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PALAEONTOGRAPHICA
AMERICANA

VOL. VI

1968-1970

PALEONTOLOGICAL RESEARCH INSTITUTION

ng a '

Le et ieee
nw it iy Ayla fy

CONTENTS OF VOLUME VI

NUMBER

38. Lycopsip STEMS AND ROOTs AND SPHENOPSID
FRUCTIFICATIONS AND STEMS FROM THE UPPER
FREEPORT COAL. OF SOUTHEASTERN OHIO
By Maxine L. Abbott

39. CENOZOIC EVOLUTION OF THE ALTICOSTATE VENERICARDS
IN GULF AND EAsT CoAsTAL NorTH AMERICA
By W. G. Heaslip

40. CARBONIFEROUS CRINOWS OF ‘TEXAS WITH
STRATIGRAPHIC IMPLICATIONS
By H. L. Strimple and W. T. Watkins

41. TRACE Fossits OF THE CINCINNATI AREA
By R. G. Osgood, Jr.

PLATES

1-19

20-29

50-56

PAGES

51-136

137-275

276-444

INDEX

No separate index is included for the volume. Each number is
indexed separately. Contents of the volume are listed in the beginning
of the volume.

Pon (5
Pi5

PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

LYCOPSID STEMS AND ROOTS AND
SPHENOPSID FRUCTIFICATIONS AND STEMS
FROM THE UPPER FREEPORT COAL OF
SOUTHEASTERN OHIO

By

MAXINE L. ABBOTT

1968

Paleontological Research Institution

Pa aMmtlTHSo Nia Ithaca, New York, U.S.A., 14850

PALEONTOLOGICAL RESEARCH INSTITUTION

1967 - 1968
PRESIDENT "5 sciseucss qattesnessasccsscksxavassansbasssucssvekscexasasasibhvedsnsaaseateisssaaaioys«at KENNETH E. CASTER
WICE=BRESIDE NT: assess sescesconteroshascspecuavuanesttansbcsneer ox ciegs ones teaeaswepewan saints WILLIAM B. HERoy
SECRETARY" REASUREBM \ ston scccisattestsssiahosssspeonsiitenvany souncetiansstvialtiacanasabine REBECCA S. HARRIS
DDERRCTOR: cos ccassscesseacesssscerosescessssxtcecsnetesbetervescesstssvncesessatenecnsates KATHERINE V. W. PALMER
MSO UENBRL: wepuccorsrene tay -asivensitonsaccuemeserontsaasanice rans isaenesart anata eee ae ARMAND L, ADAmMs
REPRESENTATIVE ALAA 'COUMNGID, | cccesscccccssctssossassasassartsersvavesassotsbaed KENNETH E, CASTER

Trustees

KENNETH E. CASTER (1966-1972) KATHERINE V. W. PALMER (Life)
Donato W. FIsHER (1967-1973) Wiuiam B. Heroy (1963-1968)
Resecca S. Harris (Life) Axe A. Oxsson (Life)
DanieL B. Sass (1965-1971) Hans G. KucLer (1963-1969)

W. Storrs Cote (1964-1970)

BULLETINS OF AMERICAN PALEONTOLOGY

and

PALAEONTOGRAPHICA AMERICANA

KATHERINE V. W. PALMER, Editor
Mrs. Fay Bricos, Secretary

Advisory Board

KENNETH E. CASTER HANS KUGLER
A. Myra KEEN Jay GLENN Marks
AXEL A. OLSSON

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PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

VOL. VI

NO. 38

LYCOPSID STEMS AND ROOTS AND
SPHENOPSID FRUCTIFICATIONS AND STEMS
FROM THE UPPER FREEPORT COAL OF
SOUTHEASTERN OHIO

By

MAXINE L. ABBOTT

March 4, 1968

Paleontological Research Institution

Ithaca, New York, U.S.A., 14850

Library of Congress Card Number: GS 68-130

Printed in United States of America
Arnold Printing Company

CONTENTS

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\

LYCOPSID STEMS AND ROOTS
AND
SPHENOPSID FRUCTIFICATIONS AND STEMS
FROM THE UPPER FREEPORT COAL
OF SOUTHEASTERN OHIO
MAxinE L. ABBotr
Western State College of Colorado

ABSTRACT

The Upper Freeport coal, uppermost Allegheny, in southeastern
Ohio, contains a shale parting which bears a diversified compression
flora. A portion of the flora consists of organ genera of | lycopsids
and sphenopsids and are herein described and figured by line draw-
ings and photographs. The lycopsid species are as follows: Lepidoden-
dron andrewsi Lesquereux, 1879-80, Lepidodendron cf. jaraczewskit
Zeiller, 1888, Lepidodendron aculeatum Sternberg, 1825, Lepidoden-
dron scutatum Lesquereux, 1879-80; Lepidophloios vaningent D.
White, 1900, Lepidophloios larcinus Sternberg, 1825; Asolanus cam pto-
taenia Wood, 1861; Sigillaria mamillaris Brongniart, 1836, Sigillaria
elegans (Sternberg), Brongniart, 1828, Sigillaria lacoei Lesquereux,
1879-80, Sigillaria laurenciana (Lesquereux ex Abbott), Sigillaria
rugosa Brongniart, 1828, Sigillaria orbicularis Lesquereux, 1879-80,
Sigillaria pileata Abbott, n. sp.; Stigmaria ficoides (Sternberg),
Brongniart, 1822. ,

The sphenopsid species, which include the fructifications of
Sphenophyllum and Calamites and the axes of Calamites, are as fol-
lows: Bowmanites incurvata Abbott, n. sp., Bowmanites magna (Les-
quereux? ex Abbott); Mazostachys compacta Abbott, n. sp., Calamo-
stachys tuberculata (Sternberg), Weiss, 1884, Calamostachys german-
ica Weiss, 1876, Calamostachys bisporangiata Abbott, n. sp., Cala-
mostachys minuta Abbott, n. sp. Calamostachys recurvata Abbott,
n. sp., Calamostachys longibracteata Abbott, n. sp., Calamostachys in-
terculata Abhott, n. sp., Palaeostachya elongata (Presl), Weiss, 1876,
Palacostachya ovalis (Lesquereux ex Abbott), Palacostachya trabe-
culata Abbott, n. sp.; Calamites undulatus Sternberg, 1825, Calamites
carinatus Sternberg, 1824, Calamites cisti Brongniart, 1828, Calamites
cistiiformis Stur, 1877, Calamites sachsi Stur, 1878, Calamites schiit-
zeiformis Kidston and Jongmans, 1913, Calamites suckowi Brongniatt,

op INTRODUCTION

The Upper Freeport, uppermost Allegheny, is more or
less continuous throughout the Appalachian Coal Basin.
One of the few areas of this coal, bearing an identifiable
compression flora, occurs near Kimberly, northern Athens
County, in southeastern Ohio. The coal was strip mined
over a period of approximately 10 years and resulted in
about 20 miles of essentially continuous exposures of the
coal. In this area, the diversified compression flora occurs
in a discontinuous shale parting in the coal. The shale part-
ing ranges from a few inches up to three feet in thickness.

The flora consists of organ genera of lycopsids,
sphenopsids, ferns, and pteridosperms. The species are so
numerous that monographic treatments of each of the
groups is necessary in order to: 1) review previously re-
ported North American species, many of which are inade-
quately described and figured; 2) to present additional new
material: 3) to describe new species; 4) to list the occur-
rence of the same species in floras of comparable age from
other coal basins of the United States and Canada. There-
fore, a few of the species discussed herein do not occur in
the Kimberly flora.

The present paper treats only the lycopsids and sphen-
opsids. The fern-pteridosperm complex will be reported at

a later date. The lycopsid flora is limited to stems and
roots, including species of Lepidodendron, Lepidophloios,
Sigillaria, Asolanus, and Stigmaria; the fructification genera
were reported previously (Abbott, 1963). The sphenopsid
flora considered, consists of fructifications and stems, in-
cluding species of Palaeostachya, Calamostachys, Mazo-
stachys, Bowmanites, and Calamites. The leaf genera were
reported in an earlier paper (Abbott, 1958) .

In order to obtain cellular detail of cone parts and
spores, transfer (Abbott, 1950) and modified maceration
techniques were used. Individual sporangia were isolated
and macerated in deep well slides and the resulting spores
and sporangial wall parts were permanently mounted in
permount.

In the synonymies only American species are consid-
ered because of the availability of the American types upon
which the original descriptions were based. I have not in-
cluded European species in the synonymies because I was
unable to study the type specimens.

All of the form genera considered in this study are
well established morphologically and taxonomically, there-
fore I have not repeated their definitions.

Lesquereux, in his Coal Flora, cited numerous fossil
plant localities in Pennsylvania. Some of the names of the
towns have been changed or are no longer in existence. In
order to relocate these localities I consulted maps dating
as early as 1858. The localities most often referred to and
thus effected are as follows:

Cited Location
Pittston

Present Location

—northeast of Wilkesbarre, Lu-
zerne County

—southwest of Scranton, Lacka-
wanna County

— Schuylkill County

—southwest of Scranton, Lacka-
wanna County

—about 12 miles northeast of
Scranton, Lackawanna County

—not located.

West Pittston

Pottsville
Archbald

Northern Anthracite
Coal Field
Alton

ACKNOWLEDGMENTS
This study was made possible through the National
Science Foundation grant numbers G9021 and G24501.
I wish to thank the directors and their personnel in
charge of the numerous paleobotanical collections for their

6 PALAEONTOGRAPHICA AMERICANA (VI, 38)

hospitality and generosity in making their specimens avail-
able for my studies. They are as follows: 1) Type speci-
mens are now deposited:

Geological Survey of Canada, Ottawa, Canada

The Smithsonian Institution, Washington, D.C.

Museum of Natural History, Springfield, Illinois

University of Ilinois and the Illinois Geological Sur-
vey, Urbana, Illinois

Harvard University, Cambridge, Massachusetts

University of Michigan, Ann Arbor, Michigan

Academy of Natural Sciences, Philadelphia, Pennsyl-
vania

University of Cincinnati, Cincinnati, Ohio;

2) Other specimens, from different localities, but of com-
parable age are deposited:
Princeton University, Princeton, New Jersey
Brown University, Providence, Rhode Island
University of West Virginia, Morgantown, West
Virginia
Michigan State University, East Lansing, Michigan
Ohio University, Athens, Ohio.

My thanks are also given to Mrs. Bettina Dalvé, Uni-
versity of Cincinnati, for inking my penciled drawings; to
Dr. Harry Muegel, Emeritus, University of Cincinnati, for
suggestions concerning new epithets as well as translations;
and to Dr. Francis Hueber, Smithsonian Institution, for
photographs of some of the type specimens.

CLASSIFICATION

Division Cormophyta
Subdivision Paleocormophyta
Class Lycopsida
Order Lepidophytales
Family Lepidodendraceae
Lepidodendron andrewsi Lesquereux
Lepidodendron cf. jaraczewskii Zeiller
Lepidodendron aculeatum Sternberg
Lepidodendron scutatum Lesquereux
Lepidophloios larcinus Sternberg
Lepidophloios vaningeni D. White
Asolanus sigillarioides (Lesquereux ex Abbott)
Asolanus camptotaenia Wood
Family Sigillariaceae
Sigillaria mamillaris Brongniart
Sigillaria elegans (Sternberg), Brongniart
Sigillaria lacoe: Lesquereux
Sigillaria laurenciana (Lesquereux ex Abbott)
Sigillaria rugosa Brongniart
Sigillaria orbicularis Lesquereux
Sigillaria pileata Abbott
Stigmaria ficoides (Sternberg) Brongniart
Class Sphenopsida
Order Sphenophyllales
Family Sphenophyllaceae
Bowmanites magna (Lesquereux? ex Abbott)
Bowmanites incurvata Abbott

Order Equisetales

Family Calamitaceae
Mazostachys compacta Abbott
Calamostachys tuberculata (Sternberg) Weiss
Calamostachys germanica Weiss
Calamostachys hisporangiata Abbott
Calamostachys minuta Abbott
Calamostachys recurvata Abbott
Calamostachys longibracteata Abbott
Calamostachys interculata Abbott
Palaeostachya elongata (Presl) Weiss
Palaeostachya ovalis (Lesquereux ex Abbott)
Palaeostachya aperta (Lesquereux ex Abbott)
Palaeostachya trabeculata Abbott
Calamites cistiiformis Stur
Calamites cisti Brongniart
Calamites carinatus Sternberg
Calamites suckowi Brongniart
Calamites schiitzeiformis Kidston and Jongmans
Calamites sachsi Stur
Calamites undulatus Sternberg

DESCRIPTION OF SPECIES
Genus LEPIDODENDRON Sternberg, 1820
Lepidodendron andrewsi Lesquereux, 1879-80

Plate 12, figure 12; Plate 18, figure 3

Lepidodendron andrewsi Lesquereux, 1879-80, Coal Flora, p. 389,

pl. 64, fig. 6.

Description. —Stems up to 5 cm in diameter. Leaf
cushions rhomboidal or rhomboidal oval, base tapering
but blunt, apex blunt, rounded; lateral angles rounded.
Leaf cushions 3.5-7 mm high, 3.5-6 mm wide. Leaf scars
in middle of leaf cushion and average 3 mm wide and 2
mm high; thickened rim outlines the outer limits of leaf
Scar; central vascular area below middle, vascular bundle
scar about one-fourth mm high, obovate; parichnos scars
rounded, extremely small but recognizable. Ligular scar
1 mm below apex of leaf cushion, horizontally elongate,
three-fourths mm long.

Entire surface of leaf cushion smooth, flat without keel
markings except rarely a cushion will have a faint central
vertical line from apex to base.

Discussion.— The type specimen at the National
Museum, number 15430, reverse side numbered 15431, is
a Mazon Creek, Illinois, nodule. The leaf cushions are 5
mm high, 4 mm wide; leaf scars 3 mm wide, 2 mm high and
centrally placed on the leaf cushion. A raised rim outlines
the leaf cushion. No ligular scar or depression is evident.
On a few of the cushions, a faint “caudal” line or ridge
extends from the apex to its base but not evident on the
very smooth leaf scar. Other specimens in my collections,
especially the numerous ones from Wheatland, Indiana,
show additional features. The ligular scar is prominent on
all the cushions and is a millimeter below the blunt apex.
The vertical central “caudal” line is missing and the en-

Oun10 FREEPORT CoAL Lycopsips AND SPHENOPSIDS: ABBOTT /

tire surface of the cushion and leaf scar is smooth. On the
smaller stems, which are 1 cm in diameter, the leaf cushions
are 3.5 mm high and 3.5 mm wide with leaf scars 2 mm
wide and 1.5 mm high. Larger stems have cushions up to
7 mm high and 4.5 mm wide.

The leaf cushions on the Ohio specimens average 6
mm high and 4 mm wide with leaf scars averaging 3 mm
wide and 2 mm high. Leaves were not seen attached to
any of the stems.

Lesquereux related L. andrewsi to L. vol!kmannainum
Sternberg but remarked on the difference in position of
the leaf scars. He noted that the leaf scar was not so broad
as apically placed in L. andrewsi. After seeing the type of
L. andrewsi at the National Museum, I agree with Les-
quereux.

Janssen, 1940 (pp. 14-16) reevaluated the Lesquereux
specimens pertaining to L. volkmannianum. These speci-
mens have nothing in common with L. andrewsi. The
American material of L. volkmannainum has leaf cushions
widely separated both horizontally and vertically, and the
surfaces are distinctively ornamented. L. wortheni also has
a distinctively wrinkled surface below the leaf scar.

Occurrence. — Kimberly, Ohio (8587, 8588); Wheat-
land, Indiana (numerous specimens) ; and Mazon Creek,
Grundy County, Illinois (Lesquereux, 1879-80) .

Lepidodendron cf. jaraczewskii Zeiller, 1888
Plate 12, figure 7

Lepidodendron jaraczewskti Zeiller, 1888, Valenciennes, p. +57, pl. 67,
figs. 3, 3a.

Description. — Leaf cushions subrhombic, raised, with
prominent groove outlining the cushions; cushions 26 mm
high, 6-7 mm wide, long tapering, widest in upper one-
third. Leaf scar slightly raised, small, 1 mm high and 1.5
mm wide, diamond-shaped with upper margin more or less
descending from lateral angles for about 2 mm; vascular
area obscured by heavy carbonization; small scar above
leaf scar might be ligular scar. Uppermost coaly surface
unmarked by cushion outlines; coarsely striated.

Discussion. —The three specimens from Kimberly are
more similar to Zeiller’s L. jaraczewskii than to other
known species. The Kimberly specimens do not show a
central line or raised area from apex to base of the leaf
cushion, but this is perhaps a minor point due to the state
of preservation. Dr. Bell (1944) recorded the species from
Nova Scotia. His figure 2 on plate 51 is more nearly like
the Kimberly species than his figure 1, although the leaf
scars on his specimens are more nearly centrally placed.
His specimens are perhaps from older portions of the
trunk than are those from Kimberly.

Occurrence. — Kimberly, Ohio (8328, 8320, 8102). In
Canada: two localities in northern Nova Scotia (Bell,
1944) .

Lepidodendron aculeatum Sternberg, 1825
Plate 12, figure 8

Lepidodendron aculeatum Sternberg, 1825, Flora der Vorwelt, vol. 1
jh, seals iy ahh, 7,

Lepidodendron ingens Wood, 1860, Acad. Nat. Sci. Philadelphia,
Proc., p. 239, pl. 6, fig. 4.

Lepidodendron lesquereuxti Wood, 1860, ibid, p. 240, pl. 5, fig. 4.

)

Description. —Trunks large, portions incomplete, 34
cm wide, 60 cm Jong, with leaf cushions spirally arranged,
separated by a prominent, 1 mm wide striated furrow out-
lining each cushion.

Leaf cushions 11-15 mm wide, 20-28 mm high, elongate,
long tapering, asymmetric; lateral margins gently curving,
lower right margin concave, lower left margin roundedly
convex. Leaf cushion surface with prominent keel ridge
from leaf scar area to base of cushion; 3-4 horizontal trans-
verse ridges or wrinkles sometimes present on either side
of keel ridge.

Leaf scar area raised, placed above the middle and in
upper one-third of cushion; 1.5-4 mm below apex of
cushion; leaf scar angular, wider than high, 3-6 mm wide,
2-5 mm high; lines from lateral angles of scar arch to mar-
gin of cushion in most specimens.

Vascular area in center of leaf scar; vascular bundle
and parichnos scars small, circular, less than 0.5 mm in
diameter. Ligular scar above leaf scar and at apex of
cushion, circular, | mm in diameter.

Discussion. — The above description is based on speci-
mens from Paoli, Indiana. The poor preservation and the
presence of an iron stain has rendered a single specimen
from Kimberly almost useless. However, the extremely large
leaf cushion, asymmetric shape, and leaf scar area make
it noteworthy.

I have been unable to locate the specimens of Wood
and of Lesquereux. The synonomy is based on that of
Lesquereux’ because he had access to Wood's specimens.

Occurrence. — Kimberly, Ohio (8622); Illinois (Jans-
sen, 1939); Paoli, Indiana, and site unknown (Canright,
1959) ; Michigan at several localities (Arnold, 1949) ; Car-
bondale, Pennsylvania (Lesquereux, 1879-80). In Canada:
two localities from the Cumberland Group, northern Nova
Scotia (Bell, 1944).

Lepidodendron scutatum Lesquereux, 1879-80
Plate 12, figure 11; Plate 18, figure 2

Lepidodendron scutatum Lesquereux, 1879-80, Coal Flora, p. 369, pl.
43, fig. 6 (figs. 6a, 6b specimens not located).

Lepidodendron setifolium Lesquereux, manuscript bound in with
Lesquereux’ copy of the Coal Flora, p. 370,

8 PALAEONTOGRAPHICA AMERICANA (VI, 38)

Description. —Stems 1 to 10 cm in diameter. Leaf
cushions rhomboidal or rhomboidal oval, base tapering but
blunt, apex rounded, lateral angles rounded. Leaf cushions
8-10 mm high, 5-6 mm wide; leaf scars above middle of
cushion with lower part or base of scar more or less in the
middle. Leaf scars transversely rhomboidal, 3-3.5 mm wide,
2-2.5 mm high. Vascular area below middle of scar, round
with central vascular bundle one-fourth mm in diameter;
two lateral parichnos scars rounded to oval but smaller
in diameter than vascular bundle.

Ligular scar narrow, irregularly shaped, sometimes de-
pressed about 1 mm long horizontally and located 0.75 mm
below apex of leaf cushion, other than scar area, smooth
except for from 3 to 5 zigzag horizontally aligned wrinkles;
wrinkles 0.5-1.25 mm long and narrow; no prominent
vertical “keel line” or ‘caudal line’’.

Discussion. —The above description is based not only
on Lesquereux’ type but also on additional material from
1) the same locality, 2) other localities in Henry County,
Missouri, 3) Kimberly, Ohio, and 4) Wheatland, Indiana.

The type of Lepidodendron scutatum which Les-
quereux illustrated on plate 43, figure 6, is in the National
Museum, bears number 15448, and is from Clinton, Henry
County, Missouri. Additional material, collected later from
the same locality, was described by Dr. White, 1899, pages
198-200.

The smaller stems, well seen on specimen number
6074 from Gilkerson’s Ford (only a part of which is shown
on Dr. White's plate 72, figure 4) has twigs 3 mm wide and
up to 13.5 cm long. The attached leaves are 15-18 mm
long, 0.5 mm wide, sickle-shaped and erect spreading.

Number 6044 from Pitcher’s mine, on Dr. White's
plate 45, figure 4, has leaf cushions 4 mm long, and slightly
over 2 mm wide. The leaf scar is 1.3 mm wide and more
or less | mm high and placed slightly higher on the cushion
than the usual central placement. A few cushions show a
faint more or less continuous ridge from base of leaf scar
to the blunt apex. This is the only specimen which shows
even a hint of a vertical ridge.

Number 6046 from Deepwater, Dr. White’s figure 2,
plate 55, has stems 2 cm wide, 15 cm long with leaf cush-
ions 5mm high and 3 mm wide. The leaf scars are 1.5 mm
wide and about | mm high.

Number 6045 from Deepwater, Owen’s coal mine, Dr.
White’s figure 1, plate 55, has several dichotomously
branched stems with and without leaves attached and a
few recognizable leaf cushions. His line drawing at figure
2, plate 65, shows too much wrinkling and, therefore, the
appearance of a ridge which is not present.

Sellards recognized L. scutatum in the flora of Kansas.

On his page 423 (plate 56, figure 3) Sellards stated: ‘“This
species seems to be represented by a single specimen of a
small dichotomous branch from Lancing, Kansas. The
stem is slender, only 5 mm wide below the dichotomy, the
bolsters elliptical, 2.5 to 3 mm long, about one third as
wide as long, acute aboye and below, rounded at the sides.
The leaf scar is placed above the middle.”

Lepidodendron scutatum is distinct from L. obovatum
and should not be included in that synonomy as indicated
by the bibliography of Fossilium Catalogus. The differences
lie in the shape, proportion of area in leaf cushion with
that of the leaf scar, size range, and the ornamentation at-
tributed to L. obovatwm is not seen on L. scutatum.

Occurrence. — Kimberly, Ohio (8589) ; Clinton, Gilk-
erson’s Ford, Pitcher’s mine, Deepwater, Missouri (Les-
quereux, 1879-80, White 1899) ; Lancing, Kansas (Sellards,
1903) ; and Wheatland, Indiana (numerous specimens in
Abbott collection) .

Genus LEPIDOPHLOIOS Sternberg, 1825
Lepidophloios vaningeni D. White, 1900
Plate 13, figure 6

Lepidophloios van Ingeni D. White, 1900, Monograph 37, pp. 205-210;
pl. 56, figs. 1-8; pl. 58, fig. 12; pl. 61, fig. 1-c; pl. 62, fig. £; pl.

68, fig. 5.

Description. — Large trunks characterized by diagonal
rows of transversely rhomboidal leaf cushions, Cushions
more than twice as wide as high, ranging from 9 mm wide
and 2 mm high to 20 mm wide and 6-8 mm high; apical
angle 135-145°, sides forming the apical angle more or
less straight to slightly convex, in order to accommodate
the rounded lower margin; lateral angles 25-23°; lower
margin concave and rounded and appears to overlap the
apex of the cushion below. Leaf scar, in lower one-third of
cushion, also transversely rhomboidal, but apical portion
gently rounded and not acute as in the corresponding por-
tion of the cushion, Leaf scar near to or touching lower
border of cushion; scar 9 mm wide and 2 mm high on a
cushion 20 mm wide and 6 mm high; others are in propor-
tion to area of cushion. Vascular bundle scar minute, cir-
cular, about 14 mm in diameter. Parichnos scars to right
and left of vascular bundle scar vertically elongated and 1
mm wide and 0.25 mm high.

Discussion. — This species is represented in the Upper
Freeport flora by a single specimen which agrees in all
aspects with the specimens used by Dr. White in his original
descriptions.

Occurrence. — Kimberly, Ohio (8826); Pitcher’s coal

Ono FREEPORT CoAL Lycorsips AND SPHENOPSIDS: ABBOTT 9

bank and Gilkerson’s Ford, Henry County, Missouri
(White, 1900) .

Lepidophloios larcinus Sternberg, 1825
Plate 12, figure 6; Plate 19, figure 4

Lepidophloios larcinum Sternberg, 1825, Flora der Vorwelt, vol. 1,
Lepidositcits acadianus Dawson, 1968, Acadian Geology, p. 489, fig.
Re seaiiiyas obcordatum Lesquereux (in part), 1866, Geol. Sur. II-

linois, vol. 2, p. 457, pl. 41, figs. 2-2a, not fig. 1.

Lepidophloios obcordatus Lesquereux, 1880, Coal Flora, vol. 2, p. 423.

Description. — Hexagonal leaf cushions 6.5 mm _ wide
and 4 mm high; leaf scars not clearly delineated; vascular
bundle scar slightly above the middle, circular, 0.25 mm
in diameter; vertically oriented, crescent-shaped parichnos
scars, to right and left of vascular scar, 0.25 mm wide and
0.75 mm high.

A bow-shaped arching wrinkle prominent in lower one-
third of cushion extending across width of cushion.

Discussion. — The specimen illustrated at figure I,
plate 41 by Lesquereux in his Coal Flora has not been lo-
cated, therefore, I shall not speculate on it.

Lesquereux’ type of Lepidophloios obcordatus, which
is illustrated at figures 2 and 2a in the Geological Survey
of Illinois, II, is in the Illinois State Museum collections
and bears number 1728. Janssen, 1940, pages 20-21 and
plate 3, figure 3, reevaluated the status of this specimen
and believed it to be a poorly preserved Lepidophloios
larcinus.

Dr. Bell, in his Memoir 238 (1944), stated that due
to the variation of leaf cushion, leaf scars, difference in
size and of branches and their mode of preservation, that
all of the Canadian specimens should be considered as L.
larcinus. Therefore, Dawson’s L. acadianus, is only one
form of Lepidophloios larcinus.

Only one specimen was collected at the Kimberly
locality, and although it is fairly well preserved, it does
not show a ligular scar as do Dr. Bell’s specimens.

Occurrence. — Kimberly, Ohio (8715); DuQuoin and
Colchester, Illinois (Lesquereux, 1866, 1880; Janssen,
1940) ; Vigo County, Indiana (Canright, 1959) . In Canada:
Riversdale Group, Cumberland Group —at several local-
ities each, northern Nova Scotia and roof of the Coalbrook
seam near Wadden Cove, Sydney Coalfield, Nova Scotia
(Bell, 1938, 1944).

Genus ASOLANUS Wood, 1861
Asolanus camptotaenia Wood, 1861
Plate 13, figure 1; Plate 19, figure 1

Asolanus camptotaenia Wood, 1861, Acad. Nat. Sci. Philadelphia,
Proce. vol} 12; p. 238, pli 4, fig. 7,

Sigillaria monostigma Lesquereux, 1866 in part, Geol. Sur. Illinois,
vol. 2, p. 449, pl. 42, figs. 1-5.

Lepidodendron cruciatum Lesquereux, 1870, Geol. Sur. Illinois, vol.

| 4s pp: 432; pl. 25; fig. 2:

Sigillarioides stellaris Lesquereux, 1870, Geol. Sur. Illinois, vol. 4, pl.

29, fig. 3.

Stigmarioides tuberosus Lesquereux, 1870, Geol. Sur. Illinois, vol.
tp 3, pl. 29, fig. 5.
Stigmaria stellaris Lesquereux, 1870-80, Coal Flora, vol. 2, p. 516, pl.

74, figs. 5, 7.

Stigmaria ficoides var. stellata Lesquereux, 1883-84, Indiana Dept.
_ Geol. Nat. History, 13th Ann. Report, Part 2, p. 96, pl. 19, fig. 4.
Sigillaria (Asolanus) camptotaenia D. White, 1899, U.S. Geol. Sur.,

Mon. 37, pp. 230-238, pl. 69; pl. 70, figs. 1, 3, 4; pl. 61, fig.

1g?; pl. 62, fig. i?; pl. 64, fig. e?.

Description. — Surface of stems marked between leaf
cushions by rough, irregular, ropelike striae or strands;
striae radiating from all sides of the leaf cushion, the most
prominent of which diagonally connect the four neighbor-
ing leaf cushions.

Leaf cushions range from 10-32 mm apart vertically
and 13-40 mm apart horizontally; cushions rhomboidal,
higher than wide with acute lateral angles and rounded
apical and basal margins; cushions 4-6 mm high and 4-5 mm
wide; leaf scar area in upper 2 mm of cushion, 2 mm high
and 3-6 mm wide; leaf scar area raised apically 2 mm, low-

ering gradually to more or less | mm high basally.

Vascular scar slightly above the middle of the leaf
scar, slightly elongated horizontally, about 14, mm; two
lateral parichnos scars linear, crescent-shaped, aligned verti-
cally, 0.5 mm high, upper and lower ends almost touching.

Discussion. — Asolanus camptotaenia is well repre-
sented in American collections from many horizons and
localities. Dr. White, 1899, in Monograph 37, discussed
many of the earlier types of Lesquereux in the National
Museum and concluded that the types listed in the above
synonomy should be considered as one and the same
species, Asolanus camptotacnia Wood.

Janssen, 1940, reviewed the Lesquereux’ specimens at

the Illinois State Museum and was in agreement with Dr.
White.

I agree with the above authors that almost all of the
specimens should be considered as one species. However,
two specimens figured by Lesquereux are different from
Asolanus camptotaenia although they do belong to genus
Aso'anus. One specimen bears U.S.N.M. number 16420 and
is shown on plate 26, figure 5 of the Geological Survey of
Illinois vol. 4, and the same figure is repeated in Coal
Flora on plate 73, figure 6. The second specimen is shown
on plate 73, figure 4 of the Coal Flora. Both of these speci-
mens were used by Lesquereux in his later description of
Sigillaria monostigma. This epithet cannot be retained nor
used for these two specimens since the first description was

10 PALAEONTOGRAPHICA AMERICANA (VI, 38)

published in 1866 and the type, shown at figure 1, plate
42, is another specimen.

Occurrence. — Kimberly, Ohio; Braidwood, Chester,
Morris and Mazon Creek, Illinois (Lesquereux, 1866, 1870,
1879-80 and Noé, 1925); Indiana (Lesquereux, 1883-84) ;
Cannelton, Pennsylvania (Lesquereux, 1879-80); St.
Charles and Grand Ledge, Michigan (Arnold, 1949) ;
Gilkerson’s Ford and Pitcher’s coal mine, Henry County,
Missouri (White, 1899); LeRoy shales near Lawrence and
the Chanute shales, Thayer, Kansas (Sellards, 1908). In
Canada: roof of Gowrie seam (Phalen seam) and shales
above Bouthillier seam, Sydney Coalfield, Nova Scotia
(Bell, 1938) .

Asolanus sigillarioides (Lesquereux ex Abbott)
Plate 13, figure 3

Sigillaria fissa Lesquereux, 1858, Geol. Pennsylvania, p. 871, pl. 13,
fig. 4 (nomen confusum).

Lepidophloios sigillarioides Lesquereux, 1879-80, Coal Flora, p. 425,
pl. 68, figs. 8, 8a.

Sigillaria (Asolanus) sigillarioides D. White, 1899, U.S. Geol. Sur.,

Mon: 37, p: 239; pl! 70; fig. 2.

Description. — Surface of trunk marked by fine longi-
tudinal striations arching around leaf cushions. Leaf cush-
ions spirally arranged, 0.5-1.5 cm distant vertically, 1.8-2.25
cm distant horizontally; no ribs or grooves separate cush-
ions into a vertical series; leaf cushions transversely rhom-
boidal, laterally acute, 5-6 mm high, 5-7 mm wide.

Leaf scar area raised up to 2.5 mm; leaf scar area
occupies the upper one-half of the cushion; 5 mm wide, 2-
mm high; vascular bundle scar in lower part of leaf scar,
circular, a little less than 0.5 mm in diameter, parichnos
scars to right and left of vascular scar, rounded to linear
crescent-shaped, 0.25 mm wide, 1.2 mm high. An additional
rounded scar sometimes present in upper one-third of leaf
scar, 0.5 mm in diameter, perhaps ligular.

Discussion. — The surface of the specimens is marked
by rope-like striations vertically aligned and arching around
the leaf cushions. The striations are not so prominent as
those of Asolanus camptotaenia and only occasionally do
the striations appear as diagonal lines between the cush-
ions. There are variations on a single specimen, although
the cushions remain the same size.

There are two pieces of shale at the National Museum
which obviously fit together and form one adjoining and
contiguous segment of stem. One of these pieces is labeled
as the type of Sigillaria fissa Lesq., while the other is
the type of Asolanus sigillarioides. The two fragments,
according to Dr. White, were either separated at the time
of collection or one was afterwards misplaced. The type
S. fissa as described and figured was supposed to have come

from Pennsylvania, while in reality, it came from Missouri,
The figure representing the type is inadequate as well as
the description of the leaf cushions. Therefore, I agree with
Dr. White, that the epithet S. fissa should be designated as
a nomen confusum and should be designated as Asolanus
sigillarioides.

Occurrence. — Kimberly, Ohio (8650, 8651), and Clin-
ton, Henry County, Missouri (U.S.N.M. No. 6173).

Genus SIGILLARIA Brongniart, 1822
Sigillaria mamillaris Brongniart, (1824) 1836
Plate 12, figures 2, 2a, 3, 3a; Plate 16, figures 1-2;
Plate 19, figure 2

Sigillaria mamillaris Brongniart, 1824—Ann. des Sciences, natur., vol.
4, p. 33, pl. 2, fig. 5 poorly preserved and not used as the type;
1836, Histoire des veg. Fossiles, vol. 1, p. 451, pl. 149, fig. 1
(type).

Description. — Ribs raised, 6-10 mm wide, separated by
zigzag or undulating furrows; where decorticated, furrows
straight. Ribs with vertical series of leaf cushions separated
by 1-4 mm of transversely wrinkled area and more or less
prominent: arc-shaped groove 0.25 mm above the apex of
leaf scar.

Leaf cushions 6-10 mm wide, delimited by width of
rib, and up to 10 mm high.

Leaf scars angularly pyriform, 5-6.5 mm wide, widest
slightly above the middle, taper to a 2 mm wide, rounded
apex; apex of leaf scar sometimes with vertical central in-
dentation.

Vascular area 2-3 mm below apex of leaf scar, rounded
vascular bundle scar 0.25 mm in diameter; lateral parichnos
scars concrescent, narrow, 1.5 mm high, more or less con-
verging upwards. Vascular area on decorticated specimens
large, 2-2.5 mm long, I-1.25 mm wide; upper margin con-
cave, 0.5 mm deep, lower margin rounded.

Discussion. — Most of the American specimens attrib-
uted to this species have raised more or less rounded ribs
with leaf cushions touching, although the lower limit of the
cushion is not prominent in all specimens. The apex of the
cushions is clearly defined by an arching transverse groove,
since the apex of the cushions stands slightly higher than
the basal area of the cushions. The leaf scars on younger
stems occupy all of the upper surface of the cushions since
the cushion is interpreted as an inverted truncated cone.
On older stems, the leaf cushion is more elongated, the leaf
scar is not elongated by remains essentially the same size
as on the younger stems, and thus does not occupy all of
the upper surface of the cushion. The surface of this part
of the cushion is inconspicuously wrinkled by transverse,
discontinuous wrinkles.

Onto FREEPORT Coat Lycorsips AND SPHENOPSIDS: ABBOTT ll

According to Dr. Bell (1944, page 92) a ligular scar
can be seen above the vascular scar on his Canadian ma-
terial.

The specimen figured by Lesquereux in Coal Flora,
(vol. 3, plate 108, figure 6) is in the National Museum and
bears Number 16520 (Lacoe 475). The specimen has un-
dulating ribs with an average diameter of 6 mm. The
cushions vary in size from base to apex of the large speci-
men. Towards the base, the cushions are not clearly de-
limited, but the pyriform leaf scars are separated by 3.5 to
5 mm of cushion which is slightly ornamented by discon-
tinuous transverse wrinkles. In the apex of the specimen
the leaf scars are smaller and only slightly separated from
one another, with their outlines angular. This specimen is
from Buctel, Ohio, which is not far from the Kimberly
locality.

The specimen also figured by Lesquereux in Coal
Flora, (vol. 2, plate 72, figure 6) is also at the National
Museum. It does not show the variation in leaf scars that
the first specimen does. However, they are the same as those
on the lower part of the previous specimen, that is, pyri-
form in outline, and separated by fine transverse wrinkles.
This specimen is from Oliphant, Pennsylvania.

I have not been able to locate the specimen which
Lesquereux illustrated on plate 72, figure 5.

Occurrence. — Buctel (U.S.N.M. 16520) and Kimberly,
Ohio (8831, 8057); Knox and Sullivan Counties, Indiana
(Canright, 1959); Christopher, Illinois (Janssen, 1939) ;
‘Tennessee and Virginia (Lesquereux, 1879-80) . In Canada:
the Cumberland Group at Springhill, northern Noya Scotia
(Bell, 1944) .

Sigillaria elegans (Sternberg) Brongniart, 1828
Plate 12, figure 5

?Favularia elegans Sternberg, 1825, Versuch, 1, 4, pp. 43-44, pl. 52,
fig. 4.

Sigillaria elegans Brongniart, 1828, Prodrome, pp. 65, 172.
Description. — Leaf cushions in a vertical zigzag series

with about 0.25 mm between them. Cushions raised, trun-

cated, hexagonal with somewhat rounded lateral margins,

upper and lower margins straight; cushions 3.5-4 mm high
and 4-6 mm wide.

Leaf scars occupy most of the surface on the truncated
leaf cushion and essentially the same shape although slight-
ly smaller; center of upper margin sometimes indented or
emarginate. Vascular bundle and parichnos scars about 1
mm below upper margin of cushion; on decorticated sur-
face, vascular area almost in middle of cushion.

Elliptical vascular bundle scar elongated horizontally
from 0.4-0.75 mm long; parichnos scars narrow, elongate,
diagonal to vertical in alignment.

Decorticated surface with vertical striations, undulating
or not so prominent zigzag furrows, vascular area 1.5 mm
wide with carbonized circular rim.

Ligular pit, in Canadian specimens, commonly incon-
spicuous, placed almost above the center of the leaf scar.

Discussion. — Zeiller, in 1886, 88, stated that Sternberg’s
figure is not accurate and that until Sternberg’s type is
reviewed, there will be some doubt as to the validity of
the species. Zeiller reviewed the specimens upon which
Brongniart based his species giving their variations, differ-
ences, and relationships.

The Canadian, as well as the Kimberly specimens,
agree with the figures and description given by Zeiller in his
Valenciennes.

Lesquereux, in Coal Flora, placed many specimens in
his consideration of S. tesse/lata. I believe some are, in all
probability, S. elegans; however, until I have occasion to
review all specimens labeled S. tessellata, 1 cannot be cer-
tain. S. tessellata does not occur in the Kimberly flora.

Dr. Bell recognized S. elegans Dawson as correct and
without comment.

Occurrence. — Kimberly, Ohio (8354). In Canada: five
localities in the Cumberland Group, northern Nova Scotia
(Bell, 1944) and roof of the Harbour seam at Old Queen
Pit, Sydney Coalfield, Nova Scotia (Bell, 1938).

Sigillaria lacoei Lesquereux, 1879-80
Plate 13, figures 4a, b

Sigillaria lacoci Lesquereux, 1879-80, Coal, Flora, p. 499, pl. 72, figs.

12- 12a- 12b.

Sigillaria discoidea Lesquereux, 1858, Geol. Pennsylvania, p. 873, pl.

14) fig. 5:

Description. — Ribs large, 4-7 cm wide, average 4 cm
with surface vertically striated except on leaf scars. Leaf
scars 12-20 mm apart vertically.

Leaf scars elongate, ovate, upper margin emarginate;
scars 17-25 mm high; 5-13 mm wide; width uniform from
near apex to rapidly tapered blunt base.

Vascular area above middle, vascular bundle 1 mm
wide crescent-shaped, aligned horizontally and centrally
between two parichnos scars; parichnos scars elongate, ellip-
tical, aligned vertically, 2 mm high, about 0.5 mm wide.

Discussion. —On decorticated ribs, the vascular area
remains prominent, especially the parichnos portions. At
first the area is entire, that is, a single unit deeply emargi-
nate at the upper margin, then deeper further below the
surface. By further emargination or indentation of the

12 PALAEONTOGRAPHICA AMERICANA (VI, 38)

upper margin, the area becomes high twin scars close to-
gether, contiguous, still vertically aligned, higher than
wide —17 mm high and each one 5 mm wide.

In his Coal Flora, plate 72, Lesquereux figured only
one rib of a large specimen from Wyoming Valley, Ply-
mouth, Pennsylvania (U.S.N.M. Number 16817) . Its preser-
vation is such that several stages of decortication is present.
Several other specimens from Plymouth are in the National
Museum collections as well as some from Pittston, Pennsyl-
vania.

One specimen is from Kimberly, Ohio, Number 8649.

None of the surfaces, either where the scar is better pre-
served or where decortication has continued, are rugose.
This species has nothing in common with Sigillaria rugosa
except that the two are of the genus Sigillaria.

Other of the Lesquereux’ species listed in Fossilium
Catalogus under the bibliography of Sigillaria rugosa
should not be considered under that species since they are
different.

Occurrence. — Kimberly, Ohio (8649), and Plymouth
and Pittston, Pennsylvania (U.S.N.M. 16811, 16806, 16810) .

Sigillaria laurenciana (Lesquereux ex Abbott)
Plate 12, figure 4; Plate 16, figure 3

Sigillaria laurenciana Lesquereux, description only, written in long-
hand (script), p. 483b and bound inside Lesquereux’ copy of the
Coal Flora (nomen nudum).

Description. — Ribs 15-30 mm wide; leaf cushions not
clearly defined; leaf scars of every other rib match as to
position or are in the same position horizontally; leaf scars
10 mm high, 11 mm wide and 10-13 mm apart vertically.
Rib surface faintly vertically striated; surface horizontally
wrinkled or reticulate between the leaf scars, sometimes
with a rounded concave deeper wrinkle 2 mm above the
leaf scar.

Leaf scars ovate, slightly eccentrically placed on the
rib, not in the middle, more to the right side of the rib,
especially noticeable in specimens with larger ribs.

Leaf scars with apical and basal margins convexly
rounded, sides with more or less rounded lateral angles.
Lowermost 2 mm of leaf scar surface faintly horizontally
wrinkled.

Vascular bundle scar and parichnos scars prominent,
situated above the middle on narrower ribs; vascular bundle
scar rounded, 0.75 mm in diameter; parichnos scars about 2
mm high vertically; bases of parichnos scars and vascular
scars connected apically to form a bracket-shaped scar. On
larger ribs, vascular scar is u-shaped and 0.75 mm wide
horizontally; parichnos scars more or less straight, 2-2.5 mm

high and 0.5-0.65 mm wide, occupying about 2-3 mm of
space horizontally.

Discussion. — The type specimen is in the National
Museum and assigned Number 16571. A second specimen is
numbered 16570. Both specimens are from the shales of
Lawrence, Kansas.

Lesquereux described this type in script or longhand,
on tissue thin pages which were inserted and bound be-
tween the pages of his copy of the Coal Flora. The writing
is beautifully executed and, with the exception of a few
words where the ink has faded, is still clearly legible. A
small outline sketch of the vascular area is also included in
the margin, so there is no doubt as to which specimen
Lesquereux was referring.

Reference to this species is made by Dr. White in his
Bulletin 211, page 106, although he did not discuss it.

Reference is also made to the species by Lesley, Rept.
of the Geological Survey, Pennsylvania, Dictionary of
Fossils, vol. 3, 1890, page 855, however, he made no com-
ment.

Occurrence. — Shales of Lawrence, Kansas (U.S.N.M.
16571) and Kimberly, Ohio (8000-1) .

Sigillaria rugosa Brongniart, 1828
Plate 12, figure 10; Plate 17, figures 3, 4

Sigillaria rugosa Brongniart, 1828, Prodrome, pp. 64, 171; Histoire,
1, 12, p. 476, pl. 144, fig. 2.

Description. — Ribs distinctly delineated, more or less
straight but irregular in width; 5-13 mm, average 7 mm
wide, with vertical series of leaf scars separated by 21-25
mm; surface of rib with low ornamentation of horizontal
wrinkles in “jet-like trail” below leaf scar, widest immedi-
ately below leaf scar and outlined for a length of 15 mm
by a straight line or wrinkle on either side of rugose area.
Surface above leaf scar also rugose and a reverse “‘jet trail”
extends upward above leaf scar about 8-10 mm.

Leaf scar pyriform, 2.5-5 mm wide, widest below mid-
dle, apex notched or shallowly grooved; 6-7 mm high, base
more or less truncated. Area below base and 1-2 mm of base
of leaf scar raised. Scars in diagonal series horizontally and
no two are in the same alignment.

Vascular area in upper one-third of leaf scar, 2 mm
below apical notch. Central vascular bundle U-shaped or
concave upwards, small, 1 mm high; parichnos scars ellip-
tical, vertically aligned, 1-2.5 mm high, 0.33-0.5 mm wide,
parichnos scars usually not same length in one vascular
area.

A circular area or scar 1 mm in diameter, situated

Ouro FREEPORT Coat Lycopsips AND SPHENOPSIDS: ABBOTT 13

immediately above apical notch, perhaps representing ligu-
lar position.

Discussion. — The Lesquereux specimen, U.S. National
Museum, Number 17463, is from the Diamond Vein, Brigg’s
shaft Scranton, Pennsylvania. The trunk is 20 cm in di-
ameter and it is not complete. The ribs are 12-12 mm wide
with leaf scars 21-22 mm apart. The leaf scars are 8 mm
high and 5 mm wide with lines off the sides trailing down-
ward for about 15 mm. The aluminum-like surface makes
the “jet-trail” wrinkling conspicuous.

The specimens, which were described by Brongniart
and upon which this species is based, were from Wilkes-
barre, Pennsylvania. :

The bibliography of Sigillaria rugosa, in Fossilium
Catalogus, includes several species, described by Lesquereux,
which do not belong to Sigillaria rugosa. Because the pres-
ent study is one of a flora and not a monograph of each
genus represented, I shall not dwell here on all of the
American species so flagrantly included in the synonomies
of S. rugosa or other species. Three species do not belong
here: Sigillaria marginata, S. lacoei, and S. pittstoniana.

Occurrence. — Kimberly, Ohio (8652, 8615, 8693, 8621) ,
and Brigg’s shaft, Scranton, Pennsylvania (U.S.N.M.
17463) .

Sigillaria orbicularis Lesquereux, 1879-80
Plate 12, figure 9; Plate 16, figure 4; Plate 17, figure 1;

Plate 18, figure 4
Sigillaria orbicularis Lesquereux, 1879-80, Coal Flora, 2, p. 491

Description. — Ribs straight, 11-12 mm wide with leaf
scars up to 12 mm apart vertically; cushions aligned hori-
zontally at every third rib. Rib surface smooth, decorticated
surface finely striated vertically.

Leaf cushion wider than high, 9 mm high, 6-7 mm
high; upper margin emarginate, rounded, sides angular,
base roundedly truncate. Scars match horizontally, every
third rib.

Circular vascular area slightly elevated above rib sur-
face, 1 mm below the apex of the leaf scar. Vascular bundle
scar circular, one-fourth to one-third mm in diameter,
parichnos scars prominent, vertically elliptical, 0.35-0.5 mm
wide; 1.5 mm high, scars almost touching apically.

Decorticated rib surface vertically striated; vascular
scar area prominent, broadly ovate, larger than on the
upper surface, 5 mm high, 2 mm wide.

Discussion. — Two specimens at the National Museum
labeled by Lesquereux are from Maltby, Pennsylvania. Both
specimens show details of ribbing and leaf scars, as well as a
decorticated rib surface area. The one bearing Lacoe col-

lection Number 616a and U.S.N.M. Number 16489, was
used by Lesquereux, in Coal Flora, page 491, in his descrip-
tion of the species Sigillaria orbicularis.

Other specimens of this species are from Pittston,
Pennsylvania.

The shape of the leaf scar, as well as its width, (be-
cause it takes up almost all of the rib width), and the
absence of prominent vertical lines trailing below the scar
area, precludes the inclusion of this species within the
circumscription of Sigillaria laevigata as is suggested in Fos-
silium Catalogus, page 892.

Occurrence.— Maltby and _ Pittston, Pennsylvania
(type — Lacoe Number 616A and U.S.N.M. Number 16489)
and Kimberly, Ohio (8800, 8626, 8654, 8831, 8829).

Sigillaria pileata Abbott, n. sp.
Plate 12, figure 1; Plate 17, figure 2

Description. — Ribs raised, straight, 4 mm wide, with
vertical series of leaf scars 14 mm apart. Surface of rib
immediately above leaf scar rugose in rounded patch 2.5-3
mm high and 2-2.5 mm wide, the remainder of surface verti-
cally striated to base of scar above; rarely is a narrow area
below scar rugose.

Leaf scar angularly pyriform, 6 mm high, 3.5-4 mm
wide, widest below middle; as wide as rib; apex of scar in-
dented, base truncated.

Vascular area 2 mm below apex of leaf scar, with con-
crescent, 0.5 mm wide, bundle scar and two parichnos scars
also concrescent in shape but vertically elongated, 1 mm
long.

Discussion. — There are five specimens from Kimberly,
Ohio, but I have not seen this species in any other Ameri-
can collections.

The species can be compared with Sigillaria rugosa in
that the two are ornamented on the rib surface by horizon-
tally oriented wrinkles; however, the surface of S. rugosa
is wrinkled both above and below the leaf scars in charac-
teristic “jet stream” areas. The species also can be compared
with S. schlotheimiana because this species also is ornament-
ed by a rugose rib surface; however, S. schlotheimiana is
a European species which is recorded from the lower Coal
Measures.

Occurrence. — Kimberly, Ohio (8719, 8796, 8790, 8722,
and 8017).

Genus STIGMARIA Brongniart, 1822
Stigmaria ficoides (Sternberg) Brongniart, 1822
Plate 19, figure 3

Variolaria ficoides Sternberg, 1820, Flora der Vorwelt, vol. 1, fase.
1, p. 24, pl. 40, figs. 1-3.

14 PALAEONTOGRAPHICA AMERICANA (VI, 38)

Stigmaria ficoides Brongniart, 1822, Mem. Mus. Hist. naturelle, vol.
_ 8, pp. 228, 239, pl. 1, fig. 7.
Stigmaria elliptica Lesquereux, 1870, Geol. Sur. Illinois, vol. 4, p. 451,

p29. ne 2)

Stigmaria verrucosa D. White, 1899, U. S. Geol. Sur. Monograph 37,

p. 244.

Discussion. — None of the American specimens, attrib-
uted to this well-known root species, show additional in-
formation which could be of aid to further its description.

The type of Stigmaria elliptica Lesq., No. 1706 in the
Illinois State Museum, is illustrated by Janssen, 1940. The
specimen is distorted slightly, perhaps by lateral pressure,
as Janssen suggested (p. 28), and the rootlet scars are more
or less elliptical in outline rather than circular. This differ-
ence does not seem significant enough to assign it to a
separate species.

Dr. White, 1900, based his synonomy of this species
upon the date 1809 at which time Martin described this
root under the name Phytolithus verrucosus.

Occurrence. — Kimberly, Ohio, numerous specimens;
Mazon Creek and Braidwood, Illinois (Lesquereux, 1870,
Noé, 1925, Janssen, 1939) ; Gilkerson’s Ford, Henry County,
Missouri (White, 1900) ; St. Charles-Garfield mine at East-
wood, Michigan (Arnold, 1949) ; several localities in Penn-
sylvania, West Virginia, Indiana, and Ohio, Alsa scattered
throughout the Morien Series of Nova Scotia, Canada (Bell,
1938).

Genus BOWMANITES (Binney), Hoskins and Cross, 1943
Bowmanites incurvata Abbott, n. sp.

Plate 1, figures 1-4; Plate 11, figure 2

Description. — Cone incomplete, 9 cm long, 15 mm
wide; raised ribs or ridges 0.75 mm wide, separated by
narrow grooves; internodes 5 mm long with axis 6 mm
wide; apical verticil sterile.

Bracts 8-10 mm long, 0.5 mm wide, taper to an acute

‘

apex; bracts united basally into a “collar” for a distance
of about 2 mm, free portion at first at right angles to then
downwardly directed, free tips spreading and 4-5 mm long.

Sporangiophores arise midway on collar portion of
bract whorl, arch upward then recurve toward cone axis;
maximum height of curve 5 mm, entire sporangiophore
length 7 mm.

Sporangia 5-6 on each sporangiophore, in vertical align-
ment, individually attached, not grouped. Sporangia 3 mm
high, 3 mm wide, circular in outline; where crowded, ovate
to pear-shaped, 4 mm long and 2 mm wide; attachment area
to sporangiophore small.

Sporangial wall cells long, narrow with thick walls,
cells elongated in the direction of the long axis of the
sporangium.

Discussion. — The axis of the cone, with its ribs and
grooves, is more nearly that of a young calamitean axis
than one of Sphenophyllum, Attachment of the sporangio-
phore to the bract whorl, rather than to the cone axis,
places the species in Bowmanites, heretofore assigned to
species belonging to Sphenophyllum.

Bowmanites also has various arrangements and orient-
ation of the sporangial grouping on the sporangiophores.
The sporangial arrangement is unique in this species be-
cause the sporangia are attached in a vertical series on the
sporangiophore.

Usually fertile specimens from Mazon Creek have the
spores excellently preserved, but in this specimen, I was
unable to procure them.

Occurrence. — The species does not occcur in the Kim-
berly flora. Mazon Creek, Grundy County, Illinois (at the
University of Illinois, Botany Department Number 8446a
and b, both halves of an ironstone nodule) .

Bowmanites magna (Lesquereux ex Abbott)
Plate 6, figures 1, 2, 4; Plate 8, figures 1, 2:
Plate 9, figures 9-16; Plate 11, figure 1

Calamostachys magna (Lesquereux?) Noé, 1925, Pennsylvania Flora
of Northern Illinois, plate 5, fig. 5.

Description. — Four incomplete cones, 8 cm long, 12
mm wide; internodes 6 mm long. Cone axis 6-7 mm wide.
Bracts 20 at a node, subtend 2-2.5 bract whorls aboye; 12-15
mm long, 0.5 mm wide, widest near base, bracts at first at
right angles to cone axis then gently arch upward. Apical
whorl not spreading, sterile. One sporangiophore attached
adaxially to each bract about | mm away from axis; spo-
rangiophore 4 mm long, 0.25-0.53 mm wide, gently arching,
stout, extends up to one-half to slightly more than the
middle of the internode.

Sporangia elliptical, terminal on sporangiophore, four
in number on sporangiophore, distal ends of sporangia
toward cone axis; unequal in size, lower pair 5 mm long,
1 mm wide; upper pair about 3 mm long, | mm wide.

Spores rounded, smooth, wrinkled, trilete, range from
42.5 u to 67.5 u in diameter; trilete apparatus with sutures
inconspicuous, 7.5-12.5 u long.

Discussion. — Noé, in 1925, on plate 5, figure 5, showed
the type specimen and gave credit to Lesquereux for its
specific epithet. Noé did not give a description or discussion
of the species. Because Lesquereux worked for many years
in Illinois, he probably labeled the specimen. However,
I have been unable to find any mention of it in the num-
erous writings of Lesquereux. The specimen is in the Uni-

Onto FREEPORT Coat Lycopsips AND SPHENOPSIDS: ABBOTT 15

versity of Illinois Collection, Number P1284, and rephoto-
graphed on Plate 6, figure 2.

The type specimen, as well as three cones from Kimber-
ly, shows the sporangiophore attachment to be on the bract
and not on the cone axis, thus belonging to the genus
Bowmanites.

The Illinois specimen did not contain spores, however,
all three of the Kimberly specimens contained the same
spore type.

Occurrence. — Mazon Creek, Grundy County, Illinois
(University of Illinois Botany Department Number P1284) ;
Kimberly, Ohio (8532, 8531, 8504, 8691 and 10 slides) .

Genus MAZOSTACHYS Kosanke, 1955
Mazostachys compacta Abbott, n. sp.
Plate 6, figures 5-8; Plate 11, figure 3

Description. — Fertile branches bear from 2-7 cones at a
node, internodes 1-1.5 cm distant, axis (0.5-1 mm wide. Cones
sessile on fertile branches; composed of a compact series
of bracts alternating with sporangiophores and verticillate.
Cones 1.2-2.5 cm long, average 2 cm; 2-3 mm wide, average
3 mm; with internodes 1-1.5 mm long between bract whorls;
central axis 0.5 mm wide.

Bracts 12-16 in a verticil, average 2.5 mm long, 0.3 mm
wide at base, acuminate; slender, narrow, strongly keeled,
somewhat reflexed near axis then abruptly curve upward;
bracts reach slightly above the base of the next higher
bract whorl and in some cases, reach as high as the second
higher whorl.

Sporangiophores 6-8 in a verticil; only one-half the
number of bracts in verticil; attached above middle of inter-
node and directed downward; each sporangiophore bears
two sporangia, one above the other.

Sporangia ovate, 0.75-1 mm long, 0.5-1 mm wide.

Sporangial wall cells typical for the sphenopsids; cells
elongate, more or less rectangular, elongated in the direc-
tion of the long axis of the sporangium.

Discussion. — The leaves of Annularia sphenophyl-
loides Zenker have been found in association with three
types of articulated cones: Calamostachys, Metacalamo-
stachys, and Mazostachys. Sterzel, in 1882, gave a text-figure,
showing the leaves of Ann. sphenophylloides attached along
the branch axis bearing several cones of Calamostachys
calathifera Weiss. The Upper Freeport coal flora bears
numerous specimens of cones as well as numerous frag-
ments of leafy shoots of Ann. sphenophylloides. Other cone
genera are also present among the fragments of this foliage
genus, signifying that mere association does not mean affin-
ity to that genus.

The species belongs within genus Mazostachys in that:
1) the sporangiophore attachment is below the bract; 2)
there are two sporangia on a sporangiophore; 3) there
are one-half as many sporangiophores as bracts in a verticil.

The species cannot be intimately compared with
Mazostachys pedunculata Kosanke because the highly car-
bonized nature of its preservation eliminates the study of
the vascular traces to the sporangiophore. However, dif-
ferences lie in the facts that there are only two cones per
node in M. pedunculata while there are two to seven cones
per node in M. compacta. The internodes are longer in M.
pedunculata and the bracts are not keeled. The sporangia
M. pedunculata lie side by side or in horizontal juxtaposi-
tion, while the sporangia are over and under or vertically
positioned in M. compacta.

Occurrence. — Kimberly, Ohio (8539, 9540 and 25
transfers) .

Genus CALAMOSTACHYS Schimper, 1869
Calamostachys tubercu/ata (Sternberg) Weiss, 1884
Plate 3, figures 1, 2, 5; Plate 9, figure 6; Plate 10, figure 2

Bruckmannia tuberculata Sternberg, 1825, Flora der Vorwelt, 4, 29,
pl. 45, fig. 2.

Calamostachys tuberculata Weiss, 1884, Steinkohlen-Calamarien 5, 2,
ps LZ8.

Description. — Articulated strobili with alternating
verticils of sterile bracts and sporangiophores. Incomplete
strobili up to 11 cm long, 15 mm wide; internodes average
5 mm long; axis averages 5 mm wide.

Bracts at axis united into a collar; bracts 1 mm wide
on separating from collar; bracts 5-9 mm long, linear lance-
olate, acuminate, reflexed, spreading away from collar then
directed upward to cover base of succeeding bract whorl
above. Bracts 14-16 in a verticil.

Sporangiophores attached in midportion of internode;
2.5-3 mm long, 0.25 mm wide; four sporangia narrowly at-
tached apically; sporangiophores fewer in number than
bracts, average nine in a verticil.

Sporangia 2.5 mm long, 2 mm wide, 1.25 mm thick;
ovate; sporangium wall cells elongated, thickened by num-
erous projections internal to each cell.

Discussion. —'This species has been reported as hetero-
sporous. I also have found large and small spores in the
macerations but not in the amounts necessary to substanti-
ate that they originated in this particular cone. Especially
since there are other spores present which obviously do not
belong to the calamitean taxa.

Occurrence. — Kimberly, Ohio (8530, 7413, 7254 and
six slides) ; Mazon Creek, Grundy County, Illinois (B2573) ;

16 ; PALAEONTOGRAPHICA AMERICANA (VI, 38)

Wheatland, Knox County, Indiana (9620, 9621). In Can-
ada: 200’ above the Tracey seam to the roof of the Hub
seam of the Morien Series, Sydney Coalfield, Nova Scotia
(Bell, 1938) .

Calamostachys germanica Weiss, 1876
Plate 7, figures 1-5; Plate 9, figures 4, 5, 7, 8; Plate 10, figure 7

Calamostachys germanica Weiss, 1876, Steinkohlen-Calamarien, 2, p.
47, pl. 16, figs. 3, 4.

Description. — Strobili 8-11 cm long, 1-1.5 cm wide,
cylindrical with alternating verticils of bracts and sporan-
giophores. Cone axis $.5-6 mm wide, internodes 3.5-6 mm
long; axis striated and with up to 14 ribs. Numerous bracts
in a verticil, 20-25 in the visible part of the verticil; only
4-5 sporangiophores in a verticil.

Bracts united for 3-4 mm into a collar at axis; collar
bends downward at juncture with axis then curves upward,
the free portion of bracts ascend and are somewhat diver-
gent from axis extending one to two nodes above. Free
portion of bracts 4-10 mm long, average 6 mm; 0.5-1 mm
wide at union into collar.

Sporangiophores attached to axis slightly below mid-
portion of internode, with varying degrees of divergence
from axis; apex of sporangiophores enlarged to 1 mm wide
with four horizontal prongs or arms. Sporangiophores 2-5
mm long, average 3 mm, and 0.25-0.33 mm wide.

Four sporangia attached narrowly at apex of sporangi-
ophore, one on each of the four prongs; sporangia ovate,
2-4 mm long, 1.25-2 mm wide.

Sporangium wall one cell layer thick, lateral walls with
many inwardly projecting ridges; cells rectangular, elon-
gated in the direction of the long axis of sporangium; near
or in attachment area, cells rounded.

Spores average 47.5 >< 60 p, tend to stick together in
tetrads, triradiate with trilete sutures; sutures thin, narrow,
7.5-10 «4 long; outer spore layers granulose,

Discussion. — Calamostachys germanica are the most
numerous of all of the cones in the Upper Freeport. The
cones are mostly incomplete, that is, some lack a base while
others lack the apex. None of them were found attached
to a larger axis.

The cones show varying degrees of maturation. Many
which are over 1 cm in width have shed their spores and
the sporangial sacs are fragmentary. Many of the bracts
are gone, as well as the sporangiophores and the small scars
of the axis can be counted to indicate the numbers orig-
inally attached in each verticil. In some instances, bases of
the sporangiophores adhers to the axis giving them the

appearance of a hook or, as Jongmans (1911) said, a “rose
prickle”’.

In specimens where the cone axis is visible, the axis
is not only striated but also ribbed with as many as eight
ribs in a “half-whorl” or the part visible.

Bracts at the node are united into a collar for from 3
to 4 mm before being freed. As many as four to five of the
free bract portions bend upward to partially surround each
sporangial grouping, thus giving the striking appearance
which immediately separates this species from others in
Calamostachys.

According to Jongmans (1911) C. thuringia Weiss is
similar to C. germanica, C. germanica being more robust.
Because there is a variation in size, perhaps attributed to
natural growth, as well as in preservation, I hesitate to
separate the Kimberly specimens into these two species.

Occurrence. — Kimberly, Ohio (numerous specimens,
the better preserved ones: 8861, 8862, 5606-26, 8045) ;
Wheatland, Knox County, Indiana (numerous specimens) ;
Chiefton Mine, Terra Haute, Vigo County, Indiana (nu-
merous iron stone nodules) ; Mazon Creek, Grundy County,
Illinois (B2573) . In Canada: Thorburn Member of Stellar-
ton Series, Pictou Coalfield, northern Nova Scotia (Bell,
1940) ; and ranges from about 1000’ below the Tracey seam
to roof of Phalen seam, Sydney Coalfield, Nova Scotia
(Bell, 1938); Cumberland Group, northern Nova Scotia
(Bell, 1944) .

Calamostachys bisporangiata Abbott, n. sp.
Plate 6, figure 3; Plate 10, figure 6

Description. — Articulated cone, incomplete with either
apex or base missing, fragments 5-6 cm long, up to 13 mm
wide, cylindrical, linear with alternating verticils of sterile
bracts and sporangiophores. Cone axis 1.5-2 mm wide, in-
ternodes averaging 4 mm long, ribbed with 3-4 striated
ribs, ribs not alternating at node. Axis below cone striated,
ribbed, up to 15 mm long without another articulation
visible.

Bracts 6-12 in a verticil, fewer near apex, 5-7 mm long,
0.5 mm wide, linear, acicular, stiff, somewhat divergent
from axis, extend to base of node above.

Sporangiophores same number as bracts in verticil,
6-12, up to 1.5-3.5 mm long, 0.5 mm wide, attached to axis
in center of internode.

Sporangia two in a group, attached narrowly at apex
of sporangiophore; ovate, 1.5-3 mm long, 1-2 mm _ wide;
6-12 groups in verticil, depending upon size of cone.

Sporangium wall one cell layer thick, cells elongated in

Onto FREEPORT CoAL Lycopsips AND SPHENOPSIDS: ABBOTT |

the direction of long axis of sporangium; cells at or near
attachment area smaller, more nearly rounded.

Discussion. — This type of cone, where the base is pre-
served, consistently showed a slender, naked, ribbed axis
below the fertile portion which is 10 to 15 mm long, but
without an articulation. This is the longest stalked species
of cone in the Kimberly flora.

The bracts and sporangiophores are of the same num-
ber in each vyerticil and the verticils are about the same
throughout one cone. However, the size of the cone varies
and the next cone may have fewer bracts or more bracts
in a yerticil. The variation is from six to as many as
twelve. The bracts and sporangiophores are in a vertical
series, one above the other.

There were no assignable spores to these cones.

Occurrence. — Kimberly, Ohio (8555, 8821, 8558, and
several transfers and macerations) ; Maumee Chiefton Mine,
Terra Haute, Vigo County, Indiana; Mazon Creek, Grundy
County, Illinois.

Calamostachys minuta Abbott, n. sp.
Plate 8, figure 3; Plate 10, figure 4

Description. —Fragments of cones up to 37 mm long,
articulated with alternating whorls of sterile bracts and
sporangiophores; cones long, slender, 5.5 mm wide.

Cone axis smooth (not ribbed), striated, 0.5-0.75 mm
wide, internodes 3 mm long.

Bracts six in a verticil, widely divergent from cone
axis, acicular, 2-2.5 mm long; cone apex terminates in six
erect bracts, 1 mm long.

Sporangiophores attached well above the middle of the
internode, in upper one-third of the length of the inter-
node; directed upward and outward; thorn-shaped, 1-1.5
mm long, peltate with enlargement 0.75 mm wide.

Sporangia four in a group on each sporangiophore, at-
tached to the adaxial peltate portion; sporangia ovate,
1-1.5 mm long, 0.75 mm wide.

Cells of sporangial wall one cell layer thick. Cells
elongated in the direction of the long axis of the sporan-
gium, cells in attachment area much smaller and more
or less rounded.

Discussion. — Parts of seven cones are in close prox-
imity on one transfer. Two of the cones bifurcate from a
common axis. The others are oriented in such a fashion
that they probably were attached to the same axis as the
bifurcating pair.

The sporangiophore attachment, in the upper one-
third of the internode, and its ascending position, places

~I

the sporangia under the protection of the bracts of the next
higher verticil.

The sporangial wall cells are about the same general
shape and size as those in the other species of articulated
cones even though the sporangia are much smaller than the
general size.

No spores, with certainty, can be assigned to the
species.

Occurrence. — Kimberly, Ohio (Transfer $25) .

Calamostachys recurvata Abbott, n. sp.
Plate 1, figure 6; Plate 10, figure 3

Description. — Three incomplete cones, 3-4 cm long,
5-6 mm wide; cones articulated with alternating verticils
of sterile bracts and sporangiophores.

Cone axis smooth, not ribbed, striated, 1 mm wide;
internodes 4 mm long, except near apex and then 2 mm
apart.

Bracts eight in a verticil, acicular, 2.5 mm long, 0.25
mm wide, widely divergent from node, droop downward
then outward.

Sporangiophores attached 1 mm above the node, well
below middle; 2 mm long, upwardly directed holding the
sporangia well away from the cone axis.

Four sporangia apically attached to each sporangio-
phore, sporangia ovate, 2 mm long, 1 mm wide, | mm in
thickness. Sporangial wall cells elongated in the direction
of the long axis of the sporangium; cells smaller than the
usual sporangial wall cells yet in proportion to the size of
the sporangium.

Discussion.— ‘This cone is one of the smallest and
slenderest of the articulated cones. Its long internode length
and small bracts and sporangia make it different from other
species. The position of the sporangiophore is likewise dif-
ferent since it originates in the lower one-third of the inter-
node. This position and the upwardly directed sporangio-
phore also makes it appear close to a Palaeostachya.

No spores, with certainty, can be assigned to the
species.

Occurrence. — Kimberly, Ohio
7089 on shale.

(Transfers 526-327),

Calamostachys longibracteata Abbott, n. sp.
Plate 2, figures 6, 7; Plate 10, figure 5

Description. — Incomplete portion
only, 9 cm long, 15 mm wide in lower part, gradually
diminishing toward apex, with alternating verticils of
bracts and sporangiophores, in vertical series. Internodes
average 7 mm long, 7 mm wide, ribbed, ribs not alternat-

strobilus, apical

18 PALAEONTOGRAPHICA AMERICANA (VI, 38)

ing at node. Bracts 18-20 in a verticil, 13-15 mm long, | mm
wide at base. Bracts acicular, stiffly divergent from axis,
extend to a little above the base of next succeeding verticil.

Sporangiophores equal in number with bracts, 2.5 mm
long, 0.25 mm wide, attached in center of internode and at
right angles to it. Each sporangiophore with two sporangia
at apex; sporangia attached along upper one-half and ex-
tend beyond the apex of the sporangiophore. Sporangia
rounded to ovate, 3.5-4 mm long, 3 mm wide, 2-2.5 mm
thick.

Apical verticil sterile with all bracts enclosing the
apex tightly, budlike in appearance.

Discussion. — This calamitean type of cone has been
reported in American literature from Mazon Creek, Illinois,
and Michigan Valley, Kansas. Dr. Baxter, 1963, in the
American Journal of Botany, described heterosporous
calamitean cones found in coal balls. These compression
cones have more or less the same general organization as
do the three dimensional cones of the coal balls. Using
maceration methods, the carbonaceous cones have the same
sporangial wall details as the better preserved coal ball
material. That is, the cells are rectangular and elongated
in the direction of the long axis of the sporangium. The
cells are large, up to 400 » long with the walls thickened
by inward projections. The projections are not unique to
this particular species because the sporangium wall of all
the species which I have been able to macerate have the
same inward thickenings.

On the theory that this might be the megasporangiate
cone of the heterosporous species, the Kimberly material
was completely used up in macerations. I found no spores
which might designate it megasporangiate.

Occurrence. — Kimberly, Ohio (was 8526); Mazon
Creek, Grundy County, Illinois (Janssen, 1939), and
Michigan Valley, Kansas (Baxter, 1963).

Calamostachys interculata Abbott, n. sp.
Plate 7, figures 6, 7; Plate 10, figure 1

Description. — Cones articulated with alternating verti-
cils of bracts and sporangiophores; bracts and sporangio-
phores of equal number, 16-20 in a verticil; bracts attached
at node, sporangiophores attached in middle of ribs about
middle of internode, attachment areas zigzag or are not in
vertical alignment; axis striated, ribbed, ribs alternate at
nodes. Cones incomplete, up to 10 cm long, 18 mm wide
with internodes 4-6 mm long.

Bracts united into a narrow, | mm wide, collar at
node, free portion at first sags downward from collar then
gently arches upward and diverges from axis, reach in

height to about the middle of the internode. Free portion
of bracts 5-6 mm long.

Sporangiophores attached at right angles to axis about
midway on internode, 2-4 mm long, about 0.33 mm wide.
Apex of sporangiophore divided into 4 “arms”, each of
which is 0.5 mm long.

Four sporangia attached narrowly at apex of sporangio-
phore, one on each arm; sporangia 3 mm wide, 3-3.5 mm
long, about 2 mm thick, ovate.

Cellular detail of sporangial wall as in Calamostachys
germanica.

Discussion. — Calamostachys interculata is one of the
larger calamitean cones in the Kimberly flora, There are
four large fragments upon which this description is based.
The cone axis is not always clearly ribbed, but with careful
dissection and partial maceration, the axis as well as other
parts may be examined.

The sporangiophores are not situated directly above
the bracts as in many of the species; the sporangiophores
alternate with the bracts and are midway on the rib. The
ribs, therefore, alternate at the node which arrange the
bracts and sporangiophores in a zigzag series rather than
a vertical one.

Calamostachys interculata is not an oversized C. ger-
manica because the bracts of C. interculata are equal in
number with the sporangial groups, while as many as four—
five bracts subtend each sporangial group of C. germanica.
Also the free bracts in C. interculata are stubby, stiff, and
short reaching upward to midway of the internode on
which they are inserted.

The cellular detail of the sporangium walls is like that
in C. germanica (see Pl. 7, fig. 5).

No spores could be assigned with certainty to this
species.

Occurrence. — Kimberly, Ohio (8513, 7413).

Genus PALAEOSTACHYA Weiss, 1876
Palaeostachya elongata (Presl), Weiss, 1876
Plate 1, figures 5, 7; Plate 10, figure 9

Palacostachya clongata (Presl), Weiss, 1876, Steinkohlen Calamarien
I, p. 108, pl. 15; II, p. 181, pl. 22, fig. 15.
Description. — Axis 13 cm long, 7 mm wide, striated,

ribbed; ribs 0.5 mm wide; six internodes 2 cm long, bear-

ing two opposite cones at each node.
Cones shortly stalked, stalk not more than 5 mm long,

8 mm wide. Cones incomplete, up to 7 cm long, 8 mm

wide with 16 internodes, bract whorls 3-4 mm apart.
Bracts average eight in a whorl, but as few as six

toward apex; bracts 6-7 mm long, 0.75 mm wide, narrowly

OnI0 FREEPORT COAL Lycopsips AND SPHENOPSIDS: ABBOTT 19

lanceolate, pointed, extend to, seldom above the succeeding
bract whorl.

Sporangiophore attached in axil of bract, ascending,
3-4 mm long, 0.25 mm wide; equal in number with bracts.

Four sporangia attached apically and narrowly to
apex of each sporangiophore. Sporangia average 3 mm long,
1.25 mm wide, ovate; sporangial wall cells similar in size,
shape and orientation as those of Calamostachys. The cells
are rectangular, elongated in the direction of the long axis
of the sporangium, and walls with inwardly directed thick-
ened projections.

No spores can be assigned to this species.

Discussion. — Shale specimen, No, 8690, bears parts of
eight cones, none of which are complete. The longest cone
is 7 cm long with 16 bract whorls. This cone tapers grad-
ually from its maximum width of 8 mm near the base to
5 mm near its apex. The lower part bears eight bracts in a
whorl and the apical part only six bracts in a whorl. The
overall preservation leaves much to be desired and un-
fortunately no spores could be found that could be as-
signed to this cone.

Occurrence. — Kimberly, Ohio (8690, 8710 and two
transfers) ; in Canada: roof of the McLean seam, Sydney
Coalfield; and several localities in the Cumberland Group,
northern Nova Scotia (Bell, 1938, 1944).

Palaeostachya ovalis (Lesquereux ex Abbott)
Plate 2, figures 2, 3; Plate 10, figure 10

Asterophyllites ovalis Lesquereux, 1858, iz Roger’s, Geol. Pennsyl-
vania, 2, p. 851, pl. 1, fig. 2.

Calamostachys ovalis Lesquereux, 1879-80, Coal Flora, 3, pp. 717-718,
pl. 89, figs. 3, 4.

Description. — Cones more than 7 cm long, incomplete,
1 cm wide near apex, 1.7 cm wide near base; cones com-
pact with sporangiophores superimposed on bracts, bracts
verticillate; axis distinctly ribbed, ribs not alternating at
node. Internodes 3 mm long near apex, 4.5 wide toward
base.

Bracts near apex six in verticil, 5 mm long, 1 mm
wide; bracts near base of cone ten in verticil, 10 mm long,
1.5 mm wide, stout; bracts acicular, narrow, carinate, taper
from base to slender apex; somewhat reflexed near axis
then curved outward and upward; bracts near apex reach
almost to base of second node above, those near base more
reflexed and reach only to base of first node above.

Sporangiophore inserted on adaxial surface of each
bract near juncture with axis; sporangiophore 5 mm long,
bears four ovate sporangia.

Sporangia narrowly attached at apex, 3.5-4.5 mm long,
2-2.5 mm wide.

No spores could be assigned with certainty to this
species.

Discussion. — While reviewing specimens which others
have referred to as Calamostachys germanica in the Ameri-
can collections, I found significant differences in many of
them. One such related group of specimens, which I here
designate as Palacostachya ovalis, had heretofore been as-
signed as a probable Calamostachys germanica [Calamo-
stachys ovalis, in Fossilium Catalogus (p. 487) .] The type,
from Alton, Pennsylvania, is Number 852f Lacoe Collec-
tion and National Museum Number 18055. This specimen
shows parts of two cones and is refigured on Plate 2, fig-
ure 3. Additional specimens from Dade County, Georgia,
also in the National Museum and later considered by
Lesquereux, bear Lacoe numbers 825a and 825b.

Due to the attachment of the sporangiophore in rela-
tion to the subtending bracts, this species of cone is placed
in Palaeostachya. Also the bracts are fewer in number and
different in arrangement from that of Calamostachys ger-
manica.

Occurrence. — Not present in the Kimberly flora. The
type is from Alton, Pennsylvania (Lesquereux, 1858). Dade
County, Georgia (Lesquereux, 1879-80) ; and questionably
from Henry County, Missouri (White, 1900) .

Palacostachya aperta (Lesquereux ex Abbott)

Plate 3, figures 3, 4, 6; Plate 4, figures 1, 2; Plate 5, figures 1, 2;
Plate 9, figures 2, 3; Plate 10, figure 8
Asterophyllites aperta Lesquereux, 1858, in Rogers, Geol. Pennsyl-

vania, p. 825, pl. 1, fig. 4.

Volkmannia fertilis Lesquereux, 1884, Coal Flora, p. 720, pl. 90, fig. 4.
Macrostachya aperta Lesquereux, 1884, Coal Flora, p. 829, pl. 3, fig.
ite ot infundibuliformis Schimper, Lesquereux, 1884, in part,

Coal Flora, pl. 3, fig. 20 only.

Volkmannia crassa Lesquereux, 1884, Coal Flora, p. 719, pl. 90, fig. 1.
Annularia tuberculata Lesquereux, 1884, Coal Flora, p. 723, pl. 89,

figss 2:

Description. — Cones long, slender, articulate; basal
portion for three internodes without leaves; next six in-
ternodes above with leaves but sterile; all internodes up-
ward to apex with leaves and fertile.

Axis striated, ribbed, 1 cm wide at base gradually di-
minishing upwards to one mm wide at apex; basal inter-
node 3.5 cm long; second internode 16 mm long; third
internode 14-17 mm long with more or less appressed
leaves, the number of leaves indeterminate, many missing;
axis 5 mm wide; 4th, 5th, 6th internodes diminish in height
with more leaves remaining at node, leaves 10-12 mm long,
longer than internode, extending outward and upward for
a height of 1.5-2 internodes. The 7th internode, about mid-
way of cone, and on to cone apex fertile.

20 PALAEONTOGRAPHICA AMERICANA (VI, 38)

Fertile zone with up to 25 internodes; internodes more
or less same height throughout, 4-4.5 mm, with leaves at
first united basally then becoming united forming a
“collar” with the outer edge toothed; collar 10-15 mm high,
extends one sometimes two internodes above; teeth on edge
of collar show that the width of about five bracts subtends
each sporangial group; six to eight sporangial groups visible
in a verticil, with two sporangia in a group; groups super-
imposed.

Sporangiophores attached in axil of bract whorl, up
to 7 mm long, slightly cruciate at distal end. Sporangia
ovoid, attached to sporangiophore distally at narrower end,
up to 5 mm long, 1.5 mm wide, widest at lower end.

Spores range from 60 to 80 » in diameter, triradiate
with thin, short trilete sutures; sutures 7:5-10 » long; outer
spore layers thin, peeling, and smooth.

Discussion. — In 1858, Lesquereux described and _ fig-
ured Asterophyllites aperta, a cone, from Archbold, Penn-
sylvania, Lacoe Collection Number 950.

In 1880, Plate III, figure 20, the same cone is figured,
but the explanation of figures shows the name as Macro-
stachya aperta. In the text, page 60, the plate and figure
number is included in Lesquereux’ Macrostachya infundi-
biliformis. In the discussion (page 62) Lesquereux had
doubts about its inclusion under this epithet.

Later in his volume III (p. 829) in the Chapter on
Additions and Corrections, Lesquereux removed this speci-
men and renamed it Macrostachya aperta.

This same specimen, Lacoe Collection Number 950, is
figured also on plate 90, figure 4, and is described, this time,
under the epithet Volkmannia fertilis.

To sum up, the first four names in the synonomy deal
with one specimen, Lacoe Collection Number 950.

The last two names in the synonomy are cones of sim-
ilar organization and are Volkmannia crassa (Lacoe Num-
ber 851) and Annularia tuberculata (Lacoe Number 914a-
f).

Lesquereux used the genus Volkmannia to show the
cone’s relationship to the foliage genus Asterophyllites. The
definition of the genus Volkmannia, as applied by Stern-
berg, 1825, is vague and may apply either to a cone or to
a sterile branch bearing crowded verticils of imbricate
bracts or leaves. Its usage, as applied here, is untenable.

To further clarify the position of the Lesquereux’
cones, terms applied by Wood, 1860, should not be con-
sidered. Wood (page 438) renamed a British specimen
(Annularia fertilis) Trochophyllum fertilis. Wood used the
generic term Trochophyllum because he believed the term
Annularia to be preoccupied by a subkingdom of Mollusca,

Schumacher, 1817. The usage of the terms by Wood, should
in no way be considered when dealing with the Lesquereux’
specimens.

Lesquereux, in 1884 (p. 723) described a cone and
believed it to show a resemblance to one which Lindley
and Hutton (1831) named Asterophyllites tuberculatus.
Because the bracts of the cone were in closer affinity to an
Annularia, Lesquereux used the term Annularia tubercu-
lata for his cone. In Kidston’s revision (1899) of the Lind-
ley and Hutton specimens, he assigned the cones to
Stachannularia, a term used to signify a section of Calamo-
stachys.

Lesquereux’ original description, as well as the study
just completed, show the sporangiophores of the cones to
be axillary in their orientation on the axis and thus in
closer harmony with genus Palaeostachya than to Calamo-
stachys. Therefore, their inclusion with the British speci-
mens is untenable.

This species, for one reason or another, does not fit
within the circumscription of the 18 “valid” species already
assigned to genus Palacostachya. In order to avoid further
confusion, the species epithet aperta is retained.

Occurrence. —The species does not occur in the Kim-
berly flora. Archbald and West Pittston or southwest of
Scranton, Luzerne County, Pennsylvania; Dade County,
Georgia (Lesquereux, 1884); Stark County, Illinois ?
(Lesquereux, 1884) .

Palaeostachya trabeculata Abbott, n. sp.
Plate 2, figures 1, 4, 5; Plate 9, figure 1; Plate 11, figure 4

Description. — Cones incomplete, base missing, 20
bract whorls; cones 3 cm long, 5-8 mm wide; apical verticil
of bracts sterile, appressed, budlike in form.

Internodes 2-2.5 mm long, axis 1 mm wide; ten bracts
in a verticil with a sporangiophore attached in the axil of
each bract; bracts up to 5 mm long, extend horizontally
just beyond the sporangial groups then curve upward
reaching to about one-half the internode. Sporangiophores
2 mm long, narrow, about 0.33 mm wide, apex slightly en-
larged, 1-1.5 mm wide. Four ovate sporangia attached
adaxially to curciate apex, 1.5-2 mm long, 1-1.5 mm wide,
widest below middle. Cells of sporangium wall rectangular,
elongated in the direction of the long axis of sporangium.

Spores range in diameter from 27.5 to 37.5 u, trilete,
sutures 9-10 » long, thin; spores wrinkled, folded, surface
smooth.

Discussion. —This cone has few bracts in a verticil,
only ten, and each bract bears a sporangiophore in its axil.

Outro FREEPORT COAL Lycopsips AND SPHENOPSIDS: ABBOTT

The apex of the sporangiophore is enlarged and in longi-
tudinal view the length is 1-1.5 mm, but the width is un-
known. Four sporangia are attached to this apical enlarge-
ment. The spores are like those described for the homo-
sporous Calamostachys and the sporangium wall cells are
buttressed also like the sporangial wall of Calamostachys.

The cones are similar to those of Palaeostachya vera
Seward, 1898, with the bracts and sporangiophores equal
in number, and sporangiophores cruciate. The course of
the vascular trace of the sporangiophore cannot be com-
pared due to the type of preservation of this cone. Also
P. vera is a larger cone generally with 18 bracts and spo-
rangiophores in a verticil.

Occurrence. — Kimberly, Ohio (8718, 8719, 7445) .

Genus CALAMITES Suckow, 1784
Calamites undulatus Sternberg, 1825
Plate 11, figure 9; Plate 14, figures 1-2

Calamites undulatus Sternberg, 1825, Versuch, 1, 4, p. 26.

Description. — Stems up to 13 cm wide, some incom-
plete, up to 34 cm long with six complete internodes. Inter-
nodes shorter than broad, 1-6.5 cm high; shortest where
branch scars occur; branch scars verticillate, 1.5-1.75 cm
apart horizontally; branch scars above nodal lines, trans-
versely oval, 5 mm in diameter. Ribs converge around
branch scars.

Ribs flattened, 1-2 mm wide, on some internodes
straight, on others flexuous, or both straight and flexuous
on same internode; some ribs do not alternate at node;
rib endings rectangular to sharply pointed.

Leaf scars on upper rib endings circular to oval, up to
1 mm in diameter; smaller scars sometimes present on low-
er rib ends.

Outer surface and pith cast in organic connection;
outer surface smooth to conspicuously transversely wrinkled.
All internodal surfaces longitudinally striated and smooth,
except for wrinkles; rib position not reflected on_ this
surface.

Discussion. —One Kimberly specimen, 8000-2, shows
the outer surface in organic connection with the ribbed
pith cast. The outer surface on the upper three internodes
is conspicuously wrinkled; while the fourth internode has
only a few wrinkles, the fifth internode is not wrinkled.
The ribbed surface shows straight ribbing on most of the
internodes; however, two internodes at the top of figure
2, Plate 14, shows the characteristic flexuous ribbing.

Occurrence. — Calamites undulatus ranges from the

Middle through the Upper Carboniferous of the United
States and Canada. Kimberly, Ohio, numerous specimens;
Michigan basin (Arnold, 1949); Danville, Illinois (Jans-
sen, 1939); also in collections from Illinois, Indiana,
Arkansas, Missouri, Pennsylvania, and West Virginia. In
Canada: Pictou Coalfield (Bell, 1944).

Calamites carinatus Sternberg, 1824
Plate 11, figure 8; Plate 15, figures 1-2

Sine be: carinatus Sternberg, 1824, Versuch, fasc. 3, pp. 36, 39, pl.
Bee he hs Artis, 1825, Antediluvian Phytol., pl. 2.

Description. — Stems up to 19 cm wide with usually
one, sometimes two, branch scars visible at a node; branch
scars variable in diameter, 1.2-2.5-3.3 cm, oval, round to
subtriangular in outline; ribs converging on margin of
branch scar, forming fan-shaped groups of 2-4 ribs. Inter-
nodes up to 24 cm long, with ribs straight, longitudinally
striated, 1-1.5 mm wide, most of which alternate at the
nodes; upper ends of ribs broadly or obtusely angular with
an elongate, ovate leaf scar, 1 mm long.

Discussion. — Kidston and Jongmans (1917, p. 143)
discussed at length the history concerning the species Cala-
mites carinatus and Calamites ramosus. Their conclusions,
based on the International Rules, are that the name
Calamites ramosus must be placed in the synonomy of C.
carinatus which was created a year sooner.

American specimens previously referred to as Cala-
mites ramosus and thus effected, are as follows: Lesquereux
(1879-1884) ; D. White (1893, 1900, 1903); Bell (1938,
1944) ; Janssen (1939).

Occurrence. — Kimberly, Ohio (7422, 7469, 8075) ;
Deepwater, Owen’s bank, Gilkerson’s Ford, Henry County,
Missouri; McClelland’s Shaft near Beeleville, and Aurora,
Lawrence County, Missouri (White, 1893, 1900); Grand
Ledge, Michigan (Arnold, 1949); Severly shales, Osage,
LeRoy shales, Lawrence, Kansas (Lesquereux, 1879-80 and
White, 1903); Mazon Creek, Grundy County, Illinois
(Janssen, 1939). In Canada: New Waterford, roof of Har-
bour seam, Sydney Coalfield (Bell, 1958); Riversdale
(rare) and Cumberland (common) groups, northern Nova
Scotia (Bell, 1944).

Calamites cisti Brongniart, 1828

Plate 11, figure 5

Calamites cisti Brongniart, 1828, Historie, 1, (2), p. 129, pl. 20, figs.
1-5.
Description. —Internodes long, up to 13 cm _ with
branching irregular to sparce, nodes not contracted; ribs
straight, alternating at the nodes; rib endings angular or

2? PALAEONTOGRAPHICA AMERICANA (VI, 38)

subacute with an oval or elliptical vascular scar on the
upper ends; lower ends usually without such scars; ribs
1-1.5 mm wide, longitudinally striated,

Discussion. — The type specimen of Calamites cisti is
from Wilkesbarre, Pennsylvania, and was originally de-
scribed by Brongniart. This specimen has been refigured
in Kidston and Jongmans (1917) on plate 95, figures 4, 5.

Since Brongniart’s description, additional material has
been figured and described by Lesquereux (1879-1883) ,
Dawson (1873), White (1900), Bell (1944), and Arnold
(1949) .

Lesquereux’ specimen, figure 16, plate 75 (1879-80)
and repeated on figure 6, plate 5 (1883), belongs to Cala-
mites schiitzeiformis and should not be placed in the
synonomy of Calamites cisti as Dr. White indicated on his
page 149, 1900.

Occurrence. — Wilkesbarre, Pennsylvania (Brongniart,
1828); Kimberly, Ohio (8075, 8028, 8064); Grand Ledge
and Corunna, Michigan (Arnold, 1949); Pitcher’s coal
bank (U.S.N.M. 5424), Henry County, Missouri (White,
1900). In Canada: several localities, Cumberland Group;
1 locality, Riversdale Group, northern Nova Scotia (Bell,
1944) .

Calamites cistiiformis Stur, 1877
Plate 11, figure 6; Plate 18, figure 1

Calamites cistiiformis Stur, 1877, Culmflora, II], Abh. K. K. Geol.
Reichsanst., 8, p. 94 (200), pl. 4 (21), figs. 5, 6.

Description. — Stems 3.3-9 cm wide with internodes 4-5
cm long; nodes slightly constricted with or without branch
scars; several branch scars may occur at one node with not
more than eight ribs effecting each scar. Ribs straight,
longitudinally striated, 1 mm wide, separated by deep fur-
rows; some ribs alternate at node, others continuous; rib
endings sharply pointed when alternating at node; leaf
scars small, rounded, on upper rib ends.

Discussion.—The species is represented in the Kim-
berly collections by several specimens although two are
larger and show more features in common with the Eu-
ropean specimens.

C. cistiiformis is separated from C, cisti in having some
of the ribs alternating at the nodes while others pass di-
rectly through. In C, cisti all the ribs alternate.

C. cistiiformis is similar to C. ramifer yet separated
on the basis of rib width and rarity in the occurrence of
branch scars. Calamites cistiiformis has much narrower and
straighter ribs.

Occurrence. — Kimberly, Ohio (8505, 7387). In Can-
ada: four localities, Cansco series, Pictou Coalfield, Nova

Scotia (Bell, 1940) and two localities, Cansco group, north-
ern Nova Scotia (Bell, 1944).

Calamites sachsi Stur, 1878
Plate 14, figure 4

Calamites sachsi Stur, 1878, Verhandl. K. K. Geol. Reichsanstalt, p.
327; 1887, Calam. Schatzl. Schichten, II, 2, p. 180, pl. 2, fig. 1.
Description.—Stem 8 cm in diameter, incomplete;

outer surface smooth with fine fibrillous, longitudinal stri-

ations. Ribs inconspicuous, noticeable only near nodal line,
slightly raised at upper ends; upper ends rounded with
small leaf scars | mm in diameter.

Branch scars whorled, five evident, 1.0 cm in diameter,
7-9 mm high, all same height, rounded.

Discussion and occurrence. — A single specimen, Num-
ber 8000-3, from Kimberly, conforms to the general descrip-
tion of the European species. This is the only specimen that
I have seen in all the American collections.

Calamites schitzeiformis Kidston and Jongmans, 1913
Plate 11, figure 7; Plate 14, figure 3

Calamites schiitzeiformis Kidston and Jongmans, 1913, forma typicus
Kidston and Jongmans in Jongmans and Kukuk, Cala. Rhein.
Westf. Kohlenb., No. 20, p. 32, text fig. 2.

Calamodendron ? species Lesquereux, 1880-84, Coal Flora, vol. I, pp.
32, 33, pl. 75, fig. 16.

Calamites approximatus Lesquereux, 1883, 13th Ann. Report Indiana,
II, p. 40, pl. 5, fig. 6.

Description. — Stems 4-6 cm wide; internodes irregular
in length, 6-13 cm; nodes contracted; ribs straight, some-
times alternating at nodes, narrow, 10-13 ribs per cm.

Leaf vascular scars on upper rib endings circular, small,
up to 0.33 mm in diameter; smaller scars on lower part of
rib occasional, only two to three per node.

Discussion. — Lesquereux’ Calamodendron species, il-
lustrated on his plate 75, figure 16, which he later called
Calamites approximatus with the same figure repeated
(1883, pl. 5, fig. 6), is from Cannelton, Pennsylvania. This
specimen conforms to the general description of Calamites
schiitzeiformis.

Because there are so few specimens representing this
species in the United States, their separation into forms is
unwarranted. Therefore the Ohio and Pennsylvania speci-
mens are lumped under the general name, as does Dr.
Arnold for his Michigan specimens (Arnold, p. 181, 1949).

Occurrence. — Kimberly, Ohio (7360, 7087); Grand
Ledge, Michigan (Arnold, 1949) ; Cannelton, Pennsylvania
(Lesquereux, 1879-80, 1833).

Calamites suckowi Brongniart, 1828
Plate 11, figure 10; Plate 15, figures 3-5

Calamites suckowi Brongniart, 1828, Histoire, 1, p. 124; pl. 15, figs.
5, 6; pl. 16, fig. 2.

Oun10 FREEPORT CoAL Lycopsips AND SPHENOPSIDS: ABBOTT 23

Calamites cannaeformis Dawson, 1871, Rept. Geol. Sur. Canada, p.
26, pl. 4, figs. 47, 48.

Description. —Stems large, up to 15 cm wide with
nodes not contracted; internode length variable from 15
mm up to 9.5 cm; branch scars not common, irregular,
large.

Ribs straight, longitudinally striated, average 2 mm
wide; some ribs pass through without interruption; other
ribs alternate at node with upper endings bluntly rounded
and conspicuous oval leaf scars; lower rib endings may or
may not show circular leaf scars.

Ribs of rhizomes as in stems but rib endings with
sharply pointed terminations. Nodes with verticils of root-
let scars.

Discussion. — Calamites suckowi is characterized by its
wide ribs, some of which do not alternate at the nodes; by
rib endings which are more or less rounded; and by the
large conspicuous round to oval leaf scars on the upper
rib endings.

Branching in Calamites suckowi is usually sparce. How-
ever, a specimen from Pitcher’s coal bank, Henry County,
Missouri, (U.S. National Museum, No. 5425) has branch
scars occurring at every two or three nodes. Number 8610
from Kimberly has an unusually large branch attached.

Calamites suckowi is common and is reported from
most of the collecting areas. Because it is so widely dis-
tributed, it, therefore, has little stratigraphic significance.
Stopes, 1914, followed by Bell, 1944, placed Dawson’s Cala-
mites cannaeformis in the synonomy of C. suckowi.

Lesquereux’ plates, pertaining to the fossil plants and
referred to in the Geological Survey of Kentucky (vol. 4,
p. 435) were not published. I do not know upon which
specimen or specimens Lesquereux based his description of
Calamites decoratus. In the absence of a figure, as well as a
specimen, I shall not place this species.

Occurrence. — Kimberly, Ohio; Illinois, several local-
ities (Noé, 1925 and Janssen, 1939) ; shales above the Grand
Ledge coal, Michigan (Arnold, 1949) ; Pitcher’s coal bank,
Henry County, Missouri (White, 1900); several localities
in Pennsylvania (Lesquereux, 1879-1880) ; Green County,
Indiana (Canright, 1959); LeRoy shales, Kansas (White,
1903). In Canada: St. John beds, St. John, New Brunswick
(Stopes, 1941); Joggins, Nova Scotia (Matthew, 1906) ;
many localities in Nova Scotia (Bell, 1938, 1940, 1944)
ranging from the Riversdale through the Pictou Series.

BIBLIOGRAPHY

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Andrews, H. N., Jr.

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1951. d new American species of Bowmanites. Bot. Gaz., 113(2),

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Arnold, C. A.

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Barthel, Manfred

1958. Stratigraphische und paldeobotanische untersuchungen im
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Baxter, Robert W.

1950. Peltastrobus reedae: a new sphenopsid cone from the
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Bell, W. A.

1938. Fossil flora of the Sydney Coalfield, Nova Scotia. Canada
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Brongniart, A.

1822. Classification. Memoirs du Museum d’histoire Naturelle,
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Browne, !. M. P.

1925. Notes on the cones of the Calamostachys type in the
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1927. A new theory of the morphology of the calamitean cone.
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Brush, Grace S., and Barghoorn, E. S.

1955. Kallostachys scottii: a new genus of sphenopsid cones from
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346-356.

24 PALAEONTOGRAPHIC:

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Onto FREEPORT CoA Lycopsips AND SPHENOPSIDS: ABBOTT 95

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PLATES

98 PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 1

Figure

1-4. Bowmanites incurvata Abbott, n. sp.
Mazon Creek nodule, University Illinois, No. 8 owing general overall aspect
of cone. 2. Reverse half of nodule shown in figure. 1. 3. Enlargement of figure 2 to
show: a, the ribbed axis; b, sporangiophore; c, bract; and d, the sporangium. 4.
Detail of one sporangium, showing the elongated wall cells, not unlike those shown
on Plate 7, figure 5, for Calamostachys germanica; X 13.3.

5. Palaeostachpa elongata (Pres), Weiss) stccccsscsccssscsceuesstecctateasev-btsteasece ss chdvaccsai ovata t Neen 18
Kimberly, Ohio, No. 8690. An enlargement of a part of the lower cone at figure 7.
The sporangiophore (b) is situated slightly above the bract (c). Two of the four
sporangia (d) in a group are also evident.

6; Galamostachys recurvata Abbott, 1. Spe .caccteacrcdosesssacsasersssnnisnvansrevencuoveycueervateeess ved tana mneanes 17
Kimberly, Ohio, No. 7089. The bracts (c) sag downward, away from the sporangial
group (d). The sporangiophore (b) is attached near the axil of the bract. The
internode is 4 mm long; X 7.5.

Palacostachya_ elongata (Presl'), WiC1SS)-0.csc.-ssscesccosssexcestesessaneseassceoenserservansnarecareasaeahaetekes kana 18
Kimberly, Ohio, No. 8690. Three cones, two of which are attached to a calamitean
axis.

ba |

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 1

nn

>
—
-
>
—
-
-

i

aa

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Plate 2

Ou1o FREEPORT Coat. Lycopsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 2

Figure Page
Dem AD SEL G/B EMEA CLO CEE eNO OLE RI GR Yo ey cy ceo nese ceuesesiecetensnfuntpraxatiisavarvevevensevekesssicarecuatspvareessnnee 20
Kimberly, Ohio, No. 8718.

Sue Lalacostachyanovalis \((besquereux Ex AbbOtt)) .c.:.:-cccasccosecesoessstashessesudscccesnsosusisconsnenenssenpuantan 19
Alton, Penna., Lacoe collection No. $52f and U.S.N.M., No. 18055. One of three cones
widely separated on one piece of shale, none of which was definitely designated
type 3. The other two cones on the piece of shale, as in figure 2.

NS QAO SIALILV LAITY EGHIGEE DOLL Dy SPier accceccnncassascasesstscsade tencssescheyavasveutes assvorspcceateasbasncsusausvevese 20
Kimberly, Ohio, No. 8719. Same cone as shown in 4; b, indicating sporangiophore ;

c, bract; and d, sporangium.
GeCalammasiachysmiomgturacteatar OW DOtt Mees pes arccccesecsrsscenas-scnce’can-sseaccstsensvysassoscarcsseccathavsstansuasese 17

Kimberly, Ohio, No. 8526. Overall aspect of cone identical to that shown by Janssen,
1939, and Baxter, 1963. 7. Enlargement of figure 6, to show at b, sporangiophore;
c, bract; and d, sporangial group.

50

Figure

1,2.

34.

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 3

Calamostachys tuberculata. (Sternberg), W189 rsscssasssvossotessccacseecacssesfucctecsventaerenseneenneeareye
Kimberly, Ohio, No. 7254, natural size, transfer. 2. Same specimen as figure 1, en-
larged & 4. At c, free portions of bracts; b, the sporangiophore; and d, the
sporangium.

Palaeostachya aperia:. (Lesquerewx: Ex ADDOtE)) sreccctessssssscnnnsstevessyerenensescnseveestets senna eee
Lacoe Collection, No. 950, natural size, type of Volkmannia fertilis Lesquereux.
4. Enlargement of a portion of the reverse side of Palacostachya aperta, Lacoe No.
950, shown in figure 3. The apical whorl shows the bracts united into a wide collar.
The projecting teeth on the margin of the collar indicate the only free portion of
the bract whorl. Scale included.

Galamostachys tuberculata (Sternberg), VWess sc.csssss.ccs-scsssceeccverdessivecsecsesscevesevevesssracsieeernete
Further enlargement, 20, of figure 1 showing sporangial groups (d) inserted mid-
way on the internode, their orientation to the sterile bracts (c) and the cellular detail
of the sporangium wall.

Palaeostachya aperta (Lesquereux ex Abbott) ......0.....scccccccsseseeccssnscsseseacsssensstorassevotssnsescaseenyan
Same specimen as shown in figure 4, but further away from the apex to show the
relationship of the bract whorl (c) to the sporangiophore (b).

Plate 3

IP ,

Woltection of R,D.. Lathe, >

me

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OHIO FREEPORT CoA Lycopsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 4

RUA OSLAC AY Ema per Iga MhESQUELEUX (6X) AUDDOtE) ict icsectessstarthessansecshstesasesest<usscveseccsuscecsoseoetvenive
The type of Lesquereux’ dAnnularia tuberculata, Lacoe No. 914b. -M., No. 18147,
natural size. The cone shows the base and six sterile internodes ‘before the fertile
area where the sterile bracts and sporangiophores are in alternating verticils. The
teeth on the margin of the collar, or united bract whorl, are longer on this speci-
men than on the one shown on Plate 3.

Lesquereux believed that both the cone and the calamitean axis on the left, in
both figures, belonged to the same plant. Photographs courtesy of the U.S.N.M.

Page

32

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 5
Figure Page

1,2. Palaecostachya aperta (Lesquereux ex Abbott) ....-sssscsesssesssssssecsnecsnscenectneenueecsicesunssnneceneccsis
Lacoe Collection, No. 914f, U.S.N.M., No. 18152. This is another specimen labeled
Annularia tuberculata Lx., from the Brewery Cut, West Pittston, Pennsylvania.
Natural size. 2. Lacoe Collection, No. 914e, U.S.N.M., No. 18151. One complete cone
with its sterile base and apical portions of two other cones of the same species. From
the Brewery Cut, West Pittstcn, Pennsylvania. Natural size.

Both photographs courtesy of the U.S.N.M.

Plate 5

PALAEONTOGRAPHICA AMERICANA, VOL. VI

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PALAEONTOGRAPHICA AMERICANA, VOL. VI

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5-8.

Onto FREEPORT CoA Lycopsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 6

BORUIMIATLILES TLAG7ain (UES QUeETE UIT EX CANDID OTE) | coretetsntcccersfedoctstiesssvatccusasscsesussancuosssnveveseusserees
Kimberly, Ohio, No. 8691, scale included. Verticils of sporangiophores b are external
to groups of sporangia d, and the bracts are not shown.

Bic ai thas ania guigom WEESO VER ENX.?) laIN OC yaccvsss seve issccecsstatetscstucvsercsteeseuesaacatenccstssstessocaddsscsadrasnees
University Illinois, No. 1284. Scale included. A new photograph of parts of two
cones originally illustrated by Noé. The bract verticil ec is divergent from the
sporangial groups.

Galammosaci Apis POnaag tara AN DDOU A Mew SP ay ecerecs sory recs“ eds ceeceanosnccine ccsansatceveseceveshscarssavesscstacere
Kimberly, Ohio, No. 8555. The sporangiophore is shown at b, midway between
verticils of bracts; c, the bract whorl; and d, the sporangium.

Baowrmanmtesmnagma (Mesqueceuxc) ex ADbOLE)| i ..-cacccescsessesecoscvts-sssnsvaccenvurestasoivere sauctanacsnessdves
Kimberly, Ohio, No. 8532, scale included. Portions of two cones are partially
skeletonized. The sporangiophores and axis remain. The darker sporangial-like areas
at e, are masses of spores, and only fragments of the sporangial wall remains.

VEG RIASACCIE SERCO MUP GLEAN DOUG IID ee SPs avscea sarees ee aia cac ak sahara asasos soscs ae vadaedaiunvassereretecaomnececeeeentesecuees
Kimberly, Ohio, transfer, scale included. Parts of two skeletonized cones show at b,
the sporangiophore and at c, the bract. 6. Kimberly, Ohio, No. 8539, scale included.
Five cones are attached at one node and parts of three cones are in close association.
7. Kimberly, Ohio, transfer, scale included. The transfer shows part of two cones
in the midst of, but not connected to, leaves of Annularia sphenophylloides
Zenker. 8. Kimberly, Ohio, No. 8540, scale included. Several cones are attached
at each of the successive nodes.

16

15

34

Figure

1-5;

6,7.

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 7

Calamostachys germanica WeASS ...c.s.secssscsssessscrssescssssecesssscssscseneansenenesenesensnenencassenessssacssanesasnenes
Kimberly, Ohio, No. 8862, scale included. Overall appearance of cone with most of
the bracts remaining. 2. Kimberly, Ohio, No. 8045; x 3. At d, position of the spo-
rangial grouping, and c, several bracts around one sporangial grouping. 3. Kimberly,
Ohio, No. 8861; 3. At b, sporangiophores inserted midway between bract whorls.
4. Kimberly, Ohio, B5606-26, 3. Several bracts enclose one sporangial grouping.
5. Kimberly, Ohio, No. 8045; 10. Enlargement of a part of figure 2 with details
of one sporangial group, especially the sporangium wall cells.

Calamostachys interculata Abbott, m. SP. ......ccssssssssecsssssssessssssssnserneesneenecsnecnseensesseanecenscsneensonssseye 18

Kimberly, Ohio, No. 7413, with Neuropteris heterophylla, scale included. At d,
sporangial grouping midway on the internode between two verticils of bracts. 7.
Kimberly, Ohio, No. 7413, same specimen shown in figure 6, scale included. At b,
the sporangiophore; c, bract; d, sporangium; and e, oval scar midway between bract
whorls and indicates insertion area of deciduous sporangiophore.

=
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PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate &

Figure

1,2.

On10 FREEPORT CoAL Lycopsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 8

larger axis belongs to genus Calamites, but because the nodal details are not
present, no specific identification can be made.

35

36

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 9

Figure Page

1.

2,3.

4,5.

Palaeostachyatrabeculata: Abbott, mais pias cccsnssesessatotersvscaisvesssusceteceasretendtttttneet ete ate ceived tr aee ee 20
Kimberly, Ohio, No. 8718. Spores smooth, 27.5-37.5 “ in diameter, show wrinkling
and a triradiate apparatus.

Palaeostachya aperta (Lesquereux ex Abbott)
See Plate 3, figure 3; spores smooth and 60-80 # in diameter.
from the same cone as figure 2.

Calamostachys germanica WEISS .........:01.+-sscscecscssss-s--sossesreeeeeeceseseoe-ee Seis ea ee 16
Wheatland, Indiana, WI-1, spore tetrads, with spores 47.5 * 60 “ in diameter and
showing a granulose surface. 5. Spores form the some cone as figure 4.

Calamostachys) tuberculatay (Sternberp)) sv eiss eee ie ee ee 15
Kimberly, Ohio, No. 8530. Sporangium wall cells showing inwardly directed buttresses

or wall thickenings. The same cell type is found in all of the sporangium walls of the
genera of articulated cones.

Calamostachys germantca’ Weiss. “sdscsccecroutseunecencunrtrencn ates taeaimerareiaientetenern ner ee . 16
Wheatland, Indiana, WI-1. Spore 50 u in diameter showing typical wrinkling and
triradiate apparatus. 8. From the same cone as figure 7.

Bowmanites; magna (Lesquereux ‘Pex CA DbOott)| miscssernestusteretectssirenseenseueerteemeectersctaesveranunsraeet 14
Kimberly, Ohio, No. 8532. Spores 42.5-67.5 in diameter, are wrinkled with a
triradiate apparatus and appear to have remnants of an outer granulose surface. 10,
Kimberly, Ohio, from cone No. 8532. 11. Kimberly, Ohio, from cone No. 8691. 12.
Kimberly, Ohio, from cone No. 8504. 13. Kimberly, Ohio, from cone No. 8532. 14.
Kimberly, Ohio, from cone No. 8532. 15. Kimberly, Ohio, from cone No. 8532. 16.
Kimberly, Ohio, from cone No. 8504.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 9

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 10

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Onto FREEPORT CoAL Lycopsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 10

Camera lucida tracings combined with reconstructions.

Figure Page

ik;

10.

Galamostachy smrmtemei lata DDOtes I. WS Psy ecescrceens cect cpecvivssscessaesteatssretveiseecessasssececsakescieoseoeteasereee 18
Kimberly, Ohio, No. 8531. At d, one of four sporangia in a group; b, the sporangio-
phore split at its apex into four tiny prongs, each of which is attached to a single
sporangium; c, is the sterile bract whorl; and e, is the oval scar indicating the pre-
vious attachment of the deciduous sporangiophore.

Calamostachys tuberculata (Sternberg) Weiss ....
Kimberly, Ohio, No. 8530. At d, the sporangial grouping of four, each of which is
attached apically to the sporangiophore b. The sporangiophore is not enlarged at its
apex. C, indicates the sterile bract whorl.

CAAT LOSEAG/Ey SITE CUTUGLE GA DORE WINS Det sescensceztoazsnesercantensesase ctvessocnee esvienedenese-ureeedon cioctanavastensree 17
Kimberly, Ohio, No. 8327 transfer. At d, the sporangial grouping, four of which are
attached apically to the sporangiophore; the sporangiophore b, is attached to the
axis below the midportion of the internode; and c, the bract whorl.

GGL Aa OS TAGHAS STULL) SOD DOCH, Te 1S Psy reeseecsreree cesesereastescvers cst nenascrasccescerersctcetorsoseaeccer an eetotreeene erates 17
Kimberly, Ohio, No. 8325 transfer. At d, the sporangial grouping, four of which are
attached apically to the b, sporangiophore. The sporangiophore is attached above
the midportion of the internode; and c, the sterile bract whorl.

GULALASLAChY LO GLUT aActeatal ADDO Hie SPs) sssercacecersesatsntedaeesecvon-2 oes cevstarsseeeccierevsrsecerscecnovdeoon 17
Kimberly, Ohio, No. 8526. At d, only two sporangia are in the sporangial grouping
and are attached laterally to the sporangiophore. The sporangia are vertically ar-
ranged, that is, one above the other. At b, the sporangiophore is split apically into
two prongs, each of which is attached to a single sporangium. C, is the sterile bract
whorl.

CClamaslaciysny tS parangtara yA DbOLty Wer SPiceeeccereertcnsetersctrasessecteetecesemnesicasterecedesterseasteteeattarevense 16
Kimberly, Ohio, No. 8821. At d, two sporangia are attached apically and straddle

the apex of the sporangiophore. At b, the sporangiophore and ec, the sterile bract
whorl.

(QIATEOSTACHASENG ELTECTILCH A NV CLS essecesercetsenaettenencenceuceterescteckeed tenieokcscascsnpseseotisevsereassasiveercavetsate 16
Kimberly, Ohio, No. 8045. At d, four sporangia are attached at the slightly en-
larged apex of the b, sporangiophore. At c, a single bract, although several bracts
protect one sporangial grouping.

Palacostachyavapenta’ (Wlesquereux ©x ADDOtE)) Vrsc.cecczscecescsocesssscerccecvnsusonsssyeteecersivecustvaetsssesests 19
Lacoe No. 950. Only two sporangia are attached to the sporangiophore and the spo-
rangiophore b, is enlarged apically into an elongated bar, the ends of which are

the attachment areas of the sporangia. Basally, the sporangiophores are attached

to the axis in the axil of the sterile bract whorl. At c, the sterile bract whorl is
united into a continuous collar; the ribs are vascular bundles which terminate in the
teeth at the upper margin.

PAAR O SACHS I POLOIMG ALG (SEALESI)) MRVWICLSS) cscesancncevexsesereorthrackoe<csex-<cencissasxetiveratue /osecsevexcessisvartsissere 18
Kimberly, Ohio, No. 8690. At d, four sporangia are attached apically to the spo-
rangiophore; b, the sporangiophore is situated in the angle of the subtending bract
whorl shown at c.

Palaeostachya awalis, (Wesquereux ex: ADbOtt) 2..-....cscccncecececcsvesecervesecsseyensscseccosscssseacotventivecese 19
Lacoe No. 852f. At d, four sporangia are attached apically to the slightly enlarged
apex of the sporangiophore b; the sporangiophore is situated in the axil of the sub-
tending bract whorl at c.

37

58

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 11

Camera lucida tracings combined with reconstructions.

Figure Page

Ls

10.

Bowmanites magna (Lesquereux ex Abbott) ......ccccsescesesessnsesesensescnsncescscneseecnsnanscsenensesenensnsnss 14
No. P 1284, Mazon Creek, Illinois. At d, the sporangia; c, the bracts, and b, the
sporangiophores. X is the transverse view of : at 1, the upper whorl of sporangia;

at 2, the lower whorl of sporangia and their relationship to the sporangiophores;
and at 3, the basal part of the whorl of sporangiophores and their attachment on

the lower part of the sterile bract whorl.

Bowmanites incurvata Abbott, m. SP. .....cscescecesceseecsssesecseretseesensensesseneassssessnensnsrsnasasneeasananensnsnys 14
Kimberly, Ohio, No. 8446. At d, four to six sporangia are singly attached along the
interior face of the sporangiophore from near its base to its apex; b, the inwardly
curved and elongated sporangiophore, note that the sporangiophore splits off a few
millimeters from the base of the bract c.

Mazostachys compacta Abbott, n. SP. .s.ssessssessecsssesssesseesescenesesestenssneestncensnseneenssscnsesensensnvansanensens 15
Kimberly, Ohio, No. 8539. At d, two sporangia are in the sporangial grouping and
are in vertical alignment. They also are apically attached to the sporangiophore. At

b, the sporangiophore is attached immediately below the sterile bract whorl; at ¢,

the bract whorl is horizontal for a few millimeters then abruptly bends upward to
protect the sporangial group.

Palacostachya trabeculata Abbott, 1. SP. ..ssccscsesssssecssereeenececsnerecsnsnensnsrersnsscsnarensvenenseseesarenensennns 20
Kimberly, Ohio, No. 8718. At d, four sporangia are in a group and are apically
attached to a bar-shaped enlargement of the apex of the b, sporangiophore. The
sporangiophore is attached in the axil of the bract whorl, c.

Galamites. ctstt! RroDGMiart <ciscccssseececcetccrectesecscess-nductancuanaspusncacvenesuvestuonnnencsynevesevebNesrnssesevsbazesines 21
Kimberly, Ohio, No. 8075. A detailed sketch of the nodal line and angular arrange-
ment of the rib endings. Note the alternating arrangement of the ribs. The small
circles near the apex of the lower ribs indicate previous leaf attachment areas, or
vascular strands. The ribs are 1-1.5 mm wide.

Calamites cistiiformis Star
Kimberly, Ohio, No. 8505. A sketch of the nodal line and angular rib endings. Small
circles indicate previous vascular areas and the larger circles indicate previous at-
tachments of larger axes; the ribs conyerge at the rim of the larger scars.

Calamites schiitzeiformis Kidston and Jongmans -
Kimberly, Ohio, No. 7360. The zigzag nodal line is formed by the a ternating ribs;
the ribs are 1-1.5 mm in diameter. Small vascular scars occur on most of the lower
ribs and occasionally on the upper rib endings as well.

Galamites carinattts, Sternberg sere ccccscceteeettenctnsscatnatersevnessecansnotonste isahapaneceesores eentecnesreatoueerenees 21
Kimberly, Ohio, No. 7422. The nodal line is zigzag with some ribs alternating while
others do not. The prominent larger scars are more or less typical for the species.
Note the one circular scar on the lower part of the upper two ribs which probably
represents a previous root attachment. The ribs are 1-1.5 mm wide.

Gallamites wupedulatis® Stern bergen csc ccecsectereereeret atin nnoaeee ities cccaet aarnee caacieaeseciae carrera nearer 21
Kimberly, Ohio, No. 5608-8. The nodal line is zigzag with ribs usually alternating.
The ribs do not always undulate and may be as straight as those of other species.
The ribs are up to 2 mm wide.

Galamites suckowi Brongmiart: .ssc.c.scscssssscscecsscccsssssscsscsscosssescnssesvenenssesnsseneneerensescenccnsnesentesesentende 22
Kimberly, Ohio, No. 7451. Although a nodal line is evident not all of the ribs alter-
nate at the node but pass directly through unaltered.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 11

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PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 12

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2

Outo FREEPORT Coat Lycorsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 12

Figure Page

1.

2,3.

10.

11.

12.

Sigillaria pileata Abbott, 1. SP. .s..sssssessssesssssecssssecsseecssnsesnvesesescecaneceenucccsuriccenssnonssssenssssnnancentseesets 13
Kimberly, Ohio, No. 8791. The leaf cushion is + mm wide and as wide as the ribs.

A small patch of wrinkling is immediately above each cushion; vertical striations
and wrinkles extend between the cushions which are 14 mm apart.

Sigillaria mamillaris Brongniart ...
U.S.N.M., No. 16500. In this specimen the leaf cushions are 5-5.5 mm wide and sep-
arated by + mm of transverse wrinkles. In 2 a with further decortication, the vascu-
lar area remains about the same size and has the same position on the rib. 3. Kim-
berly, Ohio, No. 8057. This specimen from Kimberly shows the leaf cushions so
close together that the lower one seems to distort the base of the cushion above and
causes a deeper horizontal wrinkling. The ribs appear to be undulating in order to
accommodate the closely spaced cushions. The cushions are 5 mm wide. In 3 a, with
further decortication, the vascular area is in the same position as those on the upper
surface.

Sigillaria laurenciana (Lesquereux ex TNS so § 3) Yi eee ete oe ee eo ores eer 12
U.S.N.M., No. 16571. The leaf cushion occupies over one-half the diameter of the

rib which is up to 3 cm wide. The cushions are only 10-13 mm apart vertically.
The area between the leaf is only slightly striated and wrinkled.

Sigillaria elegans (Sternberg) Brongniart
Kimberly, Ohio, No. 8354. The nearly six-sided, closely spaced, leaf cushions give
the ribs a definite undulating appearance. The leaf scar area is above the middle of
the cushion and two vascular bundle scars, as well as a parichnos scar, are well
preserved. X 5.

Lepidophloios larcinus Sternberg. --..------.----:--------e-ccpee eect ence ennnaenase 9
Kimberly, Ohio, No. 8715. The hexagonal leaf cushions are so closely fitted together
that there is little or no space separating them. A prominent horizontal wrinkle
occurs on the lower part of the cushion. The leaf scar area is slightly above the
middle of the cushion; X 5.

Lepidodendron cf. jaraczewskit Zeilher ......---sssesssisececses secs ssseennnssiceseneest 7
Kimberly, Ohio, No. 8320. The leaf cushions are greatly elongated, 26 mm, and
tapering to a long acute angled base. The leaf scar area is in the widest part and

near the apex of the cushion. The vascular scars are not clear in the Kimberly speci-
mens. The drawing is only one half the scale of the other drawings.

Lepidodendron aculeatum Sternberg ore
Kimberly, Ohio, No. 8622. The diamond-shaped leaf cushions are assymetric in out-
line. The leaf scar area is more or less triangular but the vascular scar is not clear

in the Kimberly specimens. A keel line extends from below the vascular area to
base of the cushion.

Sigillaria orbicularis Lesquereux Collection oo... recssceeensensnnnnnsesaeasssssesrsersste 13
Lacoe No. 616A, U.S.N.M., No. 16489. The cushions are wider than high with a
smooth surface. The vascular and parichnos scars are near the apex of the cushion
and are prominent. The surface of the rib between the cushions is smooth; 5.

Sigillaria rugosa Brongniart
Kimberly, Ohio, No. 8621. T
ings between the leaf cushions. Prominent horizontal wrinkling occurrs immediately
below the cushion and above the next cushion in the vertical series, although wrink-
ling occurs the full distance between the cushions; 5.

Lepidodendron scutatum Lesquere ux corssr-svscseeceesseereccssscssccunesscennnneennnspaseetissrsent sss ob oy
Kimberly, Ohio, No. 8589. The leaf cushions are diamond-shaped with four to five
prominent deep, horizontal wrinkles below the leaf scar. The leaf scar is wider
than high, above the middle of the cushion, and a ligular (?) scar is near the apex

of the cushion. The apical part of the cushion is raised; X 5.

Lepidodendron amdrewwsi Lesquere ux .n....sscssssssssscssstesesereeeeenssnnsesssssscsssneurecsseccannnnacecsssissassssine 6

Kimberly, Ohio, No. 8588. The leaf cushions are bluntly diamond-shaped with a
smooth surface except for the leaf scar area. The leaf scar area is wider than high
with vascular scars small and circular, and the parichnos scar is larger. A ligular
scar is about midway between the leaf scar and the apex of cushion; X 5.

39

Figure
1.

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 13

Asolanus Campiotaentae Wood scccsccsssvacsasccsiviseocpusssazepscndscaystiioedayobuuctns ustaeteveeeaned nance catnbedeseaevirecees
Kimberly, Ohio, No. 6106. Photographed on plate 69, David White's Monograph 37.
The leaf cushions are equidistant with diagonally placed wrinkles seemingly connect-
ing all of the cushions. The vascular area is circular with no separation of bundle
scars nor parichnos scars; x 5.

Asolanwus-monostigma. Wesquene ux tacts 2c ces ccanccscacenseerete ce ena a ee
U.S.N.M., No. 16415. The leaf cushions are closer together horizontally than in 4.

camptotaenia but further apart vertically. Wrinkles seem to connect groups of four
scars and are elliptically aligned. The specimen is illustrated in Lesquereux’ Coal
Flora, plate 73, figure 3; X 5. (See Synonomy in Text.)

Asolanus ‘sigillariotdes: (Lesquereux ex Abbott), oi.-cscecscscccaqntsecnececeececavsorecaeceesivonscsetaxpupunvasncee
Kimberly, Ohio, No. 8650. The leaf cushions are far apart, and the surface between
the cushions is smooth. The leaf scars occupy the upper one-half of the cushion.
The vascular scars and parichnos scar are prominent and the ligular scar is above
the vascular apparatus; X 5.

Stgthlaria\ Tacaet LeSquere x: <cs.c.cacusscxcsesccecssehese-stseqeesonacsia ye epiesateseansesesastsedionsananeeteehyeooen eater neentee
U.S.N.M., No. 16811 and Kimberly, Ohio, No, 8649. The leaf cushions are large and
heart-shaped, with the apex notched. The vascular apparatus scars are slightly
above the middle of the cushion. On decortication, at 4+ a, the two vascular scars are
vertically ovoid and touching and on further decortication, at + b, the vascular area
is a single one and heart-shaped.

Bebidophiloios dilatakus: UT esque xe wx: csctesc0qna5-5-suvestsscscvesenaceteatuarcarracesnaysintuevetanpers teasevuaesureesesteae
Lacoe No. 748, U.S.N.M., No. 15944. The leaf cushion is narrow vertically but
elongated horizontally. The leaf scar area occupies the lower one-half of the cush-
ion and extends across the full diameter. The vascular scar area is in its center.
Dd ey,

Lepidophloios: samtiagemt Dis Witte. <2..ch-2cseencnccssn.tacnentssevacsnetonsvaracurasereenieaccas sett rouetneaee foseeeneaereten
Kimberly, Ohio, No. 8826. The species has the largest leaf cushion area of all the
American species. The cushion is much wider than high and the leaf scar area is
in the lower one-fourth of the cushion. The two vascular bundle scars are horizon-
tally elliptical and slightly above the parichnos scar; X 5.

Lepidophloios sp.
U.S.N.M., No. 15947. s
is in the lower part of the cushion and occupies only about one tenth of the cushion
area. The vascular apparatus was not clearly marked; X 5.

Lepidophlatos iS psx x ccescscaiestacirn. channels oes eee a Rane Cee aes
U.S.N.M., No. 6047. The cushions are wider than high and the leaf scar area occu-
pies the lower one-third of the cushion area. The vascular area is centrally placed
on the leaf scar, although individual parts of the vascular apparatus are unclear;
>, SF

Plate 14

PALAEONTOGRAPHICA AMERICANA, VOL. VI

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Figure
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Onto FREEPORT Coat Lycopsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 14
Page

CMTS TAY TURTLE TTAICETI I) ANDi BOLE Seer cee eee eee oer oe cece oe ee cee 21
Kimberly, Ohio, No. B5606-8. The four nodal lines show the typical rounded leaf
scars on the upper ends of the more or less undulating ribs. The internodes are
much shorter than wide. 2. Kimberly, Ohio, No. 8000-2. The specimen is figured less
than one-half the natural size. The internodes on the upper right side in the figure
shows the cortical tissue still preserved and horizontally wrinkled. The rib positions
are not reflected on this surface.

Kimberly, Ohio, No. 8000-3. The outer surface of the axis is smooth with fibrillous,
longitudinal striations. Five branch scars occur above the nodal line as indicated by
the sunken circular areas.

4]

Figure

V2

3-5.

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 15

Calamites carimatis, Sternberg cciiescsscssccerseseesisesnssssscecssecknyyaacsvassscievessddachi eenerinsaeaneenorcomsa dae einer 21
Kimberly, Ohio, No. 7469. The specimen shows a single branch scar at the node.
The other circular area is a damaged area due to preservation. 2. Kimberly, Ohio,
No. 8075. At the nodal line, the upper part of the ribs show elongate, ovate leaf
scars over 1 mm long.

Calamates cuckoqut Brom gmiargy ccisvcorsacsssvcsesccceessusscusyevecpregensativareevesctanveneeyevissses eb esenenees eaeeeee enna 22
Kimberly, Ohio, No. 8610. This specimen is complete in its diameter and a large
branch is attached on the left side. 4. Kimberly, Ohio, No. 8006. The nodal line is
accentuated by the upper rib endings with their prominent leaf scars. For further
details see Plate 11, figure 10. 5. Kimberly, Ohio, No. 8648. A transverse section
through a small limb shows a hollow central area, an encircling rib area also
partially hollow, and an outer cortical area.

PALAEONTOGRAPHICA AMERICANA, VOL. VI

16

Plate

PALAEONTOGRAPHICA AMERICANA, VOL. VI

At <s

A

ay

Ou10 FREEPORT CoAL Lycorsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 16

Figure Page

USE GALL CAT ACaT Te LPIEEL CATE SMES GOEL TER cose dec ese eoras ohetese (etl kedstevcondevstevbvivessnccacen stecshateenctctinns
Kimberly, Ohio, No. 8831. The coaly surface shows the leaf cushions separated a
little further than is seen on Plate 19, figure 2. The decorticated surface shows the
position of the vascular apparatus areas; x 1. 2. Enlarged from K 8057, shown on
Plate 19, figure 2.

Bh = SIN ce eih Grad BHTUC AAG LTE UTES CENTS os eepee eRe bgp ccee- oats eee
Kimberly, Ohio, No. 8000-1. The leaf cushions in the Kimberly specimens are essen-
tially the same as those specimens from Pennsylvania. However, the cushions on this
specimen are about as wide as high, while the ones shown on Plate 12, figure 9,
from Pennsylvania, are wider than high.

10

43

44

Figure
iS

3,4.

PALAEONTOGRAPHICA AMERICANA (VI, 38)

EXPLANATION OF PLATE 17

Page
Sigillaria orbicularts \LESquerevix: <cccccctecssecososeiseaccscdae einstse csp svasucaeedeqoneackcaebytetis cestetnee CSE eae 13
Kimberly, Ohio, No. 8000-1. An enlargement of figure 4+, Plate 16.
Stgtllaria: Pileata ABbOtE, Ms SPs cczcacexccctsanscvicesecteruccdstechesucoqpane cous axreesenssenn chet fusaeeeterse er anv taAN ICE! 13

Kimberly, Ohio, No. 8796. The distinguishing feature of this species is the wrinkling
of the rib surface immediately above the leaf scar. For further detail see Plate 12,
figure 1.

Sigillarta rtigosa Brome niart: z.::...ccevescutsscceesscvevsacaysecustoestoneeateasentetesaresncusCeatetyesanrteesaes Exe TOE
Kimberly, Ohio, No. 8652. The reverse side of this specimen shows the same type of
surface markings and leaf cushions. The specimen is complete in diameter and rela-
tively small in comparison with one of the Lesquereux specimens which is 20 em
in diameter and incomplete. +. Enlargement of figure 3 to emphasize the “jet trail”
type of wrinkling both above and below the leaf cushions. For further detail see
Plate 12, figure 10.

17

Plate

PALAEONTOGRAPHICA AMERICANA, VOL. VI

i Se ce

fll

ip

T

ea
Fi.
“8

RAPHICA AMERICANA, VOL. VI

PALAEONTOC

eit

u]
‘ :

’ a

Bites Maem!

Pri LEL LLL Ars ayn?”

Figure
i

Oun10 FREEPORT CoA Lycopsips AND SPHENOPSIDS: ABBOTT

EXPLANATION OF PLATE 18

Calamites cistiiformis Stur
Kimberly, Ohio, No. 8505. Parts of four internodes are present. For detai
nodal line see Plate 11, figure 6.

Mea ptd od ered OemeCGgEE CA PTENTUEN GES ET UK. oeec¥ssossesececaccvectsteacecseccvenvseoscedevcneebasbidhnfiainsaseiisscigivesceuctiosse if
Kimberly, No. 8589. The figure is enlarged to show the typical keel wrinkling from

the base of the leaf scar to the base of the leaf cushion. For further detail see Plate

12) ties Ad:

Lepidodendron andrewsii Lesquereux .... 6
Kimberly, Ohio, No. 8587. The specimen shows the leaf scar occupying almost one-
half of the area of the leaf cushion. There is no special ornamentation on the sur-
face of the cushion. For detail see Plate 12, figure 12.

SGA LAIMA LGTEL PT R MAE ES VLET UK, so sec aces asscatoueesseesvops ovevteresaowtasesovsasteenl oovuecteca deer sreatbe a okexiass 13

Kimberly, Ohio, No. 8800. This specimen is decorticated and shows the circular
vascular apparatus. For detail of the leaf cushion see Plate 12, figure 9. Also shown
on the shale is Sphenophyllum oblongifolium.

45

Figure
Me

PALAEONTOGRAPHICA AMERICANA (VI, 58)

EXPLANATION OF PLATE 19

Asolanus, camplotaenta NWO cissessecusscasczcacssccssctvacacvapssnscsatnscasicsss1gashietae peeaaneee mM eee See 9
Kimberly, Ohio, No. 8828. This is a much smaller specimen than D. White’s shown

on his plate 69, Monograph 37. U.S.N.M., No. 6106. For detail, see Plate 13,
figure. 1

Sigillaria ‘mamillaris Brom Pi yark spacers sve-castcstseecstnvssetstesetsatasbas testacesahicssarensernees snare mane eaeiee ere 10
Kimberly, Ohio, No. 8057. See Plate 12, figure 3 for detail.

Shgmaria ficotdes (Sternberg) Bromgoiart Minc-ccc-csucscacasvcexexscatucsstccayescasvtncrey,venspencsateuaponenesr#ens 13
Kimberly, Ohio, No. 7466. A typical root with its circular rootlet sears.

Lepidophloios larcinus: Stentibeng yc, -scazsciccshacesecnecora eco raree eet eae tee one 9
Kimberly, Ohio, No. 8715. The coaly outer surface shows vertical rows of leaf
scars, while the decorticated lower surface shows the hexagonal closely fitted leaf
cushions. For detail see Plate 12, figure 6.

Plate 19

PALAEONTOGRAPHICA AMERICANA, VOL. VI

A
acadianus,
Lepidophloios 9
aculeatum,
Lepidodendron 2650

Alton, Pennsylvania 5,19
andrewsi,

Lepidodendron 12,18 6
Annularia 15, 19, 20
aperta,

Asterophyllites 19, 20

Calamites 22

Macrostachya 19, 20

Palaeostachya 3,4, 5,9,

0 6,19
Archbald,

Pennsylvania 5, 20
Arkansas 21
Asolanus : 9, 10
Asterophyllites 19, 20
Aurora, Missouri 21

B
bisporangiata,

Calamostachys 6,10 6,

16
Bouthillier seam 10
Bowmanites 6, 14,15
Braidwood, Illinois. 10,14
Brigg’s shaft . i153
Buctel, Ohio 11
Cc
Calamites 21-23
Calamodenron? sp. 22
Calamostachys 6, 15,
16, 18, 19, 20, 21
calathifera

Calamostachys 15
camptotaenia,

Asolanus 13,19 9

Sigillaria

(Asolanus) 9
Canada,

Bouthillier seam 10

Cansco series 22

Coalbrook seam 9

Cumberland

Group Ole

16, 19, 21, 22

Gowrie seam 10

Harbour seam bePAl

Joggins 23

McLean seam 19

Morien Series 14

New Waterford 21

Northern

Nova Scotia .. 7,9, 11,

16, 19, 21, 22

Nova Scotia 10, 11,

EPA OB

Old Queen Pit 11

Phalen seam 10, 16

Pictou Coalfield 16, 21,

PPA PR}

Riversdale Group 9, 21,

22,23

Springhill .. 11

St. Johns beds 23

St. John ........ 23

Stellarton series 16

Sydney Coalfield 9, 10,

11, 16, 19, 21

INDEX

Thorburn Member
of Stellarton

series .... 16

Tracey seam 16
cannaeformis,

Calamites . 23
Cannelton,

Pennsylvania 10, 22
Cansco series,

Canada 22
Carbondale,

Pennsylvania a
carinatus,

Calamites 11,156,

21
Chanute shales 10
Chester, Illinois 10
Chiefton Mine,

Indiana 16, 17
Christopher,

Illinois 11
cisti, Calamites 116, 21,
cistiiformis

Calamites 11, 18 6,

22
Clinton, Missouri 8, 10
Coalbrook seam 9
Colchester,

Illinois .. 9
compacta,

Mazostachys 6,11 6,

15
Cormophyta,

division 6
Corunna, Michigan 22
crassa, Volkmannia. 19, 20
cruciatum,

Lepidodendron 9
‘Cumberland

Group eo elal

16519; 21; 22
D
Dade County,

Georgia 19, 20
Danville, Illinois 21
decoratus, Calamites 23
Deepwater,

Missouri 8, 21
dilatatus

Lepidophloios 13 8
discoidea,

Sigillaria 11
EuQuoin, Illinois 9
E

Eastwood,
Michigan 14
elegans,
Sigillaria 12 6,11
elliptica,
Stigmaria . 14
elongata,
Palaeostachya 1,10 6,
18
Equisetales_ . 6
F
fertilis, Annykarua 20
Trochophyllum 20
Volkmannia 19, 20
ficoides, Stigmaria ..19 6, 13,
14

47

Variolaris . 13
fissa, Sigillaria 10
?Flavularia elegans 11

G
Georgia,

Dade County 19, 20
germanica,

Calamostachys 79°10

6, 16, 18
Gilkerson’s Ford,

Missouri 8, 9, 10,
14, 21
Gowrie seam 10

Grand Ledge,
Michigan 10, 21,
22, 23

Green County,

Indiana 23
Grundy County,

Illinois 7, 14,

15, 16, 17, 18, 21

H

Harbour seam
Henry County,
Missouri 8, 10,
14, 19, 21, 22, 23

11, 21

Illinois 23
Braidwood 10,14
Chester 10
Christopher 11
Colechester 9
Danville Pal
DuQuoin 9

Grundy County 7, 14,
15, 16, 17, 18, 21

Mazon Creek 6, 7, 14,
15, 16, 18, 21

Stark County 20

incurvata,
Bowmanites Tl 16}
14
Indiana 14, 21
Chiefton Mine 16, 17
Green County 23
Knox County 11, 16
Paoli if
Sullivan County 11
Terra Haute 16,17
Vigo County 9, 16,17
Wheatland 6, eB
infundibuliformis,
Macrostachya 19, 20
ingens,
Lepidodendron ul
interculata,
Calamostachys 7,10 6,
18
J
cf. jaraczewskii,
Lepidodendron 12) (G7
Joggins,
Nova Scotia 23
K
Kansas
Chanute 10
Lancing 8

Lawrence 12

LeRoy shales 1OS2i0
23

Michigan Valley 18

Severly shales,

Osage PA

Thayer 10

Kimberly, Ohio 7, 8, 9,
OMI asa.
15, 16, 17, 18, 19,

212220

Knox County,
Indiana 11, 16

L
lacoei, Sigillaria S265 151
laevigata, Sigillaria 13
Lansing, Kansas 8

larcinum,

Lepidophloios 12,199
larcinus,

Lepidophloios 6,9
laurenciana,

Sigillaria 12, 166,

12
Lawrence County,

Missouri 21
Lawrence, Kansas 12
Lepidodendron 6
Lepidophloios 8
Lepidophloios sp. 13 8
Lepidophytales 6
LeRoy shales LORD

23
lesquereuxii,

Lepidodendron 7
Localities in

Pennsylvania 5
longibracteata,

Calamostachys 6,17
Luzerne County,

Pennsylvania 20
Lycopsida 6

M
Macrostachya 19, 20
magna, Bowmanites 6, 8,9,

11 6,14

Calamostachys 14
Maltby,

Pennsylvania 13
mamillaris,

Sigillaria 12, 16, 19

6, 10
marginata,

Sigillaria 13
Mazon Creek,

Illinois Oi, 14,

15; 16507. Te
Mazostachys 6,15
McClelland’s Shaft

near Bellville 21
McLean seam 19
Metacalamostachys 15
Michigan

Corunna 22

Eastwood 14

Grand Ledge 10, 21,

22, 23

St. Charles —

Garfield mine 14

St. Charles 10, 14

Michigan Valley,

Kansas 18
minuta,
Calamostachys 8,10 6,
17
Missouri 21
Aurora 21
Clinton 8, 10
Deepwater 8, 21
Gilkerson’s Ford 8,9, 10,
14, 21
Henry County 8, 10,
14, 19, 21, 22, 23
Lawrence County 21
McClelland’s shaft
near Bellville 21
Owen's bank 21
Owen’s coal mine 8
Pitcher’s mine ...8, 10, 23
Mollusea 20
monostigma,
Asolanus 13
Sigillaria 13 9
Morien Series 14
N
New Waterford,
Canada 21
Northern Nova
Scotia S10.
16, 19, 21, 22
Nova Scotia TOMI:
22, 23
12)
obcordatum,
Lepidophloyos 9
obeordatus,
Lepidophloios . 9
obovatum,
Lepidodendron 8
Ohio : : 14
Buctel 7 ini
Kimberly 7, 8,9

10, 11, 12, 13, 14,
15, 16, 17, 18, 19.

21, 22, 23
Old Queen Pit : 11
orbicularis,
Sigillaria 12, 16,
L718 06303
ovalis,
Calamostachys . 19
Palaeostachya 2,10 6,
19
Owen's Bank,
Missouri es 21
Owen’s coal mine,
Missouri eee 8
P
Palaeostachya . ; (a ae
18, 20
Paleocormophyta 6
Paoli, Indiana 7
pedunculata,
Mazostachys 15
Pennsylvania 14, 21,
Alton 5, 19
Archbald 5, 20
Brigg’s shaft 13

48

Cannelton 10, 22

Carbondale 7

Luzerne County 20

Maltby 13

Pittston 5.12

Plymouth 12

Pottsville 5

Scranton 13, 20

West Pittston 5, 20

Wilkesbarre 22
Phalen seam 10, 16
Phytolithus 14
Picton Coalfield 16, 21,

22, 23
pileata,

Sigillaria 12, 17 6,

13
Pitcher’s mine,

Missouri 8, 10, 23
Pittston,

Pennsylvania Bye)
pittstoniana,

Sigillaria 13
Plymouth,

Pennsylvania 12
Pottsville,

Pennsylvania im

R
ramifer, Calamites 22.
ramosus, Calamites 21
recurvata,

Calamostachys 1,10 6,

ily
Riversdale Group... 9,21,
22, 23
rugosa, Sigillaria 12, 176,
13
Ss
sachsi, Calamites 6, 22
schlotheimiana,

Sigillaria 13
schiitzeiformis,

Calamites 11, 146,

22
schiitzeiformis

forma typicus,

Calamites 22
Scranton,

Pennsylvania . 13, 20
seutatum,

Lepidodendron 12, 18 6,

8
setifolium,

Lepidodendron 7
Severly shales,

Osage, Kansas 21
Sigillaria 9-11
Sigillarioides 9

Asolanus 13 6,10

Lepidophloios 10

Sigillaria

(Asolanus) ; 10
sphenophylloides,

Annularia : : 15
Springhill,

Nova Scotia 21... 11
Stachannularia _. 20
St. Charles —

Garfield mine 14
St. Charles,

Michigan 10, 14

St. Johns beds,

Canada .. 23
St. Johns,

New Brunswick 23
Stark County,

Illinois... 20
stellaris,

Sigillarioides 9

Stigmaria 9
Stellarton series 16
var. stellata,

Stigmaria ficoides 9
Stigmaria aa 13, 14
suckowi, Calamites 11, 15 6,

22, 23
Sullivan County,
Indiana . 11
Sydney Coalfield 9, 10,
11, 16, 19, 21
ili
Tennessee 11
Terra Haute,

Indiana . 16,17
tessellata,

Sigillaria 11
Thayer, Kansas 10

Thorburn member

of Stellarton
Series
thuringia,
Calamostachys
trabeculata,
Palaeostachya

Tracey seam

Trochophyllum

tuberculata,
Annularia
Bruckmannia
Calamostachys

tuberculatus,
Asterophyllites

tuberous,
Stigmarioides

U

undulatus,
Calamites

Upper Freeport
Coal

11,14
21

5

Vv
vaningeni,
Lepidophloios
Variolaria

vera, Palaeostachya

verrucosa,
Stigmaria

verrucosus,
Phytolithus

Virginia

Vigo County,
Indiana

Volkmannia

volkmannianum,
Lepidodendron

WwW

West Pittston,
Pennsylvania

West Virginia

Wheatland,
Indiana

Wilkesbarre,
Pennsylvania

wortheni,
Lepidodendron

13 6,8

RT-F

ei

XLIV.

XLV.
XLVI.

XLVII.

XLVIII.

XLIX.

Volume I.

II.

Il.

IV.

GINO ee B47) 99 Gilat. (TP psp ccarecesaccareceventsentecntevouscssconsesavesisvacscssccvascers

Type and Figured Specimens P.R.I.

INO pS SL RAN eps PP 059) DIS.) cccervesavsettusecavessivenacuesdvseosscsscosseoosss

Australian Carpoid Echinoderms, Yap forams, Shell Bluff,
Ga. forams. Newcomb mollusks, Wisconsin mollusk faunas,
Camerina, Va. forams, Corry Sandstone.

(UINGose 193) 1678 pps, 48 pl ai ycsatecsscscecesesccsssessageccosssscecvexoucucncussessonsovces

Venezuelan Cenozoic gastropods.

(UNioe 94198) 74270 Pps 39) PISes ccpsseceneceseancverecarexevssesnsacseaeeceuscvasess

Ordovician stromatoporoids, Indo-Pacific camerinids, Missis-
sippian forams, Cuban rudists.

COIN Gas 99=203)i0 S65; ppc, (GSuIDIS® 2-ccocschvsccsoserestesssvassecsecrsesseabonsecsien

Puerto Rican, Antarctic, New Zealand forams Lepidocyclina,
Eumalacostraca.

CIN Gp 2k) OOF DDS MOS PINs. <. cescscecocnssseccabransserstususdascabinnecesCoskponceves

Venezuela Cenozoic pelecypods

(Nos see 05-2 UD) SAAS sos, TOM PlSsi/cscncssvssscouccesaconvaceatesesascecscuecseesbuaes

Large Foraminifera, Texas Cretaceous crustacean, Antarctic
Devonian terebratuloid, Osgood and Paleocene Foramini-
fera, Recent molluscan types.

Cire 1 A 21a S84 pi Soe DIS ceases ceccteaseccae-spsescosacscscesseasstoraaceds

Eocene and Devonian Foraminifera, Venezuelan fossil
scaphopods and polychaetes, Alaskan Jurassic ammonites,
Neogene mollusks.

CON arg2 ES) ee lO S Sipps) tip Seu cecsecextstvesccottries scecsessnncsacsscxctesersvarstssesss>

Catalogue of the Paleocene and Eocene Mollusca of the
Southern and Eastern United States.

(NS 9219-224) cp G71) Tppssh S30 DIS: *csccssscccocsscsenptesescopesesoresessssovevecsacses

Peneroplid and Australian forams, North American carpoids,
South Dakota palynology, Venezuelan Miocene mollusks,
Voluta.

UN Gare ee 5-250) a5) SDD ssi om DIS piatnivrrstcccccdnrsecscrecsanreronccetescasence

Venezuela and Florida cirripeds, Antarctic forams, Linnaean

Olives, Camerina, Ordovician conodonts, Niagaran forams.

IN carpal 2 32) 420 pps LO iSe erscectencectoveccvercereseuciencosescrseuceseseseso
Antarctic bivalves, Bivalvia catalogue.

(NGatt 233-230) S87 sPP en (45) DIRS ccctrercstarecsetstssctvececssencecssecesornecss

New Zealand forams, Stromatoporoidea, Indo-Pacific, Mio-
cene--Pliocene California forams.

(Nona 237-238) uF 88 Pps AS MDISE sssovarscsccestacncancovcvesidaccassavessacasosses

Venezuelan Bryozoa, Kinderhookian Brachiopods

PALAEONTOGRAPHICA AMERICANA

See Johnson Reprint Corporation, 111 Fifth Ave., New York,

N.Y. Monographs of Arcas, Lutetia, rudistids and venerids.

(NG FNG 92) ra SS Hpi 37 Pl Ss cecrsasecrecosstesasnsecerusssevarevescecoresssosses

Heliophyllum halli, Tertiary turrids, Neocene Spondyli, Pale-
ozic cephalopods, Tertiary Fasciolarias and Paleozoic and
Recent Hexactinellida.

CONS ooh 5225) ea Ss ps, (60 DISY: i heccesccressxevssove<ceacescveesasarncacsaseaat=o

Paleozoic cephalopod structure and phylogeny, Paleozoic
siphonophores, Busycon, Devonian fish studies, gastropod
studies, Carboniferous crinoids, Cretaceous jellyfish, Platy-
strophia, and Venericardia.

(ING 82) 26-35) a0 AO ZIP iy Ae PISS, sxcrccstssvescscccshyssetesaressecveccetsveneccarbs

Rudist studies, Busycon, Dalmanellidae, Byssonychia, De-
vonian lycopods, Ordovician eurypterids, Pliocene mol-
lusks.

(NOs 4-a7)) rn 44 Sepp PLO DIS: ceccccssstrecssscanestersscncssesocosacnsseaens

Tertiary Arcacea, Mississippian pelecypods, Ambonychiidae,
Cretaceous Gulf Coastal forams.

16.00

16.00

16.00

16.00

16.00

16.00

16.00

16.00

18.00

16.00

16.00

16.00

16.00

16.00

23.00

28.00

28.00

32.00

XXVI.

XXXII.

XXXIV.

XXXV.

XXXVIII.

XXXTX.

BULLETINS OF AMERICAN PALEONTOLOGY

I-XXIII. See Kraus Reprint Corp., 16 East 46th St., New York,

N.Y. 10017, U.S.A.

(Nos. 80-87). 334 pps 27 DIB. srcssacessesvosecessseroenesasnsaseseeaeosenvenssnrs
Mainly Paleozoic faunas and Tertiary Mollusca.

(Nos. 88-9435). S06) pp, SO iss: ccscsscasteccccesscnsessxcseessronsncossassstecteats

Paleozoic fossils of Ontario, Oklahoma and Colombia, Meso-
zoic echinoids, California Pleistocene and Maryland Mio-
cene mollusks.

(Nos: 95-100). 420° pp. SS pls cc--c-cssscsstintsossscosncassyssxonsaonovessuspanen

Florida Recent marine shells, Texas Cretaceous fossils, Cuban
and Peruvian Cretaceous, Peruvian Eogene corals, and
geology and paleontology of Ecuador.

(Nos: SOUTOS) o> S76 ei BB ae ecerncesperccattigoxescnpuscecaceceesarerterass
Tertiary Mollusca, Paleozoic cephalopods, Devonian fish and
Paleozoic geology and fossils of Venezuela.

(Nog: 109-104). 2402 pps, eS Aspe escacetnarccansrctrest swsnises eaceonaseceoesns
Paleozoic cephalopods, Devonian of Idaho, Cretaceous and
Eocene mollusks, Cuban and Venezuelan forams.

(Nios 215-116) 5 FSS ip es ro ae Piha ennccertecremns tater costensnekeexcsoxs ope ice
Bowden forams and Ordovician cephalopods.

(No. 117). 5163) papas 650 Dl Se ee cccctennnccsccenssatsnanaansnebuscucessvenadsscessshpearee
Jackson Eocene mollusks.

(Nos. 118-128) = SASS pps, eo UBe cerccsseeaaessegereceesstepsnsnsececeramneer

Venezuelan and California mollusks, Chemung and Pennsyl-
vanian crinoids, Cypraeidae, Cretaceous, Miocene and Re-
cent corals, Cuban and Floridian forams, and Cuban fossil
localities.

(Nos. 129-133) 5 294 pps SO pI ae iecccssssenreseteeesecrnenscacentecerteeroteests
Silurian cephalopods, crinoid studies, Tertiary forams, and
Mytilarca.

(Nos: 134-139)... 448° pp, VSI. (plan ecaveeserecsctrneessernesseeeeriine
Devonian annelids, Tertiary mollusks, Ecuadoran strati-
graphy paleontology.

(Nos. 140-145). 400 pp, 19) pls: -.....sceccsccssessccesssbacscusnsnssnenensesas

Trinidad Globigerinidae, Ordovician Enopleura, Tasmanian
Ordovician cephalopods and Tennessee Ordovician ostra-
cods and conularid bibliography.

(Nos. 146-154). 386 pp, 30 psi cc.cccciccscscnsscscscesvecsssseecserasesesonse

G. D. Harris memorial, camerinid and Georgia Paleocene
Foraminifera, South America Paleozoics, Australian Ordo-
vician cephalopods, California Pleistocene Eulimidae, Vol-
utidae, and Devonian ostracods from Iowa.

(Nos; 255-160)... 4:12) pyr 5S 0 plea arceseeesnsanccenstasebesceresestexseseesecpars

Globotruncana in Colombia, Eocene fish, Canadian Chazyan
Antillean Cretaceous rudists, Canal Zone Foraminifera,
fossils, foraminiferal studies.

(Noss, 161-164) 2486; ppi0 37 pls wacscscrccererscncceteepsertintehssececnenseanens
Antillean Cretaceous Rudists, Canal Zone Foraminifera,
Stromatoporoidea.

(Nos: 165-176) 447 py.) 53 sles oon es sccacpnenteatasssmenteeeetenncenlvoemasaae

Venezuela geology, Oligocene Lepidocyclina, Miocene ostra-
cods, and Mississippian of Kentucky, turritellid from Vene-
zuela, larger forams, new mollusks, geology of Carriacou,
Pennsylvania plants.

(Noa. 1772183) S448 pps, 36 pISs aisccctccccasccerseces ethernet echeameevoresccee

Panama Caribbean mollusks, Venezuelan Tertiary formations
and forams, Trinidad Cretaceous forams, American-Eur-
opean species, Puerto Rico forams.

12.00

12.00

14.00

14.00

14.00

18.00

16.00

16.00

16.00

16.00

16.00

16.00

16.00

16.00

16.00

16.00

PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

VOL. VI

NO. 39

CENOZOIC EVOLUTION OF THE ALTICOSTATE VENERICARDS
IN GULF AND EAST COASTAL NORTH AMERICA

By

WILLIAM GRAHAM HEASLIP
State University College at Cortland,
Cortland, New York

1968

Paleontological Research Institution

Ithaca, New York, U.S.A., 14850

PALEONTOLOGICAL RESEARCH INSTITUTION

1967 - 1968
PRESIDE NITY. ceosscscssvecusenegs undec tt ob oinescetabraevais urtnsaepabenecomi Cant tna) ..KENNETH E, CASTER
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SECRETARY-TREASURER scscsecossssocssecossecesvssavecsssesecnssorneeasonnenenssesereneeseoees REBECCA S. HARRIS
JURE CTOR) vessnrsecectssoraovsscsscestiascsssvasesoetvesadsseenspracesacnepestetzencasiane KATHERINE V. W. PALMER
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PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

VOL. VI

NO. 39

CENOZOIC EVOLUTION OF THE ALTICOSTATE VENERICARDS
IN GULF AND EAST COASTAL NORTH AMERICA

By
WILLIAM GRAHAM HEASLIP

State University College at Cortland,
Cortland, New York

August 1, 1968

Paleontological Research Institution

Ithaca, New York, U.S.A., 14850

ae i oo)
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CONTENTS

Abstract
Introduction
Need for investigation
History of classification .
Purposes of the investigation
Results of the investigation ...

Conclusions
Acknowledgments
The alticostate yenericards
Orientation within V’enericardia
Morphology and definition of terms
Ornamentation
Dentition
Ligamental features .
Musculature
Standard dimensions
Standard ratios
Analyses and presentation of data
Standard presentation .
Bivariate analysis
Life habits of Vemericardta ........--...-.:--c0-s0-----t0--00--
General anatomy of J’. (Cyclocardia) borealis
Patterns of growth

1) Year classes ......

2) Pattern of growth curvature
Properties of the equiangular spiral
Method of study
Patterns of curvature in V’enericardia
Interpretation of curvature pattern
Evolutionary trends .....-2.:::.ceeeeeee
Dental pattern
Umbonal height increase
Size increase
Shape ratio
Ornamentation ...

Sexual dimorphism in two lineages —...- .---.--.-..-----..--

Evolutionary history of the alticostate venericards —......
Geographic scope of investigation
Stratigraphic scope of investigation ..
Sources of material studied
Alticostate evolution
Evolutionary history of Baluchicardia in North America ..
Evolutionary history of Rotundicardia in North America _.......-..--..----- 75

Evolutionary history of Claibornicardia in North America
Evolutionary history of Glyptoactis in North America
Systematics
Venericardia Lamarck, 1801 ... *
Baluchicardia Rutsch and Schenck, 1944, subgen
Rotundicardia Heaslip, n. SUbgeNUS «.........ce eects
Claibornicardia Stenzel and Krause, 1957, subgenus ..........
Glyptoactis Stewart, 1930, subgenus
Bibliography
Plates

79

ake

2%

TEXT - FIGURES

Fluctuating abundance of alticostate /’enericardia, eastern and

Golf Worthy Ame rica a i5. Se. sas tianecennacteectpivenssbass neta oattekencoreareeree eee 57
Contrasting morphology of alticostate and planicostate shells —............... 58
General phylogeny of J’enericardia
General morphology and orientation of J’. alticostata 20000. 59
Cardinal elements of heterodont Groups. ............cc.ccesesccersscsesecesseseesesessecnesseeee 60

Comparison of reduced major axes in height and length in J’.
alicastata and: 2 sillianil os enn eee
V. (Cyclocardia) borealis shell anatomy
General ‘anatomy’; BORr bab as cic cinc sessions ees cesta
General organ anatomy of J’. borealis _..
Frequency-distribution histogram of height, J”. a/lticostata ..
Equiangular spiral
Semilogarithmic protractor
Semilogarithmic plots of curvature patterns
Semilogarithmic plots of curvature in I’. complexicosta .
Semilogarithmic plots of curvature, J’. alticostata
Semilogarithmic plot of curvature, J’. rotunda
Patterns of dentition in four subgenera ....

Mean BH/H vs. time . 70
Height vs. time in Baluchicardia and Rotundicardia .. . 70
Mean H/L vs. time Si!
Mean rib number ys. time — i
Development of sexual dimorphism in Rotundicardia al
Evolution of sexual dimorphism in J’. alticostata ........ Jee
Cenozoic stratigraphy of Gulf Coastal Plain and Atlantic

Coastal Paha ies isiatrtte nee erecta eetereen OE ee ea 73
Evolutionary pattern of Baluchicardia ... : 7+
Evolutionary pattern of Rotundicardia 76
Evolutionary pattern of Claibornicardia ... yf
Evolutionary pattern of Glyptoactis 02 80

CENOZOIC EVOLUTION OF THE

ALTICOSTATE VENERICARDS

IN GULF AND EAST COASTAL NORTH AMERICA

WILLIAM GRAHAM HeEas.ip

ABSTRACT

The alticostate yenericards, united by constant characters of
shape, dentition and ornamentation, belong to four subgenera: Balu-
chicardia Rutsch and Schenck, 1944, Rotundicardia, n. subgen., Clai-
bornicardia Stenzel] and Krause, 1957, and Glyptoactis Stewart, 1930.
These form a dynamic, continuously evolving lineage on the eastern
and Gulf coasts of North America, undergoing evolutionary bursts
in the early Paleocene, middle Eocene and early Miocene, becoming
extinct in this region at the end of Miocene time.

From the primitive baluchicards of the Paleocene, the rotundicards
evolved in the early Paleocene, retaining the ancestral dental pat-
tern throughout their evolutionary history, but simplifying the primi-
tive tripartite ribs into simplified noded blades. The last known
rotundicard is I’. (R.) carsonensis Dall, 1903, of the early Oligocene
Red Bluff Formation in Mississippi.

The claibornicards, first appearing in the early Eocene, undergo
a marked expansion in the middle Eocene, becoming an important ele-
ment in coastal waters in eastern North America during that time.
Their numbers are severely reduced by the time of Jackson deposition,
and the last known species of this important lineage is V’. (C.) nasula
Dall, 1903, of the (?) middle Oligocene.

During the late Oligocene, the first glyptoactid appears in the
Gulf, giving rise to an entirely new lineage which expands in the
early Miocene to fill the ecologic roles left vacant by the extinct
claibornicards. The last known species, ’. (G.) olga Mansfield, 1939,
of the late Miocene may have given rise to the glyptoactids living
in coastal waters off California, Central and northern South America.

Several important trends are noted in the evolution of the alti-
costate venericards in eastern and Gulf North America: (1) The de-
parture from the ancestral, rugged dental pattern by Claibornicardia
and the later glyptoactids, (2) Increase in inflation of the beaks,
characteristic of the entire group, probably correlated with increased
ability to nurture developing larvae, (3) Increase in size noted in
Baluchicardia and reduction in size among the rotundicards, (4)
Relative decrease in length in all but Claibornicardia in which a
marked increase is noted, (5) An increase in rib number in all
groups but Rotundicardia, (6) The development of sexual dimorphism
noted in Rotundicardia and Claibornicardia by distinctions in curva-
ture pattern and ornamentation.

The study of growth patterns of curvature establishes two dis-
tinct patterns: A type, comprising four increasingly convex segments
and B type, consisting of the 4 pattern with an added fifth terminal
segment of lower convexity. In species characterized by the 4 pat-
tern only, protandric hermaphrodism is inferred with the final con-
vex stage representing the female “marsupial” development. In forms
exhibiting both patterns, the highly convex 4 type is associated with
“femaleness” and the Jess convex B pattern with ‘“maleness.”’ This
is documented in V’. (R.) rotunda, Lea, 1833, and V’. (C.) alticostata
Conrad, 1833, by characteristic ornamental types associated with each
pattern.

Thirty-four species are described, 28 by previous authors, and
six as new, I’. amplicrenata, V. linguinodifera, V. trapaquaroides, }
impendeocosta, V. gibberumbonata, and V. chelomodonta. Three
species are each divided into two subspecies which recognize hetero-
chronous or geographical races. One new subgenus, Rotundicardia, is
segregated.

INTRODUCTION

NEED FOR INVESTIGATION

Lamarck’s descriptions (1801) of Paris Basin Eocene
fossils first indicated the importance of the bivalve genus
Venericardia in the marine faunas of the early Cenozoic.
Later investigations of Conrad (1830-) and Lea (1833) in
the Atlantic and Gulf Coastal Plain sediments aroused in-
terest in American Tertiary paleontology, and Conrad
(1855) was so impressed by the abundance of Venericardia
that he called it the “fingerpost of the Eocene.” The value

of its species as guide fossils in sediments of that age is as
high as that of any other molluscan genus, and the excellent
preservation of venericard shells, their widespread occur-
rence, and distinctive interspecific ornamental variations
makes even fragmental material diagnostic in stratigraphic
correlation,

Despite the large body of knowledge about many
American species of Venericardia which had accumulated
by the early 1900's, a lack of interest in the systematics of
the group is evident in the abundance of distinctive species
lumped under the two form taxa V. planicosta Lamarck
and JV. alticostata (Conrad) by authors. Even Dall’s impor-
tant contribution (1898-1903) only included many new
species and failed to present a phylogenetic framework.

Unfortunately, most of the systematic treatment of
Venericardia has been concerned primarily with the “plani-
costate” species. The present investigation treats only the
American “alticostate” species of the East and Gulf Coasts
in an attempt to erect a realistic phylogenetic framework
for this equally conspicuous group within the genus.

HISTORY OF CLASSIFICATION

The first systematic treatment of Venericardia ap-
peared in Harris’ (1919) study of the early and middle
Eocene sediments of the Gulf Coast. Harris recognized the
evolutionary group
separated from that of V. planicosta and included many of
the species described in the present investigation. Harris
was much more conservative than the older paleontologists
and refused to recognize many of the species erected by
Dall (1903) as well as earlier authors. He called them only
varieties” of V. planicosta or V. alticostata. Dall (op. cit.)
commented on the evolutionary succession in the group of
V. rotunda but made no attempt to place other alticostate

“rotunda-alticostata” stock as an

species in an hierarchy.

Stewart (1930) erected two subgenera of Venericardia:
Venericor, which included all species having flattened ribs
and a high massive hinge plate, and Glyptoactis, which in-
cluded those species with sharply defined, noded ribs and
a low, more delicate hinge plate. In the latter group, he in-
cluded Dall’s Miocene species V. hadra and V. himerta, as
well as V. alticostata and V. rotunda of the North American
Eocene, and V. acuticosta Lamarck of the French Eocene.
Glyptoactis, as originally defined is only descriptive, just as
the term “alticostate” is descriptive, rather than taxo-
nomically accurate, for the group includes several distinct
lineages in North America.

Gardner and Bowles’ (1939) treatment of the “Veneri-

56 PALAEONTOGRAPHICA AMERICANA (VI, 39)

cardia planicosta group” represented an important contribu-
tion to the systematics of the genus, but those authors
failed to recognize a number of examples of extreme con-
vergence, and Venericor, as they understood it, is polyphyle-
tic (Heaslip, 1963).

Another addition to the understanding of the “plani-
costate” evolution was made by Verastegui (1953) in his
monograph of the Pacific Coast venericards. His subgenus
Pacificor is an offshoot of the early Venericor group on the
Gulf Coast which invaded the eastern Pacific waters.

The first systematic attention directed to the “alti-
costate” phylogeny was the work of Harris (1919). In 1941,
Rutsch and Schenck described the ancestral group originat-
ing in the Late Cretaceous of the Tethyan region which
they called Baluchicardia, distinguished by its primitive
ornamentation and shell form. Stenzel and Krause (1957)
described the subgenus Claibornicardia to include species
distinguishable by ornamentation and characters of denti-
tion from the ancestral baluchicards, as well as from the
more advanced glyptoactids, whose type species of Glypto-
actis is the Miocene species V. (G.) hadra Dall, 1903.

In the present investigation, another lineage is recog-
nized which includes species related to V. rotunda Lea.
Rotundicardia, n. subgen., contains species characterized by
a flattened shell with a minimum of posterior truncation,
which diverged from the baluchicards in the early Paleocene
and became extinct after the early Oligocene. This group
evolved mainly toward ornamental simplification, although
a branch is characterized by accentuated primitive charac-
ters of ornamentation.

PURPOSES OF THE INVESTIGATION

The present investigation attempts to examine all alti-
costate venericards known from the eastern and Gulf
Coastal Plain sediments in order to determine (a) the
distribution in time and space of the group, and (b) evolu-
tionary relationships among species of the group, maintain-
ing the subgeneric approach of previous authors.

The attempt is also made to evaluate important
morphological characters as aids in a realistic appraisal of
classification and as indicators of important evolutionary
trends.

Growth patterns of curvature are examined in the at-
tempt to relate these to possible ontogenetic stages in the
life of fossil venericards and as possible indicators of phy-
logeny.

RESULTS OF THE INVESTIGATION

Thirty-four species were found to be valid during the
present investigation, including 28 previously described

forms and six new. In addition, six subspecies are indicated
to include geographical or heterochronous races of three
species. These were formerly described as species, but a
lack of morphological distinctiveness makes the subspecific
rank a more meaningful appraisal of differences which do
occur.

A critical examination of the three previously pub-
lished subgeneric categories, Glyptoactis Stewart, Baluchi-
cardia Rutsch and Schenck, and Clatbornicardia Stenzel
and Krause results in a more complete understanding of
the scope, time significance, and interrelationships among
these, as well as the necessity for a new subgenus Rotun-
dicardia Heaslip.

The evaluation of morphologic characters results in the
arrangement of the species on the eastern and Gulf Coasts
of North America in a phylogeny showing a number of
striking evolutionary trends in a continuing lineage. A num-
ber of important offshoots from the main stem indicate a
constant process of evolutionary experimentation, producing
many radical forms.

The analysis of the patterns of growth in terms of the
equiangular (logarithmic) spiral model has correlated dif-
ferences in curvature growth patterns with certain intra-
specific morphological variations in V. (Rotundicardia)
rotunda Lea and other species, and points to sexual dimor-
phism in these species as a possible explanation. Interpreta-
tion of patterns of curvature in other species is not so deci-
sive, but the possible implications of protandric hermaphro-
dism is suggested for some which show an increasing shell
convexity as growth progressed.

CONCLUSIONS

The alticostate venericards of North America comprise
a dynamic, widespread, and continuously evolving group
from their first appearance in sediments of Paleocene age on
the Gulf Coast to their disappearance in eastern North
America after the Miocene. Their evolution is characterized
by several important trends in dentition, shape, and orna-
mentation, but the basic morphological unity is maintained
from earliest to latest members.

This group undergoes three “bursts” of adaptive radia-
tion during its Tertiary history in eastern North American
coastal waters (see Text-fig. 1). In each, a new subgeneric
group emerges (see Text-fig. 3). Following each burst, a
severe reduction in numbers occurs, and in some cases, only
inference can be made that a species is present to continue
a lineage found in earlier and later rocks (e.g. the absence
of any rotundicard in the late Paleocene record). The most
impressive burst of speciation is witnessed in the middle

AMERICAN ALTICOSTATE VENERICARDS: HEaASLIP

te} 5 10 15
NUMBER OF SPECIES

Text-figure 1. Fluctuating abundance through time of alticostate
species of Vencricardia on the eastern coast and Gulf coast of North
America.

Eocene, during which the claibornicards attained their peak
of diversity and geographic spread. The Miocene expansion
is a relatively local phenomenon, occurring only in the
eastern part of the Gulf (Florida) and the southern part of
the East Coast (Carolinas), and no record is found in the
western Gulf. The number of species present in the Miocene
of the Antilles and the Mediterranean Region also indicates
a burst of evolutionary activity during this time.

The decline of importance after the Miocene in marine
faunas in many parts of the world, as well as North America,
is a conclusion reached by many authors (e.g. Davies, 1935).
The reasons for this are not entirely understood, but a com-
parison to modern representatives of the group living in the
eastern Pacific from Baja California to northern Peru may
be justified. V. crassicostata G. B. Sowerby I, 1825, JV.
tricolor (G. B. Sowerby I) 1833, and V. megastropha Gray,
1825 all live in relatively deep water and extend through the
tropical and subtropical latitude zones. Although the present
Gulf Coast is characterized by subtropical climate, the
waters near the coast are shallow. However, Puri (1953)
showed that depth of deposition of Miocene sediments in
which many glyptoactid species evolved [e.g. V. (G.) hadra,
V. (G.) himerta] was also shallow. This may indicate the
abandonment of a former shallow habitat by this group and
the invasion of a deeper one.

Another possible reason for the post-Miocene decline

wn
“I

in numbers and distribution of the glyptoactids in North
America may be the increased competition by another bi-
valve group. The large venerids [e.g. Mercenaria mercenaria
(Linn.)] occur in great abundance in Miocene and later
sediments and perhaps the venericards responded by moving
to an environment in which the competition was not so in-
tense.

A number of examples of convergence between species
belonging to different alticostate subgenera seems to indi-
cate similar roles assumed by these species in the marine en-
vironment. A notable example is the high degree of simi-
larity between V. (Baluchicardia) whitei Gardner, 1923, of
the Kincaid Formation of early Paleocene age in Texas and
V. (Glyptoactis) hesperide Gardner, 1936, of the Shoal
River Formation of Miocene age in Florida. Both species
evolved a high degree of postero-ventral elongation and
simplified, rounded ribs which meet in restricted inter-
costals and have only subdued transverse costal nodes. Puri
(1953) characterized the Shoal River depositional environ-
ment as near shore, brackish, and shallow, based on environ-
mental preferences of living relatives of the microfauna of
that unit. Whether or not this relationship of shell form,
ornamentation, and habit in the Miocene species can be ex-
tended back to the Paleocene is not known but this repre-
sents a possible approach.

ACKNOWLEDGMENTS

The writer is indebted to Norman D. Newell of the
American Museum of Natural History and Columbia Uni-
versity, whose interest in bivalve evolution was a source of
inspiration. Helpful advice and criticism were received from
John Imbrie of Brown University, Andrew McIntyre of
Columbia University, Louis S. Kornicker, U. S. National
Museum, David Nicol, University of Florida, Gainesville,
Florida, Druid Wilson of the U. S. Geological Survey, and
Mrs. Katherine V. W. Palmer of the Paleontological Re-
search Institution.

Collections made in the field during the summers of
1955 and 1957 were supported by grants to the writer from
the James F. Kemp Fund of Columbia University and the
American Museum of Natural History. The writer was
accompanied and aided by Donald F. Squires of the U. S.
National Museum on the first trip and by Andrew F. Me-
Intyre on the second. Miss Winnie McGlamery, formerly
chief paleontologist of the Alabama Geological Survey, ac-
companied both field expeditions in Alabama and lent the
benefit of her knowledge of coastal plain stratigraphy and
fossil localities in that state.

7

V BASHIPLATA

V ALTICOSTATA

Text-figure 2. Contrasting morphology of alticostate (V’. alticos-
tata) and planicostate (I. bashiplata) shells. C, costal, CC, costal cord,
PC, paracostal cord, IC, intercostal. Right valve cut away to expose
interior and dentition.

Chester Tarka of the American Museum of Natural
History designed a telescopic device used in tracing curva-
ture profiles in the present investigation, and his help and
advice is appreciated.

For the loan of specimens and for assistance during
visits to their respective institutions, the writer is indebted
to Katherine V. W. Palmer, Paleontological Research Insti-
tution, G. Arthur Cooper and Joseph P. E. Morrison, U. S.
National Museum, Ellen Trumbull Moore and Druid Wil-
son, U. S. Geological Survey, Horace G. Richards, Phila-
delphia Academy of Natural Sciences, and Walter B. Jones
and Winnie McGlamery of the Geological Survey of Ala-
bama. The late John T. Lonsdale of the Texas Bureau of
Economic Geology, and Peter T. Flawn of the same insti-
tution, David M. Raup, University of Rochester, formerly
of Johns Hopkins University, and Robert O. Vernon of the
Florida Geological Survey also lent specimens for study.

Ernest Guignon of Connecticut State College, while at
Syracuse University, placed the facilities of his zoology
laboratory at the writer’s disposal, as well as gave advice
in the biological work of the present investigation. Edna
Kaneshira Kehlenbeck of Syracuse University also gave
assistance. Caryl E. Buchwald of Syracuse University plot-
ted the regression analysis in Text-figure 6.

The cost of illustrations has been met by the Research
Foundation of the State University of New York, Grant
23-0048A.

Finally, the writer acknowledges aid from his wife,

58 PALAEONTOGRAPHICA

Americana (VI, 39)

Barbara M. Heaslip, who assisted in the curve analysis,
manuscript preparation and plates, and gave encourage-
ment throughout the project.

THE ALTICOSTATE VENERICARDS
ORIENTATION WITHIN VENERICARDIA

The genus Venericardia, as commonly understood, em-
braces several distinct evolutionary lineages with common
characters of dentition and shell form, distinguishable main-
ly on the basis of ornamentation (see Text-fig. 2). The
“planicostate” lineage, comprising the subgenera Leuroactis
Stewart, 1930, and Venericor Stewart, 1930 on the East and
Gulf Coasts of North America are characterized by elevated
hinge plates with strongly developed, steeply sloping cardinal
teeth, a subtrigonal shape and smooth flattened costae
which become obsolete during the growth of certain species.
This group appears in the early Paleocene and undergoes its
entire evolutionary history during Paleocene and Eocene
time, becoming extinct at the end of the Eocene in North
America. (See Text-fig. 3.)

ALTICOSTATE

CYCLOCARDIA

5
Lei)

NO. SPECIES

PLANICOSTATE

LEUROACTIS
VENER/ICOR
eS
2—Y

J

BALUCHICARDIA

Text-figure 3. General phylogeny of Venericardia in North Amer-
ica, eastern coast and Gulf Coast.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP

Text-figure +. General morphology and orientation of /’. (C.) alticostata. A, left anterior
view: APAX, antero-posterior axis, NS, nymph support, N, nymph, LG, ligament, B, beak, LU,
lunule, LP, lunular protuberance (anterior lateral), 2, +b, anterior and posterior cardinals, 3a’,
3b’, 5b’, sockets. B, right interior view: L, length, BL, beak length, AA, anterior adductor scar,
APR, anterior pedal retractor scar, LS, anterior lateral socket, 3a, 3b, 5b, anterior, medial and
posterior cardinals, 2’, +b’, sockets, PPR, posterior pedal retractor scar, PA, posterior adductor
scar. C, right anterior view: HR, highest rib (measured for curvature), 1%2W, semiwidth, H,
height, BH, beak height. D, right exterior view: PTR, posterior truncation, HR, highest rib,
IC, intercostal, PC, paracostal cord, CC, costal cord, CN, costal node.

60 PALAEONTOGRAPHICA AMERICANA (YI, 39)

The group considered in the present investigation is the
“alticostate” assemblage, comprising the subgenera Balu-
chicardia Rutsch and Schenck, 1944, Rotundicardia Heaslip,
n. subgen., Claibornicardia Stenzel and Krause, 1957, and
Glyptoactis Stewart, 1930. These appear in the primitive
subgenus Baluchicardia on the Gulf Coast of North America
in the Paleocene from Late Cretaceous ancestors of Balu-
chistan and northwest India and continue through evolu-
tion of the other subgenera in the glyptoactids of the mod-
ern eastern Pacific. The group is characterized by primitive
tripartite, coarsely noded costae, an elongated, subelliptical
shell shape, and somewhat lower hinge plate with delicately
arched and less massively constructed cardinal teeth.

MORPHOLOGY AND DEFINITION OF TERMS
ORNAMENTATION

The radical ornamentation of the alticostate veneri-
card is one of the most distinctive among all the bivalves
and diagnostic of this group. The term “tripartite” best
describes it, from the term used in French Gothic architec-
ture in which the groins of adjacent vaults join in a three-
part column, having a partial cross-section like a subdued
fleur-de-lis.

The costal cord (CC, Text-fig. 4) stands like a crest
above the subjacent paracostal cords (PC, same Text-
figure), and is ornamented by nodes which are asymmetrical
and pointed like teeth of a saw in the primitive baluchi-
cards, becoming lingulate in the early claibornicards, and
progressively simplified to symmetrical transverse ridge-
like swellings in the advanced glyptoactids. Harris (1919)
coined the apt terms “funiculate” to describe the knotted
cordlike costals in members of V. (R.) rotunda Lea, and
“funginate” for other members of the same species in which
the nodes resembled flaring tree fungi on a limb. “Flabellate”
was used by the same author for the flaring nodes of V.
(R.) flabellum (Harris).

Paracostals resemble terraces on either side of the cen-
tral cord in the primitive baluchicards and claibornicards,
and are often accentuated by the development of a well-de-
fined thread on the apical ridge, which is occasionally noded
in concert with the central cord. The main tendency, in the
evolution of the alticostate venericards, however, is the sup-
pression of these paracostals, and they become highly sim-
plified and rounded in advanced claibornicards, disappear-
ing altogether in the later glyptoactids.

Intercostals (IC, Text-fig. 4) are generally U-shaped
in cross-section and delicately ornamented by fine concen-
tric increment lines of growth. In many species these be-

come V-shaped, or even restricted to a fine slit in forms
with highly expanded costals.

The radial ornamental is reflected on the commissural
margin of the shell by strongly developed crenulations which
provide an extra-cardinal interlock for the closed valves.

DENTITION

The dental pattern in Venericardia is heterodont, com-
posed of cardinal teeth radiating from a position under the
beak, and lateral teeth which are poorly developed. It is in-
termediate between the cyrenoid type with three cardinals
in each valve and the lucinoid type with only two in each
valve (see Text-fig. 5). Davies (1935, p. 155) compared the
venericard pattern to the lucinoid type, but “acquiring a
cyrenoid aspect through the development of an additional
tooth.” This is the anterior cardinal of the right valve (3a).

LEF@D RIGHT

A

CYRENOID

Vn 4b 2 ) 3a 3b 5b
B

VENERICARD

Cc

LUCINOID

Text-figure 5. Cardinal elements of three major heterodont groups,
noting terminology of the Bernard system. Lateral elements not in-
cluded.

The right cardinals are numbered 3a (anterior), 3b
(medial), and 5b (posterior), using the system devised by
Bernard (1895) and Munier-Chalmas (1895). This is based
on the appearance of primitive chevron-shaped lamellae in
early ontogenetic shell stages in the cardinal region under
the beak. These become progressively modified during early
ontogeny until the pattern of cardinals and laterals seen in

AMERICAN AL'TICOSTATE VENERICARDS: HEASLIP 61

the adult shell is attained. The lamellae are numbered from
ventral to dorsal, odd numbers for those on the right, and
even numbers for those appearing on the left valve. The
adult teeth are designated “a” or “b”, depending on whether
the tooth was derived from the anterior or posterior limb of
the chevron.

The cardinal elements of the left valve are numbered
2 (anterior), since this was thought by Bernard (1895) to
represent the complete chevron, and 4b (posterior).

Among the alticostate venericards, the right anterior
cardinal (3a) is highly variable in development, and appears
in the primitive subgenus Baluchicardia, as well as Rotundi-
cardia, as a semierect ridge bordering the anterior margin of
the socket (2’) for the anterior cardinal (2) of the left
valve. In Claibornicardia and Glyptoactis this tooth (3a) is
oriented on the anterior apex of the medial right cardinal
(3b) and aligned with the point, lying over the socket (2’).
This reflects an important difference in shape of the left
anterior cardinal (2) among these subgenera, and this tooth
is erect and triangular in the former two and flattened and
bladelike in the latter two. The earliest known carditid
genus, Paleocardita Conrad, 1867 first appearing in the
Triassic, had only two cardinals (2, 4b) in the left valve,
and only a single element (3b) in the right valve (Dou-
villé, 1912). Whether or not this form is ancestral to Venert-
cardia remains to be established, but the first members ap-
peared in sediments of Cretaceous age with fully developed
dentition characteristic of the genus.

The cardinal teeth are striated by minute grooves and
ridges parallel to the direction of occlusory motion on sur-
faces of contact, and these serve as guides in opening and
closing the valves.

The development of lateral dentition is not noteworthy
in Venericardia as in many heterodont groups, including the
allied genus Cardita. “Alticostate,” as well as “planicostate”
species uniformly exhibit a small protuberance at the inter-
section of the left lunule and the anterior shell margin (LP,
Text-fig. 4) which is described as a “slight puckering of the
shell” by Gardner and Bowles (1939). A corresponding
indentation (LS, same Text-figure) appears on the right
valve.

Posterior Jaterals are even less conspicuous, and appear
only as a rounded protuberance just behind, and usually
aligned with, the right posterior cardinal (4b), with a cor-
responding depression in the left valve. In no alticostate
species is the feature well developed, although there is a
general tendency toward increasing prominence in the evo-
lution of the alticostate venericards. Whether or not the an-
terior and posterior features are truly laterals in the sense
of Bernard and Munier-Chalmas is not known.

LIGAMENTAL FEATURES

The outer layer of the ligament is set in a deep groove
between the dorsal shell margin and the nymph support
(NS, Text-fig. 4), and a portion of this fibrous inner layer
is often preserved with the shell. Nymph supports are
lamellar and in the left valve, lie just above the socket (5b’)
for the posterior cardinal of the right valve, which has its
nymph support mounted directly on the tooth itself, and
discernable by a slight groove which distinguishes it from
the tooth. These features are well, but not massively de-
veloped, as in planicostate forms, and are delicate structures.

MUSCULATURE

The pattern of musculature as seen on the shell interiors
is standard in Venericardia, consisting of large anterior and
posterior adductor scars (AA & PA, Text-fig. 4). The degree
of impression of these features is proportional to the shell
thickness, and the anterior scar is generally more deeply
inset than the posterior.

Two other scars are also visible: the anterior pedal re-
tractor (APR, Text-fig. 4) is separate from the anterior
adductor and deeply impressed just behind that muscle on
the ventral edge of the hinge plate. The posterior pedal re-
tractor is a triangular projection from the posterior dorsal
edge of the anterior adductor. In only one species, V. ( Clai-
bornicardia)natchitoches Harris, does the pedal retractor
become separated from its normal position, and lies isolated
just above its former position (see PI. 25, fig. 7c).

STANDARD DIMENSIONS

A system of measurement was established which is most
conveniently applied to dimyarian rather than monomyarian
bivalves. The anterior-posterior axis (APAX) is a line con-
necting the centers of adductor scars and is assumed to be
the most reasonable orientation for the fore-and-aft ana-
tomical direction. Other lines of reference such as hinge
margin might be employed but vary with shell shape (in-
flation of umbones, height of dentary plate). Most measure-
ments were made using this line (APAX) as a datum which
could only be estimated in cases of locked valves or poorly
preserved muscle impressions.

Length (L) is the greatest dimension parallel to the
line APAX and beak length (BL) the distance of the beak
from the most posterior point.

Height (H) is the distance between the ventral margin
and the umbo, taken at right angles to APAX. Beak height
(BH) is the distance of the beak from the ventral margin.

The term “width” is used as the most anatomically suit-
able for the dimension measured at right angles to the plane

62 PALAEONTOGRAPHICA AMERICANA (VI, 39)

of length and height measurements (plane of commissure)
as suggested by Waterhouse (1960). Because of the rare
occurrence of interlocked valves or complete specimens in
collections examined during this investigation, the term
“semi-width” (1/2W) is used throughout. Because of the
interlock of the coarse marginal crenulations, there is a dis-
crepancy between half the measurement across the pair of
interlocked valves and the “semi-width”, or measurement
of a single valve. However, this error is probably less than
that obtained in measuring coarsely noded forms. No con-
venient adjustment could be found for either of these sources
of error in the present investigation, but measurements were
uniformly made.

Rib number (RN) is a “dimension” recorded for each
shell, and this was found to be a meaningful element in
comparisons among different alticostate species.

STANDARD RATIOS

The standard measurements themselves may not be
meaningful, except that they establish impression of size in
a species. It was found that by relating measurements in
ratio form, a better basis of comparison among species could
be made, as well as numerical values summarizing shell
shape for each species. Any ratio between two measurements
is only valid for a single growth stage, and ratios, as well
as linear measurements, are dynamic, changing characters.
The average ratio, like the average linear dimension, em-
braces a range of ontogenetic stages. It is assumed, however,
that most of the valves measured in this study are mature,
so that the rate of change in ratio between any two meristic
characters might be minimal, because juveniles would be ex-
pected to show the highest degree of allomorphic growth.

Standard ratios are:

a) BL/L, the ratio between beak length and total
length as an estimate of posterior elongation.

b) BH/H, the ratio between beak height and _ total
height as an estimate of umbonal inflation.

c) H/L, the ratio between height and length as a para-
meter of total shell shape.

d) 1/2W/H, the ratio between semi-width and height
as an estimate of inflation of the valves.

ANALYSIS AND PRESENTATION OF DATA
STANDARD PRESENTATION
‘The dimensions and ratios are given in standard tabular
form for each species holotype, paratype, and lectotype im-
mediately after the description of these specimens, noting
L, H, 1/2W, RN, BL/L, BH/H, H/L, and 1/2W/H.

Studied collections of more than two individuals are

summarized in tabular form under the heading “Dimensional
analysis” for each species, noting statistics for the same
dimensions and ratios. Each dimension is summarized by
values for X (mean), OR (observed range), S (standard
deviation), V (coefficient of relative variation) and Ox
(standard error of the mean). For ratios, only values of
mean and observed range are given.

For monotypic species, or those in which only the holo-
type was examined during the investigation, only type
measurements are included.

Under the heading “Curvature characteristics,” ob-
served ranges and mean values of angular segments in the
curvature pattern of each species and shell heights at which
breaks in curvature occurs are noted (see later section for
curvature analysis).

BIVARIATE ANALYSIS

In certain cases, a statistical comparison between popu-
lations was undertaken using the technique of “Reduced
Major Axes” outlined by Imbrie (1956). This entailed the
calculation of the slope (a) of the growth line of two varia-
bles, height and length, the line’s position in terms of the
joint mean of the two characters, and its intersection with
the ordinate axis. The coefficient r, as a measure of the de-
gree of dispersion of points (each H vs. L) or cluster
around the growth line, and the standard error of the slope
(Oa) were also calculated.

The correlation coefficient, Z, which involves a com-
parison between slopes and their standard errors between
the two species was calculated, giving a numerical value to
the degree of similarity between the growth lines based on
a 95 per cent probability that observed differences between
slopes are due to chance sampling errors, and members of
the two populations are actually part of the same species.
Greater values indicate that the two growth slopes are not
the result of sampling errors, but that the two populations
belong to different species.

The use of this technique is predicated on the assump-
tion that populations having similar genetic constitution
evidence similar rates of growth, despite minor differences
among geographically isolated populations. Admittedly, the
same pattern of growth might characterize distantly related,
or even unrelated animal populations, but this technique
Was not applied indiscriminately among all species described
in the present investigation, but employed only where neces-
sary to document impressions gained from morphological
comparisons.

Results of this type of analysis affirmed decisions
(based on morphological comparisons) to:

1) Unite V. (C.) alticostata Conrad and V. (C.) sil-

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 63

limani Lea as intra-population variants of the same species,
V. (C.) alticostata (See Text-fig. 6).

2) Unite V. (C.) coloradonis Harris and V. (C.) texa-
lana Gardner under the same species name, recognizing dif-
ferences in rib number between heterochronous populations
by the subspecies V. (C.) coloradonis coloradonis and V.
(C.) coloradonis texalana Gardner.

3) Unite V. (R.) diversidentata Meyer and Aldrich and
V. (R.) vicksburgiana Dall (including V. “scabricostata”
recorded by Richards (1953) from western peninsular
Florida), recognizing synchronous allopatric subspecies V.
(R.) diversidentata diversidentata and V. (R.) diversiden-
tata vicksburgiana Dall.

4) Unite V. (G.) waynensis Mansfield and V. (G.)
nodifera Kellum, recognizing heterochronous subspecies JV.
(G.) nodifera waynensis Mansfield and V. (G.) nodifera
nodifera Kellum.

LIFE HABITS IN VENERICARDIA
GENERAL ANATOMY OF V, (CYCLOCARDIA) BOREALIS

Although relationships between the subgenus Cyclo-
cardia Conrad, 1867 and the alticostate venericards are not
understood at the present time the general unity of shell
morphology among all species of Venericardia makes a dis-
cussion of the “soft” anatomy of the living V’. (Cyclocardia)
borealis Conrad propitious in the present investigation. The
shell of this species exhibits normal dental pattern and orna-
mentation resembling that of the “planicostate” forms but
exhibiting nodes in early ontogeny which progressively dis-
appear. (See Text-fig. 7).

During the summer of 1960, the writer obtained a num-
ber of living specimens of this species from the Marine
Biological Laboratory at Woods Hole, Massachusetts, and
a number of dissections provided a rudimentary outline of
the general anatomy of the soft parts. Unfortunately, there
is a dearth of information in the literature on gross anatomy
and only in a few commercially important species of mol-
lusks is this information available.

V. (C.) borealis is moderate in size, attaining a length
of 1% inches with high, blunt beaks, and radial ribs charac-
terized by nodal suppression through ontogeny (see Text-
fig. 7). A well-developed, nonbyssate foot and isomyarian
musculature with strongly inserted pedal retractor muscles
indicate a normal, active habitus. There is no development
of siphons and mantle edges are open along the ventral
margin, but a transverse septum (TS, Text-fig. 8), formed
by fusion of the mantle, separates the incurrent and excur-
rent areas. The excurrent siphon is a small, circular orifice
above the septum.

607—

Z= 1.80

n

ws |

Ww 40

Ww

=

=

aj

= al

za

yy °V.(C.) ALTICOSTATA
-

= £5 —O-JOINT MEAN

Ww °V(C)SILLIMAN/
x

> JOINT MEAN

(Seas aiet Sh cae ee ee eres

10 20 30 40 50 60 70
LENGTH IN MILLIMETERS
Text-figure 6. Comparison of reduced major axes in height vs.

length between V’. (C.) alticostata and V. “sillimani.” Correlation
coefficient Z = 1.80.

Text-figure 7. VY. (Cyclocardia) borealis shell showing dental
pattern, musculature, and ornamentation. Portion of right valve
mounted on left to show adult ornamentation and shell cross-section.
Isolated segment in center shows juvenile ornamentation (X3).

64 PALAEONTOGRAPHICA AMERICANA (VI, 39)

The branchial apparatus is the normal eulamellibranch
pattern with the suprabranchial chambers following the
umbonal ridges of the visceral mass. Part of the anterior
edge of the inner gill is fused to the anterior visceral ridge
connecting the umbo to the central part of the anterior
adductor, and on this same ridge the labial palps are con-
nected to the anterior edge of the gill by a strand of con-
nective tissue which may be a vascular tubule (see Text-
fig. 8). The dorsal margin of the gills join the posterior ad-
ductor muscle at its antero-dorsal prominence and continue,
unconnected, to the transverse septum, where the excurrent
openings of the suprabranchial chambers provide the cur-
rent for fecal discharge. The free posterior flaps of the gills
join at the center to isolate the anal atrium from the in-
current area.

The mouth (M, Text-fig. 9) is located centrally be-
tween the transverse ridges of the labial palps between the
muscular ridges of the yokelike pedal retractors which
diverge dorsally into either valve from the foot. The
straight esophagus connects the mouth and the dorsally lo-
cated stomach (ST) which is a simple sac without diverticu-
lum or crystalline style. From the floor of the stomach, the
intestine descends to the floor of the visceral cavity where
it bends to the anterior, then recurves along the floor to
the posterior, ascending dorsally, where it is surrounded by
the ventricle of the heart. From here it bends to the pos-
terior along the dorsal mid-line, and the rectum follows the

Text-figure 8. General anatomy of I’. (C.) borealis Conrad. TS,
transverse septum of mantle, EO, excurrent opening, PA, posterior
adductor, R, rectum, FM, fused mantle, PPR, posterior pedal retrac-
tor, SBC, suprabranchial chamber, RIG, right interior gill, REG,
right exterior gill, F, foot, APR, anterior pedal retractor, LP, labial
palps, AA, anterior adductor. X2%.

posterior face of the posterior adductor muscle around the
ventral surface, where the anus (A) opens to the anterior.

The digestive gland is a large, brownish organ which
surrounds the esophagus, stomach and descending ramus of
the intestine, occupying the anterior and antero-dorsal part
of the visceral cavity. The nephridia are dark-colored, paired
organs just dorsal to the upper edges of the posterior adduc-
tors and surrounding the posterior retractor muscles. These
open in pores on the ventral border of the gills and visceral
mass of the foot at the center of the pedal retractor muscle.

Only females were observed in the investigation, and
the large ovaries comprise most of the posterior part of the
visceral mass, but extending anteriorly along the floor of
the visceral cavity, surrounding the intestinal tract except
the descending ramus. Ovarian tissue ascends dorsally in
sheets which cover the dorsal lobes of the digestive gland.
These sheets have been stripped away in Text-figure 9 to
show the outlines of the digestive gland. Eggs are arranged
around stalked ovarian tubules which ramify in these sheets.
No specialized portions of the gills (“marsupia”) were ob-
served, but the subsequent development of the eggs is sub-
ject for further study.

Anterior and posterior aortae run dorsally from the
ventricle of the heart which is surrounded by the pericardial
sac just posterior to the apical lobes of the digestive gland,
branching laterally to the retractor muscles and mantle
muscles as well as the viscera, where the blood enters large

Text-figure 9. General organ anatomy of I’. (C.) borealis Conrad.
LIG, flap of left interior gill, A, anus, N, nephridium, H, heart, OV,
ovary, IN, intestine, DG, digestive gland, ST, stomach, M, mouth,
Female, X2%.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 65

sinuses and is returned to the suprabranchial veins and
the auricles via vein tubules which penetrate these organs.

Cerebro-pleural and posterior pedal ganglia and nerve
ramifications to the muscles have been observed, but more
detailed discussion must await further investigation.

The general morphological similarities of the shell
among fossil and living forms of the genus Venericardia in-
dicate the basic unity in the soft anatomy of members of
the genus.

PATTERNS OF GROWTH

1) Year classes. — In general, no distinct stages of
growth such as evident in arthropod molts can be readily
distinguished in Venericardia, but plotting a frequency-dis-
tribution histogram of a meristic character which changes in
proportion to the age of the animal in a large sample, re-
sults in a multimodal curve. Text-figure 10 shows the re-
sult of a plot of height in a sample of 165 individuals of V.
(C.) alticostata from AMNH collections 1055, 984 and
9950/1 on 3 millimeter class intervals. Five distinct modal
classes can be distinguished (A,B,C,D,E?), although only
a single individual comprises the largest class (57-60 mm.).
The modes are spaced at approximately equal intervals, and
may represent year classes. It is conceivable that a smaller
class is not represented. A bivalve of similar size range
[Mercenaria mercenaria (Linn.)| studied by Loosanoff
(1936) has a first year class average of less than 7 milli-
meters in length. This is about half modal range of 13 to 15
millimeters, and in the fossil population, the winnowing ac-
tion of currents may have carried the young shells away.

2) Patterns of growth curvature. — Characteristic
growth among most mollusks as well as certain other animal
groups is gnomonic, in which geometrically similar, but pro-
gressively larger increments are added to shell or other
structure as the animal grows. The most widely known gno-
monic pattern is the equiangular spiral whose properties are
summarized by D’Arcy Thompson (1917). From a point of
origin on the embryonic molluscan shell, calcium carbonate
is secreted by pallial glands according to a definite spiral
law subject to genetic, ontogenetic, and seasonal variations
or pathology, and becomes a permanent record of some of
the laws governing the life activity of the animal capable of
fossilization. Patterns of curvature in Venericardia were
examined during the present investigation in an attempt to:
a) distinguish stages of growth such as those reported by
Currie (1942) among juvenile ammonites, b) determine if
phylogenetic significance can be attached to curvature pat-
terns and c) determine if biologic significance may be at-
tached to ontogenetic stages of growth in fossil bivalves by
a comparison of Recent forms.

30

20

FREQUENCY

fo) ta Sl
15 30 45 60

HEIGHT IN MILLIMETERS

Text-figure 10. Frequency-distribution histogram of height in
sample of 165 individuals of /’. (C.) alticostata from AMNH locali-
ties 1055, 984 and collection 9950/1 (topotypes). A, B, C, D and E
are peaks probably representing year classes. Inferred class below A
not distinguishable. Class interval 3 mm.

Properties of the equiangular spiral. — The general
formula for the equiangular spiral is: r = e@cota, in which

r is the ratio between successive polar radii (Pn+1/Pn)
measured at equal angular increments (6) about the origin
of the curve (see Text-fig. 11). The term “e” is the root of
Napierian (natural) logarithms, having an approximate
value 2.718, a is the acute spiral angle of the curve formed
by the intersection of any polar radius and the tangent to
the curve at the point of intersection. Limiting cases are
a = 0°, in which the curve would be a straight line, and
90°, in which the curve would be a circle of no dimension.
Higher spiral angles result in highly convex curves typical of
gastropod growth, while low spiral angles characterize most
pelecypods. Values found in Venericardia were generally
less than 70°, and mostly 60° or less.

Text-figure 11. Equiangular (logarithmic) spiral showing im-
portant relationships. O, origin, p, radius, 9, angular increment be-
tween successive radii, 4, spiral angle.

66 PALAEONTOGRAPHICA AMERICANA (VI, 39)

Method of study. — An infinite number of spirals
radiating from the beaks and differing in spiral angle may
be drawn for each clam shell. Lison (1949) introduced the
concept of the “directrix rib” which is unique on each valve
and lies entirely in one plane (“directrix plane”). All other
ribs, when viewed normal to this plane, will have the same
spiral angle as that of the directrix rib. This was not sub-
stantiated in the present investigation, and, although this
rib may be readily identifiable on orthogyral forms (e.g.
many members of Pecten), its identification in strongly pro-
sogyral forms such as Venericardia proved impossible, and
all ribs curve in a turbinate (three-dimensional) sense,
agreeing with Owen’s (1953) conclusion.

In order to standardize curvature analysis for interspeci-
fic and intraspecific comparisons, the rib which defines the
greatest width of the shell was chosen to represent the
curvature of the entire shell, corresponding to Owen’s (op.
cit.) concept of the “normal axis” or line of greatest con-
vexity. The curvature of this rib is also a good indicator of
the convexity of the shell. For simplicity, the three-dimen-
sional aspect of the rib was ignored, and curves treated as
plane figures in the present investigation.

The curve of the highest rib was traced, orienting the
shell with the antero-posterior axis (APAX, Text-fig. 4) co-
incident with the line of sight of the observer in one of two
ways: a) sighting through a telescopic device equipped with
an ocular grid which allowed the curve to be transferred to
an enlarged grid on tracing paper, or b) by enlarged anterior
view photographs of the specimens. The results of both
methods were comparable.

Radii of each curve were measured at 10° intervals (4)
and plotted on semilogarithmic graph paper. (See Text-fig.
13). On a spiral angle protractor (Text-fig. 12), the value
of the spiral angle for each curve or segment of the curve
was obtained. The use of this protractor eliminated the need
for about 25 ratio calculations for the radii of each curve.

Patterns of curvature in Venericardia. — Instead of the
anticipated straight line semilogarithmic plot of radii of a
curve of constant spiral angle (C, Text-fig. 13), curves
drawn from shells of Venericardia exhibited different seg-
ments indicating growth in a sequence of stages in two dis-
tinct patterns (A and B, Text-fig. 13). Pattern A consists
of four segments increasing in value of spiral angle (a)
from beak to ventral margin. Pattern B consists of a similar
sequence of four stages on which is superimposed a fifth
stage of lower convexity, resulting in a somewhat sigmoidal
pattern. In the curvature characteristics noted for each
species in the present investigation, the spiral angle values
are recorded under S1 to S5, and the shell height at the point
of inflection between segments given as H1 to H4, from beak

Log 9,

Text-f{igure 12. Semilogarithmic protractor of Log pn ys. n@ for
whole degree values of a from 20° to 70°, 9 = 10°.

Log Pn

ne

Text-figure 13. Semilogarithmic plots of curvature patterns. A, four
segments (S1, S2, S3, $4) of increasing spiral angle. B, four seg-
ments of increasing spiral angle (S1 to $4) and fifth segment (S5) of
lower spiral angle. H1 to H4, heights at points of inflection between
segments. C, single spiral angle (S1).

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 67

to ventral margin of the shell (see Text-fig. 13). In some
shells an additional segment of extremely high convexity
(about 70°) is noted, and this is attributed to gerontism,
being recorded from only large specimens. In general, one
or the other of these basic patterns characterizes all mem-
bers of a species, but in several cases both are found in
members of a single population sample (see Text-figs. 14,
16). This has more than one interpretation. In some cases
[e.g. V. (C.) complexicostata, Text-fig. 14], the A pattern
is found only among small specimens which have not at-
tained the mature final segment. This is also noted in V.
(C.) alticostata, (see Text-fig. 15), as well as the presence of
a gerontic segment (Text-fig. 15, Nos. 7, 9, 13). In other
cases both patterns characterize fully mature individuals
[e.g. V. (R.) rotunda, Text-fig. 16]. The possible implica-
tions of this are discussed below.

INTERPRETATION OF CURVATURE PATTERN

Jackson (1890) noted four post-embryonic stages of
growth among several groups of pelecypods which he termed
(1) nepionic, or that of first formation of the true shell
(dissoconch) following the embryonic prodissoconch stage;
(2) nealogic, or “the period succeeding the nepionic and
preceding that period which may be properly considered as
the adult,’ and which may differ markedly from the (3)
ephebolic, or adult condition which is characterized by the
adult ornamentation and other features typical of the
species. Stage (4) is geratologic, or the period of old age.
These stages are not recognized in Venericardia as distinct
ornamental changes recognized by Jackson (op. cit.) or
Nicol (1952) in the genus Echinochama, nor do the inflec-
tion points in curvature patterns among venericards corre-
late with the common yarix-like interruptions in growth
seen on shells of that group.

In fact, the high variation found in shell height at each
of the characteristic inflections found in most species is not
in keeping with the regularity of curvature patterns, and
the meaning of this irregularity is not understood.

The primitive baluchicards, including V. (B.) beau-
monti (d’Archiac and Haime) have a curvature pattern of
the A type, although the most advanced form, V. (B.)
greggiana Dall of the early Eocene displays only the B pat-
tern. The claibornicards are characterized by the B pattern,
although V. (C.) alticostata Conrad of the late Claibornian
has developed the A pattern in certain members of the
population, and the monotype of V. (C.) nasuta Dall of the
(?) middle Oligocene also exhibits the A pattern. Early
glyptoactids [e.g. V. (G.) serricosta (Heilprin), V. (G.)
nodifera Kellum] exhibit both patterns of curvature, but

20644 pee:
17.5, 20645__| j3> 14.0 1362

Loc

NO

Text-figure 14. Semilogarithmic plots of curvature in V. (C.)
complexicosta. Pattern A in immature individuals BEG 20645
and (holotype) (see page 105). Larger individuals (BEG 20644, 1, 3, +)
show B pattern with addition of less convex segment. Spiral angles
indicated at left of curve segments, height in millimeters indicated
at inflection points between segments.

353-7 367 9 13
36.4372 ,36.1

Loch}

N@

Text-figure 15. Semilogarithmic plots of curvature in five in-
dividuals of I’. (C.) alticostata, AMNH 9950/1. Terminal segment
of immature individual (No. 34) equivalent to medial stage of larger
forms. Highly convex small terminal segment added to normal pat-
tern in Nos. 7 and 9 indicates gerontism. Individuals of “si/limani”
tvpe (not shown) have B pattern similar to figure 14, nos. 20644, 1,
3, 4. Spiral angles indicated at left of curve segments, height in
millimeters indicated at inflection between segments.

68 PALAEONTOGRAPHICA AMERICANA (VI, 39)

only the B type is found in more advanced members. Both
patterns are found in Rotundicardia, and V. (R.) rotunda of
the middle Eocene displays a distribution of about 50 per
cent for each pattern at the type locality in Monroe County,
Alabama.

Hermaphrodism, in which (1) a single individual un-
dergoes several male and female stages, (2) developmental
sex change, in which the same individual matures from one
sex to the other and (3) true hermaphrodism, in which an
individual fulfills simultaneously both male and female
sexual functions are well known among mollusks. Orton
(1937) documented the alternation of sexuality in members
of Ostrea edulis Linn., noting also protandric hermaphro-
dism in the calyptraeacean gastropod Crepidula fornicata
(Linn.) in which the young are born males, developing with
growth into females with the loss of male reproductive or-
gans. Loosanoff (1936) reported that the gonads of tiny
(less than 7 mm length) Mercenaria mercenaria ( Linn.) are
male in function, and after discharging mature sperm into
the water during the first summer of life, about half the
members of the generation differentiate into true males and
females. During the following summer, the remainder of
the generation also differentiate. Among fresh-water unioid

clams, a distinct sexual dimorphism characterizes most
species, and females are much more inflated, growing ac-
cording to much higher spiral angles than the males. This
is correlated with the “marsupial” habit among these forms,
and females retain maturing glochidia in the tubules of
specialized portions of the gills.

Protandric hermaphrodism is suggested as the inter-
pretation for the curvature pattern of the A type found in
Venericardia. The two early stages (S1,S2) may be juvenile
and sexually immature. The third (S3) may represent the
male stage, followed by the female highly convex stage
(S4). A similar pattern of curvature is found in V. (Cyeclo-
cardia) borealis Conrad which lives in coastal waters along
the North American east coast, in which “. . . the young in
large numbers are incubated in the ovary or its atrium in
the umbonal cavities of the valves” (Dall, 1903, p. 1407).
The terminal stage of high convexity may be correlated with
the onset of the female “marsupial” function well known
among carditids. Oddly enough, in a living sample of this
species from the waters near Woods Hole, Massachusetts,
not a single male was noted among ten individuals of over
10 millimeters in length.

The interpretation of pattern B which appears uni-

A CURVATURE

23.9

5268 5270b 5272L 5274
5269 S527iL 52

B FUNICULATE

FUNGINATE

B CURVATURE

ne

Text-figure 16. Semilogarithmic plot of curvature in type collection of I’. (R.) rotunda
showing correlation between ornamentation and curvature pattern.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 69

formly in most claibornicards is not readily apparent. This
pattern, formed by the addition of a less convex segment
to the ancestral baluchicard pattern may indicate the loss
of the marsupial function, although the A pattern is re-
developed in V. (C.) alticostata. That it signified the re-
turn to maleness is highly unlikely, since a larger shell will
accommodate more larvae than a smaller one.

The interpretation of the presence of both patterns
within a species is easily explainable. Text-figure 16 shows
the semilogarithmic plots of curvature for the type collec-
tion of V. (R.) rotunda Lea. Twelve individuals show curva-
ture of the A type, and eight of the B. Two types of orna-
mentation are represented in the population at the type
locality: (1) that which Harris (1919) called funiculate,
or somewhat simply noded, and (2) funginate, or forms
with flaring nodes (see species discussion), The distribution
of ornamental type (see Text-fig. 16) corresponds to that of
curvature pattern, the funiculate specimens having A curva-
ture, and the more ornate funginate specimens having B
curvature, with the exception of two small forms which
have not yet attained the terminal stage of lower convexity.
This distribution of ornamentation and curvature pattern
is best explained as sexual dimorphism, in which the plainly
ornamented, highly convex specimens are female and the
more ornate, flatter members of the population males. The
same explanation may be applied to V. (C.) alticostata, in
which the more primitively ornamented “‘sillimani” speci-
mens, appearing in small numbers among the common “a/ti-
costata’ forms, exhibit the B curvature pattern, and are
male, while the convex “alticostata” specimens with A curva-
ture and highly simplified ornamentation are females. It may
be significant that this development appears simultaneously
in two distinct evolutionary lineages which may have been
competing in Gosport seas.

EVOLUTIONARY TRENDS

In the evolution of the alticostate venericards, several
important evolutionary trends are noted:

Dental pattern. — (Yext-figure 17.) Although the ro-
tundicards maintain the primitive dental pattern of Balu-
chicardia, retaining the massive triangular left anterior
cardinal (2), the claibornicards evolve away from this condi-
tion to a weakened and flattened 2, and the right anterior
cardinal becomes correspondingly flattened and occupies
a position in-line with the medial tooth 3b and lying over
2 when valves are occluded, rather than on the anterior
border as in the baluchicards and rotundicards. This clai-
bornicard pattern is maintained in the evolution of Glypto-
actis in the late Oligocene, although a marked reduction in
3a takes place in the Miocene evolution of the group.

Umbonal height increase. — A plot of the average ratio
of beak height to height (BH/H) vs. time (Text-fig. 18)
indicates a phyletic trend toward a reduction in the value
of this ratio, signifying a progressive increase in the height
of the umbones over the commissural disk of the shell. This
may be correlated with expansion of the umbones to house
more efficiently the developing larvae in “marsupial” re-
production,

Size increase. —A plot of maximum observed height
vs. time (Text-fig. 19) shows a marked increase in size in
Baluchicardia and a decrease in Rotundicardia. No sys-
tematic trend is observed in Claibornicardia, although the
Gosport Sand species, V. (C.) alticostata is much larger than
any ancestral form, attaining heights of over 50 millimeters,
while other Eocene species cluster around a value of 20
millimeters. A great range of size in Glyptoactis, likewise
has no consistent pattern. (These are not shown in Text-
fig. 19.)

Shape ratio. — A plot of mean values of Height/Length
ratios vs. time (Text-fig. 20) shows trends toward increase

in Baluchicardia (B), Rotundicardia (R) and Glyptoactis,

(See sie

GLYPTOACTIS

aS

ROTUNDICARDIA

BALUCHICARDIA

Text-figure 17. Patterns of dentition in the four subgenera
Baluchicardia (V. (B.) bulla), Rotundicardia (V. (R.) rotunda),
Claibornicardia (V. (C.) linguinodifera), Glyptoactis (V. (G.) nodi-
fera nodifera), showing divergences from ancestral pattern.

70 PALAEONTOGRAPHICA AMERICANA (YI, 39)

indicating a reduction in relative elongation in the evolu-
tionary history of those groups. The meaning of this is not
clear. Claibornicardia (C), on the other hand, shows a strik-
ing decrease in this value, indicating a trend toward greater
relative elongation, reaching the zenith in V. (C.) nasuta
Dall of the (?) middle Oligocene.

Ornamentation. — Two striking trends are evident in
(a) increase in rib number (Text-fig. 21) noted in Balu-
chicardia and Claibornicardia but not Glyptoactis. In the
latter subgenus, the dominant trend (b) is the simplification
of ribs by the reduction of primitive paracostal cords, ex-
pansion of the central cord, and simplification of costal
nodes. This is seen on the phylogenetic diagram, Text-figure
28. This tendency is also sporadically found in Baluchicardia
in V. (B.) francescae Gardner and Bowles and JV. (B.)
whiter Gardner. The latter species is a close approximation
to V. (Glyptoactis) hesperide Gardner of the Shoal River
Miocene in shape and ornamentation, and may indicate con-
vergence into the same ecologic niche. The tendency is also
somewhat evident in Claibornicardia, in which several spe-
cies have vestigal paracostals [e.g. V. (C.) alticostata, V.
(C.) complexicosta Meyer and Aldrich] (see Text-fig. 27),
and even exhibit flattened costal nodes, most notable on
the posterior ribs. The reduction of paracostals is charac-
teristic also of Rotwndicardia, and the ribs in the main
lineage of this group become high and bladelike, but in the
other lineage, the paracostals are actually accentuated and
form flangelike projections over the intercostals (see Text-
fig. 26).

The meaning of trends in ornamentation is not known.
It is probably safe to assume that most of the alticostate
venericards were shallow burrowers, and that the ornate

@ GLYPTOACTID
@ CLAIBORNICARD
@ ROTUNDICARD
@ BALUCHICARD

Q
Tp
Tm
To
Te

Tf
Text-figure 18. Mean BH/H vs. time, showing general shift

toward lower value in entire alticostate group indicating increase
in umbonal inflation.

90 1.00

MEAN BH/H

ribs were anchoring deyices in the sediment. Planicostate
forms, with generally heavier shells may have lived on the
sediment surface, and trends toward flattening and simpli-
fication of the costae may indicate shallower burial or a
surface habitat. The modern V. (Cyclocardia) borealis Con-

@ ROTUNDICARD
@ BALUCHICARD

MAXIMUM HEIGHT IN MILLIMETERS

Text-figure 19. Maximum height vs. time in Baluchicardia and
Rotundicardia showing progressive increase in the former subgenus

and decrease in the latter.

AMERICAN ALTICOSTATE

Q
® GLYPTOACTID
Tp @ CLAIBORNICARD
O ROTUNDICARD
@ BALUCHICARD
;
se
Cc S R
(o}
AP e ~
e e 8 © 00
ee ef
e ¢
B
_ O®
00 ¢@ ~ e0 0 @
75 .80 85 .90 95 1.60 105 1.10
MEAN H/L

Text-figure 20. Mean H/L vs. time. Increase in value (decrease
in relative length) in Baluchicardia (B), Rotundicardia (R), and
Glyptoactis. Decrease (increase in relative length) in Claiborni-
cardia (C).

rad lives on the sediment surface and forms an abundant
food supply for cod and other large fish on the Grand Banks.
This form has highly simplified costals (see Text-fig. 7) and
a dense shell much like those of many planicostate veneri-
cards of the early Tertiary which also may have been epi-
faunal benthos.

SEXUAL DIMORPHISM IN TWO LINEAGES

In Rotundicardia, V. (R.) rotunda contains individuals
of two types in the populations of the Gosport Sand bed, as
discussed above under “curvature patterns.” In earlier
populations of the Wautubbee Marl in eastern Mississippi,
both curvature patterns are noted, but only the “funiculate”
type of ornamentation is represented. The less convex and
ornately sculpted males and the more convex females of the
Gosport Sand give rise to V. diversidentata Meyer, in which
sexual dimorphism is also evident in the two forms which
Dall (1903) referred to V. “jacksonensis” Conrad and J.
“praecisa” Dall. These occur in the same sample and are
distinguished by shape and convexity, although both have
the B pattern of curvature. It is evident that the males are
the smaller, flatter, and more “precisely” ornamented forms
(C, Text-fig. 22; Pl. 23, figs. 5a,5b,6) and the females more
convex with inflated beaks for the “marsupial” function in

reproduction (B, Text-fig. 22; Pl. 23, figs. 3a-3c,4). The

VENERICARDS:

HeEaAswip 71

@BGLYPTOACTID
@ CLAIBORNICARD
@BALUCHICARD

Q
1p
Tm
To
Te

10 15 20 25 30 35 40
MEAN RIB NUMBER

Text-figure 21. Mean rib number ys. time. Increases in Baluchi-
cardia and Claibornicardia. No trend in Glyptoactis.

Text-figure 22. Development of sexual dimorphism in Rotundi-
cardia. A, V. (R.) rotunda of the Gosport Sand showing male orna-
mentation on anterior and female ornamentation on posterior half of
cut away right valve. B, V. (R.) diversidentata female with high
beaks, inflated umbones. C, male (“fraecisa” type) of the latter
species with low umbones,

72 PALAEONTOGRAPHICA AMERICANA (VI, 39)

early Oligocene descendant, V. (R.) carsonensis, fails to
exhibit any obvious character indicating sexual dimorphism.

In Claibornicardia, all species grew according to the B
pattern of curvature, except V. (C.) alticostata, in which
most individuals exhibit the A pattern. The rare specimens
with B curvature are more primitive in characters of orna-
mentation, having tripartite ribs with well-developed para-
costals (C, Text-fig. 23; Pl. 26, fig. 11), whereas the indi-
viduals with A pattern of curvature have highly simplified
central and posterior costals, a thicker shell and somewhat
heavier cardinal teeth (B, Text-fig. 23; Pl. 26, figs. 8-10).
The primitive individuals have been traditionally assigned
to V. “sillimani” Lea, while the more convex individuals
with simplified paracostals have been assigned to V. alti-
costata Conrad. A sample of this species from Gopher Hill
in Washington County in western Alabama contains in-
dividuals showing only the “si/imant” ornamentation (A,
Text-fig. 23; Pl. 26, figs. 6,7), but both patterns of curva-
ture. This seems to be a case of sexual dimorphism similar
to that in V. (R.) rotunda and its descendant. Of interest is
the presence at Gopher Hill of V. (R.) rotunda showing only
the primitive pattern of ornamentation (funiculate) and
the absence of highly specialized funginate male ornamenta-
tion found at Claiborne, Alabama. This may indicate a
slightly earlier age than that of the Claiborne locality, al-
though the Alabama Geological Survey correlates the sand
bed in which these species occur with the Gosport Sand at
Claiborne.

In Baluchicardia, sexual dimorphism may be repre-
sented by highly ornate males which Dall called “variety
tripla” and female forms with nontripartite costate (e.g.

the holotype of V. (B.) wilcoxensis Dall) (Heaslip, 1964).

EVOLUTIONARY HISTORY OF THE
ALTICOSTATE VENERICARDS

GEOGRAPHIC SCOPE OF INVESTIGATION

The present study is limited to the eastern and Gulf
coastal plains of North America as a natural faunal pro-
vince, especially during the early Tertiary, when Eocene
species or closely related forms ranged from Texas to New
Jersey in coastal waters. Localities for studied material ap-
pear under each species. References to other distribution
data are cited under “Geographic distribution” with each
species.

STRATIGRAPHIC SCOPE OF INVESTIGATION
The main evolutionary history of the venericards in

eastern North America occurs in the period of time begin-
ning with the Paleocene and ending in the late Miocene. The

Text-figure 23. Evolution of sexual dimorphism in /’. (C.) alti-
costata. A, typical morphology in sample from Gosport Sand at
Gopher Hill. B, highly inflated female with simplified posterior
ornamentation from Gosport Sand at type locality. C, male (“silli-
mant’ type) from Gosport Sand at type locality.

stratigraphic units mentioned in the present investigation
appear in the Chart (Text-fig. 24) compiled from a number
of sources.

SOURCES OF MATERIAL STUDIED

Collections examined during this investigation are
from several sources, and these are noted in each species
description.

(1) Collections made in the field during summers of
1955 and 1957 from eastern Mississippi to Virginia which
have been acquired by the American Museum of Natural
History. These, as well as existing collections at that institu-
tion were studied. (Notation AMNH.)

(2) Type collections of Dall’s and Gardner’s species, as
well as those of Kellum and Mansfield, in the U.S. National
Museum. (Notation USNM.) U.S. Geological Survey col-
lections were also examined, (Notation USGS.)

(3) Types and figured specimens of Harris in the Pale-
ontological Research Institution. (Notation PRI.)

(4) Lea’s and Conrad’s types and other collections in
the Academy of Natural Sciences of Philadelphia. (Notation
ANSP.)

(5) Stenzel, Krause, and Twining (1957) figured and
described specimens in the Texas Bureau of Economic Geol-
ogy collections. (Notation BEG.)

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74 PALAEONTOGRAPHICA

(6) Meyer’s type of V. (R.) diversidentata in the
Alabama State Museum collection. (Notation ASM.)

(7) Meyer and Aldrich type of V. (C.) complexicosta
formerly in the Johns Hopkins University collections,
missing United States National Museum. (Notation JHU.)

(8) Florida Geological Survey collections. (Notation

FGS.)

V.(B.) BULLA

em

k hg
EY

V.(B.) AMPLICRENATA

Toe
P VB) MOA

V.(B.) HESPERIA

AMERICANA (VI, 39)

ALTICOSTATE EVOLUTION
The alticostate forms of the genus Venericardia are a
prominent element in fossil faunas from Tertiary rocks in
the eastern and Gulf coastal plains of North America, al-
though their abundance and distribution decrease markedly
after the Eocene. (See Text-figs. 1, 3.) They are naturally
divisible into four closely related, but distinct evolutionary

V(8) GREGGIANA

V.(B.) WHITE/

Text-figure 25. Evolutionary pattern of Baluchicardia in shape, ornamentation, and den-
tition. Right valves cut away to show left dentition. Dentition unknown in /’. (B.) moa, V. (B.)
amplicrenata, V. (B.) hesperia, and V. (B.) whitei, left dentition restored in V. (B.) francescae.
Double vertical line next to each species indicates stratigraphic range. Branching solid line
indicates inferred relationships.

AMERICAN ALTICOSTATE VENERICARDS: FIEASLIP 75

lineages designated as subgenera, and the evolutionary his-
tory of the entire group is best described by considering the
history of each subgenus in a time sequence.

EVOLUTIONARY HISTORY OF BALUCHICARDIA IN
NORTH AMERICA

Appearing without apparent antecedent in Maestrich-
tian and Danian beds of the Sind region in northwest India
and simultaneously in Baluchistan and Iran to the west, as
well as northern Africa, V. (B.) beawmonti (d’Archiac and
Haime) and closely related forms spread rapidly to the New
World (see PI. 20, figs. la,lb). Rutsch (1936) gave an ac-
count of the distribution and age relations of these ancestors
of the “alticostate” and, presumably, the “planicostate”
lines of evolution.

The presence of members of Baluchicardia is recorded
first in North America from rocks of early Paleocene age in
Texas (Kincaid Formation), Mississippi (Clayton Forma-
tion), and in the Prairie Bluff Beds of Alabama, although
the Alabama forms are too poorly preserved for description
here. V. (B.) hesperia Gardner most closely approaches the
Cretaceous ancestor, V. (B.) beawmonti (d’Archiac and
Haime) in shape, convexity, and ornamentation, and its
occurrence late in the early Paleocene (upper Kincaid,
Tehuacana Member) testifies to a greater stratigraphic
range than the record indicates.

From a central evolutionary stock much like that
species, the less typical baluchicards of the early Paleocene
must have radiated, producing a number of species modified
in shape or ornamentation from the ancestral pattern. Both
V. (B.) whitei Gardner and V. (B.) francescae Gardner and
Bowles evolve simplified ornamentation until only vestiges
of tripartite costal structure remain in the latter species.
The total simplification in costae in the former species is
an almost exact convergence to V. (Glyptoactis) hesperide
Gardner of the Shoal River Miocene in Florida.

Two other species of the early Paleocene (Tehuacana
Member of the Kincaid Formation) in Texas differ from
the ancestral pattern in the reduction in rib number and
the amplification of the irregular sawtooth costal nodes of
the central costal cord. V. (B.) amplicrenata, n. sp., retained
a quadrate shape similar to that of V. hesperia, but V. (B.)
moa Gardner evolved a posteroventral elongation similar to
that found in many of the more advanced claibornicards
of the middle Eocene,

Of the early Paleocene species which lived in the waters
of the Gulf Coast, only the baluchicard lineage of V. (B.)
hesperia appears to have carried into late Paleocene time,
and the two species known from the Wills Point Beds

(Texas) and Naheola Formation (Alabama) retained the
inflated quadrate form of the Kincaid ancestor. V. (B.) bulla
Dall and V. (B.) wilcoxensis Dall represent comparable ele-
ments in the western and eastern margins, respectively, of
the Mississippi embayment, but indicate a marked reduc-
tion of the former importance of Venericardia as a faunal
element. V. (B.) bulla Dall disappeared by the end of Mid-
way deposition without continuation of its lineage, whereas
V. (B.) wilcoxensis Dall appears to have continued into Wil-
cox time in V. (B.) greggiana Dall, a distinctive but geo-
graphically restricted element in the Tuscahoma fauna of
Alabama and the last of the baluchicards in North America.

The ecologic roles vacated by the extinct members of
Baluchicardia were filled during the middle Eocene by the
more advanced members of Claibornicardia which under-
went its own “adaptive radiation” during Claiborne time
on the Gulf Coast and Atlantic Coast.

EVOLUTIONARY HISTORY OF ROTUNDICARDIA IN
NORTH AMERICA

The earliest known form of the subgenus is found in
the Kincaid Formation of early Paleocene (Midway) age
in Bastrop County, Texas. V. (R.) eoa Gardner, with its
tripartite costal structure and primitive, heavy dental pat-
tern containing a well-developed almost vertical 3a (anter-
ior cardinal) in the right valve, seems to have radiated from
the ancestral baluchicards during this time. Its “rotund”
outline, lenticular shell profile and narrow, bladelike costal
cord began the lineage which later became a widely distri-
buted element in middle and late Eocene faunas.

In the primary radiation of the alticostate venericards
in the early Paleocene, another “rotund” form was produced
in V. crenaea Gardner (see Text-fig. 26), and its flattened
profile, rounded outline and low, inconspicuous beaks are
thought to indicate a side-branch of the main rotundicard
stock, even though its ornamentation is strongly developed.
This species may have given rise to the similarly orna-
mented V. (R.) eutawcolens Harris of the middle Eocene
Santee Limestone. Unfortunately, dental patterns are un-
known in both, making association here tentative.

Curiously, no rotundicard species is known from the
late Paleocene or early Eocene (Wilcoxian) sediments, but
neither are the other alticostate species belonging to the
other subgenera abundantly represented in rocks of this
age (see Text-fig. 3).

Appearing only in the middle Eocene (Claibornian ),
V. rotunda Lea exhibits shape and dentition which clearly
indicates the continuation of the lineage begun by V. eoa
Gardner, but with highly simplified bladelike paracostals,
indicating a high degree of evolutionary advance over the

PALAEONTOGRAPHICA AMERICANA (VI, 39)

V.(R.)CARSONENS/S

V.(R.)JEUTAWCOLENS

~ VR.) ROTUNDA \

es V.(R.) CRENAEA
LZ SEES

VIRJEOA

Text-figure 26. Evolutionary pattern of Rotundicardia in shape, ornamentation, and den-
tition. Right valves cut away to show left dentition. Dentition unknown in I’. (R.) crenaea and
V. (R.) eutawcolens. Both morphological types shown in I’. (R.) diversidentata, with male
(“praecisa”) on left, female on right. Double vertical line next to each species indicates strati-
graphic range. Branching solid line indicates inferred relationships.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 77

Midway species. The earliest reported occurrence of JV.
rotunda Lea is by Foster (1940) from the “lower Claiborne”
in Clarke County, Mississippi, but the species is most widely
distributed and abundant toward the latter part of Claiborne
(middle Eocene) time, when it is recorded from Cook
Mountain and Gosport Sand sediments from northeast
Mexico (Laredo Formation) to New Jersey (Shark River
Marl). The ornate “funginate” nodulation of the ribs which
characterizes the male members of the species does not ap-
pear until the Gosport Sand, and pre-Gosport forms have
fairly uniform development of the simpler “funiculate”
nodes which characterize the Gosport females of the species.

A closely related species appears in the Weches Forma-
tion in Bastrop County, Texas, and although V. flabellum
persists into the lower part of the Yegua Formation (Gos-
port Sand equivalent), its recorded geographic distribution
is only Bastrop and Brazos Counties where it appears to
have lived side-by-side with V. rotunda and was only of
local importance in the middle Eocene faunas of the Gulf
Coast.

The direct Jacksonian (late Eocene) descendant of V.
rotunda is V. diversidentata Meyer which is also widely dis-
tributed over the North American Gulf Coast during this
time. This species, however, is not reported from Texas or
the Carolinas, and its range from Louisiana and Arkansas
to Georgia and Florida is somewhat more restricted than
that of its Claiborne ancestor. Gardner (1945) described
imperfectly preserved forms of Jackson age in northeast
Mexico with a lower rib count than most topotype speci-
mens. This may be comparable to the form which Dall
named JV. vicksburgiana Dall of the Ocala Limestone in
Florida, which is reduced to a subspecies of V. diversiden-
tata, and differs from the topotype specimens only in a
lower rib count. Gardner’s Mexican specimens have not
been examined by the writer, and no comment can be made
on their relationships.

From the profusion of Claibornian alticostate species
of Venericardia, V. diversidentata of the Jackson is one of
the few which survived to continue into the Oligocene as
V. carsonensis Dall of the Red Bluff Formation. However,
the restricted geographic range (Wayne County, Missis-
sipp1) testifies to the reduced importance of Venericardia in
the marine faunas of eastern and Gulf Coastal North
America. The stringent reduction of the alticostate lines of
evolution is accompanied by a total extinction of the forms
which Gardner and Bowles (1939) assigned to the “Plani-
costa Group” at the end of the Eocene, and seems to herald
either a (1) climatic or environmental condition unfavor-
able to the life or preservation of Venericardia, or (2) in-
creased competition from a similarly adapted bivalve group.

Gardner (1945) similarly reported poorly preserved
venericards which she compared to V. carsonensis in the
lower Oligocene sequences as well as higher in the Oligocene
section in northwest Mexico. This indicates that this species,
or closely related forms may have continued in this area
during middle Oligocene time. In the Chickasawhay Beds
of Wayne County, Mississippi, and Monroe County, Ala-
bama, a species V. nodifera waynensis Mansfield appeared
in the late Oligocene. This species is certainly descended
from the middle Eocene claibornicards, but modified to the
extent that it must be associated with the subgenus Glypto-
actis, and indeed is the base for the evolutionary “explosion”
of Venericardia again in the early Miocene and the rein-
vasion of the eastern coastal regions as far north as North
Carolina.

EVOLUTIONARY HISTORY OF CLAIBORNICARDIA IN
NORTH AMERICA

The evolution of the claibornicards from the baluchi-
cards of the Paleocene is apparent in similarities in shape
and ornamentation, but the distinct dental pattern of the
descendant group is first recorded in V. linguinodifera, n.
sp. of the Bashi Member of Wilcox (early Eocene) age. The
most likely ancestor of this small species is V. (B.) wilcox-
ensis Dall of the Matthews Landing Member in Alabama,
although a severe reduction in size, as well as the radical
change in dental pattern and curvature, took place in the
transition.

From this single species of the Wilcox evolved V’. (C.)
coloradonis texalana Gardner of the lower Weches Forma-
tion in Texas, with a slight increase in rib number and a
more well-defined posterior truncation and postero-ventral
elongation. This subspecies of limited stratigraphic range
(lower Weches) and distribution (vicinity of the type lo-
cality in Burleson County, Texas) gave rise to the sub-
species V. (C.) coloradonis coloradonis Harris, characterized
by a much higher costal number and a much more exten-
sive stratigraphic range (Weches Formation to Wautubbee
Marl), spanning much of the middle Eocene. The ancestral
lower Weches subspecies also gave rise to V. (C.) natchi-
toches Harris during Weches deposition. An opposite trend
toward costal decrease is noted in this species which, unlike
V. (C.) coloradonis coloradonis, appears to end its history
in the Weches without issue, although V. (C.) perantiqua
Conrad of the Shark River Marl of New Jersey is closely
related and undoubtedly was derived from the same an-
cestor in the Weches. This species also seems to end in Shark
River deposition, and no subsequent form recorded is its
descendant. Both of these species have reduced and irregu-
larly noded ribs and indicate a trend in this direction.

78 PALAEONTOGRAPHICA AMERICANA (VI, 39)

A highly specialized lineage of Claibornicardia evolved
from V. (C) coloradonis coloradonis during Weches deposi-
tion and began in the monotypic V’. (C.) trapaquaroides, n.
sp., whose beadlike costal nodes indicate an ornamental

simplification which is continued almost unmodified in the

V.(C.) COMPLEX/ICOSTA

V.(C.) NATCHITOCHES

flattened and elongated V. (C.) trapaquara Harris of the
Stone City Beds. This species is highly specialized in its
low convexity shell and pointed postero-ventral process and
appears to represent another “dead-end” in evolution as does
V. (C.) complexicosta Meyer and Aldrich which retained

V(C)NASUTA

V(C) PERANTIQUA

VC) COLORADONIS COLORADON/S

V.(C.) COLORADON/S TEXALANA

Text-figure 27. Evolutionary pattern of Claibornicardia in shape, ornamentation, and den-
tition. Right valves cut away to show left dentition. Right dentition restored in /’. (C.) nasuta,
left dentition restored in ’. (C.) trapaquaroides. Only simplified female shown for V’. (C.) alti-
costata, Double vertical line next to each species indicates stratigraphic range. Branching

solid line indicates inferred relationships,

AMERICAN ALTICOSTATE VENERICARDS: HEasLtp 79

the normal, highly convex shell of V. (C.) trapaquaroides,
but underwent a further reduction in costal node develop-
ment until the central and posterior ribs are almost flat-
topped. These three species indicate a trend toward increase
in rib number (about 28-30) over V. (C.) coloradonts colora-
donis (about 26).

Another lineage apparently derived from V’. (C.) colora-
donis Harris is that which contains the highly specialized
V. (C.) alticostata (Conrad). V. (C.) blandingi Conrad of
the lower Santee Limestone of Vance’s Ferry in Orangeburg
County, South Carolina, exhibits all of the primitive attri-
butes of the claibornicards in ornamentation and dentition
while lacking a high definition of posterior truncation which
is common to this group. This observation is based on a
single specimen with a complete posterior margin and may
actually represent an extreme condition and not typical of
the species. Nonetheless, this might be the ancestor of /.
(C.) alticostata of the Gosport Sand in Alabama and Mc-
Bean Formation in South Carolina as well as deposits
(Castle Hayne?) in North Carolina. The Gosport Sand
species grew to ponderous size when compared to the general
lineage of Claibornicardia. It is highly specialized in other
respects, as well, and the two different patterns of spiral
growth present in the species seem to indicate that males of
the population (“sillimant” types) retained the ancestral
patterns of ornamentation and growth, while the much more
abundant females (“alticostata” types) became somewhat
simplified in costal development and highly inflated, perhaps
in connection with specialized reproductive habits.

V. (C.) nasuta Dall is the only subsequent species to
V. (C.) alticostata which is placed within Claibornicardia,
but its stratigraphic placement is open to question, being
known from a single specimen and an unknown locality in
Conecuh County, Alabama. The interpretation set forth in
the present investigation is that the species is middle Oligo-
cene in age, but future work may establish a different view.
This species shows several advances over the ancestral
Gosport Sand form. Simplification of the tripartite costal
structure involves not only posterior ribs, but the first few
anterior ones, as well. The curvature pattern is that of the
female V. (C.) alticostata, although future collecting may
also establish the male pattern, The dental pattern, while
similar to that of V. (C.) alticostata shows a development
of the left anterior cardinal (2) which is similar, if not
stronger than that in the Paleocene subgenus Baluchicardia,
being erect and triangular in lateral aspect, and even sloping
downward toward the posterior. Because of this feature of
dentition and the highly elongated shape, this species may
be ancestral to the Miocene subgenus Carditamera, repre-

senting a form intermediate between Venericardia and
Cardita.

V. (C.) nasuta is the last recorded member of the vigor-
ous subgenus Claibornicardia in the American Cenozoic, and
the ecologic niches left vacant by this group were filled
again during the resurgence of evolutionary activity in the
Miocene glyptoactids which evolved from Claibornicardia.

EVOLUTIONARY HISTORY OF GLYPTOACTIS IN
NORTH AMERICA

After the claibornicard burst of evolutionary activity
in the middle Eocene, the total absence of this important
bivalve group in sediments of late Eocene age is enigmatic.
The last known species of Claiborne age in eastern North
America is V. (Claibornicardia) alticostata, which was
widespread in the Gosport Sand and its equivalents from
western Alabama to North Carolina, but failed to leave any
record in the sediments of Jackson age. The only species of
later age is V. (C.) nasuta Dall of the (?) Glendon Lime-
stone of middle Oligocene age from an unknown locality in
Conecuh County, Alabama, but this species is much too
highly specialized in shape, dentition, and ornamentation
to have given rise to the glyptoactids.

The first glyptoactid recorded from Gulf Coastal North
America is V. (G.) nodifera waynensis Mansfield of the
Chickasawhay Marl in Wayne County, Mississippi, and
Monroe County, Alabama. This highly variable subspecies
possesses many indications of claibornicard heritage (modi-
fied tripartite costate, similar dental pattern and shape),
but also the folded lunule which becomes an extra-cardinal
“tooth” in the more advanced species of the Miocene (see
lal, A7/, inka SKE)

From this subspecies radiated several forms of the
early and middle Miocene, as well as V. (G.) nodifera nodi-
fera Kellum of the “Trent”? Marl which is different from the
parent subspecies only in minor characters. These forms
include V. (G.) impendeocosta of the “Trent,” which ex-
hibits an early trend toward costal simplification and flat-
tening, V. (G.) gibberumbonata, showing also a costal
simplification but a degree of dental simplification not
found in other species, and V. (G.) himerta Dall of the Oak
Grove.

V. (G.) hesperide Gardner, exhibits an almost total
reduction in rib nodes, as well as highly elongated shape,
and is found in Shoal River sediments. It is unlike any other
glyptoactid of this time, in shape and costal simplification,
and may have been a form highly specialized for brackish
water conditions. V. (G.) serricosta (Heilprin) of the Tam-

80 PALAEONTOGRAPHICA AMERICANA (VI, 39)

pa Limestone retained the dental pattern of the Oligocene
subspecies and the ornamentation, but is somewhat more
elongated, and like V. (G.) gibberumbonata, n. sp., evolved
the highly inflated and inrolled beaks which characterize
all of the later glyptoactids.

From V. (G.) serricosta (Heilprin) of the early Mio-

SE 7
* (G.) HESPERIDE
J Ja“

V.(G.)IMPENDEOCOSTA _ V.(G.) GIBBERUMBONATA V.(G.)NODIFERA NODIFERA {\ 77

ae

~

cene evolved V. (G.) hadra Dall of the Chipola Beds of
middle Miocene age. This illustrates a trend toward umbonal
inflation and elevation from the somewhat inflated ancestor.
Extreme costal simplification and loss of the paracostal
cords characterizes the Oak Grove species, but the Chipola
form retained relatively strong and closely spaced costal

V.(G.)CRASSICOSTATA

V(G.) OLGA

V(G.)CHELOMODONTA

N\A =
V(G.) HIMERTA /— cm SS .
(CaN

V(G.) SERRICOSTA

\ VIG) NODIFERA WAYNENS/S

Text-figure 28. Evolutionary pattern of Glyftoactis in shape, ornamentation, and den-
tition. Right valves cut away to show left dentition. Right dentition restored in J’, (G.) im-
pendeocosta, V. (G.) chelomodonta, and V. (G.) gibberumbonata, \eft dentition restored in J’.
(G.) hesperide. Double vertical line next to each species indicates stratigraphic range. Branch-

ing solid line indicates inferred relationships.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 81

nodes, as well as paracostals on the central and anterior ribs.

Similarities in shape, but paracostal reduction and ex-
pansion of the central cord and transverse nodes, characterize
the evolution from V. (G.) hadra to V. (G.) chelomodonta,
n. sp., of the Shoal River and V. (G.) olga Mansfield of late
Miocene age. The former species evolved an added dental
element on the dorsal surface of the left posterior cardinal
(4b), however, and it does not appear to have given rise
to any subsequent species. V. (G.) olga of late Miocene age
with its tightly inrolled beaks, high umbones, and develop-
ment of an umbonal ridge and somewhat vaulted posterior
slope, appears to be an ideal ancestor for the highly ad-
vanced glyptoactids of the Pleistocene and Recent faunas
of the eastern Pacific middle latitudes, but it is the last
known glyptoactid in the eastern or Gulf Coast of the
United States.

Three species of Venericardia living in modern seas off
the western coasts of South America, Central America, and
North America from northern Peru to California are evi-
dently descendants of Glyptoactis. Of the three, V. (G.)
tricolor (G. B. Sowerby I), 1833, is the most generalized,
maintaining the broad, noded ribs of the Pliocene ancestors.
V. (G.) crassicostata G. B. Sowerby I, 1825, has ribs which
are almost completely flattened and smooth, except for sub-
dued nodes appearing mainly on the anterior ribs. This
species grew to extremely large size (length to almost 70
mm, Olsson, 1961), comparable to that of certain Eocene
“planicostate” species, and perhaps the combination of
flattened ribs and large, heavy shell indicates convergence
in habitat with that group. Photographs of this species are
included for comparison (see Pl. 29, figs. 7-8). The last
species, Venericardia (Strophocardia) megastropha Gray,
1825, possesses ribbing so simplified that the fresh shell with
complete periostracum appears almost smooth. This feature
and the extremely coiled beaks justified the erection of the
new subgenus Strophocardia Olsson, 1961. However, its evo-
lution from glyptoactid ancestry cannot be denied, even
though it signifies a further simplification of ancestral
characteristics.

Both V. (G.) crassicostata and V. (S.) megastropha are
known from the Pleistocene of Baja California (Durham
1950), and it is evident that the glyptoactid ancestors in-
vaded that area before that time. The last known species in
eastern North America, V. (G.) olga Mansfield of the late
Miocene may have continued its evolutionary history in a
migration across the Gulf to the eastern Pacific before the
elevation of the Panamanian isthmus in the Pleistocene,
although no record exists to document this. It is also pos-
sible that Caribbean species migrated via the coastal waters
of northern South America to Pacific waters to give rise to

the West Coast forms. A critical study of the late Tertiary
faunas of these regions is needed before a definite conclusion
is reached.

Several evolutionary trends are noted in the history of
glyptoactids in eastern and Gulf North America.

Inflation of the umbones is evident in the sequence
from the small V. (G.) nodifera waynensis Mansfield of the
Chickasawhay to V. (G.) gibberumbonata, n. sp., V. (G.)
serricosta (Heilprin), and V. (G.) himerta Dall of the
early Miocene, and continued in V’. (G.) hadra Dall, and V.
(G.) chelomodonta, n. sp., of the middle Miocene. The con-
tinuation in V. (G.) olga Mansfield of the late Miocene to
the Pleistocene and Recent forms of the eastern Pacific is
accompanied by a strong torsion of the beaks.

The trend toward simplification of the ancestral tri-
partite costae is evident in most glyptoactid species, and
only the lineage including V. (G.) nodifera - V. (G.) ser-
ricosta-V. (G.) hadra retained simplified paracostals and
narrow central cords. All other species eliminated the para-
costals in concert with expansion of the central cord.

Flattening of costae and elimination of nodes is charac-
teristic of several lineages, the most noteworthy being those
which resulted in V. (G.) impendeocosta, n. sp., V. (G.)
hesperide Gardner of the Miocene and V. (G.) crassicostata
G. B. Sowerby I and V. (S.) megastropha Gray of Pleisto-
cene and Recent ages.

The evolutionary modification of the dentition is an-
other important trend in the glyptoactids, and three results
are noted: 1) a general reduction in size and distinctiveness
of the right anterior cardinal (3a) which is totally dimin-
ished in some forms (e.g. V. (G.) hesperide); 2) the com-
plimentary strengthening of the left anterior cardinal from
the flat, bladelike condition of V. (G.) nodifera Kellum to
the prominent erect triangular element in ’. (G.) hadra Dall
and later forms, culminating in the Recent species of the
eastern Pacific; and 3) the enlargement and strengthening
of the “tooth” formed by the flexion of the left lunule found
in incipient state in V. (G.) nodifera but increasing in
prominence in most later species.

SYSTEMATICS

Phylum MOLLUSCA
Class BIVALVIA
Order HETERODONTA

Dominantly active equivalve isomyarian ceculamelli-
branch bivalves having porcellanous shell structure and
opisthodetic, parivincular, mainly external ligament. Hinge
armature comprises cardinal teeth and opposing sockets

Q2 PALAEONTOGRAPHICA AMERICANA (VI, 39)

radiating from position under beak, and lateral teeth and
sockets parallel to hinge line and variously developed. Inter-
gripalliate or sinuipalliate, posterior mantle edges formed
or fused into siphonal structures variously developed. Shell
ornamentation radial, concentric, cancellate or divaricate.

Superfamily CARDITACEA Fleming, 1820

Prosogyral, equivalve, intergripalliate heterodonts with
maximum of three cardinal teeth in right valve, two in left;
right anterior cardinal present, vestigial or absent. Lateral
teeth variously developed. Ornamentation basically radial.

Family CARDITIDAE Fleming, 1820

Byssate or nonbyssate carditaceans with or without gill
marsupium; some with shelly marsupium.

Genus VENERICARDIA Lamarck, 1801

Lamarck, J. B., Systtme des animaux sans vertebres ... , Ist ed.,

p. 123, Paris, 1801.

Type species —Venericardia imbricata (Gmelin), 1791.
Subsequent designation by F. C. Schmidt, 1818.

Type locality —Grignon, west of Paris, Départment de
Seine et Oise, France. Calcaire Grossier, Lutetian, middle
Eocene age.

Distribution of genus. — Fossil: Tethyan region of
Europe and Asia, Caribbean, North America and South
America, South Pacific. Recent: Indo-Pacific, coasts of
North America and South America.

Stratigraphic range-——Late Cretacous (Cenomanian)
to Recent.

Generic definition—Shell nonbyssiferous, closed, sub-
ovate to trigonal in lateral outline, posterior margin general-
ly truncate. Lunule small, deeply impressed, escutcheon
lacking or poorly developed.

Cardinal dentition of three teeth on the right valve, the
anterior one small and occasionally lacking; two teeth on
left valve. Lateral teeth small, poorly developed on pos-
terior; anterior lateral dentition consists of small protuber-
ance at juncture of left lunule and shell margin, correspond-
ing indentation on right valve. Nymphs slender and well de-
veloped. Ligament external, lodged in deep groove between
nymphs and dorsal shell margin.

Ornamentation of radial costae primitively tripartite,
nodiferous, but rounded, flattened or smooth in some.
Muscle scars and pallial line usually impressed, anterior
pedal retractor distinct on antero-ventral margin of hinge
plate; posterior pedal retractor is dorsally directed triangu-
Jar extension of posterior adductor, rarely becoming isolated.
Commissural margin interlocking and crenulated in concert
with costae.

Discussion and diagnosis—The constancy of dental
characters are sufficient to distinguish species of Venert-
cardia from those of other carditid subgenera (e.g. Cardita)
in which the cardinals are much more highly extended with
the right anterior cardinal (3a) absent, and a high develop-
ment of lateral teeth. Various subgenera described under the
genus maintain a constant dental pattern which differs only
in the attitude and shape of the dental elements, and some
(e.g. Glyptoactis) trend toward the loss of the right an-
terior cardinal (3a) in some of the above advanced Miocene
species,

Ornamentation in the group is highly variable, and the
loss of the ancestral paracostals and expansion of the central
costal cord of each rib is characteristic of advanced species
of Baluchicardia. Glyptoactis shows a consistent trend in
this direction in the Miocene. Other carditid genera are also
characterized by simplified costals, but their relationships
to Glyptoactis or other alticostate subgenera are not known.

Members of Pteromeris Conrad and Pleuromeris Con-
rad which are also found in North American Tertiary de-
posits have been traditionally associated with Venericardia
by authors (e.g. Dall, 1903). Because of the prominent
lateral teeth and marked reduction of cardinals, as well as
the absence of nymphs and ligamentary fossette in some
forms, these groups are probably not to be united with
Venericardia s.s. but will undoubtedly be raised to generic
rank with further study.

Cretaceous species described by Stephenson (1947,
1952) from the Late Cretaceous of the Gulf Coast have not
been studied during the present investigation, and their sub-
generic placement is unknown. One (V. alveana Stephenson )
is found in the Woodbine Formation of Cenomanian age
and pre-dates the baluchicards.

The relationship of Cyclocardia Conrad, 1867, first ap-
pearing in the Oligocene, is not understood, but the shell
form and thickness as well as dental pattern (see Text-fig.
7) of V. (C.) borealis Conrad indicates a convergence with
the planicostate subgenera Venericor and Leuroactis of the
early Tertiary which also have flattened ribs, However,
juvenile ornamentation which is noded in this species testi-
fies to evolution from alticostate rather than planicostate
forms (see Text-figs. 2,7).

Subgenus BALUCHICARDIA Rutsch and Schenck, 1943

“Venericardia beaumonti group,” Rutsch and Schenck, 1940, Geol.
Soc. Amer., Bull., vol. 51, p. 1976 (abstract).

Baluchicardia Rutsch and Schenck in Rutsch, 1944, Eclogae geol. Helv.,
vol. 36, p. 155, (Vokes, 1967); Stenzel and Krause in Stenzel,
Krause, and Twining, 1957. Univ. Texas, Pub. 5704, pp. 106, 107.

Type species —Venericardia (Baluchicardia) beaumonti

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP

(d’Archaic and Haime), 1854, Maestrichtian and Danian of
northwest India and Pakistan.

Type locality —‘Calcaire jaune de la chaine d’Hala,” in
the Sind region of northwest India.

Distribution. — Northwest India, Pakistan, southern
Iran, northern Africa, East Indies and West Indies, South
America and North America.

Stratigraphic range —From Maestrichtian to Paleocene
in most areas; Paleocene and early Eocene in Gulf Coastal
North America.

Subgeneric definition. — Venericards have highly in-
flated, somewhat elongate shells with marked posterior
truncation, ornamented by tripartite radiating costae com-
prising noded central cord and variously developed para-
costals on either side of the central cord. Basic ornamenta-
tion simplified in some with elimination of paracostals, re-
duction of nodes and widening of costae to restrict inter-
costals. Dentition containing well-developed right anterior
cardinal and strong, erect triangular left anterior cardinal.
Spiral growth pattern of curvature in four increasingly con-
vex segments from beak to ventral margin, but addition of
less convex terminal segment in advanced forms.

Discussion and diagnosis—The name Baluchicardia
remains until this time a nomen nudum, because it was
never formally characterized by a diagnosis, and it is prob-
ably better known in the literature as the “Cardita beau-
montt” or “Venericardia beawmonti group” (e.g. Rutsch,
1936). The latter name is used by Rutsch and Schenck
(1940), and not until 1944 is the name Baluchicardia intro-
duced by Rutsch who credited it to Rutsch and Schenck,
1941 (1940) based on the abstract of the paper presented
by those authors to the Geological Society of America in
1940. Whether or not the subgeneric name was given ver-
bally in the presentation of the paper is not known to the
present investigator, but in no case does it obtain validity
on this basis. Stenzel and Krause (1957) noted the sub-
generic name Baluchicardia as well as the type species but
give no diagnosis or description.

The basic morphologic pattern of the late Cretaceous
baluchicards of strong dental armature, vigorous tripartite
costae and highly inflated shell is continued into the Ceno-
zoic of the Gulf Coastal Plain of North America in species
of the early Paleocene such as V. (B.) hesperia Gardner
which appears to give rise to the late Paleocene species I’.
(B.) bulla Dall and V. (B.) wilcoxensis Dall. The relation-
ship of these forms is obvious, but in several species of the
early Paleocene [e.g. V. (B.) whitei Gardner] the ribs
became simplified and flattened, much like the planicostate
forms. Nothing would be gained by the creation of other

o
Ge

subgeneric taxa to include such species which seem to be
“experiments” in the early adaptive radiation of this group,
and they are united to Baluchicardia in the present investi-
gation. There is no indication that their evolutionary lineage
went beyond the early Paleocene. This increases the scope
of the subgenus beyond that originally intended by the
authors but seems more a biologic reality than a strict typo-
logical approach.

Two evolutionary lineages represent exceptions to the
above statement of latitude. The subgenus Rotundicardia,
n. subgen., which represents a continuation of V.-(R.) eoa
Gardner of the early Paleocene, and Claibornicardia Stenzel
and Krause which first appears in V. (C.) linguinodifera,
n. sp., of the early Eocene. Both are obviously derived from
baluchicard ancestry, but the “rotund” lateral outline and
lack of convexity in the former species documents its rela-
tion to V. (R.) rotunda Lea and other species of the Eocene.
The flattened left anterior cardinal and aligned medial right
cardinal of the latter species of early Eocene age denote its
ancestral relation to the claibornicards of the middle Eocene,
but represents a distinct transition from the rugged denti-
tion of the baluchicards.

Etymology—The subgeneric name Baluchicardia is
geographic in honor of the type region of V. (B.) beau-
monti, Baluchistan, in northwest India (modern Pakistan)
and the Latin cor, heart.

V. (Baluchicardia) hesperia Gardner, 1923

Plate 20, figures 2a, 2b

I’. alticostata (Conrad) Harris, 1896, Bull. Amer. Paleont., vol. 1
No. 4+, p. 57 (partim).

Tl’. alticostata hesperia Gardner, 1923, U.S. Geol. Sur., Prof. Paper
131-D, p. 112, pl. 32, figs. 1,2.
Non IT. alticostata (Conrad), 1833.

I’. hesperia Gardner, Trowbridge, 1932, U.S. Geol. Sur., Bull., No.
837, pl. 31, figs. 1,2; Plummer, 1933, Univ. Texas, Bull. 3232,
pp. 547, 550; 1935, Gardner, Univ. Texas, Bull. 3301, pp. 157,
164, 165; Stenzel, et al., 1957, Univ. Texas, Pub. 5704, p. 107;
Palmer and Brann, 1965, Bull. Amer. Paleont., vol. 48, No. 218,
pp. 330, 331.

Distinguishing characters. — Dentition unknown, but
probably much like that of V. (B.) bulla Dall. General form
similar to that species but distinguishable on basis of greater
elongation with straight ventral margin, greater anterior
margin projection. Much greater development of terrace-
like paracostals giving ribs wide base; ribs more bladelike in
other species. Sawtooth nodes on central costal cord much
more irregularly placed than in other species. Lower costal
number (21) than in other similar species.

84 PALAEONTOGRAPHICA AMERICANA (VI, 39)

Type.—Holotype an interlocked pair filled with matrix,
USNM 352268, dimensions below.

Sp'n 6 H WwW RN BL/L BH/H H/L %W/H

Holo. 44.6 38.5 18.5 21 fe a 86 48

Type locality—USGS station 3180, bluff on the Frio
River, % mile below Evans’ (Myrick’s) apiary, Uvalde
County, Texas. Tehuacana Member, Kincaid Formation.

Distribution —Other localities in Uvalde and Medina
Counties, Texas, are listed by Gardner (1923) and Gardner
and Bowles (1939). Only the holotype was examined during
the present investigation.

Stratigraphic occurrence—Yehuacana Member of the
Kincaid Formation, early Paleocene age.

Dimensional swummary.—See holotype dimensions.

Curvature characteristics.—

Sample: Holotype, Right valve. Nea

S1 Hi S2 HZ ss H3 S4 H+ $5

OR 23°81 Se S489 BSCR. ISIS GD shes

* Gerontism, highly convex segment.

Discussion and diagnosis —V. (B.) hesperia, with its in-
flated shell, convex A pattern of curvature, broadly based
ribs with well-developed paracostals resembles the ancestral
baluchicard V. (B.) beawmonti, as well as bearing a resem-
blance to other baluchicards in the North American Paleo-
cene, This was recognized by Stenzel, Krause and Twining
(1957), and this species undoubtedly resembles the gen-
eral ancestor from which the more highly specialized
baluchicards originated. Gardner and Bowles’ (1939) in-
clusion of this species in a systematic treatment of the
“Venericardia planicosta group” because it “. . . seems to be
related to the earlier members . . . ” is untenable, since fully
developed planicostate species predate it in the Paleocene.
There is, however, abundant evidence suggesting that many
“planicostate” forms have arisen independently during and
after the Paleocene (Heaslip, 1963).

The two well-known baluchicard species of the late
Paleocene, V. (B.) bulla Dall and V. (B.) wilcoxensis Dall
are undoubtedly derived from V. (B.) hesperia Gardner (see
Text-fig. 25). However, in the transition to the late Paleo-
cene forms, ornamentation underwent a marked simplifica-
tion with reduction of paracostals. This trend is also char-
acteristic of other lines of venericard evolution.

Etymology—tThe trivial name hesperia is from the
Latin hesperius, western, in reference to the location of the
type locality.

V. (Baluchicardia) francescae Gardner and Bowles, 1939
Plate 20, figures 3a-3c

V'. francescae Gardner and Bowles, 1939, U.S. Geol. Sur., Prof. Pap.
189-F, pp. 183,184, pl. 46, figs. 6,7; Palmer and Brann, 1965,
Bull. Amer. Paleont., vol. 48, No. 218, p. 329.

Distinguishing characters —Dentition poorly preserved
in holotype, but general pattern normal to Baluchicardia.
Convexity somewhat lower but general shape similar to V.
(B.) hesperia. Distinguishable from this species and other
baluchicards mainly on possession of ribs becoming highly
simplified during growth with vestigial tripartite cross-sec-
tion in mature portion and more pronounced toward the
umbones.

Type.—Holotype a right valve, filled with matrix and
poorly preserved, USNM 372925, dimensions below.

Sp'n L H 1%W RN BL/L BH/H H/L %W/H

Holo. 28.0 27.8 11.8 22 69 1 99 42

Type locality—USGS 6573, 1 mile southeast of New
Albany on St. Louis-San Francisco Railroad cut, Union
County, Mississippi.

Distribution—Known only from the type locality.

Stratigraphic occurrence—Clayton Formation, early
Paleocene age.

Dimensional summary.—See type dimensions.
Curvature characteristics. —

Sample: Holotype. N=
S1 Hi s2 2) Ss H3 S4+ H4 $5
OR 18° 1:2) eat 26. SSP 4.1 50° — _

Discussion and diagnosis—The inclusion of V. (B.)
francescae in a monograph of planicostate venericards by
Gardner and Bowles (1939) is understandable in the light
of the simplified costal structure exhibited by this species,
which represents a divergence from the normal baluchicard
character, and a radiation toward the planicostate condi-
tion which is also evidenced by V. (B.) whitei Gardner.

With the exception of this ornamental simplification
and the somewhat low convexity (terminal segment, S4 of
50°), the general shape, well-developed posterior truncation
and dental pattern are distinctly baluchicard and suggestive
of these characters in other species. There is no doubt that
this Clayton species is highly advanced but should be united
to Baluchicardia.

Etymology.—The trivial name francescae is a personal
patronym in honor of Miss Frances H. Walthall of Dr. Gard-
ner’s acquaintance.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 85

V. (Baluchicardia) whitei Gardner, 1923
Plate 20, figures 4a-4c

V’. alticostata Conrad, Harris, 1896, Bull. Amer. Paleont., vol. 1, No.
ADe Oar pl. SO, cklee Se

V’. (alticostata subsp?) whitei Gardner, 1923, U.S. Geol. Sur., Prof.
Pap. 131-D, p. 112, pl. 32, fig. 3.

Non V. alticostata (Conrad), 1833.

V’. whitei Gardner, Trowbridge, 1932, U.S. Geol. Sur., Bull., No. 837,
pl. 31, fig. 31; Gardner, 1935, Univ. Texas, Bull. 3301, p. 166;
Palmer and Brann, 1965, Bull. Amer. Paleont., vol. 48, No. 218,
p. 346.

Distinguishing characters—Dentition unknown. Shell
somewhat incomplete, but general shape comparable to that
of V. (B.) moa Gardner and other baluchicards. Curvature
of A pattern as in other allied species, but distinguished
from all other baluchicards on basis of highly simplified
costals with no trace of tripartite structure, coarsely and
irregularly decorated by subdued transverse nodes and
separated by relatively narrow U-shaped intercostals.

Type.—Holotype an incomplete right valve, interior
partly filled with matrix, poorly preserved. USNM 352269,
dimensions below (estimated ).

Sp’n L Hi YW RN BL/L BH/H H/L “’%W/H
Holo. 32.0 30.1 12.0 21 94 96 97 40

Type locality —USGS station 3180, bluff on the Frio
River, % mile below Evans’ (Myrick’s) apiary, Uvalde
County, Texas. Tehuacana Member, Kincaid Formation.

Distribution —Gardner (1923, 1933) reported the dis-
tribution of this species in Uvalde and Maverick Counties,
Texas, in three localities. Only the holotype was examined
in the present investigation.

Stratigraphic occurrence.—Yehuacana Member, Kin-
caid Formation, early Paleocene age.

Dimensional analysis —See holotype dimensions.

Curvature characteristics. —

Sample: Holotype N=
S1 H1 $2 H2) $3 H3 S4+ H+ $5
OR 19° 3.1 26° SOL 4 17.2 50° _ _—

Discussion and diagnosis —The unfortunate lack of pre-
servation of critical characters such as dentition in the holo-
type makes the placement of this species with Baluchicardia
open to question, especially in the light of the extreme
simplification of ornamentation which can be compared only
to that of the Miocene glyptoactid V. (G.) hesperide Gard-
ner. The shape, as well, is reminiscent of this species, but
no direct ancestry is inferred. These two forms converge
closely into the same shell form and ornamentation, strongly
suggesting exploitation of the same ecological role by both.

The convex A pattern of curvature and its stratigraphic

occurrence suggest that V. (B.) whitei is a highly specialized
offshoot of the more primitive baluchicards, and it is not
deemed advisable to recognize this distinct form by creating
a separate subgeneric group.

Etymology.—The trivial name whitei is a personal
patronym in honor of Dr. Charles A. White, “one of the
foremost of the earlier paleontologists and one of the first
to collect from the Texas Tertiary formations.” (Gardner,

1923).

V. (Baluchicardia) amplicrenata, n. sp.
Plate 20, figures 6a, 6b
?V’. hesperia, Gardner, Gardner, 1935, Univ. Texas, Bull. 3301, p.

165, pl. 15, fig. 6.

Distinguishing characters—Dentition unknown. Gen-
eral shell shape (see Text-fig. 25) inferred from early growth
lines somewhat elongated. Similar in coarsely noded tripar-
tite costae to V. (B.) moa Gardner, but without the pos-
tero-ventral extension of that species. Distinguished from
all baluchicard species by low costal number, extremely thin
central costal cord swelling at each node and paracostals
well developed forming terraces on either side of central cord
and separated by wide intercostals U-shaped in cross-section.

Type.—Holotype a well-preserved left valve imbedded
in matrix, USNM 370912 (not 370922, typographical error

in Gardner, 1935). Estimated dimensions below.

Sp'n jt H 144W RN BL/L BH/H H/L %W/H

Holo. 31.0 27.3 ES 18 -65 eel 88 42
Type locality—U.S.G.S. station 2439, 1 mile east of
Webberville, Travis County, Texas, Tehuacana Member,
Kincaid Formation, early Paleocene age.
Distribution —Known only from type locality.

Stratigraphic occurrence.—Tehuacana Member, Kin-
caid Formation, early Paleocene age.

Dimensional summary.—See type dimensions.

Curvature charactertstics.—

Sample: Holotype Na

S1 Hi S2 H2 83 H3 S4 H+ $5

OR D5 ew eae 70 | AL OP 6:44. KOON Silla wezihoe tose

Discussion and diagnosis—The curvature pattern, as
indicated from values in the table is highly erratic, and this
represents a simplification of the original data. In the stage
$2, there are three small stages of 54°-70°-60°. The mean-
ing of this is not clear and no similar pattern is recorded
for any other species.

V. (B.) amplicrenata is probably descended from a
more generalized baluchicard such as V. (B.) hesperia but
becomes distinct by reduction of costals and amplification

86 PALAEONTOGRAPHICA AMERICANA (VI, 39)

of the sawtooth nodes characteristic of the group. Gardner’s
(1935) questionable inclusion of this species in V. (B.)
hesperia is not realistic, because that species has a higher
rib count, much finer, more regularly spaced costal nodes
and much more highly inflated beaks.

Because of the similar development of ornamentation
and similar rib number, IV’. (G.) moa is probably closely re-
lated to this species, differing mainly in its great postero-
ventral extension.

Etymology.—The trivial amplicrenata is derived from
the Latin amplus, large, and crenatus, notched, in reference
to the large costal nodes.

V. (Baluchicardia) moa Gardner, 1935
Plate 20, figures 5a, 5b

V’. moa Gardner, 1935, Univ. Texas, Bull. 3301, pp. 157, 168, 169, 170,

pl. 10, fig. 1.

(Glyptoactis?) moa Gardner, Gardner and Bowles, 1939, U.S.

Geol. Sur., Prof. Pap. 189-F, pp. 195, 196, pl. 46, fig. 1; Palmer

and Brann, 1965, Bull. Amer. Paleont., vol. 48, No. 218, pp. 334,

335.

Distinguishing characters—Ornamentation similar to
that of V. (B.) amplicrenata, n. sp., in rib number, well-
developed paracostals and coarsely noded central costal cord
but distinguishable from that and other baluchicard species
on the basis of high degree of postero-ventral elongation
much more pronounced than that of other species.

Type.—Holotype a poorly preserved pair of locked
valves, USNM 370922, dimensions below.

H WW RN BL/L BH/H H/L “4W/H
Holo. 33.8 31.4 11.0 18 .80 -80 Ah) 35

Type locality —USGS station 4398, Bibora Creek, just
below Bibora tank, about 18 miles south and east of Eagle
Pass, Maverick County, Texas. Tehuacana Member, Kin-
caid Formation, early Paleocene age.

Distribution —Gardner (1935) noted five occurrences,
including the type locality in Maverick County, Texas, and
recognized a poorly preserved form allied to this species
from four additional localities in a stratigraphically lower
position. Only the holotype was examined during the present
investigation.

Sp'n L

Stratigraphic occurrence-—YVehuacana Member of the
Kincaid Formation, early Paleocene age. The questionably
allied forms are found in the Littig Member of the same
formation.

Dimensional summary.—See type dimensions.

Curvature characteristics.—

Sample: Holotype Nis

S1 Hi) SZ H2 83 H3 S+ H+ $5

OR Ome 1.6 Ac S25 ae 530 93 59° — —

Discussion and diagnosis. — V. (B.) moa and other
species of the Kincaid, or lowermost unit in the Paleocene
record of the western Gulf Coastal Plain, are incompletely
known because of the poor quality of preservation. This is
especially unfortunate, since the primary radiation of the
ancestral baluchicards as well as other important Cenozoic
mollusks occurred during this age.

Only the poorly preserved holotype was examined in
the present investigation, but fortunately the costae near
the anterior ventral margin preserve the original tripartite
structure and coarsely noded central cord serve to identify
this species as a baluchicard, and similar to V. (B.) ampli-
crenata, n. sp., of the same stratigraphic unit.

The convex A pattern of curvature is similar to that of
other baluchicards, but this species has elongated in the
postero-ventral direction forming a “corner” at the meeting
of the posterior and ventral margins reminiscent of that
formed in many claibornicards of the Eocene.

Etymology.—The derivation of the trivial name moa
is uncertain.

V. (Baluchicardia) bulla Dall, 1903
Plate 20, figures 7, 8; Plate 21, figures la-ld

V’. bulla Dall, 1903, Wagner Free Inst. Sci., Trans., vol. 3, pt. 6, pp-
1424, 1425, pl. 56, figs. 13, 14; Plummer, 1933, Univ. Texas, Bull.
3232, pp. 811, 812, pl. 8, figs. 2a,2b; Gardner, 1935, Univ. Texas,
Bull. 3301, pp. 163, 164, pl. 13, figs. 1-4, pl. 14, figs. 4-6.

I’. (Baluchicardia) bulla Dall, Stenzel, et al., 1957, Univ. Texas, Pub.
5704, p. 107; Palmer and Brann, 1965, Bull. Amer. Paleont., vol.
48, No. 218, p. 325.

Distinguishing characters.—Similar in size, degree of in-
flation, dental pattern, and ornamentation V. (B.) wilcox-
ensis Dall but readily distinguishable from that species on
basis of inordinate shell thickness in V. (B.) bulla, in which
ratio of shell thickness to height averages .12. That of the
other species is only .06. V. (B.) bulla has much more highly
inflated umbones and more coarsely and irregularly noded
costal cords than the other species.

Type.—Lectotype, a left valve, imbedded in matrix,
poorly preserved, USNM 164556 (fig. 13), dimensions be-
low. Paralectotype 645691 (fig. 14), interlocked valves,
same collection number. Hypotypes figured by Gardner
(1935), complete individual, disarticulated valves, USNM
370913, dimensions below.

Sp’n ib H %W RN BL/L BH/H H/L “%W/H
Tech) nasties e240 0.4 “Saerinee ? 97 45
Para, ) SR0n cs} 72 sree 2 OTe mas?
Sp'n ib H %W RN BL/L BH/H H/L %W/H
Hypo. 33.8 33.6 16.2 30 73 86 99 48

Type locality —Exact location indefinite. Dall (1903)

AMERICAN ALTICOSTATE

described it as “Brown sandstone of the Midway horizon,
east of the first small creek on the road to Bastrop, Texas,
from Old Garfield in the Austin Quadrangle . . .” Gardner
(1935) noted that “. . . several unsuccessful attempts were
made to find this locality. The closest approach to it is repre-
sented in a collection made in 1932 by L. W. Stephenson
and O. E. Cook from 2.5 miles southeast of the Travis-
Bastrop County Line.” The type locality is listed as USGS
station 3309. Gardner’s hypotypes are from USGS station
5281, on the Colorado River, 5 miles below Webberville.
Wills Point Formation, late Paleocene age.

Distribution. — Gardner (1935) listed a number of
localities in Bastrop County. In addition to the type and
hypotype collections, nine individuals from AMNH collec-
tion 27677 labelled, “Wills Point formation, Bastrop County,
Texas,” were examined and measured.

Stratigraphic occurrence.—Wills Point Formation, late
Paleocene age.

Dimensional summary.—

Sample: AMNH 27677

Ni -9 Dimension

Statistic L H 14W RN
OR 28.8-31.9 27.8-34.8 13.5-16.4 26-30
x 31.2 30.1 15.7 27.4
S 5.9 2.4 5 1.5
Vv 18.9 7.8 9.4 5.5
Ox 2.0 8 & 5

Ratio
BL/L BH/H H/L Y4W/H
OR -68-.77 .83-.87 .94-1.05 46-.55
x 73 85 98 51
Curvature characteristics. —
Sample: AMNH 27677, part. Ne
S1 H1 $2 H2 $3 H3 $+ H4* S$5*

OR 30-35° 2.6-3.9 41-46° 4.8-5.6 51-56° 13.0-21.4 59-65° 22.4-26.5 63-70°
xX Bathe mses 4335 ee 58107 16.9 62.3° 24.9 67.7°
*Gerontic segment.

Discussion and diagnosis—With the exception of the
narrow, bladelike development of the central costal cord,
V. (B.) bulla is similar in shape and degree of inflation to
other baluchicards of the early Paleocene in Texas, and its
appearance corresponds to the beginning of Wills Point
deposition in that area.

V. (B.) wilcoxensis of the Matthews Landing Member
of equivalent age in Alabama resembles in many respects
the Texas species, and the two undoubtedly evolved from
common ancestry, and most likely from V. (B.) hesperia.
The two similar species of the late Paleocene, separated by
600 miles across the Mississippi embayment, undoubtedly

VENERICARDS: HEASLIP 87

represent ecologic equivalents and might conceivably be
grouped in a superspecies in the sense of Mayr, et al. (1953).

The inordinate shell thickness, reaching 6.3 millimeters
for mature individuals of 33 millimeters in height has no
parallel among other species of the present investigation. As
a result of this abnormal growth, the interior space becomes
almost parallel-sided, and the volume within the occluded
valves is severely restricted. There is no evidence to suggest
that this effect may have been pathologic or caused by
subpallial parasites restricted to the species, and the charac-
ter of shell thickness seems to be intrinsic to V. (B.) bulla.

No species of the Eocene appears to have been descend-
ed from V. (B.) bulla and the lineage of Baluchicardia. The
single early Eocene member of this subgenus probably
descended from V. (B.) wilcoxensis and V. (B.) greggiana
and restricted to a small area in Alabama, is the last balu-
chicard in Gulf or east coastal North America.

Dall’s type (USNM 164556) is poorly preserved, re-
vealing nothing of dentition or interior, and with highly
eroded ornamentation. Only in the small paralectotype (PI.
20, fig. 7) is an indication of tripartite costal structure visi-
ble. These, however, are comparable in general features of
shape and inflation to the better preserved specimens from
the Colorado River localities.

Etymology.—The trivial name bulla is taken directly
from the Latin bulla, bubble, in recognition of the high de-
gree of inflation in the species.

V. (Baluchicardia) wilcoxensis Dall, 1903
Plate 21, figures 2a-2c, 3a-3c

V’. alticostata Conrad var., Harris, 1896, Bull. Amer. Paleont., vol. 1,
No. 4, p. 171, pl. 4, fig. 12.

Non IV. alticostata (Conrad), 1833.

V’. wilcoxensis Dall, 1903, Wagner Free Inst. Sci., Trans., vol. 3, No.
6, p. 1426, pl. 54, fig. 12; Cooke, 1926, Geol. Sur. Alabama, Spec.
Rept. 14, pl. 93, figs. 4a, +b; Plummer, 1933, Univ. Texas, Bull.
3232, p. 811, pl. 8, figs. 3a, 3b; Gardner, 1935, Uniy. Texas, Bull.
3301, pp. 163, 164, pl. 14, figs. 1-3; Palmer and Brann, 1965,
Bull. Amer. Paleont., vol. 48, No. 218, p. 346.

Distinguishing characters —Similar in size, shape and
general features of ornamentation and dentition to V. (B.)
bulla, but distinguishable on the basis of lower inflation
(average 144 W/H of .43) than that species (average .51),
normal shell thickness, and smaller and more regularly
spaced sawtooth costal nodes. Paracostal development is
highly variable in V. (C.) wilcoxensis, and both tripartite
and nontripartite ornamentation is represented in popula-
tions; tripartite ribbing is more constant in V. (B.) bulla.
Many specimens of V. (B.) wilcoxensis have a small, pointed
fold on commissural surface or left lunule just posterior to
the apex of the anterior cardinal (2) forming a projecting

88 PALAEONTOGRAPHICA AMERICANA (VI, 39)

tit which fits into a corresponding socket in the right lunule.
This is absent in V. (B.) bulla.

Type.—Holotype, a left valve, well preserved, USNM
164542. Lectotype (Palmer and Brann, 1965, p. 346 which
see) of “var. tripla” Dall which was not figured, a right
valve with broken posterior, USNM 129897. Dimensions

below.

Sp’n L H YW RN BL/L BH/H H/L %W/H
Holo. 36.7 37.6 15.6 26 70 82 1.03 42
Para. ats esd 12.6 26 -67 85 1.02 40

* Estimated.

Type locality.—USGS station 3116, Matthew’s Landing
on the Alabama River in the northern part of section 12,
T 12 N, RG E, about 9 miles west of Camden and 214 miles
above Clifton Ferry, Wilcox County, Alabama. Dall’s orig-
inal description was “Upper clayey layers of the Midway
stage of the Eocene in Wilcox County, Alabama, near Grave-
yard Hill.” The locality of V. variation “tripla” is USGS
287, Dale’s Branch near Allentown, Wilcox County, Ala-
bama.

Distribution—Material from the following localities
was examined during the present investigation: Type and
paratype localities, including an AMNH collection from
locality 987, corresponding to the type locality; USGS 264
and 284, Prairie Creek, Wilcox County, Alabama, USGS 283,
Black Creek Branch of Prairie Creek, Wilcox County, Ala-
bama.

Stratigraphic occurrence——Matthews Landing Member,
Naheola Formation, late Paleocene age.

Dimensional summary.—

Sample: USNM 129746

N = 6 Dimension
Statistic IL, ZW RN
OR 23.3-36.6 23.4-34.5 10.1-15-6 26-29
xX 30.0 29.6 12.8 Dias
S 5.1 4.6 2.5 ea
V 16.9 15.4 19.9 3.8
Ox hat 1.9 1.0 4
Ratio
BL/L BH/H H/L YW/H
OR .70-.76 80-.86 -94-1.01 39-46
x vie .84 mh) 43
Curvature characteristics.—
Sample: USNM 129746 Ni 5

S1 H1 $2 H2 $3 H3 S¢ H4* S5*

OR 20-39° 1.2-4.2 35-45° 3.4-8.5 45-58° 5.3-17.5 56-63° 22.4 75°
x RA BA MESDOP ASS 52.057 IONS i Gtae en ae

*Gerontic Segment.

Discussion and diagnosis. —The variability of paracostal

development in this species led Dall (1903) to distinguish
between the members of V. (B.) wilcoxensis with only vesti-
gial paracostals (e.g. the holotype, Pl. 21, figs. 2a-2c) and
those possessing fully developed tripartite ribs like those
of V. (B.) bulla. The latter condition he gave the varietal
name “tripla” without figuring or otherwise designating a
type specimen. Palmer and Brann, 1965, page 346 desig-
nated the broken right valve (USNM 129897) in Dall’s
collection as the lectotype for the “variety,” and this course
is adopted in the present report (see Pl. 21, figs. 3a-3c).
This variability in ornamentation, however, is a characteris-
tic of the species, and it is highly likely it represents a paral-
lel to the sexual dimorphism found in V. (Rotundicardia)
rotunda and its Jackson descendants, as well as the claibor-
nicard V. (C.) alticostata. Variety “tripla” is likely the male
of the species ( Heaslip, 1964).

V.(B.) greggiana Dall of the early Eocene Tuscahoma
Sand in Monroe County, Alabama, is similar to V. (B.)
wilcoxensis in shape, dentition, and degree of inflation, and
is the logical descendant of the Midway species, differing
mainly in size and ornamentation.

The relationship of V. (B.) wilcoxensis to any other
species in the lower Eocene rocks of the Gulf Coast is not
clear, but it may be assumed that this species gave rise to
the Eocene claibornicards with certain modifications in the
dental pattern (reduction and flattening of left anterior
cardinal (2) and straightening of right anterior cardinal
(3a) in line with the medial cardinal (3b) ). Certain in-
dividuals in V. (B.) wilcoxensis and V. (B.) greggiana ex-
hibit a tendency toward partial flattening of tooth 2 and
reduction of the ridgelike 3a which borders the anterior
socket on the right valve.

Etymology.—tThe trivial name wilcoxensis is geograph-
ic in honor of the type region, Wilcox County, Alabama.
Unfortunately, the term Wilcoxian refers to rocks of the
lower Eocene stage on the Gulf and eastern coasts of North
America. V. (B.) wilcoxensis is late Paleocene and not Wil-
coxian in age.

V. (Baluchicardia) greggiana Dall, 1903
Plate 21, figures 4a-4c, 5

2V. decusata Tuomey, 1858, 2d Bien. Rept., Alabama Geol. Sur., p. 271.

Non V. decusata Lamarck, 1807.

V. alticostata Conrad var., Harris, 1897, Bull. Amer. Paleont., vol. 2,
No. 9, p. 55, pl. 11, fig. 1.

Non I. alticostata (Conrad), 1833.

V’. greggiana Dall, 1903, Wagner Free Inst. Sci., Trans., vol. 3, No.
6, p. 1425; Gardner and Bowles, 1939, (under Venericor?), U.S.
Geol. Sur., Prof. Pap. 189-F, p. 194, pl. 46, fig. 9; Harris and
Palmer, 1946, Bull. Amer. Paleont., vol. 30, No. 117, p. 68;
Palmer and Brann, 1965, Bull. Amer. Paleont., vol. 48, No. 218,
pp. 329, 330.

Distinguishing characters—Similar to V. (B.) wilcoxen-

AMERICAN ALTICOSTATE VENERICARDS: Herasuip 89

sts in dental pattern and general features of shape and con-
vexity but distinguishable from that species on the basis
of more numerous ribs with fairly regularly spaced flattened
subdued sawtooth nodes pointing in beak direction and ex-
panded. Paracostals developed to the degree that they al-
most cover the U-shaped intercostals. Posterior truncation
almost vertical to main axis of shell; in V. (B.) wilcoxensis
it descends obliquely forward. Beaks much more highly in-
flated and tightly inrolled, obscuring much of the lunule;
beaks smaller and lunule open in the other species.

Type.—Lectotype (Palmer and Brann, 1965, p. 330),
Harris’ figured specimen, PRI 137, a large right valve, per-
fectly preserved, dimensions below.

Sp'n L H ZW RN BL/L BH/H H/L %W/H
Holo. 52.3 56.0 23.6 40 73 85 1.07 42

Type locality —Greggs Landing on the west bank of
the Alabama River, Monroe County, Alabama. Greggs
Landing Member, Tuscahoma Formation.

Distribution.—Collections from the following localities
were examined in the present investigation: Type locality,
including collections from USGS station 5604 and AMNH
locality 1040, USGS 3098, Bells Landing on east bank of
Alabama River, Monroe County, Alabama.

Stratigraphic occurrence. — Greggs Landing Member
and Bells Landing Member, Tuscahoma Formation, early
Eocene age.

Dimensional summary.—Sample includes AMNH_ 1040
(1 specimen), USGS 3098 (2 specimens), USGS 5604 (1
specimen) and holotype, PRI 137.

Sample: Composite, Greggs Landing, Bells Landing

in} 5 Dimension
Statistic L H VW RN
OR 48.2-52.3 50.6-56.0 19.3-23.6 34-40
50.9 52.4 21.7 35.8
S 1.7 2.1 1.6 2.5
Vv Si8) 4.1 74 7.0
Ox o 1.0 all 11
Ratio
BL/L BH/H H/L 16W/H
OR -71-.77 -83-.86 1.00-1.07 40-43
2.4 -73 84 1.63 Al

Curvature characteristics*—

Sample: Composite, Greggs Landing, Bells Landing N=4
S1 H1 $2 H2 $3 H3 S+ H+ $5
OR = 25-40° 2.2-3.137-55° 5.2-9.8 50-60° 8.5-12.3 60-69° 14.5-22.8 53-56°

x B25 12:5) 435508 6:8 34.3° 10.7 643° 20:4 55.0°

* Curvature of B type, similar to Claibornicardia.

Discussion and diagnosis —Dall (1903) neglected to
figure or formally designate a holotype, but he referred to

Harris’ (1897, pl. 11, fig. 1) figured specimen. Palmer and
Brann (1965, p. 330) designated this specimen as a lecto-
type thus establishing a definiteness as to the type and
type locality. The possibility exists that Harris’ specimen
may be the holotype, because it is the only definite specimen
to which Dall referred. Dall probably included specimens
from. Bells Landing (USGS 3098), as well as those from
Greggs Landing (USGS 3118), in his original description,
but the “type” specimen, PRI 137 is from the latter locality,
and, therefore, this is the type locality.

The association of this species with the planicostate
subgenus Venericor by Gardner and Bowles (1939) is un-
realistic. Neither the dental pattern, the high degree of in-
flation, nor the distinctly tripartite, coarsely noded orna-
mentation of this species are characteristic of that sub-
genus, and only establish its placement in Baluchicardia.

The descent of this species from V. (B.) wilcoxensis of
the Naheola is reasonable not only on the basis of similarity
in shape and dental pattern but also on the presence in some
individuals of the small tit developed in the right lunule of
that species, and these are the only two species exhibiting
this feature, reminiscent of the flexion in the left lunule seen
in the later glyptoactids (see Teese, U7 Ab 27 fig. 3c).

The high number of ribs (up to 40) in this species is
unique among all forms studied in the present investigation,
although the accompanying emphasis of paracostal develop-
ment as seen in two rotundicards, V. (R.) crenaea Gardner
and V. (R.) eutawcolens Harris.

V. (B.) greggiana appears to be the last of the baluchi-
cards in eastern North America, and whether or not this
species gave rise to subsequent forms in the Eocene is not
established. The alticostate lineage is continued in V. (Clai-
bormcardia) linguinodifera n. sp., of the Bashi Member of
Wilcoxian age, but this tiny species is distinct from V. (B.)
greggiana in that it is likely to have arisen independently
from the Paleocene baluchicards and perhaps from V. (B.)
wilcoxe nN SIS.

The curvature pattern of the B type seen in V. (B.)
greggiana is not typical of the baluchicards and represents
a development similar to that in Claibornicardia.

Etymology.—The trivial name greggtana is geographic
in honor of the type locality at Greggs Landing.

Subgenus ROTUNDICARDIA, n. subgen.

Type species—V. (Rotundicardia) rotunda Vea, first
described in 1833 from Claiborne Bluff, Monroe County,
Alabama. Gosport Sand, late Middle Eocene.

Type locahty.—Claiborne Bluff, near top of east bank
of Alabama River, south of bridge crossing of U.S. Highway
84 and east of the town of Claiborne, T 7 N, R 5 E, Mon-

90 PALAEONTOGRAPHICA AMERICANA (YI, 39)

roe County, Alabama. Gosport Formation, late Middle
Eocene.

Distribution—East coast of North America to New
Jersey, Gulf Coast of North America to Texas and Mexico.
Other distribution unknown.

Stratigraphic range—From early Paleocene to early
Oligocene.

Subgeneric definition —Alticostate venericards having
rounded lateral outline with minimum of posterior trunca-
tion of the shell and shell of low convexity. Dental pattern
similar to that of Baluchicardia, with prominent, massive
left anterior cardinal (2) triangular and erect in shape,
massive and prominent medial right cardinal (3b) and ex-
tremely well-developed right anterior cardinal (3a) forming
a straight ridge almost erect and bordering the anterior
socket in that valve. General tendency toward reduction of
paracostal cords in main evolutionary lineage with develop-
ment of bladelike costals but well-developed paracostals in
other lineage.

Discussion and diagnosis.—Both the subgenera Baluchi-
cardia and the descendant Claibornicardia are characterized
by a high degree of posterior elongation and a marked
posterior truncation which may have been associated with a
burrowing habit and placement of the siphonal areas with
relation to the substrate surface in these groups. In contrast,
the shell outline in members of Rotundicardia is subrqunded,
or, as Lea described it, “rotund,” with height-length (H/L)
ratios from .92 to 1.10. The beak tends to be more centrally
located (BL/L from .61 to .74) and the shell more “equilat-
eral” in the terminology of previous authors, than in the
other two subgenera. Much overlap is noted in this charac-
ter, however. The beaks are small and umbones not promi-
nent, except in certain members of V. (R.) diversidentata
Meyer (BH/H from .84 to .95). The rounded outline and
relative absence of posterior truncation may reflect the ab-
sence of burrowing in Rotundicardia, although this is not
thought to be the case because of the high development of
bladelike ornamentation which may have anchored the ani-
mals in the substrate.

The development of sexual dimorphism documented by
distinct ornamentation and curvature pattern is noted in
V.(R.) rotunda and in the descendant V. (R.) diversiden-
tata.

Harris (1919) fully appreciated the close relationship
among V. (R.) rotunda, V. (R.) diversidentata and V.
(R.) carsonensis as a continuous evolutionary sequence, as
well as the common ancestry of what he termed the “rotun-
da-alticostata stock” (see ibid., pp. 84, 85). However, not
realizing the importance of dental pattern as a distinction

between the two lineages, he inadvertently brought mem-
bers of Claibornicardia into Rotundicardia under such
names as “V. rotunda varying toward trapaquara” and “V.
(rotunda?) var. coloradonis.” Most of these are conspecific
with V. (C.) coloradonis Harris of the present report.

Etymology.—R otundicardia is from the Latin rotundus,
round, and cor, heart, in reference to the rounded shell out-
line.

V. (Rotundicardia) eoa Gardner, 1935
Plate 22, figures la-lc, 2

V. eoa Gardner MS, Plummer, 1933, Univ. Texas, Bull. 3232, p. 547.
Non, p. 811, pl. 8, fig. 1.

lV’. (Glyptoactis) eoa Gardner, 1935, Univ. Texas, Bull. 3301, pp. 170,
171, pl. 15, figs. 3-5; Palmer and Brann, 1965, Bull. Amer. Pale-
ont., vol. 48, No. 218, p. 329.

Distinguishing characters—Similar to V. (R.) rotunda
Lea in outline, convexity and dentition, but distinguishable
from that and other rotundicard species in having higher
degree of posterior truncation, directed anteriorly downward,
relatively low rib number (23) and narrow central costal
cords ornamented by sawtooth nodes pointing toward the
ventral margin and not much wider than cord. Paracostals
distinct, forming low, wide terraces on either side of central
cord; intercostals shallow, broad U-shaped channels. Ribs
of V. (R.) rotunda high and bladelike, lacking distinct para-
costals.

Type.—Holotype, a well-preserved right valve with
slightly damaged anterior margin, USNM 370918, dimen-
sions below. Paratype a larger left valve lacking posterior
and ventral portion of shell, same collection number.

Sp’n 1b, H %W RN BL/L BH/H H/L Y%W/H

Holo. 22.5 21.0 7.8 23 61 91 93) 37

Para.* 28.4 26.9 10.6 23 .74 91 95 39
*Estimated

Type locality—USGS station 11914, south bank of
Colorado River 1% miles below the Travis-Bastrop County
line, Bastrop County, Texas.

Distribution—Two other localities in the vicinity of
the type locality are listed by Gardner (1935), but material
from these has not been examined in the present investiga-
tion.

Stratigraphic occurrence. — Upper part of the Kincaid
Formation, early Paleocene age.

Dimensional summary. — See type dimensions.

Curvature characteristics: —

Sample: Holotype
S1 H1 $2 H2 $3 H3 S4 H4 $5
OR 22° oii 40° 1.7 43° 74K] 47° _ —_

Discussion and diagnosis. —V.(R.) eoa is the earliest

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 91

representative of Rotwndicardia known from the fossil rec-
ord, but its evolution from Baluchicardia is evident in primi-
tive characters such as distinctly tripartite ornamentation
with sawtooth nodes unlike the ornate ribs of the Eocene
descendants. The A pattern of curvature and robust dental
pattern also ally it to this group.

Its rotund shape with small, low beaks, however,
indicates a direct ancestry to rotundicards of the Eocene,
and the primitive dental pattern is transmitted almost un-
modified to the more advanced forms.

It is interesting to note that no rotundicards are known
from the sediments of late Paleocene or early Eocene age.
However, the alticostate groups living in that time span
are in general not well represented, and depositional en-
vironments of known deposits may not have been favorable
for the life activities of the group. Paradoxically, the early
Eocene sediments contain the greatest abundance of the
planicostate species (see Text-fig. 3) in an evolutionary
burst in that group.

Gardner (1935) recognized the basic similarity be-
tween V. (R.) eoa and V. (R.) rotunda, and it appears cer-
tain that the Paleocene species is the ancestor of the Eocene
species, even though intermediate forms are unknown.

Etymology. — The trivial name eoa is from the Latin
eos, dawn, alluding to the early appearance of this form
in the early Paleocene.

V. (Rotundicardia) crenaea Gardner, 1935
Plate 22, figures 3a, 3b, 4

V. (Glyptoactis) crenaea Gardner, 1935, Univ. Texas, Bull. 3301, pp.
167, 168, pl. 15, figs. 1, 2; Palmer and Brann, 1965, Bull. Amer.
Paleont., vol. +8, No. 218, p. 327.

Distinguishing characters. — Dentition unknown. Flat-
ness and rounded shell outline with relative absence of pos-
terior truncation, small low beaks probably ally V. (R.)
crenaea to the rotundicards, but ornamentation is unlike any
other species in the group, and generally similar only to that
of V. (R.) eutawcolens Harris. Most conspicuous paracostals
of any species in the present investigation only slightly lower
than the costal cord which is ornamented by fairly regularly
spaced sawtooth nodes somewhat subdued to transverse
ridges in certain specimens (e.g. the paratype). Paracostals
noded in concert with central cord and expand over inter-
costal space until almost in contact. Intercostals constricted
and slitlike.

Type.— Lectotype (herein designated), Gardner’s
figured right valve, USNM 370911, imbedded in matrix.
Paralectotype, figured left “cotype,” same collection num-
ber, Dimensions below.

Sp’n 18 H WRN) VBE/L) BHI/E “H/L 342Wi/
ects 25.3 22.7 8.0 28 79 95 90 35
Para. 28.6 22.0 8.5 27 -80 96 “Ef 39

Type locality. —USGS station 11770, bed of the Hondo
River, 1 mile northwest of New Fountain, Medina County,
Texas. Tehuacana Member of the Kincaid Formation, early
Paleocene age.

Distribution. — Gardner (1935) listed several localities
from Medina, Uvalde, Kaufman and Guadalupe Counties in
Texas. Only the lectotype and paratype were studied in the
present investigation.

Stratigraphic occurrence. —Tehuacana Member of the
Kincaid Formation, early Paleocene age. Some questionable
localities for this species listed by Gardner (op. cit.) are in
the Littig Member of the same formation which is strati-
graphically lower than the Tehuacana.

Dimensional summary. —

Sample: Holotype and Paratype

IN) = 2 Dimension
Statistic 1B, H UW RN
OR 25.3-28.6 22.0-22.7 8.0-8.5 27-28
4 27.0 22.4 8.3 27.5
S 2.3 “5 4.3 7
cae 8.7 2.2 +4 2.6
Ox LF 4 3 5
Ratio
BL/L BH/H H/L Y4W/H
OR .79-.80 .95-.96 -77-.90 -35-.39
X 80 96 .84 EA7/
Curvature charactertstics. —
Sample: Holotype and paratype N=2
S1 H1 $2 H2 $3 H3* $4* H+ S5
OR 20-21° 1.2-1.9 40° 1.8-2.3 46-47° ? ? — —
xX 20.5° 1.5 40° 2.1 46.5°

*Terminal segment missing.

Discussion and diagnosis —Except for Gardner’s (1935)
statement, “Medial right cardinal obliquely deltoid and very
heavy,” the dentition in the species is unknown, but a
heavily constructed dentition is characteristic of Rotundi-
cardia.

The almost unique expansion of the paracostals is found
in only two other species of the present investigation: J.
(Baluchicardia) greggiana of the Tuscahoma Formation and
V. (R.) eutawcolens of the Santee Limestone. The para-
costals of the former species, however, are expanded from
the sides of basically bladelike ribs, giving a cross-shape to
the rib cross-section. Those of the latter species form solid
terraces, much as in V. (R.) crenaea, and could suggest a
direct relationship between these species.

92 PALAEONTOGRAPHICA AMERICANA (YI, 39)

Another feature atypical of Rotwndicardia possessed
by this species is the convex pattern of curvature (A) which
lacks the final, most convex stage. The meaning of this is not
clear, because the specimens examined are assumed to be
adults. This may represent an instance of paedomorphosis,
in which sexual maturity is attained at an early ontogenetic
stage with the corresponding suppression of the normally
adult character. This is not found in any other species in the
scope of this report.

Because of its rounded shell outline and relative lack of
posterior truncation, as well as small beaks and umbones and
low convexity, V. (R.) crenaea is questionably placed as an
offshoot on the phylogeny of Rotwndicardia (see Text-fig.
26), despite its distinct ornamentation and incomplete curv-
ature pattern, Its descendants, other than possibly V. (R.)
eutawcolens, are not recognized.

Etymology. — The trivial name crenaea is derived from
the Latin crenae, notches or rounded projections, alluding to
the nodes on the central and paracostal cords.

V. (Rotundicardia) flabellum Harris, 1919
Plate 22, figures 6a-6c, 7

V. rotunda var. flabellum Harris, 1919, Bull. Amer. Paleont., vol. 6,
No. 31, p. 80, pl. 29, fig. 8; Renick and Stenzel, 1931, Univ. Texas,
Bull. 3101, pp. 104, 108 (species lists); Plummer, 1933, Univ
Texas, Bull. 3232, p. 646 (species list); Harris and Palmer, 1946,
Bull. Amer. Paleont., vol. 30, No. 117, p. 71; Palmer and Brann,
1965, Bull. Amer. Paleont., vol. 48, No. 218, pp. 341, 342.

V. flabellum Harris, Plummer, 1933, ibid., pp. 811, 812, pl. 8, figs. 5, 8.

V. flabellum var. kingi Plummer, 1933, ibid, pp- 811, 812, pl. 8, fig. 9
(nomen nudum).

Distinguishing characters.— Similar only to V. (R.)
rotunda in shape, dentition, and development of a “pseudo-
lunule” by emphasis of third or second anterior rib, but dis-
tinguishable on basis of ornamentation which consists of
widened, flat-topped costae with regularly spaced nodes
which appear as transverse ridges, easily distinguished from
more delicate ornamentation of V. (R.) rotunda.

Type. — Holotype, a well-preserved valve, dimensions

below, PRI 651.

Sp’n L H %W RN BL/L BH/H H/L “%W/H
Holo. 22.0 21.6 6.9 21 71 91 98 69

Type locality. —Smithville, Bastrop County, Texas.
Weches Formation, middle Eocene age.

Distribution. — Besides the holotype, the following col-
lections were examined in the present investigation: USGS
5285 and 6088, AMNH collection 27674, all from Smithville,
Texas. AMNH 9954, “International and Great Northern
Railroad, Texas” (exact locality not listed). Plummer
(1933) described the Smithville locality as “on the Colo-

”

rado River just north of the bridge in Smithville... ”,

and another “800 to 1000 feet west of highway bridge over
the Colorado River at Smithville.” Plummer (1933) and
Renick and Stenzel (1931) listed other localities in Bastrop,
Robertson, and Brazos Counties, Texas.

Stratigraphic occurrence.— From Weches Formation
to Yegua Formation, middle Eocene age.

Dimensional summary. —

Sample: USGS 6088

N= 10 Dimension
Statistic L H LW RN
OR 14.0-22.4 14.3-21.8 4.8-7.9 22-26
X 18.6 18.3 233
NS) 2.6 2.3 1 2.4
Vv - 14.0 12.8 16.9 10.4
Ox 82 7 4 8
Ratio
BL/L BH/H H/L V“W/H
OR .66-.73 .89-.93 -95-1.02 32-41
xX 69 .90 98 36
Curvature characteristics. —
Sample: USGS 6088 Nit)

S1 H1 $2 H2 $3 H3 S4+ H+ $5

OR 19-33° .6-1.0 28-42° 1.1-2.2 42-55° 1.5-4.2 50-70° 2.9-7.5 45-55°
mx 28.08 38 34.0° 1.6 49.0° 3.1 59:0 = 53 49.0°

Discussion and diagnosis. —V.(R.) flabellum is simi-
lar to V. (R.) rotunda, and Harris (1919) recognized this on
a varietal level of distinction. Reduced major axis compari-
son between a topotype sample of V. (R.) flabellum and the
type collection of V. (R.) rotunda established a correlation
coefficient (Z) of .80, representing a high degree of similar-
ity in growth pattern of Height/Length. This might indicate
a subspecific relationship rather than distinct species. This
course has not been adopted here for several reasons: (a)
the ornamentation of V. (R.) flabellum is uniformly dis-
tinguishable from that of the other species, and ornamental
type is the commonly applied basis for species distinction in
Venericardia; (b) there exist at least two localities where
V. (R.) rotunda and V. flabellum are found in the same
stratum (Renick and Stenzel, 1931 p. 104), signifying that
these two lived sympatrically without interbreeding; (c) no
apparent sexual dimorphism in ornamentation or curvature
pattern is found in this species such as that which charac-
terizes populations of V. (R.) rotunda.

The “variety” V. (R.) flabellum var. kingi Plummer
from the Crockett Formation is a nomen nudum, because
no description exists nor distinguishing characters which
serve to differentiate it from the Weches populations of the
species. Plummer’s (1933, pl. 8, figs. 9a,9b) photographs
of unidentified specimens show forms indistinguishable from

ee

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 93

specimens in the earlier Weches Formation in the same
region of Texas.

V. (R.) flabellum is undoubtedly derived from the
same ancestry as V. (R.) rotunda, but the absence of inter-
mediate forms in the sediments of late Midway or Wilcox
age poses a significant gap between these two middle Eocene
species and V. (R.) eoa, the logical ancestor for both in the
early Paleocene.

Etymology. — The trivial name flabellum, fan (Latin),
refers to the transversely expanded costal nodes.

V. (Rotundicardia) rotunda Lea, 1833

Plate 22, figures 8, 9a, 9b;
Plate 23, figures 1, 2a, 2b

V. rotunda Lea, 1833, Contributions to Geology, p. 70, pl. 2, fig. 48;
Harris, 1919, Bull. Amer. Paleont., vol. 6, No. 31, pp. 78-80, pl. 28,
figs. 4-7 (vars. fungina, funiculus) (non var. flabellum, p. 80,
pl. 29, fig. 8, non var. coloradonis, p. 81, pl. 29, fig. 9, mon “vary-
ing toward trapaquara,” pp. 80,81, pl. 29, figs. 6,7); Renick and
Stenzel, 1931, Univ. Texas, Bull. 3101, p. 104 (list); Plummer,
1933, Univ. Texas, Bull. 3232, pp. 646, 811 (non pl. 8, figs. 7a,
7b), Gardner, 1945, Geol. Soc. America, Mem. 11, p. 93,
?pl. 7, ?figs. 7a, 7b; Harris and Palmer, 1946, Bull. Amer.
Paleont., vol. 30, No. 117, pl. 17, figs. 4, 5a, 6a; Palmer and Brann,
1965, Bull. Amer. Paleont., vol. 48, No. 218, p. 341.

?Cardita subrotunda Conrad, 1848, Acad. Nat. Sci. Philadelphia,
Proc. 1847, vol. 3, p. 298; Conrad, 1848, Acad. Nat. Sci. Phila-
delphia, Jour., vol. 1, 2d ser., p. 129, pl. 14, fig. 11.

IV’. transversa var. rotunda deGregorio, 1890, Mon. Fauna l’Eoc.
Alabama, p. 212, pl. 13, figs. 6-13.

Distinguishing characters. — V. (R.) rotunda is similar
in general form and dentition to V. (R.) diversidentata
Meyer and V. (R.) flabellum Harris but distinguished from
those species and other rotundicards mainly on ornamenta-
tion. Costal nodes on the high, bladelike ribs are of two
types: (a) subdued, irregularly spaced expansions of the
central cord (funiculate) and (b) closely and regularly
spaced flaring scales (funginate). Nodes of V. (R.) diversi-
dentata are uniformly spaced, flattened transverse ridges on
similar ribs; those of V. (R.) flabellum much coarser trans-
verse ridges on wider ribs. Lower average rib number in /.
(R.) flabellum (about 23) and much higher in V. (R.) di-
versidentata (about 30); V. (R.) rotunda average about 26.
Smaller, less inflated umbones in V. (R.) rotunda than in
V. (R.) diversidentata, but certain individuals (V. “praeci-
sa”) comparable in this respect. Development of pre-um-
bonal ridge formed by second or third anterior costa and
reflection of the shell in a “pseudo-lunule” found only in
V. (R.) rotunda and V. (R.) flabellum, not in V. (R.)
diversidentata.

Type.— Lectotype (herein designated), Lea’s illus-
trated right valve ANSP 5267, dimensions below. Para-
lectotype, figured left valve ANSP 5268, dimensions below.

Sp’n L H %W RN BL/L BH/H H/L %W/H
Lect. 24.5 22.9 8.5 26 Al -90 93 37
Para. 25.0 23.5 7:5 27 67 91 94 32

Type locality. — Upper part of Claiborne Bluff, on east
bank of Alabama River, south of the bridge crossing of U. S.
Highway 84 and just west of the town of Claiborne, T 7 N,
R 5 E, Monroe County, Alabama. Gosport Sand, late mid-
dle Eocene age.

Distribution. — Besides the type collection (ANSP
5267-5284), the following collections were examined in the
present investigation:

1. AMNH 1038, 0.8 mile north of intersection between
U. S. Highway 80 and State Highway 15, on highway 15
just south of bridge over Potterochitto Creek. Archusa Marl
Member, “Wautubbee Formation” (Lisbon Fm.), middle
Eocene.

2. AMNH 1044, cut along Gulf Mobile and Ohio Rail-
road, across from L. B. Jones’ fire tower, Newton, Newton
County, Mississippi. Potterochitto Member, “Wautubbee
Formation,” middle Eocene.

3. AMNH 9723/2, Farmingdale, Monmouth County,
New Jersey. Shark River Marl, middle Eocene.

4. USGS 4272, Saline Bayou, St. Maurice, Winn Parish,
Louisiana. Cook Mountain Formation, middle Eocene.

5. USGS 6087, Gopher Hill, bluff on the Tombigbee
River about 1/2 mile above St. Stephens Bluff, Washington
County, Alabama. Gosport Sand, middle Eocene.

6. USGS 2009, 2 miles west of the Orangeburg Court-
house, Orangeburg County, South Carolina. McBean Forma-
tion, middle Eocene.

Plummer (1933) reported this species from the Weches
Formation (middle Eocene) at Smithville, Renick and
Stenzel (1931) from the Crockett Member of the Yegua in
Texas, and Gardner (1945) reported poorly preserved simi-
lar, if not conspecific, specimens from the Laredo Forma-
tion of Cook Mountain age in northeast Mexico. Foster
(1940) reported the presence of this species in “lower Clai-
borne” beds in Clarke County, Mississippi.

Stratigraphic occurrence. —Lower Lisbon to Gosport
Formations, middle Eocene age.

Dimensional summary, —

Sample: Type collection, ANSP 5267-5284*

N = 20 Dimension

Statistic L H VA RN
OR 5.0-25.0 4.9-23.5 1.7-8.5 23-30
X 14.5 13.6 4.8 26.3
S 6.3 oie) Pall 1.6
V 43.6 43.4 43.3 5.9
Ox 1.4 1.3 5 4

94 PALAEONTOGRAPHICA AMERICANA (YI, 39)

Sample: Type collection, ANSP 5267-5284* (continued )

N = 20 Ratio

Statistic BL/L BH/H H/L VW/H
OR .59-.80 .87-.95 -90-.99 31-.38
xe 69 91 94 35

*Not including 5279 (not this species).
Curvature characteristics. —

Sample: Type collection, ANSP 5267-5284* N= 20

S1 Hi $2 H2 $3 H3 S4 Fige*) 9S5**

OR 20-33° .3-1.2 22-48° .7-2.1 40-52° 1.4-8.3 52-55° 3.8-12.0 43-53°
x 24.3° 6 37.4° 1.2 44.6° 35) 48.0° 5:5 45.4°
*Not including 5279 (not this species).
** Only seen in funginate individuals.

Discussion and diagnosis—Harris (1919) named two
“varieties” of V. (R.) rotunda from topotype specimens to
recognize two distinct types of ornamentation: V. rotwnda
var. fungina (see Pl. 22, figs. 9a, 9b), which has costal nodes
“reminding one of a series of minute, inverted funnels, or
perhaps a dead branch bedecked with a series of saucer-
shaped fungi, hence . . . the varietal name fungina. The
second, slightly more abundant and growing at times a little
larger than the first, is . . . ornamented by simple nodules,
giving a cordlike aspect, hence the varietal name funi-
culus ...” (see Pl. 22, fig. 8). The distribution of these orna-
mental types corresponds exactly to the distribution of
curvature pattern (see Text-figs. 16, 22) in the type collec-
tion of 20 individuals. The most reasonable explanation for
this is sexual dimorphism. Highly ornate males with fungi-
nate ornamentation become less convex upon reaching ma-
turity and the simpler, funiculate females become more
convex on reaching maturity in connection with the
“marsupial” habit in reproduction in which the larvae are
nurtured within the umbonal region of the mantle cavity
of the female. The ornate funginate ornamentation is not
found in collections of pre-Gosport age, although both the
male B and female A patterns are seen.

The widespread occurrence of V. (R.) rotunda on the
east and Gulf coastal plains makes it a valuable index fos-
sil for sediments of middle Eocene age and testifies to the
evolutionary success of the species. Wilcoxian and late Mid-
wayan ancestors are unknown, but its descendant in the late
Eocene is V. (R.) diversidentata.

Etymology.—The trivial name rotunda is derived from
the Latin rotundus, circular or round, in reference to the
subcircular lateral outline of the shell.

V. (Rotundicardia) eutawcolens Harris, 1919

Plate 22, figures 5a, 5b

Venericardia eutawcolens Harris, in Van Winkle and Harris, 1919,

Bull. Amer. Paleont., vol. 8, No. 33, pp. 13, 14, pl. 2, figs. 1, 2;

Palmer and Brann, 1965, Bull. Amer. Paleont., vol. 48, No. 218,

p. 329.

Distinguishing characters—Neither total shell shape,
musculature, nor dentition known, but can be distinguished
from other rotundicard species by ornamentation. Costal
cord narrow and rounded, ornamented by subdued asym-
metrical nodes pointing toward the ventral margin. Para-
costals irregularly noded, strongly developed and projecting
over intercostal space. Intercostals broadly U-shaped with
moderate constriction due to overhanging paracostals. Ribs
comparable to those of V. (R.) crenaea Gardner, but less
strongly developed.

Type.—Lectotype (herein designated), a natural mold
of nearly complete left exterior, PRI 1401. Paralectotype, a
natural external mold of left fragment, PRI 1402. Lectotype
dimensions below (estimated ).

Sp'n 1b H %W RN BL/L BH/H H/L “~AW/H
Lecto. 26.8 26.8 9.0 25 75 -90 1.00 34

Type locality —“Eutaw Springs and Centre Hill, South
Carolina” (Harris, 1919). Eutaw Springs, about 3 miles east
of Eutawville, Orangeburg County, South Carolina.

Distribution —Known only from type locality.

Stratigraphic occurrence——Santee Limestone, middle
Eocene age.

Dimensional summary.—See type dimensions.

Curvature characteristics —

Sample: Holotype N=
Shs Hd: ase Hens H3 S+ H4+ = §5
OR 20° pA Pete 2 Jie Ses 22h age

Discussion and diagnosis—The ornamentation of J.
(R.) eutawcolens can be compared only to that of V. (R.)
crenaea of the Kincaid Formation and V. (B.) greggiana
Dall of the Tuscahoma Formation. Neither of the latter
species are close to the Santee species in stratigraphic place-
ment. It is excluded from a relationship with V. (B.) greg-
giana because of the high convexity, well-defined posterior
truncation and narrowly based ribs, despite the great ex-
pansion of paracostals in that species. It is similar in con-
vexity and lack of posterior truncation as well as broadly
based ribs with noded paracostals to V. (R.) crenaea, and a
descendant relationship to that species is proposed, despite
the wide stratigraphic gap which separates the two.

The curvature pattern of V. (R.) eutawcolens is some-
what atypical and of the B type, but showing a stage of
low convexity in the third (S3) growth stage. However, the
beak is not preserved in the holotype, and its estimated
position may have involved error in drawing the curve.

AMERICAN ALTICOSTATE VENERICARDS: HEasLip 95

Etymology —Trivial name eutawcolens is geographic
in honor of Eutawville, near the type locality.

V. (Rotundicardia) diversidentata Meyer, 1885
Plate 23, figures 3a-3c, 4, 5a, 5b, 6, 7a-7c, 8

V. diversidenta Meyer, 1885, Amer. Jour. Sci, 3d ser., vol. 29, pp. 460,
467; Casey, 1903, Acad. Nat. Sci. Philadelphia, Proc., vol. 55,
p. 264; Harris and Palmer, 1946, Bull. Amer. Paleont., vol. 30,
No. 117, pp. 69-71, 72, 73, pl. 17, figs. 1-3, 5, 6, 7-10; “var.
garlandica”; Harris, ibid., p. 73, pl. 17, figs. 15, 16a, 17a;
Palmer and Brann, 1965, Bull. Amer. Paleont., vol. 48, No. 218,
p. 328.

Cardita tetrica Conrad, 1854, in Wailes, Rept. Agric. and Geol. of
Mississippi, p. 289 (nomen nudum); Aldrich, 1885, Amer. Jour.
Sci., 3d ser., vol. 30, p. 307; Dall, 1903, Wagner Free Inst. Sci.,
Trans., vol. 3, No. 6, p. 1424; Harris, 1919, Bull. Amer. Paleont.,
vol. 6, No. 31, p. 85-

Venericardia tetrica Conrad, Dall, 1903, ibid., pp. 1427, 1428, 1429;
Harris, 1919, Bull. Amer. Paleont., vol. 6, No. 31, p. 85.

V’. jacksonensis Conrad, 1866, Smithsonian Misc. Coll., vol. 7, No. 200,
p. 23 (nomen nudum); Vaughan, 1896, U.S. Geol. Sur., Bull., No.
142, p. 51; Dall, 1903, ibid., loc. cit.

V. rotunda Lea var., Harris, 1894, Ann. Rept. Geol. Sur. Arkansas
for 1892, vol. 2, p. 149.

V. praecisa Dall, 1903, tbid., pp. 1424, 1427, 1429, pl. 56, figs. 7, 8;
Harris, 1919, of. cit., loc. cit.; Harris, 1946, and Palmer, zbid.,
pp. 71, 72, pl. 17, figs. 12-14.

I’. scabricostata Guppy, Richards in Richards and Palmer, 1953,
Florida Geol. Sur. Bull., vol. 35, pp. 47, 48, pl. 10, figs. 1, 2.

Distinguishing characters—General shape and denti-
tion similar to V. (R.) rotunda but distinguishable from
that species by more numerous ribs which contain closely
spaced flattened transverse nodes and not flaring as those
of that species. Beaks somewhat higher and more inflated
in some, but low, noninflated beaks comparable in other
specimens. Anterior left cardinal more inclined on posterior
face than steeper feature in V. (R.) rotunda, with conse-
quent wider angle between right anterior and medial car-
dinal. Absence of “pseudolunule” and ridge formed by first
two anterior ribs found in V. (R.) rotunda.

Discussion —Because of the distinction between the
populations of the Mississippi region and those farther east
in Florida, two subspecies are erected.

V. (Rotundicardia) diversidentata diversidentata Meyer, 1885
Plate 23, figures 3a-3c, 4, 5a, 5b, 6

Synonymy includes all references except those under
V. vicksburgiana and V. scabricostata

Distinguishing characters. — Distinguished from the
subspecies V. (R.) diversidentata vicksburgiana (Dall) by
the somewhat more accentuated nodes which are more
numerous and closely spaced, and higher rib number, Sexual
dimorphism also pronounced in this subspecies, with flat-
tened, low beaked forms (‘“‘praecisa”) which were probably
males of the population. This was not noted in V. (R.)
diversidentata vicksburgiana.

Type——Holotype a right valve, ASM type No. 74,

dimensions below.

Sp’n L H %W RN BL/L BH/H H/L Y%W/H

Holo. 17.5 18.0 6.8 31 -66 87 1.03 38

Type locality —*Jackson, Mississippi.”

Distribution Besides the rather poorly preserved holo-
type, material from the following localities was examined in
the present investigation:

1. All Harris’ figured specimens in the Paleontological
Research Institution collections from Moody’s Branch and
Town Creek, Jackson, Hinds County, Mississippi; Garland
Creek, north of Shubuta, Clarke County, Mississippi, and
Hiwannee, Wayne County, Mississippi; Montgomery, Grant
Parish, Louisiana, and Bunker Hill on the Ouachita River,
Caldwell Parish, Louisiana; Vince Bluff on the Saline River,
Cleveland County, Arkansas.

2. USGS collections from Jackson, Hinds County, Mis-
sissippi numbers “Jackson 60,” 4270, 3746, 6467, 3735, 6466
and 10377 (Moody’s Branch).

3. USGS 2232, Sec. 32, T 10 S, R 11 W, near crossroads
church, Cleveland County, Arkansas.

4. AMNH 1045, Montgomery, Louisiana.

Stratigraphic occurrence——Moodys Branch Marl of the
Jackson Group, late Eocene age.

Dimensional summary.—

Sample: USNM 136672, Jackson 60, Jackson, Miss.

N= 7 Dimension
Statistic ib H LW RN
OR 16.4-20.6 17.1-21.8 6.1-9.8 27-34
xX 19.4 19.8 7.8 29.7
S 1.5 tee, 1.1 2.4
V HA 8.7 14.5 8.2
(OMe 6 “ij A 9
Ratio
BL/L BH/H H/L Y4W/H
OR -64-.74 .83-.90 -94-1.06 .36-.50

XG 69 84 1.02 40
Curvature characteristics.—

Sample: PRI 4275, 4276, 4278, 4274, Jackson, Miss. Nt

S1 H1 $2 H2 $3 H3 S4 H+ $5

OR 22-30° .5-.8 33-46° .9-1.8 46-52° 1.9-3.9 55-63° 5.0-9.1 49-52°
xe 2513 ae oe 1) Oe 4: 48.8° ON Sy ioe 6.6 50.0°

Discussion and diagnosis —The presence of smaller in-
dividuals which are flattened and more finely ornamented,
such as those which Dall (1903) named V. praecisa, in
most collections examined seems to indicate that sexual
dimorphism characterizes this subspecies, just as the an-
cestral V. (R.) rotunda contained two distinct ornamental
types. The difference here is more subtle, however, and both

96 PALAEONTOGRAPHICA AMERICANA (VI, 39)

forms are characterized by ribs with closely spaced trans-
verse nodes, giving a “T-rail” cross-section described by
Dall (1903). However, the higher, more inflated beaks and
umbones in most individuals seem to indicate females and
the more “rotund,” low beaked individuals, males. No dis-
tinction can be made on the basis of curvature and both are
characterized by the B pattern. Reduced major axis com-
parison between Dall’s types of V. praecisa and a topotype
collection from Jackson, Mississippi, of this subspecies failed
to indicate any significant difference.

Many of the varietal names by Harris were only varia-
tions. His (1946) “variety” “garlandica” of this subspecies
is a worn small individual with a broken anterior, giving a
spurious height to the shell.

Gardner (1945) recorded specimens from beds of Jack-
son age in northeast Mexico which are similar to V. (R.)
diversidentata but have a lower rib number than the topo-
types. These have not been examined in the present in-
vestigation, and no comment can be made on their affinities.
They may represent a subspecies restricted to that area,
just as V. (R.) diversidentata vicksburgiana lived in the
eastern part of the Gulf. Curiously, no Texas occurrences of
this species are reported, unless Plummer’s (1933) reference
to a poorly preserved “Venericardia sp.” in the Whitsett
Member of the Fayette Formation of Jackson age is con-
specific.

V. (R.) diversidentata s. l., is the last widely occurring
rotundicard in North America, and its descendant in the
early Oligocene, V. (R.) carsonensis Dall is restricted to a
small area in eastern Mississippi and represents the last
known species belonging to this subgenus.

Etymology.—The trivial name diversidentata is de-
rived from the Latin diversus, different, and dentatus,
toothed. The reference is not clear.

V. (Rotundicardia) diversidentata vicksburgiana Dall, 1993
Plate 23, figures 7a-7c, 8

V’. vicksburgiana Dall, 1903, Wagner Free Inst. Sci., Trans., vol. 3,
No. 6, p. 1428, pl. 56, fig. 6; Palmer and Brann, 1965, Bull. Amer.
Paleont., vol. +8, No. 218, p. 345.

V’. scabricostata Guppy, Richards im Richards and Palmer, 1953,
Florida Geol. Sur. Bull., vol. 35, pp. 47, 48, pl. 10, figs. 1, 2.
Distinguishing characters—Distinguished from V. (R.)

diversidentata diversidentata mainly by less numerous ribs

(18-23) than that species (24-34) and less expanded costal

nodes. Specimens with flattened shell and circular outline

characteristic of that subspecies, not evident in populations
of V. (R.) diversidentata vicksburgiana,
Type.—Holotype, silicified pair of valves from same in-

dividual, posterior margin broken, USNM 164548, dimen-

sions below.

Sp'n IE: H %W RN BL/L BH/H H/L “%W/H
Holo. 15.8 14.8 5.7 23 hi) 86 94 239

Type locality —USGS sta., Martin Station, near Ocala,
Marion County, Florida. Ocala Limestone, late Eocene age.

Distribution.—Beside the type collection, material was
examined from FGS localities L-93 and L-139, Levy County,
Inglis Formation, Ocala Group, and Syracuse University
Collection 9619, Camp Wheeler, Bibb County, Georgia,
Barnwell Formation, late Eocene age.

Stratigraphic occurrence.—‘Ocala Limestone”, Inglis
Formation, Ocala Group; Barnwell Formation, Georgia. All
late Eocene age.

Dimensional summary.—

Sample: USNM 164548, Type Collection

NS Dimension
Statistic 1 H 14W RN
OR 8.1-15.8 8.1-14.8 3.1-5.7 21-23
xX 11.0 11.0 4.1 22.0
Ss 4.2 3.4 1.4 1.0
V os 37.9 31.3 34.2 4.6
OX 24 2.0 8 6
Ratio
BL/L BH/H H/L Y4W/H
OR .69-.77 -85-.91 -94-1.10 .35-.39
be ne .87 1.01 37
Curvature characteristics.—
Sample: USNM 164548, Type collection Nis
S1 H1 $2 H2 $3 H3 S4 H+ $5

OR 24-44° 4-.7 32-40° 1.0-1.7 46-54° 2.5-4.0 57-61° 4.0-8.56 52-53°
K 30.7" 6 950° 1S 49.0° 3.4.) S910" ~ Gaeeeneee

Discussion and diagnosis—Specimens from the type
locality at Martin Station are comparable to many in col-
lections from Jackson, Mississippi, and the apparent geo-
graphic isolation of the two populations resulted in the de-
velopment of two subspecies in the eastern and central
parts of the Gulf Coastal waters during late Eocene time.
No “praecisa’-like individuals are found in Florida collec-
tions, and this, as well as the lower rib number, is a basis for
distinction.

Dall’s (1903) comment that, “As the last of its race it
is small and degenerate with the feebleness of sculpture as
an indication of senility,” may be erroneous in two respects.
V. (R.) diversidentata vicksburgiana is the logical ancestor
for V. (R.) carsonensis of the early Oligocene which has an
even lower rib count range (18-22). The “feebleness of
sculpture” may be a result of preservation as siliceous pseu-
domorphs.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 97

A comparison of Richards’ (1953) specimens of “V.
scabricostata’” (see Pl. 23, figs. 7a-7c) indicates they are
members of this subspecies, with a similar rib number but
a slightly greater shell thickness and somewhat heavier
dentition. These are found in the Inglis Formation, Ocala
Group in western Peninsular Florida, and may bear an
ecophenotypic relation to the thinner topotype specimens
found in the “Ocala Limestone.”

Etymology.—The trivial name vicksburgiana was given
by Dall under the misconception that the limestone around
Ocala, Florida, was equivalent to the limestones of the
Vicksburg Group of middle Oligocene age because of litho-
logic similarities and extensive silicification of fossils in the
Glendon Formation of that group. A specimen in Dall’s type
collection is imbedded in limestone matrix containing the
Jackson orbitoid, Lepidocyclina mortont Cushman.

V. (Rotundicardia) carsonensis Dall, 1903
Plate 23, figures 9a, 9b, 10a-10c

V. vicksburgensis Casey, 1903, Acad. Nat. Sci. Philadelphia, Proc.,
p. 264 (nomen dubium).

V’. carsonensis Dall, 1903, Wagner Free Inst. Sci., Trans., vol. 3,
No. 6, p. 1427, pl. 56, fig. 9; Harris, 1919, Bull. Amer. Paleont.,
vol. 6, No. 31, pp. 81, 84, 85; Gardner, 1945, Geol. Soc. America,
Mem., vol. 11, p. 94; Harris and Palmer, 1946, Bull. Amer.
Paleont., vol. 30, No. 117, pp. 70-73, pl. 17, fig. 9a.
Distinguishing characters—Most similar to V. (R.)

diversidentata and V. (R.) rotunda in general shape and

convexity but distinguishable from those species by orna-
mentation which consists of a thin, elevated cord with ob-
scure traces of paracostal threads. Nodes are closely spaced,
somewhat elevated rounded transverse ridges becoming
somewhat spinose on the anterior ribs. Nodes not flattened
as in V. (R.) diversidentata but not nearly so pronounced as
in V. (R.) rotunda. Costals slope into broadly U-shaped,

comparatively wide intercostals. Low rib number (18-22)

similar to that of V. (R.) diversidentata. Dental pattern

similar to those species but differs in triangular projection
on commissural face of lunule formed by the prominent an-
terior cardinal (3a) of the right valve and the lunule face,
below which lunule is excavated to the ventral margin of
hinge plate. Face of lunule on left valve correspondingly
indented and ridge joins anteroventral corner of the an-
terior cardinal (2) to the anterior lateral tooth.

Type.—Holotype, a right valve, USNM 140693, dimen-
sions below.

Sp’n L H YW RN BL/L BH/H H/L “%W/H
Holo. 13.3 12.8 4.7 20 64 91 97 37

Type locality —USGS 2633, Carson’s Creek, five to six
miles southeast of Shubuta, Wayne County, Mississippi.

Distribution—Beside the type collection, collections
from AMNH 1050, below bend in Chickasawhay River, SE
1/4, sec. 29, T 10 N, R 7 W, about 1 mile south of Hiwan-
nee station, and PRI shell (4277) from Red Bluff Station
were examined in the investigation. All localities in Wayne
County, Mississippi.

Stratigraphic occurrence—Red Bluff Formation, early
Oligocene age.

Dimensional summary.—

Sample: AMNH 1050

N = 6 Dimension
Statistic L H 1,W RN
OR 8.6-7.7 7.9-17.7 2.6-8.5 18-21
xX 12.8 12.7 4.7 20.0
S 3.3 3.6 1.3 1.3
View Aye | 28.5 28.6 6.3
Ox 1.3 1.5 6 5
Ratio
BL/L BH/H H/L Y4,W/H
OR .61-.73 .84-.94 .92-1.03 .35-.39
x .67 .89 AS av
Curvature characteristics.*—
Sample: AMNH 1050 N= 75
S1 H1 $2 H2 $3 H3 S4 H4+ S5

OR 19-30° .5-.8 39-43° .9-1.6 45-54° 1.5-3.4 49-59° 4.6-9.0 44-53°
».¢ 23.4° Opto Ol) al 50.0° 2.5 54.0° 6.5 48.0°

*B pattern only, similar to /’. (R.) diversidentata.

Discussion and diagnosis. — Casey’s name V. vicks-
burgensis, is rejected as nomen dubium because of inadequate
description and lack of identifiable specimen which might
be a holotype, even though his publication (February, 1903 )
pre-dates Dall’s (October, 1903). A left valve in the Casey
collection at the U. S. National Museum may be the speci-
men that the author planned to use as the type.

The high degree of similarity in shape to the Eocene
rotundicards, as well as dentition and ornamentation, makes
it apparent that V. (R.) carsonensts is the Oligocene descen-
dant of V. (R.) diversidentata, which was recognized by
Harris (in Harris and Palmer, 1946). The similarity in rib
number to the subspecies, V. (R.) diversidentata vicks-
burgiana seems to indicate descent from that form, rather
than the Mississippi populations, although geographic occur-
rence might imply the opposite interpretation.

V. (R.) carsonensis is the last rotundicard species re-
corded in coastal plain sediments, and its lineage seems to
have come to an end in the early Oligocene. The only species
of Venericardia known from sediments of middle Oligocene
(?) age is V. (Claibornicardia) nasuta Dall, possibly from
the Glendon Limestone in Alabama, indicating at least that

98 PALAEONTOGRAPHICA AMERICANA (VI, 39)

the claibornicards were still extant at that time, and most

likely gave rise to the late Oligocene glyptoactids.
Etymology. — The trivial name carsonensls 1s ge0-

graphic in honor of the type locality at Carson’s Creek.

Subgenus CLAIBORNICARDIA Stenzel and Krause, 1957

Claibornicardia, n. subgen., Stenzel and Krause, 1957, in Stenzel,
Krause, and Twining, Univ. Texas, Pub. 5704, pp. 106, 107;
Heaslip, 1960, Geol. Soc, Amer. Bull., vol. 71, No. 12, pp. 1885,
1886 abstract.

Type Species. — Venericardia (Claibornicardia) alti-
costata (Conrad), 1833, from the Gosport Sand at Clai-
borne, Alabama. Late middle Eocene age.

Type Locality—Claiborne bluff, on the east bank of
the Alabama River near the town of Claiborne, T 7 N,
R 5 E, Monroe County, Alabama.

Distribution —The subgenus Claibornicardia is known
from sediments of the Gulf Coastal Plain as far west as Cen-
tral Texas, and from the Atlantic Coastal Plain as far north
as New Jersey. Whether the subgenus extends into Mexico
or Central America and South America, or the west coast of
North America remains to be established.

Stratigraphic occurrence. — Recorded occurrences of
members of Claibornicardia from the Bashi Member of
early Eocene (Wilcoxian) age to the Glendon Limestone
(?) of middle Oligocene (Vicksburgian) age.

Subgeneric definition —Venericards with highly inflated
shell, high degree of posterior and postero-ventral elonga-
tion and generally inflated beaks. Ornamentation consists
of radiating costae having central costal cord with enlarged
or suppressed nodes, and a paracostal cord on either side of
the central cord as sharp or rounded terraces resulting in
tripartite cross-section. Costal nodes and paracostals sup-
pressed in advanced members of the subgenus.

Singular dental pattern with right anterior cardinal
(3a) a horizontal blade as a continuation of the dorso-an-
terior point of the medial cardinal (3b), and lying dorsal
to the anterior socket for the anterior cardinal (2) of the
left valve which is a horizontal blade or very broadly tri-
angular in lateral aspect.

Growth pattern of curvature of the “B” type with in-
creasingly convex spiral segments and a terminal mature
segment of low convexity.

Similar in convexity, shell shape, and ornamentation to
members of the ancestral subgenus Baluchicardia, but that
group is characterized by an erect triangular left anterior
cardinal (2) and a right anterior cardinal tooth (3a) which
borders anterior margin of socket for left anterior cardinal
(2), as well as a convex pattern of curvature of the “A”

type.

Discussion and diagnosis. — The subgenus Claiborni-
cardia, as originally proposed, included eight species of which
two are still undescribed (Stenzel, et al., 1957, p. 106, Nos.
5,6), and one (ibid., No. 1) is from a region so far removed
from the area of claibornicard development that the inclu-
sion of V. acuticosta Lamarck from the Calcaire Grossier
(Lutetian, middle Eocene) may be unwise before a thorough
study of total European relationships is undertaken. So
many instances of evolutionary mimicry or convergence are
noted among the American venericards, that the resem-
blance between the Paris Basin species and V’. (C.) alticos-
tata may be fortuitous and not due to direct phylogenetic
relationship. V. (R.) rotunda Lea of the Atlantic Coastal
Plain and V. (?) imbricata (Gmelin) of the Calcaire Gros-
sier are undoubtedly ecologic analogues, but their placement
in the same taxon because of “rotund” shell outline is open
to question. Dall (1903, p. 1419) noted a series of “parallel
forms” between the Eocene of America and France. V.
(Glyptoactis) hesperide Gardner of the Shoal River Miocene
in Florida assumes the same shell form and ornamentation of
the unrelated V. (Baluchicardia) whitei Gardner of the
early Midway of Texas, and closely converging with these
two species are members of the subgenus Megacardita Sac-
co, 1899, of the Alpine Miocene. That these forms arose in-
dependently is a firm conclusion and testifies against the
hasty formation of taxonomic judgments.

The statement by Stenzel and Krause (1957, p. 107)
that one lineage of Claibornicardia is characterized by
“. . . the complete loss of the right anterior cardinal hinge
tooth” (3a) is erroneous, and they have mistaken the
aligned position of this tooth with the anterior point of
the medial right cardinal (3b). In only one species is the
relation indistinct, but the highly eroded condition of all
known specimens of V’. (C.) trapaquara Harris explains this.
Plastolene impressions of the corresponding socket in the
left valve make this clear. This distinctive pattern of aligned
3a and 3b is one of the hallmarks of Clatbornicardia. In fact,
the “primitive” lineage containing V. (C.) alticostata as
described by Stenzel and Krause (op. cit., p. 107) is actually
highly advanced in the high degree of variability in develop-
ment of 3a as well as ornamental simplification and curva-
ture pattern.

The original scope of Claibornicardia is highly modified
in the present investigation, and the range extended down-
ward into the early Eocene and upward (tentatively) into
the middle Oligocene, pending certain dating of J. (Ga
nasuta Dall, recognized by Stenzel and Krause (1957) as
belonging to this group.

The main basis for distinction between the subgenera

tatiana

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 99

Baluchicardia and Claibornicardia is the dental pattern. The
ancestral former group is characterized by an erect left an-
terior cardinal (2) which is strongly suppressed in the lat-
ter to a lower triangular horizontal blade. Members of
Baluchicardia also possess a fairly well-developed right an-
terior cardinal (3a) which borders the anterior margin of
the socket (2’) in this valve, whereas this tooth is parallel
to the anterior point of the medial cardinal (3b) and lies
above the socket in Claibornicardia. General shape and
ornamentation are continued from one lineage to the other,
but patterns of curvature differ: Baluchicardia is charac-
terized by the convex “A” pattern, and Claibornicardia (ex-
cept the advanced species) by the “B” pattern.

The type species for the subgenus, V. (C.) alticostata
(Conrad) is probably the worst choice which the authors
could have made because it exhibits a high degree of mor-
phological variability in shape, ornamentation, dentition
and curvature pattern, and represents a species in a stage
of evolutionary advancement over its consistent ancestors.

Etymology—The name Claibornicardia is geographic
in honor of the type locality of the type species, Claiborne
Bluff, Alabama, and the Latin cor, heart.

V. (Claibornicardia) linguinodifera, n. sp.
Plate 24, figures la-1c, 2

V. alticostata var. Harris, 1919, Bull. Amer. Paleont., vol. 6, No. 31,
p. 84, pl. 31, figs. 1-4.

Non VF. alticostata (Conrad), 1833.

?V. cf. V. rotunda Lea, Lowe, 1933, Mississippi Geol. Sur., Bull., vol.
25, p. 103.

Non IV’. rotunda Lea, 1833.

Distinguishing characters—Similar in ornamentation
and dental pattern to V. (C.) coloradonis Harris but dis-
tinguishable on the basis of its distinctive ovate shell margin
and strongly inrolled beaks, relative lack of posterior trunca-
tion and minimum of postero-ventral elongation. Tongue-
like costal nodes of this species are more closely arranged
than those of V. (C.) coloradonis, and the more pronounced
paracostals are defined by terrace-like cords which are
noded in concert with the central costal cords.

Type.—Holotype, a right valve, USNM 644318, No. 1
of collection USNM 154544. Paratype USNM 644319, a

left valve, No. 7 of collection, dimensions below.

Sp’n L H YW RN BL/L BH/H H/L Y%W/H
Holo. 13.6 13.4 5.3 18 78 82 99 46
Para. 10.5 10.8 4.4 20 72 -80 1.03 Al

Type locality —USGS 3099, Woods Bluff on the east
bank of the Tombigbee River, Clarke County, Alabama.

Bashi Formation, early Eocene age.

Distribution —Besides the type collection of 22 individ-
uals, collections from USGS localities 20850, abandoned
road east of U. S. Highway 241, Mill Creek, southwest
corner sec. 29, T 6 N, R 28 E, Henry County, Alabama, and
Harris’ PRI figured specimens (also from Woods Bluff)
were examined in the present investigation.

Stratigraphic occurrence—Bashi Formation, early Eo-
cene age.

Dimensional summary —

Sample: USNM 154544, Type Collection

N22 Dimension
Statistic i H YW RN
OR 3.2-13.6 3.1-13.4 1.5-5.3 17-21
2.8 9:5. 9.5 3.8 19.2
S 2.5 2.5 ee) 1.1
re 26.2 26.1 23.0 5.9
Ox 5 5 2 L
Ratio
BL/L BH/H H/L 14W/H
OR -65-.83 .76-.88 -96-1.03 -37-48
axe A .84 1.00 41

Curvature characteristics —

Sample: USNM 154544, Type collection N =, 132

S1 H1 $2 H2 83 H3 S4+ H+ S5

OR 21-42° .3-.9 22-48° .9-1.9 50-61° 1.9-3.7 57-71° 3.8-6.1 47-54°
a2 eo 40:8" 1-3 544° 72:5 63i8° Bae ah i)

*Alternate individuals of entire sample, except 3 smallest in series.

Discussion and diagnosis —V. (C.) linguinodifera of the
Bashi Marl is the first recorded claibornicard, and the only
one known from sediments of Wilcoxian age. In certain
respects, it is similar to the last known baluchicard, V. (B.)
greggiana Dall of the underlying Tuscahoma. Both have
high, tightly rolled umbones, little posterior elongation and
high convexity, as well as tripartite ornamentation. The
similarity ends there, however, and the comparatively tiny
Bashi species has the characteristic in-line 3a and flattened
2 anterior cardinal teeth which mark it as a member of the
more advanced Claibornicardia, while the dental pattern of
Baluchicardia is distinguishable in the Tuscahoma species.

However, the ancestry of V. (C.) linguinodifera is most
certainly among the baluchicards, and perhaps even V. (B.)
wilcoxensis Dall, because this is the only recorded species
of this group in late Paleocene time in the eastern part of
the Gulf. V. (B.) bulla Dall, of corresponding age in Texas
is discounted as an ancestor of later forms because of the
peculiar shell thickening.

From /. (C.) linguinodifera evolved the similar V. (C.)
coloradonis Harris of Claibornian age. The subspecies J.
(C.) coloradonis texalana Gardner from the lower Weches
Formation in Texas in most similar to the Bashi species in

100 PALAEONTOGRAPHICA AMERICANA (VI, 39)

shape, ornamentation, and rib number. This subspecies is
probably the form from which the later Weches to Wautub-
bee V. (C.) coloradonis coloradonis evolved.

Specimens questionably identified as V. rotunda by
Lowe (1933) from south of Meridian, Mississippi, in the
Bashi Marl have not been examined in the present investiga-
tion, but these are probably of this species.

Etymology.—The trivial name linguinodifera is derived
from three Latin roots: lingua, tongue, nodus, knot or
swelling and feron, to bear, in reference to the tongue-like
costal nodes.

V. (Claibornicardia) coloradonis Harris, 1919
Plate 24, figures 3a-3c, 4, 5a-5c, 6

V. (rotunda?) var. coloradonis Harris, 1919, Bull. Amer. Paleont.,
vol 6, No. 31, p. 81, pl. 29, fig. 9; Harris, 1946, Bull. Amer.
Paleont., vol. 30, No. 117, p. 68; Palmer and Brann, 1965, Bull.
Amer. Paleont., vol. 48, No. 218, p. 341. he

V. (rotunda) varying toward trapaquara Harris, 1919, ibid., pp-
80, 81, pl. 29, figs. 6, 7 (fig. 7, No. 650 PRI).

lV’. rotunda Lea, Plummer, 1933, Uniy. Texas, Bull. 3232, pp. 646, 811,
812, pl. 8, figs. 7a, 7b. (Non V’. texalana Gardner, pp. 811, 812,
pl. 8, figs. 5a, 5b).

Non V. rotunda Lea, 1833. hn!

V. trapaquara Harris (partim), Harris, 1919, ibid., pp. 81, 82, pl. 30,
figs.6, 75° 9:

V’. trapaquara subsp. texalana Gardner, 1927, Jour. Washington Acad.
Sci., vol. 17, pp. 370, 371, figs. 24-27; Renick and Stenzel, 1931,
Univ. Texas, Bull. 3101, p. 108 (faunal list); Stenzel, Krause,
and Twining, 1957, Univ. Texas, Pub. 5704, p. 106; Harris,
1946, ibid., vol. 30, No. 117, p. 68 (‘‘variety texalana Gardner’).

Non V’. trapaquara Harris, 1895.

Distinguishing characters —Similar in general charac-
teristics of shape and dentition to V. (C.) linguinodifera, but
differs in higher degree of postero-ventral elongation and
more highly developed posterior truncation, smaller, lower
umbones and less closely or regularly arranged tongue-like
costal nodes.

Discussion.—Distinguishable mainly on the basis of rib
number, two subspecies are recognizable: V. (C.) colora-
donis texalana Gardner and V. (C.) coloradonis coloradonis

Harris.
V. (C.) coloradonis coloradonis Harris, 1919
Plate 24, figures 5a-5c, 6

Synonymy includes all references in species synonymy
except those cited for V. (C.) coloradoms texalana.

Distinguishing characters—Similar to V. (C.) color-
adonis texalana but differs by having higher rib number
than that found in that subspecies, as well as stratigraphic
separation.

Type.—Holotype, a left valve with part of hinge plate
missing, PRI 652, dimensions below. Paratype (herein se-
lected) right valve, PRI 650, “V. rotunda near trapaquara
Harris”, dimensions below.

Sp’n It H ZW RN BL/L BH/H H/L “~W/H
Holo. 21.0 21.1 8.1 26 76 -87 1.00 39
Para. 12.5 11.8 4.7 26 72 85 94 40

Type locality—Holotype locality “Smithville, Texas.”
Probably corresponds to Plummer’s (1933, p. 640) locality
“800 to 1000 feet west of highway bridge at Smithville,
Bastrop County, one of the most famous and most fre-
quently visited fossil localities in Texas.” Paralectotype
locality “Wautubbee and Johnson’s place, Mississippi.”

Distribution.—Besides the type, collections from the
following localities were examined: USGS 5285, type lo-
cality; USGS 13303, west bank Attayae Bayou, Nacog-
doches County, Texas; USGS 2623, 914 miles south of
Hickory, and northwest of Enterprise, Clarke County,
Mississippi; USGS 3569, Saline Bayou, Winn Parish, Louisi-
ana; AMNH 1038 and 1044, Newton, Newton County,
Mississippi (see exact locality description of Newton locali-
ties under V. (R.) rotunda distribution).

Stratigraphic occurrence—Middle Weches Formation
in Texas to Wautubbee Formation in Louisiana and Missis-
sippi, middle Eocene age.

Dimensional summary.—

Sample: AMNH 1038, Newton, Mississippi

N = 24 Dimension
Statistic L H YW RN
OR 6.9-13.4 6.7-12.6 2.4-4.4 23-28
dK 10.2 9.9 3.6 25.6
$ 1.3 1.2 4+ lees
Vee 13.1 12.6 11.6 4.7
Ox 3 5a 1 3
Ratio
BL/L BH/H H/L YW/H
OR -63-.82 -87-.91 -91-1.06 .34-.40
xX .70 89 97. 36
Curvature characteristics. —
Sample: AMNH 1038 N= 3
S1 H1 $2 H2 $3 H3 S4 H+ $5

OR 19-30° .4 30-44° 1.0-1.4 44-49° 1.6-2.3 57-66° 2.1-3.6 51°
KI S2302 ee oe3 eee 45-72 ZO ins Oe 3.0) 51.0)

Discussion and diagnosis —Harris (1919) named the
“variety” “coloradonis” under the impression that this form
was related to V. (R.) rotunda Lea, but examination of the
dental pattern clearly establishes that this form is not char-
acterized by the strong dentition of that species, or indeed,
of Rotundicardia, but represents that of Clatbornicardia.

The species V. (C.) coloradonis coloradonis is found
in the middle and upper Weches Formation at Smithville
and other localities in Texas, and extends its range into the
stratigraphically higher Wautubbee Marl of Cook Mountain

AMERICAN ALTICOSTATE VENERICARDS: HeEasiip 101

age in eastern Mississippi. It evolved from the subspecies
V. (C.) coloradonis texalana Gardner which occurs in the
lower Weches. The main evolutionary modification in the
descendant subspecies is the addition of several ribs to the
ancestral number and slight refinement in the regularity of
costal nodulation. A somewhat greater degree of postero-
ventral elongation also characterizes the descendant popula-
tions.

The range of V. (C.) coloradonis coloradonis is fairly
extensive, and the species retains its identity into Wautub-
bee time, apparently spreading from Texas across the Mis-
sissippi embayment during middle Lisbon time.

Despite the apparent continuity during much of the
middle Eocene, populations remaining in Texas accentuated
postero-ventral elongation, and simplified ornamentation to
produce V. (C.) trapaquaroides, n. sp., which in turn evolved
into the small, highly flattened V. (C.) trapaquara Harris,
as well as V. (C.) complexicosta Meyer and Aldrich (see
Text-fig. 27).

Etymology.—Trivial and subspecific name coloradonis
geographic in honor of the type locality on the Colorado
River at Smithville, Texas.

V. (C.) coloradonis texalana Gardner, 1927
Plate 24, figures 3a-3c, 4

Synonymy includes references under V. trapaquara
subsp. texalana Gardner and V. rotunda Lea, Plummer. Non
V. texalana Gardner, Plummer, 1933, Univ. Texas, Bull.
3232, pl. 8, figs. 5a,5b.

Distinguishing characters. — Distinguished from the
V. (C.) coloradonis coloradonis s.s. mainly on basis of
lower costal number (about 20) as opposed to about 25 in
that subspecies. Distinguishable from most other claiborni-
cards by spinose linguate costal nodes found only in the two
subspecies of V. (C.) coloradonis and in V. (C.) linguinodi-
fera.

Type—Holotype, right valve from unmatching pair
figured by Gardner (1927), ANSP 9860. Paralectotype
(herein designated), left valve, same collection number.
Dimensions below.

Sp’n L H %W RN BL/L BH/H H/L %4W/H
Holo. 17.3 17.0 6.9 21 74 83 98 Al
Para. 17.1 16.8 6.8 21 70 36 -98 40

Type locality. — Black Shoals, Brazos River, Texas.
Equivalent to Burleson Bluff and Colliers Ferry in the older
literature, on east bank of Brazos River on northeast corner
of Andrew Sadberry 41.33 acre tract, Robertson Survey,
Burleson County, Texas. Lower Weches Formation, middle
Eocene age.

Distribution.—The type collection and topotype collec-
tions USGS 9862 and 5288 were examined in the present
investigation.

Stratigraphic occurrence—Lower Weches Formation,
middle Eocene age.

Dimensional summary.—

Sample: USGS 5288, Burleson Bluff

N= 6: Dimension
Statistic iL, H 14W RN
OR 8.9-18.7 8.8-18.7 3.4-6.8 19-22
xX 13.8 13.5 5.1 20.3
S 4.6 4.6 1.6 1.0
Ve 33.2 33.7 30.4 5.1
Ox 19 1.9 6 4
Ratio
BL/L BH/H H/L Y4W/H
OR -65-.78 -85-.89 -95-1.00 -36-.40

».¢ 71 87 98 38

Curvature characteristics. —

Sample: USGS 5288

S1 H1 $2 2 83 3 S4 H4 $5

OR 25-35° .7-1.3 34-47° 1.2-2.2 46-57° 1.8-3.8 58-65° 2.1-7.3 51-57°
».¢ ene leis SADA il 43 51.7° 3.2 63.0° Sail Sige

Discussion and, diagnosis —Similarities in rib number,
type of ornamentation, dental pattern, and general shape
make it fairly certain that V. (C.) coloradonis texalana is
a direct descendant of V. (C.) linguinodifera of the early
Eocene. Intermediate forms are unknown, and there exists
a gap of some dimension between the deposition of the Bashi
and the Weches Formations.

This subspecies is only of local significance in Texas,
and Renick and Stenzel (1931, p. 105) noted the Weches
bed at Burleson Bluff as 0 to 20 feet above the base. Two
forms appear in the middle Weches: V’. (C.) coloradonis
coloradonis, a descendant subspecies which is statistically
comparable to the lower Weches form except in number of
ribs, and V. (C.) natchitoches which has fewer ribs and
modifications in shape, but could only have been derived
from V. (C.) coloradonts texalana.

The relationship by Gardner (1927) of this form to J.
(C.) trapaquara of the stratigraphically higher Stone City
Beds in Texas is much more realistic than the association by
Harris’ (1919) of the descendant subspecies to V. (R.)
rotunda, because dental pattern and shell form are some-
what similar among claibornicards but distinct from these
features in Rotundicardia.

Comparison between slope of reduced major axes of the
V. (C.) coloradonis texalana sample from Burleson Bluff

102 PALAEONTOGRAPHICA AMERICANA (VI, 39)

(USGS 5288) and the sample of V. (C.) coloradonis color-
adonis from Saline Bayou, Louisiana (USGS 3569), pro-
duced a correlation coefficient (Z) of .383, indicating high
degree of similarity.

Etymology.—Subspecific name texalana geographic in
honor of Texas, the state in which this form is found.

V. (Claibornicardia) natchitoches Harris, 1919

Plate 25, figures 4a-4c, 5a, 5b, 6, 7a-7¢e

V. natchitoches Harris, 1919, Bull. Amer. Paleont., vol. 6, No. 31, p.
82, pl. 30, figs. 10, 13-16; Plummer, 1933, Univ. Texas, Bull.
3232, pp. 811, 812, pl. 8, figs. 6a, 6b; Palmer and Brann, 1965,
Bull. Amer. Paleont., vol. 48, No. 218, p. 336.

Distinguishing characters —Similar in shape and general
dental pattern to other claibornicards but distinguishable
from any other species on following bases: extremely low
rib count (OR 14-17), ribs somewhat low and thin, central
cord highly irregular in arrangement of widely spaced, sub-
dued nodes. Paracostals low, rounded and vestigial in some.
Ribs broadly spaced and equally wide intercostals broadly
U-shaped. Posterior pedal retractor scar removed from its
normal position as triangular projection from posterior dor-
sal corner of posterior adductor scar, occupying an isolated
position just dorsal to that corner.

Type—Lectotype (herein designated), a left valve
with incomplete anterior-ventral margin, PRI 666. Para-
lectotype, a hinge fragment of right valve with well-pre-
served dentition, PRI 664. Dimensions below.

Sp’n Ts H YW RN’ BL/L BH/H H/L %W/H
Lect. 19.0* 18.4 Todt 17 * 89 * 42
Para.**

*Length restored, ratios based on this meaningless, since length
estimated on basis of ratio data from other samples.

**Paralectotype fragment 14.2 mm. in length, original length of
individual on order of 21 mm.

Type locality —“Natchitoches, Louisiana. Lower Clai-
borne.” Probably in Weches equivalent.

Distribution. — Harris’ collection of syntypes was
examined, as well as collections from USGS 13274, 14 mile
north of San Augustine P.O., San Augustine County, Texas;
USGS 2914 (USNM collection 147496), Provencal, Natchi-
toches Parish, Louisiana; ANSP 9915, San Augustine, Texas.
Plummer (1933) listed the San Augustine localities as

Weches.

Stratigraphic occurrence. — Weches Formation in Texas
and equivalents in Louisiana, middle Eocene age.

Dimensional summary.—

Sample: USGS 13274, San Augustine, Tex.

N77, Dimension
Statistic L H UW RN
OR 14.8-21.2 15.0-20.3 6.2-8.5 14-17
X 17.3 17.2 7.2 15.8
S 2.4 2.6 1.0 1.0
‘ee 13.8 15.1 12.5 6.3
OX 9 1.0 3 4+
Ratio
; BL/L BH/H H/L 44W/H
OR -67-.82 .83-.89 91-1.07 39-46

74 85 1.00 42

Curvature characteristics*—

Sample: USGS 13274 N= 7
Si Hi SBis. Be $3 H3 S4 H4 $5

OR 18-28° .7-1.8 32-43° 1.7-4.3 45-63° 3.0-12.5 56-68° 5.8-11.5 54-61°

x: OO ae Tht ROS © 210 53.4° 6.4 63:30 ee Lyi

*Curvature of two types, A and B, $5 only in B types. Specimen
nos. 2 and 5 with A pattern are highly inflated, probably females.
Others with B pattern probably males.

Discussion and diagnosis —V. (C.) natchitoches illu-
strates a reversal in the trend toward higher costal number
common in the early members of Claibornicardia, and the in-
ference is descent from V. (C.) coloradonis texalana of the
lower Weches. This subspecies has a somewhat low costal
number (19-22) but lower than that of V. (C.) coloradonis
coloradonis which appears in the middle Weches simultane-
ously with V. (C.) natchitoches.

Besides the distinct ornamentation, this species exhibits
a character which is not typical of Venericardia, or viewed
in any other species of the present investigation: the re-
moval of the posterior pedal retractor scar from its normal
union with the posterior adductor. This radical departure
might lead some taxonomists to create a new subgeneric
category, but there is no evidence that the species gave rise
to any later forms, despite Harris’ (1919) statement that
“It would seem quite probable that this is the form from
which alticostata was derived . . .,” and it is treated here
as a variant within Claibornicardia.

It is possible that the expanded individuals in the
measured samples having convex A type curvature are fe-
males (PI. 25, figs. 6,7a,7b), and the flattened forms with
B curvature are males (same PI., figs. 5a,5b), as is the case
in V. (C.) alticostata and some rotundicards. Ornamental
distinctiveness is not seen in V. (C.) natchitoches as it is
in those forms, however.

Etymology—tThe trivial name natchitoches is geo-
graphic referring to the type locality in Natchitoches,
Louisiana.

AMERICAN ALTICOSTATE VENERICARDS: EHLEASLIP 103

V. (Claibornicardia) perantiqua Conrad, 1865
Plate 25, figures 8-11

Cardita subquadrata Gabb, 1860, Acad. Nat. Sci. Philadelphia,
Jour., 2d ser., vol. 4, p. 303, pl. 48, figs. 22a, 22b (by error on
pl. 21); Gabb, 1861, Amer. Philos. Soc., Proc., vol. 8, p. 105;
Lea, I., 1861, Acad. Nat. Sci., Philadelphia, Proc. p. 150;
Meek, 1864, Smithsonian Misc. Coll., 8, No. 177, p. 11.

Non C. subquadrata Conrad, 1847.

Vencricardia perantiqgua Conrad, 1865, Amer. Jour. Conch., vol. 1,
p. 8; Dall, 1903, Wagner Free Inst. Sci., Trans., vol. 3, No. 6,
pp- 1423, 1429; Harris, 1919, Bull. Amer. Paleont., vol. 6, No. 31,
pp. 84, 85.

Cardita perantiqua (Conrad), Whitfield, 1885, U.S. Geol. Sur., Mon.,
vol. 9, pp. 232-234, pl. 30, figs. 8-10, (reprinted, 1886, in New
Jersey Geol. Sur., Paleontology of the Cretaceous and Tertiary,
same page, plate references). :

Cardita brittoni (?) Whitfield, 1885, of. cit., pp. 233, 234, pl. 30,
figs. 11, 12 (see also op. cit. 1886).

Venericardia (?) perantiqua Conrad, Palmer and Brann, 1965,
Bull. Amer. Paleont., vol. 48, No. 218, p. 337.

Distinguishing characters. — General shape similar to
that of V. (C.) coloradonis and other claibornicards, but
beaks extremely small and umbones low much as those in
V. (C.) trapaquara Harris. Similar only to V. (C.) natchi-
toches in ornamentation, in which thin central costal cords
are irregularly noded, with nodes somewhat subdued, point-
ing toward ventral margin. Ribs broadly based and subdued,
paracostals threadlike forming margins for broadly U-shaped
intercostals. Paracostal development somewhat stronger than
that of V. (C.) natchitoches, and costal number averages
much higher than in that species.

Type. — Conrad (1865) did not name or figure a type
specimen, and Gabb’s type of “Cardita subquadrata” from
ANSP collection 19373, external mold of large right valve,
dimensions below, is lectotype (herein designated). Dimen-

sions approximate.

Sp’n L H %W RN BL/L BH/H H/L %W/H
Lect. 25.0 22.5 5.0 20 58 89 -90 22

Type locality. — Burlington County, New Jersey.

Distribution. — Besides Gabb’s ANSP collection 19373,
the AMNH collections 9273/2 and 467 from Farmingdale,
Monmouth County, New Jersey, were examined.

Stratigraphic occurrence.—Shark River Formation,
middle Eocene age.

Dimensional summary.—

Sample: Composite, ANSP 19373, AMNH 9723/2, 467

N=12 Dimension*

Statistic 1G H 4w** RN
OR 21-27 19-26 4-7 16-22
aX 24.0 21.6 5.5 19.3
S 9 2.0 £9 1.9
Vi 7.8 9.1 16.0 9.7
Ox 5 6 2 5

*Measurements to nearest millimeter because of poor preservation,
distortion.

Sample: Composite, ANSP 19373, AMNH 9723/2, 467 (continued)

N=12
Statistic
Ratio
BL/L BH/H H/L Y%W/H**
OR .63-.77 90-.95 84-100 “.18-.30
xX oA 91 90 25

**Semi-width values highly inaccurate, much too low.

Curvature characteristics*. —

Sample: ANSP 19373 N= 1
$1 H1 $2 H2 $3 H3 S4 H+ SS
OR - — 26° — 47° — 60° — 40°

a *Distortion of specimens made them unfit for curvature analysis.
Single specimen measured for qualitative values.

Discussion and diagnosis. — Unfortunately, V. (C.)
perantiqua is known only from natural molds and casts in
the glauconitic Shark River Marl and specimens are found
in great profusion almost to the exclusion of other mollus-
can species. The external ornamentation and shell interiors
and dentition can be restored only by plastic or latex casts,
and the fine details of dentition and musculature cannot be
extracted.

The name used by Gabb was preoccupied by Conrad
(1847) for a species in the McBean Formation in the
Orangeburg District in South Carolina. Conrad also de-
scribed it as “Cardita.” On this basis, Gabb’s form, known to
belong to Venericardia, might legally retain the trivial name
“subquadrata.” Conrad’s type specimen for the Orangeburg
form is unlocated and his poor description and illustration
do not resolve the generic placement, but it is highly prob-
able that it will ultimately be placed in Venericardia, and
his name V. perantiqua is taken as the valid one for the
Shark River species. Because Conrad did not designate or
figure a type specimen, Gabb’s figured specimen is chosen
as the lectotype.

V. (C.) perantiqua is not related to V. (R.) rotunda
Lea as Whitfield (1885) believed. Both ornamentation and
dentition tie this species to Claibornicardia and because of
similarities in ornamentation, it is most closely related to
V. (C.) natchitoches of the Texas Weches, and undoubtedly
also evolved from V. (C.) coloradonis. The posterior ad-
ductor scar is faintly preserved on molds of V. (C.) per-
antiqua, and it is not known if the posterior pedal retractor
became independent of the posterior adductor, as in V. (C.)
natchitoches.

V. “brittoni” (Whitfield) is known only from highly
distorted molds, and undoubtedly represents only crushed
specimens of V. (C.) perantiqua, and the “finer” quality of

104 PALAEONTOGRAPHICA AMERICANA (VI, 39)

ribbing in former species is the result of dorso-ventral com-
pression.

The occurrence of V. (C.) perantiqua in Monmouth
County, New Jersey, represents the northernmost record
of an alticostate venericard on the east coast of North
America during the Eocene expansion of this group, and is
the last known species of Claibornicardia on that coast. A
mold V. (R.) rotunda is present in the AMNH collection
9723/2 and apparently lived sympatrically with this species.

Etymology. — Trivial name perantiqua from the Latin
per, very, and antiqua, ancient, in possible reference to its
original Cretaceous stratigraphic placement by Gabb (1860).

V. (Claibornicardia) trapaquaroides, n. sp.
Plate 24, figures 7a-7¢

Distinguishing characters. — Similar in general shape,
ornamentation and dental pattern to V. (C.) trapaquara but
distinguishable from that species on much greater degree of
inflation; V. (C.) trapaquara is highly compressed, atypical
of Claibornicardia. Higher, more inflated beaks than that
species which has low, tiny beaks and umbones. Postero-
ventral elongation much less in this species and hinge line
relatively shorter than that of V. (C.) trapaquara. Regularly
spaced transverse ridgelike nodes well developed only on an-
terior of shell; vestigial on posterior. Intercostals V-shaped,
paracostals reduced, rounded.

Type. — Holotype, a right valve, USNM 644320, di-

mensions below. Monotypic.

Sp'n L H %W RN BL/L BH/H H/L %W/H
Holo. 24.8 25.3 9.2 30 74 88 1.02 36

Ty pe locality. — USGS 9255, Percilla, Houston County,
Texas. Weches Formation, middle Eocene age.

Distribution. — Known only from type locality.

Stratigraphic occurrence. — Weches Formation, middle
Eocene age. Stenzel, Krause, and Twining (1957, p. 147)
noted the Tyus Member of Weches at a locality .67 mile
west of Percilla post office.

Dimensional summary. — See type dimensions.

Curvature characteristics. —

Sample: Holotype
S1 H1 $2 H2 $3 H3 S4 H4+ $5
OR 25° 816 By 3.4 S12 5.1 61° 10:2" 453

Discussion and diagnosis. — Despite the somewhat in-
exact stratigraphic placement of V. (C.) trapaquaroides, its
Weches occurrence is certain, even though the exact level
in that formation is not known. If this species is found in
the Tyus (lower) Member, the subdued costal nodes and
rounded paracostals indicate great advance over the more

primitive features of its nearest relative and probable an-
cestor in the Weches, V. (C.) coloradonis coloradonis. The
increase in costal number over that subspecies indicates
an evolutionary trend from the primitive subspecies in the
lower Weches, V. (C.) coloradonis texalana, and a similar
number is found in descendants of V. (C.) trapaquaroides,

The high degree of postero-ventral elongation directed
mainly toward the ventral, as well as similarities in orna-
mentation, indicate this species as the ancestor of V. (C.)
trapaquara of the stratigraphically higher Stone City Beds
in Texas, and V. (C.) complexicosta of the “Wautubbee
Marl” which is similar in shape but indicates an even greater
degree of costal node simplification.

Etymology. — The trivial name trapaquaroides means
“trapaquara-like” in reference to its resemblance to that
species.

V. (Claibornicardia) trapaquara Harris, 1895

Plate 24, figure 8; Plate 25, figures la, 1b, 2, 3a, 3b

V’. trapaquara Harris, 1895, Acad. Nat. Sci. Philadelphia, Proc. 1895,
p. 48, pl. 1, fig. 7; Harris, 1919, Bull. Amer. Paleont., vol. 6,
No. 31, pp. 81, 82, pl. 30, fig. 8 (mon figs. 6, 7, 9); Harris and
Palmer, 1946, Bull. Amer. Paleont., vol. 30, No. 117, pp. 68, 71.

V’. (Claibornicardia) trapaquara Harris, Stenzel, Krause, and Twin-
ing, 1957, Univ. Texas, Pub. 5704, pp. 106, 113-115, pl. 14, figs.
6-11; Palmer and Brann, 1965, Bull. Amer. Paleont., vol. 48, No.
218, p. 345.

V’. n. sp., Renick and Stenzel, 1931, Univ. Texas, Bull. 3101, p. 104.

Similar to V. (C.) trapa-
quaroides of the Weches in ornamentation, having tripartite
costae with central cord regularly beaded with smooth,
rounded nodes only slightly lingulate on anterior ribs. Para-
costals rounded in cross-section, restricting intercostals
which become wider and U-shaped only toward posterior.
Convexity much lower, beaks and umbones much less in-
flated than in that species, and hinge margin straighter and
longer than in that species.

Type. — Holotype, a right valve, BEG 20502, highly
eroded. Dimensions below.
Sp'n E H %W RN BL/L BH/H H/L %W/H
Holo. 11.8 11.4 3.8 28 oA 95 oF, 33

Type locality. — BEG 197-T-3, Cedar Creek, southeast
corner of Wheelock League, 200 yards north of the Brazos
County Line, Robertson County, Texas.

Distribution. — Besides the holotype, a collection of
three valves and a fragment from BEG 26-T-1, Stone City
Bluff on the Brazos River, Burleson County, Texas, were
examined. Stenzel, Krause, and Twining (1957) also re-
ported this species from BEG 113-T-36, on the Trinity
River, .85 mile above Alabama Ferry, Houston County,
Texas.

Stratigraphic occurrence.—Stone City Beds, middle
Eocene age.

Distinguishing characters.

a

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 105

Dimensional summary. —
Sample: BEG 20502 (Holo.), 20803, 20499 (26-T-1)

N= 3 Dimension
Statistic L H 14W RN
OR 11.8-18.0 11.4-17.2 Beg=Se7, 27-29
we 15.6 14.9 5.0 28.0
s 3.3 3.1 1.0 1.0
Vv 21.4 20.8 20.5 3.6
Ox 1.9 il 6 6
Ratio
BL/L BH/H H/L 14W/H
OR 671-377 94-.97 90-1.01 33
xX 73 95 96 33
Curvature characteristics. —
Sample: 20502, 20803, 20499 N=

S1 H1 S2 H2 $3 H3 S4+ H+ S5
OR = 22-30° .8-1.2 40-45° 1.3-2.0 45-51° 4.3-5.4 45-49° 7.6-8.7 43-47°

XGezorie 00 <4310"' 1:6 47.7° 5.0 47.7° 8.1 44.3°

Discussion and diagnosis. — Because of the high degree
of similarity between V. (C.) trapaquaroides and V. (C.)
trapaquara, especially in characters of ornamentation, the
former species is the most likely ancestor of the latter, but a
high degree of specialization is indicated in the Stone City
form in its flattened shell and sharp, trapezoidal outline.

The relationship of this species to the “alticostata
group” recognized by Harris (1919) and its association with
Claibornicardia by Stenzel and Krause (1957) indicate the
morphological unity among all species of the subgenus. How-
ever, the highly specialized form of V. (C.) trapaquara
seems to preclude it as a direct ancestor of V. (C.) com-
plexicosta of the Wautubbee, despite Stenzel and Krause’s
(1957) conclusion to that effect. A much more logical an-
cestor is V. (C.) trapaquaroides of the Weches with shell
form and inflation similar to that of the Wautubbee species.
V. (C.) trapaquara was probably derived from that species,
as well.

Etymology. — The trivial name trapaquara is derived
from the Latin calcitrapa or trapa, a four-pointed weapon
placed on the ground to impede cavalry, and quattuor, four,
in reference to its four-pointed or trapezoidal shape.

V. (Claibornicardia) complexicosta Meyer and Aldrich, 1886
Plate 26, figures 3a-3c, 4a-4c, 5

V. complexicosta Meyer and Aldrich, 1886, Cincinnati Nat. Hist.,
Jour., vol. 9, No. 2, p. 45, pl. 2, figs. 21, 21a; Dall, 1903, Wagner
Free Inst. Sci-., Trans., vol. 3, No. 6, pp. 1424, 1429; Harris, 1919,
Bull. Amer. Paleont., vol. 6, No. 31, pp. 84, 85, pl. 31, fig. 5.

V. (Claibornicardia) complexicosta} Meyer and Aldrich, Stenzel,
Krause, and Twining, 1957, Univ. Texas, Pub. 5704, pp. 111-114,
pl. 15, figs. 1-4; Palmer and Brann, 1965, Bull. Amer. Paleont.,
vol. 48, No. 218, p. 326.

Distinguishing characters.— General shape, outline,
and convexity similar to that of V. (C.) trapaquaroides, with

delicate, elongate hinge much like V. (C.) trapaquara but
distinguishable from all other species by extreme develop-
ment of ventrally pointing postero-ventral process of shell.
Extreme simplification of ornamentation, in which posterior
and central ribs have lost almost all vestiges of costal nodes
and appear smooth. Nodes on anterior ribs relatively widely
spaced and somewhat irregular, being much less conspicuous
than same features on other species. Paracostal cords incon-
spicuous on posterior ribs.

Type. — Holotype, a left valve formerly JHU 1362, di-
mensions below, now USNM 645940.

Sp’n L H %W RN BL/L BH/H H/L %W/H
Holo. 8.9 Sal 4.3 31 65 82 1.02 47

Type locality. —“Wautubbee” (Meyer and Aldrich,
1886). Stenzel, Krause, and Twining (1957) gave more exact
location: “Railroad cut about 1 mile north of the depot at
Wautubbee, or about .4 mile north of the overpass of U. S.
Highway 11 (Laurel-Meridian road) over the Southern Rail-
road, northwestern Clarke County, Mississippi.” Wautubbee
Formation, middle Eocene.

Distribution —Besides the type, collections from the
following were examined: BEG Miss-51, shoulder and ditch
on the north side of the public gravel road connecting Deca-
tur with Conehatta, 7.8 miles west of the intersection with
State Highway 15 at Decatur, and 0.4 mile west of a cross-
roads, southeast corner of SW 14 of NE 1% of section 3,
T7N, R 10 E, Newton County, Mississippi, Archusa Mem-
ber, ‘““Wautubbee Marl” middle Eocene. Stenzel, Krause,
and Twining (1957) noted the occurrence of this species
from Mt. Lebanon, Bienville Parish, Louisiana (ANSP
9467), and BEG 11-T-70 in Bastrop County, Texas.

Stratigraphic occurrence——Archusa Marl Member of
“Wautubbee Marl,” late middle Eocene age. Stenzel, Krause,
and Twining (1957) did not explicitly date the exposures
at BEG 11-T-70 to Bastrop County, Texas, except as “Cook
Mountain,” which is equivalent to the Wautubbee in Mis-
sissipp1. The ANSP Mt. Lebanon locality in Louisiana is
also Cook Mountain in age.

Dimensional summary.—

Sample: BEG Miss-51, 20644, 20645, Holotype JHU 1362

N= 6 Dimension
Statistic L H YW RN
OR 8.9-20.5 9.1-20.2 4.3-8.0 29-31
xX 15.5 14.7 6.2 30.3
S 4.6 4.1 15 38
V 29.5 27.9 24.9 mals
Ox 1.9 1.7 6 4
Ratio
BL/L BH/H H/L Y4W/H

R .62-.72 -82-.88 .98-1.05 -38-.47
67 85 1.01 40

106 PALAEONTOGRAPHICA AMERICANA (VI, 39)

Curvature characteristics.—

Sample: BEG Miss-51, 20644, 20645, Holotype JHU 1362 N= '6

S1 H1 S2 H2 $3 H3 S4 H+ $5

nN

OR 21-34° .8-1.9 35-50° 1.7-2.9 43-56° 2.7-5.2 57-62° 6.1-11.8 50-56°
x 270ce 1:4. 40:82 eo: 50.0° 6 59.8° 8.8 53.3°

Discussion and diagnosis. — Until the publication of
Stenzel, Krause, and Twining (1957) who discussed con-
specific forms from the vicinity of the type locality, V. (C.)
complexicosta was known only from the holotype which, as
those authors suggested, is an immature specimen. This is
substantiated by the absence of the terminal, less convex
spiral segment of the B pattern typical of Claibornicardia
(see Text-fig. 14). Another specimen of size comparable to
the holotype (BEG 20645), 11.9 millimeters in height, also
lacks this terminal segment, but the larger specimens all
contain it in their curvature patterns.

The most logical ancestral species is V. (C.) trapa-
quaroides of the Weches, and not, as suggested by Stenzel,
Krause, and Twining (1957), V. (C.) trapaquara of the
Stone City Beds. This latter species, while exhibiting some-
what simplified ornamentation, is much too specialized in
its trapezoidal outline and low convexity. V. (C.) trapa-
quaroides is decidedly similar in outline and convexity to
V. (C.) complexicosta besides haying simplified ornamenta-
tion which is further simplified in that species. Despite the
absence of stratigraphically intermediate forms, this rela-
tionship seems more reasonable than that proposed by those
authors.

V. (C.) complexicosta probably did not give rise to J.
(C.) alticostata of the overlying Gosport Sand because of
the high development of costal nodes in certain members of
this species, and a much more generalized ancestor such as
V. (C.) coloradonis coloradonis, also living in the “Wautub-
bee Marl” is much more likely. The highly simplified ribs
of V. (C.) complexicosta seem to indicate that this species
is an evolutionary end product without descendants.

Etymology.—The trivial name complewicosta is derived
from the Latin complexus, embrace or entwine, and costa,
rib in allusion to the tripartite costals on the anterior of the
shell in this species.

V. (Claibornicardia) blandingi Conrad, 1830
Plate 26, figures la-1c, 2

V. blandingi Conrad, 1830, Jour. Acad. Nat. Sci., Philadelphia, 1st
ser., vol. 6, p. 229, pl. 9, fig. 20; Dall, 1903, Wagner Free Inst.
Sci., Trans., vol. 3, No. 6, p. 1424; Harris, 1919, Bull. Amer.
Paleont., vol. 6, No. 31, pp. 86, 87, pl. 31, figs. 11, 11a.

V. (cf. Claibornicardia) blandingi Conrad, Palmer and Brann, 1965,
Bull. Amer. Paleont., vol. 48, No. 218, p. 324.

Distinguishing characters—Somewhat similar to V.
(C.) alticostata in general shape and inflation, and strongly
resembles the “sillimani” variants of that species in costal
ornamentation, in which terrace-like paracostals defined by
sharp threads and developed over most of the shell, vestigial
on the posterior-most ribs. Costal cord on bladelike ridge,
ornamented by transversely expanded nodes which are regu-
larly and closely spaced on the anterior ribs, but become
less regular and more sawtooth-shaped on central and
posterior ribs. Distinguished from V. (C.) alticostata on
basis of extremely small size but known to be adult on
basis of mature curvature pattern of the B type. Relative
roundness of posterior margin and lack of well-defined
truncation also distinguish it and rib count (23-25) much
lower than that species (25-31).

Type.—Lectotype (herein designated), a right valve,
paralectotype a left valve, both silicified, both ANSP No.
30500.

Sp’n L H %W RN BL/L BH/H H/L YW/H
Lect. 15.3 14.9 5-7 23 69 89 97 38
Para. *14.8 13.3 5.1 25 68 -87 90 38

* Restored.

Type locality—Vance’s Ferry, Orangeburg County,
South Carolina. Present town of Vance is about 41% miles
northwest of Eutawville, along the south shore of Lake
Marion (Santee Reservoir). Vance’s Ferry was a landing on
the Santee River in Conrad’s day. The site was inundated
when the present lake was formed during construction of
the Santee-Cooper dam.

Distribution—Known only from type locality.

Stratigraphic occurrence. — Lower Santee Limestone,
middle Eocene age.

Dimensional summary.—See type dimensions.

Curvature characteristics. —

Sample: Lectotype and paralectotype NS
Si* Hil $2 H2 $3 H3 S4 H4 $5
OR ? ? 38-40° 1.2-1.5 45-46° 3.1-3.2 60-61° 5.3-5.6 53-56°

x ? ? 39.0° Tem 4b5e 3.1 “6052 5.5% 54a

*Smallest stage not apparent in either valve. May be attributable
to poor preservation of umbonal ornamentation.

Discussion and diagnosis.—Harris (1919) listed V. (C.)
blandingi under “ill-defined and unrecognizable species of
Venericardia described by Conrad,” and associated it with
the “St. Maurice” (lower Claiborne) horizon. He recognized
its similarities to V. (C.) alticostata, whereas Dall (1903)
erroneously associated it with V. (R.) rotunda. The poorly
preserved dental pattern and the clearly tripartite orna-
mentation reinforce Harris’ conclusion, although the relative

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 107

rounding of the posterior margin is not characteristic of
Claibornicardia. The broken posterior on the paralectotype
produces a spurious straight margin, but the complete lecto-
type shows little truncation. Both specimens were poorly
figured by Conrad (1830).

H. S. Johnson, Jr., State Geologist of South Carolina
(personal communication, 1962) indicated that recent in-
vestigation has established that the Vance’s Ferry locality
is in the lower Santee Limestone and “‘probably middle mid-
dle or lower middle Claiborne in age.”

It is likely that this species, as well as V. (C.) alti-
costata, was derived from V. (C.) coloradonts coloradoms,
although a direct relationship between V. (C.) blandingi
and V. (C.) alticostata is also possible. If this latter were
the case, it would imply the origin of V. (C.) alticostata on
the East Coast, rather than the vicinity of the type locality
in Alabama.

Etymology.—The trivial name blandingi is patronymic
in honor of Dr. Blanding of Camden, South Carolina, who
collected Conrad’s specimens.

V. (Claibornicardia) alticostata (Conrad), 1833
Plate 26, figures 8a-8c, 9, 10a, 10b, 11

Cardita alticostata Conrad, 1833, Amer. Jour. Sci. ser. 1, vol. 23, p.
342 (Jan.).

Venericardia transvcrsa Lea, 1833, Contributions to Geology; Ter-
tiary formation of Alabama, pp. 68, 69, pl. 2, fig. 46 (Dec.).

Vl’. sillimani Lea, 1833, ibid., pp. 69, 70, pl. 2, fig. 47.

V. (Claibornicardia) alticostata (Conrad), Stenzel, Kraus?, and ‘Twin-
ing, 1957, Univ. Texas, Pub. 5704, pp. 104, 106, 107-109, pl. 13,
figs. 1-9; pl. 14, fig. 5, text figure 17; Palmer and Brann, Bull.
Amer. Paleont., voi. 48, No. 218, p. 321.

V’. (Claibornicardia) sillimani Lea, Stenzel, Krause, and ‘Twining,
1957, ibid., pp. 106, 107, 109, 110, 111, pl. 14, figs. 1-4.

Complete synonymy of this species in Stenzel, Krause, and Twining,
1957, op. cit., loc. cit.

Distinguishing characters—Certain individuals resem-
ble V. (C.) blandingi, in general form and ornamentation
but distinguished from that and all other claibornicard
species by size which is much larger than any other described
form. Some individuals measure over 50 millimeters in
length. High degree of variation in lateral outline from
prominent postero-ventral elongation to direct posterior
elongation. Well-developed posterior truncation throughout,
lacking in V. (C.) blandingi. High degree of variability in
ornamentation from bladelike central costal cords with well-
developed paracostals on central and posterior of shell to
simplified, inverted V-shaped almost anodulous ribs. An-
terior costals similar in both extremes, being broadly based
with wide, rounded paracostals and closely and regularly
spaced costal nodes, although intercostals more restricted
in those forms with simplified central and posterior costae.

Dental plate gently arched and much more elongated than
in V. (C.) blandingt. Variable development of left anterior
cardinal (2) and variable development of right anterior
cardinal (3a) which often hooks around point of anterior
socket, continuing as callus deposit bordering anterior mar-
gin of socket, resembling this feature in Baluchicardia.
Type. — Conrad’s syntype collection, ANSP 30562,
recently reported by Moore (1962) cannot be located at
the present time, and it would be desirable to choose a
lectotype from the collection, rather than a neotype. The
following data for Lea’s figured types of V. transversa and
V. sillimani are presented, but not with neotype designation.

Spn L H %WRN BL/LBH/H H/L 4W/H

ANSP 5253) 47.7 39:0 7-7 (29 80 87 82 45° transversa.
AINSP)5258/36.5) S152) 15¢8) 288 75) 8286) | 685 44 sillimani.
5259

Type locality —Claiborne Bluff, on the east bank of
the Alabama River near the town of Claiborne, T 7 N,
R 5 E, Monroe County, Alabama. Gosport Sand, upper
middle Eocene age.

Distribution—Lea’s collections of V. transversa, ANSP
5253-5257 and V. sillimani, ANSP 5258-5266 were examined,
as well as collections in AMNH collections (9950/1, 1055,
984, topotypes). USGS 6087, Gopher Hill on Tombigbee
River, about 14 mile above St. Stephens Bluff, Washington
County, Alabama; USGS 2009, 2 miles west of Orangeburg
courthouse, Orangeburg County, South Carolina (McBean
Fm.); and “Eocene of New Bern, North Carolina” (precise
information withheld at request of the U.S. Geological
Survey ).

Stratigraphic occurrence-—Gosport Sand in Alabama
and McBean Formation equivalents in South Carolina;
Eocene at New Bern, North Carolina. Late middle Eocene
age.

Dimensional summary.—

Sample: AMNH 9950/1, topotype, V. (C.) alticostata

N = 34 Dimension
Statistic L H UW RN
OR 9.9-65.7 8.6-51.7 3.1-23.3 25-30
xe 40.9 35.3 15.6 27.9
S 13.2 11.0 54 1.0
V 32.3 31.1 5.9 3.7
Ox 2.3 1.9 9 2
Ratio
BL/L BH/H H/L 4W/H
OR 64-.97 80-.95 .78-.94 31-.44

x 74 86 87 38

108 PALAEONTOGRAPHICA AMERICANA (VI, 39)

Sample: AMNH 9950/1

Topotype, V’. sillimani

N=8 Dimension
Statistic 1 H ZW RN
OR 15.7-53.2 14.1-46.7 5.2-18.4 29-31
xa 31.6 28.1 10.9 29.9
S 14.3 12.6 Sail 38
V 45.3 44.8 46.5 2.8
Ox Sal 4.5 1.8 3
Ratio
BL/L BH/H H/L 1,W/H
OR .65-.74 .83-.90 .88-.93 34-.41
x 69 .87 89 39
Curvature characteristics —
Sample: AMNH 9950/1, V. alticostata N=34
S1 H1 $2 H2 $3 H3 S4 H4* S$5*

OR 10-39° .7-4.7 22-51° 2.8-9.4 45-55° 8.6-21.2 52-64° 23.4-44.6 63-81°
me P5902. 21) 0.39)0> enGss OME Suyn SO Am GOES 70.4°

*Gerontism.

Sample: AMNH 9950/1 V. sillimant. N=8
S1 H1 $2 H2 $3 H3 S4+ H4+ $5

OR 25-42° 1.1-2.5 41-60° 1.9-8.6 45-59° 2.6-14.6 54-64° 4.2-19.3 48-53°
xX Ba ee ON ara 33.9 52.3° 6.4 60.1° 12.4 50.4°

Discussion and diagnosis —Two similar species of the
Gosport Sand have been recognized: V. (C.) sillimani Lea,
characterized by bladelike tripartite costals much like those
of V. (C.) blandingi, a relatively thin shell, well-developed
right anterior cardinal (3a) in line with the point of the
delicately arched right medial cardinal 3b. V. (C.) alticos-
tata, conversely, has been characterized by central and
posterior costae with an inverted V cross-section, absence
of paracostals on these ribs, and a rounded, irregularly
noded central cord as well as a relatively thick shell with in-
flated umbones. A somewhat heavier medial right cardinal
and highly variable development of the right anterior car-
dinal (3a) also characterize this form. The former charac-
ters are primitive, and are seen in many ancestral claiborni-
cards (e.g. V. (C.) coloradonis coloradonis); the latter,
especially the ornamentation, are advanced characters.
Stenzel, Krause, and Twining (1957) recognized distinct
species, but commented (p. 111) “if they are in precisely
the same stratigraphic level, it would not be surprising for
these two forms to be variants of one natural species,” but
that “. . . no true intergrades have been seen by us.” Inter-
grades have been observed in many characters and ratios
in the present investigation, as well as “mixtures” of charac-
ters in the two forms. Among Lea’s type collection of nine
specimens of V. (C.) sillimani, only one, ANSP 5260 (simi-

lar to PI. 26, fig. 11), could be taken for that species on the
basis of the above criteria. All others (including the figured
holotype, figs. 10a, 10b, same Plate) could objectively be
called V. (C.) alticostata. The same assortment is found
among specimens of V. transversa, and some resemble the
“silimant” form.

Another distinction, not considered by previous authors,
is in pattern of curvature. Only eight individuals could be
selected from hundreds of topotype valves in the AMNH
collections which possessed all the primitive attributes of
V. silimant. These eight all had curvature of the B type,
while a sample of 34 individuals with the advanced charac-
teristics of V. (C.) alticostata showed curvature of the A
pattern, which is noted only in the baluchicards and some
rotundicards, and in the claibornicard V. (C.) natchitoches
Harris which is highly specialized in other characters as
well. The A pattern is also found in V. (C.) nasuta Dall, a
descendant of V. (C.) alticostata.

Possible reproductive implications of the two patterns
of curvature are discussed in a previous section, and only
three species other than this, in the present investigation,
were found to consist of mature individuals exhibiting both
patterns in the same population. The conclusion of sexual
dimorphism in the case of V. (R.) rotunda might also apply
to V. (C.) alticostata, in which the more primitively ribbed,
less convex specimens having B type curvature are male,
and the highly convex, but more simply ornamented forms
with A curvature are females, highly specialized for the
“marsupial” function in reproduction. The great disparity in
numbers between the two forms might not support this con-
clusion, and the ancestral B pattern of curvature may simply
be linked to the genetic combination of other primitive char-
acters.

Other evidence of two sorts may reinforce the decision
to unite V. (C.) sillimani and V. (C.) alticostata. (1) Com-
parison of reduced major axis slopes between the two sam-
ples mentioned above, and between Lea’s type collections
of V. transversa and V. sillimani result in a high degree of
correlation between the two forms. (Z = .482 in the latter
case, 1.800 in the former — see fig. 6). (2) The second line
of evidence also supports the hypothetical sexual dimorph-
ism in the species. A sample from Gopher Hill, Washington
County, Alabama (USGS 6087), west of Claiborne, was
found to contain both patterns of curvature, but only orna-
mentation and dentition of the “sillimani” type, and lacking
the simplified ornamentation of the “alticostata” type
found at the type locality No statistically significant dif-
ference was found between these and the topotype samples.
Although the Gopher Hill locality is listed as “Gosport” by

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 109

the Alabama Geological Survey, it may represent a slightly
lower stratigraphic layer of a transgressive deposit in which
a primitive population of V. (C.) alticostata had only begun
to specialize toward female distinctiveness found fully de-
veloped at the type locality and other deposits of similar
age. Specimens from Orangeburg, South Carolina, and New
Bern, North Carolina, are indistinguishable from topotype
specimens. Similarly, V. (R.) rotunda found at the Gopher
Hill locality with both curvature patterns represented ex-
hibits only the simpler, primitive ornamentation (funicu-
late) of stratigraphically lower samples found in the Wau-
tubbee in Mississippi.

The only known descendant of V. (C.) alticostata is
V. (C.) nasuta Dall of probable middle Oligocene age. It is
enigmatic, however, that no representative of this robust
lineage is reported from rocks of Jacksonian (upper Eocene )
or early Oligocene age, but the general transition from the
sandy, glauconitic marl to dominantly limestone facies in
the Jackson rocks of the Atlantic and Gulf Coasts may
have influenced post-Gosport distribution of the immediate
descendants of V. (C.) alticostata. A possible exception may
be poorly preserved Castle Hayne forms reported by Kellum
(1926).

Etymology.—the trivial name alticostata is from the
Latin altus, high, and costa, rib, in reference to the elevated
costae of this species.

V. (Claibornicardia) nasuta Dall, 1903
Plate 27, figures la-ld

V. nasuta Dall, 1903, Wagner Free Inst. Sci., Trans., vol. 3, No. 6,
p. 1425, pl. 53, fig. 9; Kellum, 1926, U.S. Geol. Sur., Prof. Pap.
243, p. 23 (non? pl. 3, fig. 3); Stenzel, Krause, and Twining,
1957, Univ. Texas, Pub. 5704, p. 106.

V’. (cf. Claibornicardia) nasuta Dall, Palmer and Brann, 1965, Bull.
Amer. Paleont., vol. 48, No. 218, p. 336.

Distinguishing characters.—General features similar to
V. (C.) alticostata but distinguishable from that and _ all
other claibornicard species by extreme length and extended
posterior, resembling in shape Carditamera. Distinct pattern
of prominent anterior left cardinal (2) which appears trian-
gular in shape and directed ventrally forward, with its dor-
sal surface projecting into a deep socket in the opposite
valve but excavated to the level of the hinge plate on the
ventral surface. All ribs except the posterior five and the
first two anterior have basic tripartite structure with swol-
len transverse nodes on central cord; paracostals equal in
width to costal cord, forming prominent rounded terraces
noded in concert with costal cord, and almost meeting in
broadly V-shaped intercostal channels. Posterior ribs simpli-

fied like those of V. (C.) alticostata, with broadly U-shaped
intercostals.

Type.—Holotype, a left valve preserved by silicification,
USNM 164626, dimensions below.

Sp'n L H YW RN BL/L BH/H H/L “Y%wW/H
Holo. 39.6 30.0 14.2 25 77 -82 -76 fae

Type locality —USGS 393, “Eocene of Conecuh Coun-
ty, Alabama.”

Distribution.—Type locality only.

Stratigraphic occurrence.—Glendon Limestone? , middle
Oligocene? age.

Dimensional summary.—See type dimensions.

Curvature characteristics.—

Sample: Holotype, USNM 164626. N=1
$1 H1 $2 H2 33 H3 S4 H+ $5
OR 28° 1) 44° ial Sis 12.5 62° _— =

Discussion and diagnosis——The type locality in Cone-
cuh County, Alabama, is unknown, and hence the strati-
graphic placement of this species is open to question. Dall
(1903) reported that the donor of the shell, I. C. Johnson,
assigned the species to the Midway, but disagreed and
placed it in the “Eocene.” Murder Creek and its tributaries
in the central part of Conecuh County, cut through a com-
plete Eocene section as well as rocks of Oligocene age in
the southern part of the county. The Sepulga River, form-
ing the eastern county border, also exposes a similar sec-
tion. Of all the lithologies mapped in this county (Cooke in
Adams, et al., 1926) and the only one likely to yield silicified
fossils is the Glendon Limestone of middle Oligocene (Vicks-
burgian) age (Cooke in Adams, 1926, et al., op. cit., pp.
286, 287). Neither the “Ocala Limestone” of Jacksonian
age nor the Marianna Limestone underlying the Glendon in
the Vicksburg Group are characterized by high percentages
of silica, while much chert and fossil silification are known
in the Glendon.

V. (C.) nasuta “looks” Eocene, and resembles V. (C.)
alticostata in ornamentation, but its extreme elongation and
strengthened dental pattern indicate an evolutionary ad-
vance over the Gosport species, as well as the simplification
of the first two anterior ribs. Cooke (op. cit., p. 292) re-
ported an unidentified Venericardia “sp.” in the Byram
Marl which overlies the Glendon at Weavers Chute in
Escambia County, Alabama. Species lists in the same report
include “V. serricosta Heilprin” from the Byram Marl from
southern Clarke County in that state. The identification of
these Byram forms may be questionable, but that early

110 PALAEONTOGRAPHICA AMERICANA (VI, 39)

Miocene species is somewhat elongated like V. (C.) nasuta
and may be a closely related form in the upper part of the
Vicksburg Group. Kellum’s (1926, pl. 3, fig. 3) report of
V.(C.) nasuta in beds of Jackson age in Wilmington, North
Carolina, from an internal mold is probably not correct,
because comparison of his specimen with an internal mold
of the holotype indicates a species of similar elongation,
but beak inflation and prosogyration are much different.
However, Kellum’s specimen is undoubtedly a Venericardia
and may be ancestral to this species, if not conspecific, and
indicates the survival of Claibornicardia past the end of
the middle Eocene, no matter what the age of V. (C.)
nasuta.

Several growth stages are evident on the holotype of
this species, and the early ones are not characterized by the
high degree of elongation which is seen in the adult speci-
men (see PI. 27, fig. 1b). Two small stages have H/L ratios
of .85 and .84, comparable to most claibornicards. Later
stages and the terminal stage are characterized by growth
ratios of .74, indicating an allometric increase in length over
height. This is not typical of other members of the subgenus
Claibornicardia, and the extreme length, as well as the
strengthening of the dentition with the forward slope of the
anterior cardinal (2), may indicate this species to be in the
line of evolution leading to Carditamera, first appearing in
the Miocene. The statement by Stenzel, Krause, and Twin-
ing (1957, p. 106) that V. nasuta Dall “. . . might better
have remained undescribed and unnamed” is totally inap-
propriate in this light.

Etymology.—The trivial name nasuta from the Latin
nasutus, large nosed, in reference to the elongated posterior
portion of the shell.

Subgenus GLYPTOACTIS Stewart, 1930

Glyptoactis Stewart, 1930, Acad. Nat. Sci. Philadelphia, Spec. Pub.
3, pp. 151, 152; Verastegui, 1953, Paleontographica Americana,
vol. 3, No. 25, pp. 40, 41; Stenzel, Krause, and Twining, 1957,
Univ. Texas, Pub. 5704, p. 106; Heaslip, 1960, Geol. Soc. Amer.,
Bull., vol. 71, No. 12, pt. 2, pp. 1885, 1886 abstract.

Type species. (original designation). — Venericardia
(Glyptoactis) hadra Dall, 1903, of the Chipola Marl, middle
Miocene age of Florida.

Type locality—United States Geological Survey lo-
cality 2213, 1 mile below Bailey’s Ferry, banks of the
Chipola River, Calhoun County, Florida.

Distribution—Southeastern and Gulf Coastal areas of
North America, Caribbean Islands (?), west coast of Cali-
fornia, Central America, and South America to northern
Peru.

Stratigraphic occurrence.

Late Oligocene and Miocene,
in southeastern United States and Caribbean, Pleistocene

and Recent on west coast of California, Central America,
and South America.

Subgeneric definition —Venericards having highly in-
flated, somewhat elongate shells with variously developed
posterior truncation, ornamented by radiating costae which
are tripartite in primitive forms with rounded, poorly de-
veloped paracostals, and fairly wide costal cord ornamented
by symmetrical, nonspinose nodes. Costal cord becomes wide
and overhanging in advanced forms in which paracostals
are lost, and anodulous and smooth in highly advanced
species. Umbones generally high and inflated, although low
in primitive forms.

Dentition strongly developed, but right anterior cardi-
nal variously developed, and weaker than same feature in
Claibornicardia, although basically aligned with right medial
cardinal as in the former subgenus, and vestigial in highly
advanced forms.

Extra element added to dentition not appearing in other
subgenera in “tooth” developed above intersection of lunule
and anterior point of left posterior cardinal tooth (4b)
formed by flexion of the lunule. Corresponding socket form-
ed by flexion and indentation of right lunule above anterior
point of right medial cardinal tooth (3b).

Adult curvature pattern consists of five segments in
most species, four of which increase in convexity from beak
to ventral margin and the fifth or terminal segment of lower
convexity than the previous one. Early species also contain
adults having a four segment pattern, minus the terminal
fifth segment.

Discussion and diagnosis—The subgenus Glyptoactis
was originally proposed by Stewart (1930) to include per-
haps all of the “alticostate” forms of Venericardia, because
he recognized V. (Claibornicardia) alticostata (Conrad) of
the Gosport Sand and V. acuticosta Lamarck from the
Paris Basin Eocene as well as the Miocene and Recent
species. Stenzel, Krause, and Twining (1957), in a more
realistic examination of the alticostate venericards, erected
the subgenus Clatbornicardia to include the Eocene forms,
although not fully recognizing the characteristic dental pat-
tern which unites the entire group and at the same time
distinguishes its members from those of the ancestral sub-
genus Baluchicardia Rutsch and Schenck.

The early species of Glyptoactis in the late Oligocene
and early Miocene forms of that subgenus resemble Clai-
bornicardia in many respects, having similar, although modi-
fied ornamentation, dental pattern, and shell shape. The
flexion of the lunule, however, forms the new hinge element
seen in primitive development in the early forms (see PI. 27,
fig. 3c) but strongly developed in the more advanced Mio-

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 111

cene forms. This seems sufficient, in concert with ornamental
simplification, to distinguish between the glyptoactids and
the claibornicards from which they were obviously derived.

The inclusion of species from the French Eocene by
Stewart is unrealistic, and much detailed study is necessary
before forms so widely separated geographically could be
included in the same evolutionary lineage. The same may be
true of the California Eocene “Cardita” sandiegoensis M. A.
Hanna, 1927, and this species is more likely a member of
Claibornicardia, although the West Coast Tertiary species
have not been studied in the present investigation.

The relationship between Glyptoactis and the European
Glans Megerle, 1811 is not established at this time, but it is
highly probable that these two groups represent independent
lines of evolution, despite certain similarities in dental pat-
tern suggested by Stewart (1930).

Etymology.—The subgeneric name Glyptoactis Stewart
is derived from the Greek, glyptos, carved, and actints, ray,
in allusion to the sculptured nature of the costae.

V. (Glyptoactis) nodifera Kellum, 1926
Plate 27, figures 2a-2c, 3a-3c, 4, 5a-5c, 6

V’. nodifera Kellum, 1926, U.S. Geol. Sur., Prof. Pap. 143, pp. 36, 37,
pl. 9, figs. 1-3; Mansfield, 1937, Florida Geol. Sur., Bull. 15,
p. 236; Harris, 1951, Bull. Amer. Paleont., vol. 33, No. 138, pp.
16, 17 (non I’. cf. nodifera, pl. 7, figs. 9-11).

V’. waynensis Mansfield, 1940, Jour. Paleont., vol. 14, No. 3, pp. 189,
190, pl. 25, figs. 9, 10, 18,

Distinguishing characters —Similar in ornamentation to
V. (G.) serricosta (Heilprin), but distinguishable from that
species by lower degree of postero-ventral elongation and
much less inflated umbones. Somewhat similar in shape and
dentition to V. (G.) hadra Dall and V. (G.) himerta Dall
but distinguishable from these on basis of narrower ribs
with smaller and more closely spaced nodes on costal cord
and higher development of paracostals as well as less in-
flated umbones. Lunular “tooth” not so strongly developed
as in other species.

Discussion —All morphological comparisons failed to
yield consistent or significant differences, and a comparison
between reduced major axis slopes yielded the low correla-
tion coefficient (Z) of .230 in samples of V. (G.) waynensis
Mansfield of the Chickasawhay (upper) Oligocene and J.
(G.) nodifera Kellum of the “Trent” (Silverdale fauna) beds
of early Miocene age. These are united under Kellum’s
name, but certain minor differences and the stratigraphic
placement of these forms are given subspecific recognition

Etymology.—Trivial name nodifera from the Latin
nodus, knot, swelling and fero, carry or bear, in reference to
the costal nodes.

V. (G.) nodifera nodifera Kellum, 1926
Plate 27, figures 5a-5e, 6

V’. nodifera Kellum, 1926, U.S. Geol. Sur., Prof. Pap. 143, pp. 36, 37, pl.

9, figs. 1-3; Mansfield, 1937, Florida Geol. Sur., Bull., vol. 15,

p. 236 (this species?) ; Harris, 1951, Bull. Amer. Paleont., vol.

33, No. 138, pp. 16, 17 (non pl. 7, figs. 9-11).

Distinguishing characters.—Distinguishable from the
ancestral subspecies, V. (G.) nodifera waynensis by larger
size, higher average rib number, somewhat wider, evenly
and closely noded central costal cord which becomes dis-
proportionately wider toward the ventral margin on the
more anterior portion of the shell. Nodes are transverse
ridges, relatively flat-topped.

Type.—Holotype, a well-preserved right valve, USNM
353227, dimensions below.

Sp'n L H YW RN BL/L BH/H H/L

Holo. 31.2 30.8 12.3 24 67 86 eh) 40

Type locality—USGS 10655, 1% mile south of Silver-
dale, on left side of Webb’s Creek, in drainage pits and soil
banks, Onslow County, North Carolina. “Trent” (Silver-
dale) Formation, early Miocene age.

Distribution —Beside the holotype, a large collection
from AMNH 1059, about 4 mile from the type locality in
Gillette’s new (1957) marl pits was examined in the investi-
gation.

Stratigraphic occurrence —“Trent” (Silverdale) Forma-
tion, early Miocene age.

YwW/

Dimensional swmmary.—

Sample: AMNH 1059, Holotype USNM 353227

= 16 Dimension
Statistic It, H YW RN
OR 13.6-42.0 12.2-37.9 4.9-16.2 20-25
XxX 26.8 25:3 10.6 22.3
hS) 6.3 6.1 2.5 1.4
Vv 23.4 24.2 23.8 6.2
Ox 1.6 1.5 6 4
Ratio
BL/L BH/H H/L W/H
OR 64-.70 83-.89 88-99 .40-.45
> .67 86 94 42
Curvature characteristics. —
Sample: AMNH 1059, holotype USNM 353227 N = 16
$1 H1 $2 H2 $3 H3 S4 H4+* $5*

OR 20-44° .7-2.2 35-50° 2.4-8.1 50-59° 5.6-15.0 56-63° 10.0-16.9 57-65°

OLS OF CUE BI 94-2): 4NG mesic ls 9°10 Wten5 S pS ommmNTIG"O memEGUAre

*General pattern A. 2 individuals have more convex segment
(S5), 2 have lower value than previous segment.

Discussion and diagnosis —Recent investigation by the
U. S. Geological Survey has established that the marl bed
at Silverdale, North Carolina, which contains abundant

112 PALAEONTOGRAPHICA AMERICANA (VI, 39)

members of this subspecies, is not the true Trent Formation
which is a highly compacted limestone containing only molds
and casts of mollusk shells. No account of the relationship
between the two units has yet been published, and the bed
at Silverdale is referred to as “Trent” (Silverdale) in the
present report.

The Tampa Limestone occurrences of this subspecies
questionably reported by Mansfield (1937) are probably
V.(G.) serricosta which is well known from that formation.
A sample of four small (maximum length 19.6 mm) in-
dividuals from Ballast Point on Tampa Bay, Florida (USGS
16609), identified as “V. nodifera” in U. S. Geological Sur-
vey collections agreed only with samples of V. (G.) serri-
costa in growth analyses (reduced major axis) but distin-
guished from the sample of V. (G.) nodifera nodifera from
the vicinity of its type locality, and Mansfield’s identifica-
tions were probably based on immature forms of the Tampa
species.

Because of its generalized shape, but beginnings of
ornamental specialization (costal widening, transverse, flat-
tened nodes), this subspecies is the likely ancestor of the
highly specialized Oak Grove form V. (G.) himerta Dall,
and perhaps V. (G.) hesperide Gardner (see Text-fig. 28).
Forms appearing at the same time such as V. (G.) impen-
deocosta, n. sp., V. (G.) gibberumbonata, n. sp., and V. (G.)
serricosta (Heilprin) were probably derived from the late
Oligocene subspecies (see Text-fig. 28).

V. (G.) nodifera waynensis Mansfield, 1940
Plate 27, figures 2a-2c, 3a-3c, 4

V. waynensis Mansfield, 1940, Jour. Paleont., vol. 14, No. 3, pp-
189, 190, pl. 25, figs. 9, 10, 18.

Distinguishing characters. — Distinguishable from JV.
(G.) nodifera nodifera mainly on basis of lower rib number
(observed range 15-20) and somewhat thinner costal cords
ornamented by transverse nodes which are somewhat less
regularly and closely spaced than in that subspecies. Rela-
tively constant width maintained in more anterior ribs
toward ventral margin; these become disproportionately
wider in that subspecies. Adult size smaller (largest ob-
served value 24.9) than that of Miocene subspecies (largest
value 42.0), although somewhat larger, but broken and un-
measurable individuals of V. (G.) nodifera waynensis pres-
ent at USGS locality 14779. Shape more variable in this
subspecies.

Type. — Holotype, a well-preserved left valve, USNM
498538. Paratype dorsal portion of incomplete left valve,
498539, dimensions below.

Sp'n L H 14W RN BL/L BH/H H/L %W/H
Holo. 22.3 22.2 8.5 18 -76 36 1.00 38
Para 19.8 18.8 7.2 19 = += = =

*Estimated

Type locality—USGS 14204, lower bed just east of
bridge over Taylor Mill Creek on Highway 45, 114 miles
north of Waynesboro, Wayne County, Mississippi. Paratype
locality, USGS 14205, soft marl bed underlying ledge in gully
about 100 yards east of dirt road into woods about 1% mile
north of Perdue Hill, Monroe County, Alabama. Chicka-
sawhay Formation, late Oligocene age.

Distribution—Beside the type and paratype, a large
collection of this subspecies from USGS 14779 in the vicinity
of the holotype locality was examined in the present investi-
gation. Mansfield (1940) noted other localities in Missis-
sippi for the distribution of this form.

Stratigraphic occurrence. — Chickasawhay Formation,
late Oligocene age.

Dimensional summary.—

Sample: USGS 14779

N = 54 Dimension
Statistic i H 4W RN
OR 2.4-24.9 2.4-23.0 .8-9.5 15-20
x 11.9 11.4 4.7 18.1
S 6.7 6.4 2.8 1.1
Vv 56.1 56.5 59.5 6.2
Ox 9 9 4 2
Ratio
BL/L BH/H H/L YW/H
OR -56-.81 .80-.96 -88-1.05 .33-.51
BS .68 88 96 4

Curvature characteristics. —

Sample: USGS 14779 N= 22"
S1 H1 $2 H2 S3 H3 S4 H+ $5

OR 19-36° .3-2.2 35-50° 1.0-5.1 46-60° 4.2-8.3 54-62° 7.4 57°
x WSs 185, FA 256 51.6° 5.6 58.8° 74 Sin
*Every third individual of total sample.

Discussion and diagnosis. — The subspecies V. (G.)
nodifera waynensis differs only in minor characters from
V. (G.) nodifera nodifera which is so abundant in the
“Trent” locality on Gillette’s property near Silverdale, North
Carolina. All values obtained in the large sample above
(USGS 14779) from about five miles south of the type
locality (USGS 14204) compared favorably to those of the
large sample of the Miocene subspecies from Silverdale in all
characters except size and ornamental details, and these dif-
ferences are best expressed by placing the two forms in a
time-stratigraphic subspecific relationship.

Shape and degree of postero-ventral elongation are

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 113

highly variable in the ancestral subspecies. Only the holo-
type and one elongate individual in the measured sample
exhibited the B pattern of curvature, and if a comparison
can be made to V. (Claibornicardia) alticostata (Conrad),
this may signify maleness, while the highly abundant forms
with A curvature may be females of the population.

The earlier Oligocene ancestors of V. (G.) nodifera
waynensis are unknown, and an hiatus exists from the time
of Gosport Sand (late middle Eocene) deposition to the late
Oligocene. Although V. (C.) nasuta Dall is probably of mid-
dle Oligocene age, it is much too highly specialized in shape,
dentition, and ornamentation to have given rise to the
glyptoactids, and a search of the Caribbean or Central
American Oligocene faunas may fill the gap.

Mansfield (1940) noted the close relationship of this
subspecies V. (G.) serricosta (Heilprin) of the early Mio-
cene, and it is clear that this species, as well as V. (G.)
nodtfera nodifera is descended from it.

Etymology—The subspecific name waynensis is geo-
graphic in honor of the type locality in Wayne County,
Mississippi.

V. (Glyptoactis) serricosta (Heilprin), 1887
Plate 27, figures 7a, 7b, 8; Plate 28, figure 1

Cardita (Carditamera) serricosta Heilprin, 1887, Wagner Free Inst.
Sci., Trans., vol. 1, p. 117, pl. 16, fig. 64, reprint Pal. Amer., vol.
4, No. 33, 1964. Dall, 1900, Wagner Free Inst. Sci., Trans.,
vol. 3, No. 5, pl. 38, fig. 9.

V’. serricosta (Heilprin), Dall, 1903, ibid., vol. 3, No. 6, pp. 1428,
1429; Dall, 1915, U. S. Nat. Mus., Bull., vol. 90, p. 132, pl. 26,
fig. 6; Mansfield, 1937, Florida Geol. Sur., Bull., vol. 15, pp. 234,
235:

Non I. serricosta brooksvillensis Mansfield, 1937, idem., p. 235, pl.
U7, figs. 1.9;

V’. himerta Dall, Dall, 1915, U. S. Nat. Mus., Bull., vol. 90, p. 132.

Non I. himerta Dall, 1903.

Distinguishing characters. — Similar in ornamentation
and dental pattern to V. (G.)nodifera waynensis Mansfield
but distinguishable from that subspecies by greater posterior
elongation and much higher, more inflated umbones. Widely
spaced costal nodes on central shell distinctively subdued
sawtooth in shape, pointing toward ventral margin. These
become regularly and closely spaced transverse ridges on
anterior ribs, much like those of V. (G.) nodifera nodifera,
but not so distinctive as those of V. (G.) hadra. Develop-
ment of paracostals similar, rib number slightly higher than
in V. (G.) nodtfera waynensis, and lacks strongly arched
ventral margin of right hinge plate found in that form.

Type. — Heilprin’s figured holotype is not present in
ANSP collection 10640, a second specimen is the large left
valve in that collection, dimensions below.

Sp'n L H %W RN BL/L BH/H H/L “%W/H
ANSP 32.7 29.2 12.4 20 79 81 89 43
10640

Type locality. — “Silex” beds at Ballast Point, Tampa
Bay, Hillsborough County, Florida. Tampa Limestone,
early Miocene age.

Distribution.—Besides the type collection, ANSP 10640,
topotype collections from USGS 16609, USNM_ 165192,
were examined. Mansfield (1937) listed other localities for
the occurrence of this species.

Stratigraphic occurrence. — Tampa Limestone, early
Miocene age.

Dimensional summary. —

Sample: ANSP 10640 (2 spec.) USGS 165192 (8 spec.)

N=10 Dimension
Statistic L H ,W RN
OR 21.5-32.7 19.4-29.2 8.4-12.4 18-20
». 26.8 23.6 10.3 19.2
S 4.2 aan 1.5 38
Vv 15.6 15.6 14.1 4.1
Ox 1.3 1.2 5 3
Ratio
BL/L BH/H H/L “4W/H
OR .73-.83 -75-.85 -84-.95 41-48
X 76 82 89 44
Curvature characteristics.—
Sample: USNM 165192 N=8
S1 H1 $2 H2 $3 H3 S4 H4+ $5

OR 31-41° 1.6-3.8 40-53° 3.8-7.2 52-60° 4.9-11.3 59-66° 7.6-15.3 55-61°
x 36.8° 2.6 46.7° 4.9 55.0° ie. MP SS 58282

Discussion and diagnosis—The high degree of postero-
ventral elongation of V. (G.) serricosta led Heilprin to
describe it as a species of Carditamera, but the typical den-
tal pattern of Venericardia was recognized by Dall (1903).
Heilprin’s figure does not compare to any specimen in the
ANSP type collection (not in Wagner Free Institute collec-
tions). The second specimen in that collection (10640
ANSP) would be the proper shell for a neotype.

This early Miocene species is highly variable in lateral
outline, much as its ancestor V. (G.) nodifera waynensts,
and because of the low rib count and ornamental similarity
to this form this relationship is most reasonable, rather than
descent from any early Miocene form. Although much less
variable than this Tampa species in rib number, V. (G.)
hadra Dall, with an observed range of 19-22, is similar to
V.(G.) serricosta in this respect, and although less elongat-
ed, also has high and inflated umbones. It is highly probable
that V. (G.) hadra is descended from V. (G.) serricosta.

114 PALAEONTOGRAPHICA AMERICANA (VI, 39)

Etymology.—Trivial name serricosta from the Latin,
serra, saw, and costa, rib, in recognition of the suppressed
sawtooth nodes on the central ribs.

V. (Glyptoactis) impendeocosta, n. sp.
Plate 28, figures 3a-3c

Distinguishing characters. — Somewhat like V. (G.)
nodifera nodifera in general outlines and dentition but dis-
tinguishable on the basis of unique pattern of left anterior
cardinal (2) which is pointedly triangular and obliquely
sloping to the rear, with excavation on anterior face for
highly sloping anterior cardinal (3a) of right valve. Highly
simplified and flat-topped ribs which project over intercostal
space. Paracostals completely lost and intercostals form
broad “V” with slitlike line at center. Costal nodes only
slightly developed on juvenile part of shell and on anterior
ribs of mature shell.

Type.—Holotype, a left valve, USNM 644321, dimen-
sions below; paratype, USNM 644322.

Sp'n L H YW RN BL/L BH/H H/L
Holo. 25.2 25.0 9.4 21 69 -88 ee) 38

Type locality —USGS 22287, Haywood Landing on the
White Oak River, Jones County, North Carolina. “Trent”
(Silverdale) Formation, early Miocene age.

Distribution —Known only from the type locality.

Stratigraphic occurrence.—“Trent” (Silverdale) Forma-
tion, early Miocene age.

Dimensional summary.—See type dimensions.

Curvature characteristics —

“W/H

Sample: Holotype Nal
S1 H1 $2 HZ Ss H3 S4 H+ S5

OR 21° 4 365 (2.2 SO 4.3 55° — —

Discussion and diagnosis. — V. (G.) impendeocosta,
n. sp., known only from four specimens, the holotype and
three small, broken valves, is undoubtedly a departure from
V.(G.) nodifera s. 1. It appears simultaneously with V’. (G.)
nodifera nodifera in the “Trent” (Silverdale) Marl but is
known only from the type locality at Haywood Landing.*
It is more similar in general form to this subspecies than
to the late Oligocene subspecies V. (G.) nodifera waynensis.

The most distinctive character is the flattened, over-
hanging costae which bear no resemblance to those of V.
(G.) nodifera except in the spacing of the suppressed an-
terior nodes. The ribs of that species are broadly based,
showing some development of paracostals and U-shaped

*Collections made by the writer in 1967 from a new quarry near
Silverdale contained several individuals of this species. Data not in-
cluded here.

intercostal channels. This costal flattening is part of a gen-
eral trend among the glyptoactids (see Text-fig. 28).

The pattern of left anterior cardinal (2) is also distinc-
tive of this species, and is seen as well in the small frag-
mental specimens, all of which are left valves. These are
immature valves, and exhibit a greater costal node develop-
ment on all ribs. However, this is also characteristic of the
immature portion of the holotype, and signifies the transi-
tional nature of this species between the strongly noded an-
cestral forms and the advanced stage of costal flattening as
that seen in the modern V’. (G.) crassicostata G. B. Sower-
by I, 1825, which lives off the modern coast of Central
America. However, no relationship is implied, and V. (G.)
impendeocosta probably did not give rise to any subsequent
forms.

Etymology.—The trivial name impendeocosta is de-
rived from the Latin, impendeo, overhang, and costa, rib,
in reference to its wide, flattened costae.

V. (Glyptoactis) gibberumbonata, n. sp.
Plate 28, figures 2a-2c

Distinguishing characters—Somewhat similar in orna-
mentation to the sympatric V. (G.) nodifera nodifera but
readily distinguished from that form by flattened, somewhat
overhanging central costal cords on which regularly spaced,
symmetrical nodes become irregular and poorly developed
toward ventral margin. Rib number within the range of that
subspecies, but paracostal cords entirely absent, differing
from that subspecies, which has poorly developed ones. High
inflated umbones and high degree of postero-ventral elonga-
tion both far beyond the observed range for these charac-
ters in V. (G.) nodifera nodifera. Dentition more delicate
than in that subspecies, with highly reduced, horizontal left
anterior cardinal (2) surmounted by a saddle-shaped process
reflecting the convex ventral articulating surface of the right
anterior cardinal (3a). Somewhat similar to V. (G.) hadra
Dall in general shape but distinguished by its much more
delicate dentition and simplification of costals.

Type.—Holotype, a well-preserved mature left valve,
USNM 644323, dimensions below. Monotypic.

Sp'n L H %W RN BL/L BH/H H/L “%W/H
Holo. 33.2 34.0 15.2 21 76 78 1.02 AS
Type locality. — USGS 21278, Gillette’s new (1957)

marl pit on Gillette’s farm, about 14 mile south of Silverdale,
Onslow County, North Carolina. “Trent” (Silverdale)
Formation, early Miocene age.
Distribution—Known only from type locality.
Stratigraphic occurrence. —“Trent” (Silverdale) For-
mation, early Miocene age.

AMERICAN ALTICOSTATE VENERICARDS: HEaSsLIP 115

Dimensional analysis—See type dimensions.
Curvature characteristics —

Sample: Holotype. N= 1
S1 H1 $2 H2 S83 H3 S4 H+ $5
OR 20° 1.6 49° 3.9 54° 8.3 64° 21.1 62°

Discussion and diagnosis—Although similar in develop-
ment of costal nodes on mature (but not terminal) portion
of shell to V. (G.) nodifera nodifera, V. (G.) gibberum-
bonata, n. sp., is separated from that synchronous and sym-
patric form by such a wide gap in other morphological char-
acters that it must be given specific rank. Differences of
this order certainly appear in other species [e.g. V. (Clai-
bornicardia) alticostata], but their intraspecific nature is
discernable as gradation among extreme individuals in sam-
ples of any size. Among the large topotype collections of
V. (G.) nodifera nodifera examined in the present investi-
gation, no individuals comparable to this unique specimen
were seen.

Because of ornamental similarities, V. (G.) gibberuwm-
bonata is undoubtedly derived from V. (G.) nodifera, but
its umbonal height and inflation anticipates V. (G.) hadra of
the middle Miocene Chipola Beds. Because of its reduced
dentition and complete loss of paracostals, it is probably not
the ancestor for the Chipola form and appears to have be-
come extinct after the time of “Trent” (Silverdale) deposi-
tion.

Etymology —tThe trivial name gibberumbonata is de-
rived from the Latin, gibber, inflated, and wmbo, boss or
knob, in reference to the highly inflated umbones.

V. (Glyptoactis) himerta Dall, 1903
Plate 28, figures 4a-4d

Cardita sp. Dall, 1900, Wagner Free Inst. Sci., Trans., vol. 3, No. 5,
pl. 40, fig. 16.

Venericardia himerta Dall, 1903, idem., vol. 3, No. 6, p. 1430, pl. 53,
fic, 12; Dall, 1915, U. S. Nat. Mus. Bull., vol. 90, p. 132
(partim); Gardner, 1926, U.S. Geol. Sur., Prof. Pap. 142-B, pp.
90, 91, pl. 17, fig. 13; Stewart, 1930, Acad. Nat. Sci., Philadelphia,
Spec. Pub. 3, p. 152 (not by name); Mansfield, 1937, Florida
Geol. Sur., Bull. 15, p. 234.

Distinguishing characters. — Generally similar to J.
(G.) hadra Dall of the Chipola but distinguishable from that
species on the basis of central costal cords which are rounded
in cross-section but widened somewhat over intercostal
space. Intercostals U-shaped with suture-like line at center
much like the Chipola species, but paracostals absent or
only poorly visible thread on anterior ribs. Costal nodes
subdued and widely, but regularly spaced transverse ridges.
V. (G.) hadra has narrower central cord and fairly promi-
nent paracostals. Umbones lower than that species, less in-

flated, and posterior margin longer and flaring. Ventral pro-

jection of shell margin less. Dental pattern similar to that
species, but hinge plate relatively lower and less strongly
arched. Anterior left cardinal (2) flatter, less strongly de-
veloped. Anterior adductor scar much more elongated and
relatively greater in area than same feature in other species.

Type. — Holotype, complete individual (right valve
figured by Dall, 1903), from collection USNM 164555, di-

mensions below.

Sp’n LE H %W RN BL/L BH/H H/L Y%W/H
Holo. 52.0 53.0 22.8 21 wil -83 1.02 43

Type locality. —USGS 2646, Oak Grove on the Yellow
River, Okaloosa County, Florida. Puri (1953, p. 58) de-
scribed the locality as “At old sawmill near Oak Grove on
right bank of Yellow River, 300 feet south of northwest
corner of NE 4, NE 4%, Sec. 20, T 5 N, R 23 W, about
100 yards below bridge on Laurel Hill-Oak Grove Road,
Okaloosa County, Florida.”

Distribution. — Only the type collection was examined
in the present investigation. Gardner (1926) noted other
occurrences of this species.

Stratigraphic occurrence.
Miocene age.

Dimensional summary. —

Oak Grove Sand, late early

Sample: USNM 164555, type collection

INS 5) Dimension
Statistic 1b, H YW RN
O! 15.7-52.0 15.2-53.0 6.3-22.8 18-21
xX 27.6 27.0 11.6 19.3
Ss 10.4 10.6 4.7 oi
V 37.8 39.4 40.5 3.8
Ox 2.7 2.8 nee 2
Ratio
BL/L BH/H H/L “4W/H
OR .64-.73 .79-.92 90-1.04 41-.45
xe 68 185 98 43

Curvature characteristics. —

Sample: USNM 164555, type collection Nis

SI H1 $2 H2 $3 H3 S4+ H4 $5

OR 21-34° 1.1-1.9 32-54° 1.9-5.8 50-69° 4.0-10.6 51-65° 7.2-20.9 52-58°
ER NRG, PVG. Bio ae ie eG ase

Discussion and diagnosis. — The original middle Mio-
cene age for this species is revised to late early Miocene
with the work of Puri (1953) which established the facies
relationship within the Alum Bluff Group in western Florida.
The Oak Grove is associated by that author with the early
rather than middle Miocene.

The trend toward costal broadening and_ paracostal
simplification evident in several glyptoactids of the Miocene
also produced V. (G.) himerta. Similarities in shape and

116 PALAEONTOGRAPHICA AMERICANA (YI, 39)

dentition indicate a close relationship between this species
and V. (G.) nodifera nodifera of the “Trent” (Silverdale)
which shows beginnings of costal broadening but still has
poorly developed paracostals.

This Oak Grove species and V. (G.) hadra of the
Chipola are the last abundant glyptoactids in the Miocene,
although V. (G.) olga of the late Miocene is also locally
abundant. After the Miocene, the glyptoactids seem to
have vacated their habitats in east and Gulf coastal waters.

If V. (G.) himerta gave rise to any subsequent forms
they are unknown in the east and Gulf regions. V. (G.)
olga Mansfield of late Miocene age represents an even more
highly advanced stage of ornamental simplification, but its
shape and dentition are more like those of V. (G.) hadra
than to V. himerta.

Etymology. — The trivial name himerta is from the
Greek, himertos, lovely, in probable reference to the finely
sculptured concentric increment lines and the beautiful state
of preservation of shells in Dall’s type collection.

V. (Glyptoactis) hadra Dall, 1903
Plate 29, figures 2a, 2b, 3, 4

Venericardia hadra Dall, 1903, Wagner Free Inst. Sci., Trans., vol.
3, No. 6, p. 1429, pl. 53, figs. 11, 13; Gardner, 1926, U.S. Geol.
Sur., Prof. Pap. 142-B, p. 90, pl. 17, figs. 11, 12; Mansfield,
1937, Florida Geol. Sur., Bull., vol. 15, p. 234.

V. (Glyptoactis) hadra Dall, Stewart, 1930, Acad. Nat. Sci, Phila-
delphia, Spec. Pub. 3, p. 152.

V. chipolana Maury, 1910, Bull. Amer. Paleont., vol. 4, No. 21, p.
34, pl. 4, fig. 9.

Distinguishing characters. — Somewhat similar in shape
and dental pattern to V. (C.) chelomodonta, n. sp., and V.
(G.) olga Mansfield but distinguished from those species
on basis of basically tripartite costae, with rounded para-
costals well developed on anterior ribs and relatively narrow
central costal cord ornamented by closely spaced transverse
ridges. Ribs in those species broadened with expanded cen-
tral costal cord, less closely spaced transverse nodes and
absent paracostals. Beaks smaller, umbones lower than in
those species. Dentition similar to that of V. (G.) olga but
distinguishable from V. (G.) chelomodonta by absence of
neomorphic “tooth” in left valve of that species.

Type. — Holotype, a complete individual, interior filled
with matrix (figured by Dall, 1903), USNM 114730, di-
mensions below.

Sp'n L H YW RN BL/L BH/H H/L “%W/H
inion 41:2 eae ON ets Seer oT 87 83 92 Al
Type locality.—USGS 2213, 1 mile below Bailey’s

Ferry, banks of Chipola River, Calhoun County, Florida.
Chipola Formation, middle Miocene age.

Distribution. — Only Dall’s type collection of 19 indi-
viduals was examined in the present investigation. Gardner
(1926) listed numerous USGS localities in western Florida
at which this species occurs.

Stratigraphic occurrence. — Chipola Formation, middle
Miocene age.

Dimensional summary. —

Sample: USNM 114730, type collection

Ni=al9 Dimension

Statistic i, H VAs RN

OR 7.3-48.7 7.4-47.2 2.7-19.8 19-22

x 33.6 31.5 14.0 20.6

Ss 14.2 13.2 6.0 38

V 42.2 41.9 42.8 3.8

Ox 3.3 3.0 1.4 2
Ratio

BL/L BH/H H/L 14,W/H

OR .62-.87 .74-.91 .73-1.04 .37-.57

xX 74 81 95 A+

Curvature characteristics. —

Sample: USNM 114730, type collection Nis

3 H3 4+ H+ $5

OR 18-35° .9-2.6 35-38" 2.5-8.3 45-60° 5.9-15.6 61-68° 12.5-25.9 53-63"
xe 233° 1.6 460° 4:8 54.0° 10:3 65.1° S70 meeeogeee

Discussion and diagnosis.— The work of Puri (1953)
established the facies relationships among the middle Mio-
cene beds of Panhandle Florida, as well as the late early
Miocene age of the lower parts of those units. The Shoal
River, Chipola, and Hawthorne Formations are time-equiva-
lents, and the Oak Grove Sand a near shore facies of only
late early Miocene age (see Text-fig. 24). It is possible that
a close age relationship exists among V. (G.) hadra, V.
(G.) hesperide, and V. (G.) chelomodonta, The Chipola,
according to Puri, is characterized by an open water neritic
microfauna, while that of the Oak Grove and Shoal River
facies indicate more brackish conditions of deposition, in-
dicating perhaps different ecological tolerances for these
species.

Because of its more inflated beaks than other early
Miocene species, V. (G.) serricosta is presumed to be the
ancestor of V. (G.) hadra, and despite its shape similarities,
V.(G.) gibberumbonata of that age is too highly specialized
in dentition and ornamentation to have given rise to the
middle Miocene species. Similarly, the inflated beaks, high
umbones, and high dental plate of V. (G.) chelomodonta of
the Shoal River and V. (G.) olga of the late Miocene indi-
cate descent from V. (G.) hadra in a trend toward simplified
ornamentation and the high specialization of dentition in
the former species.

S1 H1 $2 H2

Dn
D

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 117

Etymology. —The trivial name hadra is from the
Greek, hadros, bulky or stout, in reference to the high con-
vexity of this species.

V. (Glyptoactis) hesperide Gardner, 1936
Plate 29, figures la-lc
V’. (Megacardita) hesperide Gardner, 1936, Florida Geol. Sur., Bull.

vol. 14, pp. 23, 24, pl. 5, figs. 16, 17.

Distinguishing characters. — Distinguishable from all
other glyptoactid species by extreme degree of postero-
ventral elongation, posterior and ventral margins drawn into
projection reminiscent of certain claibornicards of the middle
Eocene.

Right dentition much like that of other glyptoactids,
but anterior cardinal (3a) not visible. Left dentition un-
known. Highest degree of costal simplification of any species
in present investigation, costae rounded in cross-section,
expanded to constrict intercostals to slits. Nodes irregular
and vestigial, only rounded, slightly elevated ridges on more
anterior ribs, posterior ribs almost smooth.

Type. — Holotype, a right valve with part of posterior
margin missing, USNM 372890, dimensions below, Mono-
typic.

Sp’n L H %W RN BL/L BH/H H/L WY%W/H
Holo. 49.2 53.0 17.0 21 wl 89 1.07 32

Type locality —USGS 10603, Gully on east side of
White’s Creek, 200 feet south of the bridge on the road to
Knox Hill, 6.7 miles by road south of Argyle and about 1.7
miles southeast of Eucheanna, Walton County, Florida.
Shoal River Formation, middle Miocene.

Distribution. — Type locality only.

Stratigraphic occurrence. —Shoal River Formation,
middle Miocene age.

Dimensional summary.

Curvature characteristics. —

See type dimensions.

Sample: Holotype. Nea
S1 Fe Se Howes H3 S4+ H+ = $5
OR ale. 2.9 48° 4.0 53° 8.6 64° 15.7 51°

Discussion and diagnosis. —V.(G.) hesperide exhibits
the highest degree of costal simplification of any glyptoactid
on the east or Gulf Coastal regions of North America. The
expanded and almost anodulous ribs are reminiscent of the
similar development in members of Megacardita Sacco, 1899,
of the Mediterranean Miocene and seems a result of close
convergence rather than the relationship which Gardner
(1936) suggested. A similar trend in ornamental simplifica-
tion culminates in V. (Baluchicardia) whiteit Gardner of
the early Midway in Texas.

Occurring in the Shoal River facies of the Alum Bluff
stage in Panhandle Florida, this highly specialized form
apparently lived in a rather brackish environment, as inter-
preted by Puri (1953). If this has any bearing on the de-
velopment of shape and ornamental simplification in this
species, then perhaps it can also be inferred of other veneri-
cards which exhibit similar characters [e.g. V. (B.) whitei].

The ancestor of V. (G.) hesperide is probably not to be
found among the other forms of the Alum Bluff, and the low
slightly inflated umbones of this species may indicate a
direct descent from V. (G.) nodifera waynensis of the Chick-
asawhay, in which somewhat elongated forms also occur. No
subsequent species is known which might be descended from
V.(G.) hesperide, and it appears as an evolutionary experi-
ment in the Miocene radiation of the glyptoactids.

Etymology. — The trivial name hesperide is from the
Latin, hesperius, western, and the suffix -ide, like, or re-
sembling. Whether or not Gardner meant to compare this
species to V. (Baluchicardia) hesperia Gardner is not known,
but there is no resemblance between the two species. More
likely it refers to the far western position of Walton County,
the type locality, in Panhandle Florida.

V. (Glyptoactis) chelomodonta, n. sp.
Plate 28, figures 5a-5d

Distinguishing characters.— Shape similar to that of
V.(G.) hadra, V. (G.) olga, and V. (G.) gibberumbonata.
Distinguishable from the former species by total absence of
paracostals and widened, rounded costals, and from the latter
by much higher hinge plate and robust dentition. Ornamen-
tation and pattern of dentition similar to V. (G.) olga but
distinguished from that and all other glyptoactids by neo-
morphic projection from the dorsal surface of the posterior
left cardinal (4b) just behind the “tooth” formed by the
lunular flexion, with a gap or notch between the two. Shell
smaller, relatively thicker, and more compact than that in
other species.

Type. — Holotype, a small, but mature left valve,
USNM 644324, dimensions below. Monotypic.

1%W RN BL/L BH/H H/L “’wW/H
21 76 76 1.09 48

Sp’n 10, H

Holo. Pl | 22.9 11.0

Type locality. —USGS 18362 (Fla. G.S. W-25), 150
yards below Bell’s old mill site, Walton County, Florida.
Shoal River Formation. Vernon (1942) described W-25 as
“a ravine in the SW %, NW 4%, Sec. 15, T2 N, R15 W...
three foot exposure of blue-green slightly silty clay, which
weathers to light gray and yellow” and placed by the author

118 PALAEONTOGRAPHICA AMERICANA (VI, 39)

in the Choctawhatchee (late) Miocene. The collection in
which the type is placed is associated with Shoal River
collections at the U. S. National Museum. (U.S. Geological
Survey).
Distribution. — Known only from the type locality.
Stratigraphic occurrence. —Shoal River Formation,
middle Miocene age.
Dimensional summary. — See type dimensions.
Curvature characterisitcs. —

Sample: Holotype. NES

S1 Hi S82 H2 S83 H3 S4 H+ $5
OR Cpe Hie) S30 seeds) wee ale (ke = =

Discussion and diagnosis. — Unfortunately known from
only a single small left valve, this highly distinctive Shoal
River species is unique in the development of a projecting
“tooth” and notch on the dorsal surface of 4b (see Pl. 28,
fig. 5d), and although the right dentition is unknown, it
probably contained a similar structure fitting into the notch
formed by this tooth and the end of the lunule. This de-
velopment is neomorphic and not seen in any other species in
the present investigation.

The high umbones and similar shape, as well as the
strong development of the left anterior cardinal (2), point
toward V. (G.) hadra of the Chipola as the ancestor of V.
(G.) chelomodonta, but the exact stratigraphic relationship
between these two is not known, since the two “formations”
are time-equivalent facies.

The A pattern of curvature is not typical of most glyp-
toactids, and the possibility exists that the holotype is im-
mature. Ths is probably not the case, however, and the high
convexity (’% W/H = 48) may indicate the holotype to be
a female.

Because of the development of the neomorphic “tooth”
structure, this species did not give rise to V. (G.) olga of
the late Miocene, which exhibits no trace of this feature.

Etymology. — The trivial name chelomodonta is de-
rived from the Greek, cheloma, notch, and odows, tooth, in
reference to the notch formed by the neomorphic “tooth” in
the left hinge.

V. (Glyptoactis) olga Mansfield, 1939
Plate 29, figures 5a-5c, 6

Venericardia olga Mansfield, 1939, Florida Geol. Sur., Bull., vol. 18,
Pao6, pl. Qo tip Se plies tips 2) 7

Distinguishing characters.— Similar in shape and
dental pattern to V. (G.) hadra and V. (G.) chelomodonta.

Distinguished from the latter species mainly on the absence
of the neomorphic “tooth” developed in that form. Dis-
tinguished from the former by ornamentation consisting of
broadly expanded and somewhat overhanging rounded
costae lacking any trace of paracostals, and ornamented
with somewhat flattened, but prominent, closely spaced
transverse nodes similar only to those of V. (G.) chelomo-
donta. Ribs of V. (G.) hadra narrower, distinctly tripartite.
Vaulted posterior slope also characteristic only of V. (G.)
olga.

Type. — Lectotype (herein designated from syntypes
of Mansfield), a large right valve, USNM 497976, Paralecto-
type, a smaller left valve with broken posterior, USNM
645690, dimensions below.

Sp'n L H %W RN BL/L BH/H H/L %W/H
Lect. 43.3 41.4 19.6 17 75 eff) -96 A7
Para. S0:5*) 29:5 13.0 19 <2" 79 97 At

*Estimated

Type locality.—USGS 14075, dredged from the
Caloosahatchee River, one mile below Olga, Lee County,
Florida. Late Miocene.

Distribution. —Only type collection was examined in
this investigation, but it is present in other collections from
that region in the U. S. Geological Survey. (Wilson, personal
communication, 1963.)

Stratigraphic occurrence. — Mansfield (1939) indicated
a probable Pliocene age for this species based on the presence
of Chione cancellata (Linnaeus) on a chunk of limestone
from which one specimen was obtained. However, Druid Wil-
son (op. cit.) indicated a late Miocene age for this form
based on collections of known late Miocene age in the U.S.
Geological Survey not examined in the present investiga-
tion.

Dimensional summary. — See type dimensions.
Curvature characteristics. —

Sample: USNM 497976, lectotype; paralectotype, USNM 645690 N =:
$1 H1 $2 H2 $3 H3 S4+ H+ $5

OR 22-24° 1,5-1.8 45-50° 5.1-6.3 61° 6.3-10.1 68-70° 10.3-14.9 59-62
x 23.0° 1:6) 475 5.7 610° 8.2 69.0° 12.6 60.5°

Discussion and diagnosis. —It is apparent that V. (G.)
olga, because of similar shape and dental pattern, with
about the same development of right and left anterior
cardinals (3a, 2) as found in V. (G.) hadra of the Chipola
Miocene, is descended from that species. Certain similarities
are also apparent to modern venericards living off the west
coast of California, Mexico, Central America, and South
America. The record of glyptoactid evolution on the eastern

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP 119

and Gulf Coasts of North America ends with this species,
and it is highly likely that the descendants of V. (G.) olga
migrated to the eastern Pacific, perhaps giving rise to such
species as V. (G.) crassicostata G. B. Sowerby I (see Text-
fig. 28, Plate 29, figs. 7a, 7b, 8-c), known from the Pleisto-
cene and Recent seas in that region.

Etymology. — The trivial name olga is geographic in
honor of the type locality, near Olga, Lee County, Florida.

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United States, Amer. Jour. Conch., vol. 1, No. 1, pp. 1-35.

Cooke, C. W.

1926. In Adams, G. I., Butts, C., Stephenson, L. W., and Cooke,
C. W., Geology of Alabama, Geol. Sur. Alabama, Spec.
Rept. No. 14, The Cenozoic Formations. Pp. 251-297.

1952. Tertiary stratigraphy of South Carolina. U. S. Geol. Sur.,
Prof. Paper 243-B, pp. 19-29.

Cooke, C. W., Gardner, J., and Woodring, W. P.

1943. Correlation of the Cenozoic formations of the Atlantic and
Gulf Coastal Plain and the Caribbean region. Geol. Soc.
Amer., Bull., vol. 54, pp. 1713-1723, chart No. 12.

Currie, E. D.

1942. Growth stages in the ammonite Promicroceras marstonense
Spath. Roy. Soc. Edinburgh, Proc., Sect. B (Biol.), vol. 61,
pp. 344-367.

Dall, W. H.

1895. Contributions to the Tertiary fauna of Florida. Part 3.
A new classification of the Pelecypoda. Wagner Free Inst.
Sci., Trans., vol. 3, pt. 3, pp. 485-565.

1903. Ibid., Part 6, Concluding the work, pp. 1219-1564.

Davies, A. M.

1935. Tertiary faunas. vol. 1, The composition of Tertiary

faunas. Thomas Murby and Company, London, 406 pp.
Douvillé, H.

1912. Classification les lamellibranches. Soc. Géol. France, Bull.,

4th ser., vol. 12, pp. 419-467.
Durham, J. W.

1950. Im Anderson, C. A., Durham, J. W., Shepard, F. P., Nat-
land, M. L., and Revelle, R. 1940, FE. W. Scripps cruise
to the Gulf of California. Geol. Soc. Amer., Mem. 43, Part
II, Megascopic paleontology and marine stratigraphy. 216
pp., 48 pls.

Foster, V. M.
1940. Lauderdale County mineral resources. Mississippi Geol.
Sur., Bull., vol. 41, 246 pp.
Gabb, W. M.
1860. Descriptions of some new species of Cretaceous fossils.
Acad. Nat. Sci. Philadelphia, Jour., 2d ser., vol. 4, pp.
299-305, pls. 47-48 in part.
Gardner, J.
1923. New species of Mollusca from the Eocene deposits of
- southwestern Texas, U. S. Geol. Sur., Prof. Paper 131-D,
pp. 109-117, pls. 29-33.
1926. The molluscan fauna of the Alum Bluff Group of Florida.
U. S. Geol. Sur., Prof. Paper 142-B, Part II. Astartacea,
Carditacea, Chamacea, pp. 81-100, pls. 16, 17.
1935. The Midway Group of Texas. Univ. Texas, Bull. 3301,
403 pp., 28 pls.
1936. Additions to the molluscan fauna of the Alum Bluff
Group of Florida. Dept. Cons. Florida, Geol. Bull. 14,
pp. 1-82, pls. 1-10.
1945. Mollusca of the Tertiary formations of northeastern
Mexico. Geol. Soc. Amer., Mem. 11, 332 pp., 28 pls.
Gardner, J., and Bowles, E.
1939. The Fenericardia planicosta group in the Gulf Province.
U. S. Geol. Sur., Prof. Paper 189-F, pp. 143-215, pls. 33-46.

Gregorio, A. de
1890. Monographie de la Faune Eocénique de l’Alabama et
surtout de celle de Claiborne de l’Etage Parisien (Horizon
a Venericardia planicosta Lamk.). Ann. Geol. et Paleont.,
livr. 7 et 8, 316 pp., 46 pls.

Hanna, M. A.
1927. An Eocene invertebrate fauna from the La Jolla Quad-
rangle, California. Univ. California Pub., Bull. Dept. Geol.
Sci., vol. 16, No. 8, pp. 247-398, pls. 24-57.
Harris, G. D.
1919. Pelecypoda of the St. Maurice and Claiborne Staqes.
Bull. Amer. Paleont., vol. 6, No. 31, 268 pp., 59 pls.

Harris, G. D., and Palmer, K. V.W.

1946. The Mollusca of the Jackson Eocene of the Mississippi
embayment (Sabine River to Alabama River). Bull. Amer.
Paleont., vol. 30, No, 117, Part I, Bivalves, pp. 1-206, pls.
1-25.

Heaslip, W. G.

1960. Equiangular spiral and the classification of the alticostate
venericards. Geol. Soc. Amer., Bull., vol. 71, No. 12, pt. 2,
pp. 1885, 1886, abstract.

1963. Planicostate trend among alticostate venericards of North
America. Geol. Soc. Amer., Spec. Paper, No. 73, pp. 168,
169, abstract.

1964. Sexual dimorphism in alticostate venericards. Geol. Soc.
Amer., Spec. Paper, No. 76, p. 78 (abstract).

Imbrie, J.

1956. Biometrical methods in the study of invertebrate fossils.

Amer. Mus. Nat. Hist., Bull., vol. 108, art. 2, pp. 211-252.
Jackson, R. T.

1890. Phylogeny of the Pelecypoda, the Aviculidae and their
allies. Boston Soc. Nat. Hist., Mem., vol. 4, No. 8, pp.
277-400, pls. 23-30.

Kellum, L. B.

1926. Paleontology and stratigraphy of the Castle Hayne and
Trent mar!s in North Carolina. U. S. Geol. Sur., Prof.
Paper 143, pp. 1-56, 11 pls.

Lison, L.

1949. Recherches sur la forme et la mécanique de développement
des coquilles des lamellibranches. Mem. Inst. roy. sci. nat.
Belgique, 2d ser., fase. 34, 87 pp.

Loosanoff, V.

1936. Sexual phases in the Quahog. Science, vol. 83, pp.

287-288.
Lowe, E. N.

1933. Coastal Plain stratigraphy of Mississippi, Part 1, Midway
ard Wilcox Groups, Mississippi Geol. Sur., Bull. 25,
125 pp.

120

Mansfield, W. C. :

1937. Mollusks of the Tampa and Suawannee Limestones of
Florida. Florida Dept. Cons., Geol. Bull. No. 15, 334 pp.,
21 pls.

1939. Mie on the upper Tertiary and Pleistocene mollusks of
Peninsular Florida. Florida Dept. Cons., Geol. Bull. No.
18, 75 pp., + pls.

1940. Mollusks of the Chickasawhay Marl. Jour. Paleont., vol.
14, No. 3, pp. 171-226, pls. 25-27.

Mayr, E., Linsley, E. G., and Usinger,R. 1.

1953. Methods and principles of systematic zoology. McGraw-

Hill Book Company, Inc., New York, 328 pp.
Meyer, O., and Aldrich, T. H.

1886. The Tertiary fauna of Newton and Wautubbee, Missis-
sippi. Cincinnati Soc. Nat. Hist., Jour., vol. 9, No. 2, pp.
40-50, pl. 2.

Moore, E. J.

1962. Conrad’s Cenozoic fossil marine mollusk type specimens
at the Academy of Natural Sciences of Philadelphia. Acad.
Nat. Sci., Philadelphia, Proc. 1962, pp. 23-120, pls. 1, 2.

Munier-Chalmas, E. €. P. A.

1895. Deuxiéme note préliminaire sur le charniere des Mol-
lusques acéphales. Séance du 18 Mars, 1895, Compte-rendu
des séances, Soc. Geol. France, Bull., vol. 23, 3d ser., pp.
LIII-LVI.

Nicol, D.

1952. Revision of the pelecypod genus Echinochama, Jour.

Paleont., vol. 26, No. 5, pp. 803-817, pls. 118, 119.
Olsson, A. A.

1961. Mollusks of the tropical eastern Pacific, Panamic-Pacific
Pelecypoda. Paleont. Res. Inst., Ithaca, New York, 574 pp.
86 pls.

Orton, J. H.

1937. Oyster biology and oyster culture. Edward Arnold and

Company, London, 211 pp.
Owen, G.

1953. The shell in Lamellibranchia. Quart. Jour. Micro. Sci.,

vol. 94, part 1, pp. §7-70.
Palmer, K. V.W., and Brann, D. C.

1965. Catalogue of the Paleocene and Eocene Mollusca of the
southern and eastern United States, Part 1, Pelecypoda,
Amphineura, Pteropoda, Scaphopoda, and Cephalopoda.
Bull. Amer. Paleont., vol. 48, No. 218, 467 pp., 3 pls.

Plummer, F. B.

1933. In Secllards, E. H., Adkins, W. S8., and Plummer, F. B.
The geology of Texas. Vol. 1. Stratigraphy, Univ. Texas
Bull. 3232, Part 3, Cenozoic systems in Texas, pp. 519-
818, pls. 7-10.

Puri, H. S.

1953. Contribution to the study of the Miocene of the Florida

Panhandle. Florida Geol. Sur., Bull. 36, 345 pp., 17 pls.
Renfroe, C. A.

1949. Petroleum exploration in eastern Arkansas with selected

well logs. State of Arkansas, Div. Geol., Bull. 14, 159 pp.
Renick, B. C., and Stenzel, H. B.

1931. The Lower Claiborne on the Brazos River, Univ. Texas,

Bull. 3101, pp. 73-108, pls. 6, 7.

PALAEONTOGRAPHICA

Americana (VI, 39)

Richards, H. G., and Palmer K. V.W.
1953. Eocene mollusks from Citrus and Levy Counties, Florida.
Florida Geol. Sur., Bull. No. 35, 95 pp., 13 pls.
Rutsch, R. F.
1936. Beitrdge zur Kenntnis tropisch-amerikanischer Tertiar-
mollusken, 5. Ist Venericardia beaumonti auf die Ober-
kreide beschrankt? Eclogae Geol. Helvetiae, vol. 29, No. 1,
pp. 187-207, pl. 17.
1944. Die Paleocaen-Mollusken der Inseln Trinidad und Soldado
Rock (Britisch-W estindien). Ibid., vol. 36, No. 2, pp.
139-192, pls. 3-5.
Rutsch, R. F., and Schenck, H. G.
1940. Upper Cretaceous pelecypods of the Venertcardia beau-
monti group from Iran. Geol. Soc. Amer., Bull. vol. 51,
p. 1976 abstract.

Schmidt, F. C.
1818. Versuch iiber die beste Einrichtung, .
Gotha, 252 pp.
Stenzel, H. B., and Krause, E. K.
1957. In Stenzel, H. B., Krause, E. K., and Twining, J. T.
Stenzel, H. B., Krause, E. K., and Twining, J. T.
1957. Pelecypoda from the type locality of the Stone City Beds

(middle Eocene) of Texas. Univ. of Texas, Pub. 5704, 237
pp., 22 pls.
Stephenson, L. W.

1947. New Upper Cretaceous fossils from Mississippi and Texas.
U. S. Geol. Sur., Prof. Paper 210-E, pp. 161-196, pls. 31-33.

1952. Larger invertebrate fossils of the Woodbine Formation
(Cenomanian) of Texas. U. S. Geol. Sur., Prof. Paper 242,
226 pp., 59 pls.

Stewart, R. B. ;

1930. Gabb’s California Cretaceous and Tertiary type lamelli-
branchs. Acad. Nat. Sci., Philadelphia, Spec. Paper 3, 314
pp., 17 pls.

Thompson, D/A. W.

1952. On growth and form. Vol. 2, 2d Ed., Cambridge Univ.
Press, pp. 465-1116.

Verastegui, P. : ;

1953. The pelecypod genus Venericardia in the Paleocene and
Eocene of western North America. Palaeont. Amer., vol.
3, No. 25, 112 pp., 22 pls.

Vernon, R. O. : 7 ;

1942. Geology of Holmes and Washington Counties, Florida.
Florida Dept. Cons., Geol. Bull. No. 21, 161 pp.

Vokes, H. E. et :

1967. Genera of the Bivalvia: A systematic and bibliographic
catalogue. Bull. Amer. Paleont, vol. 51, No. 232, 255 pp.

Waterhouse, J. B.

1960. Some Carnian pelecypods from New Zealand. Roy. Soc.
New Zealand, Trans., vol. 88, part 3, pp. 425-442, pls.
19-25.

Whitfield, R. P. ; '
1885. Brachiopoda and Lamellibranchiata of the Raritan Clays

and Greensand Marls of New Jersey. U. 8. Geol. Sur.,
Mon. 9, 269 pp., 35 pls.

. . Justus Perthes,

Ks s
a Tl ¥ y

bo

*
PALAEONTOGRAPHICA AMERICANA (VI, 39)
EXPLANATION OF PLATE 20
Baluchicardia

Figures Page

1. :¥. (B) beanmeontt (d’Archate and Haime): 2... eee
AMNH complete individual, length 26.5 mm, Sutley, near Kulla, Punjab region of
India, Late Cretaceous. la. Left exterior. 1b. Anterior.

2 P= CB.) eeeperta. Garde 255. is ciccsscsacececcssnnctecessnsensnpargvorbacateenavntn onlneed As ndsectnstioh= Setanta eee 83
Holotype, complete individual, USNM 352268, length 44.6 mm. USGS sta. 3180,
Tehuacana Member, Kincaid Formation, early Paleocene. 2a. Anterior. 2b. Right
exterior.

3... (B:) frameescae Gardiner «io o0ccccsqteseecnsssscenceostnrncs 8+
Holotype right valve, USNM 372925, length 28.0 mm.
tion, early Paleocene. 3a. Exterior. 3b. Interior. 3c. Anterior.

Ae WCB.) spin ntes Garner Sc enc csiss checnce sees ese ae eee RE cae coer 84+
Holotype, right valve, USNM 352269, length 32.0 mm. USGS sta. 3180, Tehuacana
Member, Kincaid Formation, early Paleocene. +a. Exterior. +b. Anterior. +c. Interior.

537. .(B.) maa Gardber —. >. a ee ee 86
Holotype, complete individual, USNM 370922, length 33.8 mm. USGS sta. 4398,
Tehuacana Member, Kincaid Formation, early Paleocene. 5a. Left exterior. 5b. Anterior.

6:.F. (B.) emblicrenatia, 0. SR: SS ee 85
Holotype, left valve imbedded in matrix, USNM 370912, length (est.) 31.0 mm, USGS
sta. 2439, Tehuacana Member, Kincaid Formation, early Paleocene. 6a. Anterior.
6b. Exterior.

7,8. VF. (B.) bulla Dall .......... 86

7. Exterior view of lectotype, left valve imbedded in matrix, USNM 164556, length 23.1
mm. 8. Left exterior view of paralectotype, complete individual, length 34.0 mm. USNM
645691. USGS sta. 3309, Wills Point Formation, late Paleocene.

Plate 20

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Plate 21

PALAEONTOGRAPHICA AMERICANA, VOL. VI

RN ee

an

oS Oe 6 & ay -
9

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP

EXPLANATION OF PLATE 21

Baluchicardia
Figures Page
1. V. (B.) bulla Dall 86
Hypotype, complete ind USNM 370913, length 33.8 mm. USGS s
Wills Point Formation, late Paleocene. 1a. Left exterior. 1b. Anterior. 1c. Left
interior. 1d, Right interior.

ROO AEs Ess) MaLILL COR Ere S2S NN ALLO ore 0995525 n al Secwey Seesctancettet edie olan svssacafstteacasacssvetekavisepeeréssesonnniscseecrosocheczeods 87
2. Holotype, left valve, USNM 164542, length 36.7 mm. USGS sta. 3116. 2a. Ex-
terior. 2b. Interior. 2c. Anterior views, 3. Paratype (lectotype, “var. tripla”), right
valve, USNM 129897, length (est.) 30.9 mm. USGS sta. 287. 3a. Interior. 3b.
Exterior. 3c. Anterior. Matthews Landing Member, Naheola Formation, late Paleocene.

ee mIME ERO ENT AU GCLLIE LS ANN Merce fee Seasons oe ae a ee ce en A PN ok, 88

4. Lectotype, right valve, PRI 137, length 52.3 mm, Greggs Landing, Monroe County,
Alabama. Greggs Landing Member, Tuscahoma Formation, early Eocene. 4a.
Anterior. 4b. Exterior. 4c. Interior. 5. Uncatalogued specimen, ANSP, left valve,
length (est.) 31.0 mm. Bells Landing, Monroe County, Alabama, Bells Landing Member,
Tuscahoma Formation, early Eocene.

123

PALAEONTOGRAPHICA AMERICANA (VI, 39)

EXPLANATION OF PLATE 22

Rotundicardia

Figures Page

1,2. V. (R.) eoa Gardner Ge 90
1. Holotype, right valve, length 22.5 mm. 1a. Exterior. 1b. Interior. 1c. Anterior.

2. Paratype, left valve, interior length (est.) 28.4 mm. Both USNM 370918, USGS
sta. 11914, Kincaid Formation, early Paleocene.

3,4. V. (R.) crenaea Gardner .... 91
3. Lectotype. Right valve imbedded in matrix, length 25.3 mm. 3a. Exterior. 3b.
Anterior. 4. Paratype, left valve, exterior view, length 28.6 mm. Both USNM
370911. USGS sta. 117701, Tehuacana Member, Kincaid Formation, early Paleocene.

5. V. (R.) eutaweolens Warris ........c.cssscececnsescessnnncnsnencserneneneesansnsnss a ere ee = Ey 94
Latex cast of lectotype, external mold of left valve, PRI 1401, length 26.8 mm.
$a. Exterior. 5b, Anterior. Eutaw Springs, South Carolina, Santee Formation, middle
Eocene.

6,7. Ve (Re) flabellum Harris... .--ce-n-nn-cceeceeccnseneeececcneeeecsnessnsencsenesnennnennnennncosnanennecnansnserancnnenanennsanness 92
6. Holotype, right valve, PRI 651, length 22.0 mm. 6a. Exterior. 6b. Interior. 6c.
Anterior. 7. Hypotype, right valve, interior view, USGS 6058 No. 2, length 21.5 mm.
Smithville, Texas, Weches Formation, middle Eocene.

8,9. V. (R.) rotunda Lea ..s.ssssssssssssssesneccsesseessessnccnsennecnsecssseneanesenessnecsscenssssseneessssesscnansuenessneseeneecassesesess 93

8. Paralectotype, left valve, exterior view (funiculate ornamentation), ANSP 5268,
length 25.0 mm. 9. Lectotype, right valve (funginate ornamentation), ANSP 5267,
length 24.5 mm. Claiborne Bluff, Alabama, Gosport Formation, middle Eocene.

Plate 22

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Plate 23

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Figures

1,2.

3-6.

7,8.

. V. (R.) carsonensis Dall sve

AMERICAN ALTICOSTATE VENERICARDS: [IEASLIP

EXPLANATION OF PLATE 23

Rotundicardia

PAU AD ULETELL GG Eclita sepecescayctsuoce cuss: exanststtea¥anesecnnes stuceouasssoevscesdassntsasascisesvysesdascsevsvvceeveeveheousbaevectateras
1. Lectotype, right valve, interior view, ANSP 5267, length 24.5 mm. 2. Paralectotype,
right valve, ANSP 5268, length 25.0 mm. 2a. Interior. 2b. Anterior. Claiborne Bluff,
Alabama, Gosport Sand, middle Eocene.

V. (R.) diversidentata diversidentata Meyer
3. Holotype, right valve, ASM 74, length 17.5 mm. Jackson, Mississippi. 3a. Ex-
terior. 3b. Interior. 3c. Anterior. +. Hypotype, left valve, exterior view, USNM
136672, No. 5, length 20.6 mm. Jackson, Mississippi. 5. Hypotype, (holotype, /’.
“praecisa”), right valve, length 13.0 mm. 5a. Exterior. 5b. Interior. 6. Hypotype (/’.
“praccisa’”’), left valve, interior view, length 8.2 mm. Both 5 and 6 USNM 138638.
USGS sta. 2232, Cleveland County, Arkansas. All from Moody’s Branch Formation,
late Eocene.

eee (IR) CLIENTELE TLECCRILGRS DAL GNC) NO alll ieavcusee<ccsosccas.tccsteercovsetn eastteereessticeeeerne vector evaemere aie
7. Hypotype, left valve figured by Richards and Palmer (1953), FGS I-7533, length
12.5 mm. 7a. Exterior. 7b. Interior. 7c. Anterior. FGS loc. L-93, Levy County, Florida,
Inglis Member, Moody’s Branch Formation. 8. Holotype, complete individual, left
exterior view, USNM 164548, length 15.8 mm. USGS sta., Martin Station, Florida,
Ocala Formation. Both late Eocene.

9. Holotype, right valve, length 13.2 mm. 9a. Exterior. 9b. Interior. 10. Paratype,
left valve, length 16.9 mm. 10a. Interior. 10b. Anterior. 10c. Exterior. Both USNM
140693. USGS 2633, Carson’s Creek, Mississippi, Red Bluff Formation, early Oligocene.

Page

93

95

96

97

PALAEONTOGRAPHICA AMERICANA (VI, 39)

EXPLANATION OF PLATE 24

Claibornicardia

Figures Page

1,2.

Vs (Gs) linguinodifang) (n. Spaee se ee So eh ed reper ee Se Se a PO ean 99
1. Holotype, right valve, USNM 644318, length 13.6 mm. la. Exterior. 1b, Interior. lc.
Anterior. 2. Paratype, left valve, USNM 644319, interior view, length, 10.5 mm.
USGS 3099, Bashi Formation, early Eocene.

54. Vi (G) fcoloradonts: texalama, Sax Qer s,.ncnorasertoosedieavenscetesre tet iupensscerteceereyegccte- tare Oar Meee ek ee oa 101
3. Lectotype, right valve, length 17.3 mm. 3a. Exterior. 3b. Interior. 3c. Anterior.

4. Paratype, left valve, interior view, length 17.1 mm. Both ANSP 9860. Black shoals
(Burleson Bluff), Burleson County, Texas, lower Weches Formation, middle Eocene.

SIGs an (Ga) MCOLOTAAONES COLOTACOILS EXAUTIS | eecacdencasrntceqrsrsnvr er ar mearaneetieeeetatar ter mee ete eneeme erence eee 100
5. Holotype, left valve, PRI 652, length 21.0 mm. 5a. Exterior. 5b. Interior. 5c. Anterior.
Smithville, Texas, middle Weches Formation, middle Eocene. 6. Paratype, right
valve, PRI 650, length 13.7 mm., interior view. Wautubbee, Mississippi, Wautubbee
Formation, middle Eocene.

FOV (Ge) AN GPAQUATOLAES, | TS Pisce cr cep sass aecvancvenascgaxevsscgasaneseetonencsanatyestesatl oy ct saya oneeete Ramer t Tea atts eee 104
Holotype, right valve, USNM 644320, length 24.8 mm. 7a. Exterior. 7b. Interior. 7c.
Anterior. USGS 9255, Weches Formation, middle Eocene.

8. V. (C.) trapaquara Harris =

Hypotype, anterior fragment of left valve, showing ornamentation, BEG
original length about 14.0 mm. BEG 26-T-1, Stone City beds, middle Eocene.

Plate 24

PALAEONTOGRAPHICA AMERICANA, VOL. VI

eet il

ee W465

———

—— et TOTS
— Dy “4

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 25

Figures

1-3.

4-7.

8-11.

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP

EXPLANATION OF PLATE 25

Claibornicardia

Weal G2) MPU LAP NECA Peco AAGUN Se ces or vasa cts g ec nane cutee stec scents dente sen tasvecvenev ss onisnsna cacdeeerasnevarnersaannecananseuiicn
1. Holotype, right valve, BEG 20502, length 11.8 mm. BEG loc. 197-1-5. la. Exterior.
1b. Interior. 2. Hypotype, left valve, interior view, BEG 20499, length 17.0 mm.
3. Hypotype, right valve, BEG 20803, length 18.0 mm. 3a. Exterior. 3b. Anterior.
Hypotypes from BEG loc. 26-T-1, all specimens from Stone City Beds, middle Eocene.

Pea Ge) MIPALCHELO CHES PEM ATULS we <covscces es cetator acess Senscne anos testuteeterat de ateasesacenesavar cosucevet evesveuvenaxet sev=te
4. Holotype, left valve, PRI 666, length (est.) 19.0. 4a. Exterior. 4b. Interior. 4c. An-
terior. 5. Hypotype, left valve, USGS 13274, No. 3, length 17.8 mm. 5a, Anterior. 5b.
Exterior. 6. Hypotype, right valve, interior view, USGS 13274, No. 1, length 21.2 mm. 7.
Hypotype, left valve, USGS 13274, No. 5, length 14.8 mm. 7a. Exterior. 7b. Anterior.
7c. Enlargement of posterior hinge plate and adductor, showing isolated position of
posterior pedal retractor just above adducter. Figs. + and 5 probably males, 6 and
7 probably females with heavier, more inflated shell. Holotype from Natchitoches, La.;
hypotypes from USGS sta. 13274, San Augustine, Texas, Weches Formation, middle
Eocene.

V. (C.) perantiqua Conrad
8. Latex cast of fragment of left valve showing ornamentation,
No. 4, length of fragment, 20 mm. 9. Latex cast of lectotype, right valve, exterior
view, ANSP 19393, No. 2, length 25.0 mm. 10. Latex cast of right interior, AMNH
9723/2, No. 2, length 26.0 mm. 11. Latex cast of left interior, AMNH 9723/2. No. 3,
length 25.0 mm. Lectotype from Burlington County, hypotypes from Monmouth
County, New Jersey, Shark River Formation, middle Eocene.

102

Figures

1,2.

3-5.

a Ga ali COSt ase, Mm CONTA GIA enc ccenesecntee eee ene eteee reese ree SME etree are

PALAEONTOGRAPHICA AMERICANA (VI, 39)

EXPLANATION OF PLATE 26

Claibornicardia

Ws CG) Wl ara china ts Gorn eek hea, a aes nese pegs en case cece
1. Lectotype, a right valve, ANSP 30500, length 15.3 mm, la. Exterior. 1b, Interior. 1c,
Anterior. 2. Paralectotype, left valve, interior view, same catalogue number, length
(est.) 14.8 mm. Vance’s Ferry, South Carolina, Santee Formation, middle Eocene.

F.. (G.) complexicosta Meyer and ‘AlGricht «oo oon. cstcccesmnnsne-oconnee ro sonssee0- one censansneverseenrensesners=r
3. Holotype, immature left valve, formerly JHU 1362, now USNM 645940, length 8.9
mm. 3a. Exterior. 3b. Interior. 3c. Anterior. Wautubbee, Mississippi, ‘“Wautubbee
Marl,” middle Eocene. 4. Hypotype, right valve, BEG 20604, length 20.5 mm. 4a.
Exterior. 4b. Anterior. 4c. Interior. 5. Hypotype, left valve, interior view, BEG
Miss-51, No. 3, length 15.7 mm. Both hypotypes from BEG Miss-51, Wautubbee,
Mississippi, Archusa Member, ‘“Wautubbee Marl,” middle Eocene.

6. Hypotype, left valve, exterior view, USGos 6087, No. 4, length 22.2 mm. 7. Hypo-
type, right valve, interior view, USGS 6087, No. 6, length 19.8 mm. Both from USGS
6087, Gopher Hill, Alabama, Gosport Sand, middle Eocene. 8. Hypotype, left valve,
(figured holotype of /’. “transversa”), ANSP 5253, length 47.7 mm. 8a. Exterior. 8b.
Anterior. 8c. Interior. 9. Hypotype, right valve (/’. “transversa” figured paratype),
interior view, ANSP 5254, length (est.) 45.7 mm. 10. Hypotype, left valve (figured
holotype of V’. “sillimani”), ANSP 5258, length 36.5 mm. 10a. Exterior. 10b. Interior.
11. Hypotype, right valve, AMNH 9950/1, No. 2. Length 49.8 mm. 8-11 from Claiborne
Landing, Alabama, Gosport Sand, middle Eocene.

PALAEONTOGRAPHICA AMERICANA, VOL. VI

:
{
.

Plate 27

PALAEONTOGRAPHICA AMERICANA, VOL. VI

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP

EXPLANATION OF PLATE 27

Clatbornicardia

Figures Page

Tel eal (Cpa Tt Bs1 44 200) D1 | [i RRO hes ae eee tee een ea eet Bene eats eee eh es, EL eT as
Holotype, left valve, USNM 164626, length 39.6. la. Exterior, 1b. Exterior of smallest
recognizable growth stage. 1c. Interior. 1d. Anterior. USGS 393, Conecuh County,
Alabama, ?Glendon Formation, ?middle Oligocene.

Glyptoactis

2. Holotype, left valve, USNM 498538, length 22.3 mm. 2a. Exterior. 2b. Interior.
2c. Anterior. USGS 14204. 3. Hypotype, left valve, USGS 14779, No. 5, length 24.5
mm. 3a. Exterior. 3b. Interior. 3c. Enlargement of dentition showing flexion in lunule
over anterior cardinal. 4. Hypotype, right valve, interior view, No. 2, length 15.6 mm.
Hypotypes from USGS 14779. All from vicinity of Waynesboro, Mississippi, Chick-
asawhay Formation, late Oligocene.

Sle 1 MC CID) TRIVEDI LLL FaT PN SCO goa a en ae LP SEC
5. Holotype, right valve, USNM 353227, lengih 31.2 mm. Sa. Exterior. 5b. Interior.
Sc. Anterior. USGS 10655. 6. Hypotype, left valve) AMNH 1059, No. 9a, length 25.4
mm., interior view. Both from Silverdale, North Carolina, “Trent” (Silverdale)
Formation, early Miocene.

TR MTA (Gia) IS ETTLG OS LeZoN ELON PS LADD ao ot stncng soc rmct sc ctet canna eee ee SER cone nae Pe sae Seabe Sch sncc Sa ceuch des eemuasse
7. Hypotype, left valve, ANSP 10640, length 32.7 mm. 7a. Exterior. 7b. Anterior. 8. Left
valve, interior view, ANSP 10640, length 25.7. Ballast Point, Florida, Tampa Forma-
tion, early Miocene.

112

112

113

129

130

Wes:

. V. (G.) impendeocosta, n, sp.

. V. (G.) chelomodonta, n. sp.

PALAEONTOGRAPHICA AMERICANA (YI, 39)

EXPLANATION OF PLATE 28
Glyptoactis

(G.) serricosta Heilprin sp shar angel canncBtbeea true Seeman reeeee eae
ge AER right valve, interior view, USNM 165192, No. 4, length 22.8 mm. Ballast
Point, Florida, Tampa Formation, early Miocene.

BAG) gibbentimboratay ny YS pi cecrsec nce acces ere een

: Holotype, left valve, USNM 644323, length 33.2 mm. 2a. Exterior. 2b. Interior, 2c.

Anterior. USGS 21278, Silverdale, North Carolina, “Trent” (Silverdale) Formation,
early Miocene.

Holotype, left valve, USNM 644321, length 25.2 mm. 3a. Exterior. 3b. Interior, 3c.
Anterior. USGS 22287, Haywood Landing, North Carolina, “Trent” (Silverdale)
Formation, early Miocene.

V. (G.) himerta Dall ...........

Holotype, complete individual, USNM 164555, length 52.0 mm. Anterior. 4b. Left

exterior. 4c. Right interior. 4d. Left interior. USGS 2646, Oak sears Florida, Oak
Grove Formation, early Miocene.

Holotype, left valve, USNM 644324, length 21.1 mm. Sa. “Exterior, 5b. Anterior. 5e.
Interior. 5d. Enlargement of dentition showing lunule flexion over point of anterior
cardinal and extra dental] element on the posterior cardinal. USGS 18362, Walton
County, Florida, Shoal River Formation, middle Eocene.

114

115

117

Plate 28

PALAEONTOGRAPHICA AMERICANA, VOL. VI

“ee
——

OR OE Ms of
ae St

; ds 9

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 29

Figures

AMERICAN ALTICOSTATE VENERICARDS: HEASLIP

EXPLANATION OF FIGURE 29
Gly ptoactis

1. V. (G.) hesperide Gardner ..............--.- ee ee ee eg ene 117

5,6.

7,8.

Holotype, right valve, USNM 372890, length 49.2. 1a. Interior. 1b. Anterior. 1c. Ex-
terior, USGS 10603, Walton County, Florida, Shoal River Formation, middle Miocene.

ly (AGED! UPL Ly OEMS Bos oe es a a GE ee ee ee eee Oo ee a eo

2. Holotype, complete individual, interlocked valves, USNM 114730, length 41.2 mm.
2a. Left exterior. 3b. Anterior. 3. Hypotype, right valve, interior view, USNM 114730,
No. 2, length 40.4 mm. 4. Hypotype, left valve, interior view, USNM 114730, No. 13,
length 43.7 mm. All from USGS 2213, Calhoun County, Florida, Chipola Formation,
middle Miocene.

1 (RM AQUI NEAT Ss 01 Oe eRe eR Pr PR a OS aR eR ee ee
5. Lectotype, right valve, USNM 497976, length 43.3 mm. 5a. Exterior. 5b. Interior.
Sc. Anterior view. 6. Paratype, left valve, interior view, same catalogue number,
length (est.) 30.5 mm. Both from USGS 14075, Olga, Florida, late Miocene.

Pan Kae MOM SS LOO Staten Gris SOWELD YD litt co cete ee pc eee eee ote ee Gen eee en
7. Hypotype, right valve, USNM 517572, No. 1, length 47.3 mm. 7a. Exterior view
of uncoated shell showing color pattern of dark red, tan, and white. 7b. Interior view,
coated to resemble fossil material. 8. Hypotype, left valve, coated to resemble fossil
material, USNM 517572, No. 2, length 34.9 mm. 8a. Exterior. 8b. Anterior. 8c. Interior.
Collected between Manglar Alto and Manta, Ecuador. Recent.

116

119

131

INDEX
NUMBER 39

Note: Light face figures refer to the page numbers. Bold face figures refer

to the plate numbers.

A

acuticostata,

Venericardia 55, 98,110
Alabama Ferry,

Texas 104
Alabama Geological

Survey 109
Alabama River 88, ie
Allentown,

Alabama 88
Alpine Miocene 98

alticostata, V.
(Claibornicardia) 26 55,
62, 63, 65, 67, 69, 70,
72, 78, 79, 88, 98, 99,
102, 106, 107, 108,
109, 110, 113, 115

Alum Bluff
Group 115, 117
alveana,
Venericardia 82
amplicrenata, V.
(Baluchicardia) .20 74,
75, 85, 86
Archusa Marl Mem-
ber, Wautubbee
Formation 93, 105
Argyle, Florida 117
Attayaea Bayou 100
Austin Quad-
rangle, Texas 87
B
Baileys Ferry,
Florida .. 110, 116
Ballast Point,
Florida .......... 112, 113
Baluchicardia,
Venericardia 56, 61, 69,

70, 71, 74, 75, 77, 79,
82, 83, 84, 85, 87, 89,
90, 92, 98, 99, 107, 110
Barnwell
Formation 96
Bashi Marl Mem-
ber, Hatchetigbee

Formation . 77, 89,
98, 99, 101
Bastrop County,
Texas . Dy tls Oly
90, 92, 100, 105
beaumonti, V.
(Baluchicardia) . 20 67,
75, 82, 84
Bells Landing, Ala. 89
Bells Landing Marl

Member, Tusca-

homa Formation 89
Bell’s old mill site,

Florida . ; 117
Bernard, F. 60, 61
Bibb County,

Georgia 96
Bibora Creek,

Texas 86
Bibora tank,

Texas . ; 86

Bienville Parish,

Louisiana .... 105
Black Creek Branch,

Prairie Creek,

Alabama 88
Black Shoals , 101
blandingi, V. (Clai-

bornicardia) 26 = «78,

79, 107, 108
borealis, V. (Cyclo-

cardia) 63, 64, 68,

70, 82
Brazos County,

Texas 77, 92, 104
Brazos River,

Texas 101, 104
“brittoni,” V.

(Claibornicardia) 103
bulla, V. (Baluchi-

Cardia)! .2.0...20/ 21769)

74, 75, 83, 84, 88, 99
Bunker Hill, La. 95
Burleson Bluff,

Texas .... ; 101
Burleson County,

Texas 77, 101,

104
Burlington County,

New Jersey 103
Byram Marl

Formation 109

Cc
Calcaire Grossier,
France . ... 82,98

Caleaire jaune de
la Chaine d’Hala,

Indias 2 83
Caldwell Parish,
Louisiana ee, 95
Calhoun County,
Florida 110, 116
Caloosahatchie
River, Florida .... 118
Camden, Alabama 88
Camp Wheeler,
Georgia 96
cancellata,
Chione , 118
Carditamera 79, 109,
110, 113
Carson’s Creek,
MISS: etoehie 97, 98
carsonensis, V. (Ro-
tundicardia) ... 23 72,
76, 77, 90, 96, 97, 98
Gasey, AP aie eee 97
Castle Hayne
Formation 79, 109

Cedar Creek, Texas 104
Centre Hill, South

Carolinas 94
chelomodonta, V.
(Glyptoactis) 28 ~=80,

81, 116, 117, 118

Chickasawhay Marl
Formation . Pak)
TY, 2, ay

132

Chickasawhay
River, Mississippi 97
Chipola Marl
Formation .... 80, 110,
115, 116, 118
Chipola River,
Florida 110, 116
Choctawhatchie
Formation ‘ 118
Claiborne Bluff,
Alabama 712, 89,
93, 98, 99, 107, 108
Claibornicardia 2 DO; Gly

69, 70, 71, 72, 75, 77,
78, 79, 89, 90, 98, 99,
100, 102, 103, 104,
105, 106, 107, 110,

111
Clarke County,
Alabama ; 109
Clarke County,
Miss. . : Theos
95, 100, 105

Clayton Formation 75, 84
Cleveland County,

Arkansas ae. 95
Clifton Ferry,
Alabama 88
Colliers Ferry,
Texas nae 23 101
Colorado River,
Texas i 87, 90,
92, 101

coloradonis, s.1l., V.
(Claibornicardia) 24 68,
79, 99, 100, 101, 102,

103
coloradonis, V.
(Claibornicardia)
coloradonis 24 63,

77, 78, 79, 100, 101,
102, 104, 106, 107,

108

complexicosta, V.
(Claibornicardia) 26 67,
70, 74, 78, 101, 104,

105, 106

Conecuh County,
Alabama .. 79, 109
Conrad, T. A. . SG ERGEh
72, 103, 107

Cook Mountain

Formation .... Pr einoas
100, 105
Cooke, C. W. . _ 73, 109
Cooke, O. E. .... : 87

crassicostata, V.
(Glyptoactis )......29 57,
80, 81, 114, 119
crenaea, V.

(Rotundicardia) .22 75,
76, 89, 91, 92, 94
Crockett Member,

Yegua Formation 93
Currie EsiD: one 65
Cyclocardia .............. 63, 69,

82

D

Dale’s Branch,
Alabama ‘a 88
Dall, W. T. .... ba) DO68,

72, 82, 89, 95, 96, 97,
98, 106, 109, 113,

115, 116
Danian, India, Iran,

Africa, <s.:- 75
Davies, A. M. ..... 57
Decatur, Miss. bs 105
diversidentata, s. 1.,

V. (Rotundi-

cardia) 23 ~—«463,

“71, 74, 76, 77, 90,
93, 94, 95, 96, 97
diversidentata, V.

(Rotundicardia)
diversidentata ... 23, 63,
74, 76, 95, 96
Douville, H. ............ 61
Durham, J. W. . 81
E
Eagle Pass, Texas .. 86
Echinochama ...... 67
edulis, Ostrea .... 68
Enterprise,
Mississippi .......... 100
eoa, V.
(Rotundicardia) .22 75,
76, 90, 91, 93
Escambia County,
Alabama ....... 109

Eucheanna, Florida 117
Eutaw Springs,
South Carolina 94
eutawcolens, V.
(Rotundicardia) .22 75,
76, 89, 91, 94, 95

Eutawville, S.C. 94, 95,
106
Evan’s (Myrick’s)
apiary, Texas 84, 85
F
Farmingdale,
New Jersey . 93, 103
Fayette Formation 96

flabellum, V.
(Rotundicardia) 22 76,
7

7, 92, 93
fornicata, Crepidula 68
Foster, V. M. .......... 93
francescae, V.
(Baluchicardia) ..20 70,
74, 75, 84
Frio River, Texas... 84,85
“fungina,” V.
otendicaed a)
rotunda ............ 94
“funiculus,” V.
(Rotundicardia)
ROCUNG Ay gece 94
G
Gabb, W. M. .......103, 104
Gardner, Ae * TPA

84, 85, 86, 87, 91, 93,
96, 101, 115, 116, 117

INDEX

Gardner, J., and

Bowles, E. 5 bbe Gl
77, 84, 89

Garland Creek,

Mississippi : 95
“garlandica,”

V. (Rotundicardia)

diversidentata 96
gibbumbonata,

V. (Glyptoactis) 28 79,
80, 81, 114, 115, 116
Gillette’s farm
marl pits, North

Carolina ibhhah ee
114
Glans .. eat 111

Glendon (Lime-
stone) Formation 79, 97,
98, 109
Glyplonci= ee DDG;
1, 69, 70, 71, 77, 79,
o 82, 89, 110, 111
Gopher Hill, Ala. 72, 93,
107, 108

Gosport Sand

Formation 71, 72,
77, 79, 89, 90, 93, 98,
106, 107, 108, 110,
113

Grant Parish,

Louisiana . 95
Graveyard Hill,
Ala. ee 88

greggiana, V.
(Baluchicardia) 21 67,
74, 75, 87, 88, 89,
91, 94, 99
Greggs Landing,
Alabama : 89
Greggs Landing
Marl Member,
Tuscahoma For-

mation ... ; 89
Grignon, France 82
Guadalupe Couey.

Texas 91

H
hadra, V. (Glypto-
actis) . eh) ah

56, 57, 80, 81, 110,

phe 113, 114, 115,

116, 117, 118

Harris, G. D. 55, 56,
69, 72, 89, 90, 92,

94, 95, 96, 99, 101,

102, 105, 106
Hawthorne For-

mation ...... 116

Haywood Landing,
North Carolina 114
Heaslip, W. G. 56, 72,
84, 88
Heilprin, A. 113

hesperia, V.
(Baluchicardia) 20 74,
75, 83, 84, 85, 86,

7,117

hesperide, V.
(Glyptoactis) 74) atte
70, 75, 79, 80, 81,
85, 98, 116, 117

133

Hickory,

Mississippi . : 100
Hillsborough

County, Florida .. 113
himerta, V. (Glypto-

actis) . 28 655,

57, 79, 80, 81, 111,
112, 115, 116
Hines County,

Mississippi 95
Hiwannee,

Mississippi 95, 97
Hondo River, Texas 91
Houston County,

Texas . : 104

imbricata, Veneri-

Cardia «cc... 82, 98
Imbrie, J. . 62
impendeocosta, V.

(Glyptoactis) ......28 79,

80, 81, 112, 114
Inglis Formation 96, 97

J
Jackson,

Mississippi . . 95, 96
Jackson, R. T. . : 67
“jacksonensis,” V.

(Rotundicardia) .. 71
Johnson, H. S., Jr. .. 107
Johnson, I. C. 109
Jones County,

North Carolina 114

K
Kaufman County,

Texas 91

Kellum, L. B. 72, 109,
110

Kincaid Formation,
Texas 57, 75,
84, 85, 86, 90, 91

“kingi,” V.

(Rotundicardia)

flabellum .... 92
Knox Hill,

Florida 117

L
Lake Marion,

South Carolina 106
Lamarck, J. B. de 55
Laredo Formation 77, 93
Lea, I. 5D, 2s

108
Lee County,

Florida 118
Leuroactis,

Florida .. 55, 82
Levy County,

Florida 96

linguinodifera, V.
(Claibornicardia) 24 69,

77, 78, 89, 99,

100, 10i

Lisbon Formation — 93, 101
TeISOM eee : 66

Littig Member,

Kincaid
Formation 86, 91
Loosanoff, V. 65, 68
Lowe, E. N. .... 100
M
Maestrichtian,
India, Iran,
Africa 75
Mansfield, W. C. 72, 112,
113, 118
Marianna Lime-
stone Formation 109
Marion County, Fla. 96
Martin Station,
Fla. 96
Matthew’s Landing,
Ala. 88

Matthew’s Landing
Marl Member, Na-
heola Formation 77, 87,

88
Maverick County,

Texas : 85, 86
Mayr, E. 87
McBean Formation 79, 93,

103, 107
Medina County,

Texas 84, 91
Mediterranean 117
Megacardita 98, 117

megastropha, V.
(Strephocardia) 57, 81
mercenaria,

Mercenaria 57, 65,
68
Meridian,

Mississippi 100
Meyer, O. 74
Meyer, O., and

Aldrich, T. H. 74, 105
Midway Group 87, 88,

98, 117
Mill Creek,

Alabama 99
moa, V. (Balu-

chicardia) 20 = 74,

75, 85. 86
Monmouth County,

New Jersey .... 93, 103
Monroe County,

Alabama ore

79, 89, 93, 98,
107, 112
Montgomery,

Louisiana . 95
Moodys Branch

Formation 95
Moore, E. J. 107
mortoni,

Lepidocyclina 97
Mt. Lebanon,

Louisiana ; 105
Munier-Chalmas,

OB (Epes LS OOO
Murder Creek,

Alabatna es. .ck 109

N
Nacogdoches
County, Texas 100

INDEX

Naheola Formation 75, a
nasuta, V. (Clai-
bornicardia) ‘27 = GT,
70, 78, 79, 97, 98,
108, 109, 110, 113
Natchitoches, La. 102
natchitoches, V.
(Claibornicardia) 25 61,
77, 78, 101, 102,

103, 108
New Albany,

Mississippi 84
New Bern, North

Carolina 107, 109
New Fountain,

Texas 91
Newton,

Mississippi 93, 100
Newton County,

Mississippi 100, 105
Nicol, D. 67
nodiferas, s. l., V.

(Glyptoactis) 28, 63,

67, 81, 111, 112,
113, 114, 115

nodifera, V. (Glypto-
actis) nodifera 27, 63,
69, 79, 80, 111, 112,

114, 115, 116
ie)
Oak Grove
Formation 79, 80,
LIZ IS aa
Oak Grove, Florida 115
Ocala, Florida ... 96,97
Ocala Group .... 77, 96,
97, 109
Okaloosa County,
Florida ‘ 115
Old Garfield,
Texas 87
olga, V.
(Glyptoactis) 29 ~=80,
81, 116, 117,
118, 119
Olga, Florida 118
Onslow County,
North Carolina 114
Orangeburg,
South Carolina | 93, 107,
109

Orangeburg County,
South Carolina 93, 94,
103, 106, 107, 109
68

Orton, J. H. .
Ouachita River,

Louisiana ; 95
Owen, G. 66
P
Pacificor ; 56
Paleocardita 61

Palmer, K. V. W.,
and Brann, D.C. 88, 89
Panamanian
TSthmiis) ...:..- 81
Panhandle, Florida 116, 117
Paris Basin,
VANCE Fak... nces 55, 82,
98, 110

134

Pecten 66
perantiqua, V.
(Claibornicardia) 25 77,

78, 103, 104
Percilla, Texas 104
Perdue Hill,

Alabama 112
planicosta,

Venericardia 55
Pleuromeris 82
Plummer, F. B. 92, 93,

96, 100

“praecisa,” V.
(Rotundicardia) Ty 26}

93, 95, 96
Prairie Bluff

Formation 75
Prairie Creek,

Alabama 88
Provencal,

Louisiana 102
Pteromeris 82
Puri; os: 57, ta;

115, 116, 117
R
Red Bluff,

Mississippi 97
Red Bluff

Formation wanies MG
Renfroe, C. A. 73
Renick, B. C., and

Stenzel, H. B. 92, 93,

100
Richards, H. G. 97
Richards, H. G., and

Palmer, K. V. W. 73
Robertson County.

Texas etis or 92 a 04
rotunda, V.

(Rotundicardja) 22, 23, 55,
56, 67, 68, 69, 71,

72, 75, 76, 77, 88,

90, 91, 92, 93, 94,

95, 97, 98, 100, 101,

103, 104, 106, 108,

109

Rotundicardia 56, 61,
68, 69, 70, 71, 75,

76, 89, 90, 91, 92,

93, 100
Rutsceby Rake ys 75
Rutsch, R. F. and
Schenck, H. G. . 56
Ss
St. Maurice,

Louisiana ....... 93
St. Maurice horizon 106
St. Stephens Bluff,

Louisiana ..- 93, 107
Saline Bayou,

Louisiana .. 93, 100,

102
Saline River,

Arkansas: .5 cues 95
San Augustine,

MEXAS) comascunts he 102
sandiegoensis,

Gavia eee ae 111
Santee-Cooper Dam,

South Carolina .... 106

Santee Limestone
Formation . 75, 79,
91, 94, 106, 107

Santee Reservoir,

South Carolina 106
Santee River,

South Carolina 106
scabricostata,

Venericardia 63, ye
Schmidt, F. C. 82

Seine et Oise, De-
partment de,

France 82
Sepulga River,
Alabama i... 109

serricosta, V.
(Glyptoactis) .. 27, 28, 67,

79, 80, 81, 109,
111, 112, 113, 114,
116
Shark River Marl
Formation ees LOS
Shoal River
Formation omen,
75, 79, 81, 116, 117
Shubuta, Miss. . 95, 97
“Silex beds” (Tampa
Limestone) ..... 113

“sillimani,’ V.
(Claibornicardia) 62, 63,

69, 72, 79, 106,
107, 108
Silverdale, North
Carolina . sla a a es
114
Smithville, Texas 92, 93,
100, 101
Stenzel, H. B.,
Krause, E. K., and
Twining, J. T. 56, 72,
73, 84, 98, 104,
105, 106, 108, 110
Stewart, R. B. .. lap lily
111

Stephenson, L. W. 82, 87
Stone City Bluff,

Texas ........ 104, 105
Stone City
Formation 78, 101,
104, 106
Strophocardia 81
“subquadrata,”
Cardita . 103
T
Tampa Bay,
Florida .... 113
Tampa (Limestone)
Formation ........... 79, 80,
112, 113

Taylor Mill Creek,
IMSS Face hone, 112
Tehuacana Mem-
ber, Kincaid
Formation .. 75, 84
85, 86, 91
texalana, V.
(Claibornicardia)
coloradonis ..... 24 63,
77, 78, 100,
101, 102, 104

INDEX

Thompson, D’A. W. 65

Tombigbee River,
Alabama . 93, 99,
107

Town Creek,
Mississippi 95
“transversa,” V.
(Claibornicardia) 107, 108
trapaquara, V.
(Claibornicardia) 24 78,
100, 101, 103,
104, 105, 106
trapaquaroides, V.
(Claibornicardia) 24 78,

79, 101, 104,
105, 106
Travis County,
Texas 85, 87
tricolor, V.
(Glyptoactis) 57, 81
Trinity River,
Texas : 104
“tripla,” V.
(Baluchicardia)
wilcoxensis 88
Trent Marl
Formation 79
“Trent” (Silverdale)
Formation 111, 114,
115, 116
Tuscahoma
Formation 75, 88,
89, 91, 94, 99
Tyus Member,
Weches
Formation 104
U
Union County,
Mississippi 84
U.S. Geological
Survey 111, 112,
118
Vv
Vance, South
Carolina 106

Vance’s Ferry,

South Carolina 79, 106,
107
Venericardia 55, 58,

61, 63, 65, 66, 67,
68, 74, 75, 77, 79,

81, 82, 97, 102,
106, 110, 113
Venericor =. 105,06,
58, 82
Verastegui, P. 56
Vernon, R. O. 117
Vicksburg Group 97, 109,
110
vicksburgiana, V.
(Rotundicardia)
diversidentata 23 «63,
F 77, 95, 96, 97
Vince Bluff,
Arkansas ..... ; 95

135

WwW
Wagner Free
Institute 113
Walton County,
Florida 117
Washington County,
Alabama ... 93, 107
Wautubbee,
Mississippi 100, 105
Wautubbee Marl
Formation Leis
93, 100, 101, 104,
105, 106, 109
Wayne County,
Mississippi 77, 79,
95, 97, 112
waynensis,
V. (Glyptoactis)
nodifera 27. ~=«éG63,

77, 79, 80, 81, 111,
112, 113, 114, 117

Waynesboro,
Mississippi 112
Weavers Chute,
Alabama 109
Webb’s Creek,
North Carolina 111
Weches
Formation ... Title tis
92, 93, 100, 101,
102, 104
Wheelock League,
Texas 104
White Oak River,
North Carolina 114
whitei, V.
(Baluchicardia) .20 57,
70, 74, 75, 84, 98, 117
White’s Creek, Fla 117
Whitfield, R. P. 103
Whitsett Member,
Fayette
Formation 96
Wilcox County,
Ala. 88

wilcoxensis, V.
(Baluchicardia) 21 74,
75, 77, 84, 86, 87,

88, 89, 99
Wills Point

Formation 75, 87
Wilmington,

North Carolina 110
Wilson, D. 118
Winn Parish,

Louisiana 93, 100
Woodbine

Formation 82
Woods Bluff,

Alabama 99

Woods Hole Bio-
logical Laboratory 63, 68

‘if
Yegua Formation .. 77, 92,
93

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XLVII.

XLVIII.

XLIX.

Volume I.

II.

III.

IV.

UN Coamthes®)) ee Compa asay Leg Ss tecreseccsaccessnutbanetunbasetessesseonacascoscerspsscascessaneer

Type and Figured Specimens P.R.I.

ING See) ers Sh MD psy (ndios PlSstaccctactrascsicsseateaesscrsencaceseusensscassapasss

Australian Carpoid Echinoderms, Yap forams, Shell Bluff,
Ga. forams. Newcomb mollusks, Wisconsin mollusk faunas,
Camerina, Va. forams, Corry Sandstone.

ING hOB Ver 750 DYiech FSi Elsen csetcesceseracicetscacesiransscesavatarvesscebsneseccssssenee

Venezuelan Cenozoic gastropods.

(UNiog tL 942198) SA27 yp G9 De occssctasssssnecsssccuesesesnacsessesvsvsecorsesiens

Ordovician stromatoporoids, Indo-Pacific camerinids, Missis-
sippian forams, Cuban rudists,

Nios 99-203) S65) pe OSI DIBI crcesssntsesstncensesvastessvarcssushcosunacececeess

Puerto Rican, Antarctic, New Zealand forams Lepidocyclina,
Eumalacostraca.

(Ninny 209) ey S64 pare O Sip pol Setccersces ctecnatt-nestesoctasnesshcassuraveesavrersucsersssee

Venezuela Cenozoic pelecypods

(Nose 205-210) 5419) pps 70) Dl Ss) -sccsssssssvecsccecsssedonsousesscusesuconctnsssears

Large Foraminifera, Texas Cretaceous crustacean, Antarctic
Devonian terebratuloid, Osgood and Paleocene Foramini-
fera, Recent molluscan types.

(ipsa = 2 UVa SF oP Diag OS!) DU Soar ccocascassteuetevosnecceancasenosvssvsepaenea canst

Eocene and Devonian Foraminifera, Venezuelan fossil
scaphopods and polychaetes, Alaskan Jurassic ammonites,
Neogene mollusks.

(Nga 218) Se LOS Re ppg a ples esccsscesneecstsasurbtsnastcntcesecsecorcktaacvonsaceoeceres

Catalogue of the Paleocene and Eocene Mollusca of the
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(UNG aI 19-22%) ot G7 Usp Prem S SP kSsenacteerybncscerssotevancasneccosectecoresseeseercre

Peneroplid and Australian forams, North American carpoids,
South Dakota palynology, Venezuelan Miocene mollusks,
Voluta.

(Noms 225-250) sats S pss 42) ipl See cetoinertssssesorecevanevenemeceseatentsstatrnss

Venezuela and Florida cirripeds, Antarctic forams, Linnaean

Olives, Camerina, Ordovician conodonts, Niagaran forams.

(NOs 31-232) lo 4 2 On DprA Om pi Ssaeacecssntestcercstccrsaecvscuscoseuecncceecsssancor
Antarctic bivalves, Bivalvia catalogue.

AISoare 332250) net 80) iD GS PD Sau icctteesacesceatsrsesacstpstoevosectvessouscisiee

New Zealand forams, Stromatoporoidea, Indo-Pacific, Mio-
cene--Pliocene California forams.

(Nos:237-238) = 488. pps, 45) pls: <scccsosecesscscesusecandsessorsesasearsncscessce

Venezuelan Bryozoa, Kinderhookian Brachiopods

(TNGeaeee S48) Sri TORN ets DS icsserorcnatccrusbnesvanncasetccccossencarcsauedrease

Dominican ostracodes, Texan pelecypods, Wisconsin mol-
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CONGms 26-33) 9 2i PIs 720 DISs: ccesctesccscessenssrencenconsodsecvserscossseccesers

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lusks.

(INGghia4=d7) 2) ATI DD LOL ID18.) iccsssssrscscsstesccccestccorccocersdassroccceres

Tertiary Arcacea, Mississippian pelecypods, Ambonychiidae,
Cretaceous Gulf Coastal forams.

(Cys Crise te ye 2 SOR) eh 60 3} [Bk separ enone et nti eae Ca eee

Lycopsids and sphenopsids of Freeport Coal.

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XXX,

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XXXIV.

XXXV.

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Venezuela geology, Oligocene Lepidocyclina, Miocene ostra-
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zuela, larger forams, new mollusks, geology of Carriacou,
Pennsylvania plants.

(Nos. 177-183). 448 pp., 36 pls. .....ssscscsessesscseenesnsnecsssnsscsnsensonsnsesee

Panama Caribbean mollusks, Venezuelan Tertiary formations
and forams, Trinidad Cretaceous forams, American-Eur-
opean species, Puerto Rico forams.

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PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

VOL. VI

NO. 40

CARBONIFEROUS CRINOIDS OF TEXAS WITH
STRATIGRAPHIC IMPLICATIONS

By

Harre_t L. StRIMPLE AND WILLIAM T. WATKINS

1969

Paleontological Research Institution
Ithaca, New York, U.S.A., 14850

\

STHSUN
( JUL 2v 19/0 )

PALEONTOLOGICAL RESEARCH INSTITUTION

1968 - 1969
SPY Ta) fr ge peneee et een eneen eaee Re epee epee Ree ae A A ott KENNETH E. CASTER
VICE-PRESIDENT .....-.--000-0- ..WILLIAM B. HERoy
SERCH TARY oa ee erasers eee ReBecca S. Harris
DIRECTOR; LUREASURER cassis peacescccseesroe eae nandentrioaapnaee KATHERINE V. W. PALMER
COU NSEN: <a er ARMAND L, ADAMS
REPRESENTATIVE AAAS COUNCIL cicccssscssscsscsscsessaceseccceazooecsesscurcorccercascsenses Davw NIcor
Trustees

KENNETH E. CASTER (1966-1972) KATHERINE V. W. PALMER (Life)

DonaLp W. FisHER (1967-1973) WituiaM B. Heroy (1968-1974)

Resecca S. Harris (Life) AxeEL A. Otsson (Life)

DanizL B. SAss (1965-1971) Hans G. Kucier (1963-1969)

W. Storrs CoLe (1964-1970)

BULLETINS OF AMERICAN PALEONTOLOGY

and
PALAEONTOGRAPHICA AMERICANA

KATHERINE V. W. Patmer, Editor
Mrs. Fay Briccs, Secretary

Advisory Board

KENNETH E, CASTER Hans KuGLER
A. Myra KEEN Jay GLENN Marks
AxeL A. OLsson

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PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

VOL. VI

NO. 40

CARBONIFEROUS CRINOIDS OF TEXAS WITH
STRATIGRAPHIC IMPLICATIONS

By

HarreE.i_ L. StRIMPLE AND WILLIAM T. WATKINS

July 1, 1969

Paleontological Research Institution

Ithaca, New York, U.S.A., 14850

. - s vie® -
A 7 vy ale : a a 7
; : ~at é - ste
Me Seal
' od ¢- ore

bile ‘ser

CONTENTS

PNVRES ACER cose crocs reser elt ctv tsceanvnc ste cicwases ioauav se eeexocatoenas
WMtHO MUCHO en 2icccecroecteeteostens
Acknowledgments __............. aospoas

Contributors ‘aed
Crinoid localities —......
Morphological notes

Terminology
Associated: parasitic Wke) oa iecccececsescsecceecesscosecnscesn 0d asososterenscsasvausenesvereveperoseravatadaesbans 145
Stratierapiase 1rmip lication: Spee cece cce peace seca tence creer tree eres prcerennoecrrenncete 145
Pas Tait MU CEO rower oncom cennnn Seaisecice tate estcd ears tie deccsta teasssacet pp ondebets spracmeicovesircnecee 145
SNL ISSIMONDIANL TNE EU (0 oars ree cae os at ee aaco ta endervannsoeee cp ever reenact tacenrooe 146
CON OTTERS Fa ad OLS, par ete ee a er Pee Ae ay SP nn fr ie on ne 146
OL LA) EE CFLS nay ea eee a er oa aap ens ee eer ry eh ep Be 147
LO ee ETE 5 je ee eee nee open ee nse ne Eien ree ne ees anaes eee 147
PE CVETTE OA AMOI Ss XD ONC CD secon een rece rine osc eres rocmepr serene neers 148
III. Fauna of the Atoka Formation of Oklahoma .......000022------ 149
IV. Fauna of the Atoka Formation in Arkansas ......... ASI
V. Fauna of the Atokan Stage in Missouri ~..............-.-.---. 152

VI. Fauna of the Atokan Stage in the Llano Uplift, Texas
VII. Fauna of the Atokan Stage in Iowa ....

VIII. Fauna of the Atokan Stage in Ohio 0000000. ec seeceercececeeerenececeeeees 154
1X. Fauna of the Atokan Stage of New Mexico and extreme
OR e Mi ks OAD eases cece a crea canctoe iapeoetbvocecprnckoacnacacctien 154
X. Fauna of the Atokan Stage of Wyoming and South Dakota —.... 154
XI. Fauna of the Atokan Stage of Eastern Kentucky |... 154
XII. Fauna of the Atokan Stage of Nevada ....0002.200--cscsee-serceceeeessececereee 154
XIII. Fauna of the Atokan Stage of Western Illinois 2200000000000. 154
XIV. Fauna of the Atokan Stage in Michigan .....00000.02.::ecccccccceceecceeeeee 155
XV. Fauna of the Atokan Stage of the Canadian Arctic 2000000000000... 155
XVI. Fauna of the Atokan Stage of Mexico 2000000000002-1..cccceeceeceeceeeeeeeseeeees 155
VU. Fauna of the Atokan Stage of Spain nu. cccccccocosecsscececrccaneace 156
XVIII. Fauna of the Atokan Stage of Russia 2000000000000. 156
NOSE TYTN TE | 2 fe cee Ree RR ae Sy epee oe Pe 157
E. Desmoinesian Stage .........-2-----seccecrer--- ..158
Mts, Warmeonnricamn Stage aso oases cen ceccnnece ra cserownee one er eee re - 159
Vata rege tates (ued eT a) fe SR seek seen ROD an NE En er a ...160
Clasei Crinoidea M6 r 25 Fo see cocoa ctecsocecanicn eo roa paese ante phat cece tapepce ede MO
Subclass Inadunata Wachsmuth and Spirnger —....-...22-2.2-2:---c--ceccereeeeeeeescecereeerees 160

Order Cladoidea Moore ard Laur 22.00.2200. ceeeceeenencreecreneecececenncnsnencececesnrnee 160
Suborder Dendrocrinoidea Bather ..
Family Cromyocrinidae Jaekel
Family Sellardsicrinidae
Family Ethelocrinidae Strimple
Family Texacrinidae Strimple ..........-2-.--------c--cececeeceensrneseeoree
Braman hy) ASU SUCK AMINA Ae ERAT oom anceps acca sence tere forenoon
RCcANNIS 1 este LOC AMINA NEL NSN YN ces necro ec vp ormerevepe eer
Family Erisocrinidae S. A. Miller .................-..-.--
Family Apographiocrinidae Moore and Laudon ......000000
NAIA WE AMCTAMAICLAG, SUC WOE A NEL pce cee carte case scare cervencn ce nenastpaarmccaaretastiae
Family Agasstozocrinidae S. A. Maller ono 02o ecco ccecseceterorensarcnrerenecnciacnence
Feapmily, Ame Oeritn he MR he casa an coca cence cetera me
Subfamily Ampelocrininae Kirk 022000
Subfamily Paragassizocrininae, new subfamily
Family Cymbiocrinidae, new family ..
Family Corythocrinidae, new family -.......
Family Scytalocrinidae Moore and Laudon .....

Family. Ramulocrimidae, mew family 2062s sco ccceecccecn cree -sacec oes nemeteneeeeoe men eeros 199

Family Blothrocrinidae Moore and Laudon .............---...--2..:s:ssscessesseeeeeenees 201
Family Pirasocrinidae Moore and Laudon ................ Sa pdctdep bene owetcphnsoeetes Soe 203
aml yam rant 2G Wom yn cece eee tm ccs cnee cence naon nae ne eee 216
Order Disparta Moore and Laudon .................-... nasoust padnandecctesitj wie eee eee eee 217
Family Allagecrinidae Carpenter and Etheridge ...........2.2..-..:2:12. cecceeee 217
Subfamily? Allageerinimae, (lore) ooaceaccscccne ce peeceresan-scemeonsancenaenyaarveaanaceeeorebees 217
Subclass Camerata Wachsmuth and Springer .............---.-.::cesesseseeeeeeseseseeeesseeaene 218
Oxder (Monobathxa Moore and! Pavone re cece waren cee sae ee 218
Family Platycrinitidae Bassler
Family Acrocrinidae Wachsmuth and Springer ...........2.....::0-2-0-:-e-eeeeeeeeee 220
Family Dichocrinidae S. A. Miller ..csc. sco: o cerca n co cece se mane neee enenens 221
Subclass Mlexibilia. Zittel) 55.2.5. 25. scx ete seecceccenccecceeomecee cere acer xaene ners opee enero 223
Order Sagenocrinidae Springer, 22552 2e-acsccscckecenseanes cere ee eseaeee ero tere 223
Bamily Synerocrinidae Jaekel, 22 pcseccscs cece eccxn sec cxte ences nase eens 223
Family Lecanocrimidae Springer’ <.22 op ccceeccccrcneecenssesn-ncsasenesesntieget sh eceteeeeeameee 232
Frapmentary, Crimoid! remyayys) 22cee-.n cee ee os one cece ee erences 232
References: ciesccicsscccssscssectzsta tocoraste evans Jato ee ee ee soca ebcaaas-ou onto eeeergeoe en 234

TEXT - FIGURES

1. Chart showing probably stratigraphic relationship of various selected
formations based on known fossiliferous horizons, with ammonoids,
fusulinids and crinoids used as the key fossils -.2......2.....0.:.:0e-eeeeeteseeeee 156

2. Correlation chart of some Middle Carboniferous horizons of Russia ........ 157

CARBONIFEROUS CRINOIDS OF TEXAS WITH STRATIGRAPHIC IMPLICATIONS

Harrecy L. Strimece! anp WILLIAM T. WaTKINS2

ABSTRACT

Crinoids used in this study were collected in Texas and range
from Upper Mississippian into the Missourian Stage of the Pennsyl-
vanian. Most are from the Atokan and Desmoinesian Stages. Six new
families, one new subfamily, 12 new genera, 41 new species, and 1
new subspecies are proposed. Some forms previously only known
from Europe and Asia are reported from the United States, e.g.
Hydiocrinus Trautschold (Russia), Stomiocrinus Wanner (Indonesia)
and Sinocrinus Tien (China). The first calyx of Platycrinites of
Pennsylvanian age is reported. The range of Dichocrinus is extended
into the Pennsylvanian.

A compilation of stratigraphic data is attempted with emphasis
on the Atoka Formation and the Atokan Stage.

INTRODUCTION

The nucleus of the present study consists of three col-
lections made by one of us, Watkins, in Texas from the
Kickapoo Falls Limestone Member and Brannon Bridge
Limestone Member, Millsap Lake Formation, Desmoinesian
Stage and from the Lemons Bluff Member, Marble Falls
Formation, Atokan Stage. A small collection of crinoids was
also procured from what may be the Marble Falls Forma-
tion on Espey Creek in Lampasas County, Texas. Various
specimens were added to the study and some were acquired
from other collectors which has been acknowledged in a
special section entitled “Contributors.” At a late date in the
study, the junior author became aware of two additional
crinoidal exposures in the Llano Uplift of northcentral
Texas, one in the Marble Falls Formation on the Lambert
Ranch near San Saba and the other on the Kurt Zesch
Ranch near Mason. Much effort was expended in making
contacts, in field work, in making exchanges, and in the
preparation of materials. In the course of events there were
also some Mississippian crinoids found associated with large
axes of Archimedes and a radial plate of Pentremites.

When we attempted to correlate various formations it
soon became apparent that some serious compilations of
stratigraphic data were in order. In particular a weakness
Was apparent in the time-rock gap between the Morrowan
Stage below and the Desmoinesian Stage above. The terms
Derryan and Lampasan have been used in a more or less
correlative sense but currently the term Atokan Stage is
generally in favor. We have made an effort to clearly de-
fine the Atokan Formation and thereby establish a firm
basis for the Atokan Stage.

Much information has now become available concern-
ing crinoids of the Atokan Stage and a more comprehensive
understanding attained of crinoids of the Desmoinesian
Stage. A few specimens from Upper Mississippian strata are
included as well as some from the Middle Pennsylvanian

1 University of Iowa, Iowa City.
*San Antonio, Texas.

(Missourian), The range of Dichocrinus, a typically Mis-
sissippian crinoid genus, is extended up into the Pennsyl-
vanian (Atokan). Platycrinites has been known to be pres-
ent in the Pennsylvanian strata from columnals, and many
species are known from Mississippian and Permian rocks,
but the first calyx of Pennsylvanian age is described herein
from North America. The genus Hydriocrinus has hitherto
been restricted to the Moscovian of Russia but is now
known from the Desmoinesian of Texas. The genus Stomio-
crinus is well represented in the Permian of the Island of
Timor but is reported for the first time from the Pennsyl-
vanian of North America as S. conlini, n. sp. The genus
Sinocrinus was originally described from North China and
is now reported from the Big Saline Formation (Atokan)
of Texas. Many other important studies are included. New
taxon consist of six new families, one new subfamily, 12
new genera, 41 new species and one new subspecies.

ACKNOWLEDGMENTS

W. M. Furnish, University of Iowa, made many valu-
able suggestions and criticisms of the section on strati-
graphy as well as identifying many ammonoids used in the
stratigraphic section. We are most grateful to G. A. Cooper,
and Porter Kier, U.S. National Museum, Washington, D.C.,
for making funds available to defray the costs of the plates.
M. L. Thompson, Illinois Geological Survey, Urbana, Illi-
nois, and Garner L. Wilde, Humble Oil & Refining Co.,
Midland, Texas, were most helpful in identifying many
fusulinids used in the stratigraphic study.

CONTRIBUTORS

Mr. and Mrs. Robert L. George, members of the Bexar
County Rock and Mineral Society, San Antonio, Texas,
helped through contributions of specimens and in other
ways. One of the specimens found by them is the holotype
of Erisocrinus georgeae, n. sp., named in their honor, Mr.
George sawed many of the limestone slabs into useable
blocks.

Mrs. JHugh Shearer of Mason, Texas, contributed the
holotype of Sinocrinus shearert, n. sp. which was collected by
the late Mr. Shearer some 30 years ago. Their son, Mr.
Wilburn Shearer, also of Mason, Texas, assisted consider-
ably in field work near Mason as well as contributing many
fine specimens. One of the species involved is named
Mooreocrinus wilburni, n. sp.

Mr. Louis Todd of Lewisville, Texas, generously con-
tributed, by donation and exchange, specimens collected
near Lake Bridgeport in the Graford Formation and from

142 PALAEONTOGRAPHICA AMERICANA (VI, 40)

near Brock, Texas, in the Millsap Lake Formation. A species
has been named Anchicrinus toddi, n. sp.

A fine crown of Hydriocrinus, a genus hitherto only
reported from Russia, has been contributed by Mrs. Lor-
raine Marrs of Waco, Texas. It was collected near Brock,
Texas, from the Millsap Lake Formation. The species has
been named lorraineae. Mrs. Marrs also found many of the
splendid crowns used in the study of Texas crinoids by
Moore and Plummer (1940), which specimens are now
housed in the U.S. National Museum.

Mrs. Paul C. Frost, Cedar Hill, Texas, collected and
contributed the excellent crown of Oklahomacrinus frostae,
n. sp. from Brock, Texas.

Mr. J. F. Conlin of Fort Worth, Texas, a well-known
collector of many years standing, donated cups of the only
species of Stomiocrinus ever to be reported from North
America. The genus is not uncommon in the Permian of the,
Island of Timor, Indonesia. Mr. Conlin collected the speci-
mens from near Lake Bridgeport, Texas, in the Graford
Formation. The species has been named conlini.

Mr. C. B. Lambert, Lambert Ranch, San Saba, Texas,
extended hospitality and assistance to one of us, Watkins,
as well as contributing the excellent crown of Aewxitropho-
crinus lamberti, n. sp. to the study. A small quarry in crin-
oidal rocks of the Marble Falls Formation has been opened
on the Lambert Ranch, but crinoid crowns appear to be
rare.

Mr. Kurt Zesch, owner of the Zesch Ranch near Mason,
Texas, was most cooperative and helpful. The crinoid speci-
mens from the Soldiers Hole Member, Big Saline Forma-
tion (Atokan) were all obtained on the Zesch Ranch. A
species has been named Ulocrinus zeschi, n. sp.

The late Mrs. J. H. Renfroe of Fort Worth, Texas,
collected a few of the crowns from near Brock, Texas, in the
Millsap Lake Formation. They were obtained through ex-
change many years ago. Mrs. Renfroe was a well-known
collector of many years standing. The specimens are now
housed in the U.S. National Museum.

A major contribution of effort made to the study was
by Mrs. Florence Pierce. former Secretary, Department of
Geology, University of Iowa, who typed much of the manu-
script.

A financial contribution was made by the U.S. National
Museum, Washington, D.C. to defray the costs of the illus-
trations.

CRINOID LOCALITIES
I. Kothman Ranch

The crinoid exposure was discovered by Mr. Wilburn
Shearer of Mason, Texas, while deer hunting with bow and

arrow. It is about 2% miles southwest of Mason, Mason
County, Texas, on the Kothman Ranch. Coordinates are
30° 43.5’ N. Lat., 99° 16.0’ W. Long. This is about 5/8ths
of a mile west of the northeast end of the Pennsylvanian
exposure, 400 yards upslope northwest of the Barnett Shale
which is exposed in the creek, and 50 yards south of an
east-west fence line.

Plummer’s (1950) “Geological Map of the Carbonifer-
ous Formations in the Llano Region, Texas” indicates that
the Barnett Shale or Formation and the Chappel Lime-
stone are thin ribbons bordering the northeast side of the
more extensive Pennsylvanian exposure for its full six mile
length. A limestone, oolitic in places, was recognized in the
Barnett Formation by Plummer (1950, p. 34), quote,
“—_in Mason County west of Mason, crystalline, crinoidal,
and in some places oolitic limestone layers came into the
section to replace part of the shale —.”

The crinoids appear to be Chesterian in age, and for
this reason the rocks are thought to be from the Barnett
Formation. Two large axes of Archimedes and the radial
plate of a Pentremites were recovered together with about
15 crowns of crinoids in poor to indifferent preservation.
II. Espey Creek

The exposure is in the creek bed three miles southwest
of Lampasas, Lampasas County, Texas, about 300 yards
downstream, i.e. south and east, from the first road crossing
Espey Creek, A black crystalline crinoidal limestone is ex-
posed but the bed containing the crinoid crowns has been
washed out for several years. One of us, Watkins, is of
the opinion that the horizon is Chappel Limestone, which
is of Osagian age, but the crinoids appear to be no older
than late Chesterian and more likely of Morrowan age.
Plummer (1950, p. 53) reported that the Sloan Member,
Marble Falls Formation, had not been identified in Lam-
pasas County. If this means there is no Morrowan rock
present then the possibility that the fauna is uppermost
Chesterian (post-Pitkin of Oklahoma-Arkansas usage) is
entertained. The senior author is aware of a few crinoid
species from an undescribed fauna in the Imo Formation
of Arkansas (thought to be about equivalent to the Kinkaid
Formation of Illinois) among which there is one species
that is close to Cymbiocrinus contentus, n. sp. from Espey
Creek.

III. Lambert Ranch

The locality is 2.4 miles west of the San Saba Court
House, San Saba, Texas, on State Highway 190, left on
Farm Road 1030 (Wallace Creek Road) 6.7 miles past a
big red barn and ranch house of Pool Stock Farm, left 1.8
miles on private road to the C, B. Lambert Ranch house.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 143

Quarry is 250 yards east of the house. The rock in which
the crinoid crowns were found appears to be the Lemons
Bluff Member, Marble Falls Formation, Atokan, Pennsyl-
vanian.

IV. Kurt Zesch Ranch

The crinoidal exposure of this horizon is located 4.2
miles southwest of the Mason County Court House in
Mason, Texas, on Ranch Road 1871 and 200 yards north
of road along southeast slope of low hill on the Kurt Zesch
Ranch. The crinoidal zone is a rust-colored siliceous lime-
stone in the upper part of the Soldiers Hole Member of
the Big Saline Formation. The basal layers of the member
contain chert, and being resistant form a low but notice-
able escarpment along the northwestern side of the exposure
at this point. The whole exposure extends for over six miles
along the northwest side of a large fault, extending in a
northeast-southwest direction, The high, or southeast side
of the fault is Upper Cambrian. The downthrow of north-
west side is Ordovician, except for the area next to the fault
occupied by the Pennsylvanian and underlying thin-bedded
Mississippian strata which rest on the Ordovician. These
rocks all dip southeast to the fault.

At Honey Creek, about 3% miles on to the southwest
of the above discussed locality, Plummer (1950) listed the
Big Saline Formation as being 199 feet thick, of which the
upper 82 feet was mostly black shales which were identified
questionably as the Smithwick Shale. Of the remaining 117
feet, the lower 22 feet appear to be the Brook Ranch Mem-
ber and the balance of 95 feet of limestone the Soldiers
Hole Member, Big Saline Formation.

The exposure on the Kurt Zesch Ranch is thought by
us to be the upper part of the Soldiers Hole Member, Big
Saline Formation, Atokan, Pennsylvanian.

V. Rough Creek

Limestone escarpment on north side of Rough Creek,
about five miles northwest of Bend and near the road: cross-
ing on Rough Creek, approximately 11 miles southeast of
San Saba, San Saba County, Texas. The exposure is near
locality number 205-T-17 of Moore and Plummer (1940, p.
401). The horizon is the Lemons Bluff Member, Marble
Falls Formation, Atokan, Pennsylvanian.
VI. Kickapoo Falls

Shale exposure on north side of a creek one-quarter
mile east of Kickapoo Falls, 8.5 miles by road southwest
of the bridge at Dennis, Hood County, Texas. The exposure
is locality number 110-T-4 of Moore and Plummer (1940,
p. 395). The horizon is shale below the Kickapoo Falls
Limestone Member, Millsap Lake Formation, Desmoinesian,
Pennsylvanian.

VII. Brock

Shale exposure at road corner close to the site of old
Consolation School (no longer standing), three miles in
direct line southwest of Brock, Parker County, Texas. This
is locality number 183-T-14 of Moore and Plummer (1940,
p. 400). It is just below the Brannon Bridge Limestone
Member, Millsap Lake Formation, Desmoinesian, Pennsy]l-
vania.
VIII. Rochelle

The exposure is 2% miles east and 2% miles north of
Rochelle, McCulloch County, Texas, and is numbered 153-
T-23 in Moore and Plummer (1940, p. 396). It is a shale
in the Mineral Wells Formation, Missourian, Pennsylvanian.
IX. Lake Bridgeport

The outcrop is a road cut four miles west of the rail-
road station at Bridgeport, Wise County, Texas, and 0.01
mile north on the paved road off the Lake Bridgeport Road.
It is a variegated shale, formerly termed the Lake Bridge-
port Shale Member, currently called the Wolf Mountain
Shale Member, Graford Formation, Missourian, Pennsyl-
vanian. The most common fossils are sponges.
X. Southwest of Lake Bridgeport Dam

The outcrop is about three-fourths of a mile southwest
of Lake Bridgeport dam, Wise County, Texas, in a shale
below the Rockhill Limestone Member, Graford Formation,
Missourian, Pennsylvanian,

MORPHOLOGICAL NOTES

I. Basal concavity in the dorsal cup

A better philosophical understanding of ontogenetic
changes is possible by carefully considering the process of
growth. Normally stereom (essentially calcium carbonate)
is added more or less evenly around the perimeter of a
plate. Of course we are aware of differential growth in some
segments (e.g. first primibrachs often attain their length
at an early stage and thereafter increase substantially in
width). We wish to call attention here to a condition of the
base of the cup where a concavity is involved.

In a young specimen there may be mildly downflared
infrabasals with the basals flexed slightly into the con-
cavity and recurved into the lateral sides of the cup. The
basal concavity may be shallow but under normal condi-
tions the increased material on the distal tips of the infra-
basals and on the proximal tips of the basals will result
in a deeper, more pronounced basal concavity with matur-
ity. It is conceded that differential or unusual surficial
growth may upset the plan just outlined.

The above described development apparently only
takes place when a pronounced basal concavity is found in

144 PALAEONTOGRAPHICA AMERICANA (VI, 40)

maturity. We have noted in a form like Erisocrinus, where
there is never a decided basal concavity, that young speci-
mens often have the appearance of a pronounced basal con-
cavity due to the confined nature of the depression at a
young stage, but it becomes broad and only shallowly de-
pressed at maturity.

II. Common modes of arm division

We have used five descriptive terms to cover the modes
of arm branching. This does not concern whether the bi-
furcation takes place with the first or tenth brachial but
only the method of division. If more than ten arms are
formed there will be more than two divisions. If all divisions
are even the branching is termed “isotomous.” If the first
two divisions are even but subsequent branching is restrict-
ed to the inner half-arms it is termed “bi-endotomous.”
When the inner half-arm is appreciably reduced in size at
the second branching and subsequent branching is restricted
to the inner half-arms, it is termed “endotomous.” When the
upper branching is on the outer half-arms it is termed
“exotomous.” The term “heterotomous” is used in the broad
sense where the branching is irregular.

III. Structure of brachials

The most primitive brachial structure in this study
consists of long, uniform segments which make up a uni-
serial arm. Each brachial carries a single pinnule and the
pinnular side alternates with adjacent brachials, If some
impetus toward a more effective food gathering system
arises, causing evolutionary modifications, several changes
are possible.

A. Each brachial may develop a pinnule on both sides,
as reported by Strimple (1962c) for Zeacrinites. This is con-
sidered by us to be a specialized development among crin-
oids.

B. A brachial may develop more than one pinnule on
a side (hyperpinnulation) as found in Neozeacrinus peram-
plus Wanner and various species of Exocrinus Strimple. This
is considered by us to be a specialized condition and not
common among known crinoids.

C. Each brachial becomes low and yet has a single
pinnule. This condition is usually accompanied by elongated
arms, as found in Ulrichicrinus Springer.

D. Each brachial has one pinnule and becomes cunei-
form. This is apparently a step toward development of bi-
serial arms in some lineages.

E. Each brachial has one pinnule and is wedge-shaped.
The arms remain long and have well-rounded exteriors.

F. Same as E above but with sharply angled wedges
and flattened exteriors, as found in Erisocrinus Meek and

Worthen.

We believe the normal evolution to be from A to D
to E to F, ie. from true uniserial arms to cuneiform
brachials to wedge-shaped brachials. Even casual observa-
tion discloses the highly efficient pinnular food gathering
system afforded by F as compared to A. Some forms having
biserial arms (e.g. Bronaughocrinus Strimple or Delocrinus
(Miller and Gurley) are able to subsist with relatively short
arms.

A brachial structure known as syzygy is a specialized
condition among Paleozoic crinoids but is common in the
Recent Articulata. Brachials are paired and joined by close
suture.

IV. Differential growth in axillary first primibrachials

Young stages of Stellarocrinus texanus (Moore and
Plummer) studied by us have biserial arms and somewhat
elongated first primibrachs, With growth, the length of the
primibrach does not increase appreciably, but the width
increases considerably.

V. Interfingered pinnules

A specialized development is found among some speci-
mens of Stellarocrinus studied by us, wherein the pinnules
are found to be at right angles to the arms and interlocking
with pinnules of the adjacent arms. This is thought by us
to be a protective device, to close the gaps left beween
the arms which do not abut at any time.

A somewhat comparable development of interfingered
pinnules between adjacent arms is illustrated by Yakovlev
and Ivanov (1956, pl. 7, fig. 1) for Pachylocrinus pachy-
pinnularis Yakovlev and Ivanov, 1956.

VI. Hypertrophied arms

Some crinoids commonly have a few hypertrophied
arms (e.g. Isoallagecrinus Strimple, 1966, Anartiocrinus
Kirk, 1940). In the case of Jsoallagecrinus it appears that
the two robust arms served as support for the multiple,
threadlike arms. This condition does not hold true for
Anartiocrinus, and we suggest that locomotion might be
involved. For example, among the modern comasterids there
are forms which have arms of unequal length (e.g. Comatula
purpurea), that are regularly used in reaching out, grasping
an object, and pulling the animal along (see Hyman, 1955,
p. 102, fig. 19 A). The fossil species Lecobasicrinus kickapoo-
ensis, n. sp. has several hypertrophied arms. We believe that
most late Paleozoic crinoids were eleutherozoic, at least
in the sense that firm root systems were not developed in
a manner that would make the animals permanently station-
ary. It is visualized that the stem trailed along at times
and could loop around an object. In some instances termin-
ating cirri suggest the animal might have sat on the bottom
with the crown well above the muck. In many forms the

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 145

stem was almost certainly deciduous (e.g. Stomiocrinus
conlini). Following this line of reasoning, it is not surpris-
ing to find unusually stout arms which might be used in
some special locomotive problems.
VII. Habitat of eleutherozoic crinoids

Once a crinoid loses its stem it is obvious that it would
sit on the bottom. The nature of the base of the cup could
then indicate the composition of the bottom, i.e. whether
a hard surface or a soft surface. Such genera as Agassizo-
crinus or Paragassizocrinus usually have a more or less
pointed base which indicates a soft bottom was present.
Some species develop a wide base (infrabasal disk) but
this is thought to also involve stabilization, together with
a pronounced thickening of the infrabasal elements. We are
reminded of a toy, popular many years ago, called a “rolly-
polly” doll. The base was rounded and weighted with lead
so that no matter which position the doll was pushed or
placed it would always swing upright when the pressure
was released. The shape of the doll was much like that of
Stomiocrinus conlint, n. sp. or Globacrocrinus pirum Moore
and Plummer.
VIII. Evidence of regression

A generally accepted concept of progressive evolution
in the arms of Paleozoic crinoids is for the point of bifurca-
tion to migrate downward toward the cup. In the genus
Ulrichicrinus the arms branch once with the primibrach in
all arms and again with the first secundibrach in most subse-
quent arms. The species Ulrichicrinus ramosus, n. sp. is the
youngest representative of the genus. It has an axillary
first primibrach, but there are two secundibrachs with the
second axillary. The addition of a secundibrach is consid-
ered by us to be regressive since it reflects a condition which
was probably normal in the much older progenitor of
Ulrichicrinus.

TERMINOLOGY

The terminology used herein is mainly that developed
by Moore and Plummer (1940) in their study of crinoids
from Upper Carboniferous and Permian strata in Texas.
Symbols for crinoid parts are those proposed by Moore and
Laudon (1941). Terminology relative to modifications of
anal plates within the posterior interradius of the dorsal
cup is that proposed by Strimple (1960b).

ASSOCIATED PARASITIC LIFE

There is a surprising lack of direct evidence of para-
sitic life considering the large amount of material at hand.
One interesting example is a row of shallow pits along
the right (outer) lateral side of an arm of Stellarocrinus sp.

cf. S. angulatus (Miller and Gurley) from the Wolf Moun-
tain Shale, Graford Formation, Missourian. If the parasite
had taken up residence, as when Myzostoma burrows in,
there would be some swelling. We postulate that some organ-
ism capable of rapid cutting or rasping, as found among
some gastropods, created the depressions which did not re-
quire any appreciable effort to repair on the part of the
crinoid, The regrowth of the skinlike covering over the small
damaged areas would have been sufficient.

STRATIGRAPHIC IMPLICATIONS
A. INTRODUCTION

The crinoid faunas under study range from Chesterian
(Mississippian) to Missourian (Pennsylvanian) in age. A
small fauna in poor preservation found in Mason County,
Texas, appears to be from a limestone in the Barnett Forma-
tion.

Morrowan crinoids are reported from the Sloan Mem-
ber, Marble Falls Formation. A small fauna from Espey
Creek near Lampasas, Texas, is comparable in several re-
spects to forms from the Brentwood Limestone Member,
Bloyd Formation (Morrowan) of Oklahoma and Arkansas.
A discussion relative to a possible difference in age is given
under the section “Crinoid Localities.” One species, Dicro-
myocrinus texasensis (Moore and Plummer), was previously
known in Texas from location 205-T-43, which is about 11.5
miles by road southwest of San Saba, Texas (horizon about
ten feet above the base of the Marble Falls Formation,
which is of Morrowan age) and about 0.1 mile northeast
of the road fork near bench-mark 1317, on the Cherokee-
Chappel road (horizon from four to six feet above the base
of the Marble Falls), San Saba County, Texas.

Atokan crinoids are reported from the Lemons Bluff
Member, Marble Falls Formation, on Rough Creek south-
east of San Saba, Texas, from the Lambert Ranch south-
west of San Saba and from the Soldiers Hole Member, Big
Saline Formation, on the Kurt Zesch Ranch southwest of
Mason, Texas. The Lemons Bluff fauna on the whole is not
closely comparable to any other known crinoid fauna, Some
elements of the Soldiers Hole fauna are close to forms from
the Burgner Formation of Missouri and the Bostwick For-
mation of southern Oklahoma.

Desmoinesian crinoids are reported from the Kickapoo
Falls Limestone Member and the Brannon Bridge Lime-
stone Member, Millsap Lake Formation, near Kickapoo
Falls and near Brock, Texas, respectively. These faunas are
partially known from the study made by Moore and Plum-
mer (1940).

146 PALAEONTOGRAPHICA AMERICANA (VI, 40)

A few Missourian crinoids are included for various rea-
sons. One new species is from the Mineral Wells Formation
near Rochelle, Texas. Two important crowns are reported
from the Wolf Mountain Shale, Graford Formation, near
Bridgeport, Texas. A rare genus Stomiocrinus is reported
from the Graford Formation. The Wolf Mountain Shale is
close in faunal content to the Stanton Formation of Kansas

and to some facies of the Wann Formation of northeastern
Oklahoma.

B. MISSISSIPPIAN PERIOD

The only other reported occurrence of crinoids of Mis-
sissippian age from Texas known to us is the small] species
Synbathocrinus texasensis Moore and Ewers (1942) from
shale pockets in the Chappel Limestone (Osagian), near
Marble Falls, Burnet County, Texas. Our specimens were
obtained from rock just below a brown shale of the Barnett
Formation in Mason County, near Mason, Texas. The crin-
oids appear to be Chesterian in age. Several specimens of
a species named Phanocrinus trulleum, n. sp. were found
together with a robust Scytalocrinus sp., Culmicrinus bar-
nettensis, n. sp., a radial plate of Pentremites sp., and two
large axes of Archimedes.

A species is described as Chlidonocrinus trinodi, n. sp.
from the Fayetteville Formation, Chesterian, of northeast-
ern Oklahoma to provide a progenitor of Chlidonocrinus
echinatus, n. sp. from the Lemons Bluff Member, Marble
Falls Formation (Atokan).

C. MORROWAN STAGE

The following crinoid species have been described from
Morrowan rocks of Texas: Dicromyocrinus texasensis
(Moore and Plummer), Dicromyocrinus optimus Strimple*
(first Texas listing), Diphuicrinus croneisi Moore and Plum-
mer (reported by Plummer, 1950, p. 53), Oxynocrinus spic-
ata, n. sp.® (first recording), Cymbiocrinus contentus, n. sp.*
(first recording) Graffhamicrinus antiquus n. sp.* (first re-
cording), and Cibolocrinus punctatus Moore and Plummer.

Scytalocrinus sansabensis Moore and Plummer and
Lasanocrinus cornutus Moore and Plummer were originally
reported to be of Morrowan age but are now known to be
of Atokan age.

The fusulinid Millerella marblensis Thompson is com-
mon in some exposures of the Sloan Member.

Correlation with the Morrowan of northeastern Okla-

“One of us, Watkins, is of the opinion these specimens are from the
Chapel Limestone (Osagian).

“Rocks exposed at the type locality of the Otterville Formation, near
the abandoned village of Otterville, have been found to be Des-
moinesian. Most other exposures referred to the Otterville are Mor-
rowan.

homa and northwestern Arkansas is reasonably well estab-
lished. No special problems relative to the Morrowan are
incurred other than the age of the “Winslow” of Arkansas
which has been called Morrowan by some authors but is
considered by us to belong in the Atokan Stage. The Wap-
anucka Limestone, Gene Autry Shale and Ottervillet For-
mation of southern Oklahoma are correlatives of the Bloyd
Formation of Oklahoma and Arkansas.

Furnish and Knapp (1966) found the Morrowan am-
monoid, Diaboloceras newmeiri Quinn and Carr, in the
Kendrick Shale of eastern Kentucky along with Dimorpho-
ceratoides campbellae Furnish and Knapp. Strimple and
Knapp (1966) identified Paragassizocrinus kendrickensis
Strimple and Knapp, Paragassizocrinus cf. P. turris
Strimple, Paragassizocrinus cf. P. disculus Strimple, and a
crinoid arm as Metacromyocrinus oklahomaensis (Moore
and Plummer) from the Kendrick beds and Diphuicrinus
patina Strimple and Knapp from Magoffin beds, Breathitt
Formation of eastern Kentucky. These forms all indicated
a late Morrowan or early Atokan age.

An ammonoid similar to Diaboloceras newmeieri is re-
ported by Furnish and Knapp (1966, p. 303) from the Penn-
sylvanian sequence near Port Birmingham in Jefferson
County, Alabama. The horizon appears to be Metzger’s
Stratigraphic Interval D (1965, p. 74) to the west of his
Stratigraphic Section 34, Gastrioceras occidentale (Miller
and Faber) was found associated, as well as fragments of
crinoids close to Slenopecrinus rugosus Strimple, all of which
indicated a Morrowan age.

Lane (1964) described Polusocrinus pachyplax Lane
and P. calyculoides Lane from the lower part of the Call-
ville Formation at Sloan and Frenchman Mountains, Clark
County, Nevada, respectively and judged them to be of
Morrowan age. We have specimens from the Lemons Bluff
Limestone Member, Marble Falls Formation (Atokan),
thought to be conspecific of P. calyculoides.

In the study by Miklukho-Maclay (1960, p. 412, text-
fig. 2), Stage Cob ( Bashkirian) was divided in Central Asia
into a lower Zone a, with Pseudostafella and Eostafella
present, and an upper Zone b, Yangak horizon, with
Pseudostafella and first Profusulinella. The same division
was shown for the Ural Mountains, except the horizon name
was deleted. A disconformity was shown in the Moscow
Basin. If the Russian Profusulinella are congeneric with
the American, as seems to be the case, it appears that the
upper part of the Bashkirian is of Atokan age. In text-fig.
1 of Miklukho-Maclay, the Bashkirian was represented by
the goniatite genera Gastrioceras and Branneroceras. How-
ever, Librovitch (1960, p. 383) stated that Branneroceras

‘TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 147

flourished in the beginning of the Bashkirian and later
becomes rarer. Aisenverg, et al. (1960, p. 4) listed Brannero-
ceras branneri, together with several Gastrioceras from Zone
Cbsa of the Donetz Basin and the fusulinids Fostafella
varvariensis, Pseudostaffella antiqua, and P. praegorsku. In
North America, Branneroceras branneri is an index fossil
of the Bloyd Formation, and more specifically of the Brent-
wood Limestone Member. Aisenverg, et al. (1960, p. 4)
listed Profusulinella primitiva in Zones Chsd and Csc with
Pseudostafella still present. McCaleb (1964) correlated the
Bloyd Formation with Zones Cbsa and Cbsb (Limestones G,
and H,) of the Donetz Basin which is close to the correlation
we have proposed.

McCaleb (1964, p. 11) considered the Wangchiapa
Limestone of South China correlative in part with the
Bloyd Formation (Morrowan) of Oklahoma and Arkansas.

D. ATOKAN STAGE
I, TYPE LOCALITY

The validity of the Atoka Formation, and therein the
Atokan Stage, has been questioned because of a supposed
lack of a type location, 7. e. for the Atoka Formation. The
name Atoka Formation was proposed by Taff and Adams
(1900, p. 273) and applied to an interval of sandstones and
shales between the Hartshone Sandstone Formation and the
Wapanucka Limestone below, which they thought to have
a thickness of 7,000 feet. Taff (1901) changed the thickness
of the formation to 3,100 feet which is more realistic. It is
accepted in the U.S.G.S. Lexicon (1966, pp. 165, 166) and
elsewhere as being named for the town of Atoka, Atoka
County, Oklahoma, which is located on an outcrop of the
formation. A rock-stratigraphic name was thus established
upon a general geographic location, The Atoka Formation
extends in an unbroken belt to the north and west and
covers a considerable area in the vicinity of Barnett Hill
north of the town of Clarita in Coal County, Oklahoma.
Ammonoid, fusulinid, and crinoid faunas found in the lower
Atoka Formation of Oklahoma in Coal County establish a
sound basis for stratigraphic correlations with other areas.
The easily identified Wapanucka Limestone is in juxta-
position throughout the area.

Morgan (1924) considered the Atoka Formation of the
Stonewall Quadrangle of Oklahoma to be about 800 feet
thick and the Hartshorne Sandstone to be about 100 feet.
This outcrop is in a narrow band extending from sec. 8, T.
1 N., R. 7E., Pontotoc County to secs. 18 and 19, T. 1 N.,
R. 8 E., Coal County. The Atoka spreads out and thickens
to the south and east in Coal and Atoka Counties where

Taff (1901) noted a thickness of as much as 3,100 feet, as
stated above.

Harlton (1934) proposed that the Atoka Formation be
divided and the lower portion (which he considered to be
of Morrowan age) called the “Barnett Hill Formation.” His
primary type section was at Barnett Hill north of Clarita,
Coal County, Oklahoma; although he gave a second type
section on Goose Creek in secs. 19 and 20, T. 1 N., R. 8 E.,
Coal County. Harlton further considered the remaining
portion of the Atoka Formation to be Desmoinesian. The
name “Barnett Hill Formation” has not received general
acceptance in Oklahoma, but if the status were reduced to
Barnett Hill Member, Atoka Formation, it would be useful
and significant. Such a usage is proposed herein and will
be referred to in such a manner in the balance of this
study.

The Barnett Hill Member has a thickness of about 450
feet in the vicinity of Barnett Hill which is taken as the
type locality. Harlton reported 250 feet of dark gray shale,
fossiliferous in part at the base, a fusulinid zone at about
350 feet, and a prolific cephalopod bearing sandstone at the
top. As noted elsewhere, Fusulinella prolifica has been re-
ported from a horizon about 250 feet above the base,
a short distance west of Barnett Hill by Rowett (1963).
The species has also been reported about 200 feet above the
base of the formation on the north flank of Barnett Hill
by Thompson (1935). This species is considered to be of
Atokan age as discussed elsewhere.

The Goose Creek exposures of Harlton (1934 and 1938)
are to the northwest of Barnett Hill in Coal County. Harl-
ton has shown the Barnett Hill Member to have a thick-
ness of about 650 feet at this locality, and the cephalopod
sandstone horizon to be replaced by shale and limestone.
He noted a fusulinid horizon about 530 feet above the base
which he correlated with the horizon at 350 feet on Barnett
Hill. Thin-bedded and arenaceous limestones are reported
in the upper 150 feet of the Barnett Hill Member on Goose
Creek. It is possible that the crinoid bearing calcareous
shale and sandstone found in sec. 28 about two miles south-
east of the Goose Creek exposures represents an eastward
extension of one of the arenaceous limestones, Profusulinella
fittst is found in association with the crinoids in sec. 28 as
discussed elsewhere.

Branson (1962, p. 439) in discussion of the Atokan
Series stated, “The name (Atokan) is derived from the
Atoka Formation, and in the type area, the series and
formation boundaries are identical.” We have followed this
interpretation.

The name Atoka as a time-rock term has been used for

148 PALAEONTOGRAPHICA AMERICANA (VI, 40)

several years in an informal manner but the type area of
the Atokan Series or Stage was not formally established
until proposed by Spivey and Roberts (1946, pp. 185,
186). Their statement was, “—the writers propose that
the Atoka formation be elevated to Atoka series and defined
to include all beds from the top of the Wapanucka lime-
stone, Morrow series, to the base of the Hartshorne sand-
stone, Des Moines series.” They further stated, “The Atoka
series seems to be a better name and type locality for these
beds than the term ‘Derry,’ because the Derry at its type
locality is only about 130 feet thick, whereas the Atoka beds
reach 7000 feet in thickness.” As noted elsewhere the Atoka
beds in the type area probably only reach a thickness of
3100 feet, which does not alter the significance in any event.
Spivey and Roberts (1946, p. 186) also summarized their
reasons for rejecting the terms “Bend” and “Lampasas” in
favor of the Atokan Stage.

There has been no clear definition in the past because
there was no universal agreement on the base or the top of
the Atokan, i.e. the type section of the Desmoinesian is not
adequately understood on the one side, and some question
remained as to whether the “Winslow” of Arkansas was
Morrowan or Atokan on the other side. An attempt is made
to at least partially resolve some of these matters.

II. OTHER OKLAHOMA EXPOSURES

In northeastern Oklahoma the Atoka Formation is
divided by Wilson and Newell (1937) into six prominent
sandstone members, from bottom to top (1) Coody Sand-
stone Member, (2) Pope Chapel Sandstone Member, (3)
Georges Fork Sandstone Member, (4) Dirty Creek Sand-
stone Member, (5) Webber Falls Sandstone Member, and
(6) Blackjack School Sandstone Member. The ammonoids
of the Coody Sandstone Member are comparable to those
of the “Winslow” of Arkansas. Huffman (1958, p. 168)
included eight feet of shale and sandy shale below the Coody
Sandstone in the Bloyd Formation (Morrowan), as exposed
on Braggs Mountain along State Highway 10 in Oklahoma.

In the Ardmore Basin the name Bostwick Formation is
applied to rocks above the Otterville (Morrowan) Forma-
tion and below the Lester Limestone Member, Lake Murray
Formation. The Lester Limestone has been called Desmoi-
nesian by Cronoble and Waddell (1966) and Waddell
(1966) because the “Desmoinesian fusulinids Fusulina in-
solita and F. mutabilis” are present These species are primi-
tive representatives of Fusulina, and there is some question
in our minds that they can be called or used to designate
Desmoinesian age. F. insolita was originally found in rocks
considered to be of Atokan age. In any event the Bost-
wick is Atokan,

Some ammonoids collected in the Lester Limestone
by Allen Graffham and H. L. Strimple in 1964 from the
NE% NE% sec. 9, T. 6 S., R. 2 E., Love County, Okla-
homa, have been identified as Pseudoparalegoceras cf. P.
brazoense. They represent a stage of development somewhat
more primitive than reflected by Boggy forms but more ad-
vanced than forms from the Barnett Hill Member, Atoka
Formation. Some disarticulated crinoid spines are close to
specimens from the Bostwick Formation.

In order to understand some of the comparative fea-
tures it Is necessary to be acquainted with the Krebs and
Cabaniss Groups. The names were proposed by Oakes
(1953) with the Krebs bounded below by the Atoka For-
mation and above by the Thurman Sandstone. The Cabaniss
is bounded below by the Boggy Formation of the Krebs
and above by the Fort Scott Limestone (Marmaton). It
was noted by Oakes, the group occupied the same strati-
graphic horizon as the Cherokee rocks of southeastern Kan-
sas, except that he considered the Hartshorne Sandstone
and possibly some lower McAlester rocks were older than
any Cherokee rocks of Kansas, and the lower part of the
Senora Formation was probably not represented in the
Cherokee of Kansas because of the progressive northward
overlap in the post-Boggy rocks. The conclusion that older
rocks of the Cherokee are absent in Kansas is not under-
stood by us because the Little Cabin Sandstone (=War-
ner Sandstone of Oklahoma) is reported in Kansas (see
Moore, et al., 1951, pp. 98-99) with about 60 feet of rock
beneath it. The Little Cabin Sandstone of Oklahoma has
been correlated with the Warner Sandstone (lower McAles-
ter Formation) by Newell (1937, p. 40) and Branson (1955,
p. 66), and the term Little Cabin is no longer recognized
in Oklahoma.

Hartshorne Formation is used in the following discus-
sion in the context proposed by Branson (1955, p. 64), i.e.
to cover rocks from the top of the Atoka to the top of the
Upper Hartshorne Coal. A composite section given by
Branson from SW1{ sec. 3, T. 27 N., R. 22 E. and east of C
sec. 10, T. 28 N., R. 22 E., along with observations in ad-
jacent Craig County, shows about 25 feet composed of con-
glomerate at the base, clay, coal, shale, siltstone, capped by
less than a foot of Riverton Coal, which is all ascribed to the
Hartshorne Formation. This agrees closely with a section
given by Moore, et al. (1951, fig. 38) for a 35 foot interval,
more or less, below the Little Cabin Sandstone in Kansas.

The contact between Atokan and Desmoinesian will
probably be found somewhere below the Little Cabin Sand-
stone of Kansas and below the Warner Sandstone in Mis-
souri. Howe (1961) included the Riverton Formation of

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 149

Missouri in the Atokan which he correlated with the Harts-
horne? or the Warner Sandstone in the Forest City Basin.
Moore, et al. (1944, Chart No. 6) showed the Hartshorne
and McAlester to be upper Lampasan (= ?lower Desmoi-
nesion). If the Missouri formations are correlatives of those
of Oklahoma, 7z.e. Hartshorne, Warner, then the Riverton
Formation should be referred to the Desmoinesian.

Ill. FAUNA OF THE ATOKA FORMATION IN OKLAHOMA

The most significant fauna of the Atoka Formation is
found in the northwestern extension of the formation in
the general vicinity northwest of Clarita, Coal County,
Oklahoma. It is of the utmost importance because the for-
mation extends in an unbroken belt from its nonfossiliferous
type locality in the vicinity of Atoka, Atoka County, Okla-
homa, into the area around Clarita. The formation is bound-
ed below by the Wapanucka Limestone (Morrowan) and
above by the Hartshorne Sandstone (Desmoinesian). A
prolific cephalopod fauna is found at Barnett Hill located
just north of Clarita (SW% sec. 2, T. 1 S., R. 8 E.).
The fossils occur in sandstone exposed well below the crest
of the hill on the south side and dipping to the north, so
that the fossiliferous zone is low on the hill on its north side.
It is on the north flank of Barnett Hill that a collection was
made in April, 1965, by a field party from the University of
Iowa, made up by H. L. Strimple, W. W. Nassichuk, and
Claude Spinosa. The following species were identified:
Pseudoparalegoceras kesslerense (Mather), Paralegoceras
texanum (Shumard), Stenopronorites arkansiensis (Smith)
and Eoasiamttes sp. In addition, Miller and Furnish (1940,
1958) listed and described Paralegoceras iowense (Meek
and Worthen) from Barnett Hill, albeit from on the south
side of the hill near the old Barnett homesite. The house
no longer exists, but the barn is still standing.

W. M. Furnish called attention to a personal communi-
cation from H. L. Griley, dated May 9, 1959, wherein it is
stated that the specimen figured by Miller and Furnish
(1958, text-fig. 2 E) as Paralegoceras texanum (Shumard )
from the lower Atoka Formation near Clarita, Oklahoma
(S.U.I. No. 13996), was collected by John Britts Owen,
John Fitts, and H. L. Griley in open woods just north
of Goose Creek and just east of the Jesse-Clarite road, three
miles southeast of Jesse (NW% sec. 19, T. 1 N., R. 8 E.) in
Coal County, Oklahoma. This is the area selected by Harl-
ton as his supporting type section for the “Barnett Hill
Formation,” now called Barnett Hill Member, Atoka For-
mation, and was visited by the previously mentioned field
party from the University of Iowa in April, 1965.

McCaleb (1963, pp. 868-870) listed the previous species

from Barnett Hill and in addition the following species:
Boesites girtyi (Plummer and Scott), Bisatoceras sp., Gas-
trioceras formosum McCaleb, and Eoasianites smithwick-
ensis (Plummer and Scott).

Unklesbay (1962) listed the following species from the
“Atoka Group” of Oklahoma and those marked with an
asterisk (*) were shown as occurring at Barnett Hill:
Pseudoparalegoceras compressum, *P. williamsi, *P. kessler-
ense, *Gastrioceras adaense, *Eoasiamtes globulosus, Dia-
boloceras varicostatum, *Paralegoceras iowense, *P. texan-
um, *Boesites girtyi and *Pronorites arkansasensis.

The general area near Clarita, Oklahoma, has yielded
42 identified fossil forms listed by Mather (1917, pp. 133-
139). A listing is given by Blythe (1959, p. 38) showing
modern classifications where needed. An additional 14
speciێs were identified and listed by Blythe (1959, p. 39)
from a collection made by A. R. Denison at Barnett Hill
and reposited at the University of Oklahoma.

The center of the section line between secs. 2 and 3,
T. 1 S., R. 8 E., and about one-fourth mile northwest of
the cephalopod exposure, is one of the type localities for
Fusulinella prolifica Thompson (1935), the other type lo-
cality being on the north flank of Barnett Hill in the center
of sec. 2, T. 1 S., R. 8 E. Both of these exposures are lower
than the cephalopod horizon and are reported by Thompson
to be about 200 feet above the base of the formation.

Fusulinella oliviformis Thompson (1935) is also from
approximately 200 feet above the base of the Atoka For-
mation in the center of the line between secs. 2 and 3, T.
1 S., R. 8 E., Coal County, Oklahoma (northwest flank of
Barnett Hill) and from the north flank of Barnett Hill
(center sec. 2, T. 1 S., R. 8 E.). It is of considerable im-
portance in that Thompson considered it (F. oliwiformis) to
be closely related to F. magna (Lee and Chen, 1930) from
the basal portion of the Huanglung Limestone of Chuan-
shan, China. Thompson (1935, p. 294) listed fusulinid
species of the lower Atoka Formation which are com-
parable to those of the lower faunule of the Huanglung
Limestone as Staffella atokensis Thompson, Profusulinella
fittsi (Thompson), and Fusulinella prolifica Thompson, in
addition to the above mentioned species.

Profusulinella fittsi was originally reported from the
south branch of Goose Creek, but the description was
NW4% sec. 10, T. 1 N., R. 8 E. This apparently should
read sec. 10, T. 1 S. where there is Atoka exposed because
the Boggy Formation is exposed in sec. 10, T. 1 N. The
species was also reported in the north center of sec. 19, T.
1 N., R. 8 E. The horizon is reported to be about 100 feet
above the base of the Atoka Formation.

150 PALAEONTOGRAPHICA AMERICANA (VI, 40)

The fusulinids discussed above which were described
by Thompson (1935) were collected by R. V. Hollingsworth
(personal communication from M. L. Thompson).

Rowett (1963, p. 34) noted the occurrence of Fusulin-
ella prolifica 250 feet above the Wapanucka Limestone in
SEI4 SWI, sec. 3, T. 1 S., R. 8 E., Coal County, which is a
short distance southwest of Barnett Hill.

A crinoidal horizon found several years ago by M. K.
Elias, while making a study of the Atoka Formation in
Coal County (near center N% sec. 28, T. 1 N., R. 8 E.),
was shown to Allen Graffham. This outcrop consists of a
low ridge formed by a calcareous lentil in soft clay-shales,
and the natural ravines expose a shell-layer which is often
associated with a thin sandstone about two inches thick in
places. The clay-shales above the sandstone contain many
dissociated crinoid ossicles. A small colony of crinoids was
discovered by Allen Graffham and one of us, Strimple, in
the fall of 1960. Mr. Graffham acquired a good collection
which has been purchased by the University of Oklahoma
and is currently under study by the senior author. The
previously mentioned field party from the University of
Iowa collected at the locality in April 1965. Some crinoids
were recovered and shale samples taken. Fusulinids were
recovered from washed residues and sent to M. L. Thomp-
son and Garner L. Wilde, with the following reports ob-
tained:

Thompson (personal communication August 16, 1965)
wrote, “The collection from near the center of the N% sec.
28, T. 1 N., R. 8 E., Coal County, Oklahoma, contains what
appears to be Profusulinella that resembles closely P. fittst.
No large specimens of fusulinids were found in this collec-
tion.” He considered the material to be from near the hori-
zon from which P. fittsi was originally described.

Wilde (personal communication July 1, 1965) wrote,
“The material you sent from the ‘Griley limestone’ (sec. 28,
T. 1 N., R. 8 E. Coal County) is an advanced species
of Profusulinella and Eoshubertella sp. This would be early
Atokan in age.”

On the basis of fusulinids it thus appears that the
crinoid zone in sec. 28 is below the Fusulinella prolifica hori-
zon which is in itself below the prolific cephalopod horizon.

A preliminary examination of the crinoids collected by
Allen Graffham, M. K. Elias, W. W. Nassichuk, Claude
Spinosa, and the senior author in sec. 28 discloses the exis-
tence of the following genera: Oklahomacrinus, Alcvmo-
crinus, Stellarocrinus, Pirasocrinus?, Paradelocrinus, Aesio-
crinus, Synerocrinus, Isoallagecrinus, Decadocrinus?, Par-
ethelocrinus, Metacromyocrinus, new genus (aesiocrinid),
and new genus (affinities undetermined ). All of the genera

are known to us from younger strata with the exception of
Alcimocrinus which has not previously been reported from
above the Morrowan, The individual species are, in general,
more comparable to early Desminesian forms than to Mor-
rowan species.

Strimple (1966a, pp. 8-12) described Synarmocrinus
fundundus Strimple from the Barnett Hill Member, Atoka
Formation, near Clarita, Coal County (near C NW 14 sec.
10, T. 1 S., R.8 E, and © Ni 4 sec. 28,07 VIN Ragas

The Coody Sandstone Member, Atoka Formation, near
the top of Braggs Mountain southeast of Muskogee, Musko-
gee County, Oklahoma, is cephalopod bearing. Miller and
Furnish (1940, 1958) reported Diaboloceras varicostatum
from the location as well as from the Smithwick Shale of
Culberson County, West Texas. A specimen identified as
Diaboloceras newmeri Quinn and Carr was recovered from
an exposure on Oklahoma Highway 10 at the top of Braggs
Mountain by a field party from the University of Iowa in
the spring of 1966. The cephalopods of the Coody Sand-
stone are comparable to those of the lower “Winslow” of
Arkansas but D. newmeri indicates a slightly older age. The
two horizons are about 50 miles apart and appear to belong
to the same general structure. We consider the “Winslow”
to be a member of the Atoka Formation, but the Coody
Sandstone may be slightly older.

A prolific microfauna has been studied and reported
on by Galloway and Rynicker (1930) from upper black
shales in the Atoka Formation (500 feet below the top of
the formation) near the middle of the north line of sec.
11, T. 5 N., R. 21 E., southeast of Red Oak in Latimer
County, Oklahoma. Thirty-three species of Foraminifera
are described by Galloway and Rynicker including three
species of fusulinids, which are Staffella molleri Ozawa,
Schubertella transitonia Staff and Wedekind, and Fusulin-
ella euthusepta Henbest.

Huffman (1958, table 16) listed one species of coral,
13 species of brachiopods, three species of pelecypods, and
one species of gastropod from the Atoka Formation of north-
eastern Oklahoma.

A list of 77 fossil species collected from the Webber
Falls Sandstone Member, Atoka Formation (NE% NE%
sec. 27, T. 17 N., R. 19 E.), in Cherokee County, Oklahoma,
in the road-cut at the eastern end of Oklahoma Highway
51 bridge over Ft. Gibson Lake, is given by Blythe (1959,
p. 41, 42). One ammonoid, Pseudoparalegoceras sp., is listed.
Unklesbay (1962, p. 96) reported Pseudoparalegoceras wil-
liamsi from the Atoka Formation exposed at the east end
of the Highway 51 bridge over Ft. Gibson Lake, probably
based on Blythe’s specimen. Two species of fusulinids are

‘TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 151

listed by Blythe, Ozawainella ciscoensis? (Harlton) and
Ozawainella sp. Paragassizocrinus atoka Strimple and Blythe
(1960) is from the same locality although erroneously re-
ported to be from section 10, and as about 15 feet above
the base of the Atoka Formation. The locality should read
NEY NEY, sec. 27, T. 17 N., R. 19 E., and the horizon is
about 15 feet above the base of the Webber Falls Member,
Atoka Formation. Extensive exposures of Morrowan rock
are found just to the northwest (SE% sec. 22, T. 17 N., R.
19 E.) along the shore of Ft. Gibson Lake, especially in
what was once the face of a quarry.

Strimple (1961b) described two species of Paragassizo-
crinus from the Bostwick Formation of the Ardmore Basin
as P. bulbosus Strimple and P. elevatus Strimple. The oc-
currence of a distinctive primibrach, ascribed to the crinoid
genus Plaxocrinus, at three exposures of the Bostwick was
also noted by Strimple (1961b, p. 294).

Laudon (1937) described a species as Synerocrinus
farisht Laudon from the Bostwick conglomerate, 300 feet
above the Otterville Limestone (Morrowan), (SE% SW%
SE% sec. 9, T. 6 S., R. 2 E.), in the Ardmore Basin, south
of Ardmore, Oklahoma. The species is also present in the
lower Atoka exposure previously discussed (sec. 28, T. 1 N.,

R. 8 E.), in Coal County.

IV. FAUNA OF THE ATOKA FORMATION IN ARKANSAS

Croneis (1930, p. 135) presented three lists of fossils
collected by Collier from the upper part of the Atoka For-
mation near Van Buren, Arkansas. He preferred the older
name of Atoka Formation to the term “Winslow Formation”
used by Adams and Taff (1900). A paper by Quinn (1959)
outlined structures in the area, where Atokan rocks are ex-
posed, and furnished data on source direction for the “Wins-
low” of northwestern Arkansas. In a study of the goniatite
fauna of the lower Atokan of Arkansas, McCaleb (1963, p.
868) concluded the term “Winslow” should be reinstated,
quote “—as it can no longer be considered synonymous
with Atoka, on the basis of fauna, lithology and source of
sediments.” He further concluded that the “Winslow” was
Morrowan rather than Atokan in age. We do not believe
the fauna is sufficiently distinct to warrant removal from
the Atokan. The goniatite considered by McCaleb to be
typical of and restricted to the “Winslow” is Winslowoceras
henbesti. The form has not been identified in the Atokan
of Oklahoma, but this is negative evidence.

Gordon (1965, p. 273) accepted the identification of
Elias (1956, p. 100) of W. henbesti from the Gene Autry
Shale (Morrowan) in the southern part of the Arbuckle
Mountains near Ardmore, Oklahoma. McCaleb and Furnish

(1964) assigned specimens from Elias’ primary type locality
of the Gene Autry Shale to Axinolobus quinni McCaleb and
Furnish. The species is also found in the lower middle
Bloyd Formation (Morrowan) of northern Arkansas.
Strimple and Nassichuk (1965) reported Axinolobus quinni
from two localities in the upper Wapanucka Limestone
(Morrowan).

Eowellerites has been limited by Ruzhencev (1965) to
the Moscovian, and the genus has been reported from the.
Atoka Formation of Arkansas by Gordon (1965) as E.
discotdais Gordon and &. cf. E. moorei (Plummer and
Scott). £. moorei was previously known from the Smith-
wick Shale (Atokan) of Texas.

McCaleb (1963) used the supposed presence of Eowel-
lerites in the Atoka of Oklahoma and its presumed absence
in the “Winslow” of Arkansas as paleontologic evidence sup-
porting apparent faunal differences. But as noted above,
Gordon (1965) considered Howellerites to be of Atokan age,
as did Ruzhencev (1965). The report of EHowellerites from
the Atoka of Oklahoma was made as a footnote in Miller
and Furnish (1958, p. 26) based on communicated advice
as follows, “Also Mackenzie Gordon, Jr. has advised us that
he has encountered the genus, as well as a related form, in
the Atoka Formation of Oklahoma; this occurrence is to
be described by him in a manuscript now nearing com-
pletion.” In the comprehensive cephalopod study by Gordon
(1965) there is no reference to any occurrence of Eoweller-
ites in Oklahoma. It was probably a misunderstanding, and
Gordon meant the occurrence was in Arkansas rather than
in Oklahoma.

McCaleb (1963) noted the occurrence of the following
species in the “Winslow” of Arkansas, those marked with
an asterisk (*) are also found in the Atoka Formation in
Oklahoma: *Eoastanites smithwickensis (Plummer and
Scott), Wiedeyoceras williamsi McCaleb, *Pseudoparalego-
ceras kesslerense (Mather), *Pseudoparalegoceras compres-
sum (Hyatt), Gastrioceras formoswum McCaleb, *Steno-
pronorites arkansasensis (Smith). *Paralegoceras texanum
(Shumard) and Winslowoceras henbesti Miller and Downs.

Gordon (1965, p. 48) in his study of Arkansas cephalo-
pods defined the Atoka Formation as follows: “The lower
part of the Atoka Formation, a thick sequence of slabby to
massive medium-to coarse-grained sandstone and dark-gray
silty to sandy shale with local calcareous, ferruginous, and
conglomeratic beds, is the highest stratigraphic unit ex-
posed in the Boston Mountain part of the Ozark region. The
Atoka formation was named by Taff and Adams (1900, p.
273) for the town of Atoka in Atoka County, Okla., which
is in the belt of outcrop for the formation. The later name

152 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Winslow formation of Adams (1904, p. 29) used for the
same formation in Arkansas, has been abandoned.”

The name Greenland Sandstone Member has been
given to the basal bench-forming sandstone member of the
Atoka Formation of Washington County, Arkansas, by
Henbest (1953, pp. 1946, 1947). Gordon (1965) reported
Paralegoceras (Diaboloceras) varicostatum (Miller and Fur-
nish) and Pseudoparalegoceras kesslerense (Mather) from
the Greenland Sandstone Member.

At the north portal of the railroad tunnel at Winslow,
Arkansas, about 1214 feet above the base of the Atoka, as
exposed, there has been collected Pseudoparalegoceras kes-
slerense, Paralegoceras (Diaboloceras) varicostatum, and
Stenopronorites arkansiensis according to Gordon (1965, p.
49). About 29 feet higher in the section a fauna was de-
scribed by Miller and Downs (1948) as Knightoceras oxylo-
batum Miller and Downs, Pseudoparalegoceras williamsi
(=P. kesslerense), and Winslowoceras henbestt. To this list
Gordon (1965, p. 49) added Paralegoceras (Diaboloceras)
varicostatum which must have been a rare find because
none other has been found among several thousand speci-
mens of ammonoids collected by personnel from the Uni-
versity of Arkansas from the locality.

According to Henbest (1948, p. 673), bed 7 (Miller and
Downs’ cephalopod horizon) lies 125 to 150 feet above the
base of the “Winslow formation.” Four other collections are
reported by Gordon (1965, pp. 49, 50) from the lower Atoka
Formation in Madison, Crawford, and Conway Counties,
which add the species Stearoceras smithi Gordon to those
species previously listed from the lower Atoka. Two occur-
rences are given by Gordon (1965, p. 50), estimated to be
approximately 1,000 feet above the base of the Atoka For-
mation, These exposures are USGS locations 16350, 17690,
and 16248, about one mile northeast of Hobbs and three
miles west of there on West Cedar Creek in the southwest
corner of the Ozark National Forest. The ammonoid species
listed were Pseudoparalegoceras kesslerense, Paralegoceras
texanum, Stenopronorites arkanstensis and Eowellerites cf.
E. mooret. A collection made by Louis Graff at Van Buren
in Crawford County, Arkansas, is thought to be from the
upper part of the Atoka and is identified by Gordon (1965,
pp. 50, 51) as Pseudoparalegoceras kesslerense and Eoweller-
ites discotdalis.

Furnish and Knapp (1966, p. 306) provisionally as-
signed a specimen to Dimorphoceras campbellae Furnish and
Knapp which was reported to be from the “Winslow Sand-
stone” near Spirit Creek, in northern Franklin County,
Arkansas. The locality is not geographically near typical
“Winslow” rocks which occur in Washington County, Ar-

kansas, and we have no verification as to the Atokan (or
“Winslow”) age. We suspect the horizon is pre-Winslow
which would agree with the age for a typical representative
of D. campbellae, i.e. Kendrick beds, upper Morrowan Stage.

VY. FAUNA OF THE ATOKAN STAGE IN MISSOURI

Howe (1961) showed the Seville Formation of Missouri
above the Bluejacket Formation and at the top of the Krebs
(Desmoinesian). It is not the same horizon as the Seville
Limestone of Illinois which is in the zone of Fusulinella and
the Bluejacket Sandstone is well within the zone of Fusu-
lina.

The Riverton Formation of Missouri, together with the
Burgner Formation, comprise the exposed Atokan of Mis-
sourl according to Howe (1961). The McLouth Formation,
which is also Atokan, is subsurface. The Riverton Forma-
tion is discussed under the Oklahoma section herein and is
thought to probably be of Desmoinesian age.

The name Burgner Formation was applied to relatively
early Pennsylvanian rocks (Atokan) in the general vicinity
of Webb City, Jasper County, Missouri, by Searight and
Palmer (1957). Thompson (1953) previously reported on
two species of fusulinids from the limestone which partly
filled a depression or sink in the Mississippian limestone
near Carterville (SE%4 NE% sec. 20, T. 28 N., R. 32 W.),
Jasper County, Missouri. These were described as Fusulin-
ella clarki Thompson and F. searighti Thompson and con-
sidered to be of Atokan age. F. searighti was abundant 14
feet below the top of the limestone and F. clarki was abun-
dant in the upper one foot of the limestone cores obtained
from the sink. Thompson (1953, pp. 325-327) compared F.
clarki with F. bocku Moller from Russia and considered it
to be closely similar to F. primaeva from the Big Saline
Formation of the Llano Uplift in northcentral Texas and
with F. fugax from the basal part of the Fra Cristobal For-
mation of the “Derryan” (Atokan) of New Mexico. He
also noted that a large number of small specimens referred
to F. clarki might be conspecific with Profusulinella fittsi.
The later species is typically found 100 feet above the Wap-
anucka Limestone (Morrowan) in the Atoka Formation
(Atokan) of Oklahoma. Fusulinella searighti is reported by
Thompson to be like F. dakotensis Thompson from the Min-
nelusa Formation, South Dakota.

Unklesbay and Palmer (1958) reported one ammonoid
from the Burgner Formation which was Pseudoparalego-
ceras williiamsi = P. kesslerense.

A collection of crinoids made by E. J. Palmer from the
Burgner Formation was studied by W. D. Knapp while a
graduate student at the University of Iowa but is still

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 153

in manuscript. Detailed comparisons with other known Ato-
kan crinoids will be made at a later date, but it may be
stated that some Burgner species show a close affinity
with Atokan crinoids from the Bostwick Formation of the
Ardmore Basin.

It was stated by Hoare (1960, p. 223) that Mesolobus
striatus Hoare does not occur above the Krebs-Cabaniss
boundary in Missouri, and is Desmoinesian in age. The
species was reported to be abundant in the Seville (of Mis-
sour!) and Burgner Formations. The Inola Limestone of
Oklahoma is a correlative of the Seville Limestone of Mis-
souri and Fusulina leei is found about 15 feet below the
Inola, in the Bluejacket Sandstone. As previously noted
Fusulinella clarki and F. searighti are found in the Burgner
and are considered to be Atokan. The Seville Limestone of
Illinois is a different horizon in that Fusulinella iowensis is
present and Fusulina s.s. is absent. Weller and McGehee
(1933, p. 109) gave the range of Mesolobus straitus up into
the Vanport Limestone of Ohio where the highly advanced
Fusulinella carmam Thompson was also found.

VI. FAUNA OF THE ATOKAN STAGE IN THE LLANO UPLIFT, TEXAS

Several fusulinid studies have included forms from the
Llano Uplift, with that by Thompson (1947) being the
most comprehensive, The lower member of the Marble Falls
Formation, Sloan Limestone Member, is considered to be
Morrowan. Staffella expansa Thompson, Pseudostaffella aff.
P. needhami Thompson, Foschubertella texana Thompson,
and Profusulinella marblensis Thompson are reported from
the upper Marble Falls Formation, the Lemons Bluff Mem-
ber. Of much interest here are Thompson’s (1947, p. 163)
remarks concerning P. marblensis which he considered
closely similar to P. fittsi from the lower part of the Atoka
Formation of Oklahoma. It is also close to Profusulinella
copiosa Thompson (1948) from the Arrey Formation of the
lower “Derryan” (Atokan) in New Mexico and extreme
western Texas. It is also noted that Fusulinella llanoensis
(Thomas) is from the upper part of the Big Saline Forma-
tion (Atokan) on the west side of the Llano Uplift. Fusulin-
ella primaeva (Skinner) from the Big Saline Formation is
considered by Thompson (1947) to be younger than Pro-
fusulinella fittsi and about equal to Fusulinella prolifica
in development.

Moore and Plummer (1940) reported several crinoids
from the Marble Falls Formation as Morrowan in age. Of
these Scytalocrinus sansabensis Moore and Plummer (1940)
and Lasanocrinus cornutus Moore and Plummer (1940) are
from the Lemons Bluff Member, Marble Falls Formation,
Atokan. A large fauna is described herein from near the

same locality. The species are, Chlidonocrinus echinatus,
n. sp., Paragassizocrinus altus, n. sp., Araeocrinus bassus,
n. sp., Jsoallagecrinus erectus, n. sp., Globacrocrinus multi-
plicatus, n. sp., Dichocrinus dilatus, n. sp., Lecobasicrinus
subidus, n. sp., and Polusocrinus calyculoides Lane.

Another crinoid fauna is presented herein from the
Soldiers Hole Member, Big Saline Formation, upper Atokan
from near Mason, on the west side of the Llano Uplift. The
species are, Schistocrinus acclivis, n. sp., Sciadiocrinus
llanoensis, n, sp., Goleocrinus masonensis, n. sp., G. confos-
sus, n. sp., Synarmocrinus adornatus, n. sp., Aglaocrinus
magnus (Strimple) Mooreocrinus wilburni, n. sp., Sinocrinus
sheareri, n. sp., Erisocrinus georgeae, n. sp., Graffhamicrinus
antiquus, n. sp., and Ulocrinus zeschi, n. sp. The presence of
Sinocrinus Tien suggests affinity with the Houkou Lime-
stone of northern China,

Plummer (1950, p. 68) listed the following ammonoids
from the Lemons Bluff Member: Phaneroceras compressum
(Hyatt) (—Pseudoparalegoceras compressum), Paralego-
ceras texanum (Shumard), and Branneroceras branneri
(Smith). Upon request for loan, Dr. Peter U. Rodda, Uni-
versity of Texas, advised that he was unable to locate
any specimens from the Marble Falls Formation labeled
Branneroceras branneri in the collections studied by Plum-
mer. The species is typically Morrowan and would not be
expected in the Atokan.

Plummer (1950, p. 71) listed Pronorites lanoensis
Plummer and Scott from the Soldiers Hole Member, Big
Saline Formation and (p. 82) listed Pronorites sp., Pro-
norites arkansasensis Smith, Daralites scotti Miller and Fur-
nish, Nuculoceras swithwickense Plummer and_ Scott,
Phaneroceras compressum (Hyatt) (=Pseudoparalegoceras
compressum), Paralegoceras sp., P. shumardi ( Plummer and
Scott), Glaphurites hyattianus (Girty), G.
(Girty), G. raymondi Plummer and Scott and Gonialobo-
ceras sp. from the Smithwick Shale (Atokan),

angulatus

VII. FAUNA OF THE ATOKAN STAGE IN IOWA

A well-known primitive Fusulinella is F. iowensis
Thompson (1934), the type locality of which is in a small
creek about 200 yards south of the bridge over Cedar
Creek (NW 14 sec. 33, T. 71 N., R. 9 W.) Jefferson County,
Iowa, the horizon being about 20 feet above the Mississip-
pian limestone and about 90 feet below the White Breast
Coal, in the lower part of the Cherokee of Iowa. The Chero-
kee Shale (named for prominent exposures in Cherokee
County, Kansas) is by definition Desmoinesian in age, but
in Iowa what has been called lowermost Cherokee is Atokan
in age. The ammonoid Paralegoceras iowenese (Meek and

154 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Worthen) is from Iowa and is found in Atokan rocks else-
where.

Moore, et al. (1944, p. 691 and Chart 6) noted that the
basal part of the Iowa Pennsylvanian belonged in the zone
of Fusulinella. It was stated, “Rocks called lower Cherokee
on the chart are here assigned to the Lampasan series, al-
though they have not been differentiated previously from
the Desmoinesian.” The correlation was based on Fusulin-
ella iowensis which also occurs in the Seville Limestone of
western Illinois. F. iowensis is closely comparable to F. pro-
lifica from the lower Atokan of Oklahoma.

VII. FAUNA OF THE ATOKAN STAGE IN OHIO

Moore, et al. (1944, Chart 6) correlated the upper and
lower Mercer Limestones with rocks which are currently
considered to be of Desmoinesian age, 1.e., Little Cabin (or
Warner) Sandstone, Hartshorne Sandstone, and Pumpkin
Creek Limestone. The presence of Fusulinella iowensis in the
upper and lower Mercer Limestones indicates a somewhat
older age. We suggest correlation with the Barnett Hill
Member, Atoka Formation of Oklahoma, Seville Limestone
of Illinois, Burgner Formation of Missouri, upper Marble
Falls and Big Saline Formations of northcentral Texas,
lower Hartville Formation of Wyoming, lower Minnelusa
Formation of §. Dakota, and Apodaca Formation of New
Mexico.

1X. FAUNA OF THE ATOKAN STAGE OF NEW MEXICO AND EXTREME
WEST TEXAS

As previously discussed under the section on the Llano
Uplift, Profusulinella copiosa from the Arrey Formation of
Atokan age, near Derry, and in the Mud Springs Mountains
of New Mexico, is apparently closely related to Profusu-
linella fittst, from the lower part of the Atoka Formation of
Oklahoma. Profusulinella apodacensis Thompson (1948)
from near the top of the Apodaca Formation east of Derry,
New Mexico, and in Bed 5, Section 11, is according to
Thompson “— closely similar to, and presumably is closely
related in age to Fusulinella? primaeva from the base of the
Big Saline Limestone of central Texas.” Thompson then
noted that Profusulinella apodacensis is not so advanced
(is older?) as Fusulinella? primaeva.

X. FAUNA OF THE ATOKAN STAGE OF WYOMING AND SOUTH DAKOTA

Fusulinids of Atokan age have been reported from the
Black Hills by Thompson (1936). In the lowermost por-
tion of the Hartville Formation of Wyoming is found Fusu-
linella velmae Thompson and F. protensa Thompson which
are both considered to be closely similar to F. primaeva

from the Llano Uplift of Texas. About 130 feet above the
base of the Minnelusa Formation at Loring Siding, South
Dakota, is found Fusulinella furnishi Thompson and F.
acuminata Thompson. In horizons 60 feet above the base
of the Minnelusa and 90 to 100 feet above the base is found
F. dakotensis Thompson.

X. FAUNA OF THE ATOKAN STAGE OF EASTERN KENTUCKY

The Magoffin and Kendrick beds of the Kentucky
River Valley were shown as uppermost Atokan (Lampasan)
by Moore, et al. (1944, Chart 6), albeit as correlatives of
the McAlester Sandstone and Pumpkin Creek Limestone of
Oklahoma which are now known to be Desmoinesian. In-
vestigations by Furnish and Knapp (1966) of ammonoids
and by Strimple and Knapp (1966) of crinoids from east-
ern Kentucky indicated a Morrowan age for the two beds.
This has been discussed under the Morrowan section here-
in.

XII, FAUNA OF THE ATOKAN STAGE IN NEVADA

Lane (1964), described Synarmocrinus brachiatus Lane
from the Bird Spring Formation at Indian Springs, Nevada,
as being of Atokan Age. The form described by Lane as
Parulocrinus vetulus Lane was from the upper part of the
Rhipidomella nevadensis (Meek) Zone at the base of the
Bird Spring Formation at Arrow Canyon, Nevada. P. vetu-
lus is, therefore, older than Synarmocrinus brachiatus which
is of Atokan age, but we do not believe it is as old as Mor-
rowan.

As noted in the section covering the Morrowan Stage,
Lane (1964b) reported Polusocrinus calyculoides from the
lower part of the Callville Formation at Frenchman Moun-
tain, Clark County, Nevada, which species we have found
in the Lemons Bluff Member, Marble Falls Formation, Ato-
kan, northcentral Texas.

Cassity and Langenheim (1966, p. 937) gave the base
of the Atokan in the Arrow Canyon Section (Bird Spring
Formation) as bed 101 which was marked by the lowermost
occurrence of Profusulinella. They included beds 101 through
bed 112 for a total of 140 feet as the Zone of Profusulinella.
This is roughly equivalent to our zone of Profusulinella
fittst in Coal County, Oklahoma. They show beds 113
through 151 for about 270 feet of rock as the zone of Fusu-
linella, based to a large extent on the presence of Fusulinella
devexa Thompson.

XII, FAUNA OF THE ATOKAN STAGE OF WESTERN ILLINOIS

Thompson, Shaver, and Riggs (1959) attempted to
place Fusulinella iowensis in the zone of Fusulina, 1.e. as

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 155

being of Desmoinesian age. Those authors (tbid., p. 779)
stated under discussion of the species Fusulinella iowensis,
“In many regions it occurs above advanced forms of Fusu-
linella and immediately below primitive Fusulina and primi-
tive Wedekindellina.” Some species of Fusulinella enter into
the zone of Fusulina but this has apparently not been true
of Fusulinella iowensis, therefore, we are unable to see the
justification for considering it to be a Desmoinesian fossil,
in fact we consider it to be upper Atokan in age.

Thompson, Shaver, and Riggs (1959) called attention
to some described and undescribed transitional species of
Fusulina, e.g. F. ? insolita Thompson (1948) which had
been observed to occur with advanced species of Fusulinella.
Those authors proposed that the species be included in the
zone of Fusulina, 1.e. assigned to the Desmoinesian rather
than the Atokan Stage. We do not agree with the proposi-
tion.

Fusulinella iowensis is common in the Seville Limestone
of Illinois (not a correlative of the Seville of Missour1) and
Fusulina s.s. was not found in association. This condition,
that is the absence of Fusulina s.s., was also true in Iowa
in the lower Cherokee (Atokan) and in Ohio in the Upper
and Lower Mercer Limetones,

Another matter is the zone of Profusulinella, long con-
sidered to be lower Atokan in age. In Thompson, Shaver,
and Riggs (1959) Profusulinella kentuckyensis Thompson
and Riggs, 1959, was described as an advanced form of the
genus because it had a highly elongate shape of the shell.
Profusulinella burrensis Thompson and Shaver, 1964, was
reportedly shorter and with a more inflated shell (more
primitive?) but is from a stratigraphically higher horizon
according to Thompson and Shaver. It appears that a divi-
sion was proposed by Thompson and Shaver, which is com-
parable to lowermost Pennsylvanian (Morrowan), in the
middle of the Brazil Formation between the upper Block
and Minshall Members. The lower zone was termed zone of
Amphissites rothi and Profusulinella kentuckyensis appeared
near the top of the zone. If P. kentuckyensis is more ad-
vanced than P. fittst (the latter occurs about 100 feet above
the base of the Barnett Hill Member of the Atoka Forma-
tion in Oklahoma) and less or less advanced than Fusulin-
ella prolifica (which occurs about 250 feet above the base
of the Barnett Hill Member, Atoka Formation), then it
appears that at least some of the zone of Amphissites rothi
must certainly be of Atokan age. Profusulinella was found
in the lower part of the Brazil Formation and Fusulinella
iowensis, together with F. stowti, was found in the upper
part of the Brazil Formation and equivalent horizons of
the Illinois Basin according to those authors. On the basis

of such evidence we strongly suggest an Atokan age assign-
ment to the Brazil Formation.

XIV. FAUNA OF THE ATOKAN STAGE IN MICHIGAN

According to Shiedler and Wanless (1965), the Middle
Pennsylvanian of Michigan includes the Verne Limestone
Member which contains Fusulinella iowensis and is Atokan
or early Desmoinesian in age. This is the same horizon
called “Cyclical formation F” by Kelly (1933) from which
a fauna is obtained that is similar to the fauna of the Web-
ber Falls Member, Atoka Formation of Oklahoma.

XV. FAUNA OF THE ATOKAN STAGE OF THE CANADIAN ARCTIC

An ammonoid fauna was reported by Nassichuk and
Furnish (1965, p. 724) as being remarkably similar to index
species from the “upper Morrowan Winslow Formation” in
Arkansas (of McCaleb, 1963), “or are in a comparable
stage of evolution.” Those authors noted that some authors
regard the “Winslow” as synonymous with the Atoka For-
mation of Oklahoma, which would make the Arctic fauna
basal Middle Pennsylvanian by definition. The Arctic ma-
terial is from a locality on the north shore of Hare Fiord
(lat. 81°07.5’ N.; long. 84°17’W.) and represents part of
an unnamed formation. By present definition the Arctic ma-
terial is of early Atokan age.

XVI. FAUNA OF THE ATOKAN STAGE OF MEXICO

Murray, Furnish, and Carrillo (1960, p. 736) noted that
some Atokan specimens from Oklahoma and Arkansas,
which had been identified as Psewdoparalogoceras williamsi
(= P. kesslerense), were not significantly different from P.
amotapense (Thomas) from Caballeros Canyon, a few kiio-
meters northwest of Ciudad Victoria, capital of the State
of Tamaulipas, Mexico. Associated with the ammonoids
were specimens of fusulinids closely similar to Fusulinella
acuminata Thompson, which species is also reported from
the lower part of the Minnelusa Formation of South Dakota
and the Fra Cristobal Formation, Mud Springs Mountains,
New Mexico.

XVII. FAUNA OF THE ATOKAN STAGE OF SPAIN

Van Ginkel (1965) showed a zone of Profusulinella in
the Spanish Carboniferous extending from low in the Bash-
kirian well into the Kashiran of Russian usage. This would
cover both of Van Ginkel’s Zones A and B, which we con-
sider to be Atokan in age. He then had a zone of Fusulinella
starting in the Kashiran, where it was Subzone A, of Van

Ginkel, (Atokan), and extended through Podolskian and

156 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Myachkovian as Subzone B, of Van Ginkel, in association
with Fusulina (? Desmoinesian).

Sieverts-Doreck (1951) described Jberocrinus multi-
brachiatus from limestone of the lower Upper Carboniferous
(Westphalian B) east of Herruela, province of Palencia, as
the monotype of /berocrinus, The genus is close to the
American genus Megaliocrinus Moore and Laudon (1942)
of Morrowan age. The Westphalian B is considered a cor-
relative of the upper Bashkirian by some authors, which we
consider to be lower Atokan. Breimer (1962) apparently
considered the age to be Moscovian probably because of
the presence of Cromyocrinus sp. cf. C. simplex Traut-
schold and Paradelocrinus sp. The horizon of C. simplex
in Russia is Myachkoyian and apparently correlative of the
Desmoinesian. In our opinion the specimen figured by Brei-
mer is Atokan in age, which is born out by the age given

Other

by Sieverts-Doreck, and the presence of another form identi-
fied as Pimlicocrinus sp. by Breimer from Asturias but
also the Cotarrazzo Limestone. Pimlicocrinus is typically a
Tournaisian genus.

XVIII. FAUNA OF THE ATOKAN STAGE IN RUSSIA

The Atokan Stage as presently defined is apparently
represented in the upper Bashkirian, lower Moscovian, and
the lower portion of the upper Moscovian of Russia.

Ammonoids of the Bashkirian appear to be diagnostic.
Aisenverg, et al. (1960) reported Branneroceras branneri
from C:a, lower Bashkirian of the Donetz Basin. As pre-
viously noted, this 1s characteristic of the Morrowan and
more specifically of the Bloyd Formation in the midcon-
tinental area of North America. The fusulinids Eostaffella
varvariensis and Pseudostaffella antiqua were also listed.

NORTHWES TEXAS
ARKANSAS | MISSOURI IOWA LLANO UPLIFT

DESMOINES-

Eartshorne artshorney Cherokee Millsap Lake
IAW Hartshorne ss.| Upper Lester ? Fn. Sh. (part) Fn.

Webber

Barnett Hill Member

Prolific cepha-
lopod Zone e
Zone of Fusuli-
ATOKAN mella prolifica} 200'-250'
Fm.
Zone of Profu- i
sulinella + Coody ss.

MORROWAN yWwapanucka
OR Zn. LS.
HESTERIAN

Falls ss.

Bostwick

*Otterville

Bloyd Fm.

Atokan
anmonoids
of Gordon
(1965)
Smithwick Sh.

Basal
Burgner Cherokee

Fm.

Big Saline
(part)

Winslow
Member

Sloan Ls.
Bloyd Mi or
Barnett Ls.

Text-figure 1.- Chart showing probably stratigraphic relationship of various selected formations based
on known fossiliferous horizons, with ammonoids, fusulinids and crinoids used as the key fossils.

* The name Otterville is used here as a Morrowan horizon although the formation is of Desmoinesian age

at its type locality.

## Pennsylvanian strata in Iowa is underlain by the Pella beds which are thought to be correlative of the

lower St. Genevieve Formation of the Illinois Basin.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 157

Gastrioceras listeri with Profusulinella primitiva is listed in
C’sb which we believe to be Atokan. Profusulinella rhombi-
formis and Gastrioceras cf. reticulatum are listed from C>se,
uppermost Bashkirian.

Librovitch and Malivkin (1950), in reporting on the
Carboniferous deposits of the Urals, showed among the
characteristic forms Branneroceras cancellatum and Pseudo-
stafella antiqua from the lower Bashkirian. From the upper
Bashkirian they showed Pseudostaffella antiqua together
with Profusulinella ex gr. parva, which we take to be Atokan
in age. They divided the Moscovian into lower and upper
zones with Profusulinella marking the lower (Atokan) and
Fusulina with Wedekindellina marking the upper (Des-
moinesian ).

Miklukho-Maclay (1960), in reporting on the Carboni-
ferous of Central Asia, showed Profusulinella in the Yangak
horizon=upper Bashkirian, and Karatange horizon—Vere-
yan and Kashiran. Fusulinella was shown as characteristic
of lowermost C2mz=Upper Moscovian and specifically the
Podolskian, with a different horizon marked by Fusulina in
the upper Podoloskian. Such findings coincide with other
data indicating the Atokan Stage continues into the Po-
dolskian.

Librovitch and Stepanov (in Gorsky, Stepanov, et al.,
1960, text-fig. 1) indicated the Vereyan and Kashirian hori-
zons were correlative with the Atokan of North America,
but it appears to us that the Atokan starts in the Bash-
kirian and extends into the Upper Moscovian (lower Po-
kolskian horizon).

XIX. SUMMARY

The Atokan Stage is represented in the Llano Uplift of
northcentral Texas by the Marble Falls Formation (ex-
cluding the Sloan Member), the equivalent Big Saline For-
mation and the Smithwick Shale. Essentially the same cor-
relation was given by Plummer (1950, p. 47); however, his
attempt to establish a Marble Falls Group and his use of
the name Bendian or Bend Series has not been generally
accepted. The name Atokan Stage or Series is now in gen-
eral usage,

The type area of the Atokan Stage lies in Atoka Coun-
ty, Oklahoma, where it is represented by the Atoka Forma-
tion, which formation is a rock unit resting between the
Wapanucka Limestone (Morrowan) below and the Harts-
horne Sandstone (Desmoinesian) above. In adjacent Coal
County, north of Clarita, Oklahoma, there is a zone of
Profusulinella fittsi (see Text-fig. 1) which is more or less
correlative of the Marble Falls Formation (excluding the
Sloan Member), the Big Saline Formation and the Smith-

wick Shale of the Llano Uplift, and the Arrey Formation
of New Mexico.

Based on fusulinids, ammonoids, and crinoids, the Bureg-
ner Formation of Missouri, is above the zone of Fusulinella
prolifica (see Text-fig. 1) and is about equivalent to part
of the Bostwick Formation of the Ardmore Basin, southern
Oklahoma. The zone is more or less correlative of the upper
part of the Big Saline Formation of northcentral Texas, the
lower part of the Cherokee Formation of Iowa (not of Kan-
sas ), the lower part of the Hartville Formation of Wyoming,
the Minnelusa Formation of South Dakota, the basal part
of the Fra Cristobal Formation of New Mexico, an un-
named formation at Hare Fiord, Canadian Arctic, an un-
named formation in Caballeros Canyon near Ciudad Vic-
toria, State of Tamaulipas, Mexico, Seville Limestone of

North American
Stages

Desmoinesian

Moscow
Basin
Horizons

Russian
Stages

3
3
53
° ome!
a
g Kashirian
m
Cob
Atokan
b
2
b
4 2
3 d
en Bashkirian
nah 2
ad
fs) b
3 8
s Morrowan
b
a

Fal
Cle
Chesterian

Correlation-chart of some Middle

Namurian

Text-figure 2.

Carboniferous horizons of Russia.

158 PALAEONTOGRAPHICA AMERICANA (VI, 40)

western Illinois, upper and lower Mercer Limestones of
Ohio, upper part of the Brazil Formation and equivalent
horizons of the Illinois Basin, the Bostwick Conglomerate
and Lester Limestone of the Ardmore Basin in southern
Oklahoma, part of the Barnett Hill Member, Atoka Forma-
tion of Coal County, Oklahoma, and the Atoka Formation
near Red Oak, Latimer County, Oklahoma.

The prolific cephalopod zone at the top of the Barnett
Hill Member in southeastern Oklahoma (see Text-fig. 1) is
roughly a correlative of the Webber Falls Sandstone Mem-
ber in northeastern Oklahoma. There is an ammonoid bear-
ing sandstone in northeastern Oklahoma near the base of
the Atokan Stage, the Coody Sandstone Member, Atoka
Formation. The fauna is closely comparable to that of the
“Winslow” of Arkansas and therefore also to the Arctic
ammonoid fauna from Hare Fiord (Nassichuk and Furnish,
1965).

It is probable that crinoids reported by Tien from the
Houkou Limestone of northern China are of Atokan age.
The lower part of the Huanglung Limestone of Chuanshan,
China, appears to be a correlative of part of the lower
Atokan of Oklahoma-Arkansas. The zone of Profusulinella
and subzone A of the zone of Fusulinella of Van Ginkel from
the Cantabrian Mountains of Spain appear to be Atokan
in age. In Russia and Central Asia, it appears that the
upper part of the Bashkirian Stage is of Atokan age as well
as all the Vereyan and Kashirian Horizons and the lower
part of the Podolskian Horizon of the Moscovian Stage.

E. DESMOINESIAN STAGE

Our observations on the Desmoinesian faunas are
highly selective and based primarily on the materials at
hand. Projection of some data attained in conjunction with
the more comprehensive coverage of the Atokan Stage 1s
attempted.

In Iowa where the type locality of the Desmoinesian
is located on the Des Moines River, the Cherokee Shale
is considered to be the base of the stage or series. As pre-
viously discussed under the Atokan Stage, it is generally
recognized that the lowermost part of the Cherokee Shale of
Iowa is of Atokan age. It is our hope that use of the fusu-
linid Fusulinella iowensis as a marker of the upper Atokan
will be generally accepted, because it would clarify and
simplify the situation in Iowa and elsewhere. It is our be-
lief that F. iowensts is, indeed, of Atokan age as has been
demonstrated in the Atokan section of the present study.
Another complication of the situation in Iowa is the diffi-
culty in identifying the White Breast Coal in many areas,
although the coal is used as a stratigraphic point. It is

sufficient to know that F. iowensts is from about 20 feet
above the Mississippian (and reportedly 90 feet below
the White Breast Coal) and that Fusulina leei Thomp-
son, a good Desmoinesian index, is from about 35 feet below
the White Breast Coal. The type horizon of Fusulina leet
is immediately above the Bluejacket Sandstone of Okla-
homa.

Highly developed Fusulinella are found in Ohio, F. sero-
tina in the Putnam Hill Formation of the lower Allegheny
Series, and F. carmani in the Vanport Limestone of the
middle Allegheny Series.

In a study of rocks exposed in Arrow Canyon (Bird
Spring Formation) of Clark County, Nevada, Cassity and
Langheim (1966, p. 937) used the presence of Wedekindel-
lina sp. in bed 152 to mark the base of the zone of Fusulina,
which they consider synonymous to the Desmoinesian Stage.
It is not clear where Fusulina first appears, but F. prima
Thompson is reported in bed 182 and F. Jeei Skinner from
beds 182 and 187. As previously noted F. leet in Oklahoma
is from immediately above the Bluejacket Sandstone Mem-
ber of the Boggy Formation, near the top of the Krebs
Group (Desmoinesian).

Ammonoids of Moscovian age in Russia are rare. A
specimen of Pseudoparalegoceras was first found near Mos-
cow from the Podolsk Horizon (lower upper Moscoyian
Stage) in 1876, according to Ruzhencev (1951) and was
described as Gastrioceras russiense Tzwetaev (1888). Shvet-
zov and Yablokoy, et al. (1937, p. 512) noted that the
cephalopod fauna described by Tzwetaev probably originat-
ed in a dense limestone (bed 9) in the upper part of the
Podolskian in a quarry on the left bank of the Pakhra River
near the town of Podolsk. The species (Pseudoparalegoceras
russiense) appears to be in a stage of development some-
what comparable to forms found in the Boggy Formation,
Krebs Group, of Oklahoma (personal communication from
W. M. Furnish). P. tzwetaevae Ruzhencev (1951) was
found in the southern Urals in the Aktyubinskoi District
in a block of limestone thought to be of Middle Carboni-
ferous age (Moscovian) although the formation of the area
is in the Gielian Stage of the Upper Carboniferous. The
specimens were similar to P. russiense according to Ruz-
hencev. Anthracoceras Frech reputedly has a range from
early Namurian to late Westphalian (late Mississippian to
late Desmoinesian of Pennsylvanian) in both Europe and
Asia, but the genus has been restricted to uppermost Mis-
sissippian and lower Morrowan (Lower Pennsylvanian) in
North America, with the genus Gordonites Miller and Fur-
nish following in higher strata.

Shvetzov, Yablokov, et al. (1937, p. 53) listed Fusu-

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 159

linella cylindrica and F. bockii from bed 23 of the Podolsk
Quarry which was a horizon in C™, (Myachkovian) about
14 feet above its base, and about 36 feet above the zone
which contained Pseudoparalegoceras russiense in the Po-
dolskian.

Miklukho-Maclay (1960, text-fig. 2) showed a division
in the lower part of Stage Com2 (Upper Moscovian) in
Central Asia as between “with Fusulinella” below and “with
Fusulina” above and correlated the combined zones with
the Podolsk Horizon of the Moscow Basin. In the correla-
tion chart of Librovitch and Stepanov (in Gorsky, Step-
anoy, et al., 1960, text-fig. 1) the combined horizons of
Podolskian and Myachkovian were shown to be Desmoi-
nesian, whereas the data furnished by Miklukho-Maclay in-
dicated that at least part of the Podolskian is Atokan.

Subzone B of zone of Fusulinella of Van Ginkel (1965)
in the Cantabrian Mountains of Spain is marked by the
presence of Fusulina and is probably Desmoinesian in age.

The well-known crinoid fauna from the “Kalkbriiche
von Mjatschkowa” near Moscow, described by Trautschold
(1879), is currently ascribed to the horizon Com = Mosc-
ovian (see Yakovlev and Ivanov, 1956). The cromyocrinids
described by Trautschold in 1879 as Cromyocrinus simplex,
Cromyocrinus geminatus — Mooreocrinus geminatus and
Cromyocrinus ornatus = Dicromyocrinus ornatus, are all
haracterized by having uniserial arms and three anal plates
within the posterior interradius of the dorsal cup. In this
stage of development they could be late Atokan or early
Desmoinesian in age.

Two well-known crinoid horizons in the Desmoinesian
of Texas are the shale below the Kickapoo Falls Limestone
Member, Millsap Lake Formation and the Brannon Bridge
Limestone Member, Millsap Lake Formation. Our collec-
tions from the Kickapoo Falls Limestone were obtained be-
low Kickapoo Falls in northern Hood County and from
the Brannon Bridge Limestone from about three miles
southwest of Brock, Parker County, Texas. The following
species were reported by Moore and Plummer (1940) from
the Hood County locality: Malaiocrinus parviusculus Moore

and Plummer = Glaukosocrinus parviusculus, Paradelo-
crinus brachiatus Moore and Plummer, Paradelocrinus sub-
planus Moore and Plummer = Cricocrinus subplanus,

Pirasocrinus scotti Moore and Plummer, Plaxocrinus obesus
Moore and Plummer, Plaxocrinus perundatus Moore and
Plummer, and Sciadiocrinus harrisae Moore and Plummer.

The following species are now added to those above:
Stellarocrinus bulbus, n. sp., Lecobasicrinus kickapooensts,
n. sp., Eiromocrinus grossus, n. sp., Stenopecrinus longus,
n. sp. and Platycrinites remotus, n. sp.

The following species were reported by Moore and
Plummer (1940) from the Parker County locality: Synero-
crinus formosus Moore and Plummer, Apollocrinus florealis
Moore and Plummer = Stellarocrinus florealis, Brychio-
crinus texanus Moore and Plummer = Stellarocrinus tex-
anus, Endelocrinus rectus Moore and Plummer, Ethelocrinus
millsapensis Moore and Plummer = Parethelocrinus millsap-
ensis, Haeretocrinus magnus Moore and Plummer, Neozea-
crinus praecursor Moore and Plummer = Anobasicrinus
praecursor, Schistocrinus parvus Moore and Plummer =
Sciadtocrinus parvus, Schistocrinus planulatus Moore and
Plummer = Sciadiocrinus planulatus, Sellardsicrinus marrs-
ae Moore and Plummer, Texacrinus gracilis Moore and
Plummer.

The following species have been added to those above
in the present study from Parker County: Sellardsicrinus
fortunatus, n. sp., Aesiocrinus annulatus, n. sp., Ramulo-
crinus consectatus, n. sp., Microcaricrinus delicatus, n. sp.,
Anchicrinus toddi, n. sp., Polygonocrinus libratus, n. sp.,
Oklahomacrinus frostae, n. sp. and Ulrichicrinus ramosus,
n. sp.

Plummer and Scott (1937, p. 15) proposed a Pseudo-
paralegoceras brazoense Zone below the Kickapoo Falls
Limestone Member, Millsap Lake Formation, not far above
the lowest preserved rock of the Millsap Lake Group as ex-
posed in the Brazos River Valley. One of our crinoid faunas
was obtained from the same horizon and probably from near
the same exposure on Kickapoo Creek.

The Gastrioceras venatum assemblage or zone of Plum-
mer and Scott (1937, p. 385) is apparently from higher in
the Millsap Lake Group and, according to Beghtel (1962,
pp. 17, 18), is probably slightly older than the fauna of the
Wewoka Formation of Oklahoma. This, then indicates that
the crinoid fauna from the Brannon Bridge Limestone Mem-
ber, and more particularly the crinoid fauna from below the
lower Kickapoo Falls Limestone Member, is older than the
Wewoka Formation.

F. MISSOURIAN STAGE

A fauna composed primarily of numerous small sponges
and crinoids is found in a shale of the Captain Creek Lime-
stone Member, Stanton Limestone Formation of southeast-
ern Kansas, at many localities. The zone is apparently a cor-
relative of the horizon previously known as the “Lake
Bridgeport Shale Member,” Graford Formation of Wise
County, Texas. The shale is currently referred to the Wolf
Mountain Shale Member. Components of the crinoid fauna
are found in the Wann Formation of northeastern Okla-
homa, but the sponges are relatively rare. The exact corre-

160 PALAEONTOGRAPHICA AMERICANA (VI, 40)

lative of the Captain Creek Limestone has not been satis-
factorily established in Oklahoma. The predominantly cal-
careous formations of Kansas are to a large degree replaced
by shale in Oklahoma so that detailed comparisons are diffi-
cult. Some unusual crowns from near Lake Bridgeport,
Wise County, Texas, are figured herein.

One crinoid is figured from a shale exposure 2/4 miles
east and 2% miles north of Rochelle, McCulloch County,
Texas, as Parethelocrinus plattsburgensis (Strimple). The
exposure is numbered 153-T-23 in Moore and Plummer
(1940, p. 396), Mineral Wells Formation, from which the
species Athlocrinus nitidus Moore and Plummer, Plaxocrinus
omphaloides Moore and Plummer, P. lobatus Moore and
Plummer and Delocrinus benthobatus Moore and Plummer
were previously obtained and reported.

Plummer and Scott (1937, p. 17) proposed a “Para-
schistoceras reticulatum Zone,” in a shale member of the
Graford Formation, low in the Mineral Wells Group, Mis-
sourian. The species Glaphrites angulowmbilicatus Plummer
and Scott, 1937, was listed from the same zone and was
from the abandoned brick pit in Bridgeport, Wise County,
Texas. Specimens of Paragassizocrinus tarri (Strimple) have
been collected by the senior author from the Bridgeport
locality. It is of interest, although not considered by us
as conclusive correlative data, that specimens thought to be
conspecific with Glaphyrites angulowmbilicatus have been
found in association with Paragassizocrinus tarri in a hort
zon which appears to be a shale of the Captain Creek Lime-
stone Member, Stanton Limestone Formation, three miles
east of Wayside, Montgomery County, Kansas. The horizon
is a few feet above a crinoid-sponge horizon and was col-
lected by W. M. Furnish and H. L. Strimple in February,
1966 and is reposited in the Department of Geology, Uni-
versity of Iowa.

The following crinoids were reported by Strimple
(1951a) from the sponge bearing horizon of the Graford
Formation near Lake Bridgeport, Texas: Laudonocrinus
subsinuatus (Miller and Gurley), Delocrinus pictus Moore
and Plummer, Delocrinus subhemisphericus Moore and
Plummer, Endelocrinus tumidus Strimple, Endelocrinus gra-
fordensis Moore and Plummer, Erisocrinus sp., Athlocrinus
sp., Lecythiocrinus sp. (radial plate only), Ulocrinus sp.
(IBB and portion of BB circlet), Apographiocrinus typicalis
Moore and Plummer, Apographiocrinus cf. arcuatus Strim-
ple, Stellarocrinus sp., Elibatocrinus sp. (IBB circlets), Am-
phicrinus sp., Delocrinus sp., Utharocrinus cf. granulosus
Strimple, Graphiocrinus bridgeportensis Strimple, Piraso-
crinus invaginatus Strimple, Stellarocrinus texani Strimple,
Cibolocrinus erectus Strimple, Allagecrinus basslert nodosus

Strimple = Jsoallagecrinus bassleri (Strimple), Allagecrinus
bassleri status Strimple = Isoallagecrinus status (Strimple),
Perimestocrinus moseleyi Strimple = Stenopecrinus mose-
leyi, Plaxocrinus laxus Strimple, Plaxocrinus oeconomicus
Strimple, Euonychocrinus subservire Strimple.

A crown of Ulocrinus convexus (Strimple) is illustrated
herein as well as a crown of Stellarocrinus sp. cf. S. angula-
tus from the Lake Bridgeport locality.

SYSTEMATIC PALEONTOLOGY
Class CRINOIDEA Miller
Subclass INADUNATA Wachsmuth and Springer
Order CLADOIDEA Moore and Laudon
Suborder DENDROCRINOIDEA Bather

Family CROMYOCRINIDAE Jaekel

The family Cromyocrinidae was proposed by Jaekel
(1918) for two genera, which were Cromyocrinus Traut-
schold and Dicromyocrinus Jaekel. The number of genera
has increased until a total of eight genera were ascribed to
the family by Strimple (1961la, pp. 63-65). All of those gen-
era are included below except Cryphiocrinus Kirk and
Tyrieocrinus Wright which we do not feel are closely related.
The genera Mooreocrinus Wright and Strimple, Parulocrinus
Moore and Plummer, Paracromyocrinus Strimple, Synarmo-
crinus Lane, and Goleocrinus Strimple and Watkins have
been added to the family.

ANALYSIS OF CROMYOCRINIDS
Genus CROMYOCRINUS Trautschold, 1867

Type species. —C. simplex Trautschold.

Range. — Moscovian (Atokan and Desmoinesian), Up-
per Carboniferous, Russia.

Three anal plates, upflared infrabasals, five uniserial
arms.

Genus DICROMYOCRINUS Jaekel, 1918

Type species. —D. ornatus (Trautschold).

Range. — Moscovian, (Atokan and Desmoinesian) Up-
per Carboniferous, Russia.

Three anal plates, subhorizontal infrabasals, ten uni-
serial (cuneiform) arms, pustulose surface.

Genus MOOREOCRINUS Wright and Strimple, 1945
Type species. —M. geminatus (Trautschold).

Texan CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 161

Range. — Moscovian (Atokan and Desmoinesian), Up-
per Carboniferous, Russia, Brazil, North America.

Three anal plates, subhorizontal infrabasals without
basal depression, ten uniserial arms.

Genus PARULOCRINUS Moore and Plummer, 1940

Type species. — P. blairi (Miller and Gurley).
Range. — Missourian, Upper Carboniferous, and Lower
Permian, North America.

Two anal plates, subhorizontal infrabasals without basal
depression, arms unknown.

Genus PARACROMYOCRINUS Strimple, 1966a

Type species. —P. vetulus (Lane).

Range. — Morrowan to Missourian, Upper Carbonifer-
ous, North America.

Two anal plates, subhorizontal infrabasals, basal plates
curve into basal concavity, ten biserial arms.

Genus METACROMYOCRINUS Strimple, 1961

Type species.— M. holdenvillensis Strimple.

Range. — Morrowan, Atokan, and Desmoinesian, Up-
per Carboniferous, North America, China.

Three anal plates in older species and two in younger
species, subhorizontal to mildly upflared infrabasals, ten
biserial arms, finely postulose ornamentation.

Genus SYNARMOCRINUS Lane, 1964b

Type species. —S. brachiatus Lane.

Range. — Atokan, Upper Carboniferous, North Amer-
ica.

Two anal plates, subhorizontal to mildly downflared

infrabasals, base of cup flat?, ten uniserial arms, pustulose
ornamentation.

Genus ULOCRINUS Miller and Gurley, 1890

Type species. — U. buttsi Miller and Gurley.

Range. — Atokan to Virgilan, Upper Carboniferous and
Lower Permian, North America, India?, Timor?, and Rus-
sia.

Two anal plates, upflared infrabasal plates, ten bi-
serial arms.

Genus GOLEOCRINUS Strimple and Watkins, new genus

Type species.— G. masonensis Strimple and Watkins.

Range. — Atokan-Desmoinesian, Upper Carboniferous,
North America.

Three anal plates, downflared infrabasals, smooth sur-
face.

Genus UREOCRINUS Wright and Strimple, 1945

Type species. —U. bockschii (Geinitz).

Range. — Tournaisian, Lower Carboniferous, Scotland.

Three anal plates usually Advanced Type with radi-
anal dominant, upflared infrabasals, five uniserial arms.

Genus MANTIKOSOCRINUS Strimple, 1951le

Type species. — M. castus Strimple.

Range. — Chesterian and Tournaisian, Lower Carboni-
ferous, North America, Scotland.

Three anal plates, upflared infrabasals, ten uniserial
arms.

Genus MOOREOCRINUS Wright and Strimple, 1945

Type species.—Cromyocrinus geminatus Trautschold,
1867 M. geminatus (Trautschold),

Range. — Atokan; SSSR, Brazil, North America.

Diagnosis. — Cup medium, globular, greatest width be-
low the summit of the cup, subhorizontal infrabasal circlet,
flat base with no concavity involved (other than depression
for proximal columnal); sutures impressed, surface of plates
essentially smooth; three anal plates in cup; ten uniserial
arms branching with first primibrach.

The following species are assigned to Mooreocrinus:

Cromyocrinus geminatus Trautschold Moscovian, Desmoinesian; SSSR

(non Wright)
Dicromyocrinus mendesi Lane
zil

Mooreocrinus wilburni, n. sp. Atokan; Texas

Kemarks. — The form identified as Cromyocrinus gem-
inatus by Wright from the Lower Carboniferous of Scot-
land has been named Cromyocrinus wrighti Yakovlev and
Ivanoy, 1956 and subsequently assigned to the genus Manti-
kosocrinus by Strimple (1966a, p. 6). Ivanov (1926, p. 176)
listed several crinoids from the Upper Carboniferous near
Moscow among which were “Trautscholdicrinus geminatus
Trel. sp.” and “Trautscholdicrinus ornatus Trel. sp.” There
were no indication, description, or definition of a new generic
name and it is, therefore, nomen nudum.

MOOREOCRINUS WILBURNI Strimple and Watkins, new species
Plate 39, figures 12, 13; Plate 49, figures 4-6

Dorsal cup medium bowl-shaped with erect lateral sides
and broad flattened basal area. The posterior side of the

Itaituba Formation, Atokan; Bra-

162 PALAEONTOGRAPHICA AMERICANA (VI, 40)

cup is flattened and otherwise the cup has a rounded con-
tour when viewed from above or below. Some protrusion
of the cup is apparent in the area occupied by the large
right posterior basal and the radial. The infrabasals form a
large pentagonal-shaped platform with a large center por-
tion depressed for the reception of a round proximal col-
umnal. The perimeter of the infrabasal circlet is slightly up-
flared, though not visible in side view of the cup, so that
the circlet is strongly delimitated from the basal circlet. Five
basals are prominent in side view of the cup and are tumid.
The right posterior basal is larger than the other basals.
Five radials are wider than high and are somewhat tumid.
The outer surface of each radial terminates abruptly at the
distal limit of the cup and a horizontal area is formed to
the fore of the ligamental area. The outer ligamental fur-
row is a thin, shallow depression, and the outer ligamental
ridge is marked by denticles. The ligamental pit furrow is
thin but the ligamental pit is well defined. The transverse
ridge is not prominent and only a few denticles are pre-
served. Oblique ridges and lateral furrows are indistinct,
but a well-defined intermuscular furrow and central pit
are developed. Muscle areas are marked by finger-like shal-
low grooves. Adsutural slopes are low and narrow. Rem-
nants of low lateral ridges have been noted in some ad-
sutural areas. Three large anal plates are preserved in the
types in normal (Primitive) positions. The radianal is
large and elongated and the anal X is also relatively large.
RX is small and high in the cup.

Sutures of the cup are impressed and marked by a
series of pits although it is not so apparent along some
sutures of the large holotype as it is in the paratype.

Measurements of the holotype in millimeters:

Width of cup (maximum) 28.5
Width of cup, posterior to anterior (minimum) 25.5
Height of cup 15.1
Width of infrabasal circlet 10.0
Length of basal (right anterior) 14.4*
Width of basal (right anterior) 14.4*
Length of suture between basals 8.3*
Length of radial, to transverse ridge (right anterior) 12:5%
Length of radial to outer lip of cup (right anterior) Os
Width of radial (right anterior) 15.0*
Length of suture between radials 5.8
Length of radianal 11.9*
Width of radianal 9.0*
Length of anal X ?
Width of anal X (approximate) 6.8
Diameter of columnar scar 5.6

* Measurements taken along surface curvature.

Remarks. — Mooreocrinus wilburni is closely related to
M. geminatus (Trautschold). The latter has proportionate-
ly larger infrabasal plates and the subhorizontal facet to
the fore of the outer ligamental furrow is sloped slightly

outward and has a mildly irregular surface. In M. wilburni
the facet to the fore of the outer ligamental furrow is re-
markably smooth and is horizontal. M. mendesi Lane has
more tumid cup plates than M. wilburni.

The holotype was collected by Mrs. Robert L. George
of San Antonio, Texas, and the paratype by Mr. Wilburn
Shearer of Mason, Texas. The species is named wilburni in
acknowledgment of the generous assistance given in the
field and in the donation of specimens to our study.

Types. — Holotype USNM, No. $5125 and paratype
USNM, No. $5168 reposited in the U.S. National Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch, Mason,
Mason County, Texas.

Genus DICROMYOCRINUS Jaekel, 1918

Type species.— Cromyocrinus ornatus Trautschold.

Range. — Morrowan to Desmoinesian; SSSR, North
America and South America.

A recent discussion of this genus was given by Strimple
(1966a, p. 5) and a comprehensive history of it is to be
found in Strimple (1961la, p. 66).

DICROMYOCRINUS OPTIMUS Strimple, 1951d
Plate 40, figures 5-8

This species was referred to Ataxtacrinus Strimple
(196la, p. 90) == Tarachiacrinus Strimple (1962a), with
reservation. It is now known that the species, based on a
metatype, has ten uniserial arms. Because the cup is also
ornate, there is no longer any reason to remove it from the
genus Dicromyocrinus. The type of the genus, D. ornatus,
has a broader infrabasal circlet, and the arms tend toward
cuneiformity with more pronounced ornamentation.

The figured specimen from near Lampasas, Texas, is
close to the holotype of D. optimus as well as to the meta-
type with arms and is considered by us to be conspecific.
The Lampasas specimen does not disclose as pronounced
ornamentation, has some protrusion of the area surrounding
the radianal, and the infrabasal circlet is slightly asym-
metric.

Types. — Figured hypotype USNM, No. $5129 collect-
ed by Wm. T. Watkins, reposited in U.S. National Museum.
Metatype with arms SUI, No. 32257 reposited in Geology
Department, University of Iowa.

Occurrence. — Holotype and metatype are from the
Brentwood Limestone Member, Bloyd Formation, Morrow-
an, Pennsylvanian; spillway of Greenleaf Lake southeast of

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 163

Muskogee, Muskogee County, Oklahoma. Figured hypotype
is from the Marble Falls Formation, Morrowan?, Pennsyl-
vanian; bank of Espey Creek about 3.5 miles southwest of
Lampasas, Texas.

DICROMYOCRINUS sp. cf. D. TEXASENSIS
(Moore and Plummer)
Plate 54, figures 8, 9

The crown is long and slender. Dorsal cup is low, bowl-
shaped with a broad flattened base and erect sides. There
is a basal depression in which the proximal edges of the
basals participate, but the infrabasal circlet appears to be
subhorizontal in position. Preservation of the cup is im-
perfect in the basal area. The dorsal cup rises evenly to an
area just to the fore of the outer ligamental fossae where
a small subhorizontal shelf is formed. This development
gives a gaped appearance at the juncture with the arms
but is not interpreted as a constriction of the cup. All plates
are covered by rounded pustules and the sutures are im-
pressed. Five tumid basals form the basal plane and flex
upward to form a considerable portion of the cup walls. Five
radials are pentagonal-shaped, slightly wider than long.
Three anal plates occupy the posterior interradius, although
the RX is narrow and high in the cup. Anal X is in contact
with the posterior basal.

Ten long arms are uniserial and branch with slightly
elongated first primibrachs in all rays. There is a mild cur-
vature to the exterior of the arms and a peculiar develop-
ment wherein alternating brachials are projected to the
side and met by depressed areas in the sides of adjacent
arms. The condition continues into the distal section of the
arms and provides for an interlocking of arms which is
discussed in a special section of this study. The secundi-
brachs are slightly cuneiform in structure and are slightly
elongated.

Measurements of the hypotype in millimeters:

Length of crown 54.6
Height of dorsal cup 7.0
Width of dorsal cup 13.2
Length of basal thee
Width of basal q2*
Length of suture between basals 4.0*
Length of radial 5.0*
Width of radial 7.0*
Length of suture between radials 3.0
Length of radianal 4.0
Width of radianal 2.0

* Measurements taken along surface curvature.

Remarks. — Dicromyocrinus texasensis is more ad-
vanced than most species of Metacromyocrinus in that
three anal plates are present, albeit the RX is obviously
commencing to migrate out of the cup and does in some

specimens. The hypotype is closely comparable to some
forms from the Brentwood Limestone which have been
ascribed to Metacromyocrinus oklahomensis by Strimple
(1962b, pp. 270, 272) and more recently to Paracromyo-
crinus Strimple (1966a, p. 5). In the study by Strimple
(1962b) it was recommended that the species D. texasensis
be placed in synonymy with M. oklahomensis. There is a
large crown known from the Wapanucka Formation which
has ten thick, biserial arms that has been ascribed to
the species M. oklahomensis. In finding the Marble Falls
D. texasensis to have uniserial arms a revision of the species
is required. More study will be required before the inter-
relationships will be fully understood.

Synarmocrinus adornatus has a compact cup much like
that of Dicromyocrinus texasensis but has a highly ad-
vanced arrangement of anal plates in the posterior inter-
radius.

Type. — Hypotype, USNM, No. 85190 collected by
Wm. T. Watkins, reposited in the U.S, National Museum.

Occurrence. — Marble Falls Formation, Atokan, Penn-
sylvanian; Lambert Ranch, San Saba County, Texas.

Genus PARULOCRINUS Moore and Plummer, 1940

Type species. — Ulocrinus blairi Miller and Gurley,
1893.

Range. — Atokan to Lower Permian; North America.

Diagnosis. — Dorsal cup approximately twice as wide
as high, mildly constricted at summit; two anal plates;
subhorizontal infrabasal circlet; proximal edges of basals
form basal plane, no basal concavity; no pronounced sur-
face ornamentation; arms unknown.

Species assigned to Parulocrinus:
Missourian: Missouri

Missourian: Texas

Lower Permian: Texas
Missourian: Kansas

Ulocrinus blairi Miller and Gurley
Paulocrinus beedei Moore and Plummer
Phialocrinus americanus Weller
Ulocrinus caverna Strimple

Remarks. — The genus Parulocrinus was placed in syn-
onymy with Ulocrinus by Strimple (196la, p. 68). Lane
(1964b) accepted the genus Parulocrinus as being valid and
thereafter Strimple (1966a, p. 4) emended the generic con-
cept to comply with characters of the type species. In this
manner some of the confusion surrounding the genus has
been eliminated.

The species described as Ethelocrinus plattsburgensis
Strimple was referred to Parulocrinus by Moore and Plum-
mer (1940); however, subsequent knowledge of the arms
(more than ten) led to inclusion in the genus Parethelo-
crinus Strimple (196la, pp. 80, 82, 85).

Genus PARACROMYOCRINUS Strimple, 1966a
Type species. — Parulocrinus vetulus Lane, 1964b.

164 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Range. — Atokan to Missourian; North America.

Diagnosis. — Dorsal cup normally more than twice as
wide as high, broad base, erect lateral sides, basal plates
curve into basal concavity, infrabasal circlet subhorizontal,
two anal plates, ten broad biserial arms.

Species referred to Paracromyocrinus:

Morrowan:Nevada
Desmoinesian: Texas

Parulocrinus vetulus Lane
Parulocrinus marquisi Moore and Plummer
Parulocrinus compactus Moore and Plummer
Missourian: Texas, Kansas
Parulocrinus pustulosus Moore and Plummer Missourian: Texas
Ethelocrinus oklahomensis Moore and Plummer
Morrowan: Oklahoma, Texas

Remarks. — The genus Paracromyocrinus was proposed
for forms which are similar to Parulocrinus other than in the
nature of the base which is mildly convex or flattened in
the latter but has a concavity in Paracromyocrinus. The
genus Goleocrinus has a basal concavity but the infra-
basals are sharply downflared whereas in Paracromyocrinus
the infrabasals are more or less subhorizontal in attitude.
Forms having rugose surface ornamentation are included
with forms having smooth surfaces pending additional in-
formation on which division may be made. Paracromyo-
crinus oklahomensis is now known to normally have three
anal plates rather than two.

Genus SYNARMOCRINUS Lane, 1964b

Type species. —Synarmocrinus brachiatus Lane, 1964b.
Range. — Atokan, Pennsylvanian; North America.
Diagnosis. — Dorsal cup approximately twice as wide
as high, subhorizontal to mildly downflared infrabasals, base
of cup flat?, surface marked by stout elongate tubercles;
two anal plates in cup; arms ten, broad, uniserial, branch-
ing on first primibrach; segments low, exterior curved.
Species assigned to Synarmocrinus are as follows:
Atokan: Nevada

Atokan: Oklahoma
Atokan: Texas

Synarmocrinus brachiatus Lane
Synarmocrinus fundundus Strimple
Synarmocrinus adornatus, n. sp.

Remarks. — The genus Synarmocrinus retains primi-
tive uniseral arms, as do the Desmoinesian genera Mooreo-
crinus and Cromyocrinus but has advanced in the arrange-
ment of anal plates so that only two are left within the dor-
sal cup. Actually the RX may be identifiable at the cup
summit in S. fundundus and S. adornatus but as a rudi-
mentary element. The pustulose surface ornamentation and
flattened base are common characteristics.

SYNARMOCRINUS ADORNATUS
Strimple and Watkins, new species
Plate 37, figures 1-3

Dorsal cup is medium, bowl-shaped with broad, shallow

basal concavity and a subhorizontal infrabasal circlet. The
lateral sides of the cup are erect and some constriction takes
place as the summit is approached. Five infrabasals form a
pentagonal-shaped platform but most of the circlet is occu-
pied by a large circular depression for the columnar at-
tachment. Five basals form the lateral sides of the basal
concavity, flex into the basal plane of the cup and thence
into the lateral walls of the cup. Posterior basal is trun-
cated for the reception of the large radianal which is
obliquely placed on the distal edge of the basal. Five
radials are large, wide plates with those of the right and
left posterior having less width than the other three. The
radianal is moderately large, in full posterior position and
carries a small anal X to the left, barely within the cup
and a small RX above, out of the cup. Ornamentation of
cup plates consists of masses of pustules and elongated
tubercles which intertwine. Sutures between plates rest in
V-shaped depressions and are marked by series of depres-
sions or pits.

The species name adornatus is from the Latin word
adorno, meaning decorate, and is applicable to the pro-
nouncedly ornate surface of the cup plates.

Measurements of the holotype in millimeters:

Width of cup (maximum and minimum) 24.5
Height of cup, normal 11.3
Height of cup, left anterior side 12.0
Width of infrabasal circlet 9.2
Length of basal (right anterior) 12am
Width of basal (right anterior) 13.6*
Length of suture between basals 6.6*
Length of radial (right anterior) 8.7*
Width of radial (right anterior) 15.7*
Length of suture between radials 4.0
Length of radianal 5.2
Width of radianal 5.9
Width of columnar attachment scar 4.6

* Measurements taken along surface curvature.

Remarks. — Synarmocrinus adornatus has a more ad-
vanced arrangement of anal plates in the posterior inter-
radius than other species assigned to the genus. The de-
velopment is an excellent example of an evolutionary trend
called “Developmental Trend B” as “Special Type D-1”
by Strimple (1960b, pp. 249-250, text-fig. 2). S. fundundus
is readily separable in having a pronounced protrusion of
the cup in the area occupied by its radianal.

Holotype. —USNM, No. S5118 collected by Wm. T.
Watkins. Reposited in the U.S. National Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch, Mason
County, Texas.

Genus ULOCRINUS Miller and Gurley, 1890

Type species.— Ulocrinus buttsi Miller and Gurley,
1890.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 165

Range. — Atokan to Permian; SSSR, India?, Timor?,
North America.

Diagnosis. — Dorsal cup medium, irregularly bowl-
shaped with prominent upflared infrabasals, three anal
plates in older species but typically only two anal plates,
ten biserial arms, outer surfaces essentially devoid of orna-
mentation, round stem.

Species assigned to the genus Ulocrinus:

Ulocrinus buttsi Miller and Gurley Missourian: Missouri, Oklahoma
Cyathocrinus sangamonensis Miller and Gurley Missourian: Illinois
Ulocrinus kansasensis Miller and Gurley Missourian: Missouri
Ethelocrinus convexus Strimple Missourian: Oklahoma, Texas
Ulocrinus occidentalis Miller and Gurley Pennsylvanian: Missouri
?Cyathocrinus goliathus Waagen Permian: India
?Ulocrinus indicus Wanner Permian: Timor
?Ulocrinus conoideus Wanner Permian: Timor
Ulocrinus uralensis Yakovlev Permo-Carboniferous: SSSR
Ulocrinus elongatus Strimple Desmoinesian: Oklahoma
Ulocrinus extrorsus, n. sp. Desmoinesian: Texas
Ulocrinus zeschi, n. sp. Atokan: Texas

Remarks. — Several forms which have been considered
to belong to the genus Ulocrinus are to be found herein
under the genus Parulocrinus. The only characteristic, which
may be used with assurance in separation of the forms, is
the decidedly upflared and prominent infrabasals of Ulo-
crinus. In Parulocrinus, as currently emended, the infra-
basals are not visible or are barely visible in side view of
the cup.

A species from the Millsap Lake Formation of Texas,
which is stratigraphically nearly equivalent to the Wewoka
Formation of Oklahoma, is described as Ulocrinus extror-
SUS, N. Sp.

Specimens of Ulocrinus convexus (Strimple) from the
Wolf Mountain Shale, Graford Formation of Texas, which
is stratigraphically equivalent to parts of the Wann Forma-
tion in Oklahoma and the Stanton Formation in Kansas,
give vital information relative to the arms structure of this
species. Ten thick, broad, decidedly biserial arms are pres-
ent.

Specimens of Ulocrinus zeschi, n. sp. from the Big Sa-
line Formation, Atokan of Texas, have three anal plates,
but the shape of the cup is more like that of Ulocrinus than
of Cromyocrinus, i.e. it is more elongated. The type species
of the genus Cromyocrinus, C. simplex, is Desmoinesian in
age and is eliminated from the normal lineage of the cromyo-
crinids in having reduced the number of arms to five.

ULOCRINUS EXTRORSUS
Strimple and Watkins, new species
Plate 56, figures 6, 8

This species is represented by a single dorsal cup which
is somewhat flattened by lateral compression but all es-

sential features are readily apparent. The cup would be of
moderate height, broad, expanding from the flat columnar
attachment area to just above mid-length of the basal
plates thence constricting slightly. All cup plates are tumid
causing the sutures to appear to be deeply impressed.

There are five upflared IBB forming a low bowl. The
area for attachment of the stem is sharply impressed and
the proximal columnal does not completely fill the cavity.
Five basals are large hexagonal plates, except in posterior
and right posterior where extra facets are developed for con-
tact with the anal plates. Five radials are somewhat smaller
than basals, pentagonal, and constrict at the summit of the
cup. The right and left posterior radials are not so large
as other radials. A subhorizontal area almost 2 mm in
length is found to the fore of the outer ligamental area.
This measurement is taken from the outer face of the radial
which is somewhat below the summit of the cup. Two anal
plates are present, anal X resting on a narrow upper face
of the posterior basal and extending about 5 mm above
the summit of the cup, and RA resting obliquely on the
right shoulder of posterior basal. Anal X is elongated and
followed unequally by two tube plates, the larger being on
the right. RA is elongated, quadrangular and entirely with-
in the cup. First primibrachs are axillary, wide low plates
with sloping lateral sides which sharply reduces the width
of the plates. A few lower secundibrachs are preserved but
not in place. They have a strongly curved exterior and a
broad ambulacral groove. The two preserved proximal col-
umnals are thin, round, and marked by crenulations and
preservation is too poor for more intimate observation.

The name extrorsus is Latin for “in an outward direc-
tion” with reference to the sides of the cup.

Measurements in millimeters:

Height of cup 20.0
Width of cup 25.5e
Length of infrabasal 9:5™
Width of infrabasal 8.6*
Length of basal 14.6*
Width of basal 13.4*
Length of radia] (to transverse ridge) 14.0*
Width of radial 14.0*
Length of RA 14.6*
Width of RA 11.0*-
Length of anal X 8.0*
Width of anal X 4.0*
Length of PBriax 11.0*
Width of PBriax 10.6*
Diameter of proximal columnal 4.7

e-Estimated

* Measurements taken along surface curvature.

Remarks.— There are only two other species of Ulo-
crinus known from rocks older than Missourian. U. elongatus
Strimple, 1961a, is from the Holdenville Formation (upper-

166 PALAEONTOGRAPHICA AMERICANA (VI, 40)

most Desmoinesian). As the name implies it has a rela-
tively long cup, it has no appreciable constriction at the
summit of the cup, and the cup plates have no tumidity.
Ulocrinus zescht, new species is from the Big Saline Forma-
tion (Atokan). U. extrorsus has tumid cup plates, con-
striction as the summit is approached, and the cup is rela-
tively short. Cromyocrinus grandis Mather of Morrowan
age has three anal plates and a relatively low cup and may
be ancestral to U. extrorsus. A long anal plate in contact
with posterior basal is a character of Ulocrinus sangamon-
ensis (Meek and Worthen), the specimen figured as U.
buttst Miller and Gurley, by Wright and Strimple (1945),
and U. convexus (Strimple, 1939a). The RA has lost con-
tact with the right posterior basal in U. buttst, which is also
an elongated species. U. convewxus is probably the most com-
parable in general structure, but it does not have a pro-
nounced constriction at the summit of the cup.

Holotype. —USNM, No. S5200 collected by Wm. T.
Watkins.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

ULOCRINUS CONVEXUS (Strimple), 1939a
Plate 52, figures 1, 2, Plate 55, figures 1, 2, 7
See Bassler and Moodey, 1943, p. 718.

Presently considered material consists of two partial
crowns, which are compressed laterally but provide all es-
sential characters, and one infrabasal circlet. The cup is of
moderate height, broad, and has a low, bowl-shaped IBB
disk. There is a mild concavity at the central part of the
IBB disk surrounding the columnar attachment area but
the balance of the IBB are upflared. Decidedly depressed,
arcuate areas are formed below the articular facets of the
RR which cause a somewhat constricted summit of the cup.

There are five IBB which extend well beyond the col-
umnar attachment and curve upward to form a low bowl
which is sharply differentiated from the circlet of basals
above. Five basals are long broad plates, exhibiting a mild
tumidity in the smaller specimen but with only gentle
curvature, which follows the contour of the cup, in the
mature hypotype. The radials are somewhat wider than
long, are mildly tumid in the young paratype but have lost
the characteristic in maturity. There is an arcuate area just
below the articulating facet which is more pronounced in
the small specimen than in the large hypotype. The outer
ligamental area is compact but has all the component parts,
i.e. a well-defined outer ridge, a narrow furrow, and a pit.

The transverse ridge is prominent, with the outer side al-
most vertical, and is strongly marked by denticles. There
are deep oblique furrows, small muscle areas, and a large
intermuscular notch. The facets slope outward. In some
interrays a corner of the radial plate curves into the ad-
sutural area but does not extend for any appreciable dis-
tance. There are two large anal plates, anal X resting evenly
on the posterior basal and RA obliquely to the right sup-
ported by right posterior and posterior basals and bordered
to the left by anal X and to the right above by the right
posterior radial. The anal plate is long and has facets to sup-
port two plates above.

There are ten arms branching one time on low, broad
axillary first primibrachs. The arms of the large hypotype
were found in close association with the dorsal cup which
had broken apart but has been restored. There is no ques-
tion but that they belong to the same specimen, The outer
surface of the arms is well rounded, they are decidedly
biserial, pinnular bearing and attain a great length.

The columnar scar on the IBB circlet rests in a shallow
depression, is marked by about 44 crenulations and _ is
pierced by a large penta-lobate lumen. The crenulations
do not closely approach the lumen and leave a smooth
pentagonal-shaped area surrounding the opening.

The large hypotype has a cup height of 36.0 mm, maxi-
mum width of 38.0 mm. The arms as preserved are 98.5
mm long and are not close to terminating, 7.e. there is no
sign of tapering at the distal ends as would be found if the
termination was being approached.

Remarks. — The long prominent anal X of this species
is apparently diagnostic. It rests evenly on the short upper
facet of the posterior basal and projects above the summit
of the cup The radianal is usually long and slender appear-
ing in young specimens, but the width increases more than
the length, and it often has a broad appearance in maturity.
The plates are convex in youth, having a tumid appearance
that is lost with maturity. Cup plates are all relatively thin.
The depressed area of the IBB circlet surrounding the col-
umnar scar is not so pronounced in maturity as in young
specimens.

It seems likely the specimen from the Wann Forma-
tion, ascribed to Ulocrinus buttsi by Wright and Strimple
(1945), is a variant of U. convexus. The radianal has lost
contact with the right posterior basal which is a condition
found in U. buttsi but in other respects it appears con-
specific with U. convexus. The arm structure of the crown
from the Wann Formation was not correctly shown by
Wright and Strimple. It has cuneiform to interlocking seg-
ments,

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 167

The young crown from Lake Bridgeport is similar to
U. extrorsus from the older Millsap Lake Formation.

Types. — Holotype and paratype are in the Springer
Collection, U.S. National Museum, The three hypotypes
are USNM, No. $5177, and S5195 collected by Mr. Wm. T.
Watkins and USNM, No. $5191, collected by Mr. Louis
Todd.

Occurrence.— Holotype and paratype are from the
Mound in Osage County, just west of the city limits of
Bartlesville, Washington County, Oklahoma; Wann Forma-
tion, Ochelata Group, Missourian. Three hypotypes are from
four miles west of Bridgeport, Wise County, Texas; Wolf
Mountain Shale, Graford Formation, Canyon Group, Mis-
sourian, Pennsylvanian.

ULOCRINUS ZESCHI Strimple and Watkins, new species
Plate 37, figures 4-7; Plate 38, figures 5-7

The cup expands uniformly from the columnar attach-
ment to slightly below or about mid-height of the cup,
thereafter the lateral sides rise evenly with no further ex-
pansion of the cup. A break in the contour takes place at
the meeting of the basal and infrabasal circlets. There are
five large infrabasals readily visible in side view of the
cup. A depressed area is present for the reception of the
proximal columnal. Five basals are large prominent elements
with a tumid appearance. Five large radials are only slightly
wider than long and those adjoining the broad posterior
interradius lose some width to the anal plates. The right
posterior radial has a peculiar shape because of the im-
position of the large radianal plate. There is a slight outer
slope beyond the outer ligamental area but it is not pro-
nounced. The outer ligamental furrow is well defined and
the outer ligamental pit is broad. The inner ligamental area
is broad and subhorizontal in attitude. There are two large
muscle scars and a small innermuscular notch on each facet.
The transverse ridge is prominent and oblique lateral fur-
rows are present. In the type specimens there are three well-
defined anal plates in the posterior interradius, the dom-
inant one being the radianal. Anal X is in broad contact
with the posterior basal in both specimens. In both smaller
specimens (paratypes) the RX has lost contact with the
radianal but still extends below the summit of the cup.
The RX is narrow in the holotype but is in broad contact
with the radianal.

The columnar scar is round and marked by fine ridges.
The arms are unknown.

Measurements of the holotype in millimeters:

Width of cup (maximum) 25.7

Width of cup, posterior to anterior (minimum) 23.4
Height of cup 723

Ratio of height to width 0.67
Width of infrabasal circlet 12.3
Width of columnar scar 3.2
Length of basal (right anterior) 12.0*
Width of basal (right anterior) 14.0*
Length of suture between basals 7.0
Length of radial to transverse ridge (right anterior) 12.0*
Width of radial (right anterior) 15.2*
Length of suture between radials 5.7
Length of anal X Ta
Width of anal X 4.8
Length of RA 8.8
Width of RA 7.0

* Measurements taken along surface curvature.

Remarks. — The logical progenitor of Ulocrinus zeschi
among known Morrowan forms is the species Cromyocrinus
grandis Mather, which has a broader cup and proportion-
ately larger anal X and RX. The RX is narrowed and is
migrating out of the cup of Ulocrinus xzeschi. The type
species of Cromyocrinus, C. simplex, has less erect infra-
basals than Ulocrinus zeschi and has diverged from the
lineage leading to Ulocrinus in having reduced the number
of arms to five. Several species of Ulocrinus are known to
have ten arms and all later species of Ulocrinus have only
two anal plates in the posterior interradius of the cup, of
which RA is dominant.

Types. — Holotype, USNM, No. $5119, collected by
Wm. T. Watkins, paratype, USNM, No. $5121, collected by
Wm. T. Watkins, reposited in the U.S. National Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch, Mason,
Mason County, Texas.

Genus GOLEOCRINUS Strimple and Watkins, new genus

Type species. — Golzocrinus masonensis Strimple and
Watkins, n. sp.

Range. — Atokan and Desmoinesian; North America.

Diagnosis. — Dorsal cup medium bowl-shaped with
broad base and a basal concavity. Five infrabasals are down-
flared but the distal tips are flexed outward into a subhori-
zontal attitude. Five large basals form the basal plane and
flex sharply upward to form much of the lateral walls of
the cup. The basals and radials are tumid. Five radials
are slightly wider than high and have a flattened, subhori-
zontal face to the fore of the normal articulating area.
Three anal plates may be in normal (Primitive) arrange-
ment but the RA is large and is dominant and may take
an advanced position. Surface of the plates is essentially
smooth, Cicatrix of the proximal columnal is circular. Arms
are unknown.

Species assigned to Goleocrinus:

Atokan: Texas
Atokan: Texas

Goleocrinus masonensis, n. sp.

Goleocrinus confossus, 0. sp.

Goleocrinus ardmorensis (Strimple), new combination
Desmoinesian: Oklahoma

168 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Remarks. —Goleocrinus appears to belong to the
cromyocrinids rather than the ethelocrinids. It is advanced
for the former group in that a basal invagination is pres-
ent; however, it is to be noted that the infrabasals flex
sharply out of a concavity before meeting the basal plates.
It is our belief that the pronounced basal invagination
found in this group started as a sharp impression for the
columnar attachment in a flattened infrabasal circlet and
thereafter the infrabasals developed inwardly to produce
the concavity. In the genus Ethelocrinus the basal plates
are curved sharply into the basal concavity and there is a
pronounced surface ornamentation consisting of pustules
and granules. Mooreocrinus appears to be closely related
but differs in lacking a basal concavity.

GOLEOCRINUS MASONENSIS

Strimple and Watkins, new species
Plate 39, figures 1-4

Dorsal cup of medium height, bowl-shaped, with a
broad base and a restricted basal concavity. The basal con-
cavity is occupied by all except the distal edges of the
infrabasal plates. The lateral sides of the cup are gently
curved and do not attain a vertical position until just
before the flattened shelf to the fore of the articular shelves
is reached. No constriction of the cup takes place as the
summit is approached. When viewed from above or below,
the cup has a circular outline except for the flattened pos-
terior interradius. There is no pronounced protrusion of the
area occupied by the radianal.

Five downflared infrabasals flex sharply into the basal
plane with those of the right posterior, left posterior and
left anterior extending further out of the concavity than
the other two plates. Five large basals form the basal plane
and an almost imperceptible basal depression. They flex
sharply upward to form much of the cup walls and are
mildly tumid. Five radials are slightly tumid and have dis-
tinctive upper surfaces in that a sharp edge is formed at
the summit of the cup and a flattened area is developed to
the fore of the subhorizontal articulating facet. A well-de-
veloped outer ligamental pit is present in the fore part of
the transverse ridge. Long lateral furrows are present and
extend to the intermuscular furrow. Muscle areas are not
large and are more or less triangular-shaped.

Three anal plates are in normal (Primitive) arrange-
ment. RA is the largest and has five facets, meeting the
right posterior basal to the right below, posterior basal to
the left below, anal X to the left above, RX directly above,
and the right posterior radial to the right above. Anal X
extends above the cup summit and is in broad contact with

the posterior basal below. RX is a small plate in broad con-
tact with RA below, short contact with the right posterior
radial to the right and anal X to the left. It extends a short
distance above the distal end of anal X.

The surface of the dorsal cup is devoid of ornamenta-
tion and sutures are impressed. An occasional small pit
marks the sutures between cup plates.

Measurements of holotype in millimeters:

Width of cup (maximum) 21.7
Width of cup, posterior to anterior (minimum) 20.0
Height of cup 10.7
Ratio of height to width 0.49
Width of infrabasal circlet 6.0
Length of basal (right anterior) 11.0*
Width of basal (right anterior) 10.1*
Length of suture between basals Sve
Length of radial, to transverse ridge (right anterior) 10.8*
Width of radial (right anterior) 12.6*
Length of suture between radials 4.5
Length of radianal 8.6*
Width of radianal 5.0
Width of columnar scar 3.5

* Measurements taken along surface curvature.

Remarks. — Goleocrinus masonensis is more closely re-
lated to G. ardmorensis than to any other described species.
The former has a more primitive arrangement of anal
plates in the posterior interradius of the cup and exhibits
some asymmetry in the infrabasal circlet. Goleocrinus con-
fossus has a lower cup, with a constriction of the cup near
the summit, and has an advanced arrangement of the anal
plates.

Holotype. —USNM, No. $5122, collected by Wilburn
Shearer, reposited in the U.S. National Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch, Mason,
Mason County, Texas.

GOLEOCRINUS CONFOSSUS
Strimple and Watkins, new species
Plate 38, figures 1-4

Dorsal cup low, bowl-shaped with constriction of the
cup near its summit, and a basal concavity. The cup is
asymmetric when viewed from below in that the right an-
terior side is more erect and the plates of the left posterior
appear to be more evenly projected. The right side of the
cup is actually higher than the left side. In the infrabasal
circlet those of the left are smaller than those on the right.

Five infrabasals are downflared but the distal tips flex
sharply out of the basal concavity, especially the left pos-
terior, right posterior and right anterior. Five large basals
form the basal plane and an almost imperceptible basal

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 169

depression, then flex sharply to participate in the lateral
walls of the cup. They are tumid with well-impressed su-
tures marked by a series of small pits. The posterior basal
is broadly truncated for the reception of the obliquely placed
radianal. Five radials are tumid with well-impressed sutures
marked by series of small pits. The pits along the sutures
provided the species name confossus = full of holes. A sharp
flexing near the summit constricts the cup and terminates
with a well-defined outer marginal ridge. A shallow outer
ligamental furrow is present as well as an outer ligamental
pit. The transverse ridge is marked by fine denticles and
shallow lateral furrows are present. There appears to be two
pits adjacent to the intermuscular furrow and the muscle
areas are shallowly impressed. The facets are subhorizontal
in attitude. Within the posterior interradius, the radianal
has become completely dominant with anal X and RX ap-
parently pushed entirely out of the cup. The surface of the
cup is devoid of ornamentation and the stem is round.
Measurements of the holotype in millimeters:

Height of cup, left side 9.8
Height of cup, right side 10.0
Width of cup (maximum) 21.2
Width of cup, posterior to anterior (minimum) 20.8
Width of infrabasal circlet 6.6
Length of basal (right anterior) 10.7*
Width of basal (right anterior) 10.3*
Length of suture between basals See
Length of radial, to transverse ridge (right anterior) 10.5*
Width of radial (right anterior) 17s
Length of suture between radials 3.9
Width of radianal 5.0
Length of radianal 4.2
Width of columnar scar 3.0

* Measurements taken along surface curvature.

Remarks — Goleocrinus confossus has a lower cup than
the other two species assigned to the genus, G. masonensis
and G. ardmorensis. The reduction of anal plates to one
large radianal in G. confossus is highly advanced and in-
dicates that its progenitor is older than G. masonensis
which occurs in the same rocks but has three anal plates
in normal (Primitive) arrangement.

Holotype. —USNM, No. $5120, collected by Wm. T.
Watkins reposited in the U.S. National Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch, Mason,
Mason County, Texas.

GENUS AND SPECIES INDETERMINATE
Plate 32, figure 6

The dorsal cup of this specimen is partially destroyed
in the posterior area where there is, however, a long radianal
such as typically found in Dicromyocrinus or Parulocrinus,
but other anal elements are missing. The dorsal cup is bowl-

shaped and infrabasals are not visible in side view of the
cup. There are ten arms branching on the first primibrach
in all rays, uniserial in lower portion but biserial for most
of their length. The overall length of the crown as preserved
is 46 mm the width of the cup is 13 mm.
Material. — Crown collected by Wm. T. Watkins, re-
posited in the United States National Museum, No. $5104.
Occurrence. — Marble Falls Formation, Morrowan?,
Pennsylvanian; bank of Espey Creek about 3.5 miles south-
west of Lampasas, Lampasas County, Texas.

Family SELLARDSICRINIDAE Strimple and Watkins
new family

Genus. — Sellardsicrinus Moore and Plummer, 1940,

Range. — Desmoinesian, Pennsylvanian; Mid-continent
region, North America.

Diagnosis. — Crown elongate; dorsal cup low, truncate
bowl-shaped with slight basal concavity; five IBB, five
BB, five RR, three XX in cup (Primitive Type) and eight
to nine XX in entire posterior interradius; surface of cup
plates smooth, ends of radials (other than upper articular
facet) and basals are excavated for ligaments; arms are
slender, biserial, pinnular bearing, rounded externally, do
not abut closely when closed, first bifurcation with low
PBry, a second isotomous branching may be with SBr3-;9ax
and a third isotomous branching may occur well above the
second,

Remarks. — There are not too many inadunate genera
in the Pennsylvanian having biserial arms. The Erisocrini-
dae usually have biserial arms but they are restricted to ten
and there is usually only one anal plate in the cup (or
none). The Ethelocrinidae usually have more than ten bi-
serial arms but they only bifurcate in the most proximal
regions. It is possible for some species with three anal plates
for which the arms are not known to be mistakenly as-
signed to the ethelocrinids but belong to the sellardsicrinids.
Some cromyocrinids have as many as ten biserial arms but
none exceed that number. The Eupachycrinidae have more
than ten biserial arms but all bifurcations are restricted
to the proximal section of the arms. Some forms among the
Pirasocrinidae tend to develop biserial arms but the family
is not considered to be related. The arms of pirasocrinids
are closely abutting when closed, there is a terminating
anal platform at distal end of crown, and the base of the
cup is usually deeply concave. The arms of sellardsicrinids
do not abut, there is no terminating platform of the anal
tube, and the base of the cup is mildly concave.

We believe Scotiacrinus Wright (1945) of Viséan age
could well be in the lineage leading to Sellardsicrinus. The

170 PALAEONTOGRAPHICA AMERICANA (VI, 40)

dorsal cup is slightly higher, the cup plates more tumid
and arms essentially uniserial all which would be expected
in an ancestral form. The rounded contour of the cup when
viewed from above and below, and shallow basal concavity
with flattened IBB circlet, is much alike for the two genera.

Genus SELLARDSICRINUS Moore and Plummer, 1940

Type species. —Sellardiscrinus marrsae Moore and
Plummer, 1940.

Range. — Desmoinesian, Pennsylvanian; Texas.

This genus has only been known from the type species
but is distinct from other known forms. It was placed in
the family Eupachycrinidae by Moore and Laudon, 1943,
but it is only superficially comparable to members of that
family, The structure of the arms is somewhat comparable
to that of Stellarocrinus, or at least is closer than to other
described forms. The arms of Stellarocrinus are usually con-
siderably wider than found in Sellardsicrinus and in most
species of the former genus the axillary brachial above the
first dichotomy is a small triangular piece surrounded by
secundibrachs and tertibrachs. The primitive arrangement
of the anal plates of the posterior interradius and the gen-
eral structure of the cup of Sellardsicrinus serves to readily
distinguish it from Stellarocrinus, and anal plates in Stellaro-
crinus are advanced (Extreme Type). If the two are re-
lated the common ancestor must be remote.

As noted under familial remarks, we consider Scotia-
crinus Wright from the Viséan of Scotand as the most likely
ancestral form to Sellardsicrinus, among known genera.

A new species is described as Sellardsicrinus fortwnatus
Strimple and Watkins. It is a large form but we are fortun-
ate in having a young representative and so establish the
characters as essentially the same for large and small speci-
mens.

A large crown and a small partial crown of S. marrsae
are available for study. Some remarks concerning the species
are made.

SELLARDSICRINUS MARRSAE Moore and Plummer, 1940
Plate 45, figure 4
See Bassler and Moodey, 1943, p. 679.

The species is well described and illustrated by Moore
and Plummer. Some measurements are given of the two ex-
amined hypotypes which are larger than the holotype. A
new observation is the small spinelike nodes found on some
tertibrachs of the same nature as those which accentuate
the zigzag appearance of the arms in many decadocrinids.
Moore and Plummer (1940) noted the spinelike nature of
the upper axillaries.

Attention is directed to the shape of the radial plates

which also affects the shape of the dorsal cup, 7.e. the
tumidity and outward curvature of the radials but with a
sharp inward curvature near the edge of the articular facet.
As noted by Moore and Plummer, a view of the edge of
radials and basals or basals shows the plates to be thick.
In a hypotype it is also seen that the center of these facets
is strongly depressed, as for the reception of a ligament.
No denticulation or specialized ridges are observed. The low
axillary primibrachs are also tumid but do not form a spine.
Associated Sellardsicrinus fortunatus does not have tumidity
or inward curvature of the radials and the axillary primi-
brachs are not tumid.
Measurements in millimeters:

Hypotype
Length of crown 95.0
Maximum width of crown 46.04
Height of cup 14.5d
Width of cup 30.0e
Length of basal plate 10.6
Width of basal plate 10.6
Length of radial plate (to outer ligament area) 8.6*
Width of radial plate 16.0*

* Measurements taken along surface curvature.
d Distorted by lateral compression,
e Estimated natural width.

Types. — Holotype, Plummer Collection, No. 12,372,
Bureau of Economic Geology, University of Texas, collect-
ed by Ralph King. Paratypes, M-7, M-23, and M-34, in the
collection of Mrs. W. R. Marrs, Austin, Texas; paratypes,
H-7 and H-24, in the Harris Collection, collected by Mrs.
G. W. Harris, Waco, Texas; metatype M-22, Marrs Collec-
tion; hypotypes, USNM, No. $5157, collected by Mr. and
Mrs. H. L. Strimple.

Occurrence.— Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

SELLARDSICRINUS FORTUNATUS
Strimple and Watkins, new species
Plate 35, figures 48

This species is based on three partial crowns all of
which are laterally compressed so that the nature of the
base of the cup is not known. Curvature of the few basal
plates preserved indicates the possibility of shallow basal
concavity as in Sellardsicrinus marrsae, However, we believe
that discovery of better preserved material will show S.
fortwnatus to be another genus, albeit closely related. Our
main reason for this thought is due to the nature of the
radial plates which are almost flat and have a sharp distal
edge in S. fortunatus but are tumid with a distinct outer

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 171

shelf to the fore of the distal edge in S. marrsae. Other dif-
ferences of a specific nature are given below under remarks.

One fairly young specimen and two mature specimens
are involved. The cup was probably of a moderate height,
truncate bowl-shaped. Basal plates are known from the
young paratype and are large with the width and length
about equal. Radial plates are slightly shorter than basal
plates but are wider than basal plates. As noted above the
outer surface of the radials have a flat, sharply defined
distal edge although there is a relatively broad, deep outer
ligamental furrow and a well-defined ligamental pit. The
transverse ridge is visible on two radials and is in the nature
of a thin, well-defined, straight line. One lateral and one
proximal edge of the right anterior radial is exposed in the
young paratype and the suture faces show a series of sinu-
ous ridges. One edge of a basal plate shows similar ridges
and pits but is too poorly preserved for close observation.
The anal plates are distinctive although not too well pre-
served or understood except in the young paratype. Three
robust plates in normal (Primitive Type) arrangement are
in the cup but in the next series above there are two plates
above anal X and one above RX. The plate on the right
shoulder of X also rests on the left shoulder of RX. In the
holotype there is yet another plate preserved in the anal
pyramid above RX, and on the right shoulder of the right
X, and in the young paratype another plate rests equally
above on the two X, plates. There is also another RX, on
the right shoulder of RX and a plate above the two RX,
plates

There are 40 biserial arms in the holotype that reduce
rapidly in width. The first primibrach is a low, wide axillary
plate. A second bifurcation takes place with the third to
eighth secundibrach and a third with fifteenth to twenty-
sixth tertibrach. The exterior of the arms are strongly
rounded and pinnular development is indicated. The upper
axillaries are small, low plates, not particularly tumid.

In the young paratype the few brachials preserved are
cuneiform rather than biserial. This is to be expected in a
juvenile specimen with the arms becoming biserial with age.

Measurements in millimeters:

Holotype Paratype Paratype
Maximum width of cup 52.0 46.0 24.0
(as preserved)
Length of basal — 16.2 7.3
Width of basal —_ 18.4 7.1
Length of radial 16.0 13.8 5.7
Width of radial 24.0 21.2 8.1

Remarks. — Aside from the lack of curvature of the
distal edge of radials in Sellardsicrinus fortunatus, the
broad posterior interradius with nine anal plates forming a

pyramid serves to distinguish the species from S. marrsae
which has a broad outer shelf to the fore of the ligamental
furrow and a narrow posterior interradius restricted to four
or five anal plates. The arms of S. fortwnatus branch for
the second time with the third to eighth SBr but in S.
marrsac bifurcation is with the ninth or eleventh SBr.
Strictly speaking all primibrachs are not biserial but they
have a fundamentally biserial nature.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Types. — Holotype USNM, No. $5114, paratype
USNM, No. $5113, collected by Wm, T. Watkins, and para-
type USNM unnumbered, collected by Mr. Louis Todd.

Family ETHELOCRINIDAE Strimple, 1961a
Genus PARETHELOCRINUS Strimple, 1961la
PARETHELOCRINUS WATKINSI (Strimple), 1949d
Plate 44, figures 4,5; Plate 48, figures 3,4; Plate 55, figure 4

This species is based on a monotype from the Arnold
Limestone Member, Deese Formation, Desmoinesian of the
Ardmore Basin of Oklahoma and is a crown in poor preser-
vation. The junior author has found several excellent hypo-
types from the Brannon Bridge Member, Millsap Lake For-
mation, Desmoinesian of Texas, which gives a much better
understanding of the species.

The holotype shows an apparent bifurcation in the up-
per region of the right posterior arm that must be adven-
titious because it is characteristic of the genus for bifurca-
tions to occur only in proximal regions, 7.e. with the first
primibrachs in all rays and first secundibrachs in some half-
rays. This is verified by a well-preserved crown (hypotype)
from Texas. The mode of branching was not determined for
the left arm of the holotype but is seen to be with the first
primibrach and again with first secundibrach, in the right
arm only, of the hypotype. A total of 12 arms with rather
flattened exteriors are formed, the other “extra” arm being
in the left half-ray of the right posterior.

In the cup the RA of the holotype appears to have
been a short plate but this probably was distorted in
preservation because in all hypotypes the RA is longer
than wide. Anal X is long and in narrow contact with the
posterior basal in all hypotypes, as in the holotype. The
base of the cup is partially preserved in a small hypotype
and provides a ratio of IBB circlet to cup width of 0.37.
This is somewhat greater than the ratio in Parethelocrinus
ellipticus of 0.26. The infrabasals project well beyond the

172 PALAEONTOGRAPHICA AMERICANA (VI, 40)

round columnar scar and are subhorizontal or impercept-
ably downflared. There is a shallow basal concavity.

A small hypotype (USNM, No. $5165) is in excellent
condition and is not appreciably distorted in preservation.
There is no decided difference in the linear width of radial
plates except for the right anterior radial which is slightly
wider than the others.

A large hypotype (USNM, No. $5149) discloses in-
equality in the linear width of radial plates in that those
of the right and left posterior are narrower than the other
three, Another hypotype (USNM, No. $5193) has a nar-
row right anterior radial and the right posterior radial
might have been slightly narrower than normal. It does
not appear that any reliance can be placed on this charac-
ter in this species.

Measurements in millimeters:

Hypotype Small Hypotype
USNM, No. $5149 USNM, No. $5193
Height of crown 62.0 —
Greatest width of crown, about 25.5
Height of dorsal cup, about 11.0 6.7
Greatest width of cup, about 21.0 14.0
Length of basal 1351 ihe
Width of basal 12.8* Tae
Length of radial 8.1* 5.2"
Width of radial 14.4* 8.0*

* Measurements taken along surface curvature.

Remarks. — There are several outstanding differences
between Parethelocrinus watkinsi and P. millsapensts
(Moore and Plummer) although they both occur in the
same horizon and locality and apparently belong to the
same genus. P. millsapensis is based on a monotype, and
we have not observed any additional representatives. The
specimen is crushed so that the nature of the base and the
infrabasals is obscured although there is little doubt that
at least a shallow basal concavity existed. The cup plates
are strongly ornamented by granules, the basal plates are
wider than long, the radial plates are twice as wide as long
and lack longitudinal curvature. In P. watkinsi the cup
plates are essentially unornamented, the basal plates are
slightly longer than wide, the radial plates are less than
twice as wide as long and have a longitudinal curvature.
One metatype of P. watkinsi is only slightly larger than
the type of P. millsapensis so that differences are not due
to growth differentials because of age (size). Parethelo-
crinus variabilis (Strimple, 1949d) is also of Desmoinesian
age but is a highly ornate form exhibiting considerable
tumidity of basals and radials. P. ellipticus Strimple (1961a),
the type of the genus, is of the late Desmoinesian (Holden-
ville Formation) and in common with Missourian species
of the genus has more arms than the other Desmoinesian

species. It is a smooth form with projection of the cup in
the area surrounding the radianal and has an exceptionally
small infrabasal circlet.

Types. — Holotype, deposited in U.S National Mu-
seum, collected by Wm. T. Watkins. Hypotypes, USNM,
Nos. $5149, $5165, $5193, collected by Wm. T. Watkins.

Occurrence. — Holotype, Arnold Limestone Member,
Deese Formation, Desmoinesian, west side of Lake Murray
near Water Tower, south of Ardmore, Love County, Okla-
homa; hypotypes, Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
three miles southwest of Brock, Parker County, Texas.

PARETHELOCRINUS PLATTSBURGENSIS (Strimple), 1938
Plate 49, figures 1-3

The presently considered specimen is somewhat com-
pressed in preservation but almost certainly had a low
bowl-shape and was of medium height. The infrabasal disk
is relatively small and is subhorizontally directed. The
proximal edges of the basals are slightly flexed indicating
a slight basal depression might have existed, 1.2. the plane
of the disk might have been slightly above the basal plane
of the cup. The basal plates are large. Radial plates are
large and wide. Articular facets are wide, subhorizontal with
shallow muscular fossae, Anal plates within the cup are two,
anal X in contact with the posterior basal and the large
RA resting obliquely to the right of it. Anal X is followed
by two plates above, one of which is preserved. The dorsal
cup has a maximum width of 31 mm as preserved but is
distorted by compression.

Remarks.— The arms of this specimen are not pre-
served but the specimen is sufficiently similar to Parethelo-
crinus plattsburgensis to be considered as conspecific.

Hypotype. — USNM, No. $5167 collected by Wm. T.
Watkins, reposited in the U.S. National Museum.

Occurrence. — Shale below Rough Mountain Conglom-
erate, Brownwood Shale, Canyon Series (Missourian), 3.5
miles east and 2.5 miles north of Rochelle, McCulloch
County, Texas. Plummer locality 153-T-23.

Genus AGLAOCRINUS Strimple, 1961la

AGLAOCRINUS MAGNUS (Strimple), 1949d
Plate 32, figures 3-5

The specimen at hand, which is from the Atokan of
Texas, is so much like specimens of Aglaocrinus magnus
from the Desmoinesian of southern Oklahoma that assign-
ment is made to the species with little hesitation. The
specimen originally described as Ethelocrinus periodus
Strimple (1949d, p. 13) is more comparable. It was placed

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 173

in synonymy to Aglaocrinus magnus (Strimple) by Strimple
(196la, p. 87), although it was thought to be from the
Frensley Limestone at that time and the holotype of A.
magnus is from the slightly younger Pumpkin Creek Forma-
tion. There is no question but what a close relationship
exists between these two forms, but there is some question
of the stratigraphic and geographic data applied to the
specimen of “Ethelocrinus periodus.” It is hoped that future
studies will resolve this question.

Crowns from the Holdenville Formation, near the top
of the Desmoinesian of northeastern Oklahoma, were as-
signed to the species Ethelocrinus magnus Strimple, which
became the type species of Aglaocrinus Strimple, 1961a. The
specimens from the Holdenville Formation were not so or-
nate as those from southern Oklahoma (Frensley Ls.? and
Pumpkin Creek Ls.) but this was thought by Strimple
(1961a, p. 88) to be mainly due to weathering and abra-
sion. The specimen from the Big Saline Formation of
Texas is ornate and the V-shaped depressions between cup
plates are marked by a series of pits.

Types. —The holotype is reposited in the Springer
Collection, U.S. National Museum. Hypotypes studied by
Strimple, 1961a, are OU, Nos. 4044, 4123, 4124, reposited
in the palecntological collections of the University of Okla-
homa. Hypotype USNM No. $5103 is reposited in the
U.S. National Museum.

Occurrence. — The holotype is from the Pumpkin Creek
Limestone, Dornick Hills Group, Desmoinesian; abandoned
quarry near Tucker Tower Museum, Lake Murray State
Park, Love County, Oklahoma. Hypotypes studied by
Strimple (1961a) are from the Holdenville Formation, up-
per Desmoinesian; NE14 sec. 24, T. 15 N., R. 11 E. about
four miles northwest of Beggs, Okmulgee County, Okla-
homa. Hypotype studied herein, Soldiers Hole Member,
Big Saline Formation, Atokan; Kurt Zesch Ranch, Mason,
Mason County, Texas.

Family TEXACRINIDAE Strimple, 1961a
Genus ULRICHICRINUS Springer, 1926

Type species — Ulrichicrinus oklahoma Springer, 1926.

Range. — Mississippian and Pennsylvanian; North
America.

There are only three species ascribed to this genus at
present: U. coryphaeus (S. A. Miller, 1871), of Keokuk
age, U. chesterensis Strimple, (1949d), and U. oklahoma
Springer (1926) of Morrowan age. In all three species the
arms branch once with the first primibrach and in most
rays branch again with the first secundibrach. There is a
maximum of four arms to any one ray.

The “principal type” (holotype) of U. oklahoma as

illustrated by Springer (1926) reflected an incipient bi-
serial condition in mid-portion of the arms, but the arms
above and below were uniserial. The other figured specimen
is entirely uniserial. The brachials are unusually short,
wide elements. First branching is with the first primibrach
and another bifurcation may take place with a first secundi-

brach.

The presently considered new species U. ramosus ex-
hibits a condition that is somewhat difficult to understand.
The second branching does not take place with the first
secundibrach but rather with the second secundibrach. This
can only be interpreted as a regressive trend if our present
concept of the evolution of arms is correct. That is, with
progressive evolution, the branching in upper portions of
the arms gradually moves down the arm. A form with sec-
ond bifurcation on the eighth SBr would theoretically evolve
to a form with the second bifurcation on the seventh,
sixth, fifth, and so forth, to a form with the second branch-
ing on the first secundibrach. Because the oldest known
species has already reached the point of a second bifurcation
with the first secundibrach there is hardly any other way
to account for the presence of a nonaxillary first secundi-
brach followed by an axillary except through a reversal of
the process. There has been a substantial amount of evi-
dence showing regressive evolution in the shape of the dor-
sal cup in other phyletic lines.

The addition of arms appears to be a progressive
trend among certain phyletic lines. Examples are readily
afforded by Alcimocrinus wherein A. girtyi Springer (1926)
of the Morrowan has considerably more arms than A.
ornatus Strimple (1949d) of the Chesterian. Another ex-
ample is in Polusocrinus, wherein P. ochelataensis Strimple
has more arms than the slightly older P. avanti Strimple.
Ulrichicrinus oklahoma has more arms than U. chesterensis
and the new species, U. ramosus, has as many or more arms
as U. oklahoma. In the large paratype of U. ramosus there
is a third bifurcation with the fourth tertibrach in an
outer arm of the anterior rami.

The only other Pennsylvanian form known to us hay-
ing thin, uniserial brachials, comparable to those of Ulrichi-
crinus, 1s Texacrinus Moore and Plummer (1940), That
genus has a slightly more advanced arrangement of the
anal plates and a more advanced shape to the dorsal cup,
but it has exotomous branching and the “extra” bifurcation
in Ulrichicrinus is of an exotomous nature. Although rela-
tionship may be through a common ancestor, rather than
direct, we believe Ulrichicrinus and Texacrinus may be of
the same phyletic lineage and have placed the genus under
the family Texacrinidae Strimple (1961a).

174 PALAEONTOGRAPHICA AMERICANA (VI, 40)

ULRICHICRINUS RAMOSUS Strimple and Watkins, new species
Plate 31, figures 1, 7; Plate 55, figures 5, 6

This species is based on two long crowns in good preser-
vation. One is relatively small (not immature) and is com-
plete except for the infrabasal plates which are missing. The
specimens are slightly distorted by lateral compression
which is a standard condition for the genus.

Structure of the dorsal cup is essentially conical, and
it is almost certain the infrabasals were at least mildly up-
flared. There are five large basal plates, hexagonal-shaped
except where affected by anal plates. There is no curvature
of proximal ends of basals such as would be present if a
concave base existed. Five large radials are sharply differ-
entiated from the large axillary primibrachs, and there is
a mildly gaped suture but no longitudinal curvature of the
radials. The outer edge of the outer ligamental area is
sharply crenulated. There are six anal plates in the posterior
interradius of which three (X, RA and RX) are in the cup
in normal (Primitive Type) arrangement. Axillary first
primibrachs are almost as large, and have the same general
shape though opposed, as the radial plates. All arms
branch with the first primibrach and a second bifurcation
takes place in all arms with the second secundibrach. In
the large paratype one other branching is observed on the
fourth tertibrach in an outer arm of the anterior ray. No
such branching occurs in the holotype wherein tertibrachs
become low so that the stout, long pinnules are packed
close together even though each brachial only carries a
single pinnule and they are thus alternated. The exterior
of the arms is convex but not strongly rounded. The arms
are capable of closing tightly.

Measurements in millimeters:

Holotype Paratype

Length of crown 62.0 82.01
Width of crown - maximum 21.0 50.0
Height of cup 8.0 —

Width of cup 11.08 —_

Length of basal 4.9 —

Width of basal 4.8 12.1
Length of radial 4.0 7.5
Width of radial 6.5 12.3
Length of PBr,ax 4.0 6.6
Width of PBr,ax 6.3 12.3

i Incomplete.
© Estimated.

Remarks. — Texacrinus gracilis Moore and Plummer,
1940, is from the same type locality as Ulrichicrinus ra-
mosus and has a comparable general appearance, partially
due to the similarity of flattening. The former has anal
plates in advanced arrangement (RA in dominant anal posi-
tion) and the arms bifurcate a second time well above the

first branching and at least once again in most inner
arms of each half-ray. Other species assigned to Ulrichicrinus
branch with the first secundibrach whenever a second bifur-
cation occurs, and no other species branches a second time
in all half-rays.

Types. — Holotype, USNM, No. $5194 and paratype,
USNM, No. $5098. Collected by Wm. T. Watkins, reposited
in the U.S. National Museum.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Genus TEXACRINUS Moore and Plummer, 1940

Type species. —Texacrinus gracilis Moore and Plum-
mer, 1940.

Range. — Pennsylvanian; Midcontinent region, North
America.

Remarks.— The type species is from the Brannon
Bridge Limestone Member, Millsap Lake Formation (Des-
moinesian ) near Brock, Texas, and is represented by a mono-
type. We do not have any additional specimens from Texas.
Strimple (1952 and 196la) described several additional
species of the genus from Desmoinesian and Missourian
rocks of Oklahoma.

Family ANOBASICRINIDAE Strimple, 1961a
Genus ANOBASICRINUS Strimple, 1961a
ANOBASICRINUS PRAECURSOR (Moore and Plummer, 1940)
Plate 43, figure 13
See Bassler and Moodey, 1943, p. 572.

Synonymy. — Neozeacrinus praecursor Moore and
Plummer, 1940, Anobasicrinus praecursor Strimple, 1962a.

In the presently considered collections the species is
represented by a crown in good preservation and two speci-
mens show the bulbous nature of the anal sac. The
species was referred to Anobasicrinus by Strimple (1961a)
because of fundamental differences in the structure of the
arms which in Neozeacrinus are closely abutting when
closed, with distinctive interlocking features, but in Ano-
basicrinus do not abut after the second dichotomy. It was
assumed from the structure of the crown of A. praecursor
that a balloon-shaped anal sac was present and specimens
collected by the junior author thoroughly establish the
feature,

Small distal arms are preserved along with the bulbous
anal sacs and show the uniserial upper brachials to be
elongate and to have a medium-ridge terminating with a

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 17

small spine. Many of the brachials have two pinnules on a
side (hyperpinnulated) which feature is shared with Exo-
crinus Strimple (1949b). Exocrinus also has a median-
ridge on the arms so that the possibility of relationship
becomes apparent.

Types. — Holotype, Harris Collection, No. H-25; para-
types, Nos. H-9, H-13, H-22 and H-25, collected by Mrs.
G. W. Harris. Paratypes, Marrs Collection, Nos. M-18, M-21
and M-24, collected by Mrs. W. R. Marrs. Two additional
specimens, M-3 and M-8, collected by Mrs. Marrs, were re-
ferred to the species. Topotype, No. 75, collected by Mrs.
J. H. Renfroe; and topotype, USNM, No. S5146, collected
by Wm. T. Watkins.

Occurrence.— Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Family STELLAROCRINIDAE Strimple, 1961a

Genera. — Heliosocrinus Strimple, 195la, Stellaro-
crinus Strimple, 1940b.

Range. —Chesterian (Mississippian )—Pennsylvanian;
North America.

Discussion. — Heliosocrinus Strimple, 195la, is con-
sidered by us to be ancestral to the Pennsylvanian genus
Stellarocrinus Strimple, 1940b. The older genus, Helioso-
crinus, has three anal plates in normal (Primitive Type)
arrangement, branches on the second primibrach, and has
cuneiform arm segments. Stellarocrinus has biserial arms
(exception — Stellarocrinus distinctus Strimple, which has
cuneiform arm segments) and first bifurcation is with the
first primibrach. Stellarocrinus has an advanced arrange-
ment of anal plates (Extreme Type).

The arms of these forms are so constructed that they
are never able to abut against one another, even when
closed. There are usually wide gaps between the arms
which probably explains why several apparently protec-
tive specializations may arise:

1. Spines. (May project from the arms, the anal tube

and dorsal cup or the dorsal cup.)

2. Endeavor to fill the gaps:

a. Thick short pinnules which are projected at
right angles to the sides of the arms and may
interlock with pinnules of adjacent arms.

b. Extraordinary widening of arms in_ selected
areas.

3. Plates of anal tube become thick.

The apparent effectiveness of the widening of the arms

(2b, above) is lessened by the discovery of a row of pits

wn

along the outer lateral edge of an arm which were obvious-
ly caused by some boring organism. If the arms were closely
opposed at even moderately frequent intervals the ques-
tion arises as to how the parasite could have remained in
place long enough to bore five pits. Probably only the gaps
directly above the second axillaries are closely in this man-
ner. This arm structure is only well understood for a crown
of Stellarocrinus sp. cf. S. angulatus from the Missourian
of Texas which is figured herein.

Another specialization for Stellarocrinus recently dis-
covered is for the proximal plates of the anal tube to be
articulated with the posterior shoulders of the right and
left posterior radials, as well as with the two upper anal
plates of the dorsal cup. This is probably because no sup-
port is ever afforded the anal tube by the more or less rigid
arms.

Ligamental articulation has been found between bra-
chials of separate rami (of the same arm) just below and
above the point of bifurcation, especially in the upper arms
where the axillary is a small triangular piece entirely sur-
rounded by other brachials. This feature is apparently con-
fined to Stellarocrinus.

Genus STELLAROCRINUS Strimple, 1940b

Synonymy. — Brychiocrinus Moore and Plummer,
1940; Whiteocrinus Strimple, 1939b (not Jaekel); Apollo-
crinus Moore and Plummer, 1940.

An interesting form from the Millsap Lake Formation
is presented as S. bulbus, n. sp., wherein a curious develop-
ment of interlocking pinnules is observed. The arms of this
genus are so developed as to prevent close abutment and
thus the anal tube is exposed at all times. It is our belief
that the pinnular development was an effort to fill the gaps
left between arms as previously mentioned under the fami-
lial discussion, A form from the Graford Formation ascribed
to Stellarocrinus sp. cf. S. angulatus is also figured to show
another trend having the same effect as above, 1.2. to close
gaps between arms under alerted conditions. In it the arms
widen considerably, especially in their upper portions, after
the second bifurcation.

STELLAROCRINUS TEXANI (Strimple) 1951la
Plate 32, figures 1, 2

A review of characters which we consider to be pertin-
ent to the species is as follows:

Cup low, truncate bowl-shaped; moderately deep basal
concavity with steep sides; infrabasal sutures impressed;
small nodes mark each side of basal sutures at proximal
edges; ridges are present on radials but do not affect most
of the outer surface of the plates; mild granulation under
magnification; narrow posterior interradius,

176 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Biserial arms found by the junior author at the type
locality for the species show a strong tendency to develop
spines above the second isotomous bifurcation. The spines
are directed outward at about a 45° angle from the flattened
surface of the arms and are sporadic. A third branching is
found in the inner arms of each half-ray. Only this tri-
angular-shaped axillary is projected as a spine. A total of
30 arms is indicated. The lateral sides of the arms are at
right angles to the outer surface and are nearly flat. Pinnu-
lar facets are restricted to the inner edge along the broad
ambulacral groove but are not well preserved in the material
at hand. The lateral sides of the arms have the same appear-
ance and surface structure as the outside of the arms,
which confirms the thought that they are constantly ex-
posed except in the areas surrounding the axillary brach-
ials where there appears to be close sutures between adja-
cent arms.

Remarks.—Stellarocrinus texani is a widespread species
of the upper Missourian, most readily separable from re-
lated species by the weak markings or ridges at the proximal
edge of the tumid radials. Although the ridges are present
and are confluent with those of adjacent plates they do not
affect the bulk of the surface of the radial. S. distinctus
Strimple, which is also of Missourian age, has a higher cup,
sharper, and more distinctive ridges on the cup and the
sides of the basal cavity have a slope, rather than being al-
most vertical as found in S. texani. The species S. geometri-
cus is of Missourian age and has a shallow cup, the sides
of the shallow basal cavity are sloped and heavy ridges
mark the surface of all cup plates. S. angulatus from the
Missourian at Kansas City is devoid of granules, nodes or
ridges, and has a broad, shallow basal concavity.

Hypotype. — Figured hypotype, collected by Mr. Wm.
T. Watkins, deposited in the Springer Collection, U.S.
National Museum, No, $5102, Washington, D.C.

Occurrence. — Wolf Mountain Shale Member, Graford
Formation, Missourian, Pennsylvanian; near Lake Bridge-
port, Wise County, Texas.

STELLAROCRINUS sp. cf. S. ANGULATUS
(Miller and Gurley), 1893
Plate 49, figures 7,8; Plate 50, figures 1, 6
See Bassler and Moodey, 1943, p. 685.

We have not examined the holotype of Stellarocrinus
angulatus but the description and illustrations of the species
indicate that the present specimen, which is from the Gra-
ford Formation, is closely related but not conspecific. The
holotype is reportedly from the Kansas City Group (prob-

ably Lane Shale Formation) which is slightly older than the
Graford Formation, though both are of Missourian age.
The basal plates of S. angulatus are reported to be sharp,
pointed, like a pyramid, so that the points form the basal
plane. In the present specimen there are no sharp projec-
tions and in fact the basal plates are only mildly tumid.
The arms of S. angulatus appear to be broad but only some
of the lower extremities are preserved. A small, triangular-
shaped axillary plate is preserved and illustrated by Miller
and Gurley as the primibrach but it does not rest on the
radial and has the appearance of an axillary secundibrach to
us. If it is indeed a primibrach it is the narrowest known
to us for the genus. In the present specimen the axillary
primibrach does not fill the full width of the radial plate,
which is typical of the genus, but the secundibrachs do not
extend down on both sides to reach the radial plates. A
second isotomous bifurcation usually takes place a short
distance from the cup at about the SBrjoax, giving rise
to 20 arms. As many as 58 tertibrachs have been observed
with no further evidence of branching. A broad nature of
these biserial arms is most distinctive. Although the ex-
pansion of the arms must have been for the purpose of
closing some of the gaps left due to their inability to become
closely opposed, they did not develop flat sides but rather
the edges curve strongly inward so that the ambulacral
groove is narrow. Small pinnular facets are faintly preserved,
bordering the ambulacral groove.

One segment of arm that has been freed of the re-
mainder of the crown shows a series of five small pits along
the right (outer) lateral side. These pits are most certainly
formed by a parasitic or foreign form, and are comparable
to many others of like structure known to us, albeit they
are smaller than usual. The existence of these pits illus-
trates the presence of a foreign body in a position that
would have prevented the closing of the arms and abut-
ment or abutment against an adjacent arm, if the animal
was capable of such an action. Most associated inadunate
crinoids are capable of bringing their arms together in
close abutment but Stellarocrinus does not usually appear to
have this capability.

Crenellae and ligamental fossae are present on the
innermost lateral sides of brachials adjacent to and im-
mediately above the triangular axillary brachials as well
as on the normal articulating facets.

Remarks. — There are several pronounced differences
between Stellarocrinus sp. cf. S. angulatus and the associated
S. texani. S. texam has relatively prominent raised ridges
on the cup, especially about the sharply impressed basal
cavity, and many brachials are projected as spines. The

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 7/76

former has a smooth exterior, no projections, a shallow cup
with shallow basal concavity and the arms are exceptionally
broad. This same lack of ornate projections on cup or arms
serves to differentiate S. sp. cf. S. angulatus from other
Missourian species.

Occurrence. — Wolf Mountain Shale Member, Graford
Formation, Missourian, Pennsylvanian; near Lake Bridge-
port, Wise County, Texas.

Figured specimen. — Collected by Mr. Louis Todd,
USNM, No. $5169.

STELLAROCRINUS FLOREALIS (Moore and Plummer), 1940
See Bassler and Moodey, 1943, p. 685.

There are no specimens in the presently studied col-
lections assignable to Stellarocrinus florealis. The species is
well described by Moore and Plummer, but we are setting
forth some diagnostic features for comparison with other
material.

Cup low, truncate bowl-shaped with a small infra-
basal circlet rather deeply impressed. The basal concavity
is well marked by ridges that encompass the cavity and the
long even slope of the proximal portion of the basal plates
produces a flower-like appearance as the specific name
indicates. There is also a gentle transverse concavity of the
basal plates in their proximal region. The nodes, ridges,
and depressions on other parts of the cup are not pro-
nounced. Arms are biserial, 20 in number, having two
isotomous branchings. The second axillary is a small tri-
angular plate.

Remarks. — This species is closer to S texani than to
other described forms. The posterior interradius of S.
florealis is wider than that of S. texani, it is devoid of sur-
face granulation, and the arms have more surface curvature.
The associated S. texanus is devoid of any surface mark-
ings other than a ridge like angulation surrounding the basal
concavity.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Desmoinesian, Pennsylvanian;
about three miles southwest of Brock, Parker County,
Texas.

Type. — Holotype (a monotype) listed by Moore and
Plummer, 1940, as Marrs Collection, No. M-1, collected by
Mrs. W. R. Marrs, Austin, Texas.

STELLAROCRINUS TEXANUS (Moore and Plummer), 1940
Plate 41, figure 1; Plate 46, figure 2; Plate 47, figures 1, 4;
Plate 52, figure 13.

See Bassler and Moodey, 1943, p. 685.

This species was based on a monotype which was an
incomplete crown and served as the type species of Brychio-

crinus Moore and Plummer, 1940. The form has subse-
quently been recognized as being synonymous with Stellaro-
crinus. We are fortunate in having several crowns in ex-
cellent preservation in the currently considered collections.
Various stages of growth are involved. In order to study
the ratio of development between various ossicles several
measurements have been taken. The linear width and
length of plates are used in this particular instance because
we do not feel the intervening protrusions and ridges should
be considered in these particular comparisons.

Linear measurements in millimeters:

USNM
$5132 $5150
Height of dorsal cup (to 2.0 2.5

transverse ridge)

Width of dorsal cup 6.2 ons
Ratio of H/W 0.32 0.26
Length of basal 1.6 2.8
Width of basal 1.8 Ai
Ratio L/W 0.88 0.75
Length of radial Wee 2.5
Width of radial 2.8 5.0
Ratio of L/W 0.60 0.50
Length of PBr,ax 3.0 2.7
Width of PBriax ral 4.8
Ratio of L/W 1.45 0.56
Width of IBB circlet 2.0 3.3
Diameter of proximal 1.2 3.2

columnals

No firm conclusions can be drawn but in general the
measurements substantiate other observations:

1. The dorsal cup is proportionately higher in youth.
a. The height of the cup does not increase a great

deal with age.

2. The width of the basal and radial plates increases
more proportionately with age than the length.

3. The length of the axillary first primibrach does not
increase appreciably with age but the width is
greatly increased,

Some young specimens have biserial arms, but it is
almost certain that really immature specimens will be found
with cuneiform arm segments. There are two major bifur-
cations of the arms, the first with the PBr,ax and the
second on or about SBryjo-11;ax. There may be another di-
vision of inner half-rays. The axillary secundibrachs are
small, wedge-shaped, triangular pieces surrounded by secun-
dibrachs and tertibrachs. The full division of the ray is not
really accomplished for some distance above the axillary
although the two arms are distinct. The pieces surround-
ing the triangular axillaries are probably joined by ligamen-
tal tissue and crenellae as reported herein for Stellaro-
crinus sp. cf. S. angulatus although such cannot be stated
with certainty.

One of the most distinguishing characters of S. texanus

178 PALAEONTOGRAPHICA AMERICANA (VI, 40)

is the lack of projections or ornamentation except for the
ridgelike structure surrounding the basal concavity. The in-
frabasals are almost entirely covered by the relatively large
stem in mature specimens and the sides of the basal con-
cavity are formed by steeply sloped proximal ends of the
basals. The main anal plate (RA) is moderately large and
long so that the two plates above are almost entirely ex-
cluded from the cup.

The column is composed of series of five columnals,
starting and ending with large segments having curved
longitudinal profiles, adjacent small segments having flat-
tened profiles and the center segments slightly larger and
having some longitudinal curvature. All show the strong
interlocking crenellae along the suture lines. No cirri have
been observed.

Remarks.— None of the specimens of Stellarocrinus
texanus observed by us show even a trace of ridges, such
as found in the associated S. florealis, other than the ridge
structure surrounding the basal cavity which is much more
spectacular in the later species. One large hypotype does
have dimple-like depressions at the angles of the cup plates.
There is no evidence to suggest S. texanus could be the
young of the slightly larger S. florealis.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Types. —The holotype is reportedly in the Harris
Collection, No. H-4, collected by Mrs. G. W. Harris, Waco,
Texas; hypotypes Springer Collection, U.S. National Mu-
seum, USNM, No. $5132, $5150, collected by Mr. and
Mrs. H. L. Strimple; hypotypes Springer Collection, U.S.
National Museum; USNM, No. $5160, collected by Mr.
Wm. T. Watkins, San Antonio, Texas.

STELLAROCRINUS BULBUS Strimple and Watkins, new species
Plate 44, figures 6, 9-11; Plate 47, figures 5-7

This species is based on four partial crowns from the
Kickapoo Falls Limestone Member of the Millsap Lake
Formation. One minute crown provides interesting data as
to the nature of a truly immature form.

Large specimens have a low, broad, truncate bowl-
shaped dorsal cup with a broad, shallow basal concavity.
Five infrabasals form a pentagonal-shaped subhorizontal
disk almost covered by the large proximal columnals. Five
basals are prominent, each having an outwardly directed
spine in mid-section (posterior basal may have a pair),
with proximal portion of plate almost horizontal in position

and distal portion almost vertical in position. Anal plate
(RA) rests on the broad distal edge of posterior basal and
supports two plates above. In the large hypotype the plates
are absent but the facet to the right is slightly larger than
the one to the left. In the small hypotype both plates are
preserved and that to the right is slightly wider than the
one to the left. Both plates have four sides, are slightly
longer than wide and the upper surface is slanted from the
center to the outer lateral side. Anal plate does not extend
appreciably above the normal height of the cup. Radials are
wide, subhorizontally directed with slight convexity, or
curvature, Articular facets are outwardly directed and fail
to occupy the full width of the plates. In fact a substantial
amount of the outer surface of the radials in the adsutural
areas curve inward for a considerable distance. The facet
is somewhat short except in the posterior sides of the two
posterior plates. The intermuscular notch is broad and the
outer ligamental area is marked by fine linear crenellae
except in the sharply impressed ligamental pit. The areas
occupied by the proximal tips of the radials and the distal
tips of the basals are mildly depressed in the holotype and
are decidedly depressed in one paratype.

Comparison of various measurements and ratios of
length to width shows that basal plates appear to grow
faster than radial plates, both in their breadth and in their
length. There is also indication of proportionately greater
increase in the length of basals and conversely greater
increase in width of radials in fully mature forms. A strik-
ing reduction in the length of PBryax and increase in width
is shown between the immature paratype, with a ratio of
H/W of 2.11, as compared to 0.48 for the holotype. The
dorsal cup is proportionately much higher in immaturity.
There is a surprisingly small change in the relative width
of the IBB disk compared to the width of the cup between
youth and maturity.

The arms are biserial with well-rounded exteriors and
branch with the low, broad first primibrach in all rays.
There does not seem to be any further bifurcations. Al-
though the spines are often broken in preservation, many
brachials are produced as spines. Often every other brachial
will be spine bearing, then plain for five or six brachials,
and then the spines may appear on the opposite side of the
arm. Of considerable interest to us was the pinnular arrange-
ment. The pinnules are short, robust, and are at right angles
to the long axis of the arms. Because the arms are more or
less rigid the pinnules of one arm are interfingered with the
pinnules of the adjacent arm. This development was noted
under our discussion of the genus. We are also aware that
the arms extend outward from the cup to about mid-height

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 179

of the crown and then curve inward. There is even some
indication of a sharp incurving of the distal ends which
were probably more pliable than the majority of the arms.

The column is known in a young paratype where there
appears to be a series of five segments, one large, one
small and thin, one medium, one small and thin, and one
large. The sutures are serrated and the larger columnals
have a rimlike girdle with an uneven surface.

Measurements in millimeters:
Holotype Paratype

Height of cup* CA 1.9
Width of cup 22.0* 4.2"
Ratio of H/W 0.25 0.45
Length of basal 6.8" Wal
Width of basal et 1:35
Length of radial* 5.0" 1.4"
Width of radial (bile 2D:
Length of PBri 3.9 3.8
Width of PBr 8.0 1.8
Ratio H/W 0.48 7A lit
Width of IBB disk 5.5 1.0
Diameter proximal 3.6" 0.5
columnal

*To just below transverse ridge.
“Approximate.
*Linear measurements.

Remarks. — This is one of the most distinctive appear-
ing species of the genus. The dorsal cup, with ornamentation
limited to the subhorizontal five spines of the basal circlet,
and shallow broad basal concavity, is unique, and the
relatively narrow, rounded arms with numerous spinose
projections are distinctive.

Types. — Holotype, USNM, No, $5153, and paratypes,
USNM, No. $5161, $5162, and S5181, collected by Wm.
T. Watkins.

Occurrence. — Kickapoo Falls Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; north side of Kickapoo Creek, one-quarter
mile below Kickapoo Falls, southwest of Dennis, Hood
County, Texas.

Family ERISOCRINIDAE S. A. Miller
Genus THOLIACRINUS Strimple. 1962
THOLIACRINUS RECTUS (Moore and Plummer), 1940
Plate 44, figures 1-3, 7, 8
See Bassler and Moodey, 1943, p. 451.

This species was established on four dorsal cups from
the Brannon Bridge Limestone. Considerable additional in-
formation is afforded by four topotypes collected by the
junior author which are sets of arms and crowns. The
smallest crown has arms that remain uniserial or mildly

cuneiform to the twelfth secundibrach, which is near the
termination of the arms. One larger crown (mature) shows
the arms become biserial with about the sixth secundibrach.
Complete descriptions of presently considered material are
given below.

It is a tumid form, having sporadic nodes on the
basals, radials, and brachials. The largest specimen has a
cup width of 12.2 mm, height approximately 5.5 mm. The
PBryax is 5.5 mm wide, and the arm is 1.8 mm wide at the
third secundibrach. The arms have a known length of 18
mm for this specimen, without the termination being
reached. There are single nodes on the first four or five
SBr. The brachials are more or less wedge-shaped but do
not become interlocking (biserial) before the fourth or fifth
SBr, and in some instances are not fully interlocking even in
upper portions of the crown. The column has a width of 2.5
mm at the basal plane of the cup. Anal X is a broad plate
resting broadly on the posterior basal and extending above
the cup summit.

The holotype is a crown in excellent preservation except
for the flattened area in the posterior where the specimen
was apparently partially embedded on a limestone layer. The
radials and basals are unusually tumid, with strong curva-
ture longitudinally as well as transversely. It is obvious that
nodes were present, but they are not so readily apparent as in
the large topotype. The basal concavity is broad and deep.
Infrabasals has not been observed due to the nature of
preservation. Facets for reception of five infrabasals are
shown by the lower facets of the basals. Apparently the
upper facets of the infrabasals were very narrow.

In the large topotype the proximal columnals cover the
infrabasal area. The primibrachs of the holotype have a
node just below the pointed upper end and they are of un-
equal lengths. The heights are as follows: right posterior 3.2
mm, right anterior 2.7 mm, anterior 4.2 mm, left anterior
3.2 mm, and left posterior 2.2 mm. Some of the variances
may be partially due to the poor preservation of the plates,
but the fundamental differences are typical of the genus.
The left anterior basal has a length of 5.8 mm and width of
4.1 mm and the left anterior radial has a length of 4.0 mm,
width of 7.0 mm. Measurements are along the surface cur-
vature of the plates.

Two sets of arms which are identified as belonging to
the species are larger than the holotype and reflect a re-
markable ornamentation of brachials, especially near the
termination of the arms. A confluent median groove about
half the width of the arm is formed but near the apex of
each brachial a node is produced to interrupt the continuity
of the groove.

180 PALAEONTOGRAPHICA AMERICANA (VI, 40)

The column of the large topotype is composed of seg-
ments in series of five, although the appearance is of three.
The largest columnals are fairly thick, with flattened ex-
teriors. Thin columnals are marked by external manifes-
tations of the articular crenellae and are followed by a
central columnal that has the appearance of the larger seg-
ments but is not as wide or thick.

Remarks. — Tholiacrinus rimulatus Strimple (1962a)
from the Oolagah Limestone Formation (Desmoinesian) is
of approximately equal age but it has an unusually low
cup (ratio H/W 0.28 as compared to 0.45 of T. rectus).
The basal concavity of 7. rimulatus is also more restricted
than the present species.

T. undulatus (Strimple), 1961a, has more robust arms
which become biserial with the third or fourth secundibrach
and ridges are found to parallel suture lines of the cup.
The arms do not become biserial before the fifth or sixth
secundibrach and no ridges are formed.

The relatively large height of the infrabasal circlet,
yet small portion of basals that participate in the basal
concavity, is considered by Moore and Plummer (1940, p.
301) as a distinguishing feature of the species.

Types. — Holotype, Plummer Collection, No. P-10870,
paratypes, No. P-10869, P-10871, P-10872, University of
Texas, collected by F. B. Plummer and R. C. Moore.
Topotypes, USNM, Nos. $5148, $5202, collected by Wm.
T. Watkins.

Occurence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Genus ERISOCRINUS Meek and Worthen, 1865

Type species. — Erisocrinus typus Meek and Worthen,
1865.

Range. — Desmoinesian to Virgilian, Pennsylvanian.
and Lower Permian; North America.

Diagnosis. — Dorsal cup truncate cone-shaped with
flattened or slightly rounded base and with little or no
concavity (exceptions Erisocrinus obovatus and E. pro-
tensus which have well-defined concavities). The outline of
the cup is regularly pentagonal, as seen in dorsal or ventral
view, with angles of the pentagon clearly defined. Five
infrabasals are small but extend beyond the proximal
columnal. Five basals are large, of equal size except that
the posterior basal may be slightly longer than the others.
Five radials are large, smooth, and prominent. A single
small anal plate is present in a notch between the posterior
radial facets at their inner border and in rare instances it

may extend to the fore and enter the cup. Ten arms are
long, biserial, and composed of wedge-shaped interlocking
segments. The arms divide evenly on the first primibrach,
which plate is comparable in size to the radials and is not
spine bearing.

Specials assigned to Erisocrinus as follows:

Erisocrinus typus Meek and Worthen Missourian: Illinois,

Texas, Oklahoma,

Kansas
Erisocrinus elevatus Moore and Plummer Missourian: Texas
Missourian: ‘Texas
Lower Permian: Texas
Desmoinesian: Oklahoma
Desmoinesian: Oklahoma
Atokan: Texas

Erisocrinus erectus Moore and Plummer
Erisocrinus propinquus Weller
Erisocrinus mediator Strimple
Erisocrinus terminalis Strimple
Erisocrinus georgeae, n. sp.

Remarks. — The genus Erisocrinus has widespread dis-
tribution in rocks of Pennsylvanian age in North America,
especially in the Missourian Stage. There are two types,
those which have an outward flare of radial plates, and
those with even curvature of the radial plates. The two
forms occur together in many horizons and may represent
two lineages.

ERISOCRINUS GEORGEAE Strimple and Watkins, new species
Plate 39, figures 5-7

Dorsal cup truncate bowl-shaped with flared sides and
a moderately wide subhorizontal base. The outline of the
cup is pentagonal when viewed from above or below. In-
frabasals appear to be five, forming a subhorizontal disk
extending only slightly beyond the large, sharply im-
pressed round stem attachment area. Five basals are
slightly tumid, and flex out of the basal plane to form a
portion of the lateral sides of the cup. The radials are
the dominant cup elements in side view and widen appreci-
ably above the basal circlet of plates to form five sharp
projections which give the cup its pentagonal outline. Ar-
ticulating facets slope outward slightly. Outer ligament
furrow is thin. Ligament pit is short but well defined.
Transverse ridge is well developed and marked by fine
crenulations. Adsutural areas are closed for a considerable
distance then widen as a V-shaped notch. Oblique furrows
are ill defined. Muscle areas are large and divided by a
pronounced intermuscular notch. There is no evidence of a
special anal plate. The surface is marked by granules. Arms
are not preserved.

The holotype of this species was collected by Mrs.
Robert L. George of San Antonio, Texas, who generously
donated it to this study. The species is named georgeae in
acknowledgment.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 181

Measurements of holotype in millimeters:

Height of cup 6.5
Width of cup (maximum) 19.4
Width of cup (minimum) 19.1
Width of infrabasal circlet 6.0
Length of basal Soe
Width of basal 5.3
Length of radial to transverse ridge 8.0*
Width of radial to transverse ridge 12.3*

*Measurements taken along surface curvature.

Remarks.—It seems probable that Erisocrinus georgeae
evolved from Sinocrinus as represented by forms like S.
microgranulosus. The asymmetric S. sheareri is a specialized
derivitive of Simocrinus and not in the lineage leading to
Erisocrinus. The shape of the cup of FE. georgeae is close to
that of E. typus, the type species of Erisocrinus; however,
the latter is slightly taller and has a proportionately smaller
stem.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch south-
west of Mason, Mason County, Texas.

Types. — Holotype, USNM, No. 85123 collected by
Mrs, Robert L. George and unfigured paratype collected by
Wm. T. Watkins.

Genus EXAETOCRINUS Strimple and Watkins, new genus

Type species. — Stuartwellercrinus argentinei Strimple,
1949c.

Range.— Missourian Stage, Pennsylvanian; North
American.

Diagnosis. — Dorsal cup cone-shaped, upflared infra-
basals, contour of cup pentagonal when viewed from above
or below. Five infrabasals expand evenly from the columnar
attachment and are visible in side view of cup. Five basals
are large, hexagonal-shaped. Five radials are large, penta-
gonal-shaped, articular facets are subhorizontal, Columnar
attachment scar is subpentagonal to circular. No anal
plates. Arms unknown.

Species assigned to Exaetocrinus:

Missourian: Missouri

Missourian: Illinois
Missourian: Kansas

Stuartwellercrinus argentine Strimple
Erisocrinus conoideus Meek and Worthen
Erisocrinus lustrum Strimple

Remarks.—In order to establish clear lineages it is
necessary to isolate forms which do not find ready place-
ment in established genera. The species considered here
appear to belong to a regressive trend out of Erisocrinus, i.e.
they have evolved to a stage that is comparable to their
primitive progenitors insofar as the shape of the cup is con-
cerned. A high cone-shaped cup is considered to be “primi-
tive”. There are no specimens in the collections at hand.

The name Exaetocrinus is from the Greek word exaitos,
meaning choice.

Genus CRICOCRINUS Strimple and Watkins, n. gen.

Type species. — Paradelocrinus regulatus Strimple,
1949.

Range.— Morrowan to Missourian Stages, Pennsyl-
vanian; North America.

Diagnosis. — Dorsal cup low, bowl-shaped with circular
outline when viewed from below and a subhorizontal base
lacking any appreciable basal concavity. Five small infra-
basals confined to the base of the cup. Five medium-sized
basals, confined essentially to the base of the cup although
the distal tips may be visible in side view of the cup. Five
large radials which form the lateral walls of the cup and
with proximal ends extending into the basal plane of the
cup. Normally there is no anal plate below the summit of
the cup. There are ten pinnular bearing, biserial arms,
branching with the first primibrach in all rays. The column
is round,

Specias assigned to Cricocrinus are as follows:
Desmoinesian: Oklahoma

Desmoinesian: Oklahoma
Desmoinesian: Oklahoma

Paradelocrinus regulatus Strimple
Paradelocrinus disculus Strimple
Paradelocrinus johnstonensis

Strimple
Paradelocrinus wapanucka Strimple Morrowan: Oklahoma
Paradelocrinus subplanus Moore and Desmoinesian: Oklahoma,

Plummer Texas
?Paradelocrinus planus (White) Virgilian: Kansas
?Paradelocrinus toddanus (Butts) Missourian: Missouri

Remarks.— The excellent crown of Paradelocrinus
figured by Frederickson and Waddell (1960, pl. 1) is a
representative of P. regulatus Strimple. Due to lateral com-
pression, the basal plates and infrabasal plates have fallen
inwardly to give the illusion of concavity. The specimen was
reported to be nonpinnular, but pinnules are exposed near
the summit of the crown.

Strimple (1962a, pl. 6, figs. 1, 2) figured a specimen
of Paradelocrinus subplanus with ten biserial arms in place
from the Oologah Limestone Formation, Desmoinesian.
Stage of Oklahoma.

The name is derived from the Greek word krikos mean-
ing ring or circle and has reference to the circular outline

of the cup.

Genus PARADELOCRINUS Moore and Plummer, 1938

Type species. — Paradelocrinus aequabilis Moore and
Plummer, 1938.

Range — Morrowan, Atokan, Desmoinesian and Mis-
sourian Stages, Pennsylvanian; North America.

Diagnosis. — Dorsal cup low, height usually less than
half of the width, base characteristically strongly concave,
involving the proximal portions of the basals as well as the
infrabasals, contour of cup circular when viewed from be-
low. Five infrabasals are small, downflared; five basals are

182 PALAEONTOGRAPHICA AMERICANA (VI, 40)

large in most species but are small in forms where the five
radials become dominant and attempt to reach the infra-
basal circlet. The anal plate is usually missing from the cup
proper but may extend into the cup as a narrow wedge for
a limited distance. The arms are ten, biserial, branching on
first primibrach in all rays.

Species assigned to the genus Paradelocrinus:

Paradelocrinus aequabilis Moore and Morrowan: Oklahoma,

Plummer Arkansas
?Paradelocrinus dubius (Mather) Morrowan: Oklahoma,
Arkansas
Paradelocrinus atoka Strimple Atokan and Desmoinesian:
Oklahoma

Paradelocrinus brachiatus Moore and Desmoinesian: Texas

Plummer

Paradelocrinus iolaensis Strimple Missourian: Kansas

Remarks. —It will be noted that Paradelocrinus obo-
vatus Moore and Plummer and Paradelocrinus protensus
Moore and Plummer, both of Missourian age and from
Texas, are not listed above. The reason is that they have
decidedly pentagonal outlines when viewed from below and,
therefore, do not conform with the generic concept as given
above. They are referred to Erisocrinus with reservation.

Several forms have arrived at comparable stages of
development through the elimination of the anal plates from
the dorsal cup. We feel that Paradelocrinus aequabilis is
probably a derivative of a form like Endelocrinus matheri
(Moore and Plummer) which species is also of Morrowan
age. Paradelocrinus dubius is of the same age but has a
higher cup which is constricted near the summit, and a most
pronounced basal invagination. It is specialized and may
not be closely related. P. atoka is now known to be from
rocks of Desmoinesian age as well as from the Atokan, It is
closely related to P. aequalibilis.

Genus GRAFFHAMICRINUS Strimple, 1961la
GRAFFHAMICRINUS ANTIQUUS Strimple and Watkins, n. sp.
Plate 39, figures 14, 15; Plate 40, figures 9, 10

Dorsal cup low, bowl-shaped with an unusually deep
basal concavity. Five infrabasals are confined to the bottom
of the depressed base where they are downflared and form
a narrow cone. Five large basals flex out of the basal in-
vagination to form a portion of the lateral walls of the
cup. Five radials are wide plates and have a flattened
arcuate area, beyond the outer ligamental area, formed
primarily by a row of large nodes. Articular facets are
mainly obscured by matrix but the outer ligamental area is
well exposed. The outer ligamental furrow is almost obliter-
ated but the outer ligamental ridge is marked by denticles.
The ligamental pit is well defined. The transverse ridge is
marked by denticles and lateral furrows are present. The

adsutural slopes are closed in the outer region and thence
widen rapidly but constrict some inwardly to form a distinc-
tive outline. A somewhat similar appearance of the outer
adsutural slopes was illustrated by Strimple (1962d, text-
fig. 29) for Delocrinus ardmirensis Strimple. A large, broad
anal plate rests evenly on the posterior basal and extends
well above the cup where it flexes inward slightly. It has a
single facet above for the reception of a tube plate. The
entire outer surface of the cup is covered with large
swellings and fine granulations. Sutures between plates
above the basal plane are marked by V-shaped depressions.
The proximal columnal is in place. It is small, round,
and marked by crenulations. The arms are unknown,
Measurements of holotype in millimeters:

Height of cup 9.0
Width of cup (uniform) 23.7
Height of basal concavity 5.3
Width of basal concavity 9.5
Length of basal (right posterior) 12.4*
Width of basal (right posterior) 9.0*
Length of radial to transverse ridge 9.0*
Width of radial 14.0*
Length of anal plate 6.0
Width of anal plate 5.2

*Measurements taken along surface curvature.

Remarks. — This is the oldest species to be referred to
Graffhamicrinus. It may not have evolved directly from the
Morrowan Diphuicrinus Moore and Plummer (as emended
by Strimple and Knapp, 1966), which also has a highly
ornate surface, because the anal plate of that genus flexes
sharply inward and often for a considerable distance, and
is marked by two facets for reception of two tube plates.
Undescribed derivatives of Diphuicrinus are known to us
from Atokan and Desmoinesian or Desmoinesian rocks in
southern Oklahoma which have this characteristic. The pro-
genitor of Graffhamicrinus antiquus is probably like an un-
described species from the Wapanucka Formation which has
a deep cup, deep basal invagination, one erect anal plate,
and has a pronouncedly granular but not pustulose, surface.

Types. — Holotype, USNM, No. $5126 (Kurt Zesch
Ranch), paratype USNM, No. S5130 (Espey Creek), col-
lected by Wm. T. Watkins, reposited in the U.S. National
Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan; Kurt Zesch Ranch, Mason, Mason County,
and Marble Falls Formation, Morrowan?; Espey Creek,
Lampasas County, Texas.

Gens SINOCRINUS Tien, 1924
SINOCRINUS SHEARERI Strimple and Watkins, n. sp.
Plate 39, figures 8-11

Dorsal cup low, asymmetrically bowl-shaped with a

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 183

broad base which is shallowly depressed due partially to a
mild bulbosity of basal plates at the basal plane. In the
holotype the infrabasals appear to be fused as a solid disk,
subhorizontal in attitude, and extend a relatively short
distance beyond the large stem facet. In the paratype the
division of infrabasals to five is shown and all do not ex-
tend beyond the stem cavity. There are two small basals
and three large ones. The center basal of the large ones is
slightly larger than the other two and probably represents
the posterior. The proximal portion of a basal is in the basal
plane and the distal portion flexes sharply upward to form
a part of the lateral walls of the cup. Five radials are mildly
tumid in proximal portions and are of equal size. Outer
ligamental furrow is poorly developed but the ligamental pit
is sharply defined. The transverse ridge extends to the inter-
radial suture and projects slightly to interrupt the V-shaped
depression between radials, It is marked by fine denticles.
Oblique furrows, broad muscle scars, and a small V-shaped
inter-muscular notch are present. The articular facets are
sloped outwardly ever so slightly. No evidence of a special
anal element is preserved. The entire outer surface of the
cup is decidedly granular appearing.

Columnar cicatrix is sharply depressed, is large, round,
and the perimeter is marked by fine crenulations.

Measurements of holotype in millimeters:

Width of cup (uniform) 23.2
Height of cup (maximum = posterior) 12.0
Height of cup (minimum = anterior) 8.9
Width of infrabasal circlet 8.0
Length of basal (posterior) 10:9*
Width of basal (posterior) 11.4*
Length of radial to transverse ridge a IBC
(right posterior)
Width of radial (right posterior) 14.6*

*Measurements taken along surface curvature.

Remarks. — Sinocrinus sheareri is distinguished from
the Asiatic species, S. microgranulosus Tien, S. microgranu-
losus “var.” pentalobosus Tien, and S. linchengensis Tien in
having a proportionately lower and asymmetric dorsal cup.
S. sheareri has a proportionately larger column, and the
basals do not form as much of the cup walls as the Asiatic
species. There are two other forms described by Tien as S.
nodosus Tien and S. nodosus “var.” spinosus Tien for
which the infrabasals are not known and no effort was made
to reconstruct the dorsal cup.

The holotype of S. sheareri was found over 30 years ago
by the late JHugh Shearer. He used it and sections of
columns to decorate the surface of a diamond-shaped, three
inch thick, 40 Ib. slab of concrete which he made for one of
his children. The slab lay in the front yard under a tree for
over 30 years, until Mrs. Shearer gave the junior author

permission to remove the crinoid. The weathering must have
had little effect since the granular ornamentation was not
obliterated.

Types. — Holotype USNM, No. S5124, collected by
JHugh Shearer, and one paratype transferred by Wm. T.
Watkins for deposit in the U.S. National Museum, Wash-
ington, D, C.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch south-
west of Mason, Mason County, Texas.

Family APOGRAPHIOCRINIDAE Moore and Laudon, 1943
Genus APOGRAPHIOCRINUS Moore and Plummer, 1940
APOGRAPHIOCRINUS sp.

Plate 34, figure 5; Plate 55, figure 3

Two small crowns are referred to the genus A pographio-
crinus. They are juvenile with unusually long axillary primi-
brachs and have the usual complement of cup plates includ-
ing a single anal plate. The entire surface of the crown is
covered by minute granules and a light median ray extends
the length of the arms. The juvenile characteristics are the
same as those reported by Strimple (1938, 1959) for
Apographiocrinus typicalis which species, however, is Mis-
sourian in age and in maturity is less ornate than most
other species of the genus.

Repository. — Specimens are reposited in the U.S. Na-
tional Museum, USNM, No. $5109, $5192.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles northwest of Brock, Parker
County, Texas.

Family EXOCRINIDAE Strimple and Watkins, new family

Genus — Exocrinus Strimple, Oxynocrinus, n. gen.

Range.— Morrowan to Virgilian, Pennsylvanian;
North America.

This family is comprised of small forms with multiple
arms. The cup is shallow, cone-shaped with infrabasals
mildly upflared in the older genus and subhorizontal to
downflared in the younger genus. Five basals are small with
five radials being the dominant cup plates. In the older
Oxynocrinus there are three anal plates in more or less
primitive arrangement, 7. e. anal X is in contact with the
posterior basal, but the radianal is more vertical than di-
agonal in position. In Exocrinus the radianal has moved to
posterior position and supports two anal plates above.
Fusion of brachials is a common feature in Exocrinus but
has not commenced in Oxynocrinus.

There appears to be some affinity between the exo-
crinids and the stellarocrinids. The latter appear to be more

184 PALAEONTOGRAPHICA AMERICANA (VI, 40)

advanced in having biserial arms in most instances although
some species and some young specimens have uniserial arms
(cuneiform brachials).

Genus OXYNOCRINUS Strimple and Watkins, new genus

Type species. — Oxynocrinus spicata, n. sp.

Range. — Morrowan, Pennsylvanian; North central
Texas.

The crown is relatively short and broad. The arms are
uniserial, fundamentally isotomous branching produces
more than 40 keeled arms. Dorsal cup expands evenly from
the proximal columnals with infrabasals visible in side view.
There are five basal plates, five radial plates, and three
anal plates which are in normal (Primitive) arrangement.
First primibrach is axillary. Each brachial bears a pinnule.

OXYNOCRINUS SPICATA Strimple and Watkins, new species
Plate 36, figure 4

The crown of this species expands evenly from the
proximal columnals to above mid-height, thereafter tapering
slowly to its termination. The arms could not become
closely apposed.

The dorsal cup is of medium height and is cone-shaped
with upflared infrabasals. Five infrabasals are short ele-
ments. Five basals are of moderate size. The sutures be-
tween basals are short. Five radials are large prominent
plates. Three anal plates within the cup are normal ( Primi-
tive Type) with the radianal a prominent element but anal
X is still in contact with the posterior basal.

The uniserial arms branch with the first primibrach in
each ray. The axillary primibrach is somewhat smaller than
the radial and is not elongated except for the anterior which
is longer than the other four primibrachs. A second branch-
ing takes place with the second or third secundibrach in all
arms and another with the fourth or fifth tertibrach in all
divisions. Another bifurcation takes place in some arms.
The brachials are mildly constricted in mid-section and
are keeled. The keel is projected as a small spine at the
distal end of each nonaxillary brachial in the pinnular bear-
ing side. Pinnules alternate, one to a brachial.

Measurements of holotype in millimeters:

Length of crown 36.0
Height of dorsal cup 5.0
Width of dorsal cup (maximum 12.0

as preserved)
Length of basal 3.0
Width of basal 3.5
Length of suture between basals 0.5
Length of radial by
Width of radial 5.3

Length of suture between radials
Length of radianal

Width of radianal

Length of anal X

Width of anal X

NNVey
mnAnooren]

Remarks. — Oxynocrinus spicata is monotypic of the
genus. The arms are somewhat comparable to some arms
found in the genus Ampelocrinus except there are no syzy-
gial pairs in the former and the latter has paired primi-
brachs. The brachials are similar to those of Exocrinus
other than the fusion of some brachials found in the latter
genus. Oxynocrinus spicata has upflared infrabasals where-
as the infrabasals of the younger Exocrinus are confined to
a shallow basal concavity Stellarocrinus distinctus is atypi-
cal of the genus Stellarocrinus in having uniserial arms
rather than biserial arms and may represent a lineage lead-
ing from Oxynocrinus spicata.

Holotype. — USNM, No. $5117 collected by Wm. T.
Watkins, reposited in the U.S, National Museum.

Occurrence. — Marble Falls Formation, Morrowan?,
Pennsylvanian; Espey Creek southwest of Lampasas, Lam-
pasas County, Texas.

Family AGASSIZOCRINIDAE §S. A. Miller, 1889

The only form in the present study that might be
considered in this family is Paragassizocrinus Moore and
Plummer, 1940.

The genus Paragassizocrinus appears to be advanced
in having a single anal plate and has been considered in the
lineage of Agassizocrinus, which genus typically has four
anal plates in the posterior interradius. Actually it has been
found that Paragassizocrinus (= like-Agassizzocrinus) is
not even related to Agassizocrinus. It has some homologous
features, e. g. the ability to discard the column and fuse the
infrabasal disk which has in the past caused us to consider
it as a close relative of Agassizocrinus. A character of
Paragassizocrinus which has been troublesome under pre-
vious concepts is the existence of two primibrachs, with the
second being axillary. Agassizocrinus has a single primibrach
in all rays, which is axillary. Normal evolutionary trends
among the inadunates usually reflect that the point of bi-
furcation moves toward the cup, not away from the cup.
Fortunately we no longer need to concern ourselves with
these comparisons because we now find Paragassizocrinus
belongs to the Ampelocrinidae rather than the Agassizo-
crinidae and what previously seemed strange is found to be
perfectly normal. The family characteristically has one
anal plate and two primibrachs.

Further discussion of the matter will be found in the
section devoted to the Ampelocrinidae.

TExAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 185

Family AMPELOCRINIDAE Kirk, 1942

Range. — Mississippian to Permian.

Subfamilies.— Ampelocrininae Kirk, Paragassizocrin-
inae new subfamily.

This family is of particular interest in the present
study because of the presence in the Lemons Bluff fauna
(Atokan) of Poluwsocrinus and a closely related Paragassizo-
crinus. We considered the possibility of the smaller Poluso-
crinus being the young of Paragassizocrinus but could not
reconcile this with the fact that small (young) fused cones of
Paragassizocrinus have been found elsewhere. Polusocrinus
and Paragassizocrinus are found throughout the Pennsyl-
vanian albeit seldom together. It appears that Paragassizo-
crinus, which has formerly been considered a member of the
Agassizocrinidae, is a specialized ampelocrinid in which the
stem has become atrophied and a fusion, thickening and
elongation of the infrabasal plates, takes place. Petschora-
crinus from the Permian of Russia is apparently closely re-
lated to Paragassizocrinus. The progenitor of these genera
appears to be Ampelocrinus which genus starts with small
forms having a single anal plate resting obliquely on the
upper surface of the posterior basal and with infrabasals
readily visible in side view of the cup. A. erectus Strimple
from the Chester is of medium size. All of the forms in-
volved have in common a single anal plate, a narrow, in-
wardly directed articular facet of the radial, and two primi-
brachs, the second being axillary.

Polusocrinus was proposed by Strimple (1951d) with
Polusocrinus avanti Strimple of Missourian age as the type
of the genus. Strimple (195ld, p. 24) considered Polwso-
crinus as a derivative of Aesiocrinus, but we do not believe
they are so closely related. The range of Polusocrinus has
been extended downward into the Callville Formation of
Nevada by Lane (1964b) who considers the age of the two
species to be no younger than early Atokan and probably of
Morrowan age. One of the species, P. pachyplax Lane, has a
remarkable thickening of the infrabasal plates, and the
apparent atrophy of the column in larger (more mature)
specimens.

Kirk (1942, p. 23) recognized two lines of evolution
in the family Ampelocrinidae, based on information avail-
able to him, “No described crinoids of earlier age than St.
Louis can be referred to the Ampelocrinidae. There are un-
described genera in the lower Mississippian with a single
anal plate in the post-/R and with syzygial pairs of Br.
When described, it may be expedient to refer them to the
Ampelocrinidae, although they most certainly pertain to
another evolutionary series.” When Kirk presented Cym-
biocrinus in 1944, he stated that it was the form he had in

mind in 1942; however, he referred to “genera” in that
study. We fail to see why it is necessary to leave Cymbio-
crinus and related genera in the family Ampelocrinidae
when they belong to another evolutionary series as ex-
pounded by Kirk, and we propose a new family Cymbio-
crinidae Strimple and Watkins.

Subfamily AMPELOCRININAE Kirk

Genera. — Ampelocrinus Kirk, 1942; Polusocrinus
Strimple, 1951d; Moundocrinus Strimple, 1939a.

Range. — Mississippian to Permian; USSR, Scotland,
North America.

Diagnosis. — Dorsal cup: Five IBB, medium to large,
upflared in older forms but subhorizontal in most younger
forms; five BB, medium-sized; five RR large, articular facet
is usually wide, short and inwardly sloped; one anal plate, in
line with radials but ysually extends slightly above the
summit of the cup and is followed by two tube plates above
in older forms but may be followed by a single tube plate in
some younger forms (e¢.g. Moundocrinus); columnar a tach-
ment is of medium size to small or even atrophied, circular
outline in older forms but pentagonal in some younger
forms.

Arms: Uniserial and not apposed, pinnular with alter-
nating, long pinnules, bifurcate with the second primibrach
in all rays and there may be another bifurcation in some or
all arms. The brachials in the older forms are often syzy-
gial pairs, in which case they act as a single brachial, i.e.
have a single pinnule.

Anal Sac: The anal sac is in the form of a tube but is
seldom preserved in the group.

Column: The column is round in older forms but is
usually pentagonal in more recent forms when there is no
tendency toward atrophy as happens in some species. Cirri
in whorls are typical.

Remarks.— The Ampelocrininae are closely aligned
with the Paragassizocrininae, with the latter differing in
having fused infrabasals at maturity and complete loss of
the use of the stem which is usually atrophied. This results
in a different appearing cup in that the Paragassizocrininae
take on an elongate cone-shape as opposed to the bowl-shape
of the Ampelocrininae. All ampelocrinids have mildly to de-
cidedly upflared infrabasals except Mowndocrinus which
has arrived at a stage homologous with some advanced
cymbiocrinids wherein the infrabasals are subhorizontal.
Moundocrinus is retained with the ampelocrinids because
of the short articular facets which are a consistent character-
istic of the family. Moore and Plummer (1941), e¢ al.,
considered Mowndocrinus as synonymous with Aesiocrinus

186 PALAEONTOGRAPHICA AMERICANA (VI, 40)

but the radial articular facets of that genus are long, and
the anal plate is followed by two tube plates. In Moundo-
crinus the anal plate is followed by one tube plate.

Genus POLUSOCRINUS Strimple, 1951d
POLUSOCRINUS CALYCULOIDES Lane, 1°64
Plate 42, figure 2; Plate 43, figures 6, 7

This species is represented in the current collections
by one crown in excellent preservation and two cups with
the proximal portions of the arms attached. The cup is bowl-
shaped but has erect lateral sides and the arms have a
tendency to be recumbent. The infrabasal plates are large,
plainly separated by sutures and form a bowl with the
central portion depressed. A few segments of the minute
column are preserved and the first one or two columnals
have an angular outline but thereafter they have a circular
outline and are alternatingly expanded. Round cirri are
present a short distance from the cup. Five basals are promi-
nent elements with that of the posterior truncated evenly
for the reception of the single anal plate, Five radials are
wider than long and have narrow upper articulating facets
that fill the distal face of the plates.

There are two primibrachs in each arm, the first of
medium length and the second pointed (axillary). No fur-
ther bifurcation of the arms takes place. Secundibrachs are
slightly elongated, have even sutures and well-rounded
exteriors. Each brachial bears a pinnule which alternates
with the pinnule of the adjoining brachial.

Measurements in millimeters:

Figured Hypotype

Height of dorsal cup 4.7
Width of dorsal cup
Height of infrabasal circlet
Width of infrabasal circlet
Length of infrabasal
Width of infrabasal
Length of basal
Width of basal
Length of interbasal suture
Length of radial
Width of radial
Length of first primibrach

Width of first primibrach
Length of second primibrach

* * *

a a
GOED SIE ON > On iete ty 10nd

(axillary)
Width of second primibrach 3.1*
(axillary)
Length of first secundibrach 20
Width of first secundibrach 2.4*

eEstimated.

*Measurements taken along surface curvature.

Remarks. — Presently considered material is considered
to be conspecific with Polusocrinus calyculoides Lane
(1964b). The holotype demonstrated some asymmetry in

the infrabasal circlet wherein some plates appear to be
larger than others which condition is not found in the Texas
material.

Types. — Hypotypes USNM, Nos. $5137, $5143, and
one unnumbered specimen, collected by Wm. T. Watkins
and reposited in the U. S. National Museum.

Occurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Limestone Formation, Atokan, Pennsylvanian; 11
miles southwest of San Saba on Rough Creek, San Saba
County, Texas.

Subfamily PARAGASSIZOCRININAE Strimple and Watkins,

new subfamily

Genera. — Paragassizocrinus Moore and Plummer,
1940; Petschoracrinus Yakovlev, 1928.

Range. — Pennsylvanian to Permian; North America,
USSR.

Diagnosis. — Dorsal cup: cone-shaped, usually elongate;
five IBB decidedly upflared at maturity and usually fused
in the Pennsylvanian but often divided by suture in the
Permian, youthful stages often show sutures and columnar
attachment area; five BB medium-sized, often become elon-
gate; five RR medium size, articulating facet fills width of
plate but is short and directed inwardly; one anal plate
followed by one tube plate out of the cup; column is
only known from a few segments which appear to be
round and is usually entirely obliterated in mature Pennsy]l-
vanian specimens except in the interior of the cone where
the rudimentary remains may be seen in cross sections of
the basal cone.

Arms: The arms are uniserial, with even sutures, and
branch with the second primibrach in Paragassizocrinus
but do not branch at all in Petschoracrinus.

Remarks.— Comparison with the Ampelocrininae has
been made under that group. Paragassizocrinus is thought
to have evolved from a form like Polusocrinus of the Am-
pelocrininae. The change is essentially in the basal area of
the dorsal cup and the first step is atrophy of the column (a
feature that has been observed in some Polusocrinus) which
then becomes deciduous. A portion of the column in the
proximal region is covered by the heavy influx of stereom®
which usually also obliterates the sutures between infra-
basals and produces a more or less pointed cone-shape to
the infrabasal circlet. At noted elsewhere, the remnant of
the rudimentary proximal stem segments are visible in
cross-section, This is also a feature of Petschoracrinus, as
well illustrated by Yakovlev (1956, pl. 15, fig. 6), but in
®° Stercom, after Bather, 1891 (Ann. Mag. Nt. Hist. (6) vol. 9, “any

hard calcareous tissue forming skeletal structures in Metazoa
Invertebrata and in Protoaozoa”’.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 187

this genus the tendency toward complete fusion of the infra-
basal plates has diminshed although the plates retain a
considerable length. In the same category is the tendency to
not obliterate the column although it appears to have been
deciduous near the base of the cup.

Genus PARAGASSIZOCRINUS Moore and Plummer, 1940

Type species. — Agassizocrinus tarri Strimple, 1938.

Range. — Pennsylvanian; North America.

Remarks. — Although superficially comparable there
have been three features which cause difficulty in attempt-
ing to reconstruct the evolution of Paragassizocrinus from
Agassizocrinus. The first is with the posterior interradius
wherein some of the youngest known species of Agassizo-
crinus (e.g. A. patulus Strimple, 195le, of the Pitkin
Formation, Chesterian) has four anal plates within the cup
which is a common condition for the genus. All forms
known from the Pennsylvanian have a single anal plate.

The second feature is with the arms which in Agassizo-
crinus divide on the first primibrach but in Paragassizo-
crinus bifurcation is with the second primibrach. Branching
on the second primibrach is typical in the ampelocrinids.

The third feature concerns the base of the cup. The
older genus abolished its column and fused the infrabasals
into more or less pointed cone, yet the sutures between
infrabasals are more often present than missing even in the
most recent species of Agassizocrinus. Yet there is more
vestigial evidence of a column in Paragassizocrinus than in
Agassizocrinus, and some Morrowan species retain evidence
of the sutures between infrabasals. The reason for this is
that Paragassizocrinus is evolved from a form having a stem
and five divided infrabasals rather than from Agassizo-
crinus. The ancestral form of Paragassizocrinus is thought to
be an ampelocrinid.

Many forms assigned to Paragassizocrinus are only
known from disassociated portions of the cup so that posi-
tive identification is difficult and we know there are several
different forms involved. It is possible that Agassizocrinus,
or a derivative of the lineage, will be found in the Pennsyl-
yanian, but we believe there will be distinctive features
of the cup or arms to distinguish it from Paragassizocrinus.

A form described below as Paragassizocrinus altus,
n. sp., is the first species of the genus for which an appreci-
able portion of the arms is known.

PARAGASSIZOCRINUS ALTUS Strimple and Watkins, n. sp.
Plate 53, figure 3; Plate 56, figure 5
This species is based on two crowns in excellent preser-

vation. The cup is high and cone-shaped and the ten uni-
serial arms are long, slender, and not closely abutting.

The infrabasal plates are fused with no trace of sutures
or of a stem. The infrabasal cone is low and pointed. There
are five large basal plates which are about as wide as long.
The posterior basal is truncated for reception of a large anal
plate that rests evenly on the upper facet. Five radials are
wide, pentagonal-shaped, and have a mildly gaped suture
with first primibrachs. There is some dislocation of plates in
preservation so that it is possible to see the articular facets
of the radials. The outer ligamental area is well developed
and directed outward (which causes the gaped suture) al-
though the groove is not particularly deep. The outer liga-
mental pit is deep and well defined with the sharp crested
transverse ridge behind it. The transverse ridge does not
extend entirely to the lateral sides but is cut off by well-
developed lateral furrows. The lateral furrows meet so that
a long furrow is formed behind the transverse ridge. There
is a slight rise and then the inner facets slope inward, are
short, and the two muscle areas are sharply depressed. The
intermuscular notch is small and is flanked by two small,
elongated depressions at right angles to it. The single anal
plate extends slightly above the summit of the dorsal cup
and is faceted for reception of two tube plates above.

First primibrach is quadrangular and just about equals
the width of the radial plate, Second primibrach is some-
what shorter than the first and is axillary, The two primi-
brachs are held by close suture. None of the brachials
show any subsequent tendency toward syzygial develop-
ment but are regularly quadrangular-shaped with an occa-
sional slightly cuneiform segment. The arms of the holotype
attain a length of 48.5 mm. There is a decided outer curva-
ture of the arms.

Measurements in millimeters:

Holotype Mature Paratype

Height of dorsal cup 10.5 11.5
Width of dorsal cup 10.0¢ 11.0
Height of IBB circlet 3.7 3.6
Width of IBB circlet 6.6 7.8

Length of infrabasal — as
Width of infrabasal — —

Length of basal 5.4 5.6
Width of basal 5.5 6.0
Length of interbasal ) Bal
suture
Length of radial 4.0 4.0
Width of radial 6.3 6.4
Length of PBr 3.0 2.8
Width of PBr 6.2 6.2
Length of PBreax 2.6 —

Width of PBreax 5.3 —
Length of SBr: 2.8 =
Width of SBri 28 —
eEstimated.

Remarks. — This species is closer to Paragassizocrinus
deltoideus Strimple, 1960, than other described forms.

188 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Careful comparison discloses that P. deltoideus has pro-
portionately wider basal plates and radial plates, and the
suture between basals is shorter than in P. altws. These fea-
tures produce a shorter, broader cup in P. deltoideus. The
cone of P. caliculus Moore and Plummer is somewhat com-
parable except in having a flatly rounded base as against
the pointed base of P. deltoideus and P. altus.

Paragassizocrinus tarri Strimple, 1938, P. mcguirer
(Strimple), 1939a, P. asymmetricus Strimple, 1960c, P.
elongatus Strimple, 1960c, P. turris Strimple, 1960c, P.
hoodi Strimple, 1960c, P. springeri Strimple, 1960c, P.
ellipticus Strimple, 1961b, P. elevatus Strimple, 1961, and P.
kendrickensis Strimple and Knapp, 1966, all have consider-
ably longer cones than P. altus, which allows for quick dif-
ferentiation. P. bulbosus Strimple, 1961, is a larger form,
and although it has a comparable contour to the infra-
basal cone, it is proportionately taller. P. disculus Strimple,
1960c, is an infrabasal cone with a somewhat comparable
shape, but it is a larger form and is proportionately shal-
lower. P. atoka Strimple and Blythe, 1960, has a more
rounded contour at the base of the infrabasal cone.

Types. — Holotype, USNM, No. $5199 and paratype,
USNM, No. 85184, collected by Wm. T. Watkins.

Occurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Limestone Formation, Atokan, Pennsylvanian;
11 miles southwest of San Saba on Rough Creek, San Saba
County, Texas.

Family CYMBIOCRINIDAE Strimple and Watkins, new family

Genera. — Cymbiocrinus Kirk, 1944; Aestocrinus Miller
and Gurley, 1890 (Syn. Pentadelocrinus Strimple, 1939a;
Phialocrinus Trautschold, 1879 non Eichwald, 1856);
1856); Oklahomacrinus Moore, 1939; Lecobasicrinus Strim-
ple and Watkins, n. g.; Childonocrinus Strimple and Wat-
kins, n. g.; Allosocrinus Strimple, 1949b.

Range. — Mississippian-Pennsylvanian; North America,
USSR.

Diagnosis. — Dorsal cup: Cup shape ranges from basal-
ly impressed bowl of medium height to a flat, basally im-
pressed cone; five IBB small, confined to basal concavity
in most forms but may become subhorizontal in some
younger species; five BB large, articular facet normally
fills width of plate, slopes inward and is short,® one anal
plate, in line with radials but usually extends slightly above
the summit of the cup, is followed by two tube plates in
older forms but may be followed by one tube plate in some
of the younger forms.

Column: The column has a circular outline in older

"Exception is A4esiocrinus which typically has long facets.

forms but is pentagonal in most younger forms. Cirri are
usually in whorls.

Arms: Arms are normally slender, uniserial, not apposed
and ten in number; however, in one Missourian species
(Aesiocrinus basilicus) there are more than ten reported,
and in two species (Aesiocrinus barydactylus and A. lykinsi)
only five arms are typically present. Syzygial pairs of
brachials are common in older forms but have not been
observed in younger forms unless the paired primibrachs
are considered a syzygy. When more than five arms are
present, the bifurcation always takes place with the second
primibrach. Long pinnules are present on alternating sides
of the arms.

Anal sac: The anal sac is cylindrical and may attain
a considerable length. In one genus (Aesiocrinus) the sac
becomes rugged and is composed of thick plates with
roughened exteriors.

Remarks. — This family is characterized by and separ-
able from the associated Ampelocrinidae because the small
IBB are essentially confined to a basal concavity and never
attain a truly upflared attitude.

There are small forms in the Mermac, confined to the
genus Cymbiocrinus, which become larger in the Chester. A
specialized ornate group appears in the Chester, represented
by the genus Childonocrinus, and continues into the Atokan.
Lecobasicrinus represents another specialization found in
the Atokan and Desmoinesian, wherein the RR and IBB
atempt to contact one another and some arms may exhibit
hypertrophy. The genus Oklahomacrinus is well known in
the Demoinesian and continues on up into the Virgilian. It
is highly specialized in the development of a shallow cup
with an exceedingly deep basal concavity and has more or
less recumbent arms. Another group with a shallow cup is
represented by Aesiocrinus paucus Strimple (1951d) from
the Missourian and A. prudentia Strimple (1962d) from
the Virgilian, Other characters are distinct from those of
Oklahomacrinus. As previously noted, Aesiocrinus has a
prominent anal tube and typically has relatively large radial
articulating facets. It is mainly a Missourian form and has
the largest cup in the family with the exception of Alloso-
crinus, which form is only tentatively assigned to the family.
The base of the cup in various species of Aesiocrinus varies
from concave to subhorizontal. Allosocrinus is known from
the Desmoinesian and Missourian. It differs appreciably
from other Pennsylvanian forms in having a round stem,
five robust arms and the articulating surface of the radials,
and between brachials, are marked by radiating crenellae.

Genus CHLIDONOCRINUS Strimple and Watkins, new genus

Type species. —Childonocrinus echinatus, n. sp.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 189

Range. — Chesterian (Mississippian) to Atokan Penn-
sylvanian); Texas and Oklahoma, North America.

Generic diagnosis.— Dorsal cup low, basin-shaped
with mildly concave base; IBB obscured, probably five,
confined to basal concavity; BB five, medium-sized, pos-
terior basal truncated for reception of the single anal plate;
RR five wide, pentagonal-shaped, suture gaping; two
primibrachs syzygial paired, PBr, wide low, non-axillary,
PBr» tringular-shaped, axillary; BrBr quadrangular except
for subsequent axillaries; arms do not abut; column penta-
lobate, cirriferous (cirri elliptical to circular in cross sec-
tion); calyx and arms highly ornate.

Remarks. — This form is represented in the collections
by three crowns from the Atokan in excellent preservation
and two cups with lower arm brachials preserved from the
Chesterian, being two different species. The posterior in-
terradius is occupied by a small, narrow anal plate, extend-
ing slightly above the cup in the Chesterian species. The
habit of the calyxes and the paired primibrachs leaves little
doubt that the forms belong to the ampelocrinids. The
columns of most ampelocrinids are small. An exception is
the large, round, or mildly pentagonal stem of Lecobasi-
crinus Strimple and Watkins which does not appear to be
cirriferous in proximal portions. Older (Mississippian)
representatives (C\ymbiocrinus) of the family usually have
columns that are circular in cross-section and are cirriferous.
Younger forms (Upper Pennsylvanian) such as Aersiocrinus
and Oklahomacrinus have pentagonal-shaped stems which
are usually relatively small. The large, pentalobate stem and
the ornate exterior of Chlidonocrinus is apparently unique
for the family.

The Greek word chlidon for ornament is especially
appropriate because of the spinose nature of this genus.

CHLIDONOCRINUS ECHINATUS Strimple and Watkins, n. sp.

Plate 42, figures 1, 4; Plate 51, figure 4

This species is represented in the collections by three
magnificent crowns available through a combination of re-
markable preservation and a unique method of preparation.
When collected, these crowns, and others herein described
from the same locality, were lying, mostly buried, on the
upper surface of a ledge of dark impure limestone about
seven inches thick. Slabs of the ledge were pried out of the
weathered cliff surface from beneath a massive three-foot
ledge. The ledge was separated from above and below by
soft shales, approximately one-half inch in thickness. After
the slabs were obtained sections about one inch thick were
sawed off the upper portion which contained the crowns.

Ordinary preparation methods were too hazardous because
of the projections, delicate pinnules, spines, and cirri, Con-
tinual soaking and repeated brushing over a period of
weeks gradually removed the dark matrix and revealed the
remarkable crowns lying full-length on the slabs and still
half buried in the denser matrix. Additional brushing with
a moist (not wet) toothbrush brought out even more detail.
Unfortunately, all three crowns have the posterior inter-
radius on the buried side.

The crown is elongate, expanded, and long cirri extend
up around the crown. Surface of all cup and arm plates
marked by small, spinelike nodes. Because this is the type
species the generic diagnosis also applies to the species.
Some distortion is caused by lateral compression.

The dorsal cup is essentially low, broadly expanded,
cone-shaped. Infrabasals have not been observed but are
inferred to have been either downflared or subhorizontal
because of the curvature at the proximal end of the basal
plates. The basals are large, hexagonal-shaped. Radials are
wide pentagonal-shaped with the proximal and distal edges
marked by a row of small spinelike nodes. The lateral sides
of the radials are recurved, as well as the proximal apex,
so that depressed areas are formed in conjunction with
adjacent plates. There is a gaped suture with the first primi-
brach. A syzygial pair of first primibrachs are present,
which are entirely anchylosed in some instances, the lower
one being almost as wide as the radial and quadrangular-
shaped and the one above is axillary and triangular-shaped.
The apex of the axillary is broad and the first secundibrachs
are directed outward at a low angle so they could never
abut. A row of nodes marks the perimeter of the syzygial
pair. Subsequent brachials are often syzygial pairs. A
series of nodes mark the lateral sides of the arms accen-
tuated by alternatingly projected spines. The brachials are
usually somewhat elongated, although some are low when
involved in a syzygy. They are essentially cuneiform in
structure. A second bifurcation takes place with about the
ninth or tenth secundibrach in all rays and another may
take place with about the twentieth tertibrach. Pinnules are
present but it is difficult to establish the pattern of occur-
rence.

A crown may attain a length of 90 mm with the cup
only 4.7 mm high. The cup of the holotype has a width of
12.3 mm, but this is slightly exaggerated.

Linear measurements in millimeters:

Length of basal 2.9
Width of basal 2.8
Length of radial 3.1
Width of radial 5.0
Length of PBr 7.
Width of PBri 4.5

190 PALAEONTOGRAPHICA AmeErIcANA (VI, 40)

Length of PBr.ax 10
Width of PBrsax ai
Width of proximal] columnals 3.0

The column is pentalobate with relatively large cirri.
Near the column the cirrals are elliptical-shaped and have
a median fulcral ridge comparable to the structure found in
the stem of Platycrinites. After five or six rapidly tapered
columnals the cirri become round and are pierced by a small,
round lumen. Short spines mark the upper perimeter of
many cirri.

Remarks. — This species is not closely comparable to
any species other than Chlidonocrinus trinodi from the
Chesterian of Oklahoma. The later has spinelike nodes
across the distal end of the radial but not on the proximal
ends and the lateral sides of the radials are not depressed,
as found in C. echinatus. The axillary second primibrach of
C. trinodi has three prominent nodes marking the angles of
the triangle and providing the distinguishing character for
which the species is named.

Types. — Holotype, USNM, No. $5139, paratype,
USNM, No. $5136 and $5174 collected by Wm. T. Watkins.

Occurrence. — Lemon Bluff Limestone Member, Mar-
ble Falls Limestone Formation, Atokan, Pennsylvanian; 11
miles southwest of San Saba on Rough Creek, San Saba
County, Texas.

CHLIDONOCRINUS TRINODI Strimple and Watkins, n. sp.
Plate 43, figures 3, 4

This species is included here although of Chesterian age
because it is the only form known to us closely related to
the previously described Chidonocrinus echinatus of Atokan
age. Two cups with a few lower brachials attached are avail-
able for types of C. trinodi. They are both distorted by
lateral compression.

We believe the dorsal cup to be low, cone-shaped with
subhorizontal or mildly downflared infrabasals. The exact
nature of the base is obscured in both specimens, but the
basals formed in a manner suggesting the existence of a
flattened or subhorizontal base. The basals are large hexa-
gonal-shaped except for posterior basal which has one extra
facet and unornamented except for a bulge in mid-portion
near the proximal edge. The radials are pentagonal low wide
plates and ornamented by a row of nodes along the distal
edge of each plate. Suture is gaped between the radial and
first primibrach. A single anal plate is preserved in the
holotype although slightly displaced. It is long, wide, ex-
tends slightly above the summit of the cup, and is faceted
for reception of two tube plates.

Two primibrachs form a syzygial pair, the lower one
almost filling the distal face of the radial and the upper
one triangular-shaped and axillary. PBr; has a row of nodes
along its proximal edge. PBrzax has three prominent nodes
marking the angles of the triangle. A few secundibrachs are
of a cuneiform nature and have nodes. The column is small,
apparently mildly pentagonal and cirriferous. Cirri have a
circular outline but in proximal position appear to have a
fulcral ridge.

Linear measurements in millimeters:

Width of cup 12.0
Height of cup 7.0¢
Length of basal 35
Width of basal

Length of suture between basals
Length of radial

Width of radial

Length of anal

Width of anal

Length of PBr,

Width of PBr,

Length of PBrs

Width of PBre

Width of proximal columnal*

PAGS UBS! 03 GS aif bs oS
MWnanrtrwnounoonwn

*Estimated.
*Paratype.

Remarks.—Chlidonocrinus trinodi is apparently a
transitional form between smooth surfaced Cymbiocrinus
and the highly ornate Chlidonocrinus echinatus of Atokan
age.

The name trinodi is from the Latin word tri for three
and nodus for swelling, with reference to the three nodes
on the PBrsax.

Types. — Holotype, SUI, No. 32260, paratype USNM,
No. $5142, collected by H. L. Strimple.

Occurrence. — Fayetteville Shale Formation, Chester-
ian, Mississippian; about seven miles northeast of Vinita,
Craig County, Oklahoma.

Genus CYMBIOCRINUS Kirk, 1944
CYMBIOCRINUS CONTENTUS Strimple and Watkins, n. sp.
Plate 40, figures 11, 12

The holotype is slightly distorted through compression
but is well preserved, considerably larger than typical for
the genus. The dorsal cup is low, bowl-shaped with a basal
concavity. Five infrabasals extend slightly beyond the round
stem and appear to be downflared. Five basals are large and
extend out of the basal concavity. Posterior basal is evenly
truncated to support an anal plate above. Five radials are
large. The single anal plate extends above the summit of
the cup and has facets for the reception of two plates above.

There are ten large, uniserial arms branching with the
second primibrach in all arms. First primibrach is a low,

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 191

wide element and the second primibrach is triangular-
shaped, with no lateral sides. Secundibrachs are low, broad
with strongly rounded exteriors, and a faint rim is formed
along the distal side. Sutures are irregular with the pinnular
bearing side of the brachial high and conversely the non-
pinnular bearing side of the adjacent brachials are short-
ened. There is an overhang of the inner edge of the brachial
which prevents the pinnule from attaining a vertical atti-
tude. The first segment of the pinnule occupies an indented
notch on the inner edge of the brachial.

The name contentus has its origin in the Latin word
tensus meaning spread out.

Measurements of holotype in millimeters:

Width of cup (estimated) 22.0
Height of cup (estimated) 17.0
Length of basal 7.0*
Width of basal 7.0*
Length of radial 6.5*
Width of radial 11.4*
Length of anal X Tei
Width of anal X 4.0
Diameter of proximal] columnals 2.8

Remarks.— The only described species of Cymbio-
crinus that even approaches the size of C. contentus is C.
gravis Strimple from the Fayetteville Shale, Chesterian,
northeastern Oklahoma. An undescribed species from the
Imo Formation, uppermost Chesterian, Arkansas is of com-
parable size but has relatively smaller arms.

Holotype —USNM, No. $5131, collected by Wm. T.
Watkins, reposited in U.S. National Museum.

Occourrence. — Marble Falls Formation, Morrowan?,
Pennsylvanian; three miles southwest of Lampasas, Lam-
pasas County, Texas on Espey Creek.

Genus LECOBASICRINUS Strimple and Watkins, new genus

Type species. — Lecobasicrinus kickapooensis, new
species.

Diagnosis. — Dicyclic; crown elongate, subcylindrical.
Arms ten, uniserial, branching with second primibrach in all
rays. Dorsal cup shallow, base subhorizontal or mildly con-
cave. Five small IBB, five small BB, five large RR, one
anal plate in cup followed by more than one tube plate
above. Column circular in outline to mildly pentagonal.

Range — Atokan to Missourian, Pennsylvanian; Mid-
Continent region, North America.

Remarks. — The most readily discernible characteristic
of this genus is the tendency for and the practice of the
radials to meet the infrabasal plates. The anal plate is small
and does not extend appreciably above the summit of the
cup. The only species for which the entire arm structure is
known is Lecobasicrinus kickapooensis of Desmoinesian age.

In L. kickapooensis some of the arms are hypertrophied,
but this is not necessarily of generic significance. The ten-
dency for the arms to be closely apposed is thought to be
significant because in related Oklahomacrinus and Aesio-
crinus the arms are usually subhorizontal in attitude.

Aesiocrinus typically has a pentagonal stem, a basal
concavity, and erect lateral cup walls. Oklahomacrinus
typically has a pentagonal stem, an exaggerated basal cav-
ity, and the cup walls are subhorizontal. We have not ob-
served any tendency for the radials to contact the infra-
basals in any species assigned to these genera except for
Delocrinus convexus Strimple, Lecobasicrinus subidus, n. sp.,
and Aesiocrinus paucus Strimple. It was recognized by
Strimple (1940a) that the form described as Delocrinus
convexus was atypical of Delocrinus. Assignment was not
made to Aesiocrinus because of the large round column.
Moore and Plummer (1940) have considered the form as
Aesiocrinus. It is referred by us to Lecobasicrinus with some
reservation.

Aesiocrinus paucus is a unique form of Missourian age.
It has a large slightly pentagonal attachment scar. It has
a large anal plate, not extending above the summit of the
cup and obliquely placed on the posterior basal. The dorsal
cup is shallow but not so exaggerated as in typical Ok/a-
homacrinus. Aestocrinus paucus has the strong tendency
for the radial to contact infrabasal with the subsequent
change in size and shape of basal plates, although actual
contact is not made in all rays. It is referred to the genus
Lecobasicrinus herein with some reservation. The tendency
for radials to contact infrabasals is found in genera of sey-
eral other phyletic lines, e.g. Protencrinus Jaekel, 1918,
Schistocrinus Moore and Plummer, and Sciadiocrinus Moore
and Plummer.

Species assigned to Lecobasicrinus:

Desmoinesian: Texas
(type species)
Morrowan: Oklahoma
Atokan: Texas
Missourian: Oklahoma

Lecobasicrinus kikapooensis, n. sp.

Delocrinus convexus Strimple
Lecobasicrinus subidus, n. sp.
Aesiocrinus paucus Strimple
The name Lecobasicrinus is derived from the Greek

lekos for basin, basis for the base with the common ending
krinon for sea lily and has reference to the shallow, basin-
like cup.

LECOBASICRINUS KICKAPOOENSIS

Strimple and Watkins, new species
Plate 31, figures 5, 6; Plate 53, figure 4

This species is represented in the collections by three
crowns, two from the Brannon Bridge Limestone Member,
Millsap Lake Formation, and one from the Kickapoo Falls

192 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Limestone Member, Millsap Lake Formation. The largest
and best preserved specimen is from Kickapoo Falls and
is taken as the holotype.

The species exhibits a characteristic that is relatively
rare among crinoids which is hypertrophy of certain arms.
Springer (1926) attempted to ascribe this condition to all
species of Agassizocrinus, but his premise was rejected by
Kirk (1940a) with the establishment of the genus Anartio-
crinus Kirk, That genus has remarkable hypertrophy in
half-rays in the right and left antero-lateral radii, affecting
both width and length of the arms. These are entirely un-
related genera, belonging to another phyletic lineage.

The dorsal cup is shallow, basin-shaped, with a shallow
basal concavity. The five IBB are small plates mainly
covered by the proximal columnals. The five BB are only
slightly larger than the IBB and are triangular-shaped ex-
cept in the posterior which is truncated for reception of the
single anal plate. The anal plate is followed by three tube
plates above the cup. There are four facets present but the
two proximal facets meet at a wide angle. It is probable
that the basals were not visible in side view of an undis-
torted cup. The five radials are the dominant cup plates,
are slightly wider than long, make contact with the IBB in
most rays, and have mildly gaping sutures at junction with
the primibrachs.

There are two primibrachs with the suture well de-
fined. In combination they have a more or less hour-glass
shape. Subsequent brachials are more or less regularly quad-
rangular-shaped. Along the upper edge of most brachials
there is a row of small nodes. In the hypertrophied arms,
there is a pronounced thickening with marked longitudinal
curvature of the outer surfaces.

A long section of the column is preserved in a small
paratype, but it is distorted by lateral compression, It has
a circular outline. One would expect some evidence of angu-
lation to be preserved if the stem were pentagonal, but
there is a possibility that it has been obliterated. A few
proximal segments are also preserved with the holotype and
they also appear to be of a circular nature. In proximal
portions the column is apparently composed of alternatingly
expanded segments but at a short distance from the cup
they are seen to be arranged in series of five. The largest
segments are followed by thin segments and one slightly
larger internodal is present midway between. The larger
columnals have longitudinally curved lateral sides that are
almost keel-like.

The crown of the holotype attained a length of more
than 53.7 mm and the smallest paratype of more than 14.7
mm. The specimens are flattened by lateral compression so

that accurate measurements are not possible. The height
of the cup of the holotype could not have been more than
5 mm.

Other measurements of holotype in millimeters:

Length of radia] (anterior) 4.7
Width of radial (anterior) 6.5
Length of basal (right posterior) 41
Width of basal (right posterior) 4.1

Differential in widths of arms is reflected by the
following measurements at slightly above mid-length of the
arms:

Right anterior —left ray Y/
—right ray aut
Anterior —left ray 2.0
—right ray 2.0
Left anterior —left ray 4.0
—right ray 3.9
Left posterior —left ray 2.8
—right ray 2.1
Right posterior —left ray 3.2
—right ray 3.5

Remarks. — Comparison of Lecobasicrinus kickapoo-
ensis with related species has been covered under the generic
discussion. As far as now known, the hypertrophied arms
are a specific characteristic,

Types. — Holotype, USNM, No. S5100, paratype,
USNM, No. $5185 and unnumbered paratype, collected by
Wm. T. Watkins.

Occurrence. — Holotype, shale below Kickapoo Falls
Limestone Member, Millsap Lake Formation, Strawn
Group, Desmoinesian; about one-third mile below the falls
on Kickapoo Creek, southwest of Dennis, Hood County,
Texas. Paratypes, Brannon Bridge Member, Millsap Lake
Formation, Strawn Group Desmoinesian, Pennsylvanian;
three miles southwest of Brock, Parker County, Texas.

LECOBASICRINUS SUBIDUS
Strimple and Watkins, new species

Plate 56, figures 3, 4

This species is represented in the collections by one
specimen from the Marble Falls Formation, consisting of the
proximal columnals, dorsal cup and lower portions of the
arms. There is slight distortion due to lateral compressions
but the specimen is in good preservation. In its natural state
the cup would be low, basin-shaped, with a flattened base.
Five infrabasals form a large, centrally depressed disk and
recurve out of the depressed basal area so that their tips
are visible in side view of the cup. Five basals are of
medium size, are hexagonal-shaped (except at the pos-
terior where an extra facet is provided for reception of the
single anal plate), and have relatively short lateral sides.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 193

The five radials are the dominant cup plates, are penta-
gonal-shaped, and are wider than long. The right posterior
radial has a short side in contact with the posterior basal.
The single anal plate is a quadrangular-shaped element
that does not extend appreciably above the normal summit
of the cup. It appears to have a single facet above but
probably has two facets.

About seven, alternatingly expanded, thin columnals
are preserved. The stem appears to be circular but when
carefully viewed at a suture face it has a mildly pentagonal
outline. The lumen is round. Cirri are present.

First primibrachs are low, wide, and nonaxillary. The
second primibrach appears to be nonaxillary in most rays.
These are the only preserved brachials in the specimen,

Measurements in millimeters:

Holotype
Height of cup (as preserved) 3.2
Width of cup (estimated) 5.0

Length of basal plate fF
Width of basal plate 1.
Length of radial plate of
Width of radial plate 2.
Length of anal plate 1.
Width of anal plate 2.
Width of proximal] columnals 1.

WMA A ”& © 0 OO

Remarks. — Lecobasicrinus subidus differs from Missis-
sippian species of Ampelocrinus in having a basal area ex-
tending beyond the stem with only the tips of the infra-
basals recurved to be visible in side view of the cup. There
is a depressed central area in the infrabasal disk. It also
has a slightly pentagonal outline of the column of which
about six columnals are preserved. In Ampelocrinus the
column is round and the infrabasals rise evenly from the
columnar attachment The Missourian species Aesiocrinus
paucus Strimple and A. prudentia Strimple are closely re-
lated to Lecobasicrinus subidus. The infrabasals of Aesio-
crinus paucus are not visible in side view of the cup but
this is due to the structure of the cup and not to the attitude
of the infrabasals. A. paucus has a tendency to shorten the
sutures between basals, as found in Lecobasicrinus subidus.

The name subidus is Latin for sensible, having refer-
ence to the normal evolution from Mississippian forms.

Type.— Holotype, USNM, No. $5198 collected by
Wm. T. Watkins.

Occurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Limestone Formation, Atokan, Pennsylvanian; on
Rough Creek, 11 miles southeast of San Saba, San Saba
County, Texas.

Genus AESIOCRINUS Miller and Gurley, 1890
AESIOCRINUS ANNULATUS Strimple and Watkins, new species
Plate 31, figures 2, 3

One crown and an incomplete dorsal cup represent this

species. Lateral compression causes some difficulty in tak-
ing measurements and in making some observations.

The dorsal cup is low, bowl-shaped with a sharply
invaginated basal area. At the top of the basal concavity,
five moderately large infrabasals form a subhorizontal or
slightly downflared disk. Five large basals have proximal
portions curved sharply into the basal concavity, apparently
form a broad base to the cup and then curve upward to be
slightly visible in side view of the cup. Five radials are
only slightly wider than long, pentagonal-shaped, mildly
tumid with depressed sutures and have a broad outer-
ligamental area with a thin ligamental pit. The transverse
ridge is crenulated and although preservation is poor, the
inner articular area is seen to be relatively long. The anal
plate is narrow, does not extend above the summit of the
cup and apparently is only followed by one tube plate.

There are ten robust arms bifurcating with the second
primibrach in all rays. There is a close suture between the
two first primibrachs of a type usually termed syzygial. Al-
most complete anchylosis has taken place in some pairs.
First primibrach is large, quadrangular-shaped, and the
second primibrach is low, almost triangular-shaped albeit
there are two short lateral sides. The first secundibrach is
large but thereafter the brachials are low elements marked
by a longitudinal ridge in mid-length with even upper sur-
faces. The arms taper quickly with the proximal secundi-
brachs but thereafter taper slowly. They have rounded
exteriors. One arm shows the lateral side for a short distance
and pinnular facets are present on every other brachial.
In other words, the pinnules must alternate on each side
of the arm.

The column is large, round, and composed of alter-
natingly expanded columnals, There are longitudinal ridges
on the thickened columnals.

The species name annulatus is Latin for ringed, with
reference to the column.

Measurements in millimeters:

Holotype Paratype
Length of crown (as preserved) 46.0 _—
Height of cup 7.0€ 8.2
Width of cup 27.0¢ 25.0
Length of basal 9.0* 10.7*
Width of basal 8.8* 10.0*
Length of radial 7.0* 8.70)
Width of radial 11.6* 12.20)

¢ Estimated.
* Measurements taken along surface curvature.
(1) Left posterior radial.

Remarks.— This species is closely related to Okla-
homacrinus abruptus Strimple (196la) from the Des-

194 PALAEONTOGRAPHICA AMERICANA (VI, 40)

moinesian of Oklahoma with both species having large
tumid basal plates, relatively narrow, steep-sided basal
concavity and subhorizontal or mildly downflared, infra-
basals. The small narrow anal plate, followed by one tube
plate, of Aesiocrinus annulatus, separates it from other
species of Aestocrinus. Aesiocrinus typically has a broad anal
plate followed by two tube plates; however, such forms as
Aesiocrinus luxuris Strimple (1951d) and A. detrusus
Strimple (1951d) have the anal plate followed by one tube
plate. They are, however, atypical of Aesiocrinus. This
group of four species probably represents a lineage separable
from Aesiocrinus.

Type. — Holotype, USNM, No. S5099, and one un-
numbered paratype, collected by Wm. T. Watkins.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles northwest of Brock, Parker
County, Texas.

Genus OKLAHOMACRINUS Moore, 1939
OKLAHOMACRINUS FROSTAE
Strimple and Watkins, new species

Plate 34, figures 1, 2

A single well-preserved, albeit flattened, crown is the
only known representative of this species. The crown is
long but the dorsal cup is short. Five small infrabasals are
confined to the basal concavity. Five small basals form the
basal plane and parts of the walls of the basal concavity.
The distal tips are visible in side view of the cup. Five
radials are thin, wide, large elements. There is a single
anal plate within the cup followed above by a prominent
single series of tube plates. Another series of tube plates is
present well above the cup indicating the presence of a long,
slender, weak anal tube. Ten arms are long, slender, have
well-rounded exteriors and branch on the second primibrach
in all rays. The first primibrach is sharply tapered. Each
brachial above the axillary carries a single pinnule which
alternates with the adjacent brachials. Proximal columnals
are large, are mildly pentagonal-shaped but eventually be-
come rounded. Many cirri are present, as evidenced by
scars on the nodals. Most of the surface of the cup and arms
is granular.

Measurements of holotype in millimeters (estimates of
cup height and width are not attempted):

Length of basal
Width of basal
Length of radial

Width of radial
Length of anal

NUwwn
NCwoor

Width of anal 1.5
Length of first primibrach 1.5
Width of first primibrach 5:9

Remarks. — Oklahomacrinus frostae is the oldest re-
corded species of the genus and is apparently closely related
to Cymbiocrinus pitkini Strimple (1955) of Chesterian age.
The long slender anal plate, followed by one tube plate,
the wide, thin radial plates, long, slender, uniserial arms
and proximal pentagonal columnals of O. frostae are all
typical of Oklahomacrinus. The upper Desmoinesian species,
O. abruptus Strimple (1961a) is closely related to O. frostae.
The former appears to be slightly more rugose, has more
prominent basal plates, and the anal plate widens somewhat
more than in O. frostae.

Lecobasicrinus kickapooensis, n. sp. and L. subidus, n.
sp. have many features in common with Oklahomacrinus
frostae but differ in having rounded proximal columnals.
Lecobasicrinus kickapooensis is also specialized in having
hypertrophied arms, and the single anal plate of the cup is
followed above by three tube plates, whereas Oklahoma-
crinus frostae has a single tube plate above the anal plate.

Holotype. —USNM, No. $5107 collected by Mrs. Paul
C. Frost, Cedar Hill, Texas, and reposited in the US.
National Museum.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; about three miles southwest of Brock,
Parker County, Texas.

OKLAHOMACRINUS SPICATULUS Strimple and Watkins,
new species

Plate 31, figure 4; Plate 51, figures 1, 4; Plate 53, figure 5;
Plate 56, figure 1

This species is represented in the collections by four
partial crowns from two exposures of the Lemons Bluff
Member, Marble Falls Formation, The preservation is not
satisfactory for some observations, but the specimens are
identifiable in essential features.

The cup is low and broad with shallow basal invagi-
nation. Infrabasals are confined to the invaginated base.
Five basals form the sides of the basal concavity, curve into
the basal plane, and the distal tips are in the lateral walls of
the cup. Five radials are the most prominent cup elements,
being wide and having decidedly tapered lateral sides.
There is a flattened area beyond the outer ligamental
furrow which probably acted as a stop for a similar though
less pronounced surface on the primibrach when the arms
were fully extended in a normal feeding attitude. The outer
ligamental pit is pronounced and the transverse ridge is
well developed above it but elsewhere the transverse ridge

‘TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 195

is ill defined. One anal plate is present in the cup but is not
well preserved in any specimen studied. The first primi-
brach is moderately low with the proximal edge filling the
width of the radial but tapering rapidly. The second primi-
brach is axillary with short lateral sides and a long slender
apex. The ten arms are uniserial, slender, with well-rounded
exteriors. Each brachial is slightly protuberant on alter-
nating sides for the attachment of the pinnule and give a
slight zigzag appearance which is atypical of Oklahoma-
crinus, or even late Chesterian Cymbiocrinus. The anal
tube is long, slender, and is comprised of large, smooth, ex-
ceedingly thin plates, with roughly serrated interlocking
lateral sides.

The entire surface of the cup and lower arm segments
are granular appearing. The column is pentagonal and cirri-
ferous.

Measurements in millimeters:

Width of cup—approximate 14.0
Height of cup—approximate 2.5
Width of radial 7.0
Length of radial 3.8
Width of first primibrach at base 6.9
Width of first primibrach at summit 5.0
Length of arms as preserved 18.5

Remarks.— The pentagonal-shaped stem serves to
separate Oklahomacrinus spicatulus from Cymbiocrinus and
the slightly staggered attitude of the brachials is different
from the arms of other species of Oklahomacrinus.

The name spicatulus is a derivative of the Latin word
spica for point and has reference to the pointed axillary
secundibrachs.

Types. — Holotype, USNM, No. S5186, paratypes
USNM, No. S5196 (two specimens), paratype USNM,
No. $5101, collected by Wm. T. Watkins, reposited in the
U.S. National Museum.

Occurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Formation, Atokan, Pennsylvanian; Rough Creek,
11 miles southeast of San Saba, and (paratype, No. $5101)
C. B. Lambert Ranch near San Saba, San Saba County,
Texas.

Genus PHIALOCRINUS Trautschold, 1879, nomen nudum

In the original description by Trautschold (1879) a dia-
gram was given with a single anal plate shown and two
primibrachs as Phialocrinus patens Trautschold. The genus
Phialocrinus Eichwald (1856) was based on only stems of a
considerably older age, and although pentagonal stems are
involved in both instances, the forms are certainly not
congeneric. A generic description was given by Trautschold
based on his material.

To further confuse matters, Trautschold gave illustra-
tions (1879, pl. 4, figs. 4 a-b) supposedly two views of the
same specimen, but the side view shows one anal plate and
the basal view shows three anal plates. This could easily
have been a drafting error because the shape of the cup
appears comparable in both instances, but in any event it is
imperative to establish the single anal plate character for
the species. Most modern authors have considered the
species to belong to Aestocrinus. Kirk (1942), in discussion
of the family Ampelocrinidae, noted, “Phialocrinus Traut-
schold obviously belongs here, but that name is a homonym,
If adequate material were available for study, that form
might prove to be a new genus or to belong to Aesiocrinus.”
New material has been illustrated by Yakovlev and Ivanov
(1956, pl. 7, figs. 3-5) and it (Phialocrinus patens) is seen
to be closely related to Aesiocrinus, and was so designated
by Yakovlev and Ivanov.

The species Aesiocrinus ivanovi Yakovlev (1956)
appears to us to be close to Aesiocrinus patens and could
well be a young stage of that species. Both are from the
Moscovian, C» which is apparently equivalent to the Ameri-
can lower Desmoinesian.

The form described as Phialocrinus? elenae Yakovlev
(1930) is illustrated as Phialocrinus elenae (Yakovlev) in
Yakovlev and Ivanov (1956), but there does not appear
to be any discussion of the species in the latter work. It
is closer in structure to Polusocrinus than to the other Rus-
sian species and is here referred to Polusocrinus elenae
(Yakovlev), new combination. The age is Permian, P;. The
infrabasals are large and prominent but are only slightly
visible in side view of the cup. The articular facets are
short and the stem is diminutive. As noted elsewhere, a
tendency toward atrophy of the column is found in some
species of Polusocrinus.

Family CORYTHOCRINIDAE Strimple and Watkins,

new family
Range. — Mississippian and Pennsylvanian; North
America.
Genera. — Corythocrinus Kirk, 1946; Araeocrinus

Strimple and Watkins, n. gen.

Diagnosis. — Dicyclic; cup high conical to low saucer-
shaped; IBB five, visible from side; R facets wide, may or
may not occupy full width of radial; one large anal (RA)
in cup in line with RR, in oblique contact with posterior
basal; anal sac tall; arms uniserial, pinnulate, usually
branching two or more times isotomously, syzygial pairs
formed by brachials; thin alternatingly thickened colum-
nals, with pronounced interlocking crenulations visible from
side, stem large with large lumen.

196 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Remarks. — This unique group is characterized by hav-
ing a single, large anal plate (RA) in broad, oblique con-
tact with the posterior basal and also a narrow contact
with the right posterior basal. In arm structure, there ap-
pears to be a migration of the point of branching away from
the cup, which is diametrically opposed to previously ac-
cepted concepts concerning this feature. It has been thought
that migration of the point of branching always progresses
downward toward the cup. An exception is noted herein
for Ulrichicrinus ramosus wherein regression has taken
place. In typical Corythocrinus there are three primi-
brachs and about ten to 16 secundibrachs, but in Araeo-
crinus, which is considerably younger, there are at least
four primibrachs, and at least 30 secundibrachs prior to a
second bifurcation.

In Corythocrinus the articular facet of the radial plate
reputedly occupies the full width of the plate but in Araeo-
crinus the outer surface of the radial is projected into the
adsutural portion of the upper surface. The anal plate of
Corythocrinus, as illustrated by Kirk (1946, pl. 40, fig. 1),
has three distinct distal facets although it is stated that only
tube plates followed. In Araeocrinus there appears to be only
one facet which has the general appearance of a radial plate
even to the presence of an outer ligamental pit. The pit
is in itself unusual in having a mild rise in mid-portion so
that under close scrutiny there appear to be two pits. The
horizontal plane formed by the upper surface of the anal
plate in Araeocrinus is distinctly different from the sharply
angulated upper extremity of the anal plate in Corytho-
crinus.

The affinities of the corythocrinids are somewhat ob-
scure except for apparent relationship with Ampelocrinus
of the Ampelocrinidae. We believe Ampelocrinus may be a
derivative of a form like Corythocrinus insculptus Kirk,
1946, through slight reduction in cup height, elimination of
one primibrach and reduction in the size of the stem. They
have in common the general shape of cup (conical, with
IBB visible in side view); long, slender, uniserial arms with
cuneiform segments and syzygial pairs, which branch two
or more times isotomously; long slender anal sac; a single
anal plate (RA) resting obliquely on the posterior basal.
As a matter of record, we do not believe all forms cur-
rently assigned to the ampelocrinids to be directly related
to Ampelocrinus.

Kirk (1946, p. 269, 270) did not consider Corythocrinus
to be closely related to any described genus but he gave
a discussion of differences between it and Lebetocrinus Kirk,
1940. Fundamental differences appear to exist in the nature
of the arms of those two genera.

Genus ARAEOCRINUS Strimple and Watkins, new genus

Type species. — Araeocrinus bassus Strimple and Wat-
kins, n. sp.

Range. — Pennsylvanian; North America.

Diagnosis. — Dicyclic; cup high conical to low saucer-
shaped; IBB five, visible from side; R facets wide, do not
occupy full width of radials; one large anal (RA) in line
with RR, in oblique contact with posterior basal; anal sac
tall, composed of thin plicate plates; arms uniserial, pinnu-
late, branching two or more times isotomously, brachials
cuneiform, more or less hour-glass shaped, syzygial pairs
common; stem large, columnals thin, alternatingly expand-
ed, circular outline in older species, may be pentagonal
in younger species.

Remarks.— Comparison between Corythocrinus and
Araeocrinus is given under the familial remarks section. The
two genera appear to be closely related.

The name Araeocrinus is derived from the Greek word
araios for narrow or thin, having reference to the long
slender crown,

ARAEOCRINUS BASSUS Strimple and Watkins, new species
Plate 42, figure 3; Plate 51, figures 2, 3; Plate 53, figures 1, 2

This species is represented in the collections by two
crowns in excellent preservation and a dorsal cup. The
smaller crown does not show any juvenile characters (which
may be confirmed by comparing ratios between length and
width of various plates) and is selected as the holotype.
The crown is long and slender, the cup is tall and conical,
sutures between radials and primibrachs are gaped, there
are 20 or more arms with numerous syzygial pairs of
brachials and the anal sac is tall and slender.

There are five infrabasals that rise evenly from the
columnar attachment area. Five basals are prominent cup
elements and are hexagonal, even in the posterior. The pos-
terior basal supports the large quadrangular anal plate
above on an oblique plane and does not have an extra
facet because it has lost contact with the right posterior
radial plate. There is a truly unique situation wherein the
right proximal corner of the anal plate (RA), the left
proximal corner of the right posterior radial, the right dis-
tal corner of the posterior basal and the left distal corner
of the right posterior basal have a common meeting point.
The arrangement leaves the right posterior radial plate with
only four sides. Other radial plates have five sides. The
articular facet does not occupy the full width of the radial
plate, but the external surface of the plate is seen to ex-
tend into the adsutural area and form a low shoulder.
When the primibrach is in place this feature is somewhat

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 197

obscured. The extension of the outer surface of the radial-
like anal plate (RA) onto one adsutural slope of the upper
facet is well shown on one corner of the large paratype and
is reflected by the lack of sharp distal corners of the holo-
type. Such a condition emphasizes the similarity of the anal
plate with the radial plates. There is even an outer liga-
mental furrow, albeit the pit is composed of two depres-
sions,

There are four or five primibrachs in each ray. They
are quadrangular with even upper facets except for the
axillary which has a bluntly pointed apex. The primibrachs
have a rounded surface but they recurve along the sides
so that flattened areas are actually formed. Secundibrachs
are usually syzygial paired, cuneiform-shaped and may bi-
furcate with about the thirtieth segment. Sutures between
segments of a pair are close but between pairs they are
gaped. Each syzygial pair has a single pinnule on a side,
alternating with the adjacent pair, and has a flared upper
and lower edge.

The anal sac is long and slender, composed of thin,
wide, hexagonal-shaped plates. Each series of plates is
rounded but recurve to produce flattened sides which are
marked by raised ridges which are confluent with ridges
of adjacent plates. There is usually one series of plates to
a side although it may sporadically bifurcate to meet those
of two adjacent plates. The raised median portion of the
tube plates is granular appearing. There are apparently
five series of plates.

Ornamentation of the anal tube is discussed above.
The dorsal cup is also covered with small granulations.
The arms are marked by fine ridges which bifurcate and
merge as they pass from brachial to brachial.

The column is large, circular in outline, tapers sharply
for a short distance from the cup, but thereafter no ap-
preciable tapering takes place, at least for a considerable
distance. The segments are alternatingly thickened but are
essentially thin elements. Crenulations extend to the ex-
terior giving an interlocking appearance to the sutures. The
lumen is prominent and pentagonal-shaped. Less than one-
half inch of the column is attached to the holotype out of
a total of three feet recovered from the ledge. The column
continued into the cliff. This is the only species found at
this locality with a long column preserved.

Measurements in millimeters:

Holotype Paratype
Length of crown 41.03 132.0¢
Width of crown 17.0 44.0
Height of IBB circlet 2.3 4.6
Width of IBB circlet 4.2 8.6
Length of basal plate 3.2 7.1

Width of basal plate 4.1* 8.0*

Length of suture between basals 1-8
Length of radial plate 2.8 4.8
Length of anal plate (RA) 3.1
Width of anal plate (RA) a7
Length of arms to apex of PBriax,

left posterior ray 6.0 —
Diameter of proximal columnal 3.0 4.8

i— incomplete

© — estimated

* — measurement taken along surface curvature.

Remarks. — There are few described forms with which
Araeocrinus bassus may be closely compared. Differences
with the closely related Corythocrinus have been given un-
der the familial discussion. The slender arms with numerous
primibrachs and syzygial pairs of brachials separates it from
other known species. Lecobasicrinus paucus (Strimple) from
the Missourian has a shallower cup with infrabasals essen-
tially horizontal although the tips are curved up to be
visible in side view of the cup. As noted above, Aracocrinus
bassus has a high cone-shaped cup and the infrabasals rise
evenly from the stem attachment area.

The species name bassus is Latin for deep, having refer-
ence to the deep dorsal cup.

Types. — Holotype, USNM, No. $5183 and paratypes
USNM, No. $5138, $5175, $5176 collected by Wm. T. Wat-
kins and reposited in the U.S. National Museum.

Occurrence. — Lemons Bluff Member, Marble Falls
Formation, Atokan, Pennsylvanian; on Rough Creek, 11
miles southeast of San Saba, San Saba County, Texas.

Family SCYTALOCRINIDAE Moore and Laudon, 1943
Genus SCYTALOCRINUS Wachsmuth and Springer, 1880
SCYTALOCRINUS SANSABENSIS Moore and Plummer, 1938
Plate 41, figure 5; Plate 51, figures 1, 4

Synonomy.— Scytalocrinus sansabensis Moore and
Plummer, 1938; Scytalocrinus sansabensis Moore and
Plummer, 1940; Scytalocrinus sansabensis Bassler and
Moodey, 1943; Hypselocrinus sansabensis Moore and Lau-
don, 1944.

The species have been well described by Moore and
Plummer, 1938, and no appreciable amount of additional
information has been uncovered in the excellently preserved
material at hand.

Kirk (1940b, p. 327) considered the high and prom-
inent infrabasal plates of Hypselocrinus of considerable
diagnostic significance, and designated the characteristic as
of use in distinguishing the genus from Scytalocrinus. He
considered the arms of Scytalocrinus to be short and stout,
as compared to the long, relatively slender arms of Hypselo-
crinus. The importance of length of arms was accentuated

by Kirk (1940b, p. 325) when he noted the relative length

198 PALAEONTOGRAPHICA AMERICANA (VI, 40)

of arms of an adult specimen of the type species (Hypselo-
crinus hoveyt) as 25 cm long with a cup height of 1.7 cm,
a ratio of about 0.06. A dramatic difference is afforded in
comparison with specimens of Scytalocrinus sansabensts,
wherein the ratio is about 0.20. It is probably this differ-
ence between cup height and arm length that influenced
Bassler and Moodey (1943) to retain the later species in
Scytalocrinus. It is likely that Kirk was consulted in the
matter. The nature of the infrabasal circlet, i.e. its upflared
nature, apparently influenced Moore and Laudon (1944)
in their assignment as Hypselocrinus sansabensts.

It is not within the scope of this paper to completely
review species of the genera Scytalocrinus and Hypselo-
crinus. The latter genus is usually a more delicate, special-
ized form within which most species have an autonomous
anterior ray, and a tendency toward a second bifurcation
of the arms in later species. We are cognizant of the pos-
sibility that an undescribed genus is represented but for
the present designate the species as Scytalocrinus sansa-
bensis.

We are able to clarify some characteristics. The anal
plates of the posterior interradius are perfectly normal
(Primitive Type) in one of the excellently preserved hypo-
types available to us. Some of the drawings given by Moore
and Plummer (1938, text-fig. 9; 1940, text-fig. 17b) show
anal X to have two distal facets and RX (rt) to have only
one distal facet. Some of the drawings of Moore and Plum-
mer (1938, text-fig. 9; 1940, text-fig. 17a) show anal X has
a single facet above and RX (rt) has two distal facets. Our
specimen shows the anal X with one distal facet and RX
with two distal facets, one in contact with X, which is the
common arrangement among inadunate crinoids having
three anal plates within the cup. There are two known
Pennsylvanian genera having tall conical cups upflared in-
frabasals which are readily distinguished from Scytalocrinus
sansabensis simply because the anal X and RX have a con-
fluent distal edge which is in a subhorizontal attitude
(Hydriocrinus and Melbacrinus). There are other differences
involved.

The pinnules of S. sansabensis occur on alternating
sides, The pinnular bearing side of a brachial has a slight
tendency to be longer than the nonpinnular side but de-
cidedly cuneiform brachials are not formed. The pinnules
are large and relatively long.

Types. — Holotype, Walker Museum, No. 31721, para-
type, Walker Museum, No. 31721a, University of Chicago,
(now Field Museum of Natural History) collected by R. C.
Moore. Hypotypes, Peabody Museum, No, 15238 A-E, Yale
University, collected by Charles Schuchert, D. K. Greger,

and A. McCoy. Hypotypes, USNM, Nos. $5135, $5173 col-
lected by Wm. T. Watkins.

Occurrence. — Holotype, paratype, and hypotypes are
from the Lemons Bluff Limestone Member, Marble Falls
Formation, Atokan, Pennsylvanian; in valley of Rough
Creek, about 11 miles southeast of San Saba, San Saba
County, Texas.

SCYTALOCRINUS sp.
Plate 50, figure 5

One specimen, in poor preservation, is present among
the crinoids found in the Barnett Formation of Mason
County, Texas, which appears to belong to the genus
Scytalocrinus in that it has a high cone-shaped cup with
upflared infrabasals. The arms are unusually thick and are
only comparable to those of an undescribed species from
the Pitkin Limestone of Arkansas. The brachials are low
and the pinnules are well developed.

The dorsal cup has a height of 13 mm, width of 29 mm.
The arms have a width of 6.5 mm at a distance of 64 mm
above the cup with no evidence of tapering. The columnar
attachment area has a diameter of 5 mm.

Figured specumen.—USNM, No. 85172 collected by
Wilburn Shearer, reposited in the U.S. National Museum.

Occurrence. — The limestone slab was obtained in what
is thought to be the Barnett Formation, Chesterian, Mis-
sissippian; Kothman Ranch 214 miles southwest of Mason,
Mason County, Texas.

Genus HYDRIOCRINUS Trautschold, 1867

HYDRIOCRINUS LORRAINEAE Strimple and Watkins,
new species

Plate 34, figures 3, 4, 6, 7

Dorsal cup elongate, cone-shaped. Five infrabasals
prominent in side view of cup. Five basals are elongated.
Five radials are slightly wider than high with distal faces al-
most filled by proximal edges of primibrachs. There are
three anal plates but the area is disturbed in preservation
and their exact arrangement is not known. Anal X is in
direct posterior position and has a broad contact with the
posterior basal. It does not appear to be larger than RA.
There are ten cuneiform arms with the first primibrach
axillary in all arms. The first primibrach is constricted in
mid-section. Secundibrachs have a short and a long side,
with the long side pinnular bearing. The exteriors are well
rounded and there is no median constriction. The arms
were probably long.

There are two proximal columnals of medium size in
place which have a mildly pentagonal outline and a round
lumen.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 199

Measurements of holotype in millimeters:

Length of crown as preserved 43.4
Height of dorsal cup 12.0
Width of dorsal cup 13.0
Height of infrabasal circlet 4.0*
Width of infrabasal circlet ae
Length of basal (right anterior) 5.1
Width of basal (right anterior) 4.2
Length of radial (right anterior) 4.0
Width of radial (right anterior) 5.6
Length of primibrach—average 4.2
Width of primibrach—at base 5.3

*slightly distorted due to lateral compression

Remarks.— The shape and length of the cup of
Hydriocrinus lorraineae is close to that of H. pusillus Traut-
schold (1867) of the Moscovian of Russia. The primibrachs
and arms of H. lorraineae are more robust than those of
H. pusillus.

Several species have been assigned to the genus Hydrio-
crinus by Schmidt (1930) from Mississippian or Upper
Devonian rocks which do not belong to the genus or do not
exhibit characters of the type species of the genus, H. pusil-
lus. Species from the Lower Carboniferous of England are:
Poteriocrinus barumensis (Whidborne), Mariocrinus ? mun-
dus Whidborne, Scaphiocrinus ? plumifer Whidborne,
Scaphiocrinus salebrosus Whidborne, Poteriocrinus tensus
Whidborne, and Scaphiocrinus transcisus Whidborne.
Schmidt (1930) selected the form described by Drever-
mann (1902 from “Etroeungt” (transitional beds in Upper-
most Devonian) of Ratingen, Kettwig, Rheinland as
Poteriocrinus barumensis and formed a new species, Hydrio-
crinus ratingensis Schmidt. A specimen studied by Schmidt
had three anal plates in normal position (Primitive) and
nonaxillary first primibrachs in some rays.

Hydriocrinus ? rosei Moore and Plummer from the
Morrowan of Oklahoma has three anal plates in normal
(Primitive) arrangement. The species is not typical of
Hydriocrinus but may well be the progenitor of the genus.
The arms of H. ? rosei are not known so that no firm de-
termination may be made.

The excellent crown taken as the holotype was donated
to the study by Mrs. Lorraine Marrs and is named in her
honor.

Types. — The holotype USNM, No. $5108, paratype,
USNM, No. $5110, reposited in the U.S, National Museum.

Occurrence.— The holotype is from the Brannon
Bridge Limestone Member, southwest of Brock, Parker
County, Texas, and the paratype was collected by Wm. T.
Watkins from the Kickapoo Falls Limestone Member, near
Kickapoo Falls, Hood County, Texas, Millsap Lake Forma-
tion, Strawn Group, Desmoinesian, Pennsylvanian.

Family RAMULOCRINIDAE Strimple and Watkins, new family

Genera. — Ramulocrinus Laudon, Parks, and Spreng;
Aulocrinus Wachsmuth and Springer; Decadocrinus Wachs-
muth and Springer; Trautscholdicrinus Yakovlev and lvan-
ov.

Range. — Devonian to Pennsylvanian; North America,
SSSR.

Diagnosis. — Dicyclic; crown slender; widely expand-
ed cup; IBB five, visible from side in older forms but may
be subhorizontal in younger genera; BB five, medium size;
RR five, facets wide; three XX in cup; anal sac tall, slender;
arms branch isotomously on PBrsax in all rays in older
forms but on PBr,ax in younger forms and do not branch
again; sinuous zigzag arm brachials; stout pinnules,

Remarks.— This family is proposed for a group of
closely related forms that are removed from the Scytalo-
crinidae. All have in common the sinuous zigzag arm brach-
ials and stout pinnules. Devonian species not having zigzag
arm brachials fall under the genus Bridgerocrinus which
genus is retained under the Scytalocrinidae. Decadocrinus
has two PBrBr whereas Ramulocrinus and Aulocrinus have
a single PBr. Migration of the point of bifurcation toward
the cup is considered to be progressive evolution. The range
of Ramulocrinus is extended into the Missourian (upper
middle Pennsylvanian) in this study.

Aulocrinus is a specialized form with short brachials,
a distinctive spoutlike anal tube and a low bowl-shaped
cup with a concave base.

The inception of this family appears to be with such
forms as Decadocrinus nereus (Hall) and D. multinodosus
Goldring of Devonian age.

Trautscholdicrinus is the youngest genus (C3ks, Upper
Carboniferous of Russia and Missourian of North America)
and does not have the keeled appearance and a lessened zig-
zag appearance to the arms.

Genus RAMULOCRINUS Laudon, Parks, and Spreng, 1952

Type species.— Ramulocrinus nigelensis Laudon,
Parks, and Spreng, 1952.

Range. — Devonian, Mississippian, Pennsylvanian;
North America.

By original definition, this genus is a Decadocrinus
with a single primibrach. Both genera (Decadocrinus and
Ramulocrinus) as redefined by Laudon, Parks, and Spreng
(1952) have IBB that curve upward to form part of the
lateral walls of the cup and have sinuous, zigzag appearing
brachials but Decadocrinus has two primibrachs, the second
being axillary. There is no evolution reflected by the anal
plates of the posterior interradius (i.e. there is no evidence

200 PALAEONTOGRAPHICA AMERICANA (VI, 40)

of migration and elimination or elimination of said plates)
and the reduction of primibrachs from two to one is con-
sidered to be progressive. Strangely enough there is still no
change in the arrangement of the three anal plates of the
posterior interradius in the new Desmoinesian species,
Ramulocrinus consectatus, n. sp., or in the zigzag appear-
ance and large pinules (?ramules) of the arms. The base
of the new species is disorganized in preservation but ap-
pears not to have been concave.

RAMULOCRINUS CONSECTATUS Strimple and Watkins,
new species

Plate 44, figure 13

This species is represented in the collections by a single
spectacular crown from the Brannon Bridge Limestone
Member of the Millsap Lake Formation. The posterior side
is in excellent preservation. The proximal columnals are
preserved but the base of the cup is distorted. There does
not appear to have been any basal concavity but the base
is probably subhorizontal rather than upflared.

The dorsal cup is widely expanded and shallow. The
infrabasals are covered by other plates. The five BB are
large and their proximal portions are not appreciably
curved, which indicates the lack of a basal concavity. Five
RR are large, pentagonal-shaped, flared, and marked by a
low but distinct arcuate ridge below the outer ligamental
area. Three large anal plates, in normal (Primitive) ar-
rangement, occupy the posterior interradius. Sutures be-
tween the radials and primibrachs are slightly gaped.

Primibrachs are axillary and are in general elongated
although there is considerable variance in their lengths.
All but the left anterior are preserved. The longest is the
anterior with the right and left posterior only slightly
shorter. The right anterior is the short element and it is
almost certain the missing left anterior would be of com-
parable length. The plates are constricted but do not flare
appreciably at their upper end. There is a median ridge,
and the granulose surface of the cup continues onto the
lower arms. Only ten arms are developed. BrBr are slightly
elongate, keeled, mildly constricted in mid-portion with
each plate projecting in a direction opposed to the succeed-
ing plate. Each brachial has a stout pinnule located on a
sharply projected area with the facet directed about 80 de-
grees from the main axis of the arms. The pinnules are
long, keeled, and robust. A markedly sinuous appearance is
attained by the arms.

A few lower plates of the anal sac are preserved indi-
cating that the sac is more or less rugged appearing. Three
badly crushed proximal columnals are preserved, They ap-

pear to have a circular outline, are thick and have an
ornate exterior.

Measurements of the holotype in millimeters:

Length of crown (incomplete) 27
Height of cup (distorted) 3
Width of cup (distorted) 7
Length of basal 2
Width of basal 3
Length of radial 2
Width of radial 4.7

Remarks. — This species is readily separable from any
described Pennsylvanian crinoids because of the exaggerat-
ed sinuous zigzag appearance of the brachials. It has more
pronounced pinnules (?ramules) than typical for the genus
and the girdle-like ridge of the radial plate appears to be a
specific characteristic. Ramulocrinus repertus (Miller and
Gurley) from the Keokuk of Indiana has mildly stellate
plate markings.

The name consectatus is from the Latin word consector
for “imitate” or “relative to.”

Holotype. —USNM, No. $5152 collected by Wm. T.
Watkins, reposited in the U.S. National Museum.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; about three miles southwest of Brock,
Parker County, Texas.

Genus TRAUTSCHOLDICRINUS Yakovlev and Ivanov, 1939

Synonymy. — Trautscholdicrinus Yakovlev and Ivanov,
1939 (not Trautscholdicrinus Ivanoy, 1926; not Traut-
scholdicrinus Moore and Plummer, in Moore, 1939).

Type species. —Trautscholdicrinus miloradoivitscht
Yakovlev, 1939.

Range. — C3, Upper Carboniferous; Missourian, Penn-
sylvanian: SSSR., U.S.A.

Diagnosis. — Cup truncate, cone-shaped with five IBB
not visible in side view but not downflared. Five BB are
large plates with proximal tips curved into basal area but
mainly erect as part of the lateral walls of the cup. Five
RR pentagonal-shaped, slightly wider than high, and have
a gaped suture with primibrach. Three anal plates in nor-
mal (Primitive) arrangement. Five PBrBr,ax with slightly
constricted mid-sections. Secundibrachs are slightly elongat-
ed, have well-rounded exteriors are cuneiform and mildly
constricted in mid-portion. They are not decidedly zigzag
appearing but each brachial meets the next at an angle so
they diverge alternatingly from the ray axis. In the upper
portions of the arms the pinnules are well preserved and are
proportionately large elements in this region, one on each
brachial on alternate sides. The anal sac is mainly com-

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 201

posed of small hexagonal plates in interlocking series and
appears to have been about as long as the arms.

Remarks. — Yakovlev and Ivanov (1939) apparently
considered Trautscholdicrinus Ivanov (1926) a nomen nu-
dum. Bassler and Moodey (1943, p. 714) considered Ivan-
ov’s genus a nomen nudum. As discussed elsewhere the name
Trautscholdicrinus Moore and Plummer (1939) was ap-
plied to a flexible crinoid and has been placed in syn-
onymy.

Yakovlev and Ivanov (1956) compared Trautscholdi-
crinus Yakovlev and Ivanov (1939) with Liparocrinus
Goldring (1923) with emphasis on the nature of the anal
sac. The sacs do not appear to us to be closely comparable,
at least no more so than with several other genera. We be-
lieve Aulocrinus is a more closely related genus in that it has
a shallow cup with a mild basal concavity and the arms
branch one time with the first primibrach in all rays.

The species Decadocrinus regularis Strimple (1939a)
has a flared-shape to the cup, is a smooth form and the
secundibrachs have a mildly zigzag pattern. It does not
have a basal concavity. Assignment is made here as Traut-
scholdicrinus regularis (Strimple), new combination.

The specimen figured by Yakovlev and Ivanov (1956,
pl. 6, fig. 3) as Trauwtscholdicrinus sp. is apparently mildly
distorted in preservation and shows an infrabasal plate
in side view of the cup. In other respects it appears to
us to be conspecific with 7. miloradowitschi.

Family BLOTHROCRINIDAE Moore and Laudon, 1943
Genus MICROCARACRINUS Strimple and Watkins, new genus

Type species. — Microcaracrinus delicatus Strimple and
Watkins, new species.

Range. — Desmoinesian, Pennsylvanian: North Amer-
ica.

Diagnosis. — Dicyclic; crown tall, slender; cup low,
bowl-shaped with broad basal concavity; IBB five, confined
to basal concavity; BB five, small, distal tips visible in
side view of cup; RR five, tumid, large, pentagonal-shaped;
XX three in normal (Primitive) arrangement; PBrBr oc-
cupy the full width of RR; PBr, axillary, elongated, slender
through constriction; BrBr slightly elongated with median
section constricted, a rimlike ridge marks the top and bot-
tom of each segment, and they are more or less sinuous or
zigzag appearing; SBrBr;-; axillary, each Br bears a stout
pinnule alternating with the adjacent BrBr; column round,
columnals alternatingly expanded.

Remarks. — This genus appears to be related to and is
probably a derivative of Cosmetocrinus Kirk, 1941. The

arms are more zigzag appearing than is typical of Cosmeto-
crinus. In Cosmetocrinus the IBB are visible in side view of
the cup but in Microcaracrinus they are confined to a basal
concavity. The arms branch isotomously on PBr; in both
genera and once or twice at higher positions.

The zigzag appearance of the arms of Microcaracrinus
is less pronounced than found in associated Ramulocrinus.
The later genus has a widely expanded cup and is limited
to ten arms as compared to the bowl-shaped cup and 20
arms of Microcaracrinus.

In general appearance the arms are somewhat like those
of Pelecocrinus but in that genus the articular facet of the
radial is narrower than the normal width of the plate. In
Microcaracrinus the facet occupies the full width of the
radial. Some authors consider the reduction in width of
radial articular facets as advanced or progressive, which
assumption would eliminate the possibility of the older
Pelecocrinus being related to Microcaracrinus. There is also
a specialization in Pelecocrinus in that several PBrBr are
present in the anterior ray.

The generic name is derived from a combination of the
Greek words mtkros, for little, kara, for head, and krinon,
for sea lily.

MICROCARACRINUS DELICATUS Strimple and Watkins,

new species
Plate 44, figure 12; Plate 46, figures 1, 3, 4

This species is represented in the collections by two
small crowns in good preservation, It is the type species
and only known species ascribed to Microcaracrinus so that
the generic description, previously given, applies to the
species as well. It is difficult to visualize what characteris-
tics will prove to be of specific value when other species are
found. This is a delicate form which will only be preserved
under optimum conditions.

There is no evidence of ornamentation or raised ridges
other than the thin ridges which parallel the sutures be-
tween BrBr. This serves to differentiate it readily from the
associated Ramulocrinus consecutatus in which the BrBr
are angulated by median ridges and the radial plates have a
transverse ridge below the outer ligamental area. It also
has a widely expanded cup as compared to the decidedly
curved lateral walls of the cup of Microcaracrinus delicatus.

There are 20, long, slender, uniserial arms with well-
developed pinnules and cuneiform brachials. Each brachial
bears a pinnule which alternates with the pinnules of
adjacent brachials. The parallel ridges marking the suture
between brachials is probably a specific characteristic.

202 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Measurements of the holotype in millimeters:

Length of crown 18.8
Height of cup 2.1
Width of cup (distorted) 5.3
Length of basal (approximate) 1.8*
Width of basal 1.6*
Length of radial 17
Width of radial 2.3*
Diameter of columnal (large) 0.7
Diameter of columnal (small) 0.6
Length of stem as preserved ey

* measurements taken along surface curvature.

Types. — Holotype, USNM, No. $5151, paratypes
USNM, No. $5159, No. $5158, § 5155, collected by Wm.
T. Watkins, reposited in the U.S. National Museum.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Genus CULMICRINUS Jaekel, 1918

Type species.— Poteriocrinus regularis H. v. Meyer,
1858.

Range. — Mississippian; North America, Germany,
England?,

The type species of the genus, Culmicrinus regularis, is
from the lower part of the Upper Viséan of Germany. Cul-
micrinus elegans is from the Glen Dean Formation, Ches-
terian (Upper Mississippian). Both have cone-shaped cups
with a prominent, elongated anal tube. The arms of C.
regularts do not branch until the eighth to tenth primibrach,
The latter species is apparently the younger.

C. missouriensis (Shumard) is from the St. Louis
Limestone at St. Louis, Missouri. The species reportedly
only branches once with the tenth or twelfth primibrach.
C. jeffersonensis Laudon and Severson, 1953, is from the
Lodgepole Formation, Kinderhookian, of Montana. It is
reported to branch for the first time on the eleventh or
twelfth primibrach.

C. barnettensis, n. sp. is from the Barnett Formation,
Chesterian, Texas, and is advanced in that the first branch-
ing is on the fourth primibrach.

C. thomasi Laudon from the Gilmore City Formation,
Kinderkookian, Iowa, branches on the fourth or fifth primi-
brach. It is thought to be too advanced to be in the direct
lineage of Culmicrinus but may be homogenous.

Culmicrinus? singulocirrus Schmidt, 1930, has a penta-
gonal stem and is from the “Etroeungt” near Ratingen,
Germany, which is a zone formerly thought to be transi-
tional between the Devonian and Mississippian but is now
considered to be Lower Mississippian. The arms may branch
with about the fourth primibrach but usually they branch

lower in the arm. The species is probably not in the lineage
of Culmicrinus,

Potertocrinus tensus Whidborne, 1896, was referred to
Culmicrinus by Schmidt (1930, p. 87) and retained in the
genus by Bassler and Moodey (1943), but the preserva-
tion is too poor in the lower portion of the arms to de-
termine the point of branching. The species is from the
Pilton and Marwood beds, Lower Mississippian, England
and probably does not belong in Culmicrinus.

In the direct lineage of Culmicrinus the migration
downward of the first arm bifurcation is apparently a re-
liable evolutionary characteristic. C. thomasi is atypical
in that the number of primibrachs has been reduced.

CULMICRINUS BARNETTENSIS Strimple and Watkins,

new species

Plate 50, figure 5; Plate 52, figure 12

Dorsal cup is high, cone-shaped with infrabasals readily
visible in side view. The proximal end of the cup is relatively
wide and the sides of the cup expand evenly to the summit,
The infrabasals are prominent, the basals of moderate size,
and the radials are large. There is provision for three anal
plates in normal (Primitive) arrangement. The articular
facet fills the entire width of the radial plate, and the suture
is slightly gaped. First primibrachs are four in a series, with
the fourth axillary. The column is large at its juncture with
cup and tapers rapidly for a short distance, thereafter
maintaining an even width. The columnals are alternatingly
expanded with interlocking crenulations visible in side view.

Measurements of the holotype in millimeters:

Length of crown, as preserved 24.0
Height of cup 6.0

Width of cup 6.1
Height of infrabasal circlet 2.1
Length of basal (posterior) 2.7
Width of basal (posterior) 2.8
Length of radial (left posterior) 2.1
Width of radial (left posterior) 31
Width of proximal columnal 2.6

Remarks.—C. barnettensis is the youngest Culmi-
crinus known and has evolved to the point of only having
four primibrachs. Laudon (1941, p. 370) was of the opinion
that Carinocrinus Laudon evolved from Culmicrinus.
Carinocrinus stevensi Laudon (1941) branches with the fifth
secundibrach. A form described as Carinocrinus eventus
Strimple (1953) branches with the first primibrach. It is
from the upper part of the Pitkin Formation and apparently
represents a rapid evolution in that C. stevens is also listed
as occurring in the Pitkin Formation; however, the horizon

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 203

from which the latter was obtained is considerably lower in
the section.

Elibatocrinus Moore (1940) from the Pennsylvanian
is probably a derivative of the lineage represented by
Culmicrinus barnettensis. The infrabasals have fused with
three remaining in Elibatocrinus.

Holotype. —USNM, No. $5180 collected by Wm. T.
Watkins.

Occurrence. — The holotype was found on a thin slab
1 1/4 inch thick, part of several thin layers with brachipods,
Archimedes, crinoids, and small starfishes preserved. The
horizon is below a conglomerate that marks the top of
the Barnett Formation, Chesterian, Mississipian. The loca-
tion is near the zone that produced the other Barnett
crinoids but is slightly higher in the section (Kothman
Ranch, near Mason, Mason County, Texas).

Family PIRASOCRINIDAE Moore and Laudon, 1943

Genera. — Zeusocrinus Strimple, 1961a; Paianocrinus
Strimple, 1951e; Stenopecrinus Strimple, 1961a; Plaxocrinus
Moore and Plummer, 1938; Metaperimestocrinus Strimple,
1961a; Schedexocrinus Strimple, 1961a; Polygonocrinus
Strimple, 1961a; Pirasocrinus Moore and Plummer, 1940;
Eirmocrinus, n. g.; Sciadocrinus Moore and Plummer, 1938;
Laudonocrinus Moore and Plummer, 1940; Lasanocrinus
Moore and Plummer, 1940; Athlocrinus Moore and Plum-
mer, 1940; Perimestocrinus Moore and Plummer, 1938,
Utharocrinus Moore and Plummer, 1938; Schistocrinus
Moore and Plummer, 1940; Aatocrinus Moore and Plum-
mer, 1940; Bathronocrinus Strimple, 1962a; Anchicrinus,
n. g.

Range. — Chesterian Mississippian to Lower Permian;
North America.

Remarks.—The family Pirasocrinidae, as emended
by Strimple (1962a, pp. 17-21), is restricted to the Pennsyl-
vanian with the exception of two genera which are Ches-
terian in age (Zeusocrinus and Paianocrinus) and a few
forms that pass into the Lower Permian. The oldest species
known with arms attached from Pennsylvanian rocks be-
longing to the family is Stenopecrinus rugosus Strimple
(1962) from the Morrowan. It has 30 uniserial arms that
appear to be endotomous. As noted by Strimple (1962)
the species is atypical of the genus and probably represents
an ancestral form.

True to accepted concepts of evolution there is an
increase in the number of arms in most forms. We know
one lineage (e.g. Stenopecrinus and Plaxocrinus) where the
arms remain endotomous and increase in number to 40, yet

retaining a uniserial structure. These genera have a ter-
mination anal sac platform with from six to eight large
flat spines surrounding a fairly restricted number of small
polygonal plates (typically five to eight). The dorsal cup is
moderately high, there is a pronounced basal concavity
but substantial portions of the basal plates are visible in
side view of the cup. They have small and elongated crowns.

Metaperimestocrinus is a specialized form coming out
of a form like Stenopecrinus in which the basal concavity
becomes shallow and the terminating platform of the anal
sac has a large number of perimeter spines (15 to 20 re-
ported) and numerous center plates. There is no appreci-
able change in the number of arms (40) or the structure of
the arms (uniserial), or the cup height (relatively high
cup), or the size of the crown.

Schedexocrinus apparently evolves from a form like
Plaxocrinus (e.g. P. tumulosus Strimple, 1949d, or P.
normalis Strimple, 1961a). There is no increase in the num-
ber of the arms (40) and they remain endotomous, but
the arms become biserial in the lower portions (SBrBr)
and may sporadically produce other interlocking segments.
The basal concavity is broad and shallow but the basals
are tumid and are visible in side view of the cup. The
terminating platform is large and is typically composed of
15 long flat spines directed outward and surrounding 32
small polygonal plates. The genus attains a greater width
than previously discussed forms but does not increase pro-
portionately in height.

In Polygonocrinus the arms branch isotomously at least
three times and are irregularly branched above that point.
The arms are essentially biserial although many sections
may have only cuneiform segments, The platform at the
termination of the anal sac is large with 14 large flattened
spines marking the perimeter and surrounding some 32 small
polygonal plates. The dorsal cup is low, with deep basal
concavity. The basals are depressed in mid-section and
confined to the basal concavity together with the infrabasals
and a large portion of the radial plates. The cup is much
like that of Pirasocrinus except the radials are more tumid
and the concavity more exaggerated in the later genus.

There is another form which is apparently closely re-
lated to Polygonocrinus in having a comparable dorsal cup
and terminating platform but the arms remain endotomous
although they branch five times. The secundibrachs are
interlocking much like those of Schedexocrinus but the re-
mainder of the arm segments are all uniserial. The form
is presented as Eirmocrinus, n. g.

Pirasocrinus has a dorsal cup with IBB, BB and much
of the RR in a deep basal concavity, much like those of

204 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Polygonocrinus and Eirmocrinus except the radials are pro-
jected downward and thereby give the dorsal cup an appear-
ance of greater height in Pirasocrinus. The arms of Piraso-
crinus are uniserial with regular brachials and branch iso-
tomously at least four times to produce a minimum of 80
arms. Specimens at hand show the terminating platform to
have 10 laterally directed spines surrounding several poly-
gonal plates.

Sciadiocrinus retains uniserial arms with regular brach-
ials and the arms branch at least three times isotomously.
It also has a large terminating platform with about 15
spines and numerous small central plates. It has a shallow
basal concavity and is characterized by having shortened
sutures between BB, which may be stated as a tendency for
the radial plates to attempt contact with infrabasal plates.
All species of Sciadiocrinus for which the arms are known
have the second bifurcation, and the second secundibrach
together with all of the axillary brachials fail to exhibit any
appreciable tumidity.

These are the only genera directly involved in the
current study so we are restricting our general comments
to them.

Genus EIRMOCRINUS Strimple and Watkins, new genus

Type species. — Eirmocrinus grossus Strimple and Wat-
kins, new species.

Range. — Desmoinesian, Pennsylvanian; North Amer-
ica.

Diagnosis. — Crown wide at base to about one-third of
height where it is sharply constricted, thereafter rising
evenly to the large terminating platform; dorsal cup shallow
with only the distal ends of radials visible in side view, IBB,
BB and most of RR confined to the deep basal concavity;
IBB apparently five, small, subhorizontal or mildly down-
flared; BB five, long, mid-portion depressed; RR five, wide,
long, subhorizontal; XX three, narrow, long; PBr,ax pro-
duced as a long, stout spine; SBr; is wide, large, quadrang-
ular, other SBrBr are low, interlocking, with SBry-sax and
spinose; TBr, is large and quadrangular but subsequent
TBrBr are short and quadrangular; the inner arms are un-
branched and decrease in size rapidly for a short distance
but thereafter maintain a uniform width until approaching
their termination, in the outer arms only, another bifur-
cation takes place with about two more endotomous bifur-
cations but the axillaries are only tumid without spinose
development, the arms are closely apposed; the terminating
platform is large with 13 spinlike plates surrounding numer-
ous small polygonal plates.

Remarks. — The dorsal cup of Eirmocrinus is much like

that of Pirasocrinus except the latter has highly tumid
radials and a more exaggerated basal concavity. Separation
of crowns is more readily possible because Pirasocrinus has
isotomous arms with evenly uniserial structure but Eirmo-
crinus has endotomous arms which, although uniserial in
upper portions, are decidedly biserial in lower portions,

Polygonocrinus also has a dorsal cup much like that of
Eirmocrinus, with deep basal concavity and most of the cup
plates involved in the concavity but the arms are isotomous
(at least three isotomous divisions) and essentially biserial
as compared to the endotomous arms which are uniserial in
their upper portions. Although it may not prove to be of
generic importance, the PBr,ax of Polygonocrinus are high-
ly tumid but not spinose as compared to the large spine-
like PBriax of Eirmocrinus. The terminating platforms of
these two genera are comparable.

Plaxocrinus has a low dorsal cup, but the basal plates
are tumid and extend out of the basal concavity. The arms
are typically 40 in number, uniserial and endotomous. The
PBryax is produced as a long spine, much like that of
Eirmocrinus. The terminating platform is much smaller, and
is typically surrounded by seven spine plates.

Sciadiocrinus is the only other genus closely compar-
able to Eirmocrinus. The structure of the cup is similar ex-
cept that more of the radial plates are visible in side view
and the sides of the basal concavity are more gently sloped
with the concavity not so pronounced in Sciadiocrinus. The
PBr,ax are not particularly tumid, much less spinose, and
second bifurcation is with a nonspinose SBrsax in all arms.
A third division is known to take place in some species of
Sciadiocrinus. The terminating platform of Sciadiocrinus 1s
much like that of Eirmocrinus except the spine-plates are
typically shorter and flatter in the former genus.

The generic name is derived from the Greek eirmos
for line, with reference to the long unbroken lines formed by
the inner arms of the endotomous arm structure.

EIRMOCRINUS GROSSUS Strimple and Watkins, new species
Plate 30, figures 1-4

The description of the genus Eirmocrinus is based on
E. grossus, the type species, and repetition is not made here.
Some additional observations are as follows:

The base of the cup is disturbed in preservation but
careful study and comparison with similar forms have es-
tablished the existence of a deep basal concavity with
steeply downflared basals. The radials are not appreciably
disturbed and were clearly horizontal in attitude. While dis-
articulated a drawing was made of an articular facet which

TExAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 205

is large and directed outwardly. Several interesting features
were observed. There is a single line of denticles along the
edge of the muscle area bordering the body cavity. The
outer surface of the radial plates continues into the adsu-
tural area and extends almost to the body cavity. A semi-
rounded facet is formed at some corners of the articular
facets next to the body cavity. This feature was noted by
Strimple (1961a) for Schedexocrinus gibberellus and appar-
ently represents an attachement area for the large anal
tube. There are no other special features of the facets. The
transverse ridge is broad and marked by irregular denticles,
the outer ligamental area is well defined, the large triangu-
lar-shaped muscle areas show that long fossae and the
intermuscular notch and furrow are broad.

The long posterior basal has a narrow upper facet for
reception of anal X, which plate has fallen inwardly. The
anal plate apparently expanded considerably above the
summit of the cup and its upper surface is marked by
canals such as found in anal sac plates. The RA is relatively
small and from its structure it apparently had lost contact
with the right posterior basal.

First primibrach is axillary, of medium height, and
is produced as a long stout spine, much like the typical
primibrach of Plaxocrinus. First secundibrachs are wide and
long with a more or less regularly quadrangular outline.
There are usually three or four low, interlocking secundi-
brachs (biserial structure) which bear pinnules on their
inner edges. Bifurcation is with a spinose SBry-sax and
thereafter the only branchings are in the outer arms, 1.e. the
arms become endotomous. The axillary tertibrach is spinose
but subsequent axillaries are not developed as spines. All
brachials above the secundibrachs are quadrangular, form-
ing 50 uniserial arms. Internally, there are often thin pro-
jections, terminating with an adze-shape, which occupy the
area opposite the side which has a pinnule attached and is
thus apparently associated in some manner with the pinnule
arising from the brachial below, or with the massive anal
tube. When closed the arms are closely appressed so that an
unbroken wall is formed about the visceral area.

The terminating platform of the anal sac was in place
when found by the junior author but disarticulated and had
to be restored from the recovered plates. The large spines
have a fairly long base, comparable to those of Schedexo-
crinus gibberellus. There are 13 spines of which one does not
touch the perimeter of the central area occupied by small
polygonal plates. The unusual spine is wedged between the
base of two other spines and is projected parallel to one of
the adjacent spines. The outer edge, of the central platform
proper, is sloped slightly downward and the slope is accen-

tuated by a slight curvature of the shaft of the spines, so
that a downward slope of about 27° is formed.

There is some elusive evidence of granular ornamenta-
tion under magnification, but it is only unquestionable in
the “protected” area of the basal concavity on the basal
plates.

Measurements of the holotype in millimeters:

Length of crown 120.0
Width of crown (maximum) 70.0
Height of cup 6.0
Width of cup (estimated) 33.0
Length of basal 7.2
Width of basal 8.8
Length of interbasal suture 2.2
Length of radial 12.2*
Width of radial 22.0*
Length of interradial suture 9.0

* measurements taken along surface curvature.

The species name grossus is with reference to the large
size of the crinoid.

Holotype. —USNM, No. $5097 collected by Wm. T.
Watkins, reposited in the U. S§. National Museum.

Occurrence. — Shale below the Kickapoo Falls Lime-
stone Member, Millsap Lake Formation, Strawn Group,
Desmoinesian, Pennsylvanian; about one-third mile below
the falls on Kickapoo Creek southwest of Dennis, Hood
County, Texas.

Genus PIRASOCRINUS Moore and Plummer, 1940

Type species. — Pirasocrinus scotti Moore and Plum-
mer, 1940.

Range. — Desmoinesian, Pennsylvanian; North Amer-
Ica.

Remarks. — The classification of crinoids of Pennsyl-
vanian age is usually based on characters of the dorsal cup
and modifications of the concept are made if necessary when
complete crowns are available. The genus Pirasocrinus is a
notable exception. Several crowns of the type species P.
scotti were available; however, the original material did not
fully reflect the true structure of the termination of the
mushroom-like anal sac, or the exact structure of the dorsal
cup.

The holotype of P. scotti possesses a restricted number
of spinelike plates at the termination of the anal tube in a
compressed manner, entirely unsatisfactory for adequate
observation. Moore and Plummer thought they had a nearly
intact summit and reported that there were four, broad-
based, laterally directed spines, without intervening small
plates, which unite to form a roof over the sac. We are for-

206 PALAEONTOGRAPHICA AMERICANA (VI, 40)

tunate to have specimens that show without question the
existence of ten broad-based, laterally directed spines.

PIRASOCRINUS SCOTTI Moore and Plummer, 1940

Plate 33, figures 3, 4; Plate 35, figures 1-3; Plate 45, figures 1, 2;
Plate 56, figure 2

See Bassler and Moodey, 1943, p. 611

This species is of special interest because it is the type
of the genus and the genus is familial type of Piraso-
crinidae. Three magnificent crowns in excellent preservation
have been collected by the junior author and afford addi-
tional information,

As noted under generic remarks, the holotype of P.
Scotti possessed a restricted number of spine-plates at the
summit of the anal tube, reportedly four broad-based,
laterally directed spines without intervening small plates.
We now know there are ten large spines, of the Schedexo-
crinus type, surrounding a substantial number of small,
polygonal plates to produce a good-sized platform, though
not so large as typical of Schedexocrinus. The arms of
Pirasocrinus scotti branch evenly four times to produce 80
isotomous arms. Schedexocrinus branches twice isotomous-
ly and thereafter two times in outermost half-rays only,
to produce 40 arms.

The young hypotype has already attained the four
bifurcations of the arms although it is only two-thirds as
long as a mature form. The infrabasals are not visible, but
the basal plates of the young specimen are more prominent
than found in more mature specimens. We now understand
that the accruement of calcite on the bulbous radials in-
creases their thickness and causes the basal concavity to
appear to be deeper than is structurally the case. That is,
the basal concavity is deeper only by virtue of the increased
bulbosity of the radial plates with age and is not due to a
change in the basal elements. The attitude of the basals is
also affected by the change in the radial plates.

We are unable to verify the exact structure of the
basal concavity shown by the cross section of Moore and
Plummer (1940, text-fig. 53), but we believe the infrabasals
are more subhorizontal than downflared.

The large figured hypotype of Pirasocrinus scotti has
an over-all length of crown of 70.5 mm, maximum width
of 35.0 mm.

Types. — Holotype, Kansas Univ., No. 60261, paratype
No. 60262, collected by R. C. Moore. Paratypes, Plummer
Collection, Nos. P-8190 and P-8199, Bureau of Economic
Geology, University of Texas, collected by Gayle Scott.
Hypotypes, USNM, No. 85106, $5112, $5154, $5197, col-
lected by Wm, T. Watkins, reposited in U.S. National
Museum.

Occurrence. — Shale below the Kickapoo Falls Lime-
stone Member, Millsap Lake Formation, Strawn Group,
Desmoinesian, Pennsylvanian; Moore and Plummer (1940)
loc. 110-T-4, about one-fourth mile downstream from Kicka-
poo Falls, southwest of Dennis, Hood County, Texas.

Genus ANCHICRINUS Strimple and Watkins, new genus

Type species. —Anchicrinus toddi Strimple and Wat-
kins, new species.

Range. — Desmoinesian, Pennsylvanian; North Amer-
ica.

Diagnosis. — The dorsal cup is low, bowl-shaped with
fairly broad shallow basal concavity and a large area for
stem attachment. When viewed from above or below, the
cup has a circular outline. IBB disk composed of five plates,
large, pentagonal-shaped, downflared, mostly covered by
large stem. Five basals subhorizontal, sweep out of shallow
basal concavity with distal ends flexed upward to be visible
in side view of cup. Five radials are mildly tumid with
proximal ends in shallow basal concavity with distal por-
tions curved upward to form lateral walls of cup, Almost
imperceptible notches are present at the interradial sutures,
at the summit of the cup. The slope of the facets is slightly
outward. The posterior interradius is broad and mildly con-
vex. Three large, long anal plates in normal (Primitive
Type) arrangement are present in the cup.

The arms are long, slender, uniserial, and branch
isotomously with the first primibrach but thereafter four
times in outer arms only (endotomously). The first primi-
brach is large, tumid, axillary, and is usually nonspinose
other than for a nipple-like protrusion but may develop
a full spine. Subsequent axillaries are spinose. A platform-
like termination of the anal sac marks the summit of the
crown and is composed of a few small polygonal plates
surrounded by eight outwardly directed spinelike plates.

Remarks. — This genus appears to have close affinity
with the genera Athlocrinus Moore and Plummer, 1940 and
Pirasocrinus Moore and Plummer, 1940, and may be related
to Sciadiocrinus Moore and Plummer, 1938, or Stenope-
crinus Strimple, 1961a, The arms and tegmen are somewhat
like those of Pirasocrinus but the cup, with almost imper-
ceptible basal concavity, is entirely foreign to the deep basal
concavity with sharply downflared basals of Pirasocrinus.
The cup shape is almost identical with that of Athlocrinus
except for the mild basal concavity of Anchicrinus. The
base of Athlocrinus is flat except for the stem impression
and the surface of the plates are smooth. In Anchicrinus
the cup plates are mildly tumid causing slight impressions

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 207

to appear at the sutures. Tumidity of the radial plates is a
characteristic of the form described as Athlocrinus clarus
Strimple (1962a) from the Ooologah Formation, Desmo:
nesian, of Oklahoma which species is referred to here as
Anchicrinus clarus (Strimple), now combination. This later
species reflects some characters that are like Sciadiocrinus
and may link the two genera. The arms of Anchicrinus are
almost identical with those of typical Stenopecrinus except
for the axillary first primibrachs which are more tumid
and lower than found in that genus. There is a sharp basal
concavity in Stenopecrinus but the radials do not enter the
cavity. In Anchicrinus the proximal tips of radial plates
actually curve into the basal concavity. Typically in Sten-
opecrinus the platform at the summit of the anal sac is
composed of a restricted number of small plates surrounded
by seven large spinelike plates. There are eight large spine-
like plates and a limited number of small center plates form-
ing the platform of the type species of Anchicrinus.

The generic name is taken from the Greek word anchi
meaning near.

ANCHICRINUS TODDI Strimple and Watkins, new species
Plate 43, figures 1, 2, 8-11, 14, 15; Plate 48, figures 1, 2

Dorsal cup is subcircular in outline when viewed from
above or below and is not indented at the posterior side.
The cup is shallow, bowl-shaped with downflared infrabasal
plates. There is a broad shallow basal invagination aug-
mented by a sharply impressed, large, round columnar
cicatrix. The median portion of the columnar depression is
raised in the figured paratype which might be an incipient
columnal. Sutures between radials are impressed but there
are no pronounced interradial notches at the summit of the
cup.

Five infrabasals are of medium size, project slightly
beyond the columnar scar, and form a pentagonal-shaped
disk. The five basals are of medium size, sweep out of the
basal concavity to participate in the low lateral walls, and,
except for the blunt posterior basal, form a substellate-
shaped circlet. The five radials are wide, pentagonal, mildly
tumid elements with proximal portions curved under to
participate in the basal area. The articular facets slope
gently outward, almost imperceptibly in the young para-
type; the transverse ridge is straight and distinct but is not
raised; a few crenulations are preserved and interrupt the
transverse ridge toward the outer ends; a sharply defined
ligamental pit is just below the transverse ridge and the
outer ligamental area is separated from the outer face of
the radials by a fine ridge. The inner ligamental area is
marked by a pair of oblique furrows and a pair of triangu-

lar-shaped muscle areas. An intermuscular notch is present,
but there is no well-defined intermuscular furrow.

There is usually provision for five anal plates in the
anal pyramid of the broad posterior interradius, of which
the lower three are partially or entirely within the dorsal
cup. The three anal plates in the cup are in normal
(Primitive Type) arrangement. RA is slender, pentagonal,
contacts RPB below and rests obliquely on PB, with anal
X to the left above and RX and RPR above and to the
right. Anal X is slightly larger than RA, is hexagonal,
elongate, rests solidly on PB, and more than half of its
length extends above the cup. RX is elongate, more-or-less
hexagonal-shaped with less than one-fourth of its length
within the cup. X, is usually irregularly pentagonal-shaped
and is supported by both X and RX. There is a small
narrow RX, on the upper right shoulder of RX preserved
in place in the holotype but missing in the paratypes.

There is provision for a total of 50 long, uniserial arms.
First bifurcation takes place with PBryax in all arms.
PBry,ax is a full, tumid plate in all specimens and usually
develops a short, spinelike protruberance in the upper
median portion. The arms branch isotomously one more
time and thereafter in outer arms only (endotomously),
reducing size sharply each time, particularly in the inner
unbranched arms. The upper axillalry brachials are pro-
truded as long slender spines, usually broken off in preserva-
tion.

A platform of small, polygonal plates is surrounded by
eight outwardly directed, flat, spinelike plates and marks
the termination of the anal sac. All of the plates are not
fully preserved in the material at hand The arms extend
slightly above the platform and distal ends lie in notches
between the large spine plates.

The columnar scar is round, marked by about 36 crenu-
lations, and pierced by a small lumen.

Measurements in millimeters:

Holotype Paratype
Length of crown 41.7 —_
Width of crown 17.0 —
Width of cup 14.0 17.0
Height of cup 4.5 5.3
Width of IBB disk 5.0 4.9
Width of basal 4.6% a0"
Length of basal 4.6% 4.6"
Length of interbasal suture 1.4* 1.3*
Width of radial 9.0% 9.8%
Length of radial 6.1" 6.2*
Length of interradial suture 4.0" Beil
Length of RA 5.5" 4.6%
Width of RA 2.8" 2.4*
Length of X ao 6.0*
Width of X 3.5% 4.0%
Length of RX 4.9% 4.9*
Width of RX 2:5 4.1*

“Measurements taken along surface curvature.

208 PALAEONTOGRAPHICA AMERICANA (VI, 40)

One paratype has an estimated over-all length of 37.5
cm of crown.

Remarks. — Anchicrinus toddi is larger than A. clarus
(Strimple), the basals are more prominent, and the radials
are less prominent. The cup of A. toddi is similar to that
of Plaxocrinus crassidiscus (Miller and Gurley), except the
later species has wide notches at the interradial sutures,
at the summit of the cup, and has an advanced, narrow
posterior interradius.

The arms and tegmen of Anchicrinus toddi indicate
close relationship from Pirasocrinus scotti; however, P.
scotti has a deep basal concavity with sharply downflared
basals different from the shallow basal concavity of Anchi-
crinus toddi.

Types. — Holotype, USNM, No. 85164, collected by
Louis Todd. Three paratypes, USNM, Nos. $5140, $5144,
$5147 collected by Wm. T, Watkins, reposited in the U.S.
National Museum.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Genus LASANOCRINUS Moore and Plummer, 1940

LASANOCRINUS CORNUTUS Moore and Plummer, 1940

A single poorly preserved specimen in the presently
considered collections is identified as Lasanocrinus cornutus.
The unique prolongation of the median portion of each
radial plate in a downward direction is the character afford-
ing identification. The projections are more robust and less
pointed than found in L. daileyi (Strimple), The presently
observed specimen is designated a hypotype.

Types. — Holotype, Peabody Museum, No. 15246, Yale
University, paratype, No. 15246A, collected by C. Schu-
chert, D. K. Greger, and A. McCoy. Hypotype collected
by Wm. T. Watkins, reposited in the U. §S. National
Museum. No number assigned.

Occurrence. — Lemons Bluff Member, Marble Falls
Formation, Atokan, Pennsylvanian; on Rough Creek about
11 miles southeast of San Saba, San Saba County, Texas.
Univ. Texas Loc. 205-T-17.

Genus PLAXOCRINUS Moore and Plummer, 1938

Type species.— Hydreionocrinus crassidiscus Miller
and Gurley.

Range. — Pennsylvanian; North America.

Condensed evalution of the type species of the genus as
follows:

Dorsal cup is round, broad interradial notches, broad,
shallow basal concavity. Five IBB are mildly downflared,
five BB are subhorizontal in proximal extremities but flared
upward to be readily visible in side view, five RR are wide,
mildly tumid, with proximal tips extending into the basal
plane and distal ends curved inward. Posterior interradius
is narrow and depressed, with RA in direct posterior posi-
tion and anal X above it. RX is present but is not really in
the cup, being supported by the extended left shoulder
of the right posterior radial and the right shoulder of anal
X. The articular facets of RR are sloped slightly outward.

Remarks. — A complete study of the genus is not
attempted here, but it has become necessary to return to
the type species of the genus to attain irrevocable criteria
for differentiation of the species involved. The holotype
(monotype) of Plaxocrinus crassidiscus is apparently a
highly advanced form and probably represents the ultimate
stage of a phyletic line or near to the end stage. This is
based on the highly advanced nature of the anal plates and
the assumption that evolution is from a deep basal con-
cavity to a flat base in this lineage. Few forms presently
assigned to Plaxocrinus will be retained in the genus if the
generic concept is emended to characters of the type species.

Among known forms we are considering here the
species described as Plaxocrinus perundatus Moore and
Plummer (1940) and Plaxocrinus obesus Moore and Plum-
mer (1940) from the Millsap Lake Formation (Desmoi-
nesian) of Texas. P. perundatus is based on a holotype and
two paratypes of which the holotype and one paratype
were figured by Moore and Plummer, The holotype is from
the shale below the Kickapoo Falls Limestone Member and
the two paratypes are from the Brannon Bridge Limestone
Member, both members belonging to the Millsap Lake
Formation (Desmoinesian). The Kickapoo Falls Limestone
is the older horizon. Strimple (196la) assigned the species
to Schedexocrinus and emphasized the arm structure as a
determining factor. The paratype figured by Moore and
Plummer (1940, pl. 15, fig. 5) was the basis for the
assignment and the arms were preserved to just above the
axillary secundibrachs. The specimen is listed in the text as
being from Brock (loc. H-34) although the plate explana-
tion stated it was from Kickapoo Falls. The large primi-
brachs are bulbous, not extended as spines and are like
those of either Schedexocrinus or Polygonocrinus and we be-
lieve the specimen could be assigned to Polygonocrinus
libratus, n. sp. with little hesitation. The holotype of
Plaxocrinus perundatus is distinctly different in having a

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 209

shallower basal concavity, more prominent basal plates
that extend appreciably out of the basal concavity, and
a distinctly higher cup. The species as restricted is close
to Plaxocrinus tumulosus Strimple (1949d), from the
Pumpkin Creek Limestone Formation (Desmoinesian).
Plaxocrinus obesus is atypical of the genus as noted by the
original authors. We have placed the species in Sciadio-
crinus and the explanation for this action is given in that
section of the study.

PLAXOCRINUS PERUNDATUS Moore and Plummer, 1940

See Bassler and Moodey, 1943, p. 634.

The species is well defined in so far as the dorsal cup
is concerned in that the holotype, cross-sections of cup,
and subsequent description is applied to the dorsal cup.
Two paratypes are listed by Moore and Plummer (1940),
both of which are from a younger horizon than the holo-
type and one of which was figured, as noted under the
generic discussion herein. We have not examind the un-
figured paratype, but the illustration of the other para-
type which has portions of the lower arms attached shows
a form atypical of the species. We believe the partial crown
is probably a representative of Polygonocrinus, likely con-
specific with P. libratus.

The dorsal cup has a shallow basal concavity, the
infrabasal circlet is subhorizontal in attitude, the basals
curve out of the basal concavity to become visible in side
view of the cup and are mildly tumid, and the radial plates
have subhorizontal articular facets with proximal tips reach-
ing the basal plane but do not participate appreciably in the
basal concavity. There is a decidedly arcuate flattened
vertical area on the radials in the distal region adjacent
to the articular facet.

The species is apparently closely related to Plaxocrinus
tumulosus, as noted elsewhere. We do not have any speci-
mens of P. perundatus in collections at hand.

Types. — As restricted, the holotype is probably the
only true representative of the species. It is in the Plummer
Collection, P-11190, University of Texas, Austin, Texas.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Genus STENOPECRINUS Strimple, 1961la

Type species.— Perimmestocrinus planus Strimple,
1952b.

Range. — Pennsylvanian, Lower Permian; North
America.

Remarks. — As this genus is presently defined, it is
almost essential to have the arms preserved, at least in part,
in order to make identification with any degree of cer-
tainty. Nevertheless, at this time the genus continues to
retain a distinctive status among related forms, with the
possible exception of Metaperimetocrinus Strimple (196la).
Metaperimestocrinus is usually relatively small and is sep-
arated on the basis of having no appreciable basal con-
cavity of the cup and in possessing a highly spinose ter-
mination of the anal sac. Plaxocrinus Moore and Plummer
(1938) is usually a larger form, has longer arms, the axillary
brachials above the first primibrachs are not produced as
spines, and the cup is essentially different in contour. The
distal termination of the anal sac is identical with that of
Stenopecrinus. The genus Perimestocrinus Moore and
Plummer (1938) has been emended by Strimple (1961la)
and is considered to be a different phyletic line. Strimple
(1963) suggested possible affinity of Perimestocrinus with
Dasciocrinus Kirk (1939),

One species of the genus Stenopecrinus is presently
considered and described as S. longus, n. sp.

STENOPECRINUS LONGUS Strimple and Watkins, new species
Plate 48, figures 5, 6

This species is based on a single magnificent crown.
It is slender, long, the arms are closely apposed (as is
typical of the genus) and the terminating platform spine
plates of the anal tube are unusually small.

The dorsal cup is compressed so that exact measure-
ments and comparisons are difficult but all elements are
preserved. By deduction, the cup is known to have been
shallow, truncate bowl-shaped, with a_ basal concavity.
There are five infrabasals mainly obscured by the medium.
sized, round proximal columnals. Five basals are of medium
size and appear to be depressed in mid-portion. Five radials
are large, unquestionably formed most of the cup height
and judging from their lower curvature could not have
entered the basal plane. Five anal plates form an anal
pyramid, the lower three being elongated cup elements in
normal (Primitive) arrangement. Anal X extends well
above the summit of the cup with only the lower third
participating in the cup. The distal end is narrow. RA is
entirely within the cup and does not attain as great length
as anal X. The RX is about the same size as X but has
a proportionately broader distal end. About one-sixth of it
is in the cup. X; appears to be much smaller than X and
joins RX in supporting RX, which is a small quadrangular
piece with a bluntly pointed distal end. The arms fit snugly
against the anal pyramid in the posterior interradius.

210 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Slightly less than 40 uniserial to mildly cuneiform arms are
indicated. First primibrach is slightly elongate, with the
upper mid-portion developed as a spine, and is axillary in
all rays. A second isotomous branching takes place with a
low, spinelike eight to eleventh axillary secundibrach in all
arms, At this point the inner half-arms remain unbranched
and as many as 40 tertibrachs have been observed without
the actual termination being preserved. The outer half-arms
branch again with the twelfth to fifteenth tertibrach, with
the resultant inner half-arm remaining unbranched but the
outer half-arm usually branches again with the axillary
thirteenth to fifteenth quatribrach. All axillaries are pro-
duced as spines. Each brachial has a sharply angulated
projection to fill a wide angled recession at the suture
line of the apposed pair of brachials. This creates an im-
penetrable exterior when the arms are in repose and is a
specialization among the pirasocrinids as well as a few
other forms. When one considers that the animal would
probably suffocate if the arms were closed for any appre-
ciable period of time, it appears strange that it would be
so preoccupied with the development of a complicated inter-
locking arm system.

Three small plates are preserved at the distal end of
the crown and are obviously flat spine-plates which would
participate in forming a perimeter surrounding smaller
polygonal plates to form a platform at the summit of the
anal tube. They have a short, wide inner portion that is
shallowly depressed. The spinelike projection is broken in
all instances so it is not possible to state whether short or
long spines were formed, but based on other species of
the genus they were probably long and slender. The entire
surface of the crown is covered by small granules, about
64 to a square millimeter, which have a tendency to co-
alesce.

Proximal columnals are distorted by lateral compres-
sion but were large, apparently round, and the crenula-
lations were continued to the exterior to make an uneven
suture.

Measurements in millimeters:

Holotype
Length of crown 96.5
Width of crown-maximum VASES)

Width of cup-estimated 21.0
Height of cup-estimated 5.5
Length of basal 5.1
Width of basal 5.9*
Length of interbasal suture 3.5
Length of radial 6.8

Width of radial 12.2"
Length of interradial suture 5.7"
Length of anal X 6.7*
Width of anal X 5.0

Length of RA 5.2"

Width of RA 3.5”
Length of RX 6.8*
Width of RX 5.0*

*measurements taken along surface curvature.

Remarks. — The unusually long, slender nature of the
crown of this species is distinctive as well as the relatively
small, flat, spinelike plates, with depressed mid-portions, of
the tegmenal termination platform. The species does not
appear to be typical of Stenopecrinus.

Holotype. — USNM, No. 85166, collected by Wm. T.
Watkins, reposited in the U.S. National Museum,

Occurrence. — Kickapoo Falls Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; on Kickapoo Creek about one-fourth mile
below Kickapoo Falls southwest of Dennis, Hood County,
Texas.

Genus SCIADIOCRINUS Moore and Plummer, 1938

Synonymy. — Hydreionocrinus
Schistocrinus (part) of authors.

(part) of authors;
Type species. — Zeacrinus (Hydreionocrinus) acantho-

phorus Meek and Worthen.
Range. — Morrowan? to Missourian; North America.

Remarks. — Description of the genus was primarily
formulated about a single incomplete crinoid crown which
was the holotype of the type species of the genus. Excellent
line drawings of the specimen are given by Moore and
Plummer (1938, figs. 23 a-c). Those authors stated that
positive assignment to the genus is not possible unless the
anal sac of a specimen is known in conjunction with the
dorsal cup and arms. There is indeed some difficulty in
identification of these forms, even when the terminating
platform of the anal sac is known. We find Moore and
Plummer (1940) described species as Schistocrinus parvus
and S. confertus which have closely comparable termina-
tions on the anal tube and both species have been redesig-
nated as Sciadiocrinus by Strimple (1961a). Considering
the arms of the species involved, one finds they are not
well preserved above the second bifurcation in the holotype
of Sctadiocrinus acanthophorus but so far as known are
practically identical with the arms of the two species
ascribed by Moore and Plummer to Schistocrinus. The calyx
structure is comparable for all three forms, one of which
(S. parvus) is suppressed herein as an immature form,
conspecific with S. confertus.

Schistocrinus planulatus Moore and Plummer (1940),
also from the Millsap Lake Formation (Desmoinesian), is
monotypic and the specimen is in poor preservation, but

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS Dil)

it appears to belong to Sciadiocrinus and was placed in the
genus by Strimple (196la). Moore and Plummer (1940, p.
225) stated as follows, “The sutures between infrabasals
(IBB) and radials (RR) are slightly less than 2 mm in
length.” We presume this alludes to the interbasal suture
which means that radials do not meet infrabasals as they
do in Schistocrinus torquatus, the type species of that genus.
Other factors also indicate closer affinity of planulatus for
Sciadiocrinus than for Schistocrinus.

Two other forms are referred to the genus from the
presently considered collections, which are Sciadiocrinus
harrisae Moore and Plummer, 1940, and Plaxocrinus obesus
Moore and Plummer, 1940. We believe P. obesus is closer to
the type species of Sciadiocrinus than is Sctadiocrinus har-
risae. Moore and Plummer (1940, p. 191) recognized Plaxo-
crinus obesus was atypical of Plaxocrinus, quote “This
species is not a typical representative of Plaxocrinus in that
the radials are abnormally bulbous, their proximal tips
being slightly downflaring and thus forming part of the
basal concavity.” Radial plates of this nature are typical of
Sciadiocrinus acanthophorus although the bulbosity is not
so pronounced.

The genus Schistocrinus Moore and Plummer is consid-
ered by Strimple (196la) to belong to another phyletic
lineage, represented by the family Texacrinidae.

Diagnosis. — The genus is distinguished by having a
relatively short, broad crown; RR have a tendency to touch
IBB, and often do make contact; BB circlet is substellate
in outline (except for the posterior); base of cup is shal-
lowly concave; arms fundamentally isotomous with first
three bifurcations on PBr,ax, on SBroax and TBrs-5;ax, sub-
sequent bifurcations usually restricted to outermost
branches of the two half rays; brachials decidedly uni-
serial; umbrella-like extremity of anal tube composed of
numerous small polygonal plates surrounded by 12 to 15
outwardly directed spinelike plates (many of the peri-
pheral apines are short and blunt).

Species assigned to Sciadiocrinus are as follows:

Present
Occurrence assignment

Hydreionocrinus acanthophorus Desmoinesian; Sciadiocrinus

Meek and Worthen Ill. & Mo. Type species
Sciadiocrinus? crassacanthus Morrowan; spinelike plates
Moore and Plummer Okla. & Ark.

Sciadiocrinus disculus Missourian; Sciadiocrinus
Moore and Plummer Texas
Sciadiocrinus harrisae Desmoinesian;
Moore and Plummer Okla. & Tex.
Euphachycrinus platybasis Lower Aubrey
White Group, Utah
Schistocrinus confertus Desmoinesian;
Moore and Plummer Texas
Schistocrinus planulatus Desmoinesian ;
Moore and Plummer Texas

Sciadiocrinus
Sciadiocrinus
Sciadiocrinus

Sciadiocrinus

Plaxocrinus obesus

Desmoinesian; Sciadiocrinus

Moore and Plummer Texas
Schistocrinus parvus Desmoinesian; Syn. Sciadiocrinus
Moore and Plummer Texas “ confertus
Sciadiocrinus llanoensis, Atokan; Sciadiocrinus
n. sp. Texas

SCIADIOCRINUS HARRISAE Moore and Plummer, 1940
See Bassler and Moodey, 1943, p. 673.

On the basis of current information, we are allowing
this species to remain in Sciadiocrinus because of the mildly
down-flared nature of the basal plates and the short suture
between basal plates. The robust tumidity of radial plates
often encountered in variable representatives of the species
is not entirely foreign to Sciadiocrinus acanthophorus (Meek
and Worthen) the type species of the genus, Illustrations of
S. acanthophorus do not show this feature plainly but the
distal portions of the radials are erect and there is a sharp
flexing of the proximal portions into the depressed basal
area. Pirasocrinus has this same structure though even more
exaggerated because of the steeply downflared basals and
deep basal concavity.

S. harrisae appears to a polymorphic species and will
require considerable study to be properly understood,

Types. — Holotype, Marrs Collection M-45, collected
by Mrs. W. R. Marrs; paratype, Scott Collection 6026C,
Texas Christian University, collected by Gayle Scott; para-
type Marrs Collection M-31, M-32, collected by Mrs. W R.
Marrs; paratype, Stevens Collection 45987, Kansas Univer-
sity, Nos. 45897a-k in Bob Stevens’ private collection;
hypotype, USNM, no number, collected by Wm. T. Wat-
kins; (plesiotype) hypotype, O.U. 3963, O.U. 4559 aberrant
specimen, OU 4567, collected by H. L. Strimple, Paleon-
tological Collections University of Oklahoma; (plesiotypes)
hypotypes from Garnett Quarry collected by Mr. and Mrs.
H. L. Strimple, reposited in U.S. National Museum,

Occurrence. — Holotype, Texas Loc. 110-T-3, about
414 miles east-northeast of Lipan, Hood County, Texas;
paratype 6026C, Loc. 110-T-4, north side of Kickapoo
Creek, one-quarter mile below Kickapoo Falls, southwest
of Dennis, Hood County, Texas, shale below Kickapoo Falls
Limestone Member, Millsap Lake Formation, Strawn
Group, Desmoinesian; paratypes, M-31 and M-32, and
hypotype USNM (no number), about three miles south-
west of Brock, Parker County, Texas, Brannon Bridge
Limestone Member, Millsap Lake Formation, Strawn
Group, Desmoinesian; paratypes 45897 and 45897 a-k,
(plesiotypes) hypotypes, OU 3963, aberrant specimen OU
4567 and several (plesiotypes) hypotypes in U.S. National
Museum are from Garnett Quarry, SW14 sec. 28, T. 20 N.,
R. 14 E. Tulsa County, Oklahoma; (plesiotype) hypotype

212 PALAEONTOGRAPHICA AMERICANA (VI, 40)

OU 4559, NE14 sec. 5, T 19 N., R. 14 E., Tulsa County,
Oklahoma, Oologah Limestone Formation,
Group, Desmoinesian, Pennsylvanian.

Marmaton

SCIADIOCRINUS CONFERTUS (Moore and Plummer), 1940

See Bassler and Moodey, 1943, p. 634

This species was originally described as Plaxocrinus
obesus Moore and Plummer, 1940, and no more can be
added to the excellent description and illustrations, As
noted herein under generic discussion, Moore and Plum-
mer considered the species to be atypical of Plaxocrinus and
the features involved are typical of Sciadiocrinus as repre-
sented by the type species of the genus. The distal tips of
the basals are visible in side view of S. obesus, or to word it
differently, the distal tips of basals are curved upward
which is a character ascribed to Plaxocrinus, but we do not
consider it sufficient within itself. We believe the short-
ening of the sutures between basals, along with other per-
tinent characters serve to identify obesus more closely with
Sciadiocrinus than with Plaxocrinus.

Types. — Holotype, Plummer Collection, No. P-8187
collected by Gayle Scott; paratype, Harris Collection No.
H-30, collected by Mrs. G. W. Harris; paratype, Marrs
Collection No. M-35, collected by Mrs. W. R. Marrs; hypo-
type USNM, collected by Mrs. J. H. Renfroe; (plesiotype)
hypotype OU 3996, collected by H. L. Strimple; paratype,
Plummer Collection, No, P-664, collected by F. B. Plummer.

Occurrence. — Holotype (P-8187), paratypes (H-30,
M-35) from Texas Loc. 110-T-4, below Kickapoo Falls, north
side of Kickapoo Creek, southwest of Dennis, Hood County,
Texas; shale below the Kickapoo Falls Limestone Mem-
ber, Millsap Lake Formation, Strawn Group, Desmoinesian:
paratype (P-664) three miles southwest of Salesville, Palo
Pinto County, Texas; Palo Pinto Limestone, Canyon
Group, Missourian; (plesiotype) hypotype (OU 3996),
Garnett Quarry, SW1,4 sec. 28, T. 20 N., R. 14 E., Tulsa
County, Oklahoma; Oologah Limestone Formation, Mar-
maton Group, Desmoinesian.

SCIADIOCRINUS CONFERTUS (Moore and Plummer), 1940
Plate 43, figure 12; Plate 52, figure 14
See Bassler and Moodey, 1943, p. 672

As previously noted, this species was described as
Schistocrinus confertus but has been referred to Sctadio-
crinus by Strimple (196la). There is no new informa-
tion afforded by our material other than to re-evaluate some
previously discussed features and to incorporate S. parvus
into the species as a juvenile stage. In particular, the need

for a clearer understanding of the arm structure is indi-
cated.

The description of arm structure given by Moore and
Plummer (1940, pp. 222, 223) is as follows: “Each ray
branches isotomously on the first primibrach (IBr,), on
the second secundibrach (IIBro) and again on the third to
fifth tertibrach (IIIBrz-;); above this point there is fur-
ther bifurcation only in the two outermost and the two in-
nermost branches of the ray, or, stated otherwise, the out-
side branches of each of the two half-rays may be sub-
divided as many as three times above the axillary terti-
brachs, but the other arms do not bifurcate.” Yet in the
drawings (1940, fig. 48) a fourth bifurcation is shown in
every branch on the third to seventh quartibrach and sub-
sequent branching in some inner half rays. As drawn, the
species would have a total of 125 arms but from the word
description there could be no more than 100 arms. From
our observation there appears to be usually 60 to 80 arms
which follow the pattern of branching given in the word
description by Moore and Plummer, except there is usually
only one or two divisions above the third isotomous branch-
ing, rather than three or four as previously reported. One
fragmentary specimen, almost certainly conspecific, shows
a total of 100 arms may be attained.

The form described as Sciadiocrinus parvus was based
on a monotype, which is a well-preserved small crown.
The description was well presented; however, it is juvenile
and we consider it to be conspecific with S. confertus. Some
additional information is afforded by specimens in our
collections.

One large hypotype has 13 spine-plates preserved in
the umbrella-like termination of the anal sac. A medium-
sized hypotype has an estimated 12 spine-plates in the
termination. The spines are typical for the species in having
a spade-shape.

The mean width of the crown of the largest hypotype
is 24.5 mm, estimated height of crown is 33.0 mm, a ratio of
0.7. The mean width of the medium-sized hypotype is 18.0
mm, estimated height of crown is 22.0 mm, a ratio of 0.8
mm. The width of the holotype (after Moore and Plum-
mer) is 17.0 mm, height of 21.0, a ratio of 0.8. It is of
some interest to note the holotype of Sciadiocrinus acantho-
phorus, which is the type of the genus and also of Desmoin-
esian age, has a width/height ratio of about 0.7.

Additional arm branchings are disclosed by the me-
dium-sized hypotype although preservation is not too satis-
factory for intimate observation in most rays. The usual
branching is with PBr,ax, SBroax and TBrs-;ax in all arms.
There is usually one other bifurcation with QBry-gax in the

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 213

outer arms. If this pattern of bifurcation is consistent in
all rays, a minimum of 60 arms are produced, or 20 more
than reported for “S. parvus.” It should be noted that an
extra arm may be found in the inner arms of each half-
ray of the posterior, so that a total of 61 to 63 arms might
be found in young forms.

The brachials grew more rapidly in width than in
length and so become wider than high at maturity. The
crown described by Moore and Plummer is almost as large
as the medium-sized hypotype but is somewhat more juve-
nile in having a median line extending longitudinally up
the arms with less distinct lines parallel to the main ridge.
The surface ornamentation of the arms is reportedly faint
in the proximal areas of the original type and is entirely
absent in the two hypotypes. It is not unusual for a longi-
tudinal raised ray on the arms to fade out with growth.
Of interest in ontogeny is the increase in prominence of
the basal concavity with age. In young forms there is little
depression of the base but the increased length of the
down-flared portion of the basals in growth develops a mild
concavity. The process is outlined under the section on
Morphological Notes.

Types. — Holotype, Harris Collection, No. H-3; para-
types, Nos. H-5, H-6 and H-10; collected by Mrs. G. W.
Harris, Waco, Texas. Paratypes and one hypotype, Marrs
Collections, Nos. M-10, M-11, M-12, M-13, M-14, M-20,
M-26, M-29, M-38, and M-43, hypotype M-6, collected by
Mrs. W. R. Marrs, Austin, Texas. Hypotypes, USNM, No.
$5145, $5182 collected by Wm. T. Watkins, reposited U.S.
National Museum. Two unnumbered hypotypes collected
by H. L. Strimple reposited in U.S. National Museum.
Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; 3 miles southwest of Brock, Parker County,
Texas.

SCIADIOCRINUS LLANOENSIS Strimple and Watkins,

new species
Plate 49, figures 9, 10

Dorsal cup shallow, truncate bowl-shaped, wide, shal-
low basal concavity. Asymmetry is present in that the right-
posterior side of the cup is deeper than the balance of the
cup. Five infrabasals are confined to the center of the
basal concavity, extend only slightly beyond the round
proximal columnals and form a pentagonal-shaped disk.
Five petal-shaped basals form the gently sloped lateral
sides of the concavity, and only the distal tips of the right
posterior and right anterior basals are visible in side view

of the cup. Five radials are large mildly tumid, participat-
ing in the basal concavity, forming the basal plane and
curving upward to form the lateral walls of the cup. A
shallow notch is formed at the summit of the interradial
sutures. A shallow, subhorizontal groove is in position of
an outer ligamental furrow. The ligamental pit, located next
to the transverse ridge, is deeply impressed. The inner
articular facets as a whole are sloped outwardly. The
transverse ridge is narrow where it adjoins the ligamental
pit and ventrally it is bordered by a subtriangular depres-
sion. Well-defined oblique grooves extend from the base of
the intermuscular notch to the lateral extremities of the
transverse ridge which terminates well before the inter-
radial sutures are reached. Adsutural slopes are low and
ill preserved. Three anal plates are in normal (Primitive)
arrangment. Radianal is a long slender plate. The posterior
interradius is depressed which causes the cup to have a
hexagonal outline when viewed from above or below.
Measurements of the holotype in millimeters:

Width of cup (maximum) 23.0
Width of cup, posterior to anterior (minimum) 22.0
Height of cup 4.3
Ratio of height to width 0.18
Width of infrabasal circlet 6.4
Height of basal concavity, including stem impression 1.5
Length of basal (right anterior) 9.3*
Width of basal (right anterior) 5:5*
Length of suture between basals 1.1
Length of radial to transverse ridge (right anterior) 9.2*
Width of radial (right anterior) 5.5*
Length of suture between radials 6.1*
Length of radianal 6.3*
Width of radianal 3.0

*measurements taken along surface curvature.

Remarks. — Sciadiocrinus llanoensis is closely related
to Sciadiocrinus disculus Moore and Plummer, but the for-
mer has a proportionately lower cup. S. acanthoporus has a
deeper basal concavity than S. llanoensis. S. confertus has
more prominent basal plates than S. llanoensis. S. planulatus
has an ill-defined dorsal cup but appears to have radial
plates in contact with infrabasals in some rays.

The species name Jlanoensis has reference to the Llano
Uplift from which it was obtained.

Holotype. — USNM, No. $5170, collected by Wm. T.
Watkins, reposited in the U.S. National Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch near
Mason, Mason County, Texas.

Genus POLYGONOCRINUS Strimple, 1961a

Type species —Polygonocrinus multiextensus Strimple,

1961a.

214 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Range. — Desmoinesian, Pennsylvanian; North Amer-
ica.

Remarks. — With the discovery of another species of
this genus we are able to obtain a more comprehensive
understanding of the form. The original description of the
genus allowed for the presence of partially biserial arms but
P. libratus, n. sp. has some sections of the arms entirely bi-
serial (especially secundibrachs) yet the distal arms (terti-
brachs and quartibrachs) are composed of almost entirely
cuneiform segments. The secundibrachs are reduced to a
robust, quadrangular first secundibrach and a robust ax-
illary second to fourth secundibrach in most rays. When
present the small, intervening SBrBr appear to be thin,
wedge-shaped elements of a biserial nature. Although there
is no existing evidence to substantiate or explain the half
biserial, half cuneiform arms we believe the upper arms
reflect an arrested juvenile stage. We believe that younger
species of the genus will be likely to have completely bi-
serial arms albeit the younger P. multiextensus is only
sparodically biserial.

Some characteristics of Polygonocrinus indicate rela-
tionship with Schedexocrinus Strimple, 1961a. The later
genus has a tendency to develop biserial structure among
the secundibrachs and sporadically elsewhere but funda-
mentally has uniserial arms. Only the first two bifurcations
of the arms in Schedexocrinus are isotomous, but there are
three in Polygonocrinus. The terminating platform of the
anal tube is almost identical for Schedexocrinus and Poly-
gonocrinus and both have nonspinose axillary first primi-
brachs, Major differences lie in the structure of the dorsal
cups. In Polygonocrinus the dorsal cup is extremely shal-
low with the infrabasals and basals downflared and _re-
stricted to the large basal concavity. The radials are sub-
horizontal with the distal ends forming the lateral sides of
the cup. The radials of Schedexocrinus are erect and even
the distal tips of the tumid basals are visible in side view
of the cup.

Pirasocrinus has a dorsal cup with characters close to
those of Polygonocrinus, i. e. infrabasals and basals are
down-flared and confined to basal concavity, radials have
proximal portions participating in the basal concavity,
primibrachs are tumid but nonspinose, arms are isotomous
for three bifurcations and both have relatively large termi-
nating platforms of the anal sac. Differences are that
Pirasocrinus has a long, slender crown (compared to the
broad, short crown of Polygonocrinus), the radials are
tumid, the arms are strictly uniserial and the terminating
platform has a relatively restricted area with fewer center
and perimeter plates.

POLYGONOCRINUS LIBRATUS Strimple and Watkins,

new species
Plate 33, figures 1, 2; Plate 56, figure 7

This species is represented in the collections by two
large crowns which are in fairly good preservation.

The dorsal cup is wide, low, truncate bowl-shaped with
a broad, deep basal concavity, It has a pentagonal outline
when viewed from below. The suture lines are well defined
by being mildly impressed. Five infrabasals are confined to
the base of the concavity and are subhorizontal to slightly
downflared. The five basals are strongly downflared, com-
pletely confined to the basal concavity so that even the
distal tips are not visible in side view of the cup. The
radials are subhorizontal with proximal portions in the basal
concavity and only distal ends curved upward to form al-
most imperceptable lateral sides of the cup. The basals are
large and form a petal-like circlet. The radial plates are
broad, slightly convex transversely, pentagonal-shaped with
only a mild notch at the interradial sutures and form a
gaped suture with the first primibrachs. The posterior in-
terradius is poorly preserved and in fact almost obliterated
in preservation in both specimens, The paratype shows that
anal X was in contact with the posterior basal so it can be
assumed that three plates were present in reasonably nor-
mal (Primitive Type) arrangement.

From the arms which are preserved, it is calculated
that 60 arms were formed which are fundamentally biserial
until the quartibrachs are reached and thereafter are com-
posed of cuneiform segments. First bifurcation is on the
low, thick appearing first primibrach. It forms a point
but could not be termed a real spine. The first secundibrach
is large, quadrangular-shaped and subsequent secundibrachs
are thin or absent except for the axillary secundibrach
which is a large, tumid, spinelike element. The first terti-
brach is a large quadrangular plate, but thereafter the
brachials change drastically to become thin, wedge-shaped,
and interlocking (biserial). As few as six or as many as
13 tertibrachs precede the next axillary wherein bifurca-
tion takes place in all arms. Another bifurcation in outer
arms of each half-ray only, which arms maintain consider-
ably more width than inner arms, takes place on or about
QBry-5. Tertibrachs and quartibrachs are slightly wedge-
shaped but the arms are not biserial.

There is a large, umbrella-like terminating platform
of the anal sac. All plates are not preserved, but there
appears to have been about 28 central plates and some 14
outwardly directed, spinelike perimeter plates. Some of the

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 215

central polygonal plates are small but many attain con-
siderable size (e.g. 9.5 by 7.3 mm.)

Measurements in millimeters as follows:

Holotype Paratype

Length of crown 57.0 _—
Width of crown (excluding platform) 43.0 —
Height of cup, to transverse ridge 5.74 3.5
Width of cup 30.0 37.08
Ratio H/W of cup 0.19 0.09
Width of basal concavity 13.68 22.5
Length of basal — 8.7*
Width of basal — 7.5*
Length of radial 11.7* 10.5*
Width of radial 119:0* 20.0*

4.—approximate
d.—distorted
*.—measurements taken along surface curvature.

Remarks. — Polygonocrinus libratus is distinct from
P. multiextensus of the Holdenville Formation in having
biserial arms up to the quartibrachs, thereafter the brachials
become cuneiform. In P, multiextensus the small secundi-
brachs show tendencies toward a biserial arrangement and
other biserial developments are only sporadic. The basal
plates are more tumid in P. multiextensus and prominent
notches are formed at the interradial sutures which are
distinct when the cup is viewed from below.

The specimen figured as a paratype of Plaxocrinus per-
undatus by Moore and Plummer (1940, pl. 15, fig. 5) is
reported on the plate legend as being from the same loca-
tion as the holotype, 1.2. shale below the Kickapoo Falls
Limestone Member, about 14 mile below Kickapoo Falls,
but with a question mark. In the text they report it from
the Brannon Bridge Limestone Member, three miles south-
west of Brock, Texas, which judging from the preservation
is probably correct. The iliustration is an oblique view of
the cup and therein somewhat obscure, but we feel certain
it belongs with Polygonocrinus, especially when the lower
arm segments are compared with those of P. libratus.

Types. — Holotype, USNM, No. $5105, collected by
Louis Todd, paratype USNM, No $5201 collected by Wm.
T. Watkins, reposited in the U.S. National Museum.

Occurrence. — Brannon Bridge Limestone Member,
Millsap Lake Formation, Strawn Group, Desmoinesian,
Pennsylvanian; three miles southwest of Brock, Parker
County, Texas.

Genus SCHISTOCRINUS Moore and Plummer, 1940

Type species. —Schistocrinus torquatus Moore and
Plummer, 1940.

Range.— Atokan to Desmoinesian, Pennsylvanian;
North America.

Diagnosis. — Shallow cone-shaped cup, five large infra-
basals have a convex curvature although they are not al-
ways visible in side view of the cup, five basals are small,
essentially triangular-shaped and are separated from each
other by the proximal ends of the radials. Five radials are
large and contact the infrabasal circlet, three anal plates
are elongated, a large balloon-shaped anal sac is reported
for S. azygous from the Holdenville Formation (Desmoin-
esian).

Remarks. — As demonstrated by Strimple (196la, pp.
99-101) the genus Schistocrinus is represented by three
species, S. torquatus, S. parvisculus (Moore and Plummer)
and S. azygous (Strimple) to which is now added S. brook,
n. sp. The species described as Schistocrinus confertus
Moore and Plummer and S. planulatus Moore and Plum-
mer may be conspecific and in any event have the charac-
teristics of Sciadiocrinus.

A new form, Schistocrinus acclivis, from the Soldiers
Hole Member, Big Saline Formation, has all the appear-
ance of a typical Schistocrinus. It is closely related to
S. azygous.

SCHISTOCRINUS ACCLIVIS Strimple and Watkins, new species
Plate 50, figures 2-4

Dorsal cup is shallow, cone-shaped. Five large infra-
basals form a large, slightly convex disk with the columnar
attachment area depressed and with an irregular outline
because of the unusual contact with the radial plates. Five
basals are small and triangular-shaped except where they
are in contact with the anal plates, which require extra
facets. Five radials are large broad plates. The articular
facets are obscured but are found to be inclined sharply
outward, have a heavily crenulated outer ligamental area,
and a ligamental pit backed by a transverse ridge. The
intermuscular notch is pronounced. The cup plates are
tumid and there is a sharp delineation at the meeting of
the infrabasals and basal circlet. Three large anal plates
are in normal (Primitive) arrangement.

The stem attachment cicatrix is round and well marked
with fine ridges. A depressed area extends beyond the
columnar scar.

Measurements of the holotype in millimeters:

Width of dorsal cup

Height of dorsal cup, to transverse ridge
Ratio of height to width

Width of infrabasal circlet

Length of basal (right anterior)
With of basal (right anterior)
Length of suture between basals
Length of radial, to transverse ridge
Width of radial (right anterior)
Length of suture between radials
Length of radianal

w

*

* *

SONI EONS re O81,
Bue kON HQ

216 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Width of radianal 2.1
Length of anal X cy
Width of anal X 2.8
Diameter of stem scar 2

*measurements taken along surface curvature.

The name acclivis is Latin for steep with reference to
the steep articular facets of the radial plates,

Remarks. — Schistocrinus acchvis of Atokan age is
more comparable to S. azygous than to S. torquatus, both of
which are of Desmoinesian age. In S. azygous, anal X and
RX attained a comparable height and shared equally in
supporting a prominent plate above which is a specialized
arrangement. In S. acclivis, RX extends higher than anal
X which is a normal arrangement. In the later species,
anal X and RX share in support of a large anal plate above.

Holotype. —USNM, No. $5171 collected by Wm. T.
Watkins, reposited in the U.S. National Museum.

Occurrence. — Soldiers Hole Member, Big Saline For-
mation, Atokan, Pennsylvanian; Kurt Zesch Ranch near
Mason, Mason County, Texas.

Family PHANOCRINIDAE Strimple and Watkins, new family

Genera. — Phanocrinus Kirk; Pentaramicrinus Sutton
and Winkler.

Range. — Genevievian and Chesterian, Mississippian;
North America and Great Britain.

Diagnosis. — Crown slender, compact, cylindrical; cup
low, bowl-shaped with slight or decided basal concavity;
IBB five, downflared or subhorizontal, small; BB five,
medium to large, always visible in side view of cup but
proximal portions participate in formation of basal plane
and concavity when present; RR five, large, wide; XX
three, normal (Primitive Type) arrangement to advanced
arrangement; arms ten, uniserial, branching on first primi-
brach in all rays in Phanocrinus and reduced to five arms
in Pentaramicrinus.

Remarks. —We are cognizant of a form which has
only nine arms but have not attempted to isolate it from
those which may have less than ten arms adyentitiously.
The exact affinities of the phanocrinids are somewhat ob-
scure but we are reasonably certain it originates in the
Scytalocrinidae rather than the Eupachycrinidae as has
been commonly thought. There is some question at this
time as to what forms among the Erisocrinidae are really
derivatives of the phanocrinids and we are not prepared
to attempt a comprehensive study into that matter.

Genus PHANOCRINUS Kirk, 1937
PHANOCRINUS TRULLEUM Strimple and Watkins, new species
Plate 36. figures 1-3; Plate 50, figure 5

Crown is of medium length and is slender. Dorsal cup

is medium bowl-shaped with truncate base and a _ pro-
nounced basal concavity. Five small infrabasals are confined
to the proximal portion of the concavity and are down-
flared. Five large basals are tumid, participate strongly in
the basal concavity and flex sharply to form a substantial
portion of the lateral walls of the cup. Five pentagonal-
shaped radials are tumid and only slightly wider than long.
The right posterior radial has an extra side because of its
long contact with the radianal. Three anal plates are in
normal (Primitive) arrangement. The posterior interradius
is mildly convex so that the rounded contour of the cup is
undisturbed in this area. The proximal columnals are large
and round. There are ten uniserial arms branching with
the axillary first primibrach in all rays, The first primi-
brach is slightly smaller than the radial. The anterior bi-
furcation is wider than the posterior in both of the pos-
terior rays. First and second secundibrachs are high but
subsequent brachials are very short. Sporadic biserial areas
may be developed in the distal arms. Each brachial bears a
single thin pinnule, on alternate sides.

Measurements of holotype in millimeters (approxi-
mate):

Length of crown 50.0
Height of cup 8.0
Width of cup 17.0
Width of basal concavity iD
Length of basal (right posterior) 8.7*
Width of basal (right posterior) 7.8*
Length of suture between basals 4.1*
Length of radial (left anterior) 5.9*
Width of radial (left anterior) 235%
Length of suture between radials 4.0

*measurements taken along surface curvature.

Remarks.— This species does not follow the normal
trend found among late Chesterian species of Phanocrinus,
i.e. lowering of the cup height and widening of the cup,
e.g. Phanocrinus alexanderi Strimple. The walls of the cup
of P. cooksoni Laudon are erect and the posterior inter-
radius is normal (Primitive), but the cup is broad and pro-
portionately lower than found in P. trullewm.

The arms of P. trullewm are essentially uniserial but
are changing to a biserial structure in distal portions. On
the inner sides of the arms, the pinnular bearing side of
the brachial has an increased height for the attachment of
a pinnule, but the opposite side is short and in some in-
stances terminates before the inner edge of the arm is
reached. The species may represent the lineage leading to
forms like Erisocrinus.

The species name is from the Latin word trullewm
meaning basin,

Types. — Holotype, USNM, No. $5115 collected by

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS PAS

Wm. T. Watkins paratype, USNM, No. $5172 collected
by Wilburn Shearer, reposited in the U.S. National Mu-
seum.

Occurrence. — Limestone in the Barnett Formation,
Chesterian, Mississippian; Kothman Ranch near Mason,
Mason County, Texas.

Order DISPARATA Moore and Laudon, 1943
Family ALLAGECRINIDAE Carpenter and Etheridge, 1881
Subfamily ALLAGECRININAE Moore, 1940

The family Allagecrinidae was revised by Moore, 1940.
At that time the genus Wrightocrinus Moore, 1940 was pro-
posed with Allagecrinus jakovlevi Wanner, 1929, from the
Permian as the type species. Moore reported the species
to have a cup height from 2.9 to 6.8 mm. The species Allage-
crinus biplex Wright, 1932, from the Lower Carboniferous
(Mississippian) was also assigned to the genus Wrighto-
crinus by Moore. The genus was distinguished from Allage-
crinus on the basis of whether the right posterior radial
has one arm or more than one arm and whether mature
individuals retain the oral plates. Al/agecrinus consistently
has a single arm facet on the right posterior radial but
Wrightocrinus has two or more at maturity. The oral
circlet loses its prominence with maturity in most species
of Allagecrinus and are in fact seldom preserved in matur-
ity. Information concerning Allagecrinus biplex was inade-
quate at the time of Moore’s study (1940) because in 1941
Wright gave a revision of the species wherein a different
concept is attained. Larger specimens are now known with
more arms, and the oral circlet is seen not to be preserved
in maturity, even as found in species of Allagecrinus. A con-
sistent character of A. biplex is the presence of an anal series
resting in a notch of the left shoulder of the right pos-
terior radial plate. Wrightocrinus jakovlevi does not have
an anal notch and anal plates or anal plates and Moore
(1940, p. 93) noted, “It is not surprising that an anal
plate or plates, present in the lower Carboniferous species,
should have disappeared in the Permian species.” But he
did not consider this of generic importance, “The presence
or absence of anal plates, which is normally a character of
generic importance, does not seem to be so in this case.”
Conversely, we believe the presence of an anal plate in
Allagecrinus biplex and the absence of an anal plate in
Wrightocrinus jakovlevi is of primary generic importance.
We propose the name Thaminocrinus new genus, with Alla-
gecrinus biplex Wright the type species of the genus.

A complete study of allagecrinids from the Permian
has not been made, but it appears that no forms assigned
to Wrightocrinus or to Allagecrinus have an anal plate or

plates and that all have the oral circlet prominently pre-
served. Pennsylvanian species formerly assigned to Allage-
crinus have been referred to Jsoallagecrinus Strimple
(1966b). They lose the oral circlet with maturity, or at
least the circlet does not survive in preservation. It might
also be noted that among those species from the Permian
of Timor for which the arms are known, the arms are short
and rest in grooves of the oral circlet. The grooves between
oral plates do not appear to serve any function other than
to act as receptacles for the arms, when the arms are not
extended. Advanced Permian forms have been assigned to
the genus Metallagecrinus Strimple (1966b). A new species
of Jsoallagecrinus from the Marble Falls Formation of
Texas is described as /. erectus, n. sp.

Genus THAMINOCRINUS Strimple and Watkins, new genus

Type species. — Allagecrinus biplex Wright.
Range. — Lower Carboniferous (Mississippian); Scot-
land.

Diagnosis. —Cup low with low basal circlet, sutures
usually invisible; RR unequal in width, form most of the
cup height; LAR and RAR narrower than other RR; LAR
supports one arm only, RAR usually one but occasionally
two arms; other RR have variable number of arms; right
shoulder of RPR usually axillary and supports two up to
five arms; there may be as many as 19 arms or more; left
shoulder of RPR is beveled and supports an armlike series
of anal plates; oral dome low in immaturity, missing in
maturity; column round, tapers rapidly.

Remarks. — Comparison of Thaminocrinus biplex with
Wrightocrinus jakovlevi has been given under the familial
discussion. There is a more intimate relationship indicated
with Allagecrinus austini but from which Thaminocrinus
biplex is readily separable because although it is a smaller
form (height about 1.5 mm) it has more arms, and it has
multiple arms on the right posterior radial, which plate
also carries an anal plate in a V-shaped groove in its left
shoulder. Thaminocrinus has a cup-shape close to that of
Allagecrinus austinii, the type species of Allagecrinus, for
which the cup height of adult specimens is reported to
range from 2.5 to 3.0 mm.

The generic name is derived from the Greek word
thaminos for crowded, with reference to the multiple arms.

Genus ISOALLAGECRINS Strimple, 1966b

ISOALLAGECRINUS ERECTUS Strimple and Watkins,
new species

Plate 41, figure 2; Plate 52, figures 7-11

This species is represented in the collections by two
partial crowns and a set of arms.

218 PALAEONTOGRAPHICA AMERICANA (VI, 40)

The dorsal cup is relatively high for a Pennsylvanian
representative of the genus with deeply impressed sutures
between radials but no appreciable tumidity, Cup height of
the holotype is 2.0 mm, width 2.6 mm. The basal disk is
undivided and higher than normal for other species of
Pennsylvanian age, There are five radials forming most
of the cup height. They are unequal and two (LAR and
RPR) each carry a single arm. The RPR also supports
an elongated anal plate in a notch on its left shoulder. The
other three radials (LPR, RAR and AR) each support
two arms. A total of eight arms and an armlike anal series
is present, and it is reasonable to expect larger specimens
would have more arms. The arms are slender and some
distal brachials show a tendency to have a slight medial
constriction. There is no great differentiation in width of
the arms of the various rays.

The proximal portion of the column is rapidly tapered,
curved, and composed of thin segments having a circular
outline. The entire surface of the crown is mildly granular
appearing under slight magnification. High magnification
shows the entire surface to be marked by minute pits.

Remarks. —Isoallagecrinus erectus has a cup shape
more comparable to some Mississippian forms than to
Pennsylvanian species, 7. ¢. it has a relatively high cup, with
the infrabasals contributing appreciably to the cup height.
Mississippian species of Allagecrinus usually have an elon-
gated cup with the infrabasal circlet prominent in side
view of cup. Mature specimens of A. austin have a low
cup but the lateral sides flare outward in maturity. A
syntype (G, S. Edin. 1602) of A. austinu figured by Wright
(1952, pl. 39, figs. 1 to 5) appears to be of comparable
size and at a comparable stage of development. It has a
ratio of H/W of 0.65 and eight arm facets distributed as in
Tsoallagecrinus erectus. Wright noted that A. austinu sel-
dom exceeds a cup height of 3 mm. The age of the Scot-
tish species is Viséan (Coral Zone 3 of D. Hill), Lower
Limestone Group. Mississippian species of Adlagecrinus from
North America do not appear to attain a size comparable
to large specimens of A. austinti, or at least have not been
recorded. Specimens of Allagecrinus americanus Rowley
from the Louisiana Limestone (Lower Mississippian) have
been reported by Strimple and Koenig, 1956, with as many
as nine arm facets. One specimen with eight full facets
and one incipient facet was figured by Strimple and Koenig
(1956, text-fig. 4, 28-31) which specimen had a H/W ratio
(excluding orals) of about 1.36.

Allagecrinus sculptus Strimple and Koenig, 1956, of
Osagean age, is somewhat like /soallagecrinus erectus in
general appearance of the cup, but is proportionately

longer (H/W ratio 1.58) and is a smaller form (height of
holotype 1.69 mm., including orals), The holotype has
only six arm facets, the double facet being on the left
posterior radial.

There has been no report of Allagecrinus from Mor-
rowan rocks although Moore (1940) noted that some speci-
mens he considered to be the young of Allocatillocrinus
rotundus could not be distinguished from Allagecrinus. His
“young” specimens are more like later Pennsylvanian
species in that they have considerable bulbosity of the radial
plates and proportionately low infrabasals.

The only described species from Pennsylvanian rocks
having erect, or elongated cups are /soallagecrinus copani
(Strimple), 1949b (H/W 0.69) from the Missourian, J.
pecki (Moore), 1940 (H/W 0.60) from the Desmoinesian
and /. graffhami (Strimple) 1948 (H/W 0.65) from the
Virgilian. 7. copani has only seven arms for a specimen with
a height of 3.7 mm, whereas /. erectus has eight arms at a
cup height of 2.0 mm and is proportionately taller. 7, pecki
has seven arms at a cup height of 3.5 mm and the radials
are bulbous. /. graffhami has 12 arms at a cup height of
2.0 mm.

Types. — Holotype, USNM, No. S5178, paratype
USNM, No. $5133, collected by Wm. T. Watkins, reposit-
ed in the U.S. National Museum.

Occurrence — Lemons Bluff Limestone Member, Mar-
ble Falls Formation, Atokan, Pennsylvanian; Rough Creek,
11 miles southeast of San Saba, San Saba County, Texas.

Class CRINOIDEA Miller
Subclass CAMERATA Wachsmuth and Springer
Order MONOBATHRA Moore and Laudon

Family PLATYCRINITIDAE Basselr, 1938
Genus PLATYCRINITES Miller, 1831

Type species. — Platycrimtes laevis Miller.

Range. — Silurian to Permian; Europe, Australia, Rus-
sia, Indonesian, North America, Asia.

In Bassler and Moodey, 1943, there are 140 species
or subspecies of Platycrinites accepted as valid from Mis-
sisippian (Lower Carboniferous) rocks of Ireland, Belgium,
England, Ile of Man, Scotland, Germany, and North
America. There are 12 species or subspecies listed from older
Silurian and Devonian rocks of Germany, England, and
North America. Five species or subspecies are from Permian
rocks of Australia and the Island of Timor. The only

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 219

form listed by Bassler and Moodey from rocks of Pennsyl-
vanian (Upper Carboniferous) age is Platycrinites spitz-
bergensis (Holtedahl) from the Moscovian of Spitzbergen,
Norway. This species is based on two fragments, one of
which is a radial plate and the other a basal circlet. There
are also some stems and columnals ascribed to the species.

PLATYCRINITES REMOTUS Strimple and Watkins, new species
Plate 47, figures 8, 9

This species is based on a single crown that is horizon-
tally compressed but otherwise is in good preservation. This
is apparently the first calyx of Platycrinites known from
Pennsylvanian rocks.

Study of the shape of the cup indicates a medium high
dorsal cup with mildly upflared basals and erect lateral
sides formed by the radials. All cup plates are thin and
are mildly granular appearing under slight magnification.
There are three basals of moderately large size forming a
pentagonal disc with a large, elliptical-shaped columnar
scar in mid-portion. The outer edge of the disc has a slight
notch at the interbasal suture so that two facets are
formed on the proximal edge of the adjacent radial plate.
Where the interbasal suture is absent the adjacent radial
has an uninterrupted proximal edge. Five radial plates are
large, wide elements with lateral sides slanted out and up-
ward so the greatest width is near the summit of the cup.
The arm attachment area is notched well below the summit
of the cup and is crescentic in shape. The first primibach
is axillary and is low but with short lateral sides. The
second secundibrach is apparently axillary in all arms.
There is a distinctive sinuous suture between all preserved
brachials. A large interradial plate is found in the posterior
interray and is only separated from the anus by two series
of plates. There is a low tube or projectd area surrounding
the anus. Tegmen plates are mildly tumid and are rather
large polygonal elements.

Measurements of holotype in millimeters:

Width of cup as preserved 35:7"
Width of basal circlet as preserved 19.6
Length of basal 10.0
Length of radial (to articular facet) 10.3
Length of radial (to summit) 12.7
Width of radial 19.5

*grossly distorted.

Several disassociated Platycrinites-type columnals have
been found at the Kickapoo Falls locality but they are con-
sidered separately.

Remarks. — A search of the literature has revealed no
species of Platycrinites of Pennsylvanian age based on other
than fragmentary material. P. spitzbergensis (Holtedahl)

from the Moscovian of Norway is known from a basal
circlet and one radial plate. Some stem fragments are
ascribed to the species. The radial plate reflects constriction
of the cup summit in that the lateral sides slope toward
each other whereas in P. remotus they slope away from
each other indicating a widening of the cup at the summit.
Series of nodes are present on P. spitzbergensis and on the
radial plate they show a tendency to converge below the
arm articular facet. P. remotus is marked by granules but
they are only visible under magnification.

There are no species of Platycrinites known to us from
upper Chesterian so close comparison with antecedent forms
is not possible. An arm of P. pentcillus is figured by Wachs-
muth and Springer (1897, pl. 53, fig. 76) with sinuous su-
tures between the lower segments. This species is representa-
tive of the Genevievian Stage of the Valymeyeran Series af-
ter Swann (1963). The lateral sides of the cup are almost
vertical which is reflected by the almost vertical sides of the
radials in P. penicillus. The first primibrach in P. penicillus
has no lateral sides, whereas the first primibrach in P. re-
motus has short lateral sides.

The Permian form described as Platycrinites wachs-
mutht (Wanner), 1916, and subsequently divided to sub-
species P. wachsmuthi typicus = P. wachsmuthi, P. wachs-
muthi agnaensis (Wanner), 1937, and P. wachsmuthi fre-
guentior (Wanner), 1937 do not have as many large poly-
gonal plates in the tegmen and apparently had taller and
narrower calyxes than P. remotus. The later species is also
distinctive in having lateral sides to the axillary primibrach,
which plate is bordered by the first secundibrachs in Wan-
ner’s species with the lateral sides eliminated. There 1s
also a distinctive sinuous suture between brachials in P.
remotus not found in the Permian forms.

Platycrinites wrighti Oyens, 1940, also from the Per-
mian of the Island of Timor, has a strongly tubercled sur-
face of dorsal cup and is essentially a narrower and higher
cup than P. remotus is thought to have.

Yakovlev (1956, pl. 12, fig. 8) illustrated a radial
plate as Platycrinus sp. which has a highly ornate surface
and two raised rays connecting the arm articular facet
with the two proximal corners of the plate. It is reported
as questionably C, (Upper Carboniferous) — Missourian
or Virgilian, S.S.S.R. The lateral sides of the radial are
directed outward which would produce an erect or mildly
expanded cup at its summit. Yakovlev (1956, pl. 10, figs.
14-20) proposed a species Platycrinites permiensis (Yakoy-
lev) from P, (lower Permian) of the SSSR based on one
radial plate and several columnals. The radial plate is
thick and the lateral sides slope toward the upper center

220 PALAEONTOGRAPHICA AMERICANA (VI, 40)

of the plate which would probably produce a mildly con-
stricted summit of the dorsal cup. It is at best questionably
referable to Platycrinites. The stem fragments are compar-
able to other known columnals which have been referred
to the genus. They are thick,

The species name remotus is Latin for “far off” with
reference to the great lapse of time between it and typical
representative of the genus.

Holotype. —USNM, No. $5163, collected by Wm. T.
Watkins, reposited in the U.S. National Museum.

Occurrence. — Shale below the Kickapoo Falls Lime-
stone Member, Millsap Lake Formation, Strawn Group,
Desmoinesian, Pennsylvanian; about 14 mile east of Kicka-
poo Falls, southwest of Dennis, Hood County, Texas.

Family ACROCRINIDAE Wachsmuth and Springer, 1885

The premise followed by recent authors has been that
Acrocrinus is derived from Dichocrinus. Wachsmuth and
Springer (1897, p. 804) stated it as follows, “The introduc-
tion of a narrow belt of supplementary’ plates between the
basals and radials would be sufficient to transform any
Dichocrinus into an Acrocrinus.” There is reason to believe
that Acrocrinus, as represented by the type species, is en-
tirely independent of Dichocrinus; however, it may be that
some forms currently ascribed to Acrocrinus are derived
from dichocrinid stock.

Moore and Plummer (1938, p. 217) stated as follows:
“The structure of the calyx of Acrocrinus has been inter-
preted as indicating a very interesting reversion from a
specialized, simplified calyx of the type of Dichocrinus, its
presumed ancestor, to a primitve many-plated cystid-like
form in which the several circlets of plates lack a definite
plan, as is the rule among crinoids (Jaekel, 1918, p. 39;
Springer, 1926, p. 41.)”

Moore and Laudon (1943, p. 96) stated as follows,
“The Acrocrinidae (fig. 16) are derived from Dichocrinus.
They might almost be characterized as a group of over-
specialized crinoids that had reverted to a more primitive
type. Inasmuch as no other crinoids are known that re-
motely resemble Acrocrinus in structure, this group must be
considered as a very unusual specialized derivative of the
Dichocrinidae.”

It is beyond the scope of this paper to revise the
Acrocrinidae and Dichocrinidae but we do feel it is desir-
able to establish more realistic groupings.

7The term “intercalaries” proposed by Moore and Plummer, 1938, is
used herein.

Genus GLOBACROCRINUS Moore and Strimple, 1969

Type specites.— Acrocrinus pirum Moore and Plum-
mer, 1937 (1938):

Range. — Pennsylvanian; North America and Russia.

Diagnosis. — Dorsal cup elongated, swollen at base or
mid-section with constricted summit, base flattened but
distal tips of basals may be visible in side view of cup;
two large basals of equal size, with suture marking the pos-
tero-anterior axis; five radials and anal X form a circlet
at summit of cup, articular facets of radials narrow, horse-
shoe-shaped, anal X has a crescentic facet with an appear-
ance close to the arm articulating facets; all radials have
two intercalaries at their proximal side except the anterior
which has three; anal X has three intercalaries in contact
with the proximal side, the center one being designated as
X1; there are four or more series of plates (intercalaries)
between basals and radials; the tegmen has not been ob-
served for any known species of the genus.

Species assigned to Globacrocrinus:

Occurrence

Atokan; Texas
Morrowan; Okla.

Globacrocrinus multiplicatus, n. sp.
Acrocrinus pirum Moore and Plummer

Remarks. — This genus is primarily characterized in
having a flattened or concave base extending beyond the
columnar attachment scar but with distal edges of basals
curved up to be visible in side view in some species, in
having an expanded mid-section or proximal area, a distal
constriction and with normal radial plates in proximal con-
tact with two plates (anterior radial is in contact with
three). The Russian species, A. myjatschkowensis is from
younger rocks (Moscovian) and has almost as many in-
tercalary plate series as G. pirum but lacks the expansion
at the base of the cup. The Moscovian species differs from
both the other mentioned species in that it does not have
an uninterrupted series of anal plates. In this regard it is
comparable to Acrocrinus praecursor from the Burlington
Limestone (Mississippian) which species has three anal
plates in continuous series below anal X but the series is in-
terrupted at X*. A. praecursor is excluded from Globacro-
crinus because of the cup shape wherein a continuous ex-
pansion of the cup takes place from the columnar scar to
the summit of the cup and the basals may be properly
termed upflared. The same cup shape is found in Acrocrinus
intermedius Springer, 1926, of Chesterian age, which species
is probably a direct derivative of A. praecursor.

Acrocrinus shumardi Yandell, 1855, and A. brentwood-
ensis Moore and Plummer, 1938, of Chesterian and Mor-
rowan ages respectively, are readily separable in having the

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 2

proximal edge of a normal radial plate in contact with four
intercalaries. Acrocrinus amphora Wachsmuth and Springer,
1897, of Chesterian age, A. wortheni Wachsmuth, 1882, of
Pennsylvanian age, A. pumpkensis Strimple, 1949a, and
A. expansus Strimple, 1951c of Desmoinesian age, and A.
elegans Strimple, 1949a, of Missourian age are readily sep-
arable from Globacrocrinus in having the normal radial plate
in contact with three intercalaries. The preservation of
specimens of Acrocrinus alvestonensis Wright (1958), is
too poor for precise examination, but it appears to be close
to A. amphora. The species is reportedly of Viséan age.

Acrocrinus primitivus Laudon and Beane (1937), from
the Hampton Formation (Kinderhookian) of Iowa, is not
considered by us to be a bona fide representative of the
genus Acrocrinus.

GLOBACROCRINUS MULTIPLICATUS Strimple and Watkins,
new species

Plate 52, figures 4-6

The cup plates are smooth, There are five series of
intercalaries between radials and basals and only four inter-
calaries (anal series), and these in direct series. The base
of the cup is essentially flat and expands rapidly to just
below mid-section thereafter tapering to a constricted sum-
mit. As previously noted, all radials except the anterior are
adjoined below by two intercalaries. The arms and tegmen
are not preserved. The columnar scar is circular and im-
pressed. Greatest width of the cup is 3.2 mm, width at
distal extremity is 2.5 mm, ight is 3.7 mm.

The name is taken from the Latin word multiplico
for increase, with reference to the numerous cup plates.

Remarks. — When one considers that plates adjacent
to the basals are incipient in this genus, it becomes obvious
that increase in age and size or size, will produce more
series of intercalaries. Without some knowledge of the
growth stages of a species considerable care must be taken.
There are only five series of intercalaries between the radials
and basals of Globacrocrinus multiplicatus in G. pirum
there may be as many as ten. G. multiplicatus is readily
distinguished from G. pirwm in that the basal area of the
later species is unusually wide.

Holotype. — USNM, No. $5179 collected by Wm. T.
Watkins, reposited in the U.S. National Museum.

Occurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Limestone Formation, Atokan, Pennsylvanian;
Rough Creek, 11 miles southeast of San Saba, San Saba
County, Texas.

Family DICHOCRINIDAE S§. A. Miller, 1889

We have by no means made an exhaustive study of all

to
=

interrelationships with Dichocrinus, but we are unable to
accept the prevailing concept of its evolution from the
Hexacrinitidae. In this we are not entirely alone for Wilson
(1916, p. 680, 681) considered Dichocrinus to have evolved
from a platycrinitid and specifically mentioned “Cocco-
crinus” (now Lyonicrinus) of Silurian age as the ancestral
type giving rise to both Platycrinites and to Dichocrinus.
If this is applied to families recognized by Moore and Lau-
don (1943) in their “Evolution and Classification of Paleo-
zoic Crinoids” it means both Platycinitidae and Dicho-
crinidae evolved out of the Thallocrinidae.

We recognize the fundamental difference in Platycrini-
tidae and Thallocrinidae having a pentagonal disk but we
follow Wilson (1916) and Strimple (1963, text-fig. 28) in
the belief that forms with a hexagonal base have evolved
from forms with a pentagonal base. Without going into all
of the involved factors, we believe the pressure of the gut
on the posterior side caused the animal to secrete additional
stereom on the proximal areas of the anal plate (which plate
is present in a notch at the summit of the radials in Thallo-
crinidae and Platycrinidae) in its early stage of develop-
ment so that it becomes a full plate of the radial circlet.
There was also an ensuing slight shift of the gut from the
right posterior to the direct posterior position and the suture
followed into the antero-posterior radius. Further transi-
tion of the basals requires the anchylosis of the small left
anterior basal with the left posterior basal thus eliminat-
ing the suture, and a widening and truncation of the pos-
terior edge of the basal circlet for support of the anal plate.

Genus DICHOCRINUS Minster, 1839

Type species. —Dichocrinus radiatus Minster, 1839.

Range. — Lower Carboniferous (Mississippian) and
Upper Carboniferous (Pennsylvanian); Europe, Russia,
North America.

Remarks. — The genus Dichocrinus is represented in
Bassler and Moodey (1943) by 50 species all of which are
of Mississippian age except one [Dichocrinus imsularis
(Eichwald), 1856] which is reputedly of Devonian age.
It was originally described as Platycrinus and has been re-
ferred on occasion to Hexacrinus (now Hexacrinites) where
we believe it should rest at least until a new study of the
species is possible. A few forms have been described subse-
quent to Bassler and Moodey’s bibliographic index as fol-

lows:

Dichocrinus quadriceptalus Laudon and Severson (1953)
Lower Carboniferous (Kinderhookian), North America;
Dichocrinus rotaii Yakovlev (1956) Lower Carboniferous
(C1), SSSR; Dichocrinus tomiensis Yakovlev (1956) Lower
Carboniferous (C:), SSSR; Dichocrinus unicus Wright
(1956), Lower Carboniferous (Tournesian), Eire,

tM
me
Ld

The genus is poorly defined and somewhat ambiguous
in the light of present information. A new species is de-
scribed from the Atokan as Dichocrinus dilatus, n. sp.

DICHOCRINUS DILATUS Strimple and Watkins, new species
Plate 36, figure 5; Plate 41, figures 3, 4

Crown elongated, mildly constricted at a slight distance
above the high cone-shaped dorsal cup. Two equal-sized
basals rise evenly to form a high cone approximately one-
half the height of the cup. Five radials are of unequal size
and are protuberant in the areas adjacent to the narrow,
horseshoe-shaped arm articulating facets. Anterior radial
extends into the intersutural area of the basal circlet. Right
posterior radial is the shorter and smaller of the plates.
The large anal plate occupies the posterior interradius and
has the general appearance of a radial except that it lacks
an arm articulating facet, and the resultant protrusion, and
that it is longer than any of the radials. It narrows toward
the summit and is followed by a large tube plate which
appears to be followed by a single series of smaller tube
plates. In one young paratype there is a single prominent
interbrachial plate in each interray. These plates are not
visibly preserved in any of the older paratypes but prob-
ably they were present and participated in the tegmen
structure,

The arms are long, slender, and the brachials are slight-
ly cuneiform. There are apparently ten arms, with bifur-
cation taking place on the second primibrach in all arms.
The anterior arm of the holotype is undivided as_pre-
served but because some of the lower brachials are out of
place it is not possible to determine whether this is a normal
condition. All arms are preserved in one paratype and
branch one time. The structure of the arms and pinnules,
and their preserved state, indicate that the arms could never
be tightly closed. Strong pinnules are observed as low as
the third secundibrach and probably occur even lower in the
arms. A large facet, for the reception of a pinnule, is found
on the side of the brachials, but never more than one to
each brachial, and always on the side opposite the adjacent
brachial.

The column is composed of relatively even columnals
which possess a sharp rim. It is round and slow tapering
and has been observed in considerable length.

Measurements in millimeters:

Holotype Paratype
Overall length of crown 12.1 —
Height of dorsal cup 2.74 4,74
Width of dorsal cup 3,14 4.69

Width of radia] (right posterior) 1.3 —

PALAEONTOGRAPHICA AMERICANA (VI, 40)

Width of arm articulating facet 0.6 a
4. distorted by lateral compression.
Remarks. — In a time sense, the closest representatives

to D. dilatus are of Chesterian age. D. pentalobus has a long
cup, smooth surface, narrow arm articular facets, PBroax,
20 arms which are biserial and pendent. It is from the Glen
Dean Formation. D. superstes has a slender long cup,
smooth surface, wide arm articular facets, PBroax, ten arms
which are biserial and rugged. It is from the Glen Dean
Formation. D. girtyi has a broad cup, ornate surface, small,
narrow articular facets of radials, PBr,ax, arms not pre-
served. It is from the Shelterville and Paint Creek Forma-
tions. D. dilatus does not appear to be closely related to
any of those species in that the surface of the cup is punc-
tate appearing, radial articular facets are medium wide,
PBroax, ten arms which are uniserial and slender.

It appears that D. dilatus is of a lineage typified by a
species like D. delicatus Wachsmuth and Springer (1897)
of Kinderhookian age, having retained or reverted to a
primitive state of ten uniserial arms bifurcating on PBreax.

Types. — Holotype USNM, No. $5134 and paratype
USNM, No. $5116 collected by Wm. T. Watkins, reposited
in the U. S. National Museum.

Ocurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Formation, Atokan, Pennsylvanian; Rough Creek,
11 miles southeast of San Saba, San Saba County, Texas.

Genus STOMIOCRINUS Wanner, 1937

Type species. —Stomiocrinus subglobosus Wanner,
1937.

Range. — Pennsylvanian and Permian; North America
and Indonesia.

Remarks. — The genus has previously been recognized
only from the Permian of the Island of Timor, Indonesian,
and typically has a relatively large columnar attachment
area with basals of moderate size rising evenly from the
stem cicatrix, The presently considered Pennsylvanian
species, Stomiocrinus conlini, n. sp., has larger basals which
form a bowl and the columnar attachment is small. Neo-
dichocrinus Wanner typically has wider articular facets and
the cup has an even expansion as compared to the con-
striction of the cup toward the summit in typical Stomio-
crinus.

STOMIOCRINUS CONLINI Strimple and Watkins, new species
Plate 40, figures 1-4

This species is based on three dorsal cups collected
by J. P. Conlin for whom the species is named. The senior

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

author obtained one specimen from Frank Crane several
years ago in exchange, and in the course of events the
junior author contacted Mr. Conlin who fortunately still
retained two of the three specimens and which he kindly
donated to the sudy.

The dorsal cup is more or less barrel-shaped with a
broad basal area and a constricted summit. There are two
circlets of plates, basals, and radials. Two large basals of
equal size form a high broad cup, but the median portion
is depressed and a small, delicate, round stem occupies the
center of the basal concavity. Five radials are longer than
wide and have narrow, rounded arm articulating facets.
The anal plate is in the same circlet and has the appearance
of a radial plate except the upper surface does not have a
true articular facet, although there is a mildly depressed
area which was no doubt occupied by a tegmen plate.

The articular facets of the radials are not well pre-
served and, considering the perservation of the facets of
related forms, there probably never was any appreciable
topography. In side view of the cup only the crescentic
shape of the facet is visible. The fore facet is subhorizontal
and depressed and is backed by a transverse ridge. Internal-
ly the facets slope inwardly at a sharp degree. The anterior
radial has a shape almost identical to that of the anal plate
except that it has a decided articular facet above. Both
plates are larger than other plates of the circlet, especially
in the proximal portions, and both have a decided apex
entering the notch formed at the suture line by the two
basals. The other four radials have a curved lower edge
with no central angle or apex. Because of the feature,
radials other than the anterior have only four sides. There
is a slight constriction at the base of the radial circlet
which accentuates the division of the two circlets.

The surface of the cup is smooth with no tumidity of
plates or impressions of sutures, other than between radial
and basal circlets as discussed above. The cup plates are
thick and adjoin one another closely, with no evidence of
interplate ligaments or crenulations. The basals are almost
entirely fused in one specimen,

Two or three proximal columnals are preserved in the
holotype and are thin elements. It seems likely the species
was free-swimming. The base is not greatly different from
that of some species of Paragassizocrinus, although no
relationship is inferred by us.

Measuremens in millimeters as follows:

Holotype Paratype
Height of dorsal cup 10.2 8.1"
Width of dorsal cup (maximum) 10.2 112"
Height of basal circlet 4.2 6.3

Width of basal circlet 9.6 10.8

i)
bt
>)

Width of radial plate (proximal) 4.1" 5.4*
Width of radia! plate (distal) 3.7* 4.5"
Width of anterior radial (proximal) 5.0* 7.9*
Width of anal plate (distal) 3.4* 3.1*
Width of anal plate (proximal) 5.9% 6.4"

a. approximate

*, measurements taken along surface curvature.

Types. — Holotype, USNM, No. $5127, paratypes, un-
numbered and USNM, No. S5128, collected by J. P. Conlin
of Fort Worth, Texas, reposited in the U.S. National Mu-
seum.

Occurrence. — According to Mr. Conlin, obtained from
shales below the Rockhill Limestone Member, Graford
Formation, Canyon Group, Missourian, Pennsylvanian;
southwest of Lake Bridgeport dam about three-fourths mile,
Wise County, Texas.

Class CRINOIDEA Miller
Subclass FLEXIBILIA Zittel
Order SAGENOCRINOIDEA Springer, 1913
Family SYNEROCRINIDAE Jaekel, 1918

Significe studies of the Synerocrinidae, (Synero-
Significant stud f the Sy) Jae, (Sy
crinus and associated forms) are discussed chronologically
as follows:

I. Trautschold, 1867, pp. 31-34
Forbesiocrinus incurvus Trautschold

The species was proposed with two specimens having
isotomous arms from what is now termed (CsM) Moscov-
ian, near Moscow, USSR, illustrated by Trautschold on
plate 4, figures 4, 5 and by a text-figure on page 31. The
smaller specimen was figured (figs. 5a-b) from the posterior
side and from the base.

II. Trautschold, 1879, pp. 28-29
Forbesiocrinus incurvus Trautschold

The specimen on plate 3, figure 11, is the same as that
in 1867, pl. 4, fig. 4. There is a text-figure on page 28
which is the same as that in 1867, page 31. A side and
summit view of a set of heterotomous, or more properly
termed bi-endotomous arms, is given on plate 4, figures 3a-b

and ascribed by Trautschold to F. incurvus.

Ill. Jaekel, 1898, p. 47
Synerocrinus Jaekel (1898)
Type species. — Forbesiocrinus incurvus Trautschold.
The genus Synerocrinus was proposed in a paper read
and recorded in the minutes of the Society and is found in
Section C. Verhanlungen der Gesellschaft (Zeitschrift
Deutsch. Geol. Gesell., 49, p. 47, 1897). As with much of

224 PALAEONTOGRAPHICA AMERICANA (VI, 40)

Jaekel’s work the proposal is casual, quote “Ich halte
deshalb eine generische Sonderstellung des (Forbesiocrinus)
incurvus Tr. fiir guechtfertigit and neune ihn Synerocrinus.”
He brought out the heterotomous nature of the arms and
the large interbrachial in discussion.

IV. Bather, 1900, p. 190, fig. CXI
Synerocrinus incurvus (Trautschold)

Bather’s specimen (fig. CXI) (presumably a topotype)

has bi-endotomous (heterotomous) arms which align it with
Jaekel’s definition of the genus, but it differs from other
material in having more than one large interbrachial above
the first bifurcation and it has a small series of inter-
brachials above the second bifurcation. Bather’s diagnosis
of Synerocrinus was as follows:
“Carboniferous, Europe, has arms like Dactylocrinus, ex-
cept that no ramules branch; it also differs in having 3-8
iBr, perhaps more, with occasional ilIBr and ilIIBr; x
rests on post. B (Figs. CX.5, and CXI)”. It is thus appar-
ent that he refers to bi-endotomous (heterotomous) arms
(his fig. CX.5) and has modified Jaekel’s concept to in-
clude numerous interbrachials above the axillaries of the
arms. As will be shown next, Jaekel considered the single
interbrachial (above the first dichotomy) of primary im-
portance because he specified “I grosses Interbrachials” in
his finished diagnosis.

V. Jaekel, 1918, p. 78

Family Synerocrinidae. “Armaste ausgepragt einseitig para-
tom, | grosses Interbrachiale, 5 Basalia, 3 Infrabasalia (Fig.
74B). Synerocrinus Jkl., Oberdevon-Russland. Origocrinus
Spring., Untercarbon, Nordamerika.”

Brief as the description is there is no doubt as to the
intents of Jaekel concerning the arm structure he had in
mind for he stated the arms are distinctly unbalanced and
his figure 74B referred to a drawing of bi-endotomous
(heterotomous ) arms.

Looking back to Trautschold’s diagnostic drawings,
there was no change between that of 1867 and that of 1879
for Synerocrinus incurvus, but even so there is no way of
knowing whether he intended to depict heterotomous arms
or not, because his drawing did not go past the third terti-
brachs and the branchings in question would start at this
point. There are two isotomous bifurcations in all the forms
considered.

Jaekel’s reference to “Origocrinus” apparently con-
cerns Oligocrinus Springer, 1906 which had as its type
species Forbesiocrinus asteriaeformis Hall and has subse-
quently been placed in synonomy under Onychocrinus. It is
strange he would have placed this form in the family

Synerocrinidae because it has exotomous branching and
must have been through a misunderstanding of its char-
acters.

VI. Springer, 1920, pp. 334-336, pl..42, pl. 75, fig. 12,13; text-fig 44
Synerocrinus incurvus (Trautschold)

Springer had at hand six specimens from the type
locality and stated he had casts of additional specimens
deposited in the Geological and Paleontological Museum in
the University of Moscow. Springer also figured two speci-
mens from the Viséan of Scotland (pl. 85, figs. 12,13) as
being conspecific,

The specimen figured on plate 42, figures 6a and 6 is
important because it shows the anal tube, which is a single
series of plates, not touching the arms to the right or left.
This is exactly the type of structure to be expected because
if there had been any support from contact with the arms
the plates would have likely been preserved in the small
original type (Trautschold, 1867, pl. 4, fig. 5) where a gap is
found above the single anal plate.

Most specimens illustrated by Springer (1920, plate
42) have been referred elsewhere to Talanterocrinus. Wright
(1946, p. 33) referred one specimen (pl. 75, fig. 12 of
Springer) to Caldenocrinus and the other Scottish speci-
men (pl. 75, fig. 13 of Springer) to Talanterocrinus.

VII. Laudon, 1937, pp. 706, 708, text-fig. 2
Synerocrinus farishi Laudon, 1937

This species is based on a damaged monotype of which
the base and posterior side is largely missing but it is a
large distinctive form. It is from the Atokan (Bendian),
Bostwock Conglomerate, 300 feet above the Otterville
Limestone Formation in the Ardmore Basin of southern
Oklahoma. From the evidence presented by Laudon it
appears this form has isotomous branching with the second
first primibrach and again with the third secundibrach
although his illustration is not convincing of the second
branching in some arms. Of considerable interest is the
existence of three or more interbrachials above the first
branching and three small interbrachials above the second
branching.

The arms are strongly rounded above the isotomous
branchings and become bi-endotomous (heterotomous).
There are often ramules or armlets on the inner side of the
arms on alternating brachials. The armlets would be ob-
scured if the arms were closed.

This species has been referred to the genus Talantero-
crinus by most modern authors, but we have referred it

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 225

back to Synerocrinus pending knowledge of the structure
of the posterior interradius.

VIII. James Wright, 1939, pp. 56-58, pl. 11, figs. 9, 10, 11, 16, 17;
text-figs. 75-77

Ainacrinus Wright, 1939

This is a Viséan form from Scotland in the group but
having dichotomous arm branching. Only one species is
known, A. smithi (Wright, 1934). There are several char-
acteristics which separate it from the other forms of the
considering here, the main one being the diverging nature
of the arms. Separation of rays may take place even below
the first bifurcation and just above the single interbrachial
(exception —a series of three interbrachials are found in
one ray of one specimen, starting at a notch between RPR
and RAR). This genus is reported to have a single series of
anal plates whch are suturally united with the adjoining
brachials.

IX. Moore and Plummer in Moore, 1939, pp. 195, 196
Synerocrinus Jaekel, Trautscholdicrinus Moore and Plummer

In the above cited study, reference is made to “Traut-
scholdicrinus Moore and Plummer, 1939, Mss. Univ. Texas
Bull., (in press)” which was apparently included in
Moore’s independent study to accommodate the description
of Trautscholdicrinus mcguirei Moore (1939). It also re-
quired the premature designation of a type species for
Trautscholdicrinus which was accomplished as follows,
quote “Genotype. — Trautscholdicrinus jaekeli Moore and
Plummer, from beds near Moscow, Russia, classed as be-
longing to the upper part of the Lower Carboniferous. The
specimen figured by Springer (1920, pl. 42, figs. 4a, 4b)
from the collection in the U. S. National Museum, is desig-
nated as holotype, called by Springer Synerocrinus incurvus-
(Trautschold).” This would have been a fortunate choice
of a holotype because it is well preserved and well illus-
trated and provides a fine basis for establishment of firm
generic characters except we believe it to be conspecific with
S. incurvus. When the University of Texas Publication No.
3945 appeared in 1940 there is no mention of the name
Trautscholdicrinus but the generic name appears as Talan-
terocrinus Moore and Plummer, n.g. as will be discussed
in the next section herein. One must assume they considered
their name Trautscholdicrinus to be a junior homonym of
Trautscholdicrinus Ivanov (1926, p. 176) a nomen nudum
or perhaps of Vrautscholdicrinus Yakovlev and Ivanov
(1939, p. 66), although the latter is unlikely because of the
time element.

Moore and Plummer (1939, pp. 195, 196) called atten-
tion to the fact that Jaekel (1918) ascribed a hetero-
tomous arm structure to the arms of Synerocrinus whereas
the original type specimens of the type species (Forbesio-
crinus incurvus) had isotomous arms. It is obviously the
intent of Moore and Plummer to restrict Forbesiocrinus
incurvus to the two types figured by Trautschold in 1867.

The bi-endotomous (heterotomous) form injected into
the material by Trautschold (1879), followed by Jaekel
(1897), augmented by a specimen figured by Bather (1900)
and specimens figured by Springer (1920) would be left
without a name in the restriction and Moore and Plummer
proposed Trautscholdicrinus jaekeli, as previously noted.
All of the forms indicated above are not conspecific with
T. jaekeh. Trautscholdicrinus jaekeli Moore and Plummer
(1939) is later taken as the type species of Talanterocrinus
Moore and Plummer (1940) but is erroneously designated
(1940, p. 97) as “Talanterocrinus jaekeli Moore and
Plummer, n. sp.” It should read Trautscholdicrinus jaekeh
(Moore and Plummer).

Trautscholdicrinus mcguirei Moore, 1939 is a well-pre-
served form from the Brownville Limestone Formation (up-
permost Virgilian) of Osage County, Oklahoma. It is trans-
ferred to Talanterocrinus by Moore and Plummer (1940)
but it is not a typical Talanterocrinus mainly because there
is no appreciable differentiation between the multiple plates
of the posterior interray and the normal interrays. The
species is referred by us to Aexitrophocrinus, n.g.

X. Moore and Plummer, 1940, pp. 90-98, pl. 2, figs. 3, 4;
text-fig. 10; text-fig. 9
Synerocrinus Jaekel; Talanterocrinus Moore and Plummer

A new restricted generic concept of Synerocrinus is
proposed by these authors and 1s essentially an accurate
presentation of characters exhibited by the original defini-
tion of the type species except for the nature of the anal
tube or anal series which is not mentioned, above the anal
X, in the descriptive analysis but is demonstrated as being
composed of a double series of plates in their text figure 9,
purporting to be a diagrammatic representation of the
genus. Consideration of any arrangement other than a
single series of plates which are not suturally united with
the arms is untenable for the genus if it is to be restricted
to original characters of the type species. The small syn-
type of Trautschold clearly shows a wide open space above
anal X as he also noted in the description of the species.
The upper structure of the anal series (tube) is well shown
by the specimen figured by Springer (1920, pl. 42, fig. 6a)

226 PALAEONTOGRAPHICA AMERICANA (VI, 40)

which specimen is a bona fide representative of Synero-
crinus incurvus in the most restricted sense in having iso-
tomous arms and a restricted number of interbrachials.
There is no question as to the proper character of the anal
series in Synerocrinus.

Considering the effects of the above information fur-
ther we find the species Synerocrinus formosus Moore and
Plummer (1940, p. 94-96, pl. 2, figs. 3, 4; text-fig. 10) has
an anal series that develops a double row of plates above
the anal X and abut against and interlock with the adjacent
arm segments. This appears to us to be a fundamental dif-
ference of generic stature which with other factors has
caused us to propose segregation as Aexitrophocrinus, n. g.

Moore and Plummer (1940) discussed thoroughly their
reasons for separating forms with bi-endotomous (hetero-
tomous) arms (formerly referred to Synerocrinus) into a
new genus for which they proposed the name Talantero-
crinus Moore and Plummer. One must assume they were
able to change the generic name while the 1940 paper was at
press but could not insert a footnote as to the synonmy
incurred by their proposal of the name Trautscholdicrinus
Moore and Plummer (in Moore, 1939). In any event the
name of the type species of Talanterocrinus should have
read “Genotype. — Trautscholdicrinus jaekeli Moore and
Plummer, 1939.” As previously noted Trautscholdicrinus
Moore and Plummer is in synonymy with Synerocrinus.
There is another usage of the name Trautscholdicrinus by
Yakovlev and Ivanov (1939) which has been considered
elsewhere.

The new description of Trautscholdicrinus jaekeli er-
roneously listed as “Talanterocrinus jaekeli Moore and
Plummer, n.sp.,”, is short enough to bear repeating here,
quote,

Description. — The description of Synerocrinus incurvus (Traut-
schold), as given by Springer [1920, pl. 42] refers to the species
that is here intended to be designated by a new generic and
specific name, because, as pointed out under the discussion of
Synerocrinus, it appears that the name formerly used cannot
properly be applied. Apparently all the forms illustrated by
Springer (except, of course, his reproductions of the types of
Synerocrinus incurvus, pl. 42, figs. 1, 2a, 2b) may be assigned to
Talanterocrinus jaekeli, n.sp.

Occurrence. — Upper part, Lower Carboniferous, near Moscow,
Russia, and Scotland.

Types. — The specimen in the Springer Collection at the U. S.
National Museum that is illustrated in Springer’s figures 4a and
4b, pl. 42 (1920), is designated as holotype. This is a Russian
specimen from the vicinity of Moscow.

As we have noted elsewhere, the choice of the holotype
was good because it is a specimen in excellent preserva-

tion and could have provided a firm foundation for the
species and the genus in all respects except as concerns the

posterior interradius which is missing in this specimen. But
unfortunately it is conspecific with Synerocrinus incurvus.
The specimen figured by Springer (1920, pl. 42, figs. 6a and
6b) is a typical representative of Synerocrinus incurvus as
represented by the two syntypes and is apparently the only
specimen observed by Springer having a complete anal
series (tube). The illustration he gave (1920, pl. 42, fig. 7)
is a composite of the specimens represented by figures 5, 6,
and 8, and although the gap in the posterior interradius
indicated by the specimens represented by figures 5 and 8 is
like that of Synerocrinus incurvus there were no anal or
tube plates preserved in those specimens. Fortunately
Springer (1920, p. 334) gives a detailed written description
of the arrangement in the fragmentary specimen (his fig.
8) in the posterior interradius which is obviously for the
support of a tube. His (1920) text figure 44, purported to
be a diagrammatic representation of Synerocrinus, is modi-
fied and used as a diagram of Talanterocrinus by Moore
and Plummer (1940, text fig. 9). It is mainly based on
Springer’s plate 42, figures 4a and 4b, which is the only
specimen we have seen where there is a single interbrachial
above the first dichotomy and none above the second. This
must have been the form Jaekel (1918) had in mind when
he specified “1 grosses Interbrachiale” in his familial descrip-
tion.

The specimen figured by Springer (1920, pl. 42, fig. 5a)
has a large interbrachial followed by a small interbrachial
above the first dichotomy and a single interbrachial is
visible in the left half of the right posterior ray above the
second dichotomy. If Springer’s illustration is correct there
is no second isotomous branching in the right half of the
left posterior ray and the appearance of ramules, or arm-
lets, is sporadic.

The large specimens figured by Springer (pl. 42, fig.
3) differ markedly from the other specimens in having
numerous interbrachials at the primibrach level, numerous
interbrachials above the first dichotomy and numerous
interbrachials above the second dichotomy. In the later
area it is sometimes difficult to separate interbrachials
from brachials of the ramules, or armlets. One must con-
clude all the interbrachials were present at a younger stage
or entertain the thought that the animal added inter-
brachials with age which is of course a possibility.

As mentioned elsewhere, one of the two Scottish speci-
mens figured by Springer (1920, pl. 75, fig. 12) has been
referred to Caldenocrinus by Wright (1946), but the other
(fig. 13) was referred by him to Yalanterocrinus jaekeli
(Moore and Plummer).

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 227

XI. Moore and Laudon, 1943, pp. 73-74, text-fig. 10

Synerocrinidae Jaekel, 19, 1918; Synerocrinus, Talanterocrinus,
Ainacrinus

Moore and Laudon (1943) have modified the family
Synerocrinidae to embrace forms having either isotomous,
dichotomous, or heterotomous arms and ranging from Dev-
onian to Permian age. We are only directly interested at
this time in the genera Synerocrinus and Talanterocrinus.
Those authors considered the isotomous character of arm
branching and the presence of several interbrachials be-
tween the rays to be of a primitive nature but many of the
genera in the group exhibit these characters and are of the
same or younger age as forms of the bi-endotomous (hetero-
tomous) group, which are considered to be advanced. The
type species of Synerocrinus and the type species of Talan-
terocrinus are from the same rocks and age and are prob-
ably either conspecific or are of different lineage. We pre-
fer to believe they are conspecific. The concepts of these
genera proposed by Moore and Plummer (1940) and their
diagrams are used which perpetuates the erroneous con-
cept of Talanterocrinus, especially for the plates of the
posterior interradius which was a tube in their type species
but are figured as a multiple series of plates (having com-
mon sutures with the brachial plates) by Moore and Plum-
mer (and Moore and Laudon). This has been corrected by
us elsewhere and Talanterocrinus placed into synonymy.

The paper written in 1943 by Moore and Laudon was
the basis for their section on crinoids in Shimer and Shrock
(1944) and data were repeated.

XII. Moore and Laudon, 1944, pp. 179, 181, pl. 56, figs. 20, 30;
pl. 67, figs. 3, 6
Synerocrinus Jaekel, Talanterocrinus Moore and Plummer

Moore and Laudon, 1944, gave the same diagrammatic
drawings of Synerocrinus and Talanterocrinus as shown in
their 1943 study and, therefore, the drawing of Synerocrinus
is in error for the same reasons as previously set forth.
Synerocrinus formosus Moore and Plummer is illustrated
(pl. 56, fig. 20) but as we now know is not representative.
Talanterocrinus mcguire: Moore is illustrated (pl. 56, fig.
30), but it is no longer a representative of the genus. We
have assigned it to Aexttrophocrinus, n. g.

XIII. James Wright, 1946, pp. 33-38, 2 pls.
Caldenocrinus Wright, 1946; Talanterocrinus Moore and
Plummer, 1940
In 1946, Wright restudied the flexible crinoids in his

collections. He followed Moore and Plummer (1940) in ap-
proving the segregation of forms with bi-endotomous

(heterotomous) arms into the genus Talanterocrinus. He
concluded that another genus existed in the Scottish Car-
boniferous for which he proposed the name Caldenocrinus.
One specimen figured by Springer (1920, pl. 75, fig. 13)
is referred by him to Talanterocrinus jaekeli which is now
considered to be syononymous with Synerocrinus incurvus.

He also proposed Talanterocrinus strimple: Wright for
a form from the Viséan of Scotland based on ten cups,
several of which had two primibrachs attached.

Wright placed considerable importance on the nature
of the base, in Talanterocrinus and Synerocrinus, the basals
being visible well beyond the column; whereas in Caldeno-
crinus the basals are rarely seen outside the columnar socket.
He also noted the anal tube structure in Talanterocrinus
and Synerocrinus; whereas in Caldenocrinus the single series
of anal plates are joined by suture with adjacent arm
plates. We agree that adjacent arm plates are a primary
character and the former (nature of the base) may be of
generic significance, if growth stages are not involved. The
reason for reservation is found later under our discussion
of some growth stages of Aexitrophocrinus formosus where-
in considerable change in the base is found between young
forms and fully mature forms.

XIV. Wright, 1952, pp. 320-322, pl. 13, figs. 1, 2, 3, 7; text-figs. 1, 2
Talanterocrinus redesdalensis Wright, 1952

This species is from the Viséan (D,), Calciferous
Sandstone Series of Scotland. We agree that it has many
characters more or less like those of the Moscovian genus
Talanterocrinus (=Synerocrinus) but we do not believe it
is congeneric. There is a fundamental difference in the pos-
terior interradius where in 7. redesdalensis there is no plate
homologous to anal X, but rather just the anal tube con-
nected directly to the posterior basal. The arms of T.
redesdalensis are different to those of Synerocrinus incurvus
in that the inner arms of each half-ray do not appear to
bifurcate above the second dichotomy (are endotomous),
but the outer arms of each half-ray bifurcate irregularly
and the resultant inner arms are more prominent and are
larger than the arms of S. incurvus. In the “holotype” of
T. jaekeli the arms are bi-endotomous with small ramules,
or armlets, developed at regular intervals on the inner side
of the arms.

We believe 7. redesdalensis is distinctive enough to war-
rant separation as a separate genus for which we propose
the name Enascocrinus, n. gen. For affinties we have con-
sidered Ewonychocrinus Strimple of Desmoinesian and Mis-
sourian age (Pennsylvanian) as a probable derivative.

228 PALAEONTOGRAPHICA AMERICANA (VI, 40)

XV. Wright, 1954, pp. 151-160, pls. XLI, XLII, XLIM, XLIV

Ainacrinus, Talanterocrinus, Caldenocrinus, Synerocrinus

There is no change in the concept of Atmacrinus or in
known material from the original description (Wright,
1939) other than to make comparison with the subsequently
described Caldenocrinus Wright, 1946. The later genus has
an anal series interlocking with the adjacent brachials (as
opposed to an anal tube) which is also thought to be the
case in Atnacrinus, but Calderocrinus has more numerous
interbrachials, the basals are covered by the stem, and the
arms are heterotomous. Ainacrinus has from one to occa-
sionally three interbrachials at the primibrachial level
only and the basals are readily visible beyond the columnar
socket.

Wright (pp. 153-154) referred another crushed crown
to Talanterocrinus jaekeli (now Synerocrinus incurvus) in
addition to the specimen from the Ardross Limestone fig-
ured by Springer (1920, pl. 75, fig. 13) as previously dis-
cussed. It is not figured and from Wright’s discussion does
not provide any additional information,

Enascocrinus redesdalensis (Wright, 1952) is discussed
(as Talanterocrinus) but no new evidence is given. The
species has been referred to Enascocrinus by us.

Talanterocrinus strimplei Wright (1946) is discussed
but only in the nature of a review.

A new species is proposed as Talanterocrinus swaledal-
ensis Wright (1954) from the Namurian, E;, Woodocrinus
Limestone of Yorkshire, England. It is represented by two
poorly preserved specimens and cannot be used with any
degree of assurance. The arms are heterotomous and there
is reported to be a single row of anal plates in the posterior
interradius, quote (Wright, p. 151) “which to all appear-
ance do not connect with the adjacent rays.” This species
is referred by us to Synerocrinus.

Wright discussed Caldenocrinus Wright (1946) briefly
and reviewed previous discussions on a generic and specific
level in the “Remarks” section of the monotypic species,

C. curtus Wright (1946).

XVI. Yakovlev and Ivanov, 1956, pp. 35-38, text-figs. 9-12

Synerocrinus incurvus (Trautschold)

It appears that these authors, after reviewing most
of the studies on this form up to 1940 concluded the hetero-
tomous (bi-endotomous) and the isotomous forms from
the Moscovian rocks were all the same species. These are
forms considered to be S. incurvus (isotomous) and Talan-
terocrinus jaekeli (heterotomous), It is true the somewhat

unique anal tube is present in both forms and the base is
closely comparable, i.e. the basals extend slightly beyond
the columnar socket in figured specimens and the posterior
basal is long, truncate triangular-shaped. There is no ap-
parent consistency to the number of interbrachials even
between specimens of the same species. We agree with
Yakovlev and Ivanov (1956) that Talanterocrinus jaekeli
is conspecific with Synerocrinus incurvus and so consider
Talanterocrinus as a synonym of Synerocrinus.

Yakovlev and Ivanov (1956) considered the specimen
presented by Trautschold in 1879 (pl. 4, fig. 3a, 3b) as
original material, but it was not mentioned or figured in

the original 1867 work.

CONCLUSIONS

The family Synerocrinidae (Jaekel, 1918) is a valid
taxon but needs a complete revision under present usage
which we consider beyond the scope of the present study.

The genus Synerocrinus is firmly established based on
the original two type specimens of the type species Forbesio-
crinus incurvus Trautschold. Synerocrinus has simple iso-
tomous arms branching with the second primibrach, and
again with the third or occasionally the fourth secundibrach
in the syntypes of the type species. There may be branch-
ing in the upper arms which are in the nature of ramules
and which we term bi-endotomous. The infrabasals are en-
tirely covered by the column as also much of the basals;
however, the basals project beyond the columnar socket.
The radials are in contact except at the posterior inter-
radius into which the posterior basal is extended. A rather
large interradial plate rests in a notch at the summit of
the radials and may be followed by two or more inter-
brachials. There are usually one or two interbrachials above
the first bifurcation and there may be interbrachials above
the second bifurcation. In the posterior interradius, there
is a single series of plates which are in the nature of a tube
and do not interlock with adjacent arm plates.

Synerocrinus formosus has a large anal plate followed
by a double series of anal plates all of which interlock with
the adjacent arm segments. It is taken as the type species
of Aexitrophocrinus, n. g.

The arms of Talanterocrinus redesdalensis Wright
branch into two arms of approximately equal size. In this
form there is also a unique modification of the posterior
basal, and it is removed to a new genus Enascocrinus. The
posterior basal in Synerocrinus incurvus is long, truncate
triangular-shaped supporting a plate that fills its upper sur-
face. There is an anal tube which does not interlock with
adjacent arm segments. Talanterocrinus mcguirei has a

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 229

large anal plate followed by a double series of smaller anal
plates, all of which interlock with the adjacent arm seg-
ments, It is, therefore, removed from Talanterocrinus and
placed in the new genus Aexitrophocrinus. Talanterocrinus
strimplei Wright is reported to apparently have an anal
tube rather than plates interlocking with arm segments and
for this reason is placed in Synerocrinus. The arms are not
known above the second primibrach.

Genus SYNEROCRINUS Jaekel, 1898

Type species.— Forbesiocrinus incurvus Trautschold.

Range.— Middle Carboniferous (Moscovian), Scot-
land. Viséan; USSR.

Diagnosis. — Crown elongate, rotund; IBB horizontal,
covered by stem; BB in contact all around, PB truncated
for one anal plate, extend beyond columnar socket; RR in
contact all around except in posterior where PB is extend-
ed; anal tube in single series, is not suturally united with
arms; rays widening from RR, partly separated by sparse
IBrBr, branches isotomously two times, one to three [BrBr
above first dichotomy and one or none above the second
dichotomy.

Remarks. — The background for establishment of this
genus has been given under the familial discussion. Strict
restriction to characters of the type species of the genus
has reduced representation to the genus.

Genus AEXITROPHOCRINUS Strimple and Watkins, new genus

Type species.—Synerocrinus formosus Moore and
Plummer, 1940.

Range. — Morrowan to Virgilian; North America.

In the process of investigating the genus Synerocrinus
it became obvious that forms like Synerocrinus formosus,
which have anal plates interlocking with adjacent segments,
or to phrase it differently, have anal plates suturally united
to brachial plates, are not congeneric with Synerocrinus
which genus has a small anal tube completely separated
from the adjacent brachials. There is a closely related
Viséan genus, Caldenocrinus Wright, which is the probable
ancestor of Aexitrophocrinus, n. g. Caldenocrinus has a long
posterior basal that extends beyond the outer limits of
the radial plates and the anal plates form a single series
which is suturally united to the brachials, In Aexitropho-
crinus the posterior basal is longer than other basals but
does not reach the edge of the radial circlet and the anal
plates are in a double series above the first anal plate (X).
In other respects, Aexitrophocrinus develops more arms and
more interbrachial plates in areas above the second branch-
ing of the arms, than found in Caldenocrinus.

Ainacrinus Wright, which is also of Viséan age, could

well be in this lineage but poor preservation of the type
material has caused us to consider it with reservation. In
Ainacrinus the basals project well beyond the columnar
socket, as do the basals in the young of Aexitrophocrinus
formosus (Moore and Plummer). Basal plates and portions
of the radials are completely covered by the column in a
large specimen of A. formosus. The even division of upper
arms, although at unequal heights, is considered by us to
be more primitive than the endotomous division as small
armlets, or ramules, such as found sporadically in the Des-
moinesian Aexitrophocrinus formosus and prolifically in the
Virgilian A. meguiret. The genus Ainacrinus has a small
number of interbrachials (only one at the primibrach level
in the holotype of the type species).

Generic diagnosis. —IBB are three unequal elements
entirely covered by the column, smallest in right posterior
position; BB five, projecting beyond columnar socket in
young specimens but entirely within the depressed area in
old specimens except for the posterior basal which projects
slightly beyond the column even in large forms but does
not project beyond the perimeter of the radial circlet; RR
five, in old specimens their proximal end is in the columnar
socket but in most specimens they do not reach the de-
pressed area; a large anal plate (X) rests solidly on the
posterior basal and is followed by a double series of sub-
equal plates, all plates being suturally united with adjacent
brachial plates; arms branch isotomously two times, with
PBrsax and with SBr3-4ax, thereafter there may be spor-
adic heterotomous branching in upper arms (e.g. A. for-
mosus) or systematic biendotomous branching (e.g. A. mc-
guiret); interbrachials are four or more in primibrach area,
three or more above the first dichotomy and may be one
or more above the second dichotomy (when preserved);
proximal columnals are thin broad, rapidy tapering basil-
arids.®

Species assigned to Aexitrophocrinus
Assignment

Type species,
A exitrophocrinus

Occurrence
Millsap Lake,

Desmoinesian;

Species name
Synerocrinus formosus
Moore and

Plummer Texas

Talanterocrinus Brownville, A exitrophocrinus
mcguiret Virgilian;
Moore Oklahoma

Aexitrophocrinus Atokan; Aexitrophocrinus
lamberti, n.sp. Texas

The generic name is from the Greek Aexitrophos with
reference to the increased growth of the plates of the
posterior interradius.

’ A term proposed by Strimple (1963, p. 15) for specialized proximal
columnals which are near the base of the cup and are sharply differ-
entiated from columnals lower in the stem.

bo
we
©

Aexitrophocrinus formosus (Moore and Plummer), 1940
Plate 45; figure 3, Plate 47, figures 2, 3
See Bassler and Moodey, 1943, p. 697

This species was established on several specimens from
the Brannon Bridge Limestone (Desmoinesian) near Brock,
Parker County, Texas. We have three topotypes which pro-
vide additional understanding of the species. The speci-
mens consist of one young form, one of moderate size,
and one crown in full maturity. The most readily apparent
difference is in prominence of the basals, or more explicitly
the degree of surface exposed beyond the columnar scar.
Moore and Plummer (1940, p. 94) touched on this condi-
tion in discussion of basals, quote “In one small specimen
these plates are relatively much more prominent than in
the large ones, owing chiefly to the much smaller diameter
of the stem.” Our smallest specimen reflects a condition
comparable to that shown by Moore and Plummer (1940,
text-fig. 10a-b) for supposedly immature and mature speci-
mens, 1.¢. the basals extend beyond the columnar scar
as much and in fact more than in their “immature” speci-
men. In our medium-sized specimen only the distal points
of the basals extend beyond the stem, other than the pos-
terior basal which is typically longer than the others. At
this stage the proximal tips of the radials are becoming in-
volved in the columnar depression. In the fully mature
specimen the majority of the surface of the radials have
been covered by the stem. At this stage there is still a
distal edge of the posterior basal exposed beyond the col-
umn.

We are unable to reconcile with the small quadrangu-
lar first primibrach illustrated by Moore and Plummer
(1940, p. 94 and text-fig. 10a) and discussed as represent-
ing an immature stage. In all of our specimens the first
primibrach is as large as, or almost as large, as the second
primibrach and has two facets on each lateral side for con-
tact with both the first interbrachial and one of the second
interbrachials, The first interbrachial is followed by two
plates and usually by one or two above that. There are
usually three interbrachials, that may be in series, above
the first branching. The second bifurcation takes place with
the SBrgax. Although it is not reported by Moore and
Plummer, our large crown exhibits a series of interbrachials
starting in a notch between the first tertibrachs above the
point of bifurcation. Apparently some branching takes place
above and the segments of the armlets cannot be differ-
entiated from the interbrachials. The same condition exists
in the left half of right posterior arm but interbrachials
do not appear to develop in the anterior arms. Of much
more interest is a condition found in the left arm of the

PALAEONTOGRAPHICA AMERICANA (VI, 40)

right anterior ray where the thirteenth tertibrach is axillary
with the small brachial on the inner side encroaching on the
TBr,,. The following TBr,; is axillary with the small
brachial on the inner side and encroaching on TBrig. It ap-
pears that TBryg is also axillary with small brachial on
inner side. We have been unable to discover comparable
development in other rays except the condition already re-
ported in the posterior. Only one brachial of the ramule is
exposed in each of the upper heterotomous divisions. Our
large topotype has as many as 19 tertibrachs without evi-
dence of termination although it does appear the arms were
approaching their distal ends. The brachials are all wide
low, elements with rounded exteriors and in the upper
arms they seldom show any indication of developing a
sinuous suture that is so typical of most flexibles. There is
a development that might prove unique for these forms
which is the serrated edges of the articular surfaces. Where
slight displacement or differential weathering has taken
place the structure is well shown.

In the posterior interradius, the large anal plate rests
on the truncated distal end of the posterior basal. It has
seven sides and supports two fairly large plates above.
In the large topotype there are two staggered plates in the
next series followed by two in the next series but thereafter
there are three small plates in uniserial arrangement.

The columnar facet of the large topotype is occupied
by a fused mass of compressed, tissue-thin proximal colum-
nals.

Measurements in millimeters—
Small Medium Large

Width of crown (as preserved) — a 47.5
Length of crown (as preserved) _— —_— 69.5
Diameter of columnar scar 5.5 8.0 20.5
Width of dorsal cup 14.8 17.0 25.0
Height of dorsal cup 2.9 2.9 —
Width of basal (beyond columnar scar) 3.8 24 0.0
Width of basal (internal) Bis Bez Y

Length of basal (beyond columnar scar) 2.7 1.2 0.0
Width of basal circlet (minimum) 9.2 LE / 0.0
Width of radial 6.8 8.0 11.2
Length of radial (beyond columnar scar) 4.0 4.3 2to4
Width of PBri 7.3 9.0" 12.1
Length of PBr 3.6" 355" 6.0
Width of PBrax 7.8* 9.4% 11.8
Length of PBroax 3.4* 4.5* 7.0

a—average

*—only one preserved

Remarks. — A partial crown and a cup found in the
Oologah Limestone Formation (Desmoinesian) of north-
eastern Oklahoma, have been figured by Strimple (1962a,
pl. 3, fig. 1; pl. 8, figs. 2, 3) and designated as plesiotypes
of Aexitrophocrinus (Synerocrinus) formosus. The partial
crown is about the same size as the medium topotype fig-
ured herein and various reported measurements agree.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 231

Occurrence. — Holotype, paratypes, and figured topo-
types are from the Brannon Bridge Limestone Member,
Millsap Lake Formation, Desmoinesian; three miles south-
west of Brock, Parker County, Texas: cited (plesiotypes)
hypotypes are from the Oologah Limestone Formation,
Marmaton Group, Desmoinesian,; OU 4569 is from an
abandoned quarry in SE 14, SW 1, sec. 18, T.20 N., R.14
E. and OU 3988 is from Garnett quarry in SW \% sec.
28, T.20 N., R.14 E., Tulsa County, Oklahoma.

Types. — Holotype, No, H-1, paratype, No. H-2, both
in the Harris Collection; collected by Mrs. G. W. Harris.
Paratypes, Nos. M-2, M-5, M-16, M-30, and M-39 in Marrs
Collection; collected by Mrs. W. R. Marrs. Plesiotypes OU
4569 and OU 3988 in University of Oklahoma, Paleonto-
logical Collections; collected by H. L. Strimple. Topotypes
USNM, Nos. $5156, and two unnumbered specimens, col-
lected by Wm. T. Watkins. Plastotopotypes SUI 11920,
11922, 31872, repository University of Iowa.

AEXITROPHOCRINUS LAMBERT]! Strimple and Watkins,
new species
Plate 34, figure 8

The infrabasals are covered by the proximal columnals.
Five basals extend well beyond the columnar attachment
area with the posterior basal truncated for the reception of
a large anal plate and is not appreciably longer than other
basals. Anal X does not extend entirely to the summit of
the first primibrach and is not so long as the first inter-
radial plate of other interrays, Two small plates follow anal
X and a short series of double plates is above. The five
radials are low, wide elements. First primibrachs are wider
and longer than the radials. Second primibrachs are some-
what smaller and are axillary. The elongated interradial
plate is usually followed by a single small plate at a position
between the first and second secundibrachs. The arms usu-
ally branch again with the third or fourth secundibrach.
Isolated interbrachials are found in some areas well above
the first axillary and may form a single series in some
instances.

Measurements of holotype in millimeters as follows:

Width of crown (as preserved) 40.0
Width of dorsal cup 12.9
Diameter of proximal columnal 5.4
Width of anal X 4.0
Length of anal X 3.7
Remarks. — Aexitrophocrinus lamberti has a lesser

number of interradial and interbrachial or interbrachial
plates than found in either A. mcguirei or A. formosus.

The species is named for C. B. Lambert who found and
donated the specimen to the study.

Holotype. — USNM, No. $5111, collected by C. B.
Lambert, reposited in the U. S, National Museum.

Occurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Formation, Atokan, Pennsylvanian; C, B. Lam-
bert Ranch near San Saba, San Saba County, Texas.

AEXITROPHOCRINUS sp.
Plate 54, figure 7

Three large partial crowns have been found in associ-
ation with Aexitrophocrinus lamberti but are not thought
to be conspecific. The arms of Aewitrophocrinus sp, appear
to have divided in proximal regions and to branch several
times in distal regions, with small ramule-like divisions
appearing on the inner side of some arms. One specimen
shows the basals with the posterior basal extending to about
mid-height of the radials and followed by an anal plate that
fills its entire distal surface.

Specimens. — USNM, No. $5189 collected by Wm. T.
Watkins, reposited in the U. S. National Museum.

Occurrence. — Lemons Bluff Limestone Member, Mar-
ble Falls Formation, Atokan, Pennsylvanian; C. B. Lambert
Ranch near San Saba, San Saba County, Texas.

Genus ENASCOCRINUS Strimple and Watkins, new genus

Type species. — Talanterocrinus redesdalensis Wright
1952.

Range. — Calciferous Sandstone Series, Viséan (D,);
Scotland.

Diagnosis. — Crown elongate; base broad and rounded;
IBB entirely covered by column; five BB extend beyond
columnar socket with slender posterior basal extended but
not beyond radial circlet of plates and with upper face
oblique, the anal tube rising from its left side; five RR with
proximal tips reaching columnar scar in some rays; three or
four interbrachials at primibrach level, one or none above
first dichotomy, none above second dichotomy; arms branch
twice isotomously, the first time with PBrsax and usually
with SBrzax, the inner arm does not branch again above the
second dichotomy or do subsequently developed arms
branching off the outer arms only; the inner arms are usu-
ally more or less the same size as the main trunk, at the
point of bifurcation.

Remarks. — The reasons for separating this form out of
Talanterocrinus (now Synerocrinus) has previously been
given, but it is sufficient to note the half-arms produced
at each bifurcation, in the upper arms, are only slightly
smaller than the main trunk which is different to the small
armlets, or ramules, of T'alanterocrinus (now Synerocrinus).

>

to
we
bROo

We do not believe there is any close relationship to Synero-
crinus or Aexitrophocrinus.

Family LECANOCRINIDAE Springer, 1920
Genus CIBOLOCRINUS Weller, 1909
CIBOLOCRINUS PUNCTATUS Moore and Plummer, 1940

This species is included herein because it is from the
Marble Falls Formation. We do not have any specimens
in our collections and are thus unable to contribute more
than the observations made by Moore and Plummer. There
appears to be an intimate relationship between C. punctatus
and C. tumidus from the Brentwood Limestone, but we are
not prepared to make close comparison. The types of C.
punctatus are reported to come from about 10 feet above
the base of the Marble Falls Formation, whereas most of
the crinoid material in our collections comes from higher
in the formation. Plummer (1950, p. 55) listed C. punctatus
as occurring in the Morrowan of Oklahoma and Arkansas
as well as in Texas which strongly indicates his belief that
some specimens ascribed to C. twmidus are conspecific with
C. punctatus.

Types. — Holotype, Plummer Collection, No, P-11183,
paratypes, Plummer Collection Nos, P-11186 and P-11186A,
all at the Bureau of Economic Geology, The University of
Texas, and collected by F. B. Plummer and G. H. Fisher.

Occurrence. — About 10 feet above the base of the Sloan
Limestone Member, Marble Falls Formation, Morrowan,
Pennsylvanian; Texas Loc. 205-T-43, about 200 feet north
of a cattle tank, in the bank of a creek about 0.1 mile west
of the Wallace Creek road 11.5 miles by road southwest
of Hotel San Saba, San Saba County, Texas.

FRAGMENTARY CRINOID REMAINS
Genus PLATYCRINITES Miller, 1821

PLATYCRINITES TENUIS Strimple and Watkins, new species

Plate 54, figures 1, 3, 4, 6

Two specimens of Platycrinites tenuis are known of
which the smaller is designated as the holotype. The
columnals are large, roughly quadrangular-shaped, with
elongate, elliptical facets. The intersection angle between
the two facets is about 77 degrees. Relatively large sur-
faces are exposed beyond the articular facets and a keel-
like ridge is at the middle of the sides. The ridge is
sharp-crested and marked by a continuous series of small
spines. The fulcral ridges are prominent and may be equal

PALAEONTOGRAPHICA AMERICANA (YI, 40)

to or extend above the height of the raised edges of the
facets for about half the distance to the lumen but are
sharply reduced in height and in width for the remaining
distance to the lumen. The lateral extremities of the facets
are raised slightly above the normal, outer, nonarticular
surfaces. The fulcral ridges are marked by irregular crenu-
lations. The area surrounding the small lumen is raised but
may be destroyed in preservation in which case a large
aperture remains, as happened in the large paratype.

The specimens figured by Tien (1926, pl. 3, figs. 5d-f,
6c-d) as Platycrinus sp. are quadrangular-shaped. The first
specimen has much sharper corners than Platycrinites
tenuis. Tien’s specimen was alse figured by Moore (1939,
text-fig, 6, (13) a ?Platycrinus). The form described as
Columnal quadrangulatus Strimple (1962) is closely related
to the present form but is thinner and the keel-like ridge of
the outer surface is not marked by continuous spines.

There is a remarkable similarity between the columnals
figured by Holtedahl (1911, pl. 1, figs. 5, 6) in association
with Platycrinites spitzbergensis (Holtedahl) and Platy-
crinites tenuis. A primary difference is the pronounced
spinelike projections marking the outer surface of P. spitz-
bergensis. A columnal described as Ellipsellipsopa spicata
Moore (1938) from the Coffeyville Shale (Missourian) of
Kansas has a girdle of spines which are more like those of
the Norwegian form than of Platycrinites tenuis. The Kan-
sas specimen is lower, the articular surfaces are not so elon-
gated, and the general contour is not close to either of the
two forms described herein.

Another highly spinose columnal is figured by Weller
(1930) as Platycrinus Specimen 7, from a dark lower Penn-
sylvanian limestone near the center of the north half of
section 35, T. 21 N. R. 9 W., Warren County, Indiana.
The Weller specimen is considerably thicker and the facets
are not so disportionately elongated as found in Platy-
crinites tenuis. Weller’s specimen has a rotation of one
articulating surface with respect to the other of about 73
degrees, which in P. tenuis is slightly greater, being about
77 degrees.

Stuckenberg (1875) figured a segment of stem from
the Permian of Russia as Platycrinus schmidtii that has
some columnals somewhat like those we have called Platy-
crinites tenuis, that is the segments have rotated articular
facets. In Stuckenberg’s specimen there is a series of seven
columnals that reflect little if any rotation, but the
columnals on each end thereof are rotated. The articular
facet of Platycrinites ? schmidtu (Stuckenberg), new com-
bination, is proportionately longer than in P. tenuis, is not
so deeply impressed, the fulcral ridge is narrower, and ex-

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 233

tends almost to the center of the plate where there is a
small lumen. Viewed from above or below the columnal P. ?
schmidti is more elongate-appearing and has a more angular
outline (excluding consideration of the cirral facets) than
found in P. tenuis. The crinoid specimen figured in con-
junction with the column by Stuckenberg has been referred
by Yakovlev (1954) to Dichocrinus schmidtu (Stucken-
berg).

Yakovlev (1954, p. 77, pl. 13, figs. 4 a-b) has figured
a stem segment as Platycrinus sp. It is apparently elliptical-
shaped and the specimen has seven segments preserved with
no evidence of cirral attachment or of rotation. There are
no keel-like ridges girdling the segments. This specimen 1s
more likely the cirri of an ampelcrinid than the column of
a platycrinitid.

Under the name Platycrinus ? tuberculatus Yakovlev
(1954, pl. 12, figs. 9 a-f) several columnals are figured
from C; (Upper Carboniferous) of the SSSR. The specimen
illustrated as figure 9 a-b is three columnals of a form some-
what like Platycrinites tenuis in that the Russian columnals
have a girdle comprised of a series of nodes, but the exten-
sion of the segment does not appear to be so pronounced
as in the American form. Yakovlev’s specimen, figure 9 c,
has a narrow outline, when viewed from the articular sides,
different from any specimen we have noted. There does not
appear to be any arrangement for rotation between the two
articular facets. The two specimens figured by Yakovlev as
9 e and 9 g are almost identical with those illustrated by
Tien (1926, pl. 3, figs. 5 b-c). They are both somewhat
comparable to Columnal ellipticus Strimple (1962, fig. 1-3)
except they have more evenly rounded extremities, The
specimen figured by Yakovlev (1954, pl. 12, fig. 9f) appears
to be an entirely different and distinct form.

The species name tenuis is Latin for thin.

Measurements of holotype in millimeters:

Width of articular facets (long axis) 11.0
Width of articular facets (short axis) 5.6
Maximum width of columnal 13.3
Greatest length of segment 6.0

Types. — Holotype, USNM, No. S5187; paratype,
USNM, No. S5187-a collected by William T. Watkins, re-
posited in the U.S. National Museum.

Occurrence. — Shale below Kickapoo Falls Limestone
Member, Millsap Lake Formation, Strawn Group, Desmoi-
nesian, Pennsylvanian; about 1/4 mile east of Kickapoo
Falls, southwest of Dennis, Hood County, Texas.

PLATYCRINITES TENUIS CONTENTUS Strimple and Watkins
new subspecies
Plate 54, figures 2, 5

Two specimens consisting of an isolated columnal and
a pair of joined columnals are known. The isolated colum-
nal is designated the holotype and the pair the paratype.

The specimens are smaller than the associated Platy-
crimites tenuis and although they have a weak girdle-like
ridge in midsection and sporodic nodes, they do not have
the same appearance found in the larger form. The holo-
type of P. tenuis contentus does not even have a directional
variance between the opposite articular facets but in the
paratype, it is calculated there is an intersection angle be-
tween the two facets of about 53 degrees. The intersection
angle between exposed facets of the two joined columnals
is about 45 degrees. One of the columnals of the paratype
is a nodal. The cirral scar is indistinct but appears to be
fundamentally circular and occupies a strongly projected
area. The fulcral ridge is wide near the outer edge of the
facet and tapers toward the strongly depressed center
where the minute lumen is surrounded by a circular rim.
The perimeter of the facet is crenulated and the fulcral
ridge is unevenly crenulated,

The reason this form is considered as a subspecies of
tenuis is the nature of the fulcral ridge which is more or less
wedge-shaped on each end and is roughly crenulated The
ridge is subhorizontal for about one-fourth the length of the
facet, then curves sharply into the impressed central area
and diminishes in height as well as in width. We realize that
it is even possible for contentus to have been part of the
same columnal as tenuis although they appear to be differ-
ent. Not much is known about variants between different
areas of a crinoid stem other than the fact that differences
do exist in some forms. P. tenuis contentus is different from
P. tenuis in lacking the shelflike projections of the exterior
with their serrated edges which is considered to be typical
of the later. Comparisons with other forms have been given
under discussion of P. tenuis.

The name contentus is Latin for agreeable, having
reference to the compatibility of the subspecies to the
species.

Measurements in millimeters as follows:

Paratype

Holotype (nodal)
Width of articular facet (long axis) 9.2 8.5
Width of articular facet (short axis) 5.6 6.3
Maximum width of columnal 10.0 94
Greatest length of segment 3.4 4.4
Diameter of lumen (long) 0.05 0.2
Diameter of lumen (short) 0.05 0.1

Width of cirral attachment none 1.2

tro
w
rs

Types. — Holotype, USNM, No. $5188, and unnum-
bered paratype collected by William T. Watkins, reposited
U.S. National Museum.

Occurrence. — Shale below Kickapoo Falls Limestone
Member, Millsap Lake Formation, Strawn Group, Des-
moinesian, Pennsylvanian; about 1/4 mile east of Kickapoo
Falls, southwest of Dennis, Hood County, Texas.

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238 PALAEONTOGRAPHICA AMERICANA (VI, 40)
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1939. Echinodermata in Gorsky, Ll, The Atlas of the leading Geologisheskogo Instituta (VSEGEI), Trudy, n. s., vol. 11,
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Upper Carboniferous. Central Geol. Prospecting Inst., Yandell, L. P.
Leningrad, pp. 64-70, illus. 1855. Description of a new species of Crinoidea. Amer. Jour. Sci.,
1956. Morskie Lilii i blastoidei kamennougol nykh i permiskikh ser. 2, vol. 20, pp. 135-137, illus.

240

it,

Og ot Came

Figure

1-4"

PaLaronTocraPHica Americana (VI, 40)

EXPLANATION OF PLATE 30

Magnification is approximate

Birt chimes <QrOSStS, Wis SPs cess cect soeas ete sewn tas seer acssnpepeoeansseasereeaneses eee od Beas eechcicceoes neha te 204
Holotype. 1. Terminating platform of the anal tube; X 1 1/8. 2. Upper articulating
facet of the left anterior radial plate; X 1 1/2. 3. Lower articulating facet of the
left anterior promibrach and the inside of lower arms showing projections which
probably assisted in suporting the large proximal pinulars; x 1 1/2. 4. Right
anterior view of crown; X 1 1/8. USNM $5097.

Plate 30

PALAEONTOGRAPHICA AMERICANA, VOL. VI

ta Pe T,

a

CRRA REE

‘|

Plate 31

PALAEONTOGRAPHICA AMERICANA, VOL. VI

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 31

Magnification is approximate

Figure Page
Te Sif COU ERI KETTLES ALLTEL, SURO) ce aE cag oR Ue oa 174
Paratype. 1. Posterior view of crown. 2. Anterior view of crown; xX 1 1/8.

USNM S5098.
DS teE A ESTOCRINIESEAT rt) ALIUSmm Tots pees 8 ae ee oe ne en eh te ee 193
Holotype. Two side views of slightly distorted crown; < 1 1/8. USNM S5099.
AO RAN UTLLCN III ES PICULILIS Dee SDs, 2 scesc. coos sonst a So ehtts es hae ap sates ce obec des ecsttesse ec enstes 194
Paratype. Side view of imperfect crown; < 1 1/2. USNM $5101.
Si OmI AGODA SICEIILILSMRIGRIEPOOETESES:, STIG. OSDS\,2scasc21- 220 1-ecoceta oe ee ceca auess a peas bastve eaten 191

Holotype. Anterior and posterior views of crown respectively; x 1 1/4. USNM
$5100.

241

242 PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 32
Magnification is approximate
Figure Page

1.2.) Stellarocrinus, texange((Strigiple) aes ee ee ee 175
Arms viewed from interior and exterior; « 3. USNM $5102.

Sn5; Aglaocrinus magnus \(StrAMple) merece cesses cece heer 172
Hypotype. Dorsal cup viewed from the anterior, base, and posterior respectively;
x 3. USNM S5103.

6. ‘Genus and) speciesyindetenm inate cs. .scccee-ccss nce oestcrseeeacte tee cee ee eee ee 169
Crown with indistinguishable cup plates viewed from the side. USNM $5104.

Plate 32

PALAEONTOGRAPHICA AMERICANA, VOL. VI

“ty

ye iad
i

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|

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a4

Plate 33

PALAEONTOGRAPHICA AMERICANA, VOL. VI

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 243

EXPLANATION OF PLATE 33

Magnification is approximate

Figure

Ue QOL TOROCLITUS LUT AES, elntS poe mee ee ee te 214

Holotype. Crown viewed obliquely from right side and from above respectively ;
xX 1 1/2. USNM S5105.

3,4. Pirasocrinus scotti Moore and Plummer

Hypotype. Crown viewed from left side and from the right anterior respectively ;
xX 1 1/2. USNM S5106.

244

Figure
LZ:

3, 4.

6, 7.

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 34

Magnification is approximate

Oklahomacrinus frostae, n. sp.
Holotype. Crown viewed from posterior and anterior respectively; xX 1

USNM S5107.

Fly drigerinus lorraine, cine s py. coccessaesancn scissoeieeveacsessseacnccee eee ere a 198
Holotype. Crown viewed from left posterior and anterior respectively; X 2 1/4.
USNM S5108.

AUP OG APIAOGTINUS SP y. | osc nceseovicesheeaces ees oseeess see pesnese sae teaoa che dota 183
Side view of juvenile crown; X 7 1/2. USNM $5109.
Higdrrocrinius lorrainae Mo oSps -2-.csaccsossacc-fsntanences senor seeees eee 198

Paratype. Anterior and posterior views respectively of partial crown. USNM
$5110.

AextiroPhocrinus) Lamberti, Th. eSPs (cczsccaccessectencansetetensscestwntsaeee once cone eee tine Re eee 231
Holotype. Basal view of crown; X 1 1/8. USNM S5111.

Plate 34

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Plate 35

PALAEONTOGRAPHICA AMERICANA, VOL. VI

te

bd eee PEPE ry

ere —

ars,

vote Me

Figure
1-3.

7, 8.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 35

Magnification is approximate

ar asoGLUnisestortimNioore and) Pl wmImner ese eee eee eee 206
Hypotype. Crown viewed from left anterior, base and right posterior respectively;
X 1 1/2. USNM $5112.

SAL MCaIeL SUCHE TETA San LOT EU TEALIES © WDLo, SD svy axe eee were ee ee a en 170
Paratype. Posterior view of partial crown; X 2.

Sellardsicrinus fortunatus, n. sp.
Paratype. Side view and opposite view

ALLAMASICrIN UN OTT Alls, Non Spee tt eee ee ee ee 170
Holotype. Posterior and anterior views respectively of partial crown; Xx 1 1/8.

USNM $5114.

245

Figure
1-3.

ae

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 36

Magnification is approximate

Phanocrinus: trullews y. (1vc (Syys) soissccco cect 2n eos ec aso aa ee Sean ee 216
Holotype. Right posterior, basal and posterior (upper part deleted) views of
holotype respectively; X 1 1/4. USNM S5115.

Oxyracrinis. spicata, 10. Spis, cccsee ccc sntss cease acs ence soe ete ea cc can nase av ee 184
Holotype. Posterior view of crown; X 2 5/8. USNM $5117.

Diachacrttes:: dtl ates: Mi. Spi sec tage es soe ss wee cee oes reese ee a ce 222
Paratype. Side view of dorsal cup with appreciable portion of stem attached;
x 8 1/4. USNM S5116.

Plate 36

PALAEONTOGRAPHICA AMERICANA, VOL. VI

; 5
‘
&

5 Syd

& & Spe

¥
0s
ies
Win.
“ii /

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 37

Figure
1-3,

4-7.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 37

Magnification is approximate

Page
Synarmocrinus Mdormarils, MoS. cece ee SPREE nas ceria pee eae ee ees Seog tak cat devant eas aiecus teen ceuseved 164

Holotype. Dorsal cup viewed from anterior, posterior and base respectively;
xX 2 1/4. USNM $5118.

LUG ER UTE TU SIS SOULE MAINS DD aero ane Se cee ee ee Ae ence ie 167

Holotype. Dorsal cup from anterior, summit, base and posterior respectively;
x 2 1/4. USNM S5119.

247

248

Figure
1-4.

is Wloerinas: seschtj ims (Spe ies. .cosnseovecegecsbasaser aces ceosdepeoy po oud shitaat Meese ee eeee eee ee 167

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 38

Magnification is approximate _ ; yas |

Goléocrinus” composstus, “Die. Spo) ctessssteessty<c-cescsctvenereeesss eto
Holotype. Dorsal cup viewed from anterior, Poster
spectively; X 2 1/4. USNM S5120.

Paratype. Dorsal cup viewed from summit, base and_ posterior respectively;
x 2 1/2. USNM S$5121.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 38

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 39

A ie,

14,15:

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 39
Magnification is approximate
Page

GOL EOC UNL MITSOTLETI SES pa Tico :S 1) eres sce ee ee ee rs ee Beate Pee een 168
Holotype. Dosal cup viewed from anterior {with base up), posterior base and sum-

mit respectively; x 2. USNM S5122.

Erisocrinus georgede, 0. Spy -..----------
Holotype. Dorsal cup viewed from summit, anterior (with base up) and base

respectively; X 2. USNM S5123.

S277 CIUTELES IRS EA NLEU T MOO coat Pate a Sde ns ce ere eee ace eee ee ee ee oe 182

Holotype. Dorsal cup viewed from summit, anterior (with base up), posterior
and base respectively; 2. USNM $5124.

VETO OT OCH ATLIES COLL TUT TLE TN SkS Die oan cnc acoesosesae oo foreseteceslse-asetacchead cobsencuescansesteusa.ateseenees ee ae 161
Holotype. Dorsal cup viewed from posterior and base respectively; x 2. USNM
$5125.

Graffhamicrinus antiquus, 1. SP. «.....2--2-.--2--------eneeeeeneeeeene Rete ein feet eetre eee
Holotype. Dorsal cup viewed from anterior (with base up) and base respectively;
x 2. USNM S5126.

182

249

250

Figure
i Ie

aya.

5-8.

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 40

Magnification is approximate

Page
SLOMICEFINUS: COMME, MBP. <czosescscaedaceacessascccaneas setae ee vs 222
Holotype. Dorsal cup viewed from base and posterior; & 1 3/4. USNM $5127.
SHLOMO CTINUS CONLIML, Mis, (SP). -scssssseos.a--sjecckateccectecacecseenr sees cee toags hae Sete 222
Paratype. Dorsal cup viewed from base and posterior; & 1 3/4. USNM $5128.
Dicromyocrinus optimus. 'Strimple eececs= eee ee ee 162

Hypotype. Dorsal cup viewed from the posterior, anterior (with base up), summit
and base respectively; xX 2. USNM $5129.

Grafflamicrimus: artig tts, tim S pieces tee eae ator wanes ese erecaeneeeeese eee 182
Paratype. Dorsal cup viewed from the base and posterior; x 2. USNM $5130.

Gymibtocninus: Contents, ti. \SPih oesys ces cecstisec ence ety ornate een n 6 ak oeana chains meee eee 190
Holotype. Crown viewed from right posterior (anal plate to left) and left pos-
tero-anterior; X 2. USNM $5131.

Plate 40

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Plate 41

PALAEONTOGRAPHICA AMERICANA, VOL. VI

: See i

a

Figure
a

2.

a4

SS

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 41

Magnification is approximate

Page
Stellarocrinus texanus (Moore ands Plummen)) «2c. -<.<cqscsscee cress be sence oot cne ee reeen dance ceencee 177
Hypotype. Posterior view of young individual; x 4. USNM $5150.
LIS OAEHAB TTD. CRHS TITS TONG YSN, cece ee ea eR Se Pee a 217
Paratype. Side view of crown; X 7.5. USNM S5133.
DD ECRDCEIILIESERTLE) ATAU HeRY Mm A [0c, ene ee cnttee ne cece ee 8D 222
Holotype. Posterior view (without upper arms) and anterior view of crown;
X 7.5. USNM $5134.
Scytalocinus sansabensis Moore and Plummer ..........-....2-2-2----+-c--ssecececeocceceeeceeeceeesesceceeseueaces 197

Hypotype. Crown viewed from left posterior; x 3 3/4. USNM S5135.

251

bo
al
bho

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 42
Magnification is approximate

Figure

1. Chlidonocrinus echinatus, n. sp. ....
Holotype. Crown viewed from anterior;

2. Polusocrinus calpculosdes) Vane sc. ceccxccecesee ecnncmn ene caccnne cae pdscsnsannecoyor dp cee oe
Hypotype. Crown viewed from anterior. 1 1/2. USNM $5137.

3,, Araeocrinus BASsSUs, Mi iS, cac.:ccqs-cenceconasecsensossscstecssseccsee eines oceateasupescsusenbuneecsasiosuesep=pqnenesoearenneeem 196
Paratype. Posterior view of crown, large anal plate to the right; % 3. USNM $5138.

4. Chlidonocrinus echinatus, n. sp. ....
Paratype. Side view of crown wit
upper left; x 1 1/8. USNM $5136.

small crown of Scytalocrinus sansabensis to

PALAEONTOGRAPHICA AMERICANA, VOL. VI

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 43

Figure
1,2.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 43

Magnification is approximate

PATECHE GRIT IUSETO ACLU ES D Sytner ee FI E felhs Ee e8 207
Paratype. Young specimen viewed from posterior and from base; < 2 1/2. USNM
$5140.

SAG HICOROCTANU SEI TIT Ody Sp 2 er ee a ede VR ee 190
3. Holotype SUI 32260 from right posterior. 4. Paratype USNM $5142, viewed
from base; X 2 1/2.

Owe Roulusocrinioucalyculoides. Lk anewmnre eee lee ON ee ee ee 186
Hypotype. Young specimen viewed from the posterior and base; & 1 1/2. USNM
$5143.

Sel neeirr CHEGKUR GME OA AMEN SD f-.-s sn ee eee ee oe. EE 207
Paratype. Dorsal cup viewed from posterior, base, summit and anterior (base is
up); X1 1/2. USNM S5144.

Lames ciadivenimisscompertus (Moore) and Plurmime4n)) sect ce ee eee 212
Hypotype. View of specimen preserving uppermost arms to show mode of branch-
ing and prolific number developed; x 2.5. USNM S5145.

13. Anobasicrinus praecursor (Moore and Plummer) ........-.....---2----:s:--:c2-+-2see-0e0eeeeeseeeeeoee-- 174

14, 15.

Hypotype. Imperfect specimen showing balloon-like nature of anal sac. USNM
$5146.

FAT EHICHUNIDSMEOM ATs E I SPs. cite eee ot eee he ee en RE Oh A pene 207
Paratype. Crown viewed from base and oblique posterior position; « 2. USNM
$5147.

253

wn

Figure
1-3.

4, 5.

9-11.

12.

13%

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 44

Magnification is approximate

Tholiacrinus rectus (Moore and Plummer)
Hypotype. Crown viewed from anterior, base and right posterior;
$5148.

Parethelocrinus. watkinst (Strimple)) 2:2: -ccicscsosqcesckcccesecesese:-teenoteeoeaets ceneeeoewenarre ete eeeenre 171
Hypotype. Young specimen with first primibrachs preserved viewed from base,
and right posterior; X 2. USNM S5193.

Shellarocrinus Datl bars, omy Spi cx ccesetscrtecre tes enee tence oxcr oreo st ans eee esha sen cere peerage hanes ee 178

Paratype. Basal view of very young individual (anal plate down); 4. USNM
$5181.

Thaltacrinus rectus (Moore andiP lures)! ceccresse sere ccccs noc enen senor ses she cceasteeeereeneee eee 179
Hypotype. Crown viewed from left anterior and from right posterior; % 2. USNM
$5202.

Seellarocrimus. Dil bis, ‘mie Sys pevcccssesseee see see eco one eee etree 178
Holotype. Crown viewed from anterior (base is up), posterior and base; X 2.
USNM S5153.

Microcaricrinus delicatus, n. sp.
Holotype. Crown viewed from left postero-anterior ;

Ramulocrinus Conse ctattts, 1. SPs, <xn:a.c-eecc-nstesececcasssssetnspcsosasoneotentcs teaeenestan seem ee teeonee ners 200
Holotype. Crown viewed from posterior; <X 2 1/8. USNM $5152.

wt

Plate 4

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Plate 45

PALAEONTOGRAPHICA AMERICANA, VOL. VI

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS 255

EXPLANATION OF PLATE 45
Magnification is approximate
Figure

ee mebirasocrimibses carts Moore and) Plummer o2.=. ses eee eee aoe eee 206
Hypotype. 1. Summit view showing termination platform of the anal tube. 2. Side
view of crown with arm of Stellarocrinus bulbus attached; « 1. USNM $5154.

3. Aexitrophocrinus formosus (Moore and Plummer) 2.0...0..2.2.222..2.22---2c00+esecee0eeeeeeeeeeeeee----- 230
Hypotype. Crown viewed from anterior; < 1 1/2. USNM S5156.

4, Sellardsicrinus marrsae Moore and Plummer ...........2..:-.-:0c:--:se:ce-+-sceceseececesesaesesesneoeeceseo-oe 170
Hypotype. Left anterior view of crown; X 1 1/8. USNM S5157.

: : a
2 ty
*
256 PALAEONTOGRAPHICA AMERICANA (VI, 40)
EXPLANATION OF PLATE 46
Magnification is approximate
Figure Page
1s Microcartcrinuss delicains, 0. Spin. ;...cse ee 201
Paratype. View of young crown from left side; x 3 3/4. USNM SS5158.
2. Stellarocrinus texanus (Moore and Plummer) ) <c2..:-.cccccc:ccssscccccseccsveesosesecssesssesseesesssceesses-s 177
Hypotype. Basal view of young crown; X 3 3/4. USNM $5132.
3: Macrocartcrinus, delicatus, UW. SP. 2s nei .sho eee ee 201
Paratype. Posterior view of crown; X 3. USNM S5159. -
A Microcanicrinus deltcatus, Ds-\Sps ce ceccec- eco vsns e 201

Paratype. Crown viewed from base with arms preserved in a horizontal plane:

X 3 3/4. USNM SS155.

46

Plate

PALAEONTOGRAPHICA AMERICANA, VOL. VI

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 47

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 47

Magnification is approximate

Figure Page
1,4. Stellarocrinus texanus (Moore and Plummer) 20....cc::ccsssssssssssscssessecessssssssssseeeeeeeeeeeeeccccen., 177
Hypotype. Crown viewed from the posterior and anterior; < 1 4/5. USNM S5160.

2,3. Aexitrophocrinus formosus (Moore and Plummer) 22... ccccccceseeeeceeeesseee co. 230
Hypotype. Specimen viewed from base and summit; < 1 3/4.

Shy RSAC HAN AG ATID hs: Ch ceca | 178
Paratype. Crown viewed from base; x 1 3/4. USNM S5161.

MEMES ELLIO CTTELUSEU IE) DIESMET GES D Sipps 178
Paratype. Radial plate with first arm segments in place viewed from the exterior
and interior; & 3 3/4. USNM S5162.

SR LLaiotinite Seren Otis, alls (spi cee et A 219

Holotype. Specimen viewed from base and summit; x 1 1/3. USNM S§ 5163.

257

258 PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 48

Magnification is approximate

Figure Page
1 Di Amechicranis OAs, 0. Rpe | sccnssaepsntecsa ce ees ee aoa ance pcan eo eae 207
Holotype. Crown viewed from side and from posterior; X 1 1/2. USNM S5164.
3.4. Parethelocrinus svuatkimst. \(Strimple))” 2-.2..oscccccscessecececcescancosetecatsannseentevacnavcorcntseansonenmenenands 171
Hypotype. Crown viewed from right anterior and left anterior; 1 1/2. USNM
$5149.
BG. StEHOPECTINUS LONGUS, (IN. CRP ey sect ere cases te caec exer ac canon ve aceltorentn note faa ruet eee tee ene eoee a eet enero 209

Holotype. Crown viewed from the posterior and anterior; X 1 1/2. USNM S5166.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 48

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 49

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 49

Magnification is approximate

Figure
1-3. Parethelocrinus plattsburgensis (Strimple) ...-.0cc0c-cc-cccccccccccecesseccoseeesecessesees
Hypotype. Dorsal cup from base, summit and posterior side. X 1 1
$5167.
TG emryLO OZ OCIAIUN RDI DUGTD MES. @ So. Kes beet te et we Se es ee 161
Paratype. Dorsal cup from base, posterior side and summit; x 2 1/4. USNM S$5168.
7,8. Stellarocrinus sp. cf. S. angulatus (Miller and Gurley) 200.000 ccccccccsccecsseececseeeesseeceees 176
Section of arms removed from crown to show exterior view (7) and interior view
(8); XK 2 1/4. USNM S5169.
OMOnmS cladiocriniusallamoensts Nespas eee ee ee ee ee 213

Holotype. Dorsal cup viewed from posterior and base; 2 5/8. USNM $5170.

259

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 50

Magnification is approximate

Figure Page

1, 6. Stellarocrinus sp. cf. S. angulatus (Miller and Gurley) ...2.........-:ccc:cssccceceseceneeeseseseeeeeees 176
Crown viewed from above and oblique view from below; X 1 1/2. USNM $5169.

O-4! ISchistocrimus GGelass; TS perce eee ee eee 215

Holotype. Dorsal cup viewed from posterior, base and summit; xX 3. USNM $5171.

5. Part of slab with large specimen to left Scytalocrinus sp. _.........c:c0c0--c0000000-> 198, 202, 216
and Culmicrinus barnettensis? partially exposed to its right side, a crown of Phano-
crinus trulleum (paratype) on right edge of slab and an unidentified crinoid at the
lower right. USNM $5172.

Plate 50

PALAEONTOGRAPHICA AMERICANA, VOL. VI

wee
. or SF

go
oe

re tee 2

oP
an

‘
4s ake

Plate 51

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Figure
ile

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 51
Magnification is approximate
Page

Okiahoniariiseprcaeil iss Ney SPs eee ae te ee asc 193, 197
Paratype. Side view of crown with portion of stem attached and two specimens of
Scytalocrinus sansabensis; ~ 1. USNM SS5173.

SAREE RAIDS: (YG TGs, C115) SEND ARPES ee eno Ne gt Se BE Pe e196
Paratype. Anal sac with imperfect specimen of Dichocrinus dilatus to lower left
of sac; X 1 2/5. USNM S5175.

Araeocrinus bassus, n. sp. .....
Paratype. Anterior view of
attached; x 4. USNM S5176.

(CUATTENOAOCHATATIG. (CH IATY ARTIS) V5) eee eee cees eee oases ee recone ee ee 189, 194, 197
Paratype. Side view of crown with considerable section of large pentagonal-shaped
stem attached. The left arm of Oklahomacrinus spicatulus from figure one above

is shown in upper right as well as one of the specimens of Scytalocrinus sansa-
bensis; * 1. USNM $5174.

considerable section of the stem

orsal cup wit

261

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 52

Magnification is approximate

Ulocrinus conwexus (Strimple)! ceisescccccssc-ccccecesecssn ce cnneperenetenopetn-enensnsat aciensrnaceeeomerateenesseprmnsye=—= 166
Hypotype. Infrabasal circlet viewed from exterior and from interior; X 1 1/2.
USNM $5177.

KS Polygonocrinus? Ree earn ee eae ee eee ert rere Fee reesbers odaesicpe ete 214
Young specimen with spinose axillaries and tendency toward interlocking brachials.

426, (Globacrocrinus: naultiplrcaatiss We Bye scccsecrece exec cerns! otnane tree eee tnnteat maee erence eae seep Poteee nen 221
Holotype. Dorsal cup viewed from the posterior, anterior and base; X 10. USNM
$5179.

FUN. Usoallagecrinus, Creckws, Wis, SPs scectecec cee cenecnwcracenenstotacnevereseeerctetosteeece nea ese aoe eee 217
Holotype. Dorsal cup with primibrachs and section of stem preserved viewed from
right antero-anterior interray, base, left anterior, posterior and anterior; X 10.
USNM S5178.

12. (Gulsaicrinus Barnett emsts, Ws SPs. s2c.ccassecrc-cscencasonsseceocsenensespeeceuneseearenantmemnnesoaicunansnaxpeatesenexet chars 202
Holotype. Crown viewed from posterior; 2. USNM $5180.

13. Stellarocrinus texanus (Moore and Plummer) ........-.:...:.--::-:-:<:---:-sse-sseeesnenerenssceenensneneneesnense 212
Hyptoype. Young individual viewed from below; X 2.

14. Sciadiocrinus confertus (Moore and Plummer) .........--.-.-.---------ce+ccscseecccsssnscnenenensensorneseeneare 212

Hypotype. Young individual viewed from the right anterior; x 2. USNM S5182.

Plate 52

PALAEONTOGRAPHICA AMERICANA, VOL. VI

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 53

Figure
2:

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 53

Magnification is approximate

PAD CEM GLADE ASSIESAENN SSD ois ass ac hs. 5 RT ORT eet A 196
Holotype. Crown viewed from left posterior and posterior (anal plate to left) ;
X 2.5. USNM S5183.

Beata ES SIEOCHITIS CITES USP ae ots ei an a ee eee ey aD ee 187
Paratype. View of right side (anal plate to left) ; ; X 2. USNM S5184.

4. Lecobasicrinus kickapooensis, PASS) pene tas chee Piet Pesaro ntoan Means cee UE ay eee Ie 191
Paratype. Anterior view of crown with section of stem attached; x 1 4/5. USNM
$5185.

5. Oklahomacrinus spicatulus, SS oe ere es ee rc Se ee 194

Holotype. Side view of crown showing anal tube, upper articulating facets of
radials and lower facets of first primibrachs; x 1 1/2. USNM S5186.

263

264

Figure

BAG:

Zsa

8, 9.

PALAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 54

Magnification is approximate

USNM S5187.

Platycrinites. £énuts,. We TSpis) jecssestecstcncensecssc spect one poeta nace eee cae an ince 232
Individual columnal viewed from articulating surface and from side; & 3 3/4.
USNM S5187-a.

Platycrinites tenwis:scomtentussom. S\DSpiay sxccecssecccceec ear earns nes eae nes eee erae 233
A pair of columnals viewed from articulating surface and from side; X 3 3/4.
USNM S5188.

Aexitrophocrinus sp.
Posterior view of lower portion of one crown and the upper portion of a
which may be the same species; X 1 1/5. USNM S5189.

Dicromyocrinus sp. cf. D. texasensis (Moore and Plummer) ......--..:-.-.:.:.-:ssess--seeeeeeeon 163
A crown without upper arms illustrated from the posterior and complete crown
from the anterior; X 2 5/8. USNM $5190.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 54

Plate 55

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Figure
1,2.

5, 6.

TEXAN CARBONIFEROUS CRINOIDS: STRIMPLE AND WATKINS

EXPLANATION OF PLATE 55

Magnification is approximate

Wilganuiiemanrwaniitme (StnITple)\<.2- ato oe ad oe Re ee 166

Hypotype. Partial crown from right side (anal plates to the left) and the op-
posite side; X 1 1/8. USNM S5191.

AOG RG DUT OGL ITEMS I Se eet Ee eet ates Bee ae EY RE See aa SES 183
Juvenile crown viewed from the side; X 7 1/2. USNM $5192.

EBL LELD GHITLSE TOLER ITS Tal (OPTATEND |G) yereretecen rece on en ee eee Oe, Se eee 171
Hypotype. Young individual viewed from left posterior; & 1 1/2. USNM S5165.

CGN ANROTITAUS MN CT OWIUN ANI IES Pipetite tote Ae Seen ong SO ae 174
Holotype. Crown viewed from the posterior and anterior; &* 1 1/2. USNM $5194.

LO a GTS RECO TERA CIE Sem ((SSTISETIND LC) ie seoeecee cece cee ae oe na nn eae ee, a oo 166
Hypotype. Crown viewed from posterior side; * 1. USNM $5195.

265

266

Figure
te

Byes

6, 8.

PaLAEONTOGRAPHICA AMERICANA (VI, 40)

EXPLANATION OF PLATE 56

Magnification is approximate

Page
Oklahomeacrimus’ spictlatirsn wi. cS ps) ccs cre cee sae eee 194
Paratype. Partial crown in oblique side view; X 2. USNM $5196.
IPiradsocrinus. Scotti. Wioore Vand) SEL uarmime ness ce arco eoeeese reser 2 eee ae eee 206

Hypotype. Summit view to show terminating platform of the anal tube; X 1.
USNM S5197.

Lecobasicrinus subidus, 1. SP. .xcsconcc-cece--cecosstecrncesee=

Holotype. Small specimen viewed from the posterior and anterior;
$5198.

RR aragassieo crits: | alpUis§ st: (Spee Sescesscsecseceencesee ts soe ate eae encase ae 187

Holotype. Crown viewed from side; & 1. USNM $5199.

ULacr triats Cacti OP SiUS:y: (DMs SSD sowed cmcac Se eae esse eee eat oes a ae aaa heen ee 165
Holotype. Dorsal cup viewed from right side (radianal to left) and oblique view
of left side (radianal to right); * 1. USNM $5200.

Polygomocrimus (Iibraviis, (We (Spe eetcmsen ancestral arn eee 214
Paratype. Specimen viewed from below; x 1. USNM $5201.

Plate 56

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Note: Light face figures refer to the page numbers.

to the plate numbers.

A
Aatocrinus 203
abruptus,

Oklahomacrinus 193, 194
acanthophorus,

Hydreionocrinus 210

Sciadiocrinus Telos Lt:

212, 213

Zeacrinus 210
acclivis,

Schistocrinus 50 153, 215,

216, 260,
Acrocrinus .. 145, 220,
221

acuminata,

Fusulinella 154, 155
adaense, Gastrioceras .. 149
adornatus,

Synarmocrinus 37 153, 163,

164, 247
aequabilis,

Paradelocrinus .......... 181, 182
MESIOCHINIG! © foe scassaseaess 150
Aesiocrinus ............ 150, 159,

185, 188, 189, 191, 193, 194, 195
Aexitrophocrinus 142, 225,

226, 227, 228, 229, 230, 231, 232
Aexitrophocrinus sp. 54 231, 264,

Agassizocrinus .... .... 145, 184,
187, 192
ASIAOCKINUS: | s..:)c0c..--.- LOo, LIZ
AINA CTINUS © svesvspncscerceks 225, 227,
228, 229
Aktyubinskoi
District, Russia ........ 158
(ATCIMOGEINUS) sy.c. cestscss 150, 173
alexanderi,
Phanocrinus 216
Allagecrinus 160, 217,
218
Allegheny Series 158
Allocatillocrinus 218
Allosocrinus 188
altus, Paragassizo-
CEINUS es oe 250, DO LoS Late
188, 263, 266
alvestonensis,
Acrocrinus 221
americanus,
Allagecrinus 218
Parulocrinus 163
Phialocrinus ....... 163
amotapense, Pseudo-
paralogoceras ; 155
Ampelocrinus ... 184, 185,
193, 196
Amphicrinus 160
Amphicrinus sp. 160
Amphissites
rothi Zone 155
amphora, Acrocrinus 221
Anartiocrinus ....... . 144, 192
Anchicrinus eso Ae L5G)
203, 206, 207, 208
angulatus,
Glaphurites |.............. iG!
Stellarocrinus .......... 176, 177

INDEX

angulatus, Stellaro-
crinus sp. cf. 49,50 145, 160,
175, 176, 177, 241, 259, 260

anguloumbilicatus,

Glaphrites 160
annulatus,

Aesiocrinus 31 159, 193,

194, 241
Anobasicrinus 159, 174
Anthracoceras 158
antiqua,

Pseudostaffella 147, 156,

157
antiquus, Graff-

hamicrinus ....39, 40 146, 153,

182, 249, 250
Apodaca Formation 154
apodacaensis,

Profusulinella 154
Apographiocrinus 160, 183
Apographiocrinus

Spare 34, 55 183, 244,

265
Apollocrinus 159, 175
Araeocrinus .. 153, 195,
196, 197

Arbuckle

Mountains, Okla. 151
Archimedes 141, 142,

146, 203
arcuatus, Apographio-

crinus sp. cf. . : 160
ardmirensis,

Delocrinus 182
Ardmore Basin,

Okla. = 148, 151,

153, 157, 158, 224
Ardmore, Okla. .... 151, 172
ardmorensis,

Goleocrinus 167, 168,

169
Andross Limestone 228
argentinei,

Exaetocrinus 181
argentinei,

Stuartwellercrinus 181
arkansasensis,

Pronorites ................. 149, 153
arkansiensis,

Stenopronorites 149, 151,

152
Arnold Limestone

Member ............... 2 Ad a2

Arrey Formation .......... 153, 154,
157
Arrow Canyon,

IN@ Va dare shee tent. A 5B

Artic 155, 157,

158
Articulata 144
assymetricus,

Paragassizocrinus 188
asteriaeformis,

Forbesiocrinus .......... 224
Asturias, Spain .......... . 156
Ataxiacrinus: 2 162
Athlocrinus .. 160, 203,

206, 207
Athlocrinus, sp. ............ 160

267

Bold face figures refer

Atoka County,

Okla. 147, 149,
157
Atoka Formation 141, 147,

148, 149, 150, 151, 152, 153, 154,
155, 156, text-fig. 1,157, 158

“Atoka Group” 149
Atoka, Okla. . 147, 149
atoka,

Paradelocrinus 182

Paragassizocrinus 151, 188
Atokan Stage or

Series 141, 142,

143, 145, 146, 147, 148, 149, 150,
151, 152, 153, 154, 155, 156, 157,
158, 159, 160, 161, 163, 164, 165,
166, 167, 168, 169, 173, 180, 181,
182, 183, 185, 186, 188, 189, 190,
191, 193, 195, 197, 198, 208, 213,
215, 216, 218, 220, 221, 222, 224,

229, 231

atokensis, Staffella 149

Aubrey Group 211

Aulocrinus 199, 201
austinii,

Allagecrinus 217, 218

Austrias, Spain 156

avanti, Polusocrinus ... 173, 185
AxinOlObUS® feece ce
azygous, Schistocrinus 215, 216

B
“Barnett Hill

Formation” 147, 149
Barnett Hill

Member .. 147, 148,

149, 154, 155, 156, 158
Barnett Hill, Okla. 147, 149,
150

Barnett homesite

(Okla.) 149
Barnett Shale or

Formation 142, 145,

146, 156, 198, 202, 203, 217
barnettensis,

Culmicrinus 50, 52 146, 202,

203, 260, 262
Bartlesville, Okla. 167
barumensis,

Poteriocrinus 199
baryactylus,

Aesiocrinus 188
Baskirian (C.b) 146
basilicus,

Aesiocrinus . 188
Bashkirian

Horizon 157, text-fig. 2
Bashkirian Stage 146, 147,

155, 156, 157, 158
bassleri,

Isoallagecrinus 160
bassleri, nodosus,

Allagecrinus ... 160
bassleri status,

Allagecrinus .... 160
bassus, Araeo-

crinus ..........42, 51,53 153, 196,

197, 252, 261, 263
Bathronocrinus ............ 203

beedei,

Parulocrinus 163
Beggs, Okla. 173
Bend, Texas 143
Bend or Bendian

Series 157, 224
benthobatus,

Delocrinus 160
Big Saline

Formation 141, 142,

143, 145, 152, 153, 154, 156,
157, 164, 165, 166, 167, 168,
169, 173, 181, 182, 183, 213,

215, 216
biplex, Allagecrinus 217
biplex, Thaminocrinus 217
biplex, Wrightocrinus 217

Bird Spring Formation 154, 158
Bisatoceras ....... 149

Bisatoceras sp. 149
Blackjack School

Sandstone Member .. 148
blairi, Parulocrinus . 161,163
blairi, Ulocrinus ........ 163
Block Member .............- 155
Bloyd Formation 145, 146,

147, 148, 151, 156, 162
Bluejacket Sandstone 152, 8,

bockii, Fusulinella 152, 159
bockshii, Woes 161
Boesites ae 149
Boggy Formation eee 148, cr
Boston Mountain, Ark. ie
Bostwick Formation
(conglomerate) ..... 145, 148,
151, 153, 156, 157, 158, 224
brachiatus,
Paradelocrinus .. 159, 182
‘Synarmocrinus 154, 161,
164
Braggs Mountain,
Okla. 148, 150
branneri,
Branneroceras .. 147, me
Branneroceras 146, 147,
153, 156, 159
Brannon Bridge
Limestone Member .. 141, 143,

145, 159, 166, 170, 171, 172, 174,
175, 177, 178, 179, 180, 183, 191,
192, 194, 199, 200, 202, 208, 209,

211, 213, 215, 230, 231

Brazil Formation ee lay bys)
brazoensis,
Ecudoperalecocei
Cie 148
Brazos River Valley
(Texas) : 159
Breathitt Formation 146
Brentwood Limestone
Member . _ 145, 147,
162, 163, 232
brentwoodensis,
Acrocrinus a : 220
Bridgeport, Tex. . .. 148, 146,
160, 167
bridgeportensis,
Graphiocrinus 160
Bridgerocrinus 199

INDEX

Brock, Tex. 142, 143,
145, 159, 166, 170, 171, 172, 174,
175, 177, 178, 180, 183, 192, 194,
199, 200, 202, 208, 209, 213, 215,

230, 231
Bronaughocrinus 144
Brook Ranch Member .. 143
Brownville Limestone
Formation 225, 229
Brownwood Shale 172
Brychiocrinus 159, 175,
177
bulbosus,
Paragassizocrinus 151, 188
bulbus,
Stellarocrinus . 44, 47 159, 175,
178, 254, 257
Burgner Formation 145, 152,

153, 154, 156, 157

Burlington Limestone 220
Burnet County, Texas 146
burrensis,

Profusulinella ....... 155
buttsi, Ulocrinus .......... 161, 164,

165, 166
Cc
Cm2e (Moscovian,

Kashirian -

Podolskian) .. 157
Cm2d (Moscovian,

Podolskian Horizon) 157
Cm2e (Moscovian,

Myachkovian

Horizon) .... 157
C3 (upper

Carboniferous,

Missourian

Wairgilian)) 2. 200, 213,

233
Cb2d (Bashkirian) . 157
Cb2e (Bashkirian,

Vereyan Horizon) . 157
Cm2 (Moscovian) ....... 157, 159
Caballeros Canyon

(Mexico)) ee Slat alba
Cabaniss Group ...... 148, 153
Calciferous Sandstone

Senlests Gees PPT PA
Caldenocrinus 224, "226,

297, 228, 229
caliculus,

Paragassizocrinus 188
Callville Formation ..... 146, 154,

185
calyculoides,

Polusocrinus . 42, 43 146, 153,

154, 186, 252, 253
campbellae,

Dimorphoceras .......... 152

Dimorphoceratoides 146
cancellatum,

Branneroceras 157
Cantabrian Mountains

(Spain) nee 158, 159
Canyon Group ............ 167, 212,

223
Canyon Series .... 172
Captain Creek

Limestone Member .. 159, 160
Carinocrinus ... : 202
carmani, Fusulinella . 153, 158
Carterville, Mo. 1

268

castus,

Mantikosocrinus 161
caverna, Parulocrinus . 163
caverna, Ulocrinus 163
Cedar Creek (Iowa) 153
Chappel Limestone . 142, 146
Cherokee County,

Kans. 153
Cherokee County,

Okla. 150
Cherokee Formation

or Shale 148, 153,

154, 155, 156, 157, 158
chesterensis,

Ulrichicrinus ; 173
Chesterian Stage 142, 145,

146, text-fig. 1, 2, 161, 173, 175,
185, 187, 188, 189, 190, 191, 194,
195, 198, 202, 203, 216, 217, 219,

220, 221, 222
Chlidonocrinus 146, 153,
188, 189
Chuanshan, China 149, 158
Cibolocrinus .... 146, 160,
232
ciscoensis?

Qzawainella .... 151

Ciudad Victoria,
Mexico SnG aly
Clarita, Okla. 147, 149,
150, 157
Clark County, Nevada 146, 154,
158
clarki, Fusulinella ; 152; 153

elarus, Anchicrinus .... 207, 208
clarus, Athlocrinus ...... 207
Coal County, Okla. . 147, 149.
150, 151, 154, 157, 158
“Coccocrinus”’ ee 221
Coffeyville Shale .......... 232
Comatilar pec. 144
compactus,
Paracromyocrinus 164
compactus,
Parulocrinus 164
compressum,
Phaneroceras ........ 153
Pseudoparalegoceras | 149, 151,
153
confertus,
Schistocrinus 210, 211,
212, 215
confertus,
Sciadiocrinus 43, 52 211, 212,
213, 215, 253, 262
Goleocrinus ......... 38 153, 167,
168, 248
conlini,
Stomiocrinus . 40 141, 142,
145, 222, 250
conoideus, Erisocrinus 181
conoideus,
PUOLOCHINUS) eee eee 165
consectatus,
Ramulocrinus 44 159, 200,
201, 254
contentus,
Cymbiocrinus .......40 142, 146,
190, 191, 250
convexus, Delocrinus .. 191
convexus, Ethelocrinus 165
convexus,
Ulocrinus 52, 55 160, 165,
166, 262, 265

Conway County, Ark. .. 152
Coody Sandstone
Member x.4:0.8r.. 148, 150,
156, 158
cooksoni, Phanocrinus 216
copani,

Isoallagecrinus 218
copiosa,

Profusulinella 153, 154
Coral Zone 3

(D. Hill) 218
cornutus,

Lasanocrinus . 146, 153,

208
coryphaeus,

Ulrichicrinus = iy;
Corythocrinus 195, 196,

197
Cosmetocrinus as 201
Cotarrazzo Limestone 156
Craig County, Okla. 148, 190
crassidiscus,

Hydreionocrinus 208

PlaxOcnrinus?! fone .c.e: 208
crassidiscus,

Plaxocrinus . 208
crassacanthus,

Sciadiocrinus? _. 211
Crawford Conntys. Ark. 152
Cricocrinus ....... . 159, 181
Cromyocrinus ... 156, 159,

160, 161, 164, 165, 166, 167

croneisi, Diphuicrinus . 146
Cryphiocrinus soa 160
Culberson UB

Texas ..... oe 150
Culmicrinus .. 146, 202,

203
curtus, Caldenocrinus .. 228
“Cyclical

Formation F” 155
Cyathocrinus 165
cylindrica,

Fusulinella 159
Cymbiocrinidae Pree ie
Cymbiocrinus ............ 146,

185, 188, 189, 190, 191, ir 195
C.b (Bashkirian) _ - 146
C.ks (Kashirian) 199
C.M Moscovian _ 159, 223
Cmb (Moscovian,

Vereyan-Kashirian) ’ 157
C:M: (Moscovian) 159
Cm,a (Moscovian,

Vereyan) ....... 157
C, (Moscovian) . 195
Cb.a (Limestone G:,

lower Bashkirian) 147, 156
Cb.b (Limestone oa

ge imeter Seen LEY eG YY

(Bashkirian) . . 147, 157
D
Dactylocrinus Rete 224
daileyi, Lasanocrinus 208
dakotensis,

Fusulinella 152, 154
Daralites ......... 153
Dasciocrinus .. 209
Decadocrinus 199, 201
Decadocrinus? ep ere 150
Deese Formation . > alvath aly)

INDEX

delicatus, Dichocrinus 222
delicatus,

Microcaracrinus 44, 46 159, 201,

254, 256

Delocrinus 144, 160,

191

Delocrinus sp. 160
deltoideus,

Paragassizocrinus 187, 188

Dennis, Tex. 143, 179,

192, 205, 206, 210, 211, 212, 220,

233, 234
“Derry” 148
Derry, New Mexico 154
“Derryan” 1525103:
Derryan Stage or
Series 141
Des Moines River 158
Desmoinesian Stage
or Series 141, 143,

145, 146, 147, 148, 149, 150, 152,
153, 154, 155, 156, text fig. 1, 2,
157, 158, 159, 160, 161, 162, 164,
165, 166, 167, 169, 170, 171, 172,
173, 174, 175, 177, 178, 179, 180,
181, 182, 183, 188, 191, 192, 193,
194, 195, 199, 200, 201, 204° 205,
206, 207, 208, 209, 210, 211, 212,
213, 214, 215, 218, 220, 221, 227,

229, 230, 231, 233, 234

detrusus, Aesiocrinus 194
devexa, Fusulinella 154
Diaboloceras 146, 149,
150, 152

Dichocrinus 141, 153,
220, 221, 222, 233
Dicromyocrinus 145, 146,

159, 160, 161, 162, 163, 169

dilatus,

Dichocrinus 36, 41 153, 222,

246, 251

Dimorphoceras 152
Dimorphoceratoides 146
Diphuicrinus 146, 182
Dirty Creek Sandstone

Member 148
discoidalis,

Eowellerites 151, 152
disculus, Cricocrinus 181

Paradelocrinus 181

Paragassizocrinus 188

Paragassizocrinus

(ing 146

disculus, Sciadiocrinus 211, 213
distinctus,

Stellarocrinus 175, 176,
184
Donetz Basin, Russia .... 147, 156,
157, text-fig. 2
Dornick Hills Group 173
dubius,
?Paradelocrinus . 182
E
echinatus,

Chlidonocrinus 42, 51 146, 153,
188, 189, 190, 252, 261

Eiromocrinus .......... 159, 203,

204
elegans, Acrocrinus 221
elegans, Culmicrinus 202
elenae, Phialocrinus 195

269

elenae, Phialocrinus? 195
elenae, Polusocrinus 195
elevatus, Erisocrinus 180
elevatus,

Paragassizocrinus 151, 188
Elibatocrinus 160, 203
Elibatocrinus sp. 160
Ellipsellipsopa 232
ellipticus, Columnal 233

Paragassizocrinus 188

Parethelocrinus 171, 172
elongatus,

Parragassizocrinus 188
elongatus, Ulocrinus 165
Enascocrinus 227, 228,

231
Endelocrinus 159, 160,

182
Eoasianites 149, 151
Eoasianites sp. 149
Eoshubertella 150, 153
Eoshubertella sp. 150
Eostaffella 146, 147,

156
Eowellerites 151, 152
erectus, Ampelocrinus 185
erectus, Cibolocrinus 160
erectus, Erisocrinus 180
erectus,

Isoallagecrinus 41,52 153, 217,

218, 251, 262
Erisocrinus 141, 144,

153, 160, 180, 181, 182, 216
Erisocrinus sp. 160
Espey Creek, Tex. 141, 142,

145, 163, 169, 182, 184, 191

Ethelocrinus 159, 163,

165, 168
“Etroeungt” 199, 202
Euonychocrinus 160, 227
Eupachycrinus Patil
euthusepta,

Fusulinella 150
eventus, Carinocrinus 202
Exaetocrinus 181
Exocrinus 144, 175,

183
expansus,

Acrocrinus 221
expansa, Staffella 153
extrorsus,

Ulocrinus 56 165, 166,

167, 266
F
farishi, Synerocrinus 151, 224
Fayettevile
Formation 146, 190,
191
fittsi, Profusulinella 147, 149,

150, 152, 153, 154, 155
Fittsi, Profusulinella,

Zone of 154
florealis, Apollocrinus 159
florealis,

Stellarocrinus 159, 177,

178

Forbesiocrinus 223, 224,

225, 228, 229

Forest City Basin, Mo. 149
formosum,

Gastrioceras 149, 151

formosus,
Aexitrophocrinus
45, 47 225, 227,
229, 230, 231, 255, 257
formosus,
Synerocrinus .. 159, 226,
227, 228, 229
Ft. Gibson Lake, Okla. 150, 151
Ft. Scott Limestone 148
fortunatus,
Sellardsicrinus 35 159, 170,
171, 245
Fra Cristobal
Formation 152,155,
157
Franklin County, Ark. 152
Frenchman
Mountain, Nev. 146, 154
Frensley Limestone . 173
frostae,
Oklahomacrinus | 34 142, 159,
194, 244
fugax, Fusulinella 152
fundundus,
Synarmocrinus 150, 164
furnishi,
Fusulinella . 154
Fusulina ; 148, 153,
155, 156, 157, 158, 159
Fusulina s.s. il Bh 15s)
Fusulina Zone 152, 154,
155
Fusulinella ... -.. 147, 149,
150, 152, 153, 154, 155, 156,
157, 158, 159
Fusulinella Zone 152, 154,
155, 158, 159
Fusilinella profifica
HONG Ptcn ate 156, 157
G
Garnett Quarry 211, 212,
231
Gastrioceras 146, 147,

149, 151, 157, 158
Gastrioceras venatum

assemblage or zone 159
geminatus,

Cromyocrinus 159, 161
geminatus,

Mooreocrinus 159, 160,

161, 162
geminatus,

Trautscholdicrinus 161
Gene Autry Shale 146, 151
Genevievian Stage 216, 219
Genus and species

indeterminate 32 169, 242
geometricus,

Stellarocrinus 176
georgeae,

Erisocrinus 39 141, 153,

180, 181, 249
Georges Fork

Sandstone Member 148
gibberellus,

Schedexocrinus 205
Gielian Stage 158
Gilmore City

Formation f 202
girtyi, Alcimocrinus .... 173
girtyi, Boesites ............ 149
girtyi, Dichocrinus ...... 222

INDEX

Glaukosocrinus 159
Glaphyrites 153, 160
Glen Dean Formation .. 202, 222
Globacrocrinus 153, 220,

221
globulosus,

Eoaianites 149
Goleocrinus 153, 160,

161, 164, 167, 168
goliathus,

?Cyathocrinus 165
Gonialoboceras 153
Gonialoboceras sp. 153
Goose Creek (Okla.) 147, 149
Gordonites 158
gracilis, Texacrinus 159, 174
graffhami,

Isoallagecrinus 218
Graffhamicrinus 146, Ps.
Graford Formation 141, 142,

143, 145, 146, 159, 160, 165, 167,

176, 177; 223
grafordensis,

Endelocrinus 160
grandis, Cromyocrinus 166, 167
granulosus,

Utharocrinus cf. 160
Graphiocrinus 160
gravis, Cymbiocrinus 191
Greenland Sandstone

Member .................. 152
Greenleaf Lake, Okla. 162
“Griley Limestone” 150
grossus,

Eiromocrinus 30 159, 204,

240
H
Haeretocrinus 159
Hampton Formation .... 221
Hare Fiord ible ilsye
158
harrisae,

Sciadiocrinus 159, 211
Hartshorne Coal 148
Hartshorne Formation

(Sandstone) ................ 147, 148,

149, 154, 156, 157
Hartville Formation .... 154, 157
Heliosocrinus ' 175
henbesti,

Winslowoceras Ml aap
Herruela, Spain 156
Hexacrinites 221
Hexacrinus 221
Holdenville

Formation 165, 172,

173
holdenvillensis,

Metacromyocrinus 161
Hobbs; Ark> jo. : 152
Honey Creek (Texas) . 143
Hood County, Tex. ...... 148, 159,

179, 192, 199, 200, 205, 206,210,
211, 212, 220, 233, 234

hoodi,

Paragassizocrinus 187
Hotel San Saha ....... 232
Houkou Limestone 5 sey ast:
hoveyi, Hypselocrinus .. 198

Huanglung Limestone .. 149, 158
hyattianus, Glaphurites 153

270

Hydreionocrinus 210, 211

Hydriocrinus 141, 142,

198, 199

Hypselocrinus . 197, 198

I

Iberocrinus ; 156

Illinois Basin 155, 156,

158

Imo Formation 142,191
incurvus,

Forbesiocrinus 223, 224,

225, 228, 229
incurvus,

Synerocrinus 224, 225,

226, 227, 228, 229
Indian Springs, Nev. 154
indicus, ?Ulocrinus 165
Inola Limestone 153
insculptus,

Corythocrinus 196
insoliata, Fusulina 148, 155
insularis,

Dichocrinus 221
intermedius,

Acrocrinus 220
invaginatus,

Pirasocrinus 160
iolaensis,

Paradelocrinus ..... 182
iowense,

Paralegoceras ............ 149, 153
iowensis,

Busulinellay 2)... ee 153, 154,

155, 158
Tsoallagecrinus ....... 144, 150,

153, 160, 217, 218

Itaituba Formation . 161
ivanovi, Aesiocrinus 195
J

jackeli,

Talanterocrinus oe eee
jackeli,

Trautscholdicrinus 225, 226
jakovlevi,

Alllasecrinusm cee 217
jakovlevi,

Wrightocrinus .. Pali
Jasper County, Mo. 152
Jefferson County,

Ala... eee 146
Jefferson County,

ROWAN cctnna ess Mate 153
jeffersonensis,

Calmicniniss ees 202
johnstonensis,

Cricoenmusse sae 181
johnstonensis,

Paradelocrinus .. 181

K
Kalkbriiche von

Mjatschkowa ............ 159
Kansas City (Mo.) ........ 176
Kansas City Group ...... 176
kansasensis, Ulocrinus 165
Karatange Horizon .. 157
Kashiran Horizon ........ 155; 157,

15

Kendrick shale 0

PEGS. piiecseeeee nt 1465152;
F 154
kendrickensis, _
Paragassizocrinus 146, 188
Kentucky River Valley 154
kentuckyensis, 2
Profusulinella 155
Keokuk age 173, 200
kesslerense,
Pseudoparalegoceras 149, 151,
1525155
Kettwig, Rheinland ..... 199
Kickapoo Creek, Tex. .. 159, 179,

192, 205, 210, 211, 212

Kickapoo Falls .............. 143, 145,
159, 178, 179, 192, 200, 206,

208, 210, 211, 212, 215, 219,

220, 233, 234

Kickapoo Falls
Limestone Member .. 141, 148,
145, 159, 179, 191, 192, 199,
205, 206, 208, 210, 211, 212,
215, 220, 233, 234

kickapooensis,
Lecobasicrinus 31,53 144, 159,
191, 192, 194, 241, 263

Kinderhookian .......... 202, 221,
222
Kinkaid Formation . 142
Knightoceras ........... : 152
Kothman Ranch, Texas 142, 198,
203, 217
Krebs Group . 148, 152,
153, 158
LE
laevis,

Platyerinites ... 218

Lake Bridgeport Dam,
Southwest of ....... 143, 223

Lake Bridgeport Shale
Membeniec. tio 143, 159

Lake Bridgeport, Tex. 141, 142,
143, 160, 167, 172, 176, 177

Lake Murray, Okla. 172
Lake Murray State
Park Seer 173
Lambert Ranch, Tex. .. 141, 142,
145, 163, 195, 231
lamberti, Aexitro-
phocrinus cose. 84 142, 229,
231, 244
Lampasan Series ........ 141, 149,
154

Lampasas County, Tex. 141, 142,
169, 182, 184, 191

“Lampasas”’ (Series) 148
Lampasas, Tex. ............ 142, 145,
162, 163, 169, 184, 191

Lane Shale Formation 176
Lasanocrinus ................ 146, 153,
203, 208

Latimer County, Okla. 150, 158
Laudonocrinus ...... 160, 203
laxus, Plaxocrinus . 160
Lebetocrinus ....... 196
Lecobasicrinus .. 144, 153,
159, 188, 189, 191, 193, 194, 197
Lecythiocrinus ...... 160
Lecythiocrinus sp. f 160
leei, Fusulina . 153, 158

INDEX

Lemons Bluff Member 141, 143,
145, 146, 153, 154, 185, 186,
188, 190, 193, 194, 195, 197,
198, 208, 218, 221, 222, 231

Lester Limestone
148, 156,

Member ..
text-fig. 1, 158
libratus,
Polygonocrinus 33, 56 159, 208,
209, 214, 215, 243, 266

lichengensis,

Sinocrinus 183
Limestone G, 147
Limestone H, 147
Lipan, Tex. 211
Liparocrinus .... 201
listeri, Gastrioceras 157

Little Cabin Sandstone 148, 154
Llano Uplift or

Region, Texas .. 141, 142,
152, 153, 154, 156, 157
llanoensis, Fusulinella 153
llanoensis, Pronorites 153
llanoensis,
Sciadiocrinus 49 153, 211,
213, 259
lobatus, Plaxocrinus 160
Lodgepole Formation 202
longus,
Stenopecrinus 48 159, 209,
258
Loring Siding, S.
Dakota 154
lorrainae,
Hydriocrinus 34 142, 198,
199, 244
Louisiana Limestone 218
Love County, Okla. 148, 172,
173
Lower Limestone
Group 218
Lower Mercer
Limestone 154, 155,
158
lustrum, Erisocrinus 181
luxuris, Aesiocrinus 194
lykinsi, Aesiocrinus 188
Lyonicrinus 221
M
Madison County, Ark. 152
magna, Fusulinella 149
magnus,
Aglaocrinus 32 153, 172,
173, 242
magnus, Ethelocrinus 173
magnus, Haeretocrinus 159
Magoffin beds 146, 154
Malaiocrinus 159
Mantikosocrinus 161
Marble Falls
Formation 141, 142,

143, 145, 146, 153, 154, 157,
163, 169, 182, 184, 186, 188,
190, 191, 192, 193, 194, 195,
197, 198, 208, 217, 218, 221,

222, 231, 232

“Marble Falls Group” 157

Marble Falls, Tex. .. 146

marblensis, Millerella 146
marblensis,

Profusulinella 153

Mariocrinus ......... 199

Marmaton Group 148, 212,
231
marquisi,
Paracromyocrinus 164
marquisi,
Parulocrinus 164
marrsae,
Sellardsicrinus 45 159, 170,
171, 255
Marwood bed 202
Mason County, Tex. 142, 143,

145, 146, 164, 167, 168, 169,

173, 181, 182, 183,198, 203,

213, 216, 217

Mason, Tex. 141, 142,
143, 145, 146, 153, 167, 168,

169, 173, 181, 182, 183, 198,

203, 213, 216, 217

.. 39 153, 161,
167, 168, 169, 249

masonensis,
Goleocrinus

matheri, Endelocrinus 182
McAlester Formation
(Sandstone) 148, 149,
154
McCulloch County,
Tex. 143, 160,
172
mcguirei,
Aexitrophocrinus 227, 229,
231

Paragassizocrinus 188
mcguirei,

Talanterocrinus 227, 228
mcguirei,

Trautscholdicrinus 225
McLouth Formation 152
mediator, Erisocrinus 180
Megaliocrinus 156
Melbacrinus 198
mendesi,

Dicromyocrinus 161

Mooreocrinus 161, 162
Meramecian 156
Mesolobus 153
Metacromyocrinus 146, 150,

161, 163
Metallagecrinus 217
Metaperimestocrinus 203, 209
Microcaracrinus 159, 201
microgranulosus,

Sinocrinus 181, 183
microgranulosus “var.”

pentalobus,

Sinocrinus 183
Millerella 146
Millsap Lake Group 159

Millsap Lake Formation 141, 142,
143, 145, 156, text-fig. 1, 159,

165, 166, 167, 170, 171, 172,

174, 175, 177, 178, 179, 180,

183, 191, 192, 194, 199, 200,

202, 205, 206, 208, 209, 210,

211, 212, 213, 215, 220, 229,

231, 233, 234

millsapensis,
Ethelocrinus 159

millsapensis,
Parethelocrinus 159, 172

miloradoivitschi,

Trautscholdicrinus 200, 201
Minnelusa Formation .. 152, 154,
155.1 5if

Mineral Wells

Formation 143, 146,
160
Mineral Wells Group 160
Minshall Member 155
missouriensis,
Culmicrinus 202
Missourian Stage
or Series 141, 143,

145, 146, 159, 160, 161, 164,

165, 167, 172, 174, 175, 176,

180, 181, 182, 188, 191, 193,

197, 199, 200, 210, 211, 212,

218, 219, 221, 227, 232
Mjatschkowa (near

Moscow, Russia) 159
mjatschkowensis,

Acrocrinus 220
molleri, Staffella 150
Monobathra 218-223
Montgomery County,

Kans. : 160
moorei, Eowellerites 151
moorei, Eowellerites

cf. E. isl Wey
Mooreocrinus 141, 153,

159, 160, 161, 164, 168
Morrowan Stage cevee WAL

145, 146, 147, 148, 149, 150,

151, 152, 153, 154, 155, 156,
text-fig. 1,2, 157, 158, 161,

162, 163, 164, 166, 167, 169,

173, 181, 182, 183, 184, 185,

191, 199, 210, 211, 218, 220,

229, 232

Moscovian Stage .......... 141, 151,
156, 157, text-fig. 2, 158, 159,
160, 161, 195, 199, 219, 220,

223, 228, 229
Moscow Basin, Russia .. 146, 157,
text-fig. 2, 159
Moscow, Russia 158, 159,
223, 225, 226

mosleyi,

Perimestocrinus 160
mosleyi,

Stenopecrinus % 160
Mound, the (Oklahoma) 167
Moundocrinus 185, 186
Mud Springs

Mountains, N. Mex. 154, 155
multibrachiatus,

Iberocrinus 156
multiextensus,

Polygonocrinus 213, 214,

215
multinodosus,

Decadocrinus 199
multiplicatus,

Globacrocrinus 52 153, 220,

221, 262
mundus, Mariocrinus? 199
Muskogee County,

Okla 150, 163
Muskogee, Okla. 150, 163
mutabilis, Fusulina 148
Myachkovian horizon | 156, 157,

159

Myzostoma 145
N

Namurian 157, 158

228

INDEX

needhami, Pseudost-

affella aff. P. 153
Neodichocrinus 222
Neozeacrinus 144, 159,

174
nereus, Decadocrinus 199
neumeiri, Diaboloceras 146, 150
nevadaensis Zone,

Rhipidomella 154
nigelensis,

Ramulocrinus 199
nitidus, Athlocrinus 160
nodosus, Sinocrinus 183
nodosus ‘“‘var.” spinosus,

Sinocrinus . 183
normalis,

Plaxocrinus 203
Nuculoceras 153
Oo
obesus, Plaxocrinus 159, 208,

209, 211

obesus, Sciadiocrinus .. 209, 211,

212

obovatus, Erisocrinus 180
obovatus,

Paradelocrinus 182
occidentale,

Gastrioceras 146
occidentalis, Ulocrinus 165
Ochelata Group 167
ochelataensis,

Polusocrinus ys}
oeconomicus,

Plaxocrinus 160
oklahoma,

Ulrichicrinus Ne 173
Oklahomacrinus 142, 150,

159, 188, 189, 191, 193, 194, 195
oklahomensis,

Ethelocrinus 164
oklahomensis

Metacromyocrinus 146, 163
oklahomensis,

Paracromyocrinus 163, 164
Okmulgee County,

Okla. Dy facta 173
Oligocrinus . 224
oliviformis,

Fusulinella 149
omphaloides,

Plaxocrinus 160
onychocrinus 224
Oolokah Limestone

Formation 180, 181,

207, 212, 230
optimus,

Dicromyocrinus .. 40 146, 162,

250
Origocrinus E 224
ornatus, Alcimocrinus 173
ornatus, Cromyocrinus 159, 162
ornatus,

Dicromyocrinus 159, 160,

162

“ornatus, Trautscholdi-
crinus” . 161

Osage County, Okla. 167, 225
Osagian Stage 142, 146
Otterville, Okla. 146

Otterville Formation
(Limestone) 146, 148,
151, 156, 224

272

oxylobatum,

Knightoceras 152

Oxynocrinus 146, 183,
184
Ozark National

Forest co 152
Ozark Region, Ark. 151
Ozawainella 151
Ozawainella sp. 151

P
Pachylocrinus 144
pachypinnularis,

Pachylocrinus 144
pachyplax,

Polusocrinus 146, 185
Pahkra River

(Russia) 158
Paianocrinus 203
Paint Creek

Formation 222
Palencia, Province

of (Spain) . 156
Palo Pinto County,

Tex. : ; ; 212
Palo Pinto Limestone 212
Paracromyocrinus .. 160, 161,

163, 164

Paradelocrinus 150, 156,

159, 181, 182

Paradelocrinus sp. ...... 156

Paragassizocrinus 145, 146,

151, 153, 160, 184, 185, 186,

187, 222, 223

Paralegoceras .... 149, 151,

152, 153

Paralegoceras sp. 153

Paraschistoceras 160
“Paraschistoceras

reticulatum zone” .... 160
Parethelocrinus ate, ABO WM S9e

160, 163, 171, 172
Parker County, Tex. 143, 159,

166, 170, 171, 172, 174, 175,
177, 178, 180, 183, 192, 194,
199, 200, 202, 208, 209, 211,
213, 215, 230, 231
Parulocrinus _................ 154, 160;
161, 163, 164, 165, 169
parva, Profusulin-

ella ex gr. ... 157
pavisculus,

Glaukosocrinus 159
parvisculus,

Malaiocrinus . 159

Schistocrinus 215
parvus, Schistocrinus-

confertus,

Sciadiocrinus . 159, 210,

jotd

parvus, Sciadiocrinus. 159, 212
patens, Aesiocrinus 195
patens, Phialocrinus . 195
patina, Diphuicrinus 146
patulus,

Agassizocrinus 187
paucus,

Aesiocrinus ................. 188, 191,

193

pavcus,

Lecobasicrinus en ieals ir
pecki, Isoallagecrinus .. 218
‘Pelecocrinus , 201

Pella beds 156
penicillus,

Platyerinites ............. 219
Pentadelocrinus _.......... 188
pentalobus,

Dichocrinus : 222
Pentaramicrinus eee 216
Pentremites 141, 142,

146
peramplus,

Neozeacrinus 144
Perimestocrinus 160, 203,

209
periodus, Ethelocrinus 172, 173
Permian (P,) ,........ 142, 161

163, 165, 180, 185, 186, 195,
203, 209, 217, 218, 219, 222, 232

permicus Platycrinites 219
Permo-Carboniferous 165
perundatus,
Plaxocrinus .............. 159, 208,
209, 215
Petschoracrinus ............ 185, 186
Phaneroceras 153
Phanocrinus ........... - 146, 216
Phialocrinus! =...-.--0-7. e163: 188,
195
pictus, Delocrinus ........ 160
Pilton bed .... 202
Pimlicocrinus ......... 156
Pimlicocrinus sp. 156
Pirasocrinus .. : ~ 150, 159,
160, 203, 204, 205, 206, 208,
211, 214
pirum, Acrocrinus 145, 220,
221
pirum, Globacrocrinus 220,221
Pitkin Formation ....... 187, 198,
202
pitkini, Cymbiocrinus 194
planulatus,
Schistocrinus 159, 210,
211, 215
planulatus,
Sciadiocrinus 159, 211,
213, 215
planus, Cricocrinus 181
planus,
?Paradelocrinus 181
planus,
Perimestocrinus 209
planus,
Stenopecrinus 209
plattsburgensis,
Ethelocrinus 163
plattsburgensis,
Parulocrinus 163
plattsburgensis,
Parethelocrinus _. 49 160, 163,
172, 259
platybasis,
Eupachyerinus . 211
platybasis,
Sciadiocrinus 211
Platycrinites 141, 159,

190, 218, 219, 220, 221, 232, 233

Platycrinites sp. 232
Platycrinus 221, 232
Platyerimusial sss 232
Platycrinus sp. 219, 232,
233

Plaxocrinus .... 151, 159,
160, 203, 204, 205, 208, 209,

Pill, 212525

INDEX

plumifer,

Scaphiocrinus? 199
Podolsk Quarry, Russia 159
Podolsk, Russia ee LOG
Podolskian horizon ...... 155, 157,

158, 159

Polusocrinus _..... 146, 153,

154, 173, 185, 186, 195

Polygonocrinus ............ 159, 203,

204, 208, 209, 213, 214, 215

popes ne

OS ee ee 52 214, 262

pantatne County, Okla. 147
Pool Stock Farm,

Texas 142
Pope Chapel Sandstone

Member hac ess 148
Port Birmingham, Ala. 146
post-Pitkin eset te 142
Poteriocrinus 199, 202
praecursor, Acrocrinus 220
praecursor,

Anobasicrinus 43 159, 174,

253
praecursor,

Neozeacrinus . 159, 174
praegorskii,

Pseudostaffella ........ 147
prima, Fusulina ........ 158

primaeva, Fusulinella 152, 153,

154
primitiva,

Profusulinella 147, 157
primitivus,

Acrocrinus 221
Profusulinella ....... 146, 147,

149, 150, 152, 153, 154. 55s

157, 158

Profusulinella Zone 154, 155
Profusulinella

fittsi zone 154, 156,

157

prolifica, Fusulinella _ 147, 149,

150, 153, 154, 155

Pronorites ee, 153

Pronorites sp. ..... 153

propinquus,

Erisocrinus 180
Protencrinus 191
protensa, Fusulinella 154
protensus,

Erisocrinus 180
protensus,

Paradelocrinus 182
prudentia Aesiocrinus 188, 193
Pseudoparalegoceras 148, 149,

150, 151, 152, 153, 155, 158,159
Pseudoparalegoceras

brazoense Zone 159
Pseudoparalegoceras

sp. 150
Pseudostaffella 146, 147,

153, 156, 157
pumpkensis,

Acrocrinus 221
Pumpkin Creek

Limestone .................. 154, 173,

209

punctatus, Cibolocrinus 146, 232

purpurea, Comatula 144
pusillus, Hydriocrinus 199
pustulosus,

Paracromyocrinus 164

273

pustulosus,

Parulocrinus 164
Putnam Hill Formation 158
Q

quadrangulus,

Columinaliee.. 232
quadriceptalus,

Dichocrinus 5 221
quinni, Axinolobus 151
R
radiatus, Dichocrinus .. 221

ramosus,

Ulrichicrinus 31,55 145, 159,

173, 174, 196, 241, 265

Ramulocrinus 159, 199,

201

Ratingen, Germany 199, 202
ratingensis,

Hydriocrinus 199
raymondi,

Glaphurites .. RA 153
rectus, Endelocrinus .... 159
rectus,

Tholiacrinus ....44 179, 180,

254
Red Oak, Okla. 150, 158
redesdalensis,

Enascocrinus 227, 228,

231
redesdalensis,

Talanterocrinus .. 227, 228,

t 231
regularis, Culmicrinus 202

regularis, Decadocrinus 201
Poteriocrinus 202
Trautscholdicrinus 201

regulatus, Cricocrinus 181
Paradelocrinus 181

remotus,

Platycrinites ...47 159, 219,

257
repertus,

Ramulocrinus 200
reticulatum,

Gastrioceras cf. 157
reticulatum,

Paraschistoceras 160
Rhipidomella 154
Rhipidomella

nevadensis Zone 154
rhombiformis,

Profusulinella 157
rimulatus,

Tholiacrinus 180
Riverton Coal < 148
Riverton Formation 148, 149,

152

Rochelle, Tex. 143, 146,

160, 172

Rockhill Limestone
Member 143, 223

rosei, Hydriocrinus - 199

rotaii, Dichocrinus . 221

rothi, Amphisites .. 155

rotundus,

Allocatillocrinus 4 218
Rough Creek, Tex. 143, 145,

186, 188, 190, 193, 195, 197,
198, 208, 218, 221, 222

Rough Mountain
Conglomerate 172

rugosus, Stenopecrinus 146, 203

russiense, Gastrioceras 158
russiense,

Pseudopareloceras 158, 159

Ss
Sagenocrinoidea 223-232
St. Genevieve

Formation : 156
St. Louis Formation 185, 202
St. Louis, Mo. . 202
salebrosus, _

Scaphiocrinus 199
Salesville, eae 212
San Saba County,

Tex. 143, 145,

163, 186, 188, 190, 193, 195,
197, 198, 208, 218, 221, 222,
231, 232

San Saba, Tex. .... _ 141, 142,
143, 145, 186, 188, 190, 193, 195,
197, 198, 208, 218, 221, 222,

231, 232

sangamonensis, 3

Cyathocrinus 165
sangamononsis,

Ulocrinus 165, 166

sansabensis,

Hypselocrinus ....... 197, 198

Scytalocrinus 41,51 146, 153,

197, 198, 251, 261,

Scaphiocrinus a 199

Schedexocrinus 203, 205,

206, 208, 214

Schistocrinus 153, 159,

191, 203, 210, 211, 215

schmidtii, Dichocrinus . 233
schmidtii,

Platycrinites? 232, 233
schmidtii, Platycrinus 232
Schubertella ............ 150
Sciadiocrinus ..... 153, 159,

191, 203, 204, 206, 207, 210,

Al OA RMD:

Scotiacrinus 169, 170
scotti, Daralites 153

seotti, Piraso-

crinus . 33, 35, 45,56 159, 205,
206, 208, 243, 245, 255, 266
sculptus, Allagecrinus 218
Scytalocrinus 146, 153,
197 ,198
Seytalocrinus sp. 50 146, 198,
260
searighti,
Fusulinella 2 GRA SR)
Sellardsicrinus 159, 169,
170
Senora Formation 148
serotina Fusulinella 158
Seville Formation of
Missouri 152, 153,
155
Seville Limestone
dl.) 152, 153,
154, 155, 157

sheareri, Sinocrinus 39 141, 153,
181, 182, 183, 249
Shetlerville

Formation 222

INDEX

shumardi, Acrocrinus 220
shumardi,
Paralegoceras 153
simplex,
Cromyocrinus 156, 159,
160, 165, 167
simplex,
Cromyocrinus sp. cf. 156
Sinocrinus 141, 153,
181, 182, 183
singulocirrus,
Culmicrinus? 202
Sloan Limestone
Member 142, 145,

146, 153, 156, 157, 232
Sloan Mountain,

Nevada 146
smithi, Ainacrinus ... 225
smithi, Stearoceras 152
Smithwick Shale 143, 150,

151, 153, 156, 157
smithwickense,

Nucloceras 153
smithwickensis,

Eoasianites < 14O TS

Soldiers Hole Member 142, 1438,

145, 153, 164, 167, 168, 169,

173, 181, 182, 183, 213, 215, 216
Southwest of Lake
Bridgeport Dam,

Texas 143
Specimen 7,
Platyerinus 232
spicata,
Ellipsellipsopa 232
spicata,
Oxynocrinus 36 146, 184,
246

spicatulus, Oklahoma-
crinus . 31,51, 53, 56 194, 195,
241, 261, 263, 266

Spirit Creek, Ark. 152
Spitzbergen, Norway 219
spitzbergensis,
Platycrinites Pere Ah eps yy
springeri,
Paragassizocrinus 188
Staffella pee Gay
153
Stanton (Limestone)
Ronmation® 1a 146, 159,
160, 165
status, Isoallagecrinus 160
Stearoceras; 22.0.0... 152
Stellarocrinus ............. 144, 145,
150, 159, 160, 170, 175, 176,
177, 178, 184
Stellarocrinus sp. ........ 160
Stenopecrinus 146, 159,
160, 203, 206, 207, 209
Stenopronorites ......... 149, 151,
152
stevensi, Carinocrinus 202
Stomiocrinus .......... 141, 142,
145, 146, 222
Stonewall Quadrangle,
Okla. _. ee 3 147
Stouti, Fusulinella .. 155
Strawn Group 166, 170,

171, 172, 174, 175, 178, 179,

180, 183, 192, 194, 199, 200,

202, 205, 206 208, 209, 210,

211, 212, 213, 215, 220, 233, 234
striatus, Mesolobus 153

274

strimplei,

Synerocrinus 229
strimplei,

Talanterocrinus «22k, 220s

229
Stuartwellercrinus 181
subglobosus,

Stomiocrinus 222
subhemisphericus,

Delocrinus 160
subidus,

Lecobasicrinus 56 153, 191,

193, 194, 266
subplanus, Cricocrinus 159, 181
subplanus,

Paradelocrinus . 159, 181
subservire,

Euonychocrinus 160
subsinuatus,

Laudonocrinus 160
superstes, Dichocrinus 222
swaledalensis,

Synerocrinus 228

Talanterocrinus 228
Synarmocrinus 150, 153,

154, 160, 161, 163, 164
Synmathocrinus : 146
Synerocrinus . 150,151:

159, 223, 224, 225, 226, 227,
228, 229, 231, 232

T
Talanterocrinus ...224, 225,

226, 227, 228, 231
Tamaulipas (State

Of); M@X. o.ccccc.: ae LOO
Tarachiacrinus . 162
tarri, Agassizocrinus 187

Paragassizocrinus 160, 187,

188
tensus, Culmicrinus? .... 202
tensus, Poteriocrinus .. 199, 202
tenuis contentus,

Platyerinites ..... 54 233, 264
tenuis,

Platycrinites ......... 54 232, 233,

264
terminalis, Erisocrinus 180
Texacrinus ........ fs oOs ioe

174
texana,

Eoschubertella .......... 153
texani, Stellaro-

CrinUs) 8 anu2 L605 LTS:

176, 177, 242

texanum, Paralegoceras 149, 151,

152, 153

texanus, Brychiocrinus 159
texanus, Stellaro-

crinus ....41, 46, 47, 52 144, 159,

177, 178, 251, 256, 257, 262
texasensis,

Dicromyocrinus ........ 145, 146,

163
texasensis, Dicromyo-

crinus sp. cf. D. .. 54 163, 264
texasensis,

Synbathocrinus ........ 146
Thaminocrinus ............. 217
Tholiaerinus ses re 179
thomasi,

Culmicrinus _........... 202
Thurman Sandstone 148

Timor, Island of . 141, 142,
218, 219, 222
toddanus, Cricocrinus .. 181
toddanus,
?Paradelocrinus 181
toddi, Anchi-
erinus ... 43, 48 142, 159,

206, 207, 208, 253, 258
tomiensis, Dichocrinus 221
torquatus, Schistocrinus 211, 215,

216
Tournaisian 156, 161,

221
transcisus,

Scaphiocrinus 199
transitonia,

Schubertella ........ 150
Trautscholdicrinus .. 199, a
Trautscholdicrinus,

nomen nudum 161, 200,

201, 225, 226
Trautscholdicrinus sp. 201
trinodi, Chlidono-

crinus ........43 146, 190,

253
trulleum,

Phanocrinus 36, 50 146, 216,

246, 260,
tuberculatus,

Platycrinus? 233
Tucker Tower Museum.

(Okla.) 3 173
Tulsa County, “Okla. 5 PAPHOS AL
tumidus, Cibolocrinus 232
tumidus, Endelocrinus 160

tumulosus, Plaxocrinus 203, 209
turris,

Paragassizocrinus 188
turris, Para-

gassizocrinus cf. P. 146
typicalis,

Apographiocrinus 160, 183
typus, Erisocrinus 180, 181
Tyrieocrinus......... 160
tzwetaevae, Pseudo-

paralegoceras . 158

U
Ulocrinus 142, 153,

160, 161, 163, 164, 165, 166, 167
Ulocrinus sp. am 160
Ulrichicrinus 144, 145,

159, 173, 174, 196
undulatus,

Tholiacrinus 180
unicus, Dichocrinus 221
Upper Mercer

Limestone 154, 155,

158

Ural Mountains
(Russia) 146, 157,
158

INDEX

uralensis, Ulocrinus 165

Ureocrinus 161

Utharocrinus 160, 203
Vv

Valymeyeran sizer

or Series : 219
Van Buren, Ark. 151, 152
Vanport Limestone 153, 158
variabilis,

Parethelocrinus 172
varicostatum,

Diaboloceras ... 149, 150,

152
varicostatum,

Paralegoceras 152
varvariensis,

Eostaffella 147, 156
velmae, Fusulinella 154
Vereyan horizon . 157, 158
Verne Limestone

Member 155
vetulus,

Paracromyocrinus 161,163,

164

vetulus, Parulocrinus 154, 164

Vinita, Okla. 190
Virgilian 161,180,

181, 183, 188, 218, 219, 225, 229
Viséan 169, 202,

218, 221, 224, 225, 227, 229, 231

Ww
wachsmuthi agnaensis,
Platycrinites 219
wachsmuthi frequentior,
Platycrinites 219
Platycrinites 219
wachsmuthi typicus,
Platyerinites 219
Wallace Creek, Tex. 232
Wangchiapa Limestone 147
Wann Formation 146, 159,
165, 166, 167
wapanucka, Cricocrinus 181
Wapanucka Formation
(Limestone) ....... 146, 147,
148, 149, 150, 151, 152, 156,
157, 163, 182
wapanucka,
Paradelocrinus 181
Warner Sandstone 148, 149,
154
Warren County, Ind. 232
Washington County,
Ark. : 152
Washington County,
Okla. 167
watkinsi, Parethelo-
crinus 44, 48,55 171, 172,
254, 258, 265

275

Wayside, Kansas 160
Webb City, Mo. 152
Webber Falls Sandstone

Member 148. 150,

151, 155, 156, 158

Wedekindellina ibbwabye

158

Wedekindellina sp. 158
West Cedar Creek,

Ark. 152
Westphalian 158
Westphalian B ... 156
Wewoka Formation 159, 165
White Breast Coal 153, 158
Whiteocrinus 175
Wiedeyoceras 151
wilburni, Moore-

ocrinus 39,49 141, 153,

162, 249, 259
williamsi,

Pseudoparalegoceras 149, 150,

152, 155

“Winslow” 146, 148,
150, 151, 152, 155, 158

Winslow, Ark. 152
“Winslow Formation” 151, 152,
155

Winslow Member 156
“Winslow Sandstone” 152
Winslowoceras 151, 152
williamsi, Wiedeyoceras 151
Wise County, Tex. 143, 159,

160, 167, 176, Ibis 293
Wolf Mountain Shale

Member 143, 145,
146, 159, 165, 167, 176, 177
Woodocrinus
Limestone 228
wortheni, Acrocrinus 221
wrighti, Cromyocrinus 161
wrighti, mamtikoso
crinus 161
wrighti, Platycrinites 219
Wrightocrinus 217
Y
Yangak horizon 146, 157
Yorkshire, England 228
Z
Zeacrinites 144
Zeacrinus 210
Zesch, Kurt (Ranch),
Tex. 141, 142,

143, 145, 164, 167, 168, 169.
173, 181, 182, 183, 213, 216
Ulocrinus 37, 38 142, 153,

165, 166, 167, 247, 248

zeschi,

Zeusocrinus 203
Zone b (Yangak
horizon) 146

an.

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PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

VOL. VI

No. 41

TRACE FOSSILS OF THE CINCINNATI AREA

By

Ricuarp G. Oscoop, Jr.

1970

Paleontological Research Institution

Ithaca, New York, U.S.A., 14850

er Menine.
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PALAEONTOGRAPHICA
AMERICANA

(Founded 1917)

TRACE FOSSILS OF THE CINCINNATI AREA

By

Ricuarp G. Oscoop, Jr.

July 13, 1970

Paleontological Research Institution

Ithaca, New York, U.S.A., 14850

+
.

Library of Congress Card Number

a el

? e§ ees Ore:

CONTENTS

Page

PA Shr act mente. Ra Pete oe ene eee ceetec es Sohne opie CP eee ees oer ee, eee cao 281
Acknowledgments <..--..----.-2--::----c1ecsre-c-ccececceccecesceeeeeseeerentecennenengnscecnsenecsenenatsnaneestorecenstee 281
Ten GCS LNT CL Co Tere oS eae cee Pecans isco lane dnp csc nb cas=nnnantewonpescnreeeseny es 281
Difficulty and value of trace fossil studies ........... Pee Oe
Stratigraphy of the Cincinnati section 2.2.2.2... erect 284

I. History of invertebrate palichnology ~......-..-.------------------ cesses 286
A. Age of “£UCOIdS” ........:.-2-2cecnecccenccccecccceeteeeeeecueenenennerseneeseecenecnenensensenssneensens 286

eee eCLO GMO tee AGE ORNs c sree cree ore 5 ease oy eet epee eee ee 287

C. Development of ethological and paleoecological approaches) === 289

IJ. Preservation of trace fossils ........-...-..-.-.-:----2-csseeceecesceceeceeseseesesseeeeneneereeneeneneenes 291
A. Types of preservation —......2.2:-:-:-2-2e----: PB hams «cp sey eaeeeee toate epee 291

B. Mechanics of preservation of boundary relief traces... 292

III. Nomenclature of trace fossils .........-........2.2...----0:ec:c-ceseececeseeeceeeeeeeeeeeeeeeceeeereees 295
A. Classification at the generic and specific levels —.-...-..-.------------ 295

B. Suprageneric classification of trace fossils 2.2.22. 297

IV. History of the study of Cincinnatian “fucoids” and trace LOSSIS ee eee

Trace fossils and “fucoids” reported from the Cincinnatian —......300

Baia OS Ua End CELL Teale ate wc Sacco nad ae acnk eerie aashnhemmawagaoersenatcecneean 301
SOVATMMTED COTTE a eae ee rae eee ee ene ace ese eect tee ole eer meres ne nrseranc-nneereascce 314

Introduction to Spreite-bearing U-tubes —......--... Be tl Sree ene 314
WAT, “Roimitchnta 2 occeeqcecc en cn cnc ae cet ne cee anes tcetececen-saneennerecceneoccegsanensmnncoramcnnenenascentenscnnseannens 328
WITT. Repich ria ......--2cecce-ececsscsceseeneccecseceeeneeeseenenseeecneneceneaneecenencesscnentnanatenseenassecsss 351

General introduction to arthropod tracks ~.........---------.-.--:c:eeeseeeeeeeee 351

Synopsis of the ventral morphology of Cincinnatian trilobites ............ 354
UBC pe EDU CLC Lapierre see eae eee ee cence erence ee nree ace ote anaes cennmenn ha oe eeseche st eneeeelersees 384
X. Body fossils described as plamts -...-.- ---.-.--.--.--.---eee-ccnseseee cee eee cee eecetccntnetnnnes 387
XI. “Fucoids” of imorgamic origin ~..........-------------------cecesceeeeec eens ecec eee ceeeee escent 389
SNATCH GSe CUS ocean oe secs cease wate enn nat nce neers repeat bat smseanapchreananéaennanseaee 394
KIDD. Invalid species ....---.-.-.-2-.cecececescecee-seeeccrecerseecseneeessessneneas innnenenaqeneeneccacsesseceasenens 400
XIV. Comparative palichnology and suggestions for further study —......... 400

Comparative palichnology —...-.-.-----------.2:--..c:c:ecsee creer eeeee sence en reeetteeeceees 400

Comments on Cincinnatian paleoecology and suggestions for
additional studies

Bibliography ....---.-cseeccec--e-ncenecceenecucensenecceesscesssssseenesuecnaneecnenereeceactsnescensnanensennessscs
nid exqe sss: Proce Bea ane rectrea ree chee Pe eon ee 411

SMITHSONIAN

INSTITUTION

TRACE FOSSILS OF THE CINCINNATI AREA

Ricuarp G. Oscoop, JR.
College of Wooster, Wooster, Ohio

ABSTRACT

All previously described ‘“fucoids” (fossil seaweeds) from the
North American standard section for the Upper Ordovician are dis-
cussed and evaluated. A restudy of type materials reveals that ap-
proximately one-fourth of these are inorganic sedimentary markings,
while the remainder are trace fossils (tracks, trails, burrows). Ex-
tensive treatment is given to the early history of trace fossil studies,
generic and suprageneric classification, and preservation of trace
fossils. Thirty genera and 44 species of Cincinnatian traces are
illustrated and interpreted. These are grouped in the five categories
proposed by Seilacher (1953b). New genera proposed include A/loco-
tichnus, the trail of a large arthropod; Fascifodina, a bundled feed-
ing burrow; and Tylichnus, a trail of unknown origin. In addition,
a new species of the trilobite burrow Rusophycus is established under
the name R. cryptolithi. It represents the first known burrow of a
trinucleid. A comparison of the entire Cincinnatian trace fossil com-
munity with assemblages from different ages and locations reveals a
correlation between the types of trace fossils and the general environ-
mental setting.

ACKNOWLEDGMENTS

The author extends his sincere thanks to the faculty
and staff of the Department of Geology of the University
of Cincinnati under whose direction the present manuscript
was prepared in partial fulfillment of the degree of Doctor
of Philosophy. Especial thanks go to Kenneth E. Caster
under whose direction this manuscript was prepared. He
gave generously of his time and was a constant source of
inspiration, Thanks are also due Hans Hofmann for many
helpful discussions on Cincinnatian trace fossils and stra-
tigraphy and Warren Huff for X-ray analysis and advice on
the interpretation of X-ray data employed herein.

I am grateful to the following for loan of study ma-
terials: H. B. Whittington and B. H. Kummel, Museum of
Comparative Zoology, Harvard University; E. 8. Barghoorn,
Harvard Botanical Museum; G. A. Cooper and R. S. Board-
man, United States National Museum; R. L. Batten, Ameri-
can Museum of Natural History; C. F. Kilfoyle, New York
State Museum; J. K. Pope and R. Rinehart, Miami Univer-
sity, Oxford, Ohio; W. D. I. Rolfe, Hunterian Museum,
Glasgow; Mr. Zapfe, Natural History Museum of Vienna;
F. Westphal, Tubingen University; R. H. Flower, New
Mexico Bureau of Mines and Mineral Resources, Socorro,
New Mexico, and Eugene Richardson of the Field Museum,
Chicago, Illinois.

The writer is deeply indebted to Adolf Seilacher of
Tiibingen University and Walter Hantzschel of the Geo-
logische Staatsinstitut, Hamburg. Dr. Seilacher made
several trips to Cincinnati to discuss the material under
consideration and was most helpful in answering numerous
questions. Dr. Hantzschel repeatedly provided information
on obscure European forms and placed his then unpublished
manuscript of Fossiliwm Catalogus (1965) at the writer’s
disposal.

Thanks are due Roland Goldring of Reading Univer-
sity, Reading, England, and Scott Simpson of Essex Uni-
versity, Essex, England, for helpful correspondence, Mrs.
Elizabeth Dalvé and Mrs. Julie Multer who so ably
draughted many of the illustrations in this text. Heartfelt
thanks also go to my wife, Joanne, for the thankless task
of typing and proof-reading the entire manuscript more
than once.

The cost of preparing the manuscript was supported in
part by grants from the Nevin M. Fenneman Fund and
the Walter H. Bucher Fund of the University of Cincinnati
and the William H. Wilson Research Fund of the College
of Wooster. The author is extremely grateful to the Grad-
uate School of the University of Cincinnati for defraying
the cost of the plates. The author acknowledges the support
of the National Science Foundation through two Summer
Fellowships for Graduate Teaching Assistants and a Co-
operative Graduate Fellowship.

INTRODUCTION

This study has a two-fold purpose; first to discuss all
of the “fucoids,” or supposed marine algae, reputed to be
present in the rocks of the Cincinnatian Series in the vicinity
of Cincinnati, Ohio. Aside from being noted for its excellent,
well-preserved invertebrate fossils, the Cincinnatian con-
tains an abundance of these problematics. Since 1873 some
37 species have been recognized. Most were described by
three local amateur paleontologists, Uriah P. James, Samuel
A. Miller, and Charles B. Dyer, over a period extending
from 1874-1885. A restudy of these forms was undertaken
by still another amateur, Joseph F. James (1884, 1885,
1886); however, he lacked the type specimens and was
forced to work almost entirely from the literature and his
own observations. Nevertheless, many of his interpretations
appear to have been correct. Fortunately most of the type
specimens have been located and were made available for
the present study. With a few exceptions Miller and Dyer’s
(1878) and Miller’s (1880) types were found in the Dyer
collection at the Museum of Comparative Zoology and the
Botanical Museum of Harvard University. Types of some
forms described by J. F. James, as well as topotype ma-
terial of U. P. James, were located in the University of
Cincinnati collections. The bulk of the James collection
is the property of the Field Museum of Natural History,
Chicago, Illinois, but a search of the collection failed to re-
veal any type material. In addition, the type specimens of
several species, originally described by James Hall (1847,
1852), from the Lower Paleozoic strata of New York State,
and reported by local authors to be present in the Cincin-

282 PALAEONTOGRAPHICA AMERICANA (VI, 41)

natian, were obtained from the New York State Museum
and the United States National Museum. None of these
“fucoids” is now believed to be marine algae. Most of them
are trace fossils and the remainder are of physical origin.

The second and most important part of this study deals
with the description, illustration, and interpretation of
Cincinnatian trace fossils. This includes all named forms, as
well as previously undescribed material.

Trace fossils include fossil tracks, trails, coprolites, and
burrows as opposed to body fossils which are the actual re-
mains, casts, or molds of organisms, Admittedly there is no
rigid separation of tracks and molds, hence tracks are one
aspect of body fossils. A trace fossil may be formally de-
fined as: evidence of the activity of an organism in or on
the sediment, produced by some voluntary action of that
organism. The term trace fossil was proposed by Simpson
(1957) and is synonymous, at least in part, with “hiero-
glyph” (Fuchs, 1895), Lebensspur (Abel, 1912), trace
d’activité (Lessertisseur, 1955), vestigiofossil (Moore,
1936), Bioglyph (Vassoevich, 1952), Ichnofossil (German-
Spurenfossil-Seilacher, 1953b), and Problematica (Caster,
1938, 1957, and others). The study of such structures may
be referred to as ichnology (Buckland, c. 1830), which is
subdivided into palichnology, the study of fossil traces and
neoichnology, the study of Recent forms. Other terms will
be defined as they are encountered in the text.

The material used in this study consists of the type
specimens (when obtainable) of all named forms, supple-
mented by additional material from the Dyer collection,
the collections of the University of Cincinnati and Miami
University, as well as specimens collected by the author.
Unfortunately, precise locality data for many specimens are
lacking, and in most instances it was not possible to recol-
lect from the type localities. The following abbreviations for
repositories are used herein:

American Museum of Natural History

New Yorks New Mork (ss eccccsn coe oererat eer Sageeceeeortede AMNH
Harvard University Botanical Museum

(Gamibrid pes IMiass aGhseets cece eee tive eee nae HBM
Museum of Comparative Zoology

Varvramdl a Wit Cn 8 1byie se cece os see ee eee ees eee MCZ
Miami University

Oxfond'7 (Ohio ates een. cere ens ae ete es Ss rene ee eee MU
Hunterian Museum

Glasrow— Scotlands 2.2.6... xcccctesseceesceetecac te caterers eee HM

University of Cincinnati Museum
Gineinnatt.yt Oli omer cece eee eee ee Ba eee rs UCM
Field Museum of Natural History

Chicago, Illinois ~............... Pc la CROAT Be a LOKe:
New York State Museum

Albany, New York .. NYSM
United States National Museum

Washington, D: Cy 2. USNM

The body of the paper contains a lengthy introductory
section, giving the history of the development of inverte-
brate palichnology, the nature of the preservation of trace
fossils, and the generic and suprageneric classification of
trace fossils. This extended treatment is justified, because
much of this material has never been synthesized, and
several of the more important works are in foreign languages,
chiefly German. The generic descriptions of trace fossils
(and other forms) have been arranged by employing a
combination of the ecological classification proposed by
Seilacher (1953a) (topic IIIB herein) and the format used
by Hantzschel (1962) in the Treatise on Invertebrate
Paleontology, Part W. The trace fossils are separated into
Seilacher’s five groups: Cubichnia (traces of repose),
Domichnia (dwelling traces), Fodinichnia (feeding traces),
Pascichnia (grazing traces), and Repichnia (crawling or
walking traces). It has also been necessary to consider other
kinds of “fucoids.” Some of these are of physical origin;
some are body fossils originally described as land plants.
Unfortunately, there must also be a section on /ncertae sedis
and unrecognizable genera. The final topic is devoted to a
comparison of the entire spectrum of Cincinnatian trace fos-
sils with suites from different ages in different parts of the
world. Throughout this survey there are abundant sug-
gestions for additional studies which are necessary before
many of these forms are clearly understood.

Because of the large number of forms discussed (some
59 species), the coverage is uneven and admittedly super-
ficial in spots. This applies especially to the larger “genera,”!
such as Rusophycus, Chondrites, and Paleodictyon. Like-
wise, synonymies for these larger genera are not complete.
Only those forms are included which have been studied or
of which the descriptions and figures are sufficiently clear
and accurate to permit a meaningful diagnosis. The reader
is referred to Hantzschel (1962, 1965) for extensive
synonymies and a comprehensive bibliography. Because all
synonymies are largely subjective, the only ones the author
can vouch for are his own.

DIFFICULTY AND VALUE OF TRACE FOSSIL STUDIES

Only recently have trace fossils been recognized as use-
ful tools in the paleoecological analysis of sediments. In
large part this has been due to the work of Rudolf Richter,
Hantzschel, Seilacher, Schafer, and the other German
workers at Senckenberg am Meer. Because the value of

‘There is some question as to whether trace fossil names constitute
valid zoological taxa. However, throughout this work the taxa are
treated as if they were valid, and the rules of zoological nomen-
clature (e.g. rule of priority) are applied. A further discussion of
this problem is given in topic III.

‘TRACE FOSSILS CINCINNATI AREA; OsGoop 283

trace fossil studies has been discussed at some length in
English by Seilacher (1964), it needs only to be dealt with
briefly.

As Seilacher (1964a) pointed out, trace fossils have
certain advantages over body fossils. First, they are virtually
always biocoenosic, 1.e., they occur “in place,” while body
fossils are most often thanatocoenosic. Second, trace fossils
are most frequently encountered in rock types (silts and
sands), where body fossils are not overly common and
often are poorly preserved. Third, whereas diagenesis tends
to destroy the detailed structure of body fossils, it appears
to enhance the visibility of trace fossils. Processes such as
pyritization and minor additions of iron oxides tend to
make many trace fossil structures (¢.g., U-tubes) stand out
more clearly than their Recent counterparts.

It has been successfully demonstrated that trace fossils
can add to our knowledge in diverse ways. Broadly speak-
ing, the formation gained can be subdivided into two cate-
gories: A) Biological-morphology and ethology of the pro-
ducer and B) Physical e.g.-water depth, amount of oxygen,
facies analysis.

A) Biological—Trace fossils are the sole evidence of
numerous soft-bodied organisms. While in most cases they
provide little insight into the detailed morphology of the
producer, some imprints (e.g., Walcottia rugosa Miller, PI.
69, fig. 5) are sufficiently clear to permit diagnosis. Arthro-
pod tracks are particularly valuable in revealing the nature
of the seldom preserved distal portions of the walking legs.
For example, the tracks of Asaphoidichnus trifidum Miller
(Pl. 71, fig. 6) indicate that the originator, which most
probably was Jsotelus, possessed trifid dactyls. Similar
examples are given by Seilacher (1962b).

The only direct evidence of the behavioral patterns
(i.e., ethology) of fossil life is furnished by traces. Seilacher
(1955), through a study of trilobite morphology and the
configuration of the trails, was able to hypothesize on the
mechanics of movement in this extinct group. Caster (1938)
did the same for fossil xiphosurans, with the advantage of
being able to observe the locomotion of the existing Limulus.
Unfortunately, in most instances it is impossible to deter-
mine the origin of even the well-documented patterns (¢.g.,
Phycodes Reinhard Richter, Helminthoida Schafhautl).

It is fundamentally important to separate trace fossils
of invertebrate origin from vertebrate tracks and from im-
prints of true marine algae (fucoids s. s.). Prior to the work
of Nathorst (1881), the great mass of “fucoids” gave an
erroneous impression of early plant evolution. Such false
evidence has not been restricted to the vegetable realm.
Willard (1935) established the “Order Icthyopoda” for a
supposedly primitive group of Devonian amphibians, based

solely on the tracks of his lone genus Paramphibius. Through
neoichnological studies Caster (1938) showed that Param-
phibius represents the trails of a xiphosuran similar to the
Recent Limulus. In a more positive vein Seilacher (1956)
employed trace fossils to support an argument for ac-
celerated evolution at the beginning of Cambrian time. He
found that the small number of Precambrian traces were
of relatively simple morphology, whereas the numerous
Cambrian forms were much more complex.

B) Physical—It is generally acknowledged that the
greatest potential value of fossil traces lies in their contribu-
tion to paleoecology. They have been used as indices of
various physical factors. Trace fossils can document the
rate of sedimentation and the presence of subaqueous ero-
sion in a way that most body fossils can not. R. Goldring
(1962) demonstrated that rapid sedimentation in the
Devonian Baggy Beds of Great Britain forced a U-tube
dweller (Diplocraterion yoyo) to elevate the level of its
burrow. In contrast, truncated burrows (see Lockeia and
Corophioides herein) are proof of subaqueous erosion. The
presence of trace fossils indicates a nontoxic, oxygenated en-
vironment. Thus Rudolf Richter (1931), through the dis-
covery of burrows (mainly Chondrites) in some beds of
the Devonian Hunsriick Shale of the Rhineland, demon-
strated the presence of oxygen in one of the classic black
shale sequences. Seilacher (1964b) believed that certain
traces can be used as depth indicators. Although his argu-
ments are convincing, the hypothesis requires confirmation
by studies on Recent organisms.

Trace fossils have been utilized in facies analyses.
Seilacher (1964b) reported that the Carboniferous cyclo-
thems of the Ruhr Basin can be further subdivided on the
basis of traces. Weimer and Hoyt (1964) plotted Pleistocene
shorelines in Georgia using the burrows of the arthropod
Callianassa; and Caster (1938), using Paramphibws,
demonstrated a marine influence in the Devonian of the
Penn-York embayment in horizons where before only ter-
restrial conditions had been postulated.

In spite of the knowledge that can be gained from the
investigation of fossil traces, several difficulties are en-
countered in the course of such studies. Perhaps the greatest
problem is the inability at the present time to apply fully
the concept of Uniformitarianism to trace fossils, This is
due solely to our vast lack of knowledge on the traces of
Recent organisms. The sedimentologist is more interested
in the physical characters of the sediment and tends to ig-
nore the organisms, while the ecologist or zoologist ignores
the sediment and concentrates on the organism. The study
of traces lies midway between these two fields; and the
prospective worker must be well versed in sedimentology,

284 PALAEONTOGRAPHICA AMERICANA (VI, 41)

paleontology, zoology, and ecology. Also, the trace fossil
nomenclature has been, and still remains, in a state of con-
fusion; only recently Hantzschel (1962, 1965) attempted to
clarify the situation at the generic level. However, re-
studies of specimens, as well as the literature, are required
before Hantzschel’s work can be tested in full. In addition
to the lack of a stable classification, the absence of an
established descriptive terminology has led to many con-
flicting terms. It is hopeful to note that recent reattention
to traces has partly alleviated some of these problems.

STRATIGRAPHY OF THE CINCINNATION SECTION

The strata of the North American standard section for
the Upper Ordovician crop out on the axis and along the
flanks of the Cincinnati Arch in Ohio, Indiana, and Ken-
tucky. Basically, the section is composed of a more or less
monotonous interstratification of lutites, calcisiltites, and
calearenites, whose aggregate thickness is some 270 meters.”
The individual beds are thin (1 cm-2 meters), and the con-
tacts between various lithologies are normally sharp. Most
of the beds pinch and swell, and only rarely can they be
traced for more than a few hundred meters,

The blue-gray Cincinnatian lutites (“shales”) lack
prominent fissility and probably are better termed mud-
stones. According to Scotford (1965, p. 219), “the mean
size distribution is clay 38 per cent, silt 59 per cent and sand
3 per cent. The mean mineral composition is quartz 24 per
cent, calcite 6 per cent, dolomite 9 per cent, illite 47 per
cent, mixed-layer illite 4.4 per cent, chlorite 7.7 per cent
[and] mixed-layer chlorite 2 per cent.” Trace amounts of
feldspar, kaolinite, and pyrite are also present. Scotford
(1965, p. 219) also pointed out that the lutites “.
a persistent vertical and lateral textural, mineralogical, and
chemical uniformity.” However, his conclusions were based
on a relatively small number of samples. Neither body fos-
sils nor trace fossils are common in the lutites, although
brachiopods, trilobites, trilobite tracks, and irregular struc-
tureless burrows are sometimes encountered.

The calcisiltites (“siltstones”) are normally thin (1-10
cm) and frequently exhibit cross-lamination. They are
primarily composed of fine-grained fossil fragments, al-
though according to Weiss and Norman (1960) the in-
soluble residue content may be as high as 50 per cent. The
calcisiltites are well sorted and individual laminae stand
out in polished sections. Although well-preserved body fos-

. show

2 As used herein lutite refers to rocks having grains less than 1/256
mm in diameter, siltites to rocks having grains whose diameter is
between 1/256-1/16 mm, and arenites to rocks the grains of which
have a diameter in excess of 1/16 mm.

sils are not common, trace fossils and inorganic sole mark-
ings are abundant.

Cincinnatian calcarenites (“limestones”) are composed
dominantly of whole fossils or fossil fragments. According
to Weiss and Norman (1960) they make up some 70 per
cent of the volume of the rock, The fine-grained calcare-
nites may show cross-laminations; sorting is good. As with
the calcisiltites, trace fossils are common, and the upper sur-
face may show oscillation or interference ripple marks. The
coarse-grained calcarenites, commonly used for building
stone, are composed almost entirely of fossils and fossil frag-
ments. Sorting and stratification generally are poor, al-
though shells may exhibit a parallel arrangement. Many of
these beds are marked by large pararipples (Bucher, 1919).
Owing to the grain size, any trace fossils found lack detail
and are difficult to diagnose. Weiss and Norman (1960)
studied the Cincinnatian calcisiltites and calcarenites in
some detail, and Scotford analyzed the mineralogical com-
position of Cincinnatian lutites. Recently Weiss, et al.
(1965) gave an excellent analysis of the lithology of the
Eden rocks. The reader is referred to the above works for
more detailed lithologic descriptions.

Most of the original work on Cincinnatian stratigraphy
was done by Orton (1873), Nickles (1902, 1903), Foerste
(1903), and Cumings (1908). Because of the apparent dif-
ficulties involved in delimiting homogeneous lithologic units,
most boundaries were drawn primarily on the basis of fos-
sils. According to the North American Code of Stratigraphic
Nomenclature (1961), such designations do not constitute
valid rock-stratigraphic units. For this reason several in-
dividual workers, as well as the federal and state geological
surveys, are currently in the process of re-evaluating the
Cincinnation section.

Weiss and Sweet (1964) proposed the name Kope
Formation for rocks formerly included in the Eden or
Latonia “Group” (from Kope Hollow, Levanna, Brown
County, Ohio). According to Ford (1967) and Dr. Hof-
mann of Cincinnati (1964, personal communication), the
name Kope can be applied to Eden rocks in the Cincinnati
area. However, the upper 2.5 meters of the traditional
Eden “Formation,’ as seen at the reference section on
Clifton Avenue in Cincinnati, are now included in the re-
defined Fairview Formation. Neither Ford nor Hofmann
recognized the traditional members of the Eden and Mays-
ville “groups” as being valid rock-stratigraphic units, Like-
wise, Weiss, et al. (1965) rejected the “members” of the
Eden.

The Richmond “Group” of southeastern Indiana was
restudied by Fox (1962). Employing mainly lithologic cri-

TRACE FOSSILS CINCINNATI AREA: Oscoop

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Text-figure 1.—Columnar section of the
Upper Ordovician strata exposed near Cin-
cinnati, Ohio, This section portrays the “tra-
ditional” nomenclature applied in the Cin-
cinnati area. It is acknowledged that the
members are not recognized as valid litho-
stratigraphic by the most recent North
American Code of Stratigraphic Nomencla-
ture. (From Caster, Dalvé, and Pope, 1955,
fig. 3.)

286 PALAEONTOGRAPHICA AMERICANA (YI, 41)

teria, he established the Tanner’s Creek Formation to em-
brace the Arnheim “Formation” of Foerste (1903) and the
Waynesville and Liberty “Formations” of Nickles (1903).
He did not accept the traditional members of the three
fomations as rock-stratigraphic units.

While recognizing its contemporary inadequacies, the
author has chosen to employ the traditional section as given
by Caster, Dalvé, and Pope (1955), (see Text-fig. 1). This
course was followed for two reasons. First, many of the
specimens included in this study bear labels of the tradi-
tional formations and members, and it would be both awk-
ward and confusing to fit them into the new terminology
at this time. Second, and more important, the nomenclature
of the Cincinnatian is currently in a state of flux, Until the
works of the various survey groups have been published
and evaluated it would be premature to employ their
terminology. Nevertheless, it is acknowledged that the rock-
stratigraphic units used herein are not valid according to
the most recent Stratigraphic Code.

I. HISTORY OF INVERTEBRATE PALICHNOLOGY

The history of invertebrate palichnology has never
been adequately covered. This is especially true of the con-
troversy which raged during the 1880’s over the nature of
“fucoids.” No attempt will be made to present a complete
survey, as this would require a volume in itself.

For convenience this account is divided into three
parts. It is acknowledged that the divisional boundaries
are somewhat arbitrary, because it is difficult to subdivide
the continuium of history into artifactual bundles. The first
period extends from 1828 to 1881 and covers the “Age of
Fucoids;” it was during this period that dozens of genera
and hundreds of species of fossils, supposedly marine algae
(“fucoids”), were described. The second period, the “Period
of Reaction,” extends from 1881 to the 1920's; 1881 was the
year of the classic work of Alfred Nathorst, who was the
first to question the “fucoid” myth. Although establishment
of “fucoid” genera and species continued at a rapid rate,
this 40-year interval marks the time when several authors
of varying backgrounds began to question the algal nature
of these forms. The 1920’s saw the birth of invertebrate
palichnology. It was during this decade that Rudolf Richter
initiated his important studies at Senckenberg am Meer.
This final period, spanning the last 45 years, is entitled
“The Development of the Ethological and Paleoecological
Approaches.”

A) THE AGE OF “FUCOIDS”
Perhaps as Maillard (1887) said, the “Age of Fucoids”
began in 1828 when Brongniart erected the genus Fucoides

and placed it within the algae. As the years passed, this
genus became the receptacle for many species until it
rivaled “Orthoceras” in its nomenclatoral complexity. Cer-
tainly one of the highpoints of the period was the work of
James Hall on the Paleozoic section of New York State. He
proposed several new genera (e.g., Palaeophycus, Rusophy-
cus) which later authors (e.g., Billings, Miller) extended to
other parts of North America. However, it was in Europe
where most of the work was done by Schimper, Heer,
Saporta, and Delgado. The results of these 50 years of labor
can best be seen in Schimper in Zittel (1889), where the
“Algae Incertae sedis’ cover 25 pages and are subdivided
into 16 groups. The brief discussions of the different genera
are accompanied by elaborate reconstructions which il-
lustrate the supposed resemblance to modern seaweeds. It
should be pointed out that there were some contemporary
authors who attacked the overzealous taxonomy of the
“paleobotanists.” For example, Dawson (1865) correctly
exposed Rusophycus, one of the most sacred of all “fucoids,”
as a product of the burrowing activity of trilobites; however,
his work was largely ignored until years later.

Several factors seem to have contributed to the pro-
liferation of “fucoids” in the latter part of the 19th Century,
but none more than the spirit of the times which was over-
whelmingly descriptive. The broad outlines of the taxonomy
of hard-part fossils (mollusks, brachiopods, trilobites)
were already described, and the problematic fringe still re-
mained for naming. Both professionals and_ publishing
amateurs (especially of the “Cincinnati school”) zealously
engaged in wholesale description of these supposed seaweed
traces. The orientation of most fossil-students (mainly
geologists) was nonbiologic (even largely nonevolutionary,
e.g., James Hall, Joaquim Barrande, and Louis Agassiz) and
nonecologic. Paleontology was a study of thanatocoenoses,
and paleobiology was yet a generation in the future. More-
over, sedimentology and allied aspects of stratigraphic
geology were in their infancy. Thus many purely physical
phenomena of sedimentary rocks (e.g., rill marks, ripple
marks, as well as dendrites), which create “pseudofossils,”
were misinterpreted, and the most obvious interpretation
became first choice—with a binomial name to tie it down!
In the later phase of the “fucoid” period, the impact of
Darwin’s evolutionary works may have served as a stimu-
lus; some of the “fucoids” seemed to be links in the evolu-
tionary chain. By the 1880's there was a great body of
“fucoid” species arranged with varying confidence into
higher taxa. Only today are we beginning to emerge from the
nomenclatoral snarl which resulted from this misdirected
energy.

However, not all forms, which are recognized today as

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 287

trace fossils, were once described as “fucoids.”’ One of the
most enigmatic of all trace fossils, Climactichnites, from the
Upper Cambrian Potsdam Sandstone of New York State
and Quebec, was described by Logan in 1860 and recognized
as non-algal. Likewise, Owen established Protichnites in
1852 for trilobite tracks occurring in the Potsdam. Other
works, such as Miller’s (1880) study on “Silurian Ichnolites”
of the Cincinnatian, can be mentioned as examples of early
trace fossil studies, but these works were in the minority.

B) THE PERIOD OF REACTION

The first comprehensive attack against “fucoids,” and
in the final analysis the most devastating, was delivered
by the Swedish paleobotanist, A. G. Nathorst, in 1881. His
long article, published first in Swedish and then somewhat
abridged and translated into French, was accompanied by
several plates based on neoichnological studies on the coast
of Sweden. He made plaster-of-Paris molds of Recent traces
which were illustrated in the plates and by line drawings.
Therefrom he proceeded systematically to demolish many
“fucoids.”

Nathorst pointed out that “fucoids” almost always are
found on the surface of beds, usually on the under surface
of a layer, but never in their interior, where genuine plant
remains commonly occur. Certainly for the kinds of
“fucoids” with which Nathorst was concerned this observa-
tion was valid. He also found it puzzling that “fucoids,” in
contrast to genuine plant fossils, never show carbonaceous
or other normal organic substances.

Two of the most famous ard controversial “fucoids”
are the traces known as Cruziana (Seilacher, 1959, fig. 5D)
and Rusophycus (Pl. 58). Nathorst chose them as a special
focus of his reanalysis. He noted that both forms occur as
casts on the under surface of a stratum or as depressions
originally made in the subjacent bed. These original depres-
sions may be several centimeters deep, and he questioned
the mechanism by which such a small object sinking to the
bottom could impress itself so deeply in the underlying mud.
He said (1881, p. 62) that this “. . . would necessitate a
weight and consistency . . . which is found in no present-day
plants.”” Nathorst also mentioned that Cruziana is never
composed of a substance which differs from the host rock.
Likewise, when two cruzianids come in contact, one eradi-
cates the other. If they represented branches of plants, this
certainly would not be the case.

Following Nathorst’s challenge there shortly appeared
several attempts at rebuttal, especially centering on the
proper assignments of Cruziana. Lebesconte (1883) in an
appendix to Oeuvres Posthumes de Marie Rouault at-
tempted to re-establish Cruziana as an alga. The Marquis

de Saporta (1884) presented a 100-page attempt at free-
wheeling rebuttal in his monograph “Les Organismes Prob-
lematiques des Anciennes Mers.”’ Lebesconte (1886) re-
newed his argument, and in the same year Delgado, work-
ing with the beautifully preserved cruzianids of the Silurian
of Portugal, argued for the plant nature of Cruztana. In
1886 Nathorst answered all three authors in a monograph
in which he described how he was able to reproduce many
of the ‘“fucoids” by mechanical means. In 1886 and 1887
Saporta published two long articles, mostly concerned with
Cruziana and conceding nothing. Thus the years 1881-1887
marked the period of the great controversy.

Because most of the arguments employed by the
“algal” proponents (Saporta, Lebesconte, Delgado) are the
same, little would be gained by discussing each of the papers
in turn; rather the major points of the controversy are
synthesized below. While this is concerned mainly with
Cruziana, many of the arguments hold for other “fucoids.”

1) Nathorst (1881) claimed that “fucoids” were almost
invariably preserved only on the soles of beds. Lebesconte
(1883, 1886) remarked that he possessed specimens which
gave the appearance of arching up into the host rock.
Nathorst replied that such examples could be produced by
trilobites burrowing along the sand-mud interface. Likewise,
a false impression of postdepositional origin could be given
by a trilobite which changed from crawling to swimming.

2) Saporta (1881, 1882, 1884) recognized that most
“fucoids” are preserved as casts on the bottom surface of
sandstones or siltstones which overlie shales. To explain this
unusual mode of preservation, he postulated a special pro-
cess of fossilization entitled “‘fossilization in demirelief.” In
essense, Saporta’s theory holds that a plant falls to the bot-
tom and when covered by overlying sand is driven down
into the mud. Decay is initiated on the upper surface of the
plant and proceeds downward with the sand filling in be-
hind it. The final portion to decompose is that in contact
with the mud; thus only the lower surface of the plant is
faithfully preserved.

Nathorst (1886) attacked this hypothesis on several
fronts. Why, he reasoned, should decomposition always be-
gin on the upper surface and proceed downward? It is not
more natural to assume that those organs most sensitive to
decay and most accessible to bacteria would be the first to
undergo putrifaction? As mentioned above, he also ques-
tioned the source of the great amount of pressure required to
force the object in some cases several centimeters into the
underlying mud. Because Cruziana is always convex down-
ward, there must have been a pressure differential. Why
was the greatest amount of pressure always exerted on the

bo
S
co

central portion? A highly damaging point of Nathorst’s
argument is that if this process works for supposedly soft-
bodied plants, why does it not operate for the preservation
of soft-bodied animals as well? Why are not similarly orig-
inating casts of holothuroids and worms found in the geo-
logic record? Nathorst rejected Saporta’s explanation of the
process of fossilization of “fucoids” and maintained that
“fossilization in demi-relief”’ occurred only on rare occasions.

3) Just as Lebesconte claimed to have forms which were
found within the interior of the host rock, Saporta asserted
that he possessed specimens which occurred on the upper
surface as well as on the lower. Nathorst (1886) referred
to these as “false cruzianids” and explained them as bur-
rows which were formed slightly below the surface and
caused a bulging up of the overlying send.

4) The presence or absence of organic matter in
“fucoids” was long a subject of dispute. Although this argu-
ment was more critical in other genera (e.g., Chondrites),
it also found its way into the Rusophycus controversy.
Whereas Nathorst (1881) maintained that Rusophycus and
Cruziana contain not the slightest vestige of organic struc-
tures or organic material, both Saporta (1884) and Lebes-
conte (1883, 1886) believed that the subparallel striae on
the lobes of Cruziana represented a network of fibrous ma-
terial of plant origin. Moreover, Lebesconte (1886) claimed
to have seen a meshwork of unknown composition and
origin lying below the striae. This so impressed him that he
altered his opinion and considered Rusophycus and Cruziana
to be sponges rather than plants. Nathorst rejected Lebes-
conte’s network and declared that it represented the sagging
of the bedding within the Cruziana cast (see Nathorst,
1886, fig. 23).

5) Lebesconte (1883) stated that some bilobed traces
gradually converted into unilobed forms, Nathorst (1886,
fig. 12) explained that this could be due to a simple lateral
tilting of the trilobite’s body.

6) Not infrequently cruzianids appear to branch.
Whereas the algal adherents believed this to be a case of
simple vegetable branching, Nathorst maintained that it
was truncation. A study of Delgado’s (1910) illustrations
of the Portuguese cruzianids confirms Nathorst’s explana-
tion.

7) Many of Nathorst’s cruzianid examples were from
the Cambrian beds of Lugnas, Sweden. Here cruzianids are
found throughout the section, while trilobites occur only in
the upper part. Lebesconte (1886) was quick to point this
out, although Nathorst had offered an explanation in 1881,
remarking that trilobites were active throughout the entire
section but did not possess hard exoskeletons until they

PALAEONTOGRAPHICA AMERICANA (VI, 41)

actually appear preserved as fossils. Delgado (1886) went
one step farther and argued that trilobites were never found
in the beds containing Cruziana but, instead, were found
in the immediate adjacent beds. Nathorst (1886) replied
that the lithology of the Cruziana beds did not provide
optimum conditions for trilobite preservation. More recent
work certainly upholds Nathorst’s view; one of the main
values of trace fossils is that they commonly occur where
body fossils do not.

8) Saporta (1884) argued that Cruztana is frequently
preserved as short, disconnected fragments which appear to
represent parts of plants broken up and scattered about by
storms. Nathorst (1886) explained that such forms were
created by trilobites which settled down, crawled along the
bottom for a short distance, and then proceeded to swim,
thus losing contact with the substrate.

9) A common “argument” offered by many “fucoid”
workers expresses their skepticism as to how such surface
traces could possibly have been preserved. This was one of
the major reasons for their reluctance even to consider an
animal origin for the many “fucoids.” The problem is still
with us and is considered at greater length below.

Nathorst was fighting the entire corps of “fucoid”
adherents pretty much alone. The arguments became ex-
tremely heated, and at one point Albert Gaudry (1883)
in essence accused Nathorst of not believing in Darwinian
evolution, as the latter was unwilling to accept “‘fucoids”
as the documentation of algal life which assuredly must have
been present from very early times.

In answering Gaudry, Nathorst (1883) delivered a
stinging reply to Saporta (1882), who had accused Nathorst
of discarding all of the forms placed by Schimper (1879) in
the algae. Nathorst retorted that he too was a strong ad-
herent of the Darwinian theory and believed that there
were definite examples of true marine algae in the fossil
record. He was attacking only those forms placed by
Schimper (1879) in the “Algae Jncertae sedis.” He could
not understand why Saporta did not realize this, and he
mused, “On pourrait presque croire que c’est seulement par
suite de son ignorance de la langue suédoise” (Nathorst,
1883, p. 453).

Despite these concentrated attacks, other workers be-
gan to accept Nathorst’s arguments.* J. F. James (1884,

* Later work on Rusophycus and Cruziana have vindicated Nathorst’s
views on these two forms. Numerous examples (many from the Cin-
cinnation) show the impressions of the genal spines and pleurae. In
Cincinnati alone three specimens are known which show the trilobite
in place, thus “caught in the act” of forming the Rusophycus burrow.
These discoveries, now not uncommon from many ages and areas,
are “proof positive” of the animal origin of this long-debated trace
fossil.

TRACE FOSSILS CINCINNATI AREA: OsGoopD 289

1885), working independently and in ignorance of Nathorst
and certainly much less ambitiously, used many of the same
criteria as Nathorst had to criticize Miller and Dyer’s
(1878) Cincinnatian “fucoids.” As Hantzschel (1962) re-
marked, James’ work has long been ignored by more recent
authors on the subject. Likewise, Maillard in 1887 agreed
with Nathorst’s conclusions on the majority of “fucoids”
but still considered the chondritids to be “true fucoids.” In
1895 Fuchs published a lengthy treatment of “fucoids
which is second in importance only to Nathorst’s works of
this period. This was the first major paper in German at-
tacking “fucoids.” Fuch’s covered several genera and largely
echoed Nathorst’s beliefs.

By 1900 the arguments died down, and most “‘fucoids”
were generally accepted as being inorganic or traces of ani-
mal activity. Understandably few new forms were named
during the next two or three decades, and the general disil-
lusionment with respect to “fucoids” militated against their
study or discussion.

C) THE DEVELOPMENT OF THE ETHOLOGICAL AND
PALEOECOLOGICAL APPROACHES

By 1900 the “fucoid” ranks had been badly depleted,
and most of the organic forms were recognized as trace
fossils. However, the realization that they might be useful,
meaningful fossils was slow to gain acceptance; no doubt
the nomenclatoral problems discouraged many. Moreover,
paleontology was still in the descriptive stage and these
“works” of organisms lent themselves less conveniently to
morphology-based analysis. Nathorst had stressed the Uni-
formitarian principles in his 1881 paper. He demonstrated
that Recent traces, both physical and organic, are the key
to most “fucoid” forms; but it was not until the 1920's
that Rudolf Richter broadened and refined Nathorst’s tech-
niques. Actually Nathorst’s objective was not so much the
interpretation of fossil forms in the light of present pheno-
mena as an endeavor to prove the nonalgal nature of
“fucoids.” Thus there was a slightly greater “destructive”
element in his studies than a “constructive” one. In 1925
Richter and his associates established the Forschungsanstalt
fir Meeresgeologie und Meerespalaontologie der Sencken-
bergischen Naturforschenden Gesellschaft zu Frankfurt am
Main, better known as Senckenberg am Meer. The purpose
was to exploit the vast Wattenmeer mudflats of the North
Sea for what they might teach with respect to geologic
phenomena. This was the first careful systematic study of
Recent ecology, with the principal emphasis on invertebrate
traces. Much of Richter’s work is contained in three articles
which were issued in sections over a period of years: Flach-
seebeobachtungen zur Palaontologie wnd Geologie (1922-

1926), Marken und Spuren aus allen Zeiten (1937-1939),
and Marken und Spuren im Hunsriickschiefer (1931-
1941). The last stands as the single most important contri-
bution of trace fossil study to the general field of paleoecol-
ogy. The black Lower Devonian Hunsriick Shale of the
Rhineland has long been noted for its beautifully preserved
echinoderms and other organisms of great delicacy. It had
long been assumed that the Hunsriick represented a classic
euxinic environment and that all the fossils had been washed
in. The bottom of the Hunsriick basin itself was thought to
have been barren of life. Through careful studies, Richter
(1931, 1935, 1936, 1941) was able to show a well-developed
suite of trace fossils. The most significant form was Chond-
rites, whose branching pattern extended down several centi-
meters into the substrate. This genus had survived longer
than any other “fucoid.” Richter (1927) was able to demon-
strate that it is the mining record of a feeding animal, and,
as Simpson (1957) said, its correct interpretation was the
final death knell of the “fucoids.” The discovery of Chond-
rites and other trace fossils in the Hunsriick proved the
presence of an amount of oxygen sufficient to sustain life
at the depositional interface, and Richter was able to show
that reduction took place a few centimeters below the sur-
face. This of course means that the pyritized fauna was
more biocoenosic than had been realized.

Richter did not confine his labors to general paleoeco-
logic studies, for he contributed much to the understanding
of individual genera. His studies (1924, 1926) of “Spreite-
bau” are the most comprehensive treatment of these forms.
Likewise, he was able to demonstrate that the regularly
meandering traces in the European Flysch were Weide-
spuren or browsing traces of benthonic vermiform organ-
isms. Richter’s work stimulated others (e.g., Schindewolf,
Hantzschel); and Senckenberg am Meer became, and still
remains, the headquarters for research in fossil and Recent
traces. (See, for example, Schafer’s (1962) Aktuo-Palaon-
tologie which is the most detailed study of the traces of a
Recent environment to date.)

In 1935 Othenio Abel summarized the scattered litera-
ture on trace fossils, together with many astute observations
of his own, in his Vorzeitliche Lebensspuren. Until Hantz-
schel’s recent (1962, 1965) work on The Treatise on In-
vertebrate Paleontology, this was the standard reference
for all trace fossil studies.

By 1950 a large number of trace fossils had been re-
described and redefined by a variety of workers. Many
authors were utilizing the terms defined by Krejci-Graf
(1932), and therefore there was introduced a measure of
stability and universality. However, the study of trace fos-
sils was still considered largely as an avocational pursuit.

(Cubichnig)

FOSSILS

(Djuysisand)

D
c
Zz
z
z
roy
4
Ps)
b
°
m

PALAEONTOGRAPHICA AMERICANA (VI, 41)

Text-figure 2.—Ethologic classification of organic
traces proposed by Seilacher (1953). The outside circle
contains the five major ethologic catagories (Cubichnia,
Pascichnia) which are based primarily on the type of be-
havior exhibited by the organism. This behavior is in
turn reflected in the geometry of the trace. However, in
practice it is difficult to place some traces in this classi-
fication. (See the discussion of Palacophycus). The term
“Body Fossils” in the center of the diagram demonstrates
that trace fossils can not be separated from molds, casts,
and other hard part remains of organisms. (Translated
from Seilacher, 1953, fig. 6.)

Text-figure 3.—Ethologic classification of organic
traces proposed by Miller (1962). The classification is
basically similar to that of Seilacher’s but is more com-
prehensive. Like Seilacher’s this classification can be
applied to both fossil and Recent traces. (Translated from
Miller, 1962, pl. 21.)

TRACE FOSSILS CINCINNATI AREA: Oscoop 291

There was too little in the way of a unifying force, and
the nomenclatoral problem was a pressing one. In the last
15 years two workers, Walter Hintzschel and Adolf Seil-
acher, have made great contributions to unity and nomen-
clature. The former has almost single-handedly put the
nomenclature in a workable condition at the generic level
in his publication in Part W, Miscellanea, of the Treatise on
Invertebrate Paleontology. Even more significant is his
volume of Fossilium Catalogus (1965) which attempts com-
plete indices of each ichnogenus and contains the most
comprehensive bibliography to date.

Through numerous publications Seilacher contributed
greatly to the interpretation of trace fossils and demonstrated
their broad geologic value. His works established a pattern
for future analyses, In 1953 he initiated his Studien zur
Palichnologie and proposed an ecological classification for
trace fossils which has found acceptance among many
workers. He subdivided traces into five groups: grazing,
resting, dwelling, movement, and feeding.t Although supra-
generic classifications had been proposed before, Seilacher’s
proved to be the most workable. In the same year he pub-
lished Part II of the same study which contained an analysis
of Cubichnia or resting traces. The remainder of the series
has yet to be published, but when complete it will presum-
ably consist of a comprehensive analysis of all five ecologic
groups. Seilacher (1955) was the first to systematically
study trilobite tracks. His work on the redlichiid tracks of
the Pakistan Salt Range serves as a strong foundation for
others to build on. What may prove to be his greatest con-
tribution has yet to be fully realized. In 1955 he began a
study of comparative palichnology through an examination
of the trace fossil suites of the Lower Cambrian of the Salt
Range, the Lower Paleozoic of Portugal, the Schilf Sand-
stone of Wiirttemberg, the Lias and Dogger of Wiirttem-
berg, the Miocene marine Molasse of Europe, and the Creta-
ceous and Tertiary Alpine Flysch. The results of the com-
parison indicate that broad ecologic correlations are in-
volved. Seilacher found that grazing traces (Pascichnia)
dominate the Lower Paleozoic of Portugal and the Euro-
pean Flysch, while resting traces (Cubichnia) are almost
totally lacking. The reverse is true for the other three locali-
ties, where Cubichnia are numerous and the intricate grazing
traces virtually nonexistant. Seilacher concluded that the
suite dominated by Cubichnia is indicative of shallow, well-
lighted water, whereas the Pascichnia suites formed below
the euphotic zone. Since 1964 Seilacher has expanded this
study to include several more localities. Seilacher placed

"See topic III for a more detailed analysis of Seilacher’s classification,

invertebrate palichnology on a firm foundation by carrying
on the work of Rudolf Richter; he showed that the greatest
value of trace fossils is in paleoecological work. The mere
correlation of organism with trace fossil is but the first stage
in research of considerable scope.

The reader should not be left with the impression that
all recent work on trace fossils is of German origin, As
mentioned previously, Caster (1938) produced what is still
today the most detailed analysis of a single trace fossil genus
(Paramphibius Willard). This work was the first modern
study on arthropod tracks. Lessertisseur (1955) published
a comprehensive monograph covering both invertebrate and
vertebrate traces. Because of its scope it is understandably
superficial in places but is still the most extensive work in
the French language. It is hopeful to note that Ager’s
(1963) text on paleoecology and Approaches to Pale-
oecology (ed., Imbrie and Newell, 1964) contain chapters
on trace fossils.

Thus the study of trace fossils is gradually being
recognized as a valid and productive field of scientific in-
quiry. Yet in a sense it is still in its infancy, for Recent
traces have been virtually untouched. Such work as has
been done has centered mostly on tidal flats, for we have
little knowledge of the neritic, let alone the bathyl and
abyssal zones. However, with the current impetus in ocean-
ographic science, this void is about to be filled. Underwater
photography is proving to be a great boon, but what is
really required is a method of sampling that will allow for
study of nonsurface traces. Cores are helpful but provide
only a small sample. Several means of impregnating un-
consolidated materials have been devised, and these should
be utilized in trace fossil studies. The more that is learned
of Recent traces, the more meaningful becomes the in-
terpretation of fossil forms.

II. PRESERVATION OF TRACE FOSSILS
A) TYPES OF PRESERVATION

Terminology employed in the preservation of trace fos-
sils has been discussed by Seilacher (1953a, 1964a, 1964b )
and more recently by Martinsson (1965). Although Mar-
tinsson introduced what appears to be a workable term-
inology the author prefers to employ Seilacher’s terms with
some minor modifications. As shown by Seilacher (1964a,
fig. 1) and illustrated in Text-figure 4, there are two prin-
cipal varieties of preservation. The first of these is full re-
lief (Vollform of Seilacher, 1953a), where the entire struc-
ture is preserved. These arise when an animal burrows at
some depth below the surface through a homogeneous sedi-
ment. The burrow may then be packed with fecal material

292 PALAEONTOGRAPHICA AMERICANA (VI, 41)

or passively filled by overlying sediment or by sediment
sifting into the burrow. Generally, forms preserved in this
manner do not exhibit delicate morphological details, e.g.,
scratch marks of appendages. Two such examples of full
relief preservation are the U-tube Corophioides (Pl. 60,
figs. 5, 8), and Chondrites (Pl. 79, fig. 5).

The second and more important type of preservation
is semirelief (Halbform of Seilacher, 1953a; demi-relief of
Saporta, 1882). Records of this type are incomplete, only
the top half or bottom half being preserved. Semirelief pre-
servation is further subdivided into cleavage relief (Spalt-
relief of Seilacher, 1953a) and boundary relief (Grenzrelief
of Seilacher, 1953a).

In the first, Seilacher recognized the fact that the walk-
ing or crawling action is reflected in the bedding planes
beneath the surface—on the sedimentary cleavage planes—
of thinly laminated sediments. He also noted that these
subsurface records progressively deviate from the surface
semirelief expression with depth and can be quite mislead-
ing if misconstrued as surface records (Text-fig. 19).
Cleavage relief preservation is a common expression of
arthropod tracks. The great majority of Cincinnatian trilo-
bite tracks commonly encountered are of the cleavage re-
lief type, and both nomenclatoral and _ interpretational
confusion has resulted from failure to recognize the multi-
form subsurface expressions of the same track.

The most common trace expression is the form which
Seilacher called boundary relief. By this he referred to the
record on a stratum (usually clastic, e.g., silt, fine sand)
surface, either superior or inferior, and where the adjacent
stratum is of different lithology (e.g., lutite). These two
expressions of a trace are essentially like conventional molds
(superior surface imprints) and casts (inferior surface
fillings) of traces originally made on the surface of the sub-
jacent stratum. When they were preserved on the upper
surface of a stratum, Seilacher termed them epireliefs; when
on the bottom, hyporeliefs. These are purely descriptive
terms, Although the most common expression of epireliefs is
concave imprints and of hyporeliefs, convex fillings, either
may occur as bas-reliefs or intaglios on the stratum surface.
So far the discussion has dealt with exogene boundary re-
liefs originating on the depositional interface. However,
they may also originate as endogene reliefs made below the
depositional interface.

Seilacher’s usage (1953a, 1964a, 1964b) of the dynamic
terms endogene and exogene is somewhat confusing, be-
sause the reader forgets that Seilacher is employing endo-
gene only in reference to the track, not the organism. As-
sume, for example, that a 10 cm bed of mud is overlain by

2 mm of silt. According to Seilacher a trilobite walking on
the silt surface would exert enough force to penetrate into
the underlying mud. As the foot is withdrawn the thin silt
layer immediately fills in the depression, and the resulting
trilobite track is preserved as a convex hyporelief of endo-
genous origin. To avoid this ambiguity it seems best to
adopt the neoecologic terms epifawnal and infaunal [John-
son (1964), Petersen (1913), and Thorson (1957)].
Seilacher’s original meaning of endogene and exogene are
retained, and the two ecologic terms refer to the epistratal
or endostratal position of the organism while making the
track or trace. Epifaunal applies to an organism which was
moving on the surface, infaunal to an animal moving below
the surface. In the example of the trilobite track given
above, the track is of both endogenous and epifaunal
origin. It may appear at first that this is mere duplication
of terminology, but if it is remembered that one set of terms
refers to the organism and the other to the track, the dif-
ference becomes apparent.

Although it has not yet been seen in the Cincinnatian
rocks, an additional type of preservation, termed pseudo-
exogene, has been described by Seilacher (1962; 1964b,
p. 297) from the Eocene Flysch deposits of Spain. Here an
endogenous fecal-filled burrow has been exposed by erosion
so that the roof of the burrow and fecal filling were removed.
When the remainder of the burrow was filled and covered
by subsequent deposition, the resultant spoor resembles an
exogenous trace.

When the terminology discussed above is combined
with Seilacher’s ecological classification of trace fossils, it
is possible to give an accurate, concise definition of any
form.

B) MECHANICS OF PRESERVATION OF BOUNDARY
RELIEF TRACES

The mechanics of trace fossil preservation has long
troubled students in this field. As mentioned previously, it
was just this that prevented many early workers from even
considering “fucoids” as representing the work of animals.
The problem centers mainly around exogenous imprints;
why were they not destroyed in the normal processes of
sedimentation? Since endogenous traces are already covered
by sediment, they present less of a problem.

Both Kuenen (1957, p. 251) and Dzutinski and
Sanders (1962, pp. 84, 85) agreed that the condition of the
bottom is a most important factor in preserving either sedi-
mentary markings or traces of organic origin. Sand and silt
lack the cohesiveness of mud, and the grains move inde-
pendently, while mud acts as a unit. It is obvious that the
mud presented just the right degree of consistency for track-

TRACE FOSSILS CINCINNATI AREA: OsGoop 293

FULL RELIEF SEMIRELIEF
ORIGINAL CONCAVE CONVEX EPIRELIEF
FILLING CAVITY (TOP TRAIL)

HYPORELIEF

Meseles

H| STUFFED BY ANIMAL
FILLED BY SEDIMENTATION

PRIMARY CAST OF fa beans Raion SECONDARY CAST
SURFACE TRAIL FM!
Nee eaenee OF MUD BURROW

SAND SEDIMENTATION SAND FILLS BURROW SAND SEDIMENTATION

a0

at

EROSION

GROOVE IN MUD BURROW ALONG INTERFACE FECAL-STUFFED BURROW
IN MUD

Text-figure 4.— Various types of preservation of trace fossils. The dark stippling indicates mud; the light stippling sand. The most
common type of preservation encountered is convex hyporelief. Secondary casts are unknown in the Cincinnatian section. (From Seilacher, 1964,
fig. 1; reproduced with the permission of John Wiley and Sons, New York.)

294 PALAEONTOGRAPHICA AMERICANA (VI, 41)

making. Had it been too firm, faint records would be left;
if too soft, flowage would have obliterated the finer details.
The grain size of the overlying sediment is also critical. It
must be coarser than the substrate in order to present a
contrast in grain size and a sharp parting plane; yet if it
is too coarse, the fine detail of the trace may be lost. This
explains why the best preserved trace fossils generally occur
as convex hyporeliefs in sediment of silt size which is under-
lain by a lutite. This is the usual condition in the Cincin-
natian rocks where traces are abundant. These are generally
accepted facts; the great question is how can delicate im-
prints escape erosion from the currents and scouring in-
herent in the transportation of the supradjacent coarser
stratum?

Nathorst recognized this problem and attempted to
answer it in 1881 and 1886. While discussing Cruziana from
the Swedish Amoricain Sandstone, he (1881, p. 35) postu-
lated that these traces were formed “at the bottom of the
sea” (deep water?) and later covered by sand. Delgado
(1886) countered by pointing out that the Amoricain
Sandstone was a shallow water deposit. The currents, indi-
cated by the presence of ripple marks, should certainly have
eradicated such traces. In 1886 Nathorst presented a more
elaborate hypothesis, suggesting that the traces originated
in the intertidal zone. They were formed below water; and
then as the tide ebbed, the mud, by exposure to the air, was
to a certain degree indurated. The imprints were then
covered by both wind-blown sand and by the sand driven
forward by the rising tide. While this explanation may hold
for certain isolated examples of trace fossils, it is too
specialized to serve as a general method of preservation.

Little interest was exhibited in this problem until the
publication of Kuenen and Migliorini’s studies of turbidite
flows in 1950, Following this, many sedimentologists took
an interest in the origin and preservation of the numerous
primary sedimentary features and trace fossils found in
turbidite sequences. Dzulinski and Sanders (1959, 1962)
devised an ingenius hypothesis to explain preservation of
sedimentary markings and trace fossils under such circum-
stances. Utilizing observations of Gilbert (1914) and Bag-
nold (1956), they pointed out that in traction carpets?
grains expend their energy laterally, and as a result there
is no turbulence and hence no scouring (Dzutinski and
Sanders, 1962, fig. 7). Instead, several varieties of tool
marks (2.¢., markings caused by the gouging of the bottom
by shell fragments, and the like) are produced. These mark-

"Dzutinski and Sanders (1962, p. 88) defined a traction carpet as the
zone “... which is not invaded by turbulent eddies of any size from
the overlying turbulent flow.”

ings are almost immediately covered by gravity settling and
thus preserved from future scouring. In contrast, when the
entire load is carried in suspension and no traction carpet
is present, scouring occurs and all surface markings are
obliterated. This hypothesis was proposed to explain the
method of preservation of surface markings in sequence
characterized by graded-bedding. As yet unexplained is how
similar markings can be preserved in other environments.
The clarity of detail present in Cincinnatian tool marks
certainly rivals that found in the finest turbidite sequences.
Seilacher (1962a) sidestepped the problem of preserva-
tion by maintaining that most arthropod traces, as well as
some others, are of endogenous origin. He envisioned the
thin layer of silt mentioned previously in the example of the
trilobite track. However, it is difficult to picture such a
thin blanket consistently covering wide areas. Likewise, it
must be explained how light objects such as crinoid stems
could penetrate through this layer to leave such clear im-
prints and why there is never any mixing of the silt and
mud, especially where large tool marks are concerned.
Seilacher’s hypothesis may hold in rare instances; however,
it would seem that there is some other, more common
mechanism. Cincinnatian materials were examined in the
hope of shedding some light on the problem, First, an at-
tempt was made to determine if the traction carpet and
graded-bedding of Dzulinski and Sanders could be extended
to cover nonturbidite sequences. It was thought that
graded-bedding on a much smaller scale might be found
directly overlying the sole markings. However, thin-section-
ing of a dozen or so of the best preserved specimens revealed
no trace of graded-bedding. It is now believed that cross-
bedding provides the key, for approximately 75 per cent of
all specimens bearing traces or inorganic sole markings show
cross-lamination. Such currents possibly could deposit the
silt without scouring the mud surface. This theory has not
been tested experimentally, but it is proposed here as a
working hypothesis. The shape and angle of the cross-
laminations should be studied in detail and experimental
stream table investigations might also prove fruitful.

III. NOMENCLATURE OF TRACE FOSSILS

A) CLASSIFICATION AT THE GENERIC AND
SPECIFIC LEVEL

Until recently the nomenclatural complexity of trace
fossils was sufficient to frustrate even the most conscientious
worker. Due in large part to the efforts of Hantzschel (1962,
1965), some degree of order has been attained at the
generic level. Unraveling the great number of species, of
course, will require more time.

TRACE FOSSILS CINCINNATI AREA: OsGoop 295

The reason for the complexity of the nomenclature is
due in large part to two factors: 1) the early history of
trace fossils when nearly all forms were thought to be
plants—hence a great number of taxa were based on minute
morphologic details; and 2) the International Code of
Zoological Nomenclature (ICZN) provided no clear guide
lines — thus, different workers have employed different
methods and philosophies of taxonomy. These methods have
ranged from strict adherence to the rules governing z0o-
logical taxa to the opposite extreme of not formally naming
the forms.

The most recent Code (1961) is inconsistent with re-
gard to the legality of trace fossils. Article 24b states that
the Law of Priority applies “when, before 1931, a name
is founded on the work of an animal before one is founded
on the animal itself.” This implies that trace fossil taxa
proposed prior to 1931 are valid, whereas subsequent names
are not. This is an unacceptable situation.

It has been accepted by most modern workers in
palichnology that a form must be named if it is not to be
“Jost” in the literature. Such an example is found among
Cincinnatian trace fossils. In a penetrating and well thought
out discussion, Flower (1955) described what he considered
to be tentacle imprints and resting traces of nautiloids.
These were described in some detail but never named.
Neither Flower’s paper nor the described forms appear in
the section dealing with trace fossils in the Treatise on In-
vertebrate Paleontology or in Hantzschel’s (1965) volume
of Fossilium Catalogus. Although Flower’s conclusions are
not accepted by the present writer, there is certainly no
reason why the paper should be lost to posterity, but such
is generally the price of failing to name the described forms.®

Thus, it can be declared at the outset of this discussion
that, regardless of the individual worker’s taxonomic
philosophy, trace fossils must be named to survive. Also
eliminated are informal names such as “crawling traces of
a gastropod” or “U-tube of an annelid,” for such designa-
tions are awkward to deal with in faunal lists. Witness that
neither James Hall’s (1852) nor Charles Walcott’s (1918)
arthropod tracks appear in the body of the text of Hantz-
schel’s two compilative works. Likewise, it is not sufficient
to consider a trace fossil as a junior synonym of the body
fossil it represents, if indeed the latter can be ascertained at
all. The two should bear separate generic and _ specific
names. For example, the Cincinnatian representatives of
Rusophycus pudicum Hall can be considered with certainty

"This, of course, if not to imply that even the most poorly preserved

trace fossils should be named. Only those forms which are well
preserved and which have distinct morphology are deserving of
names.

to represent the burrow of Flexicalymene meeki (¥oerste),
because the maker of the trace fossil has been found in situ
(Pl. 58, figs. 4, 5). Technically R. pudicum is regarded as
the junior synonym, but, for reasons which will be justified
below, the two are retained as separate taxa,

While there is little disagreement today that trace fos-
sils must be named, there is a diversity of opinion on the
actual system to be employed.

Rudolf Richter (1941, p. 86; 1948, p. 163) used
“nomenclatura describens,’ such as Ichnia catenaria and
Ichnia diserta, latinized but not proposed as “formal”
zoological taxa. Many workers have employed what Seil-
acher (1953a, p. 444) referred to as “semizoological nomen-
clature.” The name appears to conform to zoological nomen-
clature, but it may have affixed to it such a designation as
“new form;” thus, according to Seilacher, losing the protec-
tion of the ICZN. The latter procedure has been followed by
Schmidtgen (1927), Desio (1940), and Caster (1944).
Other similar methods would be placing the name itself or
the word “genus” in quotes or using the terms “ichnogenus”
and “ichnospecies.”

A more elaborate and even more unorthodox method
of nomenclature was employed by Faul (1951) for tetrapod
traces. His “species” is designated by a capital letter, fol-

‘

lowed by a numeral indicating the “greatest number of feet
clearly impressed.” The last numeral indicates the number
of toe imprints in each track. For example, a four-toed
bipedal track would be A 2.4. If the same author erected
another similar species, it would be B 2.4. Such a designa-
tion may suffice for one’s personal use, but it would be im-
possible to handle large numbers of such keys, and, as
Seilacher (1953a, p. 444) noted, most invertebrate traces
would be “A 0.0.”

Desio (1950) went so far as to suggest giving uninomial
names to inorganic sedimentary markings, such as flow casts
and ripple marks and binomial names to trace fossils. The
main problem here is that this system removes trace fossils
from the biologic domain and considers them as sedimentary
markings instead of traces of life.

Because it appears that trace fossils are not recognized
as formal zoological taxa, and thus not protected by the
rules, one may employ any system of nomenclature one
desires. However, for the sake of stability it is best to follow
some set pattern, and the best possible choice seems to be
the rules of zoological nomenclature. There are two reasons
for this choice. First, most trace fossil workers are paleon-
tologists and are familiar to varying degrees with the rules.
Second, at some future date the ICZN may accept trace
fossils as valid zoological taxa or possibly even as valid
parataxa. If until then diverse schemes of nomenclature are

296 PALAEONTOGRAPHICA AMERICANA (YI, 41)

followed, a massive reshuffling will be necessary. This does
not mean that the suggestions of Seilacher (1953a) and
Hiantzschel (1962) of adding the suffix “ichnus” or “ich-
num” to generic names may not be followed. This is an
excellent proposal, for it allows the reader to quickly realize
that he is dealing with a trace fossil.

What has been discussed above is in essence a practical
problem. The deeper and more complex problem is of a
philosophical nature and deals with the principals of
taxonomy to be employed in the study of trace fossils. At
present there are two opposing schools. One holds the more
traditional zoologically oriented view that we are dealing
with the remains of an organism as shown by its activity,
and we are naming the organism as mirrored by the trace
rather than the trace itself. The latter has no intrinsic value.
Howell (1957) probably provided the best example, for
even his title, “New Cretaceous Scoleciform Annelid from
Colorado,” belies his philosophy. He discussed an annelid
rather than the U-tube. Within the text he placed his form
(Polyupsilon coloradoense) in proper taxonomic order from
phylum through the specific level. However, there is no
vestige of the remains of the maker, only its tube.

A number of workers would agree with Howell’s pro-
cedure. There is excellent reason for this; paleontologists
have long been aware of the heterogenous nature of their
material. Introductory paleontology texts list several dif-
ferent kinds of fossils grading from the finest, such as a
mammoth preserved in Pleistocene ice, to the most poorly
preserved mold. It is realized that the fossil record is highly
incomplete both in regard to numbers and preservation, yet
work must be done with what is available. Trace fossils
are not merely a side branch of fossils, for there 1s an im-
perceptible gradation between body fossils and trace fos-
sils. Plate 57, figure 1 illustrates a specimen of Rusophycus
carleyi J. F. James, which combines the features of the
mold of a body fossil with the attributes of a trace fossil.
It is the ventral mold of a trilobite in that the imprints of
the hypostome, genal spines, cephalic doublure, and walk-
ing legs are present, yet the lateral lobes of a typical Ruso-
phycus are also visible. By our definition this is a trace
fossil because it shows a trace of voluntary movement;
however, it could just as easily be considered the ventral
mold of an isotelid. If one recognizes this intergradation
between molds and trace fossils, it is reasonable to follow
the zoological philosophy embodied by Howell. It may also
be argued that many trace fossils have resulted from the
activity of soft-bodied forms of which there is only a minute
statistical chance of discovering. In this sense the main value
of trace fossils would be their contribution to our knowledge
of body fossils; all else would be secondary.

Recently an opposing point of view has arisen, cham-
pioned mainly by Seilacher but having its roots in the
studies of Rudolf Richter. This school considers that the
main value of trace fossils is ecological rather than morpho-
logic. Seilacher, Richter, and Hantzschel all pointed out that
it is a mistake to place the major emphasis on the biologic
systematics of the originator of the trace. The reason that
palichnology is still considered by some today as an avoca-
tion rather than a valid scientific pursuit is undoubtedly
due at least in part to this overemphasis. Seilacher (1953a,
p. 445) remarked, “we name the trace fossil as an inde-
pendent manifestation, i.e. not as a proxy for the originator.”
To Seilacher the trace fossil does possess intrinsic value. If
one is to orient trace fossils toward paleoecology, the pro-
cedure of taxonomy must change correspondingly, Tradi-
tionally genera and species have been based primarily on
morphological not ecological interpretation; yet where the
ecologic difference between two trace fossils is marked,
taxonomic separation is justified to emphasize this differ-
ence. For example, Seilacher (1955) separated Rusophycus
from Cruziana because the former was interpreted as a
trilobite resting trace, the latter a feeding trace. However,
it is entirely possible that the two traces were made by the
same species of trilobite. A similar example can be given
from Cincinnatian trace fossils. While it is probable that
both Asaphoidichnus trifidum Miller (Pl. 71, fig. 6) and
Allocotichnus dyert, n. gen. (Pl. 72, fig. 1) are trails of
Tsotelus, they exhibit two different forms of movement. A.
trifidum (Text-fig. 20-1) is indicative of straight-ahead
movement, while in A. dyeri (Text-fig. 21-B) the body axis
was oriented to the right of the direction of movement, and
only the first four or five pairs of walking legs were used.
This difference of behavior is judged sufficient to justify
separate generic designations.

One might believe that such a procedure would lead
to a vast proliferation of names. In actuality the opposite
is true, for modern palichnologists realize that slight changes
in behavior and preservation can lead to an apparent dif-
ference in morphology. Miller (1880) established Petalich-
nus multipartitum (Pl. 75, fig. 4), Trachomatichnus permul-
tum (Pl. 73, fig. 5), and Trachomatichnus cincinnatiensis
(Pl. 74, fig. 2) for trails of cephalopods. As will be shown,
these are trilobite tracks, and there is no evidence to in-
dicate that the trails were made by different species. The
difference in morphology between P. multipartitum and T.
permultum is due solely to the orientation of the body dur-
ing movement (see Text-fig. 22). In the former the body
was oriented to the right of the axis of movement, while the
latter shows a conversion from straight-ahead to oblique
movement. 7. cincinnatiensis differs because it is preserved

TRACE FOSSILS CINCINNATE AREA: OsGoop 297

as a cleavage relief and the body was parallel to the axis
of movement. Because none of these variations are judged
to be as significant as those separating A. trifidum from A.
dyeri, Miller’s three species are combined under P. multi-
partitum. At the same time care must be taken to avoid
excessive “lumping.” The results of such action are genera
which contain such a wide range of variation as to render
them virtually meaningless.

It has been clearly indicated by several studies, e.g.,
Seilacher (1957), that trace fossils vary more with ecologic
setting than with age. There is no reason, a priori, why a
Cambrian form should not be combined in the same genus
with one from the Tertiary if the two are morphologically
identical. In traditional taxonomy such practice is frowned
upon; in palichnological studies it is the rule rather than
the exception. However, “identity” here, as in al] taxonomy,
is a matter of “expert opinion.”

We thus have two apparently irreconcilable schools,
but it is believed that a middle ground exists which combines
the best of both views. It recognizes the fact that trace fos-
sils grade imperceptibly into body fossils, and there is a
philosophical basis for treating them as valid zoological
taxa. However, it also recognizes that an important value
of trace fossils lies in what they can contribute to the paleo-
ecologic setting, and the taxonomy must be geared to it. In
this paper trace fossils are recognized as the manifestation
of organisms (thus contributing to the knowledge of mor-
phology), but they also are recognized as possessing in-
trinsic value (thus contributing to the knowledge of paleo-
ethology and paleoecology in a way that a body fossil can
not). Usually, morphologic criteria are employed in dif-
ferentiating forms and constitute the underlying philoso-
phical nomenclatural basis. However, in certain cases the
activities of the once living creature are taken into con-
sideration and used as a taxobase. The study of trace fos-
sils requires this kind of treatment, for if the paleoecological
importance of trace fossils is not acknowledged and accepted
as a basis for taxonomic treatment of them, the science will
not progress as it should.

B) SUPRAGENERIC CLASSIFICATION OF TRACE FOSSILS

Although there may be room for argument as to
whether trace fossils constitute valid zoological taxa, it is
obvious that they do not fit easily into the zoological
heirarchy. Some authors, e.g., Rudolf Richter (1926) and
Martinsson (1965), have erected “families;” however, there
was never a pretext that these constituted valid taxa in
the zoological sense. Several authors have attempted other
means of classification.

Hundt (1931) separated trace fossils into two major
groups — those which originated on the surface and those
which formed at depth. These two categories were further
subdivided according to the geometric pattern of the trace
(e.g., stellate, pocket-like, ramified ).

Krejci-Graf (1932) presented a much more elaborate
classification based mainly on the various life activities of
organisms. He proposed three major headings: traces of
existence (coprolites), traces of repose, and traces of move-
ment. Minor categories were based on life activities such as
crawling, swimming, eating. This classification, although
more comprehensive, is so elaborate that it tends to be
cumbersome.

Seilacher (1953a) proposed a classification similar in
essence to Krejci-Graf’s but greatly simplified. His scheme
is shown diagramatically in Text-figure 2. By employing a
“pie” diagram, he stressed the inseparability of trace fossils
from body fossils. The traces are subdivided into five etho-
logical categories, and for each he coined a new latin name
with the -ichnia suffix. Seilacher’s categories are as follows:

1) Pascichnia (grazing trace). Normally two dimen-
sional, these traces are usually tightly coiled or appear as
closely packed meanders which are only rarely branched.
In a sense they follow the principals of strip-mining, 1.¢.,
systematically covering the greatest area with the least pos-
sible expenditure of energy. Rudolf Richter (1928) was the
first to conclude that such traces are the feeding trails of
vagile benthos. While Seilacher (1962a) maintained that
Pascichnia are of infaunal origin, there is no strong neo-
ichnological evidence to support his conclusion. The present
author has observed Recent examples in photographs taken
in 4000 meters of water on the Mid-Atlantic Ridge by ves-
sels of the Woods Hole Oceanographic Institution. More-
over, Seilacher (1967b, pl. 2, fig. c) illustrated a Pascichnia
of what appears to be an enteropneust in the process of
of formation. The photograph was taken by the Lamont
Laboratories at a depth of 4700 meters. Additional Recent
forms can be found in Schafer (1962, figs. 155, 171). With
one possible exception (Paleodictyon) Pascichnia are not
present in the Cincinnatian strata.

2) Fodinichnia (feeding structures). Traces of this form
are the three dimensional analogues of Pascichnia, for they
too show maximum space utilization. Whereas Pascichnia
exhibit the characteristics of strip-mining, Fodinichnia,
which are infaunal, embody the principles of mining at
depth. They commonly follow nutrient-rich layers of sedi-
ment. They may be branched and illustrate phobotaxis as
in Chondrites (Pl. 79, fig. 6), bundled as in Phycodes (PI.
69, fig. 4), or possibly U-shaped as in Rhizocorallium.

298 PALAEONTOGRAPHICA AMERICANA (YI, 41)

Fodinichnia differ because they also serve as a permanent
or semi-permanent dwelling structure for the organism pro-
ducing them. The U-tube of Arenicola marina (Wells, 1945)
is a Recent example, as are the burrows of Heteromastus
filiformis illustrated by Schafer (1962, fig. 215). Fodinichnia
are abundant in the Cincinnatian, both in kind and numbers
(Text-fig. 29).

3) Repichnia (crawling trace=K reichspur of Seilacher,
1953a). Repichnia are simple, branched or unbranched trails,
burrows, or tunnels of vagile benthos which originate either
on the surface or at depth. This category also includes
tracks of arthropods. These are simple traces of movement
made as the organism moved from one location to another,
and thus they lack the characteristic morphologic patterns
of Pascichnia. To use Seilacher’s comparison, Repichnia are
similar to modern highway systems, for they lack numerous
sharp curves and instead are linear or gently arcuate. Recent
examples are easily seen in a variety of settings — ponds,
streams, seafloors, Repichnia are common in the Cincinna-
tian and consist of arthropod tracks and gently arcuate
trails and burrows.

4) Domichnia (dwelling structure). Domichnia serve
as dwelling structures for sessile or hemisessile benthos.
Generally they are simple unbranched vertical tubes, U-
tubes, or in some cases W-tubes. The inhabitants are filter
feeders, scavengers, or predators; and since the structure is
utilized as a more or less permanent domicile, the walls
may be coated with mucus to prevent collapse. Recent
counterparts are the straight, unbranched tubes of phoro-
nids, as well as the U-tubes of numerous polychaetes. Cin-
cinnatian representatives consist primarily of U-tubes.

5) Cubichnia (resting trace). Cubichnia are shallow
burrows which mirror the outline of the producer in varying
degrees of clarity (Pl. 58). They are unbranched but may
show some signs of rotary movement and are epifaunal in
origin. The purpose of such burrows, e.g., Rusophycus (PI.
58) and Asteriacites (Pl. 62, fig. 5), in the fossil record is
not entirely clear. Based on experimental observations on
Recent materials, Seilacher (1953b) surmised that these
might well have provided temporary shelter and safety for
vagile benthos. In contrast Fenton and Fenton (1937a)
suggested that some Cubichnia, e.g., Rusophycus, may
represent egg depositories, while Caster (1938) proposed
that such burrows could be employed as a device to keep
the gills moist when the organism was exposed. The latter’s
hypothesis was based on studies of the Recent Limulus. Re-
cent examples are difficult to recognize unless the producer
is “caught in the act,” for once the organism has departed
or is buried, only a disturbed area remains as evidence.
Illustrations of Recent asteroid and xiphosuran Cubichnia

are given by Seilacher (1953b, pl. 11, fig. d) and Caster
(1938, pl. 12, fig. 2). Cubichnia constitute one of the more
common forms of Cincinnatian traces.

The boundaries between Seilacher’s categories are not
sharp. For example, both Domichnia and Fodinichnia serve
as dwellings, and it is difficult in practice to separate linear
Repichnia from the burrows of vagile selective feeders or
predators. To this extent the classification is somewhat sub-
jective but is still workable, especially when combined with
his “classification” of the preservation of trace fossils.

Lessertisseur (1955) utilized a modified form of Seil-
acher’s classification combined with the general configura-
tion of the trace. He subdivided traces into endogenous and
exogenous, and each of these in turn is subdivided into
traces of locomotion, nutrition, and repose, with the endo-
genous forms having the additional category of dwelling
trace, Each of these is again separated into units based on
shape, e.g., spiral, helminthoid,

In a more recent classification, Miiller (1962, Text-
fig. 3) greatly expanded Seilacher’s scheme. He recognized
four major categories; traces of movement, bioreaction
(sickness, parasitism, death struggles), resting, and eating.
These are divided into the groups employed by Seilacher
with the additon of such categories as swimming, flying,
gnawing and biting, and running. This is a much more
comprehensive classification which follows the general
format used by Abel (1935) in treating Vorzertliche Lebens-
spuren.

All of the attempts at classification of traces recognize
geometry or ethology as fundamental criteria. There seems
little point in classifying trace fossils merely according to
general shape. As Seilacher (1953a) showed the same
morphology is created in many instances by different
creatures, and at the same time dissimilar traces can be
made by the same organism. An ethological classification is
judged to be the most workable and also the most meaning-
ful. Seilacher’s (1953a) scheme best serves current needs.
It is possible to fit all but the most unusual traces into his
groups, and his units also provide an excellent framework
for comparison of suites of ichnofossils as are discussed in

topic XIII.

IV. HISTORY OF THE STUDY OF CINCINNATIAN
“FUCOIDS” AND TRACE FOSSILS

The first reference to “fucoids” in the Cincinnatian sec-
tion was made by Orton (1873) in a report on the geology
of southern Ohio. He briefly referred to the “plants of the
Blue Limestone” and the “dumb-bell fucoid” (Corophioides

biclavata (Miller) (PI. 61, fig. 1), and also reported (1873,

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 299

p. 268) that “stems and roots apparently referable to the
genera Palaeophycus and Buthotrephis of Hall are com-
mon.” At the same time Orton described one new form
which he provisionally named Palaeophycus radiata. It is
evident from his short discussion that “fucoids” had been
known to local collectors for many years. It is also apparent
that Hall’s work (1847, 1852) on the “fucoids” of New York
State was regarded as the general reference on North Ameri-
can forms. Nearly all workers on Cincinnatian material re-
ferred to this study at one time or another.

In 1874 Leo Lesquereux, who is better known for his
work on the Pennsylvanian coal flora of Ohio, made what
he thought was the momentous discovery of the oldest re-
mains of land plants in North America. From a section six
miles east of Lebanon, Ohio, he described two specimens
which he believed represented small stems of branches of
Sigillaria. Such a find could not long escape notice or further
study; and in the same year Newberry (1874, p. 111) cast
doubt on Lesquereux’s interpretation, reasoning that if these
were indeed land plants, they would show “
tinctness and regularity of surface markings, some coating
of carbonaceous matter and some traces of organic struc-
ture.” He also reminded Lesquereux that the Cincinnati
area, during Silurian (Ordovician) time, was so far from
land that it would be difficult to explain the presence of
land plants even by floating. Lesquereux was apparently
not convinced, for in 1878 he formally erected three new
species of Cincinnatian “land plants” based on the discovery
of new material. These were sent to Dana, Eaton, and Ver-
rill] and he reported that they concurred with his opinion.
The only subsequent reference to these forms in the local
literature was by Miller (1889) and J. F. James (1891),
who both questioned Lesquereux’s interpretation.

The description of Cincinnatian “fucoids” began in
earnest in 1878. In a three year period (1878-1881), U. P.
James, a Cincinnati bookdealer and printer, described seven
species in his own short-lived journal, The Paleontologist.
Most of these species were referred to Hall’s Clinton genera
and none were figured. Also in 1878 the most extensive work
on local “fucoids” was published by Miller and Dyer, who,
in two papers, proposed 15 new species distributed among
eight new genera. In reading these papers, one gets the
idea that the work was mostly Miller’s and that Dyer’s con-
tribution was to supply the specimens. In contrast to the
work of U. P. James, most of the forms described by Miller
and Dyer were accompanied by excellent lithographic re-
productions.

It is ironic that Miller, who adhered so strongly to the
“fucoid” notion, was the first to describe tracks as such
from the local rocks. In 1880 he created four new genera

. more dis-

and seven new track species. Because of associated body
fossils, most of the forms were assigned to cephalopods and
the remainder to trilobites.

Up to this point no one had questioned the vegetable
nature of the many “fucoid” forms described, and the Cin-
cimnatian strata had become second only to the New York
State section in the number of North American species. In
1884 and 1885 J. F. James, another Cincinnati amateur and
the son of U. P. James, attacked all of Miller and Dyer’s
forms as well as those of his father. He made no mention of
Miller’s paper on the tracks and trails. James concluded
that all Cincinnation “fucoids” were either inorganic sedi-
mentary markings or tracks, trails, and burrows. It is inter-
esting to note that James used the same tools as Nathorst
(1881), although he clearly stated that he had no knowledge
of Nathorst’s work until his own paper was near comple-
tion. In fact, James’ paper closely resembles Nathorst’s,
though on a much smaller scale. Whereas Nathorst made
extensive observations in intertidal areas, employing plaster-
of-Paris molds and photographs to support his conclusions,
James confined himself to the local ponds and streams. The
main deficiency of his work is that all the illustrations are
highly schematic. This, however, is not to detract from the
importance or quality of his paper, for he was laboring
under most difficult conditions. Although he probably
studied some of his father’s types, it is apparent that he
never saw any of Miller’s specimens. As an amateur he was
writing in an age when the plant nature of “fucoids” was
still widely accepted by professionals. James was to North
America what Nathorst and Fuchs were to Europe, although
regrettably his work has been commonly overlooked or
ignored.

Miller’s North American Geology and Paleontology
for the Use of Amateurs, Students and Scientists (1889)
showed that James’ publications had little effect on him,
for he still persisted in listing all but two of his forms as
“fucoids.” Nevertheless, it is true that in both his 1877 com-
pilation and his work of 1889, he made no effort to arrange
the “fucoids” in families. He specifically indicated in the
prologue to Plantae in the first work (1877, p. 21) that
because of the controversial nature of “fucoids,’ he was
making no attempt at suprageneric classification. It is ap-
parent that there was little love lost between Miller and
James, for the latter’s writings show only contempt for
Miller’s work. In rejecting the “fucoid” assignment of
Miller’s Walcottia, James (1885, p. 162) said, “it is safe to
consign the genus with its species to the limbo of the im-
probable and wipe from the catalogues three more useless
names.”

James’ works of 1884 and 1885 represent the last major

300

publication on Cincinnatian trace fossils. From the period
1891-1897 James undertook an extensive summary of Cin-
cinnatian paleontology, and in the preface to this work he
listed all those forms which he believed to represent the
activity of annelids, trilobites. Unfortunately the body of
the text which he promised was never completed due to
his untimely death. The only other mention of Cincinnatian
traces was by Caster (1938), who made a brief reference to
some arthropod tracks but gave them no extended treat-
ment, and by Flower (1955), who discussed what he in-
terpreted as tentacle imprints and resting traces of nautil-
oids. Hiantzschel (1962) included Cincinnatian “fucoid”
genera in the Miscellanea volume of the Treatise on Inverte-
brate Paleontology, but, lacking the type specimens and
without a first-hand knowledge of the forms, he could add
little except for limited first-hand observations of Seilacher
which were made available to him.

TRACE FOSSILS AND “FUCOIDS” REPORTED FROM THE CINCINNATION

The following chart portrays the synoptic history of
“fucoids” and trace fossils reported to be present in the
local Cincinnatian section. Column “1” represents the inter-
pretation of the original author, column “2” the interpreta-
tion of J. F. James (1884, 1885, 1891, 1892), column “3”
the interpretation of Hantzschel (1962, 1965), column “4”
the interpretation given in the present work. With regard
to the taxonomic treatment of individual forms, the first
name given is the valid name. Names placed in parentheses
represent the assignments of the original author. A key to
the symbols employed in the chart is given below.

A—alga

B—body fossil

C—Cubichnia

D—Domichnia

F—Fodinichnia

I —inorganic

L—land plant

P—Pascichnia

R—Repichnia

T—trace fossil

?—Incertae sedis
——author did not discuss species

Comments
n. gen.

ies)
res)
-

Name 1

Allocotichnus (Asaphoidichnus) R R R R
dyeri (Miller), 1880

Aristophycus ramosum Miller A I I?
and Dyer, 1878

Aristophycus ramosum var.
germanum Miller and
Dyer, 1878

Asaphoidichnus trifidum R R R R
Miller, 1880

Asteriacites (Heliophycus) A B (e (c
stelliforme Miller and
Dyer, 1878

Blastophycus diadematum A Te Vl I
Miller and Dyer, 1878

=A. ramosum

PALAEONTOGRAPHICA AMERICANA (YI, 41)

Name
Buthotrephis filciformis
U. P. James, 1878
Chloephycus plumosum Miller
and Dyer, 1878
Chondrites (Buthotrephis)
gracilis Hall, 1843
Chondrites (Buthotrephis)
gracilis var. crassa
Hall, 1852
Chondrites (Buthotrephis)
ramulosa Miller, 1874+
Chondrites (Buthotrephis)
succulosa Hall, 1847
Chondrites, type-A
Chondrites, type-B
Chondrites, type-C
Conostichnus truncatum
Ulrich, 1880
Corophioides (Arthraria)
biclavata (Miller), 1875
Corophioides cincinnatiensis,
n. sp.
Corophioides cf. luniformis
Cyathophycus silurianum
J. F. James, 1891
Cyclophycus laterale Ulrich,
1880
Dactylophycus tridigitatum
Miller and Dyer, 1878
Dactylophycus quadripartitum
Miller and Dyer, 1878
Dystactophycus mamillanum
Miller and Dyer, 1878
Fascifodina floweri, n. gen.,
Nn. sp.

Fucusopsis (Trichophycus)
sulcatum Miller and
Dyer, 1878

Lockeia siliquaria U. P.
James, 1879

Mastigograptus (Psilophytum)
gracillinum Lesquereux,
1878

Ormathichnus moniliformis
Miller, 1880

Palacophycus flexuosum
U. P. James, 1879
Palacophycus ornatum

Ulrich, 1880

Palaecophycus radiata
Orton, 1873

Palacophycus rugosum Hall,
1847

Palaeophycus sculptum Ulrich,
1880

Palaeophycus tubulare Hall,
1847

Palacophycus virgatum Hall
1847

Palaeophycus, type-A

2 3
I =
I I
B D/F
B D/F
iT =
Ts DAR
I 1k
TAS
T/L T/L
B ?
I At
B (&
1G, =
I R
I =
T
Af ar
B =

oy oy

R

Comments
=Chloephy-

cus

not present

not present

no specimens
seen
not present

nomen nudum

new report

new report
=Trichophy-
cus venosum
nomen nudum

probably=D.
tridigitatum

interpreted
by Flower,
1955, as cep-
halopod ten-
tacle imprint

one syntype
specimen in-
organic

nomen nudum
?=Phycodes or
Dactylophycus
not present

nomen nudum

not present

F/D Cincinnatian

R

forms= Tricho-
phycus veno-
sum

new report

TRACE FOSSILS CINCINNATI AREA; OsGoop 301

Name 1 2 3 4 Comments
Palacophycus, type-B - - - R_ new report
Palacophycus, type-C - - - R_ new report
Palaeoscia floweri Caster, B = oe AGIAN

1942
?Paleodictyon sp. Savi and AS = T I/P new report
Meneghini, 1850
Petalichnus multipartitum R = R R
Miller, 1880
Phycodes (Licrophycus) A B F F/D
flabellum (Miller and
Dyer), 1878
Protostigma sigillaroides L L = ? not seen
Lesquereux, 1878
Rhabdoglyphus sp. Vassoe- fin Ts R_ new report
vich, 1952
Rusophycus bilobatum A als Cc C_ not present
(Vanuxum), 1842
Rusophycus (Cruziana) Ti = ( (
carleyi (J. F. James),
1885
Rusophycus cryptolithi, n. sp. - - - C_ new report
Rusophycus pudicum Hall, Ay aE: G Cc
1852
Rusophycus subangulatum AY C C/R not present
Hall, 1852
Saccophycus intortum U. P. TUS ey ? not seen
James, 1879
Skolithos (Scolithus) deli- A Tl = D
catulus U. P. James, 1881
Skolithos (Scolithus) dispar I - - I
J. F. James, 1892
Skolithos (Scolithus) tuber- ‘iii OMe w= F?
osus Miller and Dyer, 1878
Sphenophyllum primaevum L L = ? not seen
Lesquereux, 1878
Teratichnus confertum R» = R R
Miller, 1880
Trachomatichnus cincinnatien- =Petalichnus
sis Miller, 1880 multipartitum
Trachomatichnus permultum =Petalichnus
Miller, 1880 multipartitum
Trachomatichnus numerosum R = R R
Miller, 1880
Trichophycus lanosum Miller AS Pee ety aR:
and Dyer, 1878
Trichophycus venosum Miller, A I Af S1RY/D)
1879
Tylichnus (Rusophycus) 7 \om Daa Rn. gen.
asperum (Miller and Dyer),
1878
Walcottia rugosa Miller and A/T B ? R
Dyer, 1878
?Walcottia cookana Miller 2 leon) 4 temas g ? not seen
and Dyer, 1878
Walcottia sulcata U. P. = a ? ? not seen

James, 1881

V. CUBICHNIA

RUSOPHYCUS Hall, 1852

Plate 57, figures 1, 5, 6; Plate 58, figures 1-10; Plate 59, figures 4-6;
Plate 60, figure 3; Plate 66, figure 3; Plate 71, figure 1; Plate 82,
figure 9; Text-figure 5, 29-a,b,c.

1823. Non Fucoides Brongniart, Soc. Nat. Hist. Paris, Mem. 1, p.
308, pl. 19, fig. 2.

1831. Non Bilobites Rafinesque, Enumeration and Account of Some
Remarkable Natural Objects in the Cabinet of Prof. Rafinesque
in Philadelphia, p. 2.

1839. Non Bilobites d’Orbigny, Voyage dans |’Amérique meridional,
Atlas, pl. 1.

1842. Non Cruziana d’Orbigny, Voyage dans |’Amérique meridional,
vols. 3-4, p. 30.

1842. Partim Fucoides Brongniart, Vanuxem, Nat. Hist. New York,
Geol. New York, pt. 3, p. 79, fig. 11-1.

1852. Partim Rusophycus Hall, Nat. Hist. New York, Palaeont. New
York, vol. 2, p. 23, pl. 9, figs. 1-3; pl. 8, figs. 6a,b.
1864. Rusichnites Dawson, Canadian Natur., n.s., vol. 1, p. 367.

1879. Rhysophycus Schimper, im Zittel, Handbuch der Palaont., 2d
ed., p. 54.

1885. Cruziana d’Orbigny, James, J. F., Cincinnati Soc. Nat. Hist.,
Jour., vol. 7, pp. 153-157, pl. 8, fig. 1.

1935. Cruziana d’Orbigny, Abel, Vorzeitliche Lebensspuren, pp. 253-
255, fig. 224.

1953. Cruziana d’Orbigny, Seilacher, Neues Jahrb. Geologie u.
Palaont., Abh., Bd. 98, pp. 107-115, figs. 4-5.

1955. Rusophycus Wall, Seilacher, im Schindewolf, and Seilacher,
Akad. Wiss. Lit. Mainz, Math.-nat. KI], Abh., No. 10, p. 368,
pi ZOsties: Way sep Oe tion plano. atic:

1955. Partim Cruziana d’Orbigny, Seilacher, im Schindewolf, and
Seilacher, Akad. Wiss. Lit. Mainz, Math.-nat. K]., Abh., No. 10,
p. 368, pl. 16, fig. 2; pl. 19, fig. 1; pl. 20.

1955. Rhysophycus Schimper, Lessertisseur, Soc. Géol. de France,
Mem., n.s., vol. 74, pp. 44-47, figs. 25-27, 39; pl. 6, fig. 4.

1959. Rusophycus Hall, Seilacher, Die Naturwissenschaften, vol.
12, pp. 292-293, fig. 4.

1962. Rusophycus Hall, Hantzschel, Trace-Fossils and Problematica
in ‘Treatise on Invertebrate Paleont., Moore (ed.), pt. W,
Miscellanea, p. W212, figs. 131-3, 5.

Type species — Fucoides biloba Vanuxem (1842) from
the Silurian (Clinton) of New York State.

Diagnosis — Short bilobate Cubichnia of trilobites.
The lobes are commonly covered by anterolaterally directed
striae. Imprints of cephalic and pygidial doublures or pygi-
dial doublures, genal spines, and thoracic pleurae may be
present.

Discussion — Probably the most famous of all the
“fucoids,’ Rusophycus served as the principal example in
the “fucoid-” trace fossil controversy of the 1880’s and has
been the subject of numerous nomenclatural arguments.

Rusophycus, which is typically preserved as a convex
hyporelief in silt or sand-sized material, is a common
Paleozoic genus with a widespread geographical representa-
tion. Broadly speaking the body consists of two parallel
lobes separated by a median furrow. Beyond this it is dif-
ficult to generalize, for the genus shows a wide range of
morphologic variation. The length varies from 1 cm in the
Cambrian R. didyma Salter to over 25 cm in some Cincin-
natian forms. In most specimens the width is equal to one-
half the length. The height of Rusophycus is independent
of width and length; while most forms are only moderately

3()2

PALAEONTOGRAPITICA AMERICANA (VI, 41)

Text-figure 5.— Diverse Cambrian examples of Rusophycus. A.
Reconstruction of a trilobite excavating a shallow burrow. B. Dia-
gram showing the direction of movement of the legs in the formation
of the burrow illustrated in fig. A above. C. Shallow burrow where
the lobes are separate throughout their entire length; X 1.5. D. Bur-
row with a V-shaped outline; X 1.5. E. Deeper, more typical, cordate
burrow; X 0.5. F. Interpretation of R. jennings: (Fenton and Fenton,
1937) showing that the cephalon was used as a shovel. G. R.
jenningsi showing delicate striae. H. Lateral view of fig. 6 above
showing the imprints made by the cephalon (right) and the walking
legs (left); X 0.5. I. Reconstruction of a burrow from the Lower
Cambrian rocks of Lugnas, Sweden. K. Diagram of fig. I above
showing that some of the walking legs moved the excavated material
anteriorly whereas the remaining legs moved it posteriorly. L. Lateral

view of fig. I, K above; X 0.5. (From Seilacher, 1959, fig. 4.)

convex (PI. 58, fig. 8), in some specimens (Text-fig. 51-L)
the height is nearly equal to the length. In contrast some
forms (PI. 57, fig. 1), which may be as much as 20 cm long,
have a height of only 2-3 em. Although the general outline
is elliptical, it may vary considerably. The lobes may part
slightly at one end to give a cordate appearance (Text-fig.
5E), or they may be so deeply notched that the resulting
outline resembles a “V” (Text-fig. 5D). On rare occasions
the lobes may be separate throughout their entire length
(Text-fig. 5C). Even in ellipsoid specimens the median fur-
row can take on various expressions. In most forms it is a
simple V-shaped cleft, but occasionally the lobes may part
to expose two rows of nodelike bodies (PI. 58, fig. 8). A more

extreme case is where the lobes are poorly developed and

the median area has become highly complex. Specimens of
Rusophycus with such well-developed median areas are thus
far known only from the Upper Cambrian of Poland and
the Cincinnatian.

One of the most diagnostic features of Rusophycus are
the striae which cover the lobes. As shown in Text-figure 5B
the striae are normally directed anterolaterally, although in
some forms, ¢.g., R. bilobatum Hall (1852, pl. 9, fig. 3),
they may be nearly normal to the longitudinal axis.

Striae vary in size from extremely coarse, as in R. bilo-
batum Hall and R. grenvillensis Dawson, to fine as in R.
didyma Salter. Although Seilacher (1960) generalized that
the larger the specimen, the better developed are the striae,
Cincinnatian forms do not confirm this statement. Some

‘TRACE FOSSILS CINCINNATL AREA: OsGoop 303

large forms (PI. 71, fig. 1) may have almost smooth lobes,
while much smaller specimens have well-developed striae
(Pl. 60, fig. 2).

There has been considerable disagreement regarding
the nomenclatural status of these buckle-lke burrows. The
problem revolves mainly around the usage of four names:
Bilobites Rafinesque (1831), Rusophycus Hall (1852),
Crossochorda Schimper (1879), and Cruziana d’Orbigny
(1842). D’Orbigny (1839) proposed the name Bilobites for
a “fucoid” which was illustrated in the plates of his South
American work. By the time the text was published (1842),
he realized that this name was apparently preoccupied by
Bilobites DeKay (1824), a pelecypod; in its stead he pro-
posed Cruziana. Unaware of D’Orbigny’s work, Hall (1852)
erected the genus Rusophycus for a form from the Clinton
(Silurian) beds of New York State, figured by Vanuxem
(1842) under the name Fucoides biloba. At the same time
Hall described several new Clinton species. For the next
100 years confusion reigned and Bilobites, Rusophycus, and
Cruziana were used interchangeably. To add to the confu-
sion several authors altered (illegally) the spelling of Ruso-
phycus, e.g., Rusichnites Dawson (1864), Rhysophycus
Schimper (1879). Sinclair (1951) summarized the problems
surrounding the usage of Bilobites, finding that the name
has been employed at various times for a pelecypod, trace
fossil, brachiopod, and trilobite. He concluded that Bilobites
Rafinesque (1831), for an unrecognizable trilobite, has
priority. The reader is referred to Sinclair for a more com-
plete discussion of the Bilobites problem.

Seilacher (1953b) restricted Cruziana to the short bilo-
bate buckle-like imprints, 7.e., Rusophycus in the usage of
Hall. The longer bandlike bilobate forms, such as those
described by Delgado (1886), he referred to Crossochorda
Schimper (1879). At the same time he placed /sopodichnus
Bornemann (1889) in synonymy with Cruztana. lsopodich-
nus is morphologically similar to Cruziana (in the sense of
Seilacher, 1953b) but has been employed mainly for Trias-
sic and later forms. In 1955 Seilacher reversed his opinion.
He now considered Rusophycus as the short, buckle-imprints
and in part restricted the longer bands to Cruziana. He
likewise separated Cruziana from Crossochorda. The reason
for this sudden shift was his ethological interpretation of
the various forms. Rusophycus was thought to represent a
resting trace (Cubichnia), while Cruziana was interpreted
as a grazing trace (Pascichnia). Crossochorda, which he
interpreted as a crawling trace (Repichnia), was considered
to be an intermediate stage between Rusophycus and
Cruziana, Hantzschel (1962) largely agreed with Seilacher’s
treatment of 1955, although he separated /sopodichnus
from Rusophycus. To further confuse matters, Seilacher

(1964) did not employ the generic designation Crosso-
chorda. Forms which had been included under this name in
1955 now came under Cruziana.

It seems best in this case to keep paleoethological in-
terpretation apart from nomenclatural philosophy and base
the names strictly on morphology. The following usage will
be employed in this work: Rusophycus is restricted to the
short bilobate coffee-bean-like imprints of trilobite origin.
This includes all forms described by Seilacher (1953b) under
the name Cruziana, as well as what Seilacher (1955, fig. 5,
nos. 7-8) referred to as Cruziana, Cruziana is restricted to
the longer bilobate forms also of trilobite origin, such as
those described by Delgado (1886). It is obvious that it
is difficult to draw a dividing line between these two genera.
Forms such as those on Plate 66, figure 3 illustrate this
point. However, the situation is not clarified by establishing
an intermediate genus, 1.e., Crossochorda, for this only com-
pounds the problem. A complete study of Crossochorda has
not been undertaken, but it apparently les within the limits
of Cruztana as defined herein. In this work /sopodichnus 1s
restricted to short Rusophycus-like imprints of nontrilobite
origin. Both Seilacher (1960) and Glaessner (1957) agreed
that /sopodichnus is most probably of phyllopod origin.

Interpretation — Because Rusophycus was used as an
illustration for the “fucoid-”’trace fossil controversy, there
is no need to give an extensive treatment of the history of
the genus here. For many years it was interpreted as a
“fucoid” possessing a wrinkled bilobate stem, and Hall
(1852, pl. 9, fig. 1) even illustrated one form with a “stipe”
attached. As early as 1864 Dawson challenged the “fucoid”
assignment, regarding the genus as a trace fossil represent-
ing the burrowing activity of trilobites. His arguments were
similar to those used later by Nathorst, but in addition he
appointed out the similarity of Rusophycus to the Recent
burrows of Limulus. Dawson’s writings made little impres-
sion, and not until after Nathorst’s work of the 1880's was
it generally accepted that Rusophycus was the cast of a
trilobite burrow.

However, studies of Recent burrows have demonstrated
that not all short, bilobate, Rusophycus-like bodies were
produced by trilobites. Seilacher (1960 fig. 1) maintained
that there are at least three other groups of organisms cap-
able of excavating burrows similar to those of trilobites. He
illustrated the shallow burrow of the marine polychaete
Aphrodite. Although no fossil representatives are known,
Seilacher (1960) observed Recent examples on the tidal
flats at Wilhelmshaven.

According to Seilacher (1960) several fossil bilobitids
can be assigned to gastropods (e.g., those described by
Barrois (1882) from the Upper Cretaceous of the Arden-

304 PaLAron'rocraruica Americana (VI, 41)

nes). Recent counterparts produced by Bullia have been
studied by Abel (1935) and Gétzinger and Becker (1932).
The actual mechanics of burrow excavation are unknown.
Seilacher (1960, p. 42) stated that these may be differen-
tiated from trilobite-produced burrows by the orientation of
the striae. In gastropod burrows the striae are directed
posterolaterally from the midline; in Rusophycus they are
directed anterolaterally. Although a generic designation has
not been established for short bilobed burrows of supposed
gastropod origin, longer bandlike forms have been placed in
Scolicia Quatrefages (1849) by Seilacher (1955) and
Hantzschel (1962).

As mentioned previously, the genus Jsopodichnus
Bornemann (1889) has been used for bilobate forms of pre-
sumed phyllopod origin. Jsopodichnus has been applied to
many post-Paleozoic specimens, especially those from the
Triassic of Germany, but Glaessner (1957, p. 106) men-
tioned several possible Paleozoic representatives. Some
forms are from marine environments, others from brackish
and fresh water settings. Recent examples have not been
extensively studied. The only mention of such phyllopod
activity is by Krejci-Graf (1932, p. 31), who made a brief
reference to the bilobed burrows of Artemisia salina observed
in brine pools in Romania. According to Seilacher (1960)
phyllopod burrows can be distinguished by their extremely
small width (+ 1 mm).

Several authors, e.g., Dawson (1864), Caster (1938),
have mentioned bilobate burrows of Recent Limulus. Caster
found that by sprinkling stranded young limuloids with
water, they could be induced to burrow vertically (Caster,
1938, pl. 12, fig. 2). It should be possible to distinguish
well-developed deep limuloid burrows by their general out-
line; however, initially the excavation is reminiscent of
Rusophycus.

As illustrated, several organisms can produce Ruso-
phycus-like burrows, and great care must be taken in inter-
preting them. Unfortunately with a few exceptions, e.g.,
Limulus, Recent perpetrators have not received extensive
study, and more work is required before positive identifica-
tion of fossil forms can be confidently made. This is especial-
ly true in attempting to distinquish between alleged phyl-
lopod and trilobite burrows. The only apparent difference is
size, and this is no sure criterion (¢.g., see Ormathichnus
herein).

Nevertheless many times it is possible to assign such
burrows to specific trilobites. Imprints of the cephalic and
pygidial doublures (PI. 57, fig. 1), genal spines (PI. 58, fig.
1), and thoracic pleurae (Pl. 59, fig. 6) are not uncommon.
Moreover, three specimens are known where the trilobite

has been found in situ (Pl. 58, figs. 4-5; Pl. 57, fig. 6). All

three forms are Flexicalymene meeki and were collected
from the Corryville section at Stonelick Creek, Clermont
County, Ohio. Such specimens are important, not only be-
cause they offer proof of trilobite origin, but also because
they provide a guide for anterior-posterior orientation. They
confirm that the striae are directed anterolaterally from the
midline.

Seilacher (1955) noted that there is never a trace of
the excavated sediment. Rusophycus is not found with the
raised rims one would expect if the sediment were pushed
out to the sides of the burrow. He theorized that the legs
dug toward the median line and to the rear (Text-fig. 5B)
and that the comb-shaped pre-epipodites might have set up
currents which would aid in swirling away excavated ma-
terial. On the other hand, Caster (1938) noted that the
simple up and down digging movement was sufficient to
carry away the sediment.

Seilacher (1955, 1959) discussed two additional varia-
tions of burrow excavation. He described one specimen from
the Lower Cambrian of Sweden where the striae are directed
anterolaterally as well as posterolaterally. Seilacher en-
visioned a trilobite with a strongly arched back in which
the first few pairs of appendages pushed the sediment to the
median line and forward. The remaining pairs behaved in
the more conventional manner, moving the material pos-
terolaterally to the midline.

Normally the cephalon and genal spines took no active
part in the digging process. However, Fenton and Fenton
(1937a) described a burrow (Text-fig. 5F-H), R. jenningsi
from the Lower Cambrian of Alberta in which the cephalon
played more than a passive role. Both the Fentons (1937a)
and Seilacher (1955, 1959) believed that the trilobite (most
likely an olenellid) used the cephalon as a shovel and
scooped the sediment from the anterior portion of the bur-
row. Three distinct impressions of the cephalon may be seen
on the sides of the specimen.

Opinions on the ethological significance of Rusophycus
have differed. Seilacher (1953b, 1955, 1959) believed that
the majority of specimens represent Cubichnia, where the
originator burrowed for concealment. Glaessner (1957) main-
tained that all Rusophycus, as well as the longer cruzianids,
were feeding structures where the movement of the legs set
up a “feeding current.” Finally, Fenton and Fenton (1937a)
suggested that Rusophycus might represent an egg deposi-
tory. While all three explanations are possible, the Cubichnia
assignment appears the most logical, for trilobites are known
to have had vulnerable ventral surfaces. One defense
mechanism developed by many species was that of enroll-
ment, and it is not illogical to assume that Rusophycus
represents another. In most cases the trilobite burrowed just

TRACE FOSSILS CINCINNATI AREA: OsGoop 305

deep enough to cover the dorsum with a thin layer of sedi-
ment; and, based on observation of Recent arthropods, e.g.,
the sand crab Emerita, it is probable that the organism
could completely conceal itself in a few seconds.

CINCINNATIAN REPRESENTATIVES

Most of the work on Cincinnatian representatives of
Rusophycus was done by J. F. James (1885), who under-
took a general survey of all North American species and
correctly interpreted them as trilobite burrows. James con-
cluded that two of Hall’s (1852) Clinton species, R. bilo-
batum and R. pudicwm, were present in the Cincinnatian
strata; and in addition he erected one new species, Cruztana
carleyi.* At the same time he stated that R. swbangulatum
Hall (1852), which he considered to be the senior synonym
of R. clavatum Hall (1852), was not of trilobite origin,
representing instead the trail of a gastropod similar to that
produced by the Recent Melania. According to James such
trails are present in the Cincinnatian, but since there was
no accompanying figure, it is difficult to visualize what he
had in mind. Observation of the types of R. swbangulatum
(Pl. 59, fig. 3) and R. clavatum (PI. 67, fig. 5) shows that
they are almost certainly of arthropod origin. James also
dismissed R. asperum Miller and Dyer (1878) as the trail
of an annelid. Although it is not clear what this latter form
represents, it is certainly not a Rusophycus and is discussed
under 7'ylichnus.

A restudy of Hall’s types necessitates some rearrange-
ment of James’ nomenclatural assignments. Although the
type specimen of R. bilobatum has not been located, topo-
type material and the original description indicate that this
species is not present in the Cincinnatian. R. bilobatum is
a large (15 cm long) form with extremely coarse striae
which actually are better termed rugae. Here again it is
difficult to determine what James envisioned when he dis-
cussed Cincinnatian forms of R. bilobatum. He may have
been thinking either of large isotelid-produced specimens or
perhaps of much smaller nonisotelid forms in which the
median furrow extends nearly the entire length of the
specimen. Hall had mentioned this latter feature in his
original description. In R. bilobatum the median furrow ex-
tends the length of the specimen, while in R. pudicwm it is
confined to the central area.

It is concluded that there are three species of Ruso-
phycus present in the local section. These are R. pudicum

Hall (1852), R. carleyi J. F. James (1885), and R. crypto-

7 James was of the opinion that Cruiziana d’Orbigny had priority over
Rusophycus Vanuxem and employed the former name.

lithi, n. sp. Justification for these assignments will be given
below.
Rusophycus pudicum Hall

Plate 57, figure 6; Plate 58, figures 3-5, 7, 9, 10; Plate 59, figure 5;
Plate 60, figure 2; Plate 66, figure 3; Plate 81, figure 9, Text-figure
29-a.

1852. Rusophycus pudicus Hall, Nat. Hist. New York, Palaeont. New
York, vol. 2, p. 24, pl. 8, figs. 6a,b.

1885. ?Cruziana biloba James, J. F., Cincinnati Soc. Nat. Hist., Jour.,
vol. 7, pp. 154-157.

1885. ?Cruziana pudica (Hall), James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, pp. 154-157.

Type locality — Clinton (Silurian) beds of New York
State. Type specimens: Pl. 58, fig. 7; USNM 41113; latex
mold UCM 37581; Pl. 59, fig. 5, USNM 41115.

Diagnosis — Medium-sized bilobed Cubichnia of trilo-
bites (probably calymenids). The median furrow is only
moderately expanded and the striae may be weak or well
defined,

Discussion — Hall figured two specimens of R. pudi-
cum. The first (Pl. 58, fig. 7), measuring 3.2 cm by 2.0 cm,
is strongly bilobate, and the median burrow runs nearly the
entire length of the specimen. Striae are present on the
lobes but are only weakly developed. The second specimen
(PI. 59, fig. 5) is similar to the first in dimensions (3.9 cm
by 1.8 cm) but differs in that the median furrow is less
well developed and is confined to the central portion of the
specimen, Both specimens are preserved as convex hypo-
reliefs on a coarse-grained quartz arenite, the bottom of
which is covered by a thin micaceous film, apparently in-
dicating that the burrow was excavated in mud.

All Cincinnation representatives of R. pudicum occur
as convex hyporeliefs in sediments ranging from fine silt
to coarse sand. The species is relatively common and ranges
throughout the section. Although it does not normally occur
in association with other trace fossils, it is not unusual to
find more than one specimen of R. pudicum on a given
slab, and occasionally it may occur in great numbers (PI.
58, fig. 9). Morphologically it is difficult to distinguish
Hall’s specimens from the majority of those found in the
Cincinnatian. Measurements of 33 Cincinnatian specimens
reveal that the dimensions (mean length of 3.46 cm, mean
width of 1.65 cm) compare favorably with those of Hall’s
types. Although the lobes vary in their development, nor-
mally they are widest anteriorly and taper gently to the
rear, This is in contrast to some forms of Rusophycus, e.g.,
Seilacher (1955, fig. 5, nos. 4, 5), where the lobes are con-
stant in width or may even taper slightly anteriorly. The
median furrow extends the entire length of the lobes and
does not show any marked expansion. The striae covering
the lobes are normally poorly developed just as they are in

306 PALAEONTOGRAPHICA AMERICANA (VI, 41)

the syntypes. In some forms (PI. 60, fig. 2) they appear to
be unevenly concentrated over the lobes, thus giving the
lobes an almost annulate appearance.

Interpretation — There can be little doubt that the
producer of the Cincinnatian R. pudicum was Flexicaly-
mene. The general size and shape of the trilobite conform to
the morphology of the burrow, and impressions of the
cephalic doublure and thoracic pleurae are sometimes en-
countered (Pl. 81, fig. 9), As mentioned previously, three
specimens of R. pudicum have been found with the speci-
men of Flexicalymene meeki still in place. It is indeed rare
when a trace fossil is found in direct association with the
organism that produced it, and these are the first confirmed
occurrences of a trilobite found with a Rusophycus-Cruziana
trace.®

As is true of many Cubichnia, R. pudicum not infre-
quently exhibits horizontal repetition (1.e., repetition of the
burrow in a horizontal plane). In the example seen on
Plate 66, figure 3, the trilobite burrowed, moved forward a
short distance, and then burrowed again. Specimens such
as this are similar to Cruziana but there is a basic behavioral
difference. The surface of the long bandlike furrows of
Cruziana is always even, demonstrating that the organism
was moving horizontally across the substrate. In contrast
the specimen of Rusophycus under discussion shows an
alternation of vertical and horizontal movement.

R. pudicum, as well as other Cincinnatian representa-
tives of the genus, appear to confirm Seilacher’s statement
(1955) that tracks are rarely associated with Rusophycus.
The reason for this is not clear. Although Seilacher believed
that most Rusophycus were of endogenous origin, no such
examples have been found in the local section. The lack of
tracks is probably due to a variety of factors. Some speci-
mens of Rusophycus may well be of endogenous origin,
however, it is also likely that the currents sweeping the
bottom were of sufficient velocity to eradicate the tracks,
yet were not strong enough to obliterate the burrow. It is
possible that some examples of Rusophycus are the bur-
rows of nektonic trilobites, where the organism settled to
the bottom, burrowed, and then swam out of the excavation.

It is probable that Hall’s syntypes were made by caly-
menids, for Fisher (1960) listed both Calymene magarensis
Hall and C. clintoni (Vanuxem) as index fossils for the
New York Clinton. Presumably European calymenids, ¢.g.,

§
GC. blumenbachii Green, would leave similar traces,

* Abel (1935, fig. 224) illustrated a form from the Upper Devonian

sandstones of South Africa in which the striae are faithfully pre-
served. He believed that removal of the matrix would reveal the
producer im situ.

Rusophycus carleyi (J. F. James)

Plate 57, figure 1; Plate 58, figures 6, 8; Plate 59, figure 6; Plate 71,
figure 1; Text-figure 29-b.

1885. Cruziana carleyi James, J. F., Cincinnati Soc. Nat. Hist., Jour.,
vol. 7, p. 155, pl. 8, fig. 1.

Type locality —“Cincinnatian Group near Bantam,
Clermont County, Ohio” (James, 1885, p. 155). Presumed
type specimen: PI. 71, fig. 1, UCM 37604.

Diagnosis — Large bilobate Cubichnia of trilobites. In
some specimens the lobes part mesially to reveal two longi-
tudinal series of nodes.

Discussion — James (1885, p. 155) proposed R. carleyi
for specimens which differ from all other Rusophycus “in
being entire at one end and separated into two lobes at the
other; and also, and especially, in having seven or more
pairs of elevations and depressions arranged in an oval in
the center of the longitudinal groove.” He interpreted it as
an isotelid burrow.

What is most likely the holotype is now in the Univer-
sity of Cincinnati collections (PI. 71, fig. 1). Although the
specimen bears no label, the dimensions are similar to those
given by James. He also mentioned a sudden “flaring out”
just anterior to the median furrow. This is clearly shown in
his figure (James, 1885, pl. 8, fig. 1) and is present on the
specimen in question. The presumed holotype measures 13
cm by 7.5 cm and the depth is approximately 3.5 cm. The
median furrow is 2.0 em wide in R. carleyt, and within this
area are the small ridges or nodes mentioned by James.
Close observation reveals that there are nine such pairs.
James neglected to mention one additional feature which
differentiates this species from all others — the over-all
shape. R. carleyi is ellipsoidal, while most other species have
a cordate or slightly lacrimal outline. Normally such an
outline would make orientation difficult, but the cast of
one of the genal spines eliminates this problem. With the
exception of the extreme anterior area, the lobes are re-
markably free of striations.

Additional specimens of the same species are even
more revealing. One form (PI. 57, fig. 1) from the rich trace
fossil locality at Stonelick Creek (Corryville beds) shows
the cast of 11 pairs of walking legs, the hypostome, genal
spines, and cephalic and pygidial doublures! Were it not
for the presence of the poorly developed lateral lobes, this
specimen could easily be mistaken for an actual cast, 7..,
a body fossil. (Compare this form with /sotelus maximus
(Locke) (Pl. 57, fig. 5; USNM 33458) from the Richmond
beds near Oxford, Ohio, where the actual appendages are
present. )

The morphology of the Corryville specimens is similar

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 307

to Rusophycus sp. illustrated by Radwanski and Roniewicz
(1963) from the Upper Cambrian of the Holy Cross Moun-
tains of Poland. Although the Polish forms are smaller (3-7
cm long), the lobes are weakly developed, and the series of
nodes are present in the median area. Since /sotelus is un-
known in the Upper Cambrian of Poland, the producer most
likely was a different genus.

Also included in R. carleyi are a few large specimens
from unknown localities where the lobes do not part mesial-
ly. Plate 59, figure 6 illustrates one form which has a length
of 25 cm. This is the largest Rusophycus ever reported and
approaches the upper limits of isotelid dimensions. In con-
trast to the other members of the species, the lobes bear
extremely strong striae. The casts of one of the genal spines
and the thoracic pleurae are also preserved.

Interpretation — Because of their large size it is ap-
parent that the Cincinnatian representatives of R. carleyi
are the products of the burrowing activity of /sotelus, the
only local trilobite large enough to produce such a burrow.
The series of nodes seen in the specimens represent casts of
the proximal portions of the walking legs, probably the
coxae or coxal gnathobasic extensions. It is interesting to
note that these impressions on the Stonelick specimen ( PI.
57, fig. 1) differ somewhat from the coxae of the Oxford
specimen of J. maximus. Both the specimen and Raymond's
(1920, fig. 9) reconstruction of it show exceptionally long
coxae. The coxae of the Stonelick specimen are much
shorter, and in the posterior portion of the specimen, there
is a V-shaped joint. Although these impressions are dis-
torted by the movement of the legs, the joint which moved
was shorter than the gnathobasic appendages shown by
Raymond. Because the Oxford specimen is composed of
coarse crystalline calcite (there is no original skeletal ma-
terial remaining), it is entirely possible that a joint could
have been masked during replacement.

Those forms bearing the nodes appear to illustrate
a type of digging which differs from that found in most
Rusophycus (e.g., R. pudicum) where nodes are not
present. Excavation of the burrow might have been achieved
by a broad sweeping motion of the appendages much like
a swimmer doing the breaststroke. This would account for
the weak development of the lobes and striae and would
also make possible preservation of some of the features of
the ventral surface. It is equally possible that the digging
movement was similar to that in R. pudicum but that some
of the sediment became clogged along the mid-ventral axis.

Rusophycus cryptolithi, n. sp.
Plate 58, figures 1, 2; Plate 59, figure 4; Text-figure 29-c.
Type locality —The specimen figured on Plate 58,

figure 2; UCM 37587 is designated as the type specimen.
The type locality is the Economy beds of Humphrey’s
Branch of Twelve Mile Creek on Washington Trace Road,
350 meters southeast of the intersection with Twelve Mile
Creek Road, Campbell County, Kentucky.

Diagnosis — Small, ovoid, button-like Rusophycus. The
striated lobes are subdivided into a coarser outer zone and
cordate more finely striated inner zone. Imprints of genal
spines are commonly present. This species differs from all
others in the genus by having a width which is equal to or
slightly greater than the length.

Discussion — Vo date 15 specimens of the species are
known, 11 are from the type locality, three are from the
Economy strata of Duck Creek, 1,000 meters southwest of
the intersection of Duck Creek Road and Kentucky Route
8, Campbell County, Kentucky. The remaining specimen is
from the McMicken beds of West Fork Creek, approximate-
ly 30 meters upstream from the ?mollusk trails locality. All
are preserved as convex hyporeliefs in a 2 cm thick cross-
laminated calcisiltite which overlies a thick calcilutite, In
contrast to other Cincinnatian Rusophycus, R. cryptolithi
shows positive rheotaxic orientation,

These button-like specimens average 17 mm in width
and 15 mm in length. A slightly raised rim commonly sub-
scribing an arc of 180° is preserved on some forms. A
median furrow is present in all forms but is less well de-
veloped than in the other two species. The morphology
varies with the depth of the burrow. The deeper forms,
such as those seen in Plate 58, figures 1, 2, exhibit the great-
est detail. In these two forms the anterior wall of the bur-
row is slightly undercut, and the genal spines are plainly
evident. In contrast shallower “incipient” burrows, such
as that on the right in Plate 59, figure 4, show only the two
lobes.

As the specific name implies R. cryptolithi represents
the burrowing activities of Cryptolithus. The ovoid shape,
size range, and the imprints of the genal spines all point to
this genus. The rocks in which the specimens were found
abound in Cryptolithus fragments, while other trilobites
are comparatively rare. Only two small specimens of KR.
pudicum [probably made by Flexicalymene callicephala
(Green)] were found at the first two localities mentioned
above.

Traditionally the blind Cryptolithus has been considered
a burrower, yet it appears that it was not so successful in
excavating resting traces as other trilobites were. The oval
burrow would serve only to conceal the cephalic area, for
in some of the burrows where the anterior wall is much
deeper than the posterior, the thorax and genal spines must

308 PALAEONTOGRAPHICA AMERICANA (VI, 41)

certainly have been exposed. Likewise, the undercut an-
terior wall must have presented some problems when the
organism desired to leave the burrow. Perhaps the cordate
inner area of the lobes was made as the animal backed out
of the burrow, for it is best developed on those forms in
which the wall is undercut.

LOCKEIA U. P. James, 1879

Plate 57, figure 7; Plate 58, figure 10; Plate 59, figures 1, 7; Plate
63, figure 6; Text-figures 6, 29-d.

1873. Non Dawsonia Nicholson, Monograph of British Graptolitidae,
p. 70.

1879. Lockeia James, U. P., The Paleontologist, vol. 1, p. 17.

1885. Daasonia Nicholson, James, J. F., Cincinnati Soc. Nat. Hist.,
Jour., vol. 7, pp. 161, 162, pl. 9, fig. 7.

1892. Dawsonia Nicholson, James, J. F., Cincinnati Soc. Nat. Hist.,
Jour., vol. 14, pp. 162, 163, fig. 7.

1953. Pelecypodichnus Seilacher, Neues Jahrb. Geologie u. Palaont.,
Abh., Bd. 98, pp. 105-107, pl. 10, fig. 1; pl. 12, figs. 1, 2.

1962. Pelecypodichnus Seilacher, Hantzschel, Trace-Fossils and Prob-
lematica iz Treatise on Invertebrate Paleont., Moore (ed.), pt.
W, Miscellanea, p. W208, fig. 130-4.

Type species —Lockeia stliquaria U. P. James (1879)
from the bank of the Ohio River “halfway between high
and low water” at Ludlow, Kentucky (most probably Up-
per Trenton).

Diagnosis — Small oblong bodies pointed at both ends,
which resemble the end of an almond projecting above a
surface. Preservation as convex hyporelief. Surface of speci-
mens normally smooth, although a longitudinal crest or
furrow may be present in some. Vertical sectioning reveals
a shaft of disturbed bedding conforming to the measure-
ments of the specimen overlying most forms.

Discussion —U. P. James (1879, p. 17) proposed
Lockeia siliquaria for “elongated, convex fossil sea-plants,
rounding and tapering to sharp or obtuse points, seed-like
in appearance, with or without slight longitudinal depres-
sions, attached to the surface of rocks.”

Regretably the type locality is now permanently
covered by damming of the Ohio River, and recollecting is
no longer possible. Fortunately, several slabs of topotype
material (PI. 57, fig. 7; UCM 37597) are present in the
University of Cincinnati collections. Additional material in-
cludes a single slab bearing about 15 specimens from the
Economy beds of Twelve Mile Creek, Kentucky (PI. 59, fig.
7), and a single slab with three specimens (PI. 58, fig. 10)
from the Corryville strata of Stonelick Creek.

As shown by the association with Rusophycus and as-
sorted tool marks (PI. 58, fig. 10), Lockeia is preserved as
a convex hyporelief on calcisiltites. Specimens from the topo-
type locality show a high “population density” as many as

215 individuals occur on a surface of 150 square cm! In-
dividuals may occur singly or in groups of three or four, and
truncation is not uncommon. There is no preferred orienta-
tion, although up to six individuals may be aligned one
behind the other, thus presenting a sort of pseudo-preferred
orientation. Usually a faint ridge connects the aligned forms,
although it is not always present. Plate 57, figure 7 shows
a variation of the above, where several specimens are ar-
ranged in an en-echelon pattern.

The individual specimens occur as oblong bodies which
are pointed at both ends. Forms from the topotype locality
occurring on the same slab show a wide size variation. The
length may vary from 2-9 mm, while the height ranges from
a fraction of a millimeter up to 4 mm; the width is 1-2 mm.
The surface of most specimens is smooth, although some
may bear a faint longitudinal ridge or furrow. While the
majority of the forms are symmetrical, a few are tilted
to one side or are slightly irregular in outline.

The Twelve Mile Creek specimens (PI. 59, fig. 7) are
larger than the topotype forms, as they average 1.2 cm in
length and 3 mm in width and height. Some of these are
irregular in outline, and many appear to have been broken
off, possibly by Recent stream erosion. The Stonelick Creek
specimens (PI. 58, fig. 10) resemble the topotype material in
their regular outline but are somewhat plumper.

Both longitudinal and cross-sections were prepared of
specimens from Twelve Mile Creek and the topotype local-
ity. These sections clearly reveal that there is a disturbance
of the bedding directly above the markings in the material
from the first locality. This disturbed area conforms to the
dimensions of the fessil and extends to the top of the bed.
It is noteworthy that the contiguous bedding is bent as if
something had been forced down through the sediment. One
section (PI. 63, fig. 6) shows what appears to be the outline
of a rounded object near the base of the bed. That the dis-
turbed bedding overlying the hyporeliefs is arranged in an
arcuate pattern is noteworthy.

Sectioning of the topotype material presents a more
complex picture. Of the 32 specimens of Lockeia that were
studied in this manner, 27 showed no disturbed bedding over
the specimen such as was found in those from Twelve Mile
Creek. The remaining five did show the disturbance. Where
there was no disturbed bedding, it was found that the silt
forming the almond-shaped bodies was both darker and
more coarse-grained than that of the overlying bed, and
the line of contact was exactly flush with the base of the
calcisiltite bed. This is best seen in vertical section (PI. 59,
fig. 1) but is also apparent from close observation on the
surface of the slab itself.

TRACE FOSSILS CINCINNATI AREA: Oscoop 309

J. F. James (1885, 1892) compared Lockeia to certain
forms from the Llandeilo beds (Ordovician) of Scotland
which were described by Nicholson (1873) under the name
Dawsonia. Because James’ comparison rested more on in-
terpretation than on a similarity of morphology, it will be
discussed later.

Lockeia is judged to be synonymous with Pelecypodich-
nus Seilacher (1953b) from the Braun-Jura Beta beds of
Wiirttemberg (Seilacher, 1953b, pl. 10, fig. 1 and pl. 12, fig.
1; plastotype UCM 37688). Seilacher’s forms are preserved
as convex hyporeliefs and are associated with Asteriacites
quinquefolias. Although their length is close to that of the
Cincinnatian topotype specimens, their width is consistently
greater (4 mm). In this sense they are more like the Corry-
ville specimens. As in the Cincinnatian topotype material,
individuals may occur in groups, and truncation is not in-
frequent. It is not known whether disturbed bedding is
found overlying the forms, as apparently Seilacher did not
have a sufficient number of specimens to permit sectioning.

Lastly, Lockeia resembles Linck’s (1949, pl. 8, figs. 1,
2) “Reihenhécker-Spur” from the Schilf Sandstone (Trias-
sic: Keuper) of Wiirttemberg. The most striking feature of
Linck’s forms is the degree of alignment found. The longest
observed specimen was 26 cm in length and consisted of 14
individuals connected by low ridges. Occasionally one form
will protrude out of line to break the symmetry. These too
are preserved as convex hyporeliefs and their size range
compares favorably with Lockeia.

Interpretation —J. F. James (1885) discarded his
father’s algal hypothesis and instead compared Lockeia to
“reproductive bodies” or “ovarian capsules” of graptolites
similar to those described by Nicholson (1866, 1873).
Nicholson (1873, p. 139) characterized Dawsonia as “horny
or chitinous capsules . . . furnished in most cases with a
small spine or mucro, and having a marginal filament exact-
ly resembling the solid axis of a graptolite.” James admitted
that local workers had questioned his interpretation because
Lockeia is not associated with graptolites. However, because
he erroneously believed that “Buthotrephis” gracilis Hall
represented a graptolite and because it occurs with Lockeva,
this seemed to him convincing. Although Dawsonia® is a
problematicum (Bulman (1955) in his graptolite mono-
graph, placed it under “unrecognizable genera”), Nicholson’s
description indicates that it is really different from Lockeia.
None of the Cincinnatian specimens possess an organic
covering, a “mucro,” or a “marginal filament.” Likewise,
there is no evidence to support James’ conclusion that one is

» Dawsonia Nicholson (1873) is preoccupied by Dawsonia Hartt in
Dawson (1868), an agnostid trilobite.

dealing with “ovarian capsules” or graptolites. As shown
elsewhere in this paper, “Buthotrephis” gracilis is not a
graptolite but a chondritid burrow. Moreover, James was
reasoning from the philosophy that graptolites were hydro-
zoans and thus would possess large reproductive bodies.
There is no evidence that they did. The disturbed bedding
overlying the specimens and the truncation of some forms
by others indicates that Lockeia is a trace fossil.

Neoichnological studies conducted by Seilacher (1953b)
and used by him to explain the origin of Pelecypodichnus
provide insight into the origin of Lockeia. Seilacher’s in-
terpretation is based on experiments conducted with Tivela
ponderosa (Koch), a burrowing marine pelecypod of the
Pakistan sand flats. Seilacher (1953b, pl. 12, fig. 3a) showed
a vertical section in which the anterior portion of the shell
is imbedded in mud, while the bulk of it is encased in the
overlying silt bed. His plate 12, figure 3b is a plaster cast of
the impression left in the mud after the pelecypod and the
overlying silt bed have been removed. This cast is morpho-
logically similar to both Pelecypodichnus and Lockeia.
Seilacher classified the former among the Cubichnia, as he
believed that the organism burrowed for protection and
then extended its siphons to the surface to feed. Although
he did not elaborate, he considered Linck’s forms as Pele-
cypodichnus and regarded them both to be Repichnia.

It is probable that Lockeia too was made by burrowing
pelecypods. Dalvé (1948) listed nine bivalves as being
present in the Fulton strata.” Four of these, Byssonchia
vera (Ulrich), Modiolopsis sp., Pterinea demissa (Conrad),
and Pterinea mucronata (Ulrich), are byssate pelecypods
and need not be considered here, Of the remaining five, two
forms, Cleidophora fabula (Hall) and Ctenodonta obliquata
(Hall), measure only a few millimeters in length and thus
are too small to have produced the burrow. The other three,
Orthodesma nasutum (Conrad), Cleidophorus planulatus
(Conrad), and Psiloconcha tenuistriata Ulrich, are larger
and have the relatively smooth elongated body indicative
of burrowers. Although the local representatives of Ortho-
desma nasutum (Conrad) have never been described,
Foerste (1914, pl. 3, fig. 5A, B) discussed coeval forms from
the Cincinnatian of New York State. One described speci-
men has a length of 44 cm, while the greatest height is
1.55 cm and the convexity is 4 mm. Foerste (1914, p. 287)
mentioned that “the most striking feature of the shell .. .

“Eyen though the type locality of Lockeia is most probably Trenton
in age, it is not invalid to include Lower Cincinnatian fossils in a
discussion of the probable originators of the topotype burrows. As
Weiss, ef al. (1965, p. 17) pointed out, the Cynthiana and Lower
Eden faunas intergrade.

310 PALAEONTOGRAPHICA AMERICANA (VI, 41)

is the distinct dropping of the cardinal margin anterior to
the beak.” Cleidophorus planulatus (Conrad) is listed by
Bassler (1915) and Shimer and Shrock (1944) as being
present in the local strata, but, like O. nasutwm, the Cin-
cinnati specimens are undescribed. Shimer and Shrock (1944,
p- 377; pl. 146, fig. 17) said that the species is “charac-
terized by prominent umbonal ridge extending from beak to
posterior extremity, and by conspicuous flattening of shell
hingeward from this ridge.” Foerste (1914, p. 302) men-
tioned that forms from New York State attained a length
of 2.0 cm. The greatest height is 9 mm and the convexity
16 mm. The third form, Psiloconcha tenuistrata, was
established by Ulrich (1893, p. 668; pl. 52, fig. 10) for
forms “near the low water mark of the Ohio River at
Coyington, Kentucky.” The specimens are small, having a
length of about 1.1 cm and a height of 4.5 mm. The dis-
tinguishing features are the “narrowly rounded anterior end,
and the exceeding fineness of the concentric surface mark-
ings” (Ulrich, 1893, p. 668).

While it must be admitted that there is no conclusive
proof that Lockeia represents the burrowing activity of
clams, the evidence is suggestive. Seilacher showed that
burrowing pelecypods such as Tivela can produce Lockeia-
like bodies, and there are three pelecypods present in the
Fulton strata which appear, from their shell configuration,
to have been burrowers. Moreover, the consistent symmetry
of most specimens of Lockeia indicates that they were made
by some rigid or semirigid body. [Several ostracods are
present in the Fulton rocks, but the ornamentation found
on most forms demonstrates that they were surface dwellers.
Those species with smooth carapaces are far too small to
have produced the structure. The only other known bur-
rowing form in the Fulton beds is Lingula, but as seen in
Moore, Lalicker, and Fischer (1952, fig. 6-1), the structure
of the burrow is different.] Thus it is concluded that
Lockeia most likely represents the burrowing activity of
pelecypods. Based on the different size of the specimens
from the three Cincinnatian localities, it is unlikely that all
the forms were made by one species. While Seilacher’s Pele-
cypodichnus is believed synonymous with Lockeia, not
enough is known about Linck’s forms to make an interpreta-
tion meaningful. Linck thought that they might be casts of
imprints of amphibian or fish pectoral appendages. Vertical
sectioning should be sufficient to test his theory.

It is unusual that the remains of the maker of Lockeia
are not found at the bottom of some of the structures, for
such a covering of silt greatly increases the changes of fos-
silization. However, the fact that Locketa occurs in small
clusters and shows horizontal repetition indicates that the
organism probably changed the location of its burrow from

time to time. Also, changes in the salinity or in the oxygen
content of the water will cause hemi-sessile forms to move
to the surface (personal observation made at Pacific Marine
Station, Dillon Beach, California, during the summer of
1962). Moreover, the fact that the Fulton pelecypods do
not have a well-developed pallial sinus indicates that they
did not possess long retractile siphons. This means that the
posterior end of the shell was probably located at or just
below the depositional interface. In time of rapid sedimen-
tation the organism would be forced to elevate its burrow
in order to maintain this relationship. The arcuate nature
of the disturbed bedding in some of the Twelve Mile Creek
specimens does suggest that the animal moved up in the
burrow.

Most probably some of the almond-shaped bodies
represent casts of the foot, while others are casts of the
shell itself. Normally when a pelecypod burrows, the foot
is out ahead of the shell, but the longitudinal ridges and
depressions found on some specimens lead one to believe
that these specimens are casts of the shell.

A more complete analysis of the topotype material is
now possible. As mentioned above, 27 of the 32 forms
studied do not penetrate up into the silt bed bearing them.
The remaining five show clear evidence of extending from
the base of the bed to the top, The following sequence of
events is hypothesized to explain these facts and is illus-
trated in Text-figure 6. First, several pelecypods burrowed
through the silt until they encountered the underlying mud"
(Text fig. 6A). At this point the organism withdrew, and
the overlying silt filled in the depression. The silt bed was
then completely stripped off by current action down to
(but not into) the surface of the mud (Text-fig. 6B). This
is not as implausible as it may seem at first, for once deposi-
tion has taken place, a great deal more energy is required to
move mud than is necessary for the erosion of silt (Hjul-
strom, 1939). If there were no increase in current velocity, a
given current might be able to move silt yet be incapable
of eroding mud. Because the molds of the original burrows
were already filled with silt and were below the surface of
the mud, they were not obliterated. Following this erosive
period, another silt bed, slightly finer grained and lighter in
color than the previous one, was deposited over the mud.
Finally, a few pelecypods burrowed through this silt and
nicked the surface of the mud (Text-fig. 6C). They then
withdrew, the overlying silt filled in the depression, and the
disturbed bedding remained as evidence of their activity.
During diagenesis the silt bed underwent cementation, and

“Many specimens have trapped lutite fragments along their margins.

TRACE FOSSILS CINCINNATI AREA: OsGoop 3

Text-figure 6.—Interpretation of Lockeia siliquaria. Recon-
struction showing the origin of specimens from the type locality. A.
Three pelecypods burrow through silt (dark pattern) and just nick the
subjacent mud (dashed lines) with their foot or the anterior portion
of the valves. B. Subaqueous erosion strips away the silt down to the
silt-mud interface. The dark ovoid imprints of the valves and foot or
foot have been filled with silt and are protected. C. Following de-
position of more silt (dark pattern) the sequence of events described
in A above is repeated. Observation of the mud surface now reveals
six ovoid imprints, however, cross-sections show that only three are
overlain by a disturbed column of silt.

11

312 PALAEONTOGRAPHICA AMERICANA (VI, 41)

the casts of the first group of burrowing pelecypods were
cemented to the bottom of the new silt bed. However, there
is a difference in the strength of bonding, for J. F. James
(1885, p. 162) remarked that many of the Lockeia speci-
mens could be easily removed from the rock, much like
grains of wheat. This indicates that the bonding of the
original casts to the siltstone was weak. On the other hand,
the later casts, which from an integral part of the new silt-
stone bed, are not so easily removed; here cementation was

more perfect.

ASTERIACITES Schlotheim, 1820

Plate 57, figure 2; Plate 58, figure 2; Plate 62, figure 5; Text-figure
29-e.

1820. Asteriacites Schlotheim, Die Petrefactenkunde auf ihrem jetzi-
gen Standpunkte durch die Beschreibung seiner Sammlung
versteinerter und fossiler Uberreste des Thier—und Pflanzen-
reichs der Vorwelt erlautert, p. 324.

1878. Hceliophycus Miller and Dyer, Contributions to Palaeontology
No. 2 (Cincinnati, Ohio, private publication), p. 2, pl. 3. fig. 3.

1885. Heliophycus Miller and Dyer, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 163.

1953. Asteriacites Schlotheim, Seilacher, Neues Jahrb. Geologie u.
Palaont., Abh., Bd. 98, pp. 93-105, pls. 7-10.

1962. Asteriacites Schlotheim, Hantzschel, Trace Fossils and Prob-
lematica in Treatise on Invertebrate Paleont., Moore (ed.), pt.
W, Miscellanea, p. W184, figs. 110-2, 130-4.

Schlotheim
(1820) from Lias of Coburg, Germany. Lectotype, Seilacher
(1953b).

Diagnosis —Stellate Cubichnia of starfish which may
be preserved as both concave epireliefs and convex hypo-
reliefs. Vertical repetition is not uncommon and some evi-
dence of the activity of tube feet may be present.

Type species — Asteriacites lumbricalis

Discussion — Seilacher (1953b), who monographed the
genus, recognized two species of Asteriacites: A. lumbricalis
Schlotheim (1820) and A. quinquefolius Quenstedt (1876).
The former is known from several localities, ranging in age
from Upper Silurian through the Dogger. When preserved
as a convex hyporelief, it consists of a small well-defined
central disc and five long thin arms which may be striate.
In some examples (Seilacher, 1953b, fig. 2-2b) the dise may
be poorly defined and the proximal portion of the arms
greatly expanded. When A. /wmbricalis is found as a concave
epirelief, the morphology is far different (Seilacher, 1953b,
fig. 2-1b) — the central disc is indistinct and the arms
taper gradually. The delicate striae which mark the arms in
the convex hyporeliefs are not evident.

A. quinquefolius Quenstedt (1876) is known only from
two specimens, both from the Dogger Beta Sandstone near
Ziirich. One form is preserved as both a concave epirelief
and a convex hyporelief, the other only as a concave epi-

relief. The convex hyporelief differs from A. lwmbricalis in
being somewhat larger and having a shaggy appearance.
Likewise, the central disc is not as well defined. The con-
cave epireliefs have the same stellate outline as the convex
hyporelief, but the details are blurred. As Seilacher noted, it
is difficult to distinquish the two species solely from the
concave epireliefs (compare Seilacher, 1953b, fig. 2-la, b
with pl. 10, fig. 2 and pl. 11, fig. 3).

Seilacher has undertaken an extensive study of the
literature and has worked out a detailed synonymy, especial-
ly for A. lumbricalis. No attempt will be made here to judge
his conclusions on this matter.

Interpretation — Asteriacites has most frequently been
regarded as a cast of the ventral surface of asteroids. This
was the interpretation of Schlotheim (1820), and many
workers have since echoed this belief. Although Richter
(1927, pl. 1, fig. 3) concluded that asteriacitid-like forms
from the Ordovician of Bohemia were the fillings of non-
echinoderm stellate burrows, it is not possible to support his
interpretation on the basis of his illustration.

The major problem is to determine whether a given
form is a trace fossil or body fossil. Seilacher listed several
criteria in support of his trace fossils. In the specimens he
studied there was never a vestige of organic material and
neighboring individuals may intersect. Moreover, the striae
marking the arms do not resemble a plate pattern but
instead show the imprints made by the tube feet during
digging. The most convincing evidence is provided by
vertical repetition. Seilacher (1953b, fig. 3; 1964, fig. 3)
illustrated a slab from the Lower Triassic of the Tyrol
which demonstrates the reaction of eight individuals to
rapid sedimentation. As sand washed over the organisms,
they struggled to reach the surface, and their upward path
may be traced on the cleavage planes (hence the origin of
the term vertical repetition). Another type of vertical repe-
tition involves a rotation of the body during digging (Seil-
acher, 1953b, pl. 7, fig. 2). Such behavior results in several
partial imprints.

Aquarium studies conducted by Seilacher employing
an asteroid and ophiuroid have been most illuminating. A
mud surface was covered with sand and the organisms were
allowed to burrow. The results show that ophiuroids or
ophiuroid-like forms may leave imprints which closely re-
semble those of asteroids (see Seilacher, 1953b, fig. 2). Like-
wise, by moving the distal portion of the arms, an ophiuroid
with unbranched arms can leave an impression which could
erroneously lead one to select a form with branching arms
(e.g., Euryale) as the probable originator of the trace.

Seilacher reserved A. lumbricalis for traces made by

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 31.

ophiuroids or ophiuroid-like stelleroids and A. quinquefolius
for traces of asteroids. However, based on the results of
his aquarium studies, it is apparent that one must exercise
extreme caution in making specific assignments.

He placed Asteriacites within the Cubichnia, basing
his conclusion on the known fact that many Recent stel-
leroids burrow just deep enough to cover the body with a
thin layer of sediment. The most obvious explanation for
this behavior is that the shallow burrow serves as a tem-
porary place of refuge.

CINCINNATIAN REPRESENTATIVES
Asteriacites stelliforme (Miller and Dyer)
Plate 57, figure 1; Plate 59, figure 2; Plate 62, figure 5.

1878. Heliophycus stelliforme Miller and Dyer, Contributions to
Palaeontology No. 2 (Cincinnati, Ohio, private publication),
p. 2, pl. 2, fig. 3.

1885. Heliophycus stelliforme Miller and Dyer, James, J. F., Cincin-
nati Soc. Nat. Hist., Jour., vol. 7, p. 163.

1953. Partim Asteriacites lumbricalis Schlotheim, Seilacher, Neues
Jahrb. Geologei u. Palaont., Abh., Bd. 98, p. 95.

Type locality —... “about the middle of the Cincinnati
Group” (probably Fairview? ), Cincinnati, Ohio.

Diagnosis —Cubichnia of moderate-sized stelleroids,
the radii of which bear fine chevron-shaped striae.

Discussion. — Only three specimens are available for
study — the type (PI. 59, fig. 3; HBM 3175), a specimen
from the Corryville of Stonelick Creek (PI. 57, fig. 2), and
a third form from an unknown locality and horizon (PI.
62, fig. 5). All are preserved as convex hyporeliefs in calci-
siltites. There is no trace of the associated concave epirelief
preservation found in Seilacher’s forms. The dimensions of
the forms are identical; the length of each arm is 2.0 cm,
while the greatest width is 8 mm. The most noteworthy
feature is the presence of the “rugosites” on the arms men-
tioned by Miller and Dyer. In the type specimen (PI. 59,
fig. 2) and the Corryville specimen (PI. 57, fig. 2), these
striae form a “V,” the apex of which is directed proximally.
In the other specimen (PI. 62, fig. 5) there is no well-
developed “V,” and instead the striae are nearly normal
to the main axis of the arm. Such sculpturing is also present
in some of Seilacher’s forms and the orientation of the “V”
is the same in both cases. The Corryville specimen shows
clear evidence of the vertical repetition noted by Seilacher
in some of the European forms. The other two specimens
have a simple stellate outline.

Although Seilacher (1953b) placed A. stelliforme in
synonymy with A. lumbricalis, it is believed that there is
sufficient basis for retaining Miller and Dyer’s specific

we

designation. A. stelliforme differs from A. lumbricalis in
having wider arms and chevron-shaped striae and from A.
quinquefolius in having thinner arms and lacking the shaggy
appearance.

Interpretation — Miller and Dyer (1878b, p. 2) be-
lieved that A. stelliforme was a“... starlike plant without
any stem,” while J. F. James (1885, p. 163) interpreted the
form as the mold of an asteroid, possibly Palaeaster. He ex-
plained the “rugosities” on the arms as “caused by arrange-
ment of the plates on the underside of the body, but the
surface of the mud not taking the impression perfectly and
the plates were not preserved.”

Both Seilacher (1953b) and Hantzschel (1962) con-
sidered Heliophycus to be the junior synonym of Asteriacites
and considered it a trace fossil.

Although sometimes it may be difficult to distinguish
Asteriacites from bona fide ventral casts of body fossils,
such is not the case here. All three specimens clearly show
the activity of the tube feet. The striae are analogous but
not directly comparable to the V-shaped striae found on the
lobes of Rusophycus. In the latter the digging motion was
directed toward the median line and to the rear. It is prob-
able that in A. stelliforme digging was directed away from
the median line. If the opposite were true, it is difficult to
visualize how the organism could burrow at all, for the sedi-
ment would soon become clogged in the proximal region of
the arms. The imprints seen on the specimen most likely
reflect the final movement of the tube feet as the animal
left the burrow. Further support for the trace fossil assign-
ment of A. stelliforme is offered by the vertical repetition
seen in the Corryville specimen,

In conclusion, A. stelliforme is interpreted as the resting
trace of an asteroid, possibly Promopalaeaster Schuchert
(1914), which is a Maysville and Richmond form and is of
the same general size and shape as A. stelliforme. As
Seilacher surmised, it is most probable that Asteriacites falls
within the Cubichnia and is the stelleroid analogue of
Rusophycus.

VI. DOMICHNIA
INTRODUCTION TO SPREITE-BEARING U-TUBES

One of the more common marine burrows (domiciles )
from the Cambrian to Recent has a U-shape in profile.
Some show a transverse zone of disturbed sediment (the
Spreite) between the limbs of the “U,”’ whereas others do
not. The makers of the Spreite-bearing U-tubes here under
discussion are probably responsible for most, if not all, of
the enigmatic “dumbbell” markings of ancient sediments.

314 PALAEoNnToGRAPHICA AMERICANA (YI, 41)

Before undertaking a discussion of U-tubes, it is neces-
sary to define certain terms, for although U-tubes have
received a great deal of study, there has been little uni-
formity of terminology. Most of the terms discussed below
are illustrated in Text-figure 7.

arms —the lateral cylindrical portions or limbs of the SU

aperture —the openings of the “U” on the depositional interface.

U-plane section—a vertical section which passes through the
“U” (Text-fig. 7A).

median section —a vertical section taken at right angles to a U-
plane section at a point midway between the apertures (Text-fig.
7B-C).

transverse section —a section taken parallel to the bedding (Text-
fig. 7D), i.e., normal to the U-plane and median section.

depth —in complete forms, the distance from the depositional
interface to the base of the “U.” In incomplete forms, where the
upper part of the tube is missing, the depth is the distance from
the highest point of the “U” (as seen in U-plane section) to the base
of the “U.”

length —the distance as measured from the outside of one arm
to the outside of the other arm.

breadth —a measurement taken across the Sfreite normal to the
length. In some European forms the breadth increases with depth
(see Text-fig. 7D).

Spreite — arcuate relics of U-tube excavation. As seen in U-plane
section Sfreite appear as a series of ares connecting the arms of
the “U” and generally parallel to the base of the “U.”

protrusive Spreite (Goldring, 1962) —a series of ares indicating
a deepening of the tube. As viewed in a median section (Text-fig.
7B), the breadth of the Spreite may increase with depth and the
arcs are convex upward. As viewed in a transverse section (Text-
fig. 7D1-3), the arcs have their convex face directed away from the
arms of the “U.” As seen in U-plane section (Text-fig. 7A), the
arcs extend only to the immer margin of the arms.

retrusive Spreite (Goldring, 1962) —a series of ares which in-
dicate an elevation of the bottom of the “U.” As viewed in a median
section (Text-fig. 7C, dashed lines), the arcs are convex down-
ward. As viewed in a transverse section (Text-fig. 7D-+), the ares
have their convex face directed foward the arms of the “U.” As
seen in U-plane section, the ares extend to the outer margin of the
arms. Text-figure 7A,C shows retrusive Sfreite superimposed over
protrusive Spreite.

free tube —the portion of the arm as measured from the aperture
to the first evidence of a Spreite.

COROPHIOIDES John Smith, 1893

Plate 60, figures 1-6, 8; Plate 61, figures 1-8; Plate 62, figures 1-4, 6;

Plate 66, figure 5; Plate 69, figure 3; Plate 70, figure 5; Plate 71,
figure 5; Plate 77, figure 8; Text-figures 8, 9, 29-0,p,g.

1874. ?4rthraria Billings, Canadian Natur., n.s., vol. 6, p. 467, fig. 2.

1875. Arthraria Billings, Miller, Cincinnati Quart. Jour. Sci., vol. 2,
p. 354, fig. 26.

1893. Corophioides Smith, Geol. Soc. Glasgow, Trans., vol. 9, pp. 289-
292, pl. 10.

1916. Arenicoloides Blanckenhorn, Ges. z. Beférderg. ges. Naturwiss.
Marburg, 1916, Sitz. Ber. pp. 21-43.

1921. Arenicoloides Blanckenhorn, Kolesch, Preussen Geol. Landes.
Anst., Jb., Bd. 40, Teil III (for 1919), pp. 356-366, figs. 8-15.

1924. Corophioides Smith, Blanckenhorn, Preussen Geol. Landes.
Anst., Jb., Bd. 44 (for 1923), p. 31.

1924. Corophioides Smith, Richter Rud., Senckenbergiana, Bd. 6, pp.
119-165.

1924. Arenicoloides Blanckenhorn, Sérgel, Neues Jahrb. Geologie u.
Palaont., Beil-Bl. 44, pp. 510-549, figs. 1-7, pls. 19-20.

1926. Corophioides Smith, Richter, Rud., Senckenbergiana, Bd. 8, pp.
185-219, pl. 3.

1927. Arenicoloides Blanckenhorn, Andree, Palaontol. Zeitschr., Bd.
8, pp. 120-128.

1929. Corophioides Smith, Opik, Acta Comment. Univers. Tartuensis,
A., Vi 2 pp. 31238, figs: 3)4:

1931. Corophioides Smith, Westergard, Sverige geol. Unders., Ser.
C, Arh. och Upps., No. 372 (=Arsbok 25, No. 5), pp. 3-9.

1935. Corophioides Smith, Abel, Vorzeitliche Lebensspuren, pp. 452-
456, figs. 378-379.

1940. ?Bifungites Desio, Museo. Libico Storia Naturele, vol. 2, p. 78,
pl. 8, fics 3:

1955. Corophioides Smith, Seilacher, in Schindewolf, and Seilacher,
Akad. Wiss. Lit. Mainz, Math.-nat. KI., Abh., No. 10, p. 377.

1962. Partim Arthraria Billings, Hantzschel, Trace-Fossils and Prob-
lematica im Treatise on Invertebrate Paleont., Moore (ed.), pt.
W, Miscellanea, p. W184.

1962. Corophiotdes Smith, Hantzschel, Trace-Fossils and Problematica
im Treatise on Invertebrate Paleont., Moore (ed.), pt. W,
Miscellanea, p. W189, figs. 117-1, 2.

1963. Corophioides Smith, Seilacher, Fortschr. Rheinland und West-
falia, Bd. 10, pp. 86, 87, fig. 3.

1964. ?Bifungites Desio, Dubois and Lessertisseur, Soc. Geol. de
France, Bull., (7) vol. VI, pp. 626-634, figs. 1-3, 6, 7, pl. 1.

Type species — Corophioides polyupsilon John Smith
(1893) from the Upper Carboniferous sandstones of the
Gowkha Quarrry near Kilwinning, Scotland.

Diagnosis — Vertical U-tubes with Spreite. The arms
of the “U” are parallel or diverge with depth. Spreiten are
mainly of the protrusive variety, although forms with re-
trusive Spreiten have been reported. The walls of the arms
are without striations; the apertures of the tube are not
flared into funnels.

Discussion — The genus Corophioides was established
by John Smith (1893, p. 291) for structures which begin “as
a simple U-shaped tube of about 1% inches wide by the
same in height, the animal appears to have enlarged its
tube or habitation to accommodate its increasing circum-
stances till it reached as much as 4 inches wide and five
inches high, these being the largest measurements I have
taken. Between these extremes they are to be met with all
sizes. . . . They resemble a lot of distorted and crushed up
U’s of different sizes, placed one within the other.” Smith’s
figure of the form (pl. 10, top) indeed shows that he visual-
ized Corophioides to be a series of several complete “U’s”
placed one inside the other. Smith noted that some of the
sandstone beds contain several such forms. Judging from
his diagram (Smith, 1893, pl. 10, bottom), they occur in the
?basal sands associated with cyclothems. Through the kind-
ness of Dr. W. D. I. Rolfe of the Hunterian Museum in
Glasgow, the type specimen (PI. 60, fig. 6; HM X.59) was
made available for study. The host rock is a poorly sorted,
coarse-grained micaceous quartz arenite. There is no well-
defined bedding, although carbonaceous stringers give an
incipient stratification. The U-tube, which is vertical

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 315

Text-figure 7.— Generalized U-tube Morphology. A. U-plane
section of a Sfreite-bearing tube with a combination of protrusive and
retrusive Spreiten. The protrusive Spreiten (solid lines) extend only to
the inner margin of the arm of the U whereas the retrusive Spreiten
(dashed lines) extend to the outer margin. B. Median section taken
through points a-b fig. A showing protrusive Spreiten only. The arcs
are convex upward. If the Spreiten reflect the organism’s attempt to
lengthen the tube to compensate for an increase in body size one might
expect the diameter of the tube to increase accordingly, as illustrated
herein. However, the diameter of the Cincinnatian tubes remains
constant. C. Median section taken through points a-b on fig. A show-
ing the superposition of retrusive Spreiten over protrusive Spreiten.
Specimens with retrusive Spreiten are unknown in the Cincinnatian
section. D. Four transverse sections taken at corresponding points on
fig. A. The arcs of the protrusive Spreiten (2-3) are convex inward
whereas the retrusive Spreiten are convex outward. A transverse sec-
tion taken at the mouth of the “U” (1) reveals no trace of the
Spreiten.

throughout its length, has a depth of 8 cm and a length of
6.5-7 cm. The tube is slightly longer at the base than near
the top. The breadth is not clearly shown, but the diameter
of the arms of the “U” is 1-1.2 cm. The specimen shows

D

that what Smith interpreted as a “U in U” structure is
actually a Spreite, where the arcs abut against the arms of
the “U.” Although the Spreiten are slightly irregular, they
are plainly visible. The dark carbonaceous material filling
the “U” and incorporated into the Spreite makes the fossil
stand out against the lighter gray of the host rock.
Although Corophioides has been reported from the
Devonian of Scandinavia (Richter, 1926) and Germany
(Dahmer, 1937) and from the Lower Cambrian of Estonia
(Opik, 1929) and Pakistan (Seilacher, 1955), it is best
developed in the Triassic Bunter Sandstone of southern
Germany. It occurs in large numbers in the middle and up-
per members and has also been reported from the lower
Bunter by Schindewolf (1928). Although U-tubes had been
noted as early as 1869 by Platz, not until several years
later were they named Arenicoloides luniformis by Blanck-

enhorn (1916). Richter (1924) placed Arenicoloides in

316 PALAEoNnToGRAPHICA AmerRIcANA (VI, 41)

synonymy with Corophioides in spite of the fact that he
had had no opportunity to study the type specimen and
realized that Smith had described a “U in U” structure. Al-
though some authors, e.g., Andree (1927), Bather (1925),
Westergard (1931), have reserved opinion because of the
uncertainty regarding the morphology of Corophioides
polyupsilon, most workers, e.g., Schindewolf (1928), Abel
(1935), Hantzschel (1962) have followed Richter. Even
Blanckenhorn (1924) admitted that Richter was correct in
reducing Arenicoloides to synonymy. As will be explained
below, Richter’s conclusion is accepted herein.’

C. luniformis are found in such large numbers in the
Bunter that Blanckenhorn (1916) was able to count 36 in-
dividuals in an area of 90 square cm. According to him the
tubes occur in a strong, medium-grained, quartz sandstone
which has a “siliceous mud cement.” C. luniformis has
several expression in the Bunter. As described by Blancken-
horn (1916, 1924) and Soérgel (1924), some may occur as
shallow, vertical, semilunar depression (see Sérgel, 1924,
pl. 19, fig. 1); while others appear as dumbbells. There is
no preferred orientation and the forms frequently inter-
penetrate; occasionally three will converge to form a
Hiihnertrappe-like (chicken tracks) imprint. The forms are
best seen in U-plane section (Sérgel, 1924, pl. 20, figs. 2, 3
and Abel, 1935, fig. 379). The specimens are normal to the
bedding plane and appear as gently rounded “U's.” There
is no increase in length as the tube deepens as there is in
C. polyupsilon. Even though the coarse grain size obscures
the finer details, the Spreite is plainly visible, Richter (1924,
p. 211) described the Spreite as being slightly irregular in
a U-plane section, and Sérgel (1924, fig. 2) mentioned that
the breadth of the Spreite may widen with depth, as shown
herein in fig. 7B. Although no specific mention is made as
to whether the Spreite is of the protrusive or retrusive
variety, judging from the interpretations of the various
workers, the Spreite is protrusive.™ There is no mention of
of scratch marks on the arms or the presence of an apertural
funnel. As will be shown below, both these features serve to
differentiate Corophioides from other U-tube genera. Meas-
urements on the dimensions of the Bunter forms vary and
are in part given in tabular form by Sérgel (1924, p. 545).
The length varies from 1-5 cm, the depth from 1-130 mm.,

“S$rgel (1924), who conducted extensive studies on the Bunter forms,
recognized Arenicoloides as valid; however, his conclusion was based
on his ethological interpretation and not on morphology.

However, Seilacher tentatively placed the Salt Range forms in C.
rossei instead of C. luniformis, because the latter “. . . offenbar
einen abweichenden (retrusive) Bauplan verkérpert” (Seilacher,
1955s peo 7))\e

and the breadth from 1-5 mm. Large and small individuals
can occur on the same slab. Although Sérgel believed that
many forms were incomplete and once extended into the
overlying muds, he was unable to confirm this by direct
observation. j

Several other examples of U-tubes are known in the
fossil record, and in 1926 Rudolf Richter established two
“families.” The “Rhizocorallidae” was reserved for forms
with a Spreite and the “Arenicolitidae” for genera lacking a
Spreite. Three of the better known genera, Diplocraterion
Torell (1870), Rhizocorallium Zenker (1836), and Arent-
colites Salter (1857), will be briefly compared and contrasted
with Corophioides. The problematic dumbbell genera, Ar-
thraria Billings (1874) and Bifungites Desio (1940), will
be discussed later.

Diplocraterion was erected by Torell (1870) for
forms from the Cambrian sandstones at Lugnas, Sweden.
Torell’s description was vague and the genus was not figured.
The one key point he mentioned was that the apertures of
the tube were funnel-like. Richter (1926), working solely
from the literature, believed that Diplocraterion lacked
Spreite and placed it in the “Arenicolitidae.” The true
nature of the genus was revealed by Westergard (1931),
who was able to locate what he believed to be three speci-
mens of Torell’s original material. Combining this with addi-
tional material from the Swedish section, he was able to give
an accurate diagnosis of Diplocraterion. Westergard found
that the tubes are vertical and the arms normally parallel,
although some may diverge or converge slightly with in-
creasing depth. Tubes of all sizes occur together, and the
depth and length are mutually independent. Occasionally
the base of the tube is straight instead of U-shaped. The
breadth varies from 2-15 mm, the length from 2-7 cm, and
the depth from 2-32 cm. Funnels are present as is shown
in Westergard (1931, pls. 4-7), albeit Westergard remarked
that weathering tends to exaggerate them and most un-
weathered specimens did not possess funnels. Westergard
was also able to clearly illustrate the presence of Spreiten
in Diplocraterion, thereby correcting Richter’s earlier error.

A more recent work on Diplocraterion is by Goldring
(1962), who described D. yoyo as it occurred in the Upper
Devonian Baggy Beds of Great Britain. The depth of the
tubes ranges from 1-10.5 cm, although most have been trun-
cated by subaqueous erosion. By studying the various levels
of the tubes, Goldring was able to work out several stages of
erosion. His uneroded forms show a remarkable development
of the funnels (see Goldring, 1962, pl. 3, fig. 3 and text fig.
2). A transverse section through the funnels reveals “a series
of concentric laminae composed of fecal pellets, mudstone

TRACE FossiLs CINCINNATI AREA: OsGoop 317

and sandstone surrounding each core. The distance across
the rings may be 3-8 times the diameter [of the arm]... ”
(Goldring, 1962, p. 240). Goldring was also able to illus-
trate the presence of protrusive and retrusive Spreiten in
D. yoyo. In one case both varieties of Spreiten were found
im the same specimen (Goldring, 1962, p. 240).

In summary, the only major difference between Diplo-
craterion and Corophioides is the presence of funnels in the
former. Although these are not especially well developed
in many of the Swedish forms, they were present in two of
Torell’s specimens located by Westergard, and they are
well developed in Goldring’s material. Although Seilacher
(1963) discounted funnels and placed Corophioides in syn-
onymy with Diplocraterion, it is believed herein that they
are of sufficient importance to permit maintaining the two
as separate genera.

Rhizocorallium Zenker (1836), orginally described
from the Muschelkalk (Triassic) of Germany, is a common
form and is known from the Cambrian, ? Devonian, Per-
mian, Triassic, Jurassic, Cretaceous, and Tertiary and has
been reported from Europe, North America, Asia, and
Australia. Rhizocorallium differs from the genera discussed
above, as the greater portion of the “U” may be oblique or
even parallel to the bedding plane. The burrow is initially
vertical but it then undergoes a 90° bend. Rhizocorallium
may be large; Abel (1935, figs. 375,376) illustrated two
forms which have a depth of nearly 12 cm. Moreover, the
breadth of the arms may exceed 1 cm. The genus also differs
from all other U-tubes in that both the Spreiten and the
arms are marked by numerous short striae (see Abel, 1935,
fig. 376). According to Goldring (1962) no forms with re-
trusive Spreite are known. The striated arms and the curva-
ture of the “U” suffice to separate Rhizocorallium from all
other U-tubes. The reader is referred to Abel (1935) and
Hantzschel (1962) for a more complete discussion of the
morphology and relationships of Rhizocoralliwm.

Arenicolites Salter (1857) was first described from the
Carboniferous sandstones of Lancashire, England. More
recently it has been described from the Upper Jurassic
of the Juras (Rieth, 1931), Jurassic of England (Bather,
1925), Ordovician of Thuringia (Zimmerman, 1914), and
Lower Cretaceous of Colorado (MacKenzie, 1963). Areni-
colites differs from other U-tube genera, because it clearly
lacks Spreite and thus is a simple U-tube. There have been
no reports of striae or apertural funnels. The forms described
by Bather had a depth of 20-30 cm and a length of 4-6 cm.
It is worthy of note that even though there is no Spreite in
Arenicolites, the apertures, when viewed on the bedding
plane, may be connected by a bar. This is a feature which
is normally restricted to the Spreite-bearing genera. Bather

(1925, p. 188) was able to show that this was due to a
sagging of the bedding between the arms of the “U”
following the death or departure of the inhabitant. He was
able to trace the bedding across the entire structure and
this interpretation was confirmed by Richter (1926). Bather
(1927) attempted to extend this explanation to the Spreite
of “Arenicoloides,’ but in an appendix to the paper he ad-
mitted that direct field observations of “Arenicoloides” failed
to confirm his hypothesis and that the Spreite of “Areni-
coloides” formed an integral part of the “U.”

It should be mentioned that Arenicolites may be more
common than is supposed. Spreite-bearing U-tubes are
usually easy to recognize on bedding plane surface because
of their dumbbell shape. Unless sagging had occurred,
Arenicolites (as well as Spreitebau with long free tubes —
see Text-fig. 7D) would appear only as two circular open-
ings and might well be overlooked. Moreover, the Spreiten
tend to form vertical planes of weakness, and thus jointing
commonly reveals the U-plane section. This would not be so
likely to occur in Arentcolites. Nevertheless, it must be ad-
mitted that this realization has not yet led to the discovery
of arenicolitid structures in the Cincinnatian section.

Interpretation — Smith (1893) believed that Coro-
phioides was made by an animal similar to the Recent filter-
feeding amphipod, Corophium, which is a common member
of the infauna of tidal flats in many parts of the world.
He maintained that the organism enlarged its burrow to
compensate for an increase in body size by breaking
“through the outer and under sides of its tube and [re-
forming] its habitation, in some cases as many as eight
times. It is, therefore, only the last dwelling-place or the
largest outer U that is preserved in an entire state .. .”
(Smith, 1893, p. 291). The U-tubes of the Recent Coro-
phium volutator (Pallas) have been studied by Hantzschel
(1939, 1955), Seilacher (1953a, 1957), and Schafer (1962).
These amphipods construct a U-shaped burrow in sand or
mud which may attain a depth of 4 cm. The arms of the
“U” are close together but may diverge slightly with depth.
The walls of the “U” are smooth. Hantzschel (1955, fig. 1f)
illustrated an occurrence of Corophiwm U-tubes where the
upper portion of the tubes had been eroded away and only
the base remained. As he pointed out, this closely re-
sembles Corophiotdes luniformts, as it frequently occurs in
the Bunter as well as in some Cincinnatian specimens ( PI.
61, fig. 5). In well-aerated sand the animal may construct a
single vertical shaft instead of a “U” (compare Seilacher,
1953a, fig. 4, left with fig. 4, right). The aperture of the
shaft is marked by a characteristic stellate pattern of
markings which are apparently a result of feeding. Originally
Hantzschel (1939) described the burrow of C. volwtator as

318 PALAEONTOGRAPHICA AMERICANA (VJ, 41)

a simple “U” lacking Spreite. However, Seilacher (1957, pl.
22, figs. 1-2) was able to show that, although difficult to see,
protrusive Spreiten are definitely present. This observation
was confirmed by Schafer (1962, figs. 178-180). Schafer
remarked that the Spreiten were easiest to see where there
is an interbedding of sand and mud. He also pointed out
that the Spreiten are not closely packed and that the “U”
is not always vertical. His figure 180 shows one form which
is initially vertical but then undergoes several “zig-zags” as
the depth of the tube increases. The reason behind the con-
struction of the protrusive Spreite is not clear. The animals
are small, and it is doubtful that the sole purpose was to
compensate for an increase in body length.

The only other organism known which constructs a
Spreitebaw is a small polychaete, Polydora ciliata. The
burrows are most commonly made in pelecypod shells, and
the Spreite is not destructional but constructional, As shown
by Schafer (1962, fig. 177), the organism first excavates a
pocket-shaped burrow (not a U-tube!) (Schéafer, 1962,
fig. 177, left). It then proceeds to construct a U-tube by
filling in the center of the pocket with agglutinated sand
grains. The resulting structure closely resembles a U-tube
with protrusive Spreiten (see Schafer, 1962, fig. 177, right).
As will be shown below, all fossil Sprettebauten are believed
to be of the destructional form.

In marked contrast to Smith’s interpretation of Coro-
phioides polyupsilon, Blanckenhorn (1902, 1916) originally
regarded the shallow U-shaped depressions of Corophioides
luniformis as the tracks of a three-toed dinosaur, probably
Cheirotherium, which occurs in the Bunter beds. Later
Blanckenhorn (1916) studied U-plane sections and recog-
nized his mistake. As his choice of a generic name indicates,
he believed that the genus was the U-tubes of a polychaete
similar to that of the Recent lug worm Arenicola marina
(Lamarck), which occurs in large numbers on tidal flats.
Sérgel (1924) echoed this opinion. However, Richter (1924,
1926) was quick to point out that for the most part Areni-
cola does not construct a well-defined U-tube.14 As Wells
(1945) illustrated, in loose sand Aremcola marina excavates
a J-shaped burrow which is strengthened by a coating of
mucus, The animal lies with its anterior end at the closed
end of the “J” and moves upwards, disturbing the overlying
sand. The worm then withdraws into the closed portion of
the “J,” and gravity causes subsidence of the disturbed
sand which the organism then ingests. The sediment is
passed through the gut where the nutrients are removed,

"The tube of the lug worm has received considerable study; and the
reader is referred to Richter (1924), Wells (1945), and Schafer
(1962) for a complete discussion.

and the undigested sediment is passed out of the open end of
the “J,” where it forms long ropelike fecal strings. The re-
sulting structure, which is well illustrated by Schafer
(1962, fig. 174), thus resembles a U-tube with a conical
column of disturbed sand lying above the closed end of the
“J.” As viewed on the depositional interface, the burrow
appears as a small circular opening surrounded by fecal
strings and a much larger saucer-shaped depression mark-
ing the location of the column of disturbed sand. This is
probably what led earlier workers to believe that it was a
simple U-tube. In hard-packed sand it is necessary for the
animal to lengthen the closed portion of the burrow, with
the result that the conelike column of disturbed sand is
much smaller (see Schafer, 1962, fig. 175). In neither of the
above-discussed examples is there a Spreite developed. In
contrast, when dwelling in highly cohesive mud, Aremcola
does construct a Spreite-like structure. Schafer (1962, figs.
216,217) showed that the worm forms a U-shaped burrow,
then withdraws to the base of the “U” and excavates a new
shaft, feeding on some of the removed mud and plastering
the remainder in the old shaft. This process may be re-
peated several times, with the result being one permanent
shaft and several bundled feeding shafts. It is interesting to
note that this explanation is similar to the one given by
Sérgel (1924, fig. 4) as an explanation for the Spreiten of
Corophioides luniformis. While this may be a valid explana-
tion for some of the Bunter forms, it was completely re-
jected by Richter (1924). In most of the Bunter specimens
the Spreiten are not asymmetric or crowded against one
arm, and there is no mention of multiple openings on the
depositional interface. Instead there are only two openings
to the burrow, and the Spreiten are parallel to the last-
formed base of the “U.” In other words, the deepest part of
the Spreite (see Text-fig. 7A) lies midway between the
arms. Moreover, the Bunter forms occur in a sandstone;
and, as shown by Wells (1945) and Schafer (1962), Areni-
cola builds J-tubes in sand. It is more likely that Richter
(1924, 1926) was correct when he concluded that Coro-
phioides was the Domichnia of a filter-feeding organism and
that the protrusive Spreiten represent movements by the
organism to lengthen its burrow to compensate for an in-
crease in body size. The sediment removed from the floor of
the burrow was packed on the roof probably with the aid
of a mucus cement. This is not offered as a general ex-
planation for all Spreitebauten but applies only to C.
polyupsilon, C. luniformis, and the Swedish examples of
Diplocraterion, Goldring’s study of D. yoyo showed that
some Spreiten are a response to sedimentation or erosion.
Moreover, the fact that a major part of the Rhizocoralliwm
structure lies parallel to the bedding suggests that these are

‘TRACE FossiILs CINCINNATI AREA: Oscoop 319

feeding structures where the organism has mined a particu-
larly rich area.

Opik (1929) believed that the irregular Spreiten of
C. helmerseni might represent the fillings of a pocket-like
burrow [2.e., similar to the burrow of Polydora illustrated
by Schafer (1962, fig. 177, left) ] which was filled by sedi-
ment following the departure of the inhabitant. More will be
said regarding the origin of the Spreite in the discussion of
the Cincinnatian forms.

It is impossible to assign Corophioides or fossil U-
tubes in general to a given group of organisms. As already
shown, both amphipods and polychaetes construct Spreiten-
like structures, and simple U-tubes lacking Spreiten are
formed by the echiuroid Urechis caupo, freshwater insect
larvae, several polychaetes, and the hemichordate Balano-
glossus. With the possible exception of Balanoglossus, the
animals are filter-feeders which remain stationary in their
burrows. By movement of parapodia or other mechanisms
they set up currents which draw organic detritus into the
burrow. (See Barnes, 1963, pp. 184-186, for a discussion of
the feeding habits of Chaetopterus and Rickets and Calvin,
1962, pp. 304-306, for Urechis caupo.)

Richter (1926) concluded that fossil Spreitebauten
occurring in large numbers are indicative of marine environ-
ments. Although he believed that this is the only environ-
ment which could supply a sufficient amount of plankton,
the theory has not been adequately tested.

CINCINNATIAN REPRESENTATIVES

Over the past 70 years of Corophioides there has ac-
cumulated from over the world a total of seven species, rang-
ing in age from Cambrian through Triassic. Without a de-
tailed re-study their validity can not be assured. There
does not appear to be any significant difference between C.
polyupsilon Smith (Scotland: Upper Carboniferous) and
C. luniformis (Blanckenhorn) (Germany: Lower Triassic)
as restudied by Sorgel (1924). The only apparent morpho-
logic difference is that in the former the transverse section
lengthens slightly with the depth of the “U,” whereas the
arms of C. luniformis remain parallel.

Rudolf Richter (1926) proposed C. erraticus for a
single specimen occurring in a glacial erratic boulder found
in Prussia but apparently originally derived from the Cam-
brian sandstones of Sweden. He differentiated it from C.
luniformis because the Spreiten of C. erraticus are crowded
against one of the arms. Yet, one of Sérgel’s (1924, pl. 20,
fig. 3) illustrations of C. luniformis appears to show this
same phenomenon. Richter separated C. erraticus from C.
polyupsilon because the former lacked the “U in U” struc-

ture of the Spreite in Smith’s species. It has been shown
above that Smith’s description was a misinterpretation.
Aside from the fact that C. erraticus was based on a single
specimen, both of Richter’s differentia appear to rest on
shaky ground. Westergard (1931) placed Richter’s species
in synonymy with Diplocraterion parallelum TYorell (1870)
(Sweden: Lower Cambrian).

Opik (1929) proposed C. helmerseni for a form from
the Lower Cambrian of Estonia. This differs from other
species in being slightly expanded (transverse section
lengthens) at the base and in having a few irregularly
spaced Spreiten. Nevertheless, Westergard (1931) con-
sidered it synonymy with the Swedish Dipolcraterion par-
allelum.

In 1930 Hecker erected Rhizocorallium devonicus for a
U-tube from the Devonian of Russia. Without explanation
Abel (1935) placed the species in Corophioides. Judging
from Abel’s (1935, fig. 381) illustration, the arms have the
distinctive deflection of the U-plane of Rhizocorallium;
and there appears to be no basis for placing it in Coro-
phioides.

The morphologic distinctiveness of C. balticus Andree
(1927), from the Devonian of Prussia, and C. rosei Dahmer
(1937), from the Taunus Quartzite (Lower Devonian) of
the Rhineland is questionable. The former was never figured
nor diagnosed, and the latter was figured but only briefly
discussed. One gets the impression that the sole reason for
proposing the species was because they were found in areas
where U-tubes had not previously been reported.

Kolesch (1922) described one U-tube species from the
Middle Bunter Sandstone (Upper Triassic) of Germany
under the name of Arenicolites zimmermannt. However, as
Andree (1927) and Hantzschel (1965) pointed out, the
species possesses Spreiten and belongs in Corophioides.
Kolesch (1922, fig. 7) differentiated it from C. luniformis
because the Spreiten are bowed out to one side, and thus
the transverse section takes on an arcuate shape. The species
has never been restudied, and it is not known how consistent
this pattern is.

Three forms of U-tube traces are present in the Cin-
cinnatian section, and their relationship to the forms in the
world literature is necessarily somewhat doubtful. One of
the Cincinnatian kinds closely resembles most of the forms
discussed above, 2.¢., it is a simple U-tube with Spreiten,
but without any marked characteristics which would set it
apart from the other “‘species.” It is the least common of the
three forms and will be discussed as Corophioides cf. luni-
formis. The other two kinds differ sufficiently from each
other and from al! other species of Corophioides seemingly

320 PALAEONTOGRAPHICA Americana (VI, 41)

to merit specific designations. C. cincinnatiensis, n. sp. is
characterized by a bilateral expansion of the base of the
“U.” It is somewhat similar to C. helmerseni Opik (Lower
Cambrian: Estonia) but differs in that the Spreite is more
regular and the basal expansion more abrupt in the Cincin-
natian form. C. biclavata (Miller) (1875) was originally
described as Arthraria biclavata, the “dumbbell fucoid.” It
can now be shown that this type of dumbbell represents
a modification of the base of C. cincinnatiensis, where the
arms of the “U” have been extended below the arcuate base

of the! m= U2

Median sections reveal that the Spreiten of all three
types of Corophioides of the Cincinnatian are of the pro-
trusive form and are composed of reworked host rock. In
contrast to the Bunter specimens described by Sorgel
(1924), the Cincinnatian Spreiten do not broaden with
depth but remain constant. The arms and the base of the
“U” are filled with a finer grained material which readily
weathers out or is easily removed. Presumably it represents
sediment washed into the tube after the inhabitant had
departed.

Corophioides is one of the most common Cincinnatian
trace fossils. It ranges throughout the entire section and,
when found, normally occurs in large numbers. The in-
dividual U-tubes commonly intersect one another, and there
is no preferred orientation other than vertical to the
bedding. The genus is most frequently encountered in hard,
cross-laminated, fine-grained calcarenites but has also been
found in platy calcisiltites. As will be shown below, there is
indirect evidence to show that it occurs in lutites as well.
It has never been found in the extremely coarse-grained
calcarenites.

It is noteworthy that at one locality all three species
have been found in the same bed, while in another locality
C. cincinnatiensis and C. cf. luniformis have been found to-
gether. This suggests the possibility that these three “ichno-
species” represent different expressions of the burrow of the
same organism.

Corophioides cf. luniformis (Blanckenhorn)

Plate 61, figure 5; Plate 62, figure 1,2; Plate 69, figure 3; Text-figure
29-0

The forms included under C. cf. luniformis are Spreite-
bearing U-tubes which have a gently rounded base and
thus lack the marked basal expansion of C. cincinnatiensts.
As nearly as can be determined from the literature (no
specimens were available for study), these tubes are mor-
phologically similar to many of the European “species.”
However, since C. luniformis is the best known of the Eu-

ropean forms, the Cincinnatian specimens are tentatively
allied with it. Nevertheless, it should be pointed out that
a direct detailed comparison of the Bunter forms with those
from the Cincinnatian might well reveal characteristics
that would set them apart, for it is possible that they were
made by unrelated organisms.

Three specimens of C. cf. luniformis showing U-plane
sections are known. Two of these (PI. 69, fig. 3) were found
only a few centimeters apart in the McMicken beds in West
Fork Creek (Cincinnati, Ohio) at the type locality of C.
cincinnatiensis. The host rock is a fine-grained, cross-
laminated calcarenite which has a thickness of 7.5-12 cm.
The overlying and subjacent lutites are barren of U-tubes.
The two specimens are nearly identical in measurements, as
both have a depth of 2.3 cm and a breadth of 5 mm. The
transverse length of one specimen is 3.3 cm, that of the
other 3.2 cm. No free tubes are visible, as the Spreiten
extend to the upper limit of the calcarenite, and attempts to
trace the tubes into the overlying lutite were unsuccessful.
Also available for study was a slab from an unknown
locality bearing a single specimen of C. cf. luniformts (PI.
62, fig. 1), as well as several examples of C. cincinnatiensis.
The tube has a transverse length of 4 cm, a breadth of 2.5
cm, and a depth of 3.8 cm. The Spreiten are evenly spaced
and extend to the top of the bed.

Broad arcuate impressions, which are believed to repre-
sent “molds” of the base of C. cf. luniformis, have been
found in the Corryville and Richmond rocks. The Corry-
ville specimen has a breadth of 8 mm and a depth of 1 cm;
the transverse length is 10 cm. The Richmond specimen,
which has an area of 91 square cm, contains 55 individuals,
many of which intersect. The breadth (2 mm) is the only
consistent measurement, for the transverse length varies
from 6-15 mm and the depth ranges from 1 to 3 mm. These
impressions closely resemble Blanckenhorn’s (1902, fig. 1)
illustrations and discussion (1924) of the lower portions of
some of the Bunter specimens of C. luniformis. While
admittedly there is no direct proof, the Cincinnatian speci-
mens are believed to represent the bases of C. cf. luniformis
where the upper portions of the tube have been stripped
away by erosion. The fact that no Spreiten are found
suggests that the main body of the tube lay within a lutite
and that the tubes bottomed in the calcarenite. Unfortun-
ately this can not be confirmed, as some of the specimens
were found in place.

Interpretation — Based on the discussion of Recent
U-tube dwellers given elsewhere, the specimens grouped
under C. ef. luniformis are thought to represent the Domich-
nia of sessile filter-feeding organisms. Although it has never

TRACE FOSSILS CINCINNATI AREA: Oscoop 321

been demonstrated, there may well be an optimum set of
tube dimensions for maximum efficiency in filter feeding.
If the tube is too long (as measured along the “U”), it
might be difficult for the organism to set up a sufficient
current. On the other hand, if the tube is too short, the in-
habitant might be too close to the apertures. The protrusive
Spreite might represent an attempt by the organism pro-
portionately to lengthen the tube as the body length in-
creases with age. In addition, erosion of the upper part of
the U-arms could force the inhabitant to deepen the tube.
The fact that the Spreite extends to the top of the bed sug-
gests that erosion was operative. A third possibility would
be a combination of these two factors.

The zoological affinities of the producer of C. cf. /uni-
formis are unknown. U-tubes are constructed by represen-
tatives of so many different phyla that it is unwise to specu-
late too extensively. While the lack of striae on the arms and
Spreiten would seem to rule out arthropods, C. cf. luniformis
could be the tube of any one of a number of soft-bodied
organisms. Of course, the annelids are a favorite in such
speculation.

Corophioides cincinnatiensis, n.sp.

Plate 60, figures 1,5,8; Plate 61, figures 4,6,8; Plate 62, figures 3,4,6;
Plate 63, figure +; Text-figure 8,29p

Type locality —The specimen illustrated on Plate 61,
figure 4 (UCM 37590) from the McMicken beds of West
Fork Creek, Cincinnati, Ohio, is designated as the type
specimen for C. cincinnatiensis. The type locality is 303
meters east of the intersection of Shepherds Road and
West Fork Road, Mt. Airy Forest, Hamilton County, Ohio.
The bed containing the specimen crops out on the south
bank some 50 cm above the water level and 2-3 meters
downstream from a dam.

Diagnosis — Vertical Spreite-bearing U-tubes which in-
crease in transverse length with depth. Most specimens are
marked by an abrupt flaring of the base of the “U.” The
upper part of the arms is approximately parallel but in
many cases exhibits numerous minor irregularities. Free
tubes unknown. The Spreiten are regular and parallel in
most cases but may show some overlapping.

Discussion — C. cincinnatiensis is more common than
C. cf. luniformis and is especially abundant in the upper
part of the McMicken, where it is associated with Chon-
drites and Trichophycus venosum. Two of the best expos-
ures are the type locality and a long outcrop on U.S. Route
27, 1.4 miles south of Newport Shopping Center, Newport,
Kentucky. At the latter locality the species is best seen on
the east side of the road at the top of the talus slope.

At the type locality the tubes occur in the well-bedded
calcisiltite which contains C. cf. luniformis. The basal 1-2
cm of the bed is composed of a more coarse-grained fossil
hash. While a few of the tubes penetrate to the base of
the bed (Text-fig. 8), the majority terminate at the upper
limit of the fossil hash. The contact between the host rock
and the subjacent lutite is sharp, but the upper calcisiltite-
lutite contact is more gradational, the distance of transi-
tion being 2-3 cm.

The type specimen (PI. 61, fig. 4) has a depth of 6.8
cm and a breadth of 3 mm. The arms are approximately
parallel throughout most of their length, although irregu-
larities do exist, especially in the upper portion of the right
arm. The transverse length of the specimen down to the ex-
panded base varies between 2.3-2.8 cm, and the Spreiten,
which extend to the upper limit of the tube, are parallel
and regular. At a depth of 5.1 cm both arms diverge out-
ward at an angle of 40° off the vertical axis to produce the
flared base so characteristic of the species. The transverse
length of the tube then increases to a maximum of 3.9 cm
before terminating in a gently rounded base 1 cm above the
bottom of the bed. Although the Spreiten are evenly spaced,
those of the expanded portion of the tube are more strongly
bowed (1.e., more strongly convex downward) than the
Spreiten of the unexpanded portion.

Text-figure 8 illustrates three additional topotype speci-
mens which, owing to the size of the slab, could not be re-
moved from the outcrop. Text-figure 8-1 shows a tube which
extends through the shell hash to the base of the bed. It
differs from the type specimen in that the arms gradually
diverge with depth, the basal expansion is not so pro-
nounced, and the base of the tube is slightly flattened.
Forms similar to this were seen to extend as much as 1 cm
into the subjacent lutite. Another variation of C. cincin-
natiensis is shown in Text-figure 8-2. Here the arms con-
tinue to diverge with depth, and although the base of the
tube is flattened, there is no sudden expansion. The speci-
men shown in Text-figure 8-3 closely resembles the type
specimen, except that the upper portion of the arms is more
regular, Several other forms similar to the three discussed
above occur in the same bed, along with representatives of
C. cf. lumiformis and C, biclavata.

At the Kentucky locality mentioned above, C. cin-
cinnatiensis occurs in large numbers in two cross-laminated
calcisiltites separated by some 60 cm of barren lutite. The
specimens do not differ greatly from those just discussed,
except that in all observed cases the U-tubes reach to the
base of the calcisiltites and many times extend up to 5
mm into the subjacent lutite. None of the specimens in

S)
bo
bdo

t

either bed could be traced into the overlying lutite, yet
in all cases the Spreiten extended to the upper limit of the
calcisiltite. Most of the tubes are characterized by a marked
basal expansion. The most extreme example is seen on Plate
61, figure 6, where the length as measured across the upper
part of the specimens is 2.4 cm, while the length at the
base of the calcisiltite is 6.2 cm. As in the specimens from
the type locality, the transverse length of the tubes varies
considerably. In some (PI. 61, fig. 3) the average length is
only one-third the depth, while in others (Pl. 60, fig. 5)
the length may exceed the depth. Several specimens show-
ing intermediate stages have been observed.

Although the arms of most of the tubes from both
localities are smooth or covered with secondary calcite
encrustations, two specimens are known where the last-
formed “U” bears annulations. Plate 62, figure 4 shows
the base of a “U” from the Kentucky locality which has
12 annulations per centimeter. The other specimens (PI.
60, fig. 1), from the type locality, shows an unweathered
filling of one of the arms where the annuli are larger, av-
eraging 8 cm. This latter specimen also shows the angular
irregularity of the arms which is encountered in many
forms.

Interpretation — As mentioned elsewhere, C. cincin-
natiensis may well be a behavioral variant of C. cf. luni-
formis, where the major deviations are the irregularity of
the arms and the basal expansion of the tube. As in C.
cf. luniformis the Spreiten are thought to represent older
relics of the base of the “U” where the organism deepened
the tube: a) to compensate for an increase in body length,
b) to compensate for erosion of the upper part of the tube,
or c) a combination of the above. If the inhabitant of the
tube were a filter-feeder, a steady even flow of water
through the tube was a necessity. Since neither the irregu-
larity of the arms nor the angular expansion of the base
would seem to increase the efficiency of the tube, they must
represent a reaction of the organism to some obstacle en-
countered in burrowing. In fine-grained material irregulari-
ties in the symmetry of the tube could arise through un-
even compaction or slippage, but in the forms under con-
sideration here only the arms are distorted. The Spreiten
are even and show no trace of any disturbance. Perhaps
the organism was forced to detour around slightly indur-
ated or more compact patches of sediment.

The basal expansion of the “U” is thought to repre-
sent the reaction of the organism to a physical or chemical
change in the sediment. Note that at both localities the
tubes terminate either at a silt-mud or silt-fossil hash in-
terface. At the type locality only a few of the individuals

PALAEONTOGRAPHICA AMERICANA (VI, 41)

Text-tigure 8.— Corophioides cincinnatiensis, n. sp. Three speci-
mens from the type locality, West Fork Creek, Mt. Airy Forest, Cin-
cinnati, Ohio, showing various types of basal expansions. Tubes 2
and 3 terminate at the upper limit of a coarse-grained fossil hash;
X 0.4.

(Text-fig. 8-1) were able to penetrate through the fossil
hash and into the underlying mud, Instead most forms were
forced to lengthen the tube by moving laterally. At the
Kentucky locality the organism apparently had an ayersion
to mud. Perhaps it was too hard, tended to clog the res-
piratory apparatus, or was chemically distasteful.
However, it must be admitted that a change in the
texture of the sediment does not provide the answer for
all such expanded U-tubes. U-plane sections of C. biclavata
(see Pl. 77, fig. 8) occur with C. cincinnatiensis at the West
Fork Creek locality and alone in the Mt. Hope in a creek
adjacent to Sheed Road, 200 meters northeast of the inter-
section with Gaines Road, Hamilton County, Ohio. As will
be shown, C. biclavata also expands at the base, and the
only apparent difference between it and C. cincinnatiensts
is that the arms extend below the level of the Spreite. Speci-
mens of C. biclavata have been found at these two locali-
ties where the tube ends well within the silt bed and not

‘It is interesting to note that the animal could lengthen the tube by
either expanding the length, as the Cincinnatian forms did, or by
directing the ‘“‘U” either oblique or parallel to the bedding. It is un-
usual that the latter variant is not found in at least a few Cin-
cinnatian specimens. It would also be worthwhile to re-evaluate
Rhizocorallium in this light. Perhaps the curvature of some forms
assigned to this genus is a response to a change in lithology and not
a feeding structure as Seilacher (1955) and others postulated. Like-
wise, Corophioides helmerseni Opik should be restudied to see if the
flared base can be explained in the same manner. Opik (1929) attrib-
uted it solely to the growth of the animal.

TRACE FOSSILS CINCINNATI AREA: Oscoop

at a lithologic contact. Perhaps here the animal encoun-
tered a chemical change which is not reflected in the sedi-
ment.

At noted in the discussion of C. cf. luniformis, it is
difficult to assign U-tubes to a given group of organisms.
However, annuli found in two specimens of C. cincinnatien-
sis suggest a segmented vermiform body. If C. cf. luntformis
is only a behavioral variant, this means that it too was
inhabited by a metameric vermiform organism.

Corophioides biclavata (Miller)

Plate 60, figures 3,4; Plate 61, figures 1,3,7; Plate 65, figure 6; Plate
66, figure 5; Plate 70, figure 5; Plate 77, figure 8; Text-figures 9,29q.

1872. ?Arthraria antiquata Billings, Canadian Natur., n.s., vol. 6, p.
467, fig. 2.

1875. Arthraria biclavata Miller, Cincinnati Quart. Jour. Sci., vol. 2,
p. 354, fig. 26.

1879. Arthraria biclavata Miller, James, U. P., Supplement to a
Catalogue of Lower Silurian Fossils of the Cincinnati Group,
Cincinnati, Ohio, p. 29.

1884. Arthraria biclavata Miller, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 8,9; pl. 6, fig. 6.

1940. ?Bifungites fezzanensis Desio, Museo Libico Storia. Nat. Ann.,
vol. 2, pp. 78,79, pl. 8, fig. 3.

1940. ?Bifungites halli Desio, Museo Libico Storia. Nat., Ann., vol.
2, pp. 78,79.

1964. ?Bifungites fezzanensis Desio, Dubois and Lessertisseur, Soc.
geol. France, Bull., (7) vol. VI, pp. 626-634, figs. 1,3,6,7, pl. 1.

Type locality — Miller’s figured specimen was from
“one of the numerous impressions in the slabs that make
up the sidewalk, on Torrence Road near Forest Avenue”
(Miller, 1875, p. 374). The genus is relatively common
throughout the Cincinnatian section. The “type sidewalk”
has long since been replaced by concrete.

Diagnosis — Vertical Spreite-bearing U-tubes in which
the arms are extended below the base of the deepest “U”
to form blind pouchs. The species is most frequently pre-
served as concave spireliefs with a dumbbell shape, 2.e.,
two kettle-like depressions connected by a bar. The connect-
ing bar resembles a shallow vertical “U,” and it may or
may not be directly connected to the kettles.

Discussion — The Cincinnatian dumbbells were placed
in Billings’ (1872) genus, Arthraria, first described from
the Upper Cambrian rocks of Newfoundland. As will be
shown later, there is some doubt as to whether the New-
foundland forms are the same as those from the Cincin-
natian. Miller (1875, p. 354) established Arthraria biclavata
for a “fucoid [which] consists of a straight cylindrical stem,
with an enlargement at each end, nearly as round as a ball.
It has some resemblance in shape to dumb-bells. It shows
no structure, It usually consists of the impression, only . . .
sometimes, the cylindrical stem, between the balls, is found
in tolerably good condition, but it is quite rare to find

WwW
tO
Ww

even part of a ball in the matrix, and I have never seen a
specimen free from a slab.” His highly schematic figure
moved U. P. James (1879, p. 29) to remark that “the pub-
lished figure of this species appears much as if copied from
a specimen just from the turning lathe —a remarkable
growth for a sea plant.”

The dumbbell impressions referred to by Miller are
common in the local section and occur as concave epireliefs
in calcisiltites or calearenites. Forms of varying size may
be found on the same slab and they frequently intersect.
The length (the greatest distance as measured along the
bar) varies from 5-14.5 cm. The kettle-like depressions at
either end may be round (PI. 66, fig. 5), reniform (Pl. 61,
fig 3), or cordate (PI. 61, fig. 1). In addition, J. F. James
(1884, fig. 6c) figured one form shaped like an arrowhead.
Although the kettle-like depressions on a given specimen
are normally similar in size and shape, one may be smaller
than the other or even missing entirely (Pl. 65, fig. 6). The
diameter of the depressions ranges from 1-3 cm, and they
are invariably deeper than the bar connecting them. For
example, in a specimen from the Corryville of Stonelick
Creek (PI. 61, fig. 1), the cordate depressions have a depth
of 1.4 cm, while the deepest point of the connecting bar is
only 6 mm below the surface of the slab. Although the bot-
tom of most depressions is gently rounded, in some the
center of the depression is not the deepest point. In the
specimen seen in Plate 61, figure 3, the floor of the de-
pressions slope toward the connecting bar and the inner
wall is undercut.

Likewise, the morphology of the connecting bar may
vary. The bar may actually connect the two depressions
or it may terminate some 5 mm from the depressions (PI.
61, figs. 1,3). The width of the bar averages 4-7 mm and is
independent of the diameter of the depressions. In cross-
section the base of the bar is gently rounded. In almost
all cases the bar resembles a shallow vertical “U,” where
the greatest depth is attained at the midpoint, while in
some examples the bar may be filled with sediment and
rise above the surface of the slab as seen in Plate 60, fig-
ure 3. This filling is most significant in the interpretation
of the species. Sectioning of several specimens shows that
neither the depression nor the connecting bar extend down
into the host rock. The entire structure is visible as a con-

”

cave epirelief.

Similar dumbbell-shaped forms have been described
from other geological horizons, but a discussion of them is
withheld until an interpretation of C. biclavata is given.
The difficulties of a direct comparison will then be appar-
ent.

Les)
to
pe

Interpretation — Miller (1875) interpreted the species
as a “fucoid,” while J. F. James (1884) believed that it may
be a concretion. There were no further attempts to inter-
pret the species until Fenton and Fenton (1934a, pl. 28,
fig. 2) studied the movements of the gastropod Illyanassa
obsoleta (Say) on the tidal flats of San Francisco Bay.
They observed that the snail plows along the surface leay-
ing a trail which resembles a gently rounded trough; then,
for no apparent reason, it twists and digs a shallow hemi-
cylindrical pit. The organism may remain in the pit for
several hours before again proceeding on its way. This
same behavior may be repeated several times, and the final
result is several pitlike depressions connected by rounded
troughs. However, several features of the Cincinnatian
dumbbells fail to support this conclusion. The connecting
bar is straight and does not show the irregularities seen
in the Fentons’ illustration of the trail of 7. obsoleta. More-
over, the bar is U-shaped, and in many cases does not ac-
tually connect the depressions. As already noted and shown
in Plate 60, figure 3, the bar may be filled and stand above
the surface of the host rock.

Several lines of evidence indicate that what Miller
described as A, biclavata represents the base of a Spreite-
bearing U-tube, where the arms have been secondarily deep-
ened below the base of the Spreite (see Text-fig. 9). The
connecting bar is U-shaped, and the filling of the bar seen
in Plate 60, figure 3 is the remnants of a descending Spreite
(compare Pl. 60, fig. 3 with Text-fig. 7D-4). Moreover,
some specimens of Corophioides with expanded bases show
that one or both of the arms have been extended. One
specimen (PI. 61, fig. 7) from the Fulton beds of Union
Levee, Cincinnati, Ohio, is most significant. It is preserved
as a hyporelief in calcisiltite and shows the Spreite and the
filling of the expanded arm. A latex mold of this specimen
is similar to the specimen seen in Plate 60, figure 3. U-plane
sections of tubes with modified bases have been found both
at the type locality for C. cincinnatiensis and also at the
Sheed Road locality. Note that a transverse section taken
2 mm above the base of the specimen seen on Plate 77,
figure 8 would resemble the reconstruction given in Text-
figure 9.

As already noted, the dumbbells occur in calcisiltites
or fine-grained calcarenites which are overlain by lutites.
No examples have yet been located where a U-tube has
been found overlying the dumbbell, and apparently in all
observed examples the upper parts of the tube have been
stripped away. It is believed that the organism burrowed
down through the lutite and halted excavation when the
subjacent silt or sand was encountered. This would be the
opposite of the Newport examples of C. cincinnatiensis,

PALAEONTOGRAPHICA AMERICANA (VI, 41)

Text-figure 9.— Reconstruction of Corophioides biclavata (Mil-
ler). The major portion of the tube lies within the lutite (light
shading) and only the base of the ‘“U” and the extended arm lie in
the calcisiltite (dark shading). When the lutite is stripped away by
erosion only a dumbbell-shaped depression remains; X 0.75.

where the subjacent mud was apparently in some way dis-
tasteful to the burrower. This might suggest that the maker
of C. biclavata and C. cincinnatiensis were different organ-
isms, the former being primarily a mud-dweller, the latter
an inhabitant of silts and sands. Unfortunately this can not
be accepted as a firm conclusion, since both variants have
been found together in the silts of West Fork Creek.

Why the arms were extended below the Spreite is not
at all clear; perhaps another organism used the arms as
a domicile after the death or departure of the original in-
habitant. In this case two animals could live in the tube,
one in one arm, one in the other. Also several U-tube
dwellers are known to have commensals living with them.
For example, the tube of the polychaete Chaetopterus pro-
vides refuge for the small crab Pinnixa, and, according to
Ricketts and Calvin (1962), the large tube of the echiuroid
Urechis caupo is also inhabited by the scale worm Hesper-
onoé adventor, a small crab, Scleroplax granulata, and a
fish, Clevelandia ios, Vhese organisms feed on detritus
pumped into the tube by the echinoid. Although these

TRACE FOSSILS CINCINNATI AREA: Oscoop

Recent commensals do not modify the architecture of the
tube, it is possible that the extended arms of C. biclavata are
outpocketings inhabited by Ordovician commensals.

Other dumbbell-shaped bodies have been found in
the Cincinnatian rocks. All are preserved as convex hypo-
reliefs and none extend up into the host rock. In some the
terminal enlargements are rounded (PI. 61, fig. 2), in others
they are shaped like arrowheads (PI. 71, fig. 5). The total
length of the specimens ranges from 2.7-4.8 cm. Since they
do not extend up into the host rock, they can not be the
actual bases of C. biclavata. Transverse sections of the Cin-
cinnatian Corophtoides taken at any depth reveal a dumb-
bell outline. As mentioned, the Spreiten do not broaden
with depth, indicating that while the animal may have
grown in length, the diameter of the body did not corre-
spondingly expand. However, the arms of the “U” were
used throughout the life of the tube, and constant move-
ment of the inhabitant would lead to an increase in their
diameter. For this reason the dumbbells preserved as con-
vex hyporeliefs could be the casts of the base of C. biclavata
or casts of any portion of the tube of any of the three
Cincinnatian species.

This uncertainty makes it difficult to compare C. bi-
clavata with Arthraria antiquata Billings, the holotype of
the genus Arthraria. Billings’ (1872, fig. 2) illustration of
the type specimen shows that it is a convex body, not a de-
pression as is normally the case with C. biclavata. While
there is no direct proof, A. antiquata is probably the cast
of some portion of a U-tube, but it is not known whether
the tube is a Corophioides, Rhizocorallium, or Diplocra-
terion. It may even be a tube similar to Arenicolites in
which sagging of the bedding has produced a connecting
arch. Unfortunately the type specimen of A. antiquata has
been lost and no topotype material is available. Until such
material can be studied, the species will have to remain
Incertae sedis.

The same is true for Bifungites fezzanensis Desio
(1940) from the Devonian of North Africa and B. halli
Desio (1940) from the Clinton (Silurian) beds of New
York State. B. fezzanensis, which apparently is preserved as
a convex hyporelief in a quartz sandstone, bears a close
resemblance to the convex dumbbells discussed above.
While it may be similar to C. biclavata, Desio makes no
mention of U-tubes associated with Bifungites. He believed
that it was a “fucoid” or “colonial animal.” Dubois and
Lessertisseur (1964) restudied Desio’s material and con-
cluded that the genus represented a filling of the top of a
U-tube. They postulated that the inhabitant may have
been a small trilobite. B. halli is based on a specimen fig-
ured but not named by James Hall (1852, pl. 10, fig. 6)

os)
tro
vi

from the Clinton of New York. Hall believed that this was
a “fucoid” or possibly even a bone. Without a study of
the topotype locality, a meaningful conclusion can not be
given at the present time.

Finally Ruedemann (1925) established Arthraria
magna for dumbbells from the Upper Ordovician strata of
Pulaski, New York. They are preserved “by the hundreds”
as concave epireliefs in “dark-gray mud” which forms a
film over the subjacent sandstone. U-plane sections ( Ruede-
mann, 1925, fig. 65) reveal that these are the upper ex-
pressions of U-tubes and thus are not familiar to C. biclav-
ata. Ruedemann maintained that settling of the Spreite
led to the origin of a connecting bar between the two aper-
tures. In contrast, Richter (1926) said that the connecting
bar of A. magna was entirely straight and thus could not
be due to sagging. He believed that the bar marked the
path of the organism as it slid across the surface from one
aperture to the other. Examination of U-plane sections of
Ruedemann’s material should provide the answer.

?SKOLITHOS Haldemann, 1840
Plate 62, figures 7,8; Text-figure 29r

1840. ?Skolithos Haldemann, A Monograph of the Limniades, or
freshwater univalve shells of North America, containing de-
scriptions of apparently new animals in different classes, and
the names and characters of the subgenera in Paludina and
Anculosa, No. 1, suppl., p. 3.

1847. ?Scolithos Hall, Nat. Hist, New York, Palaeont. New York,
vol. 1, p. 2, pl. 1, figs. la-c.

1881. Partim Scolithus “Hall”, James, U. P., The Paleontologist, No.
Dy ep.eaae

1892. Partim Scolithus “Hall”, James, J. F., Geol. Soc. America, Bull.,
vol. 3, pp. 32-44, figs. 1-15.

1920. ?Partim Sabellarites Richter, Rud., Senckenbergiana, Bd. 2, p.
226, pls. 1,3,4.

1921. ?Partim Sabellarifex Richter, Rud., Senckenbergiana, Bd. 3,
p. 50.

1931. ?Scolithus “Hall,’ Westergard, Sverige geol., Unders., ser. C,
Avh. och upps., No. 372 (=Arsbok, vol. 25, no. 5), pp. 14-16,
pls. 7-11.

1943. ?Skolithos Haldemann, Howell, Wagner Free Inst. Sci., Public.,
vol. 3, pp. 6-16.

1962. Skolithos Waldemann, Hantzschel, Trace-Fossils and Prob-
lematica iz Treatise on Invertebrate Paleont., Moore (ed.), pt.
W, Miscellanea, p. W215, fig. 134-4.

Type species —Skolithos linearis Haldemann (1840)
from the Lower Cambrian Chickies Quartzite between
Reading and Columbia, Pennsylvania.

Diagnosis — Closely packed, unbranched, vertical tubu-
lar penetrations which occur most frequently in arenaceous
sediments. The diameter varies from 1-10 mm; the length
from a few centimeters up to 30 cm.

Discussion — Because these tubular forms are rare in
the Cincinnatian and can only tentatively be assigned to

326 PALAEONTOGRAPHICA AMERICANA (VI, 41)

Skolithos, the genus will not be covered in detail here. For
a complete treatment the reader is referred to J. F. James
(1892), Rudolf Richter (1920, 1921), and Howell (1943).
Skolithos has received extensive coverage and is fairly com-
mon in the Lower Paleozoic rocks of eastern North America
and Europe. Specimens have been described from nearly
every continent; and it is so numerous in some localities,
e.g., Lower Cambrian of northwest Scotland, that the beds
have been called “pipe rocks.”

Skolithos was proposed by Haldemann (1840, p. 3),
who characterized it as “stem simple (never branched),
rectilinear, surface nearly even, diameter 4% - % inch,
length several feet, cylindrical or compressed.” The genus
was not figured and the precise type locality is unknown.
Howell (1943) reported that he searched in vain for a type
specimen and that probably Haldemann never selected one.
Therefore, Howell designated a neotype from Mt. Penn, a
locality which is presumably close to that of Haldemann’s
original find.

At the Mt. Penn locality S. linearis occurs in a light-
gray, poorly bedded quartz sandstone. The material filling
the tubes is either lighter or darker than the host rock and
tends to weather more rapidly, leaving pits up to 1 cm in
depth on bedding plane surfaces. The individual tubes range
from 3-7 mm in diameter and incomplete specimens attain
a length of 9 cm. All forms are normal to the bedding plane
and most are straight. Howell observed that the straight
forms occur in a fine-grained sand and that as the grain
size of the host rock increased, so did the irregularity of the
tube. Occasionally specimens appeared to branch, but
Howell believed that this was due to the intersection of
two separate individuals. The fossils are extremely close-
packed, sometimes being separated by as little as 5 mm.

Skolithos has been studied by Hall (1847), Billings
(1865), and Dawson (1890). Hall changed Haldemann’s
original spelling and described his New York forms ( Pots-
dam Sandstone) as Scolithus linearis. Most subsequent
authors erroneously believed that this is the correct spell-
ing; some workers, e.g., U. P. James (1881), even credited
the species S. linearis to Hall. However, as Howell (1943 )
and Hantzschel (1962) pointed out, Haldemann’s initial
designation prevails.

As with many fossils, the early descriptions of the
genus were so vague that Skolithos rapidly accumulated
several species; James (1892) listed 11 species from North
America alone. Rudolf Richter (1920) undertook a re-
vision and clarification of the genus. He restricted Skolithos
to straight, closely packed vertical forms best typified by
the specimens from the Cambrian of Sweden (see Richter,

1920, fig. 6). The irregular, loosely packed specimens, which
included most forms previously described under Skolithos,
were placed in a new genus, Sabellarites. The type species
was S. etfliensis from the Lower Devonian Koblenz Quart-
zite of Germany. In 1921 Richter changed the name to Sab-
ellarifex, when he realized that the earlier name was preoc-
cupied by Sabellarites Dawson (1890), an annelid. Richter’s
treatment of the genus has not gained wide acceptance.
Westergard (1931) pointed out that the Swedish specimens
from the lower Cambrian “‘Scolithus” Sandstone fail to sup-
port Richter’s conclusions. He found that some scattered
tubes were straight and some closely packed tubes were
wavy. Westergard (1931, pp. 14, 15) said, “as these differ-
ent Scolithus forms [Skolithos and Sabellarifex of Richter]
are connected by intermediate forms and occur associated,
they can hardly be distinguished .. . ”

At present the genus remains in a state of confusion.
Like so many of the larger trace fossil genera, it is badly
in need of a monographic study,

Interpretation — Most early workers, such as Halde-
mann (1840) and Hall (1847, 1852), believed that Skolithos
represented the stems of marine plants preserved in sitw.
However, shortly after the genus was proposed some au-
thors, e.g., MacCulloch (1848) and Logan (1852), suggest-
ed an annelid origin, and it was not long before the “fucoid”
theory was generally discarded. The tubes show neither
rootlets nor any trace of organic matter. Nevertheless,
over the years there have been several different hypotheses
on the origin of the genus. Perry (1872) believed that the
shorter tubes might represent brachiopod pedicle perfora-
tions, while Billings (1865) claimed to have found sponge
spicules in the filling of some tubes from the Potsdam
Sandstone of eastern Canada. Billings’ discovery has never
been confirmed and Howell (1943) was inclined to dismiss
it. Polychaetes have been frequently mentioned as pos-
sible originators. Some authors, e.g., Richter (1920, 1921),
believed that Skolithos was the tubes of filter-feeding an-
nelids, such as Sabellaria, the fan worm, Others, such as
MeNair and Reid (1929), thought that the genus was a
burrow similar to that of the Recent lug worm Arenicola.
For many years the structure of the Arenicola burrow was
not clearly understood, and not until the work of Rudolf
Richter (1924) and Wells (1945) was it realized that
Arenicola makes J- or L-shaped burrows, net simple vertical
shafts. In a convincing piece of work, Fenton and Fenton
(1934b) postulated that many Skolithos burrows could be
the tubes of phoronids. They compared the fossil forms to
the closely packed vertical tubes of Phoronopsis viridis ob-
served on the tidal flats at Elkhorn Slough, California. The

TRACE FOSSILS CINCINNATI AREA: Oscoop 3

author has seen these same tubes in Tomales Bay, Cali-
fornia, and can confirm the Fentons’ observations. The thin,
unbranched tubes are over 30 cm long and are so closely
packed that they serve to bind the sediment, producing a
topographic high. They closely resemble some illustrations
of Skolithos (e.g., see Westergard, 1931. pl. 8. fig. 1).

Some authors, for example, Hégbom (1915) and Gein-
itz (1915), concluded that Skolithos is inorganic and that
the vertical tubes mark the passage of air which was forced
to the surface when the incoming tide covered a dry sand
flat. Such phenomena are discussed at some length by
Cloud (1960).

Even though Skolithos-like forms probably have a
variety of origins, there is general agreement that the Upper
Cambrian, e.g., S. linearis, are organic. They are too straight
and in many cases too closely packed to be otherwise.
Nevertheless, some forms of inorganic origin have been as-
signed to the genus. One of these is Skolithos linearis as
described from the Cincinnatian by U. P. James (1881).
As shown elsewhere, this is a groove cast.

CINCINNATIAN REPRESENTATIVES

?Skolithos delicatulus U. P. James
Plate 62, figures 7,8; Text-figure 29-r

1881. Scolithus delicatulus James, U. P., The Paleontologist, No. 5,
pn As.

1892. Scolithus delicatulus James, U. P., James, J. F., Geol. Soc.
America, Bull., vol. 3, p. 40, fig. 15.

Type locality — Richmond strata of Dearborn County,
Indiana. The only figure of the species is that given by J. F.
James (1892, fig. 15).

Diagnosis — Small, short vertical tubes occurring in
medium-grained calcarenites which are seen on the bottom
of the slab as raised “pimples.” The “pimples,” which are
composed entirely of coarse crystalline calcite, are frequently
surrounded by a circular depression.

Discussion —U. P. James (1881, p. 33) described S.
delicatulus as “small cylindrical stems from half a line
[1/24 inch] to one line in diameter, passing vertically
through the strata irregularly arranged from ' to 4 of
an inch apart .. .” The author remarked that these oc-
curred in slabs which varied from 0.12 to over 0.5 inches
in thickness. He went on to say that “on the underside the
plants are broken off even with the surface, or leaving small,
shallow pits; on the upper side they are elevated from half
a line to over one line. Other markings on the upper sur-
face, resemble Annelid trails’ (U. P. James, 1881, p. 34).
James never figured the species, but his son, J. F. James,
1892, fig. 15) did in his paper on the nature of Skolithos.

iS)
NI

A specimen (PI. 62, figs. 7-8; UCM 37673) is present
in the University of Cincinnati collections which may well
be the form illustrated by J. F. James. It is identified as
“Scolithus delicatulus James, Cincinnati Group, Indiana”
and bears the label of the Cincinnati Society of Natural
History. The number “15” appears on one surface, and this
corresponds to the figure number in J. F. James’ paper.
Although the specimen does not exactly match James’
lithograph, it is so similar that it probably is the figured
specimen. Judging from the associated markings, it appears
that U. P. James oriented the slab incorrectly in his original
description. The raised ridges, which are fillings of trails,
show that the “pimple-"like elevations occur as convex
hyporeliefs, not convex epireliefs as he believed. The op-
posite surface represents a cleavage plane and bears the
label, “Typical character not shown on this side. Specimen
split.” Thus the upper surface expression of S. delicatulus
is unknown.

Viewed as a convex hyporelief, the specimen, which
has a surface area of 72 square centimeters, consists of over
190 raised “pimples.” The “pimples” have a diameter of
1.5 mm and are approximately 1 mm in height. They re-
semble irregular or truncated cones surrounded by a shal-
low doughnut-shaped depression which may be filled with
a soft, fine-grained material. The “pimples” themselves
are composed entirely of coarse calcite, and these same
calcite blebs can be seen on the upper surface of the speci-
men. Observation of the edge of the slab shows that the
“pimples” are actually vertical rods of calcite which com-
pletely pierce the specimen.

Interpretation. — U. P. James (1881, p. 34) believed
that S. delicatulus were plants and gave the appearance
“as if soft mud, forming the strata, had deposited gently
around the plants without disturbing their erect position.”
J. F. James (1892, p. 40) dismissed this with “. . . there
can be no question about its being a worm burrow.”

With only two small incomplete specimens from an
unknown locality to work with, it is difficult to reach a
confident decision on the nature of S. delicatulus. It is
unlikely that they represent the passageways of gas moving
up through the calcisiltite. According to Shrock (1948, p.
138) the diameter of S. delicatulus would be on the ex-
treme lower range of gas passageways. Although Twenhofel
(1921) found experimentally that gas continuously rising
through mud could maintain open tubes which resemble
those produced by worms, it is unlikely that they would
remain open long enough to be filled by calcite. More-
over, gas tubes illustrated by Cloud (1960, pl. 3) are some-
what irregular in outline.

328 PALAEONTOGRAPHICA AMERICANA (VI, 41)

It is probable that S. delicatulus is the filling of a por-
tion of a slender polychaete or phoronid tube. Many tubi-
culous polychaetes either secrete parchment-like tubes or
construct agglutinated tubes which are cemented by mucus.
Phoronids employ the latter method. Such a tube would
prevent collapse of the walls after the death or departure
of the organism, If the opening of the tube were plugged
with sediment, the void could later be filled by calcite.
In all probability the specimen at hand was underlain by
mud and represents only a portion of a much longer tube.
Because the base of the tube is missing, it is not possible
to definitely assign the species to Skolithos. It is impossible
to determine whether one is dealing with unbranched ver-
tical tubes, U-tubes, or even a portion of Chondrites. The
latter appears to be unlikely, for as shown in Plate 64,
figure 6, Chondrites tunnels many times occur in groups
and are never filled with calcite.

VII. FODINICHNIA

CHONDRITES Sternberg, 1833

Plate 63, figures 2-5; Plate 64, figures 2-8; Plate 65, figure 8; Plate
67, figure +; Plate 69, figures 1,8; Plate 79, figures 5-6; Plate 82,
figure 4; Plate 83, figure 2; Text-figures 10,11,29s,t,u

1823. Partim Fucoides Brongniart, Soc. Hist. Nat. Paris, Mem., vol. 1,
p. 308, pl. 19, fig. 2.

1833. Chondrites Sternberg, Versuch einer geognostisch-botanischen
Darstellung der Flora der Vorwelt, Bd. 7-8, p. 25.

1833. Partim Sphacrococcites Sternberg, Versuch einer geognostisch-
botanischen Darstellung der Flora der Vorwelt, Bd. 7-8, p. 28,
pl. 4, fig. 1.

1833. Partim Caulerpites Sternberg, Versuch einer geognostisch- bot-
anischen Darstellung der Flora der Vorwelt, Bd. 7-8, p. 20.

1840. Non Skolithos Haldemann, A Monograph of the Limniades, or
freshwater univalve shells of North America, containing de-
scriptions of apparently new animals in different classes, and
the names and characters of the subgenera in Paludina and
Anculosa, vol. 1, supp., p. 3.

1847. Partim Buthotrephis Hall, Nat. Hist. New York, Paleont. New
York, vol. 1, p. 8, pl. 2, fig. 6.

1849. ?Phymatoderma Brongniart, Végétaux fossiles, p. 59.

1852. Partim Buthotrephis Hall, Nat. Hist. New York, Palaeont. New
York, vol. 2, p. 18, pl. 5, figs. 1la-d.

1858. Non Dendrograptus Hall, Geol. Sur. for Canada 1857, Report
Progr., 1858, p. 143; Report figs. a-c.

1873. Non Planolites Nicholson, Roy. Soc. London, Proc., vol. 21, p.
289.

1874. Buthotrephis Hall, Miller, Cincinnati Quart. Jour. Sci., vol. 1,
peso) aie.

1878. 2Scolithus Hall, Miller and Dyer, Contributions to Palaeontol-
ogy, No. 2 (Cincinnati, Ohio, private publication), p. 4, pl. 4;
fig. 4.

1885. Partim Dendrograptus Hall, James, J. F. Cincinnati Soc. Nat.
Hist., Jour., vol. 7, pp. 159-161.

1891. Partim Planolites Nicholson, James, J. F., Cincinnati Soc. Nat.
Hist., four., vol. 14, p. 17.

1895. Chondrites Sternberg, Fuchs, Akad. Wiss. Wien, math.-nat. KI.
Denkschr., Bd. 62, pp. 407-410, figs. 10-12, pl. 9, fig. 3.

1910. Chondrites Sternberg, Reis, K. K. geol. Reichsanst., Jahrb.,
Bd. 59, pp. 615-638, pl. 17.

1948. Clematischnia Wilson, Canada Dept. Mines Resourc., Geol.
Sur. Bull., vol. 11, p. 10.

1949. Chondrites Sternberg, Tauber, Geol. Bundesanst., Jahrb., Bd.
93, pp. 141-154, 3 figs.

1951. Chondrites Sternberg, Seilacher, “Zur Einteilung und Deutung
fossilen Lebensspuren” (unpublished Ph.D. dissertation Tiibin-
gen University, West Germany).

1957. Chondrites Sternberg, Simpson, Geol. Soc.
Jour., vol. 112, pp. 475-500, pls. 21-24.

1962. Chondrites Sternberg, Hantzschel, Trace-Fossils and Problem-
atica in Treatise on Invertebrate Paleont., Moore (ed.), pt.
W, Miscellanea, p. W187, fig. 115-1.

London, Quart.

Type species. — Fucoides targionti Brongniart (1828)
from the Eocene Flysch of Switzerland. Lectotype, Andrews
(955i):

Diagnosis — Tunnel systems possessing a small number
of master shafts open to the surface which ramify with
depth to form a dendritic network. The individual branches
may or may not cross over and interpenetrate,

Discussion — Chondrites is one of the most common
and notable of all the “fucoids.” It ranges from the Ordo-
vician through the Tertiary and has been found at numer-
ous localities in North America, Europe, and Asia. The
genus is probably best developed in the Devonian and
Jurassic of western Germany and the Tertiary Flysch of
the Alps, and several forms have been described by Hall
(1847, 1852) from the Lower Paleozoic of New York State.
Although Chondrites has received extensive coverage by a
number of authors, most notably Richter (1927, 1931),
Tauber (1949), and Simpson (1957), unanswered questions
still remain, At present the genus contains such a wide
variety of morphologic expressions that a detailed analysis
would probably result in the establishment of several gen-
era.

Chondrites occurs in sediments ranging from fine-
grained Jutites to medium-grained arenites, and, as Seil-
acher (1964, fig. 7) showed it is a fossil which is common
to several different environments. It has been found in
such diverse settings as the black Hunsriick Shale of the
Rhineland Devonian and the Carboniferous calcarenites of
Great Britain. Although Chondrites can occur individually,
more frequently it is encountered in great numbers where
it forms dendritic patterns on bedding plane surfaces or
riddles the sediment at depth with a large number of tunnels
(Simpson, 1957, pl. 21, figs. 1, 2).

The tunnel system is normally either oblique or paral-
lel to planes of stratification. Simpson (1957) mentioned
that a system may proceed obliquely for a distance and
then flatten out along a bedding plane. He added that
even in forms which are oblique throughout their entire

‘TRACE FossILS CINCINNATI AREA: OsGoop 329

length, the distal portions tend to be more nearly parallel
to the bedding planes.

The diameter of the tunnels varies from 0.5 mm in
C. intracatus (Brongniart) (Austria: Eocene) to nearly 5
mm in C. gracilis var. crassa (Hall) (New York: Silurian).
As illustrated by forms occurring in medium-grained sedi-
ments, the original cross-section of the tunnels was circular.
However, specimens which are found in finer grained ma-
terials are often depressed and present an ellipsoidal out-
line. The most extreme examples are to be seen in C. bollen-
sis (Zieten) (Wiirttemberg: Jurassic) (PI. 79, fig. 6),
where the tunnels are almost paper thin. In nearly all forms
the diameter tends to remain constant throughout the en-
tire length of the tunnel system; however, unconnected sys-
tems of differing diameters can occur in close proximity
(see Rothpletz, 1896).

The branching pattern of Chondrites could best be
termed random or alternating. Regular patterns, e.g., sym-
podial, dichotomous, are unknown. Normally branching
proceeds to the second or third order, although Rudolf
Richter (1927, fig. 7) showed a form with fifth or sixth
order branching. As seen in the specimen of C. bollensis
illustrated on Plate 79, figure 6, the branching tends to be
alternating, and the angle of bifurcation varies from 25-
40°. Based on a study of several European species, Simpson
(1957) found that the angle was consistently 35-45°. Al-
though Simpson (1957, p. 483) reported that the “tunnels
are straight [giving] a characteristic rigid appearance to
the pattern” (as seen in PI. 65, fig. 8 and noted by Simpson,
1957, pl. 23, fig. 3), some of the tunnels are gently arcuate.
A remarkable fact and one most significant in the interpre-
tation of the genus is that in many forms the branches
rarely interpenetrate. This was first demonstrated by
Richter (1927) and later confirmed by Derichs (1928) and
Simpson (1957). If two branches are headed on a “collision
course,” one will terminate before contact is made.!® In ad-
dition, the authors mentioned above have shown that the
proximal branches of the system consistently abut against
the distal branches. This kind of phobotaxis is best devel-
oped in specimens preserved on bedding planes in the Lias
of southwestern Germany (PI. 79, fig. 6) and the Devonian
Hunsriick Shale (see Richter, 1931, figs. 2-5). The kind of
phobotaxis described above refers to forms occurring on one
plane only. Another type of phobotaxis is present in three
dimensional forms, where two branches cross over but do

"Richter (1927) employed the term phobotaxis (fear of touching) to
describe the avoidance of contact in these chondritids. The meaning
of the term as used in trace fossil studies is discussed in greater
detail below.

not interpenetrate. However, when Chondrites occurs in
fine-grained sediments, compaction might well close the
small gap separating the two tunnels and give the impres-
sion of “pseudopenetration.” Thus in practice it is difficult
to distinguish between crossing over and actual interpene-
tration.

Phobotaxis is not found in all representatives of the
genus. Some forms, described by Hall (1852, pl. 5a, fig. 1;
pl. 5, figs. 2B, 3C) under the name Buthotrephis from the
Silurian of New York State and observed by the present
author, exhibit interpenetration. Simpson (1957, p. 492)
mentioned that in specimens from the Lias of Great Brit-
ain, “separate systems may intersect one another but when
this happens the two systems are never of the same colour.”
As will be shown, Cincinnatian forms exhibit only an incip-
ient phobotaxis, and interpenetration is relatively common.
A search of the literature was undertaken to determine if
the two forms of phobotaxis were somehow restricted by
time or geography. While the possibility of “pseudopenetra-
tion” and the highly schematic diagrams of the earlier work-
ers makes such a determination difficult, there does not
appear to be any pattern. The phenomenon of phobotaxis is
not restricted to a certain horizon or location.

The controversy regarding the origin of Chondrites
has in large part revolved around the nature of the ma-
terial filling the tunnels. In some forms, ¢.g., those from the
Holzmaden Jurassic section (PI. 76, fig. 6), the filling is
much lighter than the host rock, whereas in others, e.g., C.
paleozoicus Richter from the Hunsriick (see Rudolf Richter,
1931, figs. 2-5), the filling is darker and stands out in
bold relief against the lighter color of the host. In Cincin-
natian forms the filling is normally lighter than the rock
enclosing it. For many years it was assumed that the black
material filling the tunnels of many European species was
carbonaceous in nature as simple tests for carbon gave posi-
tive results. Some early authors even claimed to be able
to distinguish “cells” or “traces of parenchymal tissue.”!7
Several workers, e.g., Rothpletz (1897) and Giimbel (1897),
conducted more careful chemical analyses and found that
only 2-4 per cent carbon was present, and most of this was
bound up in silicates. The most detailed analysis is that of
Tauber (1949), who analyzed the fillings of Chondrites
furcatus (Brongniart) as it occurred in fine sands, lime-
stones, and marls. His results are shown in Table I.

As evident in Table I, Tauber found that the material
filling the tunnels differed considerably from that compris-

“The quotes are those of Lessertissuer (1955, p. 66), who did not
identify the original author of the statements.

330 PALAEONTOGRAPHICA AMERICANA (VI, 41)

ing the host rock. The tunnels were poor in calcium car-
bonate and showed only a trace of carbon, Tauber attrib-
uted the black color to an enrichment in iron sulfide. In
contrast, Rothpletz (1896) found that specimens of C. fur-
catus from the Jurassic Posidion Shales would effervesce,
whereas the host rock would not.

The only species of Chondrites possessing a filling
which shows any trace of structure is C. granulatus (Schlot-
heim) (Wiirttemberg and Great Britain: Lias; Alps: Eocene
Flysch), and even this is questioned by Simpson (1957).
Because the argument is based largely on interpretation,
it will be discussed below.

TABLE [I
Composition or Chondrites furcatus
Limestone

Filling Host Rock
Carbonate as CaCO; 2.50 $3.03
Iron sulfide as FeS 16.58 2.81
Clay minerals, quartz, mica 80.92 14.16
Coal flakes trace trace
100.00 100.00

Marl

Filling Host Rock
Carbonate as CaCO; 2.73 36.66
Iron sulfide as FeS 14.54 4.83
Clay minerals, quartz, mica 72.72 58.51
Coal flakes trace trace
99.99 100.00

Fine Sandstone
Filling Host Rock

Carbonate as CaCOs 0.37 4.88
Iron sulfide as FeS 18.63 5.90
Clay minerals, quartz, mica 80.99 89.22
Coal flakes trace trace

100.00 100.00

(from Tauber, 1949, Table 1)

Reconstructions of the over-all pattern formed by the
tunnel systems of Chondrites vary considerably. Simpson
(1957, fig. 2) illustrated an idealized form of Chondrites
in which he envisioned six or seven master shafts which
are bundled together proximally. These diverge with depth
and give rise to secondary, tertiary, and quaternary
branches. The main branching pattern is slightly oblique
to the bedding plane and as viewed from above, is radial
in outline, In contrast, Tauber (1949, fig. 1) in a recon-
struction of C. furcatus (Austria: Tertiary) portrayed a
single master shaft which branches repeatedly to produce
an antler-like pattern. Going one step further, Tauber (1949,
fig. 3) showed a series of such patterns arranged in tiers.
This latter configuration is similar to a figure of C. bollensts
(West Germany: Lias) given by Rudolf Richter (1931,
fig. 1). Richter (1931, fig. 10) figured one specimen of

Chondrites sp. from the Eocene of Austria which shows
14-16 such tiers, each separated by 1.5 mm.

Fuchs (1895, fig. 11) illustrated an unnamed form
from the Cretaceous Flysch of Rignano which is unusual
in that it possesses a crude bilateral symmetry. When
viewed from aboye, it resembles a lyre with numerous sec-
ondary branches. Finally, Liberman (1900, pl. 2) and
Papp (1941, fig. 2) showed figures of Chondrites which are
arranged in a stellate pattern.

Since the orientation of many specimens is oblique to
the bedding planes, it is difficult to accurately establish the
dimensions of an entire network. The longest branch found
by Simpson (1957) was 15 cm, and he estimated that the
ratio between the length of a branch and the diameter of
a tunnel was 40:1. It would not be surprising to find single
specimens which cover an area as great as one square meter.

As shown above, Chondrites contains such a wide range
of morphologic variations that at present it is a form genus
in the widest possible sense. However, a complete restudy
of the genus will be an undertaking of some magnitude, be-
cause it contains more “species” than any other “fucoid.”
Bassler (1915) listed 29 for the North American Ordovician
and Silurian alone, and the number of species described
from Europe is much greater. As accepted by recent authors,
Chondrites is in part synonymous with Fucoides Brong-
niart (1823). In 1828 Brongniart described and figured 36
species of Fucoides, 11 of which appear to be chondritids.
The remaining 25 fall into several diverse groups, and to-
day the “genus” Fucoides has little meaning. As nearly as
can be determined from the schematic figures, Chondrites
is also partly synonymous with Caulerpites Sternberg
(1833) and Sphaerococcites Sternberg (1833). The former
genus, as originally recognized by Sternberg, contains chon-
dritids, various trails, and possibly even conifers (Hantz-
schel, 1962, p. 187). Likewise, Chondrites is partly synony-
mous with Buthotrephis Hall (1843), originally described
from the Clinton beds (Silurian) of New York State. Hall’s
type specimens are fragmentary and reveal only portions
of what were undoubtedly much larger systems. However,
a restudy of some of his types shows that Buthotrephis
gracilis (Pl. 69, fig. 8), as well as B. gracilis var. crassa
(Pl. 63, fig. 5; USNM 41134), are chondritids.18 As dis-
cussed herein under Phycodes, B. palmata (Pl. 67, fig. 7;
USNM 41079) is not a chondritid but a bundled burrow

“Over a period of nine years Hall proposed nine species and varieties

of Buthotrephis. With the exception of the forms mentioned above,
these are not considered in the present study, because they were not
listed as being present in the Cincinnatian. Judging from Hall's il-
lustrations, many of them appear to be chondritids.

‘TRACE FOossiLs CINCINNATI AREA: OscGoop 3

similar to a form described by Seilacher as Phycodes pal-
mata (1955, p. 383) from the Cambrian of the Salt Range.
Buthotrephis is discussed more completely below.

Interpretation — More than any other genus Chon-
drites is the embodiment of the term “fucoid.” Its dendritic
pattern so closely resembles some Recent algae (¢.g., Deles-
seria) and higher plants that it is not surprising that it was
the “last of the fucoids.”’ As late as the middle of the pres-
ent century, Wilson (1948) and Maubeuge (1950) still
accepted an algal origin.

Nathorst (1881) was one of the first authors to offer
a non-algal hypothesis. Basing his conclusion mainly on
forms from the Alpine Flysch, he reasoned that it was un-
likely that plants would grow in such deep water where
even body fossils are rare. In addition he remarked that
there was no yestige of organic material, and the genus
occurred in rock types which could offer no firm founda-
tion for attachment. Nathorst (1881, p. 96) interpreted the
genus as the “casts of the ramified conduits of worms” but
did not elaborate. Likewise, Fuchs (1895) believed that all
chondritids are non-algal. He interpreted them as brood
chambers which were later filled by sediment sifting into
the tunnel system from above. One of his strongest argu-
ments was that the fillings lack the high carbon content
which one would expect to find in plant fossils. According
to Fuchs the black color of the fillings was caused by coal
particles washed in from above. Rothpletz (1896) criti-
cized Fuchs’ views and, after conducting chemical analyses
on some forms, concluded that the Flysch specimens were
true algae, while the Jurassic forms from Wiirttemberg
were sponges. A sponge origin was also postulated by Mail-
lard (1884) and Walther (1904). Reis (1910), working
with Cretaceous forms from the Appenine Flysch, for the
most part confirmed Fuchs’ observations. He pointed out
the lithologic similarity between the fillings and the beds
overlying the host rock and concluded that Chondrites was
a burrow. The cowp de grace was delivered by Richter
(1927), who was able to demonstrate the presence of phobo-
taxis in C. bollensis (Pl. 79, fig. 6) as it occurs on bedding
planes in the Wiirttemberg Lias. As mentioned above,
phobotaxis was later confirmed by Derichs (1928) and
Simpson (1957).

Although Chondrites 1s now generally accepted as a
trace fossil, there is still considerable disagreement concern-
ing its ethological interpretation. Rudolf Richter (1927,
1931) concluded that it was the burrow system of a Sedi-
mentfresser which was later filled by sediment sifting in
from above. Phobotaxis, presumably due to some variety of
chemoreceptor, enabled the animal to achieve maximum
efficiency by avoiding areas which had already been

w

worked. To Richter the fact that some networks were ar-
ranged in tiers demonstrated that the nutrients were con-
centrated in thin layers rather than spread evenly through-
out the host rock.

In contrast to Richter’s theory, Tauber (1949) main-
tained that Chondrites was the burrow of a filter-feeding
annelid and that the fillings represented a combination of
fecal packings and collapse of the lining of the tube. He
believed that the tierlike arrangement was a response to
increased sedimentation which forced the organism to ele-
vate the entire system. He cited the branched burrows of
the Recent annelids Pygospio and Nereis as evidence for
his interpretation. Both these organisms employ iron oxides
to cement the slime covered walls of the burrow. Tauber
concluded that the iron was extracted from plankton in-
gested by the organism, However, Tauber’s explanation
fails to explain several facts. If, as Tauber maintained,
Chondrites was the Domichnia of a filter-feeder, why was
it necessary for the organism to construct such an elabo-
rate system which in many specimens follows the bedding
planes and exhibits phobotaxis? Moreover, Tauber’s ex-
planation of the tierlike arrangement of some chondritids
is not convincing. The burrows of neither Nereis nor Pygo-
spio appear to resemble Chondrites. According to Schafer
(1962), Nereis normally lies in a long (up to 40 cm) un-
branched burrow with its head near the depositional inter-
face. Only if the water is low in oxygen does the worm con-
struct several accessory shafts which open on the surface
to increase free circulation of water through the burrow.
As illustrated by Schafer (1962, fig. 166), this does not
have the organization of the Chondrites network. Pygospio
is an annelid which feeds either by extending its tentacles
over the substrate or by constructing a slime net to catch
plankton. Schafer (1962, p. 388) reported that the organism
dwells in a small tube which may reach a depth of 9 cm.
According to Linck (1939), the burrow is branched only
when it has been partly plucked out by current action and
is torn. The animal then constructs a secondary branch
at the site of the tear and, depending on the location of
the tear, the auxillary branch may either be a blind shaft
or open to the surface. This certainly is not homologous
with Chondrites.

Simpson (1957) took issue with the belief that Chon-
drites represented a stuffed burrow. In the British ex-
amples (Lias) of C. granulatus he was able to demon-
strate that the pellets found in the burrow were derived
from the overlying beds. Simpson’s interpretation of the
meaning of Chondrites was similar to that of Richter (1927,
1931), the major difference being the meaning of phobo-
taxis. Simpson (1957, p. 485) believed that “the key to

~
we
bo

the problem lies in the fact that the animal was working
from a fixed point on the surface. By first exploring out-
wards to the maximum distance permitted to it, and then
working sideways and backwards, the animal was able to
ensure that the unexploited territory lay always on the near
side of the last sideways exploration. Thus the phobotaxic
reaction was not employed so much in preventing the ani-
mal [from] penetrating into already exploited territory
as in enabling it to keep contact with the near limit of
such territory.” Simpson was the first to suggest that Chon-
drites might be the burrows of sipunculids, although he
admitted that his belief could not be confirmed by Recent
examples. While such a hypothesis requires an organism of
considerable length, Hyman (1959) reported that some
sipunculids attain lengths of 30 cm and enteropneusts range
from 10-50 em. Barnes (1963) stated that most annelids
are 5-10 cm long, although forms up to 3 meters in length
are known.

Recent examples which resemble Chondrites are un-
known, As mentioned above, the burrows of Pygospio and
Nereis are different, as are those of other Sedimentfressers
such as Heteromastus (see Schafer, 1962, fig. 159) and
Balanoglossus. This seems unusual since Chondrites is so
abundant in the fossil record and occurs in such a wide
range of depositional environments. It is unlikely that our
lack of knowledge concerning Recent environments provides
the entire answer. Seilacher’s (1957) belief that many trace
fossils become clearly visible only after they have under-
gone diagenetic alteration may provide a clue to the prob-
lem.

CINCINNATIAN REPRESENTATIVES

There are at least three types of Chondrites present
in the Cincinnatian. Unfortunately, none of the three can
be confidently placed in the trace fossil or “fucoid” taxa
discussed by any of the local authors. A search of the
literature reveals that the local strata contain two genera
and five species which either resemble chondritids or were
actually assigned to taxa today considered synonymous
with Chondrites. J. F. James (1885) maintained that three
of Hall’s species of “Buthotrephis,” originally described from
the Lower Paleozoic of New York State, could be identi-
fied in the Cincinnatian. These were “B.” gracilis Hall
(1843), “B.” gracilis var. crassa Hall (1852), and “B.”
succulens Hall (1852).

Although it was not possible to locate the type speci-
men of “B.” gracilis Hall (1843, p. 69, fig. 14), four hypo-
types (Hall, 1852, pl. 5, figs. la-d) from the Clinton beds
of New Hartford, Oneida County, New York, were avail-

PALAEONTOGRAPHICA AMERICANA (YI, 41)

able for study. The specimens are so similar that only one
form is illustrated herein (Pl. 69, fig. 8; USNM 41125; latex
mold UCM 37676).1° “B.” gracilis is preserved as an epi-
relief on a fine-grained, iron-stained micaceous quartz aren-
ite. The most striking feature of the species is its small
size; the branches have a diameter of only 0.5 mm and the
length of the longest branch is but 4 cm. Branching pro-
ceeds to the fourth order and the angle of bifurcation varies
between 40-55°. Interpenetration of the branches is pres-
ent but is not common, and contrary to Hall’s illustrations,
the branches do not taper distally. The species agrees with
Simpson’s observation of European chondritids as the
branches are straight and give the arborescent pattern a
rigid appearance. The material filling the tunnels is both
darker and finer grained than the host rock. There is no
evidence of fecal pellets or other internal structures. As in
all chondritids the contact between the filling and the host
rock is sharply defined.

“B.” gracilis “var.” crassa Hall (1852), also from the
Clinton beds of New Hartford, New York, is poorly known.
Although Hall (1852, pl. 5, figs. 3a-d) figured what appear
to be complete forms, the actual specimens (Pl. 63, fig. 5;
USNM 41134) are incomplete. The specimens of “B.”
gracilis “var.” crassa observed by the present author are
preserved as epireliefs on a platy, micaceous, fine-grained
quartz arenite. The irregular main branch has a diameter
of 5 mm and the angle of bifurcation varies from 35-60°.
In some cases the secondary branches have a diameter which
is slightly less (4 mm) than that of the main branch. The
filling is slightly darker and finer grained than the host
rock, and although Hall described the surface of the
branches as nodose, the surface of all observed forms was
smooth. There is no trace of granules or pellets similar to
those described in Chondrites granulatus by Simpson
(1957). The type specimens are so fragmentary that the
over-all pattern of “B.” gracilis “var.” crassa can not be de-
determined. Hall (1852, p. 19) remarked that “. . . it varies
in form so much that scarcely two specimens are precisely
alike.”

Hall (1852, pp. 18, 19) believed that “B.” gracilis and
“B.” gracilis “var.” crassa represented the end members of
a completely intergraded series of Clinton “fucoids.” Based
solely on a study of the type material, it is impossible to

“Hall first used the specific name “gracilis” in 1843 for Fucoides gra-
cilis from the Clinton strata of New York. In 1847 (pl. 21, fig. 1)
he employed the same species name for “Buthotrephis” gracilis from
the Trenton beds of Herkimer County, New York. However, in
1852 he changed the specific name of the Trenton specimens to “B.”
tenuis and retained “B.” gracilis for the Clinton forms only.

TRACE FOSSILS CINCINNATI AREA: OsGoop

support his conclusion, “B.” gracilis is small and has a rigid
arborescent pattern, while “B.”’ gracilis “var.” crassa is
much larger and more irregular. The former closely re-
sembles on a smaller scale the pattern of the “typical”
European chondritids. The latter has the general appear-
ance of a chondritid, but its over-all pattern can be re-
vealed only through a study of topotype material. As will
become evident later, neither of these two “species” closely
resembles the Cincinnatian chondritids.

It is apparent from J. F. James’ (1885, p. 159) discus-
sion that he never saw Hall’s specimens of “B.” gracilis or
“B.” gracilis “var.” crassa. He concluded that the two forms
were conspecific with Psilophyton gracillimum Lesquereux
which he placed in Dendrograptus Hall, believing them to be
graptolites. James was misled by Hall’s figures of “B.” gra-
cilis which do resemble Lesquereux’s (1878, pl. 1, fig. 2)
illustration of P. gracillimum. As the above discussion of
“B.” gracilis and “B.” gracilis “var.” crassa shows, there is
nothing to support a graptolite assignment for the species.
On the other hand, P. gracillimum is judged to be a grapto-
lite and is discussed herein under Mastigograptus Ruede-
mann.

Whereas J. F. James (1885) believed that “B.” suc-
culens Hall (1847, pl. 22, figs. 2a, b) could be identified
in the local Cincinnatian section, an examination of the
more complete of the two syntypes shows that it can not
be. The specimen (PI. 69, fig. 1; USNM 41133; latex mold
UCM 37639) from the Trenton Limestone of Glens Falls,
New York, is preserved as an epirelief on a black litho-
graphic limestone. The basal portion of the host rock is a
coarse-grained calcarenite. The specimen is large and sub-
scribes an arc of almost 180°; the length of the longest ob-
served branch is 12 cm. The nearly cylindrical individual
branches are slightly sinuous and have a diameter of 4-5
mm. Fourth order branching was observed and the angle
of bifurcation is between 30-45°. Interpenetration of the
branches is not common. Although the fillings are also com-
posed of dark, fine-grained lithographic limestone, the con-
tact between the fossil and the host rock is well defined.
Where the filling has been removed, a thin sheath of limon-
ite is exposed. “B.” succulens is best characterized by peri-
odic swellings which occur at 2-4 mm intervals and give
the branches an irregular nodose appearance. Because the
filling reveals no trace of spherical bodies, the swellings are
thought to be of prior origin.

Wilson (1948) selected “B.” succulens as the type
species for her monotypic genus Clematischnia. She rede-
scribed Hall’s figured specimens as well as additional ma-

terial from the Cobourg (Middle Ordovician) beds of On-

as)
ws
we

tario. She concluded that the genus was a “fucoid” and
cited the “. . . solid, slender stems and undulating
ridges” as the diagnostic features (Wilson, 1948, p. 11).
Other than the arborescent pattern, the present author sees
no evidence to support a vegetable origin. Instead the form
is thought to be chondritid-like burrows, and the limonite
sheath probably represents the vestiges of a mucus-coated
wall. The origin of the nodular wall structure is problematic,
for the swellings would seem to be too widely spaced and
irregular to be attributed to peristaltic movements of a
vermiform body. In general appearance the species lies
midway between “Buthotrephis” gracilis Hall and “B.”
palmata Hall.*° Whether the nodose walls of “B.” succulens
are of generic or specific importance can only be determined
through a monographic study of Chondrites. In its present
state Chondrites has sufficient latitude to permit inclusion
of “B.” succulens within the genus, In any event the form
is unknown in the Cincinnatian, for the walls of Cincin-
natian chondritids are either smooth or bear faint longi-
tudinal striae.

Miller (1874, pp. 235, 236, fig. 29) proposed “Butho-
trephis” ramulosus for a form which he described as “fossil
ramose; stems branching; branches bifurcating, with very
little change in size. Diameter of stems and branches from
1/16th to 1/10th inch. Structure not distinct, but has a
granular appearance, as if from some change in its charac-
ter.” The specimens occurred in “clay nodules, or indurat-
ed marl stones, [and] form entire layers of strata for hun-
dreds of feet in length, between six and twelve feet above
low watermark, under the bank of the Ohio River.” He in-
terpreted it as a “fucoid.”

J. F. James (1885) believed that the species was a
burrow, but from the manner of his discussion, it is appar-
ent that he never saw an identified specimen.

Both Fuchs (1895) and Reis (1910) discussed “B.”
ramulosus and compared it to the tubes of the Recent an-
nelid Lanice conchilega. Reis remarked that “B.” ramulosus
had a tunic of pelecypod shell fragments, and Seilacher
(1955) has shown that L. conchilega does construct a U- or
W-tube composed of mucus-cemented shell fragments.

Unfortunately the three localities mentioned by Miller,
which are probably Lower Eden and Upper Cynthiana, are
now permanently covered by a rise in the level of the Ohio
River due to the installation of navigation dams. Identified
specimens are not present in Miller’s collections at Harvard
or the Field Museum, nor could they be found in the Uni-
versity of Cincinnati collections. His description and figure

““See Phycodcs herein for a discussion of “B.” palmata.

do resemble Cincinnatian chondritids with the exception
that none of the local chondritids possesses a “granular ap-
pearance.” Moreover, Miller remarked that the species 1s
restricted to the one horizon. Each of the Cincinnatian
chondritids here discussed is known from various localities
and horizons. It is probable that Reis (1910) based his
interpretation on Miller’s figures. No burrows with a lining
of shell fragments have yet been discovered in the local
rocks. In conclusion, while it is possible that “B.” ramu-
losus is a chondritid which is restricted to basal Cincin-
natian, it is also possible that it is a chondritid similar
to those discussed below, except that it is filled with a
coarse-grained sediment.

Scolithus tuberosus Miller and Dyer (1878b) is also
possibly a chondritid. They described the species as “the
holes (sometimes called stems) are curved or winding,
and pass through the rock, in an irregular course, sometimes
uniting or branching, but never passing vertically through
the strata as in S. linearis from the Potsdam Group [Upper
Cambrian]. Upon the upper surface of the rock, the tubes
are prolonged into crateriform elevations which are rarely
at right angles to the surface of the rock” (Miller and
Dyer, 1878b, p. 5). They went on to say that the diameter
of the holes varies from 4-7 mm but remains constant in a
given form. According to the authors, while the form is
more abundant in the lower part of the Cincinnatian, it
occurs throughout the section. They interpreted it as a bur-
row and stated that any comparison with Scolithus linearis
was fallacious. J. F. James (1892, pp. 38, 39), in his mono-
graphic study of Skolithos, pointed out that S. tuberosus
differed considerably from the accepted concept of the
genus. He suggested that it be proposed as the basis of a
new genus.

Identified material of S. twberosus has not been locat-
ed, and Miller and Dyer’s illustration shows only the
“crateriform elevations” which they likened to “the mud
elevations, thrown up around the holes, made by the com-
mon crawfish [sic] [Cambarus diogenes] on our fresh-
water streams” (Miller and Dyer, 1878b, p. 5). While their
description of the species resembles what will be discussed
below as Chondrites, type-B, the diameter of the branches
of S. tuberosus is nearly twice as great, and no “crateriform
elevations” have ever been found associated with Cincin-
nation chondritids. Thus, while it is possible that S. tubero-
sus is a chondritid, it is just as likely some variation of
Palaeophycus. (See discussion of Palaeophycus.) Little
more can be said until identified material is located.

U. P. James (1878, p. 9) proposed Buthotrephis filci-
Jormis for specimens from “near Lebanon, Ohio, middle

334 PALAEONTOGRAPHICA AMERICANA (YI, 41)

beds of Cincinnati Group.” The species was never figured,
but it is clear from his description that this is not a chondri-
tid. Instead it is thought to be synonymous with “Chloe-
phycus plumosum” Miller and Dyer (1978b) and is dis-
cussed thereunder.

The above discussion indicates that earlier authors did
recognize the presence of Chondrites in the local section.
However, the original descriptions and illustrations are in-
adequate and prohibit a meaningful comparison with the
material at hand. It is equally as difficult to place the Cin-
cinnatian specimens studied by the author in any of the
myriad European species. It is concluded that under the
present circumstances little would be gained from establish-
ing yet additional species of Chondrites. Therefore, the
three Cincinnatian “varieties” are considered below as
Chondrites, types A, B, and C. Of these three, Chondrites,
type-A appears the most likely to stand up under a detailed
comparison with other chondritids.

Chondrites, type-A

Plate 64, figure 6; Plate 67, figure 4+; Plate 83, figure 2; Text-figure
10,29-a

Discussion — The forms included in Chondrites, type-
A are abundant in the fine-grained calcarenites of the upper
part of the McMicken member on U.S. Route 27, 1.4 miles
south of Newport Shopping Center, in Taylor Creek where
crossed by Wesselman Road, Hamilton County, Ohio, and
also of the type locality of Corophioides cincinnatiensts.*
They have also been recovered from the platy, fine-grained
calcisiltites in the Southgate beds at Newport Shopping Cen-
ter. Chondrites, type-A, has not been encountered elsewhere
in the Cincinnatian section. At the three McMicken locali-
ties mentioned above, the specimens are associated with
Trichophycus venosum, Corophioides cincinnatiensis, C. cf.
luniformis, and Chondrites, type-B. Chondrites, type-A, dif-
fers from all other Cincinnatian chondritids in that the
entire system is small and compact, having its greatest di-
mension directed vertically rather than horizontally. As
seen in vertical section (PI. 67, fig. 4; Pl. 83, fig. 2), the
system resembles a series of rootlets. Frequently the speci-
mens extend through the entire thickness of the bed and
into the subjacent lutite, although others terminate before
reaching the base of the host rock. The longest (7 cm) form
encountered was incomplete. The individual branches, which
are only slightly depressed, have an average diameter of
0.8 mm. As in the chondritids studied by Simpson (1957),

“See the discussion of Corophioides cincinnatiensis for detailed local-
ity data on these occurrences.

TRACE FOSSILS CINCINNATI AREA: Oscoop

the diameter of the branches does not change within a sys-
tem. As seen in vertical sections the tunnels vary consid-
erably in their attitude. The angle formed by the branches
and the ill-defined central axis ranges from 20-50°. While
some branches continue at this attitude for their entire
length, others flatten out and proceed parallel to the bed-
ding planes. In contrast other branches (PI. 83, fig. 2) are
deflected even more strongly downwards. The highest order
of branches observed was secondary. There is no evidence
of phobotaxis and the tunnels occasionally interpenetrate.

LA "aN

G

Text-figure 10.— Chondrites, type-A. Reconstruction showing a
complete network. There is no well-defined central axis; X 2.

When viewed on bedding plane surfaces, Chondrites,
type-A, appears as a series of numerous, closely spaced holes
arranged in a circular pattern. Plate 64, figure 6 illustrates
one slab from the Southgate strata of Newport Shopping
Center which bears seven separate systems on an area of
105 square cm. These can be readily distinguished from the
other chondritids which occur in the same bed by the close
spacing and small size of the tunnels. Serial sections taken
parallel to the bedding plane demonstrate that there is no
marked increase in the over-all diameter of the system with
depth as is found in most chondritids, The over-all diameter
of one representative form measured 8 mm at the top of the
bed and had expanded to only 1.5 cm at a depth of 4.5 cm.
Serial sections also show that there is no well-defined central
axis, although the perforations may be more closely spaced

Ss
+
wal

in this area. As seen in Text-figure 10 it appears that first
one branch then another served as the axis.

The sediment filling the tunnels is both finer grained
and lighter in color than that of the host rock, As can be
seen in Plate 67, figure 4, the fillings frequently weather
out when exposed to the elements; the contact between the
tunnel wall and the host rock is always sharply defined.
X-ray defraction analysis of fresh samples of host rock and
filling shows significant mineralogical differences discussed
in detail below.

The nature of the upper limit of the burrow are un-
known, for in all observed cases the surface of the host rock
bore patterns similar to that seen in Plate 64, figure 6, and
the tunnels could not be traced into the overlying lutite.
The fact that both Corophioides and Trichophycus venosum
are also incomplete strongly suggests that the upper portion
of the host rock was stripped away before the deposition of
the lutite. Presumably, as shown in the reconstruction in
Text-figure 10, Chondrites, type-A, had one central opening.

Interpretation — As noted in the generic discussion, the
ethological interpretation of Chondrites is still an open ques-
tion. The most critical feature is the origin of the tunnel
fillings. Richter (1931) and Simpson (1957) considered it
to be sediment which sifted into the galleries; while Tauber
(1949) interpreted it as a combination of fecal material and
collapsed lining of the burrow.

It has been pointed out that there are obvious physical
differences between the fillings and the host rock that con-
tains them. In order to ascertain if these physical differences
reflected a mineralogical variation, X-ray defraction analyses
were conducted on the filling, host rock, and overlying lutite
from the Route 27 locality. The results reveal that the filling
is composed of 75-80 per cent quartz, 10 per cent illite,
and minor amounts of chlorite, calcite, and mixed-layer clays
in decreasing order of abundance. In contrast, the host rock
contains 60 per cent calcite, 30 per cent quartz, and minor
amounts of illite, chlorite, and mixed-layer clays. The min-
eralogical composition of the overlying lutite is nearly iden-
tical with that of the filling, the only difference being that
the lutite contains even less calcite. Moreover, the data
indicate that the clay minerals of the filling and overlying
lutite are well crystallized, whereas those of the host rock
are not.

These facts would seem to indicate that Tauber’s in-
terpretation of the European chondritids does not apply
to this Cincinnatian form. While it is true that the organic
acids of the digestive system of an annelid or annelid-like
form might account for the low caleite content of the filling,
it is difficult to explain the increase in the quartz content

in this manner. Also digestive enzymes would tend to reduce
rather than increase the degree of crystallinity of the clay
minerals [Warren Huff (1965) personal communication].
This indicates that Chondrites, type-A, is not a stuffed bur-
row. It is also difficult to reconcile the texture and min-
eralogical composition of the filling with Tauber’s hypo-
thesis regarding C. furcatus. In addition, agglutinated tubes
are normally composed of relatively large particles (see Seil-
acher’s 1951 discussion of tube formation in Lanice conchi-
lega).

Chondrites, type-A, would seem to be the burrow of
an organism which fed by ingesting the sediment of the host
rock, extracting the nutrients, and excreting the waste prod-
ucts at the burrow’s aperture, where they were swept away
by currents. No trace of fecal pellets has been found in
the overlying lutite. The animal probably lay with its anus
near the burrow aperture and extended the body and pro-
boscis or proboscis into the substrate. When the limit of
reach was attained, the organism partly contracted and
excavated a new tunnel. The same process was repeated
several times until the entire network was formed. Due to
the lack of phobotaxis in the burrow, it has not been pos-
sible to determine the exact mode of burrow formation as
Simpson has done for some chondritids. If, however, the
burrow was inhabited throughout the life of the dweller,
it is likely that the upper branches were excavated first,
and as the bedy increased in length, the animal burrowed
progressively deeper. After the death or departure of the
inhabitant, the open system was filled by the same mud
which now forms the overlying lutite. The sharp contact
between the fillings and the tunnel walls probably indicates
that the walls were coated with mucus which inhibited col-
lapse.

The major unanswered question is how could sediment
completely fill such a large system without leaving voids
which would later be filled by calcite or other minerals?
Simpson (1957, p. 497) was questioned on this point and
was unable to offer an explanation. This problem is con-
sidered in detail below.

Chondrites, type-B

Plate 63, figure 3; Plate 64, figures 3,7,8; Plate 65, figure 8; Plate
79, figure 5; Text-figures 11,29-s

Discussion —The forms grouped under Chondrites,
type-B constitute the most common, yet the most poorly
understood of the Cincinnatian chondritids. They occur
throughout the Cincinnatian section in rocks ranging from
fine-grained calcisiltites to medium-grained calcarenites.
They are frequently associated with 7’richophycus venosum

336 PALAEONTOGRAPHICA AMERICANA (YI, 41)

and Corophioides. The reason that the morphology of the
form is poorly known is that most frequently the entire
system is oriented obliquely to the bedding planes and is
thus difficult to observe. Normally Chondrites, type-B, is
seen only as a loose grouping of tunnels piercing the bedding
planes (Pl. 64, fig. 8). At some localities these tunnels are
so numerous that they weaken the resistance of the host
rock to erosion.

In spite of the difficulties involved, it is possible to
piece together a rough outline of the geometry of the net-
work through fragments which are found on bedding and
joint planes. However, a complete analysis can be made
only by detailed serial] section studies.?? When this is ac-
complished, it may well be discovered that a number of
valid species are contained in what is herein considered as
Chondrites, type-B.

As viewed in vertical section, Chondrites, type-B is
seen as a series of tunnels intersecting the vertical plane at
various attitudes. The diameter of the branches varies from
1-4 mm, and, as in the smaller chondritid discussed above,
the contact between the filling and the host rock is always
clearly defined. The form is especially well developed at the
type locality of Corophioides cincinnatiensis, where the chon-
dritids occur in the same bed as the U-tubes (Pl. 79, fig. 5).
One vertical section from this locality, covering an area of
171 square cm, revealed cross-sections of 239 tunnels which
can be separated into two varieties. As seen in Plate 79,
figure 5, the lower half of the host rock contains branches
averaging 1.5-2.0 cm in diameter. These are filled with a
sediment both lighter in color and finer grained than that
of the host rock. The tunnels are loosely grouped and do
not show any preferred arrangement. In contrast the
branches in the upper half of the specimen have a smaller
diameter (0.5-1 mm), are more closely crowded, and are
darker than the host rock. While they do not show the
well-developed stratification illustrated by Richter (1931,
fig. 1), some degree of linear arrangement is present. Ob-
servation of the upper surface of the host rock shows that
only the lower portions of the systems are preserved in the
calcisiltite. Either the upper part of the network lies con-
cealed in the overlying friable lutites or has been stripped
away by subaqueous erosion.

“As a senior thesis project Miss Beverly Barrow (1965) of the

Geology department of the University of Cincinnati attempted to es-
tablish the most feasible method for serial section study of Chondrites.
She found that the tunnels of different systems are so numerous in
some areas that vertical sections taken at 7 mm intervals are not
sufficient to reveal the pattern of the burrows. It is probable that the
Bauplan can be determined only through serial grinding or X-radio-
graphy.

TRACE FOSSILS CINCINNATI AREA: Oscoop a:

A remarkable vertical section (PI. 64, fig. 3), from the
uppermost Richmond beds of Blue Bank, Kentucky, on
State Route 32, six miles southeast of Flemingsburg, pro-
vides insight into what appears to be the main axis of a
Chondrites, type-B, system. The host rock is a medium-
grained, iron-stained calcarenite which bears large amplitude
interference ripples on its upper surface. The fillings have
been washed out and the walls of the branches can be seen
to be perfectly smooth. This main axis is nearly vertical
and has a diameter of 1-2 mm. As can be deduced from the
truncated branches, both the upper and lower limits of the

af

te

—t

1 cm.

Text-figure 11. — Chondrites, type-B. The most complete C., type-
B system thus far found in the local Cincinnatian section. The overall
pattern appears to exhibit radial symmetry. Corryville beds. Stone-
lick Creek. UCM 37623.

axis are incomplete. The secondary branches measure 1.0-
1.5 mm in diameter and bifurcate from the main axis at an
angle which varies from 35-75° off a horizontal plane. While
the axis is well defined throughout most of its length, it
apparently loses its individuality in the lower portion of the
illustration. Another side of the same slab (PI. 63, fig. 3)
gives additional information on the nature of the secondary
branches. While most of the secondary branches proceed
obliquely downward, some are recurved toward the upper
surface of the bed. Close observation shows that they are
following the bedding planes of the host rock. Miss Barrow
has noted the same phenomena in some of the specimens
from West Fork Creek, although the recurvature appears

ws)
Ny

to be independent of the bedding. It is apparent from
the discussion of these vertical sections that this variety of
Chondrites differs markedly from Chondrites, type-A. The
dimensions of the individual branches and the system as
a whole greatly exceed those found in C., type-A. Moreover,
the branches of the Blue Bank specimens are gently arcuate
and do not show the sinuosity of the smaller chondritids.

Thus far Chondrites, type-B, has been discussed only
as seen in vertical sections. Bedding plane surfaces bearing
small segments of C., type-B, branches are not uncommon.
Occasionally exceptional specimens are found which reveal
the nature of the branching as it occurs in the horizontal
plane. The specimen from the Corryville strata of Stonelick
Creek illustrated on Plate 65, figure 8 is similar to the
representatives of Chondrites described by Simpson (1957),
as the branches are rigid and the angle of bifurcation varies
within a small range. The tunnels have a diameter of 2.5
mm and the angle formed by any two branches is be-
tween 30-40°. While the branches do not cross over or inter-
penetrate, neither do they demonstrate the well-developed
phobotaxis seen in the Holzmaden specimens of C. bollensis.

The largest and most complete specimen of a Chon-
drites, type-B, network discovered to date in the Cincin-
natian rocks is also from the Corryville beds at Stonelick
Creek. It is preserved as a concave epirelief on a calcisiltite
and is superimposed over interference ripples. The network
covers an area of 289 square cm and as seen in Text-figure
11 appears to be composed of five “main” branches ar-
ranged in a radial pattern. The walls of the tunnels are
smooth and their diameter varies from 2.5-4 mm. Bifurea-
tion proceeds to the fifth order, and while it follows no set
pattern, many of the branches are forked distally. The
angle of branching ranges between 20-40°. As in the other
Stonelick specimens the branches do not interpenetrate but
only in a few isolated instances do they show anything
which approaches phobotaxis.

In contrast to the above forms, specimens are known
where interpenetration is the rule rather than the exception.
Plate 64, figure 7 illustrates one such form from the Blue
Bank which occurs as an epirelief on the same bed bearing
the specimens shown on Plate 63, figure 3. The branches
interpenetrate to such a degree that they form a bewilder-
ing cross-hatched pattern. Generally the interpenetration
appears to be between branches of different systems. The
dimensions of the tunnels are similar to those seen in verti-
cal section on the same slab.

Interpretation — The origin of the Chondrites, type-
B, network is thought to be similar to that discussed for
C., type-A, 1.e., the empty burrow of a Sedimentfresser

ios)
2
loc)

which was later filled by sediment. None of the fillings
contain fecal pellets nor is there anything that could be
interpreted as the lining of an agglutinated tube. The con-
sistently larger size of the C., type-B, system suggests that
C., type-B, and C., type-A, were made by different organ-
isms. The fact that some C., type-B, networks are oblique
throughout their entire length, while others tend to flatten
out on bedding planes, is probably indicative of the food
distribution in the substrate. Planar networks indicate a

Text-figure 12.— Chondrites “apparatus”. The illustrated appar-
atus was used to determine if sediment could passively fill a Chon-
driles system. The system was constructed of glass tubing with an
internal diameter of 4 mm. The tube endings, which lay on a hori-
zontal plane, were plugged with corks or fiberglass wool packing and
the network was filled with water. Sediment was introduced at the
funnel mouth.

high concentration of nutrients in some beds, whereas the
oblique forms show that the food was evenly distributed
throughout the host rock.

As mentioned previously, a most critical factor is
whether it is possible to fill such a large system solely with
sediment sifting in from above. In order to test this hypo-
thesis, a glass model of a portion of a Chondrites network
was constructed (Text-fig. 12). Glass tubing with an inter-
nal diameter of 4 mm was used, and the various sections
were connected by plastic sleeves. The greatest length of
the system (A-G) measured 51 cm, far in excess of the long-
est known chondritid branch. The distal ends of the system
(C-G) lay on a horizontal plane, and the vertical distance
from the top of the funnel (A) to the plane on which the
distal ends lay was 17 cm.

PALAEONTOGRAPHICA AMERICANA (VI, 41)

Three types of sediment were used: 1) highly angular
silt-sized quartz, 2) a fine silt composed of pumice, and 3)
Pleistocene lake clays. Corks were placed in all the endings
(B-G), and the system was completely filled with fresh
water; care was taken to see that no air bubbles remained.
The sediment was mixed with 200 ml of fresh water and
stirred until it was in suspension. The suspension was placed
in the mouth of the funnel with an eyedropper, the volume
of suspended solution introduced at any given time varying
from 2-5 cc. In order to accelerate the process, the amount
of sediment entering the system was in excess of that which
might be expected to occur under normal geological pro-
cesses.?° It must be admitted that the above apparatus repre-
sents a crude attempt to simulate actual conditions, and,
therefore, the obtained results can be accepted only as
suggestive rather than conclusive.

It was found that the angular sand grains would fill
a maximum of only 53 per cent of the volume of the net-
work. The results achieved with the finer grained pumice
were no better, The apparent reason for this is that we are
dealing with a completely closed system and that a point
was reached where the water pressure in the distal ends of
the system prohibited further flow. It was found that once
the sediment stopped moving, no amount of sediment added
would start the process again.

Even though the chondritid network was coated with
mucus, it was probably not a closed system, and in order
to better approximate actual conditions, the entire appara-
tus was submerged in water, the corks being replaced by
fiber-glass wool packings. It was found that under these
conditions the angular sand would fill 62 per cent of the
system, and the lake clays filled 74 per cent. One addi-
tional experiment was attempted. In order to simulate bot-
tom currents moving over the mouth of the burrow, an
electric fan was allowed to blow across the tank. It was
found that the clays filled 87 per cent of the entire system,
and the sediment completely filled a segment of the tube
44 cm in length. This is nearly three times the length of the
longest chondritid branch noted by Simpson (1957).

Under actual geological processes it might be even
easier to fill the burrow. The rate of sediment inflow into
the burrow would almost certainly be less than that used
in the experiment, and intermittent and continuous cur-
rents moving across the mouth of the burrow could alter
the currents sufficiently to allow the sediment to move to

““It was noted that when a large volume of sediment was introduced
at one time, the result was prominent graded bedding. It might be
fruitful to check some of the more coarse chondritid fillings for
graded bedding which would be proof of passive filling.

TRACE FOSSILS CINCINNATI AREA: OsGoop 339

the ends of the system before more sediment was intro-
duced. Likewise, wave shock or similar phenomena might
be sufficient to jar loose any sediment which had become
clogged. Possibly most important of all is the role that
diagenetic compaction could play. Most of the Cincinnatian
chondritids are generally overlain by lutites, During com-
paction the host rock, being a silt or sand, would remain
relatively unchanged, but the finer grained lutites, which
would be greatly affected, could literally be intruded into
the open network. This would explain how arcuate chondri-
tids, such as those seen on Plate 63, figure 3, could be filled
by such a process. Thus, while it is not possible to prove
that Cincinnatian chondritids are open burrows which are
later filled by sediment, there may well be sufficient mech-
anisms by which such a process could be achieved. Addi-
tional studies should be conducted employing a more re-
fined model of Chondrites. Moreover, X-ray analysis of the
larger C., type-B networks would be worthwhile. This is
especially true for the forms from West Fork Creek which
show two different types of fillings.

Because the above hypothesis requires inspection of
the overlying beds, host rock, and fillings, the author is
not in a position to extend this hypothesis to cover chon-
dritids in general. The thesis can be tested only by direct
field observations and laboratory analysis of the actual
materials in question.

Chondrites, type-C

Plate 63, figure 2; Plate 64, figures 2,4,5; Plate 81, figure 4;
Text-figure 29-t

Discussions — Chondrites, type-C, appears to differ
from all other Cincinnatian chondritids because the branches
of the system form a dense interwoven network, the sur-
face of which is marked by delicate longitudinal striae.
Because all specimens at hand are preserved as concave
epireliefs on fine- and medium-grained calcarenites, the
composition of the filling and the nature of the upper limits
of the network are unknown, All of the specimens are from
Richmond age rocks, and most are from the Whitewater
beds of Short Creek where crossed by Indiana Route 227,
three miles southeast of Richmond, Indiana. The locality
was recollected without success, as the bed containing the
specimens could not be located.

The most striking feature of these forms is the density
of the network. Some areas have been so thoroughly mined
that the interburrow areas occur as isolated knife-like
ridges (PI. 63, fig. 2.24 The density of the system is best

“Tt was found that the network can best be seen in latex molds, and
with the exception of Plate 63, figure 2 and Plate 81, figure 4, all
illustrations are of latex molds rather than the actual specimen.

shown by the form illustrated on Plate 63, figure 7, where
the burrows in the lower half of the specimen cover an area
of 49 square cm. However, this is accomplished at the ex-
pense of considerable interpenetration, and one set of
branches appears to be superimposed over another. Sixth
order branching can be observed in at least one segment of
the system illustrated on Plate 64, figure 2, and while the
angle of bifurcation varies, it tends to be acute (20-30°).
The branches irregularly pinch and swell, and the diameter
varies from 1.5-3 mm.

Based solely on the specimens discussed above, one
would have great difficulty in justifying a chondritid assign-
ment for the Short Creek material, Yet, other specimens
from the same locality reveal a more typical chondritid
pattern. One specimen (PI. 64, fig. 4) possesses a bilateral
symmetry slightly reminiscent of the form illustrated by
Fuchs (1895, fig. 11). However, this may only be a for-
tuitous arrangement of two branches from different systems.
Note that the branches do not show the extremely close
packing of those discussed previously. Another slab (PI.
64, fig. 5) bears two specimens, one of which (lower left)
shows five tunnels which are bundled proximally but branch
distally to form the characteristic anastomosing network.
The diameter of the branches is slightly in excess of 1 mm.
The other specimen on the slab differs as it is obliquely
oriented and only portions of the system are visible, It
gives the impression of possessing a radial outline. What is
perhaps the most significant specimen is illustrated on
Plate 64, figure 2. At the proximal end of the large flabel-
late branch are nine holes arrayed in a circular pattern.
This indicates that the master plan of C., type-C, might
be a relatively small number of vertically directed shafts
which flatten out with depth and bifurcate repeatedly. If
this is the case, the specimens seen on Plate 64, figures
2, 4, 5 represent small segments of a much larger pattern.

The branches of all specimens are marked by faint
longitudinal striae which do not form a well-defined pat-
tern. Usually the larger the diameter of the branches, the
more apparent are the striae.

Interpretation — The origin of the Chondrites, type-C
network is thought to be similar to that of the two types
of chondritids discussed previously. The fact that the
branches are oriented horizontally indicates a high concen-
tration of organic material in thin layers. Presumably the
host rock was overlain but a lutite, and the tunnels were
filled with fine-grained material which has since been re-
moved by erosion. The presence of striae suggests that the
originator of the burrow may have been different from those
that excavated the other Cincinnatian chondritids. Although

340 PALAEONTOGRAPHICA AMERICANA (VI, 41)

the striae recall arthropods, it is equally possible that they
are the impressions of setae borne by a vermiform organ-
ism.

FASCIFODINA, n. gen.
Plate 66, figure 6; Plate 81, figure 8; Text-figures 13,29-v

Type species —The specimen illustrated on Plate 66,
figure 6 (UCM 37632), from the Corryville beds of Second
Creek where crossed by Cozzadale Road, Warren County,
Ohio, is designated as the type specimen for the monotypic
genus Fascifodina floweri, n. sp.

Diagnosis — Vertical bundled tunnel systems of in-
faunal origin which are arranged in a semi-elliptical pat-
tern and are preserved principally as short vermiform con-
cave epireliefs on calcisiltites.

Discussion — The genus was first described by Flower
(1955) from the Corryville beds at Stonelick Creek. He
characterized it as “. . . short curved or vermicular markings

. arranged consistently in groups, each group showing a
crescentic or horseshoe-shaped pattern” (Flower, 1955, p.
862). Flower remarked that although the markings occur
as shallow concave epireliefs on only one bed, they are so
numerous that they frequently overlap.?° In nearly all cases
the crescentic markings are oriented with their concave
face to the median line of the horseshoe. He maintained
that there is a definite bilateral symmetry, although the
total number of individual markings comprising the horse-
shoe varies from 6-13. Two forms (Flower, 1955, figs. 3A,B)
were figured.?° One is nearly equidimensional, measuring
6.0 cm by 6.5 cm, while the other is shorter (4.0 cm) and
broader (7.0 cm). Both forms show a faintly defined raised
area in the center of each horseshoe, and in one (Flower,
1955, fig. 4A) a series of linear markings leads from the
vicinity of the vermiform imprints to the raised central
area.

Additional specimens were collected from the Corryville
beds of Second Creek, which is 10.4 miles north of the
Stonelick Creek occurrence. For the most part these have
the same horseshoe pattern described by Flower, although
some are irregular and poorly developed. The Second Creek
specimens provide a clearer insight into the morphology of
the genus in that the raised area between the open ends of
the horseshoe is much better developed. Upon close exam-
ination this structure appears as a group of eight or nine

“Stream action has covered Flower’s original locality which was some

20 meters below the Ohio Route 131 bridge. At the same time a sec-
ond, slightly higher exposure has been uncovered 50 meters above the
bridge.

*“°These two specimens are in Dr. Flower’s collection.

curved cylinders, 2-3 mm in diameter, which are bundled
together at their apex but radiate out in the direction of
the imprints at the base. One of these cylinders (PI. 66, fig.
6, center right) leads directly into one of the yermiform
markings. Seria] sections of these specimens reveal that
the vermiform markings extend in a highly sinuous course
to a depth of 2-3 cm and are filled with a sediment which
is lighter in color and more fine grained than the host rock.
In contrast, vertical section of the Stonelick Creek speci-

Text-figure 13.—Interpretation of Fascifodina floweri, n. gen.,
n. sp. Block diagram showing that the major volume of the burrow
lies within the silt (dark pattern). Only the upper part of the bundled
structure and the ?master shaft lies within the lutite (dashed pat-
tern). In all Cincinnatian specimens the master shaft has been
stripped away by penecontemporaneous erosion; X 0.25.

mens show that they extend only a few millimeters into
the substrate.

Interpretation — Flower (1955) believed that the
markings were made by the appendages of some animal
grasping the bottom. He maintained that the pattern is
consistently too regular to be a “chance arrangement of
individual vermicular markings made by individual worm-
like animals” (Flower, 1955, p. 862). He mentioned holo-
thuroids as a possibility but doubted that they would be
able to produce such a regular pattern. Instead he selected
an orthoconic nautiloid, Orthonybyoceras Shimizu and
Obata, whose remains are common in the Corryvyille.
Flower went on to describe in some detail how the tentacles
might have produced such markings (Flower, 1955, fig.
4C-F), Although parallel alignment and other current in-

TRACE FOSSILS CINCINNATI AREA: OsGoop 341

dicators are lacking, he believed that the animals grasped
the bottom in order to temporarily anchor themselves
against wave-produced turbidity.

Flower’s interpretation fails to explain both the bundled
cylinders and the fillings found beneath the markings. Even
if the tentacles probed to a depth of 2-3 cm, how could
the holes remain open long enough to be filled with the
finer grained material? The Second Creek specimens occur
in a calcarenite, and any opening would certainly collapse
as soon as the tentacles were withdrawn.

Instead it is concluded that the structure represents
a portion of a bundled feeding burrow. Unfortunately the
morphology of the entire burrow is unknown. At the Stone-
lick Creek locality the surface bearing the markings is
overlain by a 2.5 cm bed of lutite which bears no trace of
any allied structure. It is probable that the upper portions
of the original burrow were stripped away prior to the
lutitic deposition. The specimens from Second Creek were
found in Corryville float, and it was not possible to locate
material in place. As shown in the reconstruction (Text-fig.
13), it is believed that the organism burrowed down
through-out an unknown thickness of lutite and laced the
subjacent calcarenite with a series of bundled burrows ar-
ranged in a semi-elliptical pattern. This differs from Phy-
codes and other bundled burrows not only in the orienta-
tion (vertical as compared to horizontal) but in the highly
irregular nature of the portion of the tunnels found in the
calearenite bed. The vermiform markings seen on the sur-
face are attributed partly to the actual mining activity of
the organisms and partly to the collapse of the tunnel sys-
tem. The material filling the tunnels is probably composed
of a mixture of the overlying lutite and the sediment
worked by the organism. There is no evidence of fecal pel-
lets or strings. The raised bundles found between the open
ends of the horseshoe in the Second Creek forms repre-
sent material dragged up the tube, thus the bundles form
a mold of that portion of the network.

The walls of the burrow are not annulated as in Phy-
codes or scratched as in Trichophycus venosum, hence there
is no direct evidence which would point to the zoological
affinities of the producer.

PHYCODES Reinhard Richter, 1850

Plate 57, figure +; Plate 65, figures 1-5; Plate 66, figures 2,4; Plate
68, figure 9; Plate 69, figure 4+; Plate 70, figure 6; Text-figures

14,15,29-x
1850. Phycodes Richter, Reinhard, Deutsche Geol. Gesell., Zeitschr.,
Bd. 2, p. 205.
1851. Non Inocaulis Hall, American Jour. Sci. Arts, 2d ser., vol. 11,
p. 401.

1862. Licrophycus Billings, Palaeozoic Fossils, vol. 1, Geol. Sur. Can-
ada, Public., p. 99, fig. 87.

1878. Licrophycus Billings, Miller and Dyer, Cincinnati Soc. Nat.
Hist., Jour., vol. 1, pp. 2,3, pl. 2, fig. 4.

1885. Inocaulis Hall, James, J. F., Cincinnati Soc. Nat. Hist., Jour.,
vol. 7, p. 164, fig. 8.

1934. Phycodes Richter, Magdefrau, Neues Jahrb. Mineral., Geol.,
u. Palaont., Beil.-Bd. 72, pp. 259-282, figs. 1-6, pls. 10-11.

1955. Partim Phycodes Richter, Seilacher, Akad. Wiss. Lit. Mainz
math.-nat. KI., Abh., No. 10, pp. 383-388, fig. 3.

1962. Phycodes Richter, Hantzschel, Trace-Fossils and Problematica
in Treatise on Invertebrate Paleont., Moore (ed.), pt. W, Mis-
cellanea, p. W208, fig. 128-1.

Type species —Phycodes circinatum Reinhard Richter
(1850) from the Ordovician “Phycodes beds” of Thuringia.

Diagnosis. — Horizontal bundled burrows of infaunal
origin preserved outwardly as convex hyporeliefs. Over-all
pattern flabellate, broomlike, or occasionally circular. Some
forms consist of a few main branches showing a Spreite-like
structure which distally gives rise to numerous free
branches. In other forms the Spreiten are lacking and
branching tends to be more random. Individual branches
are terete and finely annulate or smooth. Length of entire
burrow 2.5-15 cm; diameter of bundled branches of 1.5 cm.

Discussion — Phycodes, or comparable forms, have
been the subject of three excellent studies; Sarle (1906),
Magdefrau (1934), and Seilacher (1955) all have added to
the knowledge of these bundled structures, which are found
in the lower Paleozoic of North America, Europe, and
Asia.

Magdefrau (1934) restudied topotype material (Pl. 65,
fig. 5) and found that Phycodes circinatum is preserved as
convex hyporeliefs in quartzites. Most specimens are 10-15
em long, although individuals as small as 2-4 cm long are
known, The depth (taken from deepest branch to base of
quartzite bed) is 2.5 cm (see Text-fig. 14b). Although
branching occurs at acute angles, the branches first proceed
parallel for a distance and then fan out, some doubling
back so that they again run parallel to the main axis.
Specimens exhibit no apparent preferred orientation, as the
main axes of the various forms proceed in every conceivable
direction. The free branches, which have an average dia-
meter of 1.3 mm, are marked by delicate annulations which
were originally noted by Richter. These average 20-25/cm,
the greatest number being 38/cm. According to Magdefrau
(1934, fig. 4), the annulations are exposed only when a
smooth, thin (0.25-0.5 mm) outer covering is eroded.

Serial sectioning undertaken by Seilacher (1955) re-
vealed the Bawplan of P. cincinatum (Text-fig. 14b). The
proximal portion of the structure consists of a few main
tunnels with retrusive Spreiten. Distally each tunnel di-
vides into numerous free branches which angle back up into
the host rock. Beyond this there is no set pattern, as the
free branches may swing first one way then another.

342 PALAEONTOGRAPHICA

There is some disagreement regarding the morpho-
logical limits of Phycodes. Seilacher (1955) expanded the
genus, while Hantzschel (1962) has been more conservative.
Seilacher (1955) considered Arthrophycus Hall (1852) to
be a junior synonym of Phycodes and also placed Butho-
trephis palmata Hall (1852) (Pl. 67, figs. 3, 7) within the
genus. At the same time he proposed a new species, ey
pedum, from the Lower Cambrian of the Salt Range.

Text-figure 14.—Seilacher’s reconstruction of Buthotrephis pal-
mata Hall. The reconstruction is based on specimens from the Lower
Cambrian of the Salt Range, Pakistan. A relatively small number of
master shafts (left) branch to give a broomlike pattern. The master
shafts show a crude Sfeiten-like arrangement; X 0:75. b. Seilacher’s
reconstruction of Phycodes circinatum Reinhard Richter. The inter-
pretation is based on material from the Salt Range. The master
shafts (left) have a well-developed Spreiten-like arrangement and
that the free branches are smaller and more numerous than in
Buthotrephis palmata; X 1.5. (From Seilacher, 1955, fig. 3.)

Although there is an apparent similarity in general
pattern from Arthrophycus and Phycodes, the former dif-
fers in that it is large (branches up to 60 cm long, accord-
ing to Hantzschel, 1962), bears prominent annulations, and
has bilobate branches which can be as much as 1 cm in
diameter. Moreover, Sarle (1906, fig. 4) showed that the
bundled structure of Arthrophycus, as it occurs in the Sil-
urian Medina Sandstone of western New York, is a response
to sedimentation. The organism was forced to raise the
level of the burrow as sand repeatedly buried the aperture.
Whether this explanation applies to Arthrophycus from
other localities is not known. As will be shown below, the
bundled structure of Phycodes is of different origin. Hantz-
schel (1962) considered Arthrophycus a separate genus, and,

Americana (VI, 41)

based on the differences cited above, his decision is ac-
cepted here.

Seilacher (1955) also considered Buthrotrephis palmata
Hall (1852) (see Pl. 67, figs. 3, 7) to be a species of Phy-
codes. B. palmata Hall is not common. It was originally
described from the Clinton (Silurian) beds of New York
and has been identified by Seilacher (1955) in the Lower
Cambrian of the Salt Range of Pakistan. While B. palmata
is certainly not similar to the other species of Buthotrephis
figured by Hall (1847, 1852) (they are clearly chondritids;
see Pl. 69, fig. 8), it is highly doubtful that it should be as-
signed to Phycodes. Seilacher’s sectioning of his Salt Range
forms (Text-fig. 14a) shows that the branching pattern is
slightly different and that there is only an incipient Spreite
developed. More important is the fact that Hall’s forms
have far fewer branches than P. circinatum, and these
branches are smooth and much larger (up to 2 cm in dia-
meter). Hall’s material has never been sectioned or ade-
quately studied. When such a study has been completed, it
it will most likely be found that B. palmata merits a new
generic designation.

The specimens from the Lower Cambrian of the Salt
Range described by Seilacher (1955, figs. 4a, b) as P.
pedum differ considerably from all forms discussed above.
A gently sloping master tunnel which is open to the outside
bifurcates at depth to give rise to horizontal sickle-shaped
galleries. In turn numerous short vertical shafts branch off
the galleries. Although the form is known only from a
few specimens, it should be established as a separate genus.

Phycodes is judged to be the senior synonym of Licro-
phycus Billings (1862). Although none of Billings’ many
species from the Middle Ordovician and Upper Ordovician
of Ontario has ever been sectioned, Licrophycus is ex-
ternally similar to Phycodes (compare Magdefrau, 1934,
pls. 10, 11 with Billings, 1862, fig. 87 and Wilson, 1948, pl.
4, fig. 1; pl. 5, fig. 4). Based on Billings’ figures and de-
scriptions, the only apparent differences are that Licro-
phycus has smooth branches and these lack the recurvature
seen in some of Magdefrau’s material. The forms are simi-
lar in the diameter of the branches and the size of the
structure as a whole.

Interpretation — There have been several interpreta-
tions as the origin of Phycodes-like structures.** Most early
workers, e¢.g., Reinhard Richter (1850) and Billings (1862),
held them to be “fucoids,” and this notion persisted with
at least two authors (Hundt, 1931, and Wilson, 1948) well

“The reader is referred to Magdefrau (1934) for a more complete
discussion of the early history of the genus and its interpretation.

TRACE FOSSILS CINCINNATI AREA: OsGoop 343

into the present century. Others (e.g., Potonié 1899)
thought Phycodes to be of inorganic origin (rill marks),
while Abel (1935) considered Arthrophycus as the casts of
worm tubes thrown together during a storm. Although Abel
never mentioned Phycodes specifically, he may well have
believed that the origins of the two were similar. However,
more careful studies by later authors have shown conclu-
sively that it is a branching burrow. As has proved true
with so many “fucoides,” sectioning of the specimens pro-
vided the key. This reveals (Text-fig. 14b) that both ends
of the hyporelief structure extend up into the bed bearing
them. Magdefrau reflected that this concave-upward orien-
tation is strange preservation for algae.

The most detailed ethological analysis of P. circinatwm
was given by Seilacher (1955). He considered it a Fodinich-
nia where the organism systematically mined a nutrient-rich
layer along a silt-mud interface. The animal first burrowed
down through the silt, then moved horizontally along the
interface before once again angling up into the silt bed.
The resulting structure resembles a “U” in which one limb
is steeper than the other. Following this, the organism
withdrew from the distal portion of the burrow and proceed-
ed to excavate a new tunnel adjacent to the old one.
Periodically the unbranched proximal portion of the bur-
row was elevated by removing sediment from the ceiling
and packing it, along with waste material, on the floor.?§
This activity gave rise to the retrusive Spreite-like struc-
ture seen in Text-figure 14b. When a given area was mined
out, a new “master” tunnel was formed in another location,
and the entire process repeated again.

Although Seilacher (1955) drew an analogy between
the bundled feeding shafts of Arenicola marina and those
of Phycodes, directly comparable Recent structures are un-
known. The annulate branches of P. circinatum suggest an
organism with an annelid-like body, but the fact remains
that we are dealing with a basic pattern which could have
been made by any one of a number of organisms.

CINCINNATIAN REPRESENTATIVE,

Phycodes flabellum (Miller and Dyer)

Plate 57, figure +; Plate 65, figures 1-+; Plate 66, figures 2,4; Plate
68, figure 9; Plate 69, figure +; Plate 70, figure 6; Text-figure 15,29-x

1878. Licrophycus flabellum Miller and Dyer, Cincinnati Soc. Nat.
Hist., Jour., vol. 1, pp. 2-3, pl. 2, fig. 4.

1885. Inocaulis flabellum (Miller and Dyer), James, J. F., Cincinnati
Soc. Nat. Nat. Hist. Jour., vol. 7, p. 164.

“Neither Magdefrau (1934) nor Seilacher (1955) reported finding
fecal pellets in the tunnel.

Type locality — Licrophycus flabellum Miller and Dyer
(1878) from the “upper part of the Cincinnati Group”
(Richmond) of Lebanon, Ohio. Type specimen: Plate 65,
figure 1; HBM 3178.

Diagnosis — Bundled burrows generally arranged in a
flabellate pattern. Individual branches cylindrical and an-
nulate. Sectioning reveals no evidence of Spreite-like struc-
ture or fecal material.

Discussion — Although P. flabellum is known from sev-
eral localities in the upper two-thirds of the Cincinnatian
section (Maysville-Richmond), it is not a common form,
and only 25 specimens have been available for study. How-
ever, when the species is encountered, it is not unusual to
find more than one specimen on a slab, e.g., an area in the
Corryville beds of Stonelick Creek, measuring 45 cm by 27
cm, contained five specimens. All Cincinnatian specimens
are preserved outwardly as convex hyporeliefs in calcisiltites
or fine-grained calcarenites. There is no marked difference
between the lithology of the host rock and that of the
branching structure.

It is difficult to generalize on the over-all pattern of
P. flabellum, because the species shows much variation.
However, as Miller and Dyer indicated in their choice of a
specific name, many Cincinnatian specimens are flabellate.
This condition is best typified by a portion of the type
specimen (PI. 65, fig. 1) and the specimen illustrated on
Plate 70, figure 6. The latter form possesses 98 separate
branches which cover an area of 41 square cm while in the
flabellate portion of the type specimen, 23 branches cover
an area of 3.75 square cm. This fanlike arrangement points
up one difference between P. flabellum and P. circinatum.
The European species is more fascicular, closely resembling
a broom-shaped bundle, where the height?® is nearly equal
to the width. In many Cincinnatian forms the width of
the structure greatly exceeds the height. A vertical section
of the form seen on Plate 70, figure 6 reveals that only the
basal 2 mm of the bed is disturbed. Thus this specimen has
a width of 13 cm and a height of only 2+ mm. A variation
of the flabellate pattern (PI. 66, fig. 4) consists of five or
six bundles which radiate out from a central point in such
a manner that they nearly cover a complete circle.

Not all Cincinnatian forms show such a definite Bau-
plan. In such specimens (PI. 66, fig. 2) the branches are
bundled together, but the over-all arrangement appears to

“Height is here defined as the height of the entire structure. It is
equal to the height as used by Magdefrau (lowest branch to base
of bed) plus the added height shown by sectioning. As viewed in
Text-figure 15C it would be the distance from the lowest branch to
the dashed line.

344 PALAEONTOGRAPHICA AMERICANA (VI, 41)

be haphazard, and the various bundles cross over and inter-
penetrate.

The individual branches are terete in cross-section and
range from 1-3 mm in diameter; the diameter remains
constant throughout their length. Some branches give the
appearance of possessing a median furrow, but if the
branches are traced far enough, they are seen to bifurcate.
All branches are marked by delicate annulations which ay-
erage 6-8/cm. There is no trace of a smooth outer sheath like
that described by Magdefrau in P. circinatum.

Text-figure 15.— Reconstruction of Phycodes flabellum (Miller
and Dyer) from the Cincinnatian. Block A shows the master shaft
as seen in vertical section. Blocks B and C show that there is no
Spreiten structure in the Cincinnatian. Careful study has revealed
only an area of disturbed sediment whose contacts are gradational
with the host rock; X 0.5.

In hopes of presenting a reconstruction of P. flabellwm
as Seilacher did for P. circinatum and Buthotrephis palmata,
serial sections of several specimens were prepared. When
it became apparent that polished, etched, and stained sec-
tions would not suffice, thin sections were cut, All these
reveal that with one lone exception there is no Spreite-like
structure present in P. flabellum. One encounters only a
general area of disturbed bedding ( Pl. 65, fig. 3). Moreover,
the contact between this area and the undisturbed bedding
of the host rock is not sharp but gradational. The one ex-
ception mentioned above is the presence of a protrusive
Spreite in a small area of one form (Pl. 65, fig. 2). As will
be shown below, this lack of a definitive internal structure
is significant in the ethological interpretation of the species.

While the internal structure of most of the burrow is
not known, it is possible for the first time to shed some
light on the nature of the main shaft. Neither Seilacher nor
Magdefrau made any mention of the extreme proximal (un-
branched ) ends of Phycodes. Vwo specimens of P. flabellum
(Pl. 57, fig. 4; Pl. 65, fig. 4) show a wide (up to 1 cm)
vertical shaft along which the bedding planes are bowed
down. The original diameter of the shaft was undoubtedly
smaller, for the dark L-shaped body seen in Plate 65, figure
4 shows that collapse of the walls has led to some enlarge-

ment. An interesting point is that the main shaft is vertical.
In both specimens it extends nearly to the base of the cal-
cisiltite bed, where it geniculates at 90° to run parallel
to the base of the bed before fanning out in multiple bur-
rows along the interface (Text-fig. 15).

In gross morphology P. flabellum is not unlike P. ci-
cinatum or the many species described by Billings. How-
lever, it differs sufficiently to justify the retention of
Miller and Dyer’s specific designation. The diagnostic
features are the flabellate pattern, the lack of a smooth
outer sheath, and the general absence of Spreiten. Billings’
specimens have never been restudied, but it is likely that
further work on them will reduce the number of Canadian
species in the genus.

Interpretation — Miller and Dyer (1878) followed the
lead of earlier workers on Phycodes and assigned P. flabel-
lum to the “fucoids.”

The species reminded J. F. James (1885, p. 164)
. of the appearance that would be made in mud by the
expanded tentacles of a crinoid.” He believed, however, that
it was a graptolite and placed it in /nocaulis Hall (1852),
noting that it differed greatly from J. plumosum but was
“quite similar” to J. walkeri Spencer (1884).

“cc

As Ruedemann (1908) showed, Inocaulis plumosum
and I. walkeri (—Acanthograptus walker) are graptolites
in which the usual graptolite integument is preserved.
Clearly such forms have no genetic bearing on P. flabellum.
Ruedemann (1908, p. 191) succinctly dismissed /. flabellum
with “this is no graptolite.”

The serial sections confirm that neither Miller and
Dyer nor James made a correct interpretation. The internal
structure and the interpenetration of the branches leave no
doubt that P. flabellwm is a trace fossil. However, the. exact
ethological meaning of P. flabellum is unclear and two alter-
nate hypotheses will be presented below.

As Seilacher suggested, the inhabitant of the burrow
may have been a Sedimentfresser similar to the Recent
Arenicola marina (L.). However, Schafer (1962) showed that
Arenicola constructs bundled feeding shafts only in highly
cohesive mud. In sand the burrow more closely resembles
a J- or U-tube. Likewise, there is no trace of cone-shaped
areas of disturbed sediment associated with the numerous
branches. Moreover, if the organism has mined out the bur-
row and the filling represents a collapse structure, one would
expect to find slumping in the overlying beds of the host
rock. The serial sections show little evidence of any slump-
ing.

It is also possible that the inhabitant was a selective
feeder, i.¢., a form which was able to selectively glean the

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 34

nutrients from the sediment without ingesting a large vol-
ume of inorganic material. It may be that the organism lay
with its anus near the mouth of the burrow and systematic-
ally extended its anterior end through the sediment, selec-
tively removing the nutrients. The animal was thus dis-
turbing and displacing the sediment rather than removing
it. After the animal had extended itself to the maximum
length, it withdrew and the sediment filled in the cavity.
Because there was little net loss of sediment, extensive
slumping would not occur. As evidenced by the bundled
nature of the burrow, this process was repeated several
times. When a given area was mined out, the organism
changed directions and again proceeded to feed. Since well-
developed Spreiten are not present in Cincinnatian forms,
there is little evidence that the animal made a practice of
packing waste material on either the floor or the roof of the
burrow. Following the demise or departure of the organism,
a combination of collapse and sediment sifting in from out-
side filled the proximal portion of the burrow (z.e., the main
shaft). It must be admitted that the feeding habits of Re-
cent organisms do not provide strong support for this argu-
ment. All known selective feeders (¢.g., terebellids) live
in vertical shafts and extend their tentacles over the deposi-
tional interface, It is somewhat difficult to visualize how
this might be accomplished at depth.

DACTYLOPHYCUS Miller and Dyer, 1878

Plate 57, figure 3; Plate 63, figure 7; Plate 69, figure 2; Plate 74,
figure 3; Text-figure 29-y

1847. Non Palacophycus Hall, Nat. Hist. New York, Palaeont. New
York, Hall, vol. 1, p. 7, pl. 2, figs. 1,2,4,5.
1873. ?Palaeophycus Hali, Orton, Geol. Ohio, vol. 1, p. 387.

1873. Non Planolites Nicholson, Roy. Soc. London, Proc., vol. 21, p.
289.

1878. Dactylophycus Miller and Dyer, Contributions to Palaeontology,
No. 2 (Cincinnati, Ohio, private publication), pp. 2,3, pl. 3,
figs. 1,2.

1885. Partim Palaecophycus Hall, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 164.

1891. Partim Planolites Nicholson, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 14, p. 47.

1962. Dactylophycus Miller and Dyer, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W190.

Type species — Dactylophycus tridigitatum Miller and
Dyer (1878) from the Eden beds of Cincinnati, Ohio.

Diagnosis — Small, branching, annulate, bilobed bur-
rows of infaunal origin which are preserved outwardly as
convex hyporeliefs. Branching pattern may be radiate or
random.

Discussion — Dactylophycus was proposed for forms
with “‘a single stem, which divides, at one or both ends, into
three or more buds or short branches, each of which ter-

wat

minates more or less abruptly in a point. The stem and
branches may be smooth or rugose” (Miller and Dyer,
1878b, p. 1). Two species, D. tridigitatum and D. quadri-
partitum, were established with the former being designated
the type species. Figured specimens of both species are list-
ed in the catalogue of the paleobotanical collection of Har-
vard University, but only D. quadripartitum was located.
Therefore, judgment on the nature of D. tridigitatum must
be based solely on Miller and Dyer’s description and their
highly schematic illustration.

As James (1885) stated, it is possible that Palaeo-
phycus radiata Orton (1873) belongs here. The species,
from “Morris’ Hill, Warren County, Ohio,” was never fig-
ured and Orton’s description is vague. He characterized it
as “five or six cylindrical stems radiating from a common
center. The stems are from two to three inches long, and
agree in form with some that Hall figured as Palaeophycus”
(Orton, 1873, p. 387). While it is possible that P. radiata is
synonymous with Dactylophycus, it is just as likely that
it is a specimen of Phycodes flabellum. The only form de-
scribed by Hall under Palaeophycus which resembles either
of the above is ?Palaeophycus sp. Hall (1852, pl. 10, fig. 3).

Miller (1889) pointed out the close similarity between
Dactylophycus and Ichnophycus tridigitatum Hall (1852),
pl. 10( figs. 7a, b), described from the Clinton (Silurian)
beds of New York State. The present author has studied
the type specimen of Jchnophycus and must disagree with
Miller’s conclusion. Jchnophycus is preserved as a concave
epirelief in the thin micaceous bed which overlies a coarse-
grained quartz sandstone. It possesses three completely
smooth nonbilobate branches and closely resembles a “bird’s
foot” in outline. While it is probably a portion of a burrow,
there is no real basis for comparison with Datcylophycus.

DACTYLOPHYCUS QUADRIPARTITUM Miller and Dyer

Plate 57, figure 3; Plate 63, figure 7; Plate 69, figure 2; Plate 75,
figure 3; Text-figure 29-y

1873. ?Palaecophycus radiata Orton, Geol. Ohio, vol. 1, p. 387.
78. Dactylophycus quadripartitum Miller and Dyer, Contributions

to Palaeontology, No. 2 (Cincinnati, Ohio, private publication),
PP: 2555) Plies, sige 1.

1885. Palaeophycus radiata, Orton, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 164.

1891. Planolites radiata, Orton, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 14, p. 47.

Type locality — Eden beds, Cincinnati, Ohio. Type
specimens: Plate 57, figure 3 (HBM 3174; latex mold UCM
37600) and Plate 69, figure 2 (HBM 3174; latex mold UCM
37608 ).

Discussion — Miller and Dyer (1878b, p. 2) character-

346 ParavonrocrariicA Amrricana (VI, 41)

ized D. quadripartitum as a “plant [with] a stem, which
is divided, at one end, into four finger-like prongs or
branches, the other end is unknown . . . Each branch has a
slight longitudinal depression, in the middle, and shows
some evidence of rugose markings. The specimen illustrated,
however, is nearly smooth.”

Only six specimens of D. quadripartitum are known,
all are from the Eden and occur outwardly as convex hypo-
reliefs in platy calcisiltites. The individual branches have a
diameter of 2-4 mm and a length of 15 mm. The number
of branches varies from specimen to specimen and is not
a critical factor as Miller and Dyer believed. Each branch
possesses a well-developed median furrow which in some
specimens is more like a flat median trough, taking up one-
third to one-half the width of the branch (PI. 18, fig. 3;
USNM 40027). Well-preserved specimens* possess delicate
annulations which average 15/cm. In the specimens just
mentioned the annuli are confined to the lateral ridges and
are not found in the trough.

The pattern of branching is variable. Two specimens
show a flabellate pattern. One of these (PI. 63, fig. 7) pos-
sesses 10 branches which subscribe an are of nearly 180°;
the other (PI. 57, fig. 3) has only four branches which do
not overlap. One other specimen (PI. 74, fig. 3) is slightly
flabellate, while in another the branching is random.

D. quadripartitum differs from other bundled structures
(e.g., Phycodes, Arthrophycus), in that it shows a marked
relief off the sole of the host rock. The branches are not
seen as hemicylinders but instead are exposed as almost
complete cylinders. Many of the branches have trapped
fragments of lutite along their lateral margins. In the three
specimens discussed above what appears to have been the
main shaft (i.e, the trunk which connects the numerous
branches) has broken off, indicating that it once lay
wholly within the lutite and has since been lost. Sectioning
confirms that there are no “hidden” tunnels lying com-
pletely within the calcisiltite, thus with the exception of
the main shaft and the extreme distal portions of the
branches, the entire network is visible as a convex hypo-
relief.

In appearance Dactylophycus is most closely related
to Phycodes and Arthrophycus. It differs from Phycodes in
possessing a smaller number of branches which are bilobate.
The smaller diameter and less prominent annulations of the
branches of Dactylophycus differentiate this genus from
Arthrophycus. Moreover, it lacks the bundled structure de-
scribed by Sarle (1906).

“Miller and Dyer claimed the branches were smooth or annulate.
However, their smooth branches are the results of Recent erosion.

Two species of Dactylophycus are probably not re-
quired. The descriptions of D. tridigitatum and D. quadri-
partitum are similar, the only difference (as the specific
name indicates) being the mode of branching. However,
authors did state the D. tridigitatum branched at both ends.
If this is actually the case, D. tridigitatum may be some-
thing different. For this reason it seems wise to retain the
two species until the type specimen of D. tridigitatuwm can
be located.

Interpretation. — James (1885) considered Dactylo-
phycus to be the fragments of burrows; the same conclu-
sion is reached here. As mentioned previously, the ethologi-
cal meaning of Phycodes is in doubt. Although the general
Bauplan of Dactylophycus is similar to that of Phycodes,
the ethological interpretation is more clear-cut. The fact
that the greater volume of the burrow once lay within the
subjacent lutite indicates that Dactylophycus is the excava-
tion of a Sedimentfresser.

Presumably the organism burrowed down through the
calcisiltite into the subjacent calcilutite and then proceeded
to feed along the interface. As the animal withdrew from
the tunnels, the overlying calcisiltite sifted in to insure
preservation of at least the distal portions of the burrow.
The main shaft, 7.¢., master tunnel, is not present in any
of the specimens at hand. It has either been broken off
or was composed of lutite and thus not preserved.

TRICHOPHYCUS Miller and Dyer, 1878

Plate 60, figure 7; Plate 68, figures 1-8; Plate 70, figure 4;
Text-figures 16,29u

1847. ?Partim Palacophycus Hall, Nat. Hist. New York, Palaeont.
New York, vol. 1, p. 263, pl. 70, fig. 1.

1873. Non Planolites Nicholson, Roy. Soc. London, Proc., vol. 21, pp.
288-290.

1878. Non Blastophycus Miller and Dyer, Cincinnati Soc. Nat. Hist.,
Jour., vol. 1, p. 24, pl. 1, figs. 1,2.

1878. Trichophycus Miller and Dyer, Cincinnati Soc. Nat. Hist., Jour.,
vol. 1, pp. 1-3.

1879. Trichophycus Miller and Dyer, Miller, Cincinnati Soc. Nat.
Hist., Jour., vol. 2, p. 112, pl. 9, figs. 5,5a.

1884. Partim Blastophycus Miller and Dyer, James, J. F., Cincinnati
Soc. Nat. Hist., Jour., vol. 8, pp. 158,159.

1885. Partim Palacophycus Hall, James, J. F., Cincinnati Soc, Nat.
Hist., Jour., vol. 7, p. 158.

1885. Partim Trichophycus Miller and Dyer, James, J. F., Cincinnati
Soc. Nat. Hist., Jour., vol. 7, p. 131.

1891. Partim Planolites Nicholson, James, J. F., Cincinnati Soc, Nat.
Hist., Jour., vol. 14, p. 47.

1891. Cyathophycus James, J. F., Cincinnati Soc, Nat. Hist., Jour.,
vol. 14, p. 64, figs. 5a-c.

1962. Trichophycus Miller and Dyer, Hantzschel, Trace-Fossils and
Problematica in Treatise on Invertebrate Paleont., Moore, (ed.),
pt. W, Miscellanea, p. W219.

1964. Trichophycus Miller and Dyer, Seilacher and Meischner, Geol.
Rundschau, Bd. 54, p. 614, fig. 11.

‘TRACE FOSssiLs CINCINNATL AREA: OsGoop 347

Type species — Trichophycus lanosum Miller and Dyer
(1878) from the “upper part of the Cincinnati Group”
(Richmond) of Warren County, Ohio.

Diagnosis. — Cylindrical, gently U-shaped Fodinichnia
or Domichnia of moderate size. A small number of vertically
directed secondary branches may be present. The ventral
surface of the burrow is marked by a series of delicate striae.

Discussion. — The authors established Trichophycus
for “simple branches or stems having diagonal or longi-
tudinal markings, as if made by the folding down of hair-like
filaments” (Miller and Dyer, 1878a, p. 3).

Initially only a single species, 7. lanosum, was pro-
posed, although the same year and in Part 2 of the same
paper, the authors proposed another species, 7. sudcatwm,
and with some doubts placed it within the genus. In 1879
Miller added a third species, T. venosum.

J. F. James (1884, 1885) considered 7. sulcatum and
T. venosum to be synonymous, while he placed 7. lanosuwm
in synonymy with Blastophycus Miller and Dyer (1878).
In 1891 James proposed Cyathophycus siluriana from the
Corryville beds. As will be shown below this is believed to
be synonymous with Trichophycus venosum.

It is unfortunate that 7. lanoswm was selected as the
type species, as it does not best express the nature of the
genus. It is known only from the syntypes and appears to
be an aberrant variation of 7. venoswm which is an ex-
tremely common form. 7. sudcatwm is entirely different and
is discussed under Fucusopsts sp.

Interpretation. — Miller and Dyer interpreted all three
species as “fucoids,” while J. F. James (1884, 1885) con-
sidered both 7. sulcatum and T. venosum to be casts of rill
marks and 7. /anosuwm an annelid burrow. James (1891) be-
lieved Cyathophycus siluriana to be a sponge and remarked
that it resembled the specimens of Rauffellia Ulrich figured
by Ulrich (1889, fig. 1,2,4) from the Ordovician of Minne-
sota. Hantzschel (1962, p. 219) characterized the genus as
“burrows with delicately scraped walls,” noting that Sei-
lacher too had reached this conclusion (Seilacher, 1955,
personal communication to Hantzschel).

As will be shown, 7. venoswm is a branching burrow of
infaunal origin. Although 7. /anoswm is not thoroughly un-
derstood, it may be a behavioral variant of the same
organism that produced 7’. venosum.

TRICHOPHYCUS VENOSUM Miller

Plate 60, figure 7; Plate 68, figures 1,3,4-7; Plate 70, figure 4;
Text-figures 16,29w

1847. ?Palacophycus virgatus Hall, Nat. Hist. New York, Palaeont.
New York, vol. 1, p. 263, pl. 70, fig. 1.

1879. Trichophycus venosum Miller, Cincinnati Soc. Nat. Hist., Jour.,
vol. 2, pp. 9,10, pl. 9, figs. 5,5a.

1884. Trichophycus venosum Miller, James, J. F., Cincinnati Soc.
Nat. Hist., Jour., vol. 7, p. 131.

1885. Palaeophycus virgatus Hall, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 158.

1891. Planolites virgatum (Hall), James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 14, p. 47.

1891. Trichophycus venosum Miller, James, J. F., Cincinnati Soc.
Nat. Hist., Jour., vol. 14, p. 47.

1891. Cyathophycus siluriana James, J. F., Cincinnati Soc. Nat. Hist.,
Jour., vol. 14, p. 64, figs. 5a-c.

Type locality. —“Hudson River Group, Cincinnati,
Ohio.” The species is common throughout the Cincinnatian.

Diagnosis. — Vertically branched, gently U-shaped
structures of infaunal origin. Floor of burrow marked by
numerous fine striae which radiate posterolaterally from
midline.

Discussion. — Miller (1879, pp. 9,10) described T.
venosum as a “half cylindrical stem, covered upon the cylin-
drical surface with irregular and inconsistent elevated lines,
varying in their course from longitudinal with the stem to
diagonally radiating from a central line.” He believed that
the stem originally may have been cylindrical in shape but
that half of it has been lost.

Although it was not possible to locate the actual speci-
mens figured by Miller, several similar ones are available for
study. These are not strictly hemicylinders (i.¢., “planocon-
vex” to borrow from brachiopod terminology) as Miller
described but instead are “concavo-convex.” The convex
face bears the delicate striae of the original description.
Plate 64, figure 7 shows one such form in which the striae
radiate back from at least two central points. These in turn
are superimposed oyer other such markings. The shallow
concave surface shows no sculpturing.

J. F. James (1891, p. 64) established Cyathophycus
siluriana for “rounded or flattened cylindrical bodies, varying
from one to two and one-half inches in length, and about one
inch in width; outer surface marked by longitudinal lines
starting from a common point at one end, and radiating in
the form of a cup; numerous transverse lines crossing the
longitudinal ones and so forming an irregular net work. The
internal structure seems to be entirely lacking, the center
being filled with clay, and only a portion of the framework
remains.” His description was accompanied by three highly
schematic drawings. It is doubtful if James ever designated
a type but specimens labeled “Cyathophycus siluriana” were
found in the James Collection at the Field Museum. They
are identical with Trichophycus venosum, as described by
Miller. Bassler (1915, p. 1289) reached the same conclusion.
It is strange that James did not recognize the similarity be-

cause Miller’s (1879, pl. 9, figs. 5-5a) lithographs of Tricho-

348 PALAEONTOGRAPHICA AMERICANA (YI, 41)

phycus venosum are accurate and James discussed T. veno-
sum in his 1885 paper.

Striated hemicylinders have been recovered from the
Economy beds of Humphrey’s Branch of Twelve Mile
Creek, Kentucky (type locality of Rusophycus cryptolitht).
Field studies carried out in this area reveal that the lamellae
occur as fillings in what local collectors for many years have
referred to as “turkey tracks.” “Turkey tracks” are shallow
hemicylindrical depressions with no apparent preferred
orientation which are deepest in the middle and rounded at
both ends (PI. 60, fig. 7). They are either straight or gently
arcuate in outline, and although the surface may be slightly
irregular, they are normally devoid of sculpturing. They
have never been observed to branch laterally, yet trunca-
tion is common. The width varies from 2.5-3.5 cm, and the
length from 8.5-33 cm.*!

Although beds containing large numbers of “turkey
tracks” occur throughout the Cincinnatian section, the form
is especially well developed at the locality mentioned above
and in the Corryville section at Stonelick Creek, some 90
meters downstream from where State Route 131 crosses the
creek. At these two localities approximately 25 per cent of
the depressions are filled with the lamellae (PI. 70, fig. 4).
When the fillings are carefully removed with a knife, the re-
sulting depression exactly resembles the “turkey tracks.”
Because the lamellae are normally softer and finer grained
than the host rock, it is apparent that most of them have
been eroded out by Recent stream action.

When the lamellae are either similar to or slightly finer
grained than the enclosing rock (the usual case), there is
no evidence of the sculpturing described by Miller. If, how-
ever, the fillings are composed of a material which is more
coarse than the host rock (e.g., coarse silt), striae are
present. When the lamellae are found in place, they com-
monly rise above the general level of the bedding plane.
Closer investigation reveals that these are not just simple
fillings but instead represent several packings stacked on
top of each other (PI. 60, fig. 7; Pl. 70, fig. 4). Sectioning
confirms that they do have a definite Sprezte-like form, This

“The earliest known reference to such “turkey track” depressions in
the Cincinnati section was by J. F. James (1885, p. 158) who placed
the Cincinnatian forms in Palaeophycus virgatum Hall. Indeed the
shallow depressions do resemble Hall’s (1847, pl. 70, fig. 1) figure
of P. virgatum from the Upper Ordovician (Utica) of Union Coun-
ty, New York. However, until the type specimen of P. virgatum is
studied, it is not possible to reach a conclusion regarding the nature
of the New York form. If it is found that the Cincinnatian and the
New York forms are conspecific, the trivial name wvirgatum would
have priority. As shown elsewhere in this work, P. virgatum is un-
like the other species of Palacophycus proposed by Hall.

leads to the conclusion that Miller described only a small
portion of a much larger and more complex structure.
Attempts to follow the forms into overlying beds
have met with little success. Normally the beds containing
them are overlain by lutites which bear no vestige of any
structure, and only at one locality (Twelve Mile Creek)
was it possible to trace the lamellae into the lutites and
then only for a few centimeters. At most localities 7. veno-
sum is associated with Chondrites and Corophioides, and al-
most invariably none of the three can be followed across
lithologic boundaries. The most obvious interpretation 1s
that the upper portion of the silt and lutite has undergone
sub-aqueous erosion. However, it is possible that in some
instances structures are actually present in lutites but are
masked by diagenetic compaction and Recent desiccation.

Few unlabeled specimens in the University of Cin-
cinnati collections indicate that 7. venosum is a more com-
plex structure than Miller’s description would indicate. The
specimens in question are in the form of long cylinders
composed of calcisiltite. The length varies from 15-25 em,
the diameter from 1.5-3 cm. Because they show the same
pattern of delicate striae on the ventral face, there can be no
doubt that these are a more complete expression of
Miller’s hemicylinders. In contrast, the dorsal surface re-
veals a series of long, parallel, discontinuous striae arranged
in groups (Pl. 68, fig. 4). Two of the cylinders possess
short vertically directed secondary branches. A lateral view
(Pl. 68, fig. 1) shows that the secondary branches remain
bundled to the main cylinder for some distance beyond the
actual point of bifurcation. A cross-section of such a struc-
ture somewhat resembles the retrusive Spreite of a U-tube.

Interpretation. — The earlier interpretations have al-
ready been noted and need not be repeated here. Flower
(1955), unaware of the genetic relationship between T.
venosum and “turkey tracks,” described the latter as rest-
ing traces of a nautiloid, probably Orthonybyoceras
Shimizu and Obata. His conclusions were based on obserya-
tions at the Stonelick Creek locality. He found four or five
examples where the aperture of the shell lies beyond the end
of the depression (Flower, 1955, fig. 1A). He felt that while
one such specimen could be dismissed as accidental, four or
five could not. He maintained that the organism came in
contact with the bottom, made the imprint and then died.
Another form (Flower, 1955, fig. 1B) shows where a shell
has swung laterally partly blurring the outlines of the trace.
In addition, Flower found a number of “turkey tracks”
grouped around a mass of shell debris. To him this indicated
several nautiloids feeding on an unfortunate group of
organisms concentrated by swirling currents.

‘TRACE FOSSILS CINCINNATI AREA; OsGoop 349

It is difficult to refute the evidence cited above, but the
fact remains that at least 25 per cent of the “turkey tracks”
at Stonelick Creek contain the lamellae described by Miller.
When these lamellae are removed, the depression is an exact
replica of the average “turkey track.” For this reason
Flower’s hypothesis is not accepted as a general explanation
for the origin of “turkey tracks.”

Instead, it is concluded that 7. venoswm is a burrow,
although, the exact nature of the entire structure is un-
known. It is believed that the organism burrowed down
through an unknown thickness of mud until it encountered
a subjacent silt bed. At this point the tunnel angled back up
into the mud. Since most of the “turkey tracks” have the
same shallow depth, it is probable that the organism was a
mud dweller; and when it encountered silt in its burrowing
activity, it changed direction in order to remain in a pelitic
environment.

It appears then that the first-formed structure was in
the shape of a gentle “U” or a flattened “U” lacking the
upper portion of one of the arms. It is not known whether
the tube was open to the surface at both ends (Text-fig.
16A) or was a blind gallery (Text-fig. 16B). Following ex-
cavation of this first tunnel, the organism backed up and
initiated the construction of a second higher tunnel, partially
truncating the upper portion of the first. The sediment re-
moved from the roof of the second tunnel was packed on the
floor of the first, giving rise to the retrusive Spreite. After
following the course of the first-formed tunnel for some
distance, the new tunnel lost contact and angled obliquely
upwards. How far it proceeded is unknown, as the distal por-
tions are truncated. None of the branches show the button-
like endings so characteristic of T. Janoswm. Presumably this
process of excavating and filling could have been repeated
several times, although the greatest number of secondary
tunnels found thus far is two.

Trichophycus has also been reported from the Ordo-
vician of Newfoundland and the Oslo, Norway, area by Sei-
lacher and Meischner (1964). It is worthy of note that at
the time this manuscript was prepared the present author
was unaware of Seilacher and Meischner’s paper; thus the
reconstructions of the Trichophycus Bawplan, which are
nearly identical, were arrived at independently.( compare

“Flower is most probably correct in assuming that nautiloid resting
traces shoud be found more frequently in the geologic record. To
date, however, only one possible example has been found by the au-
thor in the Cincinnatian. The specimen is from the Eden beds, U.S.
Route 50, 0.5 miles west of Aurora, Indiana, and is illustrated on
Plate 83, figure 3. The surface of the depression is smooth, and the
edges have raised rims as if something had pushed down into the
silt.

Seilacher and Meischner, 1964, text fig. 11 with Text-fig.
16). The Newfoundland forms appear to be identical with
the Cincinnati examples of Trichophycus. As Seilacher and
Meischner pointed out, the Norwegian burrows differ in
that they are deeper and the Spreite are better developed.

The plan of 7. venoswm is similar to that of Terchichnus
Seilacher (1955) and Pennatulites de Stefani (1885). Tei-
chichnus was erected for Cambrian forms which resemble a
stacking of very flat “U’s.” They are 1 cm wide and attain
a height of 5 cm. In plan view (Seilacher, 1955, pl. XXIV,

Text-figure 16.— Interpretation of Trichophycus venosum, Mil-
ler. A. Reconstruction showing a burrow with all branches open to
the surface. The numbers (1-3) indicate the order of formation; X 0.5.
B. Alternate interpretation showing a main shaft open to the surface
and three blind-ending branches; X 0.5.

fig. 1), they closely resemble the truncated lamellae found
in the Cincinnatian “turkey tracks,” although there are
significant differences. Teichichnus lacks the vertical
branching and striae of 7. venoswm, and some examples
exhibit a lateral branching which is unknown in 7. venoswm.
Pennatulites, from the Cretaceous and Tertiary of Europe
and Trinidad, could also be generally classified as a branch-
ing Spreitebau. Its most diagnostic feature is the knobby
prosopon found on the lower face of the lamellae (see
Hantzschel, 1962, fig. 127, 1a,b).

The branching and Spreite-like nature of Pennatulites
and Teichichnus indicates that they are most likely feeding

structures. This may be the case with 7. venoswm also, but
on the other hand, it may be a Domichnia. The Spreite
could be a reaction to sediment sifting in and _ partially
filling the burrow. Perhaps the organism first constructed
a simple shallow U-tube or blind “U” in mud. Silt then sifted
in from above, forcing the animal to raise the general level
of the burrow. This would result in Spreiten and possibly
secondary branching.

The peculiar striae found on both the upper and lower
faces of the burrow could have been made by either the
dactyls of arthropods or the chaetae of annelids. Seilacher
and Meischner (1964) suggested a trilobite-like animal, but
all known trilobite burrows possess a distinct bilobate pat-
tern which is lacking here. Whatever the nature of the or-
ganism, they must have been extremely abundant and
gregarious. One rarely finds only a single or even a few
examples of 7. venoswm. When this form occurs, it covers
the bedding planes. A distant analogy may be drawn with
the Recent decapod crustacean Callianassa, which excavates
irregularly branching burrows. In these burrows, which may
be as much as one meter below the depositional interface,
the organism sifts through the sediment feeding on micro-
scopic material. Weimer and Hoyt (1964) estimated that
the density of individuals is as much as three per square
foot, and this would be in line with the occurrence of 7.
venosum observed in the Cincinnatian.

TRICHOPHYCUS LANOSUM Miller and Dyer
Plate 68, figures 2,8

1878. Non Blastophycus diadematum Miller and Dyer, Cincinnati Soc.
Nat. Hist., Jour., vol. 1, p. 24, pl. 1, figs. 1,2.

1878. Trichophycus lanosus Miller and Dyer, Cincinnati Soc. Nat.
Flist., Jour, vols 1, py 2) plod, stigs. 354:

1884. Partim Blastophycus diadematum Miller, James, J. F., Cincin-
nati Soc. Nat. Hist., Jour., vol. 8, pp. 158-159.

1891. Partim Planolites diadematum (Miller), James, J. F., Cincin-
nati Soc. Nat. Hist., Jour., vol. 14, p. 47.

Type locality.—“Upper part of Cincinnati Group”
(Richmond), Warren County, Ohio. Type specimens: PI.
68, fig. 2 (HBM 3185; latex mold UCM 37595) and PI.
68, fig. 8 (HBM 3183; latex mold UCM 37594).

Diagnosis. — Striate vermiform trails of moderate size
with a button-like anterior(?) end.

Discussion. — The species is known only from the two
specimens figured by Miller and Dyer. The larger of the two
(PI. 68, fig. 8), which is shaped like a series of “S” curves,
is preserved as a convex hyporelief. The hemicylindrical
body is entirely covered by a series of striae which give it a
shaggy appearance, and at one end there is a small button-
like depression from which the striae radiate. Truncation

350 PALAEON'roGRAPHICA Americana (VI, 41)

prohibits observation of the entire form. The smaller speci-
men (PI. 68, fig. 2), also a convex hyporelief, possesses the
same button-like depression and radiating striae but ends
abruptly in a hemispherical “bud.”

James (1885) considered T. lanosum to be synonymous
with Blastophycus. He based his decision on the button-like
ending, which he homologized with the “bud” of Blastophy-
cus. In 1891 he placed Blastophycus, along with several
other genera, in synonymy with Planolites. Unfortunately
his reasons for this action were never explained.

Interpretation. — Miller and Dyer (1878) assigned T.
lanosum to the “fucoids,” while James (1885) explained it
merely as an “annelid trail.” James was correct in his inter-
pretation that 7. lanoswm is a trail or burrow; however, it
has nothing to do with Blastophycus. As shown elsewhere,
Blastophycus is a cast of an enrolled trilobite with associ-
ated scour marks.

The exact nature of 7. lanoswm is not clear. The small
depression with radiating striae is tentatively suggested as
the anterior end. Perhaps the striae represent the impres-
sions of a circlet of appendages or extensions surrounding
the mouth. The sinuous nature of the trace suggests a highly
flexible vermiform body. For the present these two features
serve to separate 7. lanosum from T. venosum. As men-
tioned previously 7. Janoswm may only be a behavioral vari-
ation of 7. venoswm. The former is most likely a variety
of Repichnia, the latter a Fodinichnia or Domichnia. On
the other hand, 7. lanoswm may be something different.
Until additional material of 7. Janosum can be collected, the
two are retained as separate species; however, it should be
repeated that the genus Trichophycus is best typified by T.
venosum, even though 7. lanoswm is technically the type
species.

VIII. REPICHNIA
GENERAL INTRODUCTION TO ARTHROPOD TRACKS

Arthropod tracks have been known for many years and
are figured by several authors, most notably by Hall (1852,
pls. 15-16) and Walcott (1912, 1918). Yet with the excep-
tion of Caster’s (1938) study on limuloid tracks, this group
of trace fossils has not received serious notice until relatively
recently. This is unfortunate because study of such tracks
increases our knowledge of both arthropod anatomy and
behavior.

Naturally any investigation of trilobite tracks must be
conducted indirectly. Seilacher (1955, 1959) suggested that
trilobites walked employing a wave of motion which pro-
ceeded from the rear end of the body toward the front. The

‘TRACE FOssiLs CINCINNATI AREA: OscGoop 3

pygidial appendages were the first to make contact with the
substrate, the wave moved forward along the body and as it
reached the cephalic area, a new wave was initiated at the
pygidium. The result of this would be a constant wave of
motion not unlike that seen in Recent millipedes. The tracks
produced by such movement resemble a series of “V’s” in
which the two limbs do not quite join (Text-figs. 17,18).
There is reason to believe that the open end of the “V” is
directed anteriorly. The general shape of the trilobite body
(excluding agnostids and some bizarre forms) points to
this, and studies of preserved appendages show, as one might
expect, that in the pygidial area the distance between pairs
is not so great as in the thoracic region. This holds true
even for such forms as Jsotelus and Cryptolithus, which
have ovate or ellipsoidal body outlines. Seilacher (1955)
also presented evidence to support his theory that the wave
proceeded anteriorly. If the movement had been in the op-
posite direction, the distance between the first and last im-
pression in a wave measured along the axis of movement
would be short, and the angle formed by the limbs of the
“V” would be well over 90°. This is not supported by the
fossil record, for the series are drawn out and the acute
angle of the “V” is less than 90°.

It should be stressed that while Seilacher’s hypothesis
regarding trilobite movement is logical, it is nevertheless
only a theory. Unfortunately trilobite remains have never
been found at the end of a trail. This is in contrast to the
xiphosurids at Solnhoten where several corpses have been
preserved at the end of their “death march” (Walther,
1904).

It is noteworthy that in xiphosurid trails the point of
the “V” is directed anteriorly as Caster (1938) showed. At
first this is a disquieting discovery, because xiphosurids are
presumably the trilobites’ closest living relatives. However,
even early limuloids were highly specialized, having under-
gone fusion of the thoracic area and modification and re-
duction of the walking legs which are restricted to the
prosoma (cephalic shields). Moreover, unlike trilobites,
xiphosurids possess differentiated walking legs. Just as their
anatomy has become specialized, so has the method of walk-
ing. Caster (1938, fig. 5) showed that by doubling up on the
cephalic-buckler joint and alternating the placement of the
last pair of walking legs (pushers) with the remaining pairs,
the animal tends to move forward in a series of lurches. Even
though it is conceded that xiphosurids and trilobites prob-
ably walked in somewhat different ways, some of what has
been learned from the study of xiphosurid movements can
be applied to the analysis of trilobite trails.

Before the more detailed discussion can be attempted,

mn
=

it is necessary to define certain terms. Unfortunately this
has not previously been done in English and Seilacher’s
German terms defy translation. For these reasons most of
the terms illustrated in Text-figure 18 and discussed below
are new. Because most trilobite tracks are preserved, or at
least collected, as convex hyporeliefs and are best studied
this way, all further discussions are given as if the viewer
were looking up at a trilobite walking on a glass table top.
Thus, when we speak of right-hand side or left-hand side,
the reader must bear in mind that anatomically speaking
these terms are reversed. The following terms are useful in
the discussion:

trail —the whole record of one walk. It is normally composed of many
complete sets.

complete set (=Trittsatz of Seilacher, 1955) —the tracks formed by a
single wave of movement. A complete set is in the shape of an open-
ended “V”’ directed posteriorly.

series (=Trittseries of Seilacher, 1955) —either the right- or left-
hand limb of the “V” as seen by the viewer. These do not
conform to right and left in the anatomical sense.

track — an essentially undeformed mold (or cast) of a leg or foot,
not a scratch or other ethologic expressions of an appendage.

imprint (=Trittsiegel of Seilacher, 1955) —a single leg record of
any sort, deformed or undeformed. The proximal portion of an im-
print is closest to the axis of movement; the distal portion farthest
from the axis. Proximal and distal refer only to the imprint, not
to the leg itself.

pair —two imprints or tracks, one on each limb of the “V,’ which
are judged to represent the trace of a pair of legs.

length of set—the length of each set is measured along the axis of
movement.

width of set (=Trittabstand of Seilacher, 1955) —the width is the
distance from the distal termination of one imprint of a pair to the
distal end of the corresponding imprint. Where the width varies in
a set, two measurements, indicating the range of variability, are
given.

withdrawal markings —semi-lunar welts (depressions on casts)
marking the withdrawal of the legs from the sediment. These are
normally found at the proximal end of the imprints.

Trilobite tracks fall into three intergradational groups:
A) those in which the direction of movement was coinci-
dental to the body axis, B) those in which the direction of
movement was slightly oblique (+ 30°) to the body axis,
and C) those in which the direction of movement was al-
most at right angles to the body axis, which might be termed
“crab-walking.”
A) Movement parallel to body axis

As Seilacher (1955) remarked, it is indeed fortunate
that trilobites did not choose to walk by this method all of
the time, or we would know little about their tracks, for in
this manner of locomotion the various sets become super-
imposed; and it is nearly impossible to determine the num-
ber of imprints in a series (see Text-fig. 17A and PI. 73, fig.
3). Seilacher felt that this was particularly true when the
trilobites were moving rapidly, apparently using only the

ws)
wn
bhROo

Sa

A

Text-figure 17.— Morphology of trilobite tracks. A. The order
of emplacement of the legs on the substrate as viewed from below.
B. A trilobite “crab-walking” and giving rise to dimorphic tracks.
C. Strongly dimorphic tracks. Seilacher (1955) believed that these were
grazing or browsing trails (Weidespur). (A and B after Seilacher,
1955, figs. la,e; C after Seilacher, 1955, fig. 2B.)

stronger anterior legs, while the rest of the body was
dragged along with the possibility that the caudal cerci
or pygidial spine(s) (when present) acted as skids. It is
likely that straight-forward walking at any rate of speed
gave a superimposed and indistinct record, no matter how
few or how many legs were employed. At any rate, virtually
no traces of straight-forward locomotion are known in
which sets are clearly decipherable.

Such tracks have been described from several horizons
and under such a bewildering variety of names that a
monographic study is necessary to sort them out. Two of
the more frequently mentioned names are Diplichnites
Dawson (1873) and Protichnites Owen (1852).

Dawson’s (1873, p. 20, fig. 3) illustration of the type
species of Diplichnites is so schematic that it is impossible
to determine if it is a trilobite track. Dawson suggested
that the form might represent imprints of the pectoral
spines of fish. Moreover, because the genus occurs in flag-
stones associated with coals, there is some doubt that it is
even of marine origin. Ignoring these uncertainties, Seil-
acher (1955) used the name for trilobite trails from the
Lower Cambrian of the Salt Range.

Protichmtes Owen (1852), originally described from
the Upper Cambrian Potsdam Sandstone of eastern North
America, has been used by both Seilacher (1955, pl. XVI,
fig. 2) and Opik (1959, figs. 2-6) for trilobite tracks show-
ing straight-forward movement. In both cases there is a
median groove separating the right and left series. The
nature of the Potsdam Protichnites is even more enigmatic
than Diplichnites. Dawson (1873) suggested xiphosurids,
while Billings (1870) questionably assigned the tracks to
either trilobites or aglaspids. Raymond (1920) accepted a

PALAEONTOGRAPHICA AMERICANA (VI, 41)

trilobite origin, but Burling (1917, p. 387) suggested a
“short low-lying, more or less heavy set, approximately 12-
legged, crab-like animal.” Caster (1938, p. 28) maintained
that Protichnites is probably of gastropod origin.

Finally, Walcott (1918, pl. 38, figs. 1, 2) illustrated
tracks from the Tonto Shale of the Grand Canyon, Ari-
zona, which show straight-forward movement. He neither
named nor discussed them.

The above discussion illustrates that while straight-
forward tracks are known from various horizons and loca-
tions, there is no uniformity of opinion as to the interpre-
tation or the application of names. The above treatment is
limited to tracks of probable trilobite origin. For a discus-
sion of possible xiphosurid tracks, see Caster (1938, pp.
26-36).

B) Movement slightly oblique to body axis

Generally, trilobites moved in a slightly oblique man-
ner much as modern crabs, and frequently this variety
of movement was combined in the same trail with that just
discussed. A trilobite can be observed to have started with
straight-ahead movement then switched to slightly oblique
movement before changing back to straight-ahead move-
ment, thus leaving an alternating record. This oblique
locomotion produced a characteristic track morphology.
Such a record is seen in Text-figure 17B where the body
was angled slightly to the right. Note that the left-hand
series forms a line of superimposed imprints, making it im-
possible to sort them out into separate series. In contrast,
the nine imprints of the right-hand series are plainly de-
fined. If the number of imprints or tracks in a set are
known, the chances of determinating the originator of the
trail are improved,

Trilobite tracks showing oblique movement are known
from at least three different localities. Rudolf Richter
(1941, fig. 13) illustrated one such form from the Devonian
Hunsriick Shale of Germany, and Abel (1935, fig. 222)
figured another from the Silurian Clinton beds of New
York State. Seilacher (1955) described tracks of this form
from the Salt Range Cambrian under Diplichnites.

There are additional explanations for dimorphic tracks
than the one discussed above. Caster (1944)described a
xiphosurid trail from the Newspaper Rock Member of the
Chinle Formation (Triassic) of Arizona, where the right-
hand series is more deeply impressed than the left. He
theorized that the organism was struggling to keep its bal-
ance on a sloping beach. In the same manner a slippery
mud slope on the sea floor could affect a trilobite and lead
to a dimorphism of the tracks. Likewise, a moderate cur-
rent on a level surface could drive the trilobite sideways

TRACE FOSSILS CINCINNATI AREA: OsGoop 353

or momentarily upset its balance so that it might lash
out in an attempt to regain its equilibrium. Here again
the result would be a dimorphism of the imprints. It is thus
evident that not all dimorphic tracks were the result of
simple voluntary oblique, movement. When current indicat-
ors are present, they should be studied to see if they might
have had some bearing on the morphology of the trail.

C) Movement strongly oblique to body axis

Trilobite trails in which the direction of movement
was nearly at right angles to the body axis are thus far
known only from the Lower Cambrian of Pakistan and
Sweden. In such specimens (Text-fig. 17C) there is a
marked dimorphism in the configuration of the right- and
left-hand series, which suggested to Seilacher (1955) the
generic designation Dimorphichnus. In the track illustrated
in Text-figure 17C, the body axis was oriented far to the
right of the direction of movement. The imprints of the
right-hand series indicate a long raking motion. If these
are long enough, they resemble a sine curve, this type of
curve being the result of two vectors, one directed in the
line of movement, the other normal to the line of move-
ment. Combining this information with the “rule of the
V’s” it is possible to determine the direction of movement,
in this case from bottom to top. Markings such as those
seen in the right-hand series may be called “raking im-
prints,” (—Harkstegel of Seilacher). Seilacher believed that
raking imprints were the result of the legs being passively
dragged over the substrate. If this were true, they would
offer little support for the body, and it remained for the
left-hand appendages to provide such support. This appears
to be the case, as the imprints of the left-hand series show
no such raking movement and instead are short and stumpy
and commonly impressed more deeply than the raking im-
prints. Such imprints are termed “support imprints”
(=Stemmsiegel of Seilacher). Seilacher maintained that
Dimorphichnus has definite paleoecologic significance. It
was his thesis that this was not merely another form of
Repichnia but instead represented a grazing (scavenging)
trace and should be classified among the Pascichnia. This
hypothesis was based on studies of a large slab from the
Salt Range (Seilacher, 1955, fig. 3), which measured 165
cm in length and bore 23 complete sets and over 360 in-
dividual imprints. Two lines of evidence led Seilacher to
his conclusion. First, as the trilobite was employing the
strongly oblique movement, it encountered the tracks of
a second redlichid. At this point the organism turned
around and proceeded back in the direction from which it
came, being careful to avoid crossing over its own track.
Eventually it reached a point where it must cross over,

but it did so without utilizing the raking motion. The
raking motion ceased as it crossed the track and picked
up again on the other side. Finally, near the edge of the
slab the strong raking motion converted to moderate oblique
movement. As Rudolf Richter (1938, 1941) showed, vermi-
form grazing trails rarely cover the same area twice, The
tightly packed meanders of Helminthoida and other such
forms from the European Flysch never come in contact.
This is a principle of economy; there is no point in “re-
grazing” an area once it has already been picked clean.
Seilacher felt that the same is true here. This would ex-
plain why the “grazing” trilobite turned around when it
encountered another track, and it also explains why the
trilobite moved so quickly over its own track. The little
that is known of trilobite feeding habits does nothing to
discredit Seilacher’s thesis, for the fossil record indicates
that trilobites possessed neither strong mandibles nor gnat-
hobases; apparently they were not predators. Stgrmer
(1951) mentioned that the flabellate pre-epipodites, whose
main function was to serve as ctenidea, may have also acted
as sieves on which food could be collected. Employing this
idea, Seilacher pictured the nutrient-rich sediment moving
up the leg, where, due to the curvature of the telopodite,
it came in contact with the pre-epodite. From here it moved
to the mouth.

Although Seilacher’s hypothesis is an ingenious one,
he is perhaps reading too much into the record. The trail
could just as easily have been caused by an oscillatory
current (é.g., wave action) which first drove the trilobite
one way and then carried him back again. Moreover, it is
not clear how the food was carried from the legs to the
mouth, If Seilacher’s hypothesis is correct, the method
he described must have been an early experiment in feed-
ing which proved ineffective and was abandoned by later
forms. As mentioned, Dimorphichnus is known only from
a few specimens from the Lower Cambrian of Pakistan and
Sweden. There is no evidence that Cincinnatian trilobites
fed in the manner described by Seilacher.

While trilobite tracks tell us something about be-
havioral patterns, they also contribute to our knowledge
of the morphology of the walking legs, especially the distal
portions. Due to the imperfections of the fossil record, we
have relatively little direct evidence in this field. What is
known has been summarized by Raymond (1920) and
Stormer (1939).

Even if it is possible to assign a set of tracks to a
given trilobite, many times it is difficult to determine the
exact nature of the dactyls. Some forms, such as that illus-
trated in Plate 73, figure 1, present a bewildering variety

354 PALAEONTOGRAPHICA AMERICANA (VI, 41)

of imprints. Note that trifid, bifid, and unifid imprints are
all present in the same trail. There are several possible ex-
planations for such discrepancies. It is likely that the man-
ner in which the leg was placed on the substrate could effect
the morphology of the imprint. An appendage might strike
the substrate in such a way that all three dactyls make
contact, resulting in a trifid track, but if the leg is rotated
slightly to one side or the other, only two dactyls may
leave imprints. Conceivably unifid impressions could be ex-
plained in the same manner. The curved outline of the
imprints in Plate 73, figure 1 support this view.

S Withdrawal markings we
+ tA

\s mo

Width of set

d SERIES

Sars Potir =e va

Imprint
vi

SY i)

Length of set

Text-figure 18.— Morphology of trilobite tracks. See the text
for definition and discussion of the terms employed.

As shown in Text-figure 19 preservation may play a
key role in the appearance of any arthropod track. Occa-
sionally tracks are preserved as convex hyporeliefs in cal-
cisiltites which overlie clay-sized material (Pl. 74, fig. 1).
This method of preservation usually provides us with the
greatest clarity of detail. More frequently, however, the
tracks are found as cleavage reliets in fine-grained calcisil-
tites. Here the fine details are lost and, depending on how
far below the surface the imprint is found, the morphology
may vary considerably. An imprint found 2 mm below the
surface may deviate greatly from that found 4 mm below
the surface. Sometimes the vagaries of preservation make
it difficult to recognize that apparently different forms are
in reality the same thing.

SYNOPSIS OF THE VENTRAL MORPHOLOGY OF
CINCINNATIAN TRILOBITES

Something is known of the ventral surface of the more

common Cincinnatian trilobite genera. Specimens recovered

Text-figure 19. — Effect of preservation on arthropod track mor-
phology. If the worker is not aware of the effects that preservation
can have on track morphology misinterpretation will follow. The
track illustrated is hypothetical; each block is 1 mm thick.

Top. The imprint is impressed as a concave epirelief on the deposi-
tional interface. The track is quadrifid and has an arcuate posterior
fringe.

Center. The imprint is preserved as a cleavage relief 1 mm below
the depositional interface. The arcuate fringe is not preserved.
Bottom. The imprint is preserved as cleavage relief 2 mm below
the depositional interface. Only the two interior imprints are preserved.

from the Middle Ordovician black shales of Trenton Falls,
New York, have been the subject of studies by Walcott
(1881, 1884, 1918), Beecher (1893-1902), Raymond (1920),
and St@rmer (1939). Unfortunately with regard to the ter-
minal digit and setae Walcott, Beecher, and Raymond em-
ployed the artist’s license; and their reconstructions must
be used with extreme caution. In discussing the appendages
of each of the genera, emphasis is placed on those structures
which may have left their imprints on the substrate.
1) Lsotelus

The best preserved specimen of Jsotelus showing ap-
pendages is from Oxford, Ohio (Pl. 57, fig. 5), and has
been discussed previously. Twenty-six pairs of appendages
have been reported; 10 on the thorax and 16 on the pygi-
dium. Judging from their size, the thoracic appendages
must have been powerful. Nothing is known of the nature
of either the pre-epipodite or the terminal segment of the
walking legs. Walcott (1884) figured setiferous legs, but
Raymond (1920) could find no evidence to support this
conclusion.

2) Cryptolithus

The exact number of appendages in Cryptolithus is un-
known. Raymond’s (1920, fig. 20) reconstruction shows 22
with the last 10 or so being small, Stgrmer (1939, p. 211)

TRACE FOSSILS CINCINNATI AREA: Oscoop 355

noted that “the telepodites are strongly curved in the distal
portion and the position in preserved state indicates that
this part was directed downwards and slightly backwards in
the thorax and pygidium during life.” Although the pre-
epipodite is net well known, it appears to have resembled a
large fan (Stérmer, 1939, p. 25). The morphology of the
dactyls is clearly shown by Raymond (1920, pl. 7, figs. 3-4).
They resemble a tuft of bristles, although Beecher’s recon-
struction (Raymond, 1920, fig. 45) shows a large terminal
spine surrounded by slightly smaller spines. Although both
Raymond (1920, fig. 20) and Stgrmer (1939, fig. 25) show
a grouping of proximally directed spines on each segment of
the telepodite, it is not possible to confirm this from Ray-
mond’s illustrations.
3) Ceraurus

Raymond’s reconstruction of Ceraurus (fig. 16) shows
26 pairs of appendages, 18 of which are large. The nature
of the dactyls is problematic. Walcott (1881) illustrated
a single long terminal spine. Raymond (1920) agreed that
while there probably was a terminal spine, it was not as
well developed as Walcott had shown. In a later paper
(1918, pl. 34, figs. 1, 2), Walcott clearly showed three
spines, and Stgrmer (1939, pl. 11, fig. B) dotted in three
spines, illustrating that the true nature of the dactyls is
really not known. Stgrmer (1939, pl. lla) did extensive
work on the pre-epipodites and found that the unicorn-like
spirals of Walcott (1918) and Raymond (1920) are due
to preservation. Instead he pictured a small fanlike mem-

ber.
4) Flexicalymene

Stormer (1939) remarked that although he had not
studied Flexicalymene in detail, sections appear to reveal
appendages similar to those of Cerawrus. Raymond (1920,
p. 56) admitted that nothing is known of the telopodites
and that any conclusion must be based on analogies with
other forms.

5) Triarthrus

Raymond’s (1920, fig. 10) reconstruction of Triarthrus
indicates 24 pairs of appendages. He reported that one
specimen possessed a terminal segment which consisted of
two or three short hairlike spines with a ball and socket
joint. In addition, Raymond illustrated the legs as possess-
ing long setae and bearing a grouping of spines on the
segments much like those of Cryptolithus. Stormer (1939,
pp. 204-205) stated that he had not studied the distal por-
tion of the telopodite, but he believed that Raymond’s
conclusions appeared to be valid. In addition he pictured
a large flabellate pre-epipodite similar to that of Crypto-
lithus.

CINCINNATIAN REPRESENTATIVES

Although arthropod tracks are not uncommon in the
Cincinnatian section, they have never received adequate
study, and this was the one field which J. F. James neglect-
ed entirely. Miller (1880) established the genus Asaphoi-
dichnus for two species of asaphid tracks, and Caster (1938)
figured and briefly discussed four specimens, placing three
of them in Asaphoidichnus, while referring to the other as
Merostomchmites sp. In spite of this small amount of
work, there is an abundance of names, for Miller (1880) un-
wittingly erected three additional genera and five species
of arthropod tracks. These were collected from the ?South-
gate beds of “Walker Mill Road” (now State Avenue) in
Cincinnati, and because of their association with molluscan
body fossils, Miller considered them to be cephalopod
tracks. However, as will be shown, these are clearly tracks
of arthropods and most probably were made by trilobites.

As pointed out previously, there is considerable doubt
as to the true affinities of the two most commonly men-
tioned genera of supposed trilobite tracks, Protichnites Owen
and Diplichnites Dawson. Because of this uncertainty, the
Cineinnatian tracks will be discussed within the framework
of Cincinnatian nomenclature.

Five genera, all monotypic, are recognized in this
study. One of these, Asaphoidichnus trifidum Miller (1880),
was correctly interpreted by Miller as the tracks of an
isotelid. However, what Miller (1880) established as Asa-
phoidichnus dyeri becomes the type species of a new genus,
Allocotichnus. As Miller originally indicated, this too is
probably an isotelid track, but it illustrates a type of move-
ment not previously seen in trilobite tracks. Three other
Miller species, Trachomatichnus permultum, T. cincinna-
tiensis, and Petalichnus multipartitum, as well as what
Caster (1938, pl. 10, fig. 1) described as “Merostomich-
nites sp (cf. Petalichnus multipartitus),” are all considered
to be Petalichnus multipartitum. These are most likely
tracks of Flexicalymene, but since the nature of the dactyls
of most Cincinnatian trilobites is unknown, there is no way
of corroborating this. Trachomatichnus numerosum Miller
(1880) appears to be generically valid and may well repre-
sent the tracks of the brushlike dactyls of Cryptolithus.
Teratichnus confertum Miller (1880) is retained; it shows
what appears to be the impression of a terminal spine, a
feature which sets it apart from all other Cincinnatian
arthropod trails.

In order to illustrate better the number of imprints
in a series and to show the direction of movement, tracings
were prepared of some of the specimens (see Text-fig.
20-23). In some areas the imprints are so indistinct or

crowded together that they tend to defeat the purpose of
the illustrations, and they were, therefore, omitted. The
overlays represent the author’s interpretation of the trails.
The different markings are used to differentiate the oppos-
ing rows of the various series. The trilobite outlines super-
imposed over the drawings indicate the orientation of the
body and the direction of movement.

ASAPHOIDICHNUS Miller, 1880

Plate 65, figure 2; Plate 71, figure 6; Plate 72, figure 3; Plate 73,
figures 1,2; Text-figures 20-A,21-A,29-f

1880. Partim Asaphoidichnus Miller, Cincinnati Soc. Nat. Hist., Jour.,
vol. 2, pp. 217-219 (refers to unnamed specimen figured by
author zm Cincinnati Quart. Jour. Sci., vol. 1, p. 136, fig. 11).

1938. Asaphoidichnus Miller, Caster, Jour. Paleont., pl. 10, figs. 2-4.

1962. Partim Asaphoidichnus Miller, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W184.

Type species. — Asaphoidichnus trifidum Miller (1880)
(PI. 11, fig. 6; MCZ 540; latex mold UCM 37704) from
“less than 100’ above low water on Walker Mill Road, Cin-
cinnati, Ohio” (?Southgate of State Avenue, Cincinnati,
Ohio). Monotypic.

Diagnosis. — Large, trifid arthropod tracks, exhibiting
both straight-ahead and oblique movement. Width of track
varies from 6.0-15.5 cm; number of imprints per set aver-
ages nine.

Discussion. — Miller (1880, p. 217) said of Asaphor-
dichnus that “the tracks are separate and distinct, showing
that the animal lifted its feet to advance, and appearing as
if made by an articulated animal, but not by one holding
its feet in an inflexible position.” Two species, A. trifidum
and A. dyeri were named; the latter was placed in the
genus with some misgivings, which appear to be justified,
as its morphology varies greatly from any known trilobite
track. It is here removed from Asaphoidichnus and is made
the type species for the new genus Adlocotichnus.

Miller (1880, p. 218) erected A. trifidwm for tracks
where “the anterior third of each track is trifid, the outer
toe usually branching from the main stem of the track
first, and the middle being a little longer than either the
outer or inner one.” He also maintained that one paratype
specimen shows “a small toe thrown off from the posterior
elevation on the inner side of the track, having a length
about equal to the inner anterior toe” (ibid).

Although Miller remarked that several specimens of
A. trifidum were in the Dyer collection, only the type could
be located. The trail (PI. 71, fig. 6), 3.5 cm long and 6.0 cm
wide, consists of 14 impressions on the right-hand side and
six on the left. The individual impressions vary in their

356 PALAEONTOGRAPHICA AMERICANA (VI, 41)

morphology; the best preserved ones show an oblong raised
welt from which radiate three thin lines. At their point of
termination the trifid imprints, whose length varies from
1.5-2.0 cm, may be separated from each other by as much
asi3 mm! (PI 72 .stie.s.)

Examination of the right side of the trail reveals that
not all the tracks belong to one series, As shown in Text-
figure 20-A, six of the tracks are from one series, six from
another. The left-hand side of the trail can likewise be dif-
ferentiated into two series, although the imprints are not
as clearly defined. None of these are judged to represent a
complete set.

Interpretation. — The width of the trail (6.0 cm) elim-
inates all known Cincinnatian arthropods except /sotelus,
eurypterids, and possibly aglaspids. The morphology of the
best known Cincinnatian eurypterid, Megalograptus, has
been studied in detail by Caster and Waering (1964). They
have shown that the four pairs of “walking legs” are strong-
ly differentiated. The first pair is short, bears numerous
spines, and was probably used more for grasping than for
walking. The second pair is greatly enlarged and armed
with many long slender spines; it probably did not play
an active role in walking. The third pair of “walking legs”
is short and possesses a great number of spines, while the
fourth pair is slightly longer than the third, and except for
trifid dactyls, is devoid of spines. The fifth and final pair
of legs is modified for swimming, as both the fourth and
seventh joints are enlarged. Even though the fourth pair
of “walking legs” is trifid, it is extremely difficult to im-
agine the type specimen of A. trifidwm as an eurypterid
track. Other eurypterids were present in the Cincinnatian
seas, for scraps of their integument are found scattered
throughout the section. These fragments more closely re-
semble Eurypterus than they do Megalograptus, but there
is not sufficient material available to undertake a morpho-

—>

Text-figure 20.— NOTE: Text-Figures 20-23 repre-

sent the authors interpretation of several] arthropod
trails. The organisms superimposed over the tracks
indicate the direction and method of movement. In the
acetate overlays different symbols are used to portray
imprints in different series. In all cases the trails are
viewed from below, i.e. as if the viewer were looking
up at the trails through a glass table top.
A. Asaphoidichnus trifidum Miller, 1880. Repichnia of
Isotelus; interpretation of the trail illustrated on
Plate 71, figure 6. Holotype. MCZ 540; latex mold
UCM 37694, X 1.2. B. Trachomatichnus numerosum
Miller, 1880. Repichnia of Cryftolithus; interpretation
of the trail shown on Plate 73, figure 4. It is difficult
to separate the imprints into series. MCZ 544; latex
mold UCM 37671, X 3.2.

357

Trace Fossits CINCINNATI AREA: OsGoop

—

crowded together that they tend to defeat the gurpose of
the illustrations, and they were, therefore, omitted. The
overlays represent the author’s interpretation ok the Pails.
The different markings are used to differentiate the oppos-
ing rows of the various series. The trilobite vutlines super-
imposed over the drawings indicate the grier®ation of the
body and the direction of movement® 4 ?
oy )
ASAPHOIDICHNUS Milley, 1880

4
Plate 65, figure 2; Plate 71, figure @ ¥ ate 72, figure 3; Plate 73,
figures 1,2; Text-$@zures 20-A,21-A,29-f

1880. Partim Asaphoidichnus mie, Me canee Soc. Nat. Hist., Jour.,
vol. 2, pp. 217-219 (ixfags fo unnamed specimen figured by
author in Cincinnati Quart.” Jour. Sci., vol. 1, p. 136, fig. 11).

1938. Asaphoidichnus Mi er, Aster, Jour. Paleont., pl. 10, figs. 2-4.

1962. Partim Asaphoidichn® Miller, Hantzschel, Trace-Fossils and
Problematica in Treatiy on Invertebrate Paleont., Moore (ed.),
pt. W, Miscelianey. p: W184.

*
Type species. 7 gAsaphoidich nus trifidum Miller (1880)
(PI. 11, fig. 6; MICZ 540; latex mold UCM 37704) from
“less than 100’ aBove low water on Walker Mill Road, Cin-

cinnati, Ohio” (?Southgate of State Avenue, Cincinnati, §

Ohio). Monotypic.

ages nine. ss
Discussion. — Miller (1880, p. 217) said of Asapho®,

dichnus that “the tracks are separate and distinct, shoygne@

that the animal lifted its feet to advance, and appearingmas

if made by an articulated animal, but not by one hol&

its feet in an inflexible position.” Two species, A. Be

and A. dyeri were named; the latter was placed in te

genus with some misgivings, which appear to be Jjustifi l

as its morphology varies greatly from any known trilobite
track. It is here removed from Asaphoidichnus and is madef
the type species for the new genus Allocotichnus. ig

Miller (1880, p. 218) erected A. trifidwm for frge S
where “the anterior third of each track is trifid, they@yter
: : y)
toe usually branching from the main stem of the by ck
i . . . . ta
first, and the middle being a little longer thanyeither the
outer or inner one.” He also maintained that oné pgratype
specimen shows “a small toe thrown off from th posterior
elevation on the inner side of the track, havy a length

about equal to the inner anterior toe” (ibid).4

Although Miller remarked that seve Al b ecimens_ of
A, trifidum were in the Dyer collection, ondy rs type could
be located. The trail (Pl. 71, fig. 6), 3.5 erp long and 6.0 cm
wide, consists of 14 impressions on the rigpt-hand side and
six on the left. The individual impress{ is vary in their

356 PALAEONTOGRAPHICA AMERICANA (VI, 41)

morphology; the best preserved ones show an oblong raised
welt from which radiate three thin lines. At their point of
termination the trifid imprints, whose length varies from
1.5-2.0 cm, may be separated from each other by as much
as 3 mm (PI. 72, fig. 3).

Examination of the right side of the trail reveals that
not all the tracks belong to one series, As shown in Text-
figure 20-A, six of the tracks are from one series, six from
another. The left-hand side of the trail can likewise be dif-
ferentiated into two series, although the imprints are not
as clearly defined. None of these are judged to represent a
complete set.

Interpretation. — The width of the trail (6.0 em) elim-
inates all known Cincinnatian arthropods except /sotelus,
eurypteridsy and possibly aglaspids. The morphology of the
best know Cincinnatian eurypterid, Megalograptus, has
been studiéfl in detail by Caster and Waering (1964). They
h%y% shown that the four pairs of “walking legs” are strong-
ly ¥dafferentiated. The first pair is short, bears numerous
spigs, and was probably used more for grasping than for
walkgg. The second pair is greatly enlarged and armed
with many long slender spines; it probably did not play
an active role in walking. The third pair of “walking legs”
is short and possesses a great number of spines, while the
fourth pair is slightly longer than the third, and except for
trifid dactyls, is devoid of spines. The fifth and final pair
of legs is modified for swimming, as both the4fourth and
seventh joints are enlarged. Even though dy? fourth pair
of “walking legs” is trifid, it is extremel ifficult to im-
agine the type specimen of A. trifidwm as an eurypterid
track. Other eurypterids were present in the Cincinnatian
seas, for scraps of their integument arg gkqund scattered
throughout the section. These rragtiaeeee closely re-
semble Eurypterus than they do Mfgalogra tus, but there
is not sufficient material yee un ake a morpho-

/ ¥

a Pie

Text-figure 20.— NOTE: Text-Figures 20-23 repre-

sent the authors interpretation of several arthropod
trails. The organismd# superimposed over the tracks
indicate the directiond/and method of movement. In the
acetate overlays dietbrgit symbols are used to portray
imprints in different » ries. In all cases the trails are
viewed from below,y.e. as if the viewer were looking
up at the trails thréugh a glass table top.
A. Asaphoidichnud trifidum Miller, 1880. Repichnia of
Isotelus; interpretation of the trail illustrated on
Plate 71, figure 6. Holotype. MCZ 540; latex mold
UCM 37694, X 1.2. B. Trachomatichnus numerosum
Miller, 1880. Repichnia of Cryftolithus; interpretation
of the trail shown on Plate 73, figure 4. It is difficult
to separate the imprints into series. MCZ 544; latex
mold UCM 37671, X 3.2.

TRACE FOSSILS CINCINNATI AREA: OsGoop

357

358 PALAEONTOGRAPHICA AMERICANA (YI, 41)

logical study. In any event it is likely that eurypterid tracks
would more closely resemble the tracks of Recent scorpions
(Seilacher, 1959, fig. 2C) than those now under considera-
tion.

Neostrabops Caster and Macke (1952), the only
known non-Cambrian American aglaspid, was described
from the Corryville section at Stonelick Creek. Although
the unique specimen is too small (3.5 cm x 2.8 cm) to have
made the trail, Raasch (1939) described Upper Cambrian
aglaspids from Wisconsin which attained a length of 18.6
cm. Because the appendages of Neostrabops are unknown,
their morphology must be inferred from Raasch’s Cambrian
forms. Aglaspis spinifer Raasch (1939) possesses 12 and
possibly 14 pairs of “walking legs” which do not extend be-
yond the body margins. Each leg has a single terminal
dactyl, and Raasch was unable to find any evidence of
setae. It seems improbable, therefore, that A. trifidwm rep-
resents the tracks of an aglaspid. The Crustacea of the
Cincinnatian are so inadequately known that no generaliza-
tions are possible.

This leaves /sotelus as the most likely candidate to have
produced the track. Even though it can not be unequivocally
stated that A. trifidwm is an isotelid track, it is probable
that /sotelus possessed three long slender dactyls which,
when the need arose, could spread to provide a flabellate
“snowshoe-like” support. In this sense the dactyls may have
been analogous to the blades of the fifth pair of “walking
legs” of Limulus.**

Because there is no marked dimorphism in the right
and left series, the track is indicative of straight-ahead
movement. However, the morphology of this specimen dif-
fers from many trilobite trails in showing vertical emplace-
ment of the legs without any appreciable movement toward
the median line. In most trails after initial contact with
the substrate, the leg moved toward the median line, leaving
a scar with a small raised pimple marking the point of
withdrawal (Text-fig. 18). Thus, A. trifidwm represents
relatively undisturbed tracks. Moreover, the method of
walking is probably responsible for the unusual type of
preservation which was noted by Miller (1880). The pos-
terior oblong welts mark the highest point of the track.
Near the welt the trifid imprints are preserved as thin
raised ridges (1.e., “convex”), but as they are followed dis-
tally, the ridges grade into shallow depressions (i.e., “con-
cave”). For this reason, the trail shows the characteristics

“The meaning of “the small toe’? mentioned by Miller is not clear.
There is no trace of any such structure on the specimen under discus-
sion. It most likely represents one track partially superimposed over
another.

of both convex hyporelief and concave epirelief preserva-
tion. It is concluded that the trail is preserved as an epi-
relief and that the mud was drawn up by the proximal
portion of the dactyls as the weight was shifted distally.
This would be analogous to a man drawing up mud with
his heel as he shifted his weight to the ball of the foot.

Miller (1880) also believed that the trail was preserved
as an epirelief. However, he remarked that the feet might
have been webbed to aid in swimming. While this might
explain the peculiar preservation, there is no evidence that
the organism was “web-footed.”

Caster (1938, pl. 10, figs. 2-4) figured portions of ad-
ditional trails which, because of their size, were most prob-
ably made by J/sotelus. All are preserved as convex hypo-
reliefs and are from the Fulton beds of the Union Levee,
Cincinnati. One of these (Pl. 73, fig. 2) shows marked
dimorphism. The five right-hand series consist of long,
slightly sigmoidal raking imprints, while the left-hand
series are made up of superimposed support imprints. The
trail width ranges from 2.2-4.0 cm. As noted elsewhere, the
markings vary in their appearance; some are unifid, while
others are bifid or trifid. Only nine imprints can be count-
ed in any given series, but none of the series appear to be
complete, and most likely the total answer was greater
than this.

Plate 75, figure 2 illustrates another specimen from
the Fulton beds which, judging from the width (15.5 em),
is probably the trail of an isotelid. The right-hand series
consist of eight to ten long sigmoidal raking imprints, and
in contrast to the form just discussed, they show no evi-
dence of being bifid or trifid. The support imprints of the
left-hand series also show some slight sigmoidal develop-
ment. A more detailed analysis of this form is given in
Text-figure 21-A.

Both of these dimorphic trails are indicative of oblique
movement. In both cases movement was from bottom to
top as viewed by the reader. Either the animal was volun-
tarily “crab-walking” with the body angled slightly to the
right of the axis of movement, or it was being swept to
the left by current. The former appears the more likely as
neither of the specimens bear any current indicators.

ALLOCOTICHNUS, n. gen.

Plate 72 figures 1,2; Plate 73, figures 7; Plate 74, figures 6,7; Plate
81, figure 7; Text-figures 21-B,C,29-h

1880. Partim Asaphoidichnus Miller, S. A., Cincinnati Soc. Nat.
Hist; Jour, svol..2.) ps 209 apl is etigee.

1962. Partim Asaphoidichnus Miller, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W184, fig. 110-3.

TRACE FOSSILS CINCINNATI AREA: OsGoop 359

Type species — Asaphoidichnus dyeri Miller (1880)
(Pl. 72, fig. 1; MCZ 543; latex mold UCM 37693) from
?Southgate beds of “Walker Hill Road,” (State Avenue)
Cincinnati, Ohio, Monotypic.

Diagnosis — Bifid, dimorphic trails of arthropod ori-
gin. Each complete set apparently composed of four pairs
of imprints, one side being preserved as long subparallel
raking imprints, the other arranged as en-echelon support
imprints.

Discussion — Miller (1880, p. 219) established A. dyeri
for a trail with four toes “. . . which do not come together
as in A. trifidus but spring from a wider foot.”

The type specimen (PI. 72, fig. 1), which has a length
of 23 cm, consists of 10 separate groups of markings and
is preserved as a convex hyporelief in a platy fine-grained
calcisiltite. The five complete groups each possess four
long subparallel imprints, as well as four shorter imprints
which are arranged in an en-echelon pattern. For purposes
of discussion the long imprints are labeled A, B, C, D; the
shorter ones A’, B’, C’, D’ (see Plate 72, figure 1). As
viewed in Plate 72, figure 1, the longer imprints fade out on
the bottom but terminate abruptly on the top, and in some
cases withdrawal markings can be seen. The length of
these imprints decreases from left to right; the average
length of A is 6.75 mm, while the length of D is 14 mm.
In contrast, the shorter imprints increase in length from
bottom to top. Whereas A’ is small and may be preserved
only as a wartlike body, D’ has an average length of 7.2
mm. The average length of the complete sets is 1.6 cm.

Below and to the right of A’ in many of the groups
is a series of two to four rounded parallel markings which
give the appearance of being slightly arcuate. These have
a length of 6 mm and are not as sharply defined as the im-
prints described above.

Another specimen from the Dyer collection, evidently
from the same locality as the type, clarifies some of the
features mentioned above. As viewed in Plate 74, figure 7, all
of the imprints end abruptly on the right but fade out
gradually to the left. The most striking feature of the
topotype specimen is the parallel markings above and to
the left of each group of imprints. Whereas only three or
four of these markings were present in the type specimen,
as many as nine can be counted in this form. They gradually
decrease in length and prominence away from the track.
As in the type they are broad and not so sharply im-
pressed as the other markings. Instead of being arcuate as
in the type, they take somewhat of a cuneiform appearance,
both the lower and right-hand slopes (as viewed in PI.
74, fig. 7) being steeper than their counterparts. It is ap-

parent that the markings form an integral part of the
trail and that any hypothesis regarding the origin of the
trail must somehow account for these markings.

Through the kindness of George Walker of Cincinnati,
several additional specimens are available for study. These
were collected from the Eden beds of an intermittent tribu-
tary of Mud Lick Creek where crossed by Kentucky Route
14, The locality is 1,650 meters southeast of the intersec-
tion of U.S. Route 42-127 and Kentucky Route 14, Boone
County, Kentucky. Aside from being slightly smaller and
lacking the cuneiform markings, they are similar to the
specimens described above. They confirm the existence of
eight imprints in each group, although, as in the type,
A,A’, and B’ may be weakly impressed, and in a few speci-
mens they are lacking entirely (Pl. 73, fig. 7). One Mud
Lick specimen (PI. 74, fig. 6) plainly shows that the longer
imprints are bifid, a feature which is not present on the
forms from the State Avenue locality.

Examination of all the specimens discussed to this
point indicates that they are preserved as cleavage reliefs.
None of the specimens present the sharpness of detail that
is characteristic of boundary relief. As shown in Text-figure
19 cleavage relief preservation can mask details and lead
to an erroneous interpretation of the track or trail. To some
degree this is true here, for one specimen (PI. 74, fig. 1),
associated with Trachomatichnus numerosum and preserved
as a boundary relief, illustrates that the longer imprints
(A,B,C,D) are not so simple as they appear. The trail is
incomplete and contains only 13 imprints which are ar-
ranged in four groups. One of the imprints shows a peculiar
“feather-stitch” (zig-zag) pattern in which long spinelike
projections on one side alternate with shorter projections
on the other. The distance between each of the longer pro-
jections is 2 mm. Another form (Pl. 81, fig. 7) in a dif-
ferent group lacks the “feather-stitch” pattern and instead
possesses six long projections which are visible on only
one side of the imprint. The other imprints are variations
of the patterns described above. It is not surprising that
the detailed morphology should vary; a slight rotation of
the leg can greatly affect the over-all appearance of the
resulting track or imprint. Miller also noticed the spur-
like projections, remarking “some of the toe-tracks are more
or less fringed, which I attribute to the action of the
water, though Mr. Dyer is impressed with the idea that it
may indicate hairy or spinous feet” (Miller, 1880, p. 219).

Interpretation. — Allocotichnus dyeri is unlike any
other known arthropod track, yet the fact that some 15-20
specimens are known from two localities 30 miles apart
indicates that it is a recurrent phenomenon, not an acci-

360 PALAEONTOGRAPHICA AMERICANA (VI, 41)

dental expression of some other common form. The most
fundamental question is whether the specimens under con-
sideration represent the entire track or only one side of
it. Miller (1880) believed that each group of four long im-
prints represented the essentially undisturbed track of a
quadridigitate foot, while the shorter en-echelon imprints
were the indistinct impressions of the opposite foot. How-
ever, the track would appear to be much too narrow to sup-
port such a conclusion. If the four imprints represent the
marking of a quadridigitate foot, the organism must have
been extremely large, and the trail should have a width of
15-20 cm. Moreover, none of the many specimens found
from Mud Lick Creek show two sets of quadridigitate im-
prints. It seems more likely that Adlocotichnus is a strongly
dimorphic track where A,B,C,D represent raking imprints
and A’,B’,C’,D’, support imprints. This impression is con-
firmed by a specimen from Mud Lick Creek which shows
a lesser degree of dimorphism. As viewed in Plate 72, fig-
ure 2, the imprints on the right would seem to be homologous
with B,C,D, while those on the left represent B’,C’,D’. Note
that if all the imprints are rotated clockwise approximately
45°, the resulting configuration resembles the other Mud
Lick specimens of Allocotichnus. The direction of the “V”
formed by the two series of imprints, as well as the with-
drawal markings at the proximal end of each imprint, indi-
cates that the direction of movement was from bottom to
top (as viewed in Text-fig. 21-C). It follows that move-
ment in both the type (PI. 72, fig. 1) and topotype speci-
men (Pl. 74, fig. 7) was from left to right, and the body
axis was angled to the right of the direction of movement
(see Text-fig. 21-B). If the above reasoning is correct, then
each complete set consists of a maximum of four pairs of
imprints. In many sets three and sometimes only two pairs
of imprints are present.

While the producer of the trail is unknown, the small
number of imprints in each series leads one to search for
an arthropod with a relatively small number of walking
legs. The only known possibilities in the Cincinnatian rocks
are Cryptolithus, Megalograptus, and Neostrabops. Of
course, there is always the additional possibility that Adloco-
tichnus represents the trail of some yet undescribed arthro-
pod form.

The appendages of Megalograptus have already been
discussed, and it is difficult to homologize the morphology
of the walking legs with that of the tracks. Moreover, the
little that is known of eurypterid tracks in general (Sharpe,
1912) gives reason to believe that Allocotichnus dyeri is
not such a trail, It is equally as difficult to make a direct
comparison between aglaspids and Allocotichnus dyeri.
While there is no evidence to indicate that it is not an

aglaspid trail, there is nothing to indicate that it is. The
stratigraphic position of the specimens (Eden) and the
small number of appendages involved in producing the
track make Cryptolithus an attractive choice. Nevertheless,
measurements show that Cryptolithus is too small to have
made the trail. Because the trial is strongly dimorphic, the
width varies. The least width, measured from the distal
end of A to the distal end of A’, ranges from 1.5-2.5 cm, while
the greatest width, measured from the distal end of D to
the distal end of D’, is 4-5 cm. Raymond’s (1920, pl. 7,
fig. 4) illustration of a well-preserved Cryptolithus shows
that the appendages even when spread-eagled, which is
unlikely, cover only three-quarters the width of the cepha-
lon. Thirty-five specimens of Cryptolithus were measured;
it was found that the mean “three-quarters width” was 1.59
cm, while the largest form observed had a width of only
2.3 cm. These figures indicate that Cryptolithus must be
eliminated as a possible originator of the trail.

The only remaining arthropods capable of leaving such
a wide trail are the multisegmented trilobites, such as
Flexicalymene and Isotelus. However, it is doubtful that
even the larger flexicalymenids would be able to produce
the tracks, for they rarely attain a length over 6.0 cm or
a width in access of 3.0-3.5 cm. This leaves /sotelus as
the most likely producer. Nevertheless, it should be repeat-
ed that Allocotichnus dyeri is unlike any known trilobite
track, and the choice of /sotelus as the producer is by no
means certain. Asaphoidichnus trifidum, which is also at-
tributed to /sotelus, bears little resemblance to Allocotich-
nus.

Judging from the Oxford specimen of the ventral side
of Isotelus (Pl. 57, fig. 5), the first 8-10 pairs of walking
legs were probably strong, and it is possible that the first
three or four pairs of legs could propel the animal fast
enough to keep the pygidium off the substrate. While the
smooth platter-like streamlined body of /sotelus would not
produce as great a drag as the bodies of other trilobites,
it is believed that the cuneiform markings of the State
Avenue specimens (PI. 72, fig. 1; Pl. 74, fig. 7) show where
the pleurae occasionally nicked the bottom. In addition the
imprints seen on Plate 74, figure 1 indicate that some-
times nearly the entire leg was placed on the substrate, for
the spurlike bodies associated with the markings most
likely represent spines on the distal ends of each leg seg-
ment. If three or four joints of the leg made contact with
the substrate, it follows that the eight blunt pleurae of
the right side of the body were extremely close to the
bottom and could have gouged the sediment in the rhythm
of movement. It is unlikely, however, that the leg was
always placed flat on the substrate; the bifid markings of

TRACE FOSSILS CINCINNATI AREA: OsGoop

44s"

Text-figure 21.— A. Asaphoidichnus trifidum Miller, 1880. Re-
pichnia of Jsotelus; interpretation of the trail illustrated on Plate
75, figure 3. UCM 37569, X 0.75. B,C. Allocotichnus dyeri (Miller),
1880. 2. Repichnia of Jsotelus; interpretation of the trail illustrated
on Plate 72, figure 1. Holotype. MCZ 543; latex mold UCM 37693,
X 1, 3. Repichnia of Jsotclus; interpretation of the trail illustrated
on Plate 72, figure 2. The dimorphism of the series is not so marked
as it is in figure 2 above. UCM 37627, X 1.

361

360

dental expression of some other common form. The most
fundamental question is whether the specimens under con-
sideration represent the entire tracks or only one side of
it. Miller (1880) believed that each §roup of four long im-
prints represented the essentially undfisturbed track of a
quadridigitate foot, whfle-the shorter pn-échelon imprints
were the indistinct.4mpptssions of the ‘opposite foot. How-
ever, the track would appear to be much too ntarrow to sup-
port such a conclusion. If pie eur imprints represent the
marking of a quadridigit¥te {85t, *the organism nfust have
been extremely large, and the teafl should have at ‘width of
15-20 cm. Moreover, none of the“many specinfens found
from Mud Lick Creek show two sets of quadridigitate3im-
prints. It seems more likely that Allogetichpnags i is a strongly;

dimorphic track where A,B,C,D represent Yaking imprints 2

and A’,B’,C’,D’, support imprints. This impression is con
firmed by a specimen from Mud Lick Creek’ whigh shows
a lesser degree of dimorphism. As viewed in ,Platé 72, fig-
ure 2, the imprints on the right would seem to be ome eons
with B,C,D, while those on the left represent B’,C’,}¥* Note
that if all the imprints are rotated clockwise app Xingétely
45°, the resulting configuration resembles the othe Mud

Lick specimens of Allocotichnus. The direction of the “V” ,-

formed by the two series of imprints, as well as the with-
drawal markings at the proximal end of each imprint, indi-
cates that the direction of movement was from bottom to
top (as viewed in Text-fig, 21-C),- It follows that move-
ment in both the type Pl, 72, fig. 1) and topotype speci-
men (PI. 74, fig. 7) dva’s from lefg to right, and the body
axis was angled to the vight? ‘of the direction of movement
(see Text-fig. 21-B). Thay above reasopfing is correct, then
each complete set consist§ off a maximum of four pairs of
imprints. In many sets three¥and; ;Sometimes only two pairs
of imprints are present. ,*_ # 7

While the producer of the ails -urfknowwn, the small
number of imprints in each séfies Ighds.. atte to ran for
an arthropod with a relatively spel hunger of w4lking
legs. The only known possibilities ¥n t GRacingget
are Cryptolithus, Megalograptus, andy Neagptrabops. Of
course, there 1s always the additional possibility th locg-
tichnus represents the trail of some yet ae arpitto-
pod form. H '

The appendages of Megalograptus have tread been
discussed, and it is difficult to homologize the mobphology
of the as legs with that of the tracks. Moreover, the
little that is known of eurypterid tracks in general (Sharpe,
1912) gives reason to believe that Allocotichnus dyeri is
not such a trail, It is equally as difficult to make a direct
comparison between aglaspids and Allocotichnus dyeri.

‘While there is no evidence to indicate that it is not an

an rocks

-

PALAEONTOGRAPHICA AMERICANA (VI, 41)

aglaspid gaily there is nothing to indicate that it is. The
stratigraphi& pgsitfon of the specimens (Eden) and the
small number of, appendages involved in producing the
track make Cry$6lithus an attractive choice. Nevertheless,
measurements’sHow that Cryptolithus is too small to have
made the trail. Because the trial is strongly dimorphic, the
width varies. Fhe’ ‘least width, measured from the distal
end of A to the distal end of A’, ranges from 1.5-2.5 cm, while
the greatest width, measured fyom the distal end of D to
the distal end of D, is, 4-5 cm. Raymond’s (1920, pl. 7,
fig. 4) illustration oF aj ‘ well- preserved Cryptolithus shows
that the appendagts even whew spread-eagled, which is
unlikely, cover only thrge-quarters the width of the cepha-
lon. Thirty-five specypens 4 gf Cry ptolithus were measured;
it was found tha» whe mea F threg?quarters width” was 1.59
cm, while the buese forth *obseryed had a width of only
2.3 gm. These figures indicate that Cryptolithus must be
eliminated as a possible originator of the trail.

The only remainjmg arthropods capable of leaving such
a wide trail are the ahyttisegmented trilobites, such as
Flexicalymene and Isbtelus. Hoyvever, it is doubtful that
even the larger flexicalymenids yould be able to produce

ad the tracks, for they rarely attain a length over 6.0 cm or

4

ee

.’a width in access of 3.0-3.5 cfn. This leaves Jsotelus as

the most likely produter. Nevertheless, it should be repeat-
ed that Allocotigtinus* dye i is unlike any known trilobite
track, and the choite of Frotelus as the producer is by no
means certain. Asaphoidichnus tmfidum, which is also at-
tributed to Jsotelus, bears little’tesemblance to Allocotich-
nus.

Judging from the Oxford a of the ventral side
of Isotelus (Pl: 37 He), thefirst 8-10 pairs of walking
legs were probabky sttong, apd it is possible that the first
three or four pairs of legs* cquld propel the animal fast
enough to keep the pygidiurs* off the substrate. While the
smooth platter-like streamligéd body of Jsotelus would not
produce as grea tat sph the bodies of other trilobites,
it is eel a #cuneiform markings of the State
Avenue specimehs (Pl 72, fig. 1; Pl. 74, fig. 7) show where
the pleurae occasionally niekodiehie bottom. In addition the
imprints seen on Plate 74, figure 1 indicate that some-
times nearly the entire leg was placed on the substrate, for
the spurlike bodies associated with the markings most
likely represeng-spines on the djstal ends of each leg seg-
ment. If three or/four joints of the leg made contact with
the substrate, it follows that the eight blunt pleurae of
the right side of the body were extremely close to the
bottom and could have gouged the sediment in the rhythm
of movement. Jt is unlikely, however, that the leg was
always placed flat on the substrate; the bifid markings of

TRACE FOSSILS CINCINNATI AREA: OsGoop

\

NS

x

Text-figure 21.— A. Asaphoidichnus trifidum Miller, 1880, Re-
pichnia of Jsotelus; interpretation of the trail illustrated on Plate
75, figure 3. UCM 37569, X 0.75. B,C. Allocotichnus dyeri (Miller),
1880. 2. Repichnia of Jsofelus; interpretation of the trail illustrated
on Plate 72, figure 1. Holotype. MCZ 543; latex mold UCM 37693,
X 1. 3. Repichnia of /sotelus; interpretation of the trail illustrated
on Plate 72, figure 2. The dimorphism of the series is not so marked
as it is in figure 2 above. UCM 37627, X 1.

362 PALAEONTOGRAPHICA AMERICANA (VI, 41)

some specimens (Pl. 74, fig. 6) show that sometimes only
the dactyls made contact with the bottom.

In summary then, it is believed that Allocotichnus
dyeri represents the strongly dimorphic trail of Jsotelus in
which only the stronger anterior appendages were employed.
In some instances nearly the entire length of the leg was
placed on the substrate and the thoracic pleurae gouged
the bottom. On the other hand, the bifid markings of
some specimens (Pl. 74, fig. 7) illustrate that in a few
cases only the dactyls made contact with the substrate.
The reason behind such movement is, problematic.

PETALICHNUS Miller, 1880

Plate 73, figures 5,6; Plate 74, figures 2,4,5; Plate 75, figure 4;
Text-figures 22,29i,j

1880. Petalichnus Miller, Cincinnati Soc. Nat. Hist.; Jour., vol. 2,
pp. 219-221, pl. 13, fig. 5; pl. 14, fig. 3:

1880. Partim Trachomatichnus Miller, Cincinnati Soc. Nat. Hist.
Jour., vol. 2, pp. 219-221, pl. 14, fig. 2.

1938. Merostomichnites Packard, Caster, Jour. Paleont., vol. 12, p. 34,
pl. 10, fig. 1. ;
1962. Petalichnus Miller, Héantzschel, ‘Trace-Fossils and Problematica

in Treatise on Invertebrate Paleont., Moore (ed.), pt. W,
Miscellanea, p. W208.

1962. Partim Trachomatichnus Miller, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W219.

Type species — Petalichnus. multipartitum Miller
(1880) (Pl. 75, fig. 4; MCZ- 546; latex mold UCM 37697)
from the ?Southgate beds of “Walker Mill Road,” (State
Avenue) Cincinnati, Ohio. Monotypic.

Diagnosis. — Trilobite tracks showing straight-ahead
or slightly oblique movement. Each series consists of 9-12

unifid or bifid imprints. Generic désignation of producer.

unknown. 8

Discussion. — As explained previously, recent studies
on arthropod tracks (Caster, 1938; Seilacher, 1955), haye
shown that a slight rotation of the body during walking
can result in tracks which differ greatly from those produced
by straight-ahead movement. Such slight modifications
must be taken into consideration when making taxonomic
assignments, otherwise the result will be a different name
for every track. It is believed that the best taxobases for
arthropod trails are: 1) a consistent pattern of imprints
which is found in several specimens and sets that trail
apart from others, e.g., the unique dimorphism of Alloco-
tichnus dyeri, and 2) a known producer — can the origina-
tor of the track be determined with a reasonable degree
of confidence? For example, the width of Asaphoidichnus
trifidwm shows that it is more likely the trail of an isotelid;
therefore, it merits separate taxonomic treatment. Regret-
ably Petalichnus does not come under either heading. The

genus contains trails showing a wide range of morphology,
and these can not be assigned to a given trilobite. Included
in Petalichnus are forms which in Miller’s work constituted
two genera and three species. Justification for placing
them in synonymy with Petalichnus will be given as each
is discussed below. Also included in the genus is a form
which Caster (1938, pl. 10, fig. 1) referred to as “Mero-
stomichmites sp. (cf. Petalichnus multipartitus).” It is ac-
knowledged. that Petalichnus as defined herein is a form
genus and more than likely contains trails made by more
than one: genus of trilobite and conceivably may even in-
clude tracks ‘of other classes of arthropods.

Petalichnus multipartitum Miller

Plate 73, figures 5,6; Plate 74, figures 2,4,5; Plate 75, figure 4;
Text-figures 22,29i,j

1880. Petalichnus multipartitus Miller, Cincinnati Soc. Nat. Hist.,
Jour., vol. 2, p. 222, pl. 14, fig. 2.

1880. Trachomatichnus permultus Miller, Cincinnati Soc. Nat. Hist.,
Jour., vol. 2, p. 220, pl. 13, fig. 5.

1880. Trachomatichnus cincinnatiensis Miller, Cincinnati Soc. Nat.
Hist., Jour., vol. 2, pp. 220,221, pl. 14, fig. 3.

1938. Merostomichnites sp. Caster, Jour. Paleont. vol. 12, p. 34,
ple 10) figs de

Discussion. — The type specimen (Pl. 75, fig 4) is a
highly complex trail:.It° is preserved as a convex hypo-
relief and has a length of 11.3 cm and a width of 2.2 cm.
The clearest portion ‘of the trail is on the right (as viewed
in PI. 75, fig. 4). Here it is possible to make out five series
of unified raking imprints. with plainly defined withdrawal

“markings at their proximal ends. The number of imprints

in each complete series varies from 10-12. The shorter
cuneiform imprints in the central part of the trail are

judged to be support imprints. These stand out in bold

relief when compared to the more shallow raking imprints.

‘In the upper part.of the trail the imprints become so super-
- imposed that it is. not possible to make a detailed analysis.

Therefore, only a partial reconstruction of the trail is given
in Text-fig. 22-C. The position of the raking imprints rela-
tive to that of the support imprints shows that, as viewed
in Plate 75, figure 4 and Text-figure 22-C, movement was
from bottom to top, and the body was angled slightly to the
right of the axis of movement.

“Trachomatichnus permultum” (Pl. 73, fig. 5; MCZ
548; latex mold UCM 37696): In contrast to the type speci-
ment of P. multipartitum, the form designated by Miller
(1880), as “Trachomatichnus permultwm” is a picture of
simplicity. Miller (1880, p. 220) separated “7. permultum”
from 7. numeroswm, because the imprints of the former
“are not so numerous, they are destitute of the beaded

‘TRACE FOSSILS CINCINNATI AREA: OsGoop

Text-figure 22.— A-C. Petalichnus multipartitum Miller, 1880. 1.
Repichnia of a medium-sized trilobite; interpretation of the trail illus-
trated on Plate 84, figure 5. MU T86; latex mold UCM 47707, X 1.7.
2. Repichnia of a medium-sized trilobite; interpretaton of the trail
illustrated on Plate 73, figure 5. MCZ 548; latex mold UCM 37696,
X 1.4. 3. Repichnia of a medium-sized trilobite; interpretation of the
trail illustrated on Plate 75, figure +. The trail is so complex that it is
not possible to place all imprints in their proper series. Holotype.
MCZ 546; latex mold UCM 37697, BG

363

PALAEONTOGRAPHICA

some specimens (Pl. 74, fig. 6) show that sometimes only
the dactyls made contact with theybottom.

In summary then, it is believed that
dyeri represents the strongly dimorphic trail of Zsotelus in
which only the stronger anterior appendagts were employed.
In some instances nearly thg#entire lengthgof the leg was
placed on the substrate and “the thoracye pleurae gouged
the bottom. On the other hafid, the ‘bifid* markings of
some specimens (Pl. 74, fig. 2) dMlustrate thgt in a few
cases only the nae made conrattt witk’ the ,substrate.
The reason behind such movement_ is puoblematt

Rie -

Allocotichnus

a

PETALICHNUS Miter, +380 oT
Plate B neue 4;

eaee”

Plate 73, figures 5,6; Plate 74, nae 545:

Res eae) 2 pial

oe,
1880. Petalichnus Miller, Cincinnati *&o¢. Nat. Hises* Jour., vol. 2,
pp. 219-221, pl. 13, fig. 5; pl. 14, fig. “Sy ~-
Partim Trachomatichnus Millowss:Cincingati TW. Nat. Hist.
Jour., vol. 2, pp. 219-221, Phy 14. figs 2) - 4X
1938. Merostomichnites Packard, Caste four. Pilconte’ vol. 12, p. 34,
pl. 10, fig. 1.

1880.

1962. Petalichnus Miller, Hanteatbel, MP te Fogle. a PFoblematica
in Treatise on InvertebratgggPaleont., Maayan ee), pt W,
Miscellanea, p. W208. ose

1962. Partim Trachomatichnus Méley ena eTeace- -Fossils and
Problematica in Treatise on Invertebrate Pa ont, Moore (ed.)

t. W, Miscellanea, p. Wl x“ iene
p sce p > Soo00 se, ‘2 g
Type species — Petalit owe gull eum Miller

(1880) (PI. 75, fig. 4; MCZ7*546, Jatex” mold UCM 37697 )
from the ‘Southpate beds of*¥#Walker Mit? Road, és eco
Avenue) Cincinnati, Ohia. Méraryyic: x of®

AMERICANA

II, 41)

genus cohtains trails showing a wide range of morphology,
and these cgn not be assigned to a given trilobite. Included
in Petilishnws are forms which in Miller’s work constituted
two geneva ‘and three species. Justification for placing
them im sy nofymy with Petalichnus will be given as each
is discyjseed bélow. Also included in the genus is a form
which CABter ( 1938, pl. 10, fig. 1) ieee to as “Mero-
stomichniteg sp. (cf. Petalichnus multipartitus).” It is ac-
knowledgedsthat Petalichnus as defined herein is a form
genus and mere than likely contains trails made by more
than one: igen of trilobite and conceivably may even in-
clude tracks Gf other classes of arthropods.

N+
te
Wégalichnus multipartitum Miller
x
Plate 73, figures \6; Plate 74, figures 2,4,5; Plate 75, figure 4;
s Text-figures 22,29i,j
.
1880. Petalichnus multipartitus Miller, Cincinnati Soc. Nat. Hist.,
Jour., vol. 2, p. 222, pl. 14, fig. 2.
1880. Trachomatichnus permultus Miller, Cincinnati Soc. Nat. Hist.,
Jour.; voli 2) p. 220;"ple 135, tip. 5:
1880. Trachomatichnus cincinnatiensis Miller, Cincinnati Soc. Nat.
Hist., Jour., vol. 2, pp. 220,221, pl. 14, fig. 3.
1938. prides sp. Caster, Jour. Paleont., vol. 12, p. 34,
! "1p, fig. 1.
‘ sha i 2

Beet
~

se Discussion. Bee -type specimen (PI. 75, fig 4) is a
whi Fy eqn mplex trailteIt*is preserved as a convex hypo-
ee ahd has a length “of'1.3 cm and a width of 2.2 cm.
2 Theeleagest portion 1of, the trail is on the right (as viewed

“un PR Bb, fig. +). Heme* fas 4s possible to make out five series

Diagnosis. — Trilobite. trek Showing | Syraight- Poe ai unitted raking imprigtsewith plainly defined withdrawal

or slightly oblique movement. »fach sgegies eonasts of 1g”

Sia “markings at their prdximag ends. The number of imprints

unifid or bifid imprints. Genggtic designations of produ e sip geach complete senies* varies from 10-12. The shorter

unknown. aa

Discussion. — As explained previously, recent studigs
on arthropod tracks (Caster, 1938; Seilacher, 1955) gha¥e
shown that a slight rotation of the body during Walkifig

can result in tracks which differ greatly from those p¥duced

sgneiform imprints *irt the central part of the trail are
dged to be suppdrt;imprints. These stand out in bold
*$elief when compgeed jo the more shallow raking imprints.

a
es In the upper part 404 the trail the imprints become so super-
* imposed that it iss not possible to make a detailed analysis.

by straight-ahead movement. Such slight modifcations 7 Therefore, only a partial reconstruction of the trail is given

must be taken into consideration when making taxonondé
assignments, otherwise the result will be a differggt name
for every track. It is believed that the best taxobases” for
arthropod trails are: 1) a consistent pattern of imprints
which is found in several specimens and sets that trail
apart from others, e.g., the unique dimorphism of Alloco-
tichnus dyert, and 2) a known producer — can the origina-
tor of the track be determined with a reasonable degree
of confidence? For example, the width of Asaphoidichnus
trifidum shows that it is more likely the trail of an isotelid;
therefore, it merits separate taxonomic treatment. Regret-
ably Petalichnus does not come under either heading. The

in Text-fig. 22-C. The position of the raking imprints rela-
tive to that of thé Support imprints shows that, as viewed
in Plate 75, figuyest and Text-figure 22-C, movement was
from bottom to top, and the body was angled slightly to the
right of the axis of movement.

“Trachomatichnus permultum” (Pl. 73, fig. 5; MCZ
548; latex mold UCM 37696): In contrast to the type speci-
ment of P. multipartitum, the form designated by Miller
(1880), as “Trachomatichnus permultwm” is a picture of
(1880, p..220) separated “T. permultwm”

NUMETOSUMN,

simplicity. Miller
from 7’. because the imprints of the former
“are not so numerous, they are destitute of the beaded

TRACE FOSSILS CINCINNATI AREA: OsGoop

Text-figure 22.— A-C. Petalichnus multipartitum Miller, 1880. 1.
Repichnia of a medium-sized trilobite ; interpretation of the trail illus-
trated on Plate 84, figure 5. MU 186; latex mold UCM 47707, DG eI
2. Repichnia of a medium-sized trilobite; interpretaton of the trail
illustrated on Plate 73, figure 5. MCZ 548; latex mold UCM 37696,
X 1.4. 3. Repichnia of a medium-sized trilobite ; interpretation of the
trail illustrated on Plate 75, figure 4+. The trail is so complex that it is
not possible to place all imprints in their proper series. Holotype.
MCZ 546; latex mold UCM 37697, X 1.5.

364 PALAEONTOGRAPHICA AMERICANA (VI, 41)

S| ie Ta

Text-figure 23.— A. Teratichnus confertum Miller, 1880. Re-
pichnia of a medium-sized arthropod; interpretation of the trail il-
lustrated on Plate 75, figure 3. The trail is highly complex and only
a partial interpretation is given. Holotype. MCZ 540; latex mold
UCM 37570, X 1.4.

‘TRACE FOSSILS CINCINNATI AREA: OsGoov 365

appearance, they are subtriangular instead of somewhat
uniformly rounded and the rews-aare proportionately
farther apart.” In this case the last-two. criteria, are in-
significant as taxobases, but the first two, i.¢,, the aeer

of imprints and the beaded appearance, are judged*eayke

ment would be from bottom to top, as seen in Plate 74,
figure 2. The width of the trail is similar to that of the
type specimen, and there are no distinct features that would
serve to set it apart as a separate taxon.

_.Merostomichnites sp. (cf. Petalichnus multipartitus)”

important. As will be shown, there is a good possibility wage (PI 74; dig. 4; UCM 22033); Caster (1938, pl. 10, fig. 1)
that 7. nwmerosum is the trail of Cryptolithus, “T. per™*¥guybysgrated a tP4e from the Fulton beds of the Union Levee,

multum” has a length of 8.5 cm and consists of five sep-
arate sets. The three complete sets contain 10, 10, and 11
imprints. The trail width measured in the lower part of
the trail, as viewed in Plate 73, figure 5, is 1 cm. The
specimen is noteworthy because it shows a transition from
straight-ahead movement. Midway through the trail the
imprints on the right side become longer and slightly sig-
moidal in outline, while in the upper part of the trail they
are strongly sigmoidal, indicating that the animal was
oriented to the right of the axis of movement. Note that
the imprints on the left side show little change and that it
is difficult to distinguish between the various series. As il-
lustrated by the sigmoidal outline and the orientation of the
“V” formed by the opposing series, movement was from
bottom to top as seen in Text-figure 22-B.

Outwardly this specimen differs greatly from the type
specimen of P. multipartitwm, yet basically they are the
same. The track widths are similar as are the number of
imprints in a series. There is no consistent morphological
difference on which the two could be separated.

“Trachomatichnus cincinnatiensis” (P|. 74, fig. 2; MCZ
542; latex mold UCM 37571): Miller (1880, pp. 220, 221)
separated “J. cincinnatiensis” from “T. permultum” “.
by the smaller and more ill-defined form of the tracks, by

the greater number in each series, and by the greater dis-%

He differentiated it from
. by the greater distance between the two
series, by the [fewer] number of impressions in each series,
and by the simple character of the tracks instead of the
compound or beaded form” (Miller, 1880, p. 221).

tance between the two series.”
T. numerosum “

The trail, which has a total length of 16 cm and, #’,

width of 1.8 cm, is preserved as a convex hyporelief ad
is quite indistinct. This obscurity suggests that the trail
actually is preserved as a cleavage relief and not as a
boundary relief (compare with Pl. 74, fig. 1). If this is
the case, the more complete set of tracks lies beneath the
surface, There is no clearly defined sets, and the individual
imprints are small and show no evidence of raking motion.
The linear arrangement of the opposing rows suggests
straight-ahead movement; however, the criteria normally
used for determining the direction of movement are lack-
ing. The only hint of a “V” is seen in the middle of the

trail; if this actually represents one-half of a “V,” move-

Cin Innatt “Obie. which he compared to Merostomichnites
as Retted by Stdtitex: (£934). The trail is preserved as a
convex R® Srelief meet calgsiltite and has a total length of
5.7 cm and’ width, of 18 cm> The specimen, which is
incomplete, consists at ‘Teporsion of six obliquely directed
series of raking imprints Om, the left and a continuous row of
support imprints on the nights» T greatest number of im-
pressions seen in any serigs Is 10. trail is unusual in
that nearly all of the impithterar emo te scallop-like mark-
ings. However, a short, Taney m4 ridge can be seen
leading from some of theSmpry ee. left. These ridges
are oriented normal to the cent the row of support 1m-
prints. *e
Although the trail beags Bg reseihblance to Stermer’s
(1934, pl. 2, fig. 7) illustraggon Vf Werwtomichnites strandi
from the Downtonian Mivogiante weds of Spitzbergen,
there are significant differences. Ase “Stormer (1934, p. 22)
noted, each of the impritS ay thé Norwegian specimen con-
sistently contains .three or ipurymiikings grouped together.
Thi demonstyates” wig * “the *frail was made by an organism
witManesridigi tye Walking legs. In contrast, in the right-
5 sta &{ the Cincinnatian form, neither the spacing

gen, the, _ imprints nor the number of imprints is as

%, consistent aS,it is In Stgrmer’s specimen, It is instead con-

aided’ that the Cincinnatian specimen is a trilobite trail
., Where “thé body was oriented to the left of the axis of
‘Smévement; thus the superimposition of the support im-
brints in the right-hand row give a false impression of a
polydactylous leg. Stérmer believed that M. strandi was
probably the trail of an eurypterid, and his conclusion is
not questioned herein. The present author agrees with
Caster (1938) that the Cincinnatian specimen is a trilo-
bite trail. The scallop shape of the imprints indicates that
the legs probably struck the substrate in a slicing motion.
The weak preservation of the raking imprints demonstrates
that the specimen is preserved as a cleavage relief.

Two other previously undiscussed specimens which are
assigned to P. multipartitum are shown on Plate 73, figure
6 and Plate 74, figure 5. One of these (Pl. 74, fig. 5) is an
excellent example of oblique movement. While the imprints
of the left side are superimposed, the obliquely directed
imprints on the right are clearly defined, and it is possible
to count 10-12 imprints in the various series (see Text-fig.

364

PALAEONTOGRAPHICA AMERICANA (VI, 41)

7
| a ey
oe am 1

Text-figure 23.— A. Teratichnus confertum Miller, 1880. Re-
pichnia of a medium-sized arthropod; interpretation of the trail il-
lustrated on Plate 75, figure 3. The trail is highly complex and only
a partial interpretation is given. Holotype. MCZ 540; latex mold
UCM 37570, X 1.4.

— =

‘TRACE FOSSILS CINCINNATL AREA: Oscoop 365

appearance, they are subtriangular instead of somewhat
uniformly rounded and the rows are proportionately
farther apart.” In this case the last two criteria are in-
significant as taxobases, but the first two, 7.e., the number
of imprints and the beaded appearance, are judged to be
important. As will be shown, there is a good possibility
that 7. numerosum is the trail of Cryptolithus, “T. per-
multum” has a length of 8.5 cm and consists of five sep-
arate sets. The three complete sets contain 10, 10, and 11
imprints, The trail width measured in the lower part of
the trail, as viewed in Plate 73, figure 5, is 1 cm. The
specimen is noteworthy because it shows a transition from
straight-ahead movement. Midway through the trail the
imprints on the right side become longer and slightly sig-
moidal in outline, while in the upper part of the trail they
are strongly sigmoidal, indicating that the animal was
oriented to the right of the axis of movement. Note that
the imprints on the left side show little change and that it
is difficult to distinguish between the various series. As il-
lustrated by the sigmoidal outline and the orientation of the
“V” formed by the opposing series, movement was from
bottom to top as seen in Text-figure 22-B.

Outwardly this specimen differs greatly from the type
specimen of P. multipartitwm, yet basically they are the
same. The track widths are similar as are the number of
imprints in a series. There is no consistent morphological
difference on which the two could be separated.

“Trachomatichnus cincinnatiensis” (Pl. 74, fig. 2; MCZ
542; latex mold UCM 37571): Miller (1880, pp. 220, 221)
separated “7. cincinnatiensis” from “T. permultum” “.
by the smaller and more ill-defined form of the tracks, by
the greater number in each series, and by the greater dis-
tance between the two series.” He differentiated it from
T. numerosum “... by the greater distance between the two
series, by the [fewer] number of impressions in each series,
and by the simple character of the tracks instead of the
compound or beaded form” (Miller, 1880, p. 221).

The trail, which has a total length of 16 cm and a
width of 1.8 cm, is preserved as a convex hyporelief and
is quite indistinct. This obscurity suggests that the trail
actually is preserved as a cleavage relief and not as a
boundary relief (compare with Pl. 74, fig. 1). If this is
the case, the more complete set of tracks lies beneath the
surface. There is no clearly defined sets, and the individual
imprints are small and show no evidence of raking motion.
The linear arrangement of the opposing rows suggests
straight-ahead movement; however, the criteria normally
used for determining the direction of movement are lack-
ing. The only hint of a “V” is seen in the middle of the
trail; if this actually represents one-half of a “V,” move-

ment would be from bottom to top, as seen in Plate 74,
figure 2. The width of the trail is similar to that of the
type specimen, and there are no distinct features that would
serve to set it apart as a separate taxon.

“Merostomichnites sp. (cf. Petalichnus multipartitus )”
(Pl. 74, fig. 4; UCM 22033); Caster (1938, pl. 10, fig. 1)
illustrated a trail from the Fulton beds of the Union Levee,
Cincinnati, Ohio, which he compared to Merostomichnites
as figured by Stérmer (1934). The trail is preserved as a
convex hyporelief in a calcisiltite and has a total length of
5.7 cm and a width of 1.8 cm. The specimen, which is
incomplete, consists of a portion of six obliquely directed
series of raking imprints on the left and a continuous row of
support imprints on the right. The greatest number of im-
pressions seen in any series is 10. The trail is unusual in
that nearly all of the imprints are simple scallop-like mark-
ings. However, a short, faintly defined ridge can be seen
leading from some of the imprints on the left. These ridges
are oriented normal to the trend of the row of support im-
prints.

Although the trail bears some resemblance to Stérmer’s
(1934, pl. 2, fig. 7) illustration of Merostomichnites strandi
from the Downtonian (Devonian) beds of Spitzbergen,
there are significant differences. As Stérmer (1934, p. 22)
noted, each of the imprints in the Norwegian specimen con-
sistently contains three or four markings grouped together.
This demonstrates that the trail was made by an organism
with quadridigitate walking legs. In contrast, in the right-
hand row of the Cincinnatian form, neither the spacing
between the imprints nor the number of imprints is as
consistent as it is in Stérmer’s specimen, It is instead con-
cluded that the Cincinnatian specimen is a trilobite trail
where the body was oriented to the left of the axis of
movement; thus the superimposition of the support im-
prints in the right-hand row give a false impression of a
polydactylous leg. Stérmer believed that M. strandi was
probably the trail of an eurypterid, and his conclusion is
not questioned herein. The present author agrees with
Caster (1938) that the Cincinnatian specimen is a trilo-
bite trail. The scallop shape of the imprints indicates that
the legs probably struck the substrate in a slicing motion.
The weak preservation of the raking imprints demonstrates
that the specimen is preserved as a cleavage relief.

Two other previously undiscussed specimens which are
assigned to P. multipartitum are shown on Plate 73, figure
6 and Plate 74, figure 5. One of these (Pl. 74, fig. 5) is an
excellent example of oblique movement. While the imprints
of the left side are superimposed, the obliquely directed
imprints on the right are clearly defined, and it is possible
to count 10-12 imprints in the various series (see Text-fig.

366 PALAEONTOGRAPHICA AMERICANA (VI, 41)

22-A). The width of the trail is 9 mm, while the length of
each series averages 2.5 cm. The specimen differs from most
forms showing oblique movement as there are no well-de-
veloped raking imprints; however, this may be more a fac-
tor of preservation than of behavior.

The other specimen (PI. 73, fig. 6) was illustrated by
Caster, Dalvé, and Pope (1955, pl. 2, fig. 35). The trail
has a width of 2.6 cm and the length of each series is 3.4
em. The individual imprints, which average 8 series, are
bifid distally and show well-developed withdrawal markings
at their proximal extremity. As viewed in Plate 73, figure
6, movement was from bottom to top.

Interpretation. — As indicated above, the forms grouped
under P. multipartitum, although interpreted by Miller
(1880) as cephalopod tracks, are herein thought to be the
trails of moderate-sized trilobites. Unfortunately it is not
possible to assign them to a given genus. Based on the per-
centage of trilobites found in the Cincinnatian rocks, most
of the trails should be those of Flexicalymene; however,
Petalichnus almost certainly contains tracks of other trilo-
bites. Until a trilobite is found at the end of a trail or
body fossils are found with the dactyls well preserved as
in Cryptolithus, P. multipartitum must remain a form genus
for unifid or bifid tracks of moderate-sized trilobites.

Trachomatichnus Miller, 1880
Plate 73, figure 3,4; Plate 74, figure 1; Text-figure 20-B, 29-g

1880. Partim Trachomatichnus Miller, Cincinnati Soc. Nat. Hist.,
Jour., vol. 2, pp. 219,220, pl. 13, figs. 2-4.

1962. Partim Trachomatichnus Miller, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W219.

Type species. —Trachomatichnus numerosum Miller
(1880), from the ?Southgate beds of State Avenue, Cin-
cinnati, Ohio. Monotypic. The specimen (MCZ 539; latex
mold UCM 37583) illustrated by Miller (1880, pl. 13, fig.
3; herein Pl. 74, fig. 1) is designated as the type specimen.

Diagnosis. — Straight-ahead trails consisting of crowd-
ed, poorly defined imprints made by a polydactylous leg.
As best can be determined, the number of imprints per set
ranges from 9-11.

Discussion. — As noted elsewhere, Miller (1880) estab-
lished three species of T'rachomatichnus. Because no holo-
type was named by the author, Hantzschel (1962) designat-
ed T. nwmerosum as the type species. This was a fortunate
choice for, as shown previously, the specimens referred to
“T. permultum” and “T. cincinnatiensis” lack sufficient
diagnostic features to justify their retention as separate
taxa. On the other hand, the three specimens of 7. nwmero-

sum illustrated by Miller differ from all others discussed
by the author in his 1880 paper. According to Miller (1880,
pp. 219, 220), the species is characterized by tracks which
are “. . . composed of four or five parts, which present a
beaded appearance.”

The specimen herein designated as the type specimen
(Pl. 74, fig. 1) occurs as a convex hyporelief on a cross-
laminated calcisiltite. In contrast to most Cincinnatian
arthropod trails which are found on cleavage planes, the
clarity of detail indicates that this form is preserved as a
boundary relief. The length of the trail is 10.5 cm and the
width ranges from 1.3-1.5 em. While there is no dimorphism
in the two rows, the morphology of the specimen varies
along its length.** Three different expressions are present,
each occupying roughly one-third of the total trail length.

The imprints of the posterior third of the trail re-
semble cuneiform wedges which taper and fade out anter-
iorly. The posterior portion of the wedge has a denticulate
margin, thus differentiating it from all other imprints so
far discussed. The better preserved forms show three to
six denticles. The various imprints are so irregularly placed
that it is impossible to determine the number of imprints
in a set. In the center portion of the trail, the imprints
flow together to form two parallel continuous fasciculate
ridges, although some individual imprints can be seen to
the interior of each ridge. In the anterior third of the trail,
it is possible to discern separate sets, albeit the number of
imprints comprising a set is unknown. The open end of the
“V” formed by the opposing arms is directed anteriorly.
Note that at one point the rows merge briefly to form a
short bilobed Rusophycus-like body.

As indicated by the orientation of the “V’s,” movement
was from left to right, as viewed in Plate 74, figure 1. The
variations in the morphology of the trail reflect different
types of movement, which were probably caused at least
in part by the uneven surface of the bottom upon which
the organism was walking. The surface of most of the slab
slopes gently to the left, while in the anterior third of the
trail, the slope is more precipitous. The cuneiform nature
of the imprints indicates that the dactyls were not placed
flat on the substrate as in the type specimen of Asaphoidich-
nus trifidwm nor were they moved toward the median line
to form raking imprints. Instead the animal inserted the
dactyls obliquely to the rear, as if it were slipping and
struggled to gain more traction. The fasciculate ridges in

“For purposes of discussion the left side of the specimen, as viewed
in Plate 74, figure 1, is referred to as posterior; the right side as
anterior.

TRACE FOSSILS CINCINNATI AREA: Oscoop 367

the center portion of the trail would seem to confirm this.
The lack of dimorphism indicates that the body was parallel
to the axis of movement.

Another specimen (MCZ 547; latex mold UCM 37695)
of T. numerosum figured by Miller (1880, pl. 13, fig. 2) is
illustrated herein on Plate 73, figure 3. The trail is pre-
served as a convex hyporelief, and the lack of detail indi-
cates that it is a cleavage relief. The trail length is 12 cm
and the width is less than 1 cm. Miller’s illustration is
slightly deceptive, as it shows cuneiform imprints similar
to those of the type specimen. In contrast, the specimen re-
veals that the imprints were made by straight downward
movement. As in the type specimen the imprints are denticu-
late, although the 3-4 denticles are not so well preserved.
A small degree of dimorphism, evident in the lower part
of the trail, indicates that the direction of movement was
from bottom to top, as seen on Plate 73, figure 3. For the
most part the body was parallel to the axis of movement.

The final specimen (MCZ 544; latex mold 37671)
figured by Miller (1880, pl. 13, fig. 4) is shown on Plate
73, figure 4. In contrast to the others, it is preserved as a
concave epirelief. A latex mold was made in an attempt
to determine if the cavities concealed any details not visible
from the surface. However, the mold reveals imprints simi-
lar to those in the specimen just discussed. The trail has a
length of 6 cm and a width of 4-7 mm. Some of the imprints
in the right row (see Text-fig. 20-B) can be differentiated
into sets, while those of the left row are superimposed.
This demonstrates that the direction of movement was from
bottom to top, as viewed in Plate 73, figure 4, and that the
body was oriented slightly to the right of the axis of move-
ment.

Interpretation. — There is evidence to indicate that 7.
numerosum is the trail of Cryptolithus. As shown in the
introduction to this section, Cryptolithus is the only Cin-
cinnatian trilobite in which the morphology of the dactyls is
known from preserved material. Beecher’s (1895) illustra-
tions of the genus plainly show that the terminal joint con-
sists of several bristle-like digits. This type of structure is
capable of producing the multidigitate imprint of the trail.
Moreover, the dimensions of the track are well within the
range of the average Cryptolithus. As mentioned previously
in the discussion of Allocotichus, the mean “three-quarter
width” of Cincinnatian Cryptolithus is 1.59 cm. The great-
est width of the trail is 1.5 cm and normally it is less than
a centimeter. Also the crowding of the imprints suggests
the closely spaced legs seen in Cryptolithus. However, it is
puzzling that no evidence of the long genal spines is pre-
served with the trail. One might expect that the long spines
would leave linear imprints on either side of the track. It

may be that some of the markings near the trail on the
type specimen represent the intermittent contact of the
genal spines with the substrate, but it is not possible
definitely to distinguish them.

Two other trails thought to be made by Cryptolithus
were found in the University of Cincinnati collections. Un-
fortunately locality data are lacking, but Cryptolithus frag-
ments found in the slab bearing the specimens probably
indicates Eden rocks. These are not assigned to 7. nwmero-
sum and are informally discussed below. In reality they lie
closer to the bandlike Cruziana burrows than they do to
trilobite surface-walking trails. Both are preserved as con-
vex hyporeliefs in calcisiltites, At first glance the form illus-
trated on Plate 75, figure 1 appears to be a single discon-
tinuous trail, but as will be shown, the gently curving arc
on the upper right is not connected with the serpent-like
complex on the left.

The main body of the trail consists of a series of dis-
continuous chevron-shaped ridges which average 6-8/cm.
In places (PI. 83, fig. 5) the chevrons are so closely spaced
that they resemble the continuous bilobate ridges of Cruzi-
ana. Close observation shows that the ridges are covered
by delicate bundled striae. On either side of the trail are
two thin ridges which are separated from the chevrons by
a distance of 2-5 mm. These are discontinuous and may be
present on one side of the trail and lacking on the other.
The width of the entire trail, including the ridges, varies
from 0.9-1.2 cm. The other specimen (PI. 65, fig. 7) is simi-
lar to that described above except that the chevron bodies
merge to form a continuous bilobate trail. Likewise, the
thin ridges flanking the trail are better developed, and in
one area the bilobate ridges actually rest on a small ter-
race, as shown in Text-figure 24.

Text-figure 24.—Schematic cross-section of a Cruiziana-like
trail of ?Cryptolithus. The hooklike casts of markings made by the
walking legs rest on a “terrace’’ which was gouged out by the
cephalon; X 1.25.

Both specimens are judged to be the Cruztana-like
trails of Cryptoltthus. The bundled striae on the chevron-
shaped bodies resemble the imprints that could have been
made by brushlike dactyls of the walking legs seen on
Beecher’s illustration of Cryptolithus. Moreover, the narrow
width of the trail places it well within the range of most
Cincinnatian specimens of Cryptolithus. The V-shaped

368 PALAEONTOGRAPHICA AMERICANA (YI, 41)

ridges demonstrate that the legs operated in a manner simi-
lar to that used when excavating a Rusophycus-like bur-
row, i.e. the digging movement was directed toward the
median axis and to the rear. Thus, the animal was plough-
ing over the surface instead of using merely the distal por-
tions of the legs. As in Rusophycus the open end of the
“V” formed by the striae of the opposing lobes is directed
anteriorly. This indicates that movement in the serpentine
trail, shown on the left side of Plate 75, figure 1, was from
top to bottom. On the other hand, the direction of move-
ment in the arcuate trail, seen on the right side of Plate
75, figure 1, was from left to right. Movement in the speci-
men on Plate 65, figure 7 was from left to right. The smaller
secondary ridges represent places where the genal spines
gouged the surface. In the one specimen the “terrace” on
which the ridges occur reveals that at times the cephalon
acted as a shovel and skimmed off the upper millimeter or
so of the substrate.

This type of trail is similar to the classic bandlike
Cruziana which is especially well developed in the Silurian
of Portugal and has been figured by Delgado (1910). The
only difference between the two is that the Silurian repre-
sentatives form even, bilobate ridges with continuous sec-
ondary ridges on either side. The Cincinnati forms vary be-
cause the trail was shallower; thus, the markings produced
by the legs generally appear as individual chevron-shaped
markings instead of continuous bilobate ridges. The broad
cephalon of Cryptolithus, combined with the relatively
small legs, may well have impeded digging in all but the
softest sediments.

The ethologica! significance of these Cruziana-like trails
is unknown. Perhaps, as Seilacher (1959) suggested, they
are feeding trails, where the organism stirred up the mud
and filtered out the nutrients. On the other hand, they
may be a variation of Repichnia.,

Teratichnus Miller, 1880
Plate 75, figure 3; Text-figures 23

1880. Teratichnus Miller, Cincinnati Soc. Nat. Hist., Jour., vol. 2,
ey ale pale eh ainfeg Ie

1962. Teratichnus Miller, Hantzschel, Trace-Fossils and Problematica
in Treatise on Invertebrate Paleont., R. C. Moore (ed.), pt.
W, Miscellanea, p. W218.

Type species. —Teratichnus confertum Miller (1880)
(PI. 75, fig. 3; MCZ 540; latex mold UCM 37570) from the
?Southgate beds of State Avenue, Cincinnati, Ohio. Mono-
typic.

Diagnosis. — An arthropod track, consisting of nine
bifid imprints per set, which is marked by a median groove
located midway between the right and left series.

Discussion. — The genus is known only from the type
specimen and is preserved as a faintly impressed concave
epirelief on a gently undulating calcisiltite. Because of
the difficulty of photographing the actual type specimen, a
latex mold was used for illustrative purposes. The trail,
which is incomplete at both ends, is sickle-shaped and has
a total length of 26 cm. The trail width does not exceed
1.7 cm. In the lower part of the specimen, as viewed in
Plate 75, figure 3, the imprints are irregular and there are
no well-defined sets. The first discernible set of imprints
is located 2 cm from the bottom of the slab and is marked
by the letter A. These imprints do not form a “V” or modi-
fied “V,’ as in most trilobite tracks, but instead are ar-
ranged in an elliptical pattern. This suggests that the animal
lay in this spot and moved the legs toward the median
axis as if to momentarily anchor itself to the substrate.
Extending from one end of the ellipsoid is a long (1.5 cm),
sharply defined groove (C). Similar grooves are located at
positions B and D. These grooves are significant in assign-
ing a maker to the trail. In the center of the slab the im-
prints become highly confused, and as will be shown, it is
believed that in this area there was considerable rotation
of the body. In the remainder of the trail it is possible to dis-
tinguish at least portions of individual sets. As illustrated
in Text-figure 23, there is an average of nine imprints per
set, and the open end of the “V” is directed toward the top
of the slab. Withdrawal markings are plainly defined at the
proximal end of many of the imprints, and one set shows
four imprints which are bifid distally and unifid proximally
GRIG7Sy tips 3)

Interpretation. — Although Miller (1880) interpreted
Teratichnus as the trail of a cephalopod, it is evident that
it is of arthropod origin. The basic direction of movement,
as viewed in Plate 75, figure 3, was from bottom to top
This is indicated by three lines of evidence: 1) The ellip-
soidal series of imprints (Pl. 75, fig. 3B) truncate those
imprints found near the lower margin of the slab; 2) I>
the confused area in the center of the slab, the imprints lead-
ing directly into the long sweeping are are superimposed
over the earlier formed imprints, indicating that the move-
ment here was from right to left; 3) In the arcuate portion
of the trail, the open end of the “V” formed by the sets is
directed toward the top of the slab (see Text-fig. 23). It
is believed that as the organism entered from the bottom, as
seen in Text-figure 23, the direction of movement was
nearly at right angles to the body axis. As noted above, the
animal then briefly anchored itself to the substrate. After
progressing forward for a few centimeters, the orientation
of the imprints shows that the organism apparently moved
obliquely backwards, Finally the animal moved in a tight

TRACE FOSSILS CINCINNATI AREA: QOscGoop 369

arc, pivoting on the walking legs of the right side and lash-
ing out at the bottom with the legs of the left side. As
shown in Text-figure 23, the body was oriented to the left
of the axis of movement. Thus, the imprints of the left
side are well defined, while those of the right side are
superimposed and form a continuous line. Whether this un-
usual movement was voluntary or whether the animal was
being driven by current is difficult to determine. There is
no clear evidence of current action on the slab.

It is possible that Teratichnus is the trail of a trilo-
bite similar to those discussed under Petalichnus multi-
partitum. However, the three grooves (PI. 75, fig. 3, points
B,C,D) are an integral part of the trail, and these appear
to be imprints of a median posteriorly directed spine or
appendage. No known Cincinnatian trilobite possesses a
single well-developed pygidial spine. While Seilacher (1955)
has attributed twin grooves in redlichiid trails to imprints
of caudal cerci. According to Moore, Harrington, and Stub-
blefield (1959, p. 118), such cerci are found preserved only
in the Cambrian species Olenoides serratus; and it is not
known whether they were common to all trilobites. The only
other arthropod known to be present in the Cincinnatian
which has a posterior spine and is similar in size to the
track is the aglaspid Neostrabops Caster and Macke (1952).
Although the telson of this genus is unknown, most Cam-
brian aglaspids possessed a narrow lanceolate telson. The
dimensions of the sole specimen of Neostrabops compare
favorably with those of the track. Neostrabops has a maxi-
mum width of 2.8 cm, while the track width is 1.8 cm.
The length of the sets, which because of the dimorphism is
difficult to measure, is 4.9 cm. The length of Neostrabops,
which is incomplete and measured only to the end of the
10th segment, is 3.5 cm. The nine imprints present in the
sets of 7. confertuwm correlate well with the 11 thoracic
segments of the aglaspid. The only apparent difference is
that, as Raasch (1939, p. 62) demonstrated, aglaspids had
but a single dactyl; as noted above, the trail shows the
imprints of bifid dactyls.

In summary, it is concluded that 7. confertwm was
made by a relatively small arthropod which possessed bifid
dactyls and a median posterior spine or fleshy appendage
(cercus). At present it is not possible to give a more pre-
cise assignment. No known Cincinnatian arthropod exactly
matches the morphology of the track. None of the trilo-
bites has a single pygidial spine, and the nature of the cerci,
if present, is unknown. On the other hand, aglaspids have
a telson, but the walking legs apparently have but a single
dactyl. It must also be noted that Neostrabops is known
only from a single specimen.

RHABDOGLYPHUS Vassoevich, 1951
Plate 78, figures 4-6, 8, 9

1951. Rhabdoglyphus Vassoevich, Les Conditions de la Formation
du Flysch, p. 61, pl. 5, fig. 3; pl. 6, figs. 3,4.

1962. Rhabdoglyphus Vassoevich, Bouéek and Elias, Geol. Prace
Zpravy, vols. 25,26, pp. 145-151, pls. 7-8.

Type species —Rhabdoglyphus grossheimi Vassoevich
(1951) from the Cenomanian of the Caucasus, Czechoslo-
vakia. Monotypic.

Diagnosis. — Rosary-like trails or tunnel fillings of vary-
ing length, preserved either as convex hyporeliefs or con-
cave epireliefs.

Discussion. — Although Vassoevich proposed Rhabdo-
glyphus for what he referred to as a “bioglyph,” his genus
was invalid due to the lack of a formal diagnosis. Recogniz-
ing this, Bouéek and Elias (1962) formally described and
diagnosed the form, studying the original type material, as
well as additional specimens from the Paleogene Flysch of
the Czechoslovakian Carpathians. The facts related below
are taken from the plates and a short German summary.

R. grossheimi is preserved as a convex hyporelief and
consists of cylindrical tunnel fillings with periodic well-de-
fined swellings or “knots” (Pl. 78, figs. 8, 9). The diameter
of the cylindrical fillings is 4 mm, that of the swellings is
7 mm. The latter are not evenly spaced, as the distance
between them varies from 1.8-3.0 cm, and in one form two
swellings occur together. Judging from the specimen illus-
trated on Plate 78, figure 8, it seems that one is not dealing
with a simple convex hyporelief, for at both ends the hemi-
cylinder appears to extend up into the bed.

Rhabdoglyphus resembles Hormosiroidea florentina F.
X. Schaffer (1928) from the Pietraforte Limestones (Upper
Cretaceous) near Florence, Italy. Through the kindness of
Dr. Zapfe of the Natural History Museum of Vienna, a
plaster case of Hormosiroidea was made available for study
(PI. 78, fig. 7). The specimen, 9 cm long and shaped as an
open “S” curve, consists of a series of raised hemispherical
bodies connected by thin, short hemicylindrical rods. The
diameter of the hemisphere ranges from 6-9 mm; the di-
ameter of the connecting rods is 1 mm. Near the larger
specimen is a slightly shorter form consisting of only two
hemispheres. Associated with these two specimens are small
raised vermiform-like markings whose diameter is the same
as the connecting links between hemispheres. The attitude
of the latter indicates that Hormostroidea is most probably
a convex hyporelief rather than a convex epirelief as
Schaffer believed.

It is entirely possible that Rhabdoglyphus is synony-
mous with Hormosiroidea. Although the general morphology

370 PALAEONTOGRAPHICA AMERICANA (VI, 41)

of the two is similar, a decision can not be made without
additional study. Both forms should be sectioned to de-
termine if they are exclusively surface features or whether
they actually extend into the host rock as they appear to.

Interpretation. — Schaffer interpreted Hormosiroidea
as an alga comparable to the Recent Hormosira and re-
garded the swellings as spore cases. Bouéek and Elias sug-
gested that Rhabdoglyphus may represent the workings of
an amphipod, a gastropod, e.g., Bulla, or even possibly a
holothuroid similar to Leptosynapta. They rejected a pos-
sible annelid origin. In the German summary they give no
explanation for the periodic swellings nor do they give any
reason for mentioning Bulla and Leptosynapta as possible
originators.

CINCINNATIAN REPRESENTATIVES

Rhabdoglyphus sp.
Plate 22, figures 4-6; Text-figure 29]

Discussion. — Rhabdoglyphus-like forms are present,
though uncommon, in the Cincinnatian. Only 10 specimens
have been collected, eight from the Richmond and one
each from the Eden and Maysville. Unfortunately material
has never been observed in place. In contrast to the Euro-
pean forms discussed above, al! Cincinnatian representatives
are preserved in calcisiltites and calcarenites as concave
epireliefs, 7.¢., as hemispherical depressions normally con-
nected by hemicylindrical troughs. The entire pattern,
whose length ranges from 9-30 cm, is either straight or
gently curved. Both the size and shape of the hemispherical
depressions vary greatly. They may be round (PI. 78, fig.
4), cordate (PI. 78, fig. 5), bottle-lke (Pl. 78, fig. 5), or
rectangular (PI. 78, fig. 6). When only four or five de-
pressions are present, they are normally of the same size,
but when the number is greater, there is a definite gradation
from smaller to larger, One example noted in the field con-
sisted of 12 depressions. The first and largest measured 8
mm in diameter, the last only 2 mm. The depth varied
from 0.5-1.0 mm and generally speaking the larger the
diameter, the greater the depth. As in European specimens
of Rhabdoglyphus, the distance between the depressions
varies. The actual measurements are present in Table IT.

The shallow connecting troughs are only one-third or
one-half as wide as the depressions. In some cases the
troughs are deeper adjacent to the depressions and appear
to bend down into them. Where there is a definite grada-
tion in the size of the depressions, there is an associated
trend in the development of the connecting troughs. The
larger and deeper the depression, the better developed the

TABLE

Specimen

Number Hemisphere Number
1 2 3 4 5 6 ah 8 9 10 11 12
1) 5 18 20 13 0 —- ee oe
2) 8 8 if 7 9 - -—- -—- — =
3) 15 6 16 9 3 if 8 7 —- —- —
4) 1l 8 u 20 18 - = Ore >
5) 8 11 - FOF a a ae ae re
6) 8 7 8 5 - ES Oe Oe or
i) 5 7 12 Wd —-— ~—~ —> ~—> ~—> ~— —
8) 7 10 12 17 17 7 - - Fre Ohl
2))) 9 7 10 ? 6 9 6 _— —_ —_ —_
10) 9 10 10 14 10 8 11 3 7 6 17

connecting trough; on the other hand, where the depres-
sions are small and shallow, there may be no connecting
trough present (PI. 78, fig. 6). Normally the connecting
troughs appear to be simple impressions, but one specimen
(Pl. 78, fig. 6) shows raised borders, and apparently there
has been movement along this trough from left to right. No
evidence of sculpturing was found on any of the speci-
mens; both the connecting troughs and the hemispherical
depressions are smooth.

A comparison of the morphology of the Cincinnatian
forms with that of Rhabdoglyphus shows that both can be
safely placed within a single genus. The Cincinnatian speci-
mens are judged to represent molds of the same trace de-
scribed from casts in Europe. This is a difference in preser-
vation rather than morphology, and for the present there
seems no good reason for not assigning the Cincinnatian
representatives to Rhabdoglyphus.

Interpretation. — The origin of these beadlike forms
is not readily apparent. Other than bearing a superficial
resemblance to the Recent algal genus Hormosira Schaffer,
1928, figs. 1,2), there is no indication that it is a plant.
The general form recalls prod marks such as those figured
by Dzulinski and Sanders (1962, pl. XII, fig. B). The
authors illustrate a sole-marking from the Polish Flysch
which shows a series of hemispheres produced by a frag-
ment of a fish vertebra skipping along the bottom. The
problem of extending this kind of explanation to at least the
Cincinnatian representatives of Rhabdoglyphus is that there
is no evidence of linear current action; in fact one specimen
(PI. 78, fig. 5) is superimposed over interference ripples.
Moreover, the hemispherical depressions of Rhabdoglyphus
do not show the distortion one would expect if an object
had gouged the bottom at a shallow angle. For these
reasons Rhabdoglyphus is not thought to be of physical
origin.

Although the behavioral pattern reflected by its mor-
problem of extending this kind of explanation to at least the
Rhabdoglypthus is a trace fossil, most probably a variety of
Repichnia or Fodinichnia. As mentioned previously, one

TRACE FOSSILS CINCINNATI AREA: OsGoop 371

specimen illustrated by Bouéek and Elias appears to ex-
tend up into the host rock, and one Cincinnatian specimen
(Pl. 78, fig. 6) does show evidence of movement. If we
assume that these fossil-bearing calcisiltites and calcarenites
were overlain by lutite*® and that the form represents in-
faunal activity, it is relatively easy to explain why troughs
are not found between all the hemispheres. It is believed
that the organism burrowed down through an unknown
thickness of lutite, periodically altering its behavior to pro-
duce the “knots” or swellings. Only where it was burrowing
horizontally along the shale-silt interface would we find
both the swellings and connecting depressions. If the animal
veered up slightly into the mud, the connecting depressions
would be the first feature to be lost, since they are not so
deeply impressed as the “knots.” However, one would ex-
pect the swellings to be smaller and shallower than when
there is a connecting link between them. This is exactly the
case. The same would hold for the specimens described by
Bouéek and Elias, except here the calcarenite which contains
the specimen was probably underlain by lutite. The organ-
ism burrowed down through the calcarenite and along the
interface. The overlying calcarenite filled in the depression
as the animal moved along, thus producing the type of
preservation commonly encountered in trace fossils, 7.e.,
convex hyporelief. Still unexplained is the nature of the
organism which produced the trace and the meaning of the
periodic swellings.

One Recent trace has been described which may pos-
sibly offer a basis for comparison with fossil forms. In at-
tempting to explain the origin of Arthraria, Fenton and
Fenton (1934a, pl. XXVIII, fig. 2) described the peculiar
actions of the gastropod J/lyanassa obsoleta (Say) on the
mud flats of San Francisco Bay. J/lyanassa ploughs along
the surface, leaving a trail much like the connecting de-
pressions found in Cincinnatian Rhabdoglyphus, then it
abruptly twists and digs a shallow conical pit before once
again proceeding on its way. The authors give no reason
for this behavior.

The meaning of periodic swellings in Rhabdoglyphus is
uncertain. Perhaps they are infaunal analogues of the conical
pits of J/lyanassa, It is also possible that Rhabdoglyphus is
the burrow of a Sedimentfresser and that the swellings
represent periodic (peristaltic) anal constrictions and ex-
pansions. Likewise, they might be the results of localized
muscle knotting, possibly for the eversion of a proboscis.

Rhabdoglyphus bears a superficial resemblance to some

“One specimen shows a finer grained mud filling the hemispheres.

moniliform sponges, e.g. Girtyocoelia; however, there is no
evidence to indicate that Rhabdoglyphus is the mold of a
sponge or spongelike body.

TYLICHNUS n. gen.
Plate 66, figure 7; Plate 71, figure 2; Text-figures 25,29m

1842. Non Cruziana d’Orbigny, Voyage dans l’Amérique meridionale,
3) psp. 30hpla de ties

1852. Non Rusophycus Hall, Nat. Hist. New York, Palaeont. Neqw
York, vol. 2, p. 23, pl. 9, figs. 1a-c.

1873. Non Pianolites Nicholson, Roy. Soc. London, Proc., vol. 21, p.
389.

1878. Rusophycus Hall, Miller and Dyer, Cincinnati Soc. Nat. Hist.,
Jour., vol. 1, p. 2, pl. 1, figs. 5-5a.

1885. Partim Cruziana d’Orbigny, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 156, pl. 8, figs. 3.

1891. Partim Planolites Nicholson, James J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 14, p. 47.

1962. Partim Walcottia Hantzschel, Trace-Fossils and Problematica
in Treatise on Invertebrate Paleont., Moore (ed.), pt. W,
Miscellanea, p. W243.

Type species. — Rusophycus asperum Miller and Dyer
(1878) (PI. 71, fig. 2; HBM 3179; latex mold USM 37588)
from the Eden beds, Cincinnati, Ohio, Monotypic. The form
is not common and is known from only four specimens.
Regrettably precise locality data are lacking.

Diagnosis. — Faintly bilobate trails preserved as con-
vex hyporeliefs; surface of trail characterized by nine paral-
lel rows of nodes which are offset so as to form a “zipper-
like” pattern.

Discussion. — Miller and Dyer (1878a, p. 2) charac-
terized “Rusophycus” asperum as consisting “. . . of long
stems, flattened on the upper side, or having a longitudinal
depression, which gives to them a subangular outline. The
surface is made very rough by numerous papillae, many of
which are transversely elongated.” While they remarked
that “.. . it is separated by generic differences from such
forms as R. pudicum and R., bilobatum” (ibid, p. 2), they
declined to establish a new genus. As will be shown below,
the unusual papillose ornamentation serves to differentiate
this form from all known trace fossils. Accordingly the new
genus 7'ylichnus is proposed for “Rusophycus” asperum.

The type specimen of T'ylichnus asperum (PI. 71, fig. 2)
is perserved as a convex hyporelief on a fine-grained, cross-
laminated calcarenite. The slab bears short fragments of
several specimens which interpenetrate but do not branch.
Vertical sectioning reveals that the forms extend up into the
host rock and are subquadrate in cross-section, the width of
the burrow being 3.5 mm; the height, 1.5 mm. The contact
between the burrow filling and the host rock is not sharply
defined, and there is no apparent difference in lithology. As
viewed in Plate 71, figure 2, the surface of the burrow is
flattened, although one portion (upper left) is faintly bilo-

372 PALAEONTOGRAPHICA AMERICANA (VI, 41)

bate. The pustulose ornamentation varies in its arrange-
ment. As shown in Text-figure 25, the basic pattern con-
sists of nine parallel rows of transversely elongate nodes.
The rows are offset so as to give the overall pattern a
“zipper-like” appearance, In contrast, some segments of
the burrow show no preferred arrangement, and the closely
packed nodes are distributed randomly over the surface.

Text-figure 25. — Enlargement of Tylichnus asperum (Miller and
Dyer), 1878. The ornamentation of the holotype consists of alternating
rows of nodes arranged in a 5-4-5-4- pattern. In other members of the
species the arrangement is not so regular; X 7.5.

Plate 66, figure 7 shows another slab bearing six short
specimens. The burrow width is 2.5-3.0 mm, and the largest
form has a length of only 1 cm. The nodose surface of these
specimens differs somewhat from that seen in the type.
In one form the nodes are arranged in three parallel rows
which are slightly oblique to the longitudinal axis of the
burrow. The lateral rows consist of a single moniliform chain
of nodes. In opposition, the median row is slightly wider,
and along most of its length the nodes exhibit no preferred
grouping. However, in one small segment the nodes are
arranged in an alternating 1-2-1-2 pattern. The remaining
specimens on the slab are weakly bilobate, and the nodes do
not show a linear arrangement; however, the lateral nodes
are slightly larger than the median. On one of the specimens
the side of the burrow adjacent to the host rock is partly
exposed, and the same nodose ornamentation can be seen,
suggesting that the entire burrow may be characterized by
a pustulose surface. Vertical sections reveal that, like the
type specimens, the burrows are of infaunal origin.

The additional specimens available for study add
little to the knowledge of the ornamentation. However, they
do show that the burrow may attain a length of up to 8.5
cm.

Interpretation. — Miller and Dyer (1878) believed
that “Rusophycus” asperum was a “fucoid,”’ while J. F.
James (1885) concluded that it was an annelid burrow.
In 1891 James placed the form in Planolites, but as shown
elsewhere, his concept of Planolites embraced such a variety
of forms that it is virtually meaningless. It is also clear
from the above discussion that Tylichnus bears no resem-
blance to either Rusophycus or Cruztana. The nodose sur-

face of T'ylichnus is totally unlike the striae which cover the
lobes of the trilobite burrows.

The nodose surface recalls forms such as Ophiomorpha
Lundgren (1891) from the Upper Cretaceous and Tertiary
of North America, Europe, and Japan and Pennatulites de
Stefani (1885) from the Tertiary of Europe and the West
Indies. However, as illustrated by Hantzschel (1952, pls.
13,14), Ophiomorpha is oriented either normal to or only
slightly oblique to the bedding and may attain a diameter
of 2 cm. Moreover, the nodes of Ophiomorpha are much
larger and show no preferred groupings. Pennatulites is a
bilobate bundled burrow and is briefly discussed herein
under Trichophycus venosum. The specimen of Pennatu-
lites illustrated by Hantzschel (1962, figs. 127-la,b) has a
diameter of 4 cm, and while the nodes are smaller than
those of Ophiomorpha, they are arranged in parallel rows
much like kernels of corn.

Since there are no known Recent counterparts of
Tylichnus, little can be said of the originator of the burrow,
The subquadrate cross-section and the possibility that the
nodes encircle the entire burrow suggest a vermiform body
ringed by some type of papillae. Perhaps the nodes repre-
sent the impressions of a feeding apparatus surrounding the
mouth of an organism working along the lutite-calcisiltite
interface. There is no evidence of a genetic relationship be-
tween 7’. asperum and the moniliform ?fecal string seen on
Plate 66, figure 7 (center bottom).

ORMATHICHNUS Miller, 1880
Plate 78, figures 1-3

1880. Partim Ormathichnus Miller, Cincinnati Soc. Nat. Hist., Jour.,
vol, 2, p. 222, pl. 14, figs. 4,5.

1886. Partim Ormathichnus Miller, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 8, pp. 160,161.

1962. Partim Ormathichnus Miller, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore( ed.),
pt. W, Miscellanea, p. W207.

Type species. — Ormathichnus moniliformis Miller
(1880) from “Walker Mill Road [State Avenue, Cincinnati,
Ohio], less than 100’ above low water” (probably South-
gate). Monotypic.

Diagnosis. — A small trail resembling a_ series of
minute disconnected Rusophycus which grade into the con-
tinuous ridges of Cruziana.

Discussion. — Miller (1880, p. 222) described the
genus as “a single continuous beaded track or trail.” Two
specimens were figured and discussed. The first (Miller,
1880, pl. 14, fig. 4) he likened to a cast of the stem of
Heterocrinus simplex; the second (PI. 70, fig. 5) differed in
that it possessed ‘“‘a longitudinal furrow in the middle.”

An analysis of the syntypes reveals that they are two

TRACE FOSSILS CINCINNATI AREA: OsGoop 373

different things. The form iJlustrated by Miller on plate 14,
figure 4 (herein Pl. 78, fig. 3) is preserved as a convex
hyporelief and consists of a row of 30 moniliform welts
whose aggregate length is 25 mm. It fades out in one
direction and is incomplete at the other end. It is associated
with groove casts and a few scour marks, as well as a series
of parallel striae which resemble a cast of the markings
made by a rolling crinoid stem.

The other specimen (Miller, 1880, pl. 14, fig. 5; herein
Pl. 78, figs. 1,2), 1 mm wide and 6.0 cm long, is semi-
circular in outline. The morphology of the form varies along
its length. In the upper left-hand corner, as viewed on Plate
78, figure 2, it consists of three long grooves separated by
two raised ridges. These ridges are so small that it is
impossible to determine whether they are striate or smooth.
The remaining 4.0 cm of the specimen resemble a series of
minute unconnected Rusophycus (Pl. 22, fig. 1). These re-
call on a much smaller scale Seilacher’s (1955, figs. 4a-b
and 5a) illustration of Rusophycus didymus.

Interpretation. — Although Miller stated that Orma-
thichnus was probably a gastropod trail, he gave no sup-
porting evidence other than the fact that gastropods were
found in the same bed. J. F. James (1886) regarded the
genus as a cast of the impression of a rolling crinoid stem,
and this probably is a correct interpretation for one of the
specimens (PI. 78, fig. 3). The length of the raised seg-
ments (1 mm) is equal to the spacing between the parallel
striae on the same surface. The only problem is_ that
“Ormathichnus” is oriented parallel to the current direc-
tion. Normally casts of the impressions of rolling crinoid
stems are either at right angles or oblique to the current.
In any event it apparently is not a trace fossil but some
variety of an inorganic tool mark.

The other specimen (PI. 78, figs. 1,2) is judged to be
the trail of a small arthropod, possibly a trilobite. Based
on the orientation of the Rusophycus-like imprints, move-
ment was from left to right as viewed in Plate 78, figure 2.
In reality the trail is a combination of Cruztana and Ruso-
phycus. The organism initially ploughed along producing a
“micro-Cruziana’” consisting of three grooves and two
ridges. It then altered its method of movement to give rise
to a series of unconnected Ruscphycus. Because of its ex-
tremely small size, Miller’s original generic designation is
retained; but, the form is not a common Cincinnatian trace
fossil, and no subsequent specimens have been found.

PALAEOPHYCUS Hall, 1847

Plate 76, figures 1-9; Plate 77, figures 1,4-7; Plate 83, figures 1,4;
Text-fig. 29k

1847. Palacophycus Hall, Nat. Hist. New York, Paleont. New York,
vol. 1, pp. 7,8,63, pl. 2, figs. 1-5; pl. 21, fig. 2; pl. 22, fig. 1.

1852. Palaeophycus Hall, Hall, Nat. Hist. New York, Paleont. New
York, vol. 2, pp. 6,22, pl. 3, figs. 2a-b; pl. 22, fig. 10.

1873. Non Planolites Nicholson, Roy. Soc. London, Proe., vol. 2, p.
289.

1885. Palacophycus Hall, James, J. F., Cincinnati Soc. Nat. Hist.,
Jour., vol. 7, pp. 155-157.

1891. Partim Planolites Nicholson, James, J. F., Cincinnati Soc. Nat.
Hist. Jour., vol. 14, p. 47.

1962. Palaeophycus Hall, Hantzschel, Trace-Fossils and Problematica
in Treatise on Invertebrate Paleont., Moore (ed.), pt. W, Mis-
cellanea, p. W208.

Type species.— Palaeophycus tubulare Hall (1847)
from the “Calciferous Sandstone” (Beekmantown; Lower
Ordovician, marine) of the Mohawk Valley of New York
State (NYSM 241/1, 241/2, 241/3, 241/4). Lectotype, Wil-
son (1948).

Diagnosis. — Slightly irregular cylindrical burrows of
varying length which are oriented obliquely to the bedding
plane. Most forms are unbranched, although a few bifur-
cate infrequently in no apparent pattern. The genus is most
commonly found as hemicylinders which are preserved
either as convex hyporeliefs or concave epireliefs. The sur-
face of the hemicylinder may be smooth, striate, or annu-
late. Specimens vary from 3-15 mm in diameter and have
a length of 5-20 cm.

Discussion. — Hall (1847, p. 7) defined Palaeophycus
as “stem terete, simple or branched, cylindric or sub-
cylindric surface nearly smooth, without transverse ridges,
apparently hollow.” Four specimens of the type species were
figured and each of these will be discussed below.

1) P. tubulare Hall (1847, pl. 2, fig. 5; herein Pl. 77,
fig. 4; NYSM 241/2; latex mold UCM 37700). This speci-
men, which is preserved as a convex hyporelief in a blue,
medium-grained calcarenite, contains several terete or hemi-
cylindrical tubes which vary in diameter from 3-7 mm.
Apart from one form which has a median longitudinal fur-
row, the surface of the tube is smooth. None of them show
any clear evidence of branching, although interpenetration
is common; and where the tubes intersect the side of the
slab, the bedding is disturbed. Thus, these are not mere
surface features but are instead of infaunal origin. There
is no marked difference between the composition of the
tubes and that of the host rock.

2) P. tubulare Hall (1847, pl. 2, fig. 2; herein Pl. 77,
fig. 7; NYSM 241/2; latex mold UCM 37701). In essence
this specimen is similar to the one just described. It is pre-
served as a convex hyporelief in a calcarenite, and the
hemicylinders with their smooth surface have a diameter
which ranges from 3-10 mm. Only one form is seen to
bifureate; the others cross over and interpenetrate. Obser-
vation of the margins of the slab reveals that the burrows
penetrate the host rock and disturb the bedding. The fill-

374 PALAEONTOGRAPHICA AMERICANA (YI, 41)

ings of the tube are slightly more fine grained than the
surrounding calcarenite. Presumably serial sectioning of the
specimen would show that those hemicylinders which appear
to fade out on the surface in reality extend up into the host
rock.

3) P. tubulare Hall (1847, pl. 2, fig. 4; herein Pl. 77,
fig. 1; NYSM 241/3; latex mold UCM 37699). This form
is so poorly preserved that it adds little to the knowledge
of the genus. The host rock is a dense, blue, limonite-en-
crusted calcarenite. The single smooth terete tube, which
is 8 mm in diameter, appears to give off three small
branches. It is not possible to determine whether the tube
is preserved as a convex hyporelief or convex epirelief.
Likewise, it is so encrusted with limonite that its composi-
tion is unknown.

4) P. tubulare Hall (1847, pl. 2, fig. 1; herein Pl. 76,
rig. 8; NYSM 241/1; latex mold UCM 37698). Remnants
of bedding indicate that the host rock originally consisted
of thick beds of blue calcarenite interstratified with thinner
layers of pink calcarenite. However, the bedding has been
so disturbed by bioturbation that the pink calcarenite now
appears only as lenses. The numerous tubes possess the
same basic patterns as those in the specimens described
above, for there is no branching but interpenetration is
common. They differ only in that they are slightly larger
(diameter of largest form is 1.7 cm), and their surface
bears faint irregular longitudinal striae. Vertical sections
show that only about one-third of the surface of the tube
is visible; thus, here again we are dealing with cylindrical
bodies which extend up into the host rock.

In summary then, a restudy of the syntypes of P.
tubulare demonstrates that the type species consists of gen-
erally unbranched cylindrical tunnel fillings of varying di-
ameters which extend up into the host rock. Aside from
the exceptions noted above, the surface of the cylinders is
smooth, and the fillings have a composition similar to that
of the host.

Hall (1847, 1852) proposed six additional species of
Palaeophycus. In order to illustrate the wide range of mor-
phology occuring in the genus, these will be briefly dis-
cussed below.

1) P. rugosum Hall (1847, pl. 21, fig. 2; NYSM 240/1
was described from the marine Trenton (Middle Ordo-
vician) beds of Middleville, New York. The species (PI. 76,
fig. 1) is characterized by coarse irregular rugosites which
are oriented approximately parallel to the axis of the
branch. The main branch has a diameter of nearly 3 cm
and is forked at one end. As the margin of the slab shows,
the specimen is preserved as a convex hyporelief where

the composition of the filling is the same as that of the
host (coarse-grained calcarenite).

2) P. tortwosum Hall (1852, pl. 3, figs. 2a-b; USNM
41102) was proposed for forms from the red Queenston
(Upper Ordovician) strata of Irondequoit Bay, New York.
The smooth subcylindrical fragments (Pl. 76, figs. 2-4) are
composed of an ferruginous silt and have a diameter of 1
cm. As Hall noted, two of the specimens are _helicoid.
Since the specimens are completely disassociated from the
host rock, it is impossible to determine the lithology of the
latter.

3) ?P. striatum Hall (1852, p. 22, pl. 10, fig. 1-2) from
the Clinton (Middle Silurian) of Oneida County, New
York, was placed in the genus with some hesitation. Only
two of the five specimens figured by Hall were studied.*®
The specimens are preserved as convex hyporeliefs in poorly
sorted silts and can be seen to extend up into the host.
The form illustrated on Plate 76, figure 6 consists of two
subcylindrical burrows which have a width of 1 cm. Both
are marked by numerous faint longitudinal striae, and the
longer burrow is characterized by a narrow longitudinal
crest. The other slab (Pl. 76, fig. 7) bears four unbranched
subcylindrical fillings. The width of the burrows ranges
from 6-8 mm, and only one form possesses the faint striae
which characterize the species.**

4) P. wrregularis Hall (1847, p. 8, pl. 2, fig. 3) from the
Beekmantown (Lower Ordovician) beds of northeastern
New York is an irregularly branched, smooth form which
covers “the entire surface of large slabs of sandstone.” Hall
related that it is much smaller than P. twbulare, but other
than that the species is so nondescript that it is difficult
to diagnose or compare.

5) P. simplex Hall (1847, p. 63, pl. 22, figs. la-d) from
the Trenton (Middle Ordovician) Limestone of Middle-
ville, New York, normally occurs as fragments which have
a diameter of 1 cm and a length of 15 cm. Hall remarked
that he has several hundred fragments and that “. . . the
interior is filled with fragments of shells, crinoidal columns,
etc.” (Hall, 1847, p. 63). He went on to say (ibid., p. 63)
that the species “. . . appears to have grown only during
the shaly deposition, and is imbedded in this portion of the
mass; consequently it is absent or obscure where such ma-
terial is wanting as a component part of the rock.”

%>P. striatum Hall, 1852, pl. 10 fig. la; USNM 41108; latex mold
UCM 37585; herein Pl. 76, fig. 6 and ? P. striatum Hall, 1852, pl. 10
fig 1c; USNM 41111; herein Pl. 76, fig. 7.

“The remaining three species of Palaeophycus proposed by Hall were
not studied; therefore, the description is taken from the original work.

TRACE FOSSILS CINCINNATI AREA: OsGoop SHV

6) P. virgatum Hall (1847, p. 263, pl. 70, fig. 1) was
described from the “Hudson River Group” (Upper Ordo-
vician) of Washington County, New York. Hall (1847,
p. 263) remarked that “this species is only seen in frag-
ments of long rigid stems, of nearly equal diameter . . .
Fractured or weathered surfaces of arenaceous shales often
present great numbers of these fragments, imbedded in
great confusion.” Hall said that the species was similar to
P. simplex but that “it is impossible to indicate characters
either to assimilate or distinguish them” (zbid., p. 263).

In the years following Hall’s work, 15 additional species
of Palaeophycus were proposed by various authors from
the Cambrian, Ordovician, and Pennsylvanian rocks of
North America and from a variety of depositional environ-
ments. These were differentiated from Hall’s species by the
number, size, and rigidity of the branches, as well as the
size of the entire pattern.

In view of the wide range of forms included in Palaeo-
phycus, it is not surprising that the validity of the genus
has been questioned. Most frequently Palaeophycus has
been compared with Planolites Nicholson (1873), which was
originally described from the Guelph (Lower Silurian) sec-
tion of Ontario. Howell (1943) placed Palaeophycus inci-
piens Billings (1861) and Palaeophycus congregatus Billings
(1861), from the Upper Cambrian Potsdam Sandstone of
Vermont, in Planolites. J. F. James (1885) discussed sev-
eral species under Palaeophycus, but in 1892 he considered
these, and still additional forms, under the heading of
Planolites. Recognizing the confusion, Howell (1943, p. 18)
suggested restricting Planolites to “. . . all those usually
more or less horizontal but sometimes diagonal or vertical
burrows having diameters of approximately 8-12 mm that
are not marked by such prominent cross ridges as are char-
acteristic of the species of Arthrophycus . . .” However,
as seen by the above discussion of Hall’s forms, this diag-
nosis does not really serve to differentiate Planolites from
Palaeophycus. It is instead believed that Nicholson’s orig-
inal discussion (1873) as augmented by Nicholson and
Hinde (1875) may provide the key. Nicholson (1873, pp.
309-310) described Planolites as “. . . burrows [which] run
more or less horizontally as compared with the laminae
of deposition, or they penetrate the strata obliquely . . .
[These] are preserved . . . not [as] the actual burrows
themselves, but the burrows filled up with sand or mud
which the worm has passed through its alimentary canal.”
Nicholson and Hinde (1875, p. 3) expanded the definition,
stating “they appear usually in the form of cylindrical
or flattened stemlike bodies, which are often more or less
matted together and which may cross one another in every
imaginable direction.” Although the type species, P. vul-

On

garis, was not figured, Nicholson (1873) stated that the
diameter varies from 1-4 mm; and the burrow may be ir-
regularly thickened. There was no mention of branching.
While they did not elaborate, both Nicholson (1873) and
Nicholson and Hinde (1875) remarked that some of Hall’s
species of Palaeophycus should probably be placed in Plano-
lites.

Whereas neither Nicholson nor Nicholson and Hinde
flatly stated that there is a difference between the lithology
of the cylindrical tubes and that of the host rock, they cer-
tainly imply it. In addition, their description has been fol-
lowed by Rudolf Richter (1937) and Reineck (1955).
Richter (1937) established Planolites montanus for irregu-
lar compressed cylinders from the Carboniferous (sedi-
ments) of the Ruhr. The cylinders are 4 mm in diameter
and can be traced for 1-4 cm; there is no over-all pattern.
However, there is a marked difference in the lithology of
the tunnel fillings (see Richter, 1937, figs. 2-4), and in
some cases the bedding bends around these fillings (see
Richter, 1937, fig. 5). Reineck (1955) proposed Planolites
rugulosus for specimens from the nonmarine Wadener beds
(Permian) of Germany. These are cylindrical bodies over
1 cm in diameter which are marked externally by annula-
tions and longtitudinal striae. They lie obliquely to the
bedding planes, are unbranched and follow an irregular
course. Internally P. rwgulosus consists of material which
is either coarser or finer grammed than the host rock.
A cross-section of the cylinder reveals a series of “growth
rings,” while a longitudinal section shows that these are
hemispherical packings (see Reineck, 1955, fig. 2).

Based on the above discussions, there may be clear
grounds for separating Palaeophycus and Planolites. The
key factor is the lithology of the burrow filling as com-
pared to that of the host rock. As best can be determined
from a restudy of some of Hall’s type material, there is
no marked lithological difference between the fillings and
the host. In contrast, on the basis of Nicholson’s descrip-
tion Planolites appears to show a distinct difference.

Interpretation. — Hall (1847, 1852), Billings (1862),
and other earlier authors considered Palaeophycus to be
fragments of stems of once larger “fucoids.” J. F. James
(1885) was one of the first to point out that Palaeophycus
was a burrow, and he compared the genus to Recent anne-
lid trails observed on the banks of the Little Miami River
in Ohio. Most subsequent authors, e¢.g., Bassler (1915),
Hantzschel (1962), have accepted the burrow interpreta-
tion. Although no one has studied the genus in detail, other
interpretations have been offered. Wilckens (1947) applied
Abel’s (1935) interpretation of Arthrophycus to Palaeo-
phycus and thought that both were fragments of worm

376 PALAEONTOGRAPHICA AMERICANA (VI, 41)

tubes uprooted and scattered about during storms. On
the other hand, Wilson (1948) held to the “fucoid” hypo-
thesis.

The fact that the tubes of Palaeophycus interpenetrate
and extend up or down into the host rocks indicates that
they are burrows and not “fucoids” or portions of agglu-
tinated tubes. It is concluded that they are of infaunal
origin and represent the unorganized pathways of errant
organisms which were progressing through the sediment,
quite possibly in search of food. As the animal moved along
at depth, the sediment was pushed aside only to collapse
again as the organism moved past, although in some cases
mucus may have given the walls a temporary rigidity.
There is no evidence in any of the Hall specimens studied
by the author that we are dealing with a structure which
was either packed by a Sedimentfresser or mined out and
filled by sediment sifting in from outside. There is no signi-
ficant difference between the grain size or composition of
the filling and that of the host rock,

Recent counterparts of Palaeophycus are made by pre-
daceous polychaetes (e.g., Glycera), gastropods (e.g., Poly-
nices), and presumably could be made by some pelecypods.
The difference in the sculpturing seen on some of Hall’s
species would seem to indicate that the genus includes forms
made by several different organisms. In its present state
Palaeophycus contains too wide a range of morphological
characteristics, and the type specimens alone do not pro-
vide sufficient material to permit a meaningful restudy of
the genus; they are far too fragmentary. It will be necessary
to recollect original localities, noting such characteristics
as branching patterns, if any, and the continuity of sculp-
turing on a given form. This will undoubtedly result in a
reduction of the number of species, and if a given form is
sufficiently distinctive and persistent, it may well be pos-
sible to establish new genera. Moreover, “species” of
Palaeophycus have been described from such diverse en-
vironments as the marine Trenton beds of New York, the
Queenston “red beds” of New York, and the Pennsylvanian
coal swamps of Indiana.

An additional problem is how to distinguish Plano-
lites from Palaeophycus, because the morphology of the
two is similar. Both genera are cylindrical, irregular, un-
branched, and penetrate obliquely through the strata. The
only significant difference lies in the nature of the filling.
It may well be that Nicholson (1873), Richter (1937),
Howell (1943), and Reineck (1955) were correct in be-
lieving that Planolites is the stuffed burrow of a Sediment-
fresser and that the filling is host rock that has been passed
through the animal’s gut. Thus, if a restudy of Nicholson’s
type specimens confirms this, Planolites may be restricted

to the stuffed burrows of Sedimentfressers, while Palaeo-
phycus is the passageways of errant predaceous or selective
feeders. Nevertheless, it must be admitted that in some
cases it may be difficult to distinguish between these two
categories. Little work has been done on the effect that
organisms have on the mineralogical and textural proper-
ties of ingested sediment. Presumably soft particles could
be reduced by abrasion, and it is possible that organic
acids could alter both the grain size and composition of
carbonate particles. Enzymes might also produce a color
change. Obviously the most positive evidence for a fecal-
stuffed burrow is the presence of fecal pellets.

CINCINNATIAN REPRESENTATIVES

Palaeophycus sp.

Plate 76, figures 5,9; Plate 77, figures 5,6; Plate 83, figures 1,4;
Text-figure 29k

Discussion. — J. F. James (1885) reported that five
“species” of Palaeophycus could be identified in the local
Cincinnatian section As he recognized, Palaeophycus flexu-
osum U. P. James (1879) is of inorganic origin and is not
comparable to the species of Palaeophycus figured by Hall.
This form is discussed herein under “Fucoids of Inorganic
Origin.” Likewise, what J. F. James (1885) presented as
Palaeophycus virgatum Hall does not appear to belong in
the genus. He remarked that they were “. about an
inch wide, and about eight inches long, of the same width
[throughout] their whole length ... and overlying one
another in various directions” (James, 1885, p. 158). These
were tentatively interpreted by James as the molds of large
Solen (which are unknown in the Cincinnatian). The de-
scription recalls the “turkey tracks” which have been shown
elsewhere in this work to be portions of the burrow of
Trichophycus venosum.

The remaining three “species” are P. tubulare Hall,
P. rugoswm Hall, and P. simplex Hall. J. F. James consid-
ered the latter two as synonymous and believed that P.
tubulare and P. rugoswm were two different expressions of
the same burrow. To James (1885, p. 158) P. rugosum
represented the entire burrow with “the rugose surface . . .
caused by the irregularly thrown up mud; the flexuous
stem and the branching, by the windings of the worm.”
He mentioned that he had seen similar burrows on the
banks of the Little Miami River. P. tubulare was merely
a P. rugosum with the roof of the burrow removed (i.e., a
concave epirelief or hyporelief). However, the restudy of
the type specimens of the two species indicates that they
are both preserved as convex hyporeliefs, and thus James’
explanation is incorrect. In 1891 J. F. James listed P. vir-

‘TRACE FOSSILS CINCINNATI AREA: Oscoop SWHE

gatum, P. tubulare, and P. rugosum under the generic head-
ing Planolites, but gave no explanation. Because J. F. James
never figured any of these species, it is difficult to corre-
late his names and brief descriptions with the specimens
which actually occur in the Cincinnatian section. Moreover,
as shown above, the various “species” of Palaeophycus have
little meaning at this time, and nothing would be gained
from assigning Cincinnatian forms to previously existing
species of the genus. For this reason Cincinnatian material
is informally discussed below as Palaeophycus types A, B,
and C. Comparisons with various “species” of Palaeophy-
cus are drawn where possible.

Palaeophycus, type-A
Plate 76, figure 5

Discussion. — Palaeophycus, type-A (Pl. 76, fig. 5),
is known only from the Corryville beds of O’Bannion
Creek, Clermont County, Ohio. The form occurs as a con-
vex hyporelief in a well-bedded, medium-grained calcaren-
ite and consists of cylindrical tunnel fillings which are both
parallel and oblique to the planes of stratification. The
cylinders are 3-6 mm in diameter and attain lengths up to
19 cm. The tunnels are distinctly annulate and bear 3-5
annuli/cm; they show no other surface markings. The forms
are unbranched but interpenetration and crossing over are
common. P., type-A is either straight or gently curved in
outline, and in places the burrows appear to dip below
the surface of the host rock, sometimes rising again after
a few centimeters. They are not found preserved as either
convex or concave epireliefs. Vertical sections reveal a dis-
turbance of the bedding where the tunnels penetrate the
host rock, and the composition and grain size of the tunnel
filling is no different from that of the host. P. type-A
appears to differ from all known species of Palaeophycus in
the possession of annuli, In general appearance it resembles
the illustrations of Planolites annularius Walcott (1889)
(see Walcott, 1890, pl. 60, fig. 5) from the Lower Cambrian
near Greenwich, New York, and Planolites corrugatus Wal-
cott (1899, pl. 24, fig. 8) from the Precambrian Greyson
Shale near Neihart, Montana. However, examination of the
type specimen reveals that P. corrugatus (Pl. 77, fig. 2;
USNM 33796) is not annulate as Walcott believed, and the
“annuli” probably represent the cleavage planes of the
slate. P. annularius (Pl. 77, fig. 3; USNM 18360) is badly
weathered but does appear to be annulate, As nearly as can
be determined, both specimens are stuffed burrows, and
thus according to the definition employed herein are plano-
litids.

Palaeophycus, type-B

Plate 76, figure 9; Plate 77, figure 6

Discussion. —The forms grouped under P., type-B,
are not common; single specimens are known from the Mt.
Hope beds on U.S. Route 50, 0.5 miles west of Aurora,
Indiana, and the Corryville beds at Stonelick Creek. P.,
type-B, combines the longitudinal striae of Palaeophycus
striatus Hall with the annulations of P., type-A. These
cylindrical forms are gently curved and are preserved out-
wardly as convex hyporeliefs. The diameter varies from
5-10 mm; the longest form is 21 cm in length. There are
4-5 annuli/em and the striae appear as fine wavy lines.
As seen in Plate 77, figure 6, the annuli may be better de-
veloped in one area than in another. In both specimens
where this is true, the weakly annulate portion of the
cylinder has a greater diameter than the strongly annulate
section. Al] four specimens extend up into the host ro~
with a resulting disturbance of the bedding. As in P., type-A
the grain size and composition of the tunnel filling is simi-
lar to that of the host rock.

Palaeophycus, type-C
Plate 77, figure 5; Pl. 83, figures 1,4

Discussion. — P., type-C, is the most common, yet the
most nondescript variety of Palaeophycus. It occurs in
great numbers in calcarenites whose coarse grains do not
permit the preservation of fine details. If the grain size
were smaller, it is possible that sculpturing might be evi-
dent. P., type-C, is common in Eden strata but relatively
rare higher in the Cincinnatian section. It is especially
well developed in the Lower Eden near the mouth of
Brierley Creek, 3.5 meters downstream from its crossin-
by Taylor Creek Road, Hamilton County, Ohio.

The diameter of the burrows varies from 5-12 mm, and
incomplete forms have a length of 10 cm. In the same bed
P., type-C can be preserved both as a convex hyporelief
(PI. 77, fig. 5) and a more poorly defined convex or concave
epirelief (PI. 83, fig. 4). None of the specimens were ob-
served to branch, although they frequently interpenetrate.
At the Brierley Creek locality the specimens occur in a
coarse-grained calcarenite that is 2-5 cm thick. A parallel
arrangement of shell fragments gives the rock a poorly
defined bedding. There is a sharp boundary separating the
calcarenite from the overlying lutite, and inspection of this
lutite revealed several obliquely-directed, cylindrical tun-
nels which angle down toward the calearenite, Although
none of these tunnels could be traced directly into the tun-

we
~sI
oS

nels of the calcarenite, the evidence is suggestive. As in
the other varieties of Palaeophycus discussed herein, cross-
sections of P., type-C, indicate that the bedding of the
host rock is disturbed where penetrated by the tunnels.

When preserved as an epirelief, P., type-C, closely re-
sembles trails found in the McMicken beds at West Fork
Creek, Cincinnati, Ohio. The difference is largely one of
interpretation and is discussed herein under “?Mollusk
Urals:

Interpretation. — The ethological interpretation of Pal-
aeophycus has already been discussed and will only be
briefly repeated here. As shown by the disturbance of the
bedding in the host rock, all three varieties discussed above
are of infaunal origin. Most likely they are the burrows of
either selective feeders or predators. The fact that there
is no “master plan” or pattern to the structures suggests
the latter, but this is no sure criterion. There is no evidence
that they are the stuffed burrows of sediment feeders. The
annulate nature of Palaeophycus, types A and B, suggest
the peristaltic movements of an elongated annelid-like body,
while the longtitudinal striae of P., type-B, could be the
impressions of setae which occur on the ventral surfaces of
many polychaetes and other vermiform organisms.

Although such forms are difficult to study and charac-
terize because they are obliquely oriented and cut across
bedding planes in addition to occurring in coarse-grained
sediment, their importance should not be underestimated.
In reality they were probably more abundant members of
the infauna than were the more definitive and impressive
forms, such as Phycodes.

WALCOTTIA Miller and Dyer, 1878
Plate 67, figure 6; Pl. 69, figure 5

1878. Walcottia Miller and Dyer, Cincinnati Soc. Nat. Hist., Jour.,
vol. 1, p. 39, pl. 2, figs. 11,11a.

1878. ?Walcottia Miller and Dyer, Contributions to Palaeontology,
No. 2 (Cincinnati, Ohio, private publication), p. 11, pl. 3,
figs. 12,12a.

1881. ?Walcottia Miller and Dyer, James U. P., The Paleontologist,
No. 5, p. 44.

1886. Partim Walcottia Miller and Dyer, James, J. F., Cincinnati
Soc. Nat. Hist., Jour., vol. 8, pp. 161,162.

1962. Partim Walcottia Miller and Dyer, Hantzschel, Trace-Fossils
and Problematica im Treatise on Invertebrate Paleont., Moore
(ed.), pt. W, Miscellanea, p. W243.

Type species.—Walcottia rugosa Miller and Dyer
(1878) (Pl. 67, fig. 6; Pl. 69, fig. 5; MCZ 545; latex mold
UCM 37677) from “near the top of the hills at Cincinnati”
(?Corryville).

Diagnosis. — Slender vermiform trace fossils preserved
as convex hyporeliefs which possess obliquely directed paired

PALAEONTOGRAPHICA AMERICANA (VI, 41)

lobes. The closely spaced lobes are ridgelike and tend to
obscure the body. The form tapers at one end and termin-
ates at the other in a structureless ovoid mass. Other
forms of the genus are reputedly marked by a shell-like
covering composed of small plates or rings, or by a longi-
tudinal groove from which issue five oblique striae.

Discussion. — Miller and Dyer (1878a, p. 16) estab-
lished Walcottia for “. . . long, tapering, rugose, flexuous
bodies, worm-like in form ... [which] come to a point at
one end and are enlarged at the other. . . .” Originally only
a type species, W. rugosa, was proposed; although in the
second part of the same paper, they questionably assigned
an additional species, W. cookana, to the genus. In 1881
U. P. James added yet another, W. sulcata. The syntypes of
W. rugosa were located in the collections of the Museum
of Comparative Zoology at Harvard and will be discussed
below. While Miller and Dyer remarked that the type speci-
men of ?W. cookana was also in the Dyer collection, the
form could not be found at Harvard, nor is it with the
Miller collection at the Field Museum. Unfortunately the
type specimen must be regarded as lost, and any discussion
will have to be based on Miller and Dyer’s description
and figure. U. P. James never figured W. sulcata and iden-
tified material could not be found with the James collec-
tion at the Field Museum. W. sulcata is discussed herein
under Incertae sedis.

Walcottia rugosa Miller and Dyer
Plate 67, figure 6; Plate 69, figure 5

1878. Walcottia rugosa Miller and Dyer, Cincinnati Soc. Nat. Hist.,
Jour., vol. 1, p. 39, pl. 2, figs. 11,11a.

1886. Walcottia rugosa Miller and Dyer, James, J. F., Cincinnati Soc.
Nat. Hist., Jour., vol. 8, pp. 161,162.

Type locality. —The species is known only from the
two syntype specimens which presumably are from the
McMillan Formation.

Diagnosis. — Slender vermiform trace fossils preserved
as convex hyporeliefs which possess obliquely directed
parallel lobes. The closely spaced lobes are ridgelike and,
where present, tend to obscure the body. The form tapers
at one end and terminates at the other in a structureless
ovoid mass.

Discussion. — Both specimens of W. rugosa occur on
a single slab of fine-grained calcisiltite. The numerous tool
marks associated with them indicate that they are pre-
served as convex hyporeliefs, The form which probably best
typifies the species (PI. 69, fig. 5) is shaped like an open
“S” and has a length of 4 cm and a width of 3 mm. The
most striking feature is the presence of 21 paired lobes.

‘TRACE FOSSILS CINCINNATI AREA: Oscoop 379

The lobes are wide proximally and taper to a point distally.
All are obliquely directed toward the tapered end of the
specimen, and the spacing between them is less than a milli-
meter. The lobes do not extend the entire length of the
specimen but instead terminate 1 cm from the tapering
end. In this area the specimen is marked by a thin ridge
which bears a faintly impressed longitudinal groove. To one
side of the ridge are markings which may be poorly de-
veloped lobes. The other end of the specimen is charac-
terized by an ovoid body, 6 mm long and 4 mm wide. It is
free of lobes and bears no trace of any markings. Both sides
of the specimen are marked by a shallow groove which in
latex molds appears as a ridge. As will be shown later,
these are important for the interpretation of the species

The second specimen ( Pl. 67, fig. 6), 3 mm wide and 1.4
cm long, is located 2.5 cm from the first and is oriented
at right angles to it. There is no evidence of a connection
between the two. At one end is the same ovoid mass de-
scribed in the specimen above. The rest of the body con-
sists of six paired lobes which differ slightly from those in
the other form. Instead of joining the body at the sides, the
lobes join at the midpoint and thus give the appearance
of being imbricate chevron-shaped structures. As in the
first specimen the apex of the “V” formed by the juncture
of the lobes is directed toward the ovoid mass.

Interpretation. — Although Miller and Dyer (1878a,
p. 16) were uncertain whether W, rugosa was a “fucoid” or
was of animal origin, they were inclined to believe that it
was “the long lost borer of Silurian age, which has left so
many holes in the corals and the rocks.” They briefly
described another specimen from Miller’s collection which
“represents a track from one worm hole to another, 114
inches apart. The track curves from one side to the other,
and has every appearance of having been made by such an
object as figure 11 [herein Pl. 69, fig. 5] represents”
(ibid., p. 16). J. F. James (1886, p. 162) rejected Miller
and Dyer’s interpretation and instead believed that W.
rugosa was “the impression of the under side of the flexible
arm of a star-fish.”

It appears that this time Miller and Dyer’s explan-
ation is more plausible than that of James. One of the speci-
mens (PI. 69, fig. 5) closely resembles (and in fact may be)
the cast of an errant polychaete. The body proportions are
similar, and the paired lobes are reminiscent of the para-
podia seen on the Recent Nereis. Vertical sections cut
adjacent to both ends of the longer form and near the en-
larged end of the other show that there is no disturbance
of the bedding. Therefore, W. rugosa is regarded as being
of epifaunal origin. Since the lobes are so well preserved,
it is unlikely that there was much lateral movement in-

volved. It is believed that a swimming annelid or annelid-
like form settled to the bottom and left its imprint. The
depressions on either side of the body represent sediment
that was squeezed up as the organism came to rest. The ani-
mal then arched its body off the substrate and began to bur-
row into the subjacent bed. The ovoid mass would thus
represent the point of penetration. In the shorter of the two
forms it appears that only the anterior portions of the body
came in contact with the bottom and that most of the body
remained off the substrate.

?Walcottia cookana Miller and Dyer

1878. ?Walcottia cookana Miller and Dyer, Contributions to Palaeon-
tology, No. 2 (Cincinnati, Ohio, private publication), p. 11, pl.
oatech Wer

1886. ?Walcottia cookana Miller and Dyer, James, J. F., Cincinnati
Soc. Nat. Hist., Jour., vol. 8, pp. 161,162.

Type locality. — Fairview and McMillan Formations at
Cincinnati, Ohio. The only known figure of the species
is that given by Miller and Dyer (1878b, pl. 3, fig. 12,12a).

Discussion. — Miller and Dyer (1878b, p. 11) pro-
posed ?W. cookana for “. . . a small, nearly round, flexuous
body, having a shell-like covering, upon the outer side,
which is composed of plates or rings. A distinct longitudinal
depression marks the back of the specimen so as to give a
section a somewhat quadrate form.” The specimen, which
they believed to be incomplete, measured 1.8 cm in length
and had a width slightly in excess of 1 mm. The plates
were small, averaging only 2/mm. One end of the speci-
men was larger than the other, and this was regarded by
the authors as the anterior end. The lines separating the
rings or plates are directed adapically. Miller and Dyer also
believed that the rings were imbricate, thus allowing the
organism freedom of movement.

The type specimen could not be located with the
Dyer collection at Harvard, nor is it with Miller’s ma-
terial at the Field Museum. A specimen labeled W. cookana
was located in the James collection at the Field Museum.
Although it somewhat resembles the illustration of the type
there is reason to believe that it is not. Evidence given
elsewhere in this paper demonstrates that the James’ most
probably never saw any of Miller and Dyer’s material. The
identification was probably based on Miller and Dyer’s
description and figures. The specimen in question (UC
52031) is a trace fossil which can be assigned to W. rugosa.
(It is not figured herein.)

Interpretation. — Miller and Dyer believed that the
species was a fossil annelid but only questionably assigned
it to Walcottia. J. F. James (1886, p. 162) evidently never

saw the specimen but thought that it was “. . . the impres-

380 PALAEONTOGRAPHICA AMERICANA (VI, 41)

sion of some portion of a crinoid, either a part of the stem
or one of the fingers.”

With no definite specimens available for study, it is
impossible to make any interpretation of the species. Per-
haps it is similar to W. rugosa and thus possibly is of
annelid origin. Miller and Dyer’ description is also reminis-
cent of Lepidocoleus Faber, a machaeridian which occurs
in the local Cincinnatian rocks. However, Lepidocoleus is
normally a little larger than the specimen of ?W. cookana
described by the authors.

FUCUSOPSIS Vassoevich, 1932
Plate 64, figure 1; Plate 70, figure 1; Plate 71, figure 5

1878. Partim Trichophycus Miller, and Dyer, Contributions to
Palaeontolgy, No. 2, (Cincinnati, Ohio, private publication),
p. 4, pl. 4, fig. 2.

1932. Fucusopsis Vassoevich, Akad., Nauk, USSR, Trudy, Geol. Inst.,
118 Toy Sih WG Ae lec Wee

1959. Fucusopsis Vassoevich, Seilacher, Eclog. Geol. Helvetiae, Bd.
51, p. 1070, Table 2, fig. 30.

1962. Fucusopsis Vassoevich, Hantzschel, Trace-Fossils and Proble-
matica in Treatise on Invertebrate Paleont., Moore (ed.), pt.
W, Miscellanea, p. W194, fig. 120-4.

Type species. — Fucusopsis angulatus Vassoevich
(1932) from the Senonian beds (Cretaceous) of Caucasus,
U.S.S.R.

Diagnosis. — Palaeophycus or Planolites-like burrows
of infaunal origin which lie close enough to the basal sur-
face of the silt beds containing them to produce a charac-
teristic stretching of the sole of the bed.

Discussion. — Fucusopsis was established by Vassoevich
(1932) for “hieroglyphs in the form of tubes.” An illustra-
tion given by Hantzschel (1962, fig. 120-4) shows that the
genus consists of long cylindrical structures which have a
diameter of approximately 1 cm. These penetrate the
bedding of the host rock and commonly cross over and
interpenetrate. According to Seilacher (personal communi-
cation), when the genus is preserved as a convex hyporelief,
it has various modes of expression. It may be visible only
as a series of thin cracks in the sole of the host rock (PI.
70, fig. 1), or it may be preserved as a hemicylindrical
swelling marked by longitudinal striae (Pl. 64, fig. 1).
These two forms occur together and can be intergradational.

Interpretation. — According to Seilacher (1959 and
personal communication), Fucusopsis is a burrow of in-
faunal origin. As the organism burrowed a few millimeters
above the bottom of the silt bed, the pressure of the body
produced sufficient tension in the sole of the bed to cause
a series of faults. Ethologically this is of little significance,
but it does show that the host sediment was strong enough
to fault rather than flow. On the other hand, when the

organism worked along the contact of the host silt and the
subjacent mud, the specimens are characterized by a com-
plex series of longitudinal striae which probably are indica-
tive of the digging action of bristles or dactyls. It may be
argued that Fucusopsis should not be retained as a separate
genus, since ethologically it differs little from Palaeophycus
and Planolites. However, Palaeophycus is badly in need of
restudy, and little would be gained by placing Fucusopsis in
synonymy with Palaeophycus at the present time. More-
over, the fact remains that Fucusopsis does tell us some-
thing of the properties of the host sediment.

It is doubtful that the forms described by Birken-
majer (1959) from the Danian-Paleocene Flysch of the
central Carpathians belong in Fucusopsis. In the English
summary the author makes no mention of the faulting pat-
tern which distinguishes the genus. Birkenmajer concluded
that the Polish specimens were the burrows of sediment-
eating annelids working along the sand-mud interface which
were filled by the overlying sand (and ? fecal material).
This interpretation would place them in Planolites.

CINCINNATIAN REPRESENTATIVES

Fucusopsis sulcatum (Miller and Dyer)
Plate 64, figure 1; Plate 70, figure 1; Plate 71, figure 5

1878. Trichophycus sulcatum Miller and Dyer, Contributions to
Palaeontology, No. 2 (Cincinnati, Ohio, private publication),
pp. 4,5, pl. 4, fig. 2.

1884. (In partim) Trichophycus sulcatum Miller and Dyer, James,
J. F., Cincinnati Soc. Nat. Hist., Jour., vol. 7, p. 131, pl. 6,
fig. 5.

Type locality. —The type specimen (Pl. 71, fig. 5;
HBM 3182; latex mold UCM 37705) is from the Economy
beds, Cincinnati, Ohio.

Diagnosis. — A Palaeophycus-like burrow of infaunal
origin which produces tension faulting in the sole of cal-
cisiltite beds.

Discussion. — Miller and Dyer (1878b, pp. 4,5) de-
scribed F. sulcatum as a “fucoid” which “. . . consists of a
long, round flexuous stem, which has not been noticed to
divide or bifureate. The surface is longitudinally furrowed
[and] ... are not regular in their size, nor in their course.
... The ridges or elevated lines, which separate the furrows
anostomoze in places, thus suddenly cutting short the fur-
rows.” They placed the species within Trichophycus with
some reservations.

The type specimen of F. sulcatum, from “the lower
part of the Cincinnati Group, Cincinnati, Ohio,” is pre-
served both as a cleavage relief and a convex hyporelief.
Viewed as a cleavage relief, the form appears as a shallow

‘TRACE FOSSILS CINCINNATI AREA: Oscoop 381

concave furrow which completly traverses the slab and has
a diameter of 6 mm. On the sole of the slab (PI. 71, fig. 5)
is what Miller and Dyer (1878b, pl. 4, fig. 5) illustrated
as the type. The specimen is 7 mm wide at the edge of the
slab and appears to gradually taper to a point at its other
extremity. However, under favorable lighting a faint linear
bulge in the bedding can be traced from the pointed end of
the fossil to the opposite edge of the slab. The striated
hemicylinder and this convex bulge are located directly
below the concave furrow on the opposite face of the slab.
The central portion of the hemicylinder is marked by a
series of short, sharply defined, approximately parallel
ridges. The basal lamina of the host rock “laps up” on
the sides of the hemicylinder, and this feature alone
differentiates F. sulcatum from Palaeophycus. A vertical
section through the specimen shows that the bedding is not
completely obliterated as it is in Palaeophycus but instead
is bowed down, forming a “microsyncline.”

Two other well-developed forms, as well as six or
seven fragmentary specimens of F. sulcatum, have been
found in the Cincinnatian beds. One of these specimens,
from the Bellevue beds exposed in Hall’s Creek near
Morrow, Ohio, has a length of 16.9 cm and a width of 6 mm.
As viewed in Plate 70, figure 1, the upper part of the form
is hemicylindrical and bears the longitudinal striae of the
type specimen. The path of the burrow may be traced
across the slab by the series of ragged faults occurring in
the basal lamina of the host rock. The other specimen, from
an unknown locality, (Pl. 64, fig. 1) more closely re-
sembles the type specimen. It has a length of 13.5 cm and a
width of 6 mm and plainly shows the knifelike anastomosing
ridges. As in the type specimen the basal lamina of the
host rock laps up on the sides of the hemicylinder.

Interpretation. — J. F. James (1884) believed that both
Trichophycus (Fucusopsis) sulcatum and Trichophycus
lanosum were the casts of rill marks. As shown elsewhere,
T. lanoswm is a sinuous burrow marked by a button-like
ending from which radiate fine striae. On the other hand,
F. sulcatum is a Palaeophycus-like burrow which was ex-
cavated extremely close to the base of the calcisiltitic host
rock. The pressure of the body produced a series of faults
in the basal lamina of the host rock, although occasionally
the organism broke through the base of the bed to give
rise to the longtitudinal striae.

?>MOLLUSK TRAILS
Pl. 79, fig. 4; Text figs. 26,27

Discussion. —‘This section is concerned with a number
of long, gently arcuate trails which are preserved as convex

and concave epireliefs in coarse-grained calcarenites. The
single exposure occurs in the lower part of the McMillan
beds in West Fork Creek, approximately 25 meters east
(downstream) of the intersection of Shepherds Road and
West Fork Road. A dam in the stream has left a surface
of about 25 square meters relatively free of debris, and the
trails are remarkably well exposed. The host rock is a
thick coarse-grained calcarenite composed of fossil frag-
ments.** The upper 2 mm of the bed is slightly finer grained
and tends to spall off. There is a sharp contact between the
calcarenite and the thin bed (1-2 cm) of overlying lutite.
Approximately 40 per cent of the surface of the calcarenite
bed is marked by a series of irregular pararipples which
have a maximum amplitude of 9 cm. The remainder of the
bed, where most of the trails occur, has a gently undulat-
ing surface, and the relief does not exceed 3-4 cm. The
trails do not appear to be affected by the irregularity of
the surface. Upon encountering a depression, the trail does
not deviate or end abruptly but instead proceeds down one
side of the depression and up the other; the same is true for
high areas,

To take maximum advantage of this excellent expos-
ure, a grid was prepared, and some 17 square meters of
the surface were mapped, employing a scale of 1/20 (see
Text-fig. 26). As can be seen from the map, most of the
trails are gently curved and their length varies consider-
ably. While some extend for only a few centimeters before
fading out, others are much longer. The longest trail (Text-
fig. 26, center left) is in the form of a “U” and has a length
of nearly 2 meters. Although there is no over-all preferred
orientation, the trails do exhibit some degree of parallel
alignment in small areas. The trails frequently cross over
one another and some branch, generally at an acute angle.

The pattern seen in the map is not thought to be a
complete reconstruction of the trail network. Many of the
trails appear to dip into the surface of the slab and thus
are lost to view. In addition, several others are faint and
have been affected by Recent stream erosion. These two
factors account for the numerous discontinuities which
will be discussed in greater detail below.

The width of the trails ranges from 5-10 mm but re-
mains constant in a single trail, Several different varieties
are visible on the calcarenite surface and are best studied in
cross-section. Some of the trails exhibit a change in mor-
phology along their length. One form (PI. 79, fig. 4; Text-
fig. 27A) begins as a hemicylindrical depression, only a

“The calearenite was so thick that it was not possible to collect any
specimens.

PALAEONTOGRAPHICA AMERICANA (YI, 41)

“ta

yo All
: l

ony

7
we | ba

igure 26 pee i Rate a Map of trails exposed in the

os
Mc ee beds of) V eek, Cincinnati, Ohio. Many of the

trails dip below shes surface sted ng the overall pattern a disjointed
appearance. The dark, irregular patches within the map area repre-
sent eroded areas,

‘TRACE FOSSILS CINCINNATI AREA: Oscoop 383

A

Text-figure 27.— ?Mollusk trails. The diagrams (A-E) illustrate
cross-sections of the trails from the McMicken beds of West Fork
Creek, Mt. Airy Forest, Cincinnati, Ohio. The trails resemble those
forms placed in Palacophycus, however the trails here under discus-
sion were made on or just below the depositional interface; X 1.

few millimeters deep and flanked by a pair of longitudinal
ridges which in turn are flanked by two shallow depressions.
After proceeding in this manner for 10 cm, the trail gradu-
ally converts into a ridge the crest of which is marked by a
small V-shaped longitudinal rift (Text-fig. 27B). On either
side of the ridge are the two shallow depressions described
above. This type of structure can be followed for approxi-
mately 20 cm, and then the trail reverts to the hemicylin-
drical depression (see Text-fig. 27A). Another variation is
similar to the combination discussed above except that the
ridge lacks the median rift; several of these were observed.
The most common trail is either a hemicylindrical depres-
sion with a ridge on either side (Text-fig. 27E) or a ridge
with a shallow depression on either side (Text-fig. 27D).
One last type, which is not common, differs from the others
as it is a shallow flat trough with two small ridges on either
side.

Interpretation. — Undoubtedly these are trace fossils.
The branching, crossing over, and lack of a definite pattern

rule out a vegetable or inorganic origin. They somewhat
resemble Archaeonassa Fenton and Fenton (1937) which
are preserved as concave epireliefs in the Lower Cambrian
of British Columbia. These forms, which were interpreted
as gastropod trails, differ from the Cincinnatian specimens
in that they possess “rounded to subangular wrinkles which
are convex in the anterior direction” (Fenton and Fenton,
1937, p. 454). Likewise, the West Fork Creek specimens are
quite similar to Palaeophycus, type-C, and the difference is
largely one of interpretation. As shown elsewhere, Palaco-
phycus is thought to be of infaunal origin, 7.e., made at
some depth below the depositional interface. In contrast, it
is concluded that these trails were made on or just below
the depositional interface. Many of them are long, and, as
mentioned, they follow the irregular contours of the sur-
face. Two trails were observed to progress down into the
trough of a pararipple, a vertical distance of about 9 cm.

The variation in the morphology of the trails suggests
a difference in the type of movement. It is believed that
in some cases (Text-fig. 27A,E) the animals ploughed along
the surface forcing the sediment to either side. On the other
hand, the hemicylindrical ridges (Text-fig. 27B,C,D) sug-
gest that the organism burrowed in a molelike fashion para-
llel to and just below the depositional interface. The de-
pressions flanking the ridges may represent the areas where
sediment had been drawn to the ridge.

Because no organisms have been found directly associ-
ated with the trails, the selection of the trail-maker is
speculative. However, a comparison with the trails of the
Recent sand snail Littorima littorea (L.), illustrated by
Hantzschel (1941, pls. 1,2), strongly suggests gastropods.
Moreover, a number of different species occur in the Mc-
Micken. Such forms as Archinacella patelliforme (Hall),
Hormotoma gracilis (Hall), Lophospira bicincta (Hall), L.
tropidophora (Whiteaves), Sinwtes cancellatus (Hall), and
S. globularis (Miller and Faber) could have made the trail.
Pelecypods are possibility, but McMicken pelecypods are the
same as those that occur in the Economy beds, and it has
already been shown (see Lockeia) that these were most
likely burrowers. Of course, there is always the possibility
that one is dealing with the trail of an unknown soft-bodied
form.

Hintzschel’s (1941) observations on the behavior of
Littorina on the German Wattenmeer offer a possible
explanation for the branching in the West Fork Creek
trails. He found that as many as seven gastropods would
move along in a single track, one behind the other. Appar-
ently the leader was “breaking trail,” and the others found
it easier to move in a pre-existing track. However, at times

384 PaLaronrocrarnicA AmERICANA (VI, 41)

they would strike off on their own, thus giving rise to a
branching trail. It is not unreasonable to tentatively extend
this explanation to the Cincinnatian forms.

Because the trails are long, gently curved, and lack
any peculiar geometric pattern, they are classified under
Repichnia. However, it should be pointed out that they
might well represent feeding trails where the organism was
feeding on surface detritus.

? PASCICHNIA

?PALEODICTYON Savi and Meneghini, 1850

Plate 67, figure 1; Plate 69, figures 6-7; Plate 81, figure 2;
Text-figure 29

1850. ?Palecodictyon Savi and Meneghini, in appendix to Murchison;
Memoria sulla Structura geologia del Alpi, p. 484.

1863. ?Glenodictyum Marck, Palaeontog., Bd. 11, pp. 1-8 (fide
Hantzschel, 1962).

1865. Non Palaecodictyon Heer, Die Urwelt der Schweiz, p. 245, pl.
10, fig. 10.

1895. ?Pleurodictyon Fuchs, Akad. Wiss. Wien, math.-nat. Kl.
Denkschr., Bd. 62, pp. 394,396, pl. 6, fig. 1.

1935. ?Palaeodictyon Savi and Meneghini, Abel, Vorzeitliche Lebens-
spuren, p. 24, figs. 11-13, 261B, 263, 264.

1939. Partim ?Palacodictyon Herr, Sacco, Roy. Accad. Torino, Mem.
vol. 69, pp. 267-285, pls. 1-2.

1954. ?Palacodictyon Herr, Seilacher, Deutsch geol. Gesell., Zeitschr.,
Bd. 165, p. 217, fig. 2, no. 16.

1959. ?Palacodictyum Heer, Nowak, Kwartalnik Geol. T. 3, pp.
103-125, pls. 1-6.

1960. ?Palaeodictyon Savi and Meneghini, Goley, Akad. Nauk
USSR, Doklad., T. 134, pp. 175-178.

1962. ?Paleodictyon Savi and Meneghini, Hantzschel, Trace-Fossils
and Problematica im Treatise on Invertebrate Paleont., Moore
(ed.), pt. W, Miscellanea, p. W208, fig. 128-5.

1964. ?Paleodictyon Savi and Meneghini, Vialov, Jour. Sed. Pe-
trology, vol. 34, pp. 664-666, fig. 1.

Type species. — Paleodictyon strozzii Savi and Meneg-
hini from the Lower Tertiary Flysch of Tuscany, Italy.

Diagnosis. — Honeycomb-like networks generally in the
form of hexagonal polygons which are preserved as convex
hyporeliefs, 7.e., a series of ridges arranged in a polygonal
pattern, The diameter of the polygons varies from 1-50 mm
but remains constant in a given specimen..

Discussion. — Paleodictyon is one of the most un-
usual fossil forms and could be properly labeled as a “proble-
maticum.” The genus was originally estabished by Savi and
Meneghini (1850) for what was considered to be a hexa-
gonal algal network from the Lower Tertiary of Italy.
Since its original discovery the genus has been reported
from every period except the Cambrian and has been found
in Alaska, Oklahoma, Italy, Spain, Russia, Germany, Iraq,
Wales, Albania, and Japan. A recent study by Nowak
(1959) on forms from the Cretaceous and Tertiary of the
Carpathian Flysch of Poland has contributed much toward
the understanding of this difficult genus.

The genus is most frequently encountered in fine-
grained clastics, and no marked compositional or textural
differences between the fossil and the host rock have been
observed. However, Seilacher (1962, pl. 2, fig. 4) figured
one form where the network is much darker than the matrix.
Nowak (1959) reported that while most forms have a hexa-
gonal network, individuals with quadrangular, pentagonal,
or septagonal polygons are not uncommon. Superficially
these polygons appear to be nearly geometrically perfect,
but upon close examination Nowak (1959, fig. 6) was able
to reveal several minor irregularities. In some specimens the
polygons may be systematically distorted, ¢.g., elongated
parallel to the current direction.

The diameter of the polygons in the Carpathian speci-
mens varies from 1-50 mm, and although it remains con-
stant within a given specimen, forms having different
diameters may be found in close proximity. According to
Nowak there appears to be no correlation between the
diameter of the polygons and the geologic age of the speci-
men, Likewise, the width of the polygon walls can vary
from specimen to specimen. Normally the width is 1-2 mm,
but von der Marck (1876) (Nowak, 1959, fig. 1) and
Koriba and Miki (1939, pl. 5, fig. 5) illustrated forms where
the thickness of the sides overshadows the polygonal outlines
of the network. Nowak also noted that many of the polygons
are incomplete. These incomplete individuals most frequent-
ly occur along the margins but can also be seen within the
body of the specimen, as shown by Nowak (1959, fig. 1),
Vialov and Golev (1960, pl. 3, fig. 3), and Abel (1935,
tion 12).

Present in the University of Cincinnati collections are
two specimens from the Eocene beds near Trieste, Italy,
which illustrate many of the features noted above. Both
forms occur as convex hyporeliefs in a massive, medium-
grained micaceous quartz arenite. One of the specimens
(Pl. 69, fig. 7) has large hexagonal polygons which are
elongated parallel to the current direction. The length
of the individual sides varies from 0.4-1.0 cm. The length
of the polygons varies from 1.4-1.7 cm, while the width is
1 cm. The outline and dimensions remain constant within
the specimen, and the major variation is that some of the
polygons rise as much as 1.5 mm off the face of the host
rock, while others are barely visible. Associated with
Paleodictyon are irregularly winding hemicylindrical trails
which have the same diameter (1.5 mm) as the sides of the
polygon. These trails and the hexagonal network are com-
posed of a material which is only slightly darker than the
host rock.

The polygons in the other specimen (PI. 66, fig. 1)

TRACE FOSSILS CINCINNATI AREA: OsGoop 38

are much smaller and have a diameter of only 4-5 mm. The
width of the walls is 0.8 mm. While the hexagons are not
notably distorted, the walls are not straight and several
minor irregularities can be seen. However, none of the
features illustrated by Nowak (1959, fig. 6) were noted in
either of the two specimens. The regular pattern of this
second specimen is broken by large gaps, and in some places
isolated stringers extend out from the complex. As in the
larger specimen, the network is only slightly darker in
color than the host rock.

Forms similar to those above have been described
under a variety of names, Several authors have changed
Meneghini’s original spelling to Palaeodictyon or Palaeo-
dictyum. The specimen, illustrated by Fuchs (1895, pl. 6,
fig. 1) from the Eocene of Austria and referred to Pleuro-
dictyon, differs in that it possesses “interrupted nodose
walls” (Vialoy, 1964, p. 666), while as mentioned above
Glenodictyum von der Marck (1863) differs in that the
walls are thick. Hantzschel (1962) considered both of these
to be junior synonyms of Paleodictyon, while Vialoy and
Golev (1960) recognized Plewrodictyon as a separate genus
and considered Glenodictyum to be a subgenus of Paleo-
dictyon. It is not known how consistent or important the
supposed differentiating features are.

As Hantzschel (1962, p. 210) pointed out, Palaeo-
dictyon Heer (1865) from the Flysch of Switzerland is not
synonymous with Palaeodictyon Meneghini. The former
is a burrow consisting of several branched loops which are
connected by Spreite-like arcs (see Hantzschel, 1962, fig.
129-6a) and is placed by Hantzschel in Phycosiphon
Fischer-Ooster (1858). It certainly bears no similarity to
Paleodictyon.

At the specific level the picture is one of confusion,
as over 30 species have been named. Nowak (1959, p. 125)
employed a quasi-zoological nomenclature and recommended
delimiting “species” on the size and configuration of the
polygons. No attempt to evaluate the various species will
be made in this work.

Interpretation. — The enigmatic nature of Paleo-
dictyon is reflected in the variety of interpretations that
have been applied to it.** The genus has been variously
interpreted as mudcracks, raindrop imprints, and the result
of gas percolated up through the sediment. Sacco (1939),
who studied Paleodictyon at various times over a 50-year
period, finally concluded that it represented interference
ripple marks. Earlier he had been an algal adherent. The

“See Sacco (1939) and Vialov (1964) for a more complete dis-
cussion.

On

organic interpretations also have run the full gamut. It has
been believed to represent fossil honeycombs, sponges or
corals, algae, strings of eggs, gastropod larvae, tadpole nests,
and traces of invertebrate activity. Those holding to a
trace fossil origin are Fuchs (1895), Abel (1935), Sei-
lacher (1954, 1955, 1962), Nowak (1959), and Vialov and
Golev (1960).

Newak (1959) based his trace fossil assignment on
three features: the irregularities of the polygons, the num-
ber of unclosed polygons, and the fact that polygons with
varying numbers of sides may occur in the same specimen.
Nowak (1959) and others have pointed out that Paleo-
dictyon may have developed from a Beloraphe-like trail.
Beloraphe Fuchs (1895), a Mesozoic form, resembles a series
of zigzags with short projections at the angles. Two of
these placed side by side give the polygonal appearance
of Paleodictyon (Nowak, 1959, fig. 4). Further evidence is
that Beloraphe and Paleodictyon do occur together, al-
though they have not been found in direct association.
Although Seilacher has not conducted a detailed study of
the genus, he (1955) placed Paleodictyon within the Pas-
cichnia. Likewise, Nowak (1959) concluded that the form
was the feeding trace of an annelid.

There is disagreement as to whether the trail is of
epifaunal or infaunal origin. Seilacher (1962) held Paleo-
dictyon to be infaunal, while Vialov and Goley (1960)
concluded that most forms were epifaunal. Nowak (1959)
believed that both conclusions were valid.

Silvestri (1911) and Koriba and Miki (1939) pointed
out the similarity between Paleodictyon and the Recent
fresh water alga Hydrodictyon utriculatum (Roth). The
cenobiae of this species form a net of pentagonal, hexagonal,
or septagonal polygons. Koriba and Miki (1939, p. 61) re-
ported that triangular and octagonal forms are also known.
Moreover, they maintained that while the plant may attain
a length of one meter in which the polygons are several
centimeters wide, most forms have measurements which
conform to those of Paleodictyon.

CINCINNATIAN REPRESENTATIVES

?Paleodictyon sp.
Plate 67, figure 1; Plate 81, figure 2

Only two small fragmentary specimens with a_hex-
agonal network similar to that of the European Paleo-
dictyon have been found in the Cincinnati strata. These are
placed in Paleodictyon with some hesitation, since there is
a possibility that both specimens are of inorganic origin.

One specimen is from the Economy beds of Duck

386 PALAEONTOGRAPHICA AMERICANA (VI, 41)

Creek, near Silver Grove, Kentucky, 1,010 meters south-
west of the intersection of Duck Creek Road and Kentucky
Route 8. The outcrop is located on the southeast bank, 20
meters below the bridge crossing the creek. The other form
is from the McMillan beds on Ravine Street in Cincinnati.
Both localities have been recollected without success. The
two specimens are preserved as convex hyporeliefs in cal-
cisiltites.

The Duck Creek form (PI. 81, fig. 2) is poorly pre-
served and consists of nine incomplete hexagons which have
an internal diameter of 3 mm. The greatest thickness of the
walls of the polygon is 1 mm. As indicated by the numerous
tool marks, the specimen is elongated parallel to the current
direction. The relationship between the network and the
tool marks is unclear, but the former does not seem to be
truncated, There is no apparent difference in composition
between the network and the host rock.

The Ravine Street specimen (PI. 67, fig. 1) is better
developed. It is composed of 25 polygons with an internal
diameter of 2 mm. Although the pattern is more regular
than most specimens of Palaeodictyon, the network varies
in its clarity. As viewed in Plate 67, figure 1, the polygons
in the center stand out in bold relief, while those on the
right- and left-hand margins are barely visible. Likewise,
all of the polygons along the lower margin are incomplete.
As in the specimen from Duck Creek the network is elon-
gated parallel to the current direction. There is a consis-
tent variation in the thickness of the polygon walls; the
walls normal to the current are thicker (0.6 mm) than
those parallel to the current (0.3 mm). This specimen differs
from the two Trieste forms (Pl. 66, fig. 1; Pl. 69, fig. 7), as
the network of the Cincinnatian specimen blends into the
background of the host. This is due in part to the litho-
logic similarity of the host rock and the network.

Interpretation.—It is not possible to determine
whether the two specimens under consideration actually
represent Paleodictyon or instead are of physical origin.
While it is clear that the forms are not interference ripple
marks or fillings of mud cracks, as some authors have sug-
gested for Paleodictyon, it is possible that they are tool
marks. As noted above, both forms are elongated parallel
to the current direction and are associated with tool marks
(groove casts). Moreover, there is at least one fossil present
in the Cincinnatian strata which could produce such im-
prints as it rolled over the bottom. This is the problematic
genus Pasceolus Billings which is most frequently placed in
the receptaculitids. Pasceolus darwint Miller (1874), de-
scribed from the Bellevue beds near Maysville, Kentucky,
approximatey 60 miles southeast of Cincinnati, has a dis-

coid or globular body which measures up to 4 cm in diameter.
The surface is marked by hexagonal cups whose dimensions
conform to those of the polygons of the two ?Paleodictyon
networks. Specimens of Pasceolus were rolled over a plaster-
of-Paris surface to test the validity of this hypothesis. The
results (Pl. 69, fig. 6) demonstrate that under proper con-
ditions a globular specimen of Pasceolus being tumbled along
the bottom can produce the “tire-tread” imprints here under
discussion. However, to form such markings it is necessary
that the rolling specimen just nick the surface of the sub-
strate. If the sediment is too soft or the specimen too
deeply impressed, the polygons are superimposed on a
shallow ellipsoidal depressions which is not present in the
Cincinnatian specimens of ?Paleodictyon. This hypothesis
also explains the unequal development of the walls in the
Ravine Street specimen, since the walls of Pasceolus,
oriented normal to the current, would cut a wider marking
than those parallel to the current.

The major problem with this explanation is that, al-
though Pasceolus darwini occurs in large numbers at the
type locality in Maysville, it is rare in the Cincinnati area.
It has not been found at either of the two localities where
?Paleodictyon occurs and has never been reported from the
Eden strata. It is thus possible that at least the Duck
Creek specimen is a valid Paleodictyon.

While the biological affinities of Paleodictyon must
still be considered an open question, the bulk of the evi-
dence favors either a vegetable or trace fossil origin. The
over-all regularity in the pattern and size of the polygons
in a given form does suggest an algal network similar to
the Recent Hydrodictyon. However, no organic material
has ever been found in Paleodictyon, and the type of preser-
vation (convex hyporelief) is the one most frequently
exhibited by trace fossils. Also Paleodictyon is not uncom-
monly associated with trace fossils which show unusual
geometric patterns, e.g., Beloraphe and Spirodictyon (Abel,
1935, figs. 263,264). Moreover, one of the specimens from
Trieste figured herein (PI. 69, fig. 7) is judged to be most
significant, since it bears gently arcuate trails which have
the same diameter as the hexagonal network. This suggests
two behavioral variations of the same organism. This evi-
dence indicates that Paleodictyon is a trace fossil, probably
a Pascichnia. It exhibits the same characteristics as the
spiral forms with which it is associated, e.g., a maximum
coverage of area. Admittedly it is difficult to visualize how
a burrowing organism could evolve such an intricate pattern,
but perhaps this is not so surprising when one recalls
such unusual forms as Daimonhelix and Helminthoida
(Hantzschel, 1962, figs. 121-9 and 122-5).

TRACE FOSSILS CINCINNATI AREA: OsGoop 387

BODY FOSSILS DESCRIBED AS PLANTS

MASTIGOGRAPTUS Ruedemann, 1908

Plate 79, figure 3

1858. Non Dendrograptus Hall, Geol. Survey Canada for 1857, Rept.
Prog., p. 143.

1878. Partim Psilophyton, Lesquereux, American Phil. Soc., Proc.,
vol. 17, p. 164, pl. 1, fig. 2.

1883. Partim Dendrograptus Hall, Walcott, Albany Inst., Trans.,
vol. 10, p. 21.

1885. Partim Dendrograptus Hall, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 8, pp. 160,161.

1908. Mastigograptus Ruedemann, New York State Museum, Mem.,
No. 11, pp. 210-223, figs. 116-118.

Type species. — Dendrograptus tenuiramosum Walcott
(1883) from the “Utica Slate” (Upper Ordovician), Tren-
ton, Oneida County, New York.

Diagnosis. — Dendroid rhabdosomes which may attain
a length of over 20 cm. Individual branches slender
(0.1-0.8 mm). Well-preserved specimens possess irregularly
placed conical thecae, while in more poorly preserved forms
the thecae are detached and the branches appear to be
smooth.

Discussion. — Mastigograptus is most commonly found
as relatively small “monopodial” branching fragments whose
black color makes them stand out against the lighter color
of the host rock. Usually the slender branches bear no trace
of any structures, although some may show a series of
depressed conical pits (see Ruedemann, 1908, pl. 12, figs.
1-6). In addition, one specimen of M. gracillimum (Les-
quereux) was seen “. .. to contract and expand regularly
about eight times in 10 mm from 2 mm to 2.5 mm, pro-
ducing an apparent jointed structure” (Ruedemann, 1908,
p. 221). The only specimens showing the long conical thecae
are those of the type species /. tenutramosum (Walcott,
1883). Thecal-bearing specimens have been recovered both
from the type locality and from the Eden beds of Coving-
ton, Kentucky. Ruedemann believed that the point of junc-
ture of the thecae and the branch was extremely fragile and
that in most cases the thecae have become detached, He
remarked that “great numbers of such bodies have been
[found loose] on the Covington slabs” (ibid, p. 212).

The purpose here is not to completely discuss the
graptolite genus and its relationships. Instead attention will
be focused on one species which is most probably con-
specific with what Lesquereux (1878) described as a “Sil-
urian Land Plant.” Therefore, the reader is referred to
Ruedemann (1908, pp. 210-223) for a more complete
treatment of Mastigograptus and of the Lesquereux error.

Mastigograptus gracillimum (Lesquereux)

Plate 79, figure 3

1878. Psilophytum gracillimum Lesquereux, American Phil. Soc.,
Proc., vol. 17, p. 164.
1883. Dendrograptus gracillimum (Lesquereux), Walcott, Albany

Inst., Trans., vol. 10, p. 21.

1885. Partim Dendrograptus gracillimum (Lesquereux), James, Vin Uy
Cincinnati Soc. Nat. Hist., Jour., vol. 8, pp. 160-161.

1908. Mastigograptus gracillimus (Lesquereux), Ruedemann, New
York State Museum, Mem., No. 11, pp. 219-221, figs. 116-118.

Type locality.— The type specimen, which was un-
available for study, may be in the collections of the Field
Museum; it was taken from Eden strata in the bed of the
Licking River, Covington, Kentucky.

Diagnosis. — Moderate-sized dendroid rhabdosomes
with thin (0.8 mm) branches. Surface normally smooth, but

one specimen shows an apparent jointed structure. Nature
of thecae unknown.

Discussion. — Lesquereux (1878, p. 164) characterized
M. gracillimum as “stem very slender, dichotomously
branching, smooth or naked half round, slightly channeled
in the length; branches numerous, of various length, filiform.
The stem is scarcely one millimeter thick at the base; the
upper branches curved as from a spiral unfolding, are slender,
gradually attenuated and capilliform, or of the thickness of
thin thread at their extremities.”

In the course of his research on the fossils of the “Utica
Slate,’ Walcott (1883) studied specimens from Cincinnati
which had been identified by Miller as Psilophyton gracil-
liimum. Walcott placed them in Dendrograptus Hall but
did not describe the species in detail.

J. F. James (1885, p. 160) also placed the forms in
Dendrograptus, and at the same time he declared that MW.
gracllimum “precisely resembles in its essential features
the figure of Buthotrephis gracilis given by Hall.” For this
reason he likewise referred Buthotrephis gracilis to Den-
drograptus. Although he gave no specific reference to any of
Hall’s plates, he was most likely comparing Lesquereux’s
figure to that of Hall (1852, pl. 5, figs. la-c). As shown
elsewhere in this work, B. gracilis is not comparable to
Dendrograptus or Mastigograptus and instead is a chond-
ritid. There is no evidence to indicate that James ever saw
Lesquereux’s type specimen; most probably he was working
entirely from the literature.

Ruedemann (1908) studied two specimens of topotype
material obtained from Ulrich. He also mentioned that the
type specimen of M. graciulimum, which was collected by
Ulrich, was loaned to a Reverend Herzer of Cincinnati, who
in turn loaned it to Lesquereux. Apparently the type was

388 PALAEONTOGRAPHICA AMERICANA (VI, 41)

then sold with the Herzer collection to James Hall. Ruede-
mann further remarked that the specimen was probably
sold again, this time to the University of Chicago with a
portion of the Hall collection. He too was unable to obtain
it for study.

Ruedemann (1908) described the “stems,” which have
a diameter of 0.8+- mm, as being solid and showing few
effects of compression. The “stem” of one specimen was
smooth, that of the other regularly expanded and con-
tracted to give the jointed appearance mentioned above.
He said that the species was closely allied to M. tenwira-
mosum (Walcott), although it differed in that the secondary
branches were a little thicker and “the branches of a higher
order are here arranged a little closer than in M. tenwira-
mosum’”’ (Ruedemann, 1908, p. 221). He mentioned that
more complete collections of the two species might result in
synonymy.

Unlabeled material (Pl. 79, fig. 3) has been found in
the collections of the University of Cincinnati which closely
resembles the descriptions given by Lesquereux and Ruede-
mann. The blue lutite contains several black fragments,
the longest of which is 2.5 cm. The main “stem” is 0.7 mm
in diameter, while the secondary and tertiary branches have
a diameter of 0.5 mm. The specimens appear to be solid,
and the only signs of compaction are faint longitudinal
wrinkles which may be analogous to the longitudinal chan-
nels of Lesquereux. None of the specimens possess the
jointed structure noted by Ruedemann, and no_ thecae
were found associated with them.

Interpretation. — Although Lesquereux (1878) orig-
inally placed M. gracillimum in the Lycopodes, 1.e., land
plants, there has been general agreement since then that the
species is a graptolite. If the specimens described by Wal-
cott and Ruedemann are the same material as Lesquereux’s
type specimen, it appears that their graptolite assignment
was correct. In general appearance M. gracillimum is ex-
tremely close to M. tenuiramosum (Walcott), where thecae
are found on the branches (Ruedemann, 1908, pl. 11, fig. 1).
Although Ruedemann (1908, p. 221) stated that the jointed
structures of M/. gracillimum might represent thecae, it was
more likely that they were “internodes between the bases
of branches of the third order which are all broken off.”
The finding of undoubted thecae on the branches of M.
gracillimum would affirm its graptolite assignment.

Walcott (1883, p. 21) concluded that “the resemblance
of these two species of Dendrograptus [Mastigograptus]
to Lycopodiaceous plants of the genus Psilophyton is very
striking and apt to mislead the observer. Their occurrence
with algae, graptolites, trilobites and brachiopods in the

same layers of shale, in a position indicating their growth
im situ, taken with their graptolite structure, precludes the
idea of their being other than of marine origin.”

“FUCOIDS” OF INORGANIC ORIGIN*

“CHLOEPHYCUS” Miller and Dyer, 1878
Plate 80, figures 1,6

1847. Non Buthotrephis Hall, Nat. Hist. New York, Palaeont. New
York, vol. 1, p. 8, pl. 2, fig. 6.

1878. Buthotrephis Hall, James, U. P., The Paleontologist, vol. 2,
Da ae

1878. Chloephycus Miller and Dyer, Contributions to Palaeontology,
No. 2 (Cincinnati, Ohio, private publication), p. 3, pl. 4, fig. 1.

1884. Chloephycus Miller and Dyer, James, J. F., Cincinnati Soe.
Nat. Hist., Jour., vol. 7, p. 130, pl. 6, fig. 3.

1885. Partim Buthotrephis Hall, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 159.

1889. Chloephycus Miller and Dyer, Miller, North American Geol.
and Palaeont., p. 113.

1962. Chloephycus Miller and Dyer, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont.,, Moore
(ed.), pt. W, Miscellanea, p. W232.

Type species. —Chloephycus plumosum Miller and
Dyer (Pl. 80, fig. 1; HBM 3154; latex mold UCM 37617)
from the Eden beds of Cincinnati, Ohio.

Discussion. — Miller and Dyer (1878b, p. 3) described
“Chloephycus” as “simple, slender stems which throw off,
upon each side, numerous fine filaments.” According to the
authors bifurcation is never present; and while a single
“stem” resembles a feather, several of them together suggest
grass. They differentiated it from Trichophycus by the
spreading of the “filaments” and by the rigidity and delicate
nature of the “stem.”

Such forms are rare in the Cincinnatian section, as
indicated by the fact that only the type and two other
previously undescribed specimens are avaiable for study.
Both occur on calcisiltites and cross-laminations in both
specimens and load or flute casts on the second one indi-
cate that they are preserved as convex hyporeliefs. The
“type” (Pl. 80, fig. 1) consists of approximately 24 paral-
lel separate “specimens.” The “stems,” which are mostly in-
complete, range from a fraction of a millimeter to 5 mm
in width. The fine “filaments,” which issue from the “stem”
at an angle of 20°-30°, range up to 1 mm in diameter.
Distally the “filaments” grade imperceptibly into the surface
of the bed. The angle of branching is constant, and in all

“The forms included in this chapter are judged to be inorganic. Cor-
respondingly, the names are not valid and are placed in quotation
marks. It should be emphasized that the present author does not ad-
vocate quasi-legal names for inorganic forms.

TRACE FOSSILS CINCINNATI AREA: OsGoop 389

cases the orientation of the “V” formed by the inter-
section of two branches is the same. In almost all examples
the “filaments” are better developed in the upper part of
the specimen, thus obscuring the “stem.” In the lower
portion of the specimen, where the “filaments” are not
so well developed, the “stem” stands out in bold relief.

The specimen located on the reverse of the slab
bearing Trachomatichnus numerosum is similar to the type,
except that the “filaments” are best developed in the mid-
portion and fade out in either direction. Associated with
this form are what are interpreted as load or flute casts
(Pl. 80, fig. 6, right) and an area of flowage-produced
stress patterns (PI. 80, fig. 6, left).

“Chloephycus” is judged to be synonymous with
Buthotrephis filciformis U. P. James (1878). James described
B. filciformis as having a slightly sinuous stem less than 1/12
inch in width, and every 1/12-1/6 inch the stem gives off
slender branches at a 45° angle. This form, which was
collected from the Maysville beds near Lebanon, Ohio, was
never figured and it is doubtful if a type specimen was ever
designated. Nevertheless, a specimen bearing the label
Buthotrephis filciformis in the James collection at the Field
Museum is identical to “Chloephycus”.

Interpretation. — Miller and Dyer (1878) assigned
“Chloephycus” to the “fucoids,” while J. F. James (1884,
p. 7) dismissed the genus with “. . .
than a mark or series of marks produced . . . by the running
of water down a sloping bank into a stream.” Miller (1889)
was evidently convinced by James’ arguments, for although
he listed “Chloephycus” among the algae, he admitted that
it was probably of inorganic origin.

it is nothing more

Evidently “Chloephycus” is not common in the geo-
logic record as the only counterparts reported are from the
Lower Devonian Hiinsruck Shale. One such form was
illustrated by Rudolpf Richter (1941, fig. 15), who inter-
preted it as the cast of the drag marks of a xiphosurid telson.
Since the marking is flanked by tracks, Richter’s interpre-
tation appears to be plausible. Another specimen is men-
tioned by Seilacher (1960, pl. 12, figs. 2-3), who attributed
it to “fillings of pinnatulated drag-marks” (Schezfrillen),
probably originating from the dragging of a starfish arm
along the bottom, According to Seilacher the acute angle
would point downcurrent. A third specimen from the
Hiinsruck is illustrated by Seilacher and Hemleben (1966,
ple2s tie. 1)e

There is no evidence which would point to an algal or
trace fossil origin for the Cincinnatian forms, which are
thought to be modified groove casts originally made on a
mud surface. It is possible that original markings resembled

“Scolithus dispar” (a groove mark) and that the feather-like
pattern resulted from penecontemporaneous flowage of mud
into the linear depression. An alternate hypothesis is that the
structure originated during compaction after the mud
surface had been covered by the calcisiltite bed. The area
to the left of the specimen seen on Plate 80, figure 6 does
show evidence of deformation and flowage, and this may
well have been post-depositional. In either case the direc-
tion of flowage was toward the closed end of the “V.”
Whatever the exact origin, “Chloephycus plumosum” and
“Buthotrephis filciformis” are believed to be nothing more
than modified groove casts.

“BLASTOPHYCUS” Miller and Dyer, 1878
Plate 67, figure 2; Plate 80, figure 2; Plate 81, figures 1,3,5,10

1873. Non Planolites Nicholson, Roy. Soc. London, Proc., vol. 2, p. 289.

1878. Blastophycus Miller and Dyer, Cincinnati Soc. Nat. Hist., Jour.,
vol. Ivps24; pl. 1 digs. 1,2:

1885. Partim Blastophycus Miller and Dyer, James, J. F., Cincinnati
Soc. Nat. Hist., Jour., vol. 8, pp 158-159.

1891. Partim Planolites diadematum James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 14, p. 47.

1962. Blastophycus Miller and Dyer, Hantzschel, Trace-Fossils and
Problematica im Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W187.

Type species. — Blastophycus diadematum Miller and
Dyer (1878) (PI. 80, fig. 2; HBM 3181; latex mold UCM
37618; Pl. 81, fig. 10; HBM 3181; latex mold UCM 37703 )
from the Eden beds, Cincinnati, Ohio.

Description. — “Blastophycus” was described by Miller
and Dyer (1878a, p. 1) as a bilobate “fucoid” “. . .
protuberance or bud-like attachment at the junction of the
branches.” They noted that the protuberance may be
smooth or rugose and believed it to be either a bud, shoot,

with a

frond, or “‘spore.”

J. F. James (1885) compared the “bud-like protuber-
ance” with the “enlarged end” of Trichophycus lanosum
Miller and Dyer (1878) and placed the two in synonymy.
In his listing of 1891 he renamed the form in question
Planolites diadematum and considered it in synonymy with
T. lanosum and Saccophycus intortum U. P. James. —

Fortunately several specimens of “Blastophycus,” in-
cluding those figured by Miller and Dyer, were located
in the paleobotanical collections of Harvard University.
These are preserved as convex hyporeliefs (as indicated by
cross-bedding and associated sole markings) on conspicu-
ously cross-bedded, iron-stained calcisiltites. All of the speci-
mens appear to be from the same bed; although, due to the
lack of data, the type locality has not been rediscovered.

Interpretation. — Miller and Dyer (1878a) interpreted

390 PALAEONTOGRAPHICA AMERICANA (VI, 41)

“Blastophycus” as a “fucoid,’ while J. F. James (1885)
believed it to be a burrow but did not elaborate.

Hantzschel (1962) listed “Blastophycus”’ under trace
fossils and cited Seilacher (personal communication to
Hantzschel) as stating that the “genus” represented “worm
burrows projecting above surface of sediment” (Hantzschel,
1962 sp L87)):

A study of the “syntypes” (PI. 80, fig. 2; Pl. 81, fig.
10) and other specimens shows that the “bud-like protuber-
ances” are in actuality casts of an enrolled trilobite,
apparently Flexicalymene. In many forms the outlines of
the thoracic segments and the cephalic border are plainly
evident. With this in mind, the origin of the two “branches”
is not difficult to perceive. Groove casts and the cross-
laminations reveal that the branches lie in the downstream
sector, What Miller and Dyer referred to as “branches” are
the casts of scour markings around an enrolled trilobite.
Potter and Pettijohn (1964, pls. 91B-93B) have termed
such markings “current crescent casts.”

In testing the hypothesis, “Blastophycus” has been
produced experimentally in the laboratory. A tray was filled
with fine-grained, water-saturated silt, and several enrolled
Flexicalymene specimens were then dropped onto the sur-
face. The tray was tilted at an angle of 10° and a moderately
strong current of water was introduced at the upper end. The
flow of the water around the trilobites produced settling
and also led to the formation of two scour channels on
the downcurrent side (Pl. 81, figs. 1,2). The trilobites were
then removed and plaster casts made (PI. 67, fig. 2; PI. 81,
fig. 5); these casts compare favorably with the original
“Blastophycus.” The scour marks of the original were un-
doubtedly made on a mud surface, as the channels are much
deeper than those produced experimentally. As soon as the
flow of water was terminated in the experimental model,
caving began which ultimately produced a wider, shallower
channel than was the case in the original.

Enrolled trilobites are relatively common in_ the
Cincinnatian lutites and can occur in large numbers.*! Pre-
sumably enrollment reflects a life-reaction,and the lutite
occurrence, a record of rapid entombment. It is postulated
that the trilobites were freed from the lutite by erosion and
rolled along the sea floor until they temporarily came to
rest again. Currents moving past them produced settling
and scouring as described above and utlimately succeeded
in dislodging the trilobites and washing them farther down-

"At the corner of Boudinot and Montana Avenues in Cincinnati, a low
angle exposure slope (area of 1,000 square meters) in the “butter
layers” of the Waynesville beds yielded over 3,000 specimens of
Flexicalymene meeki.

stream. The mud surface was then covered by a calcisiltite
bed. Another possibility is that the current eroded down
to the level of the enrolled trilobites, produced the scour
marks, and finally washed the fossils away but left the
scour marks intact. This latter suggestion is somewhat
analogous to the “pseudoexogenous preservation” hypothe-
sized by Seilacher (1962) for forms from the Spanish Flysch,
where burrows were unroofed, the filling material washed
out, and the mold filled anew with a coarser sediment.

Although Miller and Dyer remarked that “Blasto-
phycus” occurs in the Eden beds, it is more likely that the
form will be encountered higher in the section (Corryville
to Waynesville), where enrolled trilobites are more com-
mon. However, these beds are the favorite focal point of
local collectors, and the fact that no additional specimens
of “Blastophycus” have been reported would seem to indi-
cate that the “genus” is rare. This is unfortunate for crescent
scour casts are valuable in the determination of paleo-
current velocity and direction.

“DYSTACTOPHYCUS” Miller and Dyer, 1878
Plate 70, figures 2,3; Plate 80, figures 3,4; Plate 81, figure 6

1878. Dystactophycus Miller and Dyer, Contributions to Palaeon-
tology, No. 2 (Cincinnati, Ohio, private publication), pp. 2-3, pl.
3, fig. 4.

1885. Dystactophycus Miller and Dyer, James, J. F., Cincinnati Soc.
Nat. Hist., Jour., vol. 7, p. 163.

1962. Dystactophycus Miller and Dyer, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W240.

“Type species.” — Dystactophycus mamillanum Miller
and Dyer (1878) (Pl. 80, fig. 4, HBM 3177; latex mold
UCM 37691) from the Richmond strata near Milford, Ohio.
Monotypie.

Discussion. — Miller and Dyer (1878b, pp. 2,3) charac-
terized the “genus” as a “Rhizoma or frond, mammiform, or
depressed conical, and much expanded at the base. Surface
marked by numerous closely-arranged, concentric ridges or
wrinkles. Some species may be marked, also, by radiating
lines.”

The type specimen of “D. mamuillanum” (PI. 80, fig. 4)
resembles a small truncated cone, having a diameter of 3.5
cm at the base and a height of 1.5 cm. The entire structure
is composed of 15 flattened rings which subscribe an are of
220°. The remaining 140° is either covered by sediment or
has been destroyed. Likewise, the apex of the cone is missing.
As seen in lateral view the specimen is not a perfect cone
but instead is shaped like a “liberty cap.” On the attenu-
lated slope the rings are 1.5 mm wide and well preserved, al-
though they are offset by a small fault. Most of the rings

TRACE FOSSILS CINCINNATI AREA: Oscoop 391

on this slope are arranged in an imbricate pattern where
the larger rings overlap the smaller ones. On casual in-
spection the rings are smooth, but careful study reveals a
series of faint parallel ridges on one small section of one of
the rings near the apex. On the shortened slope of the cone,
the rings are thinner (1 mm), more indistinct, and give the
appearance of having been compressed. Evidently the
authors attempted to clean this area of the specimen, but
it is difficult to determine to what degree it has been
altered. The imbricate arrangement of the rings is not so
noticeable here as it is on the attenulated slope. There is
no trace of crystalline or organic material anywhere on
the form; it is composed entirely of calcisiltite. Likewise,
there is no trace of the “radial lines” which Miller and Dyer
mentioned were present on some specimens.

A specimen (PI. 80, fig. 3) labeled “D. mamuillanum”
from the Richmond (Arnheim) near Hamilton, Ohio, has
been found in the collections of the United States National
Museum. Like the type it is a conical body which has a
diameter of 9.0 cm at the base and a height of 3.0 cm. The
specimen is composed entirely of thinly laminated calcisil-
tite which is bowed down toward the apex; the surface
opposite the apex resembles a saucer-shaped depression.
The concentric rings vary in their morphology. The flattened
rings near the base are 1.5 mm wide. There is no evidence
of imbrication, and the faint parallel lines found in the
type specimen are plainly seen. In contrast, the rings near
the apex show the same imbricate pattern of the type
specimen with the difference that here the smaller rings
appear to overlap the larger.

Interpretation. — Miller and Dyer (1878b) interpreted
“Dystactophycus” as an unbranched “fucoid” and believed
that the broken apex represented an injury to the plant.
However, there is no evidence that would point to a vege-
table origin.

J. F. James (1885) compared “Dystactophycus” with
the cryptostome bryozoan Lichenalia concentrica Hall
(1852, pl. 40F, fig. 5a), originally described from the Clin-
ton (Silurian) beds of New York State. He related that “it
can now be conclusively shown that certain species of Mon-
ticulipora*? have bases which are marked by concentric
lines very similar to those of Lichenalia concentrica, and
there can hardly be a doubt but that they are the same. A
figure of Hall’s species is given in Palaeontology of New
York, Vol. II, plate 40E, which exactly resembles the mark-
ings of Dystactophycus. This “fucoid,” then, has resulted

James’ concept of Monticulipora embraced what today are considered
to be several different genera.

from the impression of the base of a coral [sic] resting on
a mud-bank and leaving its mark in the concentric rings.
The elevated portion is the part extending up into the base
of the coral stem, and the outer rings mark the extent of
the expanded base. Comparison of Hall’s figure and the
specimens of Dystactophycus leave little doubt as to this
fact” (James, 1885, p. 163).

A study of the type specimen of Lichenalia concentrica
(Pl. 80, fig. 5; AMNH 1487/1) shows that it differs from
Hall’s figure. The form is composed of dark calcite and is
preserved as a shallow saucer-shaped depression, 7.¢., con-
cave epirelief. The flattened rings are much smaller (5/mm)
and more numerous than Hall’s figure would indicate. In
addition, the margins of the rings are not smooth as shown
by Hall but instead are crenulate. Both Hall’s figure and
the specimen show that several of the rings either pinch
out or are truncated by others. Where they are not cov-
ered by foreign material, most of the rings of “D. mamillan-
um” appear to be complete. It is judged that James was
misled by Hall’s figure and that L. concentrica bears only
a superficial resemblance to the type specimen of “D. mamil-
lanum.” While certain Cincinnatian bryozoa may have
bases which are similar to Lichenalia and “Dystactophycus,”
James gave no specific examples to support his argument,
and his voluminous works on the genus contain no mention
of any. Moreover, Dr. Richard Boardman and Dr. John
Utgaard of the United States National Museum remark
(personal communication) that they have never seen a
bryozoan base that resembles “Dystactophycus.”

It instead is concluded that the type specimen, as well
as the other specimen of “Dystactophycus” described above,
represent casts of the rotational sweepings of crinoid stems.
The attitude of the bedding in the Hamilton specimen ( PI.
80, fig. 3) shows that the correct orientation of the cone
is “apex down,” and the faint parallel ridges on a portion
of one of the rings of the type specimen could represent
casts of the secondary ridges found on many columnals. It
is believed that the stem was first buried by mud to a
depth equal to the height of the cone, i.e., 1.5 cm in the
type and 3.0 cm in the other specimen. Swirling currents
caused the stem to rotate in circular patterns, producing
a cone-shaped cavity. Since there is no remnant of the
stem or its impression on either specimen, it must have
been plucked out at this time. Where the walls of the cone
were steepest, minor flowage of the mud occurred, result-
ing in the imbricate pattern. Finally, silt-size material
was swept into the cavity thus insuring preservation. The
imbricate pattern also could have resulted from diagenetic
compaction after burial, The asymmetry and the faulting

392 PALAEONTOGRAPHICA AMERICANA (VI, 41)

seen in the type specimen show that some movement has
taken place.

Such imbricate patterns can arise by other means. Some
crinoids possess columnals which are shaped like truncated
cones, i.e., the diameter of the bottom of the columnal is
less than that of the top or vice-versa. Although columnals
of this variety are found in the Cincinnatian in Ecteno-
crinus Miller (1889), the genus is not known to occur in
Richmond rocks, and in Ectenocrinus the columnals de-
scribed above alternate with cylindrical ones. Thus, it is
not possible to explain the origin of “Dystactophycus” in
this manner.

As mentioned in the discussion of Palaeoscia, there
are concentric patterns which can be shown to be the marks
made by crinoid stems. Plate 80, figure 2,3 illustrates one
such form from an unknown Richmond locality. This speci-
men is unusual because it is preserved both as a concave
epirelief (PI. 80, fig. 3) and a convex hyporelief (PI. 80,
fig. 2). The irregular concentric markings on the lower side
have a diameter of 16.0 cm. The markings on the upper
surface have a smaller diameter (10 cm) and show a more
regular concentric pattern. Vertical sectioning through the
center of the specimens not only indicates that the bedding
is undisturbed but also reveals three or four columnals.
It is concluded that the stem first swept over the bottom
probably dragging shell fragments with it. These could ac-
count for the irregularity of the markings. The stem was
then buried by 1.5 cm of silt, and before breaking off it
again scoured the substrate, this time leaving a more regu-
lar concentric pattern.

“Scolithus dispar” J. F. James
Plate 80, figure 7

1840. Non Skolithos linearis Haldemann, “A Monograph of the Lim-
nidades, or freshwater univalve shells of North America, con-
taining description of apparently new animals in different
classes, and the names and characters of the subgenera in
Paludina and Anculosa,” Supp. to No. 1.

1881. Scolithus linearis Haldemann, James U. P., The Paleontologist,
No. 5, p. 33.

1881. Nom. nud. Scolithus dispar James, U. P., The Paleontologist,
No. 5, p. 33.

1892. Scolithus dispar James, U. P., James, J. F., Geol. Soc. America
Bull., vol. 3, p. 41, fig. 14.

“Type locality.” — From the “bed of Crawfish Run,
eastern part of Cincinnati, 80 feet or so above low-water
mark of the Ohio River” (most probably Economy).

Discussion. — U. P. James (1881, p. 33) described the
Cincinnatian representatives of “S. linearis” as “. . . strong,
raised lines, from 1/24th to over 1/2 an inch or so wide,
generally straight, and parallel to each other . . . All of

the forms here referred to are found on the under side of

the bedded shale, resting on a bed of fine, soft blue clay.
.. .” He went on to say that “if, owing to the different
Geological horizon, or otherwise, these forms shall prove
to be distinct from S. linearis, I propose the name dispar for
the species” (ibid, p. 33). Because U. P. James never for-
mally proposed “S. dispar,” he does not receive credit for
the species. Technically, the specific designation “dispar”
goes to J. F. James (1892), who figured the “species” and
was the first to use the name without reservation.

Several specimens of identified topotype material are
present in the University of Cincinnati collections (PI. 80,
fig. 7). These closely resemble the figure of the Crawfish
Creek material illustrated by J. F. James (1892, fig. 16).
The host rock is a cross-laminated calcisiltite which bears
several linear markings preserved as convex hyporeliefs.
As viewed in Plate 80, figure 7, there are two sets of mark-
ings, one proceeding from left to right, the other from lower
left to upper right. The markings specifically referred to
“S. linearis” by U. P. James (=S. dispar J. F. James) are
thin (1.5 mm), straight ridges, some of which have a longi-
tudinal furrow. A few proceed the entire breadth of the
slab, others fade out before reaching the margin. Associat-
ed with “S. linearis” are other raised markings. Some are
shaped like an open “S,” others resemble small drumlins,
gradually sloping at one end and sharply rounded at the
other. Also found on the slab are moniliform ridges and
closely spaced parallel striae. One additional structure varies
from all others for it is preserved as a concave hyporelief.
These form a branching vermiformlike network, which cuts
through both sets of ridges.

Interpretation. — U. P. James (1881) agreed with Hall
that “S. linearis” was a “fucoid,” and J. F. James (1892,
p. 40) considered it as “. . . the passage of some organism
over the surface of the mud.” There is nothing to indicate
that either of these interpretations is correct, As discussed
elsewhere, S. linearis, as described by Haldemann (1840)
and Hall (1847), is the vertical tube of a phoronid or sessile
polychaete. However, “S. linearis” as employed by U. P.
James is a groove cast. The ridges are too straight and
parallel to be otherwise. Similar forms are illustrated by
Dzutinski and Sanders (1962, pl. 11, fig. B and pl. 14)
from the Oligocene Krosno beds of southeastern Poland. As
shown by Dzulinski and Sanders (1962, p. 74) and Potter
and Pettijohn (1964, pls. 61, 66), the orientation of the
drumlin-like prod marks shows that current movement was
from left to right on the photograph. The tools which made
“S. linearis’ were probably fragments of brachiopod shells.
The S-shaped ridges show where shell fragments tore into
the bottom, twisted, and were pulled out. The parallel

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 393

striae and moniliform markings are the casts of crinoid
stems which were dragged or rolled over the substrate.
The vermiform markings are the only organic features on
the slab and apparently are molds of fecal strings of
Sedimentfressers which moved along the silt-mud interface.

“Palaeophycus flexuosum” U. P. James
Plate 79, figures 1,2

1879. Palacophycus flexuosus James, U. P., The Paleontologist, No.
Bip LS:

1884. Palaeophycus flexuosus James, U. P., James, J. F., Cincinnati
Soc. Nat. Hist., Jour., vol. 7, p. 6, pl. 6, fig. 1.

Type locality. —“Cincinnati Group, Hamilton County,
Ohio, near Milford, at a hillside cut of the Little Miami
Railroad.”

Discussion. — U. P. James (1879, p. 18) characterized
“P. flexuosum” as a “fucoid” consisting “. . . of a regular
series of slightly rounded or flattened stems, which may
have been cylindrical and become flattened by pressure.
The stems lie nearly parallel to each other in their flexuous
course across the slabs of blue shale, sometimes anastomo-
sing (very seldom bifurcating) and are of nearly uniform
size from end to end, about one line [1/12 inch] in width
and the same distance apart.”’ He mentioned that thin slabs
gave the appearance of being composed entirely of “P.
flexwoswm” specimens. The largest specimen observed by
James had an area of over 600 square cm. The species was
not figured, and the type specimen (if designated) is not
present in the James collection at the Field Museum. The
only known illustration is a highly schematic drawing
given by J. F. James (1884, pl. 6, fig. 1).

Two specimens from the University of Cincinnati
collections, as well as a slab in the James collection at
the Field Museum labeled Palaeophycus fexuosum, closely
resemble U. P. James’ description and J. F. James’ illustra-
tion. One form (PI. 79, fig. 2) occurs on a thinly laminated
micaceous siltite which has a thickness of 5 mm. The host
rock has fractured in such a way as to reveal portions
of several bedding planes and each plane is marked by the
“P. flexuoswm” structure. The markings resemble a pattern
of small, flattened, parallel ripple marks having a “wave
length” of 5 mm and an “amplitude” of 0.1 mm. The ma-
terial forming the crests is darker, more micaceous, and
slightly coarser grained than the sediment found in the
troughs. The markings are wavy and infrequently bifurcate
to form two parallel branches.

Another specimen (PI. 79, fig. 1) was collected from the
Economy beds of Twelve Mile Creek, Campbell County,
Kentucky, some 20 meters downstream from the type local-

ity of Rusophycus cryptolithi. In general pattern it is simi-
lar to the specimen just discussed. It differs in that the
markings are smaller (“wave length” of 3 mm, “amplitude”
of 0.1-0.2 mm), more wavy, and anastomose more fre-
quently. The structure covers an area of 700 square cm on
the upper surface of a fine-grained calcisiltite. The “ripples,”
which have a width of but 1.7 mm, are obliterated by
trails. It is not possible to determine whether the trails
are infaunal or exofaunal in origin.

Similar structures have been described by Walcott
(1914) from the Precambrian Belt Series of Montana under
the name Kinneyia (see Hantzschel 1962, fig. 146-3). Wal-
cott (1914, p. 107) described Kinneyia as having a “body
built up of thin, subparallel layers separated by narrow
intervals that are not much greater than the thickness of
the layers forming the body.” Desio (1940) assigned forms
from the Silurian of North Africa to Kinneyia, and Fuchs
(1895) mentioned but did not name similar forms from
the Lias Alpha of southern Germany. Specimens which are
more irregular than those found in the Cincinnatian were
described by Martinsson (1965, fig. 4) from the Middle
Cambrian of Sweden under the name of kinneyian ripples.

Interpretation. —U. P. James (1879, p. 18) inter-
preted “P. flexwosum” as a “fucoid,” although he admitted
“how [the stems] came to be laid down so evenly laminated
and with so little disturbance of arrangement is not appar-
ent.” J. F. James (1884, p. 6) believed that “P. flexosum”
represented swash marks, remarking that “the washing of
water against the bank of a stream often produces ripple
marks which extend along the shores for some distance
in regular, undulating lines.” He went on to say that U. P.
James agreed with this inorganic explanation. Fuchs (1895),
p. 444) interpreted the Lias forms as corrosive markings
etched on bedrock by the algal networks. Desio (1940)
was somewhat more cautious regarding the origin of the
North African forms. He tentatively compared them to the
tubulate coral Cladopora reticulata Hall (1851) (jun. syn. of
Coenites Eichwald) from the Niagaran (Silurian) of New
York State. He also pointed out what he believed to be a
similarity between Kinneyia and Paleodictyon. Martinsson
(1965) regarded the Swedish Cambrian forms as similar to a
form of small oscillation ripple that he has observed form-
ing in the Baltic.

The morphology of the two Cincinnatian specimens
recalls depressed ripple marks; however, according to Dr.
Paul Potter (personal communication), the smaller speci-
men (PI. 79, fig. 1) has a “wave length” which is less than
that of the smallest undoubted ripple mark. Nevertheless,
Shrock (1948, fig. 79) illustrated a form equal in dimensions

394, PALAEONTOGRAPHICA AMERICANA (YI, 41)

to the Cincinnatian specimen from the Lower Ordovician
New Richmond Sandstone of Wisconsin which he referred
to as “eroded symmetrical ripple marks.” While it is possi-
ble that “P. flexoswm” represents some diagenetic phe-
nomenon, such structures have not been reproduced in
laboratory studies.

XII. INCERTAE SEDIS

Sphenophyllum primaevum Lesquereux

1878. Sphenophyllum primaevum Lesquereux, American Phil. Soc.,
Proc., vol. 17, pp. 167-169, pl. 1, figs. 3-5.

1889. Spenophyllum primaevum Lesquereux, Miller, North American
Geol. and Palaeont., p. 141.

1892. Sphenophyllum primacvum Lesquereux, James, J. F., Cincinnati
Soc. Nat. Hist., Jour., vol. 14, p. 48.

Type locality. —“Covington, opposite Cincinnati;”
Limekiln Run, about 370’ above low water, Cincinnati,
Ohio. (The first locality could be either Eden, Fairview, or
MeMillan; the latter is probably McMillan.) No specimens
have been available for examination, the types not being
known.

Discussion. — Lesquereux (1878, p. 167) described the
species as “stems or branches slender, articulations close,
equidistant; leaves in whorls of four or five leaflets con-
nected towards the base and joined by slightly obtuse
sinuses; leaflets either truncate and crenulate at the top,
or sometimes deeply slit or lobate; nerves simple at the
base, sparingly dichotomous, forking mostly once, even
simple.” The only known figures of the species are given by
Lesquereux (1878, pl. 4, figs. 4,5). The form illustrated by
Lesquereux on plate 4, figure 3 was also thought to be
Sphenophyllum, but he could not identify it as conspecific
with S. primaevum. The specimens were found in place “im-
bedded in the clay or attached to pieces of hardened clay
of the same compound as the Cincinnati blue marl” (tbid,
p. 169).

Interpretation. — Lesquereux (1878) considered S.
primaevum, along with Psilophytum gracillomum Lesquereux
and Protostigma sigillarioides Lesquereux, to be the oldest
land plants yet discovered in North America. He allied S.
primaevum with S. scolotheimu Brongniart (1828) and S.
oblongifolium Germar (1844) which he had identified in the
coal flora of Pennsylvania. Although J. F. James (1892)
accepted Lesquereux’s interpretation without question,
Miller (1899, p. 141) was more skeptical and remarked, “I
think this is not a plant.” While Lesquereux’s description
and figures of S. primaevum resemble the generally accepted
concept of Sphenophyllwm, it is unlikely that we are dealing
with a land plant. The first undoubted Sphenophyllum is
from the Upper Devonian beds of New York State and the

Gaspé region of Canada. It would be remarkable if one of
the earliest known land plants were found in marine de-
posits hundreds of miles from the nearest shore. However,
at present the form remains a Problematicum. Despite the
fact that the present study has turned up no new specimen
nor are any in the collections studied, it was Lesquereux’s
impression that it was not rare. He mentioned that he had
heard of several specimens and that local collectors had
thought it to be “the work of insects fucoidal or
coralline productions like Oldhamia” (Lesquereux, 1878, p.
169). Oldhamia Forbes (1849) is now recognized to be a
stellate burrow, and Oldhamuia-like forms are not known
to occur in the Cincinnatian. The fact that it occurs
“attached to pieces of hardened clay” makes it more likely
that this is some sort of organic encrustation, perhaps a
sponge or crinoid rootlet system. Until an actual specimen
is found, there is little more that can be done.

PROTOSTIGMA Lesquereux, 1878
1878. Protostigma Lesquereux, American Phil. Soc., Proc. vol. 17,
pp. 169-173, pl. 1, figs. 7-8.

1889. Protostigma Lesquereux, Miller, North American Geol. and
Palaeont., p. 135.

1891. Protostigma Lesquereux, James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 14, pp. 48-49.

Type species. — Protostigma sigillarioides Lesquereux
(1878) from the “Cincinnatian Group” in Longstreet Creek,
six miles east of Lebanon, Ohio (probably Richmond beds).
Monotypic.

Discussion. — P. sigillarioides was described by Lesquer-
eux (1878, p. 169) as “branches or stems cylindrical, scarce-
ly flattened by compression; surface marked by rhomboidal
cicatrices, enlarged on the sides, contiguous and in spiral
order, with indistinct impressions of oval scars in the
middle.” The form was figured by Lesquereux (1878, pl.
4, figs. 7,8). One specimen which was not figured “. . . is
larger, seven centimeters in diameter, nine centimeters
long, nearly exactly cylindrical, with irregular more or less
distinct ribs, marked crosswise by large wrinkles or irregu-
lar protuberances, which do not show any distinct relation
of form and position between them” (ibid., pp. 169, 170).
The specimens consist of a “hard bluish clay or marl mixed
with grains of coarse sand” (ibid., p. 173).

Lesquereux mentioned the form in 1874 (without
naming it), and this prompted an immediate reply from
Newberry (1874), who had the specimens in his possession
before Lesquereux described them. Newberry (1874, figs.
1, 2) gave two figures which are far less schematic than
those of Lesquereux (1878). He failed to find any trace
of oval scars in the center of the cicatrices.

TRACE FOSSILS CINCINNATI AREA: OsGoop 395

Interpretation. — Lesquereux (1874, 1878) considered
Protostigma to be the oldest of the lycopods. Newberry
(1874) regarded them as the stems of large “fucoids,” as
did Miller (1889). Newberry pointed out that there was
no carbonaceous material anywhere in the specimens. J. F.
James (1891) questioningly placed the genus in the lyco-
pods.

Newberry’s discussion and figures cause some doubt
on Lesquereux’s interpretation. The first undoubted lyco-
pods occur in the Lower Devonian of Australia. While it is
not possible to refute Lesquereux’s hypothesis, it is equally
probable that Protostigma is the cast of a large burrow,
the internal mold of a large nautiloid, or even possibly a
specimen of Beatricea, the problematic stromatoporoid;
however, the latter does possess internal structure. Until
identified material is located, Protostigma must remain
Incertae sedis.

PALAEOSCIA Caster, 1942

Plate 71, figure 3; Plate 82, figures 1-6

1879. ?Laevicyclus Quenstedt, Petrefactenkunde Deutschlands, pt. 1,
vol. 6, Korallen, Die Réhren und Sternkorallen, p. 577, pl. 164,
fig. 35.

1912. ?Cyclozoon Wurm, Deutsche geol. Gesell., Zeitschr., Bd. 63,
pies; ple 7, ties Ld

1942. Palaeoscia Caster, Palaeontographica Americana, vol. 3, pp.
26-29, pl. 5, figs. 1-4.

1955. ?Laevicyclus Quenstedt, Seilacher, in Schindewolf and Seilacher,
Akad. Wiss. Lit. Mainz, math-nat. KI., Abh., No. 10, pp. 389-
390, pl. 18, fig. 1.

1962. ?Laevicyclus Quenstedt, Seilacher, Hantzschel, Trace-Fossils
and Problematica iz Treatise on Invertebrate Paleont., Moore
(ed.), Pt. W, Miscellanea, p. W201.

Type species. — Palaeoscia floweri Caster (1942) (PI.
87, fig. 1; UCM 24079) from the Corryville beds of Stone-
lick Creek, Clermont County, Ohio. Monotypic.

Diagnosis. — A series of regular or irregular concentric
circles marked by a central “pore.” In some forms a series
of faintly impressed grooves radiates out from the “pore.”

Discussion. — At the time of Caster’s study (1942)
only two specimens were known, both from the Corryville
beds of Stonelick Creek, some 290 meters downstream from
where the creek is crossed by Ohio Route 131. Both speci-
mens are preserved as concave epireliefs in a thin calcisil-
tite which grades down into a coarse-grained calcarenite.
The type specimen (Pl. 82, fig. 1) is composed of a series
of irregular concentric circles 7.5 cm in diameter. Many of
the circles are incomplete and some of them intersect. The
center of the specimen is marked by a small well-developed
“pore” from which radiate 16 grooves (PI. 82, fig. 3). These
were interpreted by Caster as representing eight pairs of

e

radial canals. The larger paratype (Caster, 1942, pl. 5,
fig. 4) is not so well preserved and covers an arc of 160°.
The concentric lines are faintly impressed but appear to
be more regular. There is no trace of radial grooves or a
central “pore.”

Since the original description additional specimens from
the same locality have been recovered which shed more light
on the nature of the genus. The most revealing are two
overlapping specimens neither of which is complete (Pl. 82,
fig. 5). The larger form, 6.4 cm in diameter, consists of
two strongly impressed concentric arcs as well as four
very faint ones. Radiating from the central “pore” are
numerous grooves which subscribe an are of 180°. They are
too faint to count accurately but the number appears to be
about 25. There is no hint of a regular paired arrangement.
The smaller specimen truncates the larger and has a di-
ameter of 4.6 cm. It lacks the concentric appearance of the
other specimen, as only a single are is present. Likewise,
there are no radial grooves. The “pores” of both forms are
similar; each is cone-shaped and possesses a raised rim.
Also the outer ares are similar in that the outer wall is
much steeper than the inner. In some areas the inner wall
gives the appearance of having been planed off.

Three additional specimens from the type locality
should be noted. The first (PI. 82, fig. 2) differs from those
described above in that it consists of 8-10 faint but regu-
larly spaced, nearly perfect circles. The outer circle is the
most plainly defined, and in cross-section the specimen re-
sembles a shallow dish. There is no well-defined central
“pore” nor is there any hint of radial grooves. Another
specimen (PI. 82, fig. 6), which has a diameter of 5.4 cm,
is characterized by a marked contrast in the appearance of
the circles. The circle, whose outer margin lies 9 mm from
the central “pore,” is incised 2 mm below the general level
of the specimen. The central “pore” has a diameter of 3
mm and thus exceeds the dimensions of the “pores” of other
specimens. It is uncertain whether the third specimen
(Pl. 71, fig. 3) represents topotype material. In opposition
to the other Stonelick forms, it is preserved as a convex
hyporelief, and the clarity of preservation indicates that
the subjacent bed was a lutite. All other Stonelick forms
are found as concave epireliefs in calcisiltites. It may be
that the difference in preservation is due to a lateral facies
change in the same bed, On the other hand, the specimen
may be from a different bed. The form in question is ellip-
soidal in outline; the long axis measures 5.5 cm, the short
axis, 5.0 cm. This variation in dimensions can be ascribed
to the difference in the development of the four quadrants.
The outer margins of the opposing quadrants on the long
axis are raised 3 mm off the surface of the host rock.

396 PALAEONTOGRAPHICA AMERICANA (YI, 41)

In contrast, the margins of the quadrants on the shorter
axis are nearly flush with the surface of the host. Proceed-
ing from the central “pore” to the margin of the specimen
is a shallow U-shaped trough which has a width of 2 mm.

Similar concentric markings have been found at other
Cincinnatian localities. Mr. Joseph Stocker collected a half
dozen specimens from the Corryville beds of Hall’s Creek,
some 10 miles from the Stonelick Creek locality. These forms
possess anywhere from 8-26 concentric rings which are more
regular both in outline and spacing than most of the pre-
viously described specimens. All are preserved as concave
epireliefs and have a dish-shaped cross-section. There is no
evidence of planation, radial markings, or a central “pore”
on any of the specimens.

Palaeoscia-like forms have also been collected from
the Economy beds of Duck Creek, Kentucky. A single slab
(PI. 82, fig. 4), having an area of 33.6 square cm, yielded
23 specimens which are preserved as convex hyporeliefs.
Instead of being complete or nearly complete circles, mark-
ings are found in only one quadrant. Casts of scour marks
associated with the “pore” (here a small conical pimple)
reveal that the arcs are developed only in the downcurrent
quadrant. The number of arcs varies from 4-8; they are
very sharply defined but not evenly spaced. There is no
evidence of planation.

Forms like those described above are by no means
confined to the local Cincinnatian section. Quenstedt (1879)
proposed Laevicyclus for one such form from the Angulaten
Sandstone (Jurassic) of Wiirttemberg. It is preserved as a
concave epirelief and consists of several concentric circles
with a central “pore.” Wurm (1912) described others from
the Triassic of Aragon, as has Philipp (1904) from the
Upper Triassic of northern Italy. Wurm’s specimens, which
he named Cyclozoon philippi, have a diameter of 34 mm
and possess only two concentric circles. His figure (Wurm,
1912, pl. VII, fig. 11) shows a central “pore” but no radial
grooves. Philipp’s form, which was unnamed, differs in that
it has a single circular impression. The specimens occur
as concave epireliefs on a ripple-marked surface. Instead
of being a depression, the central “pore” appears to be a
raised area (Hiantzschel, 1962, fig. 123-3a).

Seilacher (1955, pl. 18, fig. 1) described additional
specimens of Laevicyclus from the Lower Cambrian of the
Salt Range. Preserved as convex hyporeliefs, the small
(diameter of 1 cm) Salt Range forms have concentric arcs
developed only in two opposing quadrants. In one speci-
men the best preserved specimen has three arcs in one
quadrant, two in the other.

Interpretation. — Theories on the origin of these circu-
lar structures are as varied as the morphology of the forms

themselves. Quenstedt (1879) assigned Laevicyclus to the
corals, while both Philipp (1904) and Wurm (1912) con-
sidered their forms to be body fossils of unknown affini-
ties. Schmidt (1934) and Lessertisseur (1955) maintained
that such circular structures arise from gas bubbles being
driven upward during diagenesis.

Seilacher (1953b, fig. 5) believed that Quenstedt’s
Laevicyclus was a trace fossil. He based his conclusion on
the feeding methods of the polychaete Scolecolepis squa-
mata Séderstrém, which lies vertically in sand with only its
head and tentacles protruding above the surface. In calm
water the tentacles rotate in a circular pattern, gleaning
the nutrients from the substrate; however, when currents
are present, such actions are not necessary, and the organism
merely grasps food particles as they are washed within its
reach. In this case, the only markings left on the substrate
are small scour channels on either side of the tube. Seilacher
(1955) mentioned that mollusks and arthropods, as well as
polychaetes, might have produced the Salt Range speci-
mens. The smaller diameter and the fact that the markings
were developed in only two quadrants indicated to him that
the tentacles were not as mobile as they are in Scolecolepis.

Caster (1942) interpreted the two Stonelick speci-
mens, then available to him, as molds of porpitid siphono-
phores, comparing them to the Recent Porpalia prunella
Haeckel. Admittedly the Cincinnatian forms do bear some
resemblance to undoubtedly soft-bodied coelenterate fossils.
Sprigg (1947, 1949) described several such forms from the
Precambrian Ediacara beds of Australia. There can be
little doubt that these Australian specimens are bona fide
“jelly-fish.” The radial canals are regularly disposed and
encircle the entire specimen, and the forms are associated
with numerous other soft-bodied fossils (Glaessner, 1959).

In spite of the resemblance to the Australian forms,
there is reason to believe that the rims of the central
“pore,’ the numerous radial grooves, the irregularity of
the concentric arcs in some forms, and the planed-off ap-
pearance of the outer arcs all indicate that the Stonelick
Creek specimens represent the markings made by some swirl-
ing object. The holotype and the two overlapping specimens
illustrated on Plate 82, figure 5 come the closest to re-
sembling the pattern which might have been made by an
Ordovician Scolecolepis-like polychaete. There is evidence of
planation, the arcs are uneven, and the radial grooves could
well be the imprints of the tentacles. However, the other
specimens from the same locality do not support this con-
clusion; the arcs are too regular. Moreover, sectioning of
one specimen revealed a brachiopod shell immediately
underlying the center of the circle. We must thus consider
the possibility that these circular patterns are the mark-

‘TRACE FOssILs CINCINNATI AREA: OsGoop 397

ings of a partly uncovered agglutinated tube as it was being
swept around by the currents. The brachiopod shell offers no
real problem as the tube could have been anchored to it
and have been partly uncovered by erosion. There is no
evidence of crinoid holdfasts or stems in any of the speci-
mens, thus that group is ruled out.

Even though some of the specimens resemble Scoleco-
lepis feeding traces while others resemble sweep marks, it is
highly impossible that the specimens, all from the same
bed, have two different origins. However, at this point it is
not clear which of the two explanations is the correct one;
therefore, Palaeoscia is assigned to the Incertae sedis. The
origin of the Hall’s Creek forms is even more problematic.
The circular markings are too perfect in outline to represent
either a trace fossil or a siphonophore. While it is believed
that these are probably sweep marks of an eroded tube, it
must be admitted that careful sectioning failed to reveal
any trace of a tube or similar structure. This may not be as
critical as it first appears. The “pores” are so small (1 mm)
that such a tube might not even be visible in vertical
sections, unless it were filled with a material different
from that of the host rock.

Because the Duck Creek specimens (Pl. 82, fig. 4)
are preserved as convex hyporeliefs, vertical sections are
irrelevant. The scour marks on either side of the pimple-
like raised area in the center show that these most probably
are sweep marks. There seems little reason why such regular
arcs should be developed in only the downstream quadrant
if this were the feeding trace of a Scolecolepis-like organism.
Yet it is unusual that no evidence of the tube or stem is
found in any of the 23 specimens.

In summary, the generic designation Palaeoscia is
applied only to specimens from Stonelick Creek. These may
be either feeding traces or the sweep marks of an agglutin-
ated tube. The specimens from Hall’s Creek and Duck Creek
are thought to be sweep marks and they are not assigned to
Palaeoscia. Not all such markings are so enigmatic, for some
can definitely be shown to be sweep marks of crinoid stems.
These are discussed under “Dystactophycus.”

ARISTOPHYCUS Miller and Dyer, 1878
Plate 83, figures 6-8; Text-figure 28

1878. Aristophycus Miller and Dyer, Contributions to Palaeontology,
No. 2 (Cincinnati, Ohio, private publication), p. 3, pl. 4, figs.
2-3.

1885. Aristophycus Miller and Dyer, James, J. F., Cincinnati Soc.
Nat. Hist., Jour., vol. 7, p. 130.

1962. Aristophycus Miller and Dyer, Hantzschel, Trace-Fossils and
Problematica iz Treatise on Invertebrate Paleont., Moore (ed.),
pt. W, Miscellanea, p. W232.

Type species: Aristophycus ramosum Miller and Dyer

Text-figure 28.— Schematic cross-section of Aristophycus ramo-
sum Miller and Dyer, 1878. The diagram shows that the secondary
branches bifurcate from the main branch from below rather than at
right angles (i.e., below a horizontal plane bisecting the main branch)

)

(1878) (PI. 83, fig. 7; HBM 3173; latex mold UCM 37580:
Pl. 83, fig. 8; HBM 3172; latex mold UCM 37668) from the
Fairview or McMillan Formation, Cincinnati, Ohio.

Diagnosis — Pulmonate, branching structures of mod-
erate size which are preserved as convex epireliefs. As
viewed in cross-section, the smaller secondary branches
join the terete main branch from beneath, forming an anele
of 30-45° with the horizontal. The tertiary and quaternary
branches inosculate to form an anastomosing pattern.

Discussion. — Aristophycus was described by Miller
and Dyer (1878b, pp. 3,4) as a “fucoid” with a “stem
which divides into branches or fibres; these again divide
and subdivide, until they become exceedingly numerous.
The ramifications are sent off like the roots of a tree or
shrub without any determinate order, while many of the
smaller fibres inosculate like the veins in the leaf of a tree.”

One species, A. ramoswm, and the variety A. ramosum
var.” germanum were proposed. The latter differed in that
“small fibres are thrown off from the main stem between
the branches . . . [these] are finer and more completely in-
terwoven than they are in A. ramosum” (Miller and Dyer,
1878b, p. 4).

On the whole Miller and Dyer’s description is accu-
rate, but the authors neglected to mention the unusual
mode of branching. As seen in Text-figure 28, the secondary
branches do not join the main trunks of the system on a
horizontal plane but instead from beneath. The angle of
juncture varies between 30-60° off the horizontal. The same
generally holds for the relationship between the small] sec-
ondary and tertiary branches; beyond this the branches be-
come so small that observation is difficult.

The type specimen of A. ramosum (Pl. 83, fig. 7),
which is 17 cm long and 13 cm wide, consists of seven ir-
regularly winding main branches which vary from 2-6 mm
in diameter along their length. Distally these give rise to
numerous short secondary branches which have a diameter

“

of 0.5-1.5 mm. These in turn branch to produce a capillary-
like system which crosses over and interpenetrates the
“ » _ ” ‘SI & = rr h

capillaries” of a neighboring branch. The unusual method

398 PALAEONTOGRAPHICA AMERICANA (YI, 41)

of branching mentioned above is best seen in the distal por-
tions of the system. The type specimens of A. ramosum
“var.” germanum (Pl. 83, fig. 8) is slightly smaller, having
a length of 9.5 cm and width of 5 cm. The main trunk has
a diameter slightly in excess of 1 mm. Unlike the type of A.
ramosum where the anastomosing pattern is confined to the
distal regions, A. ramosum “var.” germanum shows. this
arrangement along most of its length, although it is best
developed distally. The small branches wind about in inter-
penetrate to form a system which is even more complex than
that of A. ramosum.

In the A. ramosum “var.” germanum type, there is no
difference in lithology between the host rock and the fos-
sil. Conversely the A. ramosum type appears to be finer
grained than the host rock, for it flakes off with comparative
ease.

In spite of these minor differences, it is concluded that
there is no need for separate taxa for the two specimens.
The more important features, e.g., the anastomosing pattern
and the mode of branching, are common to both; therefore,
A. ramosum “var.” germanum is placed in synonymy with
A. ramosum.

Similar structures have been reported from Europe
and were called “figures de viscosité” by Issel (1889).
Fuchs (1895, pl. 9, fig. 4) illustrated one specimen from
the Braun-Jura Beta of Wiirttemberg. As nearly as can be
determined from his figure, it is congeneric with Aristophy-
cus. The form measures 9 cm by 2.2 cm, and the main
trunk has a diameter of less than a millimeter. The exact
nature of the branching is undeterminable.

Interpretation. — James (1885) interpreted Aristophy-
cus as casts of rill marks; and apparently Miller (1889, p.
107) was convinced, for he conceded that the genus might
well be inorganic in origin, or if it were a “fucoid,” it was
too poorly known to be retained as a separate genus.
Hantzschel (1965, p. 10) reported that Seilacher (personal
communication to Hantzschel) considered Aristophycus to
be a “figure de viscosité.”

Upon casual inspection, James’ rill mark interpreta-
tion appears plausible. However, closer examination reveals
that Aristophycus occurs as convex eptreliefs, 1.e., raised
patterns, on the upper surface of the beds. There can be no
question of this, as the side of the slab bearing A. ramosum
contains a portion of a Corophioides, in which the bottom
of the “U” is 180° to the surface bearing Aristophycus.
The presence of channel fillings in the other specimen con-
firms this orientation.

Seen in this light, the origin of such a form becomes
problematic. In fact it is not possible to entirely dismiss
an algal origin, but the mode of preservation, 1.e., convex

epirelief, makes it more likely that Aristophycus is either
a burrow or some physical phenomenon.

Fuchs likened it to the anastomosing pattern which
results when a knife blade is raised off a slab of butter.
Anyone who has ever polished rock sections has seen the
same effect. Fuchs pictured a shell-less gastropod or a large
Planaria rising vertically off the bottom with the resulting
suction producing the pattern. However, it is difficult to
see how suction could produce such a regular hierarchic and
interlocking pattern where the secondary branches consis-
tently join the main trunk from below.

Because of the unique preservation it is most probable
that Aristophycus originated below the depositional inter-
face along a bedding plane surface. The clarity of preserva-
tion indicates that the overlying sediment was a lutite. It
is possible that Aristophycus is a branching feeding bur-
row, where the organism moved along the main branch and
extended a slender proboscis or tentacles down along the
mud-silt interface. The terete cross-section of the branches
and the variations in the diameter of the main trunk sup-
port such a hypothesis. However, Aristophycus differs in
many respects from all known burrows. It is difficult to
explain why the angle of branching is consistently 30-60°
off the horizontal and why the branches become progress-
ively smaller distally. Likewise, the mode of preservation
is exactly the opposite from that found in many burrows.
In most burrows, which are preserved as boundary reliefs,
e.g., Phycodes, the overlying sediment fills the cavity made
by the organism. The burrow hypothesis would be more
attractive if Aristophycus were preserved as a convex hypo-
relief or a concave epirelief.

Because of these difficulties it is equally as probable
that Aristophycus is of physical origin. Perhaps it repre-
sents some form of diagenetic flow pattern. But, just as
Aristophycus resembles no known burrow, neither does it
resemble any known diagenetic phenomenon. Moreover, the
anastomosing pattern of the branches is in opposition to the
characteristics of flowing, percolating water,

Walcottia sulcata U. P. James

1881. Walcottia sulcata James, U. P., The Paleontologist, No. 5, p. 44.

1886. Walcottia sulcata James, U. P., James, J. F., Cincinnati Soc.
Nat. Hist., Jour., vol. 8, p. 162.

Type locality. — Economy beds of the eastern part of
Cincinnati. There is no known illustration of the species.
Discussion. —U. P. James (1881, p. 44) created W.

sulcata for “ sub-cylindrical, slightly tortuous forms,
with a longitudinal groove along the center of the exposed

part; a series of fine, closely arranged transverse or gently

‘TRACE FOSSILS CINCINNATI AREA: OsGoop 399

oblique striae, commencing, apparently, at the bottom of
the groove, and passing over the higher rounded surface
to the embedded portion on each side.” He remarked that
the species had a length of 2 inches and a width of 1/12
inch and “differed materially” from W. rugosa Miller and
Dyer (1878) in“... the features of the longitudinal groove,
the uniform width and the transverse striae” (ibid, p. 44).

Interpretation. — U. P. James offered no interpretation
for W. sulcata and J. F. James (1886) merely referred to
the form as “a burrow.” Neither a type specimen nor iden-
tified material could be located in the James collection at
the Field Museum, The original description recalls small,
striate, bilobed Cruziana-like trails. Thus, it is possible that
the species is a trail similar to Ormathichnus Miller (1880).

SACCOPHYCUS U. P. James, 1879

1847. ?Sphenothallus Hall, Nat. Hist. New York, Palaeont. New
York, vol. 1, p. 268, pl. 68, fig. 1.

1878. Sphenothallus Hall, James, U. P., The Paleontologist, No. 2,
Deo:

1879. Saccophycus James, U. P., The Paleontologist, No. 3, p. 17,

1885. Saccophycus James, U. P., James, J. F., Cincinnati Soc. Nat.
Hist., Jour., vol. 7, p. 157.

1891. Partim Planolites Nicholson, James, J. F., Cincinnati Soc.
Nat. Hist., Jour., vol. 14, p. 47.

1962. Saccophycus James, U. P., Hantzschel, Trace-Fossils and Prob-
lematica iz Treatise on Invertebrate Paleont. Moore (ed.),
pt. W, Miscellanea, p. W242.

Type species.—Saccophycus intortum U. P. James
(1879) from the “Cincinnati Group, near Lebanon, Ohio.”
Monotypic.

Discussion. —U. P. James (1879, p. 17) characterized
Saccophycus as resembling “. a long tortuous bag or
sack, with the ends drawn in; undulating, wavy surface,
finely striated longitudinally; sides and ends rounded in
and under.” He further mentioned that the type (the only
known specimen!) was approximately three inches long and
had a breadth of one inch at one end and one-half inch at
the other. The specimen was “partly imbedded in a fragment
of limestone.” There is no known figure of the genus. U. P.
James (1878) had mentioned the specimen previously, ten-
tatively assigning it to Sphenothallus latifolius Hall (1847)
(Upper Ordovician: New York). He remarked that it re-
sembled the specimen illustrated by Hall (1847, pl. 68,
fig. 2e) and that the Cincinnatian form might be the stem
of S. latifolius. Hall’s figure shows a smooth truncated cone
having a height of 4.5 cm and a diameter of 2.5 cm at
the base and 1.5 cm at the apex. The specimen figured by
Hall does not resemble any Cincinnatian fossil or pseudo-
fossil.

The sole specimen of Saccophycus is not with the
James collection at the Field Museum. A search of the

literature reveals that no subsequent author has enlarged
upon the original description.

Interpretation. — U. P. James interpreted Saccophycus
as a “fucoid,’ while J. F. James (1885, p. 157) believed
it to be “the burrow of an annelid.” J. F. James (1891)
considered Saccophycus to be synonymous with Blastophy-
cus diadematum and Trichophycus venosum. Although there
is nothing to indicate that Saccophycus is synonymous with
Blastophycus, U. P. James’ original description is reminisc-
ent of Trichophycus venosum. Until the type specimen of
Saccophycus is located, the matter must rest.

XIII. INVALID SPECIES

Ulrich (1880) in his Catalogue of Fossils Occurring
in the Cincinnati Group listed four species of previously un-
described “fucoids.” Palaeophycus ornatum, Palaeophycus
sculptum, Conostichus truncatus, and Cyclophycus later-
ale. He mentioned in the preface that “many of the new
species recorded in the list, have as yet not been published.”
A search of his subsequent works reveals that he never
formally proposed nor described the above species and that
they are thus nomen nudum. A comparison of the names
with any forms actually occurring in the Cincinnatian would
be pure speculation.

XIV. COMPARATIVE PALICHNOLOGY AND
SUGGESTIONS FOR FURTHER STUDY

COMPARATIVE PALICHNOLOGY

Until recently most study of trace fossils has been
concerned primarily with description and interpretation of
individual forms, 1.c., paleoautoecological investigations. As-
semblages of traces have only sporadically received atten-
tion. Richter’s (1931) analysis of trace fossils in the Huns-
riick Shale (Devonian: Rhineland) and Linck’s (1949)
study on the Schilf Sandstone (Triassic: Wiirttemberg) are
notable pioneer works of synthesis, 1.¢., paleosynecological
analysis.

The broader aspect of the paleosynecology of traces,
an attempted integrated investigation of the entire trace
fossil spectrum and subdivisions, is new indeed. Seilacher
(1954) initiated his comprehensive work in the compari-
son of the trace fossils of the Alpine Flysch and Molasse.
More complete documentation of his 1954 conclusions came
in his 1955 paper on trace fossils of the Cambrian section
in the Salt Range of Pakistan. Here he compared the trace
community of the Salt Range with five other trace com-
munities: the Triassic Schilf Sandstone, the Lias and Dog-

400 PALAEONTOGRAPHICA AMERICANA (YI, 41)

ger (Jurassic) Sandstones of Wiirttemberg, the Swiss Mio-
cene Molasse of the Alps, the Cretaceous and Tertiary
Flysch of the same area, and the Ordovician-Devonian
Nereites beds of Portugal (Seilacher, 1955, fig. 5).

Seilacher found that the trace assemblages of the
Alpin Flysch and the Portuguese section were character-
ized by abundant Pascichnia which exhibited intricate geo-
metric patterns, While Fodinichnia and Repichnia were also
numerous, there was a complete lack of Cubichnia.

On the other hand, in the remaining four sections
Cubichnia formed the dominant group and Pascichnia were
virtually nonexistant. The grazing traces were represented
only by Dimorphichnus in the Salt Range and Helminth-
otda in the Molasse, and both of these are questionable.
As shown elsewhere herein, Dimorphichnus may be a form
of trilobite trail which exhibits strongly oblique movement.
The Molasse representative of Helminthoida is poorly or-
ganized and lacks the tight meanders of its Flysch counter-
part. The Repichnia are well represented in these same four
sections, and while several different forms of Fodinichnia
were present in the Salt Range and Lias and Dogger sec-
tion, the latter were not well developed in the Schilf and
marine Molasse strata.

The dichotomy in the trace fossil assemblages is re-
flected by a marked difference in lithology and_paleo-
ecologic setting. A capsule summary of the stratigraphy
of each section is given below.

The Cambrian section of the Salt Range is composed
of nearly 360 meters of dominantly clastic sediments. The
basal 80-100 meters of alternating beds of fine-grained
sandstones and shales are marked by imprints of salt casts,
mud cracks, and ripple marks. These are overlain by a 10
meter thick, well-rounded quartz conglomerate which grades
upward into a micaceous, glauconitic quartz sandstone con-
taining a good marine fauna. Overlying this are dolomite
beds interstratified with thin shales. The uppermost units
are thin-bedded sands and shales bearing salt pseudomorphs.
Schindewolf and Seilacher (1955) believed that the column
reflected a shallow water marine environment which became
terrestrial in the upper portion.

The Schilf Sandstone (Triassic: Wiirttemberg) varies
from 3-42 meters in thickness and is composed of well sort-
ed, fine-grained micaceous sands and shales which are ripple-
marked and bear other evidence of current activity. Locally
some of the sands have a high iron content. Although plant
fossils are common, invertebrate body fossils are almost
entirely absent. Linck (1949) interpreted the Schilf as
representing a shallow water deltaic environment which was
predominately marine but contained some brackish water
units.

According to Brinkmann (1960) the Lias of the Wiirt-
temberg area contains 70 meters of fossiliferous limestones
and marls. Most of the trace fossils occur in the thin (6
meter) Angulaten Sandstone (Lias Alpha-2). The Dogger
of Wiirttemberg is composed of over 100 meters of ferru-
ginous sandstones, limestones, and marls. The trace fossils
are encountered in the clastics of the Dogger Beta. The
Lias and Dogger are interpreted as shallow water deposits
which thicken to the north and grade into the fine-grained
deep water clastics of northwestern Germany.

The Tertiary Alpine Molasse exhibits such a wide litho-
logic range that it is difficult to characterize in a few
brief phrases. Generally speaking, the section is made up
of interbedded bituminous slates, conglomerates, sandstones,
and shales. Vertical and lateral facies changes are numer-
ous, and cross-lamination and channel fillings are common.
Current indicators, even in local areas, present a bewilder-
ing variety of directions. According to Kuenen and Carozzi
(1953) the strata are irregular in thickness and distribu-
tion, and the contacts between the various units are not
sharp. Graded bedding within individual units is generally
absent. The Molasse is thought to indicate sediments de-
posited in shallow neritic and nonmarine environments;
the fauna is well developed and is composed of marine,
brackish, and lacustrine forms. As noted above, Seilacher
confined his study to the marine beds.

While the four sections discussed above vary in details
(as do their respective trace fossil communities), they are
all regarded as being shallow water (neritic) deposits. In
contrast, the sediments of the Alpine Flysch and the
Nereites beds of Portugal are thought to reflect a far dif-
ferent environment.

Recently the Cretaceous and Tertiary Flysch of the
Alps has received considerable attention. The sequence,
whose thickness is measured in thousands of meters, includes
a monotonous succession of micaceous sandstones and shales.
The sandstones are normally graded and the contact with
the subjacent shale is sharply defined. Convolute bedding is
not uncommon, and the soles of the sandstones bear numer-
ous sedimentary markings (flute casts, tool marks). In op-
position to the Molasse, these markings indicate a uniform
current direction. Ripple marks, scour channels, and cross-
lamination are generally absent. Body fossils are rare,
and any fossils found are fragmentary and poorly preserved;
in many areas trace fossils are the only evidence of au-
tochthonous life. Although the origin of the Flysch was for
many years in doubt, the work of Kuenen, Migliorini, and
others on turbidity flows has shown that the alpine sequence
is most probably of deep water origin and that the sedi-
ments are derived from periodic submarine slides.

TRACE FOSSILS CINCINNATI AREA: Oscoop 401

The Nereites beds of Portugal, as described in part
by Delgado (1908), include over 4,700 meters of interbed-
ded graywackes, shales, and quartzites. Some carbonates
and sedimentary breccias are also present. While the mica-
ceous shales do contain graptolites, other body fossils are
not common. Like the Alpine Flysch, the Nereites beds are
thought to be deep water geosynclinal deposits.

Seilacher (1954, p. 223) pointed out that the basic dif-
ferences in the two assemblages can not be explained by
the different ages of the units. With a few exceptions trace
fossils make poor index fossils (Seilacher, 1960). Pascichnia
are as well developed in the Lower Paleozoic of Portugal as
they are in the Mesozoic and Cenozoic; likewise, Cubichnia
range from the Cambrian through the Tertiary. Seilacher
(1954, p. 223) also maintained that the lithology of the
host rock was not important. Even though the gross aspects
of the sequences may differ, most of the trace fossils occur
in similar rock types, e.g., silts or fine-grained sands. Seil-
acher concluded that water depth was the controlling factor.
He believed that Cubichnia represented burrows of con-
cealment, and while in shallow, well-lighted water such bur-
rows were an important means of protection, they were
unnecessary in the dark, deep water basins. Although Seil-
acher judged that Pascichnia reflected the concentration
of organic material in thin layers of sediment, he was un-
able to explain why Pascichnia were not found in shallow-
water environments.

In 1964 Seilacher greatly expanded his previous com-
parison to include 42 sections of varying ages and geo-
graphic locations (Seilacher, 1964, fig. 7). He renamed the
community typified as Pascichnia the “Nereites facies,” while
the Cubichnia suite became the “Cruziana facies.” In addi-
tion he recognized a third suite, the “Zoophycos facies.” He
reiterated his earlier belief that the “Cruziana facies” rep-
resented a shallow-water environment, while the “NVereites
facis” was indicative of deep water. The “Zoophycos facies”
was intermediate between the two.** In the Ordovician of
northern Iraq, Seilacher (1964, p. 314, fig. 8) demonstrated
a succession of all three facies in a single vertical section.
The lower unit, the Khabour Quartzite, is typified by
Cruxiana, Diplocraterion, Daedalus, and Phycodes. This is
overlain by Dershish Sandstone which carries Zoophycos,
Teichichnus, and Chondrites. In contrast, the uppermost
unit, the Sinat Shale, contains the Pascichnia Helminthoida,

“Seilacher did not specifically define what he meant by deep or
shallow water. For our usage, shallow water refers to environ-
ments within the zone of effective light penetration, while deep water
refers to environments below the depth of light penetration. By the
nature of his discussion Seilacher implied that shallow water is in the
neighborhood of 0-100 meters, while deep water would be 350
meters+.

Paleodictyon, and Neonereites, as well as Chondrites. To
him this indicated a transition from shallow to deep water
environment. Seilacher (1964) also remarked that a lateral
transition between the “Nereites facies” and “Zoophycos
facies” can be found in the Upper Paleozoic of the Oua-
chitas, where what he interpreted as the deep-water Johns
Valley turbidites grade north into the more shallow-water
Atoka Sandstone which carry Zoophycos. Seilacher (1964,
p. 314) concluded that “there is no doubt that still further
toward the craton this Zoophycos facies would eventually
grade into the Cruziana facies including littoral deposits.”

In subsequent publications, Seilacher (1963, 1967b)
expanded his number of trace fossils communities to six. He
proposed the “Scoyenia facies” for nonmarine clastics best
typified by the Juniata “redbeds” and Catskill “redbeds”
of the Appalachians. Characteristic trace fossils are [sopo-
dichnus, limuloid tracks, and some species of Planolites.
Seilacher established the “Skolithos facies” for marine areas
of rapid sedimentation and erosion (littoral zone) where
the sessile infauna must adapt to the changing levels of the
depositional interface. The trace fossils occurring in thi-
zone are Skolithos and U-shaped Spreite structures. He
questioned (1967b, p. 424) whether Ophiomorpha belongs
in the facies. He cited the Devonian of northern Iraq, the
Lower Silurian Tuscarora Quartzite of New Jersey, and
the basal Cambrian of Estonia, Sweden, and the Grand
Canyon as examples. The “Glossifungites facies” was intro-
duced in 1967 for environments similar to the “Skolithos
facies” but where the bottom sediments were more stable
and not subject to sudden changes. He stated that the sedi-
ments were slightly indurated and the trace fossils are
confined to single bedding planes.

To summarize briefly, Seilacher, over a period of some
years, established six trace fossil assemblages or “facies.”
These are in order of increasing water depth: ‘Scoyenia
facies,” “Skolithos-Glossifungites facies,” “Cruziana facies,”
“Zoophycos facies,” and “Nereites facies.’

The Cincinnatian ichnospectrum is illustrated in Text-
figure 29. As can be seen the Cubichnia, Fodinichnia, Dom-
ichnia, and Repichnia are well represented, while the only
Pascichnia are two specimens questionably assigned to
Paleodictyon. A comparison of the Cincinnatian forms with
those suites illustrated by Seilacher clearly reveals that the
Cincinnatian is a member of the “Cruzitana facies,” 1.e.,
shallow water.4* This shallow-water assignment of the Cin-

“This is not to imply that the Cincinnatian trace fossil assemblage is
identical to any previously described suite; as one would expect each
assemblage is unique. In this part I am comparing the relative de-
velopment of Cubichnia and Pascichnia.

402 PALAFONTOGRAPHICA AMERICANA (VI, 41)

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Ae veallf
ceo
/
thers ss
i feo
if? eas
4 4
\ 77 7
\ fl Ne Ades
t N os
y; ;
hs IS
Mel S
5 MV
ma 7 ~ =
ae \\ aS i
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si \ 7 ae
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Domichnia

bos : :

Fodinichnia

TA

Text-figure 29.—Cincinnatian Trace Fossil Suite. The chart
shows reconstruction of Cincinnatian trace fossils placed in their
ethologic categories. Cubichnia are numerous both in kind and num-
bers whereas the only Pascichnia are two specimens tentatively as-
signed to Paleodictyon. This distribution places the Cincinnatian with-
in Seilacher’s “Cruziana facies,” 1.e., a shallow water, marine environ-
ment.

a. Rusophycus pudicum (X 0.5); b. Rusophycus carleyi (X 0.25);
c. Rusophycus cryptolithi (X 1); d. Lockeia siliquaria (X 0.5); e.
Asteriacites stelliforme (X 0.3); f. Asaphoidichnus trifidum (X 0.25) ;
g. Trachomatichnus numerosum (X 0.25); h. Allocotichnus dyeri

(X 0.3); i. cf. Petalichnus multipartitum, oblique movement (X 0.5) ;
j. ef. Petalichnus multipartitum; straight-ahead movement (X 0.5) ;
k. Palaeophycus sp.; plan and cross-section views (X 0.3); 1. Rhabdo-
glyphus sp. (X 0.5); m. Tylichnus asperum (X 4); n. ?Paleodictyon
sp. (X 1); 0. Corophioides cf. luniformis (X 0.2); p. Corophioides
cincinnatiensis (X 0.2); q. Corophioides biclavata (X 0.2); r. Sko-
lithos delicatulus (X 0.5); s. Chondrites, type-B (X 0.2); t. Chon-
drites, type-C (X 0.25); u. Chondrites, type-A (X 0.25); v. Fasci-
fodina floweri (X 0.2); w. Trichophycus venosum (X 0.15); x. Phy-
codes flabellum (X 0.25); y. Dactylophycus quadripartitum (X 0.25).

TRACE FOSSILS CINCINNATI AREA: OsGoop 403

cinnatian is independently confirmed by several factors.
Hofmann (1964) reported mud cracks from the Corryville
beds at Stonelick Creek, as has Fox (1962) from the Rich-
mond of southeastern Indiana. In addition large amplitude
para-ripples are not uncommon in the Cincinnatian (see
Bucher, 1919), and these are generally considered to be
indicators of shallow water, The Cincinnatian is noted for
the tremendous numbers and variety of body fossils which,
of course, represent only a fraction of the original popula-
tions. Such diversity and numbers are encountered today
only in shallow seas. It is the author’s belief that the Cin-
cinnatian sea was less than 35 meters deep, and at times
the bottom may have been exposed to brief periods of
subaerial erosion.

The author is in basic agreement with Seilacher’s in-
terpretation of the “Cruziana” and “Neretites facies.” Cu-
bichnia most probably are burrows of concealment, and
these would be more numerous in shallow, well-lighted
water than in the abyssal deeps. Conversely Pascichnia are
indicators of organic material concentrated in thin layers.
It is reasonable to assume that thin layers of organic ma-
terial would be more likely to accumulate in deep water
where current and wave action is less apt to be as constant
than in shallow water. The lack of Pascichnia in the Cin-
cinnatian probably demonstrates that wave and current
agitation distributed the organic material evenly through the
sediment. This would explain why most Cincinnatian chon-
dritids and several other forms do not follow depositional
interfaces.

The author does not believe that the differences be-
tween the “Cruziana” and ‘“‘Nereites facies” is as clear-cut
as they appear in Seilacher’s (1964, fig. 7) chart, It is felt
that additional work on both fossil and Recent sediments
will cloud some of these sharp boundaries. For example,
Zenkevich and Birstein (1956) reported that investigations
of the Kuril Trench revealed that the relative importance
of plankton predators and detritus feeders increases with
depth, while the importance of larger predators decreases.
They found that below 10,000 meters predators are lacking
entirely. Their diagram of the number of various organ-
isms (Zenkevich and Birstein, 1956, fig. 4) shows that the
change in feeding habits is gradational. Thus, it is reason-
able to assume that Pascichnia and Cubichnia can occur
together at some intermediate depth. It is also possible,
that under proper conditions, they can coexist in either
shallow or deep water. Confirming the coexistence of Pas-
cichnia and Cubichnia in Recent sediments will not be easy,
because it is difficult to recognize Cubichnia as such on the
depositional interface (Seilacher, 1953b, pl. 11, fig. D). It

is likely that only through careful analysis of undisturbed
bottom samples can any conclusions be reached.

In spite of the evidence provided by the Iraqian sec-
tion discussed above, the author questions whether the
Zoophycos suite represents a true intermediate facies. It
is difficult to comprehend why the trace fossils found at
intermediate depths should be so markedly different from
the shallow-water and deep-water assemblages. Is it not
more reasonable to expect a mixture of the shallow-water
and deep-water communities? Moreover, at least two sec-
tions in North America bearing numerous Zoophycos are
considered shallow-water deposits. While Seilacher (1967,
p. 418) conceded that Zoophycos may occur in shallower
water “due to local channeling or restriction,” both the
sections discussed below are believed to represent regional
shallow-water deposits. Concerning the Chemung (Dev-
onian) strata of New York State, Cooper (1957, p. 271)
wrote, “the peculiar worm burrow Taonurus [Zoophycos]
is abundant in all the argillaceous sandstones and coarse
siltstones, especially in central and eastcentral New York
and parts of the Appalachians. The burrows are character-
istic of the shallow-water epineritic environment.” The
author, at present, is conducting research on the trace fos-
sils of the Waverly (Mississippian) strata of northeastern
Ohio. While the work is still in the preliminary stages some
observations are relevant to this discussion. The Cuyahoga
and Logan Formations consist of about 100 meters of inter-
bedded sandstones, siltstones, shales with some coarse con-
glomeratic beds, The lithology, paleogeographic setting, and
abundant plant fragments all point to a shallow-water ori-
gin. The most diagnostic trace fossils are Zoophycos and
Phycodes; Skohthos and Tricophycus may also be present.
All evidence indicates that this is a regional shallow-water
occurence of Zoophycos.

It is also worthy of note that Seilacher (1964, fig. 7)
reported Zoophycos from three deep-water deposits. More-
over in 1967 he confirmed the first occurance of Recent
Zoophycos in “deep sea” cores taken by the Lamont Labo-
ratory. Thus, it must be concluded that the validity of
Zoophycos as an accurate depth indicator is questionable.

It is also probable that additional studies of the
“Nereites” and “Cruziana facies” will result in the estab-
lishment of subassociations, 1.e., trace fossil suites dominated
by a few genera. In the Cincinnatian 7'richophycus veno-
sum, Corophtoides, and Chondrites are common, and they
frequently occur together in the same bed. Trichophycus
has been reported from the Ordovician of Norway and
Newfoundland, It would be interesting to know if it con-
sistently occurs in the same bed with Corophioides and

404 PALAEONTOGRAPHICA AMERICANA (VI, 41)

Chondrites. It might well be that these forms represent a
subassociation within the “Cruziana facies.”

Throughout this work the lack of information on Re-
cent traces has been stressed. While this deficiency makes
interpretation of individual fessil forms difficult, it is even
more apparent in a study of Spuren communities. It is
ironic that we have more data on fossil suites than Recent
associations. Nevertheless, the true meaning of fossil com-
munities can be learned only through an examination of
Recent environments. For obvious reasons the bulk of Re-
cent studies to date have dealt with the littoral environment,
one which is rarely preserved in the fossil record. More
research should be conducted on the physical and biological
aspects of the sediments of the neritic zone.

COMMENTS ON CINCINNATIAN PALEOECOLOGY AND
SUGGESTIONS FOR ADDITIONAL STUDIES

The value of trace fossils as paleoecological indicators
is most striking in sequences where other organic evidence
is lacking (e.g., Alpine Flysch). As mentioned previously,
the Cincinnatian has an abundance of evidence, and it all
points to a shallow water, open marine environment, The
trace fossils confirm both this and the recurrent nature of
the Cincinnatian fauna. As seen in Caster, Dalvé, and
Pope (1955), several species of body fossils reappear higher
in the local section, and much of the Corryville fauna is
again encountered in the Sunset beds. The most outstand-
ing recurrence in trace fossils is the 7richophycus-Coro-
phioides-Chondrites complex which is found in the Economy
of Twelve Mile Creek, the Southgate of Newport Shopping
Center, the Upper McMicken of Route 27 south of Newport,
West Fork Creek and Wesselman Road, the Corryville of
Stonelick Creek, the Sunset of Four Mile Creek (3.7 miles
south of Oxford, Ohio), and the Liberty of Lick Run. This
indicates a repetition of bottom conditions and preservation
favorable to these three forms.

The author has also been impressed by the amount of
subaqueous erosion as evidenced by truncated burrows.
Free tubes have never been found in any specimens of Coro-
phioides; and Trichophycus venosum most frequently oc-
curs as “turkey tracks,” z.e., only the basal portion of the
burrow. Several other examples (e.g., Lockeia, Fascifodina)
could also be cited. The evidence provided by trace fossils,
coupled with the many scour channels, indicates frequent
periods of erosive activity,

Additional research on Cincinnatian traces should prove
most rewarding. An attempt should be made to determine

if there is any vertical variation in the suite. To date the
most notable change is that Palaeophycus, type-C, is more
common in the calcarenites of the Eden and is less frequently
encountered higher in the section. Weiss, et al. (1965) and
Ford (1967) showed that the Eden rocks in the vicinity
of Cincinnati contain a greater percentage of lutite (75
per cent+) than the Maysville strata (40-65 per cent).
Conversely calcarenties are thicker and more important in
the Maysville. Although no exact data are yet available
for the Richmond, the upper part of the Cincinnatian sec-
tion is generally considered to contain a greater percentage
of calcarenties than does the lower portion. It would be in-
teresting to know if this lithologic change is reflected
by a variation in the number and kinds of traces.

Just as Seilacher (1964) traced Spuren communites
laterally in the Upper Paleozoic of Oklahoma, the local
Cincinnatian suite should be compared with the trace fos-
sils of its lateral equivalents. The most fruitful field of
investigation should be the Martinsburg Shale of the Ap-
palachian Geosyncline. The Martinsburg represents a mud-
dier, more unstable environment, and there may well be
significant differences between the two communities.

A considerable amount of work still remains to be
done for individual species. Although the local traces are
useful as guide fossils only for short distances, additional
studies may indicate that some forms can be employed for
long distance correlation, The Trichophycus-Corophioides-
Chondrites association has been found just below the top
of the McMicken at three widely separated localities. Fur-
ther work would reveal whether this is merely a coincidence
or whether a single bed covers a wide area. Likewise, Fasci-
fodina has been found only in the Corryville beds of two
localities 10 miles apart. Perhaps the lateral range of this
genus can be extended by careful collecting of the Middle
Corryville. Such work will also provide greater insight into
facies relations within the Cincinnatian.

As mentioned previously, the magnitude of the present
study has resulted in a superficial coverage of some genera.
The morphology of forms such as Chondrites, type-B, Tri-
chophycus, and Tylichnus is not completely understood. It
would be well to test techniques which have not yet been
used in palichnological studies, 1.e., radiography and amino
acid analyses. X-ray defractions analysis proved valuable
in the interpretation of Chondrites, type-A, and this method
should be used on other forms as well. Only through the
employment of a wide spectrum of geologic and paleonto-
logic tools can this difficult group be understood.

TRACE FOSSILS CINCINNATI AREA: OsGoop 405

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408 PALAEONTOGRAPHICA AMERICANA (VI, 41)

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PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 57
Page

Rusopliycus carleyeN (Ys Wo Mawes) el S 85. scecc see cane ores sace bakes cease eaeee ete teraeenecese eee 302, 306
Cubichnia of Isote/us; convex hyporelief which demonstrates the inseparability

of body fossils and trace fossils. Casts of the genal spines, cephalic and pygidial
doublures, hypostome and proximal portions of the walking legs, lateral lobes

are imprinted. Gift of George Walker. Corryville beds, Stonelick Creek,
Clermont Co., O. UCM 37598, X 0.7.

Asteriacites stelliforme (Miller and Dyer), 1878 -............s::-sscessserssecsseeecesnseenensesencenen 312, 313
Cubichnia of asteroid; convex hyporelief showing vertical repetition. Corryville
beds of Stonelick Creek, Clermont Co., O. UCM 37599, X 1.

Dactylophycus quadripartitum Miller and Dyer, 1878 —..... sSrace See ab Naste case ener cee 345, 346
Fodinichnia; convex hyporelief showing bilobation and faintly defined annuli.
Syntype. Eden beds, Cincinnati, O. HBM 3174; latex mold UCM 37600, X 1.

Phycodes flabellum (Miller andgDyer), 1878 ...........--ccecccsesessesseeeeesenneseeeeenecnnceeee 341, 343
Fodinichnia; cross-section showing filling of the proximal portion of the burrow,
i.e. the master shaft. Locality unknown. UCM 37667, X 1.

Tsotelus maximus Locke, 1838
View of ventral surface showing preserved appendages. Note that coxae are much
longer than those seen in Rusophycus carleyi (fig. 1 above). Hypotype. Richmond
beds, Oxford, O. USNM 33458, X 0.7.

Rusop hay cits) Duce cerry EMA USS 2 pee ee acne oeac ree alen nner e eee age ceetnne ares 304, 305
Cubichnia of Flexicalymene mecki; one of three specimens known from the Cin-
cinnatian where the producer of the burrow (Flexicalymene mecki) is preserved

in situ. Corryville beds of Stonelick Creek, Clermont Co., Ohio. Specimen collec-

tion of William White, Jr., Milford, O. Latex mold UCM 37641, X 1.5.

Lockeia stligwariat UW. P:) Yates yy W879 eae eee cee serene eset eee ace ener eee 308
Cubichnia of pelecypod. Note linear alignment of some forms. Paratype. Upper-
most Trenton or lower-most Cincinnatian, Ludlow, Ky. UCM 37597, X 1

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 57

Plate 58

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Figure
ze

3-6.

TRACE FOSSILS CINCINNATI AREA: OsGoop

EXPLANATION OF PLATE 58

Page
Pee CEA DEO LLL I ED RTAMIES SRE cae oe eee a ee anes seestic 304, 307
Cubichnia of Cryftolithus; convex hyporelief showing a ‘specimen with an
undercut forward wall (top). Note the casts of the impressions of the genal
spines. Economy beds, Humphrey’s Branch, Twelve Mile Creek, Campbell Co.,
Kentucky. Specimen in collection of William White, Jr., Milford, O.; latex
mold UCM 37601, X 1.25.
2. Convex hyporelief showing coarsely striate outer lobes and finely striate inner
lobes. Holotype. Locality same as fig. 1 above. UCM 37587, X 1.5.
TGP NE ELS PET creer V8 aN Gel OGY, es oe ee Be ON eR 304, 305

3. Cubichnia of Flexicalymene; convex hyporelief showing horizontal repetition.
Corryville beds, Taylor Creek between Miamitown and Dent, O. MU T88;
latex mold UCM 37709, X 2.

4. Convex hyporelief of fig. 5 below, X 1.4.

5. This remarkable specimen reveals the originator of the trace, Flexicalymene
mecki, in situ. Corryville beds of Stonelick Creek, Clermont Co., O. UCM
37574, X 1.2.

6. Cubichnia of ?Calymene; convex hyporelief of Syntype. Note that this speci-
men closely resembles those seen in figs. 9-10 below. Clinton beds, Oneida Co.,
N.Y. USNM 41113; latex mold UCM 37581, X 0.8.

ISMEMINIE ND DUS CIES mAsPL COL (eer E sie fATILGS))'y, ILS BS, csu.cccctemesee tect ecenck lvascoyoelsxecesateseesy2cscssasecievovecsvactee senate 306
7. Cubichnia of Jsotelus; convex hyporelief showing a well-preserved cephalic
or pygidial doublure. Corryville beds, Stonelick Creek, Clermont Co., Ohio.
Specimen in collection of William White, Jr., Milford, O.; latex mold UCM
37710, X 0.8.
8. Convex hyporelief showing nodes attributable to the coxae of Jsotelus. Local-
ity unknown. MCZ 551; latex mold UCM 37596, X 0.9.
GLB. RETREATED FATT CET iy UES EY A eR ones ee 308

9. Convex hyporelief which shows a dense grouping of the resting traces. It is
impossible to determine how many trilobites were involved. Locality un-
known. MU 1T$87; latex mold UCM 37641, X 0.7.

10. Convex hyporelief associated with six specimens of Lockeia siliquaria. The
traces occur at the bottom of a channel filling. Mt. Hope beds, Stonelick
Creek, Clermont Co., O. UCM 37642, X 1.1.

41

J

414

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 59

Figure Page

L: Lockeia siliquarta. UW. Bs Vannes; 0879) ccsccceseccsceccncn nna. ceauctesasasansteczsctoces toeea-acee eee aeec meee peeeateeaes 308
Cubichnia of pelecypods; etched cross-section showing that the burrow is
truncated and does not extend up into the host rock. opotype. Uppermost
Trenton or lower-most Eden, Ludlow, Ky. UCM 37675, X 4.5.

2: Astertacites stelliforme (Miller and Dyer), US78) cccecceeeecenssps eee ee 312, 313
Cubichnia of asteroid; showing striae made by the tube feet. Holotype. Maysville
beds, Cincinnati, O. HBM 3175; latex mold UCM 37568, X 1.25.

3. SRusophycus” subangulature EVAN US Se ee ae eaten tees ce neee eee sie ee ee eee ee 305
Repichnia of trilobite; convex hyporelief. Such elongated burrows are not common
in the Cincinnatian. Syntype (Hall, 1852, pl. 8, fig. 2b). Clinton beds, Oneida Co.,

N.Y. USNM 41112, X 0.6.

4.. Rusophgycus cryplolitht, ri Sp. <cccrcc-ctsc-cacdeccsssem eer ree eee eee ence ee 307
Cubichnia of Cryptolithus; convex hyporelief showing groupings of casts of four
shallow burrows showing vertical repetition. Paratype. Economy beds of Duck
Creek, Campbell Co., Ky. UCM 37567, X 1.

Be Rusophycus: pudvewm Fall), V852 2 srereccccceeecteecece es ceneetesev cere es easatenenerce snes sue eteet actrees meres eeereners 305

“I

Cubichnia of ?calymenid; convex hyporelief. Syntype. Clinton beds, Oneida
Co., N.Y. USNM 41115, X 1.

Rusophycus carleyt (Ye Be James), US 85s se ccecscececcctees co eees eet eeeseeee eres en tesco ener eee 304, 306
Cubichnia of Jsotelus; convex hyporelief showing largest Rusophycus known.

Note the well-defined genal spine and pleural imprints. ?Maysville beds,

near Bantam, Clermont Co., O. UCM 37566, X 0.4.

Locketwa siliguaria UO: Ps, James, (0879 cc ee ee se eee eer eee 308
Cubichnia of pelecypod; convex hyporelief. ?Economy beds of Twelve Mile
Creek, Campbell Co., Ky. UCM 37577, X 1.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 59

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 60

Figure

ils

3,4.

TRACE FOssILS CINCINNATI AREA: OsGoop

EXPLANATION OF PLATE 60
Page

Corophioides cincinnatiensis, 0. SP. .....c...0-0.-20-00 2-0. es Se ne Re ER Le 314
Domichnia; U-plane section showing an annulate irregular arm. Paratype.
McMicken beds of West Fork Creek, Mt. Airy Forest, Cincinnati, O. UCM 37602,

Xe1e

IR PITY DL REDICS FOYE Lie Ve EMU TRV ae os Se oi a nT 305
Cubichnia of trilobite; convex hyporelief showing horizontal repetition. South-
gate strata of Rapid Run, Hamilton Co., O. UCM 16448, X 0.6.

Gorophioidess Giclawata \(Miallers),, W875) ccc. .accccecsccecstccecencestuceecesceveoves-e-e PN ae 314, 323

3. Domichnia; epirelief showing a modified arm and the remnants of the
Spreite. Corryville beds of Stonelick Creek, Clermont Co., O. UCM 37659,
X 0.75.

4. Hyporelief showing modified arms and Spfreite. Locality unknown. UCM
37658, X 1.

GOAT A PADILLA TLON AHL 5 Sy SO), Seo ey Coe a Pa nee 314, 321
U-plane section. Paratype. McMicken beds, U.S. Route 27, 1.4 miles south of
Newport Shopping Center, Southgate Ky.: UCM 37592, X 0.8.

Corophioides polyupsilon John Smith, 1893 ............ ence re IRE Steet nee a ee 314
Domichnia; U-plane section of holotype demonstrating that the Spreiten abut
against the arms instead of forming a “U in U” structure as Smith believed.

Upper Carboniferous beds near Kilwinning, Scotland. HM X59; latex mold UCM

37572, X 0.75.

litanopigamsacaeroasam Wallet, 8 79) sec. 5.2. tees 2acee-gc¢eesvaes esse hee slnteeth eee. Beet cee tee ee 347
Fodinichnia or Domichnia. A “turkey track” truncating another individual con-
taining remnants of the burrow filling. Humphrey’s Branch of Twelve Mile
Creek, Campbell Co., Ky., some 30 meters upstream from the type locality of
Rusophycus cryptolithi. UCM 37575, X 0.5.

Corophioides cincinnatiensis, nN. Sp...
U-plane section showing marked basal expansion. Locality same as fig. 5 above.
Specimen destroyed by subsequent sectioning; X 1.5.

415

416

Figure

1-3.

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 61
Page

Corophioides, biclavata (Miller), VS875i 2. eerte eee 314, 323
1. Domichnia; concave epirelief showing that the connecting bar does not extend
to the terminal depressions. Locality unknown. UCM 37656, X 0.8.
2. Convex hyporelief. Locality unknown. UCM 37655, X 1.
3. Concave epirelief. The inner wall of the large terminal depressions is under-
cut. Corryville beds of Stonelick Creek, Clermont Co., O. UCM 37657, X 0.75.

Corophioides cincinnattensts, Wh. Spo: secon cette ees eee et eee 314, 321
Domichnia; U-plane section of holotype showing the irregular course of the

arms. McMicken beds of West Fork Creek, Mt. Airy Forest, Cincinnati, O.

UCM 37590, X 1.

Gorophtotdes cf: lumformis ((Blanckenhorn) we 320
Domichnia; concave epirelief showing numerous closely spaced U-shaped de-
pressions. Waynesville beds, Flat Fork of Caesar’s Creek, Harveysburg, Warren

Co., O. MU T89; latex mold UCM 37711, X 0.75.

Gorophiotdes cincinnatiensts, W.0Spe qstece -:esceesette enone ee 321
U-plane section showing marked basal expansion. Paratype. McMicken beds, U.S.

Route 27, 1.4 miles south of Newport Shopping Center, Southgate, Ky. UCM

37589, X 1.

Corophioides biclavata (Miller), 1875
Convex hyporelief showing filling of the dumbbel! and remnants o
Fulton beds, Union Levee, Cincinnati, O. UCM 16566, X 1.

the Spreite.

Gorophioides cincinmatiensts, ms Sp: sec cecscce-cs- cesses eee 323
U-plane section. Paratype. Locality same as fig. 6 above. UCM 37591, X 0.9.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 61

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 62

Figure
aveze

3,4.

7,8.

TRACE FOSSILS CINCINNATI AREA: OscGoop

EXPLANATION OF PLATE 62
Page

Corophioides cf. luniformis (Blanckemhorn) .......-.-2...<.:0.c:..-00sssceccsesececeecessesneesesecessees 314, 317
1. Domichnia; U-plane section. Locality unknown. UCM 37611, X 1.
2. Concave epirelief; probable mold of the base of the “U.” Locality unknown.

UCM 37654, X 1.

Corophioides cincinnatiensis, n. sp. ....

3. Domichnia; U-plane section showing details of the R
type. McMicken beds of West Fork Creek, Mt. Airy Forest, Cincinnnati, O.
UCM 37624, X 1.25.

4. Latex mold of the basal portion of the U showing close-spaced annuli.
Paratype. McMicken beds, U.S. Route 27, 1.4 miles south of Newport
Shopping Center, Southgate, Ky. UCM 37589, X 1.

SEA steLiacties sreliporme (Miller and Dyer) 1878 2ccscc.ccntcecccsec-ccceccsancctsaceseessececsoctencests 312, 313
Cubichnia of asteroid; convex hyporelief. Locality unknown. UCM 37674, X 1.
(OF OP STONE GIG TUSTIN Yeo cee oe ate Ee a EE 314, 321

U-plane section of incomplete specimen. Paratype. Locality same as fig. 4
above. UCM 37625, X 1.

MAGEE ADO: CLAN EG TARTS. Wie De TE Wa Cel Fy 03 US ep eo 326, 327

7. ?Domichnia; view of lower (hyporelief) surface of presumed hypotype (J. F.
James, 1892, fig. 15). Richmond beds of Dearborn Co., Ind. UCM 37673,
X 0.75.

8. Enlarged view of a portion of above showing calcite fillings and encircling
depressions, X 3.

417

418

Figure

~I

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 63

Page
Cf. Palaeophycus Hall, 1847 ............... eee eee Sie pO SR 328
Repichnia; concave epirelief showing the sinuous nature of the trail. Southgate
beds of Newport Shopping Center, Newport, Ky. UCM 37637, X 0.75.
Chondrites,“type-C: 24.255 2.20 ee ee, ee eek eer ...328, 339
Fodinichnia; concave epirelief’ Whitewater beds of Short Creek, three miles
southeast of Richmond, Ind. MU 193; latex mold UCM 37636, X 1.
Ghondrites, ‘ty pe=B. -<.cxc, sesscis5. 22) Bee 328, 336
Fodinichnia; vertical section showing a portion of the network. Note upward
curvature of some of the branches. Upper Richmond beds, State Route 32, six
miles southwest of Flemingsburg, Ky. UCM 37665, X 0.8.
Gonopliotdes, cincinmatiensis, Ni, Shae ee 321
Domichnia; U-plane section showing association of C. cincinnatiensis and Chon-
drites, type-A. Paratype. McMicken beds, U.S. Route 27, 1.4 miles south of New-
port Shopping Center, Ky. UCM 37680, X 0.6.
Ghondrttes gracilis evar crassa ilalle\85 2s ee 329
Fodinichnia; slab shows only a portion of the network. Syntype (Hall, 1852, pl. 5,
fig. 3c). Clinton beds, Oneida Co., N.Y. USNM 41134, X 1.
Lo cketas stliquania ).. P-ii| ares, i USi/9 ieene ere eee ee 308
Domichnia of pelecypod; vertical section showing disturbed column of sediment
overlying the almond-shaped burrow filling. ?Economy beds of Twelve Mile
Creek, Campbell Co., Ky. UCM 37679, X 1.3.
Dactylophycus quadripartitum Miller and Dyer, 1878 .....c...0:00-:cccseeeee020ceeeeeee eevee 345, 346

Fodinichnia; convex hyporelief. The master shaft presumably extended to the
upper left. Economy beds, West Covington, Ky. USNM 40027; latex mold UCM
Siig NGL

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 63

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 64

Figure

4,5.

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TRACE FOSSILS CINCINNATI AREA: Oscoop

EXPLANATION OF PLATE 64

Page
Mu ciusopsrsesiulcatzrin Wiallervand! Dyer)! L878 <.ctesc-c-<c0r-nnceveeodecctvet eeeatascestvovce dencenecexcesiee 380, 381
Repichnia; convex hyporelief. Locality unknown. UCM 37616, X 1.2.
(CLADE ELAN ., - TSN OA GS cece ce sete RR ee 328

Fodinichnia; latex mold of a concave epirelief. Note the circular grouping of the
burrows in the lower left. Whitewater beds, Short Creek, three miles southeast
of Richmond, Ind. MU 192; latex mold UCM 37615, X 1.4.

CH EDTHEF TOG ay A SNM acl Sart ee oe Cee oa SCE 328, 336
Fodinichnia; vertical section showing what is interpreted as a portion of the

main shaft of the burrow. Upper Richmond beds, State Route 32, six miles south-

east of Flemingsburg, Ky. UCM 37665, X 1.2.

(OLN LETT NESTA CI SS aoe cree oe ere ee i 328, 339

4. Latex mold of epirelief showing some degree of bilateral symmetry. Local-
ity same as fig. 2 above. MU 191; latex mold UCM 37614, X 1.

5. Latex mold of epirelief. One system of burrows (upper left) is oriented
parallel to the bedding plane; the other system (lower right) obliquely to the
bedding plane. Locality same as fig. 2 above. MU 190; latex mold UCM
37678, X 0.7.

Chondrites, type-A........... Seppe PARE ACERS PORE en er tee ee ee 328, 335
Fodinichnia; epirelief showing a compact group of vertical burrows. Southgate
beds, Newport Shopping Center, Newport, Ky. UCM 37612, X 1.2.

(UNDE TAT EH AEG i ~ CEA TIQHLE 7 SERENE es SU re ie ee ee 328, 335
7. Epirelief showing a dense grouping of burrow systems. Locality same as fig.

3 above. UCM 37613, X 0.5.
8. Typical arrangement of C., type-B structure. Locality unknown. UCM 37666,

xt

419

420

Figure
1-4.

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PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 65
Page

Phycodes flabellum (Muller ‘and Dyer); 1878)eyc cacti 341, 343

1. Fodinichnia; convex hyporelief showing portions of more than one burrow sys-
tem. Holotype. Richmond beds near Lebanon, O. HBM 3178; latex mold UCM
37621, X 1.

2. Vertical cross-section showing “retrusive Spreiten.” Note that the burrow
truncates the clay inclusions. Locality unknown. UCM 37684, X 2.7.

3. Vertical cross-section; disturbance of the bedding caused by burrow penetra-
tion (center bottom). Corryville beds, Stoneiick Creek, Clermont Co., O. Speci-
men destroyed by subsequent grinding. X 1.

4. Vertical cross-section showing master shaft extending nearly to the base of
the host rock. Locality unknown. UCM 37672, X 1.3.

Phycodes circimatum ‘(Reimbard) Richter L850) eee eee 341
Convex hyporelief of a sparsely branching specimen. “PAycodes beds” (Ordo-
vician), Saalfeld, Thuringia. UCM 37526, X 1.3.

Gorophiotdes' ‘biclavati, \((Milleri)) 08753 aces eons eee eee 314, 323
Domichnia; concave epirelief. Note that only the left end of the structure has
been modified. Locality unknown. UCM 37648, X 0.6.

Repichnia; convex hyporelief interpreted as the Cruziana-like trail of Crypto-
lithus; also see Text figure 24. Location unknown. UCM 37622, X 1.

Chomd rates, ty PO=B sesso tassseds ca seesncasesteisctcxcitpesast ete a echo aac 328, 336
Fodinichnia; latex moid showing the characteristic rigidity of the branches noted

by Simpson (1957). Corryville beds of Stonelick Creek, Clermont Co., O. UCM

37623, X 1.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 65

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Figure

. Paleodictyon sp. Savi and Meneghini, 1850

TRACE FOSSILS CINCINNATI AREA: QOscoop

EXPLANATION OF PLATE 66

Pascichnia; convex hyporelief showing a well-preserved netw ork.
near Trieste, Italy. UCM 37507, X 0.8.

eV cOden milan ell rim (MILE DEAT IDYEL)y SUB TS cecctshexqnccec-cccevasevoceenscecupezececsEasestesceesses ....341, 343
Fodinichnia; convex hyporelief showing a poorly organized s system. Note that the
flabellate pattern characteristic of many Cincinnatian representatives of the

genus is lacking. Corryville beds, Stonelick Creek, Clermont, Co., O. UCM 37635,

>-eoils

Peaasannlegn ang: prvabatiarg VEIN TS) Ee ae eee A cone nae ee ere peo ee 305
Cubichnia of moderate-sized trilobite ; convex hyporelief. The morphology exhibit-

ed by this specimen is intermediate between Rusophycus and Cruziana. Such
elongated forms are not common in the Cincinnatian. Corryville beds of Stonelick

Creek, Clermont, Co., O. UCM 37633, X 1.

Phycodess plavelium (Willer tad) Dyer), 1878) c...ccc...sce-ccecececnccsscececc-neeertec-cscseeosaresenese==> 341, 343
Fodinichnia; convex hyporelief showing a system with radial symmetry. The
large protuberance marking the center of the system is not thought to be gen-
etically related to the Phycodes burrow. Locality unknown. UCM 37631, X 0.75.

OUD NEED LICS, LANAI OAV ONG ITV 2) OU JS Eee 323
Domichnia; concave epirelief. Corryville beds of Stonelick Creek, Clermont Co.,
O. UCM 37634, X 1.

ASCE OYUN LODO ETE NIN NETS, WTS SP =e sce ccscste coe aan neo cecnceaeaenccctseecguscetsneacosuch¥ena-—ecscacenbastessetes 340, 341
Fodinichnia; epirelief showing concave vermiform markings surrounding the
convex bundled shafts. One of the branches (lower right) leads directly into

one of the concave markings. Holotype. Corryville beds of Second Creek where
crossed by Cozzadale Road, Clermont Co, O. UCM 37632, X 1.2.

igitchnmonaspermm, (Miller and) Dyer); U8 78) o.oo cscscsc. sca cscccccccecsdeceventascstnenostectsestcsececenson 372
Repichnia; convex hyporelief showing several super-imposed trails. Note monili-

form casts of ?fecal strings in bottom center. Locality unknown. MCZ 1054; latex

mold UCM 37687, X 2.7.

421

422

Figure

Ie

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 67

Page
?Paleodictyon sp. Savi and Meneghini, 1850 384, 386
Pascichnia; convex hyporelief showing unequal develop ickness
of the walls. McMillan beds of Ravine Street, Cincinnati, O. ‘UCM 174315 XS 175s
“Blastophycus diadematum” Miller and Dyer, 1878 ........c--ccecce-c-ecoeecoeessecceesseeseeessesseeeeeeos 390

Plaster cast showing scour marks associated with an enrolled trilobite, Flexicaly-
mene meeki. Experimentally produced. UCM 37640, X 1.

Buthotrephis palmata Hall, 1852
?Fodinichnia convex hyporelief. View of syntype illustrated by Hall (1852, pl.
7, fig. 1b). The specimen differs considerably from the other species of Butho-
trephis proposed by Hall. Clinton (Silurian) beds of New Hartford, N.Y. USNM
41083, X 1.

Chondrites,. ty PeaA\ secsicces occ ctuseren oe en 328, 335
Fodinichnia; vertical section showing that there is ell-defined central axis.
McMicken beds, West Fork Creek, Mt. Airy Forest, Cincinnati, O. UCM 37607,

XG:

Rusophycus- clavaturn Wall 1852) ee. sae eee ee oe ee 305

Repichnia; convex hyporelief. The podlike outline of the burrow is attributable
to a “back and forth’ movement by the originator. In eee the specimen lies
closer to Cruziana than Rusophycus. Syntype. (Hall, 1852, pl. 8, fig. 1a). Clinton
(Silurian) beds of Oneida Co., N.Y. USNM 41114, X 1.

Walcottia rugosa sMilleryand yD yer 518/78 ps eee ee 379
Repichnia; convex hyporelief showing chevron-shaped bodies probably made by
parapodian-like appendages. Ovoid body is believed to represent the anterior end

of the “body.” Maysville beds, Cincinnati, O. Syntype. MCZ 545; latex mold

UCM 37677, X 2.9.

Buthotrephis palmata’ Wally 8S2) ceo ee ce 331
?Fodinichnia; convex hyporelief. This species is not present in the local Cincin-

nation strata. Syntype. Clinton (Silurian) beds of New Hartford, N.Y. USNM

41079, X 0.5

PALAEONTOGRAPHICA AMERICANA, VOL. Plate 67

Plate 68

PALAEONTOGRAPHICA AMERICANA, VOL. VI

TRACE FOSSILS CINCINNATI AREA: OsGoop 423

EXPLANATION OF PLATE 68
Figure Page

Hh, TEFEN NED: TETAS TIED ING UNVEILS, Se cee ee IO) FON AE a sn 347
Fodinichnia or Domichnia, lateral view of burrow filling showing coarse striae.
Upper surface of the burrow is to the right. Fulton beds of Twelve Mile Creek,
Campbell Co., Ky. UCM 16574, X 2.

Damliricnopryrus lanoswmn Nillers and. sD yer, 1878) <.actccestccs.- cc! cdeccsesescs ccc bosstecenezeecseese! 347, 350
?Repichnia; convex hyporelief. View of syntype showing bulblike expansion
which led J. F. James (1885, 1891) to compare the species with Blastophycus
diadematum. Eden beds, Cincinnati, O. HBM 3185; latex mold UCM 37595,
X 0.3.

3-7. Trichophycus venosum Miller, 1879 :

3. Convex hyporelief showing delicate striae. Locality unknown. UCM 37681,
X 0.9.

4. View of the upper surface of figure 1 above showing parallel discontinuous
striae. Note the slight constrictions in the diameter of the burrow. X 0.9.

5. Convex hyporelief showing a nearly complete burrow filling. Locality un-
known. UCM 37629, X 0.8.

6. Lateral view of burrow cast showing two secondary branches. Locality un-
known. UCM 37593, X 1.

7. Ventral view of burrow filling showing at least two well-developed sets of

striae. Locality unknown. UCM 37628, V 1.25.

8. Trichophycus lanosum Miller and Dyer, 1878 ..... Es irre eee 350
?Repichnia; convex hyporelief View of syntype showing highly sinuous burrow
and button-like ?anterior end. Eden beds, Cincinnati, O. HBM 3183; latex mold
UCM 37594, X 2.

OuePhyeodes plaveliurme ((Mallenvand! Dyer): US7.8) scccccccc cece cece ctcescnceecseceesc be ccteaosercanceomsons socceatees 341
Fodinichnia; convex hyporelief showing the typical flabellate pattern. Locality
unknown. UCM 37670, X 1.

424

Figure

iE

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PALAEONTOGRAPHICA AMERICANA (YI, 41)

EXPLANATION OF PLATE 69

Page

Ghondrites: ‘succilems: \(Tiall)) (US47 sc. soctsanceccevacucosencucscssosonccuszeseesetuuseceraerercetie ere ptites- enexre are 328
Fodinichnia; convex hyporelief of syntype (Hall, 1847 pl. 22, fig. 2a) showing
nodose structure of branches (upper left). Trenton beds, Glen Falls, N.Y.
USNM 41133; latex mold UCM 37639, X 0.6.
Dactylophycus quadripartitum Miller and Dyer, 1878 -........2.:-.::::.c:0:cseseceeceereereeeneee 345, 346
Fodinichnia; convex hyporelief. View of syntype showing the bilobate, annulate
nature of the burrow. Eden beds, Cincinnati, O. HBM 3174; latex mold UCM
37608, X 1.3.
Coraphtiotdes sche Vimtpormnts, 2 cxccceses ta cceteneca nt cteceer- co ene senses oy een enorme east 314, 317
Domichnia; U-plane section of two specimens. McMicken beds of West Fork
Creek, Mt. Airy Forest, Cincinnati, O. UCM 37661, X 0.9.
Phycodes flabellum (Miller and Dyer), 1878 .........-.-.:.---..:cscscse<cosceessevensescsessensseensesenes 341, 343
Fodinichnia; convex hyporelief. Locality unknown. UCM 37609, X iL
Wealcottia rugosa Miller“and) Dyer,” 1878 oeeccccce staccato eee ere 379
Repichnia; convex hyporelief. The burrow closely resembles the impression that
might be made by a Nereis-like annelid, Anterior end is believed to be to the left.
Syntype. McMillan beds, Cincinnati, O. MCZ 545; latex mold UCM 37677, X 3.1.
Bl ae sp. Savi and) Memephimiy US5 0 2eeoscecee ca ccc ac scenee cece cee cee eee eee 384

Latex mold of impression achieved by rolling a specimen of Pasceolus darwini

over wet plaster. Compare with Cincinnatian representatives of ?Paleodictyon

(see Pl. 81, fig. 2 and Pl. 67, fig. 1). UCM 37610, X 2.2.
7. ?Pascichnia; convex hyporelief showing that hexagons are elongated parallel

to the current direction. Note arcuate trails of similar dimensions in upper

left. Eocene beds near Trieste, Italy. UCM 37505, X 0.9.
Chondrites gracilis. -(Flall)); USS 2) 2.2 2c sce cc cccec aes c screenees te are canes eee ree mee 328

Fodinichnia; latex mold of syntype showing rigidity of the branches. This form
is not directly comparable to any of the varieties of Cincinnatian chondritids.
Clinton (Silurian beds, New Hartford, N.Y. USNM 41125; latex mold UCM
S676) Xue:

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 69

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Figure

il

2, 3.

‘TRACE FOSSILS CINCINNATL AREA: Oscoop

EXPLANATION OF PLATE 70
Page

Rucusopsisesuicatum (Miller and) Dyer), 0878) -...-..ccccscecn---c0.-cccensescccececvcsceesceorcseersesces 380, 381
Repichnia; convex hyporelief showing tension cracks produced by the pressure of

the burrow on the sole of the bed. Bellevue beds, Hall’s Creek, Morrow, O.

UCM 37606, X 0.7.

“Dystactophycus mamillanum” Miller and Dyer, 1878 ...........-----::::+-s---seseseeceseseeseseseeeeseeees 391
2. Inorganic; convex hyporelief. The irregularity of the arcs was probably
caused by shell fragments which were swept around with the stem. Richmond
beds, locality unknown. UCM 37605, X 1.
3. Concave epirelief of figure 2 above showing a more regular series of mark-
ings. A yertical section taken through the center of the specimen revealed the
partially disarticulated columnals. X 1.

DP ened ay NP erTICY RADIO 7 IEW g SUEZ A) a eee ee eee ee 347
Fodinichnia or Domichnia; epirelief showing burrow packings in a “turkey
track” depression. Field photograph. Corryville beds, Stonelick Creek, Clermont

Co., O. X 0.2. (Silver coin is a quarter.)

ORO PUAOI ME API Claciata m(OMIINET)) ye UST) eee ec ssc eeveetentacacdeceovasaseonsoesecesucensosisaceenecsaesecscoreesese 314, 323
Domichnia; convex hyporelief showing the base of the “U” and the fillings

of the extended U-arms. Mt. Hope beds of Rapid Run, Hamilton Co., O. UCM

37662, X 1.

Phycodesplaneliund (Miller and Dyex)), U8 78) <tc -<ccceececccncccscsesnceeecncseoetescceceedeceectscneeect 341, 343
Fodinichnia; convex hyporelief showing a flabellate specimen. Liberty beds of
Talawanda Creek where it enters Four Mile Creek just above the Butler-Preble
County line, north of Oxford, O. MU 195; latex mold UCM 37686, X 1.

426

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 71

Figure Page

L. ‘Rusophycus carleyr: (J. ha Jamies), S85) eee 306
Cubichnia of Jsotelus; convex hyporelief. Presumed holotype showing nodes
revealed by a mesial parting of the lateral lobes. Anterior end at top. ?Maysville
beds near Bantam, Clermont, Co., O. UCM 37604, X 0.7.

2. Dylichnus asperum (Miller ‘andi Dyer), eU8 78 essen neers eee 372
Repichnia; convex hyporelief showing segments of several burrows. For an en-
largement of the nodose prosopon see Text-figure 28. Holotype. Eden beds, Cin-
cinnati, O. HBM 3179; latex mold UCM 37588, X 2.

3: Palaeoscia‘flowert: Caster, 9420x2202. tie 396
Fodinichnia or inorganic; convex hyporelief of a slightly ellipsoid specimen.

The furrow leading from the center of the specimen to the lower margin may
represent the mold of an agglutinated tube. Corryville beds, Stonelick Creek,
Clermont Co., O. UCM 37586, X 1.3.

4. @Corophkroides ‘biclawata (Muller), S185 ee eee cae 314
Domichnia; convex hyporelief. This specimen may represent a cast of the modi-
fied base of Corophioides cincinnaticnsis or it may be the cast of the upper por-
tion of the tube. Locality unknown. UCM 37660, X 1.1.

St ucusopsis sulcatum (Maller, and! Dyer), ol S7See ee ee ee 380, 381
Repichnia; convex hyporelief. Holotype of Trichophycus sulcatum. The basal
lamina of the host rock “laps up” on the side of the burrow. Eden beds, Cincin-
nati, O. HBM 3182; latex mold UCM 37705, X 1.

6, Asaphordichnus trifidwr (Miller. oVS80) ose eee 356

Repichnia of /sotelus; epirelief showing well-developed trifid imprints of the
right series. Holotype. Compare with Plate 72, fig. 3. Right-hand series was il-
lustrated by Miller (1874, p. 136, fig. 11). Eden beds of State Avenue, Cincin-
nati, O. MCZ 540; latex mold UCM 37704, X 0.8.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 71

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 72

TRACE FOSSILS CINCINNATI AREA: OsGoop

EXPLANATION OF PLATE 72
Figure Page

2s a aa Cen (UNMET) TIGR) -eeeceace er cece este eee 359
- Repichnia of ?/sotelus; convex hyporelief. Holotype. Movement was from
bottom to top and the body was angled to the right of the axis of movement.

Eden beds, State Avenue, Cincinnati, O. MCZ 543; latex mold UCM 37693,
Dslr

2. Convex hyporelief showing only three pairs of imprints in each set. If the
imprints are rotated 45° clockwise, the resulting configuration resembles
fig. 1 above. Paratype. Eden beds of Mud Lick Creek near Walton, Ky. UCM
37627, X 1.2.

EMEA SIUPNOINALGI TUG ETT LCM @ WALL emp L88 Oly secccte tees cevcceccncoe conse cSbeeencdc woe. ecascvoavevocesob (2 sosececesereevocacese 356
Enlargement of portion of holotype (see Plate 71, fig. 6) showing trifid imprints;
XZ:

427

Figure

1,2. Asaphoidichnus trifidum Miller, 1880
ile

3,4.

5, 6.

2.

Trachomatichnus numerosum Miller, 1880
3.

Petalichnus multipartitum Miller, 1880
5.

Allocotichnus dyeri (Miller), 1880

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 73

Repichnia of Jsotelus; conyex hyporelief showing a combination of unifid,
bifid, and trifid imprints. Locality unknown. MCZ 1055; latex mold UCM
37694. X 1.

Repichnia of /soteclus; convex hyporelief. Hypotype (Caster, 1938, pl. 10, fig.
2). Movement was from bottom to top and the body was oriented to the
right of the axis of movement. Fulton beds, Union Levee, Cincinnati, O.
UCM 22034, X 0.8.

Repichnia of Cryptolithus; convex hyporelief showing closely spaced casts
of the impressions of the multifid legs of Cryptolithus. Movement was from
bottom to top. Syntype. Eden beds of State Avenue, Cincinnati, O. MCZ 547;
latex mold UCM 37695, X 1.

Repichnia of Cryftolithus, concave epirelief. Movement was from top to bot-
tom. Syntype. Eden beds of State Avenue, Cincinnati, O. MCZ 544; latex
mold UCM 37671, X 1.4.

Repichnia; convex hyporelief showing conversion from straight-ahead to
oblique. Movement was from bottom to top. Specimen placed by Miller
(1880) in Trachomatichnus permultum. Hypotype. Eden beds of State Avenue,
Cincinnati, O. MCZ 548; latex mold UCM 37696, X 1.

Repichnia; concave cleavage relief showing withdrawal markings at the
proximal termination of each imprint. Movement was from bottom to top.
Locality unknown. UCM 37579, X 1.

Repichnia of ?/sotelus; convex hyporelief. Movement was from right to left
and the body was oriented to the right of the axis of movement. Paratype. Eden
beds of Mud Lick Creek near Walton, Ky. UCM 37630, X 1.

362, 365

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PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 74

Figure

1.

4, 5.

6, 7.

TRACE FOSSILS CINCINNATI AREA: Oscoop

EXPLANATION OF PLATE 74

Page
Trachomatichnus numerosum Miller, 1880 ........-------.--2.----c00-----ces00000+--00000-0- eee es ee nae | 366
Repichnia of Cryftolithus; convex hyporelief. The clarity of preservation indi-
cates that the trail is preserved as a boundary relief. Note the tracks of Alloco-
tichnus dyeri which occur near the upper margin of the specimen. Direction of
movement from left to right. Lectoholotype. Eden beds of State Avenue, Cincin-
nati, O. MCZ 539; latex mold UCM 37583, X 1.
CL ALNGH TEUNMILILLLD AT EDLIT ANAM | ests Wel 8 Ole eerenes cc eee ece cone caccence levcan cease ceosceesteeve 362, 365
Repichnia of a medium-sized trilobite; convex hyporelief. The trail is poorly
defined and is most likely preserved as a cleavage relief. Presumably a more com-
plete set of imprints lies just below the surface. This specimen was designated
by Miller as the holotype of Trachomatichnus cincinnatiensis. Eden beds of State
Avenue, Cincinnati, O. MCZ 542; latex mold UCM 37571, X 0.9.
Dactylophycus quadripartitum Miller and Dyer, 1878 ............0..c.cc2ceccceseseeeeeeceeeeeeee- 345, 346
Fodinichnia; convex hyporelief showing a flabellate specimen where the median
groove of the branches resembles a flat-bottomed trough. Locality unknown;
Eile
Petalichnus multipartitum Miller, 1880 ...................... eee cee ent eer eee eee 362, 365
4. Specimen was assigned by Caster (1938, pi. 10, fig. 4) to Merostomichnites
sp. Note the scallop-like nature of the imprints. Movement was from lower
left to upper right and the body was oriented to the left of the axis of
movement. Fulton beds, Union Levee, Cincinnati, O. UCM 22033, X1.25.
5. Convex hyporelief showing an excellent example of dimorphic imprints. The
imprints of the left series are superimposed while those of the right series
are clearly defined. Southgate beds below the east end of the Beechmont
Levee, Hamilton Co., O. MU T86; latex mold UCM 37707, X 1.
UVR EGET EROS: sCeGCT (UN OUTS) USSD ese eee oe eee eae ne 359

6. Repichnia of Jsotelus; convex hyporelief showing bifid imprints. Paratype.
Eden beds of Mud Lick Creek near Walton, Ky. UCM 37626, X 1.

7. Convex hyporelief showing deeply impressed raking imprints. Note the series
of imprints to the upper left of each series which probably represent casts
of gouge marks made by the pleurae of Jsofe/us. Movement was from left
to right and the body was oriented far to the left of the axis of movement.
Paratype. ?Eden beds of State Avenue, Cincinnati, O. MCZ 1053; UCM
37708, X 0.8.

429

3)

0

PaLAEonToGRAPHICA AmeERICANA (VI, 41)

EXPLANATION OF PLATE 75

Figure Page

1. Repichniaof \Crypioltthus 2. <5 k nc ee 368
Convex hyporelief of two Cruziana-like trails. Note the imprints of the genal
spines flanking the trail on either side. Locality unknown. UCM 37603, X 0.9.

2. Asaphoidichnus tripidum Maller. WS yee see eee ee 365
Repichnia of Jsotelus; convex hyporelief showing sigmoidal imprints in both
series. Movement was from bottom to top and the body was oriented to the
right of the axis of movement. Fulton beds, Union Levee, Cincinnati, O. UCM
37 560s (019:

8, Teratichnus confertum Miller S80) 222es ee ee ae 369
Repichnia of unknown arthropod; concave epirelief. Latex mold of holotype.
B, C, D mark the locations of Jong striae produced by a caudal cercum or ter-
minal spine. The letter A marks the locality of an ellipsoid grouping of imprints.
General direction of movement was from bottom to top. Holotype. Eden beds of
State Avenue, Cincinnati, O. MCZ 540; latex mold UCM 37570, X 1.1.

4. ‘Petalichnus qaultipartiium: Niallerse 1880s ee ee 362, 365

Repichnia of medium-sized trilobite; convex hyporelief showing a complex
superposition of imprints. A few cuneiform support imprints can be seen in
the center of the trail. Holotype. Eden beds of State Avenue, Cincinnati, O.
MCZ 546; latex mold UCM 37697, X 1.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 75

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 76

Figure
1.

2-4.

6, 7.

TRACE FOSSILS CINCINNATI AREA: Oscoop

EXPLANATION OF PLATE 76

Page
TEPC NEN EIS. GLICO V8 BAU TER 7) eo no ee 374
Repichnia; convex hyporelief showing the coarse rugosities which characterize
the species. Holotype. Trenton (Ordovician) beds of Middleville, N.Y. NYSM
240/1, X 0.75.
ELLE PECTED OGGLIE OS 2L77E ES cA) emily meme oo oa oa syn cee bees ecosc ccc occu sescscseserssaZasecenents 374

2. A small cylindrical segment of the burrow filling. Syntype. Queenston (Ordo-
vician) beds of Irondequoit Bay, New York. USNM 41106, X 0.7.

3. Burrow filling shaped like a depressed, helicoid cylinder. Syntype. Locality
same as figure 2 above. USNM 41102, X 0.7.

4. Faintly striate, arcuate burrow filling. Cross-section depressed cylindrical.
Syntype. Locality same as figure 2 above. USNM 41107, X 0.7.

TRUER DE NPIETT ARID SED Sh a oN tae RE i 374, 377

Repichnia; convex hyporelief of cylindrical burrows with well-developed annuli.

Corryville beds of O’Bannion Creek, Clermont Co., O. UCM 37646, X 1.

PPEVEUZD I MAIER” GUTERETAT i VERN URS y- eps Rr ee 374
6. Repichnia; convex hyporelief showing hemicylindrical burrow filling marked
by a narrow linear crest. Hypotype. Clinton (Silurian) beds, Oneida Co.,
N.Y. USNM 41108; latex mold UCM 37585, X 0.8.
7. Repichnia; convex hyporelief showing three intersecting burrows. Hypotype.
Locality same as fig. 6 above. USNM 41111, X 1.

PCIZED PIV IALSMT IEMA ex ied all ie USA 76 cess sec ote tok Pac act ag. ecSeadents artes ane s-benanasastedecthveccushetiecacensotcseceeoach 374
Repichnia; convex hyporelief showing several intersecting burrow fillings, some

of which are striate. Syntype. Beekmantown (Ordovician) beds of Amsterdam,

New York. NYSM 241/1; latex mold UCM 37698, X 0.5.

MECAGOMD MV GH Spat, P C= 0S acces cco oac-a-vezcesegeccec=tceaeeenuson Pee ee Ne ee eer cS ee 374, 378
Repichnia; convex hyporelief showing hemicylindrical burrow filling marked
by longitudinal striae and annuli. Locality unknown. UCM 37584, X 1.

431

Figure

1.

a |

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 77

Page
Palacophycus tubulare Wall; 1847) .25...2...0o ee ee 374
Repichnia. View of a syntype which is heavily encrusted with limonite. Beek-
mantown (Ordovician) beds of Fort Plain, N.Y. NYSM 241/3; latex mold UCM
37699, X 0.6.
Planolites corrugatus Walcott, 1899 _................ supa gas ieegt Ronee ce een ee 378
Repichnia. What Walcott described as corrugations are actually cleavage planes
deflected by the burrow. Holotype. Greyson Shale (Precambrian) near Neihart,
Mont. USNM 33796, X 1.
Planolites annularius, Walcott, 1880-2055. ee 378
Repichnia. A weathered specimen bearing annuli. Holotype. Lower Cambrian
beds near Greenwich, N.Y. USNM 18360, X 1.
Palaecophycus tibulare Wall), US47 2 ee oae e oe lee eeeeeeeee 374
Repichnia; convex hyporeliet showing several cylindrical burrow fillings. Note
that one of the forms is bilobate. Syntype. Locality same as Fig. 1 above. NYSM
241/4; latex mold 37700, X 0.4.
Palacophey cs, ty pe= Cocco sesce ck nse eke se 374, 378
Repichnia; convex hyporelief showing burrow fillings preserved in a coarse-
grained calcarenite. McMicken beds of Brierly Creek, Hamilton Co., O. UCM
37647, X 0.5.
Palacophycus, stype-Bs 222. .cesecs0 cae tecscsectcue a ees 374, 378

Repichnia; convex hyporelief showing annuli and longitudinal striae. Note
bulbous expansion. Sunset beds, Four Mile Creek, 10 miles south of Oxford,
O. UCM 37682, X 0.9.

Palacophycus tubslare, Shall, R47) oe ences ore eset eee 374
Repichnia; convex hyporelief with several flattened hemicylindrical burrow fill-

ings. Syntype. Locality same as Fig. 1 above. NYSM 241/2; latex mold UCM

37701, X 0.4.

Gorophioides biclavatay (Miller) pal S SO)is oe ere eee ee ee 314, 323
Domichnia; U-plane section showing that the arms have been extended below the

level of the Spreite. The smaller, lighter colored burrows are Chondrites, type-C.
MecMicken beds of West Fork Creek, Mt. Airy Forest, Cincinnati, O. UCM

37649, X 1.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 77

Plate 78

PALAEONTOGRAPHICA AMERICANA, VOL. VI

Figure
1-3.

4-6.

8, 9.

Ormathichnus monilliformis Miller, 1880
1.

2.

35

Rhabdoglyphus sp. Vasseovich, 1951
4.

5

6.

Hormosiroidea florentina Schaffer, 1928

TRACE FOSSILS CINCINNATI AREA: OsGoop

EXPLANATION OF PLATE 78

Enlarged view of fig. 2 showing a series of small disconnected Rusophycus-
like bodies, X 5.

Repichnia of a small arthropod; convex hyporelief. Syntype. Eden beds of
State Avenue, Cincinnati, O. MCZ 549; latex mold UCM 37689, X 3.
Inorganic; convex hyporelief. The monilliform casts are interpreted as
markings of a crinoid stem. Syntype. Locality same as fig. 2 above. MCZ
541; latex mold 37690, X 2.5.

Repichnia; concave hyporelief. Locality unknown. MU 196; latex mold UCM
B02 eke Lads

Concave epirelief showing the variation in the morphology of the hemi-
spherical depressions. UCM 37683, > rile

Concave epirelief showing raised rims of the connecting troughs. Sunset beds
of Four Mile Creek 10 miles south of Oxford, O. UCM 37653, xe.

Repichnia; ?convex hyporelief showing closely spaced hemicylinders. Note small
vermiform trails. Holotype. Upper Cretaceous of Italy. Plastotype UCM 37652,
e:Oulk

Rhabdoglyphus grossheimi Vasseovich, 1951 -....---------:---s cesses -ensenerrenceensscnnscens ics
8.

Repichnia; convex hyporelief. Vertical sections would probably reveal the
connecting bar just below the surface. Hypotype. Tertiary Flysch of Czecho-
slovakia, X 1.1 (from Bouéek and Elias, 1962, Pl. 8, fig. 2).

?Convex hyporelief showing ellipsoidal swellings. Note that two of the swell-
ings occur together. Locality same as figure 8 above. X 1.1 (from Bouéek
and Elias, 1962, pl. 8, fig. 3).

Page

ww

ws

434

Figure
ie.

PALAEONTOGRAPHICA AMERICANA (YI, 41
>

EXPLANATION OF PLATE 79

Page
“Palaeophycus jlexosum's (U- ae. fares, 1879 yee 393
1. Inorganic; epirelief showing the numerous anastomosing ridges. Humphrey’s
Branch of I'welve Mile Creek, Campbell Co., Ky. UCM 37645, X 1.
2. Inorganic: specimen reveals several layers marked by the flattened ridges.
Locality unknown. UCM 37650, X 0.7.
. Mastigograpius gractllimium i (Mesquereiixc) ws 78 meee eee ee ee 387, 388
The form was interpreted by Ruedemann (1908) as a graptolite. This speci-
men bears no trace of thecae. Locality unknown. UCM 37651, X 1.5.
BIMolluske: trail gt cccyscce fete ieee an teceean cee cndenaaact ce Rn ee a 382
Trails change from a median furrow flanked by two ridges into a raised ridge
marked by a median rift. McMicken beds of West Fork Creek, Mt. Airy For-
est, Cincinnati, O.
GHomdpite sy t¥ pe Es: cece cco-sese Saace ssa oes ese ee aoe ee Fe 328, 335
Vertical cross section showing a concentration of burrows in lower half of the
host rock. Note that one branch angles upwards. McMicken beds of West Fork
Creek, Mt. Airy Forest, Cincinnati, O. (type locality of Corophioides cincin-
natiensis). UCM 37669, X 0.6.
Chondrites\:bollensts: Zieten;, 1839. <0. ke ee 329

Fodinichnia; bedding plane surface showing a specimen which exhibits phobo-
taxis. Such specimens marked the “death knell” of the “fucoids.” Jurassic Posi-
dion beds of Wiirttemberg, West Germany. UCM 37538, X 0.8.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 79

Plate 80

PALAEONTOGRAPHICA AMERICANA, VOL. VI

TRACE FOSSILS CINCINNATI AREA: OscGoop

EXPLANATION OF PLATE 80

Figure Page
1. “Chloephycus plumosum” Miiler and Dyer, 1878 ......c----v-sscosccsscecesscesssensscncesenecreecstennenensnenns 389
Inorganic; convex hyporelief of holotype showing numerous parallel specimens.
Current flow was from top to bottom. Eden beds, Cincinnati, O. HBM 3154;
latex mold UCM 37617, X 1.25.
2. “Blastophycus diadematum” Miller and Dyer, 1878 .....--.------:-:--:-cecsseceseeeossseeesett 390
Inorganic; convex hyporelief showing cast of enrolled trilobite with casts of
scour markings. Current flow was from top to bottom. Compare with forms on
Plate 81, figures 1-3. Paratype. Eden beds, Cincinnati, O. HMB 3181; latex
mold UCM 37618, X 1.5.
3,4. “Dystactophycus mamillanum” Miller and Dyer, 1878 ....-.--------:.----etese 391

3. Inorganic; convex hyporelief representing the cast of markings made by a
sweeping crinoid stem. Richmond beds, Hamilton, O. USNM 40043; latex
mold UCM 37620, X 1.

4. Inorganic; convex hyporelief showing the apparent imbrication of the rings.
The left side of the specimen has been altered by preparation. Holotype.
Richmond beds near Morrow, O. HBM 3177; latex mold UCM 37691, > Crile

Mipnilenalimmeconcentr ina, Pally (USSQ) 2.cszeccccncecsxeceenmmncseseccernerniseee man reece coat ores 391
Bryozoa; latex mold bearing a vague resemblance to “Dystactophycus.” J. F.
James (1885) believed that the two were synonomous. Syntype. Niagaran
(Silurian) beds near Rochester, N.Y. AMNH 1487/1; latex mold UCM 37685,

NGIZE5:

“Chloephycus plumosum” Miller and Dyer, 1878 ....--------c------cscsseesrecorecssceeensrsseceosenenseneens 389
Inorganic; convex hyporelief showing a single specimen. Note flute casts to

the right of “Chloephycus.” Eden beds of State Avenue, Cincinnati, O. Speci-

men on reverse of Trachomatichnus numerosum, MCZ 544;; latex mold UCM

37692, X 1.25.

“Scolithus dispar” J. F. James, 1892 -...---------.--------c--cr
Inorganic; convex hyporelief. U. P. James (1881) assigned the long linear mark-
ings to “Scolithus linearis.” Also present on the slab are casts of rolling crinoid
stems, and groove casts made by twisting brachiopod shell fragments. Current
flow was from upper left to lower right and top to bottom. Paratype. Eden beds,
“Crawfish Creek,” O. UCM 37619, X 0.6.

435

436

Figure

il

~I

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 81

Page
“Blastophycus diadematum” Miller and Dyer, 1878 .....-...--0-:0scssssscessssseeceesecesseseesaesseeanee 390
Inorganic; scour marks around an enrolled trilobite. Current flow from upper
right to lower left. Compare with figs. 5, 10 below. Experimentally produced;
XO
?Paleodictyon sp. Savi and Meneghini, 1850 ...384, 387

?Pascichnia; convex hyporelief showing a_ poorly e specimen. } ote
that the network is oriented parallel] to the current direction. Economy beds of
Duck Creek, Campbell Co., Ky. UCM 37638, X 1.2.

“Blastophycus diadematum” Miller and Dyer, 1878
Inorganic; same as fig. 1 above. Rill marks are attributable to subaerial erosion.
Current flow from lower right to upper left, X 0.7.

(YO ELA TNA S (18) Se Oa eee cena ee RE Be ee 328, 339
Fodinichnia; concave epirelief showing two systems, one oriented Parallel, the

other oblique to the bedding plane. A latex mold of this specimen is illustrated

on Plate 64, figure 5. Whitewater beds, Short Creek, three miles southeast of
Richmond, Ind. MU T90; latex mold UCM 37678, X 0.6.

“Rlastophycus diadematum. Miller and Dyers Usc8) =. eee ee 390
Inorganic; plaster cast of experimentally produced scour marks around an en-
enrolled trilobite. Current flow from top to bettom. UCM 37664, X 0.5.

“Dystactophycus mamillanum” Miller and Dyer, 1878 .....-0..-:0.----ccceeeeeeeeeeceee sence ceeeeeeeeeeeee 391
Inorganic; convex hyporelief clearing showing the casts of markings made by a
sweeping crinoid stem. Arnheim beds, Hamilton, O. USNM 40044; latex mold

UCM 37687, X 0.8.

Allacotichnus® dyerwa (Miller, S80 see ese cerca ee cer eee eee 359
Repichnia; convex hyporeijief. An enlargement of a portion of the trail shown

on Plate 74, figure 1. The small parallel markings were probably made by
spinose projections on the distal ends of the segments of the walking legs.
Paratype. Eden beds of State Avenue, Cincinnati, O. MCZ 539; latex mold UCM

37583, X 4.

Pascifodina flonvert, nt en., A Spe. csscccepecna eee eee 340
Fodinichnia; epirelief. A small individual showing the concave imprint and the
convex, lower portion of the master shaft. Holotype. Corryville beds of Second

Creek where crossed by Cozzadale Road, Clermont Co., O. UCM 37632, X 1.

Rusophycus pudicum Hall, 1852 - 2
Cubichnia of Flexicalymene; convex < hyporelie showing a trail leading away
from the resting trace. Note the well-preserved casts of the pleurae on the
Rusophycus body. Corryville beds, Stonelick Creek, Clermont Co., O. MU 194;
latex mold 37663, X 0.7.

“Blastophycus diadematum- Miller sand Dyer, 18178, sesseectees ttetices een teen eee 390
Inorganic; convex hyporelief showing the “bud” which clearly reveals its trilo-

bite origin. Syntype. Eden beds, Cincinnati, O. HBM 3181; latex mold UCM

37703, X 1.7.

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 81

PALAEONTOGRAPHICA AMERICANA, VOL. VI Plate 82

Figure

1-6. Palaeoscia floweri Caster, 1942
aS

TRACE FossiLs CINCINNATI AREA: Oscoop

EXPLANATION OF PLATE 82

Fodinichnia or inorganic; concave epirelief showing a well-developed cen-
tral pore, “radial canals” and concentric markings. Holotype. Corryville
beds of Stonelick Creek, Clermont County, O. UCM 24079, X 1.2.

Concave epirelief of a shallow regular specimen. Topotype. Locality same as
fig. 1 above. UCM 24389, X 0.7.

Concave epirelief. An enlargement of a portion of the holotype showing the

central pore and “radial canals”, X 3.

Convex hyporelief showing several specimens. The concentric arcs are devel-
oped only in the downcurrent quadrant. Economy beds of Duck Creek, Camp-
bell County, Ky. UCM 37578, X 1.3.

Concave epirelicf showing two overlapping specimens. Note that the central
pore has a raised rim and the inner wall of the arcs appears to have been
planed off. Topotype. Locality same as fig. 1 above. UCM 37575, X 1.

Concave epirelief showing a portion of a larger specimen. Note the compara-
tively large diameter of the central pore. Topotype. Locality same as fig. 1
above. UCM 37576, X 1.3.

437

438

Figure

6-8.

PALAEONTOGRAPHICA AMERICANA (VI, 41)

EXPLANATION OF PLATE 83

Page
. Palacophycus, type-C ..........-.-.. a esas ssca ae Sesow ieee nets oo RaUe Roo a 374, 378
Repichnia; convex hyporelief showing several interpenetrating burrows. The
coarse grain size of the host rock obscures the finer details of the burrows.
Locality unknown. UCM 37582, X 2.
Chondrites, type-A ~.........-.--... es =328,935
Fodinichnia; vertical section. Note downward deflection of some of the secon-
dary branches. McMicken beds of West Fork Creek, Mt. Airy Forest, Cincin-
nati, O. UCM 37680, X 1.4.
PResting, traceof- amautilord cephtalopod ey. sce cane cee eee eee eee eee 349
Concave epirelief. This specimen differs from the “turkey track” depressions
which Flower (1955) believed to be nautiloid resting traces. The two critical
features are the raised borders and the smooth floor of the depression. Mt. Hope
beds on U.S. Route 56, 0.4 miles west of Aurora, Ind. UCM 37643, X 1.
Paldeop hy cus ity pe-Gr cence eee eee eee PER errs eee eee eps: eect eee ter aon 374, 378
Repichnia; note that trails are preserved as both concave and convex epireliefs.
Eden beds of Brierly Creek, O. UCM 37644, X 0.5.
. Enlargement of a portion of the trail shown on Plate 19, figure 1 illustrating
two Cruziana-like ridges. The delicate striae are attributed to the brushlike ter-
minal segment of the Cryftolithus walking leg; X 1.5.
Aristophycus ramosum Miller and: Diyer U8 78ers eee ee 398

6. Incertae sedis; latex mold of fig. 8 below showing the anastomosing pattern of
the secondary and tertiary branches, X 0.8.

7. Convex epirelief. Note that the secondary branches extend down into the
host rock much like Ivy tendrils. Compare with Text-figure 25. Holotype.
Maysville beds, Cincinnati, O. HBM 3173; latex mold UCM 37580, X 0.8.

8. Convex epirelief showing a single main branch. This specimen was assigned
to A. ramosum “var.” germanum by Miller and Dyer (1878). Hypotype.
Maysville beds, Cincinnati, O. HBM 3172; latex mold UCM 37668, X 1.2.

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INDEX

Note: Light face figures refer to the page numbers. Bold face figures refer to the

plate numbers.

A

Abel, O. ... ...282, 289,
298, 304, 316, 317,
319, 343, 375, 385,
405

Ager. Di ia: .... 291, 405
Allocotichnus 72-74, 81 281,
300, 355, 356, 358-

362, 367, 402, 427-

429, 436
Amoricain Sand-

stone (Sweden) .. 294
Andree; K. .....:..2...:.. 316, 405
Angulaten Sand-

stone (Jurassic:

Wurttemberg) ... 396, 400
annularius,

Planolites ....... 77 377, 432
antiquata,

FATURTATI AD aveeseees 325
Aphrodite ................ 303
Archaeonassa .......... 383
Arenicola ................318, 326,

343, 344
Arenicoloides .......... 315-317
Arenicolites ............317, 325
Aristophycus ......83 300, 397,
398, 438
Arnheim beds ....... 285, 286,
391, 436
Arthraria ...... 300, 316,
323-325, 371
Arthrophycus ....... 342, 343,
346, 375

Asaphoidichnus

65, 71-73 300, 355,
356-359, 361, 402,

426-428, 430

asperum,
Rusophycus .66,71 301,
305, 371, 372, 402,
421, 426

Tylichnus 66,71 301, 305,
371, 372, 402, 421,

426

Asteriacites 57,58,62 290,
298, 300, 312, 313,

402, 412, 413, 417

B
Baggy beds (Great

Britain:

Devonian) ............ 283, 316
Bagnold, R.. ............ 294, 405
Balanoglossus ....... 319
balticus,

Corophioides ..... 319
Bantam, Clermont

County, Ohio ......306, 414,

426

Barnes, R. 319, 322,
405

Barrois, C. 303
Bassler, R. 310, 330,
347, 375, 405

Bather, F. 316, 317,
405
Beecher, C. 354, 355,
367
Beechmont, Levee,
Cincinnati, Ohio 429
Beekmantown beds
(Ordovician) .......373, 374,
431, 432
Bellevue beds 285, 386,
425
Beloraphe 385, 386

biclavata, Coro-
phioides 60, 61, 65,
66, 70, 77 298, 300,
320-325, 415, 416,
420, 421, 425, 426,

432
Bifungites 316
Billings, E. 286, 323,
325, 326, 342, 344,
352, 375, 405
biloba, Fucoides 301, 302
bilobatum,

Rusophycus 301, 302,
305, 371
Bilobites ........ 303
Blanchester beds 285
Blanckenhorn, M. .316, 318,
320, 405

“Blastophycus”’

67, 80, 81 300, 347,
350, 389, 390, 399,
422, 423, 435, 436

Blue Bank,
Kentucky 337
bollensis,
Chondrites 79 329-331,
434
Bornemann, J. ........ 303, 304
Boucek, B., &
Elias, M. .. 369-371,
405

Boudinot Avenue,
Cincinnati ‘ 390

Brierley Creek,
Hamilton County,

Ohioy< .. 432, 438
Brinkmann, R. ... 400, 405
Brongniart, A. 286, 329,

330
Bucher, W. 284, 403,

405
Bulla : ‘ 370
BUA ete ate 304
Bunter Sandstone

(Germany:

Triassic) 316
Buthotrephis 67 299, 300,

309, 329-332, 387,
422
Cc
Callianassa 283, 350
callicephala,

Flexicalymene 307

139

carleyi, Rusophycus
57, 58, 59,71 296, 301,
304-307, 402, 412-
414, 426
281-283,
291, 295, 298, 300,
304, 350-352, 355,
358, 362, 365, 395,
396, 405

Caster, K.

Caster, K., Dalvé,
E., and Pope, J. . 285, 286,

366, 404, 405
Caster, K., and
Waering, E. 356
caupo, Urechis . 319, 324
Ceraurus .. 355
Chaetopterus 319
Chemung beds (New
York: Devonian)... 403
Chickies Quartzite
(Pennsylvania:
Cambrian) 325, 326
Chinle Formation
(Arizona:
Triassic) 352
“Chloephycus” — 80 300, 334,
388, 389, 435

Chondrites 63-65, 67,
69, 79, 82, 83 282, 289,
292, 297, 300, 328-
339, 348, 401-404,
418-421, 424, 432,
434, 436, 438
ciliata, Polydora .... 318
Cincinnatian Series 281, 284,
285, 403, 404

cincinnatiensis,

Corophioides 60-463 300,
320-324, 334, 336,
402, 415-418, 426
Trachomatichnus 74 296,
301, 355, 365, 366,

429
circinatum,

Phycodes 65 341-344,

420
Clarkesville beds 285
clavatum,

Rusophycus_ ....67 305, 422
Clematischnia ‘ 333
Climactichnites : 287
Clinton (New York:

Silurian) 299, 301,

302, 305, 306, 325,
330, 342, 345, 352,
391, 422, 424
Clinton beds,
Oneida County,

New York ..........332, 374,

413, 414, 418, 422,

431

Cloud, P. 327, 405
coloradoense,

Polyupsilon 296
concentrica,

Lichenalia 80 391, 435

conchilega, Lanice 333

confertum,
Teratichnus ...75 301, 355,
364, 368, 369, 430
congregatus,

Palaeophycus ; 375
Conostichus 300, 399
cookana, Walcottia 301, 378-

380
Cooper, G. 403, 405

Corophioides 60-62,
66, 69-71, 77 283, 292,
300, 314-325, 335,
348, 398, 402-404,
415-418, 420, 421,
424-426, 432, 434

Corophium 3S 317
corrugatus,
Planolites 77 377, 432

Corryville beds 285, 320,
340, 348, 358, 377,
390, 395-397, 403,
404, 412, 413, 420,
421, 425, 426, 431,

436, 437
Covington,
Kentucky 394, 418
Crawfish Run,
Cincinnati 392, 435
Crossochorda 303
Cruziana 287, 288,

294, 296, 301, 303,
306, 367, 368, 372,
373, 399, 401, 403,
404, 421, 422, 430,

438

cryptolithi, Ruso-
phycus ......58,59 281, 301,
305, 307, 348, 393,
402, 413-415
Cryptolithus Sel, Bil
354-357, 360, 365-
368, 413, 414, 420,
428-430, 438
Cyathophycus .... 300, 347
Cynthiana beds ...... 285, 333
Cyclophycus 300, 399
Cyclozoon . : 396

D
Dactylophycus

57, 63, 69, 74 300, 345,
346, 402, 412, 418,

424, 429
Daedalus cere 401
DAL ea eee . 809, 405
darwini, Pasceolus.. 386, 424
Dawson, C. ..............286, 303,
304, 326, 352, 405
Dawsonia a Ate ee 309

Dearborn County,
Indiana ce SAL,
Delgado, J. ....... 286-288,
294, 303, 368, 401,
405

delicatulus,

Skolithos 62 301, 327,
328, 417
Dendrograptus 333, 387,
388
Desio, A. . 295, 325,
393, 405

INDEX

devonicus, Rhizo-
corallium ... 319
“diadematum,
Blastophycus”
67, 80, 81 300, 389,
390, 399, 422, 423,

435, 436
didymus,
Rusophycus 301, 302,
373
Dimorphichnus 353, 400
Diplichnites .. 352, 355
Diplocraterion 316-318,
325, 401
dispar,
Skolithos 80 301, 389,
392, 393, 435
Duck Creek,
Campbell County,
Kentucky 307, 385,
386, 396, 397, 436,
437

dyeri, Alloco-
tichnus 72-74, 81 296, 397,
300, 355, 359-362,
402, 427-429, 436
“Dystactophycus”
70, 80, 81 300, 390-
392, 397, 425, 435,

436

Dzutinski, S. &
Sanders, J. 292, 294,
370, 392, 405

E
Economy beds 285, 348,

380, 383, 385, 392,

393, 396, 398, 404,

413, 418, 436, 437
Ectenocrinus ......... 392
Eden Group ... 284, 285,
333, 345, 346, 349,
360, 367, 370, 371,
377, 386-389, 394,
404, 412, 414, 423,
424, 426-430, 433,

435, 436, 438
Ediacara beds

(Australia:

Precambrian) 396
Elkhorn beds é 285
Emerita ........... 305
Eocene beds,

Trieste, Italy 384, 386,

421, 424
erraticus, Coro-

phioides ............. 319
Estonia (Lower

Cambrian) .......... 315, 320
Huryale ee 312

F
Fairmount beds .... 285
Fairview beds ....284, 285,
313, 379, 394, 397
JOP WU UR ee 295, 405

Fascifodina 66, 81 281, 300,
340, 341, 402, 404.
421, 436

110

Fenton, C., &
Fenton, M. ... 298, 304,
324, 326, 327, 371,
383, 406
fezzanensis,
Bifungites : 325
Field Museum,
Chicago 281, 282,
347, 378, 379, 387,
389, 393, 399
“filciformis, Butho-
trephis” 80 300, 334,
389
filiformis,
Heteromastus ...... 298
Fisher, D. ... 306, 406

flabellum, Phycodes
57, 65, 66, 68-70 301, 343-
345, 402, 412, 420,
421, 423-425
Flat Fork of
Caesar’s Creek,
Warren County,

Ohions-- } 416
Flemingsburg,
Kentucky .. 418, 419
Flexicalymene ........306, 335,
360, 366, 390, 412,
413, 422, 436
flexuosum,
Palaeophycus 79 300, 376,
393, 394, 434
florentina, Hormo-
siroidea) 2... 78 370, 433
Flower, R................295, 800;
340, 341, 348, 349,
406
floweri,
Fascifodina 66,81 300, 340,
341, 421, 436
Palaeoscia 71, 82 301, 395-
397, 426, 437
Flysch (Alps:
Tertiary) eee ee SPtiy
331, 353, 384, 385,
399, 401, 404
Foerste, A. ..............284, 286,
309, 310, 406
HOLd i Jats see OF aa
406
Fort Ancient beds.. 285

Four Mile Creek,
Oxford, Ohio ...... 404, 432,
433
ORS Wirtsccneeeee 284, 403,
406
Huchs;, WL.) tse Oa 2oo!
299, 330, 331, 333,
339, 385, 393, 398,
406
BUCOIdeS see eee 286, 330
Fuccusopsis 64,70,71 300,
347, 380-381, 419,

425, 426
Fulton beds ............285, 309,
310, 324, 365, 416,
423, 428-430
furcatus,

Chondrites .......... 330, 336

G

GaUdny, Avec. s- 288, 406
Gilbert, G. .............. 294, 406
Glaessner, M. ........303, 304,
396, 406
Glossifungites .. 401
Goldring, R. . 283, 314,
316-318, 406

gracilis,
Chondrites ....69 300, 330,

332, 333, 387, 424
gracilis ‘‘var.”
crassa, Chond-
rites ................63 300, 329,
330, 332, 333, 418
gracillimum, Masti-

gograptus .......79 300, 333,
387, 388, 434
granulatus,
Chondrites . 330-332
grenvillensis,
Rusophycus ........ 302
Greyson Shale
(Montana:
Precambrian) ...... 377, 432
grossheimi, Rhab-
doglyphus 78 369, 433
H
Haldemann, S. ........ 325, 326
eal ed nites tees: _ 281, 286,

295, 299, 302, 305,
325, 326, 329, 330,
332, 350, 373-375,
387, 388, 391, 392,

399, 406

halli, Bifungites .... 325
Hall’s Creek,

Morrow, Ohio ....381, 396,

397, 425

Hamilton, Ohio 391, 435,

436

Hantzschel, W. ...... 281, 282,
284, 289, 291, 294-
296, 300, 303, 313,
316, 317, 326, 330,
342, 347, 349, 366,
372, 380, 383, 385,
386, 390, 393, 398,

406

Harvard University
collections , 378,
379, 389
Ger MO? cee 286
Heliophycus ............ 300, 313

helmerseni,

Corophioides_ ..... 319, 320,
322
Helminthoida ... 283, 353,
386, 400, 401
Hjulstrom, F. . 406
Hofmann, H. ....284, 403,
406

Holy Cross Moun-
tains (Poland:
Cambrian) .......... 307
Holzmaden Forma-
tion (Germany:
WULASSIC) 2. eer 329

INDEX

Hormosira ...... : 370
Hormosiroidea .78 369, 370,
433
Howell, B. 296, 326,
375, 376, 406

“Hudson River
Group” 347, 375

Humphrey’s Branch
of Twelve Mile
Creek, Kentucky. 307, 348,
393, 404, 413, 415,

434
Hundt, R. . 297, 342,
406
Hunsriick Shale
(Rhineland:
Devonian) 283, 289,
328, 329, 352, 389,
399
Huff, W. 335
I
Ichnophycus 345
Illyanassa ......... 371
Imbrie, J. &

Newell, N. . 291, 406
Inocaulis 344
intortum,

Saccophycus 301, 389,

399
Iraq (Ordovician) 401
irregularis,

Palaeophycus . 374
Isopodichnus .... 303, 401
Isotelus ....283, 296,

"307, 351, 354, 356-
358, 361, 362, 412-
414, 426, 428-430

J

James, J. F. 281, 288,
299, 300, 305, 306,

309, 312, 313, 324,

326, 327, 332-334,

344-347, 350, 355,

372, 373, 375, 376,

379, 381, 387, 389-

395, 398, 399, 406,

423

James, UP. ............281, 299,
308, 323, 326, 327,
334, 378, 389, 392,
393, 398, 399, 406

jenningsi,
Rusophycus 302, 304
K
Kilwinning,

Scotland 314, 415
Kinneyia ......... 393
Kolesch, K. 319, 406
Kope Formation 284
Koriba, K., &

Miki, S. ... 384, 385
Krejci-Graf, K. 289, 297

304, 406
Krosno beds
(Poland:
Oligocene) 392

111

Kuenen, P. 292, 406
Kuenen, P., &

Carozzi, A. 400, 406
L

Laevicyclus 396

lanosum, Tricho-

phycus 68 301, 347,

349, 350, 381, 389,

423

Latonia Formation. 284, 285
laterale,

Cyclophycus 300, 399
latifolius,
Sphenothallus 399
Lebanon, Ohio 299, 334,
343, 389, 399, 420
Lebesconte, P. 287, 288,
406
Lepidocoleus 380
Leptosynapta 370
Lesquereux, L. 299, 387,
388, 394, 395, 407
Lessertisseur, J. ...282, 291,

298, 329, 396, 407
Lias and Dogger

(Wiirttemberg:

Jurassic) 291, 400
Liberty beds 285, 286,

404, 425
Lichenalia 80 391, 435
Lick Run 404
Licking River,

Covington,

Kentucky 387
Licrophycus 301, 342
Limekiln Run,

Cincinnati 394
Limulus 283, 298,

303, 304, 358

Linck, O. 309, 310,

331, 399, 400

linearis, Skolithos . 301, 326,

327, 334, 392, 435

Lingula 310

littorea, Littorina 383
Llandeilo beds

(Sweden:

Ordovician) 309
Lockeia ... 57-59, 63 283, 300,

308-312, 383, 402,
404, 412-414, 418

Longstreet Creek,
Lebanon, Ohio 394
Ludlow, Kentucky. 308, 412,
414

Lugnas, Sweden 288
lumbricalis,

Asteriacites # 312
luniformis,

Arenicoloides 315

cf. luniformis, Coro-
phioides 61, 62, 69 300,
316, 318-321, 323,
402, 416, 417, 424

M

Magdefrau, K. . 341-344,
407
magna, Arthraria 325

Maillard, G. 286, 289,
331, 407
“mamillanum,
Dystactophycus”

70, 80, 81 300, 390-
392, 425, 435, 436
marina, Arenicola..298, 318,

343, 344
Martinsburg Shale
(Appalachian:
Ordovician) 404
Martinsson, A. 291, 297,
393, 407
Mastigograptus 79 300, 333,
387, 388, 434
maximus,
Isotelus 57 306, 307,
412
Maysville Group ....284, 285,

343, 370, 389, 404,
414, 422, 426, 438

Maysville,
Kentucky 386
MeMicken beds 285, 320-
322, 334, 378, 383,
404, 415-417, 422,
424, 432, 434, 438
McMillan beds 285, 378,
379, 381, 382, 386,
394, 397, 422, 424

meeki,

Flexicalymene ....295, 304,
306, 390, 412, 413,
422
Megalograptus 356, 360
Merostomichnites | 355, 362,
365, 429

Miami University

(Oxford, Ohio)
collections 282
Milford, Ohio 390, 393
Miller, S. A. 281, 286,

287, 299, 323, 324,
333, 347, 348, 355,
356, 358-360, 365-
368, 372, 373, 389,
394, 395, 398, 407
Miller, S. A., &
Dyer, C. B. 281, 289,
299, 313, 334, 343-
347, 350, 371, 372,
378-381, 388-391,
397, 407

291, 399

Molasse (Switzer-
land: Tertiary) .
moniliformis, Orma-

thichnus 78 300, 372,
373, 433

montanus,
Planolites 375
Monticulipora 391

Moore, R., Lalicker,

C., & Fischer, A... 310, 407
Morrow, Ohio 435
Mount Auburn beds 285
Mount Hope beds _ 285, 322,

377, 413, 425, 438

Mud Lick Creek,
Boone County,
Kentucky 359, 360,
427-429

INDEX

Miller, A. 290, 298,
407

multipartitum, Peta-
lichnus 73-75 296, 297,
301, 355, 362, 363,
365, 366, 369, 402,

428-430
N
Nathorst, A. 283, 286-
289, 294, 299, 303,
331, 407
Neonereites 401
Neostrabops 358, 360,
369
Nereis SBP SE PE
379
Nereites 401, 403
Nereites beds
(Portugal:
Silurian) 400, 401
Newberry, J. 299, 394,
395, 407

Newfoundland (Up-
per Cambrian) 323
Newport Shopping
Center, Newport,
Kentucky 404, 418,
419
New York State
Museum col-
lections .. 282

Nicholson, A. 309, 375,
376
Nickles, J. 284, 286
Nowak, W. 384, 385,
407

numerosum, Tracho-
matichnus 73,74 301, 355-
357, 359, 362, 365-
368, 389, 402, 428,

429, 435
ie)
O’Bannion Creek,

Clermont County,

Ohio . MAC EAL
obsoleta, Illyanassa 324, 371
Oldhamia ae 394
Ophiomorpha 372, 401
Opik, Ay y.: 319, 352,

407
Ordovician

(Thuringia) . 317
d’Orbigny, A. : 302
Oregonia beds 285
Ormathichnus ....78 300, 304,

372, 373, 398, 433
ornatum,

Palaeophycus_.... 300, 399
Orthonybyoceras 340, 348
Orton, E. , 284, 298,

299, 407
Owen, R. ; 287, 352
Oxford, Ohio 354, 360
412
P
Palaeaster 313
Palaeophycus

76,77,79, 83 286, 299,
300, 301, 334, 345,

442

348, 373-378, 380,
381, 383, 393, 394,
399, 402, 404, 418,
431, 432, 434, 438

71, 82 301, 395,

426, 437
Paleodictyon
67, 69, 81 282, 297,
301, 384-386, 393,
401,402, 421, 436

Palaeoscia

palmata,
Buthotrephis . 67 330, 331,
333, 342, 344, 422
parallelum,

Diplocraterion 319
Paramphibius 283, 291
Pasceolus 386, 424
pedum, Phycodes 342
Pelecypodichnus 309, 310
Pennatulites 349, 372
permultum,

Petalichnus ....... 296

Trachomatichnus 73 301,
355, 362, 366, 428

Petalichnus | 73-75 301, 362,
363, 365, 366, 402,

429, 430

Petersen, C. ..... 292, 407
Philipp, H. . - 396

philippi, Cyclozoon 396
Phycodes ......57, 65,
66, 68-70 283, 297,
300, 301, 341-346,
377, 398, 401-403,
412, 420, 421, 423-

425
Phycodes beds

Thuringia:

Ordovician) . 341, 420
Planolitesy ite cos ate,

375-377, 380, 401,
432

“plumosum, Chloe-
phycus” ......... 80 300, 334,
388, 389, 435
Polydorases..ce ioe 318, 319
Polyupsilon see 296

polyupsilon, Coro-

phioides 60 315, 316,

318, 319, 415
Posidion Shale

(Wurttemberg:

Jurassic) .............. 330, 434
Potsdam Sandstone

(New York, Vt.:

Cambrian) Be PA apes

SBPA BIG)
Potter, P. Shoe 393
Potter, P., &

Pettijohn F. ...... 390, 392
primaevum,

Sphenophyllum .. 301, 394
Promopalaeaster . 313
Protichnites . 287, 352,

355
Protostigma 301, 394,
395
prunella, Porpalia 396
Psilophytum 388, 394

pudicum,
Rusophycus
57, 58, 60, 66, 81 295, 301,
305-307, 371, 402,
412, 414, 415, 421,

436
Pulaski, New York 325
Q
quadripartitum,
Dactylophycus

57, 63, 69 300, 345,
346, 402, 412, 418,

424, 429
Queenston beds
(New York:
Ordovician) 374, 376,
431
quinquefolias,
Asteriacites ...309, 312,
313
R
Raasch, G. . 358, 369
radiata,
Palaeophycus 299, 300,
345
Radwanski, A., &
Roniewicz, P. ...... 307, 407
ramosum,
Aristophycus ..83 300, 397,
398, 438

“var.” germanum,
Aristophycus . 83 300, 397,

398, 438
ramulosa,

Chondrites 300, 333,

334
Rapid Run,

Hamilton County,

OhiO7 415, 425
Rauffellia 347
Ravine Street,

Cincinnati 386, 422
Raymond, P. .... 807, 352-

355, 360, 407

Reineck, H. . 375, 376,
407

Reis, O. 333, 334

Rhabdoglyphus 22,78 301,
369-371, 402, 433
Rhizocorallium _.297, 316-
319, 322, 325

Richmond Group . 284, 285,
320, 327, 337, 343,

347, 350, 370, 390-

392, 394, 403, 404,

412, 417-420, 435

Richter, Reinhard ..341, 342,
407

Richter, Rud ...282, 283,
286, 289, 291, 295-

297, 312, 316-319,

325, 326, 328-331,

335, 336, 352, 353,

375, 376, 389, 399,

407

Rickets, E. and
Calvin, J. 319, 324,
408

INDEX

Rolfe, W. 314
rosei,

Corophioides 319
Rothpletz, A. 329-331
Ruedemann, R. 325, 344,

387, 388, 408
rugosa,

Walcottia 67, 69 283, 301,

422, 424
rugosum, Palaeo-

phycus 76 300, 374,

376-379, 431
rugulosus,

Planolites 375
Rusophycus . 57-60,

66, 71, 82 281, 282,
286-288, 296, 298,
301-308, 348, 368,
371-373, 402, 412-
415, 421, 422, 426,
436
Ss

Sabellarifex 326
Sabellarites 326
Sacco, F. 385, 408
Saccophycus 301, 389,
399

Salt Range

(Pakistan:
Cambrian) 291, 331,
342, 352, 353, 396,
399, 400
Saluda beds 285
Saporta, G. 287, 288,
292, 408
Sarle, C. 341, 342,
346, 408

Savi, F. &

Menegheni, G. 384

Schafer, W. 282, 289,
297, 298, 317-319,
331, 344, 408
Schaffer, F. 369, 370,
408

Schilf Sandstone

(Wiirttemberg:
Triassic) 291, 309,
399, 400
Schimper, P. 286, 288
Schindewolf, O. 289, 316,
408

Schindewolf, O. and

Seilacher, A. 400
Schmidtgen, O. 295
Schlotheim, E. 312
Scolecolepis 396, 397
Scolicia 304
Scoyenia 401
Scotford, D. 284, 408
sculptum,

Palaeophycus 300, 399
Second Creek,

Warren County,

Ohio 340, 341,

421, 436
Seilacher, A. 281-283,

290-298, 302-304,
306, 309, 310, 312,
313, 317, 318, 330-

143

332, 336, 341-344,
350-353, 358, 362,
368, 369, 380, 384,
385, 389, 390, 396,
398-401, 403, 404,

408
Seilacher, A. and
Meischner, D. 349, 350
Senckenberg am
Meer 282, 286,
289
serratus, Olenoides 369

Sheed Road, Hamil-
ton County, Ohio 322, 324
Short Creek, Rich-

mond, Indiana .. 339, 418,

419, 436

Shrock, R. 327, 393,

408

Sigillaria 299
sigillaroides,

Protostigma 301

siliguaria,

Lockeia 57-59, 63 300, 308-
312, 402, 412-414,

418
siluriana,
Cyathophycus 300, 347
simplex,
Heterocrinus 372
Palaeophycus 374, 375,
376
Simpson, S. 282, 289,
328-332, 334-338,
408
Sinclair, J. 303, 408
Skolithos 62, 80 301, 325-
328, 392, 393, 401-
403, 417, 435
Smith, J. 314-317,
408
Sorgel, G. 316, 318-
320, 408

South Africa
(Devonian) 306

Southgate beds 285, 334,

356, 404, 415, 418,

419, 429

Sphenophyllum 301, 394

spinifer, Aglaspis 358

squamata,

Scolecolepis 396
State Avenue,

Cincinnati 355, 356,

359, 362, 366, 368,
372, 426-430, 433,
435, 436
stelliforme, Asteria-

cites 57, 59 300, 313,
402, 412, 414, 417

Stonelick Creek,
Clermont County,
Ohio 04, 306-
308, 313, 337, 340,
341, 343, 348, 349,
358, 377, 395, 396,
403, 404, 412, 413,
420, 421, 425, 426,
436, 437
353-355,
365, 408

Stgrmer, L.

strandi, Mero-

stomichnites 365
striatum, Palaeo-
phycus 76 374, 377,
431
strozzii,
Paleodictyon 384
subangulatum,
Rusophycus 59 301, 305,
414
succulens,
Chondrites 69 300, 333,
424
suleata, Walcottia 301, 378,
398, 399

suleatum, Fucu-

sopsis .69, 70,71 300, 347,
380, 381, 419, 425,
426
Sunset beds 285, 404,
432, 433
Sweden, Lower
Cambrian . 353
AG
Talawanda Creek,
Oxford, Ohio 425
Tanners Creek
Formation 286
Tauber, A. 328-331,
335, 336, 408
Taunus Quartzite
(Rhineland:
Devonian) ... 319
Taylor Creek Rd.,
Clermont County,
Ohio 34, 377,
413
tenuiramosum,
Mastigograptus . 387, 388
Teichichnus 349
Teratichnus 75 301, 364,
368, 369, 430
Thorson, G. . ., 292, 408
Torell, O. 316,317
tortuosum, Palaeo-
phycus 76 374, 431
Trachomatichnus

73, 74 301, 362,
366-368, 402, 428,

429, 435

Trenton beds . BBeA SB RE
354, 374, 412, 414,

431

Triarthrus .. aN 355

Trichophycus
60, 68, 70 301, 346-
350, 381, 388, 389,
399, 402-404, 415,

423, 425, 426

Dactylophycus . 300, 346
trifidum, Asaphoi-

dichnus 65, 71-73 283, 296,

297, 300, 355-359,

361, 362, 366, 402,

426-428, 430

300, 399

tridigitatum,

truncatum,
Conostichus

INDEX

tuberosus,
Skolithos
tubulare, Palaeo-
phycus 76,77 300, 373,
374, 376, 377, 431,
432
66,71 281, 301,
305, 371, 372, 402,
404, 421, 426
Twelve Mile Creek,
Campbell County,
Kentucky

301, 334
Tylichnus

: 308, 310,
414, 418, 423

Type-A, Chrondrites
64, 67, 83 300, 334-
338, 402, 404, 419,
422, 438

Type-B,

Chondrites 63-65,79 300,
334, 336-339, 402,
404, 418, 420, 434

ype-C,

Chondrites 63, 64,81 300
334, 339, 340, 402,
418, 419, 432, 436

ih

Type-A,
Palaeophycus 76 300, 377,

431

Type-B, Palaeo-
phycus ..... 76,77 301, 377,
431, 432

Type-C, Palaeo-
phycus 77, 83 301, 377,
383, 404, 432, 438

U
Ulrich, E. . 310, 387,
399, 409
Union Levee,
Cincinnati 324, 365,
416, 428-430
University of
Cincinnati . 281, 282,
306, 327, 348, 367,
384, 388, 392
Upper Cretaceous
(Ardennes) . 303
Urechis 319, 324
“Utica Slate”
(New York:
Ordovician) 387
U.S. National
Museum col-
lections .... .. 282, 391
U.S. Route 27, New-
port, Kentucky ... 321, 334,
335, 404, 415-417
U.S. Route 50,
Aurora, Indiana . 349, 377,
438

Vv

Vassoevich, N. 282, 369,

380
venosum,

Trichophycus 300, 301,

321, 334-336, 341,

347-350, 376, 399,

402, 404, 415, 423,

425

Vialov, O. 385, 409

444

Vialov, O., &

Goley, B. 384, 385,
409

virgatum,
Palaeophycus 300, 348,
375, 376
viridis, Phoronopsis 326

volutator,
Corophium 317

W

Walcott, C. 295, 350,

352, 354, 355, 377,
387, 388, 393, 409

67, 69 299, 301,
378-380, 398, 399,

22, 424
Walker Mill Road
(State Avenue)
Cincinnati _..........355, 356,
359, 362, 366, 368,
372, 426-430, 433,

Walcottia

435, 436
Warren County,
Ohioiao 347, 350
Waverly Group
(Ohio: Mississip-
plan)! agree ee 403
Waynesville beds . 285, 286,
390, 416
Weimer, R. &
Hoyt, J. 283, 350
409
Weiss, M. &
Norman C. .......... 284, 409

Weiss, M., Norman,
C., & Sweet, W. ..284, 404,

409

Wells, G. 298, 318,
326, 409

Westergard, A. ......316, 317,

319, 326, 327, 409
West Fork Creek
(Sheperd Road
locality) Cin-
Cinnatl 7. eons
324, 337, 339, 377,
381-383, 404, 415,
417, 422, 424, 432,

434, 438

Whitewater beds _ 285, 339,

418, 419, 436
Wilhelmshaven,

West Germany .... 303
Willard, B. ....... 283
Wilson, A. ..... SEL SRB

342, 376, 409
Wurm, A. ee 396
Yi
yoyo, Diplo-
craterion 283, 316-
318
Z
Zenkevich, L. &

Birstein, J. . 403, 409
zimmermanni,

Arenicolites ....... 319
Zoophycos ............. 401, 403

XLVII.

XLVIII.

XLIX.

LVII.

Volume I.

II.

Ill.

MUN eetGo-192) 5 oS 8 llppi 35. 8s ac cca caaese ccc es

Australian Carpoid Echinoderms, Yap forams, Shell Bluff,
Ga. forams. Newcomb mollusks, Wisconsin mollusk faunas,
Camerina, Va. forams, Corry Sandstone.

UND se 193) snG sei ppt ASep lane ene ee

Venezuelan Cenozoic gastropods.

Noss t 45198) seeded ppeeco) Dishes

Ordovician stromatoporoids, Indo-Pacific camerinids, Missis-
sippian forams, Cuban rudists.

(Niobe 199-203) spe SG5 ppt G8) pls; 2 cea

Puerto Rican, Antarctic, New Zealand forams, Lepidocy-
clina, Eumalacostraca.

CNowm204) es Gri ppy Gs) plates ee

Venezuela Cenozoic pelecypods

(UN GeeReDS-2LD) EATON pps 070) pee, cc. seosexcceaccosc desea fo casdehscace

Large Foraminifera, Texas Cretaceous crustacean, Antarctic
Devonian terebratuloid, Osgood and Paleocene Foramini-
fera, Recent molluscan types.

(Noss 2l2-217) S84 ppsy, 83) plss, cco cctec:<crescsscee ee ee

Eocene and Devonian Foraminifera, Venezuelan fossil
scaphopods and polychaetes, Alaskan Jurassic ammonites,
Neogene mollusks.

CUNGre218)<estOS Sipps, 5) plat ct

Catalogue of the Paleocene and Eocene Mollusca of the
Southern and Eastern United States.

UNosy 2192224) 670 piss) ple: ren tea ea

Peneroplid and Australian forams, North American car-
poids, South Dakota palynology, Venezuelan Miocene mol-
lusks, Voluta.

Noss225-230) SUS) pp n 42, DIB: ccosseccacceveteecsecciceceesceveeesccsensecs

Venezuela and Florida cirripeds, Antarctic forams, Lin-
naean Olives, Camerina, Ordovician conodonts, Niagaran
forams.

UINDompi 2S 252) 2. F20) ppg (20 vp lien acces orcs cates cvecescocaeaenccsesce cece

Antarctic bivalves, Bivalvia catalogue.

ONT CAR AZSED EET Aa 0) Yop Jp) Ce sae eae a ene Cee

New Zealand forams, Stromatoporoidea, Indo-Pacific, Mio-
cene-Pliocene California forams.

(INGs2237-238).) 488) pps 45) pls, occa eno oec ease cecccsacscnveccense

Venezuela Bryozoa, Kinderhookian Brachiopods.
CONGssi2d9-245) e0510) ppr050° ples ce cence

Dominican ostracodes, Texan pelecypods, Wisconsin mol-
lusks, Siphocypraea, Lepidocyclina, Devonian gastropods,
Miocene Pectens Guadaloupe.

(Noes 246-247) 50657) pps G60) ple. 2 oe cates

Cenozoic corals, Trinidad Neogene mollusks.

NGS 2482254) 572 Day 49! PIG. aos cacccnts ccc cccan oc occ ceeccceccceeonecseese

American Foraminifera, North Carolina fossils, coral types,
Belanski types, Venezuelan Cenozoic Echinoids, Cretaceous
Radiolaria, Cymatiid gastropods.

(tis ZEEE ey A) i a

Alaskan Jurassic ammonites, Pt. II, Jurassic Ammonitina
New Guinea.

PALAEONTOGRAPHICA AMERICANA

See Johnson Reprint Corporation, 111 Fifth Ave., New York,

N. Y. 10003

Monographs of Arcas, Lutetia, rudistids and venerids.
IN eel ee ef yee I Ui) [ee EA nee
Heliophyllum halli, Tertiary turrids, Neocene Spondyli,

Paleozic cephalopods, Tertiary Fasciolarias and Pale-

ozoic and Recent Hexactinellida.

(Nog reldec5) ots ppatOLo pISiy oe
Paleozoic cephalopod structure and phylogeny, Paleozoic

siphonophores, Busycon, Devonian fish studies, gastropod

studies, Carboniferous crinoids, Cretaceous jellyfish,

Platystrophia and Venericardia.

(ONS SCR ED RG ET ela i 7A oC hermes re see Re
Rudist studies Busycon, Dalmanellidae, Byssonychia, De-

vonian lycopods, Ordovican eurypterids, Pliocene mollusks.
(ING pr 154-50) op FADD LOU lets cavers cesses. ve ccsecsm cease ccvcccmeescasten
Tertiary Arcacea, Mississippian pelecypods, Ambonychiidae,

Cretaceous Gulf Coastal forams.

(Go TeE ECT) Eel) ay Bye 0g 0) fy eerste ane ne
Lycopsids and sphenopsids of Freeport Coal, Venericardia,

Carboniferous crinoids.

16.00

18.00
16.00

16.00

18.00

18.00

18.00

18.00

18.00

18.00
18.00

18.00

18.00

18.00

18.00

23.00

28.00

28.00

32.00

20.75

BULLETINS OF AMERICAN PALEONTOLOGY

Vols. I-XXIII. See Kraus Reprint Corp., 16 East 46th St., New York,

XXIV.

XXV.

XXVI.

XXXI.

XXXII

XXXIII.

XXXIV.

XXXV.

XXXVI.

XXXVII.

XXXVIII.

XXXIX.

XL.

N. Y. 10017, U.S.A.

(Oy EA ep ELM ae Pr) Py be kno ee ee ee
Mainly Paleozoic faunas and Tertiary Mollusca.

(Nos: {88-948 306)spp,,, (30) pln coe ence

Paleozoic fossils of Ontario, Oklahoma and Colombia, Meso-
zoic ehinoids, California Pleistocene and Maryland Mio-
cene mollusks.

(Nos. 95-100). 420 pp., 58 pls. .......

Florida Recent marine shells, Texas Cretaceous fossils,
Cuban and Peruvian Cretaceous, Peruvian Eogene corals,
and geology and paleontology of Ecuador.

(ONos. LO1=108) 3. 5 S176) spe 73169 poe aes tes
Tertiary Mollusca, Paleozoic cephalopods, Devonian fish
and Paleozoic geology and fossils of Venezuela.

(Nos. 109-114). 412) pp:, 34 pls: ee
Paleozoic cephalopods, Devonian of Idaho, Cretaceous and
Eocene mollusks, Cuban and Venezuelan forams.

(Nos. 115-116). 738 pp., 52 pls. ..............
Bowden forams and Ordovician cephalopods.

(No. 117). $63. :pp:, .65\ pla. 222.2 =
Jackson Eocene mollusks.

(Nos 218-128) 5.458) pps, a7, PIS. cocsecc seen eee

Venezuelan and California mollusks, Chemung and Pennsyl-
vanian crinoids, Cypraeidae, Cretaceous, Miocene and
Recent corals, Cuban and Floridian forams, and Cuban
fossil localities.

(Nos; (129-133). -294cpp.39 spl: fe ree
Silurian cephalopods, crinoid studies, Tertiary forams, and
Mytilarca.

(Nos: 134-139)5. 498) ppg SU hae eeccsceececeecesecresece tenement
Devonian annelids, Tertiary mollusks, Ecuadoran sstrati-
graphy paleontology.

(Nos: 140=145)5, 4.00 spp. 19) pals coeccas nacre scene

Trinidad Globigerinidae, Ordovician Enopleura, Tasmanian
Ordovician cephalopods and Tennessee Ordovician ostra-
cods and conularid bibliography.

(Nos. 146-154) 386: ipp:,, 31. pls: Se

G. D. Harris memorial, camerinid and Georgia Paleocene
Foraminifera, South America Paleozoics, Australian
Ordovician cephalopods, California Pleistocene Eulimidae,
Volutidae, and Devonian ostracods from Iowa.

(Nos: 155-360). 9412). pp:. (53 ple... ee

Globotruncana in Colombia, Eocene fish, Canadian Chazyan
Antillean Cretaceous rudists, Canal Zone Foraminifera,
fossils, foraminiferal studies.

(Nos: 161-164). 486 pp3w37) plo res
Antillean Cretaceous Rudists, Canal Zone Foraminifera,
Stromatoporoidea.

(Nos, 165-176). 447-pp., (530 pls. Sereno eccccoeee ae cscs

Venezuela geology, Oligocene Lepidocyclina, Miocene ostra-
cods, and Mississippian of Kentucky, turritellid from Vene-
zuela, larger forams, new mollusks, geology of Carriacou,
Pennsylvanian plants.

(Nos: 177-183). 448pp;, 36uplsi ee eee,

Panama Caribbean mollusks, Venezuelan Tertiary forma-
tions and forams, Trinidad Cretaceous forams, American-
European species, Puerto Rico forams.

(Nos 184) 57-996 pps Cipla: -ee ee
Type and Figured Specimens P.R.I.

12.00

12.00

14.00

14.00

14.00

18.00
16.00

16.00

16.00

16.00

16.00

16.00

16.00

16.00

18.00

16.00

18.00

tae hie Re

me

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