PLEIST_QCENE PLANTS FROM NORTH CAROLINA 

By EDWARD WILBER BERRY 

INTRODUCTION 

The field work upon which this report is based was 

done in 1906 and 1907 as a part of the cooperative 

study of the Atlantic Coastal Plain, under the direction of the late William Bullock Clark. Associated 

with the writer in this work were L. W. Stephenson, 

B. L. Miller, jr., and J. E. Pogue. Preliminary accounts of the plants collected were published in 1907 1 

and 1909.2 

As has been frequently emphasized, the study of the 

Pleistocene floras in this country is in an exceedingly 

backward state as Ineasured by the volqme and precision of our knowledge of Pleistocene floras in Europe. 

Researches in Pleistocene geology in North America 

have been confined almost entirely to glaciology, and 

the problem of the correlation of the glacial . deposits 

with those outside the glaciated area has not been 

solved, nor is there any general agreement regarding the 

genesis of the Pleistocene deposits south of the terminal 

moraines. The present account of what is known of 

the Pleistocene flora of North Carolina and the conclusions that may be legitimately derived from it is 

offered in the hope that it may stimulate an interest in 

a neglected field of research and form a small part of 

the evidence upon which to base future more comprehensive conclusions and generalizations. 

A word of explanation regarding the illustrations is 

required. Nearly all of them have been made from 

leaves preserved as carbonaceous films in the peaty 

clays. . These specimens were carefully washed out, 

and blue prints were made directly from them. Outlines and as much of the venation as could be seen were 

. inked on the blue prints, which were then bleached. 

This procedure made it possible to handle a much 

larger amount of material and prevented any possible 

damage to the exceedingly fragile specimens, which 

were mounted on cards or between glass. The accompanying drawings were made from tracings of the 

original nature prints. 

1 Berry, E. W., Contributions to the Pleistocene flora of North Carolina: Jour. 

Geology, vol. 15, pp. 338-349, 1907. 

2 Berry, E. W., Additions to the Pleistocene flora of North Carolina: Torreya, 

vol. 9, pp. 7i'-73, 1909. 

GEOLOGY OF THE PLEISTOCENE IN NORTH 

CAROLINA 

The Pleistocene formations of North Carolina were 

discussed in detail by Stephenson 3 in 1912. Five 

formations, mapped largely by their topographic form 

and constituting · five terrace plains with seawardfacing scarps and reentrants up the stream valleys, 

were recognized. These are, from youngest to oldest; 

the Pamlico, Chowan, Wicomico, Sunderland, and 

Coharie. Among these formations identifiable fossil 

plants have been found only at a single locality in the 

Wicomico, at two localities in the Chowan, and at two 

localities in the Pamlico. 

The Wicomico locality is 1U miles east of Weldon, 

in Northampton County, where the following section 

is exposed, according to Stephenson: 

Section 1'}4 miles east of Weldon, N. C. 

Yell ow more or less sandy clay, largely concealed by Feet 

vegetation ________________ "' _,.. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 35 

Yellow pebbly argillaceous sand_____________________ 2 

Lenses of yellow clay with leaves____________________ ~ 

Yellow very coarse pebbly arkosic sand ____________ ·___ 2~ 

The principal Chowan locality is along the right 

bank of Neuse River 79% miles above New Bern and 

4~ miles above Seven Springs, in Wayne County, 

where the following section is exposed: 

Section on Neuse River above Seven Springs, N.C. 

Clay loam grading down into sandy loam ____________ _ 

Coarse white and yellow sand with gravel layers and 

boulders in lower portion ________________________ _ 

Tough blue clay interstratifi,ed with lignitic and leaf 

Feet 

4 

7 

layers 2 to 3 inches thick _______________ _________ _ 3 

Published accounts of the fossil plants from this 

section have alluded to it as station 850 . 

The second Chowan locality is at Dupree Landing, 

on the right bank of Tar River about 14 miles above 

Greenville, in Pitt County, where the following section 

is exposed in a depression in the Patuxent formation: 

Section at Dupree Landing, N. C. 

Sandy loam and argillaceous sand with pebbles along base ___________________________________________ _ 

Lignite __________________________________________ _ 

Feet 

6 

0-3 

a Stephenson, L. W., The Coastal Plain of North Carolina: North Carolina 

Oeol. and Econ. Survey, vol. 3, pp. 266-290, 1912. · 

97 

\ 

I 
98 SHORTER CONTR.IBUTIONS TO GENERAL GEOLOGY, 1925 

The Pamlico formation, whose surface constitutes 

the lowest terrace plain of eastern North Carolina, 

eontains fossil plants at Old Mill Landing, on the left 

bank of Roanoke River 8 miles above Williamston, in 

Bertie County, ~here the following section is exposed: 

Section at Old Mill Landing, N. C. 

Ft. in. 

Surface loam grading down into buff sand with small 

pebbles interstratified with argillaceous layers __ _ 10 Fine gravel ___________________________________ _ 8-10 Concealed ___________________________________ _ 3 6 ' Gravel _______________________________________ _ 1 

Cross-bedded sand ____________________________ _ 1 

Gravel------ --------------------------------- 6 

y ell ow sandy clay ____________________________ _ 5 

Dark micaceous clay with plants ________________ _ Q-1 

78 

niteJ.y named. The Wicomico formation in North 

Carolina is a direct southward continuation of the 

Wicomico formation of Maryland and Virginia. The 

Chowan and Pamlico together correspond to what has 

been called the Talbot formation in Maryland and 

Virginia, the Chowan corresponding to the older and 

the Pamlico to the younger part of the Talbot, and it 

is probable that detailed work would show a corresponding division of the Talbot in Virginia and on the 

Eastern Shore of Maryland. (See fig. 5.) 

CHARACTER OF THE FLORA 

The flora here described contains more or less certainly determined remains of 48 species, representing 

77 

0 

0 10 20 30 4C 50 MILES 

FIGURE 5.-Map showing the Pamlico, Chowan, and Wicomico coasts of North Carolina 

The second Pamlico locality is a low exposure on the 

right bank of Neuse River 10 or 12 miles below New 

Bern, in Craven County, from which only the remains 

of the bald cypress and pine have been recorded. · 

As regards the correlation of these formations with 

the Pleistocene formations of the adjacent States it 

may be stated that all can be traced for long distances 

in the Coastal Plain, both north and south of North 

Carolina. In South Carolina these formations and 

their corresponding terrace plains have not been defi30 ~~enera in 23 families and 17 orders. All but tw() 

conifers, representing the genera Pinus and Taxodium, 

·are angiosperms or flowering plants, ·and of these all 

belong to the class Dicotyledonae except fruits representing a grass and a pond weed. 

None of the recorded species are· forms that appear 

at all out of place in a North Carolina flora, although 

several no longer exist at or near the localities where 

the fossils were found. Two species of willows, one of 

birch, two of oaks, two of crab apple, two of hawthorn~ 
PLEISTOCENE PLANTS FROM NORTH CAROLINA 99 

one of Dendrium, and one of Vaccinium are d~scribed 

as new and consequently extinct species. These new 

forms constitute nearly 23 per cent of the whole 

flora, a proportion that is somewhat misleading, _for 

the following reasons: One of the oaks is obviously an 

abnormality or hybrid and not a true botanical · 

species; the two species of Malus and the birch are not 

very precisely differentiated; and the two species of 

Grataegus belong to a genus that appears to have undergone rapid mutation in recent times and to be of un- , 

certain value. This leaves an extinct species . of 

Quercus and one of Dendrium that I regard as satisfactory and a species of Vaccinium that may be 

authentic or may possibly represent an abnormality. 

Both the oak and the Dendrium represent· Pleistocene forms, each regarded as ancestral to two existing 

species of different environmental requirements. Thus 

the true percentage of extinct species may really be 

about 4 instead of nearly 23. 

Nearly all the species represented are trees or shrubs. 

The only herbaceous forms are Sparganium, Ohaetochloa, and Polygonum, represented by fruits and not 

by leaves. Thus in Pleistocene time, as to-day in the 

. backwaters of rivers and estuaries of the Coastal 

Plain of North Carolina, the bulk of the leaves that 

were preserved in the sediments were the leaves of 

trees. 

The most abundantly represented forms in this 

'Pleistocene flora are the bald cypress, the river birch, 

the beech, various oaks, and the button ball. The 

oaks are remarkable for both their individual abundance .and variety, ten species being recorded, representing both dry and wet soil forms. The most abundant of these are the wet-soil forms Quercus nigra, 

Q. phellos, and Q. michauxii, although Quercus predigitata is about equally abundant. 

All of North America north of the terminal moraine, 

with the exception of a few nunataks or small ice-free 

areas that projected above the continental ice sheet 

and a few ice-free regions, the largest known of which 

were in central and northern Alaska, have been colonized by vegetation since the end of the Wisconsin glaciation, which is considered to have been from 10,000 

to 20,000 years ago. 

A great deal of misconception has existed regarding 

the severity of glacial climates and their effect on the 

Tertiary vegetation of North America. There is little 

doubt that the present major floristic regions were well 

marked towar~ the end of the Tertiary period, particularly as regards the separation of the eastern mesophytic region from that of the Pacific region by the 

intervening area, consisting mainly of prairie or grass 

land, but including at the south desert or semidesert. 

During the maximum extent of the ice there must 

have been cons~derable crowding and competition between the native flora and that which formerly existed in the glaciated region. That there was an actual 

I 

zonal arrangement of climatic belts south of the ice 

front seems very doubtful. Even in the immediate 

vicinity of the ice front there was probably no continuous barren ground or tundra zone inhabited by 

arctic vegetation. None of the · evidence as yet ·disclosed in the Pleistocene floras of North America, although admittedly very incomplete, shows an appre~· 

ciable lowering of temperature at any great distance 

away from the ice front. This may, of course, be due 

to the fact that the bulk of the Pleistocene plants so 

far discovered have come from relatively low altitudes 

and mostly from the Coastal Plain. 

The plant localities in the Pleistocene area of North 

Carolina are about 425 miles south of the terminal 

moraine on the Atlantic coast, about 400 miles south 

of its position in southern Ohio, about 475 miles southeast of it in Indiana, and slightly over 600 miles eastsoutheast of its southernmost limit in southern Illinois. 

Unless there were a secular lowering of temperature 

amounting to a cataclysmal change, which is highly 

improbable, the effect of the ice sheet on the vegetation 

of low-lying coastal lands several hundred miles away 

would not be detectable. The coastal region of North 

Carolina during the Pleistocene epoch appears to have 

been densely forested and to have received an abundant rainfall, both factors that would tend to stabilize 

the climate and prevent extremes. By comparison 

with areas that have been critically studied, like the: 

Dismal Swamp, it may be concluded that the climate 

along the Pleistocene coast of North Carolina, except 

where the normal climate was disturbed by violent 

and long-continued winds, was highly favorable for a 

vigorous and dense forest cover, with slight daily 

variations ·of temperature, a long growing season, 

heavy and well7distributed rainfall, a uniformly high 

humidity, and freedom from violent frosts. 

The cypress swamps along the Pleistocene coast 

clearly indicate that it was fringed by barrier b~aches 

similar to those existing at the pre_sent time along the 

northern Carolina coast, and that in the inclosed 

sounds behind these beaches the salinity of the water 

was kept down by the volume of river water entering 

them; otherwise salt marshes would have gradually 

replaced the swamp forests. 

The plant remains-discovered in the North Carolina 

Pleistocene show no elements that can be definitely " 

interpreted as northern forms, and although the horizons at which they occur can not be precisely correlated with glacial chronology, three horizons are represented-namely, the Wicomico, Chowan, and Pamlico-and it is not likely that all the periods of coastal 

subsidence and the deposition of coastal terraces 

should have coincided with interglacial stages, particularly as most of these terrace deposits, both here 

and elsewhere along the Atlantic coast, contain what 

appear to be ice-borne boulders, including some of 

considerable size. 
100 .SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY, 1925 

These boulders at certain localities have been explained as indicating material dropped by icebergs 

from the north. This might be true for the erratics 

found in the "banks" of eastern North Carolina, 

though such an explanation of the presence of such 

erratics seems improbable. It is certainly not an 

adequate explanation of their presence at many places 

in the Pleistocene deposits, as, for example, in the 

drainage basin of Potomac River. I believe that these 

are to be explained as borne by river ice, although the 

smaller ones may have been carried in the roots of 

drifted trees. If this-explanation is sound it indicates 

a considerable severity of climate, with the formation of 

much ice, in the uplands along the Alleghany Mountains, which in fact seems to be necessary to explain 

the present stranded floras and faunas on the higher 

peaks even as far south as western North CarolinB. 

In regions where boulders are common in the subsoil, 

as is in a measure true of the Piedmont district of 

North Carolina, trees encompass many of the boulders 

in their root systems, and if such trees are overturned 

by windstorms within reach of rivers in flood periods 

such stones may be carried long distances. I collected 

a cobble in 1909 from the root of an Upper Cretaceous 

tree lignifiedin the Tuscaloosa formation of Alabama, 

which is an objective demonstration of this process 

in the past. 

After the storm of August, 1924, which uprooted 

many large trees in southeastern Connecticut, nearly 

every tree had one or more boulders tightly clasped 

by the roots, and these boulders already once glaciated 

would, if stream transportation had been available, 

have been carried out and eventually come to rest in 

fine marine muds. The largest boulder seen was in 

the roots of a tall pine 18 inches in diameter and had 

dimensions of ·55 by 19 by 20 inches. 

The following 27 Pleistocene species out of a total 

of 48, or considerably more than half, are forms that 

in the existing flora range long distances north and 

south of North Carolina: 

Sparganium sp. 

Hicoria ovata. 

Hicoria glabra. 

Populus cf. P. heterophylla. 

Carpinus caroliniana. 

Betula nigra. 

"' Fa_gus americana. 

Quercus velutina. 

Quercus prinus. 

Quercus palustris. 

Quercus alba. 

Quercus lyrata. 

Celtis occidentalis. 

Platanus occidentalis. 

Polygonum sp. 

Liriodendron tulipifera. 

Liquidambar styracifiua. 

Rubussp. -

Malus pseudoangustifolia. 

Malus coronariafolia. 

Cercis canadensis. 

Ilex opaca. 

Vitis sp. 

Xolisma ligustrina. 

Vaccinium corymbosum. 

Viburnum moUe. 

Viburnum nudum. 

Many of these species have extended their range 

northward into New England, southern Canada, and 

the States north of Ohio River since the retreat of 

the Wisconsin ice sheet, but at its maximum extent 

they appear to have approached it rather closely, at 

least in the Coastal Plain, where some of them are 

recorded as far north as New Jersey, and several were 

growing in southern Maine at a time when the rivers 

were still encumbered with valley ice. It is exceed-

.ingly difficult to evaluate the Pleistocene fossil plants 

of the Middle Atlantic States, because most of the 

species range long distances to the north and south, 

and considerable extensions or contractions of their 

range might have taken place without being registered 

in ~his middle region .. 

None of the North Carolina plants are definitely 

northern types or forms that in the existing flora are 

approaching their southern limit-of range in the. Carolina region. Quite the reverse is true. Of a total of 

48 recorded species the following 14 are at or near 

their northern limit of range as judged by the existing 

flora: 

Pinus serotina. 

Taxodium distichum. 

Hicoria aquatica. 

Quercus nigra. 

Quercus michauxii. 

Quercus phellos. 

Quercus predigitata. 

Ulmus alata. 

Planera aquatica. 

Crataegus spathulatoides. 

Persea pubescens. 

Nyssa bifiora. 

Dendrium pleistocenicum. 

Vaccinium arboreum . . 

This is nearly 30 per cent of the known Pleistocene 

flora of North Carolina and, if it means anything, 

indicates a milder climate than prevails in that area 

at the present time. 

It is true that there is considerable evidence that 

certain species, such as Taxodium distichum and 

Planera aq_-uatica, are becoming more restricted in the 

Coastal Plain at the present time, but this restriction 

appears to have followed a late Pleistocene or postglacial extension northward_, and certainly, for the 

Coastal Plain flora as a whole, the main movement has 

been one of a marked ·expansion northward on the 

heels of the retreating ice sheet. 

More than half of the fossil plants from North 

Carolina are forms that in the modern flora require 

wet habitats. These forms are indicated in the 

accompanying table. Some of them; such as Pinus 

serotina, Taxodium distichum, Sparganium, Hicoria 

aquatica, Quercus nigra, Quercus phellos, Quercus palustris, Planera aquatica, and Nyssa bijlora, prefer in 

modern times partly or permanently inundated bottoms. Others, like Salix, Betula nigra, Liquidambar 

styrac~fiua, Quercus michauxii, and !lex opaca, get 

along with a less co~stantly flooded substratum. 

Others not enum'erated among the 25 wet species~ like 

Fagus americana, Quercus lyrata, Quercus alba, L~riodendron tulipifera, and Oercis canadensis, are woodl_and 

forms of deep, moist soils. 

The dry-soil species are fewer in number and include 

Hicoria glabra, Quercus velutina, Quercus prinus, Celtis 

oc_cidentalis, and Yaccinium arboreum. There are 
PLEISTOCENE PLANTS FROM NORTH CAROLINA 101 

several forms that may be considered as occurring on 

both wet and dry soils. In this oa~egory are Quercus 

predigitata, Ulmus alata, Rubus, Vitis, X olisma ligustrina, and Dendri·um pleistocenicum. 

The Pleistocene flora of North Carolina fulfills the 

requirements that might have been expected from the 

areal distribution of the deposits in which it has been 

found. Each deposit consists of sediments laid down 

in an estuary, and most of the fossil plants are those 

that might be expected to occur in the bottoms of the 

lower courses of estuary streams. The few dryground forms represented as fossils either grew near 

at. hand on high and dry river bluffs or were brought 

from the more remote upper courses of these streams 

into .the basin of sedimentation occasionally, as 

through the agency of floods. 

The rna p showing the shore lines of the seas in which 

the formations were laid down (fig. 5) brings out with 

notable clearness the estuary character of the fossiliferous localities, especially in Bertie, Pitt, and Wayne 

counties. 

Plant geographers have always been interested in 

the representatives of the eastern North American 

deciduous-forest region which are found in the uplands 

of southern Mexico, Guatemala, and Costa Rica, and 

which obviously indicate a once continuous area of 

distribution, since broken by the conditions that produced the wide belt of arid country now found in the 

southwestern United States and northern Mexico. 

There is as yet little evidence for dating this southward extension of the mesophytic temperate flora of 

North America into the Tropics. Without predicating 

climatic changes for which there is no warrant in the 

available evidence we must conclude that it could not 

have occurred before the elevation of these Central 

American regions above the tierra caliente, or tropical 

and subtropical altitudinal zones. This event is 

believed to have occurred in . late Tertiary time, 

although the distribution of marine faunas shows 

clearly numerous earlier changes of level 

Among these upland genera occurring in Central 

America might be enumerated Abies, Pinus, Taxodium, 

Liquidambar, Quercus, Juniperus, Alnus, Salix, Populus, Celtis, Tilia, Arbutus, Morus, Vitis, Hicoria,. 

Fraxinus, Rhus, Juglans, Ulmus, Prunus, Cornus,. 

Cercis, Staphylea, Platanus, Arctostaphylos, Carpinus,. 

and perhaps others. There is good reason for believing that certain of these genera accomplished this 

southward radiation early in the Tertiary period, if. 

not in the Upper Cretaceous, and probably some, 

such as the Ulmaceae, invade~ North America from 

the south. 

Celtis occurs in the Eocene and Alnus and Rhus in 

the Pliocene of South America; Liquidambar is found 

in the Miocene of southern Mexico; and there is every 

reason to believe that of the genera enumerated above 

Pinus, Liquidambar, Alnus, Celtis, Rhus, and J uglans 

represent an old element in this southern region. 

Others that appear to be somewhat more recent are 

Taxodium, Quercus, Abies, Juniperus, Salix, Populus, 

Vitis, and possibly Ulmus. It is difficult to imagine 

physical conditions different enough from the present 

to permit Pleistocene dispersal, and on the other hand 

it is difficult to imagine species like those of Carpinus, 

Platanus, and Taxodium existing practically unchanged for a long time in two segregated regions of 

rather different environments. 

An attempt to portray the possible floristic regions 

at the period of maximum extent of the continental 

ice is shown on the accompanying sketch map (fig. 6). 

It is of course much generalized, being. based upon very 

inadequate knowledge, and it synthesizes a too long 

and too constantly changing period of time. It differs 

from previous maps of this nature in restricting the 

possible belt of northern coniferous and arctic plants 

to certain unspecified situations in the immediate 

vicinity of the ice front, and, except for a restriction 

of the central prairie region, in regarding the eastern 

and western North American mesophytic regions as 

having been much as they are at the present time, and 

with much the same association of species. 
102 

' 

SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY, 1925 

Habitat and range of Pleistocene plants from North Carolina 

Wet Dry 

Habitat 

Both wet and dry Woods 

Range 

Long 

distance 

both north 

and south 

Near 

northern 

limit of 

range 

Coniferophyta: 

Pinales: 

Pinaceae: 

Pinus serotina Michaux ______________________________ _ X -------------------------------- ax 

Cupressinaceae: \ 

Taxodium distichum (Linne) L. C. Richard _____________ _ X X X 

Angiospermophyta: 

Monocotyledonae: 

Pandanales: 

Sparganiaceae: · Sparganium sp __________________________________ _ 

Graminales: 

X 

Poaceae: . 

Chaetochloa SP----------------------------------- -------- ---------------- ---------------- --.. ----- Dicotyledonae: 

J uglandales: 

J uglandaceae: Hicoria ovata (Miller) Britton______________________ X _______________________ _ Hicoria glabra (Miller) Britton _____ _______________________ _ 

Salicales: 

Hicoria aquatica (Micha,ux son) Britton_____________ X 

Salicaceae: 

Fagales: 

Populus cf. P. heterophylla Linne __________________ _ Salix viminalifolia Berry __________________________ :. 

Salix bebbianaformis Berry ______________ _________ _ 

Betulaceae: 

X 

X 

X 

X 

X 

X 

X 

X 

Carpinus carolinian a Walter_______________________ X _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X _______ _ 

Betula nigra Linne________________________________ X ________ ________ ________ X -------- Betula pseudofontinalis Berry ______________________________ ------·-- ________________ ----------------

Fagaceae: Fagus americana Sweet_______ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X X _______ _ Quercus nigra Linne_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X 

velutina Lamarck_____________________________________ X ________ ________ X -------- abnormalis Berry ___________________ ____ _____________________________________________________ _ 

prinus Linne_________________________________________ X ________ ________ X --------

michauxii NuttalL---------------~------------ X ________________________ -------- X 

palustris DuRoL _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X _______ _ alba Linne _________________ .__________________ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X X _______ _ 

lyrata Walter________________________________ X ---------------- ________ X -------- phellos Linne_________________________________ X ________________________ -------- X 

predigitata Berry_____________________________________________ X ________ ________ X 

Urticales: 

Ulmaceae: 

Ulmus alata Michaux_____________________________________________ X _______________ _ 

Celtis occidentalis Linne__ _________________________________ X ________ ________ X -------- Planera aquatica (Walter) Gmelin (?)_______________ X _______________________________ _ 

Pla tan ales: 

X 

Platanaceae: 

Platanus occidentalis Linne _______________________ _ X X 

Polygonales: 

Polygonaceae: Polygonum __________ ___________________________ _ X X 

Ranales: 

Magnoliaceae: . Liriodendron tulipifera Linne ______________________________________________ ' X X 

&sales: 

Hamamelidaceae: 

Liquidambar styraciflua Linne ____________________ _ X X 

Rosaceae: 

Rubus SP----------------------------------------- _______________ _ Pomaceae: 

X X 

Malus pseudo-angustifolia Berry ______________________________________ -_____ ------·-- X _______ _ 

Malus coronariafolia Berry_________________________________________________________ X _______ _ Crataegus spathulatoides Berry____________________ X ________ ________ ________ ________ X 

Crataegus coccineafolia Berry _____________________________________________________________________ _ 

Caesalpiniaceae: . . Cercis canadensis Linne ________________ · __________________________________ _ 

Sapindales: 

Ilecaceae: 

· Ilex opaca Aiton~--------------------------------­ Rhamnales: 

Vitaceae: 

X 

Vi tis sp ______________ .- _________________________________________ _ 

• At northern limit 

X 

X X 

X 

-x 
PLEISTOCENE PLANTS FROM NORTH CAROLINA 

Habitat and range of Pleistocene plants from North Carolina-Continued 

Habitat 

Wet Dry 

Angiospermpphyta-Continued. 

Dicotyledonae-Continued. 

Thymeleales: 

Lauraceae: 

Persea pubescens (Pursh) Sargent _________________ _ X 

Umbellales: 

Cornaceae: 

Nyssa biflora Walter _____________________________ _ X 

Ericales: 

Ericaceae: 

Xolisma ligustrina (Linne) Britton _________________________________ _ Dendrium pleistocenicum Berry ___________________________________ _ 

Vacciniaceae: · . 

Both wet and dry 

X 

X 

Woods 

103 

Range 

Long 

distance 

both north 

and south 

X 

Near 

northern 

limit of 

range 

X 

X 

X 

Vaccinium corymbosum Linne_____________________ X _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X _______ _ spatulata Berry _______________________________________________________________________ _ 

arboreum MarshalL _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X -------- -------- -------- X 

Rubia]es: 

Caprifoliaceae: Viburnum molle Michaux (?) _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ X X 

Viburnum nudum Linne(?) ________________________ X \ X -------- -------- --------
FIGURE 6.-Map of North America showing the glaciated region, the floral geography, and source of the postglacial directions of 

dispersal (shown by arrows). 

104 
PLEISTOCENE PLANTS FROM NORTH CAROLINA 105 

SYSTEMATIC DESCRIPTIONS 

CONIFEROPHYTA 

Order PIN ALES 

Family PIN ACEAE 

Genus PINUS Linne 

Pinus serotina Michaux 

Plate XLV, Figure 9 

Pinus rigida Berry (not Miller), Jour. Geology, vol. 15 p. 339 

1907. ' ' 

Leaves in threes, sometimes with sheaths preserved. 

These leaves were ref erred by me to Pinus rigida in 

1907, as they· seemed to be shorter than those of 

Pinus serotina. Subsequently cones were collected 

which in their smaller size, thinner scales and slender . ' ' easily abraded prickles ally the leaves with P. serotina instead of P. rigida, which is more at home in the 

plant association recorded from the Coastal Plain of 

North Carolina. 

Needles from the locality on Neuse River in Wayne 

County; needles and numerous cones from the locality 

on Tar River in Pitt County. 

According to Pinchot and Ashe Pinus serotina in 

the e.x:isting flora does not extend up Tar River beyond 

Washington. It has not previously been found fossil. 

Pinus rigida Miller in North Carolina is confined to 

the western Piedmont Plateau region and the mountain counties south of French Broad River, growing 

on dry, commonly sandy or gravelly ridges. 

Pinus .sero~ina ~ichaux, the pond, swamp, loblolly, 

or. pocosin pine, Is found on low peaty or wet sandy 

smls from North Carolina to Florida near the coast. 

In North Carolina it is practically confined to the 

Coastal . Plain south of Albemarle Sound and reaches 

west to the fall line only -in the valleys of Neuse and 

Cape Fear rivers. 

Occurrence: Chowan formation, right bank of 

Neuse River, 4Y2 miles above Seven Springs, Wayne 

Coun~y; Dupre~ Landing, Tar River, Pitt ,County. 

Pamhco formatwn, below New Bern, Neuse River, 

Craven County. 

Family CUPRESSIN ACEAE 

Genus TAXODIUM Richard 

Tax odium distichum (Linne) L. C. Richard 

Plate XLV, Figures 1-8 

Taxodium distichum Holmes, Elisha Mitchell Soc. Jour., vol. 2, 

p. 92, 1885 . . 

Hollick, Maryland Geol. Survey, Pleistocene, pp. 218, 237, 

pl. 68, 1906. 

Berry, Torreya, vol. 6, p. 89, 1906; Jour. Geology, vol. 15, 

p. 339, 1907; Am. Naturalist, vol. 43, pp. 432-434, figs. 

1, 2, 1909; Am. Jour. Sci., 4th ser., vol. 29, p. 391, 1910; 

Torreya, vol. 10, p. 263, 1910; Plant World, vol. 16, pp. 

39-45, figs. 1, 2, 1911; Am. Jour. Sci., 4th ser., vol. 34, p. 

219, figs. 1, 2, 1912; Torreya, vol. 14, pp. 160, 162, 1914; 

U. S. Nat. Mus. Proc., vol. 48, p. 296, 1915; Torreya, 

vol. 15, p. 206, 1915; U.S. Geol. Survey Prof. Paper 98, p. 

195, pl. 45, figs. 1-6, 1916; Jour. Geology, vol. 25, p. 

662, 1917; Florida Geol. Survey Ninth Ann. Rept., p. 21, 

1917; Washington Acad. Sci. Jour., vol. 14, p. 15, pl. 1, 

figs. 37-42; pl. 3, 1924. 

It is probably not always possible to differentiate 

fossil specimens of the common bald cypress from 

the pond cypress, Taxodium imbricarium (for example, 

from cones or seeds alone); nevertheless it appears 

significant that among the very numerous Pleistocene occurrences of Taxodium that have been 

recorded, wherever the stumps or twigs are preserved, 

these represent Taxodium distichum and not Taxodium 

imbricarium. The numerous twigs and stumps found 

in the North Carolina Pleistocene belong to Taxodium 

distichum. It may be legitimately inferred from· this 

circumstance that Taxodium imbricarium had not yet 

been ·differentiated at that time. If we assume that 

the pond cypress was already in existence, its absence 

from the fossil record at so many localities along the 

Atlantic coast is a strong argument for the method of 

genesis of the · Pleistocene terraces advocated in the 

present report-namely, by marine control due to 

changes in relative level. If these terraces were 

purely the result of stream control and continental 

deposition, as is advocated in some quarters, it is 

almost impossible to conceive that somewhere the 

pond cypress would not have been· preserved. 

The bald cypress has a .most interesting geologic 

history, and its immediate and scarcely distinguishable 

Tertiary ancestor attained a holarctic distribution in 

the later Tertiary. 

The oldest beds in which the existing species has been 

definitely_ recognized are the Pliocene deposits along 

the Gulf coast of Alabama (Citronelle formation). It 

was exceedingly common at numerous widespread lo~ 

calities ih southeastern North America during the 

Pleistocene epoch. Unfortunately it is not possible to 

correlate the outcrops at these localities with the 

chronology of the glaciated region to the- .north. 

Several of these Pleistocene localities are farther north 

or farther inland than the existing limits of range of 

the species, which -is from so:uthern Delaware and 

southern Maryland to extreme so~thern Florida and 

Texas. In North Carolina it is restricted to the 

Coastal Plain regiqn. The leafy twigs and many seeds 

are present at the Neuse River locality in Wayne 

County; stumps, knees, and burls at the locality in 

Craven County-; and leafy twigs, cone scales, and a 

staminate ament at thelocality in Bertie County. 

· Occurrence: Ch,owan formation, right bank of 

Neuse River 4Y2 miles above Seven Springs, Wayne 

County. Pamlico formation, Old Mill ·Landing, 

Roanoke River, Bertie County, and below New Bern 

Neuse River, Craven County. ' 1 
106 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY, 1925 

ANGIOSPER:MOPHYTA 

Class :MONOCOTYLEDONAE 

Order P AND AN ALES 

, Family SP ARGANIACEAE 

Genus SP ARGANIU:M Linn~ 

Sparganium sp. 

Sparganium sp. Berry, Torreya, vol. 14, p. 160, 1914. 

A nutlike fruit of some species of Sparga'nium is 

present at the locality on Neuse River in Wayne 

County. The determination was made for 1me by 

W. L. McAtee. It probably represents the existing 

Sparganium androcladum (Engelmann) Morong. 

The genus Sparganium goes back to the Eocene and 

possibly earlier. It is common in the European Pleistocene and was presumably common in that of North 

America, although the records are few, includrng only 

this one and Sparganium eurycarpum Engelmann, recorded from the Wicomico formation at W as~ington, 

D. C. 4 . 

Occurrence: Chowan formation, right bank of 

Neuse River 4Y2 miles above Seven Springs, Wayne 

County. -

Order GRA:MINALES 

Family POACEAE 

Genus CHAETOCHLOA Scribner 

Chaetochloa sp. Berry I 

Chaetochloa sp. Berry, Torreya, vol. 14, p. 160, 1914. 

A caryopsis of some grass, identified by W. L. 

McAtee as a species of Ohaetochloa, is contained in the 

collection from the Neuse River locality in Wayne 

County. · · ' 

The genus Ohaetochloa 5 includes Setaria Beauvois 

lxophorus Nash, and various species refe~red t~ Panicum, of which there are many species 1 

in our 

existing Coastal Plain flora. · 

Occurrence: Chowan formation, right bank of 

Neuse River 4 Y2 miles above Seven Springs, 'Wayne 

County. 

Class DICOTYLEDONAE 

Order JUGLANDALES 

Family JUGLANDACEAE 

Genus IDCO.RIA 

Hi coria glabra (Miller) Britton 

Plate XLVI, Figures 1-4 

I . 

Co:rya ·porcinp, Nuttall. Mercer, Acad. Nat. Sci. Philadelphia 

Jour., 2d ser., vol. 111 pp. 277, 281, figs. 84, s, 12, 116, 1899. 

Hicoria pseudoglabra Hollick, Maryland Geol. Survey, ,Pleistocene, p. 221, pl. 72, figs. 1, 16, 17, 1906. 

4 Be~ry, E. W.j Washington Acad. Sci. Jour., vol. 14, p. 16, pl. 1, fig.l36, 1924. 6 Scnbn,er, F. ll., U.S. Dept. A.gr. Div. A.grostology Bull. 4, p. 38, 1897. 

Hicoria glabra Berry, Torreya, vol. 6, p. 89, 1906; Jour. Geology, vol. 15, p. 340, 1907; Tor'"eya, vol. 9, p. 97, figs. 1-5, 

1909; vol. 10, p. 264, fig. 1, 1910. 

Fragmentary leaflets of this species are not uncommon at the locality on Neuse River in Wayne County, 

and several of these are figured. They are associated 

with several husks of nuts and one flattened nut and 

undoubtedly represent Hicoria glabra. Nuts of this 

species are not uncommon in the Pleistocene deposits 

of the Atlantic Coastal Plain and have been recorded 

from the Talbot· formation of New Jersey and Maryland, from the Sunderland formation of Maryland, 

and from the Port Kennedy cave in Pennsylvania. 

In the recent flora this species ranges from Maine 

to Minnesota and southward to Florida and Texas. 

In North Carolina, although found throughout the 

State on dry soils of ridges, it occurs chiefly in the 

Piedmont Plateau region, being uncommon in the 

mountains and rare in the Coastal Plain. 

Hicoria is not uncommon in the Pleistocene. In 

addition to the two other species recorded from North 

Carolina, Hicoria alba occurs in Pennsylvania and 

Canada, Hicoria villosa occurs in Alabama, Hicoria 

minima occurs in Massachusetts, and several specifically undetermined nuts and leaflets have been recorded 

from Maryland and Alabama. 

Occurrence: Chowan formation, right bank of Neuse 

River 4% miles above Seven Springs, Wayne County. 

Hicoria ovata (Miller) Britton 

Plate XL VI, Figures 6, 7 

Carya ovata Miller. Mercer, Acad. Nat. Sci. Philadelphia 

Jour., 2d ser., vol. 11, p. 279, fig. 86, 1899. 

Hicoria ovata Britton. Berry, Jour. Geology, vol. 15, p . . 340, 

1907. 

Characteristic terminal leaflets of this species are 

present at the Neuse River locality in Wayne County. 

No nuts were collected, but nuts of this species found 

in the Port Kennedy cave in Pennsylvania were described by Mercer. 

In the existing flora this species ranges from Quebec 

to Minnesota and southward to Florida and Texas. 

· It is found on rich, deep, moderately moist soil near 

streams and swamps. In N 0rth Carolina it is nowhere 

common, OCC\lrring chiefly west of the fall line but in 

part across the Coastal Plain in the valleys of Roanoke 

and Cape Fear rivers. 

Occurrence: Chow an formation, right bank of Neuse 

River 4% miles above Seven Springs, Wayne County. 

Hicoria aquatica (Michaux son) Britton 

_· Plate XLVI, Figure 5 

Salix sp. Berry, Jour. Geology, vol. 15, p. 340, 1907. 

Hicoria sp. cf. H. microcarpa (Nuttall) Britton. Berry, Idem. 

Hicoria aquatica Berry, Torreya, vol. 9, p. 71, 1909; vol. 14, 

pp. 161, 162, 1914. 

This species is represented by a single small but 

characteristic lateral leaflet and several nuts from 
PLEISTOCENE PLANTS FROM NORTH CAROLINA 107 

the locality on Neuse River in Wayne County. As the 

vernacular name water hickory indicates, this is a species that inhabits frequently inundated river swamps. 

Its range at the present time extends from south-. 

eastern Virginia to Florida and Texas, extending up 

the Mississippi Valley to the mouth of the Ohio but 

there as well as elsewhere being confined to the Coastal 

Plain. In North Carolina it occurs chiefly on the 

peninsula between Albemarle and Pamlico sounds 

and in Pender and Brunswick counties in the valleys 

of Northeast 9ape Fear and lower Cape Fear rivers. 

It indicates what the areal distribution of the 

Chowan formation along Neuse River showsnamely, that at that stage of the Pleistocene a drowned 

river valley like that of the present-day lower Cape 

Fear extended across Lenoir County and halfway. 

across Wayne County. , 

Occurrence: Chowan formation, right bank of 

Neuse River 4V2 miles .above Seven Springs, Wayne 

County. 

Family SALICACEAE 

Genus POPULUS Linne 

Populus? cf. P. heterophylla Linne 

Bud scales similar to those of Populus but possibly 

referable to Fagus, leaves of which are so common, 

occur in the collections from the locality on Neuse 

River in Wayne County. They are exactly like those 

of Populus heterophylla Linne, which at the ·present 

time is found in river swamps from Connecticut to 

Georgia and Louisiana and is not uncommon in such 

situations in the Coastal Plain region of North Carolina. Because of the absence of leaves of Populus in 

the deposit and the great abundance of leaves of 

Fagus and Betula, which have rather similar resinous 

bud scales, I have queried the generic reference. In 

examining the recent river deposits of the Coastal 

Plain rivers of North Carolina I have frequently 

noticed bud scales of Fagus but have not observed 

those of Populus. Populus leaves are, however, 

recorded from the Pleistocene of Maryland and 

Alabama. -

Occurrence: Chowan formation, right bank of 

Neuse River 4 Y2 miles above Seven Springs, Wayne 

County~ 

Genus SALIX Linne 

Salix bebbianaformis Berry, n. sp. 

Plate XLVI, Figure 8 

Leaves of rare occurrence, ovate-lanceolate, with 

pointed apex and base, entire margins, thin texture. 

Midvein medium in size. Secondaries thin, ascend· 

ing, about ·'five regularly spaced, camptodrome pairs. 

This species appears to be a Pleistocene willow. 

Its relation to existing species is entirely problematic, 

and the name given to it indicates merely a resemblance in leaf form without implying any close botanic 

relationship. 

Salix bebbiana Sargent is a moist-soil species, 

chiefly of stream and swamp margins, of the northern 

United States and Canada, and if the fossil could be 

more precisely correlated with this species, it would 

afford an interesting problem of distribution. 

Occurrence: Chowan formation, right bank of 

Neuse River 4V2 miles above Seven Springs, Wayne 

County. 

Salix viminalifolia Berry 

Plate XLVII, Figures 1-4 

Salix viminalifolia Berry, U.S. Nat. Mus. Proc., vol. 48, p. 297, 

1915. 

This species was named from its resemblance to the 

osier willow, Salix viminalis Linne, a native of Eurasia, 

not rare in European Pleistocene deposits, commonly 

cultivated in our Eastern and Middle States, and here 

and there escaped in wet places. The name can not 

be taken as. indicative of relationship between the 

living and fossil species, for the position of the latter 

is uncertain. 

The fossil species may be described as follows: 

'Leaves elongated, 1inear-lanceolate, abruptly acute 

at the apex and base. Margins usually somewhat 

undulate, entire in all the specimens seen. Petiole 

stout, curved, expanded proximad, 6 to 10 millimeters 

long. Length about 15 centimeters. Maximum 

width about 1.5 to 2.25 centimeters. Midvein stout, 

somewhat flexuous. Secondaries thin, numerous, 

camptodrome. 

Occurrence: Chowan formation, right bank of 

Neuse River, 4V2 miles above Seven Springs, Wayne 

County (common). 

Order FAGALES 

Family BETULACEAE 

Genus BETULA Linne 

Betula nigra Linne 

Plate XLVII, Figures .~ 19 

Betula nigra Knowlton, Am. Geologist, vol. 18, p. 371, l896. 

Berry, Jour. Geology, vol. 15, p. 341, 1907; Am. Naturalist, 

vol. 41, p. 692, pl. 2, figs. 2-4, 1907; vol. 43, p. 435, 

1909; Am. Jour. Sci., 4th ser., vol. 29, p. 393, 1910; 

Torreya, vol. 14, p. 162, 1914. 

Characteristic leaves of the black or river birch, of 

all sizes, are common in the . North Carolina Pleistocene. Their permutations in size and outline are 

shown in the accompanying figures. The species 

inhabits swamps and river bottoms and, in the existing flora, ranges from Massachusetts to Minnesota 

and Kansas and southward to Florida and Texas. 

In North Carolina it is found along streams and 

swamp borders throughout the Coastal Plain and 

Piedmont regions. 
108 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY, 1925 

The genus goes back to Upper Cretaceous time and 

became much diversified in the early Tertiary. 

Several existing species made their appearOtnce in the 

Pleistocene of this country and Eurasia. 

Betula nigra has been reported from the Pleistocene 

of West Virginia, Virginia, Georgia, Alabama, · and 

Mississippi and was evidently a common and widespread type in the country bordering the coast of the 

Pleistocene sea in southeastern North America. 

Occurrence: Chowan formation, right bank of 

Neuse River, 4% miles above Seven Springs, Wayne 

County (abundant leaves and bark fragments); 

Patnlico formation, Old Mill Lanlling; Roanoke River, 

Pitt County (leaves). 

Betula pseudofontinalis Berry 

Plate XLVIII, Figures 1, 2 

Betula pseudofontinalis Berry, Jour. Geology, vol. 15, p. 341, 

1907. 

Leavef? broadly ovate, with a rounded or bluntly 

pointed apex and a broadly truncated, rounded, entire 

base. Margins somewhat irregularly serrate or dentate above. Texture· subcoriaceous. Petiole long. 

Midvein stout. Secondaries stout, rather openly and 

somewhat irregularly spaced, craspedodrome. 

This.is a somewhat poorly defined form, named from 

its resemblance to the leaves of the existing western 

Betula fontinalis Sargent, but the name is not intended 

to imply a direct botanic relationship with that species. 

·Occurrence: Chowan formation, right bank of 

Neuse River 4 Y2 miles above Seven Springs, Wayne 

County. 

Genus CARPINUS Linne 

Carpinus caroliniana Walter 

Plate XLVII, Figures 5--7 

Carpinus caroliniana Walter. Berry, Jour. Geology, vol. 15, p. 

340, 1907; Am. Naturalist, vol. 41, p. 692, vol. 41, p. 692, 

pl. 1, 'figs. 8, 9, 1907; Am. Jour. Sci., 4th ser., vol. 29, 

p. 395, 1910. 

Carpinus pseudocaroliniana Hollick,· Maryland Geol. Survey, 

Pleistocene, p. 225, pl. 71, fig. 10,. 1906. 

T,he hornbeam, or blue beech, was apparently a 

common member of the Pleistocene forest flora of 

southeastern North America and has been recorded 

from the early Pleistocene (Sunderland) of Maryland 

and from later Pleistocene deposits in North Carolina, 

Georgia, and Alabama. In North Carolina it is 

represented by a small number of characteristic 

leaves. 

In the existing flora it grows in moist woods in the 

vicinity of streams and swamps and ranges from 

Nova Scotia· westward to Georgian Bay and Minnesota and southward to Florida and Texas, reappearing 

in the uplands of southern Mexico and Central America. 

In North Carolina it occurs along streams and in wet 

situa~ions throughout the State. 

There are about a dozen living species in central 

and southern Europe, Asia, and eastern North America. 

The genus appears in the Eocene and is exceedingly 

common in the holarctic Tertiary floras. 

Occurrence: Chowan formation, right bank of 

Neuse River 4% miles above Seven Springs, Wayne 

County. 

Family FAGACEAE 

Genus FAGUS Linne 

Fagus americana Sweet · 

Plate XL VIII, Figures 3-13 

Fagus ferruginea Aiton. Knowlton; Am. Geologist, vol. 18, 

p. 371, 1896. 

Mercer, Acad. Nat Sci. Philadelphia Jour., 2d ser., vol. 

11, pp. 277, 281, figs. 8-15, 1S99. 

Emerson, U. S. Geol. Survey Bull. 597, p. 148, 1917. 

Fagus americana Sweet. Berry, Torreya, vol. 6, p. 88, 1906. 

Hollick, Maryland GeoL Survey, Pleistocene, p. 226, 1906. 

Berry,.Jour. Geology, vol. 15, ·p. 341, 1907; Am. Naturalist, 

vol. 41, p. 692, pl. 2, fig. 7, 1907; vol. 43, p. 435, 1909; 

Am. Jour. Sci., 4th ser., vol. 29, p. 393, 19.10; Torreya, 

vol. 14, p. 162, 1914; vol. 15, p. 206, 1915. 

The American beech, sometimes named Fagus 

g;andifolia Ehrhart, on the ground of priority, is oneof 

the commonest members of the Pleistocene flora . in 

southeastern North America. 

Leaves, nuts, or husks have been recorded from 

rather numerous localities in Massachusetts, Pennsylvania, Maryland, Virginia, Alabama, and Mississippi. 

In the North Carolina Pleistocene both leaves and 

husks are common. The leave's show considerable 

variation in size and outline, as shown by the accompanying illustrations. In general they appear to 

have been smaller than the average size of the modern 

leaves of this sp.ecies. 

In the existing flora the beech occurs chiefly in 

deep, damp woods, ranging from Nova .Scotia to 

::(.;ake Huron and southward to Florida and Texas. 

In N o:rth Carolina it occurs throughout the State and 

is said by Pinchot and Ashe to be smaller and less 

abundant in the Coastal Plain than in the Piedmont 

and mountain regions. 

Occurrence: Chowan formation, right bank of 

Neuse River 4% miles above Seven Springs, Wayne 

County. 

Genus QUERCUS Linne 

Quercus alba Linne 

Plate .L, Figures 1-5 

Quercus alba Linne. Penhallow, Roy. Soc. Canada Trans., 

vol. 10, sec, ~' p. 74, 1904; Am. Naturalist, vol. 41, 

p. 448, 1907. 

Mercer, Acad. Nat. Sci. Philadelphia Jour., 2q ser., vol. 11, 

p. 281, 1899. . 

Berry, Jour. Geology, vol. 15; p. 342, 1907; Am. Jour. 

Sci., 4th ser., vol. 34, p. 221, 1912. 

Quercus pseudoalba Hollick, Maryland Geol. Survey, Pleist(}o 

cene, p. 227, pl. 70, fig. 2; pl. 71, figs. 1-6, 1906. 
PLEISTOCENE PLANTS FROM NORTH CAROLINA · 109 

The white oak is not uncommon in the Pleistocene, 

having been recorded from the old Pleistocene (Sunderland) of Maryland, from the interglacial beds of the 

Don Valley in Ontario, from the Great Valley of 

Virginia, and from postglacial deposits in Massachusetts. Its lea:ves, both young and old forms; are 

common in the Pleistocene of North Carolina. 

In the existing flora the white oak is found in woods 

from Maine to Ontario, westward to Minnesota and 

Nebraska, and southward to Florida and Texas. 

In North Carolina it occurs throughout the State but 

is most abundant on the Piedmont Plateau, being 

rarely found in the Coastal Plain south of Neuse 

River or on the latest Pleistocene Pamlico terrace 

soils of the eastern part of the State. 

Occurrence: Chowan formation, right bank of 

Neuse River 4Y2 miles above Seven Springs, Wayne 

County. 

Quercus abnormalis Berry 

Plate LI, Figure 24 

Quercus abnormali.~ Berry, Jour. Geology, vol. 15: p. 342, 1907. 

This identification is based upon the single abnormal 

oak leaf figured. This specimen is incomplete but 

indicates a linear-lanceolate leaf divided near the 

middle into two linear-lanceolate lobes that diverge 

at an angle of about 50°. The margins are entire 

so far as seen, and the base and tips were probably 

acute. The midvein is stout and prominent, and 

numerous stout secondaries diverge from· it at wide 

angles and are camptodrome. The texture is coriaceous, and all the observed features stamp it as a form 

of Quercus, possibly a hybrid or more probably · 

an abnormality. Estimated length about 12 centimeters. Width about 1.5 centimeters. 

Occurrence: Chowan formation, right bank of 

Neuse River 4Y2 miles above Seven Springs, Wayne 

· County. 

Quercus predigitata Berry 

Plate LIII, Figures 1-6 

Quercus falcata Michaux. Knowlton, Am. Geologist, vol. 18, 

p. 371, 1896. 

Quercus predigitata Berry, Jour. Geology, vol. 15, p. 342, 1907; 

Am. Jour. Sci., 4th ser., vol. 34, p. 22, figs. 4, 5, 1912; 

Torreya, vol. 14, p. 162, 1914; vol. 19, p. 9, 1919. 

This oak is considered to be the ancestral type from 

which the Spanish oak and the water Spanish oak 

were differentiated in dry and wet environments. At 

any rate the fossil leaves show gradations intermediate 

between the leaves of these modern species, and their 

precise differentiation in the fossil state is impossible. 

The species has a considerable range in the Pleistocene of southeastern North America, being recorded 

from West Virginia, the Great Valley of Virginia, 

western Tennessee, and Mississippi. In North Carolina its leaves are common and variable in the deposits 

of the Chowan formation on Neuse River and · are also I 

present in the earlier Wicomico formation near Weldon, 

both localities near the heads of drowned Pleistocene 

streams. 

The existing Quercus digitata Sudworth is a tree of 

dry soils, ranging from southern New Jersey to central 

Florida and westward to Texas. In North Carolina 

it is not uncommon throughout the State except in the 

western mountains and in the eastern part, which is 

underlain by the Pamlico formation. 

Quercus pagodaefolia Ashe, on the other hand, is a 

species of rich wet bottom lands and ranges from 

Virginia to northern Florida and the lowerMississippi 

Valley. It is common throughout the Coastal Plain 

in North Carolina. 

The recorded range and habitats of the fossil species 

show clearly that it is either the ancestor of these two 

species or that the fossil records represent both of 

these species, which have not been discriminated. It 

appears not improbable that with the Pleistocene submergence of much of the Coastal Plain and its subsequent emergence Quercus predigitata diverged into the 

two familiar dry and wet soil types. · 

Occurrence: Chowan formation, right bank of 

Neuse River 4,Y2 mil~s above Seven Springs, Wayne 

County. . Wicomico formation, 1 ~ miles ~ast· of 

Weldon; Northampton County. 

Quercus palustris DuRoi 

Plate XLIX, Figures 4-6 

Quercus palustris DuRoi. Mercer, Acad. Nat. Sci. Philadelphia 

Jour., 2d ser., vol. 11, pp. 277, 281, fig. 8 2, 1899. 

Berry, Jour. Geology, vol. 15, p. 342, 1907; Torreya, vol. 

15, p. 207, 1915. 

This species is represented by characteristic leaves 

in the Chowan formation of North Carolina. It has 

been previously recorded from the Pleistocene deposits 

of the cave at Port Kennedy, Pa., where it is represented by a cupule, and the Talbot formation of 

Maryland. 

In the existing flora this species occurs in swamps 

and w'et places from Massachusetts westward to 

Wisconsin and southward to Florida. ·It is not 

uncommon in the Coastal Plain of North Carolina. 

Occurrence: Chowan formation, right bank of 

Neuse River 4_Y2 miles above Seven Springs, Wayne 

County. , 

Quercus nigra Linne 

Plate LI, Figures 1-7 

Quercus nig~a~·nne. Be:ry, Jour. · Geology, vol. 15, p. 342, 

· 1907, . Naturalist, vol. 41, p. 693, pl. 1, figs. 3, ~ 

1907; A . Jour. Sci., 4th ser., vol. 29, p. 394, 1910; 

U. S. Gefl. Survey Prof. Paper 98, p. 201, pl. 46, fig. 11, 

1916. . 

Quercus maryla dica Berry, Jour. Geology, vol. 15, p. 342, 1907. 

Leaves of_this species are common in the Chowan 

forma.tion, a'd some .of ~he larger ones wer~ r~ferred 

to Quercus ryland~ca In 1907, although It Is now 

J 
110 SHORTER C(\NTRIBUTIONS TO GENERAL GEOLOGY, 1925 

believed that all these represent the leaves of the 

water oak. They show considerable variation in outline and in the tendency toward a trilobate form, as, 

for example, in the leaf shown in Figure 1; others 

show an unusual narrowing and elongation of the 

base, as in Figures 3 and 6. 

The oldest deposit in which this species has been 

found is the Pliocene Citronelle formation of southern 

Alabama, and its leaves and cupules occur at several 

Pleistocene localities in Alabama. 

In the existing flora it is found on sandy wet soil 

along streams and swamp borders from southern 

Delaware southward to Florida and eastern Texas 

and up the Mississippi Valley to the mouth of the 

Ohio. In North Carolina it is common throughout 

the Coastal Plain and in certain areas in the southeastern part of the Piedmont region. 

Occurrence: Chowan formation, right bank of 

Neuse River 4 ~ miles above Seven Springs, Wayne 

County. 

Quercus phellos Linne 

Plate LI, Figures 8-23 

Quercus phellos Linn~. Berry, Jour. Geology, vol. 15, p. 342, 

1907; Am. Naturalist, vol. 41, p. 694, pl. 1, fig. 1, 1907; 

. Am. Jour. Sci., 4th ser., vol. 29, p. 394, 1910; Torreya, 

vol. 10, p. 265, 1910; vol. 14, pp. 161, 162, 1914. 

The willow oak is not uncommon in the Pleistoc'ene 

o~ southeastern North America and has been previously recorded from New Jersey, Alabama, and 

Mississippi. Both leaves and acorns have been found 

in the Pleistocene of N ~rth Carolina. 

The great variety of leaves of this species are partly 

shown in the accompanying illustrations. None 

found can be attributed to the water . oak, Quercus 

laurijolia Michaux, which might be expected to occur 

in the North Carolina Pleistocene but which has not 

been encountered. 

In the existing flora Quercus phellos is common in 

low wet situations bordering streams and swamps and 

to some extent on · moist sandy uplands from Long 

Island to Florida and thence westward to eastern 

Texas and up the Mississippi Valley to southeastern 

Missouri. In North Carolina it is exceedingly common in the Coastal Plain and is sparingly present in 

the Piedmont region. 

Occurrence: Chowan formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County (leaves, common); Dupree Landing, Tar 

River, Pitt County (acorns and cupules). 

Quercus prinus Linne 

Plate LII, Figures 5-8; Plate LIV, Figure 9. 

Quercus prinus Linne. Berry, Jour. Geology, vol. 15, p. 342, 

1907; Am. Naturalist, vol. 41, p. 693, pl. 1, fig. 1, 1907; 

Am. Jour. Sci., 4th ser., vol. 29, p. 394, 1910; Pan-Am. 

Geologist, vol. 41, p. 106, 1924. 

I am not sure that this species can be distinguished 

from Quercus michaux# by the leaves in the fossil 

state, but inasmuch as characteristic acorns are 

associated with the leaves in the Pleistocene of Alabama, I have considered the narrower, longer-petioled 

leaves of the Pleistocene of North Carolina to represent Quercus prinus. 

In the existing flora the rock chestnut oak is found 

in rocky woods from Maine and Ontario to Georgia 

and Alabama. In North Carolina it is a denizen of 

dry soils and is found chiefly in the western part of 

the State, being replaced in the Coastal Plain by the 

swamp chestnut oak, Quercus michauxii, which is also 

abundant in the Pleistocene forest flora of the State. 

Inasmuch as there was a crowding together of presentday Coastal Plain and Piedmont types during the 

Ph~istocene epoch, when considerable areas of the 

Coastal Plain were submerged, and a less clear, 

demarcation between dry and wet soil types, as shown 

in several localities, the probability is strong that 

these fossil leaves represent Quercus prinus, although 

this identification can not be said to be absolutely 

proved. Quercus prinus has been recorded from postglacial deposits near Staunton, Va. 

Occurrence: Chowan formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County . Quercus lyrata Walter 

Plate L, Figures 6-9 

Quercus lyrata Walter. Berry, Jour; Geology, vol. 15, p .. 342, 

1907. 

Characteristic leaves and four flattened acorns of 

this species, which is not otherwise known in the 

fossil state, were collected from the North Carolina 

Pleistocene. It is possible that the acorns from the 

Port Kennedy cave in Pennsylvania, identified by 

Stewardson Brown 6 as Quercus macrocarpa Michaux, 

may represent. Quercus lyrata, as the mossy cup or bur 

oak is a northern species. 

The swamp post oak or overcup oak, Quercus 

lyrata, is found in swampy habitats in the existing 

flora and ranges from New _Jersey and Missouri to 

Florida and Texas. In North Carolina it is found 

throughout the ' Coastal Plain and in the southeastern 

part of the Piedmont region of the State. 

Occurrence: Chowan formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County. 

Quercus michauxii Nuttall 

Plate LII, Figures 1-4 

Quercus michauxii Nuttall. Berry, Torreya, vol. 9, p. 71, 

1909; vol. 15, p. 207, 1915. 

··Quercus prinoides Berry, Jour. Geology, .vol. 15, p. 343, 1907. 

Quercus platanoides Berry, idem. 

Leaves and cupules of this species are found in the 

Chowan formation of North Carolina, and the species 

is also recorded from the . Talbot formation of Mary4 Mercer, H. C., Acad. Nat. Sci. Philadelphia Jour., 2d ser., vol. 11, pp. 278, 281, 

fig. 8, 1899. 
PLEISTOCENE PLANTS FROM NORTH CAROLINA Ill 

land. To it I now refer specimens from North Carolina that were formerly referred to Quercus prinoides 

and Quercus platanoides. 

The cow oak or swamp chestnut oak is a dweller 

on moist soils such as swamp borders and river bottoms. In the existing flora it ranges, chiefly in the 

Coastal Plain, from Delaware to northern Florida 

and through the Gulf States to Trinity River in 

Texas and up the Mississippi Valley to southeastern 

Missouri and the lower Wabash. In North Carolina 

it occurs abundantly throughout the Coastal Plain 

and in the swamps of the eastern half of the Piedmont 

Plateau region. 

Occurrence: Chowan formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County. 

Quercus velutina Lamarck 

Plate XLIX, Figures 1-3 

Quercus velutina Lamarck. Penhallow, British Assoc. Adv. 

Sci. Rept., 1900, p. 336. 

Quercus tinctoria Bartram. Penhallow, Roy. Soc. Canada 

Trans., vol. 10, sec. 4, p. 74, 1904. 

This species is represented in the Pleistocene of 

North Carolina by both leaves and cupules. The 

only fossil occurrence previously recorded is that 

in the interglacial deposits of the Dan Valley in 

Ontario. 

In the modern flora this species, which has probably 

the most variable leaves of all the black oaks, is found 

on dry soils from Maine and Ontario to Minnesota 

and Nebraska and southward to northern Florida 

and central Texas. In North Carolina it is a common 

tree in the Piedmont and mountain regions but is 

less abundant in the Coastal Plain. 

Occurrence: Chowan formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County. Wicomico formation, 1 U miles east of 

Weldon, Northampton County. 

Order URTICALES 

Family ULMACEAE 

Genus ULMUS Linne 

Ulmus alata Michaux 

Plate LIV, Figures 1, 2 

Ulmus alata Michaux. ?Lesquereux, Am. Jour. Sci., 2d ser., 

vol. 27, p. 365, 1859. 

Berry, Jour. Geology, vol. 15, p. 343, 1907; Am. Naturalist, vol. 41, p. 694, pl. 1, figs. 6, 7, 1907; Am. Jour. 

Sci., 4th ser., vol. 29, p. 396, 1910. 

Leaves of the wahoo or winged elm are not uncommon in the Chowan formation of North Carolina. 

This species was recorded many years ago by Lesquereux from the Pleistocene of the Mississippi bluffs near 

Columbus, Ky., but as there is some question regarding this record it is queried in the above citation. 

Ulmus alata is not uncommon in the Pleistocene of 

Alabama. 

In the existing flora this species occurs in both wet 

and dry soils from southern Virginia to western Florida 

and westward to Texas, Missouri, and Arkansas. In 

North Carolina it is found along streams and swamp 

margins and is common throughout the State, except 

in the northwestern part of the Piedmont Plateau and 

in the mountain region. 

Occurrence: Chowan formation, right bank of Neuse 

River 1:~ miles above Seven Springs, Wayne County. 

Genus CELTIS Linne 

Celtis occidentalis Linne 

Plate LIV, Figure 3 

Celtis occidentalis Linne. Berry, Am. Naturalist, vol. 43, p. 435, 

1909. 

A single fragment of a leaf of this species was found 

in the Chowan formation. In the existing flora the 

species occurs in dry soils from southern Canada south· 

ward . to Florida and Texas~ A stone of this species 

has been recorded from the Talbot formation of Virginia. Another species is recorded from the Sunderland formation of Maryland, and Oeltis mississippiensis has been found in the loess of Mississippi. The 

genus is an old one, not uncommon in the Tertiary of 

both North and South America. 

Occurrence: Chowan formation, right bank of Neuse 

River 4~ miles above Seven Springs, Wayne County. 

Genus PLANERA Gmelin 

Planera aquatica (Walter) Gmelin 

Plate LIV, Figures 4, 5 

Planera gmelin Michaux. Lesquereux, Am. Jour. Sci., 2d ser., 

vol. 27, p. 365, 1859. 

Planera aquatica Gmelin. Hollick, Torrey Bot. Club Bull., vol. 

19, p. 332, 1892. 

Berry, Jour. Geology, vol. 15, p. 343, 1907; U. S. Geol. 

Survey Prof. Paper 98, p. 201, pl. 47, figs. 1-4, 1916. 

Somewhat doubtfully determined leaves of this 

species occur sparingly in the Chowan formation of 

North Carolina. 

The water elm is one of those interesting relics of a 

genus otherwise extinct, which attained an extensive 

distribution during the Tertiary but is now represented 

by only this single surviving form, which is apparently 

less wide ranging to-day than it was during Pleistocene 

time. It appears in the Pliocene Citronelle formation 

of southern Alabama and in beds of about the same 

age in soJ,Ithern New Jersey, and it has been found in 

the Pleistocene of Kentucky and North Carolina. 

In the modern flora it is found in more or less inundated swamps and river bottoms from southeastern 

North Carolina southward in the Coastal Plain to 

western Florida and westward to Trinity River, 

Texas, and up the Mississippi Valley to southeastern 

Missouri and western Kentucky, though usually rare 

away from the coast. In North Carolina it is found 

only in the lower valley of Cape Fear River and is no 

longer present in the valley of Neuse River. 
( 

112 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY, 1925 

Occurr~nce: Chowan formation, right · bank of 

Neuse River 4_72 miles above Seven Springs, Wayne 

County. 

Order PLAT AN ALES 

Family PLAT AN ACEAE 

Genus PLATANUS Linne 

Platan~s occidentalis Linne 

Plate LV, Figures 1-9 

Platanus aceroides Hollick (not Goeppert), Maryland Geol. 

Survey, Pleistocene, p. 231, pls. 73, 74, 1906. 

Platanus occidentalis Linne. Knowlton, Am. Geologist, vol. 

18, p. 371, 1896. 

Penhallow, Roy. Soc. Canada Trans., 2d ser., vol. 2, sec. 4, 

pp. 68, 72, 1896; Am. Naturalist, vol. 41, p. 448, 1907. 

Mercer, Acad. Nat. Sci. Philadelphia Jour., 2d ser., vol. 11, 

p. 277, 1899. 

Berry, Jour. Geology, vol. 15, p. 344, 1907; , Am. Naturalist, 

vol. 41, p. 695, pl. 2, fig. 5, 1907; Am. Jour. Sci., 4th 

ser., vol. 29, p. 397, 1910; Torreya, vol. 14, p. 161, 1914; 

vol. 15, p. 207, 1915; Pan-Am. Geologist, vol. 41, p. 

106, 1924. 

Emerson, U. S. Geol. Survey Bull. 597, p. 148, 1917. 

This species is one of the commoner elements of the 

Pleistocene flora of southeastern North America 

being recorded from the oldest Pleistocene of Maryland 

(Sunderland formation), the interglacial deposits of 

~he Don Valley in Ontario, and po~tglacial deposits 

In Massachusetts and Virginia. It is also found fossil 

in the Talbot formation of Maryland, in the Port 

Kennedy cave in Pennsylvania, in West VirO'inia at 

various localities in Alabama, ·and in the bCho~an formation of North Carolina, where its leaves are 

exceedingly common and the.flattened fruits or button 

balls are not rare. 

.I~ belong~ to an exceedingly interesting genus, which 

onginated In Upper Cretaceous time and represents 

an order of ·flowering plants now largely extinct. 

Platanus has many extinct species and attained ·a 

holarc~ic distribution during the Tertiary period. 

The six or seven surviving species are confined to 

southwestern Asia and eastern and western North 

America, with survivors of its once more extended 

range in the uplands of Mexico and Central America. 

T_he American plane tree or syca~ore grows on rich 

bottom lands from New England to Kansas and 

Nebraska and southward to Florida and Texas. It is 

said by Pi'nchot and Ashe to reach its maximum of 

size and abundance in North Carolina in the alluvial 

swamps of the Piedmont •Plateau region, but observations show that it is exceedingly common along the 

Coastal Plain rivers of the State. 

_ Occurrence: Chowan formation, right bank of Neuse 

River 4_72 miles above Seven Springs, Wayne County. 

Order POLYGONALES 

Family POLYGONACEAE 

Genus POLYGONUM .Linne 

Polygonum sp. 

An achene, determined by W. L. McAtee as referable to the genus Polygonum, is pres~nt in the 

collections from the Neuse River locality in Wayne 

County. Fossil forms have been recorded from the 

Pleistocene of Ontario/ Maryland,8 the District of 

Columbia,9 and Florida.10 

Occurrence: Chowan formation, right bank of 

Neuse River 4 Y2 miles above Seven Springs, Wayne 

County. 

Order RAN ALES 

Family MAGNOLIACEAE 

Genus LIRIODENDRON Linne 

Liriodendron tulipifera Linne 

Plate LIV, Figures 6-8 

Liriodendron tulipifera Linne. . Berry, Am. Naturalist, vol. 41, 

p. 695, 1907; Torreya, vol. 9, p. 71, fig. 1, 1909; Am. Jour. 

Sci., 4th ser., vol. 29, p. 396, 1910; Torreya, vol. 15, p. 

208, fig. 1, 1915. . 

The tulip tree or so-caJ.].ed yellow poplar was another 

common type in the Pleistocene floras of southeastern 

North America, where it has been recorded from 

Maryland, North Carolina, and Alabama, and is 

represented by both leaves and fruit. 

In the existing flora it inhabits deep, rich, moist 

soil of divides and slopes from Rhode Island to 

illinois and southward to Mississippi and northern 

Florida. In North Carolina it is found throughout 

the State but reaches its largest size on the slopes of 

the western mountains. 

Liriodendron is another one of those genera of forest 

trees that have come down to us from the distant past 

in a greatly diminished way.. It was exceedingly 

abundant and varied during Upper Cretaceous time 

and attained a holarctic distribution during the 

Tertiary, which apparently it retained until the close 

of that period, having been recorded by the Reids 11 

from the Pliocene of Holland, and by Schmalhausen 12 

from the Pliocene of central Asia. To-day it is 

restricted to southeastern North .Anlerica and 

southern China. 

· Occurrence: Wicomico formation, 1 U miles east of 

Weldon, Northampton County. 

7 Coleman, A. P., Geol. Soc. America Bull., vol. 26, p. 247, 1915. 

s Hollick, Arthur, Maryland' Geol. Survey, Pleistocene, p. 231, 1906. 

_ 8 Berry, E. W., Washington Acad. Sci. Jour., vol. 14, p. 19, pl. 1, figs. 19-23,1924. 

to Berry, E. W., Jour. Geology, vol. 25, p. 662, 1917. 

u Reid, C. and E. M., Pliocene flora of the Dutch-Prussian border, p. 93, pl. 8, 

figs. 1-5, 1915. 

u Schmalhausen, J., Ueber tertilire Pflanzen aus dem Thale des Flusses Buch· 

torma am Fusse des Altaigebirges; Palaeontographica, Band 33, p. 211, pl. 21, 

figs. 20, 21, 1887, 
PLEISTOCENE PLANTS FROM NORTH CAROLINA 113 

Order ROSALES 

Family HAMAMELIDACEAE 

Genus LIQUIDAMBAR Linne 

Liquidambar styraciflua Linne 

Plate LVI, Figures 9, 10 

Liquidambar europaeum Lesquereux (not Alexander Braun), 

U. S. Nat. Mus. Proc., vol. 11, p. 36, 1888. 

Liquidambar styraciflua Linne. Hollick, Torrey Bot . . Club 

Bull., vol. 19, p. 331, 1892. 

Knowlton, Am .• Geologist, vol. 18, p. 371, 1896. 

Berry, Jour. Geology, vol. 15, p. 343, 1907; Am. Jour. 

Sci., 4th ser., vol. 29, p. 397, 1910. 

The sweet or red gum is another of our forest trees 

that was common during the Pleistocene epoch in 

southeastern North America, when it was scarcely if 

at all distinguishable in its foliar characters from its 

wide-ranging Tertiary ancestors. Its oldest occurrence in this region is in the Pliocene of southern 

New Jersey. It has been recorded from Pleistocene 

deposits in West Virginia, North Carolina, and Alabama, and in the last two States it is represented by 

both leaves and fruits. 

In the existing flora it gro:ws on the wet soils of 

bottom lands, ranging from Connecticut to southeastern Missouri, Florida, and eastern Texas. It is 

essentially a Coastal Plain species, although it reappears in southern Mexico and the uplands of Guatemala, showing that it existed before the development 

of the wide belt of arid country in southwestern Texas 

\ and central and northern Mexico which now divides 

its once continuous range. It gives to the woods of 

the South Atlantic States an autumnal brilliancy that 

is unsurpassed by that of the red maple of the Middle 

and North Atlantic States. 

In North Carolina the sweet gum is found throughout the area east of the mountains but is most abundant and of largest size in the Coastal Plain. 

The leaves decay readily, in niy experience much 

more rapidly than those of most of our other forest 

trees, but the fruits or gum balls are more resistant 

and are frequently found in the fossil state. The 

genus goes back at least as far as the Eocene, and it 

attained a holarctic distribution during the Tertiary 

period. Survivors of this Tertiary dispersal are still 

living in southeastern North America and south .. 

western and southeastern Asia. 

Occurrence: Chowan formation, right bank of Neuse 

River 4~ miles above Seven Springs, Wayne County. 

Family ROSACEAE 

Genus RUBUS Linne 

Rubus sp. 

Plate LVI, Figure 8 

Rubus sp. Berry, Jour. Geology, vol. 15, p. 344, 1907. 

Prickly twigs, referred to Rubus with some hesitation, occur in the Chowan formation of North Caro856580-26--12 

lina. The genus comprises many existing species of 

herbs, shrubs, or vines and has a wide geographic distribution. The characteristic seeds of various species 

have frequently been recorded from Pliocene and 

Pleistocene deposits in Europe, and doubtless several 

species were present during Pleistocene time in southeastern North America, but because of the neglect of 

this field of research in North America, little is known 

regarding them. Characteristic seeds were recently 

discovered in' abundance in the Wicomico formation of 

the District of ColumbiaP 

· Occurrence: Chowan formation, right bank of Neuse 

River 431 miles above Seven Springs, Wayne County. 

Family POMACEAE 

Genus MALUS Hall 

Malus coronariafolia Berry 

Plate LVI, Figure 3 

Malus coronariafolia Berry, Jo~r. Geology, vol. 15, p. 344, 1907. 

Leaves triangularly ovate, widest below the middle, narrowing to the acute apex. Base broadly 

rounded, toothed. Length about 4 centimeters. 

Maximum width about 3 centimeters. Margins 

with four or five incised lobes on each side, which 

are finely salient-serrate. Midvein relatively slender. 

Secondaries about four pairs, slender, ascending; 

they diverge from the midvein at angles ranging from 

45 ° in the basal pair to half this amount in the distal 

pair; they are rather straight in their courses and 

craspedodrome, terminating at the acute tips of the 

lateral lobes. Tertiaries obsolete. 

This species appears to be referable to the genus 

Malus, and its resemblance to the leaves of th~ existing American crab apple, Malus coronaria (Linne) 

Miller, has suggested the specific name proposed. 

The latter is a species of tnoist glades which ranges 

from Ontario to Potomac River and southward along 

the Alleghany Mountains to central Alabama. I 

am not at all sure that the resemblance between the 

fossil form and this recent species can be considered 

as an indication of close relationship. 

Occurrence: Chow an formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County. 

Malus {>Seudoangustifolia Berry, n. sp. 

Plate LVI, Figures 1, 2 

Leaves small, somewhat variable in outline, generally ovate, ranging from 2. 75 to 4 centimeters in length 

and 1.25 to 2 centimeters in maximum width, which 

is usually below the middle. Midvein stout below, 

becoming thin distad. Secondaries immersed. Apex 

acute. Base rounded. . Margins with rather prominent dentate teeth. 

1a Berry, E. W., Washington Acad. Sci. Jour., vol. 14, p. 20, pl. 2, fii. 1, 1924. 
114 SHORTER CO:KTRIBUTIONS TO GENERAL GEOLOGY, 1925 

This is a fol1ll of somewhat uncertain botanical 

affinity, named from its resemblance to the existing 

Malus angustifolia, a Coastal Plain species ranging 

from Pennsylvania to northern Florida and westward to Louisiana. The resemblance is not exact, 

and it is possible that the fossil may represent some 

Pleistocene species of Orataegus. 

Occurrence: Chowa:r;:t formation, right bank of 

Neuse River 4Y2 miles above Seven Springs, Wayne 

County. 

Genus CRATAEGUS Linne 

Crataegus spathulatoides Berry 

Plate LVI, Figure 4 

Crataegus sp('Lthulatoides Berry, Jour. Geology, vol. 15, p. 344, 

1907. 

Leaves relatively small, obovate to spatulate. 

Apex acute to rounded. Base gradually narrowed 

and acuminate. Margins rather coarsely crenate 

above the middle. Length · about 4 centimeters. 

Maximum width about 1.5 centimeters. Petiole 

short and stout. Midvein stout. Secondaries relatively stout; a few ascending pairs diverge from the 

midvein at acute angles and are camptodrome. 

This form is very similar to the leaves of the existing 

small-fruited haw, Orataegus spathulata Michaux, and 

probably represents that species. Because of the subjective features on which many of the host of recently 

described species of hawthorn are based, it has seemed 

likely to be less confusing to give the fossil a new name 

which shall suggest its resemblance to Orataegus 

spathulata. The latter is a small tree found in rich 

moist soils, usually stream banks or swamp borders, 

in the Coastal Plain from southern Virginia to northern Florida and eastern Texas. 

Occurrence: Chowan formation; right bank of 

Neuse River 4 Y2 miles above Seven Springs, Wayne 

County. 

Crataegus coccineafolia Berry 

Plate LVI, Figures 5-7 

Crataegus coccineafolia Berry, Jour. Geology, vol. 15, p. 345, 

1907. . 

This species is based upon leaves and associated 

thorns. The leaves are relatively small, ovate in general outline, widest medianly, tapering upward to the 

acute tip and downward to the narrowly cuneate or 

slightly decurrent base. Margins incised to form two 

or three triangularly acute lobations on each side, with 

prominent serrate teeth, which die out in the cuneate 

base. Length about 3.75 centimeters. Maximum 

width ab<;ut 2.25 centimeters. Petiole not preserved. 

Midvein slender, curved. Secondaries thin, three or 

four pairs, diverging from the niidvein at acute angles, 

rather straightly ascending, craspedodrome. The 

thorns, attached to broken pieces of branches, range 

from 1 to 3.5 centimeters in length and are stout, 

conically tapering, acutely pointed, and often slightly 

curved distad. ' 

Both leaves and thorns are practically identical with 

the corresponding parts of the existing scarlet thorn 

and certainly represent it or something closely related 

to it. In · view of the maze of existing species into 

which the thorns have been segregated it has not been 

deemed profitable to atten;tpt too close correlations between them and the fossil species, as many southern 

species have very similar foliar characte:rs. 

The scarlet thorn is a small bushy tree of rich, welldrained soils and a northern distribution. It ranges 

from Newfoundland to Connecticut near the coast 

and up the valley of the St. Lawrence to western 

Quebec and hence is remote from North Carolina. 

Could the identity between this existing species and 

the fossil species be established with certainty the 

fossil would constitute a conspicuous exotic elemept 

in the flora of the Pleistocene of North Carolina and 

might be considered indicative of a considerable 

climatic contrast between the North Carolina of 

Pleistocene time and that· of to-day. As such a 

contrast is not borne out by the associated fossil 

plants and as many so-called modern species of 

Orataegus have similar leaves and thorns, no dependence can be placed on the resemblances pointed 

out above. 

Occurrence: Chow an formation, right bank of Neuse 

River 4Y2 miles above Seven Springs, Wayne County. 

Family CAESALPINIACEAE 

Genus CERCIS Linne 

Cercis canadensis Linne 

Plate LVII, Figures 10, 11 

Cercis canadensis Linne. Penhaliow, A~. Naturalist, vol. 41, 

p. 446, 1907. Berry, Torreya, vol. 11, p. 72, fig. 2, 1909. 

Fossil leaves of the redbud or Judas tree have been 

found in the interglacial deposits of the Don Valley 

in Ontario and in North Ca;rolina. 

In the modern flora this species is said by Sargent, 

Britton, arid Small to range northward as far as 

southern Ontario, whereas Sudworth gives its normal 

northern range as the lower valley of the Delaware 

and southern Michigan. Thence it ranges westward 

to Nebraska and southward to Florida, Texas, and 

t]+e Sierra Madre of Nuevo Leon. · In North Carolina 

it is found in both the Piedmont and Coastal Plain 

regions. It is essentially a warm temperate type, 

reaching its largest size along Sabine River and having 

niany tropical relatives. 

The genus goes back to Eocene time and is represented by existing species in Europe and Asia, as 

well as in both eastern and ·western North America. 

The south European Oercis siliquastrum Linne, like 

its relative in southeastern North America, is represented in the Pleistocene. 

Occurrence: . Wicomico formation, 1}4 miles east o£ 

Weldon, Northampton County. 
PLEIST'OOENE PLANTS FROM NOltTH CAROLINA 115 

Order SAPINDALES 

Family ILICACEAE 

Genus ILEX Linne 

Ilex opaca Aiton 

Plate LVI. Figures 11, 12 

/lex opaca Aiton. Hollick, Torrey Bot. Club Bull., vol. 19, 

p. 331, 1892. 

Berry, Jour. Geology, vol. 15, p. 345, 1907; Am. Naturalist, 

vol. 41, p. 696, pl. 2, fig. 1, 1907; Washington Acad. Sci. 

Jour., vol. 14, p. 21, pl. 2, figs. 4, 5, 1924. 

The holly, represented by both leaves and characteristic seeds, has been recorded from the Wicomico 

formation of the District of Columbia, the Chowan 

. formation of North Carolina, and beds at a corresponding horizon of the Pleistocene in Alabama, the 

coriaceous spinous leaves being. well suited for preservation. In the modern flora it is found in drier situations toward its northern limits, but t~ the south it . 

occurs chiefly in rich moist soil of woods and bottoms. 

It ranges from Massachusetts to Florida and Texas. 

It is common throughout North Carolina except in 

the mountain region. 

The genus flex goes back to the Upper Cretaceous 

and contains many extinct species. In the existing 

flora it comprises very many species, which are widely 

distributed throughout the world. It is most abundant in Central and South America. Asia also contains numerous species, but there are comparatively 

few in Europe and Africa. 

In addition to Ilex opaca, which has been found in 

the Pliocene of southern New Jersey, several other 

species of flex have been recorded from the Pleistocene of southeastern North America, and the genus is 

also represented in the Pleistocene of Europe. 

Occurrence: Chowan formation, right bank of 

Neuse River 4Y2 miles above Seven Springs, Wayne 

County. · 

Order RHAMN ALES 

Family VITACEAE 

Genus VITIS Linne 

Vitis sp. Berry 

Plate . LVII, Figure 6 

Vitis sp. Berry, Jour. Geology, vol. 15, p. 345, 1907. 

The genus Vitis is represented in the Chowan formation by the tendril figured, which is almost certainly that of some grape. It is strange that seeds 

have not been discovered in the Pleistocene of North 

Carolina, for they are exceedingly abundant in deposits of this age in southeastern North America, 

where they have been recorded from New Jersey,14 

Maryland/5 the District of Columbia/6 and VirginiaP 

Occurrence: Chowan formation, right bank of 

Neuse River 4Y2 miles above Seven Springs, Wayne 

County. 

H' Berry, E. W., Torreya, vol. 10, p. 266, 1910. 

1s ffollick, Arthur, Maryland Geol. Survey, Pleistocene, p . 235, 1906. 

16 Berry, E. W., Washington Acad. Sci. Jour., vol. 14, p. 21, pl. 2, figs. 6-9, 1924, 

11 Berry, E. W., Torreya, vol. 6, p. 89, 1906. , 

Order THYMELEALES 

Family LA URACEAE 

Genus PERSEA Linne 

Persea pubescens (Pursh) Sargent 

Plate LVII, Figure 5 

Persea pubescens Sargent. Berry, Jour. Geology, vol. 15, p. 

345, 1907. 

The swamp bay is a slender tree of pine-barrep. 

swamps occurring from North Carolina to Mississippi 

and Florida, near the coast, and unknown in the upper 

valley of the Neuse, where it was found fossil. · 1 

The related red bay, Persea borbonia Sprengel, 

ranges as a larger tree in somewhat similar situations 

from Virginia to Florida, Arkansas, and Texas and is 

believed by some botanists to be an older species than 

Persea p11-bescens. The foliage of the two is much 

alike, and it is possible that the fossil may represent 

the former, although it agrees more perfectly with the 

leaves of the latter. Neither is otherwise known in 

the fossil state, although Persea is an old genus with 

many extinct species and has been continuously present in southeastern North America since early Eocene 

time. 

Occurrence: Chow an formation, right bank of Neuse 

River 4Y2 miles above Seven Springs, Wayne County. 

Order UMBELLALES 

Family CORNACEAE 

Genus NYSSA Linne 

Nyssa biflora Walter 

Plate LVII, Figure 4 

Nyssa caroliniana Poiret. Hollick, Torrey Bot. Club Bull., 

vol. 19, p. 331, 1892. 

Nyssa biflora Walter. Hollick, Maryland Geol. Survey, Pleistocene, p. 235, pl. 69, fig. 5, 1906. 

Berry, Torreya, vol. 6, p. 90, 1906; Jour. Geology, vol. 

15, p. 345, 1907; . Am. Jour. Sci., 4th ser., vol. 29, p. 398; 

1910; Torreya, vol. 10, p. 266, 1910. 

This species, which is often confused with the more 

widely ranging Nyssa syl1)at·ica Marsh, is said by Sargent to be restricted to pine-barren ponds near the 

coast from North Carolina to Louisiana. Britton 

gives its northern range as New Jersey, and Shreve 

records it as common in the river swamps of the 

Eastern Shore of Maryland. 

Leaves or stone~ ha;,.e been recorded from the Pliocene of New Jersey and from the Pleistocene of New 

Jersey, Maryland, and Alabama~ In North Carolina 

fra()'mentary leaves were collected at the Neuse River 

loc:lity and stones at the Roanoke River locality. 

Occurrence: Chowan formation, right bank of Neuse 

River 4Y2 miles above Seven Springs, Wayne County. 

Pamlico formation. ·Old Mill Landing, left bank of 

Roanoke River 8 miles above Williamston, Bertie 

County. 
116 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY, 1925 

Order ERICALES 

Family ERICACEAE 

Genus XOLISMA Ra:finesque 

Xolisma ligustrina (Linne) Britton 

Plate LVII, Figure 12 

Xolisma ligustrina Britton. Hollick, Maryland Geol. Survey, 

Pleistocene, p. 236, pl. 69, fig. 6, 1906. 

Berry, Jour. Geology, val. 15, p. 346, 1907; Am. Naturalist, 

val. 41, p. 696, pl. 2, fig. 6, 1907; Am. Jour. Sci., 4th ser., 

vol. 29, p. 398, 1910. 

This species, which was referred by Linne to 

Vaccinium and is a type that paleobotanists usually 

refer to Andromeda, has been recorded from the 

Talbot formation of Maryland and from the Pleistocene of Alabama. 

In the existing flora it is a spemes that inhabits 

swamps and hillsides, and it ranges from Canada to 

Florida and Arkansas. Still other species occur in 

the South Atlantic and Guif States. 

Occurrence: Chow an formation, right bank of 

Neuse River 4 ~ miles above Seven Springs, Wayne 

County. 

Genus DENDRIUM Desvaux 

Dendrium pleistocenicum Berry 

Plate LVII, Figures 1, 2 

Dendrium pleistocenicum Berry, Jour. Geology, vol. 15, p. 346, 

1907; Am. Naturalist, vol. 43, p. 436, 1909. 

Small, narrowly elliptical leaves, with a rounded 

apex and base and conspicuously undulate margins. 

Texture conspicuously coriaceous, and the venation 

consequently more or less obsolete. in the smaller 

specimens. Midvein in the larger leaves thin and 

curved, giving off numerous thin wide-angled camptodrome secondaries. 

Dendrium is a small genus of shrubs of southeastern 

North America which are known as sand myrtles. 

Dendrium buxifolium (Bergins) Desvaux is found in 

sandy pine barrens from New Jersey to North Carolina. A second species, Dendrium hungeri Small, with 

larger leaves, ·is found in the mountains of. North and 

South Carolina. The leaves of D. hungeri agree rather 

well with those of the fossil, which is regarded as the 

Pleistocene ancestor of these two modern forms and 

which inhabited the Piedmont Plateau and the part 

of the Coastal Plain then above sea level. Dendrium 

hungeri represents a modern upland type much like 

the Pleistocene form, and Dendrium buxifoli·um represents a pine-barren modification with reduced leaves. 

This same species has been recorded from the Talbot 

formation of Virginia. 

Occurrence: Chow an formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County. 

Family V ACCINIACEAE 

Genus VACCINIUM Linne 

Vaccinium corymbosum Linne 

Plate LVIIJ Figures 8, 9 

Vaccinium corymbosum Linne. Hollick, Maryland Geol. 

Survey, Pleistocene, p. 236, pl. 69, figs. 7-9, 1906. 

Berry, Jour. Geology, val. 15, p. 346, 1907; Am. Jour. 

Sci., 4th ser., val. 29, p. 398, 1910; Torreya, vol. 17, pp. 

161, 162, fig. 2, 1917. 

This species has been recorded from the Pleistocene 

of Maine, Maryland, North Carolina, and Alabama. 

In 'the existing flora it is found in meadow swamps and 

moist woods and ranges from Newfoundland to 

Minnesota and southward to Florida and Louisiana. 

Occurrence: 'Chow an formation, right bank of Neuse 

River 4~ miles above Seven Springs, Wayne County. 

Pamlico formation, Old Mill Landing, left bank of 

Roanoke River, 8 miles above Williamston, Bertie 

County. 

Vaccinium spatulatum Berry 

Plate LVII, Figure 3 

Vaccinium spatulata Berry, Jour. Geology, vol. 15, p. 346, 1907. 

Small obovate to spatulate leaves, widest above the 

middle, with broadly rounded apex and narrowly 

cuneate base. Margins entire, somewhat irregular. 

Texture subcoriaceous. Length about 2.5 centimeters. Maximum width about · 1.2 centimeters. 

Petiole short and stout, .only about 1 millimeter long. 

Secondaries remote, two or three ascending camptodrome pairs. This leaf has the features of Vaccinium 

and appears to represent an extinct form. 

Occurrence: Chowan formation, right bank of 

Neuse River 4~ miles above Seven Springs, Wayne 

County. 

Vaccinium arboreum Marsh 

Plate LVII, Figure 7 

Berbe;is sp. Berry, Jour. Geology, val. 15, p. 343, 1907. 

Vaccinium arboreum Marshall. Berry, Torreya, val. 9, p. 73, 

1909; Am. Jour. Sci., 4th ser., vol. 29, p. 398, 1910; 

val. 34, p. 222, 1912. 

A single broken specimen from the Chowan formation of North Carolina, referred originally to Berberis, 

unquestionably represents the sparkleberry. It has 

subsequently been found fossil in the Pleistocene of 

Alabama and the Great Valley of Virginia. 

In the existing flora · Vaccinium arboreum is often a 

<small tree of sandy soil, ranging from North Carolina 

to Kentucky and southward to Florida and Texas. 

It is sometimes removed from the genus V accinium 

and placed in the genus Batodendron Nuttall, as by 

Small. 

Occurrence: Chowan formation, right bank of 

Neuse River 4 ~ miles above Seven Springs. Wayne 

County. 
PLEISTOCENE PLANTS FROM NORTH CAROLINA 117 

Order RUBIALES 

Family CAPRIFOLIACEAE 

Genus VIBURNUM Linne 

Viburnum moUe Michaux 

Viburnum cf. V. molle Michaux. Berry, Torreya, vol. 14, p. 

160, 1914. 

Two stones that appear to be referable to this species 

were found in the Chowan formation, 

In the modern flora Viburnum molle ranges from 

eastern Massachusetts. to Florida and Texas near the 

coast. 

Occurrence: Chowan formation, right bank of 

Neuse River 4 ~ miles above Seven Springs, Wayne 

County . 

• 

\ - \ 

\ ... / 

Viburnum nudum Linne? 

Viburnum nudum Linne. Berry, Torreya, vol. 14, p. 160, 1914; 

Jour. Geology, vol. 25, p. 662, 1917; Florida Geol. 

Survey Ninth Ann. Rept., p. 29, 1917; Torreya, vol. 22, 

p. 11, 1922; Washington Acad. Sci. Jour., vol. 14, p. 23, 

pl. 2, fig. 25, 1924. 

Stones of this species are not uncommon in the 

Pleistocene of southeastern North America and have 

been recorded from the Wicomico formation of the 

District of Columbia, the loess, of western Tennessee, 

and the low-level terrace at V ero, Fla. Two stones 

that appear to represent this species were collected 

from the Chowan formation in North Carolina. 

In the existing flora it ranges from Long Island to 

Florida and Louisiana. 

Occurrence: Chowan formation, right bank of Neuse 

River 4~ miles above Seven Springs, Wayne County. 
• 
PLATES XLV-LVII 

119 

U. S. GEOLOGICAL SURVEY PROFESSIOKAL PAPER 140 PLATE XLV 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-8. Taxodium distichum (Linne) L. C. Richard (p. 105). (Figs. 6, 7 after Hollick.) 

9. Pinus serotina Michaux (p. 105). 

/ 

1/ 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 140 PLATE XLVI 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-4. Hicoria glabra (Miller) Britton (p. 106). 

5. Hicoria aquatica (Michaux son) Britton (p. 106). 

6-7. Hicoria ovata (Miller) Britton (p. 106). 

8, Salix bebbianaformis Berry, n. sp. (p. 107). 
U. S. GEOLOGICAL SURVEY 

l 

I 

PROFESSIONAL PAPER 140 PLATE XLVII 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-4. Salix timinalifolia Berry (p. 107). 

5-7. Carpinus caroliniana Walter (p. 108). 

8-19. Betula nigra Linne (p. 107). 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 140 PLATE XLVITI 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-2. Betula pseudofontinalis Berry (p. 108). 

3-13. Fagus americana Sweet (p. 108). 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 140 PLATE XLIX 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1- 3. Quercus velutina Lamarck (p. 111). 

4- 6. Quercus palustris Du Roi (p. 109). 
U. S. GEOLOGICAL SURVEY 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-5. Quercus alba Linne (p. 108). 

6-9. Quercus lyrata Walter (p. 110). 

PROFESSIONAL PAPER 140 PLATE L 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 140 PLATE L! 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1- 7. Quercus nigra Linne (p. 109). 

8-23. Quercus phellos Linne (p. 1 10). 

24. Q·uercus abnormalis Berry (p. 109). 
U. S . GEOLOGICAL SURVEY 

PROFESSIONAL PAPER 140 PLATE LII 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-4. Quercus rnichauxii Nuttall (p. 110). 

5-8. Quercus prinus Linne (p. 110). 
U. S. GEOLOGICAL SURVEY PROFESSIOKAL PAPER 140 PLATE LIII 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-6. Quercus predigitata Berry (p. 109). 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 140 PLATE LIV 

r f.!J)l· I 

~ ) ~- ~ 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-2. Ulmus alata Linne (p. 111). 

3. Celtis occidentalis Linne (p. 111). 

4-5. Planera aquatica (Walter) Gmelin (p. 111). 

6-8. Liriodendron tulipijera Linne (p. 112). 

9. Quercus prinus Linne (p. 110). 
U. S . GEOLOGICAL SURVEY 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-9. Platanus occidentalis Linne (p. 112). 

PROFESSIONAL PAPER 140 PLATE LV 
U. S. GEOLOGICAL SURVEY 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1-2. Malus pseudoanqustifolia Berry, n. sp. (p. 113)· 

3. Malus coronariafolia Berry (p. 113). 

4. Crataequs spathulatoides Berry (p. 114). 

5-7. Crataequs coccineafolia Berry (p. 114). 

8. Rubus sp. (p. 113). 

9-10. Liquidambar styracijiua Linne (p. 113). 

11-12. !lex opaca Aiton (p. 115). 

PROFESSIONAL PAPER 140 PLATE LVI 
U. S. GEOLOGICAL SURVEY 

PLEISTOCENE PLANTS FROM NORTH CAROLINA 

1 - 2. Dendrium pleistocenicum Berry (p. 116). 

3. Vaccinium spatulatum Berry (p. 116). 

4. Nyssa biflora Walter (p. 115). 

5. Persca pubescens (Pursh) Sargent (p. 115), 

6. Vitis sp. Berry (p. 115). 

7. Vaccinium arboreu,m Marsh (p. 116). 

8-9. Vaccinium corymbosum Linne (p. 116). 

10-11. Cercis canadensis Linne (p. 114). 

12. Xolisma ligustrina (Linne) Britton (p. 116). 

PROFESSIONAL PAPER 140 PLATE LVII