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North Carolina

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I. Michael Heyman

Secretary

Smithsonian Institution

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 83

Oligocene Echinoids of
North Carolina

Porter M. Kier

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.
1997

ABSTRACT

Kier, Porter M. Oligocene Echinoids from North Carolina. Smithsonian Contributions to
Paleobiology, number 83, 37 pages, 6 figures, 11 plates, 2 tables, 1997.—Oligocene echinoids
are rare, which makes important this material from three quarries in North Carolina. Three
species occur in the state quarry at Pollocksville: Psammechinus carolinensis, new species,
Maretia carolinensis, new species, and Agassizia sp. At the New Bern quarry occur
Rhyncholampas gouldii (Bouve) newbernensis, new subspecies, Agassizia mossomi Cooke,
Psammechinus carolinensis, new species, Dixieus dixie (Cooke), Clypeaster rogersi (Morton),
and Maretia sp. Periarchus lyelli (Conrad) is found in underlying Eocene beds. The Belgrade
quarry fauna consists of Arbia aldrichi (Clark), Gagaria mossomi (Cooke), Echinocyamus
wilsoni, new species, and Agassizia mossomi Cooke. These echinoids indicate a middle to late
Oligocene age for the Trent Formation (= River Bend) and Belgrade Formation (= River
Bend).

Official publication date is handstamped in a limited number of initial copies and is
recorded in the Institution’s annual report. Annals of the Smithsonian Institution. SERIES COVER
DESIGN: The trilobite Phaecops rana Green.

Library of Congress Cataloging-in-Publication Data
Kier, Porter M.

Oligocene Echinoids of North Carolina / Porter M. Kier.
p. cm. — (Smithsonian contributions to paleobiology ; no. 83)

Includes bibliographical references (p. 14).

1. Sea urchins. Fossil—North Carolina. 2. Paleontology—Oligocene. 3. Animals, Fossil—North Caro¬
lina. I. Title. II. Series.

QE701.S56 no. 83 [QE783.E2] 560 s—dc21 [563'.95] 97-3660

@ The paper used in this publication meets the minimum requirements of the American
National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.

Contents

Page

Introduction. 1

Acknowledgments. 1

Occurrences.1

Pollocksville Quarry.1

New Bern Quarry ... . . .2

Belgrade Quarry.2

Ecology. 3

Family Phymosomatidae Pomel.3

Genus Dixieus Cooke. 3

Dixieus dixie (Cooke) .3

Family Arbaciidae Gray.4

Genus Arbia Cooke.4

Arbia aldrichi (Clark) .4

Family Uncertain.5

Genus Gagaria Duncan. 5

Gagaria mossomi (Cooke) ... 5

Family Echinidae Gray . . 5

Genus Psammechinus Agassiz and Desor.5

Psammechinus carolinensis, new species.5

Family CASSIDULIDAE Agassiz and Desor .6

Genus Rhyncholampas Agassiz.6

Rhyncholampas gouldii gouldii Bouve . . . . ... . -6

Rhyncholampas gouldii newbernensis, new subspecies.6

Family Fibulariidae Gray. 8

Genus Echinocyamus van Phelsum. 8

Echinocyamus wilsoni, new species. • 8

Family Protoscutellidae Durham. 8

Genus Periarchus Conrad . . . . . • 8

Periarchus lyelli (Conrad) . . . . . 8

Family Clypeasteridae Agassiz. • ■ • • 8

Genus Clypeaster Lamarck ... . . • • • • • • 8

Clypeaster rogersi (Morton). .... . 8

Family SCHLZASTERIDAE Lambert. 10

Genus Agassizia Agassiz and Desor.10

Agassizia mossomi Cooke . . 10

Agassizia ..11

Family Spatangidae Gray.H

Genus Maretia Gray . . . . H

Maretia carolinensis, new species. 11

Maretia ..13

Literature Cited .14

Plates.^

m

Oligocene Echinoids of
North Carolina

Porter M. Kier

Introduction

Oligocene echinoids are rare, having less than one-third the
number of species as in the Eocene; therefore, these new
Oligocene echinoids are particularly important. Climatic
cooling is considered by McKinney et al. (1992) to be the cause
for the reduction in species diversity. The species described
herein are abundant in three quarries in North Carolina near
Pollocksville, Belgrade, and New Bern. Although some of the
species are new, others have been previously described from
elsewhere and are useful in suggesting the age of the beds
bearing the echinoids. Four of the echinoid species of the Trent
Formation (= River Bend) at the New Bern quarry occur
elsewhere in beds dated as middle to late Oligocene. Likewise,
three species from the Belgrade quarry that occur in the
Belgrade (= River Bend) Formation are found elsewhere in the
late Oligocene. The echinoids at the state quarry near
Pollocksville are new species and are of little use in age
determination.

Generally, the echinoids differ at the three quarries,
suggesting age or environmental difference between the sites.
Three of the four echinoids at the Belgrade quarry are not found
at New Bern or Pollocksville. Only Agassizia mossomi Cooke
occurs at both Belgrade and New Bern. Only two of the six
Trent species at New Bern are found at either of the two other
sites. Psammechinus carolinensis, new species, occurs at New
Bern and Pollocksville, and Agassizia mossomi occurs at New
Bern and Belgrade. None of the three species at Pollocksville
occur at Belgrade, and only one of the Pollocksville species, P.
carolinensis, occurs at New Bern.

Acknowledgments. —I thank Druid Wilson who found
most of the echinoids, took me to the outcrops, and encouraged
me to describe these echinoids. Victor Zullo, William Harris,

Porter M. Kier, P.O. Box 162, Lotts burg, Virginia 22511.

Review Chairman: William Dimichele, Department of Paleobiology,
Smithsonian Institution, Washington, D.C. 20560.

and Jerry Baum visited the quarries with me and discussed the
stratigraphy. They critically read the manuscript. Lauck Ward
also read the manuscript and collected some of the specimens.
Pete Harmatuk spent many hours in these quarries and
collected many echinoids. The photography and measurement
of specimens was by Arnold Powell, and the artwork was
rendered by Mary Parrish.

The publication of this paper has been delayed for ten years,
and I particularly thank Stephen Donovan and Rich Mooi for
making many suggestions to bring it up-to-date, and for their
other recommendations. Diane M. Tyler edited the manuscript
for the Smithsonian Institution Press.

Occurrences

Pollocksville Quarry

This state quarry, utilized by the North Carolina Department
of Transportation, is located 4 km NNE of Pollocksville on the
left bank of the Trent River in Jones County, North Carolina. It
is within the type section of the Trent Formation as proposed by
Baum et al. (1978:12), and it is part of the River Bend
Formation of Ward et al. (1978:10). As described by Baum et
al. (1979:100), the section exposes 3.2 m of the Trent
Formation, with the echinoids occurring in a sandy biosparite.

Zullo and Baum (1979:235) consider the Trent to be early to
middle Oligocene. Ward et al. (1978:12) consider these beds
(part of their River Bend Formation) to be middle Oligocene
(late Vicksburgian) on the basis of the abundance of Pecten
perplanus byramensis Gardner. Ward (pers. comm., 1984) now
thinks that these beds are late Oligocene (Chickasawhayan).

Three species occur in these beds: Psammechinus carolinen¬
sis, new species, Maretia carolinensis, new species, and
Agassizia sp. The first two species are represented by a large
number of specimens, but only two have been found of
Agassizia sp. None of the species is useful in dating the beds.
Maretia carolinensis and Psammechinus carolinensis are very
distinct from any other species of their genus in this

1

2

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

hemisphere. Agassizia sp. is too poorly represented to be able
to ascertain the degree of its affinity with the species it most
closely resembles, Agassizia wilmingtonica Cooke from the
middle Eocene Castle Hayne Formation of North Carolina. The
fact that both Agassizia and Maretia are found at the New Bern
quarry in beds referred to the Trent Formation (= River Bend
of Ward et al., 1978:10) suggests that the environmental
conditions at the Pollocksville and New Bern quarries were
similar. That both species are very different from their
congeners at New Bern suggests that the beds were not
deposited at the same time. Only Psammechinus carolinensis
occurs at both sites. Ward (pers. comm., 1984) now believes,
on the basis of the mollusk fauna, that the beds where the
echinoids occur at the Pollocksville quarry are Chick-
asawhayan, whereas the Trent or River Bend at New Bem is
Vicksburgian.

New Bern Quarry

This Martin Marietta Company quarry is northwest of New
Bem, Craven County, North Carolina, 0.8 km NE of route 55
on county road 1402. Four or five stratigraphic units (Table 1)
are recognized in the quarry. The lower beds (in ascending
order) are Castle Hayne Limestone, New Bem Formation
(Baum et al. (1978:9)) = Spring Garden Member of Castle
Hayne Limestone (Ward et al. (1978:9)), and Trent Formation
(Baum et al. (1978:12)) = River Bend Formation (Ward et al.
(1978:10)). At the top, occur beds assigned to the Yorktown/
Duplin by Baum et al. (1979:103) and to Duplin/Croatan
Formations by Ward et al. (1978:18).

Zullo and Harris (pers. comm., 1984) have found, below an
unconformity at the base of the River Bend Formation, a
bryozoan biomicrudite that they consider to be the Castle
Hayne Limestone equivalent to the Ward et al. (1978:8)
Comfort Member. Periarchus lyelli (Conrad) occurs in large
numbers in these beds. It also is present in small numbers in the

New Bem Formation = Castle Hay Formation (Spring Garden
Member). This is the only echinoid found in either formation
and confirms the Eocene age of these beds.

Echinoids are very common in the overlying Trent or River
Bend Formation. Two species occur in great numbers,
Rhyncholampas gouldii (Bouve) newbernensis, new subspe¬
cies, and Agassizia mossomi Cooke. A few specimens are
present of Dixieus dixie Cooke, Clypeaster rogersi (Morton),
and Maretia sp. One specimen was found of Psammechinus
carolinensis, new species. This species occurs in great numbers
in the state quarry at Pollocksville. The fauna appears to be
middle to late Oligocene. Rhyncholampas gouldii is known
elsewhere from the Marianna Limestone (middle Vicksburgian,
Bryam Limestone (late Vicksburgian), Suwannee Limestone
(Vicksburgian-Chickasawhayan), and Flint River Formation
(Chickasawhayan). Agassizia mossomi is know from the
Suwannee and Clypeaster rogersi from the Marianna and Flint
River. The age of Dixieus dixie is uncertain. It has been found
before at a locality where both the late Eocene Ocala and the
Suwannee Limestone occur.

No echinoids are present in the beds overlying the Trent
Formation.

Belgrade Quarry

The Martin Marietta Belgrade quarry (see Baum et al.
(1979:98) and Wardet al. (1978:19) for descriptions of section)
is east of the intersection of route 1434 and U.S. 17, Belgrade,
Onslow County, North Carolina. According to Baum et al.
(1979:98), 11 meters of section is exposed. The lower eight
meters is their type section for the Belgrade Formation (Baum
et al. (1978:13)). Ward et al. (1978:20) refer this part of the
section (Table 2) to their River Bend Formation. Baum et al.
(1979:99) assign a lower Miocene age to these beds, but Ward
et al. (1978:12) consider them late Oligocene, Chicka¬
sawhayan. Three echinoid species occur 1-45 cm below the

TABLE 1.—Stratigraphic occurrences of echinoids at the New Bern quarry.

Baum et al. (1978)

Ward et al. (1978)

Echinoid species

Early-Middle

Oligocene

Trent Formation

Middle-Late

Oligocene

River Bend Formation

Psammechinus carolinensis, new species

Dixieus dixie Cooke

Rhyncholampas gouldii newbernensis, new species
Clypeaster rogersi (Morton)

Agassizia mossomi Cooke

Maretia sp.

New Bern Formation

<D <l>

x S

Castle Hayne Formation

Periarchus lyelli (Conrad)

XI CJ

S o

2 UJ

(Spring Garden Member)

<D

C

O

Castle Hayne Formation

Periarchus lyelli (Conrad)

u

NUMBER 83

3

TABLE 2.—Stratigraphic occurrences of echinoids at the New Bern quarry.

Baum et al. (1978)

Wardet al. (1978)

Echinoid species

Early

Miocene

Belgrade Formation

Late

Oligocene

River Bend Formation

Arbia aldrichi (Clark)

Gagaria mossomi (Cooke)

Agassizia mossomi Cooke
Echinocyamus wilsoni, new species

top of this formation: Arbia aldrichi (Clarke), Gagaria
mossomi (Cooke), and Agassizia mossomi Cooke. All three of
these species occur elsewhere in the late Oligocene. Arbia
aldrichi occurs in the Chickasawhayan and Paynes Hammock
formations (Chickasawhayan). Gagaria mossomi and Agas¬
sizia mossomi occur in the Suwannee Limestone (Vicksbur-
gian-Chickasawhayan). The specimens of A. mossomi at
Belgrade are slightly different from those at the New Bern
quarry, which occur in beds considered to be late Oligocene.

Specimens of Echinocyamus wilsoni, new species, were
collected from this quarry from beds that Druid Wilson (pers.
comm., 1985) believes occur lower in the Belgrade Formation.

Ecology

These echinoids are similar enough to living species that it is
possible to infer their living habits. Periarchus lyelli (Conrad)
is found at the New Bern quarry in both the Castle Hayne
Limestone (Comfort Member) and New Bern (Castle Hayne,
Spring Garden Member) formations. These formations are a
biomicrudite (Zullo, pers. comm., 1985) and a sandy biospar-
ridute (Baum et al., 1979:102), respectively. The sediment that
formed these rocks would have provided the sandy environ¬
ment for this sand dollar.

The Trent Formation (River Bend Formation) is a sandy
biosparite (Baum et al., 1979:102). Four of the five species
found in this formation would have lived buried in sediment.
The spatangoid Maretia sp. probably lived within a burrow
with a tube leading posterior for a drainoff as indicated by the
presence of a subanal fasciole. The lack of a deep anterior
groove and the lack of crowding of the porepairs in the anterior
ambulacrum indicate a limited funnel-building ability. Agas¬
sizia mossomi Cooke most likely also lived buried in the
sediment. Modem species of this genus apparently live in
sandy environments (Kier, 1984:60). They lack the well-
developed funnel-building tubefeet needed for maintenance of
a tunnel through mud to the surface. Modem species have been
collected only on sandy bottoms. Clypeaster rogersi (Morton)
presumably lived like the modem Clypeaster subdepressus
(Gray), which it resembles. This species lives (Kier and Grant,
1965:30) in sand buried as deeply as 25 mm below the surface,
or it moves along the top of the sand with sand and shell
fragments held by its tubefeet over the top of its test. The

cassiduloid Rhyncholampas gouldii newbernensis, new sub¬
species, no doubt lived like the modem cassiduloid Cassidulus
cariboearum Lamarck. This species lives buried (Gladfelter,
1978; Kier, 1975; Mooi, 1990) in the sand, but because it
cannot build a funnel, the sediment has to be well aerated.
Dixieus dixie Cooke was a regular echinoid and lacked the
ability to bury. It probably fed on grasses growing on the
sediment.

The environment during the deposition of the sandy
biosparite of the Trent Formation (River Bend Formation) at
the state quarry at Pollocksville probably was similar to that of
the New Bern quarry. Species of Maretia and Agassizia
likewise occur here. These genera also occur in a biomicrudite
in the Belgrade Formation (River Bend Formation) at the
Belgrade quarry. The most common species is again Agassizia
mossomi, suggesting well-aerated sediments. The two regulars,
Arbia aldrichi (Clarke) and Gagaria mossomi (Cooke),
probably lived on top of the sediment and fed on grasses.
Finally, Echinocyamus wilsoni presumably lived buried in the
sediment. Modem species of Echinocyamus live buried in a
coarse- or fine-grained bottom (Mortensen, 1948:162).

In summary, the fauna at the three quarries is dominated by
species that would have lived buried or partially buried in
well-aerated sediments. The water was probably warm. Modem
clypeasterids are confined to tropical and subtropical coasts.
Agassizia has two extant species, both of which are found only
in tropical waters.

Family Phymosomatidae Pomel
Genus Dixieus Cooke

Dixieus dixie (Cooke)

Plate 1: Figures 1-4

Phymosoma dixie Cooke, 1941:17, pi. 2: fig. 15.

Dixieus dixie (Cooke).—Cooke, 1948:607; 1959:24, pi. 5: figs. 8-10.—

Mortensen, 1951:558, fig. 281.

Description. —Only one specimen (USNM 398317 from
New Bern quarry) was collected during this study. Its dorsal
side is weathered with the tuberculation partially destroyed, but
the ventral side is well preserved. Its tuberculation, number of
plates, nature of its ambulacra, size of its peristome, depth of its

4

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

peristomial slits, and shape of its test are very similar to the
holotype.

Comparison with Other Species. —This specimen differs
from the specimens referred to Dixieus cf. D. dixie (Cooke) by
Kier (1980:21) from the middle Eocene Castle Hayne
Limestone. It differs in having more plates relative to its size
than in the Castle Hayne specimen. The New Bern specimen,
which is 32 mm in diameter, has 196 porepairs in an
ambulacrum or 6.1 porepairs per mm of length. The ratio in the
holotype from Florida is also 6.1, whereas the two Castle
Hayne specimens have ratios of 4.3 and 4.5.

Material. —-USNM 398317, in float from Trent Formation,
New Bern quarry, North Carolina, Judy Raypor collector.
USGS 12747 (holotype) and USGS 1515c (paratype), from a
quarry east of the Steinhatchee River north of U.S. Highway 19
near Clara, Dixie County, Florida.

The holotype and paratype were considered by Cooke
(1959:24) to be from the late Eocene Ocala Limestone;
however, according to Druid Wilson (pers. comm., 1985) the
fauna at this locality is a mixture of fossils from the Ocala
Limestone and Oligocene Suwannee Limestone. USGS 15150
is cataloged as “Suwannee Is. and Ocala Is.”

OCCURRENCE. —Florida: Suwannee Limestone and Ocala
Limestone, Dixie County.

North Carolina: Trent Formation, New Bern quarry.

Family Arbaciidae Gray
Genus Arbia Cooke

Arbia aldrichi (Clark)

Figure 1; Plate 1: Figures 5, 6

Coelopleurus aldrichi Clark, 1915:158, pi. 73: figs. 6a,b, 7a-c.

Arbacia aldrichi (Clark).—Cooke, 1941:11.

Arbia aldrichi (Clark).—Cooke, 1948:606; 1959:21, pi. 3: figs. 15-17.—

Mortensen, 1951:558, fig. 280a-c.

Description. —Three specimens and three fragments are
available from this study. Parts of the specimens are suffi¬
ciently well preserved to permit certainty as to the identifica¬
tion of the specimens. They are similar in all respects to the
holotype and paratype of this species. Cooke (1959:22)
considered both of Clark’s figured specimens to be cotypes, but
Clark in his figure explanation (1915, pi. 73) states that the
specimen in his figures 6a and 6b is the type. Mortensen
(1951:558) regreted that no detail drawings were available
showing the ambulacral structure. I include, herein, drawings of
ambulacral plates in the figured paratype (Figure lB) and in one
of the specimens from Belgrade quarry (Figure 1a).

It has not been noted previously that some of the primary
tubercles are crenulate. These crenulations are present on
well-preserved tubercles on the holotype, on some of the
Belgrade quarry specimens, and on a specimen from the Paynes
Hammock Sand of Mississippi.

Material. —USNM 398318, 1 specimen, 45 cm below top

A

B

FIGURE 1 .—-Arbia aldrichi (Clark): A, ambulacrum and part of interambula¬
crum at ambitus of USNM 398318, from Belgrade Formation (of Baum et al.,
1978) at Belgrade quarry (x6); B, ambulacral plates near ambitus of paratype
USNM 559594B, from St. Stephens Limestone, Perdue Hill, Alabama (x8).

of indurated limestone, Belgrade Formation, Belgrade quarry,
North Carolina, Druid Wilson collector. USNM 398319, 1
specimen, near contact with indurated limestone, Belgrade
Formation, Belgrade quarry, Druid Wilson collector. USNM
559594B, 1 cotype, St. Stephens Limestone, Perdue Hill,
Alabama.

OCCURRENCE. —North Carolina: Three specimens and
fragments were collected by Druid Wilson from the south face
of Belgrade quarry toward the northeast comer in a dark bed
approximately 5-25 cm below contact with indurated lime¬
stone. According to Wilson (pers. comm., 1984), this horizon is
high in zone C of Baum et al. (1979, fig. 1).

Alabama: The type specimens (cotypes, USNM 559494)
come from the late Oligocene St. Stephens Limestone at Perdue
Hill, Monroe County, Alabama.

Mississippi: One specimen is from the late Oligocene
Paynes Hammock Sand at Patton Creek in Mississippi.
According to Cooke (1959:22), Mac Neil originally referred the
bed at Patton Creek to the upper Chickasawhay Formation, but
later he called it the Paynes Hammock Sand Formation.

NUMBER 83

5

Family Uncertain
Genus Gagaria Duncan

Gagaria mossomi (Cooke)

Plate 2: Figures 1-5

Thylechinus ( Gagaria ) mossomi Cooke, 1941:13, pi. 2: fig. 16; pi. 3: figs. 6-9;

pi. 4: figs. 3-5.

Gagaria mossomi (Cooke).—Cooke, 1959:17, pi. 3: figs. 10-14.

Description. —Six new specimens were found of this
species. One of these specimens is beautifully preserved, with
the surface of the test not weathered and with the details of the
ornamentation clearly visible. This specimen is similar in all
respects to the type specimens of the species in that it has: (1)
a similar number of plates; (2) similar crenulate, imperforate
tubercles that are arranged in the same way; (3) ambulacral
plates of similar number and arrangement; and (4) a test of
similar shape. Because this specimen is far better preserved
than any other specimens yet collected of this species, many
photographs are included.

OCCURRENCE.— North Carolina: Six specimens were

collected by Druid Wilson from the south face of Belgrade
quarry toward the northeast comer in a dark bed approximately
5-25 cm below contact with indurated limestone. According to
Wilson (pers. comm., 1984), this horizon is high in zone C of
Baum et al. (1979, fig. 1).

Florida: The species previously was known only from the
(Vicksburgian-Chickasawhayan) Oligocene Suwannee Lime¬
stone of Florida.

Family Echinidae Gray
Genus Psammechinus Agassiz and Desor

Psammechinus carolinensis, new species

Figure 2; Plate 3: Figures 1-4; Plate 4: Figures 1-5

Diagnosis. —The species is characterized by large primary
tubercles, slightly arcuate porepairs, and high ambulacral
plates.

DESCRIPTION. —Twenty-nine specimens were measured for
the following characteristics.

Shape and Size: Diameter (D) varying from 5.4 to 18.1 mm
(mean 10.9); height varying from 46.5 to 58.7 percent of D
(mean 52.5). Marginal outline of test circular; test very slightly
depressed around peristome.

Apical System: No plates preserved; diameter of opening
19.4-27.2 percent of D (mean 23.5).

Ambulacra: Narrow, width 20.0-30.1 percent of D (mean
25.7). Plates trigeminate (Figure 2), arranged in echinoid
pattern with each compound plate with three elements,
demiplate between two primary plates, adoral one being
largest; 10 compound plates in single poriferous zone of
specimen 6.9 mm in diameter, 14 in specimen 11.0 mm in

FIGURE 2 .—Psammechinus carolinensis, new species: ambulacral plates near
ambitus of the holotype, USNM 398321, from float of the Trent Formation,
Pollocksville state quarry (x20).

diameter, and 18 in specimen 18 mm in diameter. Porepairs
arranged in arcs of three (Plate 3: Figure 4) with 36 porepairs in
poriferous zone of specimen 8.3 mm in diameter, 45 in
specimen 12.8 mm in diameter, and 55 in specimen 16.2 mm in
diameter. No phyllodes or crowding of porepairs at margin of
peristome.

Interambulacra: Plates low, at ambitus same height or
slightly higher than adjacent compound ambulacral plate; 9
plates in single column of specimens 5.4 mm in diameter, 12 in
specimen 11.1 mm in diameter, 15 in specimen 18.1 mm in
diameter.

Peristome: Circular, large, diameter 35.7-50.7 percent of
D (mean 42.0). Peristomial (“gill”) slits slightly developed
(Plate 3: Figure 2).

Lantern Supports: Auricles joining to form arch (Plate 4:
Figure 5).

Tuberculation: Ambulacra with two vertical rows of
primary, imperforate, noncrenulate tubercles in each area (Plate
4: Figure 3). Two inner rows of much smaller secondary
tubercles (more distinct on larger specimens). On compound
plate, primary tubercle occupies full height of plate, secondary
tubercle forming part of inner row of tubercles occurring near
medial suture; three or four smaller tubercles irregularly
situated on plate.

Interambulacra with two vertical rows of primary, imperfo¬
rate, noncrenulate tubercles in each area (Plate 4: Figure 4).
Tubercle in middle of each plate occupying almost entire height
of plate. Two inner rows of much smaller secondary tubercles;
irregularly arranged secondary tubercle of varying size between
primary tubercle and ambulacra.

Remarks. —This species is referred to the order Echinoida
because of its nonsculptured test, shallow peristomial slits,

6

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

imperforate, noncrenulate tubercles, and ambulacral plates
compounded in the echinoid manner. Of all the genera in this
order it is most similar to Psammechinus. Unfortunately, two of
the main characters of the genus, the strongly plated buccal
membrane and the structure of the globiferous pedicellariae are
not known on this species, so it is not possible to be certain that
this species belongs to Psammechinus.

Comparison with Other Species. —This species is easily
distinguished from the Yorktown Formation (Pliocene) species
of Psammechinus, P. philanthropus (Conrad), by its larger
primary tubercles (relative to the secondary), its less arcuate
porepairs with the adoral-most porepairs in each compound
plate less shifted medially, and its higher ambulacral plates.

Material. —Hundreds of specimens occur concentrated
and jumbled in a sandy biomicrudite in the Pollockville state
quarry. The tests occur haphazardly. This fact together with the
absence of any jaws, teeth, or apical-system plates suggests that
the specimens were transported after death. Most of the
specimens are covered with angular quartz sand grains, which
are partially embedded into their tests.

Holotype: USNM 398321, from float of the Trent Forma¬
tion, Pollocksville state quarry, North Carolina.

Figured Paratypes: USNM 398322, 398323, 398474,
same data as holotype.

Occurrence. —North Carolina: Hundreds of specimens
occur in the Pollocksville state quarry. One specimen also was
collected presumably from the Trent Formation, New Bern
quarry.

Family Cassidulidae Agassiz and Desor
Genus Rhyncholampas Agassiz

Rhyncholampas gouldii gouldii Bouve

Rhyncholampas gouldii Bouve, 1846:192. [For a complete synonymy and list

of localities see Cooke (1959:37).]

Rhyncholampas gouldii newbernensis, new subspecies

Figure 3; Plate 5: Figures 1-7

Diagnosis. —The subspecies is characterized by long petals,
a large peristome, a low test, and the periproct with a high
opening.

Description. —The descriptions of the following character¬
istics are based on 34 specimens that are generally well
preserved and not distorted in shape.

Shape and Size: Length 19.2-44.7 mm (mean 33.0); width
83.4-97.5 percent of L (mean 92.1); height 41.1-50.9 percent
of L (mean 46.0). Test inflated with greatest height slightly
posterior to apical system; oral surface flat, greatest width
posterior to center.

Apical System: Anterior, distance from anterior margin
48.3-56.6 percent of L (mean 52.4); monobasal, 4 genital
pores, absent on specimens less than 20 mm long.

Ambulacra: Anterior petal short, extending three-fifths
distance from apical system to anterior margin; length
29.6-39.5 percent of L (mean 35.0). Anterior petal narrower
than other petals, greatest width 10.3-13.2 percent of L (mean
11.8). Anterior petal closing distally, with 76 porepairs in
specimen 44.7 mm long, 44 in specimen 19.2 mm long.
Anterior paired petals (II, IV) extending almost two-thirds
distance from apical system to margin; length 32.7-43.2
percent of L (mean 36.3); greatest width 11.2-13.9 percent of
L (mean 12.7). Anterior paired petals with 72 porepairs in
specimen 44.7 mm long, 48 porepairs in specimen 19.2 mm
long.

Posterior petals (V, I) extending three-fifths distance from
apical system to posterior margin; length 34.8-49.0 percent of
L (mean 40.8); greatest width 9.9-14.3 percent of L (mean
12.5). Posterior petals with 84 porepairs in specimen 44.7 mm
long, 46 porepairs in specimen 19.2 mm long. Posterior petals
closing distally. Poriferous zones equal in length in anterior
petal, but in anterior paired petals posterior poriferous zones
longer, with 1.3 more porepairs; in posterior petals, anterior
poriferous zone longer by 1-3 porepairs.

Phyllodes single pored, well developed (Plate 5: Figure 6); a
specimen 44 mm long with 22 pores in phyllode in
ambulacrum III, 26 in II, 22 in I; inner series of pores with
approximately 5 pores in each phyllode.

Interambulacra: Small, irregularly arranged pits in mid¬
zone of interambulacrum 5; a few in interambulacra 2 and 3.

Peristome: Anterior of center, distance from anterior
margin 22.9-38.2 percent of L (mean 34.1); width 10.0-17.3
percent of L (mean 13.4); height 7.5-13.0 percent of L (mean
9.6). Bourrelets strongly developed.

Periproct: Supramarginal, transverse with slight trough
extending posteriorly.

Remarks. —Specimens of this subspecies are distinguished
from species of Rhyncholampas gouldii gouldii by the
following features: (1) longer petals (Figure 3e-G); (2)
peristome slightly higher and wider (Figure 3a,B); (3) lower test
(Figure 3d); and (4) higher periproct (Figure 3c). The presence
of overlap in most of these characters between the two taxa
prevents their consideration as separate species. In all other
characters the two taxa are indistinguishable.

MATERIAL. — Holotype: USNM 398324, from float of the
Trent Formation, New Bern quarry, North Carolina.

Figured Paratype: USNM 398325, same data as holotype.

Nonfigured Paratypes: USNM 492065-492096, same
data as holotype.

Occurrence —North Carolina: Trent Formation, New
Bern quarry.

HEIGHT OF PERIPROCT WDTH OF PEHCTOME

NUMBER 83

7

LENGTH OF TEST 20 '

o Rhynchdamoas gcxida youkJn Botrve'

o Rhynchoiamoas gtxJdn ooukJ» Bouve'

• Rrryncholamoas otxiO newbemenss. new subspeoes

5 10 15 20 25 30 35 40 45 mm

LENGTH OF TEST

c

5 10 16 20 26 30 35 40 45 mm

LENGTH OF TEST

FIGURE 3.—Scattergrams showing difference between specimens of Rhyncholampas gouldii gouldii Bouve and
Rhyncholampas gouldii newbernensis, new subspecies: A, height of peristome; B, width of peristome; C, height
of periproct; D, height of test; E, length of petal III; F, length of petal IV; G, length of petal V.

8

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Family Fibulariidae Gray
Genus Echinocyamus van Phelsum

Echinocyamus wilsoni, new species

Figure 4a-e

Diagnosis. —The species is characterized by a wide test, an
average width of 82 percent of the length, the anterior petal
slightly shorter than other petals, and the periproct situated at
two-thirds the distance from the peristome to the posterior
margin of the test.

DESCRIPTION. —The description is based on eight specimens
from the Belgrade quarry. These specimens were encrusted
with a secondary coat of calcite, which has obscurred the
surface ornamentation.

Shape and Size: Length (L) 2.0-3.5 mm long (mean 2.8);
width 79.6-84.7 percent of L (mean 82.2); height 46.2-52.5
percent of L (mean 49.0). Greatest height at apical system.

Apical System: Located anterior of center at distance from
anterior margin equal to 42.6-45.7 percent of L (mean 43.4);
four genital pores.

Ambulacra: Petal III slightly shorter than other petals but
with approximately same number of porepairs; 8 porepairs in
petal in specimen 2.45 mm long, 8-10 in specimen 2.95 mm
long, and 12 in specimen 3.4 mm long; see Figure 4 a-C for
shape of petals. Distribution of accessory pores not discernible
because of poor preservation of specimens.

Peristome: Central, opening circular, diameter 16.0-20.4
percent of L (mean 18.5).

Periproct: Inframarginal, located nearer to posterior mar¬
gin than to peristome, approximately two-thirds distance from
peristome to posterior margin; distance from posterior margin
to posterior edge of peristome equal to 10.0-13.0 percent of L
(mean 11.1).

Comparison with Other Species. —-No other species of
Echinocyamus is known from the Oligocene of North America.
Only one species is known from the Miocene, Echinocyamus
chipolanus Cooke, from the lower Miocene Chipola Formation
in Florida. Unfortunately, only one specimen is known of this
species, and it was broken after being figured (Cooke,
1959:32). Cooke’s figures (1959, pi. 9: figs. 1-3) show little
about the nature of the petals, but its peristome appears to be
much larger than that of E. wilsoni.

The much wider test of Echinocyamus wilsoni easily
distinguishes the new species from E. parvus Emmons and E.
bisexus Kier from the middle Eocene Castle Hayne Limestone
of North Carolina and Lake City Formation of Georgia,
respectively. It differs from E. caribbeanensis Kier from the
middle Eocene of Barbados in having a shorter anterior petal
(III) and a more anteriorly situated periproct. Echinocyamus
meridonalis Meyer from the middle Eocene of Alabama has a
much flatter, lower test and a more posterior periproct.
Echinocyamus huxleyanus Meyer, also from the middle Eocene

of Alabama, and E. macneili Cooke, from the late Eocene of
Alabama, have narrower tests.

Material. — Holotype: USNM 398476, from low in the
Belgrade Formation, Belgrade quarry, North Carolina.

Figured Paratypes: USNM 398477-398479, same data as
holotype.

Nonfigured Paratypes: USNM 492097-492100, same
data as holotype.

Occurrence. —North Carolina: The species is known
only from Belgrade quarry (lower bed).

Family Protoscutellidae Durham
Genus Periarchus Conrad

Periarchus lyelli (Conrad)

Plate 6: Figures 1,2
Scutella lyelli Conrad, 1834:152.

Periarchus lyelli (Conrad).—Kier, 1968:40, pi. 5: figs. 4, 5; 1980:40, figs. 17,

18; pi. 13, pi. 14: figs. 1-7.—Toulmin, 1969:474, pi. 4: figs. 5-7; 1977:344,

pi. 68: figs. 4-6.—Banks, 1978:109, 146, 149. [For a pre-1968 synonymy,

see Cooke, 1959:41,42.]

Description. —The dorsal surface of the two most complete
specimens (USNM 398326, 398327) show that they definitely
belong to this species.

Material. —USNM 398326 and 398327 are from float in
the New Bern quarry, Pete Harmatuk, collector. Jerry Baum
and Victor Zullo (pers. comm., 1984) state that the matrix on
USNM 398326 is middle Eocene, Castle Hayne Limestone
(which occurs at the bottom of the New Bern quarry), whereas
the matrix on USNM 398327 is late Eocene, New Bern
Limestone of Baum et al. (1978) or Spring Garden Member of
the Castle Hayne Limestone. In addition, there are 24
fragments of specimens.

Occurrence. —North Carolina: Middle Eocene, Castle
Hayne Limestone, New Bern quarry, and late Eocene, New
Bern Formation, New Bern quarry.

The species is known elsewhere from the middle and late
Eocene of Florida, Alabama, Georgia, South Carolina, Missis¬
sippi, and Louisiana.

Family Clypeasteridae Agassiz
Genus Clypeaster Lamarck

Clypeaster rogersi (Morton)

Plate 6: Figures 3,4

Scutella rogersi Morton, 1834:77, pi. 13: fig. 3. [For a complete synonymy see

Cooke, 1959:36.]

Description. —Although only three fragments are present,
one of them (USNM 398475) shows enough of the shape of the
test and part of the petals to make identification fairly certain
(Plate 6: Figure 4). This specimen is figured for comparison

NUMBER 83

FIGURE 4. — Echinocyamus wilsoni, new species: A, dorsal view of holotype USNM 398476 (x20); B, dorsal view
of paratype USNM 398477 (x20); C,D, dorsal and ventral views, respectively, of paratype USNM 398478 (x20);
E, ventral view of paratype USNM 398479 (x20). All specimens from low in the Belgrade Formation (Baum et
al., 1978) at the Belgrade quarry.

10

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

with a specimen of C. rogersi (Plate 6: Figure 3) from the
Marianna Limestone.

Material. —-USNM 398475, from float of Trent Formation,
New Bern quarry. USNM 372897, Oligocene, Marianna
Limestone, at Whitsett’s quarry, ~3 mi south of Collumburg,
Alabama.

OCCURRENCE. —North Carolina: Trent Formation, New
Bern quarry. Known elsewhere from the Oligocene Flint River
Formation, Marianna Limestone and Suwannee Limestone of
Mississippi, Alabama, and Florida.

Family Schizasteridae Lambert
Genus Agassizia Agassiz and Desor

Agassizia mossomi Cooke

Figure 5; Plate 7: Figures 1-8; Plate 8: Figures 1-8

Agassizia ( Anisaster ) mossomi Cooke, 1942:46, pi. 5: figs. 14-17; 1959:76, pi.

32: figs. 5-9.

DESCRIPTION. —Sixteen specimens from the New Bern
quarry and 20 specimens from the Belgrade quarry can be
referred to this species. Because the original description by
Cooke was based on only two poorly preserved specimens, the
New Bern specimens are described herein. They are generally
well preserved and not distorted, but the exterior of the tests are
weathered, which has enlarged the petaloid pores and obscurred
the fascioles.

Shape and Size: Length (L) 16.9-33.1 mm; width 91.8-
100+ percent of L (mean 98.4); height 75.4-94.1 percent of L
(mean 84.9). Greatest height anterior to apical system; greatest
width central.

Apical System: Anterior, located at distance from anterior
margin, equal to 22.0-39.8 percent of L (mean 33.6). Four
genital pores.

Ambulacra: Ambulacrum III in slight groove running from
apical system to lower anterior margin. Ambulacral pores small
throughout length of ambulacrum.

Anterior petals (II and IV) long, extending to margin, length
45.8-56.1 percent of L (mean 50.8); width 10.0-14.8 percent
of L (mean 12.3). Interporiferous zones narrower than single
poriferous zone; pores elongated transversely. Petals curving
convexly anteriorly. Anterior poriferous zone with adapical
pores greatly reduced in size. First petaloid porepairs in plate
11. Specimen 25 mm long with 50 petaloid porepairs, specimen
32 mm long with 54.

Posterior petals extending two-thirds distance from apical
system to margin; length 36.2-47.0 percent of L (mean 39.6);
width 11.3-14.8 percent of L (mean 12.7). Specimen 25 mm
long with 44 petaloid porepairs, specimen 32 mm long with 46.
Not possible to determine which plate bears first petaloid
porepair.

Peristome: Anterior, located at distance from anterior

A

C

FIGURE 5. —Agassizia mossomi Cooke: A,B,C, dorsal, right side, and rearviews,
respectively, of USNM 398328 from float of the Trent Formation, New Bern
quarry (x2).

NUMBER 83

11

margin equal to 21.5-28.7 percent of L (mean 24.7). Wider
than high, with width 20.0-28.5 percent of L (mean 22.5);
height 8.0-15.3 percent of L (mean 9.8).

Periproct: Located high on vertical posterior truncation;
width greater than height.

Fascioles: Peripetalous fasciole (Figure 5) indented deeply
into posterior interambulacra; passing around test low on
margin at great distance below end of anterior petals.
Lateroanal fasciole with deep indentation below periproct.

Oral-plate Arrangement: Plate sutures not clear, but
labrum appears to be very short.

REMARKS. —Twenty specimens were collected at the Bel¬
grade quarry, which appear to be conspecific with the
specimens of A. mossomi from New Bern. Their relative
dimensions overlap those of the New Bern specimens, and in
general appearance they are almost indistinguishable. The
petals in the Belgrade specimens are slightly shorter, with petal
II having a length of 37.9-47.8 percent of L (mean 44) in the
Belgrade specimens as opposed to 45.8-56.1 percent of L
(mean 50.8) in the New Bern specimens. Petal V has a length
in the Belgrade material of 28.8-36.2 percent of L (mean 33.2)
as opposed to 36.2-47.0 (mean 39.6) in the New Bern
specimens. The petals also are slightly narrower in the Belgrade
specimens. These differences appear to be too slight to warrant
specific or subspecific differentiation. Five of the Belgrade
specimens are illustrated on Plate 8: Figures 1-8.

Material. —USNM 398328, 398329 are from float of Trent
Formation, New Bern Quarry, North Carolina. The Belgrade
specimens (USNM 398330-398334) were collected by Druid
Wilson from the south face of the quarry toward the northeast
comer in a dark bed approximately 1-45 cm below the contact
with an indurated limestone. According to Wilson (pers.
comm., 1984), this horizon is high in zone C of the Belgrade
Formation of Baum et al. (1979, fig. 1).

OCCURRENCE. —North Carolina: Trent Formation, New
Bern quarry and zone C of the Belgrade Formation of Baum et
al. (1979, fig. 1), Belgrade quarry.

Florida: Cooke’s type specimens, Oligocene Suwannee
Limestone, Florida Rock Products quarry, 3 A mi SW of
Brooksville, Florida.

Agassizia sp.

Plate 9: Figures 1-7

Description. —Two specimens (one partially cmshed) with
exterior well preserved, showing details of ornamentation.

Shape and Size: One specimen (USNM 398335), with
original shape preserved, having length (L) 14.4 mm, height
11.4 mm (height 79.2 percent of L), and width 12.7 mm (width
88.2 percent of L). Greatest height at apical system.

Apical System: Four genital pores, ethmolytic, located

posterior to center at distance from anterior margin of 65.2
percent of L.

Ambulacra: Anterior ambulacrum III not petaloid, adapi-
cally in very faint groove not continuing to margin. Anterior
petals (II and IV) long, length 37.5 percent of L; petal
depressed in slight groove, petal narrow, nearly straight.
Porepairs of anterior poriferous zone greatly reduced in size; in
one specimen two enlarged porepairs at end of one petal, four
in other anterior petal; in other specimen one or two enlarged
porepairs in each anterior zone; 16 enlarged petaloid porepairs
in posterior poriferous zone of anterior petal in specimen 14.4
mm long; 14 in specimen 13.3 mm long.

Posterior petals (V and I) short with length 24.3 percent of L.
Petals wide, width 8 percent of L, in shallow groove; 23
petaloid porpairs in petal in specimen 14.4 mm long, 20 in
specimen 13.4 mm long. Interporiferous zone narrow, one-half
width of single poriferous zone.

Peristome: Anterior, located at distance from anterior
margin to anterior edge of peristome equal to 21.5 percent of L;
opening wider than high, width 21.2 percent of L, height 11.9
percent of L.

Periproct: Located high on posterior truncation; opening
wider than high, width 23.3 percent of L, height 7 percent of L.

Fascioles: Peripetalous fasciole narrow, curving sharply
into interambulacra 4 and 1, straight across interambulacrum 5;
from ends of anterior petals fasciole curves sharply adorally
passing around anterior of test below margin. Lateroanal
fasciole passing in deep lobe below periproct crossing
ambulacra V or I. Not possible to determine specific plate
crossed by fasciole.

REMARKS. —These specimens resemble Agassizia wilming-
tonica Cooke from the middle Eocene Castle Hayne Formation
of North Carolina. They differ in having a higher peristome,
and a test that is wider posteriorly with its posterior margin less
pointed, and a more vertical, less overhanging, posterior
truncation. With only two small specimens (one slightly
crushed), it is not possible to know the significance of these
differences.

Material. —USNM 398335, 398336, from float of the
Trent Formation, Pollocksville state quarry.

Occurrence. —North Carolina: Trent Formation, Pol¬
locksville state quarry.

Family Spatangidae Gray
Genus Maretia Gray

Maretia carolinensis, new species

Figure 6; Plate 9: Figures 8, 9; Plate 10: Figures 1-7;

Plate 11: Figures l, 2

DIAGNOSIS.—The species is characterized by a wide test
with width equal to length and by wide petals.

12

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Description. —Twenty-four specimens (most fragments)
are referred to this species. All specimens are badly weathered,
but one shows a fasciole.

Shape and Size: Five specimens show the dimensions of
the test. Length (L) 27.0-52.2 mm (mean 37.7); width
approximately same as length; width 99-103 percent of L
(mean 101); greatest width central to slightly anterior; height
39_44 percent of L (mean 42.7) with the greatest height near
posterior margin. Adorally, test depressed around peristome.

Apical System: Located anterior at distance from anterior
margin equal to 40-45 percent of L (mean 44.9). Four genital
pores, ethymolytic with genital 2 extending posteriorly (Figure
6a).

Ambulacra: Anterior ambulacrum III not petaloid, in
groove extending from apical system to peristome; pores
minute in slight peripodia. Anterior petals long, wide; length
36.6-42.6 percent of L (mean 40.1); width 11-13 percent of L
(mean 12.6). Pores deeply conjugate; adapically (approxi¬
mately one-half length of petal) pores in anterior poriferous
zones greatly reduced in size; petals closing distally with 50
petaloid porepairs in specimen 52.2 mm long, 50 in specimen
47.5 mm long, 44 in specimen 36 mm long, and 42 in specimen
27 mm long. Not possible to determine which plate bears first
petaloid porepair.

Posterior petals with length 35.6-43.6 percent of L (mean
39.3), width 11.9-14.8 percent of L (mean 13.2); 50 porepairs
in specimen 52.2 mm long, 50 in specimen 47.5 mm long, 44
in specimen 36 mm long, and 42 in specimen 27 mm long.

Peristome: Anterior, located at distance from anterior
margin equal to 33 percent of L (measurable on one specimen).
Labiate, wider than high with width (measurable on two
specimens) 19 percent of L and height 8 percent of L
(measurable on one specimen).

Periproct: Situated high on vertical posterior truncation
(Plate 10: Figure 3), opening with height equal to width, height
12.2-12.5 percent of L (mean 12.4). Occurring in plates 5-9.

Fascioles: No peripetalous fasciole. Subanal fasciole very
broad, width of area circumscribed by fasciole 51 percent of L,
height approximately 15 percent of L. Not possible to
determine on which plates fasciole occurs.

Tuberculation: Large tubercles on dorsal surface in all
interambulaca except posterior interambulacrum. Tubercles in
deep pits. Ventrally, large tubercles in ambulacra and interam¬
bulacra except posterior paired ambulacra and posterior
interambulacrum where tuberculation absent anteriorly from
peristome to midway to posterior margin, tubercles small from
midway to posterior margin.

Oral-plate Arrangement: Labrum long and narrow (Figure
6b; Plate 9: Figure 9) with length 16.1-20.3 percent of L (mean
17.7), extending posteriorly to or just posterior to suture
between second and third adjacent ambulacral plates. Sternal
plates long and narrow, length on single measurable specimen
34 percent of L, width of paired plates equals 20 percent of L.
Comparison with Other Species. —This species is easily

A

B

FIGURE 6. —Maretia carolinensis, new species: A, apical system of paratype
USNM 398339 (x20); B, labrum and plastron of paratype USNM 398337 (xl).
Both specimens from float of the Trent Formation at Pollocksville state quarry.

distinguished from the other two species of Maretia known
from the United States. It differs from Maretia subrostrata
(Clark) from the middle Eocene Castle Hayne Limestone of
North Carolina in its wider test, with width 99-103 percent of
L versus 89 percent of L in M. subrostrata. Its petals are much
wider, with the width of the anterior petals 13 percent of L and
the width of the posterior petals 15-18 percent of L versus 10
and 11 percent, respectively, in the Eocene species. Finally, in
M. carolinensis the greatest width of the test is central whereas
in M. subrostrata it is anterior.

Maretia carolinensis differs for the above same reasons from
M. arguta (Clark) from the Winiona Sand of Lower Lisbon age
of Mississippi. As noted by Cooke (1959:81) and Kier
(1980:50), M. arguta may be conspecific with M. subrostrata.

Material. — Holotype: USNM 398338, from float of the
Trent Formation, Pollocksville state quarry.

NUMBER 83

13

Figured Paratypes: USNM 398337, 398339-398341,
same data as holotype.

Nonfigured Paratypes: USNM 492101-492124, same
data as holotype.

OCCURRENCE. —North Carolina: Trent Formation, Pol-
locksville state quarry.

Maretia sp.

Plate 11: Figures 3-5

DESCRIPTION. —One specimen and several fragments can be
referred to this genus. The specimens are too fragmentary to

permit specific identification. They are easily distinguished
from Maretia carolinensis, new species from Pollocksville by
their much narrower petals and narrower test. They resemble
Maretia subrostrata (Clark) from the middle Eocene Castle
Hayne Formation from North Carolina, but they appear to
differ in having more deeply sunken dorsal tubercles. Until
more specimens are found, it is not possible to determine
whether these specimens are M. subrostrata.

Material. —USNM 398472, 398473, from float of Trent
Formation at New Bern quarry.

OCCURRENCE. —North Carolina: Trent Formation. New
Bern Quarry.

Literature Cited

Banks, R.

1978. Stratigraphy of the Eocene Santee Limestone in Three Quarries of
the Coastal Plain of South Carolina. Geologic Notes, 21 (3):88—149,
figures 1-15, tables 1-3, appendix 1-7.

Baum, G., W.B. Harris, and V. Zullo

1978. Stratigraphic Revision of the Exposed Middle Eocene to Lower
Miocene Formations of North Carolina. Southeastern Geology,
20(1): 1-25, figures 1-5.

1979. Structural and Stratigraphic Framework for the Coastal Plain of
North Carolina. In Gerald R. Baum, W. Burleigh Harris, and Victor
A. Zullo, editors, Carolina Geological Society Field Trip Guide¬
book, 119 pages. Raleigh, North Carolina: Geological Survey
Section, Department of Natural Resources and Community Develop¬
ment.

Bouve, T.T.

1846. Pygorhynchus gouldii, a New Echinus from the Millstone Grit of
Georgia. Proceedings of the Boston Society of Natural History,
2:192.

Clark, W.B.

1915. In W.B. Clark and M.W. Twitched, The Mesozoic and Cenozoic
Echinodermata of the United States. Monographs of the United
States Geological Survey, 54: 341 pages, 107 plates.

Conrad, T.A.

1834. Observations on the Tertiary and More Recent Formations of a
Portion of the Southern States. Journal of the Academy of Natural
Sciences of Philadelphia, series 1, 7:116-154.

Cooke, C.W.

1941. Cenozoic Regular Echinoids of Eastern United States. Journal of
Paleontology, 15(1): 1-20, plates 1-4.

1942. Cenozoic Irregular Echinoids of Eastern United States. Journal of
Paleontology, 16(1): 1-62, plates 1-8.

1948. Arbia and Dixieus, Two New Genera of Echinoids. Journal of
Paleontology, 22(5):606-607.

1959. Cenozoic Echinoids of Eastern United States. United States
Geological Survey Professional Paper, 321: 106 pages, 43 plates.

Gladfelter, W.B.

1978. General Ecology of the Cassiduloid Urchin Cassidulus caribbearum.
Journal of Marine Biology, 47:149-160.

Kier, P.M.

1968. Echinoids from the Middle Eocene Lake City Formation of Georgia.
Smithsonian Miscellaneous Collections, 153(2): 45 pages, 44
figures, 10 plates.

1975. The Echinoids of Carrie Bow Cay, Belize. Smithsonian Contribu¬
tions to Zoology, 206: 45 pages, figures 1-8, plates 1-12.

1980. The Echinoids of the Middle Eocene Warley Hill Formation, Santee

Limestone, and Castle Hayne Limestone of North and South
Carolina. Smithsonian Contributions to Paleobiology, 39: 102 pages,
26 figures, 22 plates.

1984. Fossil Spatangoid Echinoids of Cuba. Smithsonian Contributions to
Paleobiology, 55: 336 pages, 45 figures, 90 plates, 6 tables.

Kier, P.M., and R.E. Grant

1965. Echinoid Distribution and Habits, Key Largo Coral Reef Preserve,
Florida. Smithsonian Miscellaneous Collections, 149(6): 1 —68, fig¬
ures 1-15, plates 1-16.

McKinney, M.L., K.J. McNamara, B.D. Carter, and S.K. Donovan

1992. Evolution of Paleogene Echinoids: A Global and Regional View. In
D.R. Prothero and W.A. Berggren, editors, Eocene-Oligocene
Climatic and Biotic Evolution, pages 349-367. Princeton, New
Jersey: Princeton University Press.

Mooi, Rich

1990. A New “Living Fossil” Echinoid (Echinodermata) and the Ecology
and Paleobiology of Caribbean Cassiduloids. Bulletin of Marine
Science, 46(3):688-700, 6 figures.

Mortensen, T.

1948. Clypeastroida. In T. Mortensen, A Monograph of the Echinoidea,
4(2): 471 pages, 257 figures, 72 plates. Copenhagen: C.A. Reitzel.

1951. Spatangoida. In T. Mortensen, A Monograph of the Echinoidea,
5(2): 593 pages, 286 figures, 64 plates. Copenhagen: C.A. Rietzel.

Morton, S.G.

1834. Synopsis of the Organic Remains of the Cretaceous Group of the
United States. 88 pages, 19 plates. Philadelphia. [Reprinted from
American Journal of Science, volumes 17, 18 (1830)].

Toulmin, L.D.

1969. Paleocene and Eocene Guide Fossils of the Eastern Gulf Coast
Region. Transactions of the Gulf Coast Association of Geological
Societies, 19:465-487, figures 1-5, tables 1, 2, plates 1-4.

1977. Stratigraphic Distribution of Paleocene and Eocene Fossils in the
Eastern Gulf Coast Region. Geological Survey of Alabama,
monograph 13:2 volumes. [Volume 1 consists of x + 602 pages, 72
leaves of plates; volume 2 consists of 20 maps and tables inserted in
two pockets.]

Ward, L.W., D.R. Lawrence, and B.W. Blackwelder

1978. Stratigraphic Revision of the Middle Eocene, Oligocene, and Lower
Miocene—Atlantic Coastal Plain of North Carolina. Bulletin of the
United States Geological Survey, Contributions to Stratigraphy,
1457-F:F1-F23, figures 1-3.

Zullo, V.A., and G.R. Baum

1979. Paleogene Barnacles from the Coastal Plain of North Carolina
(Cirripedia, Thoracica). Southeastern Geology, 20(4):229-246,
figures 1, 2, plates 1-4.

14

Plates

16

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 1

Dixieus dixie (Cooke)

1, 2, 3, 4. Dorsal, ventral, and side views, respectively, of USNM 398317 from Trent
Formation float at New Bern quarry (x2).

Arbi a aldrichi (Clark)

5. Dorsal view of USNM 398318, from the Belgrade Formation (Baum et al.,
1978) at Belgrade quarry, 45 cm below top of indurated limestone (x2).

6. Ventral portion of test of USNM 398319, from the Belgrade Formation
(Baum et al., 1978) at Belgrade quarry, near contact with indurated
limestone (x5).

18

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 2

Gagaria mossomi (Cooke)

1, 2, 3, 4. Dorsal, ventral, and side views, respectively, of USNM 398320, from the
Belgrade Formation (Baum et al., 1978) at Belgrade quarry, approximately
30 cm below indurated limestone (x4).

5. Ambulacrum of the same specimen (x8).

NUMBER 83

19

20

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 3

Psammechinus carolinensis, new species

1, 2. Dorsal and ventral views, respectively, of the holotype, USNM 398321,

from float of the Trent Formation, Pollocksville state quarry (x8).
Additional views of this specimen are on Plate 4: Figures 1-4.

3, 4. Interambulacral and ambulacral views of paratype USNM 398322 from the

same locality as the holotype (x 15).

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SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 4

Psammechinus carolinensis, new species

Side views of the holotype, USNM 398321, from float of the Trent
Formation, Pollocksville state quarry (x8). Additional views of this
specimen are on Plate 3: Figures 1, 2.

Ambulacral and interambulacral views of USNM 398474 (xl5).

Auricles in USNM 398323 from same locality as holotype (xlO).

NUMBER 83

24

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 5

Rhyncholampas gouldii newbernensis, new subspecies

1, 2, 3, 4. Dorsal, ventral, rear, and right side views, respectively, of the holotype,
USNM 398324, from float of the Trent Formation, New Bern quarry (x2).
5, 6, 7. Dorsal, ventral, and right side views, respectively, of paratype USNM

398325 from the same locality as the holotype (x2).

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NUMBER 83

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SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 6

Periarchus lyelli (Conrad)

Dorsal view of USNM 398326 (x 1). Specimen collected by Pete Harmatuk
from float in the New Bern quarry.

Dorsal view of USNM 398327 from float in the New Bern quarry (x2).
Clypeaster rogersi (Morton)

Dorsal view of USNM 372897 from Oligocene, Marianna Limestone, at
Whitsett’s quarry, about 3 mi south of Collumburg, Alabama (xl.5). This
photograph is included for comparison with a specimen from the New Bern
quarry figured on this plate, Figure 4 (x 1.5).

Dorsal view of fragment (USNM 398475) from float from the Trent
Formation at New Bern quarry (xl.5).

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NUMBER 83

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28

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 7

Agassizia mossomi Cooke

1, 2, 3, 4. Dorsal, ventral, rear, and right side views, respectively, of USNM 398328
from float of Trent Formation, New Bern quarry (x2).

5, 6, 7, 8. Dorsal, ventral, rear, and right side views, respectively, of USNM 398329
from float of Trent Formation, New Bern quarry (x2).

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30

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 8

Agassizia mossomi Cooke

1. Dorsal view of USNM 398330 from the Belgrade Formation (Baum et al.,
1978), 45 cm below indurated limestone (x2).

2. Dorsal view of USNM 398331 from same locality (x2).

3. Dorsal view of USNM 398332 from same locality (x2).

4. Ventral view of USNM 398333 from same locality (x2).

5. 6, 7, 8. Dorsal, rear, right side, and ventral views, respectively, of USNM 398334

from the Belgrade Formation (Baum et al., 1978), near contact with
indurated limestone (x2).

32

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 9

Agassizia sp.

1, 2, 3, 4. Dorsal, rear, right side, and ventral views, respectively, of USNM 398335
from float of Trent Formation, Pollocksville state quarry (x4).

5, 6, 7. Dorsal, right side, and ventral views, respectively, of USNM 398336 from

same locality (x4).

Maretia carolinensis, new species

8, 9. Dorsal and ventral views, respectively, of paratype USNM 398337 from

float of the Trent Formation, Pollocksville state quarry (x 1.5).

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NUMBER 83

34

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 10

Maretia carolinensis, new species

1, 2, 3, 4. Dorsal, ventral, rear, and right side views, respectively, of the holotype,
USNM 398338, from float from the Trent Formation at Pollocksville state
quarry (xl.5).

5, 6, 7. Dorsal, right side, and ventral views, respectively, of paratype USNM

398339 from same locality as holotype (x2).

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NUMBER 83

SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

Plate 11

Maretia carolinensis, new species

Dorsal view of paratype USNM 398340 from float of the Trent Formation
at Pollocksville state quarry (xl.5).

Dorsal view of paratype USNM 398341 from same locality (xl.5).
Maretia sp.

Dorsal and ventral views, respectively, of USNM 398472 from float from
the Trent Formation at the New Bern quarry (x3).

Dorsal view of USNM 398473 from the same locality (x3).

NUMBER 83

37

Tr U.S GOVERNMENT PRINTING OFFICE: 1997 405-261/20020

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