A Study of the Effects of a Wave Gradient on
Intertidal Zonation a.t Yaquina Head,
Lincoln County, Oregon
Michael w. Davis
Patrlcio Arana
J<~JT105 Held
Riclwrd Schrock
June 30,1978
A Study of the Effects of a Wave Gradient on
Intertidal Zonation at Yaquina Head,
Lincoln County, Oregon
ABSTRACT
A study of the effect of wave exposure WaS made on the
outer rocky coast of Oregon. We found a definate effect on
intertidal zonation produced by the wave gradient. This
suggests that waves are a determining force in the intertidal
habitat of the outer rocky coast.
•
Il\:TRODUCTICN
Intertidal zonation is a common theme among marine
ecologists of the world. The Stephensons (1949 1.1972), outlined
the universal features of intertidal zonation in a comparative
review, although the first de£inative work on intertidal
zonation is attributable to Lorenz(l863j.
Intertidal zonation of British shores was desc~ibed by
Lewis(l9SS,l964) and a scheme of three major zones was postulated to be the Littoral, Eulittoral, and Sublittoral zones,
as defined by dominant organisms. l'1any such schemes have been
proposed(eg. Ricketts et ll•' 1968; Carefoot, 1977 forr review)..
All such schemes are subject to limited generality due to the
unique nature of intertidal sites as well as their heterogeneity.
Intertidal zonation is determined primarily by the tidal
environment, as modified by: !)wave action; 2) topography and
substrate; 3) cl-imatic factors; 4) period of immersion; and 5)
biotic factors(Lewis, 1964). In this study we have chosen to
investigate the effect of wave action on intertidal zonation,
assuming that all other factors are held constant(Southward and
Orton,l954l. Two study sites were chosen which satisfied thia
assumption for the most part. These sites differed primarily,
in the degree of wave action and were described as stronq and
moderate wave action respectively.
~
I
DESCRIPTION OF STUDY AREA
The area of this study wns Yaquina l~ad, a prominant
basaltic headland on the open coast located four miles north
of Newport, Oregon. The specific study sites were on the
eastern surface of a point which receives a great deal of
winter wave action (Figures 1 and 2). \ve chose a long,. near
vertical rock surface with a compass bearing of 30°N ~y 210°S
along its length. It has an exposed southeastern face that is
( 1}
parallel to the direction of wave trDvel and boardered by a
surge channel. The seaward end is the most exposed to wave
action. The shoreward end, with a shallow surge channel and
tidal shelf receives less wave action.
The two study sites were chosen with respect to the wave
gradient; site one at the seaward end and site two at the
shoreward end. Both sites were similar as to slope, compass
directio~, degree of shading, amount of wind exposure, and
substrate type. A comparison of transect heights and tidal
heights is given in Figure 2b, as determined by methods below.
MATERIALS
AND
l'-1ETHOOS
At each site a vertical line transect with one foot
intervals was established by measuring from the tow low tide
level with a stadia rod and hand level. Sampling quadrats were
established by randomly placing a thirty centim~ter square grid
in each interval along the transect. From within the grid,
£~mples were tuken to determine the species encountered and
counts were made of number of individuals and the percent cover
of each species. Drawings were made depicting the.types and
arrangement of organisms within the grid area(Figure 3}.
Photogr&phs were also taken at the location and study sites.
Sampled or~;,;.nisms were identified to species when possible,
using taxonomic keys of the eastern Pacific coast(Light,1975;and
Ricketts~~., 1968). The data were recorded and tabulated to
determine the species composition and widths of visually observed
zones. Because of the many factors affecting organisms existing
together as associations,(as in the mussel community) and
because many org~nisms were found only rarely and in small
quantities, only the dominant organisms encountered were considered for determining zonation. We considered organisms to
be dominant if they occured as greater than two percent of the
( 2)
.,.
i
•
cover in any one interval.
RESULTS
Relative. abundance expressed as perce t cover for flora and
fauna found in our two transect areas are g ven in Flgure5 4-7.
Common organisms found in the more wave-exp. sed transect were1,
crustoae coralline algae, ~nt~op!eurR xanth
ammi , 4i&Qngxlgcentrotus 12ureurat1.:1..!t Corallina vancouve ie i , .f;loc:g&um
violacium, ~til.us californianus, Pollici e E?2lXfrlfUi\li~: BaJtn\!1 ·
cariosus, ~· g*a~ula , £atha~al~! £~1Ji, A
~iQ~tal&lt an4
Littorina sp. £• dalli was not recorded in
due to taxonomic confusion, but .was present S• ~al:ti was
recorded on the second transect after prope identification was
made.
On the second transect common organis s were ~· aanthog~ammica,
.§. .. ;eurpuratus, crustose coralline algae, !!!l ocladia murieata,, !•
californianus, ~· sariosus, ~- glandula ' • a~gitalis, £· dalli,
and Littorina sp.
On the basis of visual obsorv.Jtion
determined that the
study sites had three major zones of org.:mi ms present. These zones
were also apparent in our quantitative data The zones are
summarized in Figure a.
Greater Wave Exposure - The transect ith more wave exposure
had a lower zone(zone !)composed primarily f exposed rock,
crustose coralline algae, !• ~anthog[a~mi~t !• ~urpu{atug, and
and P.
in the upper regions, a subzone of £• vance veriens
violacium. A middle zone(zone 2)was compose almost! exclu.si vel:y
of M. cal'ifornianu.s and associated P. ol m
mussel dominated the area of zone two and h d many species of
animals associated with the cracks between he shells .tiS they
grew out from the rock substrate. This comm nity woul~make an
excellent place to study on the micro-habit .t level, something
outside of the scope of this study • .fisaste pctf£jaCSU! was also
-
-
-
( 3)
found here and is a predator of small !:!• californianus(Dayton, 197J").
An upper zone(zone 3)was composed primarily of ~~lanus spp., £.•
dalli, ~· digitalis, and Littorina sp. This community is
dominated by prostrate organisms adapted to high wave action.
Lesser Wave Action - The transect with less wave ~xposure
had modifications of the above zonation structure. Zone one
was similar to the higher wave action site but did not include the
upper subzone of coralline red alg,•e. 'rhe abundance of A.
!anthogrammica and !• EUrQuratu! was greater in this transect
of lower wave exposure. Zone two was very disimilar, having a
predominance of £!• muricata, with a high subzone of !:1• californianus
in low abundance which did not form an extensive micro-habitat
as described above. The third zone was similar to the more wave
exposed transect, although we noted that the characteristic
organisms had a generally lower range of distribution along
tidal height. This zone was dominated by .§• gl~f!dUl!, ~·
£1gital!;!, and f• ~lJ;i.
A comparison of ranges of flora and fauna \'IdS made over
the two transects(Figures 9,10). Inspection shows that distribution
of plants and animals is genertJlly lower in transect two, the
moderate
wave
exposed site. Exceptions are found in the M.
.
.
£alifornianus-£:_. muricata interactions of zone two, where ~·
muricata tend• to displace ~· californianus i~ the moderate
wave exposed site. ~· ~nthog£_~~ also occurs hfgher up in
the moderate wave exposed site, displacing the eoralline red
algae subzone found in zone one of the strong wave expossed site.
-
-
DISCUSSION
Intertidal zonation was influenced by a wave gradient.
From our scheme of zonation, we noted that a wave gradient was
typified by a change of either: dominant organisms(eg. ~·
californma~-E· Eolymerus to f• ~icata in zone twol; relative
abundance of organisms(eg. !:.• ~:2;nthog,r~mmica in zone one); or
( 4)
relative tidal height occurance of organisms(eg. Barnacles~· digitalis in zone three).
It is apparent from the above possible change$ in intertidal zonation that many factors are involved in the observed
changes of intertidal zonation along a wave gradient. Physical
factors are paramount in the intertidal habitat(Lewis,l964;
Stephenson, ·1949) •. Our study sites were chosen to minimize the
inter-study site variation of slope, substrate, temperature,: light,
dessicatiGn,: wind, and food availability, ie. two sites along a
rock wall boardered by a connecting surge channel. Thus we
addressed ourselves to the effect of wave exposure and its two
components,, surge and shock. The surge and shock of wave action
affect the organisms through physical damage and wetting
(EifionJones and Demetropoulos,l968). Organisms have developed
adaptations to cope with wave action and we have assurr.ed that
the presence or absence of these adaptations has determined their
relative distributions along the wave gradient.
The semi-diurnal tide crea.tes the be1sic milieu of eonditfons
in the intertidal. \AJave action modifies these conditions and
thus creates further habitat pattern which we observed in our
study. The observed zonation of intertidal organisms is a direct
result of the physical and biological fuctors in which these
organisms must live(Lewis,l964).
Biological factors have been found to be of importance in
determining the zonation of intertidal organisms. We have no way
of testing these factors in this study, but merely suggest them
as interesting alternatives and modifications of the effect of
wave action. Organisms have specific physiological adaptations
which dllow them to live in specific environments. For example
E.• Eol;y:merus is adapted to high wave shock,by clinging to rocks
or mussel~with a hydrodynamic morphology and feeds with cirri
modified for wave aurge. The crustose red algae are adapted
with a prostrate morphology and are incrusted with Caco in order
3
to resist grazing and high wave exposure on steep intertidal
( 5}
areas.
Organisms may also be coadapted. We note that in the case
of ~·· californianus, a concurrent micro-community is established.
Thus once one type of organism colonizes an area, others will
move in; which are adapted to live with the original colonizer
in its modified environment. These events are modified during
succession u~~~lafeltf~~S)community is reached in time and space
(Dayton,l97St~ ~redation, both herbivory and carnivory as well .
as competition for space and food, have been found to also be
significant factors determining intertidal zonation(Connell,
196la,l96lb; Dayton,l97~; Paine,l974).
In this study it is interesting to note the large change
in zone two between the strong wave action and the moderate wave
action(transects one and two respectively). ~· californianus
dominated transect one whereas f• ~E!£Qi~ dominated transect
two. i·v'e hypothesize that there is an important biologivally
determined exclusion occuring here. However the importance of
wave action cannot be overlooked in the exclusion ~f ~· muricata
from the exposed site versus the exclusion of ~· £~l&f9fnianus
from the less exposed site~ Perhaps the mussels cannot settle
properly in the ~· muricata community, or there is a specific
predator effect of ~~ QChraceus(Dayton,l97Si.•
Specific experiments have been designed and are applicable
to the biological causes of intertidal zonation relative to
physical factors(Dayton,l97S; Menge,l976). The scope of this
study was limited to the evaluation of wave exposure effects
upon intertidal zonation, hence cannot answer the above vital
question.
(c)
BIBLIOGRAPHY
Carefoot, T. 1977 • .facific Seashores: a guide to intertidal
ecology. Univ. Wash. Press. Seattle.
Connell,J.H. 196la.The influence of interspecific competition
and other factors on the distribution of tne barnacle
f.hthamacus stellatus. Ecology 42:710-723.
Connell,J.H. l9.6lb. Effects of competition, predation by
!.hais !aEill£2 and other factors on the natural populations of the barnacles Qalanus P~.lendid~!·
•• Ecol.Monogr. 31:61-104.
Lewis,J.R. 1955. The mode of occurance of the universal
intertidal zones in Great Britaig. J. Ecol .. 43:.270-290 ..
Lewis,J.R. 1964. !he ecology 2f F.QC~X !h£~· London:English
Univ. Press. 323p.
Light, S.F. 1975. ~ight's !'lanual; intertidal inVeftebrates
of the central California coast. Univ. Calf •. Press.
Berkeley.
Lorenz,l873. In Between Pacific tides. IVedition. Ricketts,a.r.
Calvin,J. and Hedgpeth,J.w. Stanford Univ.
Press. Stanford. 614p.
f-1argalef ,R. 1968. Perspectives in ecological theory. Univ.
Chicago Press. Chicago. lllp.
Menge,B.A. 1976. Organization of the New England rocky
intertidal community: role of predation, competition
and environmental heterogeneity. Ecol. Monog. 46:355·393.
Paine, R.J. 1974. Intertidal community structure: experimental
studies on the relationship between a dominant competitor
and its principal predator. Oecologia 15:93-120.
Ricketts,E.F.,, Calvin,J. and Hedgpeth,J.W. Stanford Univ.
Press. Stanford. Fourth Ed. 614p.
Southward,A.J. and Orton, J.H. 1954. The effects of wave
action on the distribution and numbers of the commoner
plants and animals living on the Plymouth. breakwater ..
J. Mar. Biol. Assoc. U.K. 33:1-19.,
Stephenson,T.A. and Stephenson,A. 1949. The universal features
of zonation between tide-marks on rocky coasts. J •. Ecolt
37:289-305 •.
Stephenson,T.A. and Stephenson,A. 1972. !:!ill P.ftween tidemark§.
on rocky shores. W.H. Freeman Co. San Francisco •
••
Dayton,P.K. 1975. Experimental evaluation of ecological
dominance in a rocky intertidal algal community.
Ecol. ~onog.45:137-159.
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Figure 4. Percent cover of animals at the strong wave
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Figure
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Percent cover of animals at the moderate wave
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Figure 6. Percent cover of plants at the strong wave
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Figure 7. Percent cover of plants at the moderate wave
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Figure 9. Comparison of the animals at transect one and
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Figure 10. Comparison of the plAnts at transect cne and
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