Atlantic Continental Shelf and Slope 

of the United States 

Ostracode Zoogeography in the Southern Nova Scotian 

And Northern Virginian F annal Provinces 

GEOlOGICAL SURVEY PROFESSIONAL PAPER 529-E 

Atlantic Continental Shelf and 

Slope of the United StatesOstracode Zoogeography in 

The Southern Nova Scotian 

And Northern Virginian 

Faunal Provinces 

By JOSEPH E. HAZEL 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 529-E 

Geographic distribution and temperature and 

depth tolerances of Recent Arctic-North 

Atlantic benthonic ostracodes with particular 

emphasis on the sublittoral species off the 

northeastern United States 

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1970 
UNITED STATES DEPARTMENT OF THE INTERIOR 

WALTER J. HICKEL, Secretary 

GEOLOGICAL SURVEY 

William T. Pecora, Director 

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

Washington, D.C. 20402 
CONTENTS 

Page 

Abstract___________________________________________ E1 Historical development of faunal provinces ____________ _ 

Introduction_______________________________________ 1 North America ________________________________ _ 

Acknowledgments___________________________________ 1 

Previouswork______________________________________ 1 

Presentwork_______________________________________ 3 

Water temperatures from Nova Scotia to Long Island___ 3 

Substrate from Nova Scotia to Long Island_ _ _ _ _ _ _ _ _ _ _ _ 5 

Europe _______________________________________ _ 

Temperature control of species distribution ___________ _ 

Ostracodes and the Virginian-Nova Scotian boundary __ 

Nova Scotian and Virginian amphiatlantic species ______ _ 

Nova Scotian and Virginian endemic species ___________ _ 

References cited ___________________ -_-_-------------

ILLUSTRATIONS 

[For ease of identification, all plates in each group are the same color, and each group is a different color. Plates are in pocket] 

PLATES 1-6. Surface- and bottom-temperature maps: 

1. Surface isotherms for August, Nova Scotia to Long Island (from Schroeder, 1966; Emery, 1966). 

Page 

E5 

5 

9 

11 

13 

13 

16 

20 

2. Bottom isotherms for the warmest part of the year (mid-September through October), Nova Scotia to Long 

Island (from Schopf, 1967). 

3. Surface isotherms for February, Nova Scotia to Long Island (from Schroeder, 1966; Emery, 1966). 

4. Bottom isotherms for the coldest part of the year (mid-February through March), Nova Scotia to Long 

Island (from Schopf, 1967). 

5. Mean ice line and surface isotherms, in °C, for the North Atlantic-Arctic region in August (from Defant, 

1961; U.S. Navy Hydrographic Office, 1944, 1958; Schroeder, 1966). 

6. Mean ice line and surface isotherms, in °C, for February (same sources as for pl. 5). 

7-14. Virginian province distribution of nine endemic species restricted or nearly restricted to areas south of Cape Cod and 

the Northeast Channel (blue) : 

7. Aurila aff. A. amygdala (Stephenson). 

8. Actinocythereis aff. A gomillionensis (Howe and Ellis). 

9. Puriana rugipunctata (Ulrich and Bassler) and "Haplocytheridea" bradyi (Stephenson). 

10. Hulingsina spp. 

11. Bensonocythere aff. B. whitei (Swain). 

12. Munseyella atlantica Hazel and Valentine. 

13. Echinocythereis margaritifera (Brady). 

14. Cytherella sp. 

15-25. Nova Scotian province distribution of 12 amphiatlantic cryophilic species restricted to north of Cape Cod and 

Georges Bank, at least at sublittoral depths (green) : 

15. Baffinicythere howei Hazel ( = Cythere costata Brady). 

16. Cytherura? undata Sars. 

17. Cytheropteron latissimum (Nor man). 

18. Elofsonella concinna (Jones). 

19. Muellerina abyssicola (Sars). 

20. Patagonacythere dubia (Brady). 

21. Finmarchinella angulata (Sars). 

22. Cythere lutea Mueller. 

23. Cytheropteron angulatum Brady and Robertson. 

24. Palmenella limicola (Norman) and Eucytheridea bradii (Norman). 

25. Thaerocythere crenulata (Sars). 

26-30. Distribution of five amphiatlantic species occurring in both the Virginian and Nova Scotian provinces (purple): 

26. Baffinicythere emarginata (Sars). 

27. Eucythere declivis (Nor man). 

28. Hemicytherura clathrata (Sars). 

29. Finmarchinella finmarchica (Sars). 

30. Loxoconcha impressa (Baird). 

ni 
IV CONTENTS 

PLATES 31-47. Distribution of 16 amphiatlantic species found in the Nova Scotian to Arctic provincesandoftheechinocythereidine 

genera Rab1'limis and Echinocythereis (red) : 

31. Baffinicythere emarginata (Sars). 

32. Elofsonella concinna (Jones). 

33. Palmenella limicola (Norman). 

34. N ormanicythere leioderma (Norman). 

35. Baffinicythere howei Hazel ( = Cythere costata Brady). 

36. Callistocythere? cluthae (Brady, Crosskey, and Robertson). 

37. Robertsonites tuberculata (Norman). 

38. H emicytherura clathrata (Sars). 

39. H eterocyprideis sorbyana (Jones). 

40. Cytheropteron latissimum (Norman). 

41. Finmarchinella angulata (Sars). 

42. Cytheropteron angulatum Brady and Robertson. 

43. Eucytheridea punctillata (Brady). 

44. Eucytheridea bradii (Nor man). 

45. Cytherura? undata Sars s.l. 

46. H emicythere borealis (Brady). 

47. Rabilimis mirabilis (Brady), R. septentrionalis (Brady), Echinocythereis echinata (Sars) s.l., E. planibasalis 

procteri (Blake), and E. margaritifera (Brady). 

48-.C)4. Distribution of seven subfrigid to cold-, mild-, or warm-temperate amphiatlantic species (ocher): 

48. Thaerocythere crenulata (Sars). 

49. Loxoconcha impressa (Baird). 

50. Cythere lutea Mueller. 

51. H emicythere villosa (Sars). 

52. Patagonacythere dubia (Brady). 

53. Finmarchinella finmarchica (Sars). 

54. Muellerina abyssicola (Sars). 

55-69. Distribution of 14 species endemic to the western Atlantic in the Virginian, Nova Scotian, Labrador, and Arctic 

provinces (turquoise): 

55. Cytherura? mainensis Hazel and Valentine. 

56. Muellerina canadensis (Brady). 

57. Muellerina aff. M. lienenklausi (Ulrich and Bassler). 

58. Leptocythere angusta Blake. 

59. Munseyella mananensis Hazel and Valentine. 

60. Actinocythereis dawsoni (Brady), Cytherura? mainensis Hazel and Valentine, Leptocythere angusta Blake, 

Muellerina canadensis (Brady), Munseyella mananensis Hazel and Valentine. 

61. Pseudocytheretta edwardsi Cushman. 

62. Pterygocythereis americana inexpectata (Blake). 

63. Actinocythereis vineyardensis (Cushman). 

64. Actinocythereis dawsoni Brady). 

65. Bensonocythere arenicola (Cushman). 

66. Bensonocythere americana Hazel. 

67. "Sahnia" spp. 

68. C11shmanidea spp. 

69. Echinocythereis planibasalis procteri (Blake). 

Page 

FIGURE 1. Index map of the Atlantic coast from Long Island, N.Y., to Nova Scotia_______________________________ E2 

2. Map showing location of collecting stations_________________________________________________________ 4 

3. Map showing median grain size of bottom sediment__________________________________________________ 5 

4-8. Charts showing: 

4. Biogeographic schemes used for the Atlantic coast of North America_____________________________ 7 

5. Distribution of ostracodes within the faunal provinces of the western North Atlantic______________ 8 

6. Comparison of nomenclature used for the marine climatic zones of the northern Atlantic coast of North 

America________________________________________________________________________________ 9 

7. Comparison of nomenclature used for western European faunal provinces________________________ 10 

8. Biogeographic range of amphiatlantic species in Europe________________________________________ 12 

9. Diagram showing the four types of temperature zonation of species____________________________________ 12 

10. Graph showing equatorward emergence of Ba.ffinicythere emarginata_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 15 

11. Graphs showin~ depth "preference" of Muellerina canadensis and M. aff. Mlienenklausi__________________ 17 
CONTENTS v 

TABLES 

Page 

TABLE 1. Sample localities and identified taxa, southern Nova Scotian and northern Virginian faunal provinces ____ In pocket 

2. North-south limits, climatic-zone range, zonal type (Hutchins, 1947), temperature range, and depthrangeof 

26 amphiatlantic species______________________________________________________________________ E14 

3. North-south limits, climatic-zone range, temperature-tolerance range, and depth range of 19 species endemic 

to the western Atlantic_______________________________________________________________________ 19 

ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE 

UNITED STATES-OSTRACODE ZOOGEOGRAPHY IN THE 

SOUTHERN NOVA SCOTIAN AND NORTHERN VIRGINIAN 

FAUNAL PROVINCES 1 

By JosEPH E. HAzEL 

ABSTRACT 

Ostracodes were identified from 236 bottom samples taken in 

the region from Nova Scotia to Long Island. There i·s a distinctive difference between the assemblages in the northern and 

southern parts of this region. Many sublittoral cryophilic species 

are not present south of Cape Cod or the Northeast Channel, 

and several thermophilic species a.re not .found north of Cape 

Cod or Georges Bank. The ostracode assemblages in the southern 

part of the cold-temperate Nova Scotian province are a mixture 

of amphiatlantic cryophilic species and endemic, mainly thermophilic species; European forms make up approximately 50 percent of the species. Less than 25 percent of the species known 

from the mild-temperate Virginian province have been reported from European waters. More endemic species pass Cape 

Cod from the south than amphiatlantic cryophilic species do 

from the north. This is consistent with the fact that summer 

isotherms are more compressed than winter isotherms in the 

Cape Cod region, and the southern limit of distribution of most 

amphiatlantic species is regulated by summer high temperatures. The distribution data on which these interpretations are 

based are presented in a series of .figures and over 60 species 

distribution maps. 

INTRODUCTION 

This study is concerned with the geographic distribution of diagnostic, mainly podocopid, ostracode species 

obtained from offshore bottom grab samples taken on 

the Atlantic shelf and slope in the southern Nova 

Scotian and northern Virginian faunal provinces (Long 

Island to Nova Scotia ; fig. 1) . This research was carried out as part of the U.S. Geological Survey-Woods 

Hole Oceanographic Institution program to study the 

marine geology of the Atlantic continental margin of 

the United States (Emery, 1966). 

In this study the geographic spread of the taxa treated 

is of primary importance; temperature tolerances and 

depth ranges are also presented. The relationship of the 

taxa to substrate conditions in the study region (mostly 

1 Contribution 2156 of the Woods Hole Oceanographic Institution 

based ~n 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. 

sand) must a wait completion of sedimentological 

studies now in progress (Schlee and Pratt, 1966; Trumbull and others, 1966). 

The Recent ostracodes of the Atlantic coast of the 

United States are poorly known (Hazel, 1967a), and 

the purpose of this report is to contribute to the establishment of a Recent datum to which Pleistocene and 

older distributions can be compared. It is assumed that 

the collections, supplemented by the available literature, are adequate to delineate the Recent teil provinces-the geographic range of a species during a small 

part of geologic time (Valentine, 1961) -of most of the 

taxa treated. The distribution of the species treated are 

presented in a series of maps, and their tolerances with 

respect to provinces, climatic zones, depth, and temperature are given on several plates. 

ACKNOWLEDGMENTS 

I am indebted to members of the U.S. Geological 

Survey and Woods Hole Oceanographic Institution 

who are assigned to the continental margin program 

for discussions of the results of their studies. W. M. 

Walker and P. C. Valentine of the U.S. Geological 

Survey deserve special thanks for their technical assistance in processing samples and organizing data. T. J. M. 

Schopf of Lehigh University provided unpublished temperature data. I am grateful to A. H. Cheetham of the 

U.S. National Museum and John Pojeta and P. C. 

Valentine of the U.S. Geological Survey for critically 

reading the manuscript. 

PREVIOUS WORK 

Brady ( 1866, 1868b, 1870) , Brady and Norman 

(1889), Norman, (1877), Sars (1909), Stephensen 

( 1913, 1936), Elofson ( 1941), Neale ( 1959, 1961, 1964), 

Hazel (1967a), Hazel and Valentine (1968), Whiteaves 

(1901), and Kindle and Whittaker (1918) have described, listed, or reviewed podocopid ostracode occurEl 
E2 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

•'{t 

\))(90 

1717o 

10 fori, 

FIGURF. 1.-Index map of the Atlantic coast from Long Island, N.Y., to Nova Scotia showing the more important geographic 

localities mentioned in this report. 

rences along the Canadian and western Greenland 

coasts from the Gulf of St. Lawrence to about lat 80° N. 

Cushman (1906), Williams (1907), Blake (1929, 1933), 

Puri (1958a, 1958b), Williams (1966), Hazel (1967a), 

and Hazel and Valentine (1968) have described or listed 

ostracodes from the Gulf of Maine region to Long Island. In comparing the western Atlantic distributions 

of amphiatlantic species to those of the eastern Atlantic, 

the work of Elofson ( 1941) was invaluable; most of the 

eastern Atlantic map points for the species were taken 

from Elofson's maps and text or from references given 

therein. Elofson (1943), Wagner (1957), Vos (1957), 

Lange ( 1956), Neale ( 1959, 1964), and Hagerman 

( 1965) were useful later references. 

In the first \vork in the region of this study, Cushman 

(1906) described or listed 26 species from the Woods 

Hole, Mass., area. Most of the species were at Fishhawk 

station 7723 (see Sumner and others, 1913, for detailed 

localities of the stations) in Vineyard Sound at 24 

meters. Unfortunately, Cushman's specimens are not 

available for restudy ( Puri, 1958a) . This is particularly 

troublesome because it is unlikely that some of the species Cushman reported as occurring in Vineyard Smmd 

actually live in the area. In his study of the genus Oushmanidea, Puri (1958a) had Fishhawk 7723 and other 

stations reoccupied and, with the resulting material, 

revised the species Cushman assigned to Oytheridea. 

Through the courtesy of Dr. Puri, I was able to examine 

his faunal slide from Fishha wk 7723. The following 

is a list of the species present (Neocytherideidinae 

from Puri, 1958a) and the names Cushman seems to 

have used for them: 

Reoccupied Fishhawk station 7723 Cushmom, (1906) 

Pseudocytheretta edwardsi______ Pseudocytheretta edwardsi. 

JJI uellerina canadensis (Brady)_ Oythereis oanadensis. 

aft'. Jf. lienenklausi 

Ulrich and Bassler) 

Bensonocythere arenicola (Cushman) 

a:I:I. B whitei (Swain) 

arenicola. 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E3 

Reoccupied Fishhawk station 77'23 

Aurila aff. A. amygdala 

( Stephenson) 

H emicythere villosa ( Sars) ____ _ 

Finmarchinella finmarchica 

(Sars) 

Leptocythere angusta Blake 

Oyprideis SP-------------------

Actinocythereis vineyardensis 

(Cushman) -----------------

dawsoni (Brady)----------

aft. A. gomillionensis 

(Howe and E'llis) 

Puriana ntgipunctata 

(Ulrich .and Bassler) 

Cushman (1906) 

Oythereis albomaculata 

(Baird). 

villosa. 

?Oytheridea punctillata 

Brady. 

Oythereis vineyardensis. 

Oytherc dawsoni Brady. 

Oushmanidea elongata (Brady)_ Oytheridea rubra Mueller. 

seminuda (Oushman) _______ seminuda. 

Hulingsina americana americana. 

(Cushman) 

In addition to the a;bove taxa from the Vineyard 

Sound area, Cushman also reported several other species which are treated in the present study, that is Baffinicythere errnarginata, Loxoconcha impressa, Etwytheridea bradii ( = Oytheridea papillosa), Elo fsonella 

·concinna, and Robertsonites tuberculata. Baffinicythere 

emarginata and Loxoconcha irnpressa have been identifled in my samples from the general area, thus confirming Cushman's identification. Cush~an's Oytheridea 

papillo8a is a [{ rithe ( Puri, 1958a). It is questionaJble if 

Cushman actually found Elofsonella concinna and Robertsonites. tuberculata/ they do not occur in any of my 

samples from the Vineyard Sound area and have not 

been reported south of the Gulf of Maine. For the time 

being, Cushman's report of these taxa is considered to 

be based on misidentifications. 

PRESENT WORK 

Work on samples collected by U.S. Bureau of Commercial Fisheries ships and the RV Gosnold in connection with the U.S. Geological Survey-Woods Hole 

Oceanographic Institution program has added 236 more 

localities to the list of stations where ostracodes have 

been found in the Gulf of Maine-Georges Bank-Cape 

Cod region. For comparative purposes 25 new Arctic 

stations were also added in this study. These samples 

were collected by Captain R. A. Bartlett in the 1930's 

and 1940's and were used by Loeblich and Tappan 

( 1953) in a study of the Arctic Foraminiferida. All the 

new stations and the identified ostracodes are listed on 

table 1. 

Basically, the Nova Scotia to Long Island region was 

sampled on a 10-mile grid (Hathaway, 1966). About 

600 samples, mostly from depths greater than 20 m, 

were ·processed, and ostracodes were found in 236 sampies (fig. 2). Large areas in the central Gulf of Maine, 

middle part of Georges Bank, and on Nantucket Shoals 

were virtually devoid of ostracodes. Ostracodes occur 

principally on the outer parts of the shelf south of New 

England, outer parts of George Bank, Browns Bank, 

southwest of Nova Scotia, southern Bay of Fundy, and 

northern and western parts of the Gulf of Maine. 

Over 100 species have been identified to the present 

time; ho·wever, many of these are rare and (or) have 

not yet been treated taxonomically or are in need of 

further taxonomic treatment. In this study, over 40 taxa 

are used which are considered to be important, that is, 

species which are useful in delineating the Virginian 

province from the Nova Scotian province, or species 

which occur commonly over the entire region and are 

therefore diagnostic for the region as a whole. 

The samples used in the study, the amount of material 

processed, and the number of specimens of each species 

are given in table 1. Living and dead specimens were 

counted together. In most samples containing many 

ostracodes, specimens with appendages or at least some 

chitin remaining were observed for most species. In 

many samples, however, only dead specimens were observed, but living specimens were frequently found in 

nearby samples and the maps presented herein are 

considered to depict adequately the areas where the 

species live. 

WATER TEMPERATURES FROM NOVA SCOTIA 

TO LONG ISLAND 

Water temperatures in the region (Fritz, 1965; 

Schroeder, 1963, 1966; Emery, 1966; Schopf, 1967; 

Hicks, 1959) range from a minimum of less than 2°C 

on the Scotia Shelf to a maximum of over 20°C in inshores areas such as Narragansett Bay (pl. 1). The average surface temperature (pl. 1) for the warmest month 

in the Gulf of Maine__:Scotia Shelf-Browns Bank area 

ranges from l2°C in the northern Gulf of Maine and 

lower Bay of Fundy to about l8°C in Cape Cod Bay. 

It is l2°C at the surface over most of the Scotia Shelf 

and averages as high as l5°C over Browns Bank. Average bottom temperatures (pl. 2) in the warmer months 

are usually between 6° and 7°C on Browns Bank and 

on the Scotia Shelf; however, temperatures as high as 

l0°C on the Scotia Shelf and 8°C on Browns Bank have 

been recorded (Fritz, 1965). Bottom temperature 

maps for depths less than 100 m are not available for 

the western Gulf of Maine, but temperatures are between 5° and 7°C in the warmest months at 100 m 

(Schopf, 1967) and, if the end point of a species range 

was found to be in this area and at depths less than 100 

m, the temperature was interpolated between the surface and 100m. Bottom temperatures in summer and 
E4 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

FIGURE 2.-Location of samples used in the Long Island to Nova Scotia area ( Q, indicates ostracodes ab8elllt; e, indicates 

ostracodes present). See table 1 for the localtion of the numbered ostracode-bearing sampl~s. See Hathaway (1966) for 

additiona'l information abourt these and other stations\ used in the U.S. Geological Survey-Woods Hole Oceanographic 

Institution program. 

winter are about 6°C in the Northeast Channel and 

deep basins of the Gulf of Maine. 

In the coldest months the average surface temperature in the Gulf of Maine region north of Georges Bank 

is 3 °-4 ° C (pl. 3) . Average bottom tern peratures on 

Browns Bank are between 3° and 5°C and on the Scotia 

Shelf between 2° and 4°C. In the western Gulf of Maine 

bottom temperatures are about 4°C (pl. 4). 

The average surface temperature over Georges Bank 

in the warmest month is about 16°C and increases southwestward to 22°C south of Long Island. The maximum 

bottom temperature on Georges Bank is about 16°C 

and the average bottom temperature is 15°C on the 

shallow central part in the warmest months. To the 

south, east, and north, this average decreases to about 

7°C between depths of 100 and 200m. Another center 

of offshore high temperature is at middle shelf depths 

south of Martha's Vineyard where the temperature 

averages 14°0 in the warmer months. South of Long 

Island the temperature averages 14°-15°C at 30m. In 

some years temperatures in waters under 100 m may 

average 9°-10°C on the Scotia Shelf during this period 

(Fritz, 1965), and the region south of Long Island is 

warmest in November, averaging 14°-15°0 at 30 m 

(Schopf, 1967; Fritz, 1965) . At 200 m along Browns 

Bank the yearly temperature average is about 9°C with 

about a 5°0 observed range, and at 200 m south of 

Northeast Channel the yearly average is .about 10.5°C 

with an observed range of 5°C (Schroeder, 1963). 

In the coldest months the average bottom temperature 

on Georges Bank is 4°0; in the area south of Martha's 

Vineyard, 3 °C; and at depths of 100 m, from eastern 

Georges Bank to south of Long Island, 6°-7°C. At 

depths of 200 m, on the shelf edge south and east of the 

Cape Cod area, the yearly average is between 10° and 

11 oc with an observed range of 5 oc, and on the Atlantic 

side of Browns Bank the yearly average is about 9°0, 

also with a 5°0 range. 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E5 

SUBSTRATE FROM NOVA SCOTIA 

TO LONG ISLAND 

Studies of the sediments in the region are in progress, 

and preliminary reports have appeared (Schlee and 

Pratt, 1966; Trumbull and others, 1966; Hiilsemann, 

1967) . For a summary of the sedimentologic studies 

prior to the initiation of the U.S. Geological SurveyWoods Hole Oceanographic Institution program, see 

Uchupi (1963). Over much of the region (fig. 3) where 

ostracodes were found, the substrate is mainly quartzose 

sand. Large patches of gravel are present in several 

areas, particularly Georges Bank, Nantucket Shoals, 

Northeast Channel, Browns Bank, and the Scotia Shelf. 

The deep basins in the Gulf of Maine are floored mainly 

with clay and silt; the western Gulf of Maine is largely 

silt, and a large area of silt-sized material is present 

on the shelf south of Martha's Vineyard. 

HISTORICAL DEVELOPMENT OF 

FAUNAL PROVINCES 

NORTH AMERICA 

The Atlantic coast of North America was divided 

into biogeographic provinces as early at 1838 by MilneEdwards. Working with Crustacea, he divided the coast 

into a polar regioi1 (Newfoundland northward), a 

Pennsylvanian region (Newfoundland to the Carolinas), and a Caribbean region (Carolinas southward). 

In classic works, Dana ( 1853a, b) proposed a worldwide scheme of biogeographic kingdoms and provinces 

and climatic zones. The provinces were named after 

places, and the names of climatic zones were formed 

from the adjectives torrid, temperate, and frigid, used in 

the ancient Greek classification. From a nomenclatorial 

standpoint Dana's scheme for provinces and climatic 

zones is in many ways preferable to that used by some 

25 0 25 50 75 100 KILOMETERS 1""1 I I I I 

PT H CONTOURS IN METER 

FIGURE 3.-Median grain size of bottom sediment (from Hiilsemann, 1967). 
E6 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

authors today. It avoids such confusing terminology as 

"Boreal" and "Arctic" which have been used both for 

provinces and climatic zones. Dana's concept of biogeographic provinces, however, differs from that generally accepted by modern biogeographers (for example, 

see Valentine, 1961). Dana seems to have first set up 

his global scheme of climatic zones based on isocrymes 

at 6°F intervals (except for the lowest isocryme which 

was 9°F below the penultimate). The coasts and islands 

of the world therefore were first placed in a globeencircling climatic zone, the coastal areas named, and 

then the crustacean assemblages found in the provinces 

discussed. Thus, Dana's provinces seem to be based on 

temperature and not on the distribution of animals. 

Dana (1853a, p. 1488) stated, however, that one of the 

objects of his study was "to compare different geographical positions in similar regions with one another, in 

order to arrive at their resemblances and differences, 

and deduce the several distinct zoological provinces * * *." He may have studied the assemblages 

found in the temperature-based climatic zones, judged 

that the zones did contain distinct assemblages in different areas, and then accepted them as provinces. 

In most other works marine climate zones are not 

really based on temperature. Temperature is the tmderlying factor controlling the distribution of organisms, 

but it is the distribution of kinds of organisms that 

determines the provinces, and it is the boundaries between provinces upon which the climatic zone boundaries have been based. 

The fact that temperature is the basic factor controlling the distribution of organisms should make an objectiv-e scheme based solely on temperature similar to 

that based on province boundaries. Compare Dana's 

(1853a) chart showing his climatic zones based on 

isocrymes to Hall's (1964) map showing marine climates "based on the duration of marine temperatures at 

the limits of molluscan species." Along the east and 

west coasts of North America, Dana's boundaries almost coincide with Hall's, although the nomenclature 

is different. It might be rtdded here that Hall (1964, p. 

232) chose Edward Forbes' (in Johnson, 1856) province 

terminology "because of priority." However, many of 

Dana's and Milne-Edward's terms arc different from 

and predate those of Forbes. 

Along the Atlantic coast, Dana ( 1853a) recognized 

a Floridian province f,rom Key West to lat. 27°30' N., 

a Carolinian province from there to Cape Hatteras, a 

Virginian province from Cape Hatteras to Cape Cod, 

and a Nova Scotian province from there to Cape Race, 

Newfoundland (fig. 4). Without comment, Dana 

(1853b) changed the name Nova Scotian to Acadian. 

'rhe Floridian province was considered to be in the subtorrid region, the Carolinian in the warm-temperate 

region, the Virginian in the cold-temperate region, and 

the Nova Scotian in the subfrigid region. It is interesting to note that Dana recognized five temperate regions- warm-temperate, temperate, subtemperate, coldtemperate, and subfrigid- and that the temperate and 

subtemperate regions were not considered to be present 

on the American coast. 

Dana considered the coasts north of Cape Race, Newfoundland, to belong to his Arctic kingdom (north 

frigid region) but applied no provincial terminology to 

this region. He ( 1853b) divided his Arctic kingdom 

into the north pola.r, Camtschatican, and Norwegian 

provinces, but did not mention the North American 

coast north of N ewfotmdl and. 

'Voodward (1851- 56) first used the term Transatlantic province for the Atlantic coast of the United States 

from some undescribed point in Florida to Cape Cod; 

the Gulf of Maine- Nova Scotia region was placed in 

the Boreal province; and the shores of the Gulf of St. 

Lawrence and Newfoundland and regions to the north 

were placed in the Arctic province. 

Woodward (1851-56, p. 379) stated that Forbes considered the Atlantic coast of the United States to consist 

of two provinces, the Virginian and the Carolinian. He 

was apparently referring to Forbes' plate 31, a map of 

the distribution of marine life, and the accompanying 

discussion in ,Johnson (1856). Here Forbes used Carolinian province for the region from about the GeorgiaFlorida State line to Cape Hatteras, Virginian province 

for the region from Cape Hatteras to Long Island, 

Bostonian province from Long Island to Cape Race, 

Newfoundland, and Arctic province north of this and 

apparently including the Gulf of St. Lawrence. From 

the literature availabl-e to me it seems that the first use 

of Carolinian and Virginian should be attributed to 

Dana ( 1853a), although Forbes did not mention Dana's 

use of the terms. 

Packard (1863, 1867) proposed that on the American 

coast there exists a characteristic fauna between the 

true circumpolar Arctic province and the Nova Scotian 

province. He stated that this fauna bears the same relationship to theN ova Scotian ·as the fauna of the Finnmark region of Norway does to the North Sea and 

Scottish seas. Packard called this fauna th-e Syrtensian 

or Labrador fauna. The name Labrador is preferable 

though Syrtensian has priority, because all the provinces along the American coast would have a geographic 

term as the root word. Packard considered the Labrador 

province to include the coast of Labrador, Hudson's 

Bay, the northern coast of Newfoundland, and the 

northern ;parts of the Gulf of St. Lawrence, with "outliers" to the south in the region of Dana's Nova Scotian 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E7 

Packard Stephenson Milne- Forbes (1863, oc 

N. Edwards Dana Woodward 1867) and Coomans Hall This kin Johnson lat. (1853a, b) (1851-56) Stephenson (1962) (1964) paper (1838) 1856) and Ganong Feb. Aug. (1954) (1890) 

-

-

-

- 80°- Arctic -

- Arctic -

- Arctic 

70°-

-

-

-

- Polar Arctic Arctic Arctic Arctic Arctic 

~ f- oo 

600- r-------

- Labrador 

- Subarctic 

- f----?---- or ~50 

- Labrador Syrtensian f- oo f---100 or 50°-- Syrtensian 

- Nova Nova Nova - Scotian Bostonian Boreal Acadian Acadian Scotian Scotian r-15° - Pennsyl- Boreal f---20° 40°- van ian 

- Virginian Virginian Virginian "Overlap" Virginian Virginian f- 50 f---25° - 10° - Trans-

- Carolinian atlantic CaroHnian Carolinian Carotin ian 

30°- Caribbean 

Carolinian Not Not f-15° 

treated treated 1---20° - Caribbean ! Caribbean f--30° - Floridan "Tropical" Caribbean 1-25° 

FIGURE 4.-Summary of the biogeographie schemes used by various workers on the Atlantic C{)ast of North America plotted 

against latitude. Average sUJrface-water temperatures fur August and }"'ebruary given in 5°0 intervals. 

province. Ganong ( 1890, p. 170, 171) stated that it is 

uncertain how far south on the northeast coast of 

Labrador the circumpolar Arctic fauna extends, "but 

certainly it does not on the surface reach the Straits of 

Bell Isle * * * ." He considered the Syrtensian province 

to include the southern part of Greenland, all the 

northern part of the Gulf of St. Lawrence as far west 

as Anticosti Island, and the deeper part of the of the 

gulf north of a line drawn from Gaspe to the Magdalen Islands and thence to the north of Cape Breton. 

All the Newfoundland coast, except possibly a part 

of the southern shore, is included. Ganong also included 

some areas (Packard's "outliers") farther south in the 

Syrtensian that Dana and most modern workers would 

include in theN ova Scotian province. 

Johnson (1934) stated that the Arctic province is 

bounded on the south by floating ice which descends as 

far south as Newfoundland. The Boreal province was 

said to en com pass the region from northern Nova 

Scotia to Cape Cod, and the transatlantic province extends from the southern shore of Cape Cod to the 

vicinity of Cape l{ennedy. The northern limit of the 

Caribbean province was placed at Jupiter Inlet or perhaps Cape l{ennedy. 

In 1954, Stephenson and Stephenson summarized 

their thoughts on the provinces of the North American coast after study of the intertidal fauna and flora 

frmn Florida to Labrador. They stated that they are 

in general agreement with the older workers on the 

biogeographic provinces but introduce some revisions 

based on new data. The Stephensons recognized an 

Arctic province "* * * whose southern limits cannot 

yet be defined, but which probably lies north of Labrador." Their subarctic province ( = Packard's Labrador 

or Syliensian province) occupies Labrador and southern Greenland, much of Hudson Bay, and the northern parts of Newfoundland and the northern Gulf of 

St. Lawrence. They stated that further work is needed 

in these regions in order to determine how distinct the 

Labrador province may be. The northern limits of 

ostracodes (fig. 5) and many other Crustacea (Stephensen, 1936; Dunbar, 1964; Squires, 1966) suggest that 

the northern boundary of the Labrador province is at 

about Disko Bugt in western Greenland and at the 

southeastern Baffin Island in eastern Canada. 
E8 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

Climatic 

zone 

Biogeographic 

province 

FIGURE 5.-Generalized diagram showing the distribution of 

ostracodes within the biogeographic provinces of the western 

~orth Atlantic. The northern limits of several species of ostracodes (for example, Patagonaoythere dttbia, Thaerocythere 

crenulata, H emicythere villosa, Oythere lutea, and perhaps 

lfluellerina abyssioola) and other Crustacea ( Stephensen, 

1936; Dunbar, 1964; Squires, 1966) suggest that the ArcticLabrador provincial boundary falls at about Disko Bugt in 

Greenland and southeastern Baffin Island in Canada. The 

boundary between the Labrador and Nova Scotian provinces 

is less well marked by the ostracodes, but this is probably the 

result of incomplete collections as the ostracodes have not 

been described from the Atlantic coast of Newfoundland and 

are known from the Gulf of St. Lawrence almost entirely from 

The Stephensons' cold-temperate or Acadian province is virtually the same as Dana drew it (1853a), excepting the modifications of Packard (1867), Ganong 

( 1890), and the Stephensons in the Gulf of St. Lawrence. They preferred, however, not to recognize the 

Virginian province because it does not seem to possess 

"* * * a distinctive population of its own, but to represent an overlap !between the cold-temperate population of Cape Cod and the warm-temperate one south of 

Cape Hatteras." 

The Stephensons recognized a warm-temperate 

fauna, the Carolinan province, from Cape Hatteras 

to Cape l{ennedy (lat 28°30' N.) and suggested 

that no marked subtropical province seems to be extant. They considered the tropical fauna to begin about 

at Jupiter Inlet (lat27o N.). 

Coomans (1962) took a different view of the Virginian province. He stated that only 10.5 percent of the 

Virginian mollusks are endemic to the province, 

whereas "it is agreed that this should be at least 50 percent for an autonomous zoogeographical province." 

Therefore, the Virginian cannot stand as a province. 

Species Boundary 

Disko Bugt 

S. E. Baffin 

Island 

Gulf of St. 

Lawrence 

Cape Cod 

Northeast 

Channel 

Cape 

Hatteras 

the work of Brady (1870). Oytherura? mainensis, Actinocytheresis dawsoni, and Muellerina canadensis are not known 

north of the Gulf of St. Lawrence, and Leptocythere angusta 

and Loxoconcha impressa extend only to southeastern Labrador. The Nova Scotian-Virginian boundary is well marked by 

the termination of a large number of equatorward-expanding 

cryophillic and poleward-expanding thermophilic species. 

Study of collections from the area of the Virginian-Carolinian 

boundary is incomplete ; however, several species seem not to 

extend south of the Cape Hatteras area, ·and several warmtemperate or tropical and subtropical genera, such as 

Orionina, Neocaudites, Oaudites, and Oytherelloidea, do not 

seem to occur north of Gape Hatteras. 

The marine climatic-zone nomenclature is modified from Dana 

(1853a). 

Because 129 or 62 percent of the Virginian mollusks 

are arctic or boreal species, Coomans suggested extending the Boreal province ( Coomans used the term 

"Boreal" for both the European and North American 

cold-temperate faunas) south to Cape Hatteras. He con-. 

sidered the Arctic province to extend south to Cape 

Race, Newfoundland, and the southern boundary of 

the Carolinian province was placed at Cape Kennedy. 

Southern Florida was placed in the Caribbean province. 

In his analysis of molluskan distributions, Hall 

( 1964) suggested that the number of consecutive days 

or Inonths when the water is at temperatures required 

for reproduction and early growth determines the limits 

of marine shallow-water provinces and, therefore, climatic zones. On this basis the Atlantic coast of North 

America is divided into five provinces-an Arctic province that extends south to about lat 47° N. (southern 

Newfoundland), the Nova Scotian from there to Cape 

Cod, the Virginian from Cape Cod to Cape Hatteras, 

the Carolinian from Cape Hatteras to lat 30° N. (near 

St. Augustine, Fla.), and the Caribbean from there 

south. In terms of climatic zones, the Arctic was desig-
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E9 

nated "cold"; the Nova Scotian, "cold temperate"; the 

Virginian, "mild temperate"; the Carolinian, "outer 

tropical"; and the Caribbean, "inner tropical." Hall 

considered the warm-temperate zone to be rubsent on 

the North american east coast (see Dana, 1853a). 

Figure 4 shows that province boundaries at southern Newfoundland (lat 47° N.), Cape Cod (lat 42° N.), 

and Cape Hatteras (lat 35° N.) have been generally 

accepted. The position of the southern boundary of the 

Carolinian province varies from one author to another, 

and most workers have not considered it necessary to 

recognize a Syrtensian or Labrador province between 

southern Newfoundland and the circumpolar Arctic 

province. 

The above summary does not purport to be a complete review of the history of faunal provinces along 

the North Ameriqan Atlantic coast. It only attempts to 

show the systems used by some workers in the last 

century and offers a comparison with the schemes used 

by some current workers. 

In contrast to the rather stable nomenclature for the 

provinces, at least from Newfoundland to some point 

in Florida, the climatic zone terminology used for the 

Atlantic North American provinces has been quite variable. Not all authors who have treated the whole coast 

have presented opinions. Figure 6 sho,vs a comparison 

of the terminology used by four authors. Dana ( 1853a), 

whose climatic zones are based on isocrymes, called the 

east coast sub frigid region (sub frigid being the coldest 

subdivision of his temperate regions) the Nova Scotian 

province. Stephenson and Stephenson (1954), following the scheme of Stephenson (1947), placed the Nova 

Scotian fauna in the cold-temperate zone. Hedgpeth 

(1957) used Boreal for the Nova Scotian on his map of 

the littoral provinces of the world, which is more in 

the nature of a map of faunas occurring in particular 

climatic zones than a map of provinces. Hall (1964) 

used cool-temperate for the Nova Scotian. Dana considered the middle two of his five subdivisions of the 

temperate region to be absent on the American coast 

(see p. E6) because of pinching out at Cape Hatteras. Hall stated that the warm-temperate zone is 

absent on the American coast, with mild-temperate 

assemblages meeting outer-tropical assemblages at Cape 

Hatteras. 

Because these authors apply different terms to the 

same climates and most agree as to the placement of 

the major biogeographic boundaries, the different 

usage of climatic terms is basically a confusing battle 

of words. 

The marine climatic zone terminology of Dana 

(1853a) is a modification of the ancient Greek terminology, has priority as a worldwide scheme, and is the 

~tephenso11 

N. Dana and Hedgpeth Hall This 

lat. (1853 a, b) lstephensol1 (1957) (1964) paper 

(1954) 

80"'-

Arctic Frigid 

70°-

Frigid Arctic Cold 

~ 

60"'- 1-------

Subfrigid 

Subarctic 

so--

SubfrigkS Cold "Boreal" Cool Cold 

temperate temperate temperate 

4<r'- Cold "lnterme- Mild Mild 

temperate diate" Warm temperate temperate 

Warm temperate Outer 

Warm 

temperate temperate tropical Not 130"- treated ., !------- Inner 

Sub torrid Tropical Tropical tropical 

FIGURE 6.-Comparison of nomenclature used by some authors 

for the marine climatic zones of the northern Atlantic coast 

of North America. 

most consistent as far as separating climatic terms from 

the geographic terms normally used for provinces. The 

Arctic and the Tropics are, after all, defined as places 

lying between certain parallels of latitude. The word 

arctic is suitable for a province because it is a place and 

at least the high-Arctic fauna tends to be circumpolar. 

The term "tropic," however, presents problems; it denotes a place but has usually been used in a climatic 

sense. In contrast to "arctic" it is not suitable as a province term because1 with some rare exceptions, the 

shallow-marine species found in the Tropics off the 

various continents are not identical. "Torrid" has 

priority for marine climatic zone usage, but tropic is so 

entrenched in the literature that it probably will continue to be used as a climatic zone term. 

Dana's fivefold division of his temperate regions is 

probably excessive for general usage; a threefold 

division that includes cold temperature and warm temperate, which are in general usage and Hall's (1964) 

intermediate mild-temperate seems satisfactory. 

EUROPE 

Various schemes have been proposed for western 

European biogeographic provinces. Some of these are 

shown in figure 7. The Arctic circumpolar fauna seems 
ElO ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

to begin about at N ovaya Zemlya and includes Franz 

,Josef Land and most of Spitsbergen. The region from 

the Lofoten Islands, or perhaps North Cape, Norway, 

to N ovaya Zemlya probably represents a transition 

region (see Ekman, 1953, p. 101; Packard, 1863; Hedgpeth, 1957) that probably also includes Bear Island, 

northern Iceland, and perhaps southwestern Spitsbergen and is equivalent to the A1nerican-West Greenland Labrador province. The term "boreal" has been 

considerably misused and does not have a geographic 

root. The term "Norwegian" (proposed by Dana, 1853b) 

or "Scandinavian" (proposed by Milne-Edwards, 1838) 

is probably better nomenclature for the faunas of the 

N. Milne-Edwards Dana Forbes 

Norwegian coast from at least the Lofoten Islands to 

and including the Skagerrak and the Shetland Islands. 

The Celtic province (proposed by Milne-Edwards, 

1838) has been used by most authors for the faunas of 

the British Isles and the North Sea coast of Europe; 

its southern boundary has been placed either at the 

entrance to the English Channel (Forbes, in Johnson, 

1856; Ekman, 1953; Hall, 1964) or, less likely, near 

Cape Finisterre (Dana, 1853a; Coomans, 1962). Dana 

(1853a, b) used the term "Caledonian" for the faunas 

of the northern part of the Celtic province. Ekman 

( 1953) placed the faunas found from the southern 

entrance to the English Channel to northern Norway in 

the Boreal province. 

Woodward Ekman Coomans Hall 

lat. (1838) (1853a, b) (in Johnson, (1851-56) (1953) (1962) (1964) 1856) 

75°-

Polar Arctic Arctic Arctic Arctic 

Arctic 

Arctic ------------ ---

70°-

Norwegian 

65°- Boreal Scandinavian Boreal 

Boreal 

Norwegian 

------ / 

/ 

600- Boreal / 

Caledonian / 

/ 

/ 

/ 

------ v/ 

55°-

Ceftic 

Celtic Celtic Celtic 

Celtic 

50°- Ceftic 

Lusitanian 

45°-

Lusitanian Lusitanian Lusitanian 

Lusitanian Lusitanian 

Mediterranean Mediterranean Mauretanian 

Moroccan 

FIGURE 7.-Compari.soo of nomenclature used for western European faunal provinces by various authors. 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES Ell 

The difficulty in assigning province boundaries in 

western Europe as compared to North Ameriea stems 

in part from the different hydrographic situations. The 

clockwise circulation of waters in the North Atlantic 

sends the warm waters of the Gulf Stream towards 

Europe and mitigates severe winters. The temperate climatic zones are much wider on the European coast than 

on the North American coast, where the warm waters 

of the Gulf Stream flow northward and the Labrador 

Current flows southward. This 'vestern Atlantic water 

circulation is coupled with the monsoon effect of general 

offshore winds in winter and onshore winds in summer 

(see Fle.ming, 1957). This makes for cold winters and 

hot summers and causes convergence of winter and summer isotherms at several points, which establishes strong 

temperature barriers for benthonic organisms. The absence of these strong isotherm compressions in western 

Europe allows species to disperse more widely, and the 

province bounda:ries are therefore conside~rably more 

gradational. 

Correlation of European with North American provinces is difficult b~ause of this hydrographic difference. 

In the Gulf of Maine region, the change in August surface-water temperatures from 12° to 19°C is accomplished in only 270 miles (approximately from northern 

coastal Maine to Cape Cod). Many cryophilic organisms 

which are expanding their range southward and are 

controlled by summer survival temperatures in this 

range will drop out in a rather short geographic distance. In Europe during the warmest month, it is 2,000 

miles between the 12° and 19°C isotherms, that is, from 

northwest Norway to the Bay of Biscay. Figure 5 shows 

the biogeographic range of the amphiatlantic and 

endemic species in North America, and figure 8 gives the 

range of the amphiatlantic species in Europe. 

The shallow waters off Cape Cod in February average about 4°C; however, it is also 4°C 300 miles to the 

south. Average water temperatures for the coldest 

month do not reach as low as 4°C off the British Isles. 

This temperature is reached off the Low Countries, and 

most of the coastal waters of Norway are ·about 4°C. 

Thus, thermophilic shallow-water species that are possibly prevented from passing Cape Cod by the 3 °C 

winter temperatures along the Massachusetts coast 

north of the Cape could live far north of the British 

Isles, if they occur in European waters. Cold-water 

species that live in the Gulf of l\tiaine and are prevented 

from ·passing the Cape Cod area by the high summer 

temperatures on Georges Bank and in the Cape Cod 

Bay area, could extend to southern Ireland and England, if they live in European waters. Therefore, species 

that would not live sympatrically in North American 

waters could he found together in European waters. 

37,8-564 0-!70-2 

In contrast, other cryophilic species that are controlled in their southern expansion by winter temperatures (that is, temperatures must be low enough for 

reproduction and repopulation to take place) may extend far down the American coast past Cape Cod if the 

critical temperature is 4°-5°C. The same species in 

Europe should not be found in the British Isles south of 

at least the Shetland Islands or the deep and cold lochs 

of Scotland. 

Thermophilic species expanding to the north, whose 

northern limits are controlled off North Ame~riea by the 

compression of srnnmer isotherms at Cape Cod, would 

not, if they lived in Europe, reach as·far north as the 

British Isles and would probably drop out at about Cape 

Finisterre or the Bay of Biscay. Thus, cryophilic species 

controlled equa.torward by winter low temperatures and 

warm-water species controlled poleward by summer 

high temperatures can live together on the American 

coast, but theoretically would be allopatric on the coast 

of western Europe. 

The last is a difficult theory to test with ostracodes 

simply because all the thermophilic ostracode species 

found only south of Cape Cod seem to be endemic to the 

coast of the United States. Many of the cryophilic, 

summer-controlled ostracode species which do not pass 

Cape Cod extend to southern Ireland and northern England and Europe, whereas a winter-controlled species, 

such as Baffinicythere emarginata, does not pass Scotland. This suggests that the Virginian-Nova Scotian 

boundary cannot be "correlated" faunally with any of 

the European temperate province boundaries because, 

depending on which types of species chosen, it could be 

the Scandinavian-Caledonian or the Caledonian-Celtic 

boundary. The bulk of the cryophilic ostracodes are 

controlled in their southern extent by summer survival 

temperatures, and most of those that stop before Cape 

Cod extend to southwestern Ireland and northern 

England or Scotland. 

TEMPERATURE CONTROL OF 

SPECIES DISTRIBUTION 

Ifutchins (1947) proposed that species were controlled 

by temperature basically in two ways, survival temperatures and temperatures needed for reproduction and 

repopulation. To be sure, the duration of certain temperatures (Hedgpeth, 1957; I-Iall, 1964) is of importance to some if not 1nost species. However, species 

extend north and south to points where certain maximum or 1ninimum temperatures are reached. It is these 

temperatures, usually in August or September and February, that are most critical, at least for the assignment 

of temperature limits. 
E12 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

Climatic Biogeographic 

zone province 

Frigid 

Subfrigid 

Cold 

temperate 

... 

Mild 

temperate 

Warm 

temperate 

Arctic 

"Transitional" 

Norwegian 

Caledonian 

Celtic 

Lusitanian 

I 

I 

I 

i 

I 

Species Boundary 

W. Spitsbergen 

Novaya Zemlya 

N. Norway 

Iceland 

Shetland Is. 

Skaggerak 

S.W. Ireland 

N. England 

Channel 

Islands 

FIGURE B.-Biogeographic range of amphiatlantic species in Europe. For comparative purposes the arrangement of species 

is the same as in figure 5. 

Hutchins (1947) showed that there were four zonal 

types of species (fig. 9) : ( 1) species which are controlled 

poleward and equatorward by killing winter and summer temperatures, (2) those which are controlled by 

Poleward 

Summer 

Repopulation 

l 1/1 

Winter 

Survival 

Equatorward L-S_u_rv_iv_a_I ____________ R_e::__po_p:_u_l_a_ti_o__Jn 

FIGURE 9.-Diagram showing the four types of temperature 

zonation of species. From Hutchins (1947). See text for 

discussion. 

reproductive repopulation temperatures poleward 

(summer) and equatorward (winter) , ( 3) those which 

are controlled equatorwa,rd by summer survival temperatures and poleward by summer reproductive temperatures, and ( 4) species which are controlled 

poleward by winter survival temperatures and equatorward by winter reproductive temperatures. 

He considered all species which extend into the Arctic, 

where the water is of uniform coldness, Type 2 or Type 

3 species, reasoning that if these taxa can tolerate continued cold conditions, then the only temperature thrut 

could be critical poleward would be in summer. 

Because there are both maximum and minimum temperatures at the end points of ,a species range, it is not 

obvious which of the temperatures is controlling a particular species. Hutchins suggested that if a species occurs on more than one coast and if its distribution is 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E13 

well known, the control1ing temperatures can be delineated because the maximum and minimum temperatures at a point on one coast will not occur together on 

another. A large number of the ostracode species found 

on the American coast at least as far south as the Cape 

Code area also occur in Europe and, if they are well 

known in both regions, the zonal types they represent 

can be determined by this method. lTsing Hutchins' 

principles of correlrution, the zonal types represented by 

these species have been determined as closely as possible. 

For determination of the zonal type of sublittoral species, exact temperature tolerances are not usually necessary because the maximum and minimum temperatures 

occurring at a point on the American coast are usually 

geographically far apart on the European coast, and 

surface temperatures often are sufficient. For accurate 

determination of temperature tolerances, data on exact 

maximum and minimum bottom temperatures, at least 

in the area of the end points of a species range, are required but are often not available. On the basis of temperature data presented by Elofson ( 1941) for several 

of the taxa treated and data gathered in the present 

study, the tolerances for the amphiatlantic species have 

been determined as closely as possible. The zonal types 

represented by the amphiatlantic species (mostly Type 

3), their temperature tolerances, and their depth range 

are given in table 2 (see section under Nova Scotian and 

Virginian amphiatlantic species, p. E13-E16, for 

further discussion; also see pis. 31-54 which are distribution maps of the amphiatlantic species from which 

the zonal types were determined) . 

For endemic species the determination of controlling 

temperatures is more difficult. However, along coasts 

where there is strong compression of isotherms in summer or winter at certain points, the controlling temperature can sometimes be inferred. If a species ceases its 

expansion, for example, at a point where winter isotherms are compressed and summer isotherms are not, 

it is likely that the species is controlled by winter 

temperatures. 

OSTRACODES AND THE VIRGINIAN-NOVA 

SCOTIAN BOUNDARY 

The Virginian and Nova Scotian faunal provinces 

are easily differentiated in the inshore and offshore areas 

on the basis of ostracodes (pis. 7-69; fig. 5). The shallow-water species A1-trila aff. A. mnygdala (pl. 7), Actinocythereis aff. A. gmrt.illionensis (pl. 8; a new species,= 

A. gomillionensis of WiHiams, 1966), and Puriana 

rugipunctata (pl. 9) occur on the south side of Cape 

Cod but are absent to the north. Enchinocythereirs margaritifera (pl.13), Oytherella 8p. (pl.14), andMun.rseyella atlantica (pl. 12) occur mainly at outer sublittoral 

depths on southern Georges Bank 'and the shelf south 

of New England but are a;bsent to the north. Bensonocythere aff. B. whitei (pl. 11; a new species,= B. 'Whitei 

of Hazel, 1967a) does not occur north of Georges Bank. 

The genus Hulingsina (pl. 10) is restricted to the region 

south of Cape Cod, with one exception. 

Some cryophilic species extend southward into the 

western Gulf of Maine but do not reach Cape Cod. 

These include Baffinicythere howei (pl. 15; B. costata 

of Hazel, 1967a; see Hazel, 1967b), Oytherura ~ undata 

(pl. 16}, Elofsonella concinna (pl. 18), Oytherura~ 

mainensis (pL 55), and Oytheropteron latissimum (pl. 

17}, Palmenella limicola (pl. 24), Oytheropteron arngulatum (pl. 23}, Finmarchinella angulata (pl. 21}, 

Oythere lutea (pl. 22), Eueytheridea bradii (pl. ·24), 

and Patagonacythere dubia (pl. 20}. The above cryophilic species and some others extend south of Nova 

Scotia but do not pass and, for the most part, do not 

penetrate the Northeast Ohannel (pis. 15-25). 

The warm summer and fall temperatures in the Cape 

Cod Bay-Massachusetts Bay area (Fritz, 1965; 

Schroeder, 1966} seem to be a barrier to many equatorward -expanding cryophilic species. On the outer shelf, 

the deep Northeast Channel and high summer temperatures on adjacent Georges Bank provide barriers to 

these species. 

The thermophilic poleward-expanding shallow-water 

species may be halted by the low 3°C temperatures 

(Schroeder, 1963) in Cape Cod Bay and Massachusetts 

Bay or by the rapid change in summer temperatures 

in the Cape Cod area. On the outer shelf, thermophilic 

deep-water species are apparently halted by cold winter 

temperatures on eastern Georges Bank and in the Northeast Channel. More cryophilic than thermophilic species 

drop out at Cape Cod, and this is consistent with a 

stronger compression of summer than winter isotherms 

in the area. 

The small assemblage, mainly Muellerina abyssicola 

(pl. 19), M. canadensis (pl. 56}, and Thaerocythere 

crenulata (pl. 25), present in the deep Northeast Channel and Georges Basin area in zoogeographically more 

closely related to Nova Scotian province assemblages 

than to assemblages of the Virginian province, but is 

probably more appropriately included with what might 

be called an "Atlantic bathyal" province. 

NOVA SCOTIAN AND VIRGINIAN 

AMPHIATLANTIC SPECIES 

M~any of the species discussed in the present study are 

also weU known in Europe. All the amphiatlantic species occur in the Nova Scotian province, and most do not 

extend into the Virginian province (pis. 15-30). 
E14 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

TABLE 2.-North-south limits, climatic-zone range, zonal type (Hutchins, 1947), temperature range, and depth range of26 amphiatlantic 

species 

[All the species for which the zonal type could be determined are Type 3 (controlled equatorward by summer temperatures that can be tolerated) except the two species of 

Bafjinicythere which seem to be Type 2 species (controlled equatorward by winter temperatures which must be low enough for reproduction). Seventeen of the species 

occur in the frigid climatic zone and the rest, except for the common deep-water inhabitant Echinocythereis echinata s.l., occur at least in the subfrigid climatic zone. None 

of the amphiatlantic species extend into the subtropical climatic zone, and only five have been recorded from tne warm-temperate zone. Normally sublittoral species 

which have submerged to bathyal depths at thetr southern extent (for example, Robertsonites tuberculata in the Bay of Biscay) are not considered to belong to the sublittoral 

climatic zone of that area] 

North America Europe 

Amphia.tlantic species Climatic zones 

Northern Southern Northern Southern 

Bafjinicythere emarginata .. Kane Basin _________ Block Island ________ Northern Firth of Clyde ______ Frigid-mild temperate __ _ 

Spits bergen. howei__ _____________________ do _______________ Western Gulf of Franz Josef Land. __ Shetland Islands ____ Frigid-cold tempreate __ _ 

Maine. 

Callistocythere? cluthae _________ .do _______________ Northern Gulf of ____ _do _______________ North ChanneL ________ .do __________ -----------

Maine. 

Cythere lutea •• ____________ Cape Dyer _____________ .. do _______________ Murmansk. _________ Finistere. ___________ Subfrigid-mild 

temperate. 

Cytheropteron angulatum ___ Ungava Bay ________ Browns Bank _______ Franz Josef Land ___ Southwestern Frigid-cold temperate ___ _ 

Ireland. 

latissimum _____________ Dlsko Bugt_ ________ Western Gulf of _____ do _______________ Southern England._ Frigid-mild temperate. __ 

Maine. 

Cytherura? rudis ___________ Smith Sound ________ Ungava Bay ________ Stor Fjord __________ Varanger Fjord ______ Frigid-subfrigid _________ _ 

undata. --------------- Kane Basin ______________ do _______________ Northern Devonshire.-------- Frigid-cold temperate ___ _ 

Spitsbergen. 

Echinocgthereis echinata s.L Georges Basin._____ (1) Lofoten Islands. ____ Kattegat ____________ Cold temperate.----------

ElofsoneUa concinna _______ Kane Basin _________ Western Gulf ot Spitsbergen _________ Southwestern Frigid-Cold temperate .. 

Maine. Ireland. 

Eucythere declivis __________ Bay of Fundy _______ Southern Virginia ___ Varanger Fjord ______ Gulf of Naples ______ Subfrigid-warm 

temperate. 

Eucytheridea bradii. _______ Wolstenholme Roseway Bank ______ Franz Josef Land ___ Thames Estuary ____ Frigid-mild temperate __ _ 

Island. 

punctillata _____________ Kane Basin _________ Gulf of St. Northern Spits- Suffolk ______________ Frigid-mild temperate ••. 

Lawrence. bergen. 

Finmarchinella angulata ___ North Star Bay _____ La Have Basin ______ Northwestern Southwestern Frigid-cold termperate __ _ 

Spitsbergen. Ireland. 

finmarchica Disko Bugt _________ Long Island _________ North Cape _________ Bay of Biscay _______ Subfrigid-warm 

temperate. 

Hemicgthere borealis ________ Kane Basin _________ Southwestern Nova (1) Durham.----------- Frigid-cold temperate ___ _ 

Scotia. 

villosa _________________ Cape Dyer __________ Martha's Vineyard •. Bear Island _________ Bay of Biscay _______ Subfrigid-warm 

temperate. 

Hemicgtherura clathrata ____ Kane Basin _________ Georges Bank _______ Northern Isle of Wight ________ Frigid-mild temperate ••. 

Spitsbergen. Heterocgprideis sorbgana ________ do _______________ Northern Gulf of _____ do _______________ Southwestern Frigid-cold temperate ___ _ 

Maine. Ireland. 

Loxoconcha impreua _______ Southern Labrador •• Southern Virginia. __ Varanger Fjord ______ Madeira _____________ Subfrigid-warm 

temperate. 

Muellerina abgssicola _______ Disko Bugt_ ________ Slope, 40° N., 68° W_ Southwestern Skagerrak ___________ Subfrigid-cold 

Spitsbergen. temperate. 

Normanicythere leioderma •. Kane Basin _________ Western Gulf of Kong Karls Land ___ Isle of Skye _________ Frigid-cold temperate ___ _ 

Maine. 

Palmenella limicola ________ Disko Bugt_ ________ Northern Gulf of Franz Josef Land ___ Southwestern Frigid-cold temperate .•.. 

Maine. Ireland. 

Patagonacythere dubia ______ Holsteinborg ________ La Have Bank _____ _ (1) Shetland Islands ____ Subfrigid-cold 

temperate. 

Robertsonites tuberculata. _ _ Kane Basin__ _ _ _ _ _ _ _ Northern Gulf of Franz Josef Land ... Bay of Biscay _______ Frigid-mild temperate. __ 

Maine. 

Thaerocythere crenulata ____ Disko Bugt_________ Northeast ChanneL Trondheim Fjord ____ Skagerrak __________ Subfrigid-cold 

temperate. 

1 Known only from one locality. 

Zonal Temperatype ture range 

(o C) 

2 <G-20 

2 <G-14 

3 <G->7 

3 0±-22 

3 <G->11 

3 <G-22 

<G-8 

3 <G-18 

? 2-11 

3 <G-19 

3-25 

3 <G-18 

3 <G-18 

3 <G->11 

3 0±->15 

3 <G-13? 

3 <G->15 

3 <G-14 

3 <G-18 

3 0±-25± 

3 0±-11 

3 <G-19 

3 Q-12 

3 0±-10 

3 <G-18 

0±13 

Depth 

range 

(m) 

7-200 

24-146 

2Q-300 

Q-89? 

24-203 

Q-150 

Q-380 

Q-51 

53-4,700 

G-200 

Q-225 

3-750 

5-435 

Q-116 

Q-122 

Q-91 

o-? 

Q-170 

0-435 

Q-235 

4Q-1,60 

3-150 

5-200 

.5-205 

9-400 

2Q-801 

Baffinicythere emarginata (pl. 31) occurs in the 

Arctic province and thus tolerates temperatures below 

0°C. It is a cmnmon constitutent of assemblages along 

the coasts of Canada and Greenland, and its distribution 

is well known in Europe. It occurs commonly in the 

Nova Scotian province and extends into the Virginian 

province as far south as Block Island and the eastern 

pa~rt of Georges Bank (pl. 26). The species occurs commonly in Narragansett Bay (Williams, 1966) in areas 

where bottom temperatures reach a maximum of about 

20°C and a minimum of about 4°C (Hicks, 1959). In 

studying the European occurrences of the species, Elofson (1941) gave a temperature range o£ below 0°C to 

about l9°C. Thus, the species has approximately the 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E15 

same temperature tolerances on both sides of the 

Atlantic. 

Is the southern Emit of the species controlled by high 

summer temperatures it can tolerate or by low winter 

temperatures it must have for reproduction and repopulation? If its southern limit is controlled by summer 

temperatures too high for survival, theoretically 

Baffinicythere mnarginata could be found as far south 

as the Bay of Biscay off Europe. However, its southernmost occurrence is at a depth of 50 111 in Loch Fyne in 

the Firth of Clyde, Scoltand (Robertson, 1875). Average surface temperatures in August in this area are 

about 14°C, and in February the average surface temperature is about 7°C. I have no data on maximum or 

minimum botton1 or surface temperatures in the area, 

but it seems clear that Batfin.icythere ema.rginata is controlled equatorward not by high survival temperatures 

but by lmv winter temperatures needed for reproduction (apparently less than about 5°C). In Hutchins' 

(1947) terminology, B. enwrginata is a zonal Type 2 

species. 

This species is interesting also because it may be an 

example of equatorward emergence, a theoretical possibility with Hutchins' Type 2 and 4 species which are 

governed equatorward by winter temperatures needed 

for reproduction. The deepest known occurrence of this 

species is in the Arctic province off eastern Greenland 

at 200 m (Elofson, 1941). In this general area the bottom temperature at 200m is below 0°C in the warmest 

months (Schroeder, 1963). In the southern Nova 

Scotian and Virginian provinces (the species also 

occum in the Gu1f of St. Lawrence (Brady, 1870) hut 

no depth data are given), B. emarginata occurs in six 

areas; Browns Bank-Roseway Bank, Scotia Shelf, 

northern Gulf of Maine, western Gulf of Maine, northeastern Georges Bank, and the Cape Cod-Narragansett 

Bay area (pl. 26). Figure 10 shows deepest occurrence 

and the warmest and co1dest ·average bottom temperature in each of these areas. The lower depth limit decreases from 200 to 38 m equatorward as the bottom 

average high temperature increases from about 0° to 

about 14°'C; there is only about a 5.5°C difference in the 

bottom average low temperature. On the basis of the 

a vail able data, B affin.icythere e1narginata is emerging 

toward the equator, the reason being that it is eurythermal but apparently dependent on winter temperatures lower than about 5 oc for reproduction and early 

growth. 

This emergence of the lower depth limit is apparently 

in response to the particular present hydrographic-climatic situation along the At~antic coast. The outer sublittoral areas of the shelf are under the influence of the 

Gulf Stream, and the bottom temperature is always 

above 5°C south of Cape Cod. However, in inner sublittoral areas the bottom temperature falls to 4°C in 

winter. 

In the western Atlatnic, Elofsonella coneinna (pis. 

18, 32) is distributed from Cape Frazer to 43°19' N. in 

the Gulf of Maine (table 1, Gos 1021, 37m). Here the 

average surface temperature in August is about 16°C 

and in February about 3°C. The bottom temperature in 

February here is about 4°C and in August probably 

near 14°C. In Europe (pl. 32), E. concinna occurs in 

the Arctic and as far south as off southwestern Ireland 

(Norman, 1891, no depth given) which is far south of 

the 4 oc winter surface isotherm. The a¥erage surface 

temperature in the area in August is about 15°C and in 

February 9°C. The yearly average bottom temperature 

at 200 m off southwestern Ireland is 10.5°C with only a 

2 o C range (Schroeder, 1963) . Elo fsonella concinna is 

most commonly found at inner sublittoral depths, and 

the bottom temperature where E. concinna occurs off 

Ireland is probably between 11° and 15°C. These data 

indicate that the species is controlled equatorward by 

summer temperatures and, in Hutchins' (1947) terminology, is a Type 3 species. The temperature 

tolerance in terms of averages for the coldest and warmest months is probably between <0° and 14°C, although 

the absolute maximum is higher and Elofson (1941) 

gives an upper limit of 19°C. 

200 

190 

180 

170 

160 

150 (/) 

140 0::: 

I.LJ 130 

120 ~ 

110 ~ 

100 z 90 

80 :i 

70 t 60 I.LJ 

50 c 40 

30 

20 

10 

0 

"C VI -~ ..... ..... ..... c:..:.:: >. c: c: ..:.:: 15v "S "S .... c: tU 

.... tU "Cc: (.!JG> (.!JG> G) tU G)- I III 

:;;~ eX~ c: c:.!: t)III .... c: C:·- "C+" VI G) .... tU .... tU tU VI 

w 

tU ... G) VI>, 

c: tU ~~ ~~ .C:bl) 

G) G) 8~ G)c: (.!) ~ ~ ......... VI,._ ........ 50 G) 0 .... 0 a. IV 0 G) ~ 0 G) robll "- VI IIIo z ZC!J (.)~ 0::: tU 

z 

FIGURE 10.-The apparent equatorward emergence of the lower 

depth limit of Baffinicythere emarginata (Sars). The maximum depth decreases from 200 to 38m from eastern Green-

·land ( Elofson, 1941) to south of Block Island ( ta·ble 1, Gos 

1258) ; the bottom average high temperature increases from 

about ooc to 14°0, but the hottom average l<YW temperature 

increases only from a:bout -1 oc t'O 4.5°0. 
El6 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

Palmenella limieola (pls. 24, 33) occurs as far south 

in the western Atlantic as off Muscongus Bay, Maine, at 

64 m. The average surface temperature here in August is 

14°0 and in February 4°0. The average bottom temperature at 100 In in this area is about 7°0 in the 

warmest months and 4 °0 in the coldest months. By 

interpolation, the average bottom temperature at 64 min 

the warmest months ·would be about 10°0. In Europe 

(pl. 33), P. li1nicola occurs as far south as off southwestern Ireland at 200m (Norman, 1905). The yearly 

average bottom temperature off southwestern Ireland at 

200 m (Schroeder, 1963) is about 10.5°0 with a 2°0 

range. Thus, the temperature tolerance of the species, in 

terms of seasonal average temperature, is about <0°-

11 °0. Elofson (1941) gave an upper maximum limit of 

l2°C for the species. Palmenella limicola appears to be a 

Type 3 species, controHed by summer temperatures to 

the south. The species is commonly ,found in shallow 

waters in frigid-subfrigid regions, at depths of 20 111 in 

the waters off Kong Karls Land, Svalbard, 5-30 m in 

the fjords of Finnmark, Norway, and 13-18 m off eastern Greenland. At its southernmost occurrence it seems 

to inhabit only deeper waters and thus may be undergoing equatorward submergence. 

Baffinicythere mnarginata, Elofsonella corwinna, and 

Palmenella lim.icola were used to demonstrate the procedure for determining the zonal type (Hutchins, 1947) of 

a species. Determination of zonal type is best if bottom 

temperatures are available, at least at the critical southern or northern limits of the range of a species. However, these may not be required to determine the zonal 

type because the August and February surface temperatures coincident on one coast are far apart on the other. 

Therefore, if the species is common and reasonably well 

collected, the zonal type is easily determined. For an 

accurate estimate of the actual limiting temperatures, it 

is imperative to have bottom temperature data. 

Most of the amphiatlantic species in theN ova Scotian 

and, in fewer numbers, in the Virginian province also 

occur as far north as the Arctic province. Most of 

these are Hutchins' Type 3 species, limited southward 

by summer high temperatures. The fact that they do 

occur in the circumpolar Arctic province easily explains 

their presence on both sides of the Atlantic.. The amphiatlantic species treated are listed in table 2 with their 

geographic end points on either side of the Atlantic and 

their zonal type. The distributions of most of these 

species in both Europe and North America are shown 

on plates 31-54. 

Several of the amphiatlantic species do not occur in 

the Arctic province (pls. 48-54). These are Thaerocythere crenulata (pl. 48), Oythere lutea (pl. 50}, Loxoconcha impressa (pl. 49), Hemieythere villosa (pl. 51), 

Patagonacythere dubia (pl. 52), Finmarchinella finmarchica (pl. 53), and, possibly, 11/ullerina abyssicola 

(pl. 54) (depending upon whether the Spitsbergen 

loeality given by Brady and Norman, 1889, is on the 

southwestern coast of Spitsbergen or not). These forms 

tolerate 0°0 or slightly lower temperatures and therefore are not limited northward by winter temperatures. 

They only occur, however, in areas where the summer 

surface temperatures are above freezing at least 3 or 4 

months of the year and, therefore, are possibly dependent on above-freezing temperatures for reproduction 

and early growth. 

With the possible exception of Eucythere declivi8 

(the distribution of this species is difficult to map because G. S. Brady and his associates considered E. argus 

( Sars) a synonym), none of the species studied that 

occur only in the temperate provinces of the western 

Atlantic are amphiatlantic. 

NOVA SCOTIAN AND VIRGINIAN 

ENDEMIC SPECIES 

Oytherura? 1nainensis Hazel and Valentine, has been 

found in several inner sublittoral samples from the 

Gulf of Maine and Scotia Shelf (pl. 55, 60). Its southern limits seems to be about 43°19' N., and it has not been 

found north of the Gulf of St. Lawrence ( Oytherura 

undata, var., of Brady, 1870). Thus, it may be an endemic North American species restricted to the Nova 

Scotian province. Of the common species in the Nova 

Scotian province, this is the only one apparently endemic 

to eastern North America that does not also occur in the 

Virginian province. 

Several endemic species occur commonly in both the 

Virginian and Nova Scotian provinces. These include 

the two most common and, usually, most abundant 

species found, Muellerina canadensis (pl. 56) and lJf. 

aff. Jtf. lienenklau.s-i (pl. 57) (a new species=M. lienenklausi of Hazel, 1967a). The species are closely related 

and, because they occur commonly together, attention 

must be paid to details in order to consistently distinguish them (see Hazel, 1967a). 

11/uellerina canadensis was described from the Gulf of 

St. Lawrence by Brady (1870) and subsequently was 

reported from Godhavn and Holsteinborg Harbors, 

Greenland, and the Davis Strait (Norman, 1877; Brady 

and Norman, 1889). However, the illustrations and description in Brady and Norman ( 1889) seem to be of 

another genus, probably a M unseyella (rather than a 

Leptocythere as suggested by Blake, 1933). I am confident that what I (Hazel, 1967a) illustrated as Muellerina camadensi.'3 is conspecific with what Brady ( 1870) 

illustrated. Thus, M. canademis is known from at least 

as far north as the Gulf of St. Lawrence and possibly, 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E17 

but not probably, to Disko, Greenland. Muellerina a:ff. 

M. lienenklau:si occurs at least as far north as the Bay 

of Fundy. Southward ltf. aff. M. lienenklau.n has been 

in samples taken off Cape Hatteras. A single specimen 

of what may be M1.tellerina canadenr·Jis has also been 

found off Cape Hatteras (11/turrayina ca:nailen8is of 

Hulings, 1966, from off Virginia is actually 11/ uellerina 

aff. M. lienenklausi). lJfrttellerina canadensis is known 

from samples taken at depths from 7 to 801 m. It occurs, 

however, in a higher percentage of samples taken from 

depths of 100 to 250 m than in samples from greater or 

shallower depths. Mu,.ellerina aff. M. lienenklausi was 

not found in samples taken deeper than 235 m, and it 

occurs in a higher percentage of inner and middle shelf 

samples than outer shelf samples (fig. 11). 

In the study area, both species occur in areas where 

minimum to maximum bottom temperatures are 2°C to 

about 20°C. M1.tellerina aff. M. lienenklausi does not 

occur in the central parts of the Gulf of Maine, in the 

Northeast Channel, or on the continental slope, but M. 

canadensils does. Otherwise, the geographic distribution 

of these species is very similar. These two endemic 

species seem to be mild-temperate to cold-temperate 

forms with M. canadensis possibly, but doubtfully extending north into the subfrigid Labrador province. 

Two of the more eurythermal endemic species are 

Leptocythere au.gusta Blake (pl. 58, 60) and Mun8eyella 

> (.) 100 z 

LLJI- :::::lz 90 OL&J 

LLI(.) 

O:::o::: 

LLL&J 80 

oa.. 

zz 

c(- 70 

S'"' 1.&.10 60 01.&.1 

ceo 

l:c( 

(/):I: 50 

-(/) (/)z 

1.&.1:::::1 40 

a..z ..... -:EO 30 cc-

(/)~ LL.Il) 

o- 20 0:::0::: 

LLJti) 10 a:l-

::Ec 

:::::1 0 z 10 0 10 0 10 0 10 8 LO 0 LO N 10 ,..... 0 N 10 " N LO " 0 ~ I .-t .-I .-t .-I N C\1 N N .-t 

~ I ~ I ~ I ~ I N 10 .-t .-I .-t .-I 

" 0 C\1 10 " 0 N LO .-I .-t .-I .-I C\1 N N 

mananensi8 Hazel and Valentine (pl. 59, 60). Leptocythere angu.sta is found from at least as far south as off 

Virginia ( Calli8tocythere reticulata of Hulings, 1966) 

to southern Labrador. It seems to be basically a mildand cold-temperate species that extends into the southern part of the subfrigid Labrador province. M. 

mananesi8 Inay extend north into the Arctic province, 

but perhaps as a different subspecies (Hazel and 

Valentine, 1969). 

The large cytherettid Pseudocytheretta edwanbd 

Cushman (pl. 61) is common in shallow waters in the 

Vineyard Sound-Buzzards Bay-Narragansett Bay area 

(Cushman, 1906; Williams, 1966). It is rare in my samples, however, which were taken mostly at depths 

greater than 20 m. P. edwardsi may be an indicator 

for the Virginian province if it is not conspecific with 

P. tracyi (Blake) which was described from Mount 

Desert Island, Maine. I have not been able at determine 

how to distinguish the two species. Juveniles of a large 

smooth cytherettid were found in only one of my samples from the Nova Scotian province (taJble 1, Gos 1062, 

taken off Yarmouth, Nova Scotia). Tentatively, P. 

tracyi is considered to be a synonym of P. edwardsi. 

Types of neither species are available for study. 

Pterygocythereis americana inempectata (Blake) (pl. 

62) is a rather common form, particularly as juveniles, 

LO 

N " A 

LO 

N 

0 

Muellerina aff. M. lienenklausi 

(Ulrich and Bassler) 

uency distribution of the species in all 

samples collected within a depth interval 

LO 

,..... 

.-I 

I 

.-I 

10 

.-I 

10 

N 

.-I 

~ 

0 

N 

0 

10 

N 

~ N 

N 

10 

,..... 

N 

I 

.-I 

10 

N 

DEPTH, IN METERS 

FIGURE 11.-Depth "preference" of Muellerina cwnadensis and M. ·aff. M. Uenenklausi (=M. lienenklausi of Hazel, 1967a). 

The shaded histograms show the number of samples containing the species at 25-m depth initervals. This plot is similar 

for both S!peC'ies (although M. canadensis has a greater depth limit, 801 versus 235m), but this similarity is merely a 

reflection of the preferential sampling at middle shelf depths. The unshaded histograms, however, based on the frequency' distribution of the species in all samples collected within a 25-m depth interval, indicate that, although these 

species are common at neartly all depths in the sublittoral zone, the "preferred" habitat of M. canadensis is 100-225 m, 

whereas M. aff. M. lienenkl,a,U8i is JOOst commonly found at 50-150 m. 
ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

in samples from the Virginian and Nova Scotian 

provinces. 

Two species of 1Actinocytherei8 are rather widely distributed over the area. These are A. ·vineyardenJ3is 

(Cushman) (pl. 63) and A. da1JJsoni (Brady) (pl. 64); 

A. Dineyardensis tends to occur in deeper waters. A. 

vineyardensis does not occur in the ostracode-rich samples taken off Nova Scotia where bottom temperatures 

are as low at 2°-3°C in winter; however, A. dawsoni 

occurs frequently in this area. A. dawsoni ·also occurs 

in the Gulf of St. Lawrence (Brady, 1870), and the 

species seems more tolerant of colder temperatures. A. 

vineyardensis occurs at least as far south as the Cape 

Hatteras area, whereas A. dawsoni may not occur further south than the northern pa·rt of the Viginian 

province. 

Two speeies of the apparently endemic genus Bensonocythere occur in both the Nova Scotian and Virginian provinces. B. arenicola (Cushman) (pl. 65) 

and B. americana Hazel (pl. 66) occur in several samples 

but are never abundant in any one sample. They range 

from off Nova Scotia to south of Long Island. 

The neocytherideidine genera, "Sahnia" (a genus with 

no valid type species, pl. 67) , H uling sina (pl. 10) , and 

Oushmanidea (pl. 68) are fairly common, primarily 

in inner sublittoral samples, in the area. These are incompletely studied at present, and several species of 

Hulingsina and Oush-manidea may be represented. 

"Sahnia" is represented by at least two species, "S." 

foveolata Brady and "S." fasciata of Cushman ( 1906) . 

There was only one occurrence of "Sahnia" in the 

Nova Scotian province, off St. Mary Bay, Nova Scotia. 

This form, larger than the ones to the south, could be 

"S." fm-'eolata (Brady), described from the Gulf of St. 

Lawrence. The smaller species occurs commonly in the 

Virginian province and is probably what Cushman 

(1906), Williams (1966), and Hulings (1966) refer to 

"S." fasoiata (Brady and Robertson). 

Except for one occurrence, H ulingsina is confined to 

the Virginian province (pl. 10). 

E chinocythereis plmnibaBalis procteri (Blake) was 

found chiefly in the Gulf of Maine area (pl. 69), but it 

also occurs on Georges Bank (table 1, W81 ; in table 1 

of Hazel (1967a), this occurrence is erroneously indicated as E. margaritifera). Fossils of this species, but 

perhaps not the same subspecies, have been found in the 

late Miocene of Virginia and North Carolina. Off 

Europe, E chinocytherei8 does not extend north of the 

Lofoten Islands (pl. 4 7) on the Norwegian coast. Off 

N ort.h America, no E chinocythereir.; has been reporb~d 

north of the occurrences of E. planib(l8alis procteri 

(pl. 69). Thus, Echinocythereis seems to be a primarily 

tropical to cold-temperate genus. In contrast to Echinocythereis, the closely related genus Rabilimis seems to 

be chiefly a frigid and sub frigid taxon (pl. 47). It is 

known from only two localities in cold-temperate regions; one is off the Isle of Lewis, Scotland (Brady, 

1868a), and the other from the Skagerrak ( Elofson, 

1943). Each locality yielded only one empty valve. 

Except for these two occurrences, there is no overlap 

between the distribution of Echinocythereis and 

Rab·ilimis. 

Actinocytherei.s aff. A. gomillionensis is rare in offshore samples but occurs commonly inN arragansett Bay 

(Williams, 1966) (pl. 8) and has been seen in samples 

from Buzzards Bay, Vineyard Sound, Hadley Harbor, 

and off Cape Hatteras. It is a southern form related 

to A. bahamensis (Brady) and is a common fossil in 

upper Miocene, Pliocene, and Pleistocene deposits along 

the Atlantic coast. 

Recent species referred to the genus H apwcytheridea, 

probably erroneously, do not seem to occur north of 

Cape Cod. Williams (1966) reported what is probably 

"Haplocytheridea" bradyi (Stephenson) (as "H." setipunctata (Brady)) from Narragansett Bay (pl. 9). I·t 

vvas not in my offshore samples. 

The thaerocytherine genus Puriana seems restricted 

to tropical to mild-temperate waters. P. rugipunctata 

(pl. 9) occurs in Narra;gansett Bay (Williams, 1966), 

Vineyard Sound, Hadley Harbor, and off Nantucket. 

The genus A urila is represented in the study area by 

Aurila aff. A. amygdala (pl. 7). Williams reported it as 

common in Narragansett Bay. The species also is known 

from Pamlico Sound, N.C. (Williams, 1966), and the 

west coast of Florida (Benson and Coleman, 1963; 

Hulings and Puri, 1964). Oythere albomaculata reported by Cushman ( 1906) is undoubtedly this species, 

as I have seen Atttrila aff. A. amygdala in a sample from 

Cushman's reoccupied Fishhawk Station 7723. It also 

occurs off Woods Hole (Cushman, 1906) and Martha's 

Vineyard. 

B ensonocythere aff. B. whitei (Swain) seemingly is 

restricted to the Virginian province and in this study 

was found in several samples (pl. 11) from Georges 

Bank and areas south of New England. 

Only one platycopid species was found in the present 

study, a Oytherella which could not be assigned with 

any confidence to a described species. It occurs primarily 

on the outer shelf south of New England (pl. 14). 

Also characteristic of the deeper areas of the northern 

Virginian province is 11/ un.rseyella atlantica Hazel and 

Valentine (pl. 12) and Echinocythereis margaritifera 

(Brady) (pl. 13). These species have been found on the 

south side of Georges Bank and in the Hudson Canyon 

area south of New England. 
OSTRACODE ZOOGEOGRAPHY, NOVA SCOTIAN AND VIRGINIAN FAUNAL PROVINCES E19 

E chinocythereis mm·garitifer·a (pl. 13) is interesting 

because the available data, much less than desired, suggest that the species is submerging poleward. E. margaritifera occurs in the Gulf of Mexico off Louisiana 

and Mississippi (Curtis, 1960) and western Florida 

(Benson and Coleman, 1963), off Cape Hatteras (Duke 

University collections). It was found on the shelf and 

upper slope south of New England in the present study. 

Curtis reported the species from her offshore biofacies 

at depths greater than 27 m. Based on her data, the 

bottom temperature at 27 m would be about 20°C in 

February and 26°C in August. Benson and Coleman 

found the species at depths as shallow as 20m, at which 

depth the bottom temperature would be about 21 oc in 

winter and about 2'6.5°C in summer (interpolation from 

the surface temperatures and the 200-m temperatures 

given by Schroeder, 1963). 

Off Cape Hatteras (34°24' N., 76°05' W.), E. margaritifera was found at depths as shallow as 35 m. The 

bottom temperature in February would be near 11 oc 

and in August about 24.5°C. In the area of the present 

study the species was found only along the edge of the 

shelf south of New England at depths greater than 

110 m (the 82-m occurrence given in Hazel, 1967a, 

table 1, should be under E. planibasalu). The bottom 

temperature in the warmest months in the area of the 

shallowest occurrence (table 1, AB3 W34) averages 

about soc (Schopf, 1967) and in the coldest months 

about 6.5°C. Although these data from four scattered 

areas are not entirely convincing, they nonetheless suggest that the shallowest depths that E. margaritifera 

can inhabit are deeper toward the north. 

The presence of thermophilic species such as E. margaritifera and Munseyella atlantica in deeper waters 

in the northern Virginian province is no doubt related 

to the fact that the outer shelf south of New England 

is under the influence of the Gulf Stream. In such 

hydrographic situations, poleward submergence is not 

an uncommon phenomenon. Cerame-Vivas and Gray 

( 196'6) have shown that assemblages characteristic of 

three biogeographic provinces are represented in the 

Cape HaUeras region. On the outer shelf, assemblages 

characteristic of the tropical Caribbean province are 

present; in inner shelf areas south of Cape Hatteras, 

assemblages characteristic of the Carolinian province 

are dominant; and in inner shelf areas north of Cape 

Hatteras, assemblages characteristic of the Virginian 

province are present. 

Table 3 summarizes the distribution, temperature, and 

depth data available for the endemic species treated 

herein. 

TABLE 3.-N orth-south limits, climatic-zone range, temperature-tolerance range, and depth range of 19 species endemic to the western Atlantic 

[Only one species is known to occur in the frigid zone and one in the subfrigid zone. The remainder are restricted to cold-temperate or warmer climatic zones) 

Distribution 

Endemic species Climatic zones 

Tempera- Depth 

ature 

Northern Southern range (°C) range (m) 

Actinocutherei& daw&oni---------------------------- Georges Bank _______________ Hudson Canyon ____________ Mild temperate.---------------------- 0±-19 3-185 

aff. A. gomillionen&ilf __________________________ Narragansett Bay ___________ Florida _____________________ Mild temperate-subtropicaL _____ ---_- 0-25.± 2-136 

vinevarden&ilf ____________ ---------------- ______ Mount Desert Island ________ Cape Lookout. _____________ Cold temperate-mild temperate ______ - 4->20 26-231 

Aurila aff. A. amygdala ____________________________ Narragansett Bay ___________ Florida Bay ________________ Mild temperate-tropicaL _____________ _ 4-29 0-50 

Bensonocuthere americana_------------- ____________ Bay of Fundy ______________ 40° N., 73° W ________________ Cold temperate-mild temperate ______ _ 4-16 13-111 

arenicola ___________________________________________ do ___________________________ do. __________________________ do .• _____________________________ _ <4-20 2-139 

aff. B. whiteL _ ----- ___________________________ Georges Bank _______________ ?Cape Hatteras. ____________ Mild temperate. __________ -------- ___ - 4->20 23-114 

Cutherella sp ___ ---- __ ------ ____________________________ do •• ____________________ Hudson Canyon _________________ do. _______________________ ----- __ _ 5-13 82-185 

Cutherura? mainensis ___________ ------------- ______ Gulf of St. Lawrence ________ Western Gulf of Maine .• ____ Cold temperate.---------------------- 0.±-14 24-87 

Echinocuthereis margaritifera. ______________________ Georges Bank _______________ Veracruz, Mexico ____________ Mild temperate-tropical_ _________ -- __ _ 6-27 20-235 

planibasalis procteri. _ ------------ _____________ Bay of Fundy ______________ Georges Bank ______________ Cold temperate-mild temperate ______ _ 3-20 12-89 

Leptocuthere angusta _____ -------------------------- Southern Labrador _________ Southern Virgnia ___________ Subfrigid-mild temperate_------------ 0..±:->15 3-170 

Muellerina canadensis_- ___ -_______________________ Gulf of St. Lawrence .. ______ Cape Hatteras. _____________ Cold temperate-mild temperate .• _- __ _ 0->20 7-801 

aff. M.lienenklausi.. -------------------------- Bay of Fundy ______________ Northern Florida ___________ Cold temperate-subtropicaL---------- 2->20 3-240 

Mun&eyella atlantica _______________________________ Georges Bank _______________ Hudson Canyon ____________ Mild temperate ___________ ------------ 6-13 86-235 

mananensis-- -------- _________________________ Baffin Bay _______________________ do. _____________________ Frigid-mild temperate. ___ ----- __ -___ - <0->13 20-201 

Pseudocutheretta edwardsi. _________________________ Scotia ShelL ________________ Southern Virginia.__________ Cold temperate-mild temperate. _____ _ 3->20 3-123 

Pterugocuthereis americana inexpectata ______________ Bay of Fundy ______________ ?Cape Kennedy _____________ Cold temperate-subtropicaL _________ _ 2-20 3-245 

Puriana rugipunctata __ -- __ -_______________________ Narragansett Bay ___________ Caribbean. _________________ Mild temperate-tropicaL ••. _______ -_-- 4-29 2-73 
E20 ATLANTIC CONTINENTAL SHELF AND SLOPE OF THE UNITED STATES 

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