Estuaries

Vol. 28, No. 2, p. 173-189

April 2005

Historical Ecology of a Central California Estuary: 150 Years of

Habitat Change
ERIC VAN DYKE # a n d

KERSTIN ~ae~ASSON

Elhho~ Slough .N~tion~l Estua~vine Research P~eserue, .l 700 Elhhorrt Road, Watsonville, Calitk:~d(~
95076
. ~ S T I L ~ C T : We investigatedthe h i s t o r i c a l e c o l o g y of FAkhoru Slough~ a 1~200 ha tidM wetland system in cenn'al California, T h e goal of ~his study was to identify patterns of change in the extent a n d distribution of wetland habRats during
a 150-yr p e r i o d and to investigate the causes of these changeso Using a geographic information system (GIS)~ we interp r e t e d historic maps~ charts, and aerial photographs, We created a series of s a m t n a r y m a p s to illnsdrate a~d quantR'y
changes in tidal flow a n 4 habitat types at six representative historicM perio4.,,. With the aid of custom s o f t . a r e tools, we
perforrued, semi-automated spatial anal.ysis of historic aerial p h o t o g r a p h s to quantify changes in mai~h cover at fixed
quadrat~ an4 tidal creek widdl at fixed cross sections, O u r mnlti.~cNe analysis 4 o e u m e n t s dramatic shifts in die distribution of habitat types resulting f r o m antbropogenic modifications to the hydrology of the slough, More than haft of
tile marsMands were dike4~ and. m o r e than two third.~ have either degraded, or been converted to other habitat types~
T h e constrnction of an artificial m o u t h abruptly t r a n s f o r m e d the wetland s}~tem f r o m depositional to higlfly erosional~
enlarging channeN~ widening creek.% a n d converting m a r s h to intertidal mudilat or o p e n water~ Increased tidal amplitude
and vdocity are the likely causes. In recent decades~ levee failure and intentional breaching have restored the a c r e a g e
u n d e r tidal influence to nearly historic levels, but recolonization of f o r m e r wetlands by salt marsh vegetation has been
minimM, Degraded f o r m e r marshland and nnvegetated mudflat are now the 4 o m i n a n t habitat types at Flkhorn Slough,
The rate of habitat char~ge remains high, suggesting that a new equilibrium may not be reached f o r many decades, This
s i n @ can help tidal wetlan4 managers identify p a t t e r n s an4 mechanisms of habitat change and set appropriate conservation and restoration goals,

Introduction

in tidal energy due to hydrologic manipulations,

increase in relative sea level due to land subsidence, and altered s e d i m e n t i n p u t levels due to
c h a n g i n g land use practices (Adam 2002). On a
global scale, eustatic sea-level rise, accelerated by
global[ climate change, can result in long-term estuarine habitat c h a n g e (Scavia et al. 2002).
T h e ecological history of m o s t of the world's tidal wetlands has n o t b e e n studied. At those estuaries
that have b e e n investigated, habitat changes have
often b e e n drmnatic. A widely r e p o r t e d e x a m p l e is
the rapid erosion of salt m a r s h at Venice L a g o o n
in n o r t h e a s t I t @ (Day et al. 1998). T h e r e d i r e c t i o n
of rivers that historically supplied s e d i m e n t s to the
estuary, in c o m b i n a t i o n with eustatic sea level rise,
subsidence resulting f r o m g r o u n d w a t e r withdrawal, a n d increasing tidal e n e r g y due to an enlarged
tidal prism, have caused the m a r s h edge to retreat
as m u c h as 2 m yr -4 (Day et al. 1998). T h e loss of
coastal wetland along the Louisiana Gulf Coast, a
region that includes a significant p e r c e n t a g e of
U.S. salt marsh acreage, :is a n o t h e r highly visible
example. R e d u c e d s e d i m e n t i n p u t due to levees
along the Mississippi River, the c o n s t r u c t i o n of extensive networks of canals, a n d subsidence rates as
high as 1.5 cm yr -4 have c o n t r i b u t e d to a r e p o r t e d
100 k m ~- yr-* loss of tidal wetlands, a l t h o u g h the

T I D A L "~ETLANDS &INTDt-{ABITAT Ct-bkNOE

Estuaries a n d coastal lagoons are a m o n g the

E a r t h ' s m o s t biologically productive ecosystems
a n d provide essential[ habitats for birds, :fish, crustaceans, a n d m a W o t h e r species (Little 2000)~ rFidal wetlm~ds are also s o m e of o u r m o s t ihighly altered landscapes, and their conservation lags beh i n d that of o t h e r terrestrial a n d m a r i n e systems
(Edgar et aL 9000). Rates of coastal wetland loss in
the United States resulting f r o m humm~ activities
e x c e e d e d 8,000 h a yr--* in r e c e n t decades a n d are
currently estimated to be 400 h a yr--* (NOAA 1990;
Dahl 2000).
Tidal wetlands are dynamic, r e s p o n d i n g to m a n y
ffpes of e n v i r o n m e n t a l changes, including h u m a n
activities. A p a r t f r o m direct losses due to construction a n d r e c l a m a t i o n , the principal a n t h r o p o g e n i c
forces driving tidal wetland habitat c h a n g e at the
local scale include diking, ditching, dredging, and
similar activities that alter tidal flooding regimes
a n d rnodii3r s e d i m e ~ t i n p u t and m a r s h accretion
rates (Kennish 2001). O n a regional scale, important causes of estuarine habitat c h a n g e are c h a n g e
* C o r r e s p o n d i n g author; tele: 831/728-2822; fax: 831/7281056; e-mail: vandyke@elkhornsloughrorg
9 2005 Estuarine Research Federation

173

174

E. Van Dyke and K. Wassen

relative i m p o r t a n c e of each factor remains controversial (Boesch et al. 1994; T u r n e r 1997; Day et al.
2000). At Chesapeake Bay, the largest estuary in
the U.S., significant coastal wetland losses are attributed to a combination of subsidence and sedim e n t deficit (Stevenson et al. 1985; Kearney et al.

1988~.
~[&BITs

GHANGE AT (--~LIFORNIA ESTUARIES

Vgetlands are relatNely small and i n f r e q u e n t

along California's topographically complex and
seismically active coastline, occurring mainly within occasional p r o t e c t e d estuaries and lagoons (Eram e t t e t al. 2000). "While d e c r e ~ e d riverine sedim e n t inputs and relative sea level rise are major
drivers of estuarine habitat change elsewhere,
these thctors are likely- less i m p o r t a n t in California
where drainage basins are relatively small and uplift typically outpaces sea level rise (Patrick and
D e L a u n e 1990; E m m e t t et al. 2000). Subsidence
resulting from groundwater extraction has b e e n a
significant factor at some sites, particularly in the
Sau Francisco Bay area (Patrick and D e L a u n e
1990). Subsidence events resulting from seismic actMty may also be regionally i m p o r t a n t along the
tectonically active California coast, as they are m
the Pacific Northwest (Atwater et al..[977; Atwater
1987). T h e p r e d o m i n a n t cause of tidal wetland
habitat change in California has been direct human alteration (Larson 2001).
As m u c h as 91% of California's coastal wetlands
(2 million ha.) were lost during the 150 yr following
statehood and settlement by E u r o p e a n Americans,
arid nearly all that remain are altered or degraded
(Larson 2001). Diking, draining, dredging, and fi11ing for residential, commercial, and agricultural
d e v e l o p m e n t have eliminated about 85% of tidal
wetlands in the San Diego region (Zedler 1996a)
and at least 78% in the San Francisco Bay area
(Nichols et al. 1986; Goals Project 1999).
Because conversion of tideland to agriculture or
salt evaporation p o n d s typically does not involve
filling and is potentially- reversible, these f o r m e r
wetlands provide i m p o r t a n t candidate sites for habitat restoration (Goals Project 1999). Tidal flow has
recently b e e n restored t h r o u g h either accidental
or intentional breaching of levees at several of California's drained f o r m e r wetlauds (Zedler 1996b;
"Williarns and Faber 2001; Williams and O r r 2002)
and additional projects are p l a n n e d (Steere and
Schaefer 2001).

ing the past 150 yr, a p e r i o d of major modifications

to the landscape. Historical ecolog3~ is a relatively
new b r a n c h of the environmental sciences that integrates historic sources to analyze and characte>
ize past changes in natural communities (Swetnam
et al, 1999). D e v e l o p m e n t of a historical perspective is f u n d a m e n t a l to efforts toward conservation
and restoration of estuarine ecosystems (Goals Proj e c t 1999)..Although predicting future conditions
t h r o u g h simple extrapolation of past trends can be
risky, knowledge of past conditions may suggest hypotheses that can be tested with c o n t e m p o r a r y
data and can supply the parameters for retroactive
testing of predictive models (Swetnam et al. i[999).
Although it may ,lot be meaningful to define ideal
r e f e r e n c e conditions based on o n e fixed point m
time, historical ecological studies can identify the
spatial and temporal r a a g e of variability in naturally dynamic systems and assist in setting ecologically justifiable, achievable, m~d sustainable mana g e m e n t and restoration goals (Swetnam et al.
1999). Problems associated with analyzing the hi>
torical record include the f i a g m e n t a r y nature of
individual source materials as well as the subjectiv~
ity- i n h e r e n t in the interpretation process (Swetn a m e t al. 1999; Grossinger 2000). Historical materials have b e e n applied to ecological analyses at
several West Coast estuaries (e.g., Berquist 1978;
Niemi and Hall 1996; Goals Project 1999; B o r d e et
al. 2003; Foxgrover et al. 2004).
A key goal of this project was to develop accurate, repeatable m e t h o d s and tools for p e r f o r m i n g
habitat classification and long-term change analysis
from historic maps and aerial photographs. We
employed a dual approach, combining broadscale
manual interpretation of the entire wetland ~'stem
with h i g h e r resolution, semi-automated analysis of
replicated fixed quadrats and cross sections. T h e
f o r m e r m e t h o d resulted in estuary-wide habitat
classification maps for several representative periods tlhat were subjected to quantitative analysis.
'The latter yielded detailed habitat data comparable to that obtained flora long-term field studies
of m o n i t o r i n g plots and transects. These two geospatial techniques c o m p l e m e n t e d each o t h e r and
provided i n d e p e n d e n t evidence for the observed
trends. O u r rigorous, multiscale a p p r o a c h to identifying and quantiiymg estuarine habitat change is
applicable to historical ecological studies of o t h e r
t h r e a t e n e d ecosystems.
Method~

I~tISTORIGAL ECOLOGY AND ]-[ABITAT

CHANGE ANALYSIS

STUDY SITE

T h e purpose of' this study was to d o c u m e n t eaL~

iier habitat conditions and to quantif}- trends of
wetland change at a central California estuary dur-

Elkhorn Slough is a 1,200 ha tidal wetland syste:m adjoining M o n t e r e y Bay in central Califbrnia
(Fig. 1). T h e climate is m e d i t e r r a n e a n , xMth m e a n

Elkhorn Slough Historical Ecology

'I 7 5

E l k h o r n Slough's extensive intertidal m u d f l a t s

are i n h a b i t e d by a diversity of invertebrates and are
heavily used by birds for foraging and b r e e d i n g
(Harvey and C o n n o r s 2002; Wasson et al. 2002).
T h e slough's d e e p e r c h a n n e l s serve as nurseries
for n u m e r o u s species of fish and as feeding or refuge areas for two m a r i n e m a m m ~ species (Yoklavich et al. 1991; B a r r y et al. 1996).
Nearly 700 h a of E l k h o r n Slougtl's wetlands are
m a n a g e d for xa~ldlife arid conservation p u r p o s e s by
the California D e p a r t m e n t of Fish and G a m e . Elkh o r n Slough National Estuarine Research Reserve
e n c o m p a s s e s 567 h a of the slough's wetlands and
adjacent uplands~ O t h e r land uses in the vicinity
of E l k h o r n Slough include cultivation of strawberries and o t h e r crops as well as rural residential and
industrial d e v e l o p m e n t (Silberstein et aL 2002).
J)~'LKHOR_N SLOUGH ]3NWIRONMENTAL ].[ISTORY

121 ~ 4 g W

121~ 42' W

Fig. 1. Location of Elkhorn Slough and the SalLnas River

syst~-m adjoimng Monterey Bay in cenkra.1 California. Numbers
refer to mm'sh quadrat and tidal creek cross seetic,n regions
(regions 1 and 2 comprise die lower ,slough, region ,g the n:iid
slough, and regions 4 and 5 the upper slough).

m o n t h l y t e m p e r a t u r e s r a n g i n g f r o m 11.1~ in the
winter to 15~4~ in the s u m m e r a n d m e a n a n n u a l
rainfall of b5~2 cm [allmg mainly in the winter
m o n t h s (Caffrey 2002). Tides are sen:lidiurnal with
a m e a n diurnal r a n g e of 1.7 rn (Caffrey a n d Broenkow 2002).
T h e d o m i n a n t vegetation in E l k h o r n Slough's
m a r s h l a n d s is Sa~icor~ia virg~r~ic~, which almost exclusively d o m i n a t e s the intertidal z o n e b e t w e e n app r o x i m a t e l y 0.4 a n d 1.2 m above m e a n sea level
(Atwater and H e d e l 1976; M a c D o n a l d 1988). Several additional species, including .Dist~chtis @icata,
Ja~raea carnosa, b~anM.r~a, salina, a n d At~zp~x spp~,
are also p r e s e n t at the u p p e r intertidal or infratidal
zones. Sci'rp~xs and Tyi+ha species are c o m m o n m
b r a c k i s h w a t e r l o c a t i o n s . 2partina jbt~osa, w h i c h
dorninates the lower intertidal z o n e at m o s t California salt n~larshes, is conspicuously absent f r o m
E l k h o r n Slough and n e a r b y marshes, as are nonnatwe c o n g e n e r s ( Z i m m e r m a n and Caffrey 2002).
I n t r o d u c e d terrestriN plants, in eluding CSn~t~m.ma
c~alatu,m a n d C~,~obroms ed~llia, are locally a b u n d a n t
and invading the m a r s h f r o m adjacent u p l a n d s
('~Vasson u n p u b l i s h e d data)~

Rising sea levels d r o w n e d a coastal valley- app r o x i m a t e l y 10,000 years ago, converting it to a tidal e m b a y m e n t . InitiaUy a high-energy m a r i n e s3,stern, several t h o u s a n d years of s e d i m e n t deposition
and m a r s h accretion gradually t r a n s f o r m e d Elkh o r n Slough into a low-energy estuary. Broad expanses of Sdicor~ia developed, flanking a network
of tidal channels (Schwartz et al. 1986). T h e slough
r e m a i n e d largely a saltwater system due to the absence of m a j o r riverine inputs, a l t h o u g h s e d i m e n t
cores r e c o r d intervals of localized freshwater dominance ( H o r n b e r g e r 1991;.Jones 2002), likely corr e s p o n d i n g to episodes of increased flow or changes in the course of the n e a r b y Salinas River sTsteni.
Many areas of transitional brackish, freshwater, and
riparian habitat d e v e l o p e d n e a r occasional seeps
and springs and at tlhe slongh's u p p e r reaches.
Native Aniericans lived m the vicinity of the
slough for p e r h a p s 10,000 yr (Dietz et aL 1988;
.Jones a n d J o n e s 1992). Reports f>om early explorations indicate that intentional b u r n i n g o c c u r r e d
during this time, yet sedinient cores do not suggest
that this practice resulted in significant erosion
( G o r d o n 1996). Before the mid 19th century, the
p r e d o m i n a n t land use by E u r o p e a n i m m i g r a n t s in
n o r t h M o n t e r e y County, was cattle grazing, which
also a p p a r e n t l y h a d a m i n i m N effect on the slough
(King 1981; G o r d o n 1996). T h e era of m a j o r ant h r o p o g e n i c wetland changes b e g a n shortly after
the Gold Rush and California's statehood. T h e earliest m a p s included in this study were p r o d u c e d
during this period.
With the arrival of A m e r i c a n s during the latter
half' of the n i n e t e e n t h century, large areas of woodland and scrub were cleared for fuel wood and for
the cultivation of hay and barley ( G o r d o n 1996).
O n the u p l a n d sarldhills ad:jacent to E l k h o r n
Slough, the thin topsoil eroded, depositing large

E. Van Dyke and K, Waaaon

176

TABLE 1. Historic maps and chea-ts.

]3 <,v:

D escripti0 n

1853

General Map of Explorations and Surveys in CaKfornia

1854

Part of the Coast of Ca[. frorn Pajaro River Southward topographic sheet (T473)
Map of the \,icinity of Monterey Bay
Monterey Bay hychographic cheat (H5498)
Rand~o Boise. de San Cayetmlo plat
Rancho Cameros plat
Township 13 Range 2E plat
Rancho Bolas Nueva y Moro Cojo plat
Southern Pacific Railroad Pajaro Branch/Elkhorn Slough
Map of Tm-npike Road fIorfl CasCroville to "Ware House" on
the Elkhorn Slough
'l~pogra.phical Map of Central California Together with a Part
of Newda
Map of the Count}, of Monterey
Map of Moss, Salines, ea~d Watsonville Landings Belonging t o
the Pacific Coast Steamship Company
Map of Watsonville Landing Belonging to the Pacific Coast
Steamship Company
Point Buchon to Point Pinos hydrographic chart (H5400)
Official Map of Monterey Cuunty
Lower Salinas ~v'alleysoil survey
Map of Monterey Com~ey
Moss Lan4ing and its Vicinity Contiguous m Monterey Bay
Monterey Bab Pajea'o Ri,~er Southward topographic sheet
(T473a)
Monterey Be) hydrographic cheat (H5403)
Laxlds of the Empire Gun Club
Tunapike Road Between lIudson Landing Bridge a n d j , Henry
Meyer Gate.
Capitola topographic quadrangle
San Juan gautista topographic quadrangle
Salines ALes soft survey

1855
1857
1859
1859
1867
1872
1872
1873
1873
1877
1885
1885
1893
1898
1901
1908
1909
1910
1911
1913
1913
1914
1917
1925

Ori~.m

............................................................................................

a m o u n t s of sediment into the wetlands (King 1981;

Silberstein et el. 2002). Maps from the 19th century show a broad, deepwater basin at the m o u t h
of the slough between Moss Landing aud the old
Salines River m o u t h . Steamboats provided regular
service all the way to H u d s o n Landing (Fabing and
H a m m a n 1985), and maps show the broad mare
channel continuing for a n o t h e r 4 kin, well b e y o n d
the present head of the slough. By the 1880s, sedi m e n t deposition had m a d e the ciqannel too shallow to p e r m i t navigation by steamers (Van Dyke
unpublished data). Intertidal mudflats and shoals
in the lower slough likely first a p p e a r e d during this
period, T h e channel above H u d s o n Landing rapidly filled and converted to marsh (Va.n Dyke unpublished data,), By the time the first aerial photographs used in this study were taken in the early
19S0s, Elkhorn Slough was a sluggish lagoon with
limited tidal exchange for m u c h of the year due
to a persistent sandbar at its m o u t h (MacGinitie
19~%). Episodes of increased sediment deposition
during the same period at o t h e r coastal California
locations have also b e e n attributed to a n t h r o p o genie disturbance (e.g,, gerquist 1978; Nichols et

U.S. War DeparUnent/Pacific Railroad

Surveys
U.S. Coast Survey
W, R glake/U.S. Coast Survey
U,S. Coast Survey
U.S. Sta'veyor General
U,S Staveyor General
U.S. Sta-veyor General
U,S Sta'veyor General
Southern Padfic Railroad
Ltnl~Jo~n

Califbrnia Geological Survey

St. John Cox
J. H. Gerber, surveyor

j. H, Gerber. surveyor
U,S. Coast mid Geodetic Survey
Lou C. Hare, Monterey Comity surveyor
U.S. Department of AgrieuJmre
Lou C, Hare, Monterey Colmty surveyor
Lou G. Hare, Monterey Cotmtv sm'veyor
US. Coast and Geodetic Smwey
US. Coast and Geodetic Sur~,t-y
~-M~noldM. Baldvdn, licensed surveyor
Lou C. Hare, Monterey County, surveyor
U.S. Geological Survey
U.S. Geological Survey
US. Department of Agrict~ttae
........................................................

a.1, 1986; Niemi and Hall 1996; Cole and Wahl

2000).
HISTORIC MAPS AND AERIAL Pi~tOTOGRAPHS
Because of' Elkhorn Slough's coastal location
near the historic city of' Monterey, a wealth of materials was available for analysis, ~,~e~ obtained, converted to digital format, georectified, rnosaiced,
and i n t e r p r e t e d 26 historic maps and charts dating
from 1858 to 1925, and 18 aerial p h o t o g r a p h
:flights taken between 1981 and 2003 comprising
more than. $00 individual photos, Table 1 lists the
historic maps and Table 2 lists the aerial photographs used in this study,
w e scanned aerial photographs at resolutions selected to yield pixels of" approximately 0~6 m after
:rectification~ Mosaics were assembled by extracting
only the least-distorted effective area from the
overlapping p h o t o g r a p h s of each flight, Effective
areas were identified using the proximity-function
of the ArcView Geographic Information System
(GIS) Spatial Analyst extension (ESRI, Redlands,
California). To m i n i m i z e distortion, individual
photographs were resampled using the plane pro-

Elkhorn Slough Historical Ecology

'I TT

TABLE 2. Historic aerial photographs.

Date
'i~yp e
..................................................................................

Co uni:

Scale

]May 1931

panchromatic

24

November 1 9 3 7

panchromatic

17

August 1 9 4 9

panchromatic

16

May-Jmle 1 9 5 6

panchromatic

14

May-J~ly 1966

panchromatic

15

May 1971

panchromatic

14

April 1976

panc~rornatic

28

April 1980

color infrared

28

April 1987

color infrared

21

May 1989

trne color

40

l :19,500
0.65 rn pixel '~
1:20,800
0.66 m pLxel-:
1:21,100
O.66 m pixel-i
1:24,500
0.6 m pixel<
1:20,800
0.67 m pixel-~
1:24,300
0.67 m pixel-~
1:9600
0.4 m pixel-~
1:12,44)0
0 . 5 2 iTi pixel-:
1:12,100
0.51 re, pixel-:
0.4 m pixel

May 1992

color infrared

19

December 1 9 9 9

Aplql 2000

panchrc)matie
digital
ortho
n'ae color digital

1:12,400
0.53 re* pixel--:
0.6 m ptxd

40

0.45 m ptxel i

May 2001

color mtiared digital

0.6 m pixel -'

Cafffbrnia Department of Fish and Garne >Mr Services

Coung~"of Monterey/HjW GeoSpatial, 1no.

0.~ m ptxei

California Deparnnent of Fish and Game Air So>

Orlsm

Western GuK CA1Co./FaiK.b~d Aerial Surveys, Inc.

U.S. Depm-tment of Agricaltm-e/Faff'chJldAerial
Sta'w-ys, Inc.
U.S. Department of Agricuimre/Pm'k Aerial Su.rre}s, huc.
U.S. Department of AgTkUltuce/Aero Settee

CorlJ.
U.S. Department of Agrieultm-e/Cara*TightAerial
8ta"w-y_%Inc.
U.S. Deparnnent ofAgriculture/W%stern Aerial
Coil tt-a ctol'S

Califbrnia Department of '~Dansportation

California Coastal Colmnis~ion/WesternAerial Photographs, Inc.
Moss Landing Marine Labs./Western Aerial Photographs, Ix,c.
California Department of Fish m~d Game Air Services
Elkhorn Slough Foundafion/Aaial Data Systems
Courtw of Monterey/HjW GeoSpatial, -Inc.

~AI'ttlO

April 2003

true cc,lor digital

44)

...........................................................................................

j e c t i v e m o d e l to g r o u n d c o n u o l p o i n t s selected
n e a r the p e r i m e t e r of e a c h i d e n t i f i e d effective
area~ G r o u n d c o n t r o l p o i n t s were o b t a i n e d f r o m
r e c e n t 0.6 m p i x e l i digital o r t h o p h o t o g r a p h s ,
R e c t i f i c a t i o n a n d r e s a m p l i n g was p e r f o r m e d u s i n g
T N T Mips (MicroImages, L i n c o l n , N e b r a s k a ) . Mosaics were t h e n a s s e m b l e d f r o m a c u t - l i n e t e m p l a t e
u s i n g T N T Mips,
A variety of factors may- have c o n t r i b u t e d e r r o r
to o u r digitally p r o c e s s e d a e r i N p h o t o g r a p h m o saics. Aircraft tilt, t e r r a i n relief, a n d c a m e r a geo m e t r y are p o t e n t i a l s o u r c e s of d i s t o r t i o n o n the
original photography, and contact printing and
scanning introduce additional distortion (Moore
2000). T h e p r o c e s s of m a t c h i n g g r o u n d c o n t r o l
p o i n t s d u r i n g r e c t i f i c a t i o n is also a s o u r c e of e r r o n
We p e r f o r m e d a spatial a c c u r a c y a s s e s s m e n t by
r a n d o m l y s e l e c t i n g 20 p o i n t s w i t h i n t h e study a r e a
and then locating identifiable features near each
p o i n t o n e a c h p h o t o mosaic. T h e s e l o c a t i o n s were
c o m p a r e d with t h e i r c o r r e s p o n d i n g l o c a t i o n s o n
the digital o r t h o p h o t o s . A l t h o u g h r e s i d u a l s r e p o r t ed d u r i n g r e c t i f i c a t i o n were c o n s i s t e n t l y less t h a n
twice the r e s a m p l e d p i x e l size, o u r a c c u r a c y assessm e n t i d e n t i f i e d a m e a n e r r o r of 4.8 m for the l g
aerial p h o t o mosaics. P o s i t i o n a l e r r o r o n the base
o r t h o p h o t o s , w h i c h w o u l d n o t b e i d e n t i f i e d i n this
e r r o r a s s e s s m e n t , was p r e s u m e d to b e m i n o r .

vices
................................................

Historic m a p s a n d charts were s c a n n e d at vario u s r e s o l u t i o n s a c c o r d i n g to the q u a l i t y of' the image a n d t h e n g e o r e c t i f i e d to g r o u n d c o n t r o l p o i n t s

s e l e c t e d f r o m d i g i t a l U.S. G e o l o g i c a l S u r v e y
(USGS) t o p o g r a p h i c q u a d r a n g l e s u s i n g a first-order p o l y n o m i ~ model. Rectification and resamp i i n g was p e r f o r m e d i n A r c V i e w with t h e I m a g e
Analysis e x t e n s i o n . B e c a u s e the r e l i a b i l i w of historic r n a p s varies a n d is g e n e r a l l y u n k n o w n , we
o v e r l a i d all available m a p s f r o m e a c h t i m e p e r i o d
to p r o d u c e a c o m p o s i t e i n t e r p r e t a t i o n ( G r o s s m g e r
2000). ~ a s s u m e d a level of spatial a n d r e p r e s e n t a t i o n a l a c c u r a W ['or e a c h s o u r c e a c c o r d i n g to o u r
u n d e r s t a n d i n g of the m a p ' s i n t e n d e d p u r p o s e . F o r
e x a m p l e , early U S G S m a p s w e r e p r e s u m e d to b e
h i g h l y a c c u r a t e for r e p r e s e n t i n g t o p o g r a p h y , b u t
less r e l i a b l e for d i s t i n g u i s h i n g w e t l a n d h a b i t a t
t},pes.
HABH;AT AND TIDAL IFLOW ~r
We i n t e r p r e t e d h i s t o r i c m a p s a n d aerial p h o t o graphs and p r o d u c e d a spadNly accurate chronology of six L e p r e s e n t a t i v e years (1870, 1913, 1931,
1956, 1980, 2000) to c h a r a c t e r i z e t h e overall seq u e n c e of w e t l a n d c h a n g e s at E l k h o r n S l o u g h . T h e
s t u d y a r e a wm~ l i m i t e d to E [ k h o r n S l o u g h ; t r i b u t a r y
w e t l a n d s at C a r n e r o s C r e e k to the east a n d M o r o
Cojo S l o u g h a n d t h e old S a l i n a s River c h a n n e l to

178

E. Van Dyke and K. Was~on

TABLE 3. Habitat classificat[on system.

!~4.1sitattype
Saltamter channel
Seagrass b e d
Restricted salt~'ater channel
M u d or degraded salt marsh (<25% vegeeation cover)
Resn-icted m u d or degraded salt m a r s h ( < 2 5 % vegetation cover)
Diked m u d or deg'm.ded salt iriarsh ( < 2 5 % -~ge~ation cover)
Degraded salt r n m s h or m u d ( 2 ~ 7 5 % vegetatic~n cover)
Restricted degraded salt rua~sh or m u d (25-75% ,,egetation co,,er)
Diked degraded salt m a r s h or 133ud (25-75% vegetation cover)
S/it m a r s h ( > 7 5 % vegetatioi~ cow-r)
Resnqcted s i t m e a ~ ( > 7 5 % vegetation cover)
Diked salt inarsh ( > 7 5 % vegetation cce~-r)
R e d a b n e d f o r m e r tidal wetbarld
Brackish&'esh m a x ~ h o r c h a n n e l
Restricted brac~sh-fresh mm-sh or c h a n n e l (behind tidegate)
I m p o u n d e d b~ackish-~esh m a r s h o r channel (beLmd Ievee)
R i p a r ~ wooc[land

the south were excluded. I n t e r p r e t a t i o n :['or the

first two periods was based on maps and is less detailed than the four s u b s e q u e n t periods where aerial p h o t o g r a p h s were used. We developed a pair
of GIS layers for each time period: habitats (digitized polygons delineating generalized land use
and land cover cla~sses) and tidal flow (digitized
polygons delineating areas with either unrestricted, restricted, or excluded tidal flow). T h e complete classification s c h e m e is listed in Table 3, A
consistent scale of 1:1,200 was m a i n t a i n e d during
digitizing for consistency. Digitizing was p e r f o r m e d
using ArcView GIS.
For the habitats layers, we selected a set of land
use and land cover classes based on what was indicated on historic maps or clearly distinctive on
the o l d e s t p a n c h r o m a t i c aerial p h o t o g r a p h s .
Marsh habitat indicated on maps was assumed to
represent areas with vegetation cover > 75%. \~rith
aerial p h o t o g r a p h s , marsh habitat was classified by
estimating the p e r c e n t a g e of vegetation cover within. a 100 m square grid at our standard 1:1,200
scale. No distinction was m a d e between sparsely
vegetated marsh ( < 2 5 % cover) and unvegetated
p a n n e or mudflat because the}, are visually indist i n g u i s h a b l e at this scale. B r a c k i s h - f r e s h w a t e r
marsh was distinguished ['rom salt marsh by- its
greater textural variance (SaSco~rda m a r s h is uniformly gray on parmhromatic aerial p h o t o g r a p h s )
and an absence of tidal channels~ No attempt was
m a d e to differentiate subtidal from intertidal areas
as tidal heights varied between aerial p h o t o g r a p h
series. T h e main c h a n n e l b o u n d a r y was determ i n e d either by- tlhe water line (maps and p h o t o graphs at higher tides) or by a visible line on the
mudflats at approximately meat, high water (photographs at lower tides)~
For the tidal flow layers, we b e g a n by digitizing

Tq_-abitatc!as~

Tidal flow

saltwater
salt'water
salb,~'ater
mud
mud
mud
salt m a r s h
salt m a r s h
salt m a r s h
salt mal~h
salt m a r s h
salt marsl,
reded
Desh m a r s h
flesh m a r s h

u.nresttJcLed
tmrestdcted
restricmd
m~resU:fcted
restricmd
no.ntidal
unresn'icted
resta'icted
n o n tidal
m~restdcted
restricted
no.ntidal
nontidal
nontidal
nontidal

fresh ilia~sh

nondd~

ripmian

nontidal

lines represen ring either intact or b r e a c h e d levees.

Levees were interpreted as b r e a c h e d if an aerial
p h o t o g r a p h showed evidence of tidal channels
flowing t h r o u g h or a r o u n d the levee. We also located road arid railroad e m b a n k m e n t s and tidegates. We then digitized polygons to delimit areas
with unrestricted tidal exchange (e.g., u n d i k e d ) ,
areas with r e s t r i c t e d tidal flow (e.g., b e h i n d
b r e a c h e d levees and tidegates), and nontidal areas
(e.g,, b e h i n d intact levees). Areas with brackishfreshwater habitat b e h i n d intact levees or gates
were classified as i m p o u n d e d ; a r e ~ without wetland vegetation belhind intact levees or gates were
interpreted as reclaimed.
To assess the accura W of our interpretation of
habitats and tidal flow from aerial p h o t o g r a p h s , we
visited the 20 locations that we r a n d o m l y selected
for spatial a c c u r s tT assessment and c o m p a r e d
g r o u n d truth observations wffh our interpretation
of the 2000 aerials. In all but one case, field observation and p h o t o interpretation, m a t c h e d exactly~ T h e single disagreement involved p e r c e n t cover
in an. area of deteriorating marsh, and may represent a c h a n g e between the 2000 p h o t o date and
the 2004 field visit~ In m~y case, because we could
n o t p e r f o r m similar accuracy assessment with ea>
lier aerial p h o t o g r a p h s and maps, we did n o t modify, our interpretation as a result of this g r o u n d truthing.
!V~c.RSH AND '['IDAL C R E E K T I M E SERIES

We p e r f o r m e d detailed quantitative analysis of

fixed quadrats using aerial p h o t o g r a p h s take~ at
12 different dates between 1931 and 2003, and of
tidal creek cross sections using aerials taken at 13
dates betwee:n 1981 and 2003, Five regions of Elkh o r n Slough's tidal wetlands were studied, encore-

Elkhorn Slough HistoricN Ecology

passing all areas that have r e m a i n e d u n d i k e d

t h r o u g h o u t the 7 2 5 r study p e r i o d (Fig. 1).
To quantify c h a n g e s m vegetated m a r s h cover,
we divided the study a r e a into 196 100 X 100 m
quadrats~ Within each q u a d r a t (and each date), we
d e t e r m i n e d the p r o p o r t i o n of salt m a r s h vegetation versus u n v e g e t a t e d habitat ( m u d and water).
We d e v e l o p e d a custom, interactive ArcView Spatial Analyst application to p e r f o r m s e m i - a u t o m a t e d
i m a g e interpretation. This tool allowed us to rapidly d e t e r m i n e the precise grayscale value that
would trace isoline b o u n d a r i e s between vegetated
a n d u n v e g e t a t e d portions of each q u a d r a t and automatically p r o d u c e the c o r r e s p o n d i n g set of polygons in an ArcView shapefile. Grayscale values
were adjusted for every q u a d r a t to a c c o u n t for contrust variations within and b e t w e e n p h o t o g r a p h s .
For i n t e r p r e t a t i o n consistency, all analysis was p e >
f o r m e d on grayscale imagery; for years wt,ere color
or color infrared i m a g e r y was available, we perf o r m e d red-green-blue to hue-saturation-intensity
conversion (Jonson 1996) and i n t e r p r e t e d only the
intensi~ component.
Tidal creek changes were quantified by measuring the width of 19(; cross sections within the
m a r s h d r a i n a g e network. T h e s e s a m p l i n g locations
were chosen to include every m a j o r s e g m e n t of every creek that was visible on the 19S1 p h o t o g r a p h s .
Cross section width was defined as the distartce between vegetated creek banks and was n.easured at
each p o i n t (and each date) with the aid of a custom, interactive ArcView GIS script. In s o m e areas,
m a r s h l a n d has completely converted to unvegetated m u d f l a t in r e c e n t years. In these cases, cross
section width was defined as the distance to the
nearest r e m a i n i n g recognizable f r a g m e n t of m a r s h
v e g e t a t i o n in the m o s t r e c e n t p h o t o g r a p h where
such r e m n a n t s were still visible.
To d e t e r m i n e the significance of changes in salt
marsh cover, we p e r f o r m e d r e p e a t e d m e a s u r e s
analysis of variance (ANO~v\A) on m e a n p e r c e n t a g e
of vegetated area within the quadrats with year as
within-sample factor a n d region as between-sample
fiacton Covet- p e r c e n t a g e s were arcsin t r a n s f o r m e d
to m e e t the a s s u m p t i o n s of .~N'O\(~ (Sokal a n d
q
R o h l f 1~9a).
To test the significance of changes m
tidal creek cross section widths, we p e r f b r m e d rep e a t e d m e a s u r e s A N O V A o n log t r a n s f o r m e d
creek width with year as within-sample factor a n d
region as between-sample factor, To d e t e r m i n e the
significance of changes b e t w e e n pairs of years, we
p e r f o r m e d Fisher's p r o t e c t e d least significant diff e r e n c e (PLSD) post-hoc c o m p a r i s o n (p < 0.05)~
Statistical analysis was p e r f o r m e d with StatView
software (SAS Institute Inc., Cary, N o r t h Carolina),

'179

ResMts

HABITAT AND TIDAL FLOW .%~PPINO

E l k h o r n Slough has u n d e r g o n e dramatic changes in the extent and distribution of wetland habitat
types during the past 150 yr. T h e s e changes are
illustrated t0y six pairs of thematic m a p s (Fig. 2)
a n d s m n m a r i z e d by a c o r r e s p o n d i n g pair of charts
(Fig. 3). Two m a j o r trends are apparent: an initial
decrease and s u b s e q u e n t recovery of total acreage
u n d e r tidal influence and a c o n t i n u i n g decrease of
salt m a r s h acreage~ Since 1870, m o r e than two
thirds of the slough's salt m a r s h has either degraded or c o n v e r t e d to o t h e r habitat types. T h e m a p >
ity of this loss o c c u r r e d during the m i d d l e third of
the study period, c o n c u r r e n t with a period of extensive diking that either restricted or completely
excluded tidal e x c h a n g e f r o m m o r e than half of'
the slough's wetlands. Marsh loss has slowed somewhat during the Final decades of the study, conc u r r e n t with an era of b r e a c h i n g levees and restoration of tidal flow to f o r n . e r wetlands, a l t h o u g h
the extent of d e g r a d e d f o r m e r m a r s h a n d unvegetated m u d f l a t continues to i n c r e ~ e . T h e total
wetland area decreased slightly tlhrough the study
period due to the conversion, of m a r s h to u p l a n d
vegetation.
During the period between 1870 a n d 1956, m o r e
than 60 k m of levees a n d e m b a n k m e n t s were constructed, r e d u c i n g tlhe r a n g e of u n o b s t r u c t e d tidal
influence by 59%. D u r i n g the s a m e period, the extent of' intact salt m a r s h habitat (vegetation cover
> 75%) decreased by 66%. Within this era of extensive diking and salt m a r s h loss, the acreage of
fbur habitat types increased. Between a b o u t 1900
a n d 191,8, m o r e tha.a 90 h a of salt m a r s h was converted to fresh and brackish habitats t h r o u g h the
i m p o u n d m e n t of freshwater within created p o n d s
a n d marshes, a l t h o u g h tlhe extent of these habitat
types eventually decreased after m a W of' the levees
were a b a n d o n e d . Between a b o u t 1900 and 1956,
97 h a of the slough's salt m a r s h converted to unvegetated m u d f l a t (vegetation cover < 25%), primarily within a c o m p l e x of diked salt evaporation
ponds, and a n o t h e r 95 h a c o n v e r t e d to d e g r a d e d
.marsh (vegetation cover < 75%). Between 1931
a n d 1956, 275 ha, m o r e than 30% of E l k h o r n
Slough's :remaining salt marsh, were d r a i n e d a n d
r e c l a i m e d for agricultural use.
A f t e r 1956, the p a t t e r n o f tidal r e s t r i c t i o n
abruptly reversed as a result of accidental a n d intentional levee breaches~ As little as 4 km of levees
r e m a i n e d intact by 2000, and the acreage u n d e r
tidal influence (with either unrestricted or partially
:restricted flow b e h i n d culverts, t i d e g a t e s , or
b r e a c h e d levees) slightly e x c e e d e d what it ihad
b e e n in 1913. Despite this reversal, salt m a r s h c o n -

E. V~n

Dyke

and

K. Wasson

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Elkhorn Slough Historical Ecology

~ Riparian
Habitats

15oo

Woodland
Fresh I Brackish Marsh
Reclaimed Wetland
Salt Marsh (> 75% cover)
Salt Marsh / Mud (25% - 75% cover)
Mud f SaR Marsh (< 25% cover)
Saltwater Channel

100

80
%

6O

40
O region4 i

12oo

20

9oo

Ha

t930

600

'181

O,
"%

v region 5 J
,

1950

1970

1990

F v

2010

Fig. 4. Mean pereenrage of salt marsh vegetation cover vdthin 196 qua~a~s m 5 regions, 1931-2003. Line is mean for all
regions. Error bars represent 1 stmldard error.

3O0
0
1875

1925

1900

Tidal Flow

1950

1975

2000

Nondidat
ReslrfctedFlow

1500

UnrestrictedFlow

1200
900
Ha
600
300
0

'_

.,

I875

1900

1925

1950

1975

2000

Fig. 3. Habitat and tidal flow change summary, 1870-52000.

t i n u e d to d e t e r i o r a t e . D u r i n g tlhe p e r i o d b e t w e e n
1980 a n d 2000, 45 h a of f o r m e r l y v e g e t a t e d m a r s h
c o n v e r t e d to m u d f l a t a n d s h a l l o w w a t e r at the
s l o u g h ' s u p p e r east side, a r e g i o n t h a t n e v e r exp e r i e n c e d e x t e n s i v e diking. By 2000, f46% of Elkh o r n S l o u g h ' s tidal w e t l a n d s h a d c o n v e r t e d to u n v e g e t a t e d m u d f l a t a n d a n a d d i t i o n a l 21% c o n v e r t ed to d e g r a d e d m m s h , a h a b i t a t c a t e g o r y that was
n o t p r e s e n t p r i o r to i[913, T h e e x t e u t of h i g h - q u a l it?" salt m a r s h i n 2000 was 207 ha, 23% of w h a t it
was a c e n t u r y earlier.
MARSH AND TIDAL CREEK TIME SERIES
V e g e t a t i o n cover across E 1 k h o r n S l o u g h ' s m a r s h es has d e c r e a s e d d r a m a t i c a l l y s i n c e 19,31, a n d this

t r e n d a c c e l e r a t e d d u r i n g the final d e c a d e s of o u r
study. C l e a r d i f f e r e n c e s are a p p a r e n t b e t w e e a the
lower a n d m i d s l o u g h ( r e g i o n s 1, 2, a n d S) a n d
the u p p e r s l o u g h ( r e g i o n s 4 a n d 5). M u c h of the
u p p e r s l o u g h , w h i c h was o n c e d e n s e l y v e g e t a t e d ,
is n o w c o m p l e t e l y u n v e g e t a t e d , T i d a l c r e e k w i d t h
has also i n c r e a s e d s i n c e 19,81 a n d e x h i b i t s a s i m i l a r
rate of acceleration~ I n the u p p e r s l o u g h , m a n y form e r creeks a n d p a n n e s have c o m p l e t e l y d e g e n e r ated i n t o o p e n m u d f l a t , T h e s e r e s u l t s c o n f i r m the
t r e n d of salt m a r s h d e g r a d a t i o n a n d loss t h a t was
a p p a r e n t [~:om o u r b r o a d s c a l e h a b i t a t m a p p i n g ,
T h e m e a n p e r c e n t a g e of salt m a r s h v e g e t a t i o n
w i t h i n 196 q u a d r a t s i n five r e g i o n s d i s t r i b u t e d
throughout Elkhorn Slough's undiked marshlands
d e c r e a s e d f r o m 89.6% i n 1931 to 46.4% in 2003
(Fig. 4). D i f f e r e n c e s i n v e g e t a t i o n cover were sign i f i c a n t b e t w e e n the five r e g i o n s ( r e p e a t e d m e a s u r e s ANO~vT\, F - 9,2, p - 0.0001), b e t w e e n the
12 yr (F - 469.9, p - 0.0001), a n d for the intera c t i o n b e t w e e n r e g i o n s a n d y e a r s (F - 42,4, p 0.0001), V e g e t a t i o n cover c h a n g e s were s i g n i f i c a n t
b e t w e e n all pairs of years e x c e p t t h o s e i m m e d i a t e l y
s u r r o u n d i n g a t e m p o r a r y p e r i o d of r e c o v e r y
(1956-1976, 1956-1989, and i[976-1980). The
m a g n i t u d e of m a r s h loss i n c r e a s e d with i n c r e a s i n g
d i s t a n c e [i~om M o n t e r e y Bay. M e a n v e g e t a t i o n cover for r e g i o n s 1, 2, a n d S i n the l o w e r a n d m i d
s l o u g h was 89,6% i n 1931, d e c r e a s i n g to 60.8% by
2003, T h e rate of m a r s h loss i n t h e l o w e r s l o u g h
was relatively g r a d u a l , M e a n cover for r e g i o n s 4

Fig, 2. Tidal flow and ihabitat mapping. 1870 (a,b): Unrestaicted tidal flow; extensive salt marsh; natural salt pan~,es present at
lower slough. 1917 (c,d): Levees ihave been constructed at lower slough fbr salt production and east of the raih-oad to create flesh
or brackM1 ponds and marsh withixl artifidal irnpoun~nent~. 1931 (e,f): 71des ha,~e been restricted or excluded from expanded salt
production ponds and reclaimed salt marsh; marsh has stm'ted to degrade at areas ~vJt5 restricted or excluded tides, i[956 (g,h): More
than 60 1~1,of levees exclude tides from 59% of wetlands; salt ponds and reclamation ha,~e reduced salt marsh acreage by 66%; lmge
areas of marsh degrading at mldiked regions of lower arid mid slough. 1980 (ij): Breaching of levees rean'ning [low to diked or
reclaimed former wetlands; undiked areas of lower and mid slough show noticeable salt mm'sh recovery; degraded marsh expanding
at upper slough. 2000 @,I): Less than 4 km of hltact levees remain; tidal flow returned to most former wetlands; 77% of original salt
marsh degraded or converted to mudflat; losses great<-st at eastern and upper slough (Red lines represent ixltact levees, blue lines
breached levees).

182

E Van Dyke and K. Wasson

a n d 5 in the u p p e r slough was 89.9% in 1931, b u t

d e c r e a s e d to 21.1% at region 4 a n d 4.1(~) at region
5 by 2003. T h e rate o f m a r s h loss in the u p p e r
slough a c c e l e r a t e d in the 1970s, slowed s o m e w h a t
in the 1980s, and t h e n accelerated rapidly t h r o u g h
t h e 1990s. All r e g i o n s e x h i b i t e d a p e r i o d o f
m a r k e d d e t e r i o r a t i o n in the 1950s fbllowed by a
p e r i o d of recovery in the 1960s; this t r e n d was
s o m e w h a t m o r e p r o n o u n c e d in the lower slough.
Figure 5 illustrates tlhis loss a n d r e c o v e r y within region 1.
A m o r e c o m p l e x p a t t e r n e m e r g e s w h e n changes
in vegetation cover are d e p i c t e d geospatially. Figure 6a,c,e illustrates the a n n u a l i z e d c h a n g e at each
q u a d r a t d u r i n g the intervals 11931-1956 (2b yr),
1956-1980 (24 yr), a n d 1980-2003 (23 yl), respectivelF D u r i n g the first third o f the study period,
the rate o f m a r s h loss was generally high (>0.5~}.,
yr 1) to very high ( > 1 . 0 % yr 1) in the lower slough
(regions 1 a n d 2) and m o d e r a t e ( > 0 . 2 5 % yr 1) in
the m i d a n d u p p e r slough (regions 3, 4, a n d 5).
For m u c h o f this first interval, the overall rate o f
c h a n g e was low, but a c c e l e r a t e d rapidly after 1949.
A b o u t o n e third of the quadrats in tile u p p e r
slough did n o t follow this trend; these quadrats,
which were typically adjacent to large tidal channels, e x p e r i e n c e d m i n i m a l c h a n g e before the
1960s. During the middle third o f the s t u @ period,
differences between the lower a n d u p p e r slough
b e c a m e even m o r e stIiking. T h e m~-gority o f quadrats in the lower Slough, the m i d slough, a n d the
eastern third o f the u p p e r slough showed either
little c h a n g e or an increase in vegetation cover.
A b o u t half' of" the lower slough quadrats experie n c e d m o d e r a t e to r a p i d recovery ( > 0 . 2 5 % yT x).
In conlrast, eveIy quadral in the western p o r t i o n
o f the u p p e r slough e x p e r i e n c e d high or very high
rates of m a r s h loss d u r i n g this s e c o n d interval. By
the final third of the s t u @ period, m o d e r a t e to
high rates o f m a r s h loss had r e t u r n e d to the lower
slough and losses in the u p p e r slough were uniti)rmly very high.
T h e m e a n cross section width o f ] 96 tidal creeks
in u n d i k e d areas increased f l o m 2.5 in in 19311 to
12.4 m in 2003 (Fig. 7). Dift'erences in creek width
were significam between the 13 yr ( r e p e a t e d measures A N O \ ~ , F = 257.3, p = 0.0001) a n d for the
interaction b e t w e e n regions a n d years (F = 3.1, p
= 0.0001), a l t h o u g h not significant between regions (F = 0.7, p = 0.(5). (;reek width changes were
significant between all pairs of' years e x c e p t imInediately successive pairs (1937-i949, 1956-1966,
1978-1980, 1987-1992, and 2000-2001). Increasing tidal creek width was related to distance f r o m
M o n t e r e y Bay. In the 1930s, m e a n creek width was
bem-een 2 a n d 2.5 m within all five regions. By
2003, m e a n (:reek width in all f o n t regions of" the

Fig. 5. Rapid salt marsh loss and temporary recovery within

region 1. (a) Intact salt marsh, 1937 aerial photo. (b) Degraded
vegetation in marsh interior, 1956 aerial photo. (c) Temporary
vegetation recovery, 1980 aerial photo.

lower and m i d slough (1, g, 3, a n d 4) was between

9 a n d 11 m. At the far u p p e r slough (region 5),
m e a n width h a d increased to 17 m. At a n u m b e r
of sampling points m region 5, vegetated banks
were n o longer p r e s e n t after 2000. Fig. 8 il]ustrates

E!khorn Slough Historical E co!ogy

Mamh Vegelation Ox~v~(above)

Very HighLoss
~ Modl~a~ RBOD'~Bry
I~gh t ~ s
~j~ High Recove~
Moderate Loss
[ ] ve~ High

"tidal Creek Wid~ (t:~ow)

Q Very High t n ~
~ High I n ~
.::::::N~xlertt~ I n c : n ~

'~ 8 3

9:-i.:::I V l ~ e r ~ Recovery

t High Recovery
0 Very High F~Br.,ov6~'y

Fig. 6. A n n u a l i z e d change, vegetation cover a n d tidal creek width. 19Bl-1996: (a) I-{igh to very h g h m a r s h loss at lower slough;
m o d e r a t e loss at m~d a n d u p p e r siou,~h. (b) Low to m o d e r a t e overall tidal creek v,~dth increase. 1956-!980: (c) Little c h a n g e or
m a r s h recovery at lower a n d mid slough- very h i g h loss at u p p e r slough. (d) P~igh to very h g h creek width increase at u p p e r slough
a n d s o u t h e r n part of lower slo~gh; low to rr, oderate increase elsewhere, i98&-200B: (e) J o d e r a t e to hivgh m a r s h loss at lower a n d
m i d slough; very h i g h loss at u p p e r s]o'Agh. (~ bfoderate to very high creek width increase at io;ver and m i d slough; very h g h increase
at u p p e r slough.

this evolution ~rorn a. network os tidal creeks to

o p e n mudflat within region 5. N o p e r i o d ot- sigmficant recovery was evident in any of the regions.
A geospafial depictior, os a.nnualized chartges in
creek width d u r i n g the intervals 1931-1956, 19~6-

1980, and 1980-2003 is shown in Fig. 6b,d,f. T h e

rate of c r e e k widening was generally low to moderate (<0.1 m yT-~) d u r i n g the first third o f the
study period, with a somewhat lower overall rate in
the u p p e r slough. A small n u m b e r os creek cross

184

E. Van Dyke and K. Wasson

20
16
12
m

~est'r~ct~ons to T~dal Flow

I7
[] region 1
0 region 2
& region 3
0 region4

regio.

w~
_

lg30

1950

1970

lggo

f'

' I

2010

Fig. 7. Mean cross section w i d t h of 196 tidal creeks in und i k e d areas, 1951-2005. Line is rnean fD:r an regions. Error bars
r e p r e s e n t 1 stm~dard erro~t

sections e x p e r i e n c e d high (>0,1 m yr--*) or very

high (..'>-0.25 m yr-*) rates of' change during this
initial i n t e r v a l ; t h e s e l a r g e i n c r e a s e s o c c u r r e d
a l o n g several of the slough's widest channels. During the n,iddle third of the study period, differences between the lower, mid, a n d u p p e r regions
of the slough b e c a m e increasingly a p p a r e n t . Alt h o u g h tlhe rate of c h a n g e r e m a i n e d low to m o d erate t h r o u g h o u t m o s t of' the study area, a n u m b e r
of creeks in the s o u t h e r n p o r t i o n of the lower
slough ( r e g i o a 2) and the western p o r t i o n of the
u p p e r slough (regions 5 a n d m u c h of region 4)
widened at high to very h i g h rates. By the final
third of the study period, rates of tidal creek widening were uniformly m o d e r a t e to very h i g h across
the lower a n d mid slough a n d p r e d o m i n a t e l y very
high in the u p p e r slough.
Discussio~l
(7~AUSES OF T I D A L V~TETLAND HAg.[TAT G t I A N G E

V?e have d o c u m e n t e d dramatic shifts in the extent a n d distribution of wetland habitat types at
E l k h o r n Slough during the past 150 yr. T h e s e
changes ca.l.t la:rgely be attributed to contrasting ant h r o p o g e n i c influences on the slough's hydrologT:
restrictions to the r a n g e of tidal flow that o c c u r r e d
earlier in the study p e r i o d and e x p a n s i o n of tidal
range, a m p l i t u d e , and velocity- that have o c c u r r e d
m o r e recently,
Tidal wetlands adjust to a dynamic equilibrium
of erosional and depositional processes t h r o u g h a
u n i f o r m distribution of c h a n n e l bed shear stress
a n d a balancing of m o u t h cross,sectional a r e a to
tidal v o l u m e (Allen 2000). R e d u c t i o n of tidal
prism v o l u m e d u e to restricted tidal flow (e.g., diking) can result in channel shoaling and m o u t h closure, w.hile an e n l a r g e d tidal p r i s m d u e to e x p a n d ed tidal flow (e.g., levee breaching) drives c h a n n e l
erosion ( O ' B r i e n i[981; Williams et ak 2002).

Diking and draining of wetlands was the key driver of e s t u a r m e habitat c h a n g e during the initial
100 yr of the study period. In. 1872, a raised emb a n k r n e n t for the S o u t h e r n Pacific Railroad was
constructed t h r o u g h m a r s h l a n d s on the east side
of E l k h o r n Slough (Fabing and H a m m a n 1985).
This linear feature separated m o r e than o a e third
of the slough's wetlands f r o m the main channel.
Despite their physical separation, these wetlands
r e m a i n e d largely intact fbr several decades, likely
due to the c o n s t r u c t i o n of' bridges and cuNerts
that p e r m i t t e d c o n t i n u e d tidal flow u n d e r the railroad,
In the early 20th century', l a n d o w n e r s b e g a n to
isolate wetlands east of tlhe railroad e m b a n k m e n t
f r o m tidal flow by- blocking cuNerts a n d creeks under bridges. During the same period, tidal exc h a n g e was excluded f r o m additional wetland acreage as levees were constructed for various p u r p o s e s
(Silberstein et al, 2002). Beginning a r o u n d 1900,
s p o r t s m e n p u r c h a s e d tracts of tideland and m a n aged a b o u t 120 h a as waterfowl habitat by- irnp o u n d i n g fi~eshwater b e h i n d dams across inlets
a a d levees a r o u n d artificial p o n d s (Grinnell et aL
1918). In the following decades, an additional 120
h a of m a r s h were diked and r e m o v e d f r o m tidal
influence to create salt e v a p o r a t i o n p o n d s for the
M o n t e r e y Bay- Salt Vgorks (Vet Planck 1958). Between the 1920s and 1940s, a p p r o x i m a t e l y 600 h a
of f o r m e r tidal wetlm~d were converted to agricultural uses, particularly p a s t u r e l a n d for dairy o p e >
ations (King 1981). Several additional wetland areas on the p e r i p h e r y of the slough were isolated
fi~om tidal flow by the construction of roads. By
1956, these p r q e c t s had resulted in a 45% decrease in tidal r a n g e a n d a 60% loss of salt m a r s h
acreage.
F,xpansioa o/ TidaI Flow
Prior to 1947, E l k h o r n Slough was a depositional
system with r e d u c e d tidal volume, the result of extensive diking and r e c l a m a t i o n of tidelands and
tlhe clearing of adjacent uplands. Tidal energy was
m u t e d due to shoaling in. the lower c h a n n e l and a
persistent sand bar at the natural m o u t h into Monterey Bay,, 0,5 km n o r t h of the slough on the Salinas River ( G o r d o n 1996).
In 1947, the U.S. Army C o r p s of E n g i n e e r s constructed an artificial channel to a c c o m m o d a t e vessel traffic into a newly created iharbor at Moss
L a n d i n g (Silberstein et al. 2002). This deeper, ~dder m o u t h is directly in line with the slough's main
c h a n n e l and is k e p t clear with jetties and periodic
dredging. T h e result was an i m m e d i a t e inccease in
the velocity- a n d a m p l i t u d e of tidal e x c h a n g e within

Elkhorn Slough Historical Ecology

the slough (\,Vong 1989). S t r o n g e r tidal flow, greater tidal reach, a n d a m i s m a t c h b e t w e e n the larger
o p e n i n g a n d the estuary's shallow, m e a n d e r i n g
c h a n n e l s a n d creeks a b r u p d y t r a n s f b r m e d the
slough into a highly erosional system. In the years
since 1947, the m a i n c h a n n e l has rapidly increased
in b o t h width a n d depth, resulting in an increase
in v o h n n e of over 200% ( C r a m p t o n 1994; Malzone
1999). Field m e a s u r e m e n t s r e c o r d b a n k erosion
rates m~eraoing 0.5 m vr ~ between 2000 a n d 2004
(Wasson u n p u b l i s h e d data).
G r e a t e r tidal energy, increased tidal a m p l i t u d e ,
a n d e x t e n d e d periods o f m a r s h i n u n d a t i o n resulting f r o m the 1.~4 t o p e n i n g are ahnost certainly the
principal causes o f m a r s h d e g r a d a t i o n a n d tidal
creek widening d u r i n g the m o s t r e c e n t five decades o f the study period. O u r m a r s h q u a d r a t a n d
tidal c r e e k analyses show that conversion o f salt
m a r s h habitat to n m d f l a t a n d the widening of tidal
creeks a c c e l e r a t e d to significant levels only after
1949, timing that coincides with the artificial char>
nel o p e n i n g .
D u r i n g the final two decades o f o u r study, i m e n tional a n d u n i n t e n t i o n a l b r e a c h i n g of levees allowed tidal flow to r e t u r n to m a n y o f the slough's
diked f b r m e r wetlands. I t a b i t a t restoration ret u r n e d full flow to a b o u t 120 ha at E l k h o r n Slough
National Estuarine Research Reserve's South
Marsh in 1983, Rfllowed by the r e t m ' n of" partial
flow ( t h r o u g h tidegates) to an additional 40 ha at
N o r t h Marsh in 1985. D u r i n g the same period, levees at Pro'sons Slough a n d the a b a n d o n e d salt
works failed, as did n u m e r o u s smaller levees. In
less t h a n a decade, E l k h o r n Slongh's tidal prism
e x p a n d e d b v a b o u t 30% (Malzone 1999). T h e result was significantly h i g h e r tidal velocities (V~r
1989), accelerating the rate of tidal erosion in
c h a n n e l s a n d creeks. E x p a n d i n g channels a n d
creek networks drNe a positNe f e e d b a c k l o o p by
f u r t h e r enlarging the tidal p r i s m a n d by e x t e n d i n g
the reach of tidal flow d e e p e r into the marsh.
Patterns o f m a r s h d e g r a d a t i o n a n d tktal creek
widening varied by region a n d by p e r i o d (Fig. 6).
In the d e c a d e following the 11947 o p e n i n g , the rate
o f vegetation loss was greatest in regions nearest
the new artificial m o u t h . D u r i n g the s u b s e q u e n t 24
yr, losses were extremely ~ high in regions thrthest
f r o m the new m.outh, while the lower a n d m i d
slough e x p e r i e n c e d m i n i m a l m a r s h loss and, in
m a n y cases, significant recovery, This p e r i o d o f ren e w e d accretion in the lower slough p r o b a b l y resulted f r o m the onset of high erosion rates f a r t h e r
up the slough, as large v o l u m e s o f s e d i m e n t b e g a n
to be dislodged a n d t r a n s p o r t e d . In any case, the
recovery was short-lived. D u r i n g the final 23 yr o f
the stud)5 high rates o f marsh loss a n d tidal creek
widening r e t u r n e d to the lower slough. At the

185

Fig. 8. Evolution of" salt marsh to mudflat within region 5.

Dark areas are salt marsh, light areas umegemted. (a) Tidal
creek network and growing imerior pannes, 1980 aerial photo.
(by I)eteriorated marsh largely converted to open mudflat, 200 [
aerial photo.

same lime, rates o f loss have accelerated to unii'orufly very high levels t h r o u g h o u t the u p p e r
slough.
T h e exact m e c h a n i s m by which increased tidal
amplitude, velocity,, and v o l u m e have caused m a r s h
vegetation to d e g r a d e is unclear. Surface erosion
m a y be r e d u c i n g elevations to b e n e a t h the level
w h e r e SaLicornia can snrvNe, a l t h o u g h c u r r e n t velocities within the m a r s h are typically too low to
e r o d e the substrate (Lowe 1999). S t r o n g e r c m :
rents may result in d e c r e a s e d s e d i m e n t deposition,
gradually lowering the m a r s h plain ( O r r et al.
2003). Vegetation t h i n n i n g a p p e a r s to progress
f r o m the interior o f the marsh, initiating the f o >
m a t i o n of growing m u d p a n n e s (Fig. 8). In time,
only a fringe o f vegetation r e m a i n s a l o n g the banks
o f c h a n n e l s and creeks; eventually these banks deteriorate as well, leaving an e x p a n s e of mudflat.
T h e p a t t e r n is consistent with o t h e r reports o f
m a r s h d e g r a d a t i o n resulting f r o m relative lowering

18(}

E. Van Dyke and K. Was~on

of the m a r s h plain and an a c c o m p a n y i n g increase

in the f r e q u e n t T a n d d u r a t i o n of i n u n d a t i o n (Philiips 1986; Kearney et al. 1988; D e L a u n e et al. 1994;
Downs et al. 1994; H a r t i g et al. 2002).

Add~tior~a2 Po.s.sible Cayuses of Change

A l t h o u g h we attribute the changes we have docu m e n t e d largely to diking, c h a n n e l construction,
levee breaching, a n d o t h e r a n t h r o p o g e n i c modifications to tidal flow, several additional factors
m i g h t c o n t r i b u t e to wetland habitat c h a n g e at Elkh o r n Slough.
Before 1909, the Salinas River shared a c o m m o n
m o u t h with E l k h o r n Slough. Redirection of the river directly into M o n t e r e y Bay in 1909 eliminated
this seasonal source of freshwater and s e d i m e n t
(Go~don 1996). Decreased s e d i m e n t i n p u t is rep o r t e d to be a principal cause of salt m a r s h loss at
s o m e locations, and r e d i r e c t i o n of the Salinas River likely r e d u c e d the supply- of s e d i m e n t to Elkh o r n S 1 o u g h ' s tidal w e t l a n d s , a l t h o u g h h i g h l y
erodible soils, steep slopes, and extensive agricultural and residential d e v e l o p m e n t adjacent to the
slough c o n t i n u e to provide large quantities of sedi m e n t (Diekert and Turtle 1985).
Relative sea level increase is a cause of wetland
habitat change m m a n y regions, a l t h o u g h the rate
of eustatic sea Ievel rise on the central California
coast is relatively low a n d n o t likely to o u t p a c e the
rate of" m a r s h accretion (Atwater et al. 1977). If a
c h a n g e in relative sea level is c o n t r i b u t i n g to we>
land loss at E l k h o r n Slough, it is m o r e likely the
result of" land subsidence (Patrick and D e L a u n e
1990). Lowe (1999) suggests that rapid salt m a r s h
ioss in the u p p e r slough may have been due to a
d r o p in the m a r s h plain following the 1989 L o m a
Prieta e a r t h q u a k e , a l t h o u g h o u r results reveal not
a single episode, but a trend of m a r s h loss that
b e g a n decades b e f o r e the 1989 event. Subsidence
of the m a r s h plain m i g h t also result f r o m g r o u n d water overdraft. G r o u n d w a t e r levels in the E l k h o r n
Slough area have b e e n falling since the 1950s, alt h o u g h the m a g n i t u d e of associated land subsid e u c e is unkn.owTt (Fugro Wrest 1995).
Biotic factors such as disease and h e r b i v o r y are
also potential causes of wetland habitat c h a n g e
(e.g., Miller et al. i[996). Marsh d e g r a d a t i o n is occurring rapidly in u n d i k e d area,s at E l k h o r n Slough
while areas v~qth extremely- limited tidal flow are
relatively unaffected. It is unclear h o w disease or
h e r b i v o r y m i g h t be linked to these t,ydrological
conditions.
EFFECTS OF HABITAT CHANC.E ON
BIOLOGIC&~L COMMUNITIES

T h e c o n s e q u e n c e s to E l k h o r n Slough's plant
a n d animal c o m m u n i t i e s from 150 yr of hydrologic

alteration and habitat c h a n g e are poorly u n d e r stood. Additional studies a i m e d at u n d e r s t a n d i n g

the effects of a dramatic decrease in salt m a r s h
a c r e a g e , c o r r e s p o n d i n g i n c r e a s e s in i n t e r t i d a l
trmdflats a n d p a n n e s , a n d enlarged subtidal channels a n d creeks will be [iandamental to future conservation planning.
Only a b o u t 8% of c o n t e r m i n o u s U.S, salt m a r s h
acreage occurs along the Pacific Coast (Field et al.
1991), so the d e g r a d a t i o n or loss of m o r e than two
thirds of" E l k h o r n Slough's salt m a r s h is extremely
significant. In addition to providing a variety, of key
e c o w s t e m services, such as t r a p p i n g s e d i m e n t s a n d
filtering nutrients f r o m u p l a n d runoff, California's
tidal m a r s h e s provide fbod or habitat for a variety
of organisms including s h o r e crabs (Mer~igrait~susoregonensis) a n d song sparrows (Meto@fza ~dodia), as
well as s u p p o r t i n g various t h r e a t e n e d animals, including the California clapper rail (Raff'us goregirost~s obsdet~s) a n d California brackishwater snail
(Tryonfa imitator) (Zedler 1996b; Wasson et al.
2002)..As E l k h o r n Slough's m a r s h l a n d s have deg r a d e d or converted to other habitat a/pes during
the past century, salt m a r s h associated faunal comm u n i t i e s have u n d o u b t e d l y declined in a b u n d a n c e
a n d distribution..
A less a p p a r e n t c o n s e q u e n c e of E l k h o r n
Slough's ihistory of hydrologic modifications has
b e e n the loss of transitional v e g e t a t i o n c o m m u n i ties. Before dikes were constructed to segregate tidal f r o m a o n t i d a l habitat types, wetlands at the m a >
gins of the slough were subjected to e x t r e m e variations of' saliniw and i n u n d a t i o n r e g i m e s as a result
of periodic wcles as well as occasional, episodic
events. During the decades since tlhe 1940s, num e r o u s surface streams, springs, and seeps in the
vicinity of E l k h o r n Slough have d i s a p p e a r e d , presurnably due to lowered g r o u n d w a t e r levels resulting f r o m agricultural and domestic p u m p i n g (Van
7D/ke u n p u b l i s h e d data). A l t h o u g h our habitat
m a p s delineate the slough's vegetated wetlands as
u n i f o r m l y salt m a r s h during the earliest periods, a
variety of small patches of freshwater-influenced
vegetation u n d o u b t e d l y existed in the viciniw of
f'reshwater features (Hayward 1931). Vegetation
types a d a p t e d to e x t r e m e salini W fluctuations are
n o w very u n c o m m o n within E l k h o r n Slough's
modified tidal wetlands. Because these brackish
m~d trar~sitional wetIm~d habitat ~ p e s are increasingly- 9u n c o m m o n , species associated with these
conditions (e.g., the tidewater goby, E'uc},cgogoSius
r,#~berr),i), which were likely once relatNely comm o n at E l k h o r n Slough, are now very rare (Yoklavich et al. 2002)~
.As E l k h o r n Slough's intertidal habitats have und e r g o n e changes, adjacent s u b t i d ~ c o m m u n i t i e s
r
have b e e n affected by the slough
s altered hydrol-

Elkhorn Slough Historical Ecology

ogT e~s well. Physical p a r a m e t e r s , such as water

m o v e m e n t , sa,linity, a n d s e d i m e n t size, are k n o w n
to influence estuarine faunal c o m m u n i t i e s (Edgar
et al. 2000; Little 2000). Extensive eelgrass (Zvstera
wer/,~a) beds were p r e s e n t along m u c h of the
slough's lower m a i n c h a n n e l in the 1920s; only a
few small p a t c h e s r e m a i n today (MacGinitie 1985;
Z i m m e r m a n and Caffrey 2009). I n c r e a s e d turbidi t , / a n d c h a n n e l d e p t h resulting f i o m h i g h e r tidal
energy are likely causes~ This decline is a significant conservation c o n c e r n because eelgrass is a
m a i o r c o n t r i b u t o r to productivity m California estuaries a n d provides :important habitat for m a W
invertebrates a n d fish species (Ricketts et al. 1985;
Yoklavich et al. 2002). T h e widening and d e e p e n ing of' E l k h o r n Slough's m a i n c h a n n e l and tidal
creeks has also enabled the slough to be colonized
by large resume fish a n d m a m m a l s . Several species
that would only have b e e n p r e s e n t n e a r the m o u t h
a c e n t u r y ago, such as l e o p a r d sharks (7~riahis ser~
ifascfata), bat rays (MySobatfs cagijornica), h a r b o r
seals (Phoca vit',Srea), a n d sea otters (]~)nt(ydr~ Z'~O'is), are now a b u n d a n t t h r o u g h o u t m u c h of the
estuary (Harvey and C o n n o r s 2002; Yoklavich et al.

2002).
T I D A L ~ V E T L & \ D {~ONSER\C~TION A~ND
HABITAT CHANGE

C o n s e r v a t i o n and restoration of estuarine ecosystems have e m e r g e d as m a j o r e n v i r o n m e n t a l concerns in r e c e n t decades (Kennish 2002), As our
study demonstrates, E l k h o r n Slough's tidal web
lands have u n d e r g o n e m o r e than a century of habitat change. T h e majority of these wetlands are now
owned a n d m a n a g e d for conservation purposes,
and a n t h r o p o g e n i c modification of the slough's
hydrolog3~ has h r g e l y c e d e d . Yet rates of conversion f r o m salt m a r s h to rnudflat or o p e n water and
e x p a n s i o n of tidal channels and creeks r e m a i n
high a n d may be accelerating, suggesting that a
new equilibrium m a y n o t be r e a c h e d fox m a n y decades. C o n s e r v a t i o n p l a n n i n g is difficult within this
context of u n c e r t a i n t y eaid rapid change.
Wetland m a n a g e r s face a dual challenge of developing and i m p l e m e n t i n g strategies that not only
slow the rate of change to p r o t e c t existing intact
habitats, but also restore a n d e n h a n c e d e g r a d e d
wetlands in o r d e r to m a i n t a i n an a p p r o p r i a t e diversity of' habitat types. Restoration of e s t u a r m e
habitats fk~equently fails to m e e t desired goals (Zedler 1996a; Z e d l e r and Callaway 1999), M o r e than
200 h a of diked a n d d r a i n e d f o r m e r salt m a r s h at
E l k h o r n Slough National Estuarine Research Reserve were r e t u r n e d to tidal influence t h r o u g h restoration projects u n d e r t a k e n during the 1980s,
T h e s e newly created tidelands s u p p o r t rich communities of birds, fish, a n d invertebrates within rid-

187

al lagoons and mudflats, yet restoration to the fo>

m e t landscape of' salt marsh, p a n n e s , and tidal
creek networks has not succeeded.
"The likelihood of restoration success is increased
wt.en plans imitate the c o m p l e x structure of natural tidal wetlands and m a i n t a i n connectivity with
intact wetland habitats as well as with ad:joining
subtidal and u p l a n d habitats (Williams and Zedler
1999; D e s m o n d et al, 2000), T h e currently rapid
rate of habitat c h a n g e at E l k h o r n Slough is the
result of a long history of deliberate tidal alteration
and habitat isolation t h r o u g h the c o n s t r u c t i o n of
levees, channels, and tide gales. Ironically, slowing
the rate of habitat c h a n g e :may require these same
t o o l s - - t h e c o n s t r u c t i o n and m a i n t e n a n c e of" levees, channels, a n d g a t e s - - t o m u t e tidal energy,
r e d u c e erosion, and ei:~hance m a r s h accretion.
Wetland m a n a g e r s m u s t balance the n e e d to mitigate for the effects of historic alteration.s (e.g.,
m u t e tidal flow by building new dikes and gates)
with the n e e d to r e i n t r o d u c e natural tidN flushing,
salinity a n d i n u n d a t i o n variability, a n d hat0itat connectwity (e.g., by rernovmg existing dikes and
gates).
C o n s e r v a t i o n p l a n n i n g at o t h e r West Coast estuaries has b e e n s t r e n g t h e n e d by studies that docu m e n t past habitat conditions and historic p a t t e r n s
of change (e.g,, Zedler 1996b; Goals Project 1999;
Borde et al. 200g). We believe that our analysis of'
150 yr of habitat change can. similarly i n f o r m cons e r v a t i o n a n d r e s t o r a t i o n e f f o r t s at E l k h o r n
Slough. Historical ecolog T may n o t supply- easy answers to E l k h o r n Slough's c o m p l e x habitat conservation questions, but a thoughtful analysis of the
historical r e c o r d can h e l p g u i d e the d e v e l o p m e n t
of feasible and sustainable restoration goals.
A G K N O ~ L E , D GMENTS
Pioneer4mg studies by Jobm Oliver a n d c o l l e a . ~ e s at Moss
L a n d i n g Marine Laboratories a n d by Rikk K~'itek a n d c 4 1 e a g u e s
at the California State University, M o n t e r e y Bay Seafloor Mapp i n g Lab respired ttzts project, _&ndt ea 1&bolfolk, Robin Crossinger, a n d Mark Silberstein p r o v i d e d early direction, a n d A n d r e a ' s
conk[nued collaboration h a s st:tengthened tlxis work, T h a n k s t~3
Kristin Byrd, Jobm CNla~my, J o s h Collins, Mike Foster. J o h n Oliver, Barb Peichel, D o u g S~r,ith, a n d two a n o n y m o u s re~oiewers
fbr t h o u g h t ~ . l discussions or c o m m e n t s on the r n a n u s a ' i p t .
l l ~ a n k s to the University o f Califbrnia, Santa Ciuz Map Root21,
M o n t e r e y C o u n t y Sm-veyors Inc., a n d m a n y o t h e r archives that
p r o v i d e d access to i m p o r t a n t historic m a p s a n d fax,ages. We are
gratefifl to the E s m a r i n e Reserves Division o f the National Oceanic a n d Atll~ospheIic AchlEnistration :for f t m d m g a n d m t l i e
California D e p a r t m e n t of Fish a n d G a m e a n d the E l k h o m
Slough F o u n d a t i o n fbr cmgofng s u p p o r t ~f historical ecology
studies at E l k h o m Slough National E s m a r i n e Research Reserve.
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Received, A%g...~st18, 2004

Accepted, O~ob~" 28, 2004