U.S.A
See. er

(Ab oe 78)
Vegetative Study at the Duck Field
Research Facility, Duck, North Carolina

by
Gerald F. Levy

MISCELLANEOUS REPORT NO. 76-6
APRIL 1976

DOCUMENT
COLLECTION ye,

Approved for public release;
distribution unlimited.

Prepared for
U.S. ARMY, CORPS OF ENGINEERS
COASTAL ENGINEERING
RESEARCH CENTER

Kingman Building
Fort Belvoir, Va. 22060

Reprint or republication of any of this material shall give appropriate
credit to the U.S. Army Coastal Engineering Research Center.

Limited free distribution within the United States of single copies of
this publication has been made by this Center. Additional copies are
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National Technical Information Service
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The findings in this report are not to be construed as an official
Department of the Army position unless so designated by other
authorized documents.

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REPORT DOCUMENTATION PAGE BEFORE COMPLETING FORM
1. REPORT NUMBER 2. GOVT ACCESSION NO.}| 3. RECIPIENT'S CATALOG NUMBER
MR 76-6

4. TITLE (and Subtitle) 5. TYPE OF REPORT & PERIOD COVERED

VEGETATIVE STUDY AT THE DUCK FIELD RESEARCH
FACILITY, DUCK, NORTH CAROLINA

Miscellaneous Report

6. PERFORMING ORG. REPORT NUMBER

8. CONTRACT OR GRANT NUMBER(e@

- AUTHOR(e)

DACW72-74-C-0019

Gerald F. Levy

- PERFORMING ORGANIZATION NAME AND ADDRESS 10. PROGRAM ELEMENT, PROJECT, TASK
. . . AREA & WORK UNIT NUMBERS
Department of Biological Sciences
Old Dominion University G31265
Norfolk, Virginia 23508
- CONTROLLING OFFICE NAME AND ADDRESS 12. REPORT DATE
Department of the Army April 1976
Coastal Engineering Research Center (CERRE-EC) Ey COE Oe PAIS
80 % ye

Kingman Building, Fort Belvoir, Virginia 22060
. MONITORING AGENCY NAME & ADDRESS(if different from Controlling Office) 15. SECURITY CLASS. (of thie report)

UNCLASSIFIED

1Sea. DECL ASSIFICATION/ DOWNGRADING
SCHEDULE

. DISTRIBUTION STATEMENT (of this Report)

Approved for public release; distribution unlimited.

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17.

- SUPPLEMENTARY NOTES

- KEY WORDS (Continue on reverse side if necessary and identify by block number)

Coastal dunes Floristics Productivity
Duck Field Research Facility Phytosociology Taxonomy
Duck, North Carolina Plant ecology Vegetation

. ABSTRACT (Continue an reverse side if necesaary and identify by block number)

A vegetative study of the Duck Field Research Facility of the U.S. Army
Coastal Engineering Research Center, at Duck, North Carolina, was conducted
from March 1974 through June 1975. A vegetation map of this area was pre-
pared using aerial infrared photographs and ground-truth surveys. Eleven
different plant communities were delimited. Adequate stratified random
sampling of these communities produced frequency and biomass data for 10 of
the communities and frequency and density data for the eleventh community.

DD ; aren 1473 EDITION OF 1 NOV 65 IS OBSOLETE

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Abstract. (Continued)

Biomass data were obtained using the clip quadrat method. Ordination tech-
niques confirmed the distinctiveness of the foredune, wetland, oceanside
shrub, sound-side shrub, and sound-side disturbed communities. The
remaining six communities were floristically similar but quantitatively
distinct. Two of these latter communities appear natural while the
remaining four are manmade. The natural communities were designated low
dune grass and oceanside intershrub communities. The beachgrass and
bitter panicum communities were deliberately established. The spurge-
sandgrass and sandgrass-buttonweed communities were affected by sandgrass
planting and fertilization. Permanent plots were located and mapped in
each of the designated community types. Floristic collections made
throughout the study period revealed a flora of approximately 178 species
in 132 genera representing 58 families.

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PREFACE

This report is published to provide coastal engineers with a vegeta-
tive study of the Duck Field Research Facility at Duck, North Carolina.
The work was carried out under the coastal ecology research program of
the U.S. Army Coastal Engineering Research Center (CERC).

The report was prepared by Gerald F. Levy, Associate Professor of
Biological Sciences, Old Dominion University, Norfolk, Virginia, under
CERC Contract No. DACW72-74-C-0019.

The author expresses appreciation to Drs. Paul W. Kirk and Lytton
J. Musselman, Department of Biological Sciences, Old Dominion University,
for assistance in designing the research program and in species identi-
fication, respectively, and to R.W. Tyndall and J.W. Usher, graduate
students in the Department of Biological Sciences, for their assistance
in the field. The assistance of Dr. Donald W. Woodard, formerly of CERC,
is gratefully acknowledged.

Mr, Robert M. Yancey, Chief, Ecology Branch, was the CERC contract
monitor for the report, under the general supervision of R.P. Savage,
Chief, Research Division.

Comments on this publication are invited.

Approved for publication in accordance with Public Law 166, 79 th
Congress, approved 31 July 1945, as supplemented by Public Law 172, 88

Congress, approved 7 November 1963.

th

AMES L. TRAYERS
Cotonel, Corps of
Commander and Dire&tor

10

CONTENTS

I LONDURO DO CIMUOM 6g 6 61600 6 6 10) bo a)a Bua ado 0
lo @emeral 9 6 66 606656600005 016

Bo Oreaeatiny o ova reid’ odio lo eo oo ener aos

So GIT 6 6 65 0 FB ooo oO 8 Ooo
AUS t ONG Yapitoutonl/oniMel in cael oilieyiterisiuiceunion Doulsevuelikouars

5. Floristic Composition . 6.0.00 6.00.0

6. Plant Succession and Clears 06 806 6 6

7. Environmental Factors Governing Floristic
Composition and Distribution . . «. « e ©

II PROCEDURE ° ° e eo e ° e e e ° ° e e eo oe eo °o °o e
1. Floristics CG MOMeOUOMIe eal cues onc oll (hohe
4o WeEetaciomal StUEHES 555100000006
III RESULTS e °o e e ° e e e e oe e e e e ° ° e e e °
Ilo) Wilomalseles 66 6 6 6 )all6'6 16 'o!6o..0 6 6 55
25 Weuereclomell Seueles 56°64 516 0/0 6 60 6
IV DISCUSSION e @ °o oe e e e e e °o “e@ e e e eo e e e e
IG) PILOMPAS LCS Gd og gdilal Gd. Gia) 0) G40) ol 0

Ao) Wereecrelomel Seuckles 55 6 6) o)6 616 14 a0

MINNIE CIMID! 6 6 6 a,0 0116 610 0 610 6 0 6 6

TABLES
Duck Field Research Facility floristics list ...
FORSGILINE COMNBNIY Gig 5 6 6066066000066
Oceanside intershrub community . ... oo o o e e
Oceanside shrub community . . . . o « © © 0 © 0 © ©
Planted American beachgrass community . . . o o e o
Sandgrass-buttonweed community . . o « o « «© o o o
LON Ging) (ESS Coymmbmlbiey G 616 6 6 66.6606 0c
SOPING SIGS Sloywoilo} CGonmommiiay 66 So S56 6 6 Gg Bla 6

Planted bitter panicum community .. . . « o « « o

Sound-side disturbed community . . « « o »« o o e o

21

28

29

30

31

32

33

34

35

36

11

12

13

14

CONTENTS - Continued
TABLES
Weelamads Commins? 5 6 oo) 6 8 loo alo o lod '0)6)\a 5 0 0 €
Spunge-Sandora ssi Commun tyguontaly ei irelll eielnicniial hl eiiteliita citer tevicnire

A summary of community types at the Duck Field Research
Facility e e e ° e e e e e e e e e e e e e ° e ° e ° ° e e

Location of permanent quadrat stakes at the Duck Field
RESCH JEVEMITEY 6 66 oo Oo Oeb) 010.0 d 606.6 0 616

FIGURES

Location of the Duck Field Research Facility . . . o « « e
Vegetation map of the Duck Field Research Facility ... .

Ordination of plant communities at the Duck Field Research
Facility showing the X and Y axes perspective ..... e

Ordination of plant communities at the Duck Field Research
Facility showing the X and Z axes perspective ....e-.-.

Ordination of plant communities at the Duck Field Research
Facility showing the Y and Z axes perspective ....e.

Foredune community permanent quadrat 1 . . « « © «© © «© © e
Fornedune community pexmanene) Guadrat 2) hes iiisill/alvell si lentel ails
Foredune community) permanent) quadrat) Sie reiile, Wiel aile lice ive
Oceanside intershrub community permanent quadrat 1... .
Oceanside intershrub community permanent quadrat 2... .
Oceanside intershrub community permanent quadrat 3... .
Planted American beachgrass community permanent quadrat 1
Planted American beachgrass community permanent quadrat 2
Planted American beachgrass community permanent quadrat 3

Sandgrass-buttonweed community permanent quadrat 1... .

°

41

70

40

53

54

16

Ld

18

9)

20

Ail

22

23

24

25

26

27

28

29

30

CONTENTS = Continued

FIGURES

Sandgrass-buttonweed community permanent quadrat

Sandgrass-buttonweed community permanent quadrat

Low dune grass community permanent quadrat 1 .
Low dune grass community permanent quadrat 3 .
Sound-side shrub community permanent quadrat 1
Sound-side shrub community permanent quadrat 2

Sound-side shrub community permanent quadrat 3

Planted bitter panicum community permanent quadrat

Planted bitter panicum community permanent quadrat

Planted bitter panicum community permanent quadrat

Sound-side disturbed community permanent quadrat 1

Sound-side disturbed community permanent quadrat

Sound-side disturbed community permanent quadrat

Wetlands community permanent quadrat ... e« e

Spurge-sandgrass community permanent quadrat .

Zi

3

Page
55
56
57
58
59
60
61
62
63
64
65
66
67
68

69

VEGETATIVE STUDY AT THE DUCK FIELD RESEARCH FACILITY,
DUCK, NORTH CAROLINA

by

Gerald F. Levy

I. INTRODUCTION
Iho General.

The Duck Field Research Facility is located on the Outer Banks of
North Carolina, a series of offshore sandbars extending from the Virginia-
North Carolina border southward about 200 miles to Bogue Inlet, North
Carolina. These bars lie almost parallel to the mainland and are
separated from it by shallow sounds of varying widths.

The study area is included in Currituck Bank (North Bank), which
extends from the Virginia-North Carolina boundary southward to the town
of Nags Head, a distance of about 45 miles (Fig. 1). The bank was
originally demarcated by two inlets; Old Currituck Inlet, which was
located just south of the Virginia-North Carolina line until it closed
around 1730, and Nags Head Inlet, which was located just south of the
present town of Nags Head until it closed sometime between 1780 and 1810
(Stick, 1958).

Le Origin.

Most investigators cite a theory developed by Johnson (1919) for
the origin of this barrier island system. Johnson postulated that on a
gradually shallowing sea bottom, "...small waves break at the initial
shoreline and excavate a marine cliff and beach while large waves break
further out and proceed to excavate the same forms in the offshore
bottom."" Excavated material was deposited landward of the breaking
waves, eventually forming a submarine bar of significant height and
indefinite length. Continuing growth irregularly raised the bar above
water, forming a chain of islands separated by inlets.

The longshore current causes both shifting and closing of inlets.
Deposition on one side of an inlet is succeeded by erosion on the other
Side, producing inlet migration. If deposition exceeds erosion the
inlets eventually close unless tidal currents and outflowing -fresh-
waters are sufficient to dominate shoaling (Johnson, 1919).

Stick (1958) cited testimony from inhabitants which suggests that
hurricane winds raise the water level in the sound several feet above
normal, After storm passage, winds shift to the southwest and force the
impounded waters back over the banks, sometimes forming inlets.

Val Norfolk

i)
7
= a

WEIS. Be a a pale
soot) <
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DS x,
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\ » ¢
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SS of. \ ‘ SE,
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4 & ‘ .
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Mon 7
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i 30 miles

Figure 1. Location of the Duck Field Research Facility.

Sie Climate.

The area has a moderately temperate maritime climate (Engels, 1952).
Climatic data were collected at Hatteras, which is more under the
influence of the Gulf Stream than the Duck area. These data represent
maxima, but differences are probably not significant.

At Hatteras, the normal mean temperature during the 6 consecutive
hottest weeks of summer is 78.6° Fahrenheit. The coldest month, January,
has a mean temperature of 45.7° Fahrenheit, while July, the warmest, has
a mean of 77.9° Fahrenheit. The average daily temperature range is
greatest in March, 12.9° Fahrenheit, and least in August, 9.2° Fahrenheit.

The absolute maximum temperature recorded was 102° Fahrenheit, in
June, and the minimum, 8° Fahrenheit, in December. The average date of
earliest killing frost is December 13. February 27 is the latest date
of killing frost (Burk, 1968). The mean annual percent of sunshine,

54 percent, is typical for the southeastern states. The mean annual
humidity, 81.4 percent, is higher than any other continental station in
the United States with the exception of Puget Sound. Monthly differences
in humidity vary no more than 5 percent.

Mean annual precipitation is 62.76 inches, with July (6.06), August
(5.98) and September (6.06) having the most. Periods of drought are rare.

The average annual maximum wind velocity, 13.3 miles per hour, is
relatively high, with average maximum velocities greatest from December
through April. The prevailing winds are northerly in the winter and
southwesterly in the summer.

4. History.

The first permanent English community on the Outer Banks was
established in 1664 (Stick, 1958). Soon after settlement, cattle,
horses, hogs, and sheep were turned free to graze. Around 1773 settlers
were reportedly converting Currituck Banks to stock range and attempting
to prevent the Indians from hunting there. Grazing continued, to some
extent, into the 1900's. Guild (1947) reported finding pigs feeding on
mole crabs in the surf near False Cape; recent observations have con-
firmed that a viable "wild" hog population still exists.

The vegetation and topography of this region were drastically
changed after settlement. Stratton (1943) reported that grass and
shrubs were uprooted by hogs, and forests were cut for fuel, building,
and ship construction. Once vegetation was disrupted, the sandy soils
became susceptible to wind and storm tides. The blowouts and sand dunes
seen today behind the foredune are the results of these forces. Blowouts
almost reach the water table and dunes rise to over 100 feet just north
and south of the Duck Field Research Facility. Unstable dunes migrate
southwesterly across the banks, toward the sound, burying everything
in their paths.

Gradually rising sea levels (Marmer, 1951) have caused the beaches
to recede landward. The Cape Hatteras Lighthouse, constructed in the
late 1860's about 3,200 feet from the beach, is today approximately 350
feet from the water's edge (Dolan, Godfrey, and Odum, 1973). In 1935
the Works Progress Administration (WPA) and the Civilian Conservation
Corps (CCC) began stabilizing the foredune from the Virginia border to
approximately the middle of Ocracoke Island (Stratton and Hollowell,
1940). Some foredunes now exceed 25 feet in height. Foredune
Stabilization has narrowed beach width, and increased the oceanside
slope of the foredune and beach. Eventually the stabilized dunes will
be destroyed by natural oceanic processes (Dolan, 1972; Dolan, Godfrey,
and Odum, 1973).

The ocean beach, foredune, arborescent zone, and sound side marsh
are the most characteristic features of the Outer Banks profile, although
the widths and elevations have changed significantly in the past 300
years. The most variable area is between the foredune and the
arborescent zone. This area possesses either large migrating dunes,
intertidal salt marshes, and sandflats, or a narrow heterogeneous
transitional zone.

During World War II, Duck field was used as a gunnery range
(personal communication, D. Woodard, Marine biologist, CERC, 1974). The
area was subsequently decontaminated, which leveled considerable parts
of it. Effects of bombing are still apparent; the area remains littered
with bomb fragments and some cratering is still evident.

A series of planting experiments were recently conducted on the
study area. Extensive sandflats were planted with Ammophtla brevtligulata
(American beachgrass) or Pantcum amarum (bitter panicum), and other areas
were treated with fertilizer applications. Some areas were disturbed by
construction and the use of heavy equipment, both during and after the
experiments.

Se Floristic Composition.

Burk's (1968) study of the Outer Banks flora reported 462 native
species comprising 277 genera, 106 families, and 43 orders. He reported
the largest families to be Poaceae, Asteracea, and Cyperaceae. The
native plants included 14.7 percent Poaceae, 13.4 percent Asteraceae
and 8.0 percent Cyperaceae. Included in his lists were 95 exotic
Species comprising 55 genera and 33 families, which made up 17.1 percent
of the total flora.

a. Ocean Beach Community. Little or no vegetation exists between
the dunes and the ocean. This is in contrast to the Physalts vtscosa-
Croton marttimus community reported for Ocracoke Island by Kearney
(1900). Engels (1942, 1952) reported no sea beach vegetation on either
Ocracoke Island or Shackleford Banks; however, neither of these study
areas had stabilized dunes.

b. Foredune Community. Untola pantculata (sea oats) is the most
abundant plant in this community. Although not always dominant (Wagner,
1964), it is found throughout the Banks. Next in abundance is usually
A. breviltgulata which occurs as far south as Hatteras (Brown, 1959)
and is most abundant toward the Virginia border. Eventually it replaces
U. paniculata farther north. The abundance of these two species is
partly due to WPA and CCC planting activities.

The distributions of Andropogon glomeratus (bushy beachgrass) and
Andropogon ltttoralts are variously cited in the literature. Brown
(1959) reported A. Ztttoralis was absent from Outer Banks dunes. Burk
(1962) stated that A. Ztttoralts is dominant in the dune grass community,
and A. glomeratus is rarely a dominant dune species. Johnson (1900)
noted that A. martttmus (most likely A. glomeratus) was the dominant
foredune species on Bogue Bank.

Panteum amarum is a widespread subdominant from Virginia to
Ocracoke Island, North Carolina (Brown, 1959; Kearney, 1900).
Panteum amarulum is most abundant from Bodie Island, North Carolina
to Virginia (Brown, 1959). Both are considered poor sand binders.

Other plants of less ecological significance reported include:
Croton punctatus (croton), Cenchrus pauctflorus (field sandbur) ,
Cenchrus trtbulotdes (dune sandbur), Eragrostis spectabilis (purple
lovegrass), Triplasis purpurea (purple sandgrass), Caktle edentula
(sea rocket), Erigeron canadensts (horse weed), Euphorbia polygontfolta
(seaside spurge), Oenothera humtfusa (seabeach evening primrose),
Physalis viscosa (maritime ground cherry), Solidago sempervirens
(seaside goldenrod), Heterotheca subaxillarts (camphorweed) ,
Heterotheca gossyptna (camphorweed), Hudsonta tomentosa (beach heath),
Yueca glortosa (Spanish bayonet), Spartina patens (saltmeadow
cordgrass), and Opuntia compressa (prickly pear), (Brown, 1959; Burk,
1961; Johnson, 1900; and Kearney, 1900).

c. Migrating Dune Community. No plants are found in areas of
excessive sand drift. In areas of only moderate drift fast growing
species (e.g., U. paniculata and A. breviligulata) occur throughout the
year. During the less windy summer months, in areas of low drift,
annuals such as £. polygontfolta, T. purpurea, and Diodta teres
(buttonweed) are found along with S. patens and C. trtbulotdes
(Brown, 1959).

d. Sandflat Communities. On flat sandy areas various community
types occur depending upon moisture availability and salinity. Brown
(1959) found Typha lattfolta (cattail), Spartina alterntflora (smooth
cordgrass), Juncus roemertanus (spike rush), Cladtum jamatcensts (saw
grass), Pluchea sp. (marsh fleabane), Centella repanda, Setrpus
amertcana (three square), and an occasional S. patens growing on a wet
sandflat south of the Cape Hatteras Lighthouse.

Burk (1961) found the one-quarter-mile-long dry sandflats on
Portsmouth Island, North Carolina, totally barren. Johnson (1900)
reported similar areas on Bogue and Shackleford Banks, North Carolina
shifting wastes of windblown sand with only #. polygontfolta and
Andropogon sp.

Brown (1959) described the sandflat vegetation on Hatteras Island
near Rodanthe, North Carolina as dwarf, sparse, and composed of mixed
grasses and sedges. However, a sandflat near Hatteras Village possessed
a thick growth of S. patens, Fimbristylts castanea, and S. sempervtrens.
Sandflats north of Corolla on Currituck Bank had a good growth of S.
patens while some flats on Hatteras Island had only a fringe of
vegetation, with a center of packed sand coated with salt crystals.

e. Arborescent Communities. These communities include those
dominated by shrubs as well as trees. An open canopied mesic forest
dominated by Pinus taeda (loblolly pine), Carya spp. (hickories),

Quercus falcata (Spanish oak), Quercus nigra (water oak), and
Iiqutdambar styractflua (sweet gum) is described by Burk (1962) as
existing behind the active dunes near Nags Head, Kitty Hawk, and Duck.

P. taeda was the dominant species (37 percent of the total). Carya spp.,
L. styractflua, and Q. falcata with values of 16, 15, and 14 percent
respectively were next in importance. Burk (1962) found 2 percent

Q. virginiana but Brown (1959) found none.

Brown (1959) reported Q. virgintana and P. taeda as the major
species in Buxton Woods on Hatteras Island. He found 55-year-old pines
and an oak with a 3-foot diameter. Burk (1962) reported that 98 percent
of the trees were P. taeda, with Q. nigra, Q. virginiana, and L.
styractflua; Sabal minor (stemless palmetto) was abundant. Other woody
species reported by Brown (1959) included: Callicarpa americana (French
mulberry), Carpinus earolintana (ironwood), Cornus florida (flowering
dogwood), Ilex opaca (American holly), Ilex vomitoria (yaupon), Juntperus
virginiana (red cedar), Myrica cerifera (wax myrtle), Myrtca pensylvantca
(bayberry), Persea borbonia (red bay), Q. nigra, Q. phellos (willow aak),
Rhus copallina (winged sumac), and Zanthoxylum clavahercults (Hercules'-
club).

Kearney (1900) concluded that I. vomttorta was the most abundant
woody species on Ocracoke Island, followed by Q. virginiana, M. certfera,
Z clava-hercults, and J. virginiana.

Engels (1952) listed @. virginiana, J. virginiana, and I. vomttoria
as the dominant trees in a wooded area west of Whale Creek Bay on
Shackleford Bank. Other trees of less abundance included: P. taeda,

P. borbonta, and I. opaca.

On the basis of relative shrub-tree importance, Lewis (1917)

divided the arborescent zone on Shackleford Bank into three areas:
(a) thicket formation, (b) thicket woodland formation, and (c) woodland

l2

formation. The thicket formation was dominated by I. vomttorta, with
I. opaca, J. virgintana, M. centfera, and others. The thicket woodland
was characterized by P. borbonta, Osmanthus americanus (devilwood) ,

Q. nigra, P. taeda, and J. virginiana. Shrubs included: JI. vomitorta,
M. cerifera, C. americana, I. glabra, and Sabal glabra (S. minor?).

Johnson (1900) described the Bogue Bank arborescent zone as located
about 100 yards behind an advancing dune. Q. nigra, with I. opaca,
Morus rubra (red mulberry), Persea carolinensis (?), C. carolintana,

J. virginiana, and P. taeda were reported by him. Myrica gale (7),

I. glabra, and I. cassine (I. vomttoria), were noted as important
shrubs with the latter being most important. Apparently this area was
subsequently logged, or otherwise drastically altered as Burk (1962)
found 58 percent of the trees to be P. taeda, 15 percent Q. falcata,
and 12 percent Q. virgintana. The previously important water oak was
no longer present and the pines increased to major importance.

6. Plant Succcssion and Climax.

Wells (1928), the only investigator to describe the plant
successional pattern on the Outer Banks, suggested the following
pattern for the xeric dune community: Salsola (upper beach) dune
formation, Untola-Iva dune recession, Oenothera-Panitcum soil improvement,
and Quercus-Myrica. He characterized this habitat as haying a short
hydroperiod, little humus, strong winds, and little shade. His pattern
for the xeric maritime forest was: Ilex-vomitorta-Juniperus-Myrtca,
@. virgitntana, and oak-hickory associes. The habitat was described as
having a very short hydroperiod, perennial winds, and older landward-
lying dunes.

Wells (1938) reported an extensive forest of live oaks on Smith
Island, previously recorded in 1805. He concluded that live oak (@.
virgintana) was the dominant tree of the climax maritime forest, citing
as the reason the natural elimination of competitive hardwoods which
are not as tolerant of salt spray. He called this community a Salt
Spray Climax. Wells felt succession would be: U. paniculata, M.
certfera, I. vomitorta, J. virginiana, and Q@. virginiana. Bordeau and
Oosting (1959) concluded that the normal components of the oak-hickory
association are excluded by their intolerance to salt, thus only live
oak will dominate. This supports Wells' earlier conclusions.

Whe Environmental Factors Governing Floristic Composition and

Distribution.

a. Salt. Viewing storm effects on plants in England, Salisbury
(1805) noted that greater damage resulted when rain did not accompany
high winds. Beck (1819) noted that storms on the northeast U.S. coast
had little effect on plants protected from the wind. These observations
precipitated a controversy over the importance of wind effects versus
salt spray effects on plants. Coulter, Barnes, and Cowles (1911)

13

summarized this controversy as follows: "One maintains that the
stronger transpiration on the seaward or windward side causes the
branches to die earliest there, while the other theory maintains that
salt particles carried in the spray account for branch destruction."
They speculated that both points of view were valid because the two
phenomena supplemented one another.

Boodle (1920) considered salt spray to be the less important factor.
Weaver and Clements (1938) and Martin and Clements (1939) supported the
contention of Boodle (1920). Doutt (1941) and Davis (1942) found
evidence to support the contention that salt spray was of minor
importance.

Wells and Shunk (1937) observed the injury of young windward side
shoots of M. certfera, I. vomttorta, and Q@. virgintana on the lower
Cape Fear Peninsula following 19 hours of 30-mile-per-hour winds.
Shrubs farther from the ocean exposed to the same wind velocity and
time periods were not damaged. They concluded that salt was the
primary cause of deformed coastal shrubs and proposed that these
shrubs be called "spray forms" rather than "wind forms".

Oosting and Billings (1942) concluded that Bogue Bank vegetational
plant zonation patterns were not due to (a) ‘soil moisture, temperature,
pH or salt content, (b) air moisture, temperature, or (c) evaporation.
They also concluded that salt spray was the only significant factor
governing zonation. Oosting (1945) stated that salt spray effects were
most damaging when storms produced insufficient rain to wash salt from
leaves.

The amount of salt collected from the windward side of the foredune
on Bogue Bank was twice as great as at any other side (Oosting and
Billings, 1942). The crest of the foredune had the second largest
amount followed by the crest of the hind dune. The leeward side of the
foredune and the base of the hind dune received successively less
amounts of salt spray.

&

Boyce (1951) showed that significant amounts of salt spray are not
derived from the surf because the drops formed are too large to be
windborne. Most salt spray particles are from the bursting bubbles in
the swash and white caps of small waves (Boyce, 1954).

Boyce's (1954) study of the coastal vegetation of Brunswick County,
North Carolina, found Iva tmbricata (marsh elder) was subjected to an
average of 1900 droplets of salt spray per square centimeter per minute
on the windward side and only 490 droplets per square centimeter per
minute on the leeward side. The droplets averaged 51 micrometer in
diameter windward and 22 micrometer leeward.

Wells and Shunk (1938) concluded that the concentration of
chlorides in salt spray deposited on plant leaves was higher than

that for seawater because of evaporation, and leaf death was due to
cell plasmolysis. However, Boyce (1954) demonstrated that chloride ion
toxicity caused leaf death. He discovered other ions were seldom
present in high enough concentrations to produce necrosis. Since
necrosis always occurred first at the tip regardless of where the
chlorides entered the leaf, Boyce concluded that damage occurred

after salt translocation. Salt entrance was increased through wind-
induced leaf lacerations, which explained why plants with sessile
leaves or leaves densely covered by hair are damaged less by wind
action than plants with long petioles and smooth surfaces. He also
discovered that increased soil nitrates resulted in decreased cuticular
thickness and number of epidermal hairs allowing more frequent
lacerations, increased chloride uptake, and reduced salt spray
tolerance.

Wagner (1964) found that salt spray decreased the growth rate of
U. panteulata but did not kill the plant. This he attributed to the
thick cuticle and other xeromorphic characters of its leaves which
reduce surface chloride ion uptake.

Numerous investigators have classified strand plants according to
salt spray tolerance (Wagner, 1964; Oosting and Billings, 1942; Wells,
1939; and Wells and Shunk, 1938). Oosting (1945) studied the salt
spray tolerance of 14 species. He found S. patens and Atriplex arenaria
(seabeach orack) unaffected by salt spray, U. paniculata the next most
tolerant, and S. sempervirens and C. punctatus slightly less tolerant.
Iva imbricata, I. frutescens (marsh elder), and Borrichta frutescens
(sea-ox-eye) were moderately tolerant. EF. polygontfolta, Chenopodium
anbrostaides (Mexican tea), and Cynodon dactylon (Bermuda grass) had
some tolerance, while Strophostyles helvolva (wild bean) and H. subaxtl-
laris were less tolerant. Lepttlon canadensts (E. candensts) showed
virtually no tolerance to salt spray.

Other species rated as highly or moderately salt tolerant include
A. ltttoralts (Oosting and Billings, 1942), Baccharts halimifolta
(groundsel-tree), I. vomttoria, and M. certfera (Wells and Shunk, 1938).

Seneca (1969) found that maximum germination tolerance to salt
(NaCl) for A. brevtligulata and U. pantculata was between 1 and 1.5
percent. Pantcum amarulum had an upper tolerance limit between 1.5 and
2 percent, while S. patens was about 4 percent. The latter had an
average of 53.3-percent germination in 3-percent NaCl. Decrease in
germination was attributed not to salt toxicity but to the osmotic
differential. Based upon their germination experiments the order of
tolerance (from most to least) was S. patens, P. amarulum, U.
paniculata, and A. brevtligulata.

All the above species showed stress in the seedling state but at
different levels (Seneca, 1972). Symptoms included an initial twisting

15

and rolling of leaves, followed by chlorosis on the outer older leaves
with leaf tips becoming yellowish. Necrosis later proceeded from the
tips downward, affecting the newest leaves last. Ammophila
brevtltgulata and U. panteulata grew moderately well in salinities

up to 1 percent while P. amarulum showed no signs of salinity stress
until subjected to 2—percent NaCl. Seedlings of S. patens showed
salinity stress only at the 4-percent level. It was the only species
to actively secrete what was thought to be NaCl from its leaves. All
species lived for at least 10 days when exposed to 4—percent NaCl.

b. Nutrients. Willis, et al. (1959) reported a low level of
nitrogen in the sandy soil of the coastal dune system of Braunton
Burrows, North Devon, England. After further studies Willis (1963)
attributed the sparse vegetation on this dune system to low levels of
nitrogen and phosphorus, which appeared to be due to the high
leachability of the coarse sandy soils.

Wave deposition of organic detritus has been suggested as the
source of nutrients for the lower beach (Ranwell, 1972). Wagner (1964)
cited the freshly deposited beach sands as a source of nitrogen, while
Morris, et al. (1974) suggested ocean spray-carried neuston as a
possible source.

Hassouna and Warling (1964) isolated bacteria from the root soil
and roots of A. arenarta and found that Azotobactor, a nitrogen-fixing
bacterium, constituted a large proportion of these organisms. Their
studies indicated that the root surface micro-organisms in the dune
sand could stimulate significantly higher plant growth in the absence
of added nitrogen when root exudates (a carbon source) was added. They
concluded, "there seems little doubt that the fixation of atmospheric
nitrogen is an important factor in the nutrition of A. arenarta."'

Stewart (1967) showed that nitrogen fixed by blue-green algae in
soil was utilized by higher plants. Most fixed nitrogen remained in
the first centimeter of the soil unless leached.

Morris, et al. (1974) have shown that root nodules on M.
pensylvantca contain a bacterial endophyte. They found large numbers
of nodules on main and adventitious roots. Nitrogen fixation was
vigorous. They concluded that the success of M. pensylvantca as an
early successional species on dunes and other impoverished soils was
due, at least in part, to the nitrogen-fixing capacity of their
nodular bacteria. ;

Gorham (1958) found the mobile dunes in Norfolk, England, richer
in soluble calcium, nitrate and silica and the stable phase dunes
richer in soluble carbon and phosphate due to humus accumulation.
The oldest stabilized dunes were richest in potassium, sulfate,
and phosphate. Magnesium, sodium, and chlorine were always
highest in the embryo: dunes due to salt spray. Potassium was least
in the embryo dunes, and greatest in the stabilized dunes. Calcium

16

and bicarbonate concentrations were greatest in the youngest dunes
where shell fragments were abundant. Willis and Yeamon (1961) stated
that there is a sufficient supply of micronutrients (boron, iron,
manganese, zinc, copper, and molybdenum) in coastal soils to support
vigorous plant growth.

c. Soil Moisture and Water Table. Oosting and Billings (1942)
found no significant difference between soil water moisture at depths
of 10 and 20 centimeters, except in moist depressions. They could not
correlate moisture factors between the surface and 20 centimeters with
vegetational zonation. Kelley (1925) found that the beach was not
salty, because the movement of ground water maintained unmixed fresh-
water up to a few centimeters from the ocean's edge. Ranwell (1972)
found the water table below dunes convexed and unaffected by tidal
fluctuations, suggesting that the ground water table is independent of
the sea's influences.

d. Other Factors. Oosting and Billings (1942) reported that
temperature, relative humidity, and evaporation varied in different
topographic and vegetational zones of the coastal system. However,
no relation between these factors and vegetational zonation was found.

e. Conclusions. Many of the factors thought to affect species
behavior in the coastal dune system have been studied, but several
aspects of this complex ecosystem are still controversial. Careful
studies must be conducted before these contradictory points of view
can be resolved.

II. PROCEDURE
Ws Floristics.

Plant collections were made frequently throughout the period March
1974 through May 1975. Diagnostically mature specimens were collected
in duplicate, identified, mounted, and labeled.. Labeling information
included collection location, date, collectors, common associated
species, and a brief habitat description (Radford, Ahles, and Bell,
1968). The duplicate collection has been deposited in the Old
Dominion University Herbarium.

2. Vegetational Studies.

The objectives of these studies were to (a) delimit and characterize
the number and diversity of plant communities that occurred on the
Study area, (b) determine various phytosociological parameters of these
communities through randomized sampling procedures, (c) produce an
accurate vegetational map of the study area, (d) characterize the
relationships between the delimited communities, and (e) locate and
map a representative series of permanent quadrats.

17

a. Plant Community Identifications. Initially, the study area

was traversed extensively to gain a general overall impression of its
vegetational diversity. An east-west hase line was then established
along the southern boundary of the study area,using a transit and
Stadia rod (elevations were not determined). Nine north-south line
transects were than run from the base line at 300-foot intervals.
Vegetational descriptions were made along these transects, and nine
distinct community types were defined; two additional community types
were later added.

b. Community Sampling. The subjective community identifications
verified observations made prior to the initiation of this investigation.
Only two physiognomically distinct vegetational were identified: (a)
Areas dominated by grasses and forbs, and (b) areas dominated by shrubs
(some with trees associated).

Previous experience with similar vegetational types has demonstrated
that 0.2- by 0.2-meter-square quadrats arranged in a stratified random
manner will produce statistically valid results in grass-dominated and
forb-dominated vegetation (Levy, 1970). Good results can be obtained
using 1- by l-meter-square quadrats in shrubby vegetation and 4- by 4-
meter-Square quadrats in areas dominated by shrubs and small trees.

Several techniques were used to determine the number of plots
required for a statistically adequate sample. A running mean was
determined for important species in each community by plotting.the
value obtained for a dominant species in the first sample quadrat
and the mean of plots 1 and 2, 1, 2, and 3, etc. A sample was judged
adequate when the regression line generated by this procedure no longer
varied more than 10 percent from previously obtained mean values for
the population. Five plots were sampled in shrub-dominated communities
and 20 in grass-dominated communities prior to running mean analyses;
this procedure was conducted for two or three dominant species in each
community. Enough samples were obtained to ensure adequate sampling
of all dominant species. In all cases, rarer species (those with low
frequency of occurrence in the sample plots) were inadequately sampled.
Results were verified by examining variance, mean, and standard error
of the populations. All methods indicated that adequate data had been
collected for a reliable quantitative description of the various
vegetational types.

The sound—side shrub community appeared to require only five quadrats
for an adequate sample. Frequency data were collected from five
additional random plots to ensure sampling adequacy. The results of
the five initial plots were compared to the data collected for all ten
plots, using a chi-square test. The value obtained was less than the
0.05 level of confidence, demonstrating that no significant difference
occurred between these two sets of data. Therefore, the first five
samples were used to represent this community.

Quadrat frequency and species' standing crop data were collected
for all communities except the sound~side shrub community. For the
latter type, frequency for all species and rooted stem density were
determined. Standing crop was determined in grams of aboveground
oven-dried tissue. Each quadrat was clipped, separated by species,
and individual species oven-dried to constant weight at 105° Celsius,
Five 1- by l-meter plots and 330, 0.2- by 0.2-meter plots were examined
and clipped.

c. Vegetational Mapping. Three overflights of the study area were
made at different seasons. Infrared aerial photos obtained were used
with the ground truth from the nine previously described transects
and the quadrat data to produce a scale vegetative map of the study
area. The accuracy of this map is better than 90 percent.

d. Ordination of Stands. The 11 community types were arranged
in an ordination model according to the method of Bray and Curtis
(1957). In this method each community's frequency values were summed.
Each individual species frequency in the community was divided by the
total for all species and the result multiplied by 100 to yield
relative frequency expressed as a percentage. The relative frequency
values were then used to compare the species composition of each
community with the other communities, using the Index of Similarity
(IS), IS = 2w/(a + b) (Bray and Curtis, 1957). The IS values were
then subtracted from 100 to yield an Index of Dissimilarity (ID),

ID = 100 - IS, and used to locate the communities along an axis by
means of Beals' (1960) adaptation of the Pythagorean Theorem,

2 PAIN 2
X= en (ny (ee ,» and the technique of Bray and Curtis (1957).

The usual procedure for axis extraction is to sum the ID values for
each vegetational unit. The stand with the highest sum is deemed the
end of an axis; the opposite end of this axis is the stand least like
it (i.e., having the highest ID in relation to it). The units are
represented as points separated by a scale distance equal to the ID
value. All other points are then located between the end points.

In this study, however, one stand had ID values of 100 with five
other stands, and a second stand had the same relationship with four
other stands. To produce a model of acceptable geometry it was
necessary to modify the standard axis extraction technique in the
following manner. The first end point of the X axis was the community
having the highest ID sum. The second end point was the stand with
the highest ID value in relation to it, excluding those stands with
values of 100. The Y axis was selected in a similar fashion from a
matrix of ID values composed of the remaining nine communities. The
Z axis was selected using a matrix from the remaining seven stands.

e. Permanent Quadrats. Three permanent 5- by 5-meter quadrats
were located in each of the nine defined community types and three

I)

were located in the barren dune areas. Only one quadrat was located in
both the wetlands community and the spurge-sandgrass community due to
their small extent and uniformity. The quadrats were laid out
diagonally north and south and oriented with a permanent survey

marker in the east corner. Locations of the permanent quadrat

markers were determined by measuring direction and distance to the
nearest geographic monument marker. Vegetative pattern, gross
composition, and percent ground cover for each permanent quadrat were
determined and mapped on graph paper.

III. RESULTS
ike Floristics.

The flora is composed of approximately 178 species and 132 genera,
representing 58 families. The list of species collected (with their
common names when applicable) is presented in Table 1.

2 Vegetational Studies,

a. Phytosociology. Studies indicate that 11 different community
types can be delimited. Some of the community types are very distinct,
i.e., foredune, oceanside shrub, sound-side shrub, and wetlands. The
sound—side disturbed community is relatively distinct, but the
remaining six community types are qualitatively, and in some cases,
quantitatively similar. The phytosociological data for these stands
are presented in Tables 2 through 12.

The sound-side shrub community (Table 8) has the greatest number
of species (28), followed by the sound-side disturbed community (24)
(Table 10), and the wetlands community (22) (Table 11). The oceanside
intershrub community (Table 3) had 16, while the planted American
beachgrass (Table 5) and low dune grass (Table 7) communities each had
10; the remaining communities had 7 species each except for the
planted bitter panicum community (Table 9), which had only 6.

The oceanside shrub community (Table 4) had the greatest biomass
with 2779.3 grams per square meter dry weight. Though biomass data
were not obtained for the sound-side shrub community (Table 8) due to
the impracticality of cutting down the few trees and large shrubs, its
density of 36.9 individuals per square meter for all species, and
especially its density of M. pensylvantca (15), Prunus serotina (black
cherry) (8), Sassafras albtdwn (sassafras) (2.4), and Drospyros
virgintana (persimmon) (4.4) suggest that the standing crop biomass
of this community is of the same magnitude as that of the oceanside
shrub community. The wetland community (Table 11), which had the
second highest measured biomass, had only 104.9 grams per square meter.
The remaining values ranged from 68.8 grams per square meter for the
sandgrass-buttonweed community (Table 6) to 5.6 grams per square meter

20

Table 1. Duck Field Research Facility floristics list.

Family and Species Common Name

Family Aceraceae
Acer rubrum L. Red maple

Family Aizoaceae
Mollugo vertictllata L. Carpet weed

Family Alismataceae
Sagttarta graminea var. weatherbiana
(Fernald) Bogin Arrowhead

Family Amaranthaceae
Alternanthera phtloxerotdes
(Martins) Grisebach Alligator weed

Family Anacardinaceae
Rhus copalltna L. Winged sumac
R. radicans L. Poison ivy

Family Apiaceae
Centella astattea (L.) Urban
Eryngtum aquatteum L. Eryngo
Hydrocotyle umbellata L. Marsh pennywort
Iilaeopsts carolinensts C. & R.
Ptiltmntum captllaceun (Michaux) Ref.
Stum suave Walter Water parsnip

Family Aquifoliaceae
Ilex opaca Aiton American holly
I. vomttorta Aiton Yaupon

Family Asclepiadaceae
Ascleptas lanceolata Walter Milkweed

Family Aspleniaceae
Asplentum platyneuron (L.) Oakes Ebony spleenwort

Family Asteraceae

Achillea millefolium L. Yarrow
Ambrosta artemtsitfolta L. Ragweed
Aster tenutfolius L. Aster
Baecharts halimtfolta L. Groundsel tree
Btdens mitts (Michaux) Sherff Beggar ticks
Carduus sptnostssimus Walter Yellow thistle
Crepts vestecarta ssp taraxtfolia

(Thuillier) Thellung Hawk's beard
Ecltpta alba (L.) Hasskar Yerba-de-tago

2 |

Table 1. Duck Field Research Facility floristics list-Continued.

Family and Species

Family Asteraceae (concl'd.)

Erigeron canadensts var. canadensis L.

E. candensts var. pustllus (Nuttal1)
Ahles

Eupatortun captllifoltum var.
captllifoltwn (Lam.) Small

E. serotinum Michaux

Gatllardia pulchella Foug.

Gnaphaltum obtustfoltum L.

Hteractum gronovitt L.

Heterotheca adenolepts (Fernald)
Ahles

H. gossyptna (Michaux) Shinners

Iva frutescens L.

I. tmbricata Walter

Krigta virginica (L.) Willd.

Lactuca canadensts L.

Mikanta scandens (L.) Willd.

Pluchea foetida (L.) D.C.

P. purpurascens (Swartz) D.C.

Pyrrhopappus carolintanus var.
earolintanus (Walter) D.C.

Soltdago rugosa var. rugosa Miller

S. sempervirens L.

S. tenutfolta Pursh

Xanthium strumartum var.
strumarium L.

Family Bignoniaceae
Campsts radicans (L.) Seemann

Family Brassicaceae
Cakile edentula (Bigelow) Hooker
Leptdtum virginteum L.

Family Cactaceae
Opuntta compressa (Salisbury) Macbride
0. drwmnondit Graham

Family Campanulaceae
Lobelta elongata Small
Specularia perfoltata (L.) A. D.C.

Family Caprifoliaceae

Lontcera japontca Thunberg
L. sempervirens L.

22

Common Name

Horseweed
Horseweed

Dog fennel
Thoroughwort
Blanket flower
Rabbit tobacco
Hawk weed

Marsh elder
Seashore elder
Dwarf dandelion
Wild lettuce
Climbing hempweed
Marsh fleabane

Salt marsh fleabane

False dandelion
Goldenrod
Goldenrod
Goldenrod

Cocklebur

Trumpet vine

Sea rocket
Peppergrass

Prickley pear
Fragile prickley pear

Marsh lobelia
Venus! looking glass

Japanese honeysuckle
Coral honeysuckle

Table 1. Duck Field Research Facility floristics list-Continued.

Family and Species Common Name

Family Chenopodiaceae
Chenopodium ambrosiotdes L. Mexican tea

Family Cornaceae
Cornus florida L. Dogwood

Family Convolvulaceae
Calystegia septum (L.) R. Brown Hedge bindweed

Family Cucurbitaceae
Melothria pendula L. Creeping cucumber

Family Cyperaceae
Carex alata Torrey Sedge
Cyperus dentatus Torrey Sedge

erythrorhtzos Muhl.

. ftltictnus Vahl

haspan L.

ovularts (Michaux) Torrey

rtvularts Kunth

. sesquiflorus (Torrey) Mattfeld and

Kukenthal

C. strigosus L.

C. surtnamensts Rottboell

aAgagaa

Eleocharts tuberculosa (Michx.) R. & S. Spike rush
Fimbristylts autumalts (L.) R. & S. Sand rush

F. dtichotoma (L.) Vahl

Futrena squarrosa Michaux Umbrella grass
Scirpus amertcanus Persoon Chair maker's rush

Family Ebenaceae
Dtospyros virginiana L. Persimmon

Family Euphorbiaceae
Croton glandulosus var. septentrtonalts

Muell.-Arg. Croton
C. punetatus Jacquin Croton
Euphorbta polygontfolta L. Beach spurge

Family Fabaceae
Aptos americana Medicus

Cassta fasciculata Michaux Partridge pea
Centrosema virginianum (L.) Bentham Butterfly pea
Desmodium pantculatum (L.) D.C. Beggar lice
D. pauctflorum (Nuttall) D.C. Beggar lice
D. strtetum (Pursh) D.C. Beggar lice
Lespedeza capttata Michaux Bush claver

23

Table 1. Duck Field Research Facility floristics list-Continued.

Family and Species

Family Fabaceae (concl'd.)
L. cuneata (Dumont) G. Don
L. strtata (Thunberg) H. § A.
L. virginica (L.) Britton
Strophostyles helvola (L.) Ell.

Family Fagaceae
Quercus vitrgintana Miller

Family Gentianaceae
Sabatita dodecandra var. dodecandra
(bs) BoS. 2B.

Family Hamamelidaceae
Itquidambar styractflua L.

Family Hypericaceae
Hyperteum genttanotdes (L.) B.S.P.

Family Juncaceae
Juncus cortaceus Mackenzie
J. megacephalus M.A. Curtis
J. roemertanus Scheele

Family Juncaginaceae
Trtglochin strtata R. & P.

Family Lamiaceae
Monarda punctata L.
Salvta lyrata L.
Stachys nuttalltt Shuttlew

Family Lauraceae
Persea borbonta (L.) Spreng.

Family Liliaceae
Smilax bona-nox L.
Yucca ftlamentosa L.

Family Linaceae
Linum virgtntanum var. medtwm Planchon

Family Loganiaceae
Polypremum procumbens L.

24

Common Name

Japanese clover

Wild bean

Live oak

Sea pink

Sweet gum

St. John's wort

Rush
Rush
Black rush

Arrow grass

Horsemint
Sage
Hedge nettle

Red bay

Greenbrier
Bear grass

Flax

Table 1.

Family and Species

Family Lycopodiaceae
Lycopodium appressum (Chapman) Lloyd and
Underwood

Family Lythraceae
Lythrum lineare L.

Family Malvaceae
Htbtseus moscheutos L.
Kosteletskya virginica (L.) Presl.

Family Myricaceae
Myrtca certfera var. certfera L.
M. pensylvanteca Loisel

Family Onagraceae
Oenothera btennis L.
0. fruttcosa L.
0. humtfusa Nuttall

Family Orchidaceae
Sptranthes cernua (L.) Richard

Family Pinaceae
Pinus taeda L.

Family Phytolacaceae
Phytolaecea amertcana L.

Family Plantaginaceae
Plantago lanceolata L.

Family Poaceae
Andropogon elltottit Chapman
AL vergentcus: Lb.
Ammophila breviligulata
Bromus secalinus L.
Cenchrus trtbulotdes L.
Cynodon daectylon (L.) Persoon
Digttarta filtformts var. villosa
(Walter) Fernald

D. tsehaemum (Schreber) Schreber ex Muhl.

D. sangutnalts (L.) Scopoli
Echtnochloa waltert (Pursh) Heller
Eleustne indica (L.) Gaertner
Elymus virgtnicus L.

Eragrostts elltotttt Watson

25

Duck Field Research Facility floristics list-Continued.

Common Name

Club moss

Loosestrife

Rose mallow
Sea shore mallow

Wax myrtle
Bayberry

Evening primrose

Sundrops
Evening primrose

Nodding ladies' tresses

Loblolly pine

Pokeweed

Plantain

Broom straw

Broom sedge
American beachgrass
Brome grass
Sandspurs

Bermuda grass

Crab grass

Crab grass

Crab grass

Walter's barnyard grass
Goose grass

Wild rye grass

Love grass

Table 1. Duck Field Research Facility floristics list-Continued.

Family and Species Common Name

Family Poaceae (concl'd.)
E. spectabtlts (Pursh) Steudel
Ertanthus gtganteus (Walter) Muhl.
Festuca seturea Nuttall
Leptoloma cognatum (Schultes) Chase
Panteun amarulum Hitchcock and Chase

amarum Ell.

datehotomtflorum Michaux

seopartum Lam.

vagtnatum Swartz

. virgatum L.

Polypogon monspeltensts (L.) Desf.

Sacctolepts striata (L.) Nash

Setarta gentculata (Lam.) Beauvois

Sorgum halepense (L.) Persoon

Spartina cynosurotdes (L.) Roth

S. patens (Aiton) Muhl.

Sphenopholts obtusata (Michaux) Scribner

Trtplasts purpurea (Walter) Chapman

Trtsetum pensylvantcum (L.) Beauvois
Gre Io G So

Love grass
Beard grass
Fescue

Witch grass
Bitter panicum
Panic grass
Fall ronieum

ry fo tO NO fo

Switch grass
Rabbit foot grass

Fox tail grass
Johnson grass
Giant cord grass
Salt meadow grass
Wedge grass

Sand grass

Untola pantculata L. Sea oats
Zea mays L. Corn
Family Polygonaceae
Polygonum hydroptperotdes var. opelousanum
(Riddell ex Small) Stone
P. pensylvantcum L. Knot weed
P. sagtttatum L. Tear thumb

Rumex acetosella L. Sheep sorrel
R. vertictllatus L. Swamp dock

Family Pontederiaceae
Pontederta cordata L. Pickerelweed

Family Primulaceae
Samolus parviflorus Raf. Water pimpernel

Family Ranunculaceae
Ranunculus sardous Crantz Buttercup

Family Rosaceae
Amelanchter arborea var. laevis

(Wiegard) Ahles June berry
Prunus serotina var. serotina Ehrhart Black cherry
Rubus betultfoltus Small Blackberry

26

Table 1.

Family and Species

Family Rubiaceae
Diodia teres Walter
D. virgintana L.

Family Rutaceae
Zanthoxylum elava-herculis L.

Family Salicaceae
Saltx ntgra Marshall

Family Scrophulariaceae
Agalints purpurea (L.) Pennel
Linarta canadensts (L.) Dumont
Verbascum thapsus L.

Family Solanaceae
Physalts viscosa ssp maritima
(M.A. Curtis) Waterfall
Datura stramontum L.

Family Urticaceae
Boehmerta cylindrica (L.) Swartz

Family Verbenaceae
Calltearpa americana L.
Ltppta nodtflora (L.) Michaux

Family Vitaceae
Parthenoctssus quinquefolta (IL.) Planchon
Vitis aestivalis var. aesttvalts Michaux
V. rotundtfolta Michaux

Family Xyridaceae
Xyrts juptcat Richard

27

Duck Field Research Facility floristics list-Concluded.

Common Name

Buttonweed

Hercules! club

Black willow

Gerardia
Toad flax
Mullein

Ground cherry
Jimson weed

False nettle

French mulberry
Frogbit

Virginia creeper
Summer grape
Muscadine

Yellow-eyed grass

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zoyoj naLiupd v7101Un

1979W
ezenbs
zed sueiry

(queodz0d)
Aduenberj
SATIETOY

(queozed)
Aduenber

(suezs )
2YSTOM TeIO]

satoeds

7° A2tunuuo0s eunpedzoy; °7 oTqRy

28

Table 3.

Species

Triplasts purpurea
Heterotheca gossypina
Untola paniculata
Ammophila brevtltgulata
Cenchrus trtbulotdes
Cyperus ovularts
Ertgeron canadensts
Euphorbta polygontfolta
Panteum amarum
Soltdago sempervirens
Spartina patens
Physalts vtscosa
Oenothera humifusa
Diodta teres

Oenothera betnnts
Opuntta drummondtt

Total

Oceanside intershrub community. !

Frequency

(percent)

Relative
Frequency
(percent )

1 Based on 51, 0.2-meter-square quadrats.

29

Grams per
Square
Meter

€°6LL°Z
z°0

ZO

9°Z
9°ST
lee

8°L
SSL?

LI319N

ezenbs
zed swer9

°szeipenb orenbs-19}0uI-9°[ ‘Ss uo peseg

T° 968 ‘ST

T°69L°ST

(suei3)
2YSTOM TeI0]

(queo10ed)
Aduenber4
SATIELOY

(qued10d)
Aduenberj

7 Az Tunwuos qnzys eptsuescg9

T

TeIOL
snzjof1qnzeq snqny
xOU-DUog xDI IWS
purqzotes snundg
sapio1soaiquo untpodouey)
suadtazeduas obpp1j0g
pupatdieuw v20D107/AYg

potupayhsued vorthy

setoeds

“vy eT9PL

30

°sqeipenb ozenbs-s1ojoW-7°0 “gf UO peseg

98°8t 0°LL9 Teq0L
c°0 pyoynoL1espf DLssD)
vst un] niDuo wnarUDg
£0°0 stapjnao snxedhg
La unapun una Lung
£°0 p1,0f1u0bhyod viqroydng
T°6 psnf[iuny Diteay,zOUaQ
£0°0 Sadie. DLpo1g
SO sap1o71ngidz snqyouag
5 GG 0401nb111AeNq D1 1Y4douuy
£°0 peundind s1sp07 dit],
1339W (swez3) (quedzed) (qus9 20d)
erenbs VYSTOM [eIO]L Aduonber4 Aduoenberj sotoeds
zed sweig OATICTOY

, A} Tunuu0S ssersyoveq UedTIowy peqwetTd °S e1q¥L

3|

6°Z2
¢°¢s
1929;

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TeIoL
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snsob1azs snizedhg
psnfIumy DLEYZOUeO
unIDUIO UuNnoLUDg
Sap1071Nnqiaiz sntyouay
Sadeq DLpo1g

peindind s1isp7 dit],

(swe) (queozed) (qued.210d)
JYUSTOM TeIOL Aduenbary Aduenber4 setoeds
SATIETOY

, 42 TunwUOS peemuoz3Ng-ssexrspues °O 9 qPL

Ba

6Si
£°0
T°0
L°0
T°0
T°0
T°0
T's
v°6
9°0
L°0
19790W

azenbs
zed swear

c° 9ST

(suez3)
1YSTOM TeI0L

°sqzerpenb orenbs-19290UI-7°9 “Sp uo paseg I

1210],
untou WndLUuD_
s1apjnao sniadhg
sisuaepouvs uodebritq
psnfiuny DLoYyZOuUaQ
pijzofiuobhiod vaqazoydng
Sadie} DIpoIg
pypjyno1und v701UN
0qp1nby1 Laeaq v1 2y4douuy
poundind sisv7.d1a],

Sep10jnqidy sniqyoua)

(que0.10d) (}ue 020d)
Aguenberj Aduenberj sotoeds
DATIETOY

, A}tunumuos ssez3 ounp MoT °/ 9eTqQeL

35

Table 8. Sound-side shrub community. !

Relative
Species Frequency Frequency
(percent) (percent )

Myrtca pensylvantea
Prunus serottna

Soltdago tenutfolta
Smilax bona-nox

Rubus betultfoltus
Ambrosia artemisttfolta
Rhus copatltna

Soltdago sempervirens
Melothria pendula

Rhus radticans
Chenopodium ambrostotdes
Ammophtla breviltgulata
Dtospyros virgintana
Vitts aesttvalts
Parthenoctssus quinquefolta
Ditodta teres

Panteunm amarulum
Achtllea mtillefoltum
Panteum amarum

Sassafras albtdum
Trtplasts purpurea
Eupatortum captlitfoltum
Cenchrus trtbulotdes
Cassta fasctculata
Zanthoxylum elava-hercults
Juncus megacephalus
Euphorbta polygontfolta
Oenothera humtifusa

1 Based on 10, 4.0-meter-square quadrats.

34

Density

10.

m NH
oo oO

po
RP NO FP FP HRP KY KF DH N

oN 2 mS) Ss Cc oo Ss YP WS Oo eo eo 2S Ss SHS oo Ss 2 AQ eo WY EH CC) &

So oo oo bo PP Y& Ww ey

*sqzeipenb oxenbs-1ejou-7°9 “GT uo peseg |

p07 nb111AaAeAaq D7 1y4douuy

Z°0 psnfiuny DitaYyzOUeC
8°0 Satie, v1po7g
8°0 Sap1o7nqiiy sniyouag
8°T peandind sisp7 dij
ST uncpuno wnoLUDdg

1019W
azenbs
zed swery

(swex3) (queoz10ed) (qued10d)
VYUSTOM TeIOL Aduoenbe tJ Aduonbsr4 sotoeds
OATIETOY

; Adtunuumos umotued 10331q pequetd °6 STGeL

35

Table 10.

Species

Trtplasts purpurea
Erigeron canadensts*
Solidago tenutfolia*
Spartina patens
Panteum scopartum
Oenothera humifusa
Diodia teres
Ambrosta artemisstfolta
Juneus cortctous
Panteum amarum
Rubus betultfoltus
Soltdago sempervirens
Ammophila breviligulata
Myrica pensylvanica?
Euphorbia polygontfolta
Cenehrus trtbulotdes
Cynodon dactytlon
Strophostyles helvola
Leptdtum virginteum
Cassta fasctculata
Setrpus amertcanus
Juncus megacephalus
Lespedeza striata
Saltx nigra :

Total

1

Frequency
(percent)

Relative
Frequency
(percent)

Based on 37, 0.2-meter-square quadrats.

2 Counted together because of Similarity.

3

Biomass of trees not determined.

36

Sounc-side disturbed community. +

Grams per
Square
Meter

Table 11. Wetlands community. !

Species

Setrpus amertcanus
Aster tenutfoltus
Disttchlts spteata
Eleocharis sp.
Hydrocotyle umbellata
Cyperus spp.

Juncus roemertanus
Spartina patens
Bidens mitts

Lythrum lineare
Soltdago sempervirens

Centella astatica

Polygonum pensylvantcum

Sagtttarta gramtnea var.

weatherbtana

Fimbristylts autwmaltis

[Itlaeopsts carolinensts

Pontederta cordata
Oenothera fruttcosa
Sacectolepsts strtata
Stum suave

Soltdago tenutfolta

Pontederta cordata

Total

Relative Grams per
Frequency Frequency | Weight Square

(percent) (percent ) Meter

VoZ
14.5
11.8

2.6

0.6

Boll
23.4
Sie
14.1

3.0

Bow

0.2

0.2

2.8
0.8
0.02
0.3
1.6
0.02
hoe
0.3
0.3

422.6 105.74

1 Based on 20, 0.2-meter-square quadrats.

37

8° IT

10195~W
azenbs
zed sweiry

°szerpenb-orenbs-19}0ul-7°9 “QZ uO poseg

(swez3) (queozed) (quesd1ed)
2YSTIOM [BIOL Aduoenbe Lj Aduoenbery
OATIET OY

yA FunuM0S sseispues-oa3inds

[e201
unIDun unoLUDg
Sap1ojnqiiz sntyoua)
suaqod vu1zands
psnfiuny Diey4zoUuEeQ
$aie4 DLpo1g
poandind sisv7 diay,

p1z0fiuobhj0od vaqaoydny

setoeds

“CT 9T9PL

38

for the planted bitter panicum community (Table 9). The sand dune areas,
which were devoid of vegetation, had no measurable biomass.

b. Vegetational Map. The vegetational map of the study area is
presented in Figure 2. A summary of the approximate number of acres of
each community, the barren sand dune areas, and roadways is presented
in Table 13. The Duck Field Research Facility covers approximately
150 acres which includes 3 acres of roadways and 27 acres of barren
sand dunes. The low dune grass community had the largest area of any
community type, approximately 40 acres, and the planted American
beachgrass community was second with 30 acres. The remaining values
ranged from 14 acres for oceanside intershrub to about 1 acre for the
wetlands community.

c. Community Ordination. The results of ordination techniques
are presented in Figures 3, 4, and 5. The three-dimensional aspects
of this model have been depicted by graphing two axes at a time. The
distances between individual communities are related to the relative
differences between them. It should be emphasized that ordination
serves merely as a tool, which when carefully used, can yield a variety
of information. By itself, however, it provides only an interesting
arrangement of points and serves little purpose.

The distinctness of the foredune and wetland communities is borne
out by the X-Y and X-Z perspectives (Figs. 3 and 4), which indicate the
distinctness of the sound-side shrub and oceanside shrub communities.

The X-Z and X-Y axes suggest that the oceanside and intershrub community
is distinct, but it is closely allied with many stands on the X-Y aspect.

The sound-side shrub community is distinct on all three axes,
although it shows some similarities to the large centrally located
group of communities on the X-Z axis.

In contrast to the clearly distinct community types, six community
types (planted American beachgrass, sandgrass-buttonweed, low dune grass,
planted bitter panicum, sound-side disturbed, and spurge-sandgrass )
have strong similarities to each other.

d. Permanent Quadrats. Figures 6 through 30 show most of the
permanent quadrats. Low dune grass quadrat 2 has not been diagramed
as it was vacant. The three permanent plots located in the oceanside
shrub community were omitted as they had 100 percent M. pensylvantca
coverage. Only one plot was located in the wetlands community (Fig.
29), and spurge-sandgrass community (Fig. 30). The three permanent
quadrats which were located on barren dune sites have not been
represented as figures since they are featureless. Table 14 lists
separately the location of each quadrat's permanent survey marker.
The usefulness of these plots will be realized as their compositional
changes are mapped over the years.

39

uesdQ 9TIUeTIYV

punos yonqtTzIn9

Foot

ine)
oO
La)
uy
2) n
‘m) Yn
7) au)
“d 4
uU tof)
mo)
ma) iS)
ue) 3
own NN
nud 1
la oO
Uo ley)
Gov Hy
Dees
oon a,
WNEznNW

el
i
ig
iS

but tonweed
trex

Low dune grass

side shrub
i

Planted b

n
7)
i)
u
lor)

uU
8
Ww

Sound—

fe)
3}
uy
SS
7) S
HO
os 0
Psd
eC
oH YW oO
oof
oud
GH U
ee)
CS_-ee 2)
oadgd
4H oo a
000
[Eq (S) (6) (2%

.
i
NY

panicum

beachgrass

f the Duck Field Research Facility.

ion map o

Vegetat

Figure 2.

40

Table 13, Summary of community types at the Duck Field
Research Facility with approximate acreages.

Acres

Community Types (approximate)
Foredune (FD) 3.5
Oceanside intershrub (OIS) 14.0
Oceanside shrub (OSS) 11.0
Planted American beachgrass (PBG) 30.0
Sandgrass-buttonweed (SG/BW) 5.0
Low dune grass (LDG) 40.0
Sound-side shrub (SSS) 7.0
Planted bitter panicum (PBP) 3.0
Sound-side disturbed (SSD) 4.0
. Wetlands (WL) 1.0
Spurge-sandgrass (S/SG) os
Barren sand dunes 27.0
Roads and trails 3.0
Total 150.0

4|

0 10 20 30 40 50 60 70 80 90
X AXIS

Figure 3. Ordination of plant communities at the Duck Field
Research Facility, showing the X and Y axes
perspective. (See Table 13 for definition of
community type designation. )

42

100

100

0 10 20 30 40 50 60 70 80 90 100
X AXIS

Figure 4. Ordination of plant communities at the Duck Field
Research Facility showing the X and Z axes
perspective. (See Table 13 for definition of
community type designation.)

43

100

@oOss
90 @S/SG
80
SG
pReO@r cet
70
ssD@ lg BBC
60 ae
<
<

© 10 20 30 40 50 60 70 80 90 100
Y AXIS

Figure 5. Ordination of plant communities at the Duck Field
Research Facility showing the Y and Z axes
perspective. (See Table 13 for definition of
community type designation. )

44

Ma Ma
Me,

Ma,
Mn,

Mi

en,

Rr
we"

ow,

WM Me.

Me
My,

MAM,

cat

90 percent Ammophila breviligulata

10 percent Untola panteulata

Total ground cover - 95 percent

Figure 6. Foredune community permanent quadrat 1.

45

tle

Mee or, Ate B A . F Me QV
SY ¥;

Mi Me
Me

Mis yy

Ms
Mh, Me

Me vt

SR

Me a,
My,

My,

Me
aM, Niles
Me
as
Ms

Mu
My

Mh

M
ote lt ally,

AM

Me

0

Me
aM,

80 percent Untola pantculata

20 percent Pantewun amarum

Total ground cover - 90 percent

Figure 7. Foredune community permanent quadrat 2.

46

».
Mi,

lh,

at.
Ma

ake
Me yu Mi Se
a,

iM

ae

40 percent Panicum amarum
40 percent Ammophtla brevtlitgulata
20 percent Untola panteulata

Myrtea pensylvantca

Opuntta compressa

Total ground cover - 90 percent

Figure 8. Foredune community permanent quadrat 3.

47

<

MM Me,

MM.
aw

wv.

ae,

at Me

alu,
Ma
aM

a7
Sh

Macey

we MO aM,

‘ Me
Me ot My

Ma Ma, M
aM, lM

a oF
NYA a.

wM My,
MM,
orn
th

sb re
Me Me
Me whe

My,

100 percent Untola pantculata

Myrtea pensylvanica

Total ground cover - 90 percent

Figure 9. Oceanside intershrub community permanent quadrat 1.

48

Me
M

100 percent Untola paniculata

Total ground cover - 40 percent

Figure 10. Oceanside intershrub community permanent quadrat 2.

49

Ma ple
aie at, ott

he

Figure 11.

Ma atte

100 percent Untola paniculata

Total ground cover - 40 percent

Oceanside intershrub community permanent quadrat

50

Total ground cover - 30 percent

Figure 12. Planted American beachgrass community permanent
quadrat 1.

S|

“Ma
Me

oh,

M.
ah
MMe

Me
My, Me

Tae,
a, Me

ae My,

ln Me

«on f 100 percent Ammophtla breviligulata

M4,

m

Total ground cover - 30 percent

Figure 13, Planted American beachgrass community permanent
quadrat 2.

OZ

Total ground cover - 20 percent

Figure 14. Planted American beachgrass community permanent
quadrat 3.

SS)

Me,
Mi, ate
a,

MeMe

Ma

Me

90 percent Trtplasts purpurea
10 percent Panteum amarum

Total ground cover - 70 percent

Figure 15. Sandgrass-buttonweed community permanent quadrat l.

54

90 percent Trtplasts purpurea
10 percent Panteum amarum

Total ground cover - 75 percent

Figure 16. Sandgrass-buttonweed community permanent quadrat 2.

55

aM,
we

MM,
oi, a7
ly, eM,
ate, Miiths

oh
Me

a
te

95 percent Triplasis purpurea
5 percent Ammophtla breviltgulata

Total ground cover - 95 percent

Figure 17. Sandgrass-buttonweed community permanent quadrat 3.

56

<n
f silh
on, ae
Pe orn
om

ay,

100 percent Untola panteulata

Total ground cover - 35 percent

Figure 18. Low dune grass community permanent quadrat 1.

SU

Me
ny

2 Ys Me
i Mh My,

100 percent Untola paniculata

Total ground cover - 35 percent

Figure 19. Low dune grass community permanent quadrat 3.

58

C)
0,0°o
o © eo

0°o

oo
o.,90

°
0°,
O04
0,0 °
oo ° (i) ° oo.
2,0 ©,° 9,0
©0 4 00 6 09 5
250° 259° 95°
0 P92 6 9° g 0°
6 ° 2,0
°

°° Ce °°
©,0°0,0°0, 0

°

O96
°° EX) 0°
0, FRE 0% 0 0°

°
°

©,°0

oo oo
°
@,° °0O

oo
2,0

oo
9,°

eo oo oo 0°

°

°o.90
o
° oo

Prunus serotina

X

Myrica pensylvanteca

lax bona-nox

mL

Rubus betultfoltus

Si

- 95 percent

Total crown cover

Total ground cover - 15 percent

Sound-side shrub community permanent quadrat 1.

Figure 20.

D)e)

°
°

a?
bar)
oo

oo
eo
°o,°0

°o

°
°

- ee)

°
CO 4
°

°°

0° 0,0° 90
00 8 00 §
o 08,0

.)
)

0.0°0,0°0,0
co ° 0c 2 oo

00 8 00 6 00 &
0,9. 0,° 0,0

Eg Myrtca pensylvantca

X

Prunus serotina

Rubus betultfoltus
Ammopht la breviltgulata
Smilax bona-nox

ta

folt

Parthenoe

tSSUS Guinque

- 95 percent

Total crown cover

Total ground cover - 10 percent

Sound-side shrub community permanent quadrat 2.

Ble

igure

IP

60

Rhus copatlina
Myrtca pensylvantca

Rubus betultfoltus
Vitts aesttvalts
Anmophtla brevtligulata
Smilax bona-nox

Total crown cover - 70 percent

Total ground cover - 10 percent

Figure 22, Sound-side shrub community permanent quadrat 3.

6l

Me Ale
Mee

ot,

99 percent Panteun amarum
1 percent Ammophila brevtligulata

Total ground cover - 45 percent

Figure 23, Planted bitter panicum community permanent
quadrat 1.

62

Mn, ale,
ifn,
Me

Me
M

Mn,

wal Mate,

Me Mee gM
Mi

Me

My,

Me,
Me

AM al Mee
ate
My

eM Me

S77
SAU ed

95 percent Pantewn amarum
5 percent Ammophila breviltgulata

Total ground cover - 35 percent

Figure 24, Planted bitter panicum community permanent

quadrat 2.

63

~

80 percent Panicum amarun AN
20 percent Ammophila brevtltgulata

Total ground cover - 10 percent

Figure 25, Planted bitter panicum community permanent
quadrat 3.

64

eae
Ma, i,

Mn,
ee uy

Ma,
Me MM
Mh aM

Mh
<D
als,
sn,

Thy of
Min

Mos, oMae ah,

a, Vi,

Ah Me

Ms
Mi

an alt,

80 percent Panteun scopartum
20 percent Rubus betultfoltus

97 percent Spartina patens
3 percent Pantcum scopartum

95 percent Panteum amarum
4 percent Rubus betultfoltus
1 percent Spartina patens

Total ground cover - 80 percent

Figure 26. Sound-side disturbed community permanent
quadrat 1.

65

oth,
woe

mh,
Yr
Mh,

Me Mee
Mu.

we

MM,

aM
ls,

Mie
Wn.

AV,

50 percent Spartina patens
25 percent Juneus megacephalus and Juneus cortaceus
25 percent Cynodon dactylon and Panteum scopartum

Myrica pensylvantea

Total ground cover - 75 percent

Figure 27. Sound-side disturbed community permanent quadrat 2.

66

Pd Myrtca pensylvantca

Panteum amarum

Total ground cover - 10 percent

Figure 28. Sound-side disturbed community permanent
quadrat 3.

67

@
ar)
00
°°?
°
0°

?
q
q
]

o0® © © © © CO e e@,
0 ®@ © © @ OC ee 0°00

@eeeeee8 @
°

or. 8 ©@ © @ 6 6 @

eee 8 @ © © O @:6°%:.
°

°

©-e® @@6@0@006 80 @ @ ¢0™
e©eee © © © CO eB
°
°

e@eee@e8 &6hmUhOO OC 8 C&G OO
(+)

@eeeeeegc®ee ee
ee ee)
ee ee ee ee

o0®

°
°
1)

°
ce © eee 6&6 & © eo eo
oo
0,9
0°o
oo
0,0

°
°
°
°
°

oo
°
°
e
°

oo
°
°

oo

Spartina cynosurotdes
Sagtttaria graminea

80 percent Juncus coriaceus
20 percent Hydrocotyle umbellata

Juneus roemertanus

Total ground cover - 95 percent

Figure 29,

Wetlands community permanent quadrat.

68

ov L
aaanee ba
phe a bbb,
yee be
bokeh
veut! a
wl Vb bay

, iy, Me
Mh,
Mw,

Mn Mn
Me at. Ma,

Ma,

An Mn,

MW,

MaMa,

Mu
Mn,

Mi,

My,

Ma,

aM.

50 percent Panteum amarum
50 percent Triplasts purpurea

Spartina patens

Total ground cover - 70 percent

Figure 30. Spurge-sandgrass community permanent quadrat.

69

Table 14.

Quadrat Designation

Foredune 1

Foredune 2

Foredune 3

Oceanside intershrub 1

Oceanside intershrub 2

Oceanside intershrub 3

Oceanside shrub 1

Oceanside shrub 2

Oceanside shrub 3

Planted American beachgrass 1

Planted American beachgrass 2

70

Location of permanent quadrat stakes at the Duck Field
Research Facility (DFRF).

Readings are magnetic.

Location

137°, 97 feet from U.S. Army,
Corps of Engineers (USACE)
survey marker 16.

12°30', 696 feet from USACE
survey marker located in the
southeast corner of the Duck
Field Research Facility (DFRF).

45°, 224 feet from USACE survey

marker located in the southeast

corner of the DFRF.

350°10', 335 feet from USACE
survey marker 11.

350°, 209.2 feet from a point
located 290°, 242.5 feet from
USACE survey marker in the

southeast corner of the DFRF.

330°, 181 feet from USACE
survey marker in the southeast
corner of the DFRF.

320°, 220 feet from USACE survey
marker located in the southeast
corner of the DFRF.

345°, 230 feet from USACE survey
marker located in the southeast
corner of the DFRF.

137°, 293.5 feet from USACE
survey marker 16.

278°, 864 feet from USACE survey
marker ll.

192°, 781 feet from planted
American beachgrass survey
marker 1.

Table 14. Location of permanent quadrat stakes at the Duck Field
Research Facility - Continued.

Quadrat Designation

Planted American beachgrass 3

Sandgrass-buttonweed 1

Sandgrass-buttonweed 2

Sandgrass-buttonweed 3

Low dune grass 1

Low dune grass 2

Low dune grass 3

Sound-side shrub 1

Sound-side shrub 2

Sound-side shrub 3

Planted bitter panicum 1

Location

148°, 408 feet from planted
American beachgrass survey
marker 2.

349°, 233 feet from planted
American beachgrass survey
marker 1.

301°, 332 feet from planted
American beachgrass survey
marker 1.

277°, 545 feet from planted
American beachgrass suryey
marker 1.

349°, 759 feet from planted
American beachgrass survey
marker 1.

320°, 691 feet from planted
American beachgrass survey
marker 1.

326°, 807 feet from planted
American beachgrass survey
marker 1.

280°, 1004 feet from planted
American beachgrass survey
marker 1.

272°, 813 feet from planted
American beachgrass survey
marker 1.

92°, 240 feet from sound side
disturbed survey marker 1.

349°, 374 feet from planted
American beachgrass survey
marker 1.

Table 14.

Location of permanent quadrat stakes at the Duck Field

Research Facility - Concluded.

Quadrat Designation

Planted bitter panicum 2

Planted bitter panicum 3

Sound—side disturbed 1

Sound-side disturbed 2

Sound-side disturbed 3

Wetlands

Barren sand dune 1

Barren sand dune 2

Barren sand dune 3

Spurge-sandgrass

U2

Location

320°, 405 feet from planted
American beachgrass survey
marker 1.

301°, 567 feet from planted
American beachgrass survey
marker 1.

180°, 545 feet from USACE
survey marker 64.

143°, 274 feet from sound side
disturbed survey marker 1.

182°, 283 feet from sound side
disturbed survey marker l.

216°, 230 feet from sound side
disturbed survey marker 1.

290°, 579 feet from a point
2902, 242.5) feet) tromUSACE
survey marker in the southeast
corner of the DFRF.

290°, 890 feet from a point
290°, 242.5 feet from USACE
survey marker in the southeast
corner of the DFRF.

347°, 569 feet from barren sand
dune survey marker 2.

255°, 210 feet from USACE
survey marker 13.

IV. DISCUSSION
1. Floristics.

a. General. Poaceae (grass family) was represented by the most
species, making up 19 percent of the total flora. The Asteraceae
(daisy family) made up 16 percent and Cyperaceae (sedge family) 8
percent. The remaining families represent 2 percent or less of the
total flora. This compares with Burk's (1968) figures for the total
Outer Banks of 14.7 percent for the Poaceae and 13.5 percent for the
Asteraceae.

Among the species collected, Radford, Ahles, and Bell (1964) cited
five as infrequent, three as rare, and one as very rare. The infrequent
plants were Lilaeopsts carolinensts, Eupatoriwn serotinum (throughwort),
Heterotheca adenolepis, Desmodium pauctflorum (beggar lice), and Stachys
nuttallit (hedge nettle). The rare species were Eragrostis elltotttit
(love grass), P. amarulum, and A. breviligulata. The A. brevtligulata
is locally very abundant at Duck due to plantings. Cyperus surinamensts
was noted as very rare. This species was represented by a large clump
of individuals near the edge of the sound in the extreme southeastern
part of the study area.

b. Triplasts purpurea. This species was the most frequent plant
in areas of moderate sand drift which suggests that it may have
importance as a secondary soil stabilizer. If this is the case, this
species, an annual, must show salt-spray tolerance and function as a
soil binder even during the winter when it is dead.

Seed and seedling tolerance to salt and the effect of cold
treatment on seed germination was studied. The seedlings exhibited
a high tolerance to both a salty substratum and to salt spray. Seed
tolerance was as great as that of U. pantculata and A. breviligulata,
the two most important primary dune stabilizers on the Outer Banks
(Seneca, 1969).

Observations-made during the winter of 1974-75 indicate that when
this species occurs in abundance, as in the sandgrass-buttonweed
community, it acts to stabilize sandy areas during the winter months.
The usefulness of this species as a sand binder should be carefully
evaluated with appropriate test plantings.

Die Vegetational Studies.

The plant communities at the Duck Field Research Facility exist
because of the natural environmental forces characteristic of the
Outer Banks and a long history of disturbance. Some types of
disturbance are common to the Outer Banks in general, others are
unique to this site, e.g., its use as a gunnery range.

Ve)

As shown by the ordination (Figs. 3, 4, and 5), some plant
communities are strongly delimited while others are similar, The
distinct communities include the foredune community, which has been
established on the artificially stabilized foredune and enriched by a
series of plantings. This community, though quantitatively unique,
has its component species represented in almost half of the other
communities (49 percent). Hence, the species growing on the foredune
are NOt GEestrrcted) there.) but) represent, those species able Coyresist
the harsh environmental conditions of this habitat. The selecting
factors appear to be strong winds, the accompanying salt spray, sand
abrasion, and evaporative stress, as well as low nutrient levels and
extreme temperatures.

The wetlands community on the sound side of the study area was the
most unique community. There were 21 species in this community, of
which 16 were found nowhere else, and 3 others in only one additional
community. The aquatic nature of these sites has produced the
environmental factors so extremely different from those generally
found at Duck, which easily explains its uniqueness as well as its
species abundance.

Two shrub-dominated communities, the oceanside shrub and sound
side shrub communities, occur at Duck. In the X-Y aspect of the
ordination (Fig. 3), these two communities occur close to one another
and widely separated from all other communities. A similar impression
is given in the Y-Z.aspect (Fig. 5), although they are further apart,
while the X-Z aspect (Fig. 4) shows them at opposite ends of the Z
axis. These communities are unique, but six of the seven species
comprising the oceanside shrub community also occur in the sound-side
shrub community. The seventh,. though not sampled was observed growing
in the sound-side shrub. The major difference between these communities
is the richness of the sound-side shrub community (28 species) and the
dominance of M. pensylvanteca in the oceanside shrub community.

It appears that the oceanside shrub community exists where it does
due to the sheltering effect of the foredune. Conditions here are
still severe enough to exclude all but the hardiest members of the
sound—side shrub community. The oceanside shrub, which appears to be
an early successional stage of what could become like the sound-side
shrub community, is likely to remain in an arrested condition
until it is destroyed by prevailing shoreline dynamics.

The sound-side shrub community is a middle successional stage of
what could become a maritime forest (Wells, 1938). The only significant
species not included in the samples was Quercus vtrgintana (live oak)
which was represented in the community with a small number of scattered
saplings.

The sound-side disturbed community's samples included 24 species,
of which 14 were also found in the sound-side shrub community. Six of

74

the species exclusive to the sound-side disturbed community were those
often associated with highly disturbed locations. This community was
once the site of military housing which was demolished and the site
bulldozed. Prior to the establishment of these buildings the site may
have been partly covered by sound-side shrub and partly by a wetlands
community. At the present time it appears to be almost an ecotone
between sound-side shrub and low dune grass, with the weedy component
relatively stable. As long as these species are not overtopped by
heavy shade-casting competitors they will remain important. M.
pensylvanica is beginning to slowly expand into this area (frequency
5.4 percent).

The two shrub-dominated communities and to some extent the sound
side disturbed community are being enriched by nitrogen-fixing
bacteria on the nodules of legumes and on the roots of M. pensylvanica.
Several excavations revealed the presence of such nodes on M.
pensylvanica at Duck, supporting the results of Morris, et alee @SZA))i

Growing behind the foredune, in front of, between, and immediately
behind the stands of M. pensylvantca which compose the oceanside shrub
community is the oceanside intershrub community. Figures 4 and 5 show
this community to be quite distinct. Of the 16 species included in
the samples of this community, 4 were exclusive; however, the other
species occurred in 43.6 percent of the other communities. The
components of this community are not shade tolerant and the oceanside
shrub community seems to be expanding to its detriment.

Two of the remaining six communities appear basically natural
while the other four are manmade. ~ The low dune grass community resembles
the moderate drifting dune community of Brown (1959). The harsh
conditions of these exposed, unstable sand dunes result in either a
sparse plant community or no vegetation. There was no vegetation on
about 27 acres at Duck (Table 13). Eleven different species were
included in the samples of the community; however, the biomass was
only 15.2 grams per square meter (Table 7). The annual nature of
many of the low dune dominants suggests great variability in its extent
and productivity from year to year.

As indicated by their names the planted American beachgrass and
the planted bitter panicum communities were artificially established
on the most heavily bombed part of the study area. Decontamination
and extensive preparation prior to planting greatly modified the area's
topography (personal communication, D. Woodard, 1974).

Though A. brevtligulata has the highest biomass in the planted
American beachgrass community, 7. purpurea is the more widely distributed
species (Table 5). The composition of this community, as shown by the
ordination, is not very unique. In fact, the planted American
beachgrass, planted bitter panicum, sandgrass-buttonweed, and low dune
grass communities are all phytosociologically similar. The major

75

differences between these communities relate to their physiognomy,
which is a reflection of the biomass of one or two key dominants in
each community, not the presence of any unique plant assemblages.

The planted bitter panicum community had one of the most depauperate
floras of any of the community types and its biomass (5.6 grams per
Square meter) was the lowest measured in this study (Table 9). This
community appears to have been a generally unsuccessful planting
experiment.

The sandgrass-buttonweed and the spurge-sandgrass communities differ
in the relatively high biomass of the former (68.7 grams per square
meter) and the low biomass of the latter (11.8 grams). The species
composition of the two communities agrees for six out of the seven
species sampled. The major differences between them however was quan-
titative (Tables 6 and 12). Euphorbia polygonitfolta (frequency 75
percent) was the most frequent species in the spurge-sandgrass community
while it ranked next to last (6.7 percent) in the sandgrass-buttonweed
community. Trtplasts purpurea and D. teres were, however, well dis-
tributed in both types. Whether these two types are separable into two
distinct communities is open to argument. Woodard (1974) has indicated
that the sandgrass community was experimentally fertilized. This could
be the reason for the measured biomass differences. All six communities
repeatedly occur near each other on all axes of the ordination, and two
probably belong to a continuation of types which show local variation
due to environmental or historical variability.

It is felt, however, that for an intensive study on a relatively
small area such as the Duck Field Research Facility it is better to
draw out differences which might seem minor than to lose detail in
overgeneralization. For this reason, each of the 11 defined communities
has been kept separate.

The results of these studies have raised many interesting questions.
Our observations suggest that the successional theories of Wells (1928
and 1938) be closely examined and perhaps reformulated. Questions have
also been raised as to the future development of each of the delimited
communities. Which species are likely to increase in importance, and
which will decline? Especially interesting is the future of the
planted and the sound—side disturbed communities.

76

LITERATURE CITED

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7

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78

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79

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80

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qnoysnory, spew suoTRIeT[OD ITISTIOTYA “peqyTWTTep 2v1eM sat} TunmoD
queTd queteszIp usaeTqA “G/6L euNG Y8no07yI 4/6, YOITeW WOAF peqonpuod
sem *eutToie9 yqIoN ‘yong Ae TejuUeD YyOIeesSay BuTAseuTsuq TeIseoDg Aw1y
“S*n aya jo AQTTPOe™ yOIeaesey pTety yong aya Jo Apnqjs saTIeIa3eA VY
“08-22 *d : AydeazotTqt¢
(9-92 ‘ou $ TaqUeD
yoreesoy Buyiseupz3ugq [TeqJseop — Jioder snosueT[TeosTtW) “TTT : *d 0g
“9L6L
*jaqUueD yo1easey BuTrseuTsuq Tejseon : “eA SATOATOG J10q — “euTTOIeD
yjza0n Syong SAVET FORA yOTeesey pTeta yond ey; ye Apnqzs saTIeIaBaA
“d prezep ‘hna7T

aw 1ggn* L729 OyL SOEs aug con” €070L

"6L00-0-"1-ZLMOVG J0BI}U0D “1eqUeD YOTeESsdy BuTAsvoUTSUq
TeqseoD *S'N “III °9-92 “ou jao0de1 snosueTTelsTW “leqUeD YyoIPeseYy
Sup~isoutsug [TPISeOD “S*n : SeTAaS “II ‘“eTITL “I “sesseiy *h
*uotjzeqJe3eA *E *eUTTOIeD YIION ‘HONG *Z ‘“*sSeuNp pues “|
*soTTTwey g¢ Supqueseidei etsued 7E|
u}z setoeds g/| ATeqewtxoidde jo e1oTy e peTeaaet potied Apnqjs ay
Jnoysnoryy epew suOoTRIETTOO DTAISTAOTYA “*paaWTTep e1eM set Tunumos
queTd quszesJTp ueAeTyY *S/6, euNL Yy8noryI 4/6, YOIeW worz pojonpuoo
SPA feUTTOIeD YIAON ‘fyOnG 7e AejqUeD YOIeeSey BuTAseuTZuq TeIseoD Aw1y
*s*n 949 Jo ATT TOeY yOTeeSey PTeTA yond ey Jo Apnqs saTIeRa30A VY
“08-22 °‘d : Aydeas0tTqQTg
(9-9f "Ou $ raqUaD
yoiresey SupyiseuTsug ~Teqseog — yiodex snosueTTa9sTW) “TTF : *d og
“9L6L
*‘JequeD yoreesey BuTiseuTsugq Teqyseop : “eA SATOATeY WAOY — *euTToOIeD
yqaon SyONG SAITTPOeA yoreesey plTety yonq aya ze Apnis saTIeIesa,Q
‘a preteg ‘Anay

aul gon £79 QW, “Ou ami gon” £€0ZOL

°6L00-0-72-ZLMOVG 3981}U0D *Aa}UaD) YoITeeSey BufArveuT3uq
TeyseoD *S°N “III "9-9/7 ‘ou 310deX snosueTTeDsSHI *1taqUeD YyoreeSey
SuTiooutTzug TeIseoD °S*N : SeTIeS “II “eTITL “I “sesseiy “yh
*uoTqzejaZaA *E€ ‘“‘eUTTOIeD YFION ‘yONG *Z “*seUNp pues *f
“soT[TFwey gc 3upjueseider eirsued 7]
uf setoeds g/j ATezeutxoidde jo e1ioTy e patTesaez potied Apnqs ayy
qnoysnory spew suOTIDeTTOO ITISTAOTY *payrwtpfTep stem seyzzfunuuwos
quetd quertesstp uaaeTq °G/6, euNr YysnorYI 4/6, YoAeW wory paejonpuosd
sem SeuTpToie9 YyqION SyOng Je Iaej}UeD YOTeasey ZuTisvsuT3uq [Te yseog Aw1ay
"s°n ayy Fo AATTTOe_ yoTeasey pTetA yong. ey. Fo Apnys aazqeja30a V
“08-22 *d : Aydeasottqrtg
(9-9£ *ou £ 19qUeD
yoreasey ZuTiseutsuq [TeqIseoD — JAodeaA snosueTTeosTW) “TTT : *d og
“9L6L
*jajue) Yyo1essoy BuTAveuTZuq TeqseoD : “eA SATOATeG JAOG — *BUTTOIeD
yjga0n S3onq SAQTT TOR, yOAeasay PTeTA yong 9yQ ye Apnqs sat IeWe3a,A
‘aT prerteg ‘Aaay

aw gon” L09 Oceye Fis are gcn” €0720L

*6100-0-"2-ZZMOVG 2981}U0D *1a}UeD YOARaSey BuTrseUuT3ug
TeIseoD "S'N “III ‘*9-9/ ‘Ou jtodeI snosueTTeDSTW *AaqueD YyOTResay
suTiseuTsugq TeqyseoD *S*N : SeTteS “II “eTITL “I “Sesseizy “yh
suoTqeqeseq *¢ ‘“‘eUTTOIeD YyIAON SYOnG *Z “SauUNp pues “1
*soT[TTwey gc Sutqueseidei e1sues 7E|
uf~ sotoeds gy, ATeqewtxoidde Jo e10Tj e pateaAeir potized Apnqjs 9y3
qnoysnoly} speU SUOTIIETTOO ITASTAOCTY “*paqtwTTep e1eM set Tunumos
queTd JueiresjiIp ueaeTy “°G/Z6| eunr yS8nory 4/6, YAW wory pezOnpuod
seta SeutpTore) yyIoN Syong 3e JajUueD Yyoieesey BuTrveuTSsuq TeqJseoDg Awiy
“Ss°n aud Jo ARTTTOeY YOARPasey PTeTA yong ey} Jo Apnqs eat IeIesaaA Vy
“og-22 *d : Aydeassottqta
(9-9L ‘ou $ 1aqUaD
yoreosey ButisveuTsuq Te yseog — J1oder snosueTTeosT) “TTF : °*d 0g
“9L6L
*zaque) yoreeseay BuTaveuTsugq TeqyseoD : “eA SATOATAgG JA0q — *eUTTOIeD
yqIon Syong SAQTTTOe, yOTeeSey PTeTA YON eyW qe Apnqs aaT IeIa8EA
“gd preaze9 *Aaay

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