Climatic Implication of a 

Late Pleistocene Ostracode 

Assemblage from Southeastern

Virginia

By PAGE C. VALENTINE

CONTRIBUTIONS TO PALEONTOLOGY

GEOLOGICAL SURVEY PROFESSIONAL PAPER 683-D

Determination of the inner sublittoral marine 

climate represented by the ostracode assemblage 

of the Norfolk Formation of southeastern 

Virginia^ by a comparison of fossil and living 

ostracode assemblages

o

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1971
UNITED STATES DEPARTMENT OF THE INTERIOR 

ROGERS C. B. MORTON, Secretary

GEOLOGICAL SURVEY 

W. A. Radlinski, Acting Director

Library of Congress catalog-card No. 75-176115

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

Washington, D.C. 20402 - Price 70 cents (paper cover)

Stock Number 2401-1133
CONTENTS

Abstract.._________________________________________

Introduction. ______________________________________

Acknowledgments, ______________________________

Previous studies.___________________________________

Pleistocene collections.______________________________

Pleistocene stratigraphy and age of the Norfolk Formation 

as used by Oaks and Coch (1963)________________

Holocene collections_______________________________

Holocene species distributions._______________________

Faunal provinces and climatic zones of the North 

Atlantic Ocean_______________________________

Latitudinal ranges of Holocene species_____________

Holocene shelf assemblages defined by cluster 

analysis.____________________________________

Page

Dl

1

1

1

2

2

13

14

14

14

15

Holocene species distributions Continued

Biofacies of the shelf-____________________________ D15

Faunal provinces in the Cape Hatteras area______ 20

Comparison of late Pleistocene and Holocene assemblages. 21

Late Pleistocene marine climate of southeastern Virginia, _ 21

Holocene marine bottom temperatures.____________ 21

Late Pleistocene bottom temperatures.____________ 22

Late Pleistocene sea-surface temperatures-_________ 24

Late Pleistocene faunal province and climatic zone of

southeastern Virginia._____________________________ 24

Summary. _____________-__----_-______----------___ 25

Pleistocene localities. ________________________________ 25

References cited.______________-_______--------_____ 2fi

ILLUSTRATIONS

[Plates follow references]

PLATE 1. Bensonocythere, Cushmanidea, Campylocythere, Actinocythereis, and Aurila.

2. Cytherura, "Sahnia," Megacythere, Eucythere, Puriana, Pellucistoma, Haplocytheridea, Protocytheretta, and Pseudocytheretta.

3. Hulingsina, Finmarchinella, Paracytheridea, MueUerina, Cytheropteron, and Neocaudites.

4. Microcytherura, Paradoxostoma, Cytheromorpha, Leptocythere, Sclerochilus, Neolophocythere, Loxoconcha, Propontocypris, 

Proteoconcha, and Hemicythere.

Page 

FIGUKE 1. Map of southeastern Virginia indicating important geomorphologic features and localities sampled-___________ DIO

2. Measured sections at late Pleistocene localities PI, P2, and P6 in southeastern Virginia.

3. Geologic section across southeastern Virginia from the Suffolk Scarp to the Atlantic coast. ________________

4. Map showing locations of Holocene sample stations ____________________--_------------------------------

5. Chart showing modern latitudinal ranges along the middle Atlantic Coast of the United States of 68 ostracode species__

6. Dendrogram of Holocene sample clusters._____________________-_-_______---_--------------------------

7. Similarity matrix showing relation between clusters of samples, and between the Norfolk cluster and an artificial 

sample grouping.-___________________________________________-------_---------------------------

8. Map showing the geographic extent of four ostracode biofacies_________________--------------------------

9. Bottom-temperature map for the shelf off the middle Atlantic Coast of the United States for February _____

10. Bottom-temperature map for the shelf off the middle Atlantic Coast of the United States for August-September.

11. Chart showing temperature ranges of 66 ostracode species common to the Holocene and the Norfolk Formation 

(upper Pleistocene)_________________________________________-_-_-----_---------------------------

11

12

13

16

17

18

18

21

22

23

TABLES

TABLE 1. Occurrences of 171 ostracode species in Holocene samples from the shelf off the middle Atlantic Coast of the 

United States and in Pleistocene samples from the Norfolk Formation as used by Oaks and Coch (1963) of 

southeastern Virginia.________________________________________--------_--_-----------------------

2. Principal Holocene species of shelf biofacies; their occurrence in the Norfolk Formation (upper Pleistocene); 

their occurrence north and south of Cape Hatteras area._______________-___-------------------------

in

Page

CONTRIBUTIONS TO PALEONTOLOGY

CLIMATIC IMPLICATIONS OF A LATE PLEISTOCENE 

OSTRACODE ASSEMBLAGE FROM SOUTHEASTERN

VIRGINIA'

By PAGE C. VALENTINE

ABSTRACT

The Norfolk Formation, as used by Oaks and Coch (1963), 

(upper Pleistocene (Sangamon)) of southeastern Virginia contains a large and well-preserved ostracode assemblage 82 

species, 70 of which are still living.

A comparison of the Norfolk ostracode assemblage with Holocene assemblages collected on the continental shelf off the 

middle Atlantic Coast of the United States indicates that no 

modern shelf assemblage is closely similar to that of the Norfolk Formation (which contains some forms that today are 

found only north or only south of Cape Hatteras). A cluster 

analysis of 91 samples containing 159 ostracode species from the 

continental shelf between about 32°30'N. and 40°N. reveals the 

existence of four major biofacies representing two faunal provinces. A study of the temperature tolerances of the species 

common to the Norfolk Formation and the Holocene indicates 

that the Norfolk assemblage lived in waters where the bottom 

temperature in the coldest month was 12.5°-15°C and in the 

warmest month 20°-22.5°C, a more equable marine climate than 

now prevails in the inner sublittoral off southeastern Virginia 

where minimum and maximum average monthly temperatures 

range from 5°-7.5° C to 22,5°-25° C, respectively. The Late Pleistocene assemblage is indicative of a sublittoral faunal province 

and a warm-temperate climatic zone which do not exist today 

along the Atlantic Coast of the United States. At present, the 

convergence of cool- and warm-water masses at Cape Hatteras 

prevents reestablishment of a warm-temperature faunal province in this region.

Seventy-seven of the species constituting the latest Pleistocene 

assemblage are illustrated.

INTRODUCTION

The purpose of this study is to determine the marine 

climate represented by the ostracode assemblage of the 

Norfolk Formation as used by Oaks and Coch (1963) 

(upper Pleistocene) of southeastern Virginia. The Nor1 Contribution 2531 of the Woods Hole Oceanographic Institution, 

based on work done under a program conducted jointly by the U.S. 

Geological Survey and the Woods Hole Oceanographic Institution and 

financed by the U.S. Geological Survey.

folk ostracode assemblage is large and well preserved, 

and most species are still living in the sublittoral off the 

Atlantic Coast of the United States. The distributional 

ranges, temperature tolerances, and associations of these 

living species are used to infer marine climatic conditions under which the Pleistocene assemblage of southeastern Virginia lived.

ACKNOWLEDGMENTS

I am sincerely grateful to Joseph E. Hazel of the U.S. 

Geological Survey who proposed the problem, guided 

and encouraged my efforts and critically read the manuscript, and also generously provided me with working 

space and allowed me to use his equipment. I appreciate 

the advice and assistance of A. G. Coates of George 

Washington University who reviewed the manuscript, 

and I thank T. E. Waller of the U.S. National Museum 

who provided me with many samples from the Pleistocene of southeastern Virginia. I also thank W. O. Addicott and J. P. Owens of the U.S. Geological Survey 

for critically reading the manuscript. I am grateful to 

Lanci Valentine who assisted with the original illustrations and typed the manuscript.

PREVIOUS STUDIES

Most descriptive and distributional studies of ostracodes of the Atlantic continental shelf of North America 

have been restricted to the region from Long Island, 

N.Y., northward to the Arctic (see Hazel, 1970) ; only 

a few investigations of marine ostracodes occurring 

south of this region have been undertaken. Hulings 

(1966, 1967) described four new species and provided 

information on ostracode distribution for the shelf off 

Virginia and for the shelf and slope from Cape Hatteras, N.C., southward to Jupiter Inlet, Fla. Hazel and

Dl
D2 CONTRIBUTIONS TO PALEONTOLOGY

Valentine (1969) described two new species occurring 

south of Long Island along the shelf break. Grossman 

(1967) studied the ecology of the ostracodes of Pamlico 

Sound, N.C., and described two new species. Benson 

(1967) listed 36 ostracode species from stations on a 

transect extending from Cape Lookout, N.C., to the 

shelf edge.

Hazel (1968) compared Holocene ostracode assemblages to those found in Pleistocene sediments dredged 

from submarine canyons off the eastern United States. 

Distributional ranges and temperature tolerances of 

Holocene species also found as fossils in the canyon 

sediments indicate that a climatic change and a corresponding northern shift of cold-temperate assemblages have occurred since the glacial-age Pleistocene 

sediments were deposited. To date there are no published studies of ostracode assemblages from Pleistocene sediments of the Atlantic Coastal Plain of the 

United States.

The following investigations of Holocene and fossil 

ostracode assemblages from the eastern and southern 

United States, the Caribbean Sea, the western and 

northern Atlantic Ocean, and northern Europe were 

used in identifying the species found during this study: 

Baird (1850); Benda and Puri (1962); Benson and 

Coleman (1963); Blake (1929, 1933); Bold (1946, 

1963); Brady (1868a,b 1869,1870,1872) ; Brady, Crosskey, and Eobertson (1874) ; Brady and Norman (1889); 

Cushman (1906); Darby (1965); Edwards (1944); 

Grossman (1967); Hall (1965) ; Hazel (1967a,b); 

Hazel and Valentine (1969); Howe and others (1935) ; 

Hulings (1966, 1967); Hulings and Puri (1964) ; King 

and Kornicker (1970); Haddocks (1969); Malkin 

(1953) ; McLean (1957, 1966) ; Morales (1966) ; Morkhoven (1963) ; Norman (1865); Plusquellec and Sandberg (1969); Pooser (1965); Puri (1952, 1953, 

1958a,b, 1960) ; Puri and Hulings (1957) ; Sandberg 

(1964, 1969) ; Sars (1865) ; Stephenson (1938) ; Swain 

(1951, 1955); Triebel (1960); Ulrich and Bassler 

(1904); and Williams (1966).

PLEISTOCENE COLLECTIONS

Pleistocene samples for this study were collected from 

blue-gray clayey and silty fine fossiliferous marine 

sands exposed in borrow pits in the Norfolk, Va., area 

(see figs. 1, 2, and Pleistocene locality data). Nine 

samples were processed from 6 localities, and 82 species 

were identified, of which only 42 have been formally 

described. Seventy of these species are represented in 

the Holocene. Ostracode carapaces and valves containing remnants of soft parts were found in all samples. 

The geographic location and species content of all samples are presented in table 1. Seventy-seven of the 82 

Pleistocene species are illustrated on plates 1-4. Illustrated specimens have been deposited in the collections 

of the U.S. National Museum.

The borrow pits were opened to provide road fill 

during construction of interstate highways in the area1. 

The pits from which samples were collected are now 

flooded, with the exception of locality P6 which was 

being worked in June 1970. Other pits in the area are 

open and being worked but are still too shallow to offer 

good exposures of fossiliferous Pleistocene sediments.

PLEISTOCENE STRATIGRAPHY AND AGE OF THE 

NORFOLK FORMATION AS USED BY OAKS AND 

COCH (1963)

The coastal plain of Virginia is characterized by a 

series of gently eastward-sloping plains delineated by 

low scarps which trend generally north and south. Eastward from the Fall Line three major plains are separated by the Surry Scarp, the Suffolk Scarp, and the 

present coast. The region is underlain by sediments of 

Cretaceous to Holocene age resting on older consolidated rocks.

The stratigraphy of the post-Miocene units of the 

Virginia coastal plain has been studied since 1839 when 

W. B. Kogers described post-Miocene sediments extending from the Fall Line to the sea. Oaks (1964) presented a comprehensive review of many conflicting interpretations of the origins of these sediments. The alternating plains and scarps have been described as forming 

during successive stages of shoreline retreat from the 

Fall Line, or as a result of headland erosion and sediment deposition during repeated transgressions by the 

sea, a view which maintains that each plain is underlain 

by a single formation deposited on sediments of late 

Miocene age or older. A third interpretation is that each 

major plain is underlain by several formations which is 

indicative of deposition during more than one episode 

of sea-level fluctuation. Transgressions and regressions 

of the sea have been ascribed both to glacioeustatic control and to tectonically-controlled movements of the 

land.

The latest and most detailed studies of the stratigraphy of the coastal plain were conducted by R. Q. 

Oaks, N. K. Coch, and K. F. Bick (Oaks and Coch, 

1963; Oaks, 1964; Coch, 1965, 1968; Bick and Coch, 

1969) in southeastern Virginia in an area extending 

from the Fall Line to the present coast. Their studies, 

which support some earlier findings and also present 

new information, indicate that: (1) the major scarps 

and plains of the region are the result of a complex series of sea-level fluctuations; (2) more than one 

formation underlies each plain; (3) some formations
TABLE 1. Occurrences of 171 ostracode species in Holocene samples from the shelf off the middle Atlantic Coast of the United States an

Norfolk Formation, as used by Oaks and Coch (1963), of southeastern Virginia 

[Sample sources Gos and Ast = current studies of the continental shelf by the U.S. Geol. Survey and the Woods Hole Oceanographic Institution. Locality numbers PI through 

Sample

Locality

1

3

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

47 

48

Source

Gos 1378------

Gos 1377... .---

Gos 1316------

Gos 1319.. 

Gosl322.- -

Gos 1346 --

Gosl345_ ....

Gos 1340-..---

Ast 1965. ------

Gos 1351. 

Qosl338- 

Gos 2026 - 

Qosl354 .

Gosl355- 

Ast 2027-... -----

Gos 1415-- --

GOS2032- -

Gos 1417-.-. -----

Gos 2036--- 

n no 1 ftQ1

Ast 2001.-.. -----

Gos 2041. .

Gos 1420 ------

Gos 2045-----

Gos 1886-- 

Gos 1885....... -

Gos2047.______-.

Gos 1422--.---

Gosl883---..----

A of 1 QQ7

Gos 2048--.--- .

Gos 1423-.... ....

Gos2051._._.....

n no 1 J.9J.

Gos 1880-... -_-.

Gos 2057------

Gos2052. .

Gosl425- 

Gos2054 . -

Gos2053.. ....

Gos 1426-- 

Location

Latitude 

north

deg. min.

39 38 

39 31 

39 21 

39 20 

39 20 

39 09 

39 10 

39 10 

39 00 

39 01 

39 01 

38 50 

38 51 

38 49 

38 40 

38 40 

38 40 

38 30 

38 30 

38 31 

38 20 

38 10 

38 00 

37 59 

38 00 

37 49 

37 50 

37 40 

37 39 

37 31 

37 31 

37 27 

37 19 

37 20 

37 20 

37 10 

37 09 

37 05 

37 00 

37 00 

37 02 

36 57 

36 49 

36 50 

36 45 

36 39 

36 40 

36 39

Longitude 

west

deg. min.

73 59 

73 45 

74 14 

73 31 

72 46 

74 31 

74 15 

73 15 

74 45 

74 00 

73 01 

74 45 

73 32 

73 15 

74 45 

74 14 

73 30 

74 45 

74 00 

73 26 

74 30 

74 15 

75 00 

74 29 

73 59 

75 25 

74 45 

75 14 

74 42 

75 30 

74 30 

74 29 

75 16 

75 00 

74 46 

75 44 

75 15 

75 01 

75 30 

75 15 

75 00 

74 45 

75 45 

75 15 

74 59 

75 31 

75 15 

74 58

Depth 

(meters)

24 

28 

20 

48 

84 

23 

21 

62 

15 

38 

80 

18 

62 

81 

26 

39 

69 

29 

57 

89 

36 

41 

18 

49 

129 

12 

41 

24 

49 

16 

64 

406 

29 

38 

56 

12 

30 

38 

30 

36 

44 

82 

22 

29 

36 

20 

35 

32

5

X

|

s

1

1

1

X

X

X

X

X

X

X 

X

X 

X

X

X

--

X

aft. A. gomittionensis (Howe and Ellis, 1935) \

*

X

--

X

--

X

X

X

X

vineyardensis (Cushman, 1906)

^

X

X

X

X

X

--

--

ft

3 

1

..

--

§

--

X

--

-

ft

^

--

--

-

o

e

^

--

..

--

Q 

ft

^

--

-

irila floridana Benson andColeman, 1963

X

X 

X

X

X

X

laevicula (Edwards, 1944)

^

--

-

iffinicythere emarginata (Sars, 1865)

PQ

X

X

X

sslerttes miocenicus (Howe, 1935)|

BQ

--

ft

BQ

--

--

1

1

(3 

J

I*.

i

*

X

v

X

X

X

X

arenkola (Cushman, 1906)\

0)

X

X 

X

X

X

s

e 

2

a

"o

0)

X

X

"

whitei (Swain, 1951) _ \

*

X

X

X

X

X 

X

X

X 

X

X

X

X

ft

«)

X

X

X

v

X

X

v

v

X 

X

X 

X

X 

X 

X 

X

X

X

« 

ft

«)

X

X

X

X

X

X

v

X

X

X

X

X

X 

X

X 

X

X 

X 

X

X

X

X

o 

ft

«q

X

X

X

X

X

X

X

X 

X 

X 

X

X

Q 

ft

ttl

X

X

X

X

X

X

X

X

w 

ft

0)

X

X

X

X

1

i

1,1

1

aq

X

X

X

o

ft

U)

-

--

/thocypris sp. A

0)

/thocythere sp. A

si1

X

W 

ft

0)

mpylccythere laeva Edwards, 1944

O

X

X

--

udites nipeensis van denBold, 1946

C

1

W(3

£

.a

s 

e

e

X

X

X 

X 

X 

X

X

seminuda (Cushman, 1906)

O

X

X

X

X

X

X

X

X 

X 

X 

X 

X

X

ft

o

X

X

X 

X 

X 

X 

X

ft

o

--

o 

ft

o

--

Q

e

fc

--

Cyprideis sp. A

--

therella spp.

Q1

therelloidea sp. A

-

itheridena sp. A

C'

X

ithercis sp. A

tf

--

--

Itheromorpha aft. C. surta Edwards, 1944

0'

-

--

C. warneri newportensis Williams, 1966

X

X

O
TABLE 1. Occurrences of 171 ostracode species in Holocene samples from the shelf off the middle Atlantic Coast of the United States and 

Norfolk Formation, as used by Oaks and Coch (1968), of southeastern Virginia Continued

Sample

Locality

49 

50 

61 

62 

53 

54 

65 

66 

57 

58 

69 

60 

61 

62 

63 

64 

65 

66 

67 

68 

69 

70 

71 

72

Source

Gosl878------._-

Gos 2060.. .......

Gos 2062.. .......

Gos 1428.----.-..

Gos 2063.. _-_._..

Gos 1430.. .......

Gos 14,31--------

Gos 1432.. .......

Gos 1871--------

Gosl433_ -.---

Gosl870__--....

Ast 2324.. --.--.

Gbsl434_-.--_-

Gos 1869-----.-..

Ast 2323-.--. __-

Gos 1435_. --__-_.

Ast 2321- --------

Gos 2320.-. ......

Gos2322_--.--_-

Gosl436-. ......

Ast 2319. -_......

Gos 1437---......

Location

Latitude 

north

deg. min.

36 40 

36 20 

36 20 

36 20 

36 10 

36 09 

36 00 

35 51 

35 50 

35 40 

35 41 

35 31 

35 29 

35 26 

35 20 

35 21 

35 16 

35 11 

35 09 

35 08 

35 07 

35 07 

35 03 

34 59

Longitude 

west

deg. min.

74 45 

75 45 

75 30 

75 14 

75 30 

74 58 

75 14 

75 15 

75 00 

75 15 

75 02 

75 15 

74 58 

75 27 

75 15 

74 59 

75 28 

75 15 

75 38 

75 48 

75 29 

75 21 

75 57 

75 31

Depth 

(meters)

86 

16 

30 

32 

26 

43 

35 

32 

46 

34 

43 

30 

42 

15 

25 

70 

16 

27 

16 

16 

16 

26 

17 

44

g

00

>,

 a 

2 

B

T3

£? 

5

1

o 

S

H5

X 

X

X)

X

X

X

X 

X

X

afl. A. gomillionensis (Howe and Ellis, 1935)

X

--

X 

X 

X 

X 

X

X 

X 

X

X 

X

X

--

vineyardensis (Cushman, 1906)

X

X

--

--

--

<<

d

«o 

1

S 

"§

X

--

--

-

--

rgittoecia sp. A

X

X

--

--

--

M

d

X

--

--

O 

d

X

-

--

--

--

--

P 

d

X

--

M 

1

d

*<o 

o 

O 

 a

g 

M

i

o

 2

X

X

X 

X

X

X 

X

X 

X

X

X

laevicula (Edwards, 1944)

X

--

--

--

--

X

affinicythere emarginata (Sars, 1865)

01

--

asslerites miocenicus (Howe, 1935)

0)

--

--

--

X

X

<< 

d

01

X

--

--

X

'.nsonocythere americana Hazel, 1967

0)

X

X

X

X

X 

X

X

X

arenicola (Cushman, 1906)

0)

X

X

X

X 

X

X

X 

X

3

M

-a

1

oi 

"3

0)

X

--

X

--

-

whitei (Swain, 1951)

0)

X

X 

X 

X

X

X

X 

X

--

<{

d

0)

X

X

X

X

X

--

M

d

01

X

X

X 

X

X

X

o

d

0)

X

X

X

X

X

-

0 

d

0)

--

--

w 

d

01

X

X 

X

--

X 

X

--

i 

s

3

M 

 3

«o 

S

I

0)

--

--

X

X

--

I

0)

--

-

--

ythocypris sp. A

01

--

--

--

\jthocythere sp. A

0)

X

X

--

X

X

M 

d

0)

--

--

X

--

impylocythere laeva Edwards, 1944

O

X

X

X

X 

X

X

indites nipeensis van denBold, 1946

O

-

-

.-

jLshmanidea magniporosa Hall, 1965

0

X

X 

X

X

X

X

seminuda (Cushman, 1906)

0

X 

X

X

X 

X

X

X 

X

X

 <

d

O

X

X

X

X 

X

X

X 

X

M 

d

0

-.

X

--

o 

d

O

--

--

Q

d

O

--

--

-

-

Cyprideis sp. A

--

..

--

-

Cytherella spp.

X

--

--

X

Cytherelloidea sp. A

--

X

--

--

Cytheridea sp. A

X

X

X

--

--

 <

d 

 3

 G 

§

--

--

--

--

Cytheromorpha afl. C. curta Edwards, 1944 C. warner

newporteWilliam1966

X

X

X

X

X

X
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D5

;

XX

X

X

XX

X

I

1 X

X

X

X

X

; X

X

X

!

X

X

I

!

X

X

XX

X

X

X

i X

i X

X

;

!

;

X

X

X

X

X

X

X

X

X

> f

X)

x:

X

X

XX

XX

X

XX

X

X

x>

X

X

X

X

X

X

X

X

X

X

X

X

X

X

>

e>

>

X)

X)

X)

x:

x

x

x

x

x

x

ex

f \.

XXX

XXX

X X

X X

X

X

X

XX

X

XXX

X

>

X X

X

xx>

X

X

XX

X

X

X

XX

X

X

X

f

X

1

X

X

X

XX

X

X

x

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TABLE 1. Occurrences of 171 ostracode species in Holocene samples from the shelf off the middle Atlantic Coast of the United States an

Norfolk Formation, as used by Oaks and Coch (1963), of southeastern Virginia Continued

Sample

Locality

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

47

Source

Gos 1375 ---.

Gos 1377. .... .

Gos 1316 

Gos 1319... 

Gos 1322 ......

Gos 1346 .. ....

Gos 1345..-. ....

Gos 1340 .... .

Ast 1965.... .....

Gos 1361.. .......

Gos 1338... ..

Gos 2026 . ....

Gos 1364.. .......

Gos 1355-.-. ....

Ast 2027 ... ...

Gos 2029..... .

Gos 1360.. ...--. -

Gos 2106... ......

Gos 1415... .... .-

Gos 1366.. .......

Gos 2032 ... ..

Gos 1417 .... ..

Gos 2036.... -

Gos 1418. .... .

Gos 1891... ......

Ast 2001-..- . .

Gos 2038.. .......

Gos 2041 ...

Gos 2045...... ...

Gos 1886-.... .

Gos 1886. ..

Gos 2047 -----

Gos 1422------

Gosl883.___ .... .

Ast 1997.... ... ..

Gos 2048-. .

Gos 1423.-.. ..

Gos 2050- - 

Gos 2051.- ......

Gos 1424 ......

Gos 2057... ......

Gos 1425- .- 

Gos2054.___. _

Gos 2053-..-...

Location

Latitude 

north

deg. min.

39 38 

39 31 

39 21 

39 20 

39 20 

39 09 

39 10 

39 10 

39 00 

39 01 

39 01 

38 50 

38 51 

38 49 

38 40 

38 40 

38 40 

38 30 

38 30 

38 31 

38 20 

38 10 

38 00 

37 59 

38 00 

37 49 

37 50 

37 40 

37 39 

37 31 

37 31 

37 27 

37 19 

37 20 

37 20 

37 10 

37 09 

37 05 

37 00 

37 00 

37 02 

36 57 

36 49 

36 50 

36 45 

36 39 

36 40

Longitude 

west

deg. min.

73 59 

73 45 

74 14 

73 31 

72 46 

74 31 

74 15 

73 15 

74 45 

74 00 

73 01 

74 45 

73 32 

73 15 

74 45 

74 14 

73 30 

74 45 

74 00 

73 26 

74 30 

74 15 

75 00 

74 29 

73 59 

75 25 

74 45 

75 14 

74 42 

75 30 

74 30 

74 29 

75 16 

75 00 

74 46 

75 44 

75 15 

75 01 

75 30 

75 15 

75 00 

74 45 

75 45 

75 15 

74 59 

75 31 

75 15

Depth

(meters)

24 

28 

20 

48 

84 

23 

21 

62 

15 

38 

80 

18 

62 

81 

26 

39 

69 

29 

57 

89 

36 

41 

18 

49 

129 

12 

41 

24 

49 

16 

64 

406 

29 

38 

56 

12 

30 

38 

30 

36 

44 

82 

22 

29 

36 

20 

35

/therura sp. A

O

--

--

--

X

--

X

X

X

X 

X

X

--

« 

ft

o

--

--

-

X

--

X

--

--

--

0

ft

ta

o

--

--

--

--

--

--

X

--

Q

&

o

--

--

X

--

--

--

X

X

--

w

Q.

o

--

-

--

-

--

--

--

-

h 

ft

o

--

-

--

-

-

--

--

--

o

ft

o

--

--

--

--

--

--

--

--

--

-1

w

ft

CO

o

--

--

-

-

-

--

"

--

--

ft

o

--

--

--

X

--

--

X

--

--

Hj

ft

o

--

-

--

--

--

--

X

--

M

&

o

--

--

-

--

--

--

--

--

--

(OZ8I '^P^-ia) JU9ftJtMSjimu si3J3^}fi3omyt

*

-

-

--

-

X

--

--

--

E. planibasalis (Ulrich and Bassler, 1904)

--

--

--

X

X

X

--

--

 4

ft

^

--

--

--

--

X

X

X

--

--

X

Eucythere declwis (Norman, 1865)

--

X

X

X

X

X

X

X 

X

X

X 

X

X

X 

X 

X 

X

X

X

E. gibba Edwards, 1944

--

-

-

X

X

X

--

--

--

E. triangulata Puri, 1954

--

--

--

--

--

..

--

--

 < 

ft 

K|

--

--

--

--

--

..

--

--

(8981 'sxes) mtipjanamf nipuiipjmwuig

X

X

X

X

X 

X

X 

X 

X 

X

--

X

(8E6I 'uosu8tid8}S) tfipvjtq mpu^tnMid'Djj

-

--

X

X

-

X 

X 

X

X

--

H. setipunctata (Brady, 1869)

--

--

--

--

--

-

--

--

--

3

Q

§

1 

F?

fl

--

-

"

--

--

--

-

--

--

(9061 'umnqsno) ouoouauio DUjsOutpifj

X

--

X

X

X

X 

X

X 

X 

X 

X

X

--

. glabra (Hall, 1965)

&3

--

--

-

--

--

--

--

--

--

. rugipustulosa (Edwards, 1944)

fl

X

X

X

X

--

X 

X

X 

X

X

 <

ft

&3

X

--

X

X

X

X 

X

X 

X 

X

X

X

«

ft

as

--

--

--

--

X

--

X 

X

--

o

ft

as

X

X

X

X

-

X

X 

X 

X 

X

X

X

X

Q 

ftM

as

--

--

-

--

-.

X

--

Jonesia cf. J.acuminata (Sars, 1865)

--

-

--

--

--

-

--

--

Krithe sp. A

--

-

-

-

--

--

--

--

« 

ft

fcd

--

--

--

-

--

--

--

--

--

Leptocythere angusta Blake, 1935

X

X

X

X

X

X

X

X

X 

X

X

X

X 

X

X

L. cf. nikraveshae Morales, 1966

--

--

--

--

--

..

--

X

--

 < 

ft

^'

-

--

--

--

--

-

--

--

Loxoconcha off. L. granulata Sars, 1865

X

-

X

--

X

X 

X 

X

X

L. impressa (Balrd, 1850)

X

X

X

X

X 

X

X 

X 

X 

X

X

X

L. matagordensis Swain, 1955

--

--

-

--

--

--

--

--

--

L. reticularis Edwards, 1944

-

--

--

--

--

--

--

--

--

L. sperata Williams, 1966

X

X

X

X

X

X

X 

X 

X 

X

X

X

Loxocorniculum postdorsolatum (Puri, 1960)

--

--

-

--

-

--

--

MacrosA

-

-

-

-

-

-

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LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D7

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TABLE 1. Occurrences of 171 ostracode species in Holocene samples from the shelf off the middle Atlantic Coast of the United States and 

Norfolk Formation, as used by Oaks and Coch (1963), of southeastern Virginia Continued

Sample

Locality

1

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

2,2 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41

Source

Gos 1378- 

Gos 1377 -...--

Gos 1316 

Gos 1319-. ...

Gosl322_. ------

Gos 1346 

Gos 1345 -----

Gos 1340- .

Ast 1965- 

Gos 1351------

Gos 1338 

Gos 2026---------

Gosl354- ------

Gos 1355- 

Ast 2027 -

Gos 1360. ___ 

Gos 2105. ------

Gos 1415. --------

Gos 1365.- 

Gos 2032------

Gos 1417 

Gos 2036--- 

Gos 1418 

Gos 1891 

Ast 2001. 

Gos 2038------

Gos 2041. -------

Gos 1420------.

Gos 2045-- --.

Gos 1886 

Gos 1885. 

Gos 2047. - 

Gos 1422-------

Ast 1997.- 

Gos2048 ...

GOS1423 

Gos 1424--- -

Location

Latitude 

north

deg. mln.

39 38 

39 31 

39 21 

39 20 

39 20 

39 09 

39 10 

39 10 

39 00 

39 01 

39 01 

38 50 

38 51 

38 49 

38 40 

38 40 

38 40 

38 30 

38 30 

38 31 

38 20 

38 10 

38 00 

37 59 

38 00 

37 49 

37 50 

37 40 

37 39 

37 31 

37 31 

37 27 

37 19 

37 20 

37 20 

37 10 

37 09 

37 05 

37 00 

37 00 

37 02

Longitude 

west

deg. min.

73 59 

73 45 

74 14 

73 31 

72 46 

74 31 

74 15 

73 15 

74 45 

74 00 

73 01 

74 45 

73 32 

73 15 

74 45 

74 14 

73 30 

74 45 

74 00 

73 26 

74 30 

74 15 

75 00 

74 29 

73 59 

75 25 

74 45 

75 14 

74 42 

75 30 

74 30 

74 29 

75 16 

75 00 

74 46 

75 44 

75 15 

75 01 

75 30 

75 15 

75 00

Depth 

(meters)

24 

28 

20 

48 

84 

23 

21 

62 

15 

38 

80 

18 

62 

81 

26 

39 

69 

29 

57 

89 

36 

41 

18 

49 

129 

12 

41 

24 

49 

16 

64 

406 

29 

38 

56 

12 

30 

38 

30 

36 

44

Munseyella atlantica Hazel andValentine, 1969

X

I

 8 

5

 a 

B

1

[g,

 5

§

o

a

Neolophocythere subquadrata Grossman, 1967

X

a 

fe;

X

X

Neonesideaf sp. A

pq 

a

fe;

Orionina bradyi van de Bold, 1963

0. vaughni (Ulrich and Bassler, 1904)

Paracypris sp. A

pq 

a

Paracytheridea altila Edwards, 1944 j

P. rugosa Edwards, 1944

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D10 CONTRIBUTIONS TO PALEONTOLOGY

76 "30' 76° 15'

36°45'

FIGURE 1. Map of southeastern Virginia indicating important geomorphologic features and localities sampled (P1-P6) in the

Norfolk Formation as used by Oaks and Coch (1903).

underlie more than one plain ; and (4) both marine and 

nonmarine formations are present. Five major periods 

of submergence before the post-Wisconsin submergence 

have been postulated to explain the stratigraphic relations of the eight post-Yorktown formations which they 

recognize in this region (the Yorktown Formation is 

the youngest unit of the Chesapeake Group and in this 

region is generally considered to be of late Miocene and 

early Pliocene age).

The post-Yorktown, pre-Holocene sediments which 

underlie the coastal plain of southeastern Virginia between the Suffolk Scarp and the sea (fig. 3) were designated the Late Columbia Formation and "Pliocene" by 

Darton (1902); were divided into the Dismal Swamp 

and Princess Anne Formations by Wentworth (1930) ; 

and were designated the Nansemond Formation by 

Moore (1956). These sediments were divided by Oaks 

(1964) and Coch (1968) into five formations of

Pleistocene age. With the exception of one sample from 

the overlying Kempsville Formation, the ostracode assemblages for this study were collected from the Norfolk Formation, which is the most widespread and fossiliferous of these formations (see figs. 1, 2, and 3).

The Norfolk Formation was described by Clark and 

Miller (1906) as consisting of fossiliferous clays and 

sands of Pliocene age exposed in deep cuts along the 

Dismal Swamp Canal and. buried elsewhere in southeastern Virginia under thick Pleistocene sediments. No 

type section was designated. The formation was assigned a Pliocene age on the basis of earlier identifications of mollusks (presumably collected from the same 

beds) made by L. Woolman (in Darton, 1902). Subsequently, Clark and Miller (1912, p. 175, 187-188), on 

the basis of J. A. Gardner's identification of a molluscan 

assemblage from the Dismal Swamp Canal, considered 

these fossiliferous beds to be part of the Talbot Forma-
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA Dll

EXPLANATION

 f o Locality PI

Locality P2

Serpulid worm tubes

Fe concretions

P2-1

Locality P6

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FIGUEE 2. Measured sections at late Pleistocene localities PI, P2, and P6 in southeastern Virginia.

tion of Shattuck (1901) of Pleistocene age. Oaks (1964, 

p. A10) described a boring in the Dismal Swamp (1.5 

miles west of the Dismal Swamp Canal) and designated it the type section of the Norfolk Formation.

The redefined Norfolk Formation extends east and 

west of the Suffolk Scarp (figs. 1, 3) and consists of 

two members. The lower member extends in the subsurface eastward from the Suffolk Scarp as a bed of 

medium to coarse sand and gravel of beach origin. The 

upper member, west of the Suffolk Scarp, consists of 

clayey sands of fluvial-estuarine origin exposed at an 

altitude of 45 to 50 feet or 15-16 m (meters) ; along 

the scarp this member crops out as fine to coarse dune 

and beach sands which reach an altitude of 70 feet. In

the area immediately south of the James River, the 

upper member consists of estuarine silty sands. Marine 

clayey and silty sands of this member underlie the area 

farther south of the James River and east of the Suffolk Scarp; the top of the formation in this area is 

0 to 10 feet (0-3 m) above present sea level, except in 

the Fentress Rise where these sediments are exposed 

at altitude of 17 to 25 feet (5-8 m).

The Norfolk Formation, according to Oaks (1964), 

represents the marine phase of a late Pleistocene interglacial transgression of the sea. This transgression began 

with the deposition of the Great Bridge Formation, 

which comformably underlies the Norfolk Formation 

and contains a restricted molluscan fauna indicative of
D12 CONTRIBUTIONS TO PALEONTOLOGY

A'

EXPLANATION

*

Undifferentiated

Sand Bridge Formation

8 10 12 14 MILES 

I I I I

Bacons Castle Formation

Sedley Formation

Londonbridge Formation | |

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Kempsville Formation | S I Yorktown Formation

Norfolk Formation

VERTICAL EXAGGERATION X 840

INDEX MAP

Great Bridge Formation 

Windsor Formation

FIGURE 3. Geologic section across southeastern Virginia from the Suffolk Scarp to the Atlantic coast. Adapted from Oaks (1964,

figs. 25, 28) and Coch (1968, fig. 5).

an estuarine or open bay environment of lower than 

normal salinity (H. G. Kichards in Oaks, 1964, p. 109). 

Although Norfolk sediments are principally marine, 

estuarine deposits of Norfolk age occur west of the 

Suffolk Scarp and along the south edge of the James 

River east of the scarp. The occurrence of beach sands, 

of Norfolk age at an altitude of 45 feet (15 m) dune 

sands occur up to 70 feet (23 m) (Coch, 1968) along 

the erosional Suffolk Scarp, and estuarine deposits west 

of the scarp at 50 feet (16 m) indicate that the sea extended approximately 35 miles (56 km) westward from 

the present coast and was 45 to 50 feet (15-16 m) above 

present sea level. The altitude of the collected samples

relative to present sea level indicates, if possible minor 

tectonic effects are discounted, that these marine sediments were deposited at depths of 20 m or less. The 

thickness of the Norfolk Formation (10-50 ft, about 

3-16 m), of both estuarine sediments west and marine 

sediments east of the Suffolk Scarp, indicates that the 

transgressive sea of Norfolk time was stable for a long 

period (Oaks, 1964). According to H. G. Richards 

(in Oaks, 1964, p. 122), Pleistocene molluscan assemblages of the Norfolk Formation are similar to nearshore assemblages which today occur only farther south 

in waters warmer than the present coastal waters of 

southeastern Virginia. Radiocarbon dating of wood
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D13

found in the Great Bridge, Norfolk, and Kempsville 

Formations gives an age greater than 40 X103 years B.P. 

(before present) (Oaks, 1964, table 2), and the Great 

Bridge-Norfolk transgression is thought to represent 

the Sangamon Interglaciation.

The maximum development of the last glacial stage 

in North America, the Wisconsin Glaciation, has been 

dated at about 16 X103 years B.P. on the basis of radiometric ages of shallow-water marine deposits associated with a maximum regression of sea level to about 

 425 feet ( 130 m) relative to present sea level (Milliman and Emery, 1968). The preceding Sangamon Interglaciation, representing a significant rise in sea level, is 

much more difficult to date. On the east coast of the 

United States a Sangamon age has been tentatively 

assigned to relict shorelines which parallel the present 

coast and appear to represent the last major stabilized 

rise of the sea above its present level. Radiocarbon dates 

of fossil wood in sediments apparently contemporaneous with these shorelines in southeastern Virginia 

have given ages of greater than 40X103 years B.P. 

(Oaks, 1964). Recent radiometric dating of shorelines 

that are at higher elevations than present sea level 

suggests that the Sangamon Interglaciation reached its 

maximum development between 75 X103 years B.P. and 

125 X103 years B.P. (Broecker, 1965; Broecker and 

others, 1968; Hopkins, 1967; Osmond and others, 1965; 

Veeh, 1966, Veeh and Chappell, 1970).

HOLOCENE COLLECTIONS

Holocene samples for this study were collected from 

the continental shelf off the middle Atlantic Coast of 

the United States from about 32°30' to 40° N. (fig. 4) as 

part of the joint U.S. Geological Survey-Woods Hole 

Oceanographic Institution program of study of the Atlantic continental margin of the United States (Emery, 

1966). The geographic location, depth and species content of all samples used in the present study are presented in table 1 (also see Hathaway, 1966). Of the 

133 samples processed, 115 contained ostracode assemblages, and 159 species were identified by the author 

of this report. Only 66 of the species have been formally described; however, several of the undescribed 

forms have been misidentified and are illustrated in 

the literature. Nearly all species were represented by 

living as well as dead specimens. Eighty percent of 

the samples were collected from depths of 10 to 50 m; 

only three samples were collected at depths greater than 

100 m. The 18 barren samples were composed of very 

coarse sand or worn shell fragments and were generally 

from shallow water near the coast (fig. 4).

0 25 50 75 100 Kilomete 

,1 I I 1 I

10 0 10 20 30 40 50 60 Nautical Miles 

I I I I 1 I I I 1

CONTOURS IN METERS

FIGURE 4. Map showing locations of Holocene sample stations. 

For exact location see table 1. Symbol X indicates that sample 

did not contain ostracodes.

The shelf sediments in the study area are principally 

relict upper Pleistocene quartzose silts and sands

426-209 O - 71 - 3
D14 CONTRIBUTIONS TO PALEONTOLOGY

(Emery, 1968, fig. 8). The carbonate content of the 

sediments north of Cape Hatteras is low (generally less 

than 5 percent) but increases significantly south of the 

cape and locally reaches values greater than 50 percent 

(Milliman and others, 1968, fig. 3).

HOLOCENE SPECIES DISTRIBUTIONS

FAUNAL PROVINCES AND CLIMATIC ZONES OF THE 

NORTH ATLANTIC OCEAN

The geographic distribution of marine organisms indicates that the sublittoral areas of the world can be 

divided into regions that are characterized by certain 

assemblages of species. The boundaries of these regions, 

or faunal provinces, are areas in which the geographic 

ranges of assemblages terminate or overlap with assemblages of neighboring provinces. Geographic ranges 

of marine organisms are influenced by water temperature, salinity, substrate, available food and light, and 

myriad other ecological relations. The effect of water 

temperature on the metabolism and reproduction of 

organisms is considered the fundamental factor determining species and assemblage distribution and therefore the geographic extent of sublittoral faunal provinces (Gunter, 1957; Kinne, 1963; Hall, 1964). The 

marine temperature of a region of the continental shelf 

is determined chiefly by its latitudinal position and the 

configuration of oceanic currents. Coastal topography at 

many places interferes with marine circulation patterns 

and causes an abrupt change in the climatic regime, thus 

creating a province boundary (Valentine, 1963).

Along the Atlantic coast of North America the northerly drift of warm water and the southerly drift of 

cool water interact at several points and cause isotherm 

convergences. At Cape Hatteras, warm southern shelf 

waters, influenced by the offshore Florida Current, meet 

the cool northern shelf waters of the Virginian Coastal 

Current. Farther north along the coast a similar situation prevails at Cape Cod, Mass., where cold northern 

waters influenced by the Labrador Current interact with 

the warm southern water of the Gulf Stream. The convergence of isotherms creates a temperature gradient 

which is a formidable barrier to species distribution; as 

a result the faunal provinces of this part of the western 

North Atlantic are sharply delineated. The North 

Atlantic Drift, of which the Gulf Stream is a part, 

proceeds northeast from off Newfoundland, warms the 

higher latitudes of the eastern North Atlantic Ocean, 

and prevents isotherm convergences along the European 

coast. Consequently, there are no prominent temperature barriers along that coast, and faunal provinces are 

relatively poorly delineated, the areas of overlapping 

species ranges being very large.

The sublittoral faunal provinces of the North Atlantic 

Ocean have been established by many workers on the 

basis of the distribution of shallow-water and intertidal 

organisms. The geographic limits of the provinces recognized by various authors differ little, but confusion 

has been caused by the varied nomenclature of f aunalprovinces and climatic-zones. Hazel (1970) reviews the 

development of the concept of faunal provinces and 

climatic zones in the North Atlantic Ocean. Six faunal 

provinces have been delineated along the east coast of 

North America and are, from north to south: Arctic, 

Labrador, Nova Scotian, Virginian, Carolinian, and 

Caribbean (Hazel, 1970). Holocene samples used in the 

present study were collected from the southern Virginian province (Cape Cod to Cape Hatteras) and from 

the northern Carolinian province (Cape Hatteras to 

Cape Kennedy).

The importance of the marine climate on the formation of faunal provinces is expressed in the establishment of sublittoral climatic zones whose boundaries are 

coincident with those of faunal provinces. Climatic 

zones, unlike faunal provinces, are characterized by and 

equated in such qualitative terms as frigid, subfrigid, 

cold-temperate, mild-temperate, warm-temperate, subtropical, and tropical. The configuration of currents in 

the North Atlantic and North Pacific ocean basins 

causes isotherm convergences and temperature gradients 

in the western parts and produces a more equable marine climate in the eastern parts of the basins; therefore, 

similarly named climatic zones in eastern (or western) 

parts of different ocean basins are more likely to resemble each other thermally than similarly named zones 

on opposite sides of the same basin. The temperature 

regimes on opposite sides of the same basin are so different that even if the same species were present off both 

the eastern and western coasts of an ocean basin, faunal 

provinces on one side would have little similarity to 

those on the other side. Along the east coast of North 

America the sublittoral faunal provinces and equivalent 

climatic zones are: Arctic frigid; Labrador subfrigid; Nova Scotian cold-temperate; Virginian 

mild-temperate; Carolinian subtropical; Caribbean 

tropical. At present there is no warm-temperate faunal 

province in this region.

LATITUDINAL RANGES OF HOLOCENE SPECIES

The occurrence of Holocene ostracodes on the continental shelf off North America from 32°30' to 40°N. 

indicates that the marine climatic conditions that pre< 

vail at Cape Hatteras, N.C., form a significant barrier 

to ostracode species distribution. Of the 159 species 

identified, 105 occur at more than five stations, and 54
LATE PLEISTOCENE OSTRACODE S, SOUTHEASTERN VIRGINIA D15

species are rare. The following is a list of 19 well-known 

species found only north of Cape Hatteras and 28 wellknown species found only south of the cape:

North of Cape Hatteras

Actinocythereis dawsoni

vineyardensis 

Bensonocy there arenicola

sp. A

sp. B

sp. C

sp. D

Bythocythere sp. A 

Cushmanidea seminuda 

Cytheridea sp. A 

Cytheropteron pyramidale 

Cytherura wardensis 

Finmarchinella finmarchica 

Leptocythere angusta 

Loxoconcha impressa

sperata

Muellerina canadensis 

Neolophocythere sp. A 

Puriana rugipunctata

South of Cape Hatteras

Bythocythere sp. B 

Cytherellolda sp. A 

Cytherura reticulata

aff. C. sablensis

sp. F

sp. G

Eucythere triangulata 

Hulingsina glabra 

Loxoconcha reticulata 

Loxocornicuhim postdorsolatum 

Macrocyprina propinqua

sp. A

sp. B

Neocaudites sp. A 

Neonesidea? sp. A

sp. B

Orionina bradyi 

Paracytheridea altila

rugosa

sp. A 

Paranesidea? sp. A

sp. B

sp. C 

Pellucistoma magniventra

sp. A

Proteoconcha multipunctata 

Puriana sp. A

sp. B

The late Pleistocene ostracode assemblage from southeastern Virginia comprises 82 species, TO of which are 

still living. In order that we may determine if a relation exists between the Pleistocene and Holocene assemblages of the study area, the modern latitudinal ranges 

of 68 Pleistocene species are plotted in figure 5. The

ranges of 64 of the species overlap between 35° and 

35°30' north latitude. This area is just off Cape Hatteras, where the shelf is narrowest for the region (30-35 

km), and the shelf-slope break occurs at about 50-m 

depth. It appears from this evidence that the Pleistocene assemblage may be represented by the same species 

in a Holocene assemblage in this small area. A comparison of Pleistocene and Holocene samples on the basis 

of their constituent species was undertaken to explore 

their relation more fully.

HOLOCENE SHELF ASSEMBLAGES DEFINED BY 

CLUSTER ANALYSIS

In order to establish relations between Holocene shelf 

samples, the author arranged for Dice coefficients of 

similarity [(2C/Ni+N2 ) X100, where C=the number of species in common between the two samples containing Ni and Nz species; see Cheetham and Hazel, 

1969] to be calculated and the samples clustered using 

the unweighted pair-group method of Sokal and Michener (1958). The matrix was 159 species by 91 samples 

(24 small samples were eliminated). The dendrogram 

(fig. 6) indicates that the samples segregate into two 

major clusters, A and B, on the basis of species content. 

Clusters A and B contain only samples occurring north 

or south of Cape Hatteras and provide further evidence 

that there is a boundary between the Virginian and 

Carolinian faunal provinces at the cape. Each of the 

major sample clusters has two subclusters. Cluster A 

contains subclusters 1 and 2 and includes a minor group 

of samples (12, 28, 30, and 26) which exhibits very low 

similarity internally between its constituent samples as 

well as externally to samples in subclusters 1 and 2. 

Cluster B is composed of subclusters 3 and 4 and also 

includes samples 83 and 104 which show low similarity 

to the other samples in cluster B.

A shaded similarity matrix (trellis diagram) (Sokal 

and Sneath, 1963; Valentine and Peddicord, 1967) 

using 20-point classes was constructed. The midpoint 

of each class was used to calculate average similarity 

coefficients within and between subclusters. See figure 7, 

a synoptic similarity matrix which provides a somewhat quantitative representation of the relations indicated by the dendrogram (fig. 6).

BIOFACIES OF THE SHELF

The geographic distribution of the 105 most wideO ~ JL

spread of the 159 Holocene ostracode species was compared to the distribution of the sample subclusters. 

Each subcluster contains a characteristic assemblage 

or biofacies that indicates certain environmental conditions (fig. 8. table 2). Biofacies 1 contains chiefly
D16 CONTRIBUTIONS TO PALEONTOLOGY

MODERN LATITUDINAL RANGES OF 68 SPECIES FOUND IN THE PLEISTOCENE OF THE NORFOLK, VA., AREA

;^ls

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fill I**-!

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i ^ ^ "s> « 3 tj "Sj ,' DHCqCjCqasi; Cji40-

39-

38-

37-

36-

35-

34-

33-

FIGUBE 5. Modern latitudinal ranges along the middle Atlantic Coast of the United States of 68 ostracode species common to 

the Holocene and the Norfolk Formation as used by Oaks and Ooeh (1963).

northern forms that do not occur south of Cape Hatteras. Biofacies 2 contains the northern forms of I in 

addition to species that occur north and south of the 

cape. Biofacies 3 contains species occurring north and 

south of the cape. Biofacies 4: is composed of species 

occurring north and south as well as species restricted 

to south of the cape. Biofacies 2 and 4: have large assemblages, whereas 1 and 3 have less species diversity. 

Biofacies 2 contains most of the principal species of 1 

and 3 ; biof acies 4: contains most of the principal species 

of only 3. These relations are shown in figure 7.

The composition of each biofacies is controlled by 

environmental factors. Temperature is the principal 

factor in areas as large as those being examined, but

in the area occupied by biofacies 3, substrate and energy 

conditions are important. Temperatures characteristic 

of the four biofacies were derived from maps showing 

average monthly bottom temperatures in 2.5°C contour 

intervals (Walford and Wicklund, 1968). Biofacies 1 

experiences a yearly temperature range of 2.5°-5°C to 

12.5°-15°C, biofacies 2 a range of 2.5°-5°C to 20°- 

22.5°C, and biofacies 3 and 4 a range of 12.5°-15°C to 

higher than 27.5 °C.

The effect of substrate on the composition of the 

biofacies is difficult to determine. Detailed mapping of 

bottom sediments has been done only south of Cape 

Hatteras (Cleary and Pilkey, 1968; Milliman and 

others, 1968, fig. 8). Most shelf sediments are Pleisto-
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D17

100 90 70

DICE COEFFICIENT OF SIMILARITY

60 50 40 30

CAPE HATTERAS (lat 35°15'N.)

FIGURE 6. Dendrogram of Holocene sample clusters. Samples are compared on the basis of their ostracode 

species composition using the Dice similarity coefficient (2C/Ni-\-Na) X100; clustering was by the 

unweighted pair-group method.
D18 CONTRIBUTIONS TO PALEONTOLOGY

33

32-'

32

10

10

23

32

32

28

49

10

32

42

38

10

28

42

34

Average of similarity

coefficients from 

intersections of subclusters

23

49

38

34

46

0 - </i

a) .y c

00 £ °

46

48

FIGURE 7. Similarity matrix using Dice coefficient (2C/Ni+Nz) 

XlOO showing relation between clusters of Holocene and Norfolk Formation samples and also between the Norfolk cluster 

and an artificial sample grouping based on latitudinal overlaps of species.

cene relict quartz sands, but the carbonate content increases significantly south, of the cape. Biof acies 1 and 2 

are associated with quartz sands. Biof acies 4 is generally 

restricted to sediments containing greater than 5 percent calcium carbonate. Biofacies 3, a small assemblage 

characterized by many robust forms (Aurila floridana, 

Campylocythere laeva, Cushmanidea magniporosa, 

Cytheromorpha warneri, newportensis, Haplocytheridea 

bradyi, Hulingsina amerieana, Hulingsina rugipustulosa, Hulingsina sp. A, Hulingsina sp. B, Hulingsina 

sp. C, Proteoconclia gigantica, Proteoconcha nelsonen5/5, Proteoconcha tiiberculata, Protocytheretta daniana, 

Protocytheretta aff. P. sahnii, Puriana floridana, and 

Puriana sp. A) is restricted to areas of coarse quartzose 

sediment generally containing less than 5 percent 

calcium carbonate. These areas are best developed off 

Cape Hatteras, at Cape Lookout Shoals (loc. 86), Frying Pan Shoals (loc. 98), in the nearshore areas of bays 

(loc. 68, 74, 101), and off inlets in barrier islands (loc. 

71, 81, 93) (fig. 8). Biofacies 3 is best developed in the 

area off Cape Hatteras and appears discontinuous to 

the south. Because its species can tolerate temperatures

greater than 27.5°C, biof acies 3 may exist intermittently 

in nearshore areas south of the study area.

10 0 10 20 30 40 50 60 Nautical Miles 

I I 1 I I I I I I

CONTOURS IN METERS

FIGURE 8. Map showing the geographic extent of four ostracode 

biofacies delineated on the shelf off the middle Atlantic 

Coast of the United States. Symbol X indicates that sample 

did not contain ostracodes; solid dot indicates that sample 

contained an assemblage too small to be indicative of one 

of the biofacies.
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D19

TABLE 2. Principal Holocene species of shelf biofacies; their occurrence in the Norfolk Formation (upper Pleistocene); their occurrence

north and south of Cape Hatteras area

[X, characteristic species of the biofacies; o, species rarely occurring in the biofacies; P, species present]

aff. A. gomillionensis _ ________ __ __________

Aurila floridana. .__ ______ ______ ______ ____ ____

laevicula-.-- _ ________ _____ ___ ________ _ __

Bensonocythere americana _ ____ ______________

arenicola

cf. B. sapeloensis^ _ ________ ___ _________ _____

whitei- ______________ ______ ___________ ____

sp. B. _ _ _ _ _ _ _____ ________

or> P

sp. D____ _____ ______________ _ ______________

sp. E-___ __________ _________ _ ____ _________

sapeloensis.. _ _________ ______ _________ _____

sp. B_ _ __ _ _ _ __ __ ___ ___ _ _ _ _____

Campylocythere laeva.. _____ __ _____ __ _____ _ __ _

Cushmanidea magniporosa ___ _______ _______________

sp. A _ ____ ____ __________________ _ _____ __

Cytherelloidea sp. A ____ _____ ______ __________

Cytheromorpha warneri newportensis _ _________________

Cytherura elongata.. ____ ____ ______ _______________

foru lata

howei.. _________ ____ _ ____ ____ _ _____ ____

pseudostriata- ________ ______ _ _______________

reticulata

aff. C. sa6Jerms_____ _ _ _ _ _________________

sp. A__________ __ ____________________________

sp B

sp. D_____ ________ __ ___ ___ ___ _ _ _

sp. F__________________________________________

sp. G___ ___ _ __ __ _ _ ___ __ _ ____ ____

Echinocythereis margaritifera __________ __________ __.

planibasalis.- __________________ ______________

sp. A. _ _______ ___ ____ ______ ____ ___ _ ___

Eucythere declivis _ ____ ____ ___ _________ ____

gibba. ____________ __ _____ __ ______ _________

triangulata _ _ _ __ _____ _____ ___________ ______

sp. A______ _ _ _ _ _ _ ____ ______ _ _ ___ __

Finmarchinella finmarchica- _____ ______ ____________

Haplocytheridea 6rodyi.___ _ __ ______ ___ ___ ____

Hulingsina americana. ___ ________________________ __

glabra- _______ ______ _______________________

rugipustulosa. _______ _ ________________ _______

sp. A _ ________ ___ _____ ____________________

sp. B__ _______________________________________

sp. C__. _______________________________________

sp. D __ ______ _ __ ____ _ ____

Loxoconcha impressa. _ _____ ______ _____ ________

reticularis... ________ _ _ __

aff. L. granulata. _ __ _____ _ _ __ __________

Loxocorniculum postdorsolatum..,.. _______ _______ __

Macrocyprina propinqua-. ____ ___ _________ ________

sp. A__ _ _ ______ __ ______ ___ __ ______ ___

Microcytherura choctawhatcheensis ___ _______________

sp. A _ ___________________________________ ___

sp. C _______________________________________

aff. M . licnenklausi.. __ _________ ____ ____

Neocaudites sp. A___ _ ___ ______ _ ___ _

Biofacies

1

______ __ _______ X

_________________ X

_________________ X

_______ _ _ ___ X

_________________ o

v

V

v

_______ _____ -_ O

v

___________ _____ 0

__________________ X

_____________ ____ o

__________________ X

______ ___________ o

.-_______-_ _- -_ O

__________________ X

__________ _____ _ o

.____-__ -_- _____ O

v

__________________ X

_____ ____ ______ o

_________ ____ _ o

___ ___ ______ o

._________-_--.-_- X

2

X

o

X

X

o

X

X

X

X

v

X

X

X

v

X

X

X

X

o

X

v

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X

o

o

V

o

X

X

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o

X

X

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X 

X

v

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0

o

0

o

X

X

x

v

X

v

3

0

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0

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o

o

o

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X

o

X

o

X

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o

o

X

o

o

o

0

o 

X

X

o

X

X

X

X

o

o

o

X

o

o

o

Cape Hatteras area

4 North

X

X

X _

o

o

o

X

o

X

X

X _ X

X

X -

x . X

X - X

X

X

X

x . X

o

X

x . x . x .

o

x .

o

X

x . x .

X

X

X

X

X

X

X

X

x .

o

x . x . x . x . X

X

o

0

X

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

_ P

P

P

_ P

P

P

P

P

P

P

- P

P

P

P

- P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

_ P

P

P

P

P P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

South

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

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P

P

P

P

P

P

P

P

P

P

P

P

P

. P

P

P

P

P

P
D20 CONTRIBUTIONS TO PALEONTOLOGY

TABLE 2. Principal Holocene species of shelf biofacies; their occurrence in the Norfolk Formation (upper Pleistocene); their occurrence

north and south of Cape Hatter as area Continued

Species

Biofacies

Pleistocene

Cape Hatteras area

North South

Neonesidea ? sp. A______ ______________ _ __ __ _ __. 

? sp. B___. _____________________________________

Orionina bradyi.-.- _ _____ ________ __________ _ __.

Paracytheridea altila.-. _____ __________ _ ______

rugosa ._ ____ __________ _ ____ _______ _______

sp. A__.. _______ _ _______ ___. _____ _ ____ _

Paradoxostoma delicata _________________ __ ____ __.

sp. B_ ___ _ ___ _ _ _ _-___.___ _______ _ _

sp. D __ __ ____ __ _ ___ _____ __ _ __ _ _

Paranesidea ? sp. A__ _______ _ __ _______ _ __ ____

? sp. B____ _ __ _ ____ _ _ _ __ __ _ _ __

? sp. C . _ __ _ _ ___ -_ _ ___ _ _ __

Pellucistoma magniventra ___ _____ ____ _ _ _ ___ __

sp. A__ ___ __________ ___ __ _______ _ ___ _

Phlyctocythere sp. A__ __________ __ _______ _ _______

sp. B___ ________________________________________

Pontocypris sp. A __ __ ______ __ __ ___ _________

Propontocypris edwardsi... _______ __________ ___ ____

aff. P. howei- _________ ____ ___ __________ ____

Proteoconcha gigantica _ ____ __ ____ __ __ __ _ ____

multipunclata.--. ___ ________ ____________ _____

nelsonensis _ ___ _ __ __ _ _ _____ _____ _______

tuberculata __ _ _ _ _____ _ __ _ __ ______ __

Protocytheretta daniana _ ____ _ ___ _ _ ______ ____

aff. P. sahnii _ ___ _ __ _ __ _____ _ ___ _ __ __

Pseudocytheretta edwardsi.. _ _____ ____ _____ _ ___

sp. A_ _______ ___ _ __________________ ____ __

Puriana floridana-.---- __ ___ __ _____ _ ________ _.

sp. A _ _______ _______ _______ _________ _ __

sp. B___ _ _ _ __ _ ___ __ ____ _______ _

"Sahnia" cf. "S" subulata. _ _. ____ ____ __ ___ _ __

sp. A _ __ __ _ _____ __ _ _________ __ ____ _

sp. B_ _ ___ , _ _ __ __ ____ __ __ ._ __

________ _ __ _________ o

__-__-_-_ ________ 0

.__ _______ ____ __ __ o

__ ___ __ -___ _____ _ 0

_______ _ __ _____ ___ 0

__ __ _______ _______ o

______________________ X

______________________ X

____ _ __ _ __ ______ o

_ __ _________ _ __ 0

__ _____ _____________ 0

________ _ _____ _ - 0

__ ____ _____ _ _____ 0

___ __ _ -___ __ _____ O

__ _____ ____ _ __ _ o

___ _ __ _ _ __ o ______

___________ __________ o

________________ o X

______________________ X

____ .__ _ _______ ___ o

_ __ ____ __ _____ o

_________________ o X

___-_-____--___-___-_. X

________________ X X

_________________ X X

___ ___ ____ _____ o

________-__-___--__-__ X

________________ o X

_ __ _ __ __ __ ____ o

______________________ X

_________________ X X

_________-____--_--__-. X

X

o

o

X

o

o

X

o

o

o

X

0

0

X

X

X

0

o

X

X

X

0

o

o

X . X .

X .

X

X .

X

X

0

X .

X .

X .

X .

X

X - X - X .

o

X

X .

X

X -

0

X

X - X

0

X .

X

o

X .

X -

0

0

X

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P 

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

P

Thirty samples were not included in the four biofacies 

by the cluster analysis (24 small samples not included 

in the cluster program and six samples which showed 

low similarity to samples in the major clusters A and 

B). The species content of these samples was compared 

with the typical assemblages of the biofacies, and 21 

samples (5, 7, 12, 14, 18, 23, 25, 26, 28, 30, 33, 44, 46, 4V, 

50, 59, 65, 68, 83, 96, and 104) were assigned to a particular biofacies on this basis. Four samples (10, 13, 27, 

and 42) are transitional between biofacies 1 and 2, and 

five samples (2,16, 21, 94, and 111) contained an assemblage too small to be indicative of one of the biofacies 

(fig. 8).

FAUNAL PROVINCES IN THE CAPE HATTERAS AREA

Two recent zoogeographic studies have been conducted in the Cape Hatteras area. Cerame-Vivas and 

Gray (1966) investigated the benthic invertebrate assemblages of a relatively small area from Cape Lookout to north of Cape Hatteras. They recognized three 

areas characterized by distinctive assemblages and

equated these areas with parts of three faunal provinces : 

the Virginian north of Cape Hatteras, the Carolinian 

on the inner shelf south of the cape, and the Tropical or 

Caribbean on the outer shelf south of the cape and north 

to about 36° N. (Cerame-Vivas and Gray, 1966, figs. 3, 

5). Maturo (1968) studied the bryozoan distribution on 

the shelf from the Florida Keys to 40° N. and concluded 

that he could recognize only two faunal provinces near 

Cape Hatteras, the Carolinian plus Tropical south of 

the cape, and the Virginian to the north.

The geographic extent of the four ostracode biofacies 

established in this study is indicated in figure 8. Biofacies 2, 3, and 4 all occur in the Cape Hatteras area. 

The dendrogram (fig. 6), on which a delineation of the 

Virginian and Carolinian faunal provinces may be 

based, indicates that biofacies 3 and 4 constitute the 

northern part of the Carolinian province, and biofacies 

1 and 2 the southern part of the Virginian province. 

Biofacies 3, although an entity, is closely related to 

biofacies 4; in fact, all major species characteristic of 

3 are found in 4.
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D21

COMPARISON OF LATE PLEISTOCENE AND 

HOLOCENE ASSEMBLAGES

The Dice coefficients of similarity between late Pleistocene samples are very high. The average of similarity 

coefficients within the Pleistocene samples is 78 and 

indicates the very close relation between the samples, 

on the basis of their species composition. The similarity 

matrix (fig. 7) illustrates the relation between the 

Pleistocene sample cluster and the Holocene sample 

subclusters. The Pleistocene assemblage is most similar 

to biof acies 2, 3, and 4, in descending order and least 

similar to biof acies 1, an assemblage composed principally of species restricted to the area north of Cape 

Hatteras. Although the Pleistocene assemblage is most 

similar to biofacies 2, the two assemblages are not 

equivalent and show less similarity to each other than 

the Holocene biofacies show internally.

The author attempted to create an artificial assemblage which would resemble that of the late Pleistocene. 

Holocene samples were selected from the area in which 

there are overlaps of latitudinal ranges of Holocene 

species which also occur in the Pleistocene samples (figs. 

5, 8). This artificial group includes samples from biofacies 2 and 3 from latitudes between 35° and 30°30' N. 

(localities 61, 62, 63, 64, 67, 69, 70, 71, and 73). The 

similarity between the artificial group and the Pleistocene samples is somewhat lower than the similarity between biofacies 2 and the Pleistocene samples, and the 

internal similarity of the artificial group is lower than 

that of any of the biofacies, Holocene or Pleistocene 

(fig. 7). The Pleistocene ostracode assemblage does not 

closely resemble assemblages extant in the study area, 

but this does not prevent determining marine climatic 

conditions prevailing during deposition of the Norfolk 

Formation (upper Pleistocene).

LATE PLEISTOCENE MARINE CLIMATE OF 

SOUTHEASTERN VIRGINIA

HOLOCENE MARINE BOTTOM TEMPERATURES

The distribution of marine organisms, though easily 

mapped, is often difficult to explain except in terms of 

the climatic tolerances of the species involved. The existence of major f aunal provinces has been explained on 

temperature tolerances of organisms. Limiting temperatures, however, do not include extremes of short 

duration. According to Hall (1964), the number of consecutive days or months that shallow sea water is at temperatures required for reproduction and early growth 

determines the distribution of organisms and therefore 

faunal provinces and climatic zones.

The configuration of average monthly bottom temperatures in the study area, as illustrated by Walford

and Wicklund (1968), shows that Cape Hatteras is a 

point of isotherm convergence during the coldest 

(February) and warmest (August-September) months 

of the year (figs. 9,10). At Cape Hatteras in February

10 0 10 20 30 40 50 60 Nautical Miles 

111 II I I I I

CONTOURS IN METERS

FIGURE 9. Bottom-temperature map for the shelf off the middle 

Atlantic Coast of the United States for February, the coldest 

month of the year (redrawn from Walford and Wicklund, 

1968, pi. 13). Temperature in °C.
D22 CONTRIBUTIONS TO PALEONTOLOGY

10 0 10 20 30 40 50 60 Nautical Miles 

I i | I I I I I I

CONTOURS IN METERS

FIGURE 10. Bottom-temperature map for the shelf off the middle Atlantic Coast of the United States for August-September, the warmest part of the year. Bottom temperatures are 

highest in August immediately south of Cape Hatteras and 

highest in September immediately north of the cape (redrawn 

from Walford and Wicklund, 1968, pis. 14, 15). Temperature 

in °C.

the bottom temperature is about 12.5°C, ranging from 

10° to 15°C. North of the cape temperatures of the waters 

off the coast of New Jersey may reach as low as 2.5°C, 

but in general temperatures of the waters in the north 

ern area range from 5° to 12.5°C. South of Cape Hatteras, temperatures in February may reach as low as 

12.5°C in the shallow waters of Long Bay, but range 

from 15° to 20°C over most of the shelf. Bottom temperatures are highest in August south of the cape and 

in September north of the cape, but the temperature 

regime in the cape area is identical for both months 

when isotherms ranging from 17.5° to 25 °C converge at 

Cape Hatteras. North of the cape, temperatures reach a 

maximum of 25 °C in nearshore waters off the North 

Carolina and Virginia coasts, but generally the bottom 

temperatures on the shelf range from 10° to 22.5°C. 

South of Cape Hatteras, bottom temperatures of the 

shelf range from 12.5°C to greater than 27.5°C off the 

coast of North and South Carolina.

LATE PLEISTOCENE BOTTOM TEMPERATURES

Biofacies 1, 2, 3, and 4 are indicative of certain maximum and minimum average monthly water temperatures. Although the late Pleistocene assemblage represents none of these biof acies, the marine climate of the 

Pleistocene can be inferred by determining temperature 

tolerances of species common to the Holocene and Pleistocene. The geographic distribution of these species was 

plotted on bottom-temperature maps for the warmest 

and coldest months of the year, and species temperature 

tolerances were determined (fig. 11). Temperature 

ranges were plotted in 2.5°C classes to adjust for inherent errors in sea-water-temperature maps which are all 

based on nonsystematic sampling and relatively few 

data points for the large areas mapped.

The overlap of species temperature tolerances (fig. 

11) permits determining marine bottom temperatures in 

southeastern Virginia during that part of the Pleistocene represented by the Norfolk Formation. On the basis 

of the temperature tolerances of eight ostracode species, 

the minimum bottom temperature during the coldest 

part of the year was between 12.5° and 15 °C. All these 

species occur only south of Cape Hatteras, and the distribution of four, Hulingsina gldbra, Paracytheridea 

altila, Paracytheridea sp. A, and Pellucistoma magniventra, is very well known in the study area. In fig. 11, 

the temperature tolerances of five species, Actinocythereis dawsoni, Cytheropteron pyramidale, Cytherura 

sp. B, Cytheridea sp. A, and Neolophocythere sp. A, suggest that the bottom temperature in the warmest month 

was between 17.5° and 20°C. However, the first species 

is very rare in the Pleistocene, and the distribution of 

the last three is not well documented in the Holocene.
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D23

TEMPERATURE RANGES OF 66 HOLOCENE SPECIES WHICH OCCUR IN THE PLEISTOCENE OF THE NORFOLK, VA., AREA

to ^

. «M 

' Q &'«

1^ 1 O H '

S5 1

3| s §

s S

 S v * 'J

2 H w »4'

; a & t & § s

I f^ <3 t e § fe

I 11 ? I 13

' S S .g ' > * ti

0 § S

! s -I I e C 3 -O 3>

1 00 *

.i £

<* I. I S 2 e g I a e

i e S

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' to

0-2.5 

2.5-5.0 

5.0-7.5 

7.5-10.0 

10.0-12.5 

12.5-15.0 

15.0-17.5 

17.5-20.0 

20.0-22.5 

22.5-25.0 

25.0-27.5 

>27.5 

FIGURE 11. Temperature ranges of 66 ostracode species common to the Holocene and the Norfolk Formation (upper

Pleistocene).

Five other species, Bensonocythere arenicola, Bensonocythere sp. A, Bensonocythere sp. B, Leptocytliere 

angusta, and Muellerina canadensis have upper limits 

between 20° and 22.5°C; these taxa are common in the 

Pleistocene, except for Muellerina canadensi-s, and their 

Holocene distribution is well documented. These data 

strongly suggest that the bottom temperature in shallow 

water during the warmest part of the year was between 

20° and 22.5°C. Thus, the bottom-temperature range

indicated for the late Pleistocene in this area was between 12.5°-15°C minimum and 20°-22.5°C maximum, 

a yearly range of about 7.5°C. These temperature values 

are contrasted with those of inner shelf waters off southeastern Virginia which today have bottom temperatures 

ranging from a minimum of 5°-Y.5°C to a maximum of 

22.5°-25°C (very near shore), a range of about 1T.5°C, 

or 10 °C greater than during the late Pleistocene in 

southeastern Virginia. Most of this increase in the yearly
D24 CONTRIBUTIONS TO PALEONTOLOGY

temperature range results from the difference between 

the Pleistocene and Holocene winter minima and indicates that during the late Pleistocene interglaciation, the 

marine environment of southeastern Virginia had 

warmer winters than now occur off southeastern Virginia, whereas summer maxima differed little.

In a study of the ostracodes of some Texas bays and 

lagoons (depth to 9 ft, salinity 10-50°/oo), King and 

Kornicker (1970) observed temperature ranges for 

several species that occur in the late Pleistocene of 

southeastern Virginia. Temperature ranges of Aurila 

floridana (= Aurila conradi littorala of King and Kornicker), Cytherura forulata, Hulingsina rugipustulosa 

(= Hulingsina sandersi of King and Kornicker), Haplocytheridea bradyi, and Haplocytheridea setipunctata 

(a species which does not occur in the Holocene samples 

of this study but is common in the Pleistocene) indicate 

that these species could live within the temperature 

limits of 12.5° to 22.5° C established for the late Pleistocene of southeastern Virginia.

Hemicythere villosa occurs commonly in the Pleistocene samples but is absent in the Holocene samples 

from the shelf area studied. This species occurs today 

in the Bay of Biscay, off the coasts of northern Europe 

and eastern Canada and south to Vineyard Sound, 

Mass., in depths to 37 m (Hazel, 1970). Elofson (1941) 

describes this species as an algal feeder living in shallow water at temperatures from 0° to 22° C. The modern known geographic range of Hemicythere villosa 

does not extend south of Vineyard Sound, but this may 

be due to inadequate collecting in shallow nearshore 

areas or to some unknown ecological factor. The temperature and depth tolerances of this species would 

allow it to live in the Pleistocene environment characterized by bottom temperatures of 12.5° to 22.5° C and 

depths of approximately 15 to 20 m or less.

LATE PLEISTOCENE SEA-SURFACE TEMPERATURES

In a comparison of average monthly bottom temperatures (figs. 9, 10) and average monthly sea-surface 

temperatures north of Cape Hatteras off southeastern 

Virginia (Schroeder, 1966, figs. 2, 8), it is noted that 

the sea surface is coldest in February (5°-12.5° C) 

and that the surface and bottom isotherms are nearly 

coincident. Therefore, the late Pleistocene sea-surface 

temperatures for nearshore areas during the coldest 

part of the year were possibly equivalent to the bottom 

temperatures and ranged from 12.5° to 15° C. The sea 

surface is warmest in August, and the configuration 

of the surface isotherms differs markedly from the bottom isotherms for August and September. Sea-surface 

isotherms trend northeast-southwest, range from 25° 

to 27.5° C, and average about 26° C. In contrast, bottom isotherms for this part of the inner shelf parallel 

the coastline, range from 20° to 25° C, and average 

22.5° C, or approximately 4° C lower than the seasurface temperature. Therefore, the late Pleistocene seasurface temperatures for nearshore areas during the 

warmest part of the year ranged possibly from 24° 

to 26.5° C, or 4° C higher than the bottom-temperature 

range of that time (20°-22.5° C). If these postulated 

sea-surface maxima (24°-26.5° C) and minima (12.5°- 

15° C) are correct, the range of the late Pleistocene 

sea-surface temperatures in southeastern Virginia was 

about 11° or 12° C.

LATE PLEISTOCENE FAUNAL PROVINCE AND 

CLIMATIC ZONE OF SOUTHEASTERN VIRGINIA

Modern or contemporaneous sublittoral faunal provinces of the middle latitudes in different ocean basins 

(the North Atlantic and North Pacific, for example) or 

off opposite coasts of the same ocean basin contain different species and assemblages and therefore cannot be 

equated. Because populations, communities, and provinces are constantly evolving (Valentine, 1968), provinces existing at different times in the same region will 

not be equivalent even under similar climatic conditions, and similarity between provinces decreases as 

difference in age increases. Late Cenozoic fossil f aunal 

provinces can, however, be fairly closely equated with 

modern provinces in the same region if many constituent 

species are extant and if the past and present marine 

climates are similar. Although 85 percent of the species 

of the late Pleistocene ostracode assemblage of the Norfolk Formation are still living, the assemblage is unlike 

any found today on the inner shelf off eastern North 

America. The species still exist but are found in 

different associations because the climatic regime has 

altered.

The Norfolk ostracode assemblage is indicative of a 

marine climate which may be described in modern climatic zone terminology as warm-temperature, a climatic regime which does not now exist off the middle 

Atlantic Coast of North America. If sea level rose about 

15 m above present sea level, a very wide shelf would be 

created in the Virginia-North Carolina area (where the 

mild-temperature Virginia faunal province and the 

subtropical Carolinian faunal province meet now on the 

present narrow shelf) ; in Virginia the shoreline would 

be at the Suffolk Scarp (fig. 1). This shelf would be 

approximately 160 km wide off southeastern Virginia 

and about 135 km wide off North Carolina at the latitude of Cape Hatteras. The path of the Florida Current, which follows the continental slope and extends to 

a depth of 800 m over the Blake Plateau (Pratt, 1966), 

would probably be little altered by a rise in sea level
LATE PLEISTOCENE OSTRACODES, SOUTHEASTERN VIRGINIA D25

of 15 m; however, the cold Virginia Coastal Current 

which flows southward over the shelf would be diffused 

over a much larger area. A broad expanse of shallow 

water extending inland from the present coast would 

therefore be less influenced by warm southern and cool 

northern waters than are present shelf waters in the 

area, and the marked convergence of isotherms at Cape 

Hatteras would be eliminated.

A rise in sea level (to an altitude of +15 m relative to 

present sea level), a consequent increase in shelf area 

and diffusion of thermal currents, and a general atmospheric warming all apparently contributed to establish 

a Pleistocene marine climate with a very narrow annual 

range in temperature and, therefore, markedly different 

from any now found in the western North Atlantic. 

Ostracode species that today are restricted to subtropical and warmer waters or mild-temperature and cooler 

waters coexisted and suggest that along the middle Atlantic Coast of the United States during the Sangamon 

Interglaciation there was a distinctive faunal province 

and corresponding warm-temperate climatic zone 

which have since been eliminated. The convergence of 

cool- and warm-water masses at Cape Hatteras profoundly influences the marine climate and consequently 

the distribution of marine organisms and at present 

prevents the reestablishment of a warm-temperate 

faunal province in this region.

SUMMARY

1. The Norfolk Formation (upper Pleistocene (Sangamon) ) of southeastern Virginia contains a large 

and well-preserved ostracode assemblage, 82 species, 70 of which are still living.

2. No modern shelf assemblage off the middle Atlantic 

Coast of the United States is closely similar to that 

of the Norfolk Formation; the Norfolk assemblage 

includes forms which today are restricted to the 

area north or south of Cape Hatteras.

3. The boundary between the modern Virginian and 

Carolinian faunal provinces at Cape Hatteras, already well established on the geographic distribution of other organisms, is also reflected in the 

distribution of ostracode species on the continental 

shelf. From the cluster analysis of 91 Holocene 

samples containing 159 ostracode species, four biof acies are recognized on the shelf between 32°30'N 

and 40 °N. Temperature is the principal factor controlling the composition of the biof acies, although 

substrate and energy conditions are important in 

defining the species composition of biofacies 3.

4. The Norfolk ostracode assemblage lived at depths of 

20 m or less, and temperature tolerances of ostracode species common to the Pleistocene and the

Holocene indicate that the assemblage lived in 

waters where the bottom temperature in the coldest month was between 12.5° and 15°C and in the 

warmest month between 20° and 22.5°C.

5. The late Pleistocene marine climate of the inner sublittoral was more equable than now exists on the 

inner shelf off southeastern Virginia where minimum and maximum monthly temperatures range 

from 5°-7.5° C to 22.5°-25° C, respectively. The 

inner sublittoral of southeastern Virginia had 

warmer winters during the late Pleistocene interglaciation than now occur on the inner shelf off 

southeastern Virginia, whereas summer maxima 

differed little.

6. During the Sangamon Interglaciation a rise in sea 

level ( + 15 m relative to present sea level), a consequent increase in shelf area, diffusion of cool and 

warm oceanic currents over a broad area, and a 

general atmospheric warming apparently all contributed to establish a warm-temperate marine climate off the middle Atlantic Coast of the United 

States.

7. The late Pleistocene ostracode assemblage is indicative of a sublittoral faunal province and warmtemperature climatic zone which has no modern 

counterpart off the Atlantic Coast of the United 

States. At present, the convergence of cool and 

warm water masses at Cape Hatteras prevents 

there being a warm-temperate faunal province in 

this region.

PLEISTOCENE LOCALITIES

Locality PI. Borrow pit 0.4 mile southeast of Acredale 

(Bonneys or Mears Corner; intersection of Virginia State Route 

190 and Indian River Road) on south side of Indian River 

Road; Virginia Beach, Va.; USGS Kempsville 7.5-min quadrangle 

(1965); 36°47'30" N., 76°10'13" W. USGS Cenozoic loc. 24796. 

Section measured and collected in southwest corner of pit in 

October 1968. Pit being flooded by ground water in January 1970. 

Unit and description Thickness (ft) 

Sand Bridge Formation:

1. Silty clay, unfossiliferous______-_-_-___________ 3.0

Kempsville Formation:

1. Interbedded sand and gravel, cross-bedded,

unfossiliferous________________-__-_----_--_ 4. 5

2. Fine-medium sand, crossbedded, contains wood_ _ 5. 5

3. Fine-medium sand; burrowed, bivalve ghosts;

wood_-_____________--__---_----_--------- 4. 0

4. Medium-coarse sand, very fossiliferous; nested,

disarticulated bivalves (Spisula) (sample Pl-1) _ 2. 0 

Norfolk Formation:

1. Fine sand, argillaceous, very f ossilif erous; articulated bivalves in living position (Mercenarid); 

serpulid worm tubes very abundant (sample 

Pl-2, collected 5 ft. from top)_____________ 10. 5

Total Exposed-_--__-_-------__----------.------- 29. 5
D26 CONTRIBUTIONS TO PALEONTOLOGY

Locality P2. Borrow pit 0.6 mile east of Davis Corner (intersection of U.S. Route 58 and Newtown Road) on south side 

of U.S. Route 58 at the end of Toy Ave., in Boulevard Manor 

subdivision of Virginia Beach, Va.; USGS Kempsville 7.5-min 

quadrangle (1965); 36°50'30" N., 76°10'20" W.; (USGS Cenozoic loc. 23780). Section measured and collected by T. R. Waller 

in July 1966. Pit flooded in May 1968. 

Unit and description Thickness (ft)

Kempsville (?) Formation:

1. Coarse sand and gravel, unfossiliferous__________ 6. 0

2. Medium-coarse sand mixed with gravel, very 

f ossilif erous; disarticulated worn bivalves 

(Spisula) __________________________________ 7. 0

Norfolk Formation:

1. Fine sand, argillaceous, very f ossilif erous; articulated bivalves in living position (Mercenaria), 

serpulid worm tubes very abundant (sample 

P2-1 collected from top 3 ft; P2-2 collected 

5 ft from top; P2-3 collected from bottom 3 ft) _ _ 16. 0 

Great Bridge (?) Formation:

1. Sand, argillaceous; contains Crassostrea-MyaRangia fauna; bivalves in living position 

(Crassostrea virginica and Mya) ______________ 2. 0

Total exposed-________________________ 31. 0

Locality P3. Borrow pit 0.4 mile north of Davis Corner 

(intersection of U.S. Route 58 and Newtown Road) 0.2 mile west 

of Newtown Road, Virginia Beach, Va., USGS Kempsville 7.5- 

min quadrangle (1965); 36°51'30" N., 76°11'00" W.; (USGS 

Cenozoic loc. 23800). Section collected by T. R. Waller in July 

1966. Pit flooded in July 1966.

Unit and description Thickness (ft) 

Norfolk Formation:

1. Sand, very f ossilif erous; contains Mercenaria and

serpulid worm tubes (sample P3-1)_________ 5. 0

Total exposed________________________ 5. 0

Locality P4- Borrow pit 0.4 mile southeast of Powells Crossroads (intersection of Holland Road and Virginia State Route 

646) on north side of Holland Road and south of Interstate 264; 

Virginia Beach, Va.; USGS Kempsville 7.5-min quadrangle 

(1965); 36°50'00" N., 76°08'00" W.;(USGS Cenozoic loc. 23801). 

Section collected by T. R. Waller in July 1966. Pit flooded in 

May 1968.

Unit and description Thickness (ft) 

Kempsville (?) Formation:

1. Sand, unfossiliferous__________________________ 5. 0

2. Coarse sand and gravel, very f ossilif erous; disarticulated bivalves (Mercenaria, Spisula)_____ 2.5

3. Coarse sand, unfossiliferous__________________ 3. 0

Norfolk Formation:

1. Sand, argillaceous, very fossiliferous (Ensis,

Busycon, sand dollars) (sample P4-1)_________ 10. 0

Total exposed________________________ 20. 5

Locality P5. Borrow pit 0.6 mile east of Greenbriar Road; 

on a farm road that intersects Greenbriar Road 0.2 mile south of 

intersection of Greenbriar Road, and Interstate 64; Chesapeake, 

Va.; USGS Kempsville 7.5-min quadrangle (1965); 36°47'00" N., 

76°13'30" W.; (USGS Cenozoic Iocs. 23802, 24797). Section 

measured and collected by T. R. Waller in July 1966. Recollected in May 1968. Pit flooded in October 1968.

Unit and description Thickness (ft) 

Norfolk Formation:

1. Sand and clay, unf ossilif erous__ __ ______________ 1-2

2. Sand, argillaceous, very fossiliferous; large serpulid 

worm tube concentrations, articulated Mercenaria in living position (sample P5-!)__________ 12. 0

Total exposed____________________ 13. 0-14. 0

Locality P.6 Borrow pits 0.5 mile southwest of Yadkin, Va., 

and 0.3 mile south of Norfolk and Western Railroad near the 

intersection of Portsmouth Ditch and powerlines; 0.6 mile south 

of Yadkin on Gallberry Road, turn right at cemetery, turn right 

at first dirt road (0.2 mile), 0.5 mile to pit; Cheseapeake, Va.; 

USGS Norfolk South 7.5-min quadrangle (1965); 36°45'30" 

N., 76°22'00" W. (USGS Cenozoic loc. 24798). Section measured 

and collected in cuts between north and south pits in May 1968 

and April 1969. Pits being worked in June 1970.

Unit and description Thickness (ft) 

Sand Bridge (?) Formation

1. Sand; large-scale crossbedding; interbedded with

thin layers of clay and peat___________________ 4. 0

Norfolk Formation

1. Fine-medium sand; contains fossiliferous beds 

(Ensis) and scattered shell fragments; thin bed 

at bottom contains broken shell fragments, 

iron concretions, and cobbles (sample 6-1 

collected 3.5 ft. from top)___________________ 7. 5

Yorktown Formation

1. Medium sand; iron cemented in part; very 

f ossilif erious; broken shells and bivalved 

Corbicula________________--_--_--_---_--_ 3. 5

2. Sandy shell hash_______--_-----_-_-------- 4.0

Total exposed..__________-_____--_-_----_-----_ 19. 0

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Florida panhandle; Part 3, Ostracoda : Florida Geol. Survey 

Bull. 36, p. 217-345 [1954].

 1958a, Ostracode genus Cushmanidea: Gulf Coast Assoc. 

Geol. Socs. Trans., v. 8, p. 171-181.

 1958b, Ostracode subfamily Cytherettinae: Gulf Coast 

Assoc. Geol. Socs. Trans., v. 8, p. 183-195.

 1960, Recent Ostracoda from the west coast of Florida: 

Gulf Coast Assoc. Geol. Socs. Trans., v. 10, p. 107-149.

Puri, H. S-, and Hulings, N. C., 1957, Recent ostracode facies 

from Panama City to Florida Bay area: Gulf Coast Assoc. 

Geol. Socs. Trans., v. 7, p. 167-189.

Rogers, W. B., 1884, Report of the progress of the geologic survey of the State of Virginia for the years 1835-1840, in 

Rogers, W. B., A reprint of annual reports and other papers 

on the geology of the Virginias: New York, D. Appleton 

and Co., 832 p.

Sandberg, P. A., 1964, The ostracod genus Cyprideis in the 

Americas: Stockholm Contr. Geology, v. 12, 178 p.

 1969, Appendages and family placement of the ostracod

genus Pellucistotna: Jour. Paleontology, v. 43, no. 5, pt. 1,

p. 1174-1178. 

Sars, G. O., 1865, Oversigt of Norges marine ostracoder:

Vidensk.-Selsk. Christiania Forh., 130 p. 

Schroeder, E. H., 1966, Average surface temperatures of the

western North Atlantic: Bull. Marine Sci., v. 16, no. 2,

p. 302-323. 

Shattuck, G. B., 1901, The Pleistocene problem of the North

Atlantic Coastal Plain: Johns Hopkins Univ. Circ., v. 20,

no. 152, p. 69-75. 

Sokal, R. R., and Michener, C. D., 1958, A statistical method

for evaluating systematic relationships: Kansas Univ. Sci.

Bull., v. 38, p. 1409-1438, 

Sokal, R. R., and Sneath, P. H., 1963, Principles of numerical

taxonomy: San Francisco, Calif., W. H. Freeman, 359 p. 

Stephenson, M. B. 1938, Miocene and Pliocene Ostracoda of the

Genus Cytheridea from Florida: Jour. Paleontology, v. 12,

no. 2, p. 127-148. 

Swain, F. M., 1951, Ostracoda from wells in North Carolina;

Pt. 1, Cenozoic Ostracoda: U.S. Geol. Survey Prof. Paper

234-A, 58 p.

 1955, Ostraeoda of San Antonio Bay, Texas: Jour. Paleontology, v. 29, no. 4, p. 561-646. 

Triebel, Erich, 1960, Die taxionomishe Stellung und die Gattungen der Unterfamilie Macrocypridinae (Ostracoda) :

Senckenbergiana Biol., v. 41, p. 109-124. 

Ulrich, E. O., and Bassler, R. S., 1904, Systematic paleontology,

Miocene Arthropoda: Maryland Geol. Survey, Miocene

(volume), p. 98-130. 

Valentine, J. W., 1963, Biogeographic units as biostratigraphic

units: Am. Assoc. Petroleum Geologists Bull., v. 47, no. 3, p.

457-466.

 1968, The evolution of ecological units above the population level: Jour. Paleontology, v. 42, no. 2, p. 253-267.

Valentine, J. W., and Peddicord, R. G., 1967, Evaluation of 

fossil assemblages by cluster analysis1 : Jour. Paleontology, 

v. 41, no. 2, p. 502-507.

Veeh, H. H., 1966, Th^/U238 and u^/U238 ages of Pleistocene 

high sea level stand): Jour. Geophys. Research, v. 71, no. 14, 

p. 3379-3386.

Veeh, H. H., and Chappell, John, 1970, Astronomical theory of 

climatic change : Support from New Guinea : Science, v. 167, 

p. 862-865.

Walford, L. A., and Wicklund, R. I., 1968, Monthly sea temperature structure from the Florida Keys to Cape Cod: Am. 

Geog. Soc. Serial Atlas of the Marine Environment, 

Folio 15.

Wentworth, C. K., 1930, Sand and gravel resources of the 

Coastal Plain of Virginia: Virginia Geol. Survey Bull. 32, 

146 p.

Williams, R. B., 1966, Recent marine podocopid Ostracoda of 

Narragansett Bay, Rhode Island: Kansas Univ. Paleont. 

Contr., Paper 11, 36 p.

U. S. GOVERNMENT PRINTING OFFICE : 1971 O - 426-209
PLATES 1-4

[Contact photographs of the plates in this report are available, at cost, from U.S. Geological 

Survey Library, Federal Center, Denver, Colorado 80225]
PLATE 1

[All figures X 50]

FIGURES 1, 6, 11, 12. Bensonocythere sapeloensis (Hall, 1965).

1. Lateral view left valve, female. Sample P5-1. USNM 168641. 

6. Lateral view right valve, female. Sample P4-1. USNM 168642.

11. Lateral view left valve, male. Sample Pl-2. USNM 168643.

12. Lateral view right valve, male. Sample Pl-2. USNM 168644. 

2, 3, 7, 8. Bensonocythere americana Hazel, 1967.

2. Lateral view left valve, female. Sample P4-1. USNM 168645.

3. Lateral view left valve, male. Sample P2-2. USNM 168646.

7. Lateral view right valve, female. Sample P5-1. USNM 168647.

8. Lateral view right valve, male. Sample P4-1. USNM 168648. 

4, 9, 15, 20. Bensonocythere sp. E.

4. Lateral view left valve, female. Sample P4-1. USNM 168649.

9. Lateral view right valve, female. Sample Pl-2. USNM 168650. 

15. Lateral view left valve, male. Sample P4-1. USNM 168651. 

20. Lateral view right valve, male. Sample P5-1. USNM 168652. 

5, 10, 16, 21. Bensonocythere sp. D.

5. Lateral view left valve, female. Sample P4-1. USNM 168653. 

10. Lateral view right valve, female. Sample P4-1. USNM 168654. 

16. Lateral view left valve, male. Sample Pl-2. USNM 168655. 

21. Lateral view right valve, male. Sample P2-2. USNM 168656.

13. 18. Bensonocythere sp. A.

13. Lateral view right valve, female. Sample P4-1. USNM 168657. 

18. Lateral view left valve, male. Sample P4-1. USNM 168658.

14. 19. Bensonocythere whitei (Swain, 1951).

14. Lateral view left valve, female. Sample P5-1. USNM 168659. 

19. Lateral view left valve, male. Sample P5-1. USNM 168660. 

17, 22, 30. Bensonocythere sp. B.

17. Lateral view left valve, female penultimate. Sample P4-1. USNM 168661. 

22. Lateral view right valve, female. Sample Pl-2. USNM 168662. 

30. Lateral view left valve, male. Sample P4-1. USNM 168663. 

23. Benscnocythere sp. G.

23. Lateral view right valve, female. Sample P6-1. USNM 168664. 

24, 25, 26, 27. Cushmanidea sp. B.

24. Lateral view left valve, male. Sample Pl-1. USNM 168665.

25. Lateral view right valve, male. Sample Pl-1. USNM 168666.

26. Lateral view left valve, female. Sample Pl-1. USNM 168667.

27. Lateral view right valve, female. Sample Pl-1. USNM 168668. 

28, 29. Bensonocythere cf. B. sapeloensis (Hall, 1965).

28. Lateral view left valve, female. Sample Pl-2. USNM 168669.

29. Lateral view left valve, male. Sample Pl-2. USNM 168670. 

31. Bensonocythere arenicola (Cushman, 1906).

31. Lateral view right valve, female. Sample P4-1. USNM 168671. 

32, 33, 36; 37. Campylocythere laeva Edwards, 1944.

32. Lateral view left valve, female. Sample P2-2. USNM 168672.

33. Lateral view left valve, male. Sample Pl-2. USNM 168673.

36. Lateral view right valve, female. Sample P2-2. USNM 168674.

37. Lateral view right valve, male. Sample Pl-2. USNM 168675. 

34, 38. Cushmanidea seminuda (Cushman, 1906).

34. Lateral view left valve, female. Sample P4-1. USNM 168676. 

38. Lateral view right valve, female. Sample P4-1. USNM 168677. 

35. Actinocythereis dawsoni (Brady, 1870).

35. Lateral view left valve, female. Sample Pl-1. USNM 168678. 

39, 40, 44, 48. Actinocythereis aff. A. gomillionensis (Howe and Ellis, 1935).

39. Lateral view left valve, male. Sample P6-1. USNM 168679.

40. Lateral view right valve, male. Sample P2-2. USNM 168680. 

44. Lateral view left valve, female. Sample P6-1. USNM 168681. 

48. Lateral view right valve, female. Sample P2-2. USNM 168682. 

41, 42, 45, 46. Cushmanidea magniporosa Hall, 1965.

41. Lateral view left valve, female. Sample P4-1. USNM 168683.

42. Lateral view left valve, male. Sample P4-1. USNM 168684.

45. Lateral view right valve, female. Sample P4-1. USNM 168685.

46. Lateral view right valve, male. Sample P4-1. USNM 168686. 

43, 47. Aurila fioridana Benson and Coleman, 1963.

43. Lateral view left valve, female. Sample P6-1. USNM 168687. 

47. Lateral view right valve, female. Sample P6-1. USNM 168688.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 683-D PLATE 1

BENSONOCYTHERE, CUSHMANIDEA, CAMPYLOCYTHERE, ACTINOCYTHEREIS, AND AURILA
PLATE 2

[All figures X 50]

FIGURES 1, 7. Cytherura howei (Puri, 1954).

1. Lateral view left valve, female. Sample Pl-2. USNM 168689. 

7. Lateral view right valve, female. Sample Pl-2. USNM 168690. 

2, 8. Cytherura sp. E.

2. Lateral view left valve. Sample Pl-1. USNM 168691. 

8. Lateral view right valve. Sample P2-2. USNM 168692. 

3, 4, 9, 10. Cytherura sp. B.

3. Lateral view left valve, female. Sample Pl-2. USNM 168693.

4. Lateral view right valve, female. Sample Pl-2. USNM 168694.

9. Lateral view left valve, male. Sample Pl-2. USNM 168695.

10. Lateral view right valve, male. Sample Pl-2. USNM 168696. 

5, 6, 11, 12. Cytherura reticulata Edwards, 1944.

5. Lateral view left valve, female. Sample Pl-2. USNM 168697.

6. Lateral view right valve, female. Sample Pl-2. USNM 168698.

11. Lateral view left valve, male. Sample Pl-2. USNM 168699.

12. Lateral view right valve, male. Sample Pl-2. USNM 168700.

13. 19. Cytherura wardensis Howe and Brown, 1935.

13. Lateral view left valve, female. Sample Pl-2. USNM 168701. 

19. Lateral view left valve, male. Sample Pl-2. USNM 168702.

14. 20. Cytherura pseudostriata Hulings, 1966.

14. Lateral view right valve, female. Sample P4-1. USNM 168703. 

20. Lateral view right valve, male. Sample Pl-2. USNM' 168704. 

15, 16, 21, 22. Cytherura sp. A.

15. Lateral view right valve, female. Sample Pl-2. USNM 168705.

16. Lateral view left valve, female. Sample Pl-2. USNM 168706.

21. Lateral view right valve, male. Sample Pl-2. USNM 168707.

22. Lateral view left valve, male. Sample Pl-2. LSNM 168708. 

17, 18, 23, 24. Cytherura sp. D.

17. Lateral view left valve, female. Sample P2-1. USNM 168709.

18. Lateral view right valve, female. Sample P2-1. USNM 168710.

23. Lateral view right valve, male. Sample P2-1. USNM 168711.

24. Lateral view left valve, male. Sample P2-1. USNM 168712. 

25, 26. Cytherura forulata Edwards, 1944.

25. Lateral view left valve, female. Sample Pl-2. USNM 168713.

26. Lateral view right valve, male. Sample P2-1. USNM 168714. 

27, 28. "Sahnia" cf. "S" subulata (Brady, 1868).

27. Lateral view left valve, female. Sample Pl-2. USNM 168715.

28. Lateral view right valve, female. Sample Pl-2. USNM 168716. 

29. Megacythere sp. B.

29. Lateral view left valve. Sample Pl-1. USNM 168717.

30. 35. Eucythere declivis (Norman, 1865).

30. Lateral view left valve, female. Sample Pl-2. USNM 168718. 

35. Lateral view left valve, male. Sample Pl-2. LSNM 168719.

31. 36. Puriana rugipunctata (Ulrich and Bassler, 1904).

31. Lateral view left valve. Sample P4-1. USNM 168720. 

36. Lateral view right valve. Sample P4-1. USNM 168721.

32. 37. Puriana floridana Puri, 1960.

32. Lateral view left valve, disarticulated carapace. Sample P5-1. USNM 168722. 

37. Lateral view right valve, disarticulated carapace. Sample P5-1. USNM 168723.

33. 38. "Sahnia" sp. A.

33. Lateral view right valve, female. Sample Pl-2. USNM 168724. 

38. Lateral view left valve, female. Sample Pl-2. USNM 168725. 

34. Megacythere stephensoni (Puri, 1954).

34. Lateral view left valve. Sample P2-2. USNM 168726. 

39. Pellucistoma magniventra Edwards, 1944.

39. Lateral view left valve, male. Sample Pl-2. USNM 168727.
PLATE 2 Continued

FIGURES 40, 41, 44, 45. Eucythere gibba Edwards, 1944.

40. Lateral view left valve, female. Sample P5-1. USNM 168728.

41. Lateral view left valve, male. Sample 5-1. USNM 168729.

44. Lateral view right valve, female. Sample P5-1. USNM 168730.

45. Lateral view right valve, male. Sample P6-1. USNM 168731.

42. 46. Haplocytheridea bradyi (Stephenson, 1938).

42. Lateral view left valve, female. Sample P6-1. USNM 168732. 

46. Lateral view right valve, male. Sample P6-1. USNM 168733.

43. 47. Protocytheretta aff. P. sahnii (Puri, 1952).

43. Lateral view left valve. Sample P6-1. USNM 168734.

47. Lateral view right valve. Sample P6-1. USNM 168735. 

48, 49. Haplocytheridea setipunctata (Brady, 1869).

48. Lateral view left valve, female. Sample P6-1. USNM 168736.

49. Lateral view left valve, male. Sample P6-1. USNM 168737. 

50. Pseudocytheretta edwardsi Cushman, 1906.

50. Lateral view right valve, penultimate. Sample P5-1. USNM 168738.

PROFESSIONAL PAPER 683-D PLATE 2

CYTHERURA, "SAHNIA," MEGACYTHERE, EUCYTHERE, PURIANA, PELLUCISTOMA, 

HAPLOCYTHERIDEA, PROTOCYTHERETTA, AND PSEUDOCYTHERETTA
PLATE 3

[All figures X 50] 

FIGURES 1, 2, 6, 7. Hulingsina sp. D.

1. Lateral view left valve, female. Sample P6-1. TJSNM 168739.

2. Lateral view left valve, male. Sample P6-1. TJSNM 168740.

6. Lateral view right valve, female. Sample P6-1. USNM 168741.

7. Lateral view right valve, male. Sample P6-1. USNM 168742. 

3. Hulingsina glabra (Hall, 1965).

3. Lateral view right valve, female. Sample P4-1. TjSNM 168743. 

4, 5, 9, 10. Hulingsina sp. B.

4. Lateral view left valve, female. Sample P5-1. USNM 168744.

5. Lateral view left valve, male. Sample P5-1. USNM 168745. 

9. Lateral view right valve, female. Sample P5-1. USNM 168746. 

10. Lateral view right valve, male. Sample P5-1. USNM 168747. 

8, 12, 13, 14. Hulingsina americana (Cushman, 1906).

8. Lateral view left valve, female. Sample P2-2. USNM 168748.

12. Lateral view right valve, female. Sample P2-2. USNM 168749.

13. Lateral view right valve, male. Sample P2-2. USNM 168750.

14. Lateral view left valve, male. Sample P2-2. USNM 168751. 

11, 15, 16. Hulingsina sp. A.

11. Lateral view left valve, female penultimate. Sample P4-1. USNM 168752.

15. Lateral view right valve, female penultimate. Sample P4-1. USNM 168753.

16. Lateral view right valve, female. Sample P3-1. USNM 168754. 

17, 18, 21, 22, 25, Hulingsina rugipustulosa (Edwards, 1944).

26, 29, 30. 17. Lateral view left valve, female. Sample P5-1. USNM 168755. 

18. Lateral view left valve, female. Sample P5-1. USNM 168756.

21. Lateral view right valve, female. Sample P5-1. USNM 168757.

22. Lateral view right valve, female. Sample P5-1. USNM 168758.

25. Lateral view left valve, male. Sample P5-1. USNM 168759.

26. Lateral view left valve, male. Sample P5-1. USNM 168760. 

29. Lateral view right valve, male. Sample P5-1. USNM 168761. 

 > 30. Lateral view right valve, male. Sample P5-1. USNM 168762.

(Note pronounced development of pustules in specimens illustrated in figures 18, 22, 26, and 30). 

19, 20, 23, 24. Hulingsina sp. C.

19. Lateral view left valve, female. Sample P6-1. USNM 168763.

20. Lateral view right valve, female. Sample P6-1. USNM 168764.

23. Lateral view left valve, male. Sample P6-1. USNM 168765.

24. Lateral view right valve, male. Sample P6-1. USNM 168766. 

27,28. Finmarchinella finmarchica (Sars, 1865).

27. Lateral view right valve, female. Sample P4-1. USNM 168767.

28. Lateral view left valve, male. Sample P4-1. USNM 168768. 

31, 32, 33, 34 Paracytheridea altila Edwards, 1944.

31. Lateral view left valve, female. Sample P6-1. USNM 168769.

32. Lateral view right valve, female. Sample P6-1. USNM 168770.

33. Lateral view left valve, male. Sample P2-2. USNM 168771.

34. Lateral view right valve, male. Sample P5-1. USNM 168772.

35. 39. Muellerina canadensis (Brady, 1870).

35. Laterial view right valve, male. Sample P4-1. USNM 168773. 

39. Lateral view right valve, female. Sample P4-1. USNM 168774.

36. 40. Muellerina aff. M. lienenklausi (Ulrich and Bassler, 1904).

36. Lateral view right valve, male. Sample P2-1. USNM 168775. 

40. Lateral view right valve, female. Sample P4-1. USNM 168776.

37. 41. Cytheropteron pyramidale (Brady, 1868).

37. Lateral view left valve, male. Sample P4-1. USNM 168777. 

41. Lateral view left valve, female. Sample P4-1. USNM 168778.

38. 42. Neocaudites sp. A.

38. Lateral view right valve, female. Sample P2-1. USNM 168779. 

42. Lateral view right valve, male. Sample P3-1. USNM 168780.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 683-D PLATE 3

HULINGSINA, FINMARCHINELLA, PARACYTHERWEA, MUELLERINA, 

CYTHEROPTERON, AND NEOCAUDITES
PLATE 4

[All figures X 50]

FiGUBES 1, 7. Microcytherura sp. C.

1. Lateral view left valve, female. Sample Pl-2. TJSNM 168781. 

7. Lateral view right valve, female. Sample Pl-2. TJSNM 168782. 

2, 3, 8, 9. Microcytherura sp. B.

2. Lateral view left valve, female. Sample Pl-2. TJSNM 168783.

3. Lateral view right valve, female. Sample Pl-2. TJSNM 168784.

8. Lateral view left valve, male. Sample P6-1. TJSNM 168785.

9. Lateral view right valve, male. Sample P6-1. TJSNM 168786. 

4. Paradoxostoma delicata Puri, 1954.

4. Lateral view left valve. Sample P2-1. TJSNM 168787. 

5, 6, 11, 12. Cytheromorpha warneri newportensis Williams, 1966.

5. Lateral view left valve, female. Sample P4-1. TJSNM 168788.

6. Lateral view right valve, female. Sample P4-1. TJSNM 168789.

11. Lateral view left valve, male. Sample Pl-2. TJSNM 168790.

12. Lateral view right valve, male. Sample Pl-2. TJSNM 168791. 

10. Paradoxostoma sp. B.

10. Lateral view left valve. Sample Pl-2. TJSNM 168792. 

13, 14, 19, 20. Microcytherura sp. A.

13. Lateral view left valve, female. Sample Pl-2. TJSNM 168793.

14. Lateral view right valve, female. Sample Pl-2. TJSNM 168794.

19. Lateral view left valve, male. Sample Pl-2. TJSNM 168795.

20. Lateral view right valve, male. Sample Pl-2. TJSNM 168796. 

15, 16, 21, 22. Paradoxostoma sp. F.

15. Lateral view left valve, male. Sample Pl-2. TJSNM 168797.

16. Lateral view right valve, male. Sample Pl-2. TJSNM 168798.

21. Lateral view left valve, female. Sample Pl-2. TJSNM 168799.

22. Lateral view right valve, female. Sample Pl-2. TJSNM 168800. 

17., Leptocythere sp. A.

17. Lateral view left valve. Sample P4-1. TJSNM 168801. 

18, 24. Leptocythere cf. L. nikraveshae Morales, 1966.

18. Lateral view right valve, female. Sample Pl-2. TJSNM 168802. 

24. Lateral view left valve, female. Sample Pl-2. TJSNM 168803. 

23. Paradoxostoma sp. G.

23. Lateral view right valve. Sample Pl-1. TJSNM 168804. 

25, 30. Microcytherura choctawhatcheensis (Puri, 1954).

25. Lateral view left valve, female. Sample Pl-2. LSNM 168805. 

30. Lateral view left valve, male. Sample P4-1. TJSNM 168806.

26. Sclerochilus sp. D.

26. Lateral view right valve. Sample P5-1. LSNM 168807.

27. Sclerochilus sp. C.

27. Lateral view left valve. Sample Pl-2. TJSNM 168808.

28. Paradoxostoma sp. C.

28. Lateral view left valve. Sample Pl-1. TJSNM 168809. 

29, 34. Leptocythere angusta Blake, 1933.

29. Lateral view left valve, female. Sample Pl-2. TJSNM 168810. 

34. Lateral view left valve, male. Sample Pl-2. TJSNM 168811. 

31, 32, 35, 36. Neolophocythere subquadrata Grossman, 1967.

31. Lateral view left valve, female. Sample Pl-2. USNM 168812.

32. Lateral view right valve, female. Sample Pl-2. TJSNM 168813.

35. Lateral view left valve, male. Sample Pl-2. TJSNM 168814.

36. Lateral view right valve, male. Sample Pl-2. USNM 168815.
PLATE 4 Continued

33, 37. Neolophocythere sp. A.

33. Lateral view right valve, male. Sample P4-1. USNM 168816.

37. Lateral view left valve, female. Sample P2-1. USNM 168817. 

38, 39, 43, 44. Loxoconcha matagordensis Swain, 1955.

38. Lateral view left valve, male. Sample Pl-2. USNM 168818.

39. Lateral view left valve, female. Sample Pl-2. USNM 168819.

43. Lateral view right valve, male. Sample Pl-2. USNM 168820.

44. Lateral view right valve, female. Sample Pl-2. USNM 168821.

40. 45. Loxoconcha aff. L. granulata Sars, 1865.

40. Left lateral view of carapace, female. Sample P3-1. USNM 168822. 

45. Right lateral view of carapace, female. Sample P3-1. USNM 168822.

41. 46. Loxoconcha sperata Williams, 1966.

41. Lateral view left valve, male. Sample P5-1. USNM 168823. 

46. Lateral view left valve, female. Sample P5-1. USNM 168824.

42. 47. Propontocypris edwardsi Cushman, 1906.

42. Lateral view left valve. Sample Pl-2. USNM 168825.

47. Lateral view right valve. Sample Pl-2. USNM 168826. 

48, 49, 53, 54. Proteoconcha tuberculata (Puri, 1960).

48. Lateral view left valve, male. Sample Pl-2. USNM 168827.

49. Lateral view right valve, male. Sample Pl-2. USNM 168828.

53. Lateral view left valve, female. Sample Pl-2. USNM 168829.

54. Lateral view right valve, female. Sample Pl-2. USNM 168830. 

50, 55. Proteoconcha nelsonensis (Grossman, 1967).

50. Lateral view left valve, male. Sample Pl-2. USNM 168831. 

55. Lateral view left valve, female. Sample P5-1. USNM 168832. 

51, 52, 56, 57. Hemicythere villosa (Sars, 1865).

51. Lateral view left valve, male. Sample Pl-2. USNM 168833.

52. Lateral view right valve, male. Sample Pl-2. USNM 168834.

56. Lateral view left valve, female. Sample Pl-2. USNM 168835.

57. Lateral view right valve, female. Sample Pl-2. USNM 168836. 

58, 59, 60, 61. Proteoconcha gigantica (Edwards, 1944).

58. Lateral view left valve, female. Sample 5-1. USNM 168837.

59. Lateral view right valve, female. Sample P4-1. USNM 168838.

60. Lateral view left valve, male. Sample P2-2. USNM 166839.

61. Lateral view right valve, male. Sample P4-1. USNM 168840.

GEOLOGICAL SURVEY PROFESSIONAL PAPER 683-D PLATE 4

MICROCYTHERURA, PARADOXOSTOMA, CYTHEROMORPHA, LEPTOCYTHERE, SCLEROCHILUS,

NEOLOPHOCYTHERE, LOXOCONCHA, PROPONTOCYPRIS,

PROTEOCONCHA, AND HEMICYTHERE