The Ecology of the New England High Salt Marshes: A Community

FWS/OBS-81/55 Plarch 1982

THE ECOLOGY OF NEH ENGLAND HIGH SALT MARSHES:

A Corrnrunity Profi l e

Scott W . Mixon Graduate School of Oceanography

University of Rhode Is land Kd ngs ton, Rhode Island 02881

Project Officers

W i ley M. Kitchens Edward C . Pendleton

National Coastal Ecosys tews Team U.S. Fish and Mildlife Service

11310 Gause Boulevard S l i d e l l , Louisiana 70458

Prepared fo r National Coastal Ecosys tens Team

Off ice of Biological Services Fish and Wildlife Service

U.S. Department of t h e I n t e r i o r Washington, D.C. 20240

DISCLAIMER

The f i n d i n g s i n this report are n o t t o he construed as an o f f i c i a l U.S. Fish and K i l d l i f e S e r v i c e position unless so designated by other authorized documents.

L.ibrary o f Congress Number 82-600533

Nixen, S.W. 1982, The ecology o f New England high s a l t marshes: a conmunity profile. U.S. Fish and Wildlife Service, Of f i ce o f Biological Services, h a s h i n g t s n , D,C. FbJS/OES-81/55, 70 pp.

P R E F A C E

In his c lassic description of the New England shore1 ine, Douglas Johnson (1925) recognized the coastal s a l t marshes found from Maine through Long Island as a type d i s t inc t from those t o the north ("Fundy Type"') or south ("Coastal Plain Type"). His distinction i s s t i l l considered useful, and I have t r ied to confine th i s community prof i l e t o observations, measurements, and experiments which have been made in New England Type marshes. A1 t hough i t i s widely recognized tha t New Eng- land marshes are characterized or distinguished by a higher organic content of the marsh peats, to my knowledge no one has yet shown rationale for not extrapolating many of the concepts gleaned from the much more extensively researched Coastal Plain marshes to this area.

The focus of th is profile i s primarily on the h i g h marsh i n New Eng- land rather than the low, creekbank, or regularly flooded areas which have received most of the attention i n the ecological l i t e ra ture . All of the marsh i s in te r t ida l , and i t must be understood and managed as a geomorpho- logical and ecological unit. I hope i t will be useful to those working in coastal planning, management, and research to bring together much of the information tha t has been developed on th is less frequently discussed, b u t important area of the marsh.

emphasis i s on the Spartina patens- Distichlis spicata community, b u t i n several cases I have included information from the stunted S. a l te rn i f lora zone. The developmentof marshes and the zonation of different species, especially plant, receive more attention in th is profile than do animal populations or community metabolism. This largely re f lec ts the re la t ive abundance of information rather than

own biases. I can only hope tha t the gaps which are so evident in th i s profile might stimulate future work in these areas.

Sa l t marshes of the New England Type comprise less than 2 % of the marsh area along the Atlantic coast of the United States (Reimold 19771, and the high marsh may amount to only 25% to 50% of that 2%. The r a t i o of people to wetland, however, i s the highest i n the country (Gosselink and Baumann 1980), and there i s a long t radi t ion in New England of using and valuing the marshes. I hope th i s profile will contribute to that t radi t ion,

S.W. Nixon Kingston, Rhode Island June 1980

Any questions or comments about, or requests for th is publication should be directed to:

While the h i g h marsh i s commonly Information Transfer Specialist thought s f as Tying betweeri Gean high Nationaf Coastal Ecosystems Team water and spring high water, the pro- U.S. Fish and Wildlife Service f i l e drawn here has not always fol- NASA-Slide11 Computer Complex lowed such s t r i c t , and somewhat arbi- 1010 Gause Boulevard t rary, 1 irnits. Similarly, the major Sl i det 1, Louisiana 70458

i i i

Number

FIGURES (Continued)

8 Typ ica l curves o f t h e p red i c ted d a i l y t i d e d u r i n g December 1970 f o r s t a t i o n s on the A t l a n t i c , G u l f o f Mexico, and P a c i f i c coasts o f the Un i ted S ta tes . . . . . . . . . . . . . . 13

9 Height o f mean s p r i n g t i d e s o f f s h o r e a long t h e A t l a n t i c and G u l f o f Mexico coasts o f t he Un i ted S ta tes . . . . . . . . 14

10 R e l a t i o n o f t h e t i d e range a t Boston, Masschusetts, t o t h e phases o f the moon over an annual c y c l e . . . . . . . . . . 14

11 Cuniulative d i s t r i b u t i o n o f g r a i n s i z e i n the sediments from the mouth t o t h e head o f Barnstab le Harbor and m a r s h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

12 E l e v a t i o n i n t en ths o f f e e t o f the h i gh marsh su r f ace r e l a t i v e t o mean h igh water a t Barnstable, Cape Cod . . . . . . 17

13 Dark areas represen t the d i s t r i b u t i o n o f pond holes o r pannes and t i d a l creeks on t he h igh marsh a t Barnstab le , Cape Cod . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

14 Surface m i c r o r e l i e f across t he t r a n s i t i o n from low t o h i g h marsh a t Farm Creek, near New Haven, Connect icut . . . . . . . 20

15 Some common h igher p l a n t s o f t h e New England marshes . . . . . 22

16 R e l a t i v e d i v e r s i t y , dominance, and major species composit ion o f vege ta t ion zones descr ibed by M i 11 e r and Eg le r (1950) a t the Wequetequock-Pawcatuck marshes i n C o n n e c t i c u t . . . . . . . . . . . . . . . . . . . . . . . . 2 5

17 General ized t r a n s e c t from the uplands t o the low i n t e r - t i d a l i n a " t y p i c a l " New England s a l t marsh showing t he common vege ta t ion types . . . . . . . . . . . . . . . . . . . . 2 7

18 V e r t i c a l d i s t r i b u t i o n o f r oo t s , rhizomes, and dead ma t te r on the h i gh marsh a t Great S ippewisse t t Marsh on Cape Cad . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . 39

19 Amounts o f aboveground vege ta t i on and r o o t s f rom A p r i l through November on the h i gh marsh a t Great S ippen i sse t t . . . . . . . . . . . . . . . . . . . . . . . M a r s h o n C a p e C o d 40

20 The accumulat ion o f o rgan ic carbon, t o t a l n i t rogen , and i no rgan i c phosphorus i n t h e sediments o f a marsh c a l c u l a t e d f o r d i f f e r e n t a c c r e t i o n ra tes , sedirrient dens i t i es , and sediment composit ions . . . . . . . . . . . . . . . . . . . . . 43

v i

FIGURES (Continued)

Page

21 Decomposition of various kinds of p l a n t material on the ground or submerged in water a t different s i t e s . . . . . . . . 46

22 Locations of New England towns tha t were sett led by 1650 adjacent to fresh or s a l t hay marshes . . . . . . . . . . . . . 49

23 Top: Salt hay on staddles t o keep i t above the tide. Bottom: Gundalow loaded with s a l t hay to be floated out on the flood tide . . . . . . . . . . . . . . . . . . . . . . . 5 1

24 A m o u n t of coastal wetlands in the Northeastern United . . . . . . . . . . . . . . . . . . . . . . . . . . . . States 52

25 Amount of time the grasses and surface sediments a t Farm Creek, Connecticut, are exposed t o the atniosphere a t different elevations across the rnarsh . . . . . . . . . . . . . 5 4

26 Concentrations of copper a t various depths in the sediment under Spartina - patens a t Farm Creek Marsh, . . . . . . . . . . . . . . . . . . . . . . . . . . Connecticut 56

27 Concentrations of manganese a t various depths in the sediment under the Spartina patens a t Farm Creek Marsh, Connecticut . . . . . . . . . . . . . . . . . . . . . . . . . . 56

28 ttistorical variation in the anthropogenic fluxes of copper, zinc, and lead recorded in the h i g h marsh sediments a t Farm Creek, Connecticut . . . . . . . . . . . . . . . . . . . . . . 5 7

29 Aboveground biomass of 3 a r t i u a a l te rn i f lora in experimental plots treated with soluble iron, copper, and chromium a t Great . . . . . . . . . . . . . . . . . Sippewissett Marsh, Cape Cod 59

TABLES

Number

1 Rates of sea level r i s e re la t ive t o the land i n the Northeastern United S t a t e s . . . . . . . . . . . . . . . . . . 3

2 Estimates of accretion rates i n s a i t marshes along the Northeastern United States . . . . . . . . . . . . . . . . . . 4

TABLES (Continued )

Number

.3 Annual marsh flooding a t various elevations, volume of water, and period of submergence . . . . . . . . . . . . . . . 18

4 Relative amount (%) of coverage of high and low marsh in various New England s a l t marshes . . . . . . . . . . . . . . . 2 6

5 The r a t i o of high marsh t o low marsh along the Atlantic c o a s t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

6 Principal epibenthic algal species on the Cape Cod marshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

7 Dominant macroinvertebrates in different vegetation zones on a high marsh near Stonington, Connecticut . . . . . . . . . 3 1

8 Average number of Aedes mosquito larvae per dip of water on the marshes of Egg Island, New Jersey . . . . . . . . . . . 3 2

9 Birds on the high niarsh a t Cape Ann, Fllassachusetts . . . . . . 3 3

10 Estimates o f aboveground primary production of vascular plants on New England high marshes . . . . . . . . . . . . . . 3 7

13 E f f e c t af nitrogen additions on the production of high marsh and low marsh vegetation a t Great Sippewissett Marsh,CapeCod. . . . . . . . . . . . . . . . . . . . . . . . 3 S

12 Comparison of sediments found on the high marsh a t Farm Creek, Connecticut, with those of Long Island Sound and a short S. - al ternif lora ~ a r s h a t Barnstable, ~ a s s a c h u s e t t i . . . . . . . . . . . . . . . . . . . . . . . . . 42

13 Metal added to each plot and amount of each element found in the top 2 cm o f marsh sediments . . . . . . . . . . . . . . 5 8

14 Metal concentrations of l ive Spartina a l t e r n i f l o ~ and . . . . . . . . . * . . . . . . . . . . . . * . . - 0 0

ACKNOWLEDGMENTS

I am grateful to a number o f a n d Ralph Tiner. Their help eliminated reviewers who read and critica'zed errors and obscure passages. Far ear l ie r drafts of th i s prof i le , those that remain, I arr: responsible. including Virginia Lee, bJi ley Germai'ne kebb typed and retyped the Kitchens, Ed Pendleton, John Gallager, manuscript with her usual s k i l l , Frank Da-iber, Joseph Ustach, Rick patience, and good humor, and I thank Linthurst, Curt Laffin, Ralph Andrews, her for i t .

Racks i n the high marsh a t Wequetequock-Pawcatuck, Connect icut , where Miller and Eg le r (1950) c a r r i e d out t h e i r benchmark study of zona t i on an a New England r37t niarsh during the l a t e 1940's.

CHAPTER 1

ORIGIN AND DEVELOPMENT OF THE KARSHES

I t may seem odd to begin with an f o r ~ a l sc ien t i f ic inquiry a t least 125 observation about rocks, b u t they were years ago. the f i r s t things I noticed when I went o u t across a New England s a l t n-arsh. They seenled out of place; marshes were EVOLVING CONCEPTS OF MARSH DEVELOPPIEMT sedimentary environments, not high energy places l ike sea c l i f f s or Subsidence and Sea Level Rise--the cobble beaches. B u t my images had Mudge Model been formed in Coastal Plain marshes to the south, where larger supplies of In 1857, B.F. Rudge (1862) s e d i ~ e n t from the rivers developed presented a paper to the Essex, mineral so i l s around the grasses and Connecticut, Ins t i tu te in which he helped to build the marshes outward described his findings from a core across accreting shoals. The northern taken i n the Romney Marsh, near Lynn, marshes f i l l ed with peat were Eassachusetts, a t a s i t e "about one different , and they made some f o o t above ordinary h i g h water ark processes that were operating a l l and only overflowed by the higher. along the coast more conspicuous. spring tides. " The remarkable feature There had been other oddities, 1 ike of th i s core was that i t showed the t ree stumps, in many marshes I had roots and rhizomes of the marsh grass seen, b u t for some reason the rocks extending d o ~ n unifortr;ly t o a deptn caught my attention more forcefully. well below normal low t ide. Because They were a dissonant note--cold, the grasses grew only above the noruial inorganic, gray, and unmoving amidst high water level, Mudge concluded that a l l tk,at green and hindblown grass. the marsh had been subsiding and tha t I t was a useful lesson to see them i n the subsidence had been counter- the middle of a Spartina meadow, and a balanced by a upward accretion from reminder of the complexity of marsh grass growth and subsequent sediment devel opment . deposi tion, The process responsible

for the subsidence o f the ~ a r s h was Since the present-day marshes are n o t known a t that t i ~ e , and Mudge

s t i l l responding to the forces that speculated that i t might be due to produced them, i t i s of more than erosion beneath the marsh caused by a historical or academic interest to "current of' water i n the d i ' l uv iu~ investigate marsh develop~ent in some under the clay. '' deta i l , particularly with respect t o the Northeastern United States , The As more cores were examined from story o f haw the rocks cane to be Rany marshes, however, i t became clear there, or rather of how t h e ixarsh that Eudge k sfindings were too grass came t o grow around the rocks, common Is be explained by such a i s an 3nteresting one that began as a local phenomenon (Johnson 1925). The

1

growing acceptance of glacial theory and sea level r i s e soon provided a more sat isfactory general explanation for the thick deposits of marsh peat !Davis 1910). The development of 41: dating techniques shortly a f te r

the Second World War made i t possible t o begin study of the quantitative relationship between marsh development, sea level r i se , and land subsidence since the fas t glaciation-- the Holocene Transgression.

Using radiocarbon-dated material from present-day s a l t marsh peat as we11 as from re l i c peat deposits and other materials on the continental shelves, workers have developed curves relat ing sea level to the land over the past 35,000 years. While there i s some uncertainty in the data and various versions of the curve are offered from time to time (see Emery and Uchupi 1972), the general picture

suggests a rapid f a l l i n sea level which began about 20,000 years ago and continued for same 5,00C years. T h i s rapid f a l l was followed by a rapid r i s e in sea level until about 7,OCO or 8,600 years ago when the rate of r i s e began t o s l o w appreciably (Figure 1). Virtually a l l present-day marshes found in the Northeastern Uni ted States appear to have become established only during the p a s t 3,000 t o 4,000 years (Redfield 1972; Rampino and Sanders 1980). The average ra te o f relat ive sea level r i s e during t h i s recent period o f marsh building has been about 1 m f y r i n this region (Table I ) , and i t i s commonly thought that marsh development can only take place when the r a t e of sea level r i s e i s slow. B u t r e l i c salt-marsh peat has also been found on the continental shelf ranging i n age f r o m 5,000 t o 11,QOO years; a period when the average ra te of sea level r i se may

THOUSANDS OF YEARS AGO

a coralltne algae

Figure 1. Sea level relative to the present level on the Atlantic continental shelf during the past 36,000 years (Emery and Uchupi 1972, based on Ewry 8 n d Milliman 1971).

2

-.

Table 1. Rates of sea level r i s e re la t ive t o the land in the Northeastern United Sta tes .

Long- ern ra tes f o r the past 2000 t o 30430 years estimated from age-depth curves for '&-dated material i n marshes and continental shelf sedirrents (Bloom and Stuiver 1963; Redfield 1067; Keene 1971; Oldale and O9ara 1980; Rampino and Sanders 1980)

New Hampshire 1.1 Northeastern MA,(probably a lso NH and ME) 0.8 Southeas tern FiA 1.0 Cape Cod to Virginia 1.1 Connecticut 0.9 Long Island, N Y 1.0

Short-term ra tes for the past 35 years (1948-75) from t i de gauge records (Wicks 1S78)

Eastport, ME Port land, f i E Portsmouth, NH Boston, PA Hoods Hole, MA Ncwport, RI New London, GT New York, N Y

a The pub1 ished value sf 0.02 m/100 y r i s a typographical er ror in Oldale and B'Hara 1980 (Charles O 'Hara, personal communication),

have been 16 mn/yr (Energy and Uchupi those periods were times of roarrh 1972). dcvelop~ent .

A t l e a s t two possible explana- Analyses o f t i d e gauge records tions ex i s t for thSs apparent discre- (Nicks 1978) have shown tha t the pancy. The f i r s t is t h a t marsh re la t ive r i s e of sea level along the cievel opment: is possible d u r i n g No~thcastern Un'i t e d S ta tes during the times s f more rapid sea level r i s e past 35 years has been two to three t h a n has been experienced, on the times the recent long-term average average, during the past several (Table 1)- Studies of salt-marsh thousand years, The second i s accretion ra tes i n this area have tha t there have been times when sea shown t h a t the marshes a r e capable level r i s e was much slower than i t of "keeping up" w i t h t h i s ra te o f has been on the average, and t ha t r i s e (Table 21, and Redfield (1972)

3

Table 2 . Estimates of accretion rates i n s a l t marshes along the Northeastern U n i t e d States

---

Accretion Location Vegetation type J ~ t : l I y r )

Barnstable Marsh, Cape Cod Spartina -- a1 ternif lora "Young marsh" 118-3 p i ~ ~

"Older marsh" 1.5-2.7

b Barn Island, CT Great Island, CT Hammock River, GT

0 Stony Creek, CT Nells Island, CT Farm River, CTC Flax ~ o n d , ~ ~ o n ~ Island,

N Y ~

S . patens - S. patens 3, - patens-Distich1 is g i c a t a Phragmi tes communis S. patens-dwarf S. a l ternif lora -" P - -- S. al ternif lora + 2, p z t e ~ ~ 3. &ens (n;ean for top l T c r ~ ) - S . a l t e m i f ~ o r a

" ~ e d f ield (19 72). b ~ a r r i s o n and Bloom (1977). " ~ e ~ a f f r e ~ (1977). d ~ l e s s a e t a l . (1977).

found that a young, actively growing portion of the marsh a t Barnstable, an Cape Cod, Kassachusetts, increases in ctevation a t a ra te exceeding 50mrr)lyr. A detailed comparison of accretion rates and sea level r i s e aver time was carried o u t by KcCaffrey (1977) on the high marsh a t Farm Greek, near New Haven, Connecticut, by using a pk2nda ted core. The results showed t h a t sea level r i s e was clasely matched by marsh accre- t ion, and that accretion continued even during short periods of relat ive sea level f a l l (Figure 2 ) . Nswever, the more rapid recent rates o f sea level r i s e along the northeast coast are s t i l l considerably slower than the average 16 mm/yr that may have occurred during ear l ie r sarsh develop~ent.

Along the Louisiana coast, where recent subsidence rates have been about 1 2 mmlyr, e x t ~ n s i ve n~easurenlents by Baun~ann (1980) have shown that streamside marshes have had sedimentation rates of 15 n;m/yr, b u t that more inland rnlarsh areas have had rates of only 9 mm/yr. As a resu l t , there h a s been a substantial loss of wetland. I t i s hard to know i f this suggests a natural upper l i m i t of about 10 t o 12 mmpgir beyond which marshes cannot, on the average, keep pace. The more correct cenclusisn may be that , given an adequate sediment supply, the marsh grasses themselves are capable o f dealing w i t h rapid rates of sea f e v ~ 3 r i se , I do not know haw the past sediment supply on the northeast coast compares with the present day supply a long the Gulf of Mexico; t ida l

AGE , Y

0 40 80 120 160 200 r

Smoothed Tide Goge Record *o-.-+.

Salt Morrh F'b2'' Record - . - . .5 .. * .* - 7 . - _ . a .

3. s.: *. -

- - - - 3 ' : ' " " : " " : ' " ' 4

4% 6' Q0 9

e0 -?

oQ fJ 9 2? \ A?

Y E A R

Figure 2. Variation i n apparent sea level a t tdcw York City as shown by a smoothed t ide gauge record and the elevation of a x a r t i n a atens marsh in New --h the sediment Haven, Connecticut, calculated from the distribution of Pb (McCaf frey 1977).

regimes are very different in the two areas. Harrison and Bloon: (1977) found a positive correlation between t idal range and accretion rates i n Connecticut marshes ; Bau~ann (1980) found greatest sedimentation in Louisiana marshes during winter, when the wetlands were innundated for less time than tbieg/ were during summer.

The secand explanation for past mars h-bui ldi ng has been developed recently by Rampino and Sanders (1980), who suggested that marsh development i n the Northeastern United States has been episodic during the l a s t 10,000 years, taking place only when relat jve sea level remained constant for a time or went th rough a transient lowering i n response to s horter-term cl fmatic events. Sea level has risen over the past 15,000 years w i t h different average rates of

increase for different time increments (Figure 1). These data and other information have also been interpreted as showing osci l la t ions i n sea level, and the available i n f o r ~ a t i s n i s apparently not detai 1 ed enough now t o resol e the question. On the basis of l!C-dated peat samples from the inner continental she l f , Rampino and Sanders (1980) concluded that there were six previous periods of marsh growth about 1,006 years apart, the most recent of which began some 4,780 years ago.

Changes i n r e l a t ive sea level (see Figure 1 and Table 1) are thought t o ref lect two components : i sas ta t ic processes t h a t r a i s e or lower the land surface and e u s t a t i c processes due to changes i n the vaiun;e of the ocean from glacier formati on and me1 t ing . The relat ive con t r ibu t ion of these

components to the observed sea level THOUSANDS OF YEARS AGO change varies from place to place, h u t 5 4 3 2 1 Q

Redfield (1S6T) and Emer and hichupi (1972) were able to use 1i~-dated peat from s a l t marshes along the Atlantic and Gulf of Eexico coasts t o arrive a t an estimate of about 0.8 mnl/yr for the eustatic r i se in sea level during the past 4,OCO years (Figure 3 ) . Thus, most of the change in sea level observed in the Northeastern United States during the l a s t 2,OCO t o 3,000 years (Table 11) appears t o be due t o a n absolute increase in the level of the sea rather than t o land subsidence. 5

dati on and-Accretion--the Shaler V) OC W

Abaut 30 years a f te r Rudge i- offered his explanation for the t h i ck 2 salt-marsh peat accumulation he observed, M . S . Shaler (1886) developed a rf~odel for marsh forniation based on a different set of observations. Shaler eniphasired the gradual accuniulat i on of sediments in shallow coastal waters, particularly where seagrasses might accef erate the depori Lionel process. As the water Lrecarrre shallower, the seagrass beds would be replaced by mud f l a t s which would, in turn, be colonized by a & r & % ~ a r z j z , the only yrass t o survive in the low inter t idal zone. The presence of the -5 grass would further enhance sediment depositt'on, and the roots and rhizomes Figure 3 , Age and depth of salt-marsh would contribute t o peat forrcation, peat in different areas: A (Cape Hat- This process would continue until the teras t o Flexico plus Bermuda), B (Cape sediment accuntu"ftte dlnn!ost to the Cod "c Cape Hatteras), C (eastern lims"t of the h i g h t i de . By "cis Flassachusetts), D (Bay of Fundy). I n precess, the rrtarsh would build u p and area A , the l a s t 3,500 years are out from the share as sedirrrents were assumed to ref lect the eustat ic r i s e redistsi buted a long the coast, in sea level, so that the deviation of

each of the other curves fran; A i s due I n his classt'c descriptl'on arrd Lo local land subsidence. For example,

analysis o f the hew England coast1 ine, in area C focal subsidence continued Biahrrson (1923) discussed the problem uti t- i f about 2,520 years ago rats ra te of s a l t marsh formation in some of 0 - 3 mm/yr.. When corrtbined w i t h the detail, and established "cr i ter ia eustatic r i s e o f 0.8 mm/yr, the resu l t far testing the 8"tucfg end Shaler i s the 1.1 n i ~ j y r of rela"civc r i se fheori es . '"ccordi ng to !<udge, s hawrt (Regif ?el d 1961, as n:odi f i ed by sections through the marsh should show Emery and Uchupi 1 9 1 2 ) ,

6

L U 3 c=.

%+-' I--LS ' L c t l C r : m a w cu w 7 Tr U

h r n c n E * ~ r r o m 'r O L U

y_ 24 *.r 't- .r-

" E % , = ; QJ Q J Q J - ,n a) *EL'- o = v) _cC, L O +3 4-a+'

I n Q E c 13 . -6:

patens had becoirie es tab1 i shed, the high ~ a r s h peat kept accreting i n response to rising sea level. T h u s , a thick section of S. @tens - peat lay over a thin layer -of S . al ternif lora peat which was underlaif; by a mud f l a t deposit. I t was not until A . C . Redfield (1972) carr ied out his extensive studies o f the Barnstable Marsh in Cape Cod that a well- documented and comprehensive picture of marsh development on the northeast coast emerged.

Data collected from the Barns- table marsh confirmed the model proposed ear l ie r by Redfield (1965; Redfield and Rubin 1962) in which the sequence of events described by Shaler was placed i n the context of a r is ing sea level. I t was c l ea r ly shown tha t the different views of Mudge and Shaler arose, a t l e a s t i n par t , because Mudge had Focused on the upland side of the marsh while Shaler had been looking primarily a t the seaward s ide . W i t h a rising sea level and a s u f f i c i e n t sediment supply, Redfield (1972) found tha t

the inter t idal S. a l t e r n i f l ~ peat extended progressively out from shore and a t an upward slope over an aggrading sand and mud deposit. The h i g h marsh peat then formed over the inter t idal peat as a wedge which thinned as i t expanded toward the upland and toward the seaward edge of the marsh (Figure 5). Cores taken in areas where the marsh had overgrown the upland in response to r is ing sea level would contain only a uniform deposit of high marsh peat, as reported by Mudge, while cores froni the outer portions of the marsh would appear as described by Schaler. Despite a l l his care and ef for t s , the many cross sections examined by Johnson (19251, and which confirmed only Fludge's views, happened to come from areas i n the marshes that had formed over old upland s i t e s or from marshes i n which erosion had removed areas of recent marsh accretion.

The marshes are growing out and over the sand and mud f l a t s as we11 as up and around the rocks.

Figure 5. Redfield ' S wx!et fa r aa1 t-marsh cfevelop~ent over acctarrulating sediment on a sand f l a t and over the upland under the influence of r is ing sea level (Redfield 1972). refers to miln high water a t various times during devel op~ent . 8

CHAPTER 2

RATER LEVELS, SEDIMENT DEPOSITION, AND THE FORM OF THE MARSH

FLUCTUATIONS IN klATER LEVEL Short-term Changes in Mean Sea Level

While the long-term secular r i s e i n sea level due t o glacial melting and land subsidence has played an i ~ ~ p o r t a n t role i n salt-marsh development, other processes influence water level s along the coast. Although people often think of sea level as a fixed da tum, continuous water level records (such as those obtained from tide gauges) have shown that sea level varies on vir tual ly every time scale. There are wind- generated waves which may have periods of seconds or ~ i n u t e s , as well as semidiurnal or diurnal t idal Maves. The passage of atmospheric fronts w i t h d i fferent barometric pressures and wind f ie lds influences coastal water levels for hours or days. Seasonal and yearly changes i n temperature, s a l in i ty , and barometric pressure influence the density and volume o f coastal waters, making them r i s e and f a l l re lat ive to the land. And changes in coastal geomorphology tha t may take place relat ively rapidly (dredging, breachway opening) or aver a nuvber of years (develop~ent of a barrier spi t ) , may influence water levels and t idal ranges. The name "t idal marsh" re f lec ts the widely recognized importance of th i s component of water level changes, and i t will receive particular attention in the next sectiorr. F i rs t , i t s"s worthwhile t o consider some other processes that influence water levels i n the marsh on time scales longer than a t idal cycle O u t considerably shorter than the melting sf glaciers.

Sea level i s usually calculated as the arithmetic mean of hourly water level measurements collected over the period of in te res t . Usually, water level data are taken from t ide gauges tha t are designed to f i l t e r out short period changes due t o waves. The elevation of the gauge i t s e l f i s usually leveled to U.S. Geological Survey bench marks on land which can, in turn, be related to the zero elevation of the National Geodetic Vertical Datum (Wicks 1978).

Examinations of water level data reveal a bewildering array of nontidal changes, some of which are irregular while others appear to be cyclical. I t i s we1 1 known that there i s a r i s e in sea level associated w i t h storms which may reach 3 to 5 m (10 to 16 f t ) above normal t ide in the extreme case of a hurricane. More commonly, the effect of winter storms along the northeast coast will increase water levels by less than a meter. This increase i s due t o a short-term "surge" o f water moving into the area because of the low barometric pressure associated with the passage of the s ta r% front (the '7nverse barometer effect ," Smith 1979) and t o a longer-tern? accumulation o f water against the coast due t o wind s t ress . Killer (1358) studied these two processes on the New England coast; and found that there was a time lag of I. t o 14 hr w i t h an average o f 5 to 6 hr , between fiaximurn wind and maximum ""st-cnp'bar

Ti 0 2 m c , r Q, LC, CT-f- 0 L T r o X Q - 7 -'a& C Q , 'A-

m t - - a + e= I,L G 3 a 2 a o - m m o 0 > L E i-- .f-

er a a s c, S - r - a Q J c,ua ", *z < -7 2 .z

m u L Q , - m Q, a =m,> Wv, L L Q , c (U @ m m Z W p f F-,aJc,L2

C m " m o o 'P"- ,wwcc V)

Y- O m $ ( U g

' A * ZL' 'A 'A % E 3.: m

Q, 0)- L V -a m z s V)= Q, 3c., -7 ='= 4 ' m . c ) t - u c s L O

a o r - g~gKLur( m aJi--<z 3

EOJZZ E U * E 5 ,

Q , a J r s;* ' A C , d t - l L

c, cb 0 Z

La,==' a J w o a c , ~ Q,Y VI S C1.r C T K 3 . r ' A a 3 O c J 4 ' L

Figure 6. Departure from the annual mean of average monthly sea level, water densi ty, temperature, and sal ini ty a t three primary t ide s tat ions (Emery and Uchupi 1972) . Departures fron the monthly mean for nearby river discharge (bot tom l ine ) are given for comparison,

I n a detailed study of sedimentation on a Louisiana marsh, Baurnann (1980) found that sediment deposition was not correlated with Rean sea level or duration of submergence, b u t with the concentration of sediment in the flooding water. I t i s l ikely that many mars h-water interactions a re variable and complicated, and that the strength of any particular coupling may not be a simple function of the duration o f submergence.

Tides - ----- In contrast t o water level

changes discussed earl ie r , variations caused by the tides are remarkably regular and their influences are tbmought to be "the most significant

11

environmental factors responsible for the segregation of sa7 t-marsh vegetation" (Redf ield 1972). Tides along the Atlantic coast of the United States are semidiurnal and symmetrical w i t h a period of 12 hr 25 pin, i n marked contrast t o those of the G u l f of biexico and Pacific coasts (Fl'gure 8) . The tides along the northeast and much of the southeast coast are also o f considerably greater range (1 l o 3 m or 3.3 "i 10 f t ) than those along the Gulf o f Mexico coast (Figure 9) . There i s a substantial and regular variazion i n the t i d a l range, not only w i t h the lunar cycle as shown i n Figure 8, b u t over the annual cycle as we7 1. Rhile the t idal range -0s greatest d u r i n g f u l l and new moon, the h ighes t and 1 owest tides occur nearest

the summer and winter so ls t ices (Figure 10). As Emery and Uchupi (1972) pointed out, there i s also a "fortunate circumstance" such that the times of lowest low tide come during the night or early morning, so that animals and plants accustomed to l i f e

BOSTON, MASS below the tide l ine are not exposed to excessive heating and desiccation

1920 1930 1940 1950 1960 $970 (Figure 10).

While the National Ocean Survey (NOS), National Oceanic and Atmos- pheric Administration, publishes annual predictions of the dai 'ly t ide

20 heights and times for many primary and 2 1

secondary locations along the coast, the t idal pattern found in a marsh may

22 often be quite different from tha t ,, observed or predicted a t the nearest reference s tat ion. Generally, the t idal signature found inside a narrow opening, behind a barrier sp i t , or u p a winding t idal creek will show a

26 reduced t idal range and a delayed time 27 of high water; flood t ide will be

shorter than ebb w i t h a fas te r Rean * current speed,

2s::

3 o The t ide heights published i n the NOS tables are given with reference t o

3 I mean sea level, an elusive datum we have spent some time discussing in ear l ie r sections. To eliminate or a t least reduce much of the short-term variabi 1 i ty i n sea level measurements, the NOS uses a 19-year average of hourly t ide gauge records for most of i t s t idal work. The choice af this averaging interval has

Figure 7 , Annual variation i n mean some astronomical significance and sea l eve1 a t Boston, Massachusetts, represents a practical choice, given cornpareef w i t h mean annual water the lengths of records available for dens i ty (inverted scale), temperature, most stations and the level of arid saf ini ty a t the t ide gauge variabi l i ty in the yearly data (Hicks s ta t ion , p l u s annual discharge of the 1968). Sea level, however, i s often Charles River including urban waste estimated by N@S from a t idal record water (En~ery and Uchupi 1972), that i s only 1 month to 1 year

long by co~parjng 1 t t o a nearby station wi k h a 19-year record and siaiilar t idal characteristics. Sima"'lar'ly, i t is relat ively easy t o develop yearly t idal predictions

12

DECEMBER 1970 2f 22 23 24 25 21 87 28 29 30 31

4

Figure 8. Typical curves of the predicted daily t ide during December 1970 for stations on the Atlantic, Gulf of Nexico, and Pacific coasts of the United States ( E ~ e r y and Uchupi 1972).

for a particular marsh area by using a measured t idal record as short as 15 days (Palmer e t a l . 1980).

Despite our long-standing ab i l i t y to predict the t ides with reasonable accuracy (except when winds, baro- ~ e t r i c pressure, or fresh water influences dominate), i t i s d i f f i cu l t to predict water level i n a s a l t marsh or other coastal ernbayment because water level resul ts from a complex s e t of interactions. !dater level, especially as affected by t ides , has traditional ly been considered the major influence is? determining mrsh ecology. Water level also has been used often for classifying wetlands in inventories

9 3

and in legal descriptions protecting or regulating various portions of the coastal environment, particularly descriptions of wetlands (Kavenagh 1980). There may be good ecological reasons, however, for doubting tha t marsh vegetation is finely tuned t o the t ides and water levels (Lagna 1975) because concepts l ike mean sea level, mean high water, and irean low water are arbi t rary simplificatisns tha t depend on the time interval chosen.. The relat ive position of the land-water-air i n t e r f a c e over a defined time interval is a physical r ea l i ty ; i t can be readily measured, and i ts importance -in the marsh has been a central theme i n coastal ecology for a t l eas t 50 years.

Figure 9. Height o f mean spring tides offshore along the Atlantic and Gulf o f Mexico coasts of: the United States (E~ery and Uchupi 1972). Contours are i n meters with extra contours f o r 0.5 and 1.5 m.

F igure 10. Relation of the t ide range a t Boston, iilassachusetts, t o the phases of the moon over an annual cycle (Enery and Uchupi 1932) . Upper envelope encloses the daily t idal range w i t h levels of mean h igh t ide, mean tide level, and Rean low t ide sftown. Lower portions show the r e l a t i o n o f t ides 'lower than -30 crs, (wide diagonal 'lines) with respect to the tirite o f sunrise and sunset.

14

THE FORM OF THE KARSH

The shape and appearance of each fias-sh resul t from unique and complex interactions of local topography and bathymelry, sea level r i s e , t ides , sedin,ent supply, and vegetation. Whi f e practical considerations force us to consider each of thcsc subjects in t u r n , i t i s their interaction that makes a narsh. For example, the s a l t - marsh plants play a vajor role in trapping and s tabi l iz ing inorganic sediments as well as in producing the organic vat ter that fornls the marsh peat. B u t marsh vegetation, espe- c i a l 7y i t s zonation and productivity, has received much study and a separate chapter will be devoted to th i s problem af te r a brief discussion of the development and form of the marsh substrate,

Harsh Development, Topography, and blorphol ogx

Piarshes usual ly devef op behind barrier sp i t s or in the mouths of t idal river estuaries where there i s sorne protection from waves. The ma,ior problem with waves appears to be that they prevent the sediments from forniing a s table substrate, rather than inechanica'l ly damage the marsh grasses (Redfield 1972) . As t idal currents carry water and sediments into these areas, they become progressively slower due to constrictions and bottom f r ic t ion . As a resu l t , their ab i l i ty to keep part ic les i n suspension decreases, so that sands become deposited near the mouth of the embayment w i t h s i l t s and clays toward the head of t idal creeks and meanders (Figure 11). Redfield ( 5 9 7 2 ) distinguished between marshes that developed on slopins foreshores, in which the distribution of sediment had been relat ively uniform and the drainage a t low t ide was reasonably complete, and those that devef oped across sdnd or mud f l a t s where:

"The pat tern of development appears to have depended on the vagaries of the sedimentary processes which bui l t up the sand f l a t s to the c r i t i c a l level above which 2. a l te rn i f lora can grow. The drainaqe pattern of the hiqh marsh has -been fixed by that of the channels which f inal ly drained the f l a t s i n the broad sounds enclosed by the developing narsh. Such channels s h i f t their position continually until stabilized by the turf of the marsh, which then fixes their f inal position."

I n the former case, the resulting marsh has a more or less uniform appearance, and the sloping surface of the landform makes i t possible for the hiah marsh to deve lo~ independently of t h e inter t idal spa r t i na a? tern%' f l r a . I f the marsh accretes and aggrades across f l a t s , however, i t s appearance i s more interest ins and i t s deve1o~- ment follows the" general patteb.n described by the Shaler model.

In sp i te of i t s appearance on casual inspection, the surface of the h i g h marsh i s not absolutely f l a t , b u t i s elevated s l ight ly toward i t s inner and ol der portions because of the longer period over which this area has been able to accumulate sediment and peat (Figure 12). Low natural levees, perhaps 5 t o 15 crr; ( 2 t o 6 inches) h i g h and several meters wide, sometimes occur along the major marsh creeks; these levees develop because a re lat ively l a r g e r amount o f sedin~ent i s deposited there when the r is ing t idal water f i r s t overflows the creek banks and slows down as i t spreads out across the marsh.

The greater elevation of the older marsh means t h a t i t will be less frequently flooded by t-ides, that i t will be? submerged for shorter periods o f lime, and that less water will need

G R A I N S I Z E F u r 1 C u ~ u i a t i v e d i s t r i b u t i o n o f p a i n s i z e i n the sediaents froi?! t he nrouth t o the head of Barnstzblt. h'arbiar and c:arsh (Redfield 1972). Coarser r::aterials drop out quickly as cur ren ts slow i ns ide the barrier s p i t ,

10

TON CREEK

ELEVATlQN IN TENTHS OF FEET RE MHW

Figure 12. Elevations in tenths of fee t o f the high marsh surface re la t ive to Rean high water (FIHK) a t Barnstable, Cape Cod (Redfield 1972). Older portions of the marsh are higher.

to be drained from i t s surface (Table 3 ) . As a resu l t , there will be progressively fewer and smal l e r drainage creeks, pond holes, and "rotten spots" (Figure 13). The reduced amount of water reaching the high marsh will also bring in less s e d i ~ c n t , so that the rate of vertical accretion w i l l decline relative to the young portions of the marsh (Johnson 1525; Redfield 1972; Harrison and Bloorr, 1377; Baumann 198C).

The pannes and pond holes or "rotten spots" shown i n Figure 13 are common featurcs of the New England marshes which have been studied i n detai l (Hiller and Egler 1950; Chapnan

17

1960; Redfield 1972). Sometimes the pannes or shallow depressions of the marsh surface may be quite large and represent areas within the marsh which, for various reasons, d i d not receive enouah sediment t o shoal w

suff ic ient ly for Spartina t o grow. Eany of the pannes contain round, shdlow holes or small pools ("primary pannes")) w i t h a depth sopewhat greater than the thickness of the Spartina turf. They are f i l l e d by the higher t ides , though some may even have small drainage systems. The lack of a peaty turf in m r e southern mrskes may explain the absence of deeper pools in those areas (Redf ield 1972)- I n other pannes the standing

Figure 13. Dark areas represent the distribution of pond holes or pannes and tidal creeks on the h i g h irarsh a t Barnstable, Cape Cod (Redfield 1972). The marsh on the l e f t an in the foreground where fewer of these features are found i s 01 der accord to l f C dating.

water may evaporate, leaving s a l t deposits that liniit the vegetation. Several processes txay be responsible for these features, including the blockage of drainage creeks by slumping of the banks, the evolution of h i g h marsh from patches or lines of s laugh marsh growing together, the decay o f surface turf because of poor drainage ("rotten spots "1, and the accun~ulation of "trash. '"he pond holes appear to be relatively stable over the short tern? because the depth of standing water i n them (0,s t o 1 m or 1.6 t a 3.2 f t ) i s uscral?y great enough to prevent the spread of Spart ina rhizomes (Redf ield 19721, but the i r disappearance from the older portions of the marsh (Fisure 13) suggests that clost of them are an

ephemeral part of the marsh (Chapman 1960).

MARSH SEDXMENTS

Ciarshes along the Atlantic coast may receive sediment from rivers, from the nearshore zone, and from re1 i c mud deposits on the continental shelf (Mead 1969; Phleger 1977). The l a t t e r sediment source i s particularly important for many of the coastal rnarshes along the Northeastern United States, where the supply of new terrigenous s e d i ~ e n t i s l o The mechanism responsible i s a landward flow o f bottom water across the shelf. In areas where marshes have formed behind barrier sp i t s , large amounts of

sand may also be carried onto the ~ a r s h by wind and storm overwash. A recent review by Frey and Basan (1978) gave a detai led description of the rxechanisms responsible for the movement and deposition of sedirnent i n marshes, and 1 have drawn comparisons between some aspects of the chemical composition of marsh sedin:ents and those o f fresh water and nearshore marine waters (Nixon 198C).

Usual ly h i g h marsh sediments consist o f a fine, s i l t - l i k e inorganic fraction and a coarse organic fraction made u p 1 argely of S ar t ina roots and rhizomes. As McCaf frey "r 1 3 7 ) pointed out, the organic content of ~ a r s h so i l s i s only s l ight ly higher than that of many estuarine and nearshore sediments, although their bulk density i s much lower. On a dry weight basis, the bulk density of Far81 Creek marsh was only C . 2 g/cm (1.01 g/cm wet), while t h a t of the adjacent Long Island Sound sediment was about 0.6 g/cm (McCaffrey 1977). Despite the use of the term "peat" i n connection with New England s a l t marshes, the organic

content i s much lower than freshwater peat bogs, and i t would be a cold home that tr ied to burn high ~ a r s h peat in the fireplace. Nevertheless, the dense growth of the Spartina patens roots and rhizomes greatly accelerates the vertical accretion of the trarsh through their own volume as we11 as through sediment trapping ; where dense tussocks of the grass develop, the vicrorelief of the marsh surface i s affected (Fi yure 1 4 ) - The continual input of new sediment onto the marsh i s c r i t i ca l not only for the system to keep u p with rising sea level, b u t nitrogen, phosphorus, and other elements associated with the sediment f e r t i l i z e the vegetation to maintain the remarkable productivity (DeLaune e t a l . 1979; Wixon 1980). Though less spectacular than the annual flooding of the great r iver system l ike the Nile or the Mississippi, the daily r i s e of the tides and the sediments they carry may be jus t as in;portant for the productivity of these systems as those riverine s e d i ~ e n t s were for man's early floodplain agricul ture.

0 .

sediment surf o c s _.I .-. - . - . .- .. . . -J*..".-- : ..-,

. . ...... ...... ..+.-:'-" i ." *..- .-,...-- . - .-...-*.-. . ".---- .ae.... ..-/-+. ..C" . d5 . - --.' -.$

. .- -.<-

30 cm

. ...- 4 "

2 0 . " . - I 0 ' . .*. ar I o- 0

Ver ZieaO Exaggeration : 2.54X

Figure 14. Surface microrelief across the t ransi t ion from low to high marsh a t Farm Creek, near Rew Haven, Connecticut (McCaffrey 1977). Note the e f fec t o f =rtdna - paten? tussock development ori sedfment surface.

20

CHAPTER 3

ZONATION ON THE MARSHES

Striking patterns of plant zonation on the New England marshes attracted the attention of coastal ecologists and, beginning with the classic studies of Johns~n and York (1915), there has been a continuing e f fo r t t o understand the n;echanisrr,s responsible for the distribution and groupings of higher plants, as we11 as algae and animals, on these marshes (e.g., Nichols 1920; Johnson 1925; Taylor 1938; Chapnian 1340; Frliller and Egler 1950; Webber 1967; Daiber 1377). Most of the early e f for t s were descr ipt ive, and we now have a reasonably complete picture of the various marsh species (for e x a ~ p l e , a taxonomic guide for the Northeastern United States was prepared by Moul 1973). Progress has also been made i n under- standing the significance (or lack of i t ) of various plant groupings and i n appreciating the other factors in addition to t ides that influence marsh vegetation (Niering and Warren 1980).

H I G H E R PLANTS

The working definiton of a New England high ~ a r s h i s often a taxonomic one, encompassing the area dominated b y s a l t marsh hay or fox grass a;tina patens) and spike grass '%EXEiilis spicata) , i n contrast to the r e a u l a r l ~ flooded marsh on which cordgrass - (Spartina- a f te rn i f lora) i s vir tual ly a monospecif ic doninant. Along the upland border, the high niarsh often develops large areas of black grass ( J u ~ ~ u s a and switch grass

2 1

(Panicurn vi rgatum), a1 though where fresh water enters the marsh, c a t t a i l s (Typha spp. ) or reeds (Phragmi t ez ) often dominate (Figure 15). B u t the situation i s complicated, As Redfield (1972) observed, "The distinction between high marsh and the upper levels of the inter t idal marsh cannot be clearly drawn." Many writers seem t o consider a t least a portion of the stunted 2. &terniflora, which grows away from the creekbanks, as belonging t o the high marsh, and the characterization of any portion of the marsh as " inter t idal" may be ambiguous. While i t has often been reported that 2. a l te rn i f lora grows u p t o the level of rrrean hiqh water, and .. that this should define the "intertidal " marsh, a careful analysis of this proposition by Lagna (1375) has sh~wn that i t has l i t t l e merit except as a rough approximation. Because the level sf KWW is an arbitrary datum based on a 19-year record, i t would be a remarkable coincidence i f MHW was a finely tuned botanical indicator,

Our mdel o f s a l t marsh vegetation must include a certain amount of overlap in boundaries. Factors other than t ida l range may also influence the vegetation patterns. The more prominent factors have recently been summarized by Niering and Warren (1980), including sa l in i ty (Taylor 1938; Adams 1963; Parrondo e t aS. 19781, nutrients (Adams 1963; Kendel ssohn 1979 1, and soil oxygen (Linthurst 1979). A77 factors contribute in varying degrees

Figcare 25. Some contmon higher plants of the New England marshes. ( a ) 5 m c s t h cordgrass, 5pa)rtina c t e r n i f lor8 ( b ) SaT drneadow grass, Spartina patens ( c ) Blackgrass, ----- Juncus E r d m i k e g r a s s , -- -- Distich1 i s spica ta .

F i g u r e 15. (Continued). (e ) Sai toiarsh b u l r u s h , Scirpus robustus ( f ) M~rsh eider, & frutescens ( g ) S w i tchgrass panicup v i rgatua 3) Common reed-grass, P h r a g m i E comuni?. Drawings from Tidal Marshes of Lyme, Connecticut, - pub] ?shed b y m l d Lyne ~onservation'-=issje;tn, 19-68.

2 3

t o the height differences between t a l l creekbank S. a l ternif lora and the stunted f o k on the more poorly drained back marsh. The greater oxygenation of pore waters by the tides may also be the mechanism resrtonsible for the correlation betkeen creekbank S . a1 ternif lor2 production and t idal Kanqe reported by iteever e t a l . (1976), " On .the high marsh, however, the s i tua t i on i s even more complex; no simple or direct relationship has yet been found between the distribution and abundance of high narsh species and a particular environmental variable. En spi te of some exciting advances in our under- standing o f marsh zonatian and niarsh plants, Miller and Egler's (1950) comment s t i 11 seems appropriate:

"One i s tempted to feel that this remarkable mosaic should be interpreted in terms of ecologic factors. I f so, our present knowledge is yet far from suff icient , I t i s very l ikely t ha t contenlporary concurrently acting factors are only partly responsible fo r present d i s l r i - butians. I n other words, simple abnorn~a'f ca tastrophic factors rnay have produced effects ? a s t i ng into the present; and general past condi t ians may have been such that the vegetation s t i l l ref 1 ects them. '"

The picture developed by Hiller and Egler (1950) from their work in Connecticut i s probably the n,ost useful general ritodel of vegetation on the New England s a l t marshes, I n the i r studies of the Wequetequack- Pawcatuck marshes, M i 7 8er and Egler fzund scKe 150 species of higher plants distributed in five belts or zones classified rougt.11~ accordi ng to elevation. The number of species was greater, and t h ~ r e were nore coman species ( 80% occurrence in

2 4

t e s t quadrats), on the upper borders and slopes of the marsh than on the lower portions or on the creekbanks (Figure 1 6 ) . Their general upland- to-bay sequence consisted of a --- Panicum virgatum Upper Border, a --A- Juncers gerardi iipper Slope, a Spartina Fatens- Lower Slope, and a Spartina --.-..--- a l te rn i - flora Lower Border, B u t F4i91er and Egler also devoted considerable attention to the shallow pannes a n d pond holes on the rrarsh surface, Flany Danncs were characterized by s tunted Spartina ----- al terniflora or by" colorful forbs such as --- Limoniuni carol inianurn, TricJochin maritima, Aster Genuifol- - ---- ius and Flantaga decipiens which grew ____' around their edges. Toward the inner portions of the pannes , evapara t i on and poor drainage produced areas with s a l t accumulation that were colonized by succulents such as --------- Salicornia e u r o E or remained unvegetated. Pond -- holes w.i t h the subnjerged a~acrcrphyte,

maritima, and various algae occurred in other areas.

The extent of these zones varies considerably in individual n~arshes. I n general, the ---- Panicun; Upper Border and the Juncus Upper Slope are narrow and separate the marsh proper from u p l a n d trees and shrubs. The high mirsh, c o n ~ i sting o f Spartins patens, Distichlis skicata-, and s h o r t s . ------ - a ? terniflora i n varjous con~binat io~s --- o f pure stands and mixtures, appears to comprise the largest area of most unfilled marshes. The regularly flooded or low slarsh consisting of the t a l l form of S. ----- al ternif lora -- often amounts t o 10%-ta 20% of the area of emergent grasses (Table 4) . I n the nast, the coveraae of the hiqh marsh 5. &ens, h c u < , and ~ i s t i c h ~ i s nay - ------ have been even qreater. In comparing their nore recent ntargh surveys on Long Island w i t h those B G ~ E 34 years ear l ie r by Taylor (1938), 0 'Connor and Terry (1992) noted that :

"Taylor described Sparti na patem as %by far the mos"eommsn Grass

UPPER BORDER UPPER SLOPES LOWER SLOPES STUNT ED SPARTINA LOWER BORDERS I08 Specres 16 S p e c ~ e s 10 Species 12 Species 5 Species

Ciudium moriscoides,

Ponicurn

RELATIVE OCCURRENCE , % of t es t quadrats

Figure 16. Relative diversity, dominance, and major species cornpasitisn o f vegetation zones described by Mi 1 l e r e n d Egler (1950) a t the Wequetequock- Pawcatuck narshcs i~ Connecticut,. I n each zone, species l i s ted are those present i n 80% to 50% or 90% t o 100% o f the sample quadrats. For example, in the upper border 1C8 plant species were found; 5 species occurred in 80% to 90% o f the quadrats sanipied, and 4 species occurred i n 90% to 100% of a l l quadrats. Alvost 50 specl'es were rare and only found in 1% to 1 0 U f the quadrats.

on the marshesk whereas we now estimate i t covers only 16 percent of the varshes, or i s less than onc-quarter as cormon as -- 5. -- a l te rn i f la ra . Taylor a l s o described -- Juncus germrdi as %ndoubtcdlv the nex t nost "

prorinent p l a n t , . . , A w k e r e a s there now appears t o be fewer than 30 acres of - 3. s r a r d i -- + "

25

The loss o f h i g h marsh appears Lo be due t o i t s suscept ibi l i ty t o f i l l i n g and development, and i t may be t h e lack of man" -influence as much as any feature af geography t h a t -is rezpsns i ble for the re? a t i ve abundance o f high marsh in northern New England (Table 5 ) - In their recent summary of &eLland loss in the United States, Gossel ink and B~altmann (l980) reported

Table 4. R ~ I a t j v f ai:rount ( $1 of coverdgf o f h i g h and low ?:tir~h i n various Fdew England sa l t marshes. I___*-- _ _ _.-_I-_ I_---- X-----I_------------------

Tall >partin8 S h o r t S . Fiixed S. paten? Marsh lacat ion _-. altcrniflora ~-Iw..II_I- -L- --- a1 te rn i -- flbra a n d E i s t 7 ' e : ~ j j ~

Barnstahi~, W I ~ , . . 3 % . . . . . .11. . . 53 Barnstable, MA , ti . * .S%.. . Bissel Cavc, R J 7 ?% 19 Cottrell Marsh, GT . . .45. s

6 5 H ~ n p s t e a d Bay, 1 t: hi 7 2 3 Flax Pond, k.1. * . . 6 0 . + . Iron Point , 1 .1 . . . .56.. , Nassau & S u f f o l k , L . I . ' 14 45 2 3

_I_______11_ -- - - _ __-_ . . . I _ _ " ~ ^ - ~ - - - ----11-------------- - - -

a ~ l u n ~ l f 1968). ''ltedfjcld (147%) $ o f ciii;ergt~nt slarsh. C Nixon a n d OvfakC (19X3a), '~ttscvrr ( 1 0 1 % ) . p ~ d e l f ct dB. ( 1 9 G S ) .

~ a ~ n a (141!1). ~ ~ l ~ a n n a r - artd Tpu.ry (1$?2)), averrrcjp f o r virhtial ly a! 1 o f the rnarsfles -in the t a ~ ~ count if:^, alxcept i t i s not c f c a r i f t h e r~tarshes o f t.icrnpslrctd arid south Oys tur Bays w r c 17xclrrd~d as be ing " ' a t y p i ca? . "

C i s t i c h l i s , Juncusf to l o w marsh Tdble 5 , TFK r a t i o o f h igh r:%i'lrsf.~ (1. git_ig%, --__-_---- -- I S . d l t c r n i f l s r a ) i + l u ~ y t l ; ~ Atlantic coas t . Tl ie ~ ~ t e n t of each wet land type was nF6i<t;t:y68'f?Iy"" Spinrler ( i (16cj) based on 1454 U. S. Curcau of S p o r t F i sileri es and W4lclZlile d a t d and bq t l t e % Kairw D~pdrtri lcrat of I n l a n d Fisheries and Garle. *a*. -*_--*__ 1- - 1*""11"4- * "" .̂ I - --- -- ." "_. ~ll______--ll_l

SLa tc* i r i g h r:iarsh/low marsh *- _ "-_^_ .-__*. ̂- _ _ I _ _ _ 1 1 1 * -" _---- .ll_.~-lllll---_l.__-.-----l

I nca 11. l S4ew Haiiipshr r p 1 4 , X Eiassathusrtts 4.3 Rhodc I s l a n d 2. J Cc~nrrtjetjcut 3 .3 FSew Y or k 3 - 1 New Jersey 7 , 2 Delaware: a 1.1 Mary1 &n"d, 7.4 Visga'nia C. 5 North Carolina" 1 . 7 bnutR t a r n l r na' 0 , 3 Georgia 0.3 Florida {east ad st)^ 3 , s

that the rates o f loss in Naine and New Ha~npshire during 1954-74- (and presumably earl i e r ) were much 1 ower than for most of the other Northeast- ern States.

Ears h vegetation, however, changes in response to Elany other factors besides those re1 ated to human devel opslent. For exanp 1 e , Njering and karren (1980) descri bed sh i f t s that have taken place during the past 25 years on the Wequetequock- Pawcatuck narshes, including a loss of the Juncus be l t and a reolacement of the 5. patens by short '2. a l te rn i - flora.. After studying more than 10Q marshes on tons Island Sound, Nierins and Warren f e l f that while:

"

"the Miller and Egler pattern was found to be generally valid ... t idal marsh vegetation i s highly dynamic, and our f i e ld observations and peat core studies have shown tha t traditional succes- sional concepts are of limited

value i n terms of interpreting vegetation changes."

As a resul t of their work, they developed a revised version of the often reproduced generalized cross section of the vegetation on a New England s a l t marsh that was f i r s t pub1 ished by Ril l e r and Egler (1950). The resulting diagram (Figure 17) i 7 lus trates the complex distribution that may resuf t from a sequence o f changes that Chapman (1940) t r ied to represent w i t h an involved web o f potential vegetation sequences on the New England marshes. After struggling to understand salt-marsh succession, Chapman commented with understatement:

"This scheme may appear somewhat bewildering as i t i s very complex, b u t the present author has been forced to the conclusion that s a l t marsh succession i s by no means the simple phenomenon seen by ea r l i e r authors, and that i t can only be represented

/ UPPER BORDER ,! I I

I

HIGH MARSH i LOW MARSH 1

Figure 2 7 . Generalizer! tranzecr, f ror the u p l a n d s "c the ;ow i n t e r t i d a l i n a "typical " New England s a l t marsh showing the common vesetation types. Key to symbols: Sat = t a l l Spartina- a l te rn i f l s ra : , Sp = Spartina patens; US = Distichlis 2picaka; Sas = short Sgartina a i t e rn i f l a ra ; I f = - I v a frutescens-; dg = Juncus -- ; Pv = P a n i c ~ ~ virgatum; Pa = - Phragmites - aus t ra l i s . From Niering and W 1980).

schec.s~atically by a canrplex d i a - gram.

I think few would disdgree with such a canclusion, b u t bliller and Egler p u t i t another way:

'\ . . the present nlosiac may be thought of. as a r;~ol~entary expression, d i f fe ren t in the past , destined to be di f ferent in the future , and yet as typical as kdould fae a photograph of rlroving clouds. "

EPIBENTt I IC ALGAE.

Specic?s conlposi t i o n and dis t r ibu- t i o n p a t t ~ r n s of epibenthic d l g a e u n the rriarsh surfdce a rc not well known. Afgac a rc n o t as conspicuous as ilawering platlts and have r~c t? ived less attrrrtiort; the lower plants a rc s l~or tcr-1 i vcd a n d helve less specid1 - i r e d growth requi re?ni~nts,

I n ddd i t i on to h is investigations on the zuccesxiot.1 n l yrdsscs on Koninclly a near 8ostun. chdptriatl ( 1 ~ 4 0 ) also identifird d 1 y d l G I I I I I ~ ~ U I I ~ ~ ~ ~ S wirich he recoyn~zed on tile bdsis of spccl'cs ccrinposition, t i d a l rartge, artd r,@ac;on, R l terripts t o i t~iposc: tdxonorrii c order 011 CFDC ~t~arsh a1 yae, however, wer+e n u t w r y ca!~virici rry or uscaful , As recerrtly c ~ s IY67, Webber rioted that no s p e c i f i c a e ~ v u n t s of New E n y l d n d c ~ r s h algae hdd been publ ished i n the 27 years f o r lowing Chdprndri's pdpcr.. ijer own work urr Ctle bluu-green a l g a e o f s 1tl;4rsh a%. Zpswich, Mcassachlast..tts, i d e n t i f i e d u v w 30 species t h a t app~dred t o be dssociated w i t h tile various Friyher plants ( 3 w i r h ~ J ~ I M C I I S ~ ---- .%<

13 w i t h --- Spartina @ t e n s , 20 with S, -. al tcrni f ' tora , and 4 Frorn the subiix- -.---. - trrrikl^r; although no algal curf*+ ,riuni t i e s up. zones were real ly defined. Kithin ;a year of kebber" pubfiication, John B i t d m k (1968) rr:onograph, " S a l t Fkrsh Spartinas arid Associated A1 yae, " appeared, While recognizing t h a t n:ang, of t h e dsc;iriant algae ide r~r i f i ed by Chapman were characterist-ic of t i le

2 8

ri?asshes, B'i un, concluded t h a t : "'r~ss t species of s a l t marsh algae appear t o grow i n r i s ce l l a n ~ a u s rnixturcs with other species. Observations of nurrierous marshes in~presses me with the paucity of i:iixed co~~nitlni t i tls which dre constapt i n n ,ake-up. 'VeencraI observations, however, could be imde about the algae on the Cape Cod ;ilarsh~s. For exari:pIe, the a 1 g a l laver under the t a l l creek-Sank S. -- a l t e rn i f l a r a -- -- - consisted mainly of d'r'atorns qrowinq on the niud surface, while t h e high nlarsh s tur~ted S . a l t e rn i f lo ra was associated w i t h ------- ---- filaaientous algae (Table 6 ) tha t grew upward on the grass leaves and culnis to a height where the h~mid i t y became too low to support algal growth, H i g h iiiarstl algae a lso were found associated with other plants such as Linioniunl, --- -- Fqantaao, and Salicorn-ia and in -* .----- .".,-- .---A-p.- unvc~eta ted uannes, With the e x c e ~ t i o n of ----- ~ i ! a t h r i < which grows u p an the S . paterrs ----dm mat, there &as v i r tua l ly no d l q a l layer bolow tile h i q i r i~iarskt S .

w -- ggter~s -- -" - and ------ Oistichl j s .

The l a c k of algal cover over the lar'gh rriarsh doriiinated by these spec? es i s clue t o the shading of the ntarsh surface by the dense S. ~ a t e r t s mat (Rlu r : , 1968). On a s p r i n g d a y x l y 2% t o 32 of the incidrnt 11 'gh t reached the so i l beneath the S. patens- ----- Dist ichl is -.-- mat studied by Bluni, whi le 5C1'; to 55% reached the a l g a l layer under stunted and creek-bank S. dIterniflorc3. The growth of epr- ----- -- - - - .- " benthic a l g a e under S. a l t e rn i f fo rd

----a .--- is l i g t l t dependent and appears t o be greatest during spring and fa1 l when the grass cover i s not as dense (Sullivan a n d Daiber 1975; Van Raaf t e e t a ] . 1970). High l i gh t intensit- ies appear to favor the growth of f i 1an;errtous a l g a e whereas diatorns dorilinate i n law 1 S ~ h t areas (Sullivan 1974; Sulf i v a n and Daiber 1975).

EiARSII AN SMAPS

I n the sea, the det-rsity and kinetic energy o f the water provide

C X T 3 . c u ( n a J v 1

L* L m r g

;F.G 7 O w l l i s .,.- IT- ,--- &'TI fa 'TI "2"

I E Q I L A 2 7 . r =I C O - W O I Q m o C - ' - C F W ' F -v E a - Q

'" L. W a a;.'+= sr 02 L- 9, L 3 5

rJ O M W > a 3 . L i - C

X s-r-, q-v .& L.

43 0 in Q

:C - ' gLg " m ~ 0 E $524 0

C a J L rd rrn 5 rnc rn a ClL N

LO f 0 0 " 34A+ b-

2 u *2* 28- L ' P U

' P- o n -C,o C - 0 L O v QJ aJ . 3 . , m >, -0 ,o

LnLL W C T b a U r o n r..c U G r l (Ll t

a c * + c c 'P fa m 'P-

C, 'P L W-J-trp-. v, t u - 0

O m 0 .r C - ' r U a J Q C , ---as= a

43 u+i -5:. k f-3 *-a ( f . z -a C T Q )

f% g -Q - . w mLO "dp;

3 C- U $I$.- u,., L L m C n +J $ $ 3 "

C B , - QJ 0 EL-= as.- fa r. g

a m 0

N 3 iJs.p E ~ * , " , " i - a ? s= v,zIC, E

"c t -. L O - u> Ifla

h:

Qi a

a m - 6 J S z Y ) > > @ a -

L , * w c -(U LC., no a e m . - . r - - ~ ~ E h w Q i n o V ) cn L S I1S L m'k- cr Q6JX C n r 0

+3- C rJ a h i L I W r-r ir- %- e~

,- 31 U C

'?--

-%? 5' (U V

4=J .r rV +-J L U a (Li

CJJ s +" E: L 5 CJJ ti > C fi *r 61 p 0 L C )

0 G? 'TJ c C *P

7 &

fa 0 =r'W c m a C L

't-- K3 5 g CLt -

I/, . t

Table 8. Average number o f Aedes masqui t o larvae per dip of water on the marshes a f Egg Island, New JG-SG (Ferrigno 1958).

--- --------a -- --- Species -----

A . cantor A . sol l ic i tans T o t a l - -- -- -+

- 6-.

S. patens-S. a l ternif lora mix - ..... --- S. a1 terniflora ."". -

d i f f i cu l t , even when these areas are f l o o d e d . I n the case of the green- head f l i e s (Tabanidae), i t appears that the larvae drawn i f they are subjected La more than 2 days of subn~eryence.

N i t h the exception of Dexter's (1947) monograph on the i ntert ida 1 i t n i ~ ~ a l s of Cape Ann, Massachusetts, and Tiner's (1974) thesis on a niarsk~ near Stoningtan, Connecticut, there ayjpears t o have been l i t t l e work o n insects or other artinla15 i n New England marshes. " f h e n~ust extensive g e n e r a l study sf insects on a s a l t marsh was carried out in North Carolina by Davis and Gray fT966) , wha found a ratarked zonation that correlated with vegetation, particular ly on the high marsh, I n general, bot.f.1 S, g j Q e r n L f f s arid gjsticb& had maye insects than d i d 2. p3.J.r12 or Juncus roerrierianus. Because izos t -- ----- insects can escape the t ide by P l y i n g or hopping , i t i s likely t h a t titel. abundance of insects on the n1arst.1 i s regulated more by food a n d shelter than by the hydroperiod,

Crabs and Snails ------- .a T

Severdl crab species rive i n marshes; most i n h a b i t the lower $, a1 terniflora rone rather t h a n the high marsh. On the Farn: Creek, Connecticut, niarsh studied by EcCaffrey ( l9773, the density o f

3 2

fiddler crab (Uca w n a x ) --- burrows declined fro"~ 2 5 4 . 4 40 ( H i s)/nJ in the cree -bank S . alterrliflora t o -- --- - 64 -+ 20/mS a t a " ' s i te 2 rr. i t ) i n l a n d i n S. &ens t o 2 + 3/n;? in the nliddle of--the C-laJcn: rone, The relative lack ofl crabs and burrowing animals on the h i g h marsh nray be due directly to the lower frequency of flooding (especially for those species which dre active only under water, Teal 1959) as well as t o the dense root and rhizarne mat of S. ~ t e r t s (Dal'ber 1977; Frey and ~asan-l3787-.*-- One con- sequence i s that there is considerably less hioturhation or r r l jx iny of the high marsh seditilents (McCaffrey 1 9 7 7 ) .

While nlost a i the work on 111arsh crabs has beer] perfar~ed in southern marshes, Dex te r (1942, 1944, 1945) published a series of detailed studies on the n:ot luscs o f Cape Ann, Massachusetts, including those of the rr:arshes. in coritrast t o the distr-ibulion o f crabs, he found t h a t h igh icarsh S. patens --- was the most isngortant h a i t a t for the conimon cnf f ee bedn snai 1 , !;el amps- biden- tatus* Dexter ( 1 9 v - identified --- a Spartina patens-Pielail! us-Orchesti a (ai:~phipud, beach " '&ssociation as one ot the seven ma;or ~ lar ine coixmmi t i e s o f the Cape A n n t-egion. The comnjon r;iarsh sna i l s , i i t t o r i n a 7 i t to r ia (periwinkle) and L. ---- saxa- t i l i s , were also a b u n d a n t on-the high ---- marsh.

m 2 w * - C 0% a L

%-", g 3 E

G a 2 r . e - 0 a L T3 nS -r

E U & + ? g s

.,-- E Qd

x J * S o ? i - c

'+" g I-' .P

ij-1 n/ .;: 4 2 Z g E 8 o n r w a r n 2 m 0

Far cxa~:ple, the clapper rai 1 (Rallus lan~i ros t r . i s ) is often conspicuous by ..",- ---- j t s faniiliar c a l l i n southern New England rriarshes duri n y sumnier, and ottrer birds incl uding great black- backed gull s (idLa2 tnarqj~j2-), herring gulls (L. g e&_i"';usrlaughing gulls (_i. a t r i c i i l a 7 coai~~ion terns (Sterna h1rur2drF---and l eds t terrrs----E. *---.--. albifrosrs) also use thal h i g h rr~arsh for -- - -- - - nestlny ( l -ucid 1971; Nixon and Oviatt 1973a; Burger a n d Shisler 1978). The relatively high diversity of birds on the h i g h titarsh is l a r ~ e l y due t o the ""e!j!i e f fec t " of the njarsh-upland ecotonr where shorebirds and water birds r l ~ i x wit11 f i e l d and Forest spec ies . I-I~cause !;)any species appear t u ~ e s t i n areas with l i t t l e or no tidal flooding, the high marsh may a 'fsa he cons idcrdbly nmre a t t rac t ive ds a nest s i t { . thdr r thza 5. --- alterni- .f lord ~utzet . l i~rger ist~d Shrster 0978J -- *-"-.. p o i need n u t , however, "Dcspi te the ex tens i v r rtbc,en t work orr shoreb i rds , l i t t l e i n t c ~ r r ~ ~ d t i o n t x i ~ t s either on gerrvrd 1 h d b ~ L d t prcfert.nccs, or on spce,i fic: nest-si te preferences." Thei t- f ~ d r h icular study was cor~ccrnrd w i t h r . r e ~ t - s i l e selection by t h p

w 7 I r l: t (La -*- - r -" ~p 2._ri1irtgLgs ~g$r ! :~p~j~&~, ;~ ) , a r,onrrnon 18~drsI'i bird off erl assncia ted with Sp$:J.j~i?~ &atens, - -- While wi'llcts d i d build ttmelr nests f'r-on) S. pdtens, --- - *.."... the irnpor-tdrrt env-irctntilentaf variable i r i nest-site i ;e'icctioll war; eleva2,ion rdther t k d n vcqetd t i u n .

t.lhrie few, i f any, birds arc c o ~ f i n e d t u the h i g h c~arsh habi t d t , rildrty ~ l j e c f e s usc the h i t j h m r s h for ctr:t.r or r;gou-r d c t j v'i t i rl; : f ced i nt:, cover, xzesti ny, o r r e a r i n g young, P h c $01 1abtin;r habi t d t ust?-species assoc ia t i ons i n Rew Engjand high rr:at-sh were ~re.vi ded by Ra 1 ph Anbrews and colifragues of the U*S. Fish and W i f d l i f e S ~ r v i c e i n Ka~sa'chu~etts,

Rest a n d feed ir, h i g h marsh:

Sharp-tailed sparrow Loncj-bil led marsh wren (Lypha -- or

Pjlragirli tes ) ~eadowl ark Savannah sparrow (highest areas) Marsh hawk Short-eared owl (local ) Black rai l ( ra re)

Nest i n high marsh, b u t feed i n pools of 2. al ternif lora zone:

Clapper r a i l W i 1 l e t Black duck Bluc-winged teal Canada goose Seaside sparrow

Nest in high marsh, b u t feed in open water :

Gulls Terns

Nest in u p l a n d edge, bu t feed in h igh marsh:

Yellowthroat Song sparrow Cathi rd K i ngbi rd Aedw i n g Grackle

N e s t on woody islands; feed i n the rnarsh :

tJcrons Egrets Glossy ibis

Nest elsewhere; feed on insects over :!;a r s h :

Swa 1 1 ow Chimney swift

It i s d i f f i c u l t t o quantify the importance o f different rnarsh plants and p lan t parts i n the diets of the

various bird species. Most waterfowl and shorebirds ea t a great variety o f plant or animal material or both, and the i r g u t contents may ref lec t re lat ive food abundance a t a particu- l a r t ine rather than food preference or requirement (Cronan and Halla 1968).

A1 t h o u g h no large grazing animals l ive on the New England s a l t marshes as they do (or did) on prairies and savannas, many smaller mammals feed or l ive there or both (Daiber 1977). The dense mat of Spartina patens and Distichlis spicata provides excellent habitat far the meadow or f ie ld niouse (Microtus pennsylvanicus) ; other small mammals frequent or l ive in the high marsh including the meadow jumpi ng mouse (Zapus hudsonius) , the whi te-footed mouse -us

-+i leucopus), the house mouse - musculus), and the masked shrew (Sorex cinereus). -

Larqer animals such as raccoon mink (RusteJa vison), mephi tis),d=el

on the shel l f ish, bird eqqs, and mice of the marsh, a1 though- their homes are usually in upland trees (raccoon), upland dens (skunk), or under fallen logs or i n hollow stumps (mink and weasel ). One of the most conspicuous anisals on many marshes i s the muskrat, Ondatra zibethica, whose diet consists almost e n t i r e l y of vegetation, including roots and tubers. The muskrat favors lower sa l in i ty ~ a r s h e s with less t idal variation. Nany New England muskrats use bank dens or burrows rather than the familiar large "house" made from marsh vegetation. The average house i s a mound from 1 to 2 m (3 to 7 f t ) in diameter and 0.5 do 1.5 m (1.5 t o 5 f t ) high. Generally, the mammals of the New England high marsh remain invisible to a l l b u t the very patient or fortunate observer, although many will leave some t rac ts of their pass- ing in the soft mud.

CHAPTER 4

COFiFrUNITY I4ETABOi ISM

Earshes havc attracted the attention of systems ecologists w h o are interested in the transfers of energy and matter in natural systems. The s a l t marshes of Georgia were among the f i r s t ecological systems to be studied as systen's; Teal (1962) synthesized information fronr studies conducted a t Sapelo Island under the overall guidance of E . P . Odum. The work of the Georgia group and others studying the mid-Atlantic, southeast and Gulf of Mexico coasts of the United States has docinated our thinking about wetlands, and only recently have results of ecosystem- level studies become available from the New England marshes (Nixon and Oviatt 1S73a; Woodwell e t a l , 1977 ; Valielr? and Teal 1579; Ide'lsh lG80; t-fowarth and Teal 198C). No one yet has compared s y s t e ~ a t i c a l l y the different types of niarshes. Probably the differences in t idal signature (Figure 81, t idal range (Figure 51, freshwater i n f l o ~ f hixon 1981) and sediment type (Hi17 and Shearin 1970; Cotnoir 1974) along the coast will influence the n~etabolism as we1 l as the species conposi tion of marshes. Reviews of the amunt of new aboveground production by W r t i n a a l ternif lora have already d e s c r m north-south gradients correlated with solar energy input (Turner 1976) and tidal rang? (Steever e t a l . IS76), Most of the work on eco9ogical energetics and nutrient cycl i n g has emphasized the regularly flooded S. a l ternif lora zone (3ow marsh), - - b u t some inforGtion i s available on the New England I-ri gh niarsh,

3 6

P R I R A R Y PRODUCTION

The marsh in sunimer i s a great sward of green; productivity of the grass is high. Ever since R.M. Harper (1918) made what appears t o be the f i r s t measurements of Spart ina growth on the marshes of Lonq Island, countless quadrats of vegetation have been clipped and weighed a11 along the U.S. coast (see reviews by Keefe 1972; Turner 1976; and a bibliography conipiled by the U.S. Fish and Wildlife Service 1977). Whi l e researchers in New Ensland have no t been as busy with productivity treasurenlents as the i r colleagues t o the south, even on the high tzarsh (which has been less inten- sively studied than the creek-bank areas) enough ~easurements have been made to establish that an impressive anlount of carbon i s fixed each year during the relatively short Kew England growing season (Table 10) .

E u t production, measured by harvesting the grass, i s an underestinate of the total energy or carbon fixed by the plants. Some growth h i l l have been eaten; sonle w i 1 l have been lost as leaf f a l l , seed dispersal, and organic exudates. A1 1 k i l l be missed i n an end-of-the-season harvest. There are various ways to try t o account for such losses (see Turner 1976), and some of them have been used by those work~ng i n New England. Unfortunately, i t appears frorr a cornparati ve study of commonly used techniques that the choice of a method fo r estirnating psoducti on will have a large influence on the resul ts

2 Table 10. Estirrates of aboveground primary production ( g dry weight/m l y r ) of vascular plants on New England high marshes.

Location -- - - . - . - .

Longa ~ h o d ~ a ~ e ~ N . Island ~ o n n . Island Cod ass.^ Claine f

Spartina ------ a1 t c rn i f lora ( sho r t ) 51C 250 43C 510 48Cr 705

Spartina - patens ---. 500 3f)O 430 1,160 2,740 920g

Salicornia at-opea --- - - 24C

S. Latens - g. s i c a t a n i i x - --- 446 680

Juncus gerardi . 570 450 425

Typha l a t - i f o l i a - - 1, 3 6 0 ~ 690 580

Phrag~i i t e s corn~r~uni s - -- --- 2,6130' go0

a Udel l e t a1 , 1969 (from end-of-the-season t o t a l bioslass).

b ~ t e c v e r 1572 (fron sequential riteasurements of 1 i ve and dead standing vegetdtion) .

C Nixon and Oviatt 1973a (fron, end-of-the-growing season t o t a l bionass) . d ~ a l i e l a e t a l . 1975 (fron; sequential ineasuren~ents of l i v e and dead standing

vegetat ion).

'~uber e t a7. 1961 (frorrr sequential harvests of l i ve and dead vegetation and assur;,ed corrections fo r grazing and decomposition. Data reported as ash-free weight; values given here have been increased by I N ) .

f ~ i n t h u r s t and Reimold 1978 (mean of f i ve techniques).

g ~ a r p e r 1918 (from end-of-the-growing season t o t a l biomass, probably a i r d r ied) .

(Linthurst and Reimold 1578). Grazins losses are sn:all on marshes, and the relatively shor t and d i s t i n c t growing season for Syartind itr the Xor th~as t makes the ha t -ve s t t e chn ique more apprayriate there than alclng 'the southern coast where grass grows continuously. B u t even in t h e Great Sippewissett %iarsh on Cape Cod a

37

3-year analysis of production and standing crop showed t h a t annual aboveground production may be as much as t w i c e t h c a:sxieurc standint ; crop (Va l j e l a e t a ] . 1975). The grea tes t difference between t o t a l primary production and the harvest of green vegetation appears t o be due t o belowground growth, however,

I n 1976, Valiela e t a l . published the results of the f i r s t ~easurernents of tire underground production of 2. al ternif lora and z. patens roots and rhizon~es on a Flew England marsh. Their remarkable finding a t Great Sippewissett, Massachusetts, was that belowground production on the high marsh was about four times greater than the green abovegr~und production. I n addition to the 630 gdw/m2/yr produced above ground, they calculated a production of some l,ClOgdw/2/yr of rhizones and 910 gdw/m$yr of roots (Table 11). Host of this production took place in the f i r s t 5 cm ( 2 inches) below the marsh surface (Figure 1E), began ear l ie r i n the season, and proceeded

fas te r than leaf growth (Figure 19). The hish ra t io of below~round t o aboveground growth i s surprisl'ng, b u t i t may ref lec t the fac t that *rtina i s water stressed. As Valiela e t a l . (1976) noted, the plants on the marsh appear greener and show increased growth fol lowing heavy rai nfal I .

In addition t o the production of t he higher plants, sane carbon and energy i s fixed by the mrsh algae growing on the sediment surface. This must be a verv small amount under the dense - S. --. mat, b u t in the lower ~ o r t i o n s of the hiqh marsh there i s a s ignif icant an:ountV of production by algae in the stunted 2. alterniflora- zone. The algal productivity i s trlore

Table 11. Effect of ni trogena (H) additions on the production (g dry wcight/mZ/yr) of h i g h tmrsh and l ow marsh vegetation a t Great Sippewissett Marsh, Cape God (af ter Valiela e t a ? . 1976).

-------

Fiarsh type -------- N application --- rates and No N addition

biomass conrpartment (control ) 2 2 +O.$gN/m/wk + 2 . 5 g N / ~ / w k

Low Marsh

Abovegraund Rhizomes Roots

Total 3, $26

High Marsh

Abavegrsund 630 Rhizac?es 1,610 Roots 9 10

Total 3*150 4,990 4.800

a ~ h e sewage sludge f e r t i I izer used a1 so contained phosphorus and other materials , b u t addi tionai expericxents demons tt-ated that nitrogen was the effective ingredient.

38

E R O O T S R H I Z O M E S DEAD M A V T E A

Figure 18. Vertfcal distribution of roots, r h i z o ~ e s , and dead ciattcr on the high niarsh a t Great Sippcirdissett Marsh on Cape Cod (Valieta e t a ] . 1976) - b{ost of the 'living n!aterial i s f o u n d within 5 cri? ( 2 inches) of the marsh surface,

d i f f i cu l t t o masure than that s h o ~ n above ground by grasses, and Great Sippewissett Karsh is the only s i t e in New England that has been studied for th is aspect (Van kaalte e t a l . IS76). There is soKe uncertainty i n tkc resu l t s , b u t j'ie apprartd t h a t algal production has greatest in spring, before the grass canopy shaded the s e d i ~ e n t , with a secondary peak i n fa7 7 . When integrated over. the ycar, a l s a l production a~ounted to SOPF? 10C gdw/ i~2 or about 20% of ti.?€ average aboveground S. al ternif lora production. A s i m i T a r - - i a m c ( r e p i b e n t h i c a n d epiphytic algae was also found in a l o n g Island rrarsh (lu'oodwcll c t a l . 197% ).

Thcrc i s also some production i n mrsh pools by phytoplankton, rnacroalgar, a ~ d , in soae cases, rooted n-iacrophytes such as widgeongrass,

Rup~ia maritima. This aspect of rarsh ecology- has n o t been adequately studied, t h o u g h recent r-~easurements o f phytoplankton and Cladophora p a t s in pools on a northern f4azachusetts n,arsk: si.ok,ed production c f about 2 550 gdw/n: /yr (Ruber e t a ? . 1L?8l). Because pools usually cover a sciall portion of the! rrarsh, however, their contribution t o total ritarsh production will 5c considerably lower.

All tkesc production figures are rough apprsxintations that vary considerably accordirac, to the ~ e t h o d used fer measurevent (Linthurst and Reir-(lold 1578) as well as f ro r year t o &ear and frois; place to palce, even w i t h i n 2 restr2cted 2rea. FOP exarple, in three consecutive years a t the Great Sippewissett Pkrsh, Va'lfela e t a l . ('1575) ca l cu la t ed the foliowing values far the h i g h rarsh:

@ ABOVE GROUND

800

400

0

Figure 19, Amounts o f aboveground v e g e t a t i o n and r o o t s f rom A p r i l t h rough November on the high marsh a t G r e a t S i p p e w i s s e t t Marsh on Cape Cod (Valiela e t a l . 1976),

Year Peak biotxass

gdw/m2 Met aboveground production

gdw/n~2/yr

1971 1972 1973

3-year mean

Although the authors concluded tha t the differences among the years were not s t a t i s t i c a l l y s ignif icant , a simple estimate of the maximum standing crop of grasses may appear to vary by over 48% of the overall mean during jus t 3 years of sampl in$. I n comparinq the end-of-the season biomass' of 'reek-bank 2. a1 ternif lora durinq one year on 12 marshes in Rhod6 1slan2 (a very small a rea) , we found a range in that one year from 430 t o 1,380 gdw/o~2 (Nixon and Oviatt 1973b).

NUTRIENTS AND PRIMARY PRODUCTION

Of a l l the environmental parameters that may influence primary production on the New Ensland high marsh (see Chapter 3 ; Valiela and Tea1 1974; Niering and Warren 198C), the most convincing evidence concerns the importance of nitrogen as a limiting factor. In replicated f i e ld f e r t u i z a t i o n experiments carried out over a number of years a t Great Sippewissett Marsh, Teal, Valiela, and their coworkers have developed data which show that nitrogen additions a t leas t as low as 0 .8 g ~ / d / w e e k during the growing season more than double the aboveground production of S. patens and Distichlis on the hiGh marsh. A similar effect was observed with the low marsh S. a l te rn i f lora (Vaiiela and Teal 1974: b1aTieI; e t a ? . 1975, 1976). Phosphorus additions had no effect on the production of any of the species, In terms of belowground production, the addition of nitrogen reduced the development o f roots by

about 75%, b u t more than doubled the production of rhizomes (Tab1 e 11). Overall, the production of low and high marsh appeared remarkably similar.

Since the f e r t i l i z e r input was maintained for about 6 months out of every year, the total nitrogen supplement in the Sippewissett Marsh experiments amounted to about 20 g f4/m2/.yr and 60 g ~ / r n z / ~ r for low and high treatment expe r i~en ta? plots , respec- t ively, on both regularly flooded and h i g h n;arsh. These i n p u t s are large compared with other nitrogen sources and sinks on the marsh. Bacterial nitrogen fixation on the high marsh a t Sippewissett i s less than 5 g ~ / r n ~ / ~ r (Teal e t a l . 1979), b u t there i s a net loss of nitrogen t o the atmosphere of about 4 g N/m2/yr from deni tr i f icat ion (Kaplan e t a l . 1979).

FATE OF THE PRIMARY PRODUCTION

Discovering the f a t e of the organic carbon and associated nitrogen, phosphorus, and other nater ials that are fixed on the high marsh each year i s not simple. Since there appear t o be few grazers feeding on the grass, l i t t l e i s transferred direct ly into secondary production of t e r r e s t r i a l animal t issze. Usually, the primary production e i ther accumulates in the sediments as peat, deconposes in the marsh, or i s exported by the t ides to more open estuarine and coas ta1 waters,

kccumuf a t ion i n the Sediments ----- -------

Surprisingly, feb studies of the scdiriients a n d peat found on keki England h igh narshes have becn conducted. Nith the notable exception of KcCaffrey's (1977; FcCaffrty and "Tonson 158Q) analysis of zpac3_?g patens peat a t Fartri Creek Harsh in ~ s n n c c t i c u t , the 1 i n ~ i ted inforcratian available is based largely on s tudies of the stunted 5. --------..-+..- a l t e rn i f lo ra - zones of two niarshes on Gape Cod, Because the conlpos i t i an of ri~ars h st:ditner\l: appears t o be somewhat higher in organic carbon and nitrogen than nearshore subtidal s c d i r ~ ~ ~ n t s (faille 121, some frdction of the biologicdl ly accun;ulated carbon and nl troqer~ on the nlarsh must also be bur ied alotig with the vinerrrl and organic r:ralerial deposi tr?d by t h e t ida l waters. Tfie role of phospftorus i s n ~ t as c lca r because there is s a w silc;ge'; t i on that th i s c:l.eii~ent may be released by ar~oxic marsh sedirrients. Ptic rtlrriobil izcd phosphorus rnay then bc exchanged ~CP"Q)P;S Lht: 5edin:ent-water i n t e r f a6.e

and rel!:oved fror: the carsr, on e b b t j de s (Nixor3 iSi;C).

Eased on a rrasnr>ablc r a n y i n density a n d cbcrwical ciiirpcsition of s a l t rrarsi-1 sccijv~t?-t, arld t i l e r a n y o f accretion ra tes sbrl-flar~zcd i n l a h 7 g 2 , sor~ewRerc between 75 to .aC'C: g Cjcs lyr and 5 t o 26 2 r n,y be dccurulated in rarsh p e a t ( P i yure 2 C ) . A consSderaliot.i o f the tor- yos sill on o f estuarine s e d i r ~ n t stic;ycsks tha tssor ; ,~ 35 t o 7 5 g o f the c a r b o n a n d 2 t o 4 a o f the r ~ i t r n q e n tiray bc a s s o c i a t e d k i t h the materidl that i s Ycrilovcd fro[*$ the t i da l w a p r . I h e rcc1ain117tj O , t o 365 ~4 C f r "lyr and 1 t o I& 3 f l / r5 . / y r w o u l d t h e n be dtic l o the b u r i a l o f S ~ a r t i n a and ic:arsh a l g a t i , though t h r - ----- contribution o f the 1att.er nus t kw very s~lidll.

i t sccrt (, ajlparent t h d t th~? source of this organic na t te r i s the 7arcie d s a u n t of he1 nwgraur~ci prodticl,~ori o f roots d n d r h ~ ~ o r l e s , dlthough i t i c s t i l l not c lea r khat 1s k a y p ~ n i r t q bclow the rlarsh iur tdcc .

l a b l c 12, Cornparl'sori at scdiriients fotrtld on ttte h i g h r,arsr> a t Far-r Creek, Connecticut;, w i t h t,tiasc. o f Long island Sorind and a s??ort 5 , a ? tcrni f:rtrd marsh a t Barns t d b l e , lilassachusetts. CaZa fror ."-- --- *-- - --"-,

McCaf frey 1 1 ? 7 7 ) and Redficld (1965).

S . c l t c rn i f fo rd - - --- --- -- -- *- i, i .Sound r:larsl-I, Mass.

3 Lret bulk densl t y '4/ci1!3 laO11 1.15 Dry bulk clensj ty 9 / c ~ : ~ 0,2 G , G 5 0 .25 Inorganic ifiatter g/rm3 V, 135 i,:* 624 G . 15 Organic content g /cr @,05i3 0.04 6. Cis Organic content, 2 dw 2 8 6 5.2 - ,, __l^_l____p_ --_-. __.-I ----." ---- ^ - .-----.----

a~veraged o v e r 1 n.

' ~ v e r a ~ e d over 5 n.

F i g u r e 20. The accumulation o f organic carbon, total nitrogen, and inorganic phosphorus in the sediaients o f a narsh calculated for d i f f e r e n t acc re t i on rates , sedil-ent densit ies, and sedirrent coiaposi tions (#ixon 1SSC). T h e upper and lower lines represent approxic;ateiy maninurn a n d n i n i n u ~ estimates based on the l i t e r a tu re ,

43

The helowgrsund processes my also vary i n d i f ferent areas of a ~ :a r sh , dependjng on water table and groundwater flow, t idal inundation, or sedii~ient input. I n analyzing a 1-m ( 3 . 3 - f t ) long core from the 2. patens zone, McCaffr~y ( 1 9 7 7 ) f o u n b - d rcmarkat31y uni form organic content w i t h dcptk and a varying inorganic cor:iponertt. Siruilarly, Redfield (1905) rof>arted 011 a 5-111 (16.4-ft) Core froll? the $, a l t e rn i f lo ra zone i n which the organic content a f the peat was rela- t ivcly constant w i t h depth while the dstt content vdricd by a factor of 10. Ne'ttk~er of these authors concentrated on the Sine structur-e o f the top 10 cm ( 4 incha~t;) or so, where the production of - Sjar~tina .."----- roots and rhizomes i s gredtest d n d niost variable (Figure 18; Vdf i e f a e t d l , 1 9 1 0 ) .

i f theh iinder$lrnur~d flroduction r d d ~ s nkeasurc1d by Valiela e t 31. ( T " i 6 ) on Gripe Cod are representative ~ 7 f ot i ier NPW E.ngland n~arshes, less than B third of ~ I I C J belowgr-ound frrcsdctctiot~ I S buried. f n the S. d l s " - t ~ * r t ~ i f "". "* ,. 11~r.a - "- zonei of the Great i~ftixaw i s % ~ h ' t t a i t appc!ars that orily aborit 5'; of the to ta l - . - Spartina jtr.ilcfair*t'ron ( 7 of below<;raund pro- d u t t f o n ) i s dscumulatc.ad in peat; the 3t1ttg(.r pdrt I \ consufiled acr-obicdl7y ori Ilrrh n~arsh sut'fdce ur through sulfate,. re:dur..tir~rr irr the dnoxic sedirrlcnt .

ile>cc!rx;us - - x - - - - - " r' t , ~ - - - - - orr

I t i s n u t su rp r r ' i i n y t h a t rriost of the or-yanic- rXiatt.er p i t belobt, ground by &!it> - 9yar"tina - -- --- does not rcrrlain t o forr;: p w t , I f i t d i d , Valieia e t a ? . (1976) cgtlculattd that i t alone wcruid ra ise the level of Great Sippewissett low riarsh by about I. cni each year, Moreover, the distrillution of orgariic siatter ~iti.1 d e p t h i n the sediment (Figure 18) suggests that nluci.1 o f the orgarric rratter produced near the marsh surface i s not buried, A t f i r s t , the r e ~ o v a l of such a l a r ~ e annual

increment i n belowground organic matter seerred d i f f i c u l t to explain. As Valie'la e t a l . p u t i t in 1976:

"We did not expect the marked decay in dead ma t t e r . , . . , s ince we supposed tha t d~composition i n anoxic sediments would be sfow. However, dead parts s t i l l attached t o the l iv ing plant would be supplied with oxygen from the p l an t ' s a i r spaces.. . , so tha t aerobic oxidation could occur. I '

Later work a t Great Sippewissett, however, showed that su l f a t e reduction by the niicrobial cornmuni ty in the peat appqgu'ed t o oxidize some 1,800 g C/rnL/yr in the 2. a l t e rn i f l o r a zone, an arr~ourlt roughly con~parable t o - - the be1 owground production (Wowarth and Teal 1980). I t i s a l sa possible tkla t belo~ground production measure- i;~ents can be confounded by the overwintering storage of organic n;atter in basal ~ o r t i o n s of arasses. In work with S. ' a ~ t e r n i f l o r d , Lytle and t4ul l (1988') G u n d tha t a large Pract i a n of late-season photosynthate was translocdted t o rhizonies and t ha t th i s trtatcrial was then used i n spring t o support n:uch of the growth of the plants through the fourth or f i f t h leaf s tage , Even i n midsurr?l;ier, "new rh-izorrles were regenerated largely usi t ig errergy stored in over-wintered rhizoit~es. " Unfortunately, s i m i l d r s tud ies dre not yet available for the 5. p?Lk~$ high marsh, nor do we ye t - have d i r ec t nieasurenlents of the decon:position ra te in the - S . patens -- zone.

The above~round primary produc- t i o n can be decoaposed on the r,,arsh surface or i t can be carried off the ir:arshe I f i t i s carr-n'ed off the marsh, i t m y accun:ulate on the? bottonl o f r!?arsh creeks and e~bayments o r i t may ren3ain suspended i n the water

colutnn and, perhaps, be transported into adjacent estuarine and nearshore waters. In general, the h i g h marsh S. patens i s n o t usually thought of a s contributing significantly to the export of organic matter from the marsh. There are a t least three reasons for th is opinion: the h i g h marsh i s much less frequently exposed to the t idal waters, the grasses are far ther from t idal creeks, and 2. patens forms a dense interwoven mat rather than an open stand of vegetation (Blum 1968). I n general, decomposition of the high marsh vegetation appears to be relatively slow. This may be true not only because decomposition i s usually slower on the ground than i n water, b u t because marsh plants (with the exception of Sa'l icornia, a succulent) are relatively resistant to decay compared with a number of other marine and te r res t r ia l plants (Figure 21).

Organic Export

Salt marshes are often valued more for their contribution to other environments than for their intr insic value. Nowhere i s this more evident

than i n the "outwell i n g ' h o n c e p t developed by E.P. Odum (1968, 1980), i n which the export of organic matter and/or nutrients to coastal waters from marshes has often been considered a major part of wetlands valuation (Gosselink e t a l . 1974). The rea l i ty , magni tude, and significance of "outwelling" and i ts role i n valuation have been reviewed by Wal ker (1973), Hai nes (1979), W . E. Odum e t a l . (1979), E.P. Odum (1980), Nixon (1980), and Shabman and Batie (1980), and l i t t l e will be gained by doing so again here. The high marsh i s not usually considered an important source of organic or nutrient exchange w i t h the t ida l waters. Upper portions of the inter t idal zone with stunted 5. a7 terni flora may show variable uptake or release of nutrients (Lee 1979), and some of the aboveground production of the grass may be carried off the emergent marsh into t idal creeks. I t would be d i f f i cu l t , however, to make a convincing argument that the export of organic matter or nutrients from h i g h marshes in general plays an important role i n the ecology of New England coastal waters,

DECOMPOSITION 1N WAT E W SECOMPOSiT!ON ON THE GROUND

Q 50 IOQ 150 200 0 50 100 150 2 00

T I M E , d a y s

F i g ~ r ~ 21, Oec~~i~positjon of various k i r ~ d s o f p l a r l t niiaterial on the ground o r subr;i.ergc;d in water a t d i f f e r e n t s i t e s . Individual points arc froni s ing le rrreasurcn~~nts while lines are shown for sequential measurements.

I n water: - - -- --- t: .- _Jun_cg (dc l a Cruz and A = S ~ r t i n a ..-..- - cynosuruides ----.--.- ---- Gabriel 1974) S = T j i s t i c i ~ l i s s&ata F = w i l l o w leaves (Salix)

-c frgrrde l a Cruz G = bi rch leaves (Betula) -- --- (Chanlie a n d Richardson 1978)

U = - Slartina ---- alternif l o r d (Wood H = 9 ~ - r L f 52~223r3j_dg5 et a l . ~9%37------- 1 = Dis t i th l i s sf icata

E = *---- lastera ----- marina (Burkhaldcr J ---. duncus ----*-- roert~erianus ( H , I ,J and Doheny 1368) from de la Cruz 1375)

F = ----- Juncus ree~terianus -- (de ? a K = - Cistichlis g i c a t a - ---- (Oduii~ Crux and Gabriel 1974) and de l a Gruz 1967)

C = Pel Lartdra virgi nica (Oduci and L += S. - -- al iernif lora .- - - -- (Odunj and RFirTTs7V c!e l a Crua 1967)

H = nrarine plankton (Gairbcr 1981) M = S c i r US dlr~ericat~us 1 = -. Ulva --- lactuca (Burkholder and 72;%-cFE-TgET-

Doheny 196t3)- Fd - i~ i l law On land: *- -- 0 = rhododendron

A = f i f t e r paper P = oak l.3 = fern ( P t e r i d i u a ? , d a t a o f Q = ash

rankl land 1966 i n F r a n k l a n d * r-. l \

R = oak 1 S j J " b J ; 5 = b i r c n

= c0n-i ferous leaf l i t t e r (data T = m p l c of Cl-ikola i n M511ar 1934) i! = e?r;i

D = sedge (Carex) V = a lde r (N-V t r ee leaves from E = --- Stincus s ma Cruz a n d rr~till s i t e s , "aocock 1564)

Gabriel 1574) W - ---- SaSicornia (Odun and de l a Gruz 19m

46

CHAPTER 5

HUF?Al'4 IliiPACT ON TWE i iXGIi KAR5I.I

Lying between the t ide l ine and the uplarid, the high s a l t rnarshes have been pushed in both directions by !]urnan ac t iv i t i e s . Since the mid IGOO's, the marshes -in New Englarrd have been Flooded or drained, intpounded or diked, ditched or f i l l ed . They have been converted into fresh or brackish water meadows as nell as fdndfil ls , parking lo ts , and housing efevel opi;ients. They have been prr39'sed for growing hay that saved 1 ivcstcrck and damned for breeding rnosyui toes that brought discomfort and disease, Human ac t iv i t ies have pol luted thein with aietals, o i l , cherricals, a n d trash. Recently they have been protected and preserved with environn;ental legislation. I t i s an interesting pattern of changing pcr~eptrans aqd values. In this envj row-.er?t, pf~rhaps more than in any other. :11ar-i rle ecosys tem, man has been bat31 riz6raager and n~anl'gulator.

SALT MARSH HAY

Before the s a l t c~arshes were considered wastelands i n need of "reclkimation," a n d even longer before they were elevated to the rank of a "sacred cok" i n the environmer~tal r:invcnent, the rnarshes were clearly, a n d i n t i ~ a t e l y , a part of the early New Enqiander's " l i f e support system. '"

While the cutting s f Spartina -- a t ~ n s or s a l t marsh hay is a recent E&

enough act ivi ty to be part o f the boyhood r;ienior-ies a f iyany preser~t-day hew E n g l a n d caas ta i farmers, il is

d i f f i cu l t t o appreciate the importance of this resource in the f i r s t 100 years or so of the agricultural econoi-y of the area. I n the recent past, s a l t narsh hay was a supplement used more for animal bedding, n;ulchirt~, and "topping" hay stacks t o keep f ie ld grasses dry, than as a staple feed. B u t a t one time the mrsh hay was a major food source which nlade the keeping of l ivestock possible and practical. And i t was livestock that fornied the mainstay of Nek; England agriculture in the ear ly years (Russel 1 5976).

The presence, a t leas t in southern coastal New England, of large areas of land cleared by the Indians helped the f i r s t colonists greatly, as did the open freshwater n:eadows a long the river floodplains. B u t i t was d i f f i cu l t to obtain suitable forage for a large number- o f animals, and predators, especia7l.y wolves, were a great p r o b l e ~ floiood 1634; Russell 1976) . As Bidwell and Falcotaer noted 1192if in their cl ~ ~ r i c u l t u r - e i n the s t a t e3m- I%@ :--- P -------.

assic History of Northern United - - --

"A condi t ion o f prii;;e importance for the successful raising o f livestock i s of course an abundant supply of native forage p l an t s . I n th is respect the Rorth Arrerican contr nent was strikingly deficient. The Ind jans of the region kept no herbivorous d~fiiestic animals and hence had developed no forage p l a n t s . * .. I n the face of such

di f f icu l t ies i t was a noteworthy accomplishment of New England and the Middle Colonies in the seventeenth century to have become not only independent of outside sources of supply, b u t even to have developed a surplus of ca t t le , horses, and meat products for export. "

The use of salt-marsh hay contri- buted substantially to this success and helped to determine the pattern of sett lepent along the New England coast. I n describins the history of New England agriculture, "Russel 1 (1976) has shown that the presence of fresh and s a l t hay marshes was a major factor in s i t e selection of many towns set t led before 1650 (Figure 2 2 ) . As he described i t :

"All along the winding Kassa- chusetts Bay shore, wherever s a l t grass caught the eye, exploring stockfiien were petition- ing the General Court to be allowed to se t up new townships. The adjoining lap? and n:ight be only moderately f e r t i l e , even chiefly ledges and woods, yet cattlemen brought u p amid England's grassy vales and t idal marshes coveted the s a l t hay in the lowlands. In Plymouth Colony the same magnet drew ambitious rnen toward new locations. Reluctantly the Plymouth authorit ies permitted their neighbors to leave the close-knit n:other town and i t s scant f e r t i l - i t y and set up new and distant farnisteads beside inviting hay l ands. Duxbury, Green Narbar (Marshf ie ld) , and Ningham, their t idal narshes rich in s a l t hay, drew planters northwzrd. The miles of green s a l t readow on the Cape Cod shore and Indian f ie lds there open for ti7tage beckoned s t i l l others to plant Sandwich, Barnstable, and Yarmouth, and t o

move inland to Taunton a t the head o f K t . Hope Bay. I'

On Long Island, and perhaps in other areas as well, the "sal t meadows" were owned by t h e town and the right to mow and carry off the hay was auctioned off early in the spring of each year (Kavenagh 1380). The same practice probably applied to "thatch grass" 0; S. alterni'f'lora. I t i s hard to know Tf this species was really used as thatch or as feed, heddi n g , or soniething else. Present- day farn:ers I have interviewed never recal l any use For i t , and Kavenagh (1980) concluded that i t was probably not used for roofing:

"Very early in the colonial experience in both Plymouth and Boston the colonists found to their sorrow that thatch grass for roofing quickly dried in th is c'l.imate, in contrast to Old England with i t s more moist climate and abi l i ty to keep the outer grass damp and less fire-prone. Here wood and ~ u d chimneys caught f i r e easi ly , sparks flew, and a dried thatch roof did n o t l a s t very long. Ordinances were soon passed to prohibit them. "

By 1700, "English grasses" had been introduced and spread throughout New Ena l and for pasture (Bidwell and Fa1 coner 1925), b u t salt-marsh hay continued to be used in large quantity throughout the coastal resion until the early 1900". Russell (1576) described the situation as i t was in the la te 1700's:

iiCountl ess stadd les (wood under- p j n n s n g ) for s a l t hay s t i f 7 dotted seacoast niarshes from southern Eaine to Cape Cod, along the shores of the Sawnd, and up the Connecticut and similar estuaries. f n the f a l l ,

F i g u r e 2 2 , Locations of Ne~j England towns t h a t were settled by 1650 ad jacen t t o f r e s h or s a l t hay mrshes (Russell 1 9 7 6 ) -

49

gundalows ferried the hay u p every s a l t creek to the home farm. On such provender thousands of ca t t l e and horses were wintered every year. The resulting manure, supplemented where possible from other sources, nourished merchantable crops of corn, potatoes, tobacco, flax, onions and other produce. ''

Staddles and gundalows (gondola) were s t i l l in use in Maine where they were photo raphed in the la te 1800's (Figure 23 3 . Fisreover, the value of the high marsh black grass (Juncus gerardi) had been discovered, and this species was also being harvested (Russet 1 1576) :

"'Salt hay from the thousands of acres of coastal marshes retained i t s importance, This 'harvest of the sea ' actual 'iy improved in quality, as the nutritious 'black grass, ' good fodder even far m i lkers, spread triore widely. Black grass, cutting about a ton per acre, made up ha l f the crop a l o n g Massachusetts Bay" sorth

The continuing i~portance c f s a l t hay through the l .800'~ i s reflected in i t s inclusion in the agricultural census data f o r the New Engiand States. For exrarr:ple, in 1875 farmers in Khode Island cut 1,717 tons o f s a l t hay from 2,506 acres o f marsh, for an average. yield of 0.7 tonssacre or 160 g/mL (Anony~,ous 18G7), The yield was comparable t o conventional hay fields a t the time, b u t low, relative to modern measupentents of the production of high mrsh vegetation (see Table 10) . Some of this discrepancy ~ s y ke dzc ts differences in harvest technique, or because sat t hay was usually harvested early in the season, before i t bent over and formed a mat that was hard t a cut (Kavenagh P98Oj, Even by 1815, the value o f s a l t hay harvested i n Rhode Island

was only $16,000 compared with s seaweed f e r t i l i z e r harvest (frop d r i f t on the beaches) of QGCI,OCC and a marine fishery of almost $450,00G. The impor tance of s a l t hay declined along with the fortunes of New England farming as agriculture roved w s t ,

CHANGES IN THE AEOUNT Or HIGH Ic'lAASH

For a tinre, the at t ract ion of s a l t hay may have drabhin s o w coastal farmers t o t ry to increase the acreage of high marsh. I n t i i s 17413 ---- E s s a ~ s -- upon Field Husbandr in New %'land, J a r e d ~ I i o ~ ( ' ? i 7 d ) 7 e E i b e d - % - ? s successful e f fo r t t o cortvert a '%holly unprofitable" low-lyina piece of swamp into a s a l t meadow, and sarg~ested that others might do the sdme since he had seen "'sundry such places upon the Sea coas t . I '

"Last F a l l I began upon i t and drew [dug] a Ditch 04 four f o o t wide from a large Salt Creek, a n d carried i t u p i n the nl iddlc of the Cove seventy Rods, in order t o turn i t into Sal t Meadow, that being the best t h a t ! c o u l d do with i t : I t so f a r answers the design, t h a t the h i d e flows regularly r 'nto i t , t o the upper end o f i t ; the Tide now flowing, where 1 suppose i t never reach'd before. l'

I t seerr;s irx~pass ib le t o dets rmi rie how much high s a l t rrlarsh m i g h t have been created i n this way, b u t i t must have been a very small arrlount. The more conimon procedure was fo r farmers t o dike the r~arshes i n a n attempt 'to convert then : t o f r e s h ~eadow or w i t h the hope of draining therr~ for growing tradi tional craps.

The expanding maritinie econotrY of New England during the 17CCis san'd the impact of the industrial r evo? ution during the 18CGk sust have rescllted i n rcore widespread f i l l i n g o f coas t a l marshes, particularly i n southern

Figure 23. Top: S a l t hay on stadales t o keep i t above the t ide . B~tts~i : Gunda'lcaw loaded w i t h s a l t hay t o be floated o u t on t he f l o o d t i d e . of the S o c i e t y f u r the Preservation o f New England A n t - i q u i t i e s , Boston, Pass .

5 1

parts of the region. Ejut no fys ten~a t ic inventory appears to have been compiled, and i t may be impossible to make one. S o ~ e appreciation for the extent of wetland loss can usually be gained by examining deta i led niaps of coastal urban areas a t various tiillcs in the past, The f i l l i n g involved in the creation of harbors (using dredge s p o i l ) as well as r:1i11 and factory s i t e s , roads, railways, and housing i s usual ly dramatic.

Data f a r r~~ore recent years are available from various sources l-isted by Spinner (1969) and Gosselink and Baumann (1980). Accordin5 t o the l a t t e r authors, wetland loss in New England s ince 1886 was greates t from 1922 t o 1954 (Figure 241, "probably [as] a r e su l t of public works projects of the 193C1s, the construction s f n:ajor a i r po r t s , the increase i n n i i l i - tary ins ta l Iationr; during World War IT, and a post-klorlti k'ar 11 housinq

boor.:' 4s discussed i n Chapter 3 , i t appears t h a t a disproportionate par-t of t h i s loss involved high marsh areas since they a r e l e ss often fioodcd, easier t o f i l l , and close t o the uplands <OiConnor and Terry 1972). F!uch of the r e n d i n i n s r~iarsh ' land i s in pub1 i c ownership, however, a n d l eg i s la t ion in the Eew E n y l d n d Sta tes now protects s a l t marshes, so i t i s l i k e l y t h d t the r a t e a t wetland loss due to hunan a c t i v i t i e s w i l l continue to slow. Eiut the dynanic nature of the 111arshes k i l l continue to r e su l t in vegetation chan~es a n d i n s h i f t s of s i ze and shape o f the coastal wetlands;.

I4OSOUITO BITCHES

Atalong the rr,ost consp-icuuus s i y r ~ s of human ac t iv i ty on the Rew Englana rirars tics are the ckdr-acteri s t i c patterns of s t r a i gh t para l le l ditches running froril the uplar~d edge of the r;!arsh or frorc old pond h u l ~ s

COASTAL MARSH AREAS- NEW ENGLAND & ATLANTIC N.V.

0 i ---- -- - - . q

1886 1906 "1922 41354 1968 1976

YEARS

Figure 24. Amount of coastal wetlanzis in the f;arLI:eastern United States;. (Gosse? i nk and Bauniann 1980. )

52

t o the larger tidal creeks. Spaced 35 to 7C n, (115 t o 230 f t ) apart, the shal low, narrow ditches were designed to reraove pools and standing water from the marsh and thereby prevent the brcrding of mosquitoes. Ditching as a riethod of ntosquito control appears to have begun in New Jersey a t the turn of the century (Smith 1902, 19071, b u t i t was practiced nost hr'dely during the Depression years of the 1930's with support from the Works Progress Adniinistration and the Civilian Conservation Corps. This atterrrpt a t "nlanaging" the marshes was so thoroush that by 1938 almost SO% of the t idal wetlands between Maine and Virginia had been ditched (Bourn and Cottan 1950).

The ecological in~pact of such a widespread a1 teration of the marshes i s surprisingly d i f f icu l t t o describe with any certainty (Daiber 1974). Much of the early l i terature appears t o be based on casual impressions and anecdotal information a n d often reflects the biases of "'rrrosquito controllers" or conservationists. The Findings of a widely cited study of ditching effects in a Eelaware marsh by Bourn and Cottam (1950) pay have been influenced by dredging in a nearby river (Lesser c t a l . 1976).

Inforniation is lacking about the effects c t f di t c h i n ~ on Nerv England high marsh; most of the work on this prob1en1 has been done in Uelawarr a n d NEW Jersey, though one of the better early studies on the effect of ditching on shorebirds and waterfowl was carried o u t in the Duxbury, b?assachusetts, ~ a r s hes (Bradbury 1938). I n Duxbury, the niarshcs had supported abundant and diverse waterfowl before ~osqu i to control r3per-il t ions were co~pleted, b u t a f te r di tchi ng, the ndrshes becacrie "dry and devoid of birds" (Daiber 1374).

Ditching can enhance the growth of high marsh plants a t the expense of

5 3

S ~ r t i n a alterni f lo ra , a1 though the - ---- -*-

ta'i 1 creek-hank 2. al ternifJora of ten srcrws alons the banks of the ditches i f the spo"i1 fronl ditch construction has n o t been l e f t there. Gdhere S. a l ternif lora does develop, the nest6-g and production of clapper rai 1s ( ~ a l l u ' s > ~ i r o s t r i s ) rriay be enhanced (Stewart 1951 ; Ferri yno 1966; Shisler and Shulre 1C76). Where spoil i s deposited, high marsh grasses or woody vegetation suc11 as Iva fructescens and Bacchat-i s - - -- -- halircifolia become established ( ~ ? ~ r - a ~ ^ ~ c ; l e r 1950; Daiber 1974). These species are generally of taw value, b u t s a w birds (e.9. hoat-tailed grackle, w i d i x ~exicanus ; red-wi nged b l a c k b m Age1 a i u s r k hoeniceus) may use then, for nesting ~ e a h T v n d Webb 19C3; Post 1974), Because the ditches are often dug to drain pond holes and other shal low depressians, submerged aquatic plants such as are usually eliminated. The lass of these plants as well as the protected open water makes the marsh less a t t ract ive t o waterfowl and other birds.

Gradbury's (1938) study of the Duxbury iriarshes suggests that %any of these changes can be reversed. Daiber (1974) sun~:arized i t in his review as follows:

"The technique of restoration was based on the premise that mosauito larvae would be eaten by

heteroclitus ~undulus - -_, ----- t h e mui;,michog nrinnuw. The job was to create a - h a b i t a t where fish could l i v e a t low tides and high tefiperaturcs. Forper potholes were restored by darnrcing outlets w i t h sod. Care war taken to keep Lhr water level about nine inches beloahi the marsh surface, t h u s , keeping i t free: of water. Sone potholes were deepened t o assure sufficient water for Fundulus t o l ive i n during dvy periods. Control led burning of s a l t hay made a variety of insects

available for shore birds and i t f i s h which feed on rrlosquito larvae. helped control niosquitoes by Studies have been carried o u t t o enhancing standing water evapora- docunlent the effectiveness of OF:i*:fit i n tion. Ditches were part ia l ly control l i n g rvasqui toes and i n blocked so water was retained enktancin~ the wildl i fe va lue uf the b u t did not flow out over the nlarsh ( T e r r i g n o 1979; Fcrrigno marsh surface. Bird use was e t a l . 1975). This contrasts w i t h ? reported t o inrmediately increase traditional parallel or grid ditch- without any loss in ~ o s q u i t o ing--a practice that has keen of control. questionable value in cont ro ' l l in~

n~osquitoes and that is t h o r i g h t to have Some of these techniques fornl had varyi nc; (and of ten undesirable)

p a r t of "open marsh water nlanagernent" i ~ p a c t s on overall mrsh ecology (OEWM), an a1 ternative to parallel (Daiber 1574). ditching and insecticides for mosquito contra? that has been developed by the New Jersey Departnlenmf Environmental POLLI!TION Flanirgement (Ferrigno and Jobbi ns 1968; Ferrigno e t a l . 1975) - Using this Because the h i g h nlarsfies are approach involves selective ditching above n1ear-i h i g h t ide, rrtnst of the tirw of major mosquito breeding depres- they are exposed t o the atrfiospbrere sions, f i 11 in$ shallow depressions rather t h a n t o tidal waters ( F i c~ure on the marsh surface, and care- 2 5 ) . As a resul t , deposition of ful ly crrnslruct.r'ng sotile ditches to particulate riiatter frorr: the a i r or i n c ~ l l e c t water in ponds that are deep precipitation can he rnaior patl~kidys enough a t a l l times to contain sri.:all for pollutant inputs. Ketals, toxic

A V E R A G E F R A C T I O N OF T i M E 1 M M E R S E B 1 ,Q 0.8 0.6 0.4 0 -2 0.3

E 2.6 - "J u7 z 1-0 M 3

2 o 0,Q - e- <e 3 I ~ P -i.Q .".J Lcl

- 2.0

Figure 25. Anount o f time the grasses and surface sedii~ents a t Farsi Creek, Connecticut, are exposed to the atniosphere a t different elevations across the marsh (McCaffrey 1 C 7 7 f .

organics, petroleunl hydrocarbons, and just plain j u n k also iijay be brought o n t o the high niarsh each rrionth with the highest tides. Junk i s often evident along the d r i f t line a t the upland edge of the marsh; when l a r ~ e amounts accunlula te the vcgetati or1 rnay be sn;othered and the visual quality of the area decreased. The accut~ul a t i on and effects of other anthropo5enic rrlaterials are usually more subtle.

Petroleunl Wydrocdrbons ----- Of the various o i l sp i l l

incidents in New England cotiipiled by Hyland (1S77), few appear t o have had a rrajor impact on s a l t narshes j n general or on the high crarsh in particular. F l e~c r th~ lcs s , the poten- t i a l i s there. Because urban scwase effluents are the ii3?lor source of petro7eur1 hydrocarbons i n coastal waters (Van Vleet and Quinn IS??), many rTlarshes in rrore developed estuarine areas r:iust be exposed t o t ida 1 waters with el eva ted concentrations of dissolved and particulate petroleut;~ co~rpounds. The effects of chronic, relatively low-level (conlpared t o spi 1 Is ) concentrations of these riatcrials on r;,arshes have never been assessed, however. The few marsh-oil studies which are available have been concerned with the impact of single or repeated c i l s p i l l s , and most of this work has been carried out in Europc or the Southern United States (Cow11 1971; Bender e t a l , 1977; Baker 1979). The only rtia:or study of the impact of an o i l sp i l l on a New England sa l t marsh appears to be the work of tia~pson and Moul (15178), who docut~iented the ir;:pact o f No. 2 fuel o i l on a riiarsll in Cjuzzards Bay, Nassachusetts. Their observations indicated that, in g e r ~ ~ r a l , perenniai plants such as spar't-ina cind Gistichlis were nrore resistant than ---- annuals 1 i k e Salicornia. B u t even for S. a1 ternif lora , the-Eiorr,ass, height, - --- and nuniber of plants were markedly reduced in oi?ed areas 3 years a f t e r

5 5

the s p i l l , As inight be expected, they also found t h a t plants higher u p in the rrsarsh recovered Elore quickly because their exposure t o the o i l was less. tiowever, petroleuc~ conlpounds vary widely i n coc;posi tion and toxicity, and their impact must also be a function of other factors including teo,perature and season. A t th is point, i t i s ii:;possible to rnake a kery useful speculation about the response of the New England high rriarsh comn~uni ty t o o i l s p i l l s or to a large nuinher of other possible perturbations.

Fiunierous researchers have investigated the various aspects of cfbundance, distribution, biological uptake, and effects of heavy metals i n New E n g l a n d high frirarsh conmunities (Nixon 198C'J, Because there are few burrowing anin~als l i v i n g under the dense S ~ r t i n a gatens mat, there i s --- I i tt 'le-bioturbation and the sediments appear to provide a relatively undisturbed record of metal input t o the oarsh surface. The higher concentrations usua 1 ly found near the surface clay ref lect an anthrapogenic ir~fluence (Figure 26) or my be the result o f rer:iobiJization of the niaterial a t depth, For exan!ple, in the case of Pn i t appears that manganese oxide i s reduced in the anoxic sedisientri, and the soluble En i s lost from the solid ptiase by diffusing through the pare waters and across the sedirient water interface (Figure 27 , RcCaffrey 1977; Lord 1980). For cietals l ike C u , Z n , and Pb, which are relatively stable i n the sedintents, i t i s possible to combine their vertical distribution with measure- riients of the sedifiwnt accretion rate t o ?air: 2:: estiz,at,e a f the history of antljropoc,enic inputs (Figure 29). I t i s a l s o possible t o compare the accumulation rates of different ~ e t a f s ~ i t h est i ra tes o f their input rates to calculate the degree to which the

COPPER , y g / g ash

Figure 26. Concentrations of copper a t various depths in the sediment under Spartina atens a t Farm Creek E1 fiarsh, Connecticut. he increase from 30-cm depth to the surface i s due to anthropogenic inputs, largely from the atmosphere (McCaffrey 1977).

MANGANESE , p g / g ash

0 250 500 750 1000 1250

Figure 27. Concentrations o f manganese a t various depths in the sediment under the S ar t ina patens a t +- Farm Creek Marsh, onnecticut. The rapid increase a t the surface i s largely due to a reniobilization a t Mn a t depth and i t s subsequent loss across the sediment-water interface (McCaffrey 1977).

5 6

-2 -1 E X C E S S F L U X , p g cm y

Figure 28. Historical variation in the antjlropogenic fluxes of copper, zinc, and lead recorded i n the h i g h riarsh sediments a t Farr:~ Creek, Connecticut (E;cCaffrey 1 S 7 7 ) ,

rnarsl? functions as a sink for various pollutants. General IS,, i t appears that Pb, Cu, and Fu are held very t ightly in the h i g h r~arst-i, with E n , Z n , and Cr skuwin2 only about 50% retention drtd Cd sorncwhat less (Siccama and Porter 1572; Banus c t a l , 1574, 1975; EcCaffrey 1977; Gibl in e t a l . 198C). The retention of nletals by the lower intertidal r;,arsk pay be considerably less coriplete (65blin e t a l . I98C).

I n addition t o providin~ us w i t h a record of pollution irrputs, i t also has been suggested that niarsf.~es niight serve as "biological f i l t e r s " for urban sewage. To ~xplarc, the ecological consequencss o f th i s i d e a , a long-term experintental s tudy of the effects of nutrient enrichrlent and heavy ri;etaf s was conducted a t Great Sipp~ivfssett $;at-sfT 03 Cc?ptt Cscl hy groups at Woods Hole Cceanagraphic Insti tution and The Maritit fjialoyical Laboratory a t koods Kale. Kuch of that work has been cited tkroughout th is con;i~unity profile.

5 7

TIE cxpcrin~ent i nvo lved the application of n;etals i n cornnier- cia1 sewage s ludge f e r t i l i z e r (Table 13) and i n dissolved form w i t t t u u t associated nutrients (Fe -- 650 nig/t;12/week; Cu and Gr = 20 mg/r;z/week) t o plots o f low and h i g h riiarsh. I n bo th treatr;,ents, the rlietats did n o t appear to have any effect on the growth of a a r t i n a patens --... -- or - 5, alterni f lora ( F i g u G 7 q accordi ny t o -Timm= a 7 . (1980). !lowever, both grasses becar~~e enriched in Cd, Cr, Cu, and Zn in plots treated kith large doses of the sludge rvixture (Table 14) . The fate of these n:etals i s s t i l l uncertain and, as Ciblin e t a l . (1986) concluded a t the end of their paper, "The role of the grasses i n ciaking riietals available t o marsh organisns is presently being investigated."

In 31;C years we seem t o have cove f u l l c i rc le , frsn~ viewins the New England 17;arshes as a source of food t o exploring their value as sewage treatrent p l a n t s . I suppose i t i s

1 ( j n sewage fer t i lizcrj added t o each p l o t aa?d arjaurat cf cdcl- the top 2 csi of r3arsh sedi~gents (Gib l i r e e t a l . 1960).

Aniount added Trea t r~ t~n t to p ' ig t t

Ar o v n t detected -- "."* -- - - m w -" * -& >- Ke ta 1 p l o t a [n,g Jc, 2) .ow r ~ d r s h i g h rarsr: - -[{- -- - - - - - a" -

2 f ~ y u r c 25. Abovcgrrst/nd bioniass ( g dry ~ t P m ) of > g a r t i n g ---~T--- a l b e r n i f l a r a i n exper~r:antdl p lo t s treated wi th soluble i r o n , copper, apmFchrar;irum tit great Sipl lewisset t : F:arrpk, Cape Cad (Gibt irr e t a l , 1580).

slnsed-sys tena dgr - lcu l ture of a s o r t , though h i t k i a lcsng tiirie2 135; sekarjae t ~ - t i ; x t n - & r r a ~ I S & S r:*uch a p a r t of' aolr present "imfe suppor t systen" a s s a l e hdy producl~on ever was. P ~ r h a p s sotrc*day, af ter a hundred years or sa of wors td ly id hdve a lnc~o~l id ted , hew Englanders nSay l u s k at, photosraphs o f

spray nunzles s tand ing i n the n~srrshes wi th the sacr fee l ings we have now i n looking a t t he o l d salt, hay s taddl es and gunda lows , Perhaps t h e y w i f l be as puzzled as X was when I cane fror:, t h e south and f i r s t saw the rucks sitting out i n the grass o f a Netd England marsh.

eta1 concentrations (ppm, oven dry weight) of Jive Spartina - and 2. patens (Gi bl in et a1 . 1980).

Treatwent Metal plot a S. alterniflora

b -

Fe C 2,500. 0Od 80. 0Od X F 1,700.00 65.00

P b C 26. OOd 25. OGd X F 20.00 21.00

Nn C 48. OOd 37.OOd X F 47.00 50.00

Z n C 31.00 30.00 X F 150.00 11O.OQ

- a~=control plot; XF = metal-containing sewage sludge plot.

'&an of four replicates (each replicate being a pool of three samples).

'Nean of four replicates (each replicate being a pool of six samples).

d~ and XF means are not significantly different at the (2.05 level.

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