (3). 

Type locality.-Both Milne-Edwards and Haime 

(1851) and Hall (1877) gave Skaneateles Lake, N.Y., as the 

source of their specimens. This is almost certainly the 

Staghom Point coral bed in the Otisco Shale Member, 

Ludlowville Formation, Hamilton Group (Givetian). The 

coral bed crops out for more than 2 km along the east side 

of Skaneateles Lake and has been a notable source for 
Hamilton corals for 150 years (see description and discussion in Oliver, 1951). The following description is based 

primarily on collections from the coral bed at Willow Point 

and Staghom Point on the lakeshore. The bed is a mass of 

hom corals from 0. 5 to 1. 5 m thick; Oliver ( 1951) listed 13 

species of rugose corals from the coral bed (at all known 

localities), of which B. halli (including B. cista n. sp.) was 

the most abundant. 

Diagnosis.-Medium to large, cylindrical Breviphrentis with a moderately deep calice and well-developed 

siphonofossula in which is a distally short cardinal septum. 

Major septa extend from half to two-thirds the distance to 

the axis; those adjacent to the cardinal septum bend toward 

it, but most are radially arranged. Minor septa short. 

Tabulae thin, closely spaced, M-shaped with axial part 

nearly flat. 

External features. -Ceratoid to cylindrical but 

mostly cylindrical when fully developed; common cylindrical stage diameters are 30 to 50 mm; maximum measured 

length is 25 em. One ceratoid specimen is 65 mm in 

diameter and 14 em long; it seems to have reached cylindrical growth only in its last 2 em. Some specimens are 

straight or hom shaped, but longer ones tend to be irregular, 

curving or abruptly bending away from the initial plane of 

curvature. The position of the cardinal septum relative to 

the curvature varies, but it tends toward the convex side in 

the early stages. Exteriors gently rugose with shallow septal 

grooves. Calice depth approximately half the diameter; flat 

or gently concave axially. Siphonofossula deep, formed by 

downbent tabulae and a short cardinal septum. 

Internal features.- In cylindrical stage, major septa 

amplexoid, extend 0.4 to 0. 7 of the radius (a few may be 

longer), and number 40 to 63 in sections having diameters 

of 30 to 50 mm (text-fig. 7). Minor septa short, approximately 0.2 of the radius. Septa attenuate to only moderately 

thickened except at periphery where they thicken to form a 

wall (septotheca) 1 to 4 mm thick. In transverse sections 

through conical stages, major septa may be longer and 

thicker or still short and attenuate; numbers of septa are 

proportionately smaller (text-fig. 7). At diameters smaller 

than 10 to 15 mm (30-36 major septa), the lumen may be 

nearly filled by dilated septa. Minor septa are short. 

Bilateral septal arrangement is based on recognition 

of cardinal and counter septa. C is short and in a siphonofossula; one or two major septa on either side may bend 

toward the C, but most septa project radially. The K is 

commonly slightly shorter (longer in some sections) than 

the other septa but is flanked by long minors. The C and K 

septa can be recognized in almost any section by these 

characters. 

In longitudinal section, tabulae are thin and closely 

spaced (see pls. 14-16), commonly complete; axial area is 

without septal traces, flat or gently concave. Peripherally 

the tabulae tum down a short distance, then out to the wall; 

there may be a slight upturn at the margin, and some 

individuals have thin, short, descending tabellae in the 

upturn. 

Microstructure is as described for the family. 

There is considerable variation in length and thickness of major septa, in wall thickness, and in the shape and 

spacing of tabulae and their marginal configuration. Range 

of variation can best be judged from the photographs. 

Variation in number of major septa at given diameters (n:d 

ratio) is graphed in text-figure 7. 

Discussion. -B. halli differs from B. cista, with 

which it is associated, in its shorter and thinner septa and 

thinner, flatter, and more closely spaced tabulae. These are 

qualitative differences that are hard to define convincingly, 

but in the mixed sample of 56 Breviphrentis from Skaneateles Lake, only 1 or 2 were difficult to place. Both B. 

invaginata and B. pumilla are smaller and have thinner 

septa and tabulae. 

Occurrence.-Hamilton Group, New York. B. halli 

and (or) B. cista range through the Skaneateles, Ludlowville, and Moscow Formations, but individual occurrences are yet to be worked out. Numerous nominal species 

from the Middle Devonian of the Falls of the Ohio area (see 

Stumm, 1965, p. 23-25) and from the Columbus Limestone, Ohio, may or may not be synonymous with the 

Staghom Point species. 

Material.-This description is based on a sample of 

20 specimens from the Staghom Point coral bed, on the east 

shore of Skaneateles Lake (see discussion of type locality, 

above, and locality data at end of paper); the sample was 

extracted from much larger collections made by the author. 

The sample was supplemented by the holotype and four 

Hall specimens, all of which are from Skaneateles Lake and 

probably from the same bed. 

Holotype, EM 15173. Illustrated specimens of Hall, 

1877, pl. 20 (fig. no. in parentheses): AMNH 39405 (1), 

NYSM 6262 (3), AMNH 4942 (5), AMNH 39404 (8). 

Staghom Point coral bed sample, illustrated: USNM 

454105-454107, 454110, colln. USGS 7543-SD; 454108, 

454109, 454112, 7542-SD; 454111, 8763-SD; unillustrated (measured, text-fig. 7): USNM 454113-454118, 

USGS 7542-SD; 454119, 454120, 7543-SD; 454121, 

454122, 8763-SD; 454123, 454124, 8781-SD. 

Breviphrentis cista n. sp. 

Plate 14, figures 7-9; plates 17, 18 

part 1877. Zaphrentis halli Milne-Edwards and Haime. Hall, 

pl. 20, figs. 4, 6, 7 (not figs. 1-3, 5, 8-9, for 

which see B. halli). 

Type specimens.-Holotype, NYSM 6264; paratypes, NYSM 6263 and 6265 and USNM 454125-454153. 

Type locality. -Skaneateles Lake, N.Y. The holotype and other Hall specimens were almost certainly collected from the Staghom Point coral bed on the east shore of 

the lake as were the other paratypes. See locality data at end 

of paper. 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 821 
75 .-----.------.-----,.-----.------.-----,.-----.------.-----,-----/70------.-----.------, 

70 

65 

60 

55 

:5: 

<( 50 Ia... 

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15 ~----~-----L------~----~-----L----~L-----~-----L----~L-----~-----L----~L---~ 

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DIAMETER (d), IN MILLIMETERS 

Figure 7. Number of major septa (n) plotted against diameter (d) for Breviphrentis halli (Milne-Edwards and Haime). 

Data are from 39 sections of a sample of 20 individuals 

from the Staghorn Point coral bed, Skaneateles Lake, 

N.Y., plus 1 section from the Milne-Edwards and Haime 

(1851) holotype and 4 sections of 3 Hall (1877) specimens. 

Diagnosis. -Medium to large, cylindrical Breviphrentis with a moderately deep calice and well-developed 

siphonofossula in which is a short cardinal septum. Major 

septa extend two-thirds the distance to the axis, are somewhat dilated, and are radially arranged except near the 

fossula. Minor septa short. Tabulae thickened, widely 

spaced, M-shaped with axial part strongly concave. 

External features. -Ceratoid to cylindrical but cylindrical when fully developed; common cylindrical diameters 

822 Shorter Contributions to Paleontology and Stratigraphy, 1992 

The holotype and Hall specimens are from Skaneateles 

Lake and probably the same coral bed. See figure 2 for 

meaning of symbols; holotype marked by a square; Hall 

specimens marked by +. Bounding lines are identical in 

figures 7 and 8 to facilitate comparison of B. halli with B. 

cista. 

are 20 to 40 mm, maximum measured length is 30 em. 

Longer specimens tend to bend or curve irregularly. Exteriors gently rugose. Calice depth half or more the total 

diameter; axially concave. Siphonofossula deep, formed by 

downbent tabulae and a short cardinal septum. 

Internal features. -In transverse sections through the 

cylindrical stage, major septa extend 0.6 to 0. 7 of the radius 

(although a few may be longer) and number 40 to 58 in 

sections having diameters of 25 to 50 mm (text-fig. 8). 
60 

20 

16 

55 

22 

21 

+ 

50 16 20 6 9 4 18 

22 

~ 45 8 7 3 19 10 21 

1 <( 

Ia.. 

LU 

(/) 2 

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0 10 19 12 I 

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~ 

LL.. 7 X 15 0 35 5 17 a: X 4 13 11 LU 

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::::> 

z 13 1~ 30 14 

11 

25 

20 

15 ~----~----~------~----~----~------~----~----~----~ 

5 10 15 20 25 30 35 40 45 50 

DIAMETER (d), IN MILLIMETERS 

Figure 8. Number of major septa (n) plotted against diameter (d) for Breviphrentis cista 

n. sp. Data are from 51 sections from a sample of 29 individuals from the Staghorn 

Point coral bed, Skaneateles Lake, N.Y., plus 3 sections of 3 Hall (1877) specimens, 

also from Skaneateles Lake and probably the same coral bed. See figure 2 for meaning 

of symbols; Hall specimens marked by+. Bounding lines are identical in figures 7 and 

8 to facilitate comparison of B. cista with B. halli. 

Minor septa short, 0.2 to 0.3 of the radius. Septa moderately to quite dilated, commonly thickest in middle part, 

rarely in contact with adjacent major septa where thickest. 

Septotheca is 2 to 6 mm thick. In transverse sections 

through conical stages, major septa are longer and thicker; 

numbers are shown in text-figure 8. Sections smaller than 

10 mm are not known. 

Cardinal and counter septa are as in B. halli; C is 

short and in a siphonofossula, and K is flanked by long 

minors. 

In longitudinal sections, tabulae tend to be thickened, 

widely spaced, and strongly concave axially; most are 

complete. Peripherally the tabulae turn down, then curve 

out to the wall; there may be a slight upturn at the margin. 

Microstructure is as described for the family. 

There is considerable variation in length and thickness of major septa, in wall thickness, and in the shape, 

thickness, and spacing of tabulae; the range of such variation is best judged from the photographs. Variation in septal 

number at given diameters (n:d ratio; text-fig. 8) is 

essentially the same as in B. halli. 

Discussion.-B. cista differs from B. halli in its 

tabulae, which are widely spaced, thickened, and axially 

concave in the B. cista but are closely spaced, thin, and 

axially flat in B. halli (compare illustrated longitudinal 

sections). In addition, major septa tend to be longer and 

thicker in B. cista. Both species are variable in all of these 

characters and there is some overlap, and there are even 

some morphologically intermediate individuals. Because of 

this variability, I have questioned the probability of there 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 823 
being one or two species. My decision to recognize two 

species is based on three reasons: (1) most specimens are 

easily assigned to one species or the other; (2) characteristic 

forms are very different; and (3) if merged, the one species 

would embrace such extensive variation that it would be 

impractical to describe other EAR species. Variation in B. 

halli and comparisons with other Breviphrentis are discussed in the description of B. halli. 

Occurrence. -Staghom Point coral bed, Givetian, 

New York. Presence in other parts of the Hamilton Group 

and equivalents in the EAR is yet to be determined. 

Material.-Holotype, NYSM 6264, paratypes 

NYSM 6263, 6265 (all three illustrated by Hall, 1877, pl. 

20, as Z. halli M.-E. and H.). In addition to the 3 Hall 

specimens, the species description is based on 29 specimens 

from the coral bed along Skaneateles Lake: illustrated 

paratypes: USNM 454125-454130, 454132-454134, colin. 

USGS 7542-SD; 454131, 7543-SD; unillustrated paratypes 

(measured, text-fig. 8): USNM 454135-454142, USGS 

7542-SD; 454143-454145, 7543-SD; 454146-454148, 

8763-SD; 454149-454153, 8781-SD. 

Breviphrentis pumilla n. sp. 

Plate 19 

Type specimens.-Holotype, USNM 454154; paratypes, USNM 454155-454180. Centerfield Member (Givetian), Ludlowville Formation, Hamilton Group, western 

New York. 

Diagnosis. -Small, cylindrical Breviphrentis with a 

deep calice and weak siphonofossula. Major septa extend 

half to two-thirds the distance to the axis, are radially 

arranged, and are moderately attenuate; minor septa short. 

Tabulae thin or thickened, closely spaced, axially flat. 

External features. -Individuals are ceratoidcylindrical tending to cylindrical; coralla are straight, irregularly curved, or contorted; the maximum diameter in some 

individuals is 5 to 15 mm below the calice margin, 

reflecting a size contraction late in life (gerontic?). The 

exterior is very rugose, commonly masking the overall 

form; septal grooves and interseptal ridges are subdued. 

Common cylindrical diameters are 13 to 18 mm; total 

lengths are 40 to 50 mm; maximum diameter and length in 

type population, 21 and 60 mm. Calice depth approximately 

half the diameter; cardinal fossula formed by short cardinal 

septum and downbent tabulae but not strongly marked; 

septal traces extend nearly to axis on uppermost tabula. 

Internal features.- In cylindrical stage, major septa 

amplexoid, extend 0.4 to 0.6 of the radius, longer on the 

tabular surfaces; they number 24 to 32 in sections having 

diameters of 13 to 18 mm (text-fig. 9). Minor septa short, 

commonly 0.3 the radius. Septa attenuate to moderately 

thickened except at periphery where they dilate to form a 

wall (septotheca) 0.8 to 3.5 mm thick. In transverse 

sections through early (conical) stages, major septa tend to 

be longer and relatively thicker, but with plentiful intersep824 Shorter Contributions to Paleontology and Stratigraphy, 1992 

40 ~------~~---------~~------~--------~ 

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DIAMETER (d), IN MILLIMETERS 

Figure 9. Number of major septa (n) plotted against diameter (d) for Breviphrentis pumilla n. sp. Data are from 43 

sections of a sample of 27 individuals from the Centerfield 

Member, Ludlowville Formation, in New York. See figure 

2 for meaning of symbols; 1-9 and the X's represent the 

East Alexander population; 10-15 and the plus signs represent specimens from other localities. 

tal space at diameters of 8 mm, where minor septa are 

commonly 0.2 the radius. At smaller diameters, some 

apical regions are virtually filled by the thickened tabulae 

and septa (seen in longitudinal sections). 

Septa are radially arranged except immediately adjacent to the cardinal septum. In most transverse sections, the 

cardinal-counter plane is marked by septa that differ from 

the other major septa. The cardinal septum is generally 

short with the two adjacent major septa curved toward it. A 

weak siphonofossula is commonly marked by the U-shaped 

interception of one or two tabulae. The counter septum is 

commonly long but may be short or differ in thickness from 

the other septa; where short, it is not necessarily distinguishable from the cardinal septum. 

In longitudinal section, tabulae are thin or thickened, 

commonly complete; they are flat or slightly concave 

axially, downtumed peripherally; tabulae in smaller coralla 

tend to be more strongly arched. Axial ends of major septa 

commonly project up from the tabular surfaces and may 

provide much of the thickening. 

Microstructure is as described for the family. 
There is considerable variation in most of the characters described. This is illustrated in plate 19 to the extent 

feasible. Variation in numbers of major septa at given 

diameters (n:d ratio) is shown in text-figure 9. 

Discussion. -B. pumilla differs from other described 

Breviphrentis in its small size, short minor septa, and weak 

siphonofossula. 

Occurrence. -Common in the Centerfield Member, 

Ludlowville Formation, western New York. 

Material. -The description is based primarily on a 

sample of 19 specimens, including the holotype, from a 

creek locality 2.9 km east of East Alexander, N.Y. Eight 

additional specimens from five localities were used to 

supplement the description. Holotype, USNM 454154, 

colln. USGS 4678-SD. Illustrated paratypes: USNM 

454155-454164, USGS 4678-SD; 454165-454166, 

5692-SD; 454167, 11197-SD; 454168, 5705-SD. Unillustrated paratypes (measured, text-fig. 9): USNM 

454169-454176, USGS 4678-SD; 454177, 5682-SD; 

454178, 5686-SD; 454179-454180, 5692-SD. 

Genus SIPHONOPHRENTIS O'Connell, 1914 

1914. Siphonophrentis O'Connell, p. 187, 190, 191. 

part? 1938. Siphonophrentis O'Connell. Stewart, p. 25-27. 

1940. Siphonophrentis O'Connell. Lang, Smith, and 

Thomas, p. 120. 

part 1944. Siphonophrentis O'Connell. Shimer and Shrock, p. 

93 (? pl. 28, lower left fig.). 

1947. Siphonophrentis O'Connell. LeMaitre, p. 28-29. 

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

? 1950. Siphonophrentis O'Connell. Wang, p. 214 (?pl. 5, 

fig. 26a-b; not pl. 9, fig. 70). 

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

not 1960a. Siphonophrentis O'Connell. Oliver, p. 87-89 

( = Breviphrentis). 

1963. Siphonophrentis O'Connell. Smith and Thomas, p. 

163, 167-168. 

part 1965. Siphonophrentis O'Connell. Stumm, p. 23-25. 

? 1970. Siphonophrentis O'Connell. Brice, p. 259-261. 

? 1974a. Siphonophrentis (Siphonophrentis) O'Connell. 

Merriam, p. 23. 

197 4b. Siphonophrentis ( Siphonophrentis) 0' Connell. 

Merriam, p. 41--42. 

? 1975. Siphonophrentis O'Connell. Joseph and Tsien, p. 

196. 

1978. Siphonophrentis O'Connell. Birenheide, p. 69. 

1981. Siphonophrentis O'Connell. Hill, p. F147-F148. 

? 1982. Siphonophrentis O'Connell. Yu and Kuang, p. 

53-55. 

1989. Siphonophrentis O'Connell. Wang and others, p. 

22-23. 

The above synonymy is tentative because none of the 

descriptions of Siphonophrentis or its nominal species has 

included pre-ephebic morphology. At present, judgment 

and comparisons can be made only on the basis of 

cylindrical-stage characters: large size; cylindrical form; 

short amplexoid septa; complete, axially flat tabulae; and 

presence of a siphonofossula. 

Type species. - Caryophyllia gigantea Lesueur, 

1821, p. 296-297; by original designation. According to 

Lesueur, the species was found "a Waren, a trente mille[s] 

d'Utica" (New York). Warren, N.Y., at the southern tip of 

Herkimer County, is 39 km (24 airline miles; perhaps 30 

miles by road in the early 1800's) from Utica, and this is 

commonly assumed to be the locality of the specimen(s). If 

this assumption is correct, the source is most likely the 

Edgecliff Member of the Onondaga Limestone. The village 

of Warren is in an area of thick glacial cover (Rickard and 

Zenger, 1964, geologic map), but three Edgecliff coral 

bioherms crop out 1.8 to 2.4 km to the north (bioherms 18, 

19, and 20 of Oliver, 1956), and these and other Edgecliff 

outcrops are possible sources. Even if the specimen is from 

drift, the Edgecliff is the most likely provenance, because 

(1) the Edgecliff underlies an extensive area just north of 

Warren and a large coral is unlikely to have traveled far, 

and (2) no such large coral is known from other possible 

source rocks farther north. 

Lesueur (1821, p. 296-297) described the coral in 

French. The translation that follows was made with the 

assistance ofP. Semenoff-Tian-Chansky, Paris, who was of 

particular help in determining that Lesueur's "mille" was 

probably the American mile and that his "pouce" was 1112 

of the early 19th century French "pied" of 32.4 em (thus, 1 

pouce=27 mm). 

Caryophyllia gigantea. This is the largest of all 

[caryophyllids] that I have seen. Its height is approximately 

5 to 6 "pouces" [135-162 mm, see above], with an upper 

part diameter of 2 to 2.5 "pouces" [54-67 mm]; it has the 

form of a curved cone, longitudinally striated, with slight 

transverse swellings. A section toward the base shows 

transverse partitions, from which rise the septa. These 

septa are divided into two parts [? groups] in length 

towards the margin. This division is developed from the 

base to the summit. The septa are fairly thick; they are 

connected to each other by cellular oblique partitions, 

arched, medium and irregular in a vertical section, but 

seen transversely they are quite regular, and thin toward 

the margin so that the widest are at the center. 

When decorticated, so that the small cells are 

uncovered, one can mistake them for pieces of bone. 

The description is hardly definitive, as it would fit 

almost any large rugose coral. The "cellular oblique partitions" may be tabulae but might more logically be dissepiments. However, stability is served by assuming the best, 

rather than the worst. Lesueur did not illustrate his species, 

and no original specimen is known to exist; thus, the 

description is the only basis for its recognition, and considerable latitude is available to the reviser. 

Turbinolia buceros var. elongata Rafinesque and 

Clifford, 1820, from the Jeffersonville Limestone at the 

Falls of the Ohio, is considered to be a senior subjective 

synonym of S. gigantea Lesueur. This was first suggested 

by Stumm (1965), who selected a neotype for S. elongata; 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 825 
however, this did not stabilize Siphonophrentis because the 

genus is ultimately based on S. gigantea. 

In order to provide a solid basis for Siphonophrentis, 

I select a neotype for S. gigantea from the Onondaga 

Limestone in New York. The specimen fits current concepts 

of the genus and is considered conspecific with the neotype 

of S. elongata (R. and C.) as designated by Stumm. The 

two neotypes are described and illustrated from thin sections, and the species is discussed, but a full synonymy and 

description of the species are beyond the scope of this work. 

Diagnosis. -Large, cylindrical rugose corals with 

short, amplexoid major septa and very short minor septa; a 

short cardinal septum occupies a commonly well developed 

siphonofossula. Septa tend to be very thin in the cylindrical 

(mature) growth stage, but they are strongly dilated in parts 

of the earlier, expanding stages and may be in later stages as 

well. A septotheca is formed by the dilated peripheral ends 

of the septa; epitheca thin and not necessarily preserved. In 

early stages, the wall can be quite thick, particularly in the 

cardinal quadrants. Tabulae complete, arched, axially flat 

or concave; central area free of septal ridges. 

Discussion.-My description and discussion of 

Siphonophrentis elongata are based on the neotype specimens of S. gigantea and S. elongata, both of which are 

illustrated from thin sections, plus the sectioned and illustrated specimen of Stumm (1965) and several additional, 

mostly fragmentary, specimens from eastern North America. A tentative synonymy is given, but this is based on 

published illustrations. 

At this time, S. elongata is the only species assigned 

to Siphonophrentis. Many nominal species of both Siphonophrentis and Breviphrentis must be reinvestigated (for 

example, those described, illustrated, and synonymized by 

Stumm, 1965, p. 23-25). Several of the specimens illustrated by Stumm (1965, pls. 9, 12-15) seem to be typical 

Siphonophrentis, but few have been sectioned, and those 

that have been are not the name-bearing types. As with S. 

elongata/gigantea, the problems are largely ones of type 

specimens and priority of publication. 

Distribution. -All confirmed S. elongata may be 

from limestones of Edgecliff age (early Eifelian) in eastern 

North America, but good Siphonophrentis sp. range up 

through much of the Onondaga, Columbus, and Jeffersonville Limestones (early to late Eifelian). 

Most of the corals that have been described and 

illustrated as Siphonophrentis from areas outside of the 

EAR either are not this genus or are indeterminate from the 

published illustrations and need to be restudied. Specimens 

that appear to be Siphonophrentis on the basis of illustrations of the ephebic stage are from Morocco (LeMaitre, 

1947), Spain (Barrois, 1882; Birenheide, 1978), England 

(Smith and Thomas, 1963), and the former U.S.S.R. 

(Ivania, 1965, as Heterophrentis). Specimens illustrated 

from western Canada, the French Pyrenees, Bulgaria, 

Poland, and Afghanistan; most of those from the former 

826 Shorter Contributions to Paleontology and Stratigraphy, 1992 

U.S.S.R.; and the numerous ones from China either are not 

Siphonophrentis or are indeterminate. 

Siphonophrentis elongata (Ratinesque and Clifford), 1820 

Plates 20-22 

1820. 

1821. 

? 1847. 

1851. 

1859. 

1860. 

1874. 

not? 1875. 

not? 1875. 

part 1876. 

not 1882. 

not 1883. 

not 1884. 

part? 1887. 

part? 1887. 

1887. 

not? 1890. 

1899. 

1901. 

? 1904. 

1906. 

1909. 

1914. 

? 1938. 

1944. 

part 1949. 

? 1950. 

part 1956. 

not 1956. 

? 1956. 

? 1961. 

1965. 

1981. 

Turbinolia (Turbinolia) buceros var. elongata R. 

and C., p. 233. 

Caryophyllia gigantea Lesueur, p. 296-297. 

Cyathophyllum gigas Clapp in Yandell and 

Shumard, p. 8. 

Zaphrentis gigantea (Lesueur). Milne-Edwards and 

Haime, p. 340-341, pl. 4, fig. la--c. 

Z. gigantea (Lesueur). Billings, p. 121. 

Z. gigantea (Lesueur). Milne-Edwards, p. 346. 

Z. gigantea (Lesueur). Nicholson, p. 22, pl. 3, 

figs. 1-la. 

Z. eriphyle Billings, p. 233. 

Z. hecuba Billings, p. 234. 

Z. gigantea (Lesueur). Rominger, p. 146-147, pl. 

52, 3 figs. The specimen on the right is the 

neotype of S. elongata; the upper left specimen is 

probably conspecific, but the lower left specimen is 

questionable. 

Z. ponderosa Hall, p. 27. 

Z. ponderosa Hall. Hall, p. 288, pl. 19, fig. 7. 

Z. ponderosa Hall. Hall, p. 431. 

Z. immanis Davis, pl. 138, fig. 2; pl. 139 (not pl. 

80, fig. 18). 

Z. maconathi Davis, pl. 136, fig. 3 (? figs. 1, 2, 

4). 

Z. gigantea (Lesueur). Davis, pl. 137, figs. 1-2; 

pl. 138, fig. 1. 

Z. gigantea Rafinisque [sic]. Lesley, p. 1272. 

Z. gigantea (Lesueur). Lambe, p. 252-254. 

Z. gigantea (Lesueur). Lambe, p. 125, pl. 9, figs. 

3-3a. 

Z. callosus Greene, p. 198, pl. 58, figs. 1-2. 

Z. gigantea (Lesueur). Grabau and Shimer, p. 193. 

Z. gigantea (Lesueur). Grabau and Shimer, p. 

56-57' fig. 80. 

Siphonophrentis gigantea (Lesueur). O'Connell, p. 

187' 191. 

S. gigantea (Lesueur). Stewart, p. 25, pl. 4, fig. 1. 

S. gigantea (Lesueur). Shimer and Shrock, p. 93, 

pl. 28, figs. 25-26, 30-31. 

S. gigantea (Lesueur). Stumm, p. 12, pl. 5, figs. 

1, 3 (? figs. 2, 4). 

S. gigantea (Lesueur). Wang, pl. 5, fig. 26a-b; pl. 

9, fig. 7. 

S. gigantea (Lesueur). Hill, p. F270 (? fig. 

183:7a-b). 

S. cf. gigantea (Lesueur). Warren and Stelck, pl. 

4, fig. 8. 

S. gigantea (Lesueur). Ma. p. 68-69 (? pl. 57, fig. 

10). 

S. gigantea (Lesueur). Fagerstrom, p. 12, pl. 4, 

fig. 5. 

S. elongata (R. and C.). Stumm, pl. 13, figs. 1-4. 

S. elongata (R. and C.). Hill, p. F147-F148, fig. 

79:4a--c. 
? 1989. S. gigantea (Lesueur). Wang and others, p. 23, pl. 

2, fig. 8a-b; pl. 20, figs. 7-12. 

Type specimens. -Neotype of S. elongata, UMMP 

8616, designated by Stumm, 1965, p. 24, the original of 

Rominger, 1876, pl. 52, right-hand fig. (reversed), and 

Stumm, 1965, pl. 13, figs. 1-2. Thin sections are illustrated 

here for the first time (pl. 21, figs. 1-4). Jeffersonville 

Limestone, Falls of the Ohio River, at or near Louisville, 

Ky. 

Neotype of S. gigantea, here selected, USNM 

454181 (pl. 20, figs. 1-4). The specimen was collected in 

1895 by Charles Schuchert; it is labeled simply "Onondaga, 

Leroy, New York." Most of the members of the Onondaga 

Limestone crop out in or near Leroy, but the matrix 

lithology indicates a probable origin in the Edgecliff Member. Leroy is 250 km west of Warren, the presumed locality 

of Lesueur's specimen. I would have preferred to select a 

neotype from the Warren area but felt it most important to 

have a good specimen with which to secure the name. 

Diagnosis. - Szphonophrentis with short septa and 

early stage septal dilation in cardinal quadrants. 

Description.-Large, ceratoid becoming cylindrical, 

solitary corals as much as 75 em long (high) and 8 em in 

diameter; the neotypes of S. gigantea (USNM 454181) and 

S. elongata (UMMP 8616) are (respectively) 42 and 25 em 

long and 6. 0 and 4. 2 em in maximum diameter; the largest 

single specimen that I have measured is 73 em long and 7.5 

em in diameter (USNM 454182, Falls of the Ohio, pl. 22). 

Most specimens (including the types) are gently curved but 

not necessarily in a single plane. The expanding, ceratoid 

stage is approximately half the total length of each neotype. 

Exteriors are commonly worn but marked by gentle rugae; 

when well preserved there are shallow septal grooves and 

broad, low interseptal ridges. Calice broad, flat bottomed 

with a prominent cardinal fossula formed by a short cardinal 

septum and strongly downbent tabulae (siphonofossula). 

The calice floor is the uppermost tabula; septa project only 

a short distance from the calice wall but farther on the floor; 

an axial area is commonly smooth, without septal traces, 

but traces extend to the axial area in some individuals. 

In transverse sections through the cylindrical parts of 

coralla, major septa amplexoid, extend from 0.3 to 0.5 of 

the radius; minor septa extend less than 0.1 the radius. 

Numbers of major septa at various diameters are shown in 

text-figure 10. A wall, less than 1 mm thick, is of uncertain 

structure but may be formed by the dilated peripheral ends 

of the septa. The cardinal septum is short and is in a fossula 

that is most clearly defined by the U-shape of downbending 

tabulae that are cut by the section. In longitudinal sections 

through the cylindrical part, tabulae are numerous, commonly complete, flat over a broad axial area, and strongly 

downbent at the periphery; there may be uparching between 

the flat and downbent parts. 

In lower parts of coralla, septa are partly dilated. In a 

transverse section through the upper ceratoid part of USNM 

454181 (pl. 20, fig. 3), the peripheral parts of major septa 

100 

90 

80 

:£ 

<( 

1-

a... 

UJ 

(/) 70 a: 

0 

...., 

<( 

~ 

LL 

0 60 

a: 

UJ 

a:l 

~ 

::::> 

z 50 

40 

30 ~----~----~----~----~------~--~ 

20 ~ w ~ M m 80 

DIAMETER (d), IN MILLIMETERS 

Figure 10. Number of major septa (n) plotted against 

diameter (d) for Siphonophrentis elongata (Rafinesque 

and Clifford). Data are from 20 sections of 7 individuals. 

USNM 454181 (circles), neotype of S. gigantea, Onondaga 

Limestone, New York. UMMP 8616 (squares; neotype of S. 

elongata) and all the other specimens are from the Jeffersonville Limestone, Falls of the Ohio, Ky.-Ind. Sections 

from the same individual are either connected by lines 

(USNM 454182, pl. 22) or represented by the same numbers (1-3). 

near the cardinal fossula are fully dilated and in lateral 

contact forming a septotheca up to 1.0 em thick at the 

fossula; this thins and disappears toward the counter quadrants; parts of septa inside of the septotheca are very thin 

and the change from thick to thin is abrupt. The fossula is 

clearly marked and the minor septa are very short. In a 

section 15 em lower (d=39 mm; pl. 20, fig. 4), dilation is 

evident but markedly asymmetrical; major septa are wedge 

shaped, and there is space between most of them. In the 

lowest section observed (d=approx. 20 mm), dilation is 

marked; thickened wedge-shaped septa are in the cardinal 

quadrants. In a longitudinal section taken just above the first 

section described, the cardinal thickening disappears 

upward in a distance of 3 to 4 em (pl. 20, fig. 2). 

UMMP 8616 has comparable septal dilation in the 

ceratoid stage (pl. 21 , figs. 1-4). A section through the 

middle of the ceratoid stage has considerable cardinal 

quadrant dilation; septa are wedge shaped with much 

interseptal space (pl. 21, fig. 3); this is comparable to the 

lower section of USNM 454181. A lower section (d=21 

mm, pl. 21, fig. 4) has attenuate septa with a relatively thin 

wall. 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 827 
A third specimen, USNM 454182 (pl. 22), has 

varying amounts of dilation extending through the cylindrical and conical stages. The circumferential position of 

greatest dilation varies along the length of the coral but 

tends to be in the counter quadrants rather than the cardinal, 

as in the two neotypes. 

Microstructure is as described for the family. 

Discussion.-Early stage dilation in the cardinal 

quadrants is common to pervasive inS. elongata although, 

as indicated, it varies in extent. More study is needed to 

understand the limits of the species, but specimens with 

both more and less dilation are tentatively included; in 

some, the dilation extends to, or is centered in, the counter 

quadrants. The parts of specimens with thin walls and 

attenuate septa are commonly crushed; in many specimens, 

only the dilated side of the coral is preserved. USNM 

454181 illustrates the contrasting strength of dilated and 

attenuate sides of the ceratoid stage (pl. 20, figs. 3, 4), but 

the same specimen is uncrushed in its thin-walled cylindrical stage (pl. 20, fig. 1). 

Occurrence. - Edgecliff Member of the Onondaga 

Limestone (early Eifelian) and age-equivalent parts of the 

Columbus and Jeffersonville Limestones in New York, 

southwestern Ontario, Ohio, and Indiana-Kentucky. The 

same species, or a similar one, occurs in the Moorehouse 

Member of the Onondaga (middle Eifelian) and may occur 

in the higher parts (early to late Eifelian) of all three 

formations in the area indicated. 

Material.-Neotype of S. gigantea, USNM 454181, 

Onondaga Limestone, Leroy, N.Y. Neotype of S. elongata, 

UMMP 8616; illustrated specimen of Stumm, 1965, pl. 13, 

figs. 3-4, UMMP 26423; additional illustrated specimen, 

USNM 454182; additional measured specimens, USNM 

455881-455883; the last six specimens are from the Jeffersonville Limestone, Falls of the Ohio, Ky.-Ind. Additional 

specimens in the USNM collections from the Edgecliff 

Member of the Onondaga Limestone in New York, mostly 

crushed and incomplete, and additional silicified specimens 

from the Jeffersonville Limestone have been used to supplement the description. 

LOCALITY DATA 

Onondaga Limestone Localities 

Specimens of Metaxyphrentis prolifica (Billings), 

except the types, are from the Edgecliff Member of the 

Onondaga Limestone at the following nine localities, all in 

New York. The list is arranged as follows: locality number, 

name of USGS topographic map (7.5-minute quadrangle), 

locality description. The omitted locality descriptions are in 

Oliver, 1976 (p. 140-141). 

108, Buffalo NE quad. 

828 Shorter Contributions to Paleontology and Stratigraphy, 1992 

124, East Springfield quad.; quarry, 0.6 km northnortheast of highway intersection in Springfield 

Four Corners. 

137, Cobleskill quad.; abandoned quarry on east side 

of Cobleskill. 

154, Sharon Springs quad. 

164, Sprout Brook quad. 

206, East Springfield quad. 

286, Gallupville quad. 

338, Buffalo NE quad.; roadcut southeast of Bennett 

High School, Buffalo. 

388, Akron quad.; large quarry in Akron, south of 

Central School. 

Hamilton Group Localities 

Most of the specimens described in this paper are 

from various members of the formations of the Hamilton 

Group in New York. In the following list, collections are 

grouped by members, which are listed from oldest to 

youngest. Each description includes collection number 

(from the USGS Silurian-Devonian catalog), name of 

USGS topographic map (7 .5-minute quadrangles), and the 

locality description. 

Skaneateles Formation, Delphi Station Member, Delphi 

Coral Bed 

One or more 15-cm-thick coral beds occur in the 

siltstones of the upper part of the Delphi Station Member. 

Four exposures are known, all within 10 km of each other. 

Oliver (1951, p. 719-721) described the southernmost 

outcrop as the Delphi coral bed in the Delphi (now Delphi 

Station) Member. The more northerly outcrops were called 

to my attention by J.C. Brower, Syracuse University (oral 

commun., 1976) and were recently cited by Brower and 

Nye (1991, p. 73; localities 9, 10, and 12). The beds in the 

four outcrops are very similar, and all seem to contain the 

same species of corals; they may or may not represent a 

single, continuous bed and be at the same horizon. However, the age difference, if any, is insignificant, and the 

corals are early Givetian in age. Corals are described from 

two of the outcrops: 

9132-SD, De Ruyter quad.; small outcrop beside 

private farm road, 0.95 km south, and 2.6 

km west of north and east edges of quadrangle. Delphi coral bed locality of Oliver, 

1951 (p. 719). 

9745-SD, Oran quad.; quarry on east side of road, 

1.6 km north of Highway 20 at 1240 

intersection, and 2 km northeast of Pompey Center. Locality 9 of Brower and Nye 

(1991' p. 73). 
Ludlowville Formation, Centerfield Member 

The Centerfield Member is dated by conodonts as 

Lower varcus Zone, Givetian age (Kirchgasser and others, 

1985). Thickness of the Centerfield coral bed is 1 to 2 m in 

the Genesee Valley. Localities are listed from west to east. 

5705-SD, Alexander quad.; Murder Creek, just 

west of Harper Road, 2.0 km north of 

Darien. 

4678-SD, Batavia South quad.; creek bed and bank 

0.1 km north of D., L. and W. Railroad 

tracks, 0. 6 km east of Francis Road and 

2.9 km east of East Alexander. 

11199-SD, Batavia South quad.; in railroad cut, 

approximately 0.2 km east of locality 

4678. 

5716-SD, Batavia South quad.; railroad cuts east of 

Center Road and 3. 2 km north of Bethany Center. 

5692-SD, Stafford quad.; White Creek, south of 

D., L. and W. Railroad, 0.4 to 0.5 km 

west of Transit Road, and 1.8 km east of 

East Bethany. 

5324a-SD, Geneseo quad.; northwest corner of 

quadrangle; Browns Creek, 0.2 to 0.3 km 

west of G. and W. Railroad and approximately 1.0 km east of York (Leicester 

quad.). 

5714-SD, same locality as 5324a, collected in different year. 

11197-SD, same locality as 5324a, collected in different year. 

5686-SD, Canandaigua quad.; Schaeffer Creek, 1.4 

km north of Centerfield, just north of 

point where tributary enters from west. 

5682-SD, Geneva South quad.; east shore of Seneca Lake near south edge of quadrangle, 

just north of mouth of Wilcox (Wilson) 

Creek. 

Ludlowville Formation, Otisco Shale Member, Staghorn 

Point Coral Bed 

The Staghorn Point coral bed was described by Oliver 

(1951). It extends some 24 km east-west and is known from 

more than 60 exposures; thickness ranges from a few 

centimeters near the margins to a maximum of 5. 6 m. The 

finest outcrops are along the east shore of Skaneateles Lake 

(Spafford quadrangle), where exposures are nearly continuous for more than 2 km. The museum specimens labeled 

"Skaneateles Lake" almost certainly came from this bed. 

The newer material described is from collections made at 

Willow Point, near the south end of the exposure, and 

Staghorn Point, some 0.6 to 0. 7 km to the northwest. 

7542-SD, Spafford quad.; Willow Point, specimens 

collected loose. 

7543-SD, Spafford quad.; Willow Point, specimens 

collected in place. 

8763-SD, Spafford quad.; Staghorn Point. 

8781-SD, Spafford quad.; Staghorn Point (collected 

in different year). 

Moscow Formation, Windom Shale Member, Fall Brook 

Coral Bed 

The Fall Brook coral bed was named and described 

by Baird and Brett (1983). All Fall Brook corals described 

in this paper are from a 30- to 40-cm-thick interval at one 

locality, Little Beards Creek; the five collections were made 

at different times or in different ways. Corals collected 

loose are almost certainly from the coral bed because 

similar corals are not known in the overlying and underlying 

shales. See Brett and Baird (1983, p. 163-165) for locality 

description. The conodont age of the unit is Middle varcus 

Zone (Kirchgasser and others, 1985, p. 250), Givetian. 

5700-SD, Leicester quad.; cliff on east side of 

Little Beards Creek, east and southeast of 

end of private road east of N.Y. Highway 

36, 1. 3 km north of Leicester. Specimens 

collected loose. 

570 1-SD, same locality, but specimens collected in 

place. 

8770-SD, same locality, specimens collected in 

place. 

11190-SD, same locality, most specimens collected 

in place. 

11476-SD, same locality, most specimens collected 

loose. 

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2, Palaeontology, p. 181-242. 

O'Connell, Marjorie, 1914, Revision of the genus Zaphrentis: 

New York Academy of Sciences Annals, v. 23, p. 177-192. 

Oliver, W.A., Jr., 1951, Middle Devonian coral beds of central 

New York: American Journal of Science, v. 249, p. 

705-728, 2 pls. 

---1956, Biostromes and bioherms of the Onondaga Limestone in eastern New York: New York State Museum Circular 

45, 23 p. 

---1958, Significance of external form in some Onondagan 

rugose corals: Journal of Paleontology, v. 32, p. 815-837, 

pis. 104-106. 

---1960a, Rugose corals from reef limestones in the Lower 

Devonian of New York: Journal of Paleontology, v. 34, p. 

59-100, pis. 13-19. 

---1960b, Devonian rugose corals from northern Maine: U.S. 

Geological Survey Bulletin 1111-A, 23 p., 5 pls. 

---1963, A new Kodonophyllum and associated rugose corals 

from the Lake Matapedia area, Quebec: U.S. Geological 

Survey Professional Paper 430-C, p. 21-31, pls. 9-14. 

---1967, Stratigraphy of the Bois Blanc Formation in New 

York: U.S. Geological Survey Professional Paper 584-A, 

8 p. 

---1976, Noncystimorph colonial rugose corals of the Onesquethaw and lower Cazenovia stages (Lower and Middle 

Devonian) in New York and adjacent areas: U.S. Geological 

Survey Professional Paper 869, 156 p., 108 pls. 

---1977, Biogeography of Late Silurian and Devonian rugose 

corals: Palaeogeography, Palaeoclimatology, Palaeoecology, 

v. 22, p. 85-135. 

---1980, Corals in the Malvinokaffric Realm: Miinstersche 

Forschungen zur Geologie und Palaontologie, v. 52, p. 

13-27. 

---1987, Middle Devonian coral faunules from Illinois and 

their bearing on biogeography: U.S. Geological Survey 

Bulletin 1690-D, 9 p., 6 pis. 

---1989, An early Middle Devonian coral faunule from the 

Needmore Shale in south-central Pennsylvania and adjacent 

areas of West Virginia and Virginia: U.S. Geological Survey 

Bulletin 1860-C, 11 p., 7 pls. 

---1990, Extinctions and migrations of Devonian rugose 

corals in the Eastern Americas Realm: Lethaia, v. 23, p. 

167-178. 

Oliver, W.A., Jr., and Pedder, A.E.H., 1979, Rugose corals in 

Devonian stratigraphical correlation: Palaeontological Association (London), Special Papers in Palaeontology, no. 23, p. 

233-248. 

---1989, Origins, migrations, and extinctions of Devonian 

Rugosa on the North American plate: Association of Australasian Palaeontologists Memoir 8, p. 231-237. 

Owen, D.D., 1847, Review of the New York geological reports: 

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Pedder, A.E.H., 1983, Systematic paleontology, in Pedder, 

A.E.H., and Goodbody, Q.H., New Devonian rugose corals 

of probable late Dalejan age from the Bird Fiord Formation of 

southwestern Ellesmere Island, Northwest Territories: Geological Survey of Canada Paper 83-1B, p. 337-349. 

Rafinesque, C.S., and Clifford, J.D., 1820, Prodrome d'une 

monographie des Turbinolies fossiles du Kentucky (dans 

1' Ameriq. septentr.): Annales Generales des Sciences Physiques, Bruxelles, p. 231-235. 

Rickard, L.V., and Zenger, D.H., 1964, Stratigraphy and paleontology of the Richfield Springs and Cooperstown quadrangles, New York: New York State Museum Bulletin 396, 101 

p., map. 

Rominger, Carl, 1876, Fossil corals: Michigan Geological Survey, Lower Peninsula, v. 3, pt. 2, Palaeontology, 161 p., 55 

pis. 

Scrutton, C.T., 1973, Palaeozoic coral faunas from Venezuela. II, 

Devonian and Carboniferous corals from the Sierra de Perija: 

British Museum (Natural History) Geology Bulletin, v. 23, 

no. 4, p. 223-281, 10 pls. 

Shimer, H.W., and Shrock, R.R., 1944, Index fossils of North 

America: New York, John Wiley, 837 p., 303 pis. 

Simpson, G.B., 1900, Preliminary descriptions of new genera of 

Paleozoic rugose corals: New York State Museum Bulletin 

39, p. 199-222. 

The Siphonophrentidae (Rugose Corals, Devonian) of Eastern North America 831 
Smith, Stanley, and Thomas, H.D., 1963, On Amplexus coralloides Sowerby and some ampleximorph corals from the 

English Devonian: Annals and Magazine of Natural History, 

ser. 13, v. 6, p. 161-172, pls. 7-9. 

Stewart, G .A., 1927, Fauna of the Silica Shale of Lucas County: 

Ohio Geological Survey, ser. 4, Bulletin 32, 76 p., 5 pls. 

---1938, Middle Devonian corals of Ohio: Geological Society 

of America Special Paper 8, 120 p., 20 pls. 

Stumm, E. C., 1937, The lower Middle Devonian tetracorals of the 

Nevada Limestone: Journal of Paleontology, v. 11, p. 

423-443, pls. 53-55. 

---1942, Fauna and stratigraphic relations of the Prout Limestone and Plum Brook Shale of northern Ohio: Journal of 

Paleontology, v. 16, p. 549-563, pls. 80-84. 

---1949, Revision of the families and genera of the Devonian 

tetracorals: Geological Society of America Memoir 40, 92 p., 

25 pls. 

---1965, Silurian and Devonian corals of the Falls of the 

Ohio: Geological Society of America Memoir 93, 184 p., 80 

pls. (Title page dated 1964 but published January 27, 1965.) 

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coral faunas of northeastern British Columbia: Geological 

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light of their minute skeletal structures: Royal Society of 

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832 Shorter Contributions to Paleontology and Stratigraphy, 1992 

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evolution and biogeography of the Palaeozoic corals of 

China: Beijing, Science Press, 391 p., 81 pls. (In Chinese 

with English summary.) 

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western Canada: Geological Association of Canada Special 

Paper 1, 15 p., 29 pls. 

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corals of the Falls of the Ohio: Washington University 

Studies, new ser. , Science and Technology 7, p. 113-122. 

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Part 3, Beginning with the Oriskany Sandstone: American 

Museum of Natural History Bulletin 11, p. 189-356. 

Willard, Bradford, Swartz, F.M., and Cleaves, A.B., 1939, The 

Devonian of Pennsylvania: Pennsylvania Geological Survey, 

ser. 4, Bulletin G-19, 481 p., 40 pls. 

Winchell, Alexander, 1886, Geological studies: Chicago, S.C. 

Griggs, 513 p. 

Yandell, L.P., and Shumard, B.F., 1847, Contributions to the 

geology of Kentucky: Louisville, 36 p. 

Yu Chang-min and Kuang Guo-dun, 1982, Biostratigraphy, biogeography and paleoecology of Devonian rugose corals from 

the Beiliu Formation in Beiliu, Guangxi: Nanjing Institute of 

Geology and Palaeontology Bulletin 5, p. 41-82, 7 pls. (In 

Chinese with English summary.) 

Yu Chang-min and Liao Wei-hua, 1978, Lower Devonian rugose 

corals from Alengchu of Lijiang, northwestern Yunnan: Acta 

Palaeontologica Sinica, v. 17, p. 245-266, 3 pls. (In Chinese 

with English summary.) 
PLATES 1-22 

Transverse sections are oriented with the cardinal position (or what I interpret to be the cardinal 

position) toward the top of the plate and are photographed from the top, that is, from the calicinal 

end of the coral. Most of the longitudinal sections are in the plane of curvature, rather than the 

cardinal-counter plane. Sections are thin sections unless otherwise indicated. Depository abbreviations used in the plate captions are explained in the text under "Depositories." Contact photographs 

of the plates in this report are available, at cost, from U.S. Geological Survey Photographic Library, 

Federal Center, Denver, CO 80225. 
PLATE 1 

[Exteriors, x 1] 

Figures 1-4. Heterophrentis spatiosa (Billings) (p. B5). 

1, 2. Lectotype, GSC 3451. See also pl. 2, fig. 2. 

3, 4. Paralectotype, GSC 3451a. See also pl. 2, fig. 1. 

Port Colbome, Ontario; Bois Blanc Formation or Onondaga Limestone. 

5-7. Triplophyllum terebratum (Hall) (p. B7). 

Holotype, NYSM 341. Jeffersonville Limestone, Falls of the Ohio, near Louisville, Ky. 

8-15. Metaxyphrentis prolifica (Billings) (p. BlO). 

8. Paralectotype, GSC 3449i. 

9-12. Lectotype, GSC 3449h. See also pl. 2, figs. 3-5. 

13-15. Paralectotype, GSC 3449k. 

Port Colbome, Ontario; Bois Blanc Formation or Onondaga Limestone. 

16--18. Kionelasma? sp. cf. Heterophrentis spatiosa (Billings) (p. B6). 

USNM 455880. Calice (cardinal septum down) and side views of specimen with boss and distally spatulate 

septa as in H. spatiosa. Bois Blanc Formation, Town of Clarence, N.Y.; collection USGS 4672-SD. See 

Oliver, 1967, p. 7, for location. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 1 

HETEROPHRENTIS, TRIPLOPHYLLUM, METAXYPHRENTIS, AND KIONELASMA? 
PLATE 2 

Figures 1, 2. Heterophrentis spatiosa (Billings) (p. B5). 

1. Paralectotype, GSC 3451a; longitudinal section, x 1.5. See also pl. 1, figs. 3, 4. 

2. Lectotype, GSC 3451; transverse section, x 1.5. See also pl. 1, figs. 1, 2. 

Port Colbome, Ontario; Bois Blanc Formation or Onondaga Limestone. 

3-10. Metaxyphrentis prolifica (Billings) (p. B10). 

3-5. Lectotype, GSC 3449h; transverse section, x 2, and cardinal and counter areas of same, x 10. See also 

pl. 1, figs. 9-12. Port Colbome, Ontario; Bois Blanc Formation or Onondaga Limestone. 

6-10. USNM 454008, X 2. Transverse sections and one longitudinal section numbered in order from 

uppermost (just below calice) to lowermost; longitudinal section is through fossula (to left). Edgecliff 

Member, Onondaga Limestone, New York (loc. 154). 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 2 

9 

HETEROPHRENTIS AND METAXYPHRENTIS 
PLATE 3 

Metaxyphrentis prolifzca (Billings) (p. B 10). 

[All figures X 2] 

Figures 1, 2. USNM 454009. Longitudinal section (including cal ice) and immediately underlying transverse section. 

Locality 154. 

3. USNM 454006. Transverse section from just below calice. Locality 137. 

4--6. USNM 454003. Transverse and longitudinal sections arranged in descending order. Locality 108. 

7-9. USNM 454004. Longitudinal section through calice and two lower transverse sections. Locality 286. 

10. USNM 454007. Longitudinal section including base of calice. Locality 124. 

11-13. USNM 454005. Longitudinal section (including calice) and two lower transverse sections. Locality 338. 

All specimens from Edgecliff Member, Onondaga Limestone, New York. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 3 

METAXYPHRENTIS 
PLATE 4 

Compressiphyllum davisana (Miller) (p. B9). 

[Figures x 2 except as noted] 

Figures 1, 2. USNM 453997. Transverse ( x 2) and long-diameter longitudinal ( x 1.5; cardinal fossula to left) sections. 

3, 4. USNM 453998. Long-diameter longitudinal (fossula to right) and transverse sections. 

5-7. USNM 453996. Transverse sections from just above and below long-diameter longitudinal section (fossula to 

left). Collection USGS 4723-SD, Jeffersonville Limestone, Aemulophyllum exiguum Zone, Falls of the Ohio. 

8, 9. USNM 453999. Transverse sections just below base of calice and approximately 1.5 em lower (showing 

particularly well developed siphonofossula). 

10-12. Short -diameter longitudinal sections. 

10. USNM 454000. 

11. USNM 454001. 

12. USNM 454002. 

All specimens from Jeffersonville Limestone, coral zone (probably lower part, Emsian), Falls of the Ohio, near 

Louisville, Ky. See more definitive data for USNM 453996. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 4 

COMPRESSIPHYLLUM 
PLATE 5 

Enallophrentis simplex (Hall) (p. B 12). 

[Exteriors, x 1] 

Figures 1--4. Holotype, NYSM 360. See also pl. 6, figs. 1, 2. The specimen was illustrated by Hall (1843) and was cut prior 

to 1877 when the polished transverse section was illustrated; the upper part of the specimen is apparently lost. 

5, 6. AMNH 39407, the original of Hall, 1877, pl. 21, figs. 6, 7. See also pl. 6, figs. 5, 6. 

7-9. NYSM 352, lectotype of Zaphrentis amp/a Hall, 1877, pl. 21, figs. 2, 3. See also pl. 6, fig. 3. York, N.Y., 

probably the Centerfield Member, Ludlowville Formation. 

10, 11. NYSM 6266, the original of Hall, 1877, pl. 21, fig. 5. See also pl. 6, fig. 7. 

12-14. AMNH 39409, the original of Hall, 1877, pl. 21, fig. 10. 

15, 16. AMNH 39408, the original of Hall, 1877, pl. 21, fig. 9. 

17-19. NYSM 6267, the original of Hall, 1877, pl. 21, fig. 11. See also pl. 6, figs. 13, 14. 

See text discussion of the provenance of Hall's specimens. Except as noted for NYSM 352, all are from the Genesee 

Valley and probably from the Moscow Formation near Leicester, N.Y. The crosses and grids seen in several of the 

figures were scratched on the specimens as a guide to Hall's delineator (G.B. Simpson). 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 5 

ENALLOPHRENTIS 
PLATE 6 

Enallophrentis simplex (Hall) (p. B 12). 

[All figures x 2] 

Figures 1, 2. Holotype, NYSM 360; transverse polished section (the same as is shown on pl. 5, fig. 1) and thin section (taken 

approximately 1.5 em lower). See also pl. 5, figs. 1-4. 

3. NYSM 352, lectotype of Zaphrentis ampla Hall; transverse section. See also pl. 5, figs. 7-9. 

4. NYSM 351, transverse section of paralectotype of Z. ampla Hall, 1877, pl. 21, fig. 1. Darien, N.Y. 

5, 6. AMNH 39407; transverse section and lower longitudinal section. See also pl. 5, figs. 5, 6. 

7. NYSM 6266; transverse section below calice. See also pl. 5, figs. 10, 11. 

8, 9. USNM 454017. Transverse section and underlying longitudinal section. Collection USGS 5700-SD, Fall 

Brook coral bed, Leicester, N.Y. 

10-12. USNM 454024. Transverse sections from immediately above and below the longitudinal section. Collection 

USGS 8770-SD, Fall Brook coral bed, Leicester, N.Y. 

13, 14. NYSM 6267; transverse and lower longitudinal sections. See also pl. 5, figs. 17-19. 

See text discussion of the provenance of the Hall specimens (AMNH and NYSM numbers). Except as noted for NYSM 

352 and 351, all are from the Genesee Valley and probably from the Moscow Formation near Leicester, N.Y. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 6 

3 

6 

12 14 

ENALLOPHRENTIS 
PLATE 7 

Enallophrentis simplex (Hall) (p. B 12). 

[All figures x 2] 

Figures 1-4. USNM 454018. Transverse and longitudinal sections of gerontic individual, numbered in descending order; 1 

and 2 show gerontic characters, 3 and 4 are neanic. Collection USGS 5700-SD. 

5-7. USNM 454025. Transverse sections taken immediately above and below longitudinal section. Collection 

USGS 8770-SD. 

8-10. USNM 454026. Transverse sections taken immediately below and above longitudinal section. Collection 

USGS 8770-SD. 

11. USNM 454027. Longitudinal section of lower middle part of coral. Collection USGS 8770-SD. 

12, 13. USNM 454019. Transverse section and longitudinal section immediately below. Collection USGS 5700-SD. 

All specimens from Fall Brook coral bed, Windom Shale Member, Moscow Formation; Little Beards Creek near 

Leicester, N.Y. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 7 

9 

ENALLOPHRENTIS 
PLATE 8 

Enallophrentis simplex (Hall) (p. B 12). 

[All figures x 2] 

Figures 1--4. USNM 454031. Transverse and longitudinal sections numbered in descending order. Collection USGS 

11476-SD. 

5-7. Longitudinal sections. 

5. USNM 454028; gerontic?; collection USGS 8770-SD. 

6. USNM 454021; collection USGS 5701-SD. 

7. USNM 454029; gerontic; collection USGS 8770-SD. 

8-11. USNM 454020. Transverse and longitudinal sections numbered in descending order. Collection USGS 

5701-SD. 

12. USNM 454022. Longitudinal section of middle part of coral. Collection USGS 5701-SD. 

13-15. USNM 454023. Transverse sections taken immediately above and below longitudinal section. Collection 

USGS 5701-SD. 

16. USNM 454030. Longitudinal section of upper middle part of gerontic individual. Collection USGS 8770-SD. 

All specimens from Fall Brook coral bed, Windom Shale Member, Moscow Formation; Little Beards Creek near 

Leicester, N.Y. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024---B PLATE 8 

4 

14 16 

ENALLOPHRENTIS 
PLATE 9 

Enallophrentis simplex (Hall) (p. B 12). 

[All figures x 2] 

Figures 1--4. USNM 454038. Transverse and longitudinal sections numbered in descending order; 1 is through calice and 

shows gerontic characters of septa; 2 is immediately below and is mostly gerontic; 3 and 4 are through ephebic 

and neanic parts of coral. 

5. USNM 454839. Transverse section. 

6-8. USNM 454040. Longitudinal section through calice and lower transverse sections. 

9-11. USNM 454041. Transverse sections taken immediately above and below longitudinal section. 

All specimens from collection USGS 5686-SD, Centerfield Member, Ludlowville Formation; Schaeffer Creek, just 

north of Centerfield, N.Y. 

PLATE 10 

Enallophrentis spp. 

[Figures X 2 unless otherwise indicated] 

Figures 1-7. E. simplex (Hall) (p. B12). 

1-3. USNM 454042. Transverse sections immediately above and below longitudinal section in gerontic 

individual; 3 and the lower part of 2 are in ephebic part. Collection USGS 5692-SD. 

4, 5. USNM 454043. Transverse section and lower longitudinal section. Collection USGS 5692-SD. 

6, 7. USNM 454044. Longitudinal section and overlying transverse section. Collection USGS 4678-SD. 

Specimens shown in figures 1-7 from Centerfield Member, Ludlowville Formation, N.Y. 

8-10. E. broweri n. sp. (p. B16). 

USNM 454102. Longitudinal section ( x 1.5) and two half transverse sections; longitudinal section is in plane 

of curvature; transverse sections show that the cardinal-counter plane is consistently different. Collection 

USGS 9132-SD, Delphi coral bed, Skaneateles Formation, New York. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024--B PLATE 10 

5 6 

ENALLOPHRENTIS 
PLATE 11 

Enallophrentis broweri n. sp. (p. B16). 

[All figures x 2] 

Figures 1-4. Holotype, USNM 454068. Transverse and longitudinal sections numbered in descending order. 

5. Paratype, USNM 454070. Transverse section through lower middle part of corallum. 

6-9. Paratype, USNM 454069. Longitudinal and transverse sections numbered in descending order; fig. 6 shows the 

lower part of the calice. 

10-12. Paratype, USNM 454071. Transverse sections from just above and below the longitudinal section. 

13-15. Paratype, USNM 454072. Transverse sections from just above and below the longitudinal section. 

All specimens from collection USGS 9745-SD, Delphi coral bed, Pompey Center, N.Y. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 11 

15 
PLATE 12 

Enallophrentis broweri n. sp. (p. B16). 

[All figures x 2] 

Figures 1-3. Paratype, USNM 454073. Transverse sections from just above and below the longitudinal section; numbered 

in descending order. 

4-6. Paratype, USNM 454074. Transverse sections from just above and below the longitudinal section; numbered 

in ascending order. 

7-9. Paratype, USNM 454075. Transverse section through the lower calice, longitudinal section including lowest 

calice, and underlying transverse section. 

10-12. Paratype, USNM 454076. Transverse section from between the two longitudinal sections (oriented to appear 

to have been photographed from the same side). 

13-15. Para type, USNM 454077. Transverse sections from just above and below the longitudinal section; numbered 

in descending order. 

16. Paratype, USNM 454078. Longitudinal section through upper middle part of corallum. 

All specimens from collection USGS 9745-SD, Delphi coral bed, Pompey Center, N.Y. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 12 

ENALLOPHRENTIS 
PLATE 13 

Breviphrentis invaginata (Stumm) (p. B19). 

[All figures x 2] 

Figures 1, 2. Lectotype, USNM 94443; thin sections cut from the original of Stumm, 1937, pl. 53, fig. 2 (labeled 

"holotype"), and illustrated by Hill, 1981, fig. 95:1a,b. See text discussion; this is not the same specimen as 

the original of Stumm, 1937, pl. 54, fig. 2a, also labeled "holotype." 

3. Paralectotype, USNM 454103, the original of Stumm, 1937, pl. 54, fig. 2a (labeled "holotype"). 

4, 5. USNM 94442, holotype of Heterophrentis nevadensis Stumm, 1937; the transverse section is the original of 

his pl. 54, fig. 1a; the longitudinal section was prepared later. 

6-16. Specimens of Merriam, 1974b, pls. 15 and 16; Merriam illustrations are cited by plate and figure numbers; see 

synonymy and discussion in text. 

6-8. USNM 159306, 16, 5-7. 

9, 10. USNM 159297, 15, 3, 4. 

11. USNM 159302, 15, 11. 

12. USNM 159311, 16, 15, 16. 

13. USNM 159300, 15, 9. 

14. USNM 159305, 16, 4. 

15. USNM 159304, 16, 3. 

16. USNM 159307, 16, 8. 

All specimens from Nevada Formation, Eurekaspirifer pinyonensis Zone, Emsian, Nevada. Merriam's (1974b) thin 

sections are reillustrated because of the importance of the species in defining this genus of the Eastern Americas Realm 

and in understanding the morphology of the family. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 13 

6 

7 

BREVIPHRENTIS 
PLATE 14 

Breviphrentis spp. 

[All figures x 1 unless otherwise indicated] 

Figures 1-6. B. halli (Milne-Edwards and Haime) (p. B20). 

1-3. Holotype, EM 15173. Side (X 0.8) and calice views. See also pl. 15, figs. 1, 2. 

4, 5. AMNH 39405, the original of Hall, 1877, pl. 20, fig. 1. Side view and broken transverse section 

showing an axially flat tabula and siphonofossula. 

6. AMNH 39404, the original of Hall, 1877, pl. 20, fig. 8. Broken longitudinal section showing closely 

spaced, axially flat tabulae. 

7-9. B. cista n. sp. (p. B21). 

7. Holotype, NYSM 6264, original of Hall, 1877, pl. 20, fig. 6. Front view, X 0.7; see also pl. 17, figs. 

1, 2. 

8, 9. Paratype, NYSM 6265, original of Hall, 1877, pl. 20, fig. 7. Calice (X 0.9) and side (X 0.8) views; 

see pl. 17, figs. 11, 12. 

All specimens from Skaneateles Lake, N.Y., presumably the Staghorn Point coral bed (see discussion in text). The 

crosses scratched on the specimens shown in figures 4 and 7 were placed there as a guide to Hall's delineator (G.B. 

Simpson). 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 14 

BREVIPHRENTIS 
PLATE 15 

Breviphrentis halli (Milne-Edwards and Haime) (p. B20). 

[All figures x 1.5] 

Figures 1, 2. Holotype, EM 15173. Transverse section taken just below longitudinal section; see also pl. 14, figs. 1-3. 

3-5. NYSM 6262. Transverse sections taken just above and below the longitudinal section. Specimen illustrated by 

Hall, 1877, pl. 20, fig. 3. 

6-8. USNM 454107. Transverse sections taken immediately above and below longitudinal section. Collection 

USGS 7543- SD. 

9-11. USNM 454112. Transverse sections taken immediately above and below longitudinal section. Collection 

USGS 7542-SD. 

12-14. USNM 454111. Transverse sections taken immediately above and below longitudinal section. Collection 

USGS 8763-SD. 

All specimens from Skaneateles Lake, N.Y. , presumably the Staghorn Point coral bed (see discussion in text). 
U.S. GEOLOGICAL SURVEY BULLETIN 2024--B PLATE 15 

6 

7 

2 

9 

13 

BREVIPHRENTIS 
PLATE 16 

Breviphrentis halli (Milne-Edwards and Haime) (p. B20). 

[All figures X 1.5] 

Figures 1-5. USNM 454105. Transverse and longitudinal sections in descending order. Collection USGS 7543-SD. 

6, 7. USNM 454106. Transverse section from just above longitudinal section. Collection USGS 7543-SD. 

8, 9. USNM 454109. Transverse section from just above longitudinal section. Collection USGS 7542-SD. 

10-12. USNM 454110. Longitudinal sections from just above and below the transverse section; the upper longitudinal 

includes the lower part of the calice. Collection USGS 7543-SD. 

13-15. USNM 454108. Transverse sections from just above and below longitudinal section. Collection USGS 

7542-SD. 

All specimens from the Staghom Point coral bed, Otisco Shale Member, Ludlowville Formation; Skaneateles Lake, 

N.Y. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 16 

2 

3 

BREVIPHRENTIS 
PLATE 17 

Breviphrentis cista n. sp. (p. B21). 

[All figures x 1.5] 

Figures 1, 2. Holotype, NYSM 6264. Transverse section taken just above longitudinal section. See also pl. 14, fig. 7. 

3, 4. Paratype, USNM 454125. Transverse section taken just above longitudinal section. Collection USGS 

7542-SD. 

5. Paratype, USNM 454126. Longitudinal section. Collection USGS 7542-SD. 

6, 7. Paratype, NYSM 6263. Transverse section taken just below longitudinal section, which includes the lowest 

part of the calice. Specimen illustrated by Hall, 1877, pl. 20, fig. 4. 

8-10. Paratype, USNM 454127. Transverse sections from just above and below longitudinal section. Collection 

USGS 7542-SD. 

11, 12. Paratype, NYSM 6265. Transverse section taken just above longitudinal section. See also pl. 14, figs. 8, 9. 

All specimens from Skaneateles Lake, N.Y., presumably the Staghorn Point coral bed (see discussion in text). 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 17 

9 

BREVIPHRENTIS 
PLATE 18 

Breviphrentis cista n. sp. (p. B21). 

[All figures x 1.5] 

Figures 1-3. Paratype, USNM 454128. Transverse and longitudinal sections; upper transverse section from just above 

longitudinal section; lower transverse section approximately 1.5 em below the longitudinal section. 

4-6. Paratype, USNM 454129. Transverse sections from just above and below longitudinal section. 

7-9. Paratype, USNM 454130. Transverse sections from just above and below longitudinal section. 

10-12. Paratype, USNM 454131. Transverse sections from just above and below longitudinal section. Collection 

USGS 7543-SD. 

13-15. Paratype, USNM 454132. Transverse sections from just above and below longitudinal section. 

16-18. Paratype, USNM 454133. Transverse sections from just above and below longitudinal section. 

19-21. Paratype, USNM 454134. Transverse sections from just above and below longitudinal section. 

All specimens from the Staghom Point coral bed, Otisco Shale Member, Ludlowville Formation; Skaneateles Lake, 

N.Y. Collection USGS 7542-SD, except as noted. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 18 

11 

20 

12 21 

BREVIPHRENTIS 
PLATE 19 

Breviphrentis pumilla n. sp. (p. B24). 

[All figures x 2] 

Figures 1-3. Holotype, USNM 454154. Transverse sections from just above and below longitudinal section. 

4, 5. Paratype, USNM 454155. Transverse section from just above longitudinal section. 

6--8. Paratype, USNM 454156. Transverse sections approximately 3 mm apart and lower longitudinal section; 

uppermost section in lower calice. 

9-11. Para type, USNM 45415 8. Transverse sections from just above and below longitudinal section. 

12-14. Paratype, USNM 454157. Transverse sections approximately 3 mm apart and lower longitudinal section. 

15, 16. Paratype, USNM 454159. Transverse section from just above longitudinal section. 

17, 18. Paratype, USNM 454161. Transverse section from just above longitudinal section. 

19-21. Paratype, USNM 454167. Transverse sections from just above and below longitudinal section. Collection 

USGS 11197-SD. 

22, 23. Paratype, USNM 454160. Transverse section from just above longitudinal section. 

24, 25. Paratype, USNM 454162. Longitudinal section from just above transverse section. 

26, 27. Paratype, USNM 454163. Transverse section from just above longitudinal section. 

28, 29. Paratype, USNM 454166. Longitudinal section from just below transverse section. Collection USGS 

5692-SD. 

30, 31. Paratype, USNM 454168. Longitudinal section from just above transverse section. Collection USGS 

5705-SD. 

32. Paratype, USNM 454164. Longitudinal section. 

33-35. Paratype, USNM 454165. Longitudinal section, transverse section from just below longitudinal, and second 

transverse section (approximately 12 mm lower). Collection USGS 5692-SD. 

All specimens from the Centerfield Member, Ludlowville Formation. Collection USGS 4678-SD, unless otherwise 

indicated. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 19 

30 21 

BREVIPHRENTIS 
PLATE 20 

Siphonophrentis elongata (Rafinesque and Clifford) (p. B26). 

[All figures x 1.5] 

Figures 1--4. USNM 454181, neotype of Siphonophrentis gigantea (Lesueur), type species of genus. Transverse and 

longitudinal sections numbered in descending order. Onondaga Limestone, Leroy, N.Y. See discussion in text. 
BULLETIN 2024-B PLATE 20 

SIPHONOPHRENTJS 
PLATE 21 

Siphonophrentis elongata (Rafinesque and Clifford) (p. B26). 

[All figures x 1.5] 

Figures 1--4. UMMP 8616, neotype, selected by Stumm, 1965; thin sections cut from specimen illustrated by Rominger, 

1876, and Stumm, 1965 (see discussion in text). Sections numbered in descending order. 

5, 6. UMMP 26423, illustrated by Stumm, 1965, pl. 13, figs. 3, 4. Longitudinal section taken just below transverse 

section. 

Both specimens from Jeffersonville Limestone, Falls of the Ohio, near Louisville, Ky. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 21 

2 

SIPHONOPHRENTIS 
PLATE 22 

Siphonophrentis elongata (Rafinesque and Clifford) (p. B26). 

[Sections x 1] 

Figures 1-7. USNM 454182. 

1-6. Polished sections numbered in descending order; transverse sections are consistently oriented to show how the 

position of septal dilation changes along the length of the coral; the longitudinal section is in or close to the 

cardinal-counter plane with the cardinal fossula to the right. 

7. Specimen before cutting, x 0.4 (length, 73 em); the positions of the polished sections are labeled by the 

corresponding figure numbers; the cardinal fossula is on the left. 

Jeffersonville Limestone, Falls of the Ohio, near Louisville, Ky. 
U.S. GEOLOGICAL SURVEY BULLETIN 2024-B PLATE 22 

SIPHONOPHRENTIS 



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