Population Dynamics and Habitat Suitability of the Intertidal Sea Anemones

Anthopleura elegantissima and A. xanthogrammica

Kenneth P. Sebens

Ecological Monographs, Vol. 53, No. 4. (Dec., 1983), pp. 405-433.

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 Ecoiogicai Mortogrrrphs, 53(4), 1983, pp. 405-433
0 1983 by the Ecological Society of America

POPULATION DYNAMICS AND HABITAT SUITABILITY

OF THE INTERTIDAL SEA ANEMONES
ANTHOPLEURA ELEGANTISSIMA AND

KENNETHP. SEBENS
Museuin of Cornprrrutive Zoology, Hcirvcird University,
Cciinbriclge, 1Mussachusetts 02/38 U S A

.4bstruct. Population dynamics of the intertidal sea anemones Anthopleura elegantissiina (Brandt)
and A . xanthogrrrmmict~ (Brandt) were investigated by long-term monitoring and expe~imentalma-
nipulation of populations in Washington State. Sizes of individual A . xanthogrciininica and A . ele-
grrntissiinci fluctuated seasonally, increasing during the spring and early summer and shrinking or
reaching a pleateau the rest of the year. Data from marked individual A . xunthogrrrmmicu showed
that adults (26.5 cm) moved very little and may persist for at least several decades. Only 3 of 160
mapped adult anemones disappeared over 2 yr. Juveniles moved more often and thus appeared and
disappeared at much higher rates. Growth of A . xcinthograrnmictl in control and experimental removal
areas was compared by a new technique. One-year growth increments were used to compare growth
in each size class by analysis of variance and were also used to plot long-term growth trajectories
and expected reproductive success (habitat suitability, Fretwell 1972). Growth was much more rapid
in areas with experimentally reduced density than in control areas. The control area with the lowest
density had the greatest individual growth rates. Differences in growth trajectories and expected
reproductive success between control areas and the experimental areas indicate that adult density
affects growth of most size classes. The mechanism is probably both noninterference competition for
prey and direct interference by adult tentacle crowns blocking capture by smaller individuals.
Clonal aggregations are formed by longitudinal fission of A . eleguntissima. ( A . xrrnthogutlminica
never reproduce asexually.) Monitored clonal aggregations of A . eleguntissiinrr showed that fission
rate is greatest during the fall and winter, averaging 0.17 divisions per clonal individual per year in
all areas. Only the larger individuals in each clone divided and a single individual never divided more
than once in a given year. Rate of fission was independent of mean individual size within clones.
Therefore, clones in more suitable habitats produced larger individuals and thus had greater gonad (off-
spring) production. Clones in the high intertidal were composed of very small individuals, had the
lowest rates of fission, and were clearly occupying a marginal habitat.
Key ~vorcls: Anthopleura elegantissima; Anthopleura xanthogrammica; nse.xual reprotluction;
habitat suitubilitj~;rocky interticlal population dynrrmics; sea onernones.

also be a function of population density once habitats

Populations usually exist over a range of habitat types
and individuals may have several optional responses
to such habitat variability. Animals which make an
initial selection and occupy a particular area over long
periods of time often show a plastic individual re-
sponse to changes in the habitat characteristics, while
highly mobile animals may be less plastic but more
selective. Few field studies of populations have con-
sidered the relationships among habitat variability,
density, and individual success. Fretwell (1972) de-
fined habitat suitability as the expected reproductive
success of all individuals in a particular habitat. It is
also equal to the mean reproductive success of all in-
dividuals in a particular habitat. Although physical and
biological characteristics may define an a priori "basic
suitability" for a habitat before its occupation by in-
dividuals of the population in question, suitability will
Manuscript received 10 November 1981; revised 23 June
1982; accepted 7 October 1982; final version received 7 De-
cember 1982.
are occupied. Thus, all other things being equal, a
crowded habitat with more prey may be less suitable
than a sparsely populated habitat with few prey avail-
able.
The present study considers the concept of habitat
suitability as it relates to populations of intertidal sea
anemones (Anthozoa; Actinaria). Few studies have in-
vestigated the population ecology of sea anemones
(Ottaway 1979b, Sebens 1982b, c ) , although they are
conspicuous members of rocky intertidal communities
(Dayton 1971, 1973), tropical coral reefs (Fishelson
1970, Sebens 1976b), and certain soft-substratum ben-
thic communities (Fager 1968). Field investigations
have concentrated on anemone agonistic behavior
(Francis 1973n, b , 1976, Purcell 1977), reproductive
cycles (Ford 1964, Chia and Rostron 1970, Dunn 1975,
Jennison 1978, 1979~1,b , Sebens 1981b, 1982b), distri-
bution, and physiological tolerances (Sassaman and
Mangum 1972). There have also been recent studies
of asexual division and spatial extent of clones (Hoff-
man 1976, Shick 1976, Sebens, 1982b).
406 KENNETH P. SEBENS
Several factors have limited potential investigations
of anemone life history characteristics and basic nat-
ural history in temperate areas: the impossibility of
aging individuals, the difficulty o f marking individuals,
and the relative inaccessibility of exposed coast sites
during winter seasons. The present investigation was
designed to quantify individual and population char-
acteristics and to use such information to analyze the
process of habitat selection, the factors contributing
to habitat suitability (expected reproductive success,
Fretwell 1972),and the degree of behavioral, morpho-
logical, and reproductive plasticity available to the
anemones over a range o f habitats. A comparison of
species which differ substantially in their reproductive
mode can also provide insight into the adaptive sig-
nificance of asexual reproduction among marine in-
vertebrates (Sebens 1979, 198%).
The congeneric sea anemones, Anthopleura xantho-
grammicu (Brandt) and A . eleganti~sima(Brandt),are
common and conspicuous members of rocky intertidal
communities along much of the western coast of North
America. The large solitary A. xanthogrammica is most
abundant in the Mytilus californianus-dominated com-
inunity of exposed outer coast shores, where it feeds
primarily on mussels released by heavy wave action
and by the foraging activity of the starfish Pisaster
ochmceus (Dayton 1971, 1973). A. elegantissima is
much smaller and divides by longitudinal fission to
form extensive clonal aggregations which compete ag-
onistically for space along their borders (Francis 1973~1,
b, 1976).
Sea anemones are long lived and relatively immo-
bile. They are often extremely plastic in morphology
and behavior, occupy a wide range of habitats, and
persist while habitat conditions change. The intertidal
anemones A. xanthogrammica and A. elegantissima
can be considered spatial dominants in their respective
assemblages (Dayton 1971, Sebens 1981c, 1982, c ) .
Individual A. xanthogrammica, once situated in a suit-
able site near a source o f prey, have the potential to
occupy that site for many years, changing size as hab-
itat conditions change. Clones of A. eleganti~sima,
established by longitudinal fission, are also likely to
persist indefinitely given normal levels o f disturbance
(Sebens 198%). In fact, the likelihood of a given dis-
turbance completely removing a well-established clone
containing several hundred individuals or more is re-
mote.
I f this general impression of anemone population
characteristics is true, monitored areas or populations
might be expected to show little change. Important
occurrences such as migration, mortality, and individ-
ual growth could happen on a scale far too long for
reasonable study. An experimental perturbation, how-
ever, can bring about radical changes in the rates of
such processes. T o that end, these investigations o f
population characteristics and habitat suitability in-
Ecological Monographs
Vol. 53, No. 4
clude both control population monitoring and experi-
mental manipulations.
Population characteristics including size-dependent
fecundity (Sebens 1981b), growth rates, immigration
rates, sizes of immigrating individuals, disappearance
of individuals from a given area, and timing of recruit-
ment were investigated by a combination of quadrat
sampling, individual marking, photographic mapping,
removal, and transplant experiments. Experimental
manipulations and monitoring programs were used to
analyze: ( 1 ) individual growth and seasonal size
changes in control and experimentally altered areas,
(2) size and season at which asexual division occurs
in mapped A. elegantissima and rates of clone growth,
(3) recolonization rates into experimentally cleared
areas, (4) mortality rates, and (5) differences in such
characteristics as a function of habitat type. The in-
formation on population dynamics from unaltered con-
trols and from experiments is ultimately used to ex-
amine the concepts of habitat selection and habitat
suitability as they relate to anemone populations. The
study used portions of the populations located in both
extreme and more normal habitat types to compare
expected reproductive success.
Anthopleura xanthogrammica (Brandt 1835) is com-
mon from Alaska (and possibly Japan, Uchida 1938)
to Baja California(Hand 1955) in close association with
the Mytilus calijornianus-dominated community on
exposed coasts. It occurs as small individuals (0.65.0
cm diameter) within mussel beds (Sebens 1981c, 198%)
and as large individuals (to 25 cm diameter) in surge
channels, tidepools, and rock walls below mussel beds
(Fig. 1). It continues subtidally to at least 15 m , di-
rectly below intertidal areas with M. californianus, but
is absent from isolated rocks and submerged pinnacles
that lack such an intertidal zone (Washington State,
Sebens 1977).
A. xanthogrammica is dioecious and spawns in ear-
ly autumn (Sebens 1981b). Fertilization and subse-
quent larval development to a swimming plankto-
trophic planula larva were described from Washington
State specimens (Siebert 1974). Dayton (1973) con-
ducted experiments demonstrating that mussel disturb-
ance during foraging by the seastar Pisaster ochraceus
increased prey capture (primarily Mytilus californianus)
by anemones in an experimental seastar addition area
and that prey escape response to the presence o f the
seastar Pycnopodia helianthoides increased the num-
ber of sea urchins (Strongylocentrotus purpuratus)
captured by the anemones. Although A. xanthogram-
mica possesses acrorhagi morphologically suitable for
agonistic behavior (as occurs in A. elegantissima), they
are observed in use only rarely (Francis 1973b).Func-
tional morphology and the relationship between me-
chanical properties of the anemone and water flow have
December 1983 SEA ANEMONE HABITAT SUITABILITY

F I G . 1 . Anthopleura xnnthograrn~nica(Tatoosh Island, Washington). (A) Expanded individuals in a low intertidal pool
with coralline algae and the hydrocoral, Allopora crtlifornica (scale 10 cm). (B) Crowded individuals on the floor of a surge
channel (scale 10 cm). (C) Individuals on a vertical wall just below a mussel bed and in a low tide pool. This is on the
permanently shaded side of a rock wall (scale 10 cm). (D) The exposed side of the same rock wall as in C (2-3 m distant).
Note that the anemones are present but have migrated farther downward into the pool (scale: 10 cm).

been investigated by Koehl (1976, 1977u, b, c ) . Else-
where (Sebens 1981c, 1982c) I have reported on rates
of recruitment into cleared areas, and Batchelder and
Gonor (1981) have described the population structure
of A. xunthogvutntnicu in Oregon.
The smaller anemone, Anthopleziru eleguntissimct
(Brandt 1835), (Fig. 21, is morphologically similar to
A. xunthogrummicu, differing primarily in the ar-
rangement of adhesive verrucae on the column and in
the relative thinness of the mesoglear layer (Hand
1955). A . eleguntissitnu is dioecious and shows a clear
annual reproductive cycle, spawning primarily during
late summer and early fall (Ford 1964, Sebens 1977,
198 lb, Jennison 19790, b). Spawning is epidemic, ini-
tiated by males; fertilization and development of the
planktotrophic planula larva occur externally (Siebert
1974). Francis (19730, b) determined that the aggre-
gations observed in the field were true clones and that
the anemones possess structures (acrorhagi) capa-
ble of inflicting damage (tissue necrosis) on other in-
dividuals. Individuals are able to recognize clonemates
from nonclonemates and engage in agonistic encoun-
ters only with the latter, resulting in an anemone-free
strip at the boundary of two adjacent clones. Individ-
uals along borders in contact with other clones devel-
op much larger and more numerous acrorhagi than do
internal clonemates. This may be a costly alteration
resulting in reduced individual gonad production com-
pared with internal clonemates (Francis 1976). Single
clones can occupy pools and channels over areas of
 KENNETH P. SEBENS Ecological Monographs
Vol. 53, No. 4

F I G . 2 . A n t h o p l e ~ o uelegilntissiinn from various habitat types along the Pacific coast. (A) A clone of small individuals in
a depression at an inner coast site, Iceberg Point, Lopez Island, Washington (scale: 1 cm). (B) A large contiguous clone in a
protected location just below a mussel bed at an outer coast site, Tatoosh Island, Washington (scale: 10 cm). (C) Extremely
large clonal individuals on a boulder at the mouth of Bodega Bay, California. The area, located behind a breakwater, appears
to collect floating material which may be an important food source (scale: 10 c ~ n )(D) . Clonal individuals with much adherent
shell material (C) and large solitary individuals (S) in a channel below mussel beds at the moderately exposed outer coast
site, Arroyo Hondo, California. Note the spacing between large solitary individuals (scale: 10 cm).

many square metres (Sebens 198%). I n southern Cal-
ifornia, A . elegc~ntissiinn is a n important community
member and competes with the tubeworm Phrc~grna-
topoina crrlijorniccl for space in areas experiencing
temporary sand burial (Taylor and Littler 1982).
Desiccation very likely sets the upper limit to inter-
tidal distribution of both species. Under moist winter
conditions, wandering A . elegc~ntissirnacrawl onto ex-
posed surfaces where they are later removed by des-
iccation during summer months when low tides and
midday sun coincide (Dayton 1971). Shell debris and
gravel retained on the anemone column by the adhe-
sive verrucae give some protection from desiccation
(Roberts 194 1, Hart and Crowe 1977).
Both A . xantlzog~.nrntnicaand A. elrgantissiinn con-
tain symbiotic algae (zooxanthellae: Sytnbiodiniutn sp.
or zoochlorellae [undescribed sp.] [McLaughlin and
Zahl 1966, O'Brien 19801). Individuals with the sym-
b i o n t ~lose mass more slowly during starvation than do
those without the symbionts (Muscatine 1961) o r those
kept in darkness (Sebens 1980), and the symbionts are
necessary for positive phototactic behavior (Pearse
1974, Fredericks 1976).
A . xrrntlzogrnrnrniccr is absent from a series of sites
in southern California but appears again in Baja Cali-
fornia, where coastal upwelling produces lower tem-
peratures and the community composition is more like
that in Oregon and Washington (Sebens 1977). A large
solitary form of A . rlrgantissiinn (to 20 cm basal di-
ameter; Fig. 2) occurs in the lower intertidal and main-
tains clear space around itself by agonistic behavior
(Francis 1979, Sebens 19810). This form was found
from Baja California north to at least Arroyo Hondo,
California, but no further, corresponding roughly to
the zone of higher surface water temperatures. The
large solitary form, which feeds primarily on mussels
(Sebens 1981a ) , presents several interesting questions.
I t may b e one of the clonal forms that has ceased to
undergo longitudinal fission (Sebens 1979, 1980), or it
may be a different morph or variety of the species that
lacks the ability to divide (Francis 1979).
Tatoosh Island (48"24'N; 124'48'W) (Figs. 3, 4) is
an extremely exposed outer coast site protected from
human disturbance (United States Coast Guard p r o p -
erty). Within this cluster of islands and rock platforms,
there is a gradient of exposure from the most exposed
December 1983 SEA ANEMONE HABITAT SUITABILITY 409

southwest (Strawberry Island) and northwest (North

Island) points, to the most protected areas around the
central sheltered beach (Fig. 3B). The community is
dominated in the middle intertidal by large beds of the
mussel Mytilus cillifornitltzus ( t 0 . 9 to t 3 . 4 m) (Paine
1974), often with a band of IW. edulis along the upper
border (Suchanek 1978) and with patches of algae and
Semibalirnus cariosus in recently cleared areas.
A well-developed algal association occurs below the
mussel bed, dominated by the large perennial kelp
Lessoniopsis littornlis. Below the kelp canopy the rock
surface is occupied by A. xantlzogramtnica, hydro-
corals (Alloporc~ccllifornica), sponges (Halichondria
spp.), bryozoans, and hydroids. On the exposed points,
A. eleguntissima occur within mussel beds as solitary
individuals a t densities up to =500 individuals/m2 (Se-
bens 1982~).I t is rarely found in the exposed tide-
pools, channels, or on rock walls below the mussel
beds.
A . xatzthogrntnlnicn often carpets the floor of tide-
pools and surge channels in exposed areas (Fig. 1).
Small individuals are present within the mussel bed,
and individual size increases from the lower edge of
the mussel bed, where anemones are =3-10 cm in di-
ameter, to the subtidal areas, where they can get as
large as 33 cm across the basal disc. Small individuals
(<4 cm) are found clustered around the bases of larger
ones and on the sidewalls of pools in areas where large
anemones fill the available central space. A . xatztho-
grrimmica density and the presence of small individ-
uals both decline rapidly with increasing distance from
the extensive mussel beds toward the more protected
areas.
I n more protected locations on Tatoosh Island
Selnibalatzus cariosus dominates much of the upper
intertidal (+ 1.2 to +2.1 m), interspersed with large
clonal aggregations of A. elegclniissirna (Fig. 2), and
with Baltrnus gland~flii,F ~ f c u sclistichus, and other
macroalgae. Mussel beds are fewer and occur in iso-
lated patches. The algal association is composed pri-
marily of Hedophyllum sessile and Lavlzinariu setch-
ellii. The few A . xiiizthogramtnictr are in the low
intertidal and subtidal. F o r a more complete descrip-
tion of the exposed outer coast community along the
Olympic Peninsula and the inner coast areas of San

FIG. 3. A. Study sltes on the outer coast of the Olympic

Peninsula and in the San Juan Archipelago (map modified
from Dayton 1971). Tatoosh Island (TI), Mukkaw Bay (MB),
Shi Shi Beach (SS), Iceberg Point (IP), Iceberg Cove (IC),
Cattle Point (CP), Eagle Point (EP), Friday Harbor (FH). 4B
is the location of the photograph in Fig: 4B, taken facing
west.
B. Tatoosh Island, Washington. Stippled areas are subject
to heavy wave action (exposed locations). Study sites and
other important locations on the island are indicated: A . xrrr7-
thogrrrnnnicc~transplants (TR), A. rcinthogrnmmicci mapped
areas ( M I , M2, M3), A. scrnthogrc~mmicciremoval areas (1-
lo), A . elegrintissimrr quadrat survey area (AE), Dermclste-
rins experiments (DE), beach draw area (BD), Cape Flattery
Light Station (CFL). (Map drawn from several aerial pho-
tographs taken by the author.) 4A is the location of the pho-
tograph in Fig. 4 A, taken facing northwest.
410 KENNETH P. SEBENS Ecological Monographs
Vol. 53, No. 4
and pools that dissect the bench area (to +1.38 m)
although individual anemone size and population den-
sities are much lower than on the exposed points of
Tatoosh Island. A . elegantissinza form large clonal ag-
gregations similar in extent and appearance to those
in the protected Strawberry Draw and South Draw
areas on Tatoosh Island (+0.57 to + 1.64 m, Fig. 2).
Some of the largest A. elegantissima (7-8 cm diame-
ter) are present at the same level as A. x-anthograrn-
mica, just below the mussel beds.
At the enclosed sound (inner coast) sites of the San
Juan Archipelago, A. xanthogrammica drops out al-
most entirely except for a few isolated individuals at
moderately exposed areas such as Eagle Point, San
Juan Island (48'27'N; 123"02'W), and Iceberg Point,
Lopez Island (4S025'N; 122'53'W). The same is true
of Mytilus calijornianus. A . elegantissima is very
common and can usually be found as clones or isolated
small individuals on any of the protected shores. It
reaches its largest individual size and population den-
sity on the moderately exposed points such as Eagle,
Iceberg, and Cattle Points (48'27'N; 122"57'W), San
Juan Island (Figs. 3, 4).
Such areas are also notable for dense populations
of the large barnacle Semibalanus cariosus in the mid-
dle and upper intertidal (0 to f 2 . 0 m). Dayton (1971)
suggested that S. cnriosus and A. elegantissima com-
pete for space in such areas. Anemones, barnacles,
FIG. 4. A. An exposed surge channel at Tatoosh Island, and the few mussels are limited to concave surfaces
Washington, with a well-developed mussel bed (mb), and al- (Fig. 2) since drift log scraping and summer exposures
gal zone (az). Depressions along the lower edge of the mussel keep raised surfaces cleared except for annual blooms
bed contain At~thop/eur(i~ut~thogr(irntnicu and the bottom of of ephemeral algae. Drift log scraping appears more
the channel itself is carpeted with them. Anthopleura ele-
gnntis~itna is common within the mussel bed. The photo- important in such locations than on the open coast,
graph was taken September 1974. since logs collect in the upper intertidal of inner coast
B. A moderately exposed inner coast location, Cattle Point, sites and are moved around by very high tides, scrap-
San Juan Island, facing the Strait of Juan de Fuca. Antho- ing over intertidal surfaces (Dayton 1971). In the
pleurri elegantissirnn is abundant and limited to channels and
depressions among the barnacles, Setnibalur~uscnriosus. A . lower intertidal (below -0.3 m), Hedophyllum sessile,
snnthogrummica is absent. Note the large concentration of Laminnria setchellii, Costaria costata, and a variety
drift logs which float periodically at very high tides and scrape of other algal species cover the surfaces, and both
the intertidal. The photograph was taken September 1975. exposure and drift log scraping are of lesser impor-
See Fig. 4 for the exact locations of the areas in these two
photographs.
tance.
Iceberg Point, Lopez Island, was chosen as a study
area because of the dense A. elegantissima popula-
tions on distinct upper and lower intertidal bench areas.
Juan Archipelago, Washington State, see Dayton (1971, A . elegnntissimn is abundant from + 1.9 m to a sharp
1975). lower boundary at +0.4 m and ranges in size from
The north end of Shi Shi Beach (48"17'N; 127"411W, aggregations of small individuals (0.8 cm mean diam-
Fig. 3B) is a gently sloping bench and boulder area eter) on the high bench to aggregations of larger in-
and is therefore less wave exposed, conforming to the dividuals (1.8 cm mean diameter) at the lower bound-
description of a protected outer coast site (Ricketts et ary. The area is exposed to westerly swells from the
al. 1968). Mussel beds are limited to the tops of several Strait of Juan de Fuca and is thus a relatively exposed
large vertical protrusions of the bench (+ 1.35 to + 1.64 inner coast site. Around the point to the north, Iceberg
m). Because of the low inclination of the bench, the Cove is protected from the heavy wave exposure and
algal zone (Hedophyllum s e ~ s i l eand Laminuria setch- is shaded by a vertical cliff to the south. This unique
ellii) begins much farther seaward than at Tatoosh Is- site has a group of high intertidal pools (+ 1.5 to f 2 . 1
land, where it is directly below the mussel beds. Cer- m) where the temperature remains essentially the same
tain tide pools harbor dense urchin populations as that of the surface water, and a series of mid-inter-
(Strongylocentrotu~purpuratus). tidal pools and channels (+0.5 to + 1.3 m) which are
A . santhogrammicn occurs in the many channels not shaded and can be several degrees warmer than
 December 1983 SEA ANEMONE HABITAT SUITABILITY 411

the surface water when exposed. This site also has a (Strawberry Draw area, high + 1.8 to t 2.5 m , low + 1.1
very high diversity of anemones (nine species, Sebens to + 1.4 m ) and 12 in the San Juan areas (Iceberg Point,
1977). Cattle Point, high + 1.6 to + 1.9 m , low +0.2 to +0.8
Cattle Point is almost identical to Iceberg Point in m ) , with three replicates for each combination of vari-
community composition but the study site faces north, ables (intertidal height, concavity, exposure).
exposed only to waves that turn the point. A . elegan- T o begin monitoring an experimental clone, the un-
tissirna occurs from +0.2 to + 1.8 m in a pattern similar altered condition was photographed and a central group
to that at Iceberg Point. The San Juan study sites, of 15 individuals was chosen as the nucleus to be stud-
ranked from most to least exposed by wave action ied. All anemones surrounding the nucleus within a
(noted on visits from July 1973 to March 1977) are: radius of 30 cm were measured and removed. Clones
Iceberg Point, Cattle Point, Iceberg Cove. For a de- were situated in topographically isolated areas, pools,
scription of the other anemone fauna in this area see channels, or rock faces, surrounded by areas uninhab-
Kozloff (1973, 1974) and Sebens and Laakso (1977). ited by anemones so that immigration was unlikely. A
map o f the isolated nucleus was drawn, each individ-
ual measured, and the whole aggregation photo-
graphed. A small yellow enamel paint spot was applied
Anthopleura elegantissima population monitoring
to the rock surface near the clone to aid relocation.
Quadrat sampling was used to determine the size- At each subsequent visit, all anemones found in the
frequency distribution of Anthopleura elegantissima cleared area around the edge were measured and re-
populations at each study site over a 2-yr period. Such moved so as not to be confused with those in the nu-
sampling was necessary to determine whether or not cleus. The anemones' tendency to aggregate (possibly
changes in mean individual size were the result of sig- as desiccation protection or to defend acquired space)
nificant changes in the distribution of individuals over meant that anemones within the nucleus remained there
size classes. Massive settlement of larvae and subse- and shifted position only slightly if at all between sam-
quent growth, or the removal of large individuals (by pling periods.
mortality or asexual division) could be indicated by Each clone was mapped and compared to the pre-
such sampling. vious map, anemones were measured, and the clone
High and low intertidal areas at Tatoosh Island (+ 1.8 was photographed (with suitable scale) at each visit.
to +2.5 m and +0.5 to +1.1 m ) , Iceberg Point, Lopez Anemones in the process of longitudinal fission or those
Island (+ 1.8 to + 1.5 m and +0.5 to +0.9 m ) , Iceberg recently divided (with division scars) were noted. At
Cove, Lopez Island (+0.5 to + 1.2 m ) , and Cattle Point, the end of the study, tracings were made from pho-
San Juan Island (+ 1.7 to +1.8 m and +0.3 to f0.9 m ) tographs o f the clones using an overhead slide projec-
were sampled bimonthly (March 1974 to May 1976) tor on a f exible arm (adjustable for parallax and scale),
and monthly during the period o f March to September anemone measurements were noted on the maps, and
1975. Blind throws of a 10 x 10 c m wire quadrat were field maps were used to verify the number o f anemo-
taken in each defined area (30-50 m2) and basal di- nes and the identity o f individuals in the photograph.
ameters of all anemones within the quadrat were mea- Anemones noted as dividing or having divided in the
sured. This was repeated until 150 anemones had been field were located on the maps, and any large anem-
sampled. ones replaced on the photo from the next visit by two
smaller ones in the same position (without other rear-
Mapped clonal aggregations of rangement in the clone) were also considered to have
A. elegantissima undergone fission. From this information clonal growth
Although individual marking of A. elegantissima is rates (AN) and clonal biomass changes (from ash-free
possible (Sebens 1976), it is exceedingly laborious in dry mass to basal diameter regression [Sebens 1979,
the field. However, given their tendency to aggregate, 1981~1) were calculated. Numbers and sizes o f individ-
it was possible to monitor individual clones (and in- uals migrating inward at the edge o f the clearing and
dividuals within clones) by long-term field and pho- of individuals missing from the nucleus were recorded.
tographic mapping. With this technique, the following Timing of asexual division and size prior to and fol-
were examined: ( 1 ) size changes o f individual anem- lowing division were also determined.
ones, (2) division of individual anemones (size and
seasonality), (3) actual clonal growth rates (number, Anthopleura xanthogrammica [napped areas
biomass) in a variety o f habitat types, (4) immigration Three areas on Tatoosh Island were chosen as un-
and settlement into the cleared space around clones, altered control populations in which to examine indi-
( 5 ) emigration from the experimental clone. vidual size changes, immigration, emigration, and in-
Clones to be monitored were selected to represent dividual movement using marked individuals. The three
the range o f high and low intertidal distribution, flat areas represent a range of habitat characteristics (in-
or concave substratum (susceptibility to desiccation tertidal height and distance from mussel bed prey
and log scraping), and inner or outer coast location. source). All individuals at the site were within well-
Twelve clones were designated on Tatoosh Island protected concavities in the substratum. The areas
412 K E N N E T H P. S E B E N S Ecological Monographs
Vol. 53, No. 4
chosen were: (1) high intertidal exposed point (Straw- mapped, measured, photographed, and given dye spots
berry Island, +2.0 to +2.6 m), (2) low intertidal ex- as they entered the area. Rates of immigration are giv-
posed point (Strawberry Island, + 1.4 to + 1.8 m), (3) en in Sebens (1981c, 1982~).Each immigrant was
low intertidal protected area (Strawberry Draw, +0.6 mapped and measured on every subsequent sampling
to + 1.0 m) distant (20 m) from mussel beds. Workable visit. Growth increments could thus be compared to
areas containing 7&100 individuals were delimited and those for individuals in the three control areas by the
mapped by sketch and photographic mosaic during method outlined for those areas.
September 1974. Each anemone was mapped, mea-
sured, and given a dye spot with neutral red vital dye
paste (Sebens 1976a). The dye was left on for 20 min, Size distributions in control areas (both species) and
after which excess was washed off with sea water. Dye in experimental clones of Anthopleura elegantissima
spots persisted at least 12 mo and the treatment was were compared using analysis of variance (ANOVA),
repeated after substantial fading had occurred (Sep- followed by Student-Newman-Keuls (SNK) multiple
tember 1975). However, because of high wave expo- comparisons test if there were more than two samples
sure, there were several visits when it was impossible (Sokal and Rohlf 1969). Growth increments for each
to monitor the areas. Successful monitoring averaged size class of Anthopleura xnnthogramtnica were com-
trimonthly (more often in the spring and summer and pared by ANOVA and SNK tests between the three
less often in the fall and winter). At each monitoring control areas and the removals. Assumptions of the
visit, the anemones were located, compared to the ANOVA were tested according to the methods out-
previous visit's map, measured, and the area was pho- lined in Sokal and Rohlf (1969) (independence, nor-
tographed. New arrivals (without the dye spot) were mality by examination of size-frequency distributions
noted, usually in locations not previously containing for skewness, and homogeneity of variances by the
an anemone. Disappearance was recorded when an F,,, test).
individual was missing from a previously mapped lo- Pearson's product-moment correlations were used
cation during all subsequent sampling visits and no to compare sizes of anemones in clones to those that
individual matching its general size and dye spot could underwent division, and principal axis lines were de-
be found in a nearby location within the mapped area. termined (Sokal and Rohlf 1969). Regressions were
Such disappearances indicated that an anemone had performed on division frequency, clone biomass
either died or moved out of the area completely (usu- change, gonad production, and rate of change in gonad
ally a distance of several metres). Since anemones production as functions of mean individual size in
larger than 6 cm diameter move very rarely and over clones of A. elegnntissima (Sokal and Rohlf 1969).
very short distances, the former is more likely. How-
ever, smaller anemones are capable of much greater
movement and are more likely to relocate (Sebens A. elegantissima population monitoring
1981c, 1 9 8 2 ~ ) Disappearances
. of smaller anemones The populations in all quadrat survey areas showed
probably included mortality and some emigration. In- a seasonal change in mean individual size with a max-
dividual size and growth increments were calculated imum size in July-August of each year and minimum
over the 2-yr period (September 1974 to October 1976) during January-February (Fig. 5). Size-frequency his-
for individuals of all size classes. For each anemone tograms (Fig. 6) indicate that all size classes shifted
measured at a sampling visit, the size after I yr was re- upward or downward rather than changing by influx
corded. If the I-yr period ended between two sampling or removal of individuals of particular sizes. The pe-
visits, final size was calculated as an interpolation be- riod of size increase (April-July) coincides with known
tween sizes at the two visits. For each anemone in the increases in zooplankton abundance (Johnson 1932,
study, up to nine such annual growth increments were Hebart 1956, Tully and Dodimead 1957, Hameedi 1974,
calculated. Winter et al. 1975), particularly of settling invertebrate
An area at Shi Shi Beach was chosen for A. xan- larvae on which the anemones feed (Sebens 1 9 8 1 ~ ) .
thogrnrnmicn size monitoring which included groups During August to March of each year the mean diameter
of individuals on a horizontal high platform (+ 1.2 m), of individuals showed a general decrease of 10-40%.
an adjacent vertical wall (+0.4 to +0.9 m), and a low The initial rapid size decrease (August-October) coin-
channel below the wall (-0.4 m). The nearest mussel cides with spawning (Sebens 1981b) (also see Ford 1964,
bed was =30 m distant. Areas to be monitored were Jennison 1978, 1979a, b, for spawning periods in Cal-
designated on a map and included 50 individuals each ifornia). Part of this decrease may also be accounted
for by fission within the population.
A. xanthogrammica removal areas There is some indication that the onset of size de-
All A . xnnthogratntnica were removed from 10 tide- crease is earlier on the outer coast. Note also that
pools and portions of channels on Tatoosh Island. Each there is a sharp temporary size decrease during May
removal consisted of 14-36 individuals from areas ~ 0 . 5 1975 that appears in all of the San Juan area samples
m2 each. New arrivals in the cleared areas were but not in those from Tatoosh Island. This correspond-
December 1983 SEA A N E M O N E HABlTAT SUITABILITY 4 13

j e r q Cove. iow 1 I 1 1 Tatooih Island I 1

FIG.5. Seasonal size changes in high and low intertidal populations of Attthopleur.ci elegantiaairnci at Iceberg Point, Lopez
Island. Cattle Point, San Juan Island, Tatoosh Island, and the low intertidal population at Iceberg Cove. Lopez Island.
Populations in the middle intertidal mussel bed at Tatoosh Island are also included. Each point is the mean basal diameter
( t s ~ in
) quadrat samples of 150 individuals. Solid lines at the bottom of each graph depict significant differences between
time periods. Samples above the end points of any one of these lines are significantly different from each other; all other
samples spanned by the line are not significantly different by ANOVA and Student-Newman-Keuls (SNK) tests (P < .05
at least): the end points thus indicate periods of significant size change. H = high intertidal, L = low intertidal, M = middle
intertidal mussel bed samples. Note that the ANOVAs were carried out o n the entire samples of 150 individuals in each
sampling period for 2 yr and that S N K comparisons were made only within a particular tidal height and site.

ed to a week of abnormally high temperatures in the
San Juan areas.
Included with the high and low intertidal samples
for Tatoosh Island are results of 10-cmQuadrat throws
within a mussel bed located between the Strawberry
Draw high and low population sampling areas (Fig. 5).
They show a general increase in mean size (individual
growth) over the sampling period (see also Sebens
198 lc,) rather than the fluctuations observed in anem-
ones on open surfaces. Whereas the low intertidal
population in the San Juans is composed of larger in-
dividuals, the Tatoosh Island low and high groups are
nearly identical in individual size. This may result from
the very protected habitats formed by the deeply dis-
sected high intertidal bench a t Tatoosh Island, which
keep the anemones cool and their metabolic rate low
(Sebens 1980). None of the histograms (Fig. 6) shows
a significant increase in the number of very small ( 2 -
4 mm) A . elegcrntissi/ncr which would occur following
heavy settlement.
.Mcrpp~dcloncil crggi.egcltions of
A. elegantissima
Experimental clones allow a comparison of individ-
ual and clonal size changes in several distinct habitat
types. Figs. 7 and 8 illustrate such changes as number
of individuals in the clone nucleus, biomass change of
the entire clone, and number of divisions occurring
between each two monitoring visits. Number in-
creased in 17 of the 24 clones, decreased in 3 and did
not change in 4. Biomass changes all show the sea-
sonal fluctuation already noted in the population sam-
ples, rapid size increases in the spring and decreases
beginning in late summer o r early fall. Longitudinal
fission occurred almost exclusively during periods of
individual shrinkage (reduction in total clone biomass
 KENNETH P. SEBENS Ecological Monographs
Vol. 53, No. 4

ANTHOPLEURA ELEGANT1 S S I M A
ICEBERG POINT
HIGH LOW

k 10 JAN 76 10 MAR 7 6

0 1 2 3 4 5

;r:L
0
16 MAY 76

I 2 3 4 5

BASAL DIAMETER ( cm )
FIG.6 . Size-frequency histograms of anemones in high and low intertidal populations of Anthopleura eleyantissima at
Iceberg Point, Lopez Island. Mean diameter ( t s ~for ) each quadrat sampling date appears in Fig. 5. Note that changes in
mean individual size involve no alteration of the form of the size-frequency distribution but simply shifts to the left or right.

also), most often in the late fall just following the onset
of shrinkage (Sebens 1982). The larger individuals in
the population were the ones that subsequently divid-
ed (Sebens 1980, 1982). When the comparison is made
for individual clones (Fig. 9) it is evident that it is the
larger individuals in each clone, rather than the largest
in the population per se, that have divided (Sebens
1982). Size at division is strongly correlated with mean
individual size in clones (Fig. 10) and is thus a habitat-
dependent characteristic.
Clones located in cracks and depressions experience
enough microhabitat variability that individuals situ-
ated only centimetres apart experience important dif-
ferences in the physical environment, and possibly
feeding interference from clonemates. In the deeper
center portions of crevices, individuals are often larg-
er, having failed to divide at as small a size as did their
clonemates. If size at division depends partially on
variable physical conditions (Sebens 1979), microhab-
itat heterogeneity could be the cause of the high with-
in-clone variability in size at division.
Of the 113 daughter individuals produced during the
first season of the experimental clone study, 8 divided
again during the second season. In no case did a sec-
December 1983 SEA ANEMONE HABITAT SUITABILITY 415

THE l
p 1 THE 7 1 ~ 0

THP l 30 1 THP5 THP 6

20

0
IL
0
S
4 -

j J ~ i ~ ~ u j j ~ 0 ~ F " Jh1974
J ma F Mh A M
1975
J j % & D J F M A M O O JJASONDJFMAMJJASONDJFMAM
1974 1975 1976 1976 1974 975 1976

TLE l

- 10

TLP 3 TLP 5 30 y
+)
/
20

O - O - - O - O - O - O - - ~ ~ o

J J A S b N D J I F M ~ M J J i s a ~ i * O JJiSONbJ!MAMjji;'ibhbjkhih O O J ~ ~ ~ O N D J ~ ~ A M ~ ~ AO ~ O N O J ~ M
1974 1975 1976 1974 1975 976 1974 975 976
 KENNETH P. SEBENS Ecological Monographs
Vol. 53, No. 4

[ SHE 8

lot
SHES
f[;
, I
120

SLE I

130
SLP 2
I0 I

FIG.8. Biomass changes, numerical changes, and divisions in numbered experimental clones of Anthopleura elegnntissimn
in San Juan Islands (S) high (H) and low (L) intertidal sites. Legend as in Fig. 7.

ond division follow during the same season as the first.
Clonal growth rates (divisions per individual per year)
were quite variable, although divisions occurred in all
but three of the experimental clones. The San Juan
high exposed areas and the Tatoosh low exposed areas
(the most and least desiccation-stressed areas, respec-
tively) had the lowest division frequencies. These areas
also had the very smallest, and the largest, mean in-
dividual sizes, respectively.
Immigration rates (anemones per square metre per
month) (Table 1) indicate that there are wandering
anemones available for immigration in all areas a t all
times of the year. The lowest immigration rates appear
to be in the fall and winter months, but this is com-
 December 1983 SEA A N E M O N E HABITAT SUITABILITY 417

plicated by the fact that monitoring visits were less

frequent during those months and wandering individ-
uals may have moved in and out again before the next
visit. Spring and summer have the highest rates, pos-
sibly because desiccation drives anemones into the
more protected sites. There is also a large peak im-
mediately following creation of the experimental areas,
which probably represents a response to the newly
cleared space by bordering anemones, subsequently
removed. The sizes of immigrating individuals (1.2 r
0.3 cm SD, N = 262) are significantly smaller than sizes
of anemones within the experimental clones (1.6 r 0.4
cm SD, N = 432), or the population in general (AN-
OVA, F, = 195, P < .001,df = 1,692).
Disappearances from the clone nuclei were not com-
mon and were not seasonal (Table 2). The sizes of such
individuals (1.6 & 0.3 cm SD, N = 35) were not sig-
nificantly different from those in the experimental
cones, or the population in general (ANOVA. F , = 0 I 2 3 4
3 x P > .05, df = 1, 465). Disappearances were
CLONAL INDIVIDUAL DIAMETER (cml
more common in the two high intertidal areas than in
the low areas, but there was a total of only 35 disap- FIG. 10. Mean diameter of individuals dividing in a given
pearances among the 787 anemones monitored ( ~ 2 . 2 % year (U,) plotted against the mean diameter of all clonal in-
disappearance per year). dividuals (D) at the annual peak of individual size. The cor-
relation is highly significant ( r = .94, P < ,001) and the prin-
+
cipal axis line is U, = 1.1 10 0.097. The 95% confidence
limits for the slope are 1.30 and 0.97.

lor TAT HIGH PR r I

The formation of experimental clones involved a lo-

cal density reduction. The presence of surrounding
clonemates may reduce prey availability to each in-
dividual and thus inhibit individual growth. During the
period July 1974 to May 1976 in the San Juan areas,
and September 1974 to May 1976 a t Tatoosh Island,
101 TAT L O W PR r r
there was no significant change (ANOVA, P > .05) in
mean size for control population samples (Table 3).
However, mean individual size increased significantly
(ANOVA, P < .05) in 8 of the 24 clones and de-
creased in only I (Table 3). Size increased to a lesser
degree in 12 more clones. Biomass and number of in-
dividuals also increased in most clones (Figs. 8, 9),
but it is not certain that there were not corresponding
changes in unmanipulated areas. I t was not possible
to keep track of clone growth in crowded, contiguous
clones.
Anthopleura xanthogrammica
population monitoring
There appears to be a seasonal pattern of A . ,win-
thogrammicu size change similar to that in A . elegun-
tissitnu populations, but of reduced magnitude (Fig.
1 1). Largest sizes were observed during the summer
O 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 months and smallest in winter and spring. Histograms
DIAMETER ( c m ) (Fig. 12) indicate no major seasonal changes in the
size-frequency distribution of the populations. Nearly
FIG.9. Experimental clones of Anthopleurtr elagtrrltissi-
mcl; size-frequency histograms of all individuals at both an-
all large individuals occupied their exact locations over
nual peaks of individual size (1974, 1975) indicating those the entire 2-yr study period. The rest of the population
individuals that divided in the subsequent year (dark bars). consisted of smaller and more mobile individuals.
418 K E N N E T H P. S E B E N S Ecological Monographs
Vol. 53, No. 4
TABLE1. Immigration rates of Anthopleura elegantissima .nental removals had growth rates significantly greater
into cleared areas surrounding experimental clones (1974- than the grouped controls in 6 of 11 size classes. I n 5
1976), expressed both as individuals per square metre per
month and as individuals in the entire area around six clones of the size classes they also differed from one or more
(-0.40 m2 area) since the last sampling visit (in parenthe- of the individual control areas. The mean final size
ses). Totals are immigration rates over the 2-yr period. drops to the zero growth isocline a t 8 cm in map area
2, a t 11.5 cm in map area 1, and a t 12.6 c m in map
Anthopleura elegantissima immigration area 3. These are the asymptotic sizes (size plateau)
San Juan San Juan Tatoosh Tatoosh for individuals in each of the control areas. There is
Date high low high low no plateau for individuals in the removal areas, be-
N .,,-2. mo-l cause even the largest individuals had not reached their
(total anemones since last sample) final size ( > I 5 cm) during the study.
June 1974 ... ... ... ...
Growth trajectories, plotted in Fig. 14, were calcu-
August 1974 14.0 (10) 10.0 (7) ... 11.3 (10)
September lated by summing growth increments (g [D,], Fig. 13)
1974 33.6 (12) 19.8 (8) 7.7 (3) 24.9 (11) over n years:
February 1975 5.1 (9) 2.5 (5) 1.1 (2) 4.1 (9)
April 1975 5.6 (4) 1.2 (1) 2.5 (2) 3.3 (3)
May 1975 8.4 (3) 24.8 (10) ... .,.
June 1975 2.8 (1) 17.4 (7) 2.5 (3) 5.6 (5)
August 1975 11.2 (8) 6.2 (5) 10.4 (8) 9.0 (8) where D, is diameter at year x after settlement. Time
January 1976 3.3 (6) 5.4 (1 1) 3.6 (7) 4.9 (1 1)
March 1976 8.4 (6) 21.1 (17) 7.7 (6) 14.6 (13) for growth from settlement to 1 cm diameter has not
May 1976 8.4 (6) 13.7 (11) 2.5 (2) 11.3 (10) been measured and is assumed to be =2 yr based on
Totals 181.8 (66) 207.5 (83) 66.5 (33) 174.8 (80) data for Anthopleura elegantissima (Sebens 1 9 8 2 ~ ) .
Trajectories plotted using mean growth increments
(Fig. 14A) are probably correct above 6 cm diameter,
but below 6 cm they include the growth of many un-
At Shi Shi Beach, anemones on the high platform, successful individuals which may have died o r emi-
vertical wall, and low channels (Fig. 11) showed a grated. Plotting most rapid growth (Fig. 14B) as the
marked seasonal size change, increasing in diameter mean plus one standard deviation for the growth in-
during summer months and decreasing over the fall crement may illustrate trajectories typical of the fast-
and winter. Sizes of individuals on the high platform est growing third of anemones in all size classes. HOW-
and vertical wall were much smaller than in the low ever, it is probable that no single anemone attains that
channel. Size-frequency distributions indicate no
change in the size structure and no appearance of many
small individuals (recruits) over the study period. TABLE2. Disappearances ofAnthopleura elegantissima from
The numbers of individuals that appeared or dis- experimental clones (1974-1976). Disappearances are ex-
appeared from the areas were extremely small com- pressed as number of individuals disappearing from each
pared to the overall population, and were of similar clone (clone size was 15.3 5 1.6 SD individuals during the
1st yr and 17.9 2 4.1 SD individuals during the 2nd yr) per
size in the three areas (Table 4). Immigration rates month and as the total disappeared since the last sampling
(number per square metre per year) in the two exposed visit. Average monthly disappearance values are given for
areas were indistinguishable, but in map area 3, where the 2-yr period, followed by the total disappeared for the
entire period (in parentheses).
anemone density was much lower, rates were lower
than in the other two areas (chi-square test, P < Anthopleura elegantissitna
.05). disavvearance from clone nuclei
Growth increments of individuals in the map areas
San Juan San Juan Tatoosh Tatoosh
grouped by size classes (each 1 c m diameter) are given Sampling date high low high low
in Fig. 13 and are compared to growth in experimental
N .clone-' mo-I
anemone removal areas. Final size for each size class (total N since last sample, for six clones)
was compared by ANOVA among the three control June 1974 ... ... ... ...
areas and the removal area, followed by a multiple August 1974 ... ... ... ...
comparisons test (SNK). The removals were also September
1974 0.33 (2) ... ... ...
compared to the three controls grouped, since remov- February 1975 0.17 (5) ... ... ...
als spanned the area between the controls. Of the two April 1975 0.08 (1) ... 0.17(2) ...
exposed areas, map 3 (density = 7 anemones!mz) had May 1975 0.04 (1) 0.04 (1) ... ...
the greatest individual growth, while map 2 (density = June 1975 0.04 (1) 0.04 (1) 0.50 (3) ...
August 1975 ...~,
...~,
0.17 (2) ...
51 anernones!m2) had the least, and map 1 (density = Januarv 1976 0.03 (1) 0.17 (5) 0.17 (5) 0.03 (1)
20 anemonesJm3 was intermediate. Differences in March-1976 0.08 (1) ... ... ...
growth rate were significant ( P < .05 a t least) in only May 1976 ... ... 0.17 (2) 0.08 (1)
3 of 14 size classes between controls (7, 8, and 11 cm 2-yr average
(total) 0.087 (12) 0.050 (7) 0.101 (14) 0.014 (2)
initial diameter, by S N K test). However, the experi-
December 1983 SEA ANEMONE HABITAT SUITABILITY 419

TABLE3. Sizes of Anthopleura elegantissirna at beginning (July 1974, San Juans; September 1974, Tatoosh Island) and end
of the monitoring period (May 1976) in experimental clones and in control populations.

Direction
of
1974 diameter (mm) 1976 diameter (mm) Direc- ANOVA signif-
tion of Fs icant
Area x * SD n x i SD n chanee (df = I . n-2) P chanee
Controls:
Cattle Pt. low
Cattle Pt. high
Iceberg Pt. low
Iceberg Pt. high
Tatoosh Is. low
Tatoosh Is. high
Ex~erimentalClones exposed; P, protected)
an Juans low
SL5E

SL2P
SL4P
San Juans high
SH4E
SH8E
SH9E
SHSP
SH7P
SH IOP
Tatoosh low
TLlE
TL2E
TL4E
TL3P
TLSP

Tatoosh high
THZE

high rate year after year. These growth rates are thus
likely to be higher than most of those experienced by
individuals in the population.
One useful compromise to provide a typical growth
trajectory is to use the most rapid growth of juveniles
(to 6.5 cm), assuming that only the fastest growing
animals survive, followed by the average growth of
adults, since 99.4% of those survived each year. This
comparison (Fig. 14C) yields trajectories intermediate
between the first two. Table 5 gives estimates of time
to maturity, time to the size plateau, and diameter at
the plateau, for the three control areas and the removal
area, calculated by the three methods outlined above.
Time to maturity differs by 2-3 yr between estimates,
but by only 0.5-1 yr between control areas. Individ-
uals in removal areas matured 1-2 yr earlier (4.5-6.5
yr) than in controls (5-8.5 yr).
Time to final adult size (plateau) differs even more,
by 7-9 yr, between control areas (11-20 yr by this
method). The final adult diameter decreased with in-
creased density (Table 5), with a difference of 2.4-4.4
cm between controls, and at least 7 cm between the
smallest final size in controls (8 cm) and the removals
(>I5 cm).
Growth of fish (reviewed in Ricker 1975, Jones 1976)
and gastropods (Frank 1965, 1969) have been modelled
using a Ford-Walford plot (Ford 1933, Walford 1946)
of initial size (x axis) to size 1 yr later Cv axis). The
intersection of a regression line through all such data
points and the y = x line estimates L,, the length as-
ymptote in the von Bertalanffy growth model (see
method in Ricker 1975, Jones 1976):
where L, is the length at time t , k is a constant, and
t o is the time when size would be zero. The slope of
the regression line estimates the value ck, and thus
length at any time can be calculated from the above
 420 KENNETH P. SEBENS
'CHANNEL
VERTICAL W A L L
-
----
-
CHAWWEL
TOP
ALL
1 possible by the iterative method). The projected size
' - "A1974
F4T N~
1975 1976
FIG. 11. A. Seasonal size changes of Anthopleurci xiin-
thogrtrmmicci (mean diameter rso) in three mapped areas at
Tatoosh Island (1974-1976) and in experimental anemone re-
moval areas. Each point represents 54 to 116 individuals in
each mapped area and 9 to 48 individuals in the removals.
Map area 1 is a mid-intertidal, high wave exposure area, map
area 2 is a low intertidal, high wave exposure area, and map
area 3 is a low intertidal, low wave exposure area. Note the
general increase in mean size of anemones that have migrated
into the experimental removal area.
B. Seasonal size changes (mean diameter t s o ) of Alltho-
pleura xnnthog~.atn~nicrrat Shi Shi Beach (1974-1976). Each
point represents 50 + 4 individuals. The three study areas
are on the same rock outcropping 30 m from the nearest
mussel bed. Note that there is a clear seasonal size change;
individual size increases during the spring and summer and
decreases over the fall and winter. At this location, where
mussels form a smaller part of the anemone's diet, A . xan-
thogrci~nmictiare generally smaller than at Tatoosh Island
and their seasonal size changes are very similar to those of
A . e1egcintis.sirna.
Solid lines at the bottom of each graph depict significant
differences between time periods. Samples above the end
points of any one of these lines are significantly different
from each other; all other samples spanned by the line are not
significantly different by ANOVA and Student-Newman-Keuls
(SNK) tests (P < .05 at least); end points thus indicate periods
of significant size change. ANOVAs were carried out on the
entire samples at each sampling period and the S N K test was
used to compare samples from different times within the same
site and intertidal height.
equation. This method suffers from several major
drawbacks. First, there is little reason to assume that
the data are arranged linearly and are appropriate for
Ecological Monographs
Val. 53, No. 4
such a regression. For example, a species may go
through an early period of very slow growth, produc-
ing an exaggerated sigmoid growth curve. Second, sta-
tistical comparisons between such regressions are pos-
sible only when there are extreme differences in slope.
It is not possible to differentiate between growth dif-
ferences within particular size classes. Third, if the
analysis includes all individuals studied, differential
mortality by size class can significantly bias the result
(Ricker 1975). Nonlinear modifications of the von Ber-
talanffy model can remove the first problem (Ebert
1980), but not the latter two.
The Ford-Walford method for fitting the von Ber-
talanffy model was applied to the data on A . .vnrzt/zo-
graininica annual size changes. The size plateau (di-
ameter L,), slope, and y intercept (estimate of size at
1 yr if size at birth or settlement is = zero [I-2 mm
here]) were calculated. These values along with the
coefficient of determination are given in Table 6. The
growth equation was also used to calculate time to
maturity at 6.5 cm diameter and time to 95% of L,.
By comparison with the values in Table 5, the Ford-
Walford method predicts diameter plateaus similar to
those calculated using average growth increments, and
also predicts a final diameter in the removal areas (not
plateaus (Table 6 ) differ from the directly observed
values (Table 5) by up to 2 cm, or 16% of the observed
value. Time to maturity is similar between the two
methods, but time to plateau differs by up to 65% from
the estimates using average growth (Table 5). Esti-
mates of size at I yr (1.3-1.4 cm) are probably high,
based on a comparison with field data (Sebens 1982).
The iterative method (Table 5 , Figs. 12 and 13) thus
gives a more accurate description of growth and allows
statistical comparison between areas. The method de-
pends on a large data set, however, whereas the Ford-
Walford method can give a rough estimate of growth
from a very few points. Any method that uses a regres-
sion to fit a particular growth model to the observed
data will probably give a reasonable depiction of the
growth curve, but cannot be as good a description of
growth as is the size-class comparison. Since this tech-
nique makes no assumptions about the form of an un-
derlying growth equation, it gives a description of
growth as it actually occurred in the population during
the study period. The method also allows the investi-
gator to compare either the average size change or the
size change of those individuals growing most rapidly
(any fraction of individuals within a size class). This
could be the upper third (as in this case) or a smaller
fraction if mortality in those size classes is very high.
The most crowded area (map area 2) had the lowest
growth rate, size plateau, and expected reproductive
success as measured by this analysis. As density de-
creased, the growth rate and size plateau increased.
How does intraspecific crowding affect growth rate?
One possibility is that probability of prey capture de-
creases with crowding. Mussels released from the
December 1983 SEA ANEMONE HABITAT SUITABILITY

ANTHOPLEURA XANTHOGRAMMCA
TATOOSH IS.
MAP AREA l

MAP AREA 2
1 28 MAR 75

MAP AREA 3
hl 5 M A Y 76 23 O C T 76

'Ol 2 9 SEPT 74 1 25 J A N 75 1 28 M A P 75 1 9 JUNE 751 22 JULY 75

4 SEPT 75 ( 2 F E E 76 / 5 M A Y 76 / 23 O C T 76

B A S A L DIAMETER (cm

FIG. 12. Size-frequency histograms of Anthop1rzir.a xclnthogrammica in mapped areas at Tatoosh Island (1974-1976).
Mean size and standard deviations appear in Fig. 11. Note that there is essentially no change in the form of the distributions
over the study period. Small individuals wandered into and out of the area while nearly all large individuals (>6 cm) remained
in place.

mussel beds by wave action or by predators are washed
into pools and channels where they continue to move
around until captured by an anemone or until they
lodge in a crevice. Clearly, more anemones in a pool
or channel will mean less prey capture per individual.
The following observation illustrates the effect of
crowding near the mussel bed prey source. In July
1973, 30 ~Mytiluscnlifornianus (3-4 cm length) were
collected and injected through the byssal gap with 1
m L each of concentrated neutral red dye solution. The
mussels were replaced onto the bed on the exposed
side of Strawberry Island during the same low tide and
left to b e washed away by the high tide wave action.
At the next low tide, all Anthopleura xnnthogrurnrnicn
in the vicinity were examined and 10 were found with
red tentacles where the dye had been released within
the coelenteron. Those 10 were within 2 m of the re-
lease point, and no others were found within a 6 m
radius of the release point. Thus, 33% of the released
mussels were captured within 2 m of the release point,
and the rest of the mussels were either washed into
the subtidal or into crevices and thus missed by the
anemones. The experiment was repeated in June 1975,
with 6 stained anemones observed in the same area.
In March 1976, 500 mussels, each =3 cm long, were
marked with a file groove across the byssal gap, frozen
to prevent reattachment, and distributed back into
mussel beds on Tatoosh Island. The first 100 anemo-
nes encountered within 2 m of the release point were
examined for mussels, which were then removed from
the coelenteron and checked for file marks. One
hundred anemones captured 134 of the 500 mussels.
The 50 closest to the mussel beds had captured 83,
and the 50 further from the bed had captured only 46.
This illustrates the possibility that anemones closer to
mussel beds may be getting more prey and possibly
making less available to those further away as well.
Comparisons of habitat suitability for
Anthopleura xanthogrammica
Using the information on individual growth, growth
increments of each size class (Fig. 13), and gonad pro-
duction as a function of size (Sebens 198 l b ) , it is pos-
sible to estimate expected reproductive success in a
422 KENNETH P. SEBENS Ecological Monographs
Vol. 53, No. 4
TABLE4. Immigration and disappearance in mapped populations of Anthoplezira xanthogrammicn, Tatoosh Island (1974-
1976). For each sampling visit, numbers of new individuals appearing and numbers of disappearances are given (in paren-
theses) beneath the mean diameter (2 2 SD) of anemones appearing in or disappearing from the mapped area. Note that
there has been an increase in the total number of individuals in each mapped area over the 2 yr; total immigrations are
two to three times as great as disappearances. The study occurred over a period of steady immigration of small individuals
from nearby mussel beds with no catastrophic disturbance to remove established adults.

All areas
(changes at each sampling visit)
1975 1976
-
1974-1976 changes
12 Feb
All areas Area 1 Area 2 Area 3 25 Jan 28 Mar 9 Jun 22 Jul 4 Sep or 1 Apr 20 May 23 Jan
Immigrants Mean diameter (cm)
Total 3.3 t 1.4 3.4 t 1.6 3.1 2 1.3 3.4 t 1.2 ... 3 . 6 t 1.8 3.0t0.8 2.5 3 . 3 t 1.4 3.622.2 3.2 t 0 . 9 2 . 8 t 1.3
(41) (22) (12) (8) (12) (4) (2) (7) (4) (5) (8)
Juveniles 3.1 t 1.4 3 . 0 t 1.0 3.1 2 1.3 3.4 t 1.2 ... 3.0 t 0 . 8 3.0 t 0 . 8 2.5 3 . 3 t 1.4 3.6 t 2.2 3.2 t 0 . 9 2.8 A 1.3
(40) (20) (12) (8) (10) (4) (2) (7) (4) (5) (8)
Adults 7.0 7.0 ... ... ... 7.0 ... ... ... ... ... ...
(2) (2) (2)
Disappearances
Total 3.822.2 4.322.9 4.3t1.8 3.121.1 2.0 1.2 3.3 2 2 . 4 6 . 0 t 1.4 5.0 4 . 5 ~ 2 . 4 3.520.7 ...
(22) (1 I) (2) (9) 1) (1) (9) (2) (1) (6) (2)
Juveniles 3.1t1.5 2.9tl.7 4.311.8 3.121.1 2.0 1.2 2.6 t 1.3 5.0 5.0 3.7 t 1.6 3.5 t 0.7 ...
(19) (8) (2) (9) (1) (1) (8) (1) (1) (5) (2)
Adults 8.221.0 8.2t1.0 ... ... ... ... 9.0 7.0 ... 8.5 ... ...
(3) (3) (1) (1) (1)
Population changes (number.m-'.yr-')
Immigrants 1.5 2.4 3.5 0.5 ... 5.2 1.2 1.8 3.0 0.6 2.2 1.4
Disappearances 0.8 1.2 0.6 0.6 0.2 0.4 2.6 1.7 0.4 1.0 0.6 ...

comparative manner for each habitat type, given some
important assumptions and limitations.
I f we assume that A. xnnthograrnmicu survival
probabilities (1,) are similar in each habitat type (as
indicated in Table 3), we can examine the prospective
growth and gonad production o f an individual surviv-
ing to full adult size for each habitat. Years ( n ) to
sexual maturity (at 6.5 cm diameter, Sebens 1981b)are
given by:
where D, is diameter at year x, g(D,) is the yearly
habitat-dependent growth increment o f an individual
of size D (from data in Fig. 13). I f gonad production
begins at year n , cumulative gonad production W(D,)
to some year m, weighted by survivorship to yearx(l,),
is given by:
where G(D,) is gonad production at size D in year x.
G(D)values are derived from an analysis o f gonad vol-
ume at the annual peak o f maturity (Sebens 1981b ) ,pre-
sented as mean gonad volume for each size class in Fig.
15. Gonads are almost entirely gametes with little sup-
port tissue, and reduce to a volume o f zero after spawn-
ing. Thus gonad volume at peak is a good estimate o f the
production of gametes. The cumulative gonad produc-
tion for a hypothetical individual entering each of the
three mapped areas. and the experimental removal
areas, is plotted in F i g . 16. The intermediate growth
trajectories of Fig. 14C were used for this calculation.
Lifespans of 150 yr are used for all habitat types and
mortality is assumed equal in each (0.122 deaths.in-
dividual-l.yr-l [ ~ cm 6 diameter], 0.006 depths.indi-
~idual-l.yr-~ [>6 em diameter]). These age-specific
mortality rates were used to calculate 1, values in the
analysis.
How realistic is it to consider lifespans of over a
century? Table 4 shows that o f 160 anemones (26.5
em diameter) monitored in the three mapped areas,
only 3 disappeared over the 2 yr of the study. This
equals 6 individuals per 1000 per year disappearing
from the study areas. I f this represents mortality (very
likely for large individuals), then the minimum esti-
mate of the half life (Ti) o f a group of anemones on
Tatoosh Island ( ~ 6 . 5em) would be 110 yr ( N , =
Noe-0.006t, T + = In 0.510.006) and mean lifespan (5 +
110.006, given 5 yr to reach 6.5 em diameter) would
be 172 years.
Cumulative gonad production is much less in the
most crowded control area (map 2 ) , and greatest in
the removals. The least crowded control area (map 3 )
had the highest value among the controls, and Map
area 1 was intermediate. Both growth rate and final
adult size determine this index of expected reproduc-
tive success. Offspring are initially planktonic planula
larvae and are assumed to settle over all habitat types
in proportion to the frequency of habitat types by area.
December 1983 SEA ANEMONE HABITAT SUITABILITY
If we assume that habitats such as the control mapped
areas 1 , 2, and 3 occur in equal frequency by area,
and that density will be similar to that during the 2 yr
of this study (no change in population size, R = 1)
then, by Euler's equation:
wheref, is the frequency of individuals in habitat type
i (based on area occupied by that habitat type and
density of individuals in that habitat type), l,, is sur-
vivorship to age x in habitat type i, m,, is gonad pro-
duction (volume) at age x in habitat type i, R is the
geometric rate of increase, and a is a constant relating
gonad volume to actual number of offspring surviving
to recruit into the population as juveniles. Total gonad
produced per individual per lifetime over all habitat
types is thus 1456 cm3 in this case. This is also the
amount of gonad (including support tissue associated
with gametes) equal to the production of one surviving
offspring assuming population stability (a = 0.00069).
For a mature oocyte diameter of 140 pm (Sebens
1981b) there are 3.64 x 1 0 j oocytes per cm3 gonad,
assuming immature gametes and support tissue occu-
py negligible space just before spawning. The replace-
ment value of 1456 cm3thus represents approximately
5.3 x lo9 eggs produced per female to result in one
new female offspring (and one male).
Cumulative gonad production and expected female
offspring for each habitat type are given in Table 5.
These values were also calculated for removal areas,
assuming that such areas include a small fraction of
the population such that 1456 cm3 still represents one
surviving offspring. That is to say, all offspring pro-
duced in removal areas will land in one of the control
habitat types. The most crowded control area (Map 2)
contributes little to the population's reproductive out-
put. The least crowded of the map areas (Map 3) con-
tributes half again as much as the area that is twice its
density (Map 1). Individuals fortunate enough to find
newly cleared areas, as in the experimental removals,
contribute more than twice the offspring of even the
best control area.
Comparisons of habitat suitability for
Anthopleura elegantissima
Habitat suitability is best compared slightly differ-
ently because of asexual reproduction of A . elegantis-
sima. For clones of equal size that are not growing, a
comparison of mean gonad production for individuals
in the clone would suffice (Fig. 17C), but for growing
clones the comparison is more complicated. Division
frequency is independent of mean size within clones
(Fig. 17A) and clones thus increase in number of anem-
ones at a rate independent of mean individual size.
Assume that an individual has succeeded in estab-
lishing itself in a given habitat type and proceeds to
form a clone by asexual fission. Let M, be the clone's
423
151 REMOVALS 1 ,
/ ,GI MAP AREA 3 Y 1
ORIGINAL DIAMETER (cm)
FIG. 13. Annual growth increments of individual Antko-
plerlra xanthogramtnica over the 2-yr study period (1974-
1976), mean ~ S DThe . diagonal line is the zero-growth iso-
cline. Numbers below points are sample sizes. Results of
ANOVA and SNK tests are given at the bottom of each
graph. Numbers refer to areas with significantly different growth
rates ( P < .05 at least; map areas l , 2 , 3 ;removals R; all control
areas T), (0= P < .05, A = P < .01, = P < ,001).
Anemone densities within the areas were: map area 1 (20
individuals/m2), map area 2 (51 individuals/m2), map area 3
(7 individuals/m2) and experimental removal areas (3 individ-
ualsim3. Areas I and 2 are on the exposed side of Strawberry
Island (Tatoosh Island) while area 3 is in the protected draw.
Removal areas include exposed and somewhat protected sites.
Note that the greatest growth rates occurred in the experi-
mental removal areas (significant for 6 of 11 size classes)
where growth of immigrating individuals was not inhibited
by the presence of established adults. Among the three con-
trol areas, area 3 (lowest density) showed the fastest growth,
followed by area 1, and area 2 (greatest density) showed the
least. Control areas differed significantly in only one to three
of 15 size classes.
average individual mass in habitat type i,fi the number
of divisions per individual in the clone per year, ei the
disappearance rate as disappearances per individual in
the clone per year. The rate of biomass change of the
clone over time, as it produces more individuals, is
given by:
where Md is clone mass and N o is the initial number
of individuals in the clone (here, N o = 1).
The rate of change in annual gonad production (AGI
at),is given by:
for a clone starting at N oindividuals where G is annual
gonad production of an individual at size A&.
 KENNETH P. SEBENS Ecological Monographs
Vol. 53, No. 4

'-1 A A L L INDIVIDUALS 1 B MOST RAPID GROWTH C. MOST RAPID INITIAL GROWTH

MAP 3
t

MAP 2

0 2 4 6 8 10 12 14 16 18 20 22 24 26

YEARS
FIG. 14. Growth trajectories for Anthopieura xanthogranzmica in control and removal areas. A. Growth calculated from
mean annual size increments (from Fig. 13). This represents the average for juveniles and adults and thus includes unsuc-
cessful juveniles. B. Growth calculated from mean annual size increments plus one standard deviation. This represents the
fastest growing juveniles and adults and may b e more rapid than any one individual actually grew. C. Growth calculated
from mean annual size increments plus one standard deviation until 6.5 cm diameter (maturity), then by the mean. This
represents growth of the fastest growing juveniles, then the average for adults.

Gonad production as a function of individual size is

given for A. elega~ztissitna in Fig. 15, from data in
Sebens (19810). G was calculated from peak annual
diameters for 2 yr given M, = 0.019 diameter3,27(Se-
bens 1979, 1981a ) . AM,/At is plotted for all experi-
mental clones as a function of mean individual mass
(/Gi) (Fig. 17B). For a clone without spatial limitations,
as in the experimental clones, growth can be contin-
uous so that the rate of increase in gonad production
(AGIAt) (assuming that the clone grows by producing
more individuals of the mean size) is the best com-
parative measure, and its integral is equivalent to ex-
pected reproductive success, and thus habitat suita-
bility (Fig. 17D).
Small individuals (<1.3 cm diameter, 0.05 g M,) were
not producing gonad (Sebens 1981b); so few individ-
uals < 1.6 cm (0.09 g 124,) were doing so that clones of
small individuals (note the nine smallest in Fig. 17D),
such as those in the San Juans high intertidal, are con-
tributing almost nothing to the gamete production of
the population, while those with large individuals pro-
duce much greater volume (Fig. 17D). Because asex-
ual division occurs in clones of large as well as small
individuals, the increase per year in gonad production
is much higher for clones of large individuals. There
is no estimate of the longevity of clones or of the rate
of establishment of new ones. Therefore, estimates of
expected surviving offspring production are not pos-
sible. However, expected offspring production of a
clone should be directly proportional to the summation
of AGIAt over the lifetime of a clone.
Population characteristics of
Anthopleura elegantissima
Anthople~lrl~ elegantissitntl is limited to protected
outer coast sites and moderately exposed inner coast
sites. Desiccation effects are more extreme in calmer
areas of the San Juan Archipelago (where transplanted
anemones failed to survive the midday summer des-
iccation [Dayton 19711). Desiccation very likely sets
the upper limit of intertidal distribution (Dayton 197I),
limits the local geographical distribution to moderately
exposed shores, and restricts the habitat distribution to
shaded crevices in the San Juan areas. The presence of
dense barnacle populations probably influences the
local geographic distribution as well, since cyprids tend
to settle where adult barnacles already exist (Crisp 1961,
1974), providing a major input to the anemones' diet.
Experimental results suggest that the sharp lower
limit of distribution in both the San Juan Islands and
the outer coast could be set by the upper foraging limit
of the subtidal seastar Demastrrias irnbricritcr, which
feeds readily on Anthoplrura spp. (Sebens 1977). The
presence of a dense algal canopy, and its shading of
the substratum, may also limit A. elegctntissinza to the
upper barnacle and mussel zone. Shading would re-
duce the contribution of zooxanthellae to anemone en-
ergetics (Sebens 1980) and overhangs may interfere
with prey capture. The positive phototactic response
of A. elegantissirnci (Pearse 1974, Fredericks 1976)
could cause individuals not to wander below the algal
December 1983 SEA ANEMONE HABITAT SUITABILITY 425

TABLE 5. Life history characteristics of Anthopleura xanthogrammica, calculated from growth trajectories (Fig. 14) and
gonad production (Fig. 15). Areas 1, 2, and 3 are mapped population areas, RE refers to all removal areas. AVE are
average growth trajectories in Fig. 14A; RA, most rapid trajectories in Fig. 14B; and I, intermediate trajectories in Fig.
14C. Cumulative gonad production is weighted by survivorship (see text).

Years to Plateau Cumulative Expected

maturity Years to plateau (cm) gonad offspring
Density production production
Area (N/mY) AVE RA,I AVE RA I RA I , AVE (cm3) ( 9 9)
RE 3 6.5 4.5 >30 >8 >20 >I5 > 15 4000 2.74
1 10 7 5 13 II 20 14.8 11.5 1095 0.75
-7 51 8.5 5.5 16 1I 11 12.6 8 50 0.03
3 7 8.5 5.5 20 11 16 15 12.4 1795 I .23

canopy or to move out from areas where such a can- clones. However, divisions and subsequent growth
opy develops. within clones of large individuals produced much
Within the upper and lower limits of intertidal dis- greater increases in clonal biomass than did divisions
tribution, spatial competition appears to be important in clones of small individuals. Therefore, clones of
for clonal aggregations. The elaborate agonistic sys- large individuals produce more individuals of large re-
tem possessed by A . elegrtntissiina functions to pro- productive size, such that the annual increase in gonad
tect clonal boundaries and to extend the clone in areas production from these clones, which are growing in
occupied by clones of lesser competitive ability (Fran- number of individuals, will be much greater than for
cis 19730, b). The extensive clones on protected outer clones of smaller individuals.
coasts, containing thousands of individuals per clone The clones with the greatest individual size, rate of
(e.g., Shi Shi Beach, Washington; La Jolla, California; mass change, and annual gonad production were those
Arroyo Hondo, California; Bodega Bay, California in protected (deeply concave) areas independent of
[Sebens 19771) clearly occupy all available space and intertidal height or location (Fig. 17). Clones of large
are limited at their borders by the presence of other individuals which are not dividing can be expected to
clonal aggregations. The smaller aggregations in the produce a similar volume of gonad each year, whereas
San Juan areas appear to be limited by the size of the clones whose individuals are undergoing fission have
acceptable habitat (tidepools, crevices) rather than by the potential to produce a much greater volume of
contact with different clones. In fact, aggregations oc- gonad as clone size increases. Thus the two most im-
cupying many separate pools may all belong to a single portant measures of reproductive success are size-de-
clone (Sebens 19826). The fact that numerical in- pendent gonad production (Sebens 198 16) and rate of
creases occurred in 8 of the 12 experimental clones in increase in gonad production by the asexual produc-
the San Juans (decreases in 2, no change in 2) indicates tion of new individuals.
that a spatial limitation would be reached in a rela- The mean number of divisions per individual per
tively few years. What happens when a clone reaches year for all areas was 0.17 *
0.09 SD. If we assume
the limits of available space? Individuals must either that none of the disappearances from clones represent
cease dividing asexually and grow to some individual mortality but simply emigration, such a division rate
size limit, or asexual products must leave the clone
and become wandering solitary individuals. Densities
TABLE6. Calculations of A. xanthogrammica life history
in Dayton's (1971) control areas (with aggregations) characteristics from Ford-Walford plots. Areas as in Table 5.
remained near a constant value over 3 yr (1966-1969) t,,, is the time to 95% of asymptotic size; t,,; is the time
while removal of barnacles caused average densities to maturity at 6.5 cm diameter; t , , the y intercept, is esti-
mated size at 1 yr; r2 is the coefficient of determination for
to increase, indicating a release from spatial limita- the regression; and slope ( e - 9 is the regression coefficient
tions imposed by the presence of barnacles. Reduced where k is the constant in the von Bertalanffy equation.
density within clones allowed both numerical increase
and. in some cases. individual size increase in this
study (Table 3). inter-
Plateau cept
Individual size changes and numerical change in ex- (Lm) f , , ~ ~ t6.5 Slope (t,)
perimental clones showed that annual cycles of bio- Area (cm) (yr) (yr) N (e-9 (cm) r2
mass increase and decrease were occurring at the level 23.6 48 5,2 282 0,94 ,4 0,86
of the individual, and that size at division was a hab- 1 10.4 20 6.5 579 0.86 1.4 0.88
itat-dependent characteristic rather than an absolute 2 9.1 20 8.2 419 0.86 1.3 0.89
(Sebens 19826 and Fig. 10). Actual frequencies of di- 14.4 33 6.6 367 0.91 0.91
All 11.0 23 7.0 1365 0.88 1.3 0.89
vision do not correlate with mean individual size in
 KENNETH P. SEBENS Ecological Monographs
Vol. 53. No. 4
mT
-
0
70-
A. -
- xanthogramm~ca
4
A. ejegantissima .
w
I
60-
3
. 2

3 50-
0
13

>
40-
0
u

20

. I2

I 2 3 4 5
B A S A L D I A M E T E R (cm) 8 A S A L DIAMETER (cm)
FIG. 15. Mean gonad volume in each size class of Anthopleura xanthogrammica and A. elegantissima. Data are from
Sebens (1981b) and standard deviations are approximately equal to the means. Numbers below points are the number of
individuals examined.

would fill a 100 m2 area with close-packed 2 cm di- (higher temperature) and occurs more frequently in
ameter anemones in 81 yr, given exponential growth. larger members of a clone (Sebens 1980, 198%). Min-
However, if we assume that all disappearances rep- asian (1976), and Minasian and Mariscal (1979) have
resent mortality, the rate of increase is 0.10 _t 0.06 shown high rates of reproduction by fission for Hali-
individuals per individual per year, and the same feat planella luciae (to 40 divisions126 d) and Johnson and
would be accomplished in 134 yr. In other words, A . Shick (1977) have found that this species produces
elegantissima has the potential to colonize and cover larger individuals under conditions of greater physio-
large areas of intertidal in ecologically meaningful time
periods. Actual clone growth rates in the presence of
competing clones or individuals will be somewhat less,
depending on the cost of such competition in terms of C xonthqramm~co REMOVALS A 3
individuals, or growth rate, lost due to agonistic en-
counters (Francis 1976).
Larval settlement of A. elegantissima occurs into
crevices in the rock or into mussel beds. While settle-
ment may be an annual event, there can also be less
frequent waves of dense recruitment, such as occurred
in Tatoosh Island mussel beds in 1972-1973 (Sebens
1982~).Individuals growing to adult size in mussel beds
or in intertidal crevices become wandering individuals
migrating downward through the intertidal, and some-
times successfully establishing new clonal aggrega-
tions. In this process, they face interference compe-
tition from established clones which can consist of
many thousands of individuals occupying all usable
space (Francis 19730, Sebens 198%). Solitary wan-
dering individuals are at a decided agonistic disadvan-
tage compared to established clones, and probably
succeed in establishing new clones only in areas where
frequent disturbance keeps existing clones from mo-
nopolizing space.
Asexual proliferation of A. elegantissima differs from
lb
I

, 0 io i o i oo-o i
YEARS
I

that reported for other anemone species in many re-
spects. It is an annual, highly seasonal, phenomenon
(Sebens 198%), and can be initiated by a period of
starvation in the laboratory (Sebens 1980). Division
occurs more rapidly with greater physiological stress
FIG. 16. Projected cumulative gonad production (cm3) of
Anthopleura xanthograrnmica (logarithmic scale), based on
growth trajectories in Fig. 14C and weighted by age-specific
survivorship. The expected number of surviving offspring
corresponding to a given gonad volume is also given along
the y axis. See text for calculations.
December 1983 SEA ANEMONE HABITAT SUITABILITY 427

0.6
1 A, A. elegantissima
0 CONCAVE SURFACE
F L A T SURFACE
11
S SAN JUAN SITE 1
T TATOOSH S I T E 1
H
L
HIGH NT.
LOW N T 1

0 THl

Md (grams) Md (grams)

Md (grams) Md (grams)

FIG. 17. Habitat suitability of Anthopleura elegantissima. A. Division frequency Cf, in divisions per individual per year)
of experimental clones of Anthopleura elegantissima plotted against mean individual mass per clone (M,, grams ash-free dry
mass) calculated from basal diameter (see text). Numbers next to data points are clone identification numbers: Tatoosh
Island (T), San Juan Islands (S) (Cattle Point, San Juan Island, Iceberg Point, Lopez Island), high (H), and low (L) intertidal.
There is no significant correlation (r = ,098). B. Rate of change in total clone ash-free dry mass (grams per individual per
year) plotted against M,. The regression line without TL6 is AM,/At = 0.17 M, - 0.008 (r2 = .80, P < ,001) and with TL6
is: hM,lAt = 0.12 M, - 0.003 (r2 = .SO, P < .001), C. Annual individual gonad production (cubic centimetres of gonad
volume) plotted against M,,,The regression line is: G = 1.27 M,, - 0.035 (r2 = .9, P < .MI). D. Annual rate of increase in
clonal gonad production (AGIAt, cubic centimetres of gonad volume per individual per yr) plotted against M. The regression
line without TL6 (on graph) is: AGIAt = 0.19 M , + 0.008 (r2 = .79, P < .001), and with TL6 AGIAt = 0.15 M, + 0.0007
(r2 = .53, P < ,001).

logical stress. Smith and Lenhoff (1976) showed that
pedal laceration of an acontiarian anemone also in-
creased with greater physiological stress. Clones of A.
elegcrtitissitncr can cover many square metres of inter-
tidal area (Sebens 1980), as can clones of H. luciae
(Shick 1976), and clones produced by pedal laceration
of Metridium s ~ n i l e(Hoffmann 1976).
Mortality of A. elegcrnti.r.ritntr is difficult to estimate
in the field. In the experimental clones, disappearance
from clone nuclei averaged 0.07 & 0.06 individuals S D
per original individual per year, very likely a combi-
nation of mortality and emigration (Table 2). Given the
tendency of the anemones to remain aggregated, and
the large clear space maintained around experimental
clones, the likelihood that such disappearances rep-
resent mortality is high. Desiccation effects and drift
428 K E N N E T H P. S E B E N S Ecological Monograph,
Vol. 53. No. L
log scraping (Dayton 1971) probably account for a cer- thus also suffer the effective loss of potentially repro-
tain amount of this mortality. I n fixed control areas (1 ductive individuals a t regions of contact.
m2 each), differences between peak winter and lowest
summer densities averaged 23.8 2 lo.% SD (N = 6) Populatioti characteristics of
of peak density for the years 1967-1969, whereas in Anthopleura xanthogrammica
two Setnibttlttnu~cerrio~usremoval areas ( 1 mYeach) Atitlzopleurtr xtrtithogrcrtntniccr is most common on
the corresponding years showed differences of 70.8 t exposed rocky shores where mussel beds (.2?\.tilu.tCNI-
19.6% ((N = 6) of the peak density (Dayton 1971). The ifornitinus) and the starfish P i ~ t c ~ t eorc h r t r c e ~ sare
large difference in survivorship between the two groups common. The anemones are limited to very protected
can be explained if the control areas contained pri- microhabitats where their low profile, and sometimes
marily established clonal aggregations, and the remov- the presence of adjacent anemones, makes them less
al areas became occupied by wandering individuals. subject to shear forces from extreme wave action
I n addition to potential physical sources of mortal- (Koehl 1977cr). This orientation also provides reduced
ity, there are several known predators. The seastar water velocities where the anemones are located, while
Dertnci.tteria~itnbricccta, although rare intertidally, can maximum water velocities occur a t the level of the
remove anemones in the low intertidal and those trans- mussel beds. Anemones can thus capture mussels from
planted t o the subtidal (Sebens 1977). T h e nudibranch the water at its reduced velocities (Koehl 1 9 7 7 ~ )Dis- .
Aeoliilier pnpillo.~eractually prefers Anthopleurcc species tance from the mussel bed prey source appears to be
over other local anemone species (Waters 1973). Aeo- one of the most important factors determining habitat
liciiin are highly cryptic, with the same color and gen- suitability and affecting recruitment rates (Sebens
era1 shape as A. elegcrtitissitncc. Nevertheless, only four 198 Ic). However, in areas directly adjacent to a well-
individual Aeoliditc were encountered at all of the study developed mussel bed, intense crowding may depress
sites during the 3-yr period. In one such case, an A. both juvenile growth rates and maximum adult size
elegccnti.s.titntr ( 1.6 c m diameter) had just released its (Fig. 14).
attachment in response to an Aeolidicr pcrpilloscc. Many A. .rttnthogrertnmiccr mean diameters are reduced
anemones are able to inflate the pedal disc and release during winter months when Pistester is less active. Al-
themselves from the substratum after being contacted though winter storms release mussels from the bed in
by predatory nudibranchs (A. elegerntissitner , Waters large patches, such prey are available to anemones
1973. Edmunds et al. 1974; A. nigrescens, Rosin 1970, only in short pulses and in clumps often too large to
reviewed in Ottaway 1977). b e handled, rather than continuously, as occurs when
In laboratory aquaria, a single individual A. pccpil- Piscrster is active. Smaller anemones may be subject
lo.ttr (2.5 cm long) consumed three anemones (1-1.5 to the same prey fluctuations that affect A. elegcrntis-
c m diameter) per week over a 4-wk period, laying egg sitner, especially if the anemones are not large enough
masses intermittently. At such rates, a single A. ptrp- t o capture mussels. Small A. .rt1nthogrtztnt71ictron the
illo.scc could account for -80 anemoneslyr, assuming high platform and vertical wall monitoring areas at Shi
that it fed equally during 6 mo and negligibly the other Shi Beach showed such annual size changes (Fig. 11).
6 mo. The predator density would not have to be very The seastar Piscc.rter ochrwceu.r used the removal
great to have a significant effect on the population. A t areas as low tide refuges (up to 25 Pisccster/O.S m2

-
a division rate of 17 anemones per 100 per year, A. removal area) during its active months, and was less
prrpillo.cer densities need only be l per 470 anemones apparent the rest of the year when it is essentially
to consume all of the newly produced anemones. The inactive in the intertidal (Mauzey 1966, Paine 1979).
close scrutiny and the numerous visits t o experimental Numbers of Piscr~terin the removal areas were re-
areas and to monitoring areas indicate that the real corded from July 1974 to May 1976 (Sebens 1977).
densities of the nudibranch are far below that figure. Periods of anemone size increase and of Pistc.rter pres-
Predatory gastropods (Epitoniidae) occur subtidally ence (indicating activity) coincided, as did the corre-
but have not been encountered with A. elegcrntissitner sponding decreases. These results agree with Dayton's
at the study sites. T h e nudibranch Hermissetidu crer.r- (1973) experimental results indicating that Pi.rcister
.ticornis appears to be a cnidarian generalist taking a foraging can supply A. xttntlzogrcrtntnicu with a large
wide variety of small polyps and zooids (Birkeland percentage of its prey.
1974), and could be a source of mortality for newly Small A. xcrtithogrcrtntnicer changed location more
settled anemones in tidepools. frequently than did large individuals. This indicates a
Finally, agonistic encounters between individuals and period of active juvenile (secondary) habitat selection.
at clonal boundaries may result in mortality or relegate Given the depressed juvenile growth rates in very dense
unsuccessful individuals to areas of the habitat where areas, it is likely that small anemones are actively
mortality from other sources (e.g., desiccation) is more searching for suitable locations (where adult prey
likely. Francis (1976) indicated that more of the ago- [mussel] capture is likely), but are interfered with by
nistic individuals at the edge of clones in contact with the presence of adults. Another part of this study in-
other clones lacked gonad during the peak summer dicated that A. .ucrtzthogrtont~~icec settlement occurs into
period. I n addition to loss by mortality, clones may beds of M. ccilifornicrtiur (Sebens 1 9 8 1 ~ .19820, an
 December 1983 S E A ANEMONE HALBITATSUITABILITY
appropriate larval choice, since this will be their adult
prey. A period of growth in the mussel bed is followed
by downward migration into pools, walls, and chan-
nels where juveniles must compete for space with large
established adults. Successful location of suitable sites
results in increased growth, attainment of reproduc-
tive maturity, and cessation of extensive movement.
Anemones >6 c m diameter rarely altered location
and appear to be extremely long-lived, if the 2-yr study
period is a t all representative. Lifespans of many de-
cades are not unrealistic. In fact, anemones collected
in the late nineteenth century were kept alive in aquar-
ia for up to 80 yr (Ashworth and Annandale 1904,
Strehler 1961, Ricketts et al. 1968) and died only when
their care was neglected. P. K. Dayton (personal cotn-
mutiicution) observed mortality of several large A.
.uanthogratntnicu a t Shi Shi Beach when large rocks
collected in a tidepool and wave action caused them
to scour the walls of the pool. In three instances I have
observed tidepools and channels become roofed over
with a bed of Mytilu.~culiforniutius when large clumps
of mussels fell from nearby beds and reattached in
these depressions. Such entrapment of large anemones
may result in mortality either by occluding the feeding
apparatus or by abrasion (as noted by Paine 1974).
Predation on large A. .uanthogrcrtntnica was never
observed during this study, although Paine (in Mauzey
et al. 1968) observed the seastar Dertnusterias itnbri-
cata feeding on small specimens in the low intertidal.
Experimental transplants and caging studies indicated
that Dertnusterias can consume only small A. xuntho-
gratntnicu, allowing an escape in size for large indi-
viduals ( 2 6 cm) (Sebens 1977). The subtidal size range
a t Tatoosh Island ( 6 3 3 cm diameter) may be a result
of predation by Dertnasteria.~on small individuals that
move to the lower intertidal and subtidal or are carried
there attached to released mussels. A pycnogonid
(Pycnogotiutn stearti.~i)and an epitoniid gastropod
(Opulia crenitnargitiata) are known to feed on A. xan-
thogratnmicu in California (Ricketts et al. 1968), prob-
ably as ectoparasites. The local Opalia chaseii is also
frequently found attached to anemones by the anem-
one's adhesive verrucae, with the proboscis implanted
in the body wall of the anemone. The importance of
such feeding by ectoparasites is unknown. Aeolidia
p a p i l l o . ~will
~ feed readily on A. xatithogratnmica in
the laboratory (Waters 1973), but the nudibranch is
very uncommon on the exposed coasts (two have been
found on Tatoosh Island during all visits since 1967).
Cotnparisons n~ithother specie.^
The population dynamics of sea anemones had been
almost completely neglected before this study and that
of Ottaway (1979b) on Actiniu tenebrosu in New Zea-
land. A. tenebrosa broods its young (Ottaway 1974),
although it is unclear whether young are produced
asexually (Black and Johnson 1979) or sexually (gon-
ochorism. Ottaway and Kirby 1975. Ottaway 1979b).
Persistence of adult A. tenebrosa (94% survival of
429
adults over 3 yr) was similar to that of adult A. xcrn-
thogrcrmmicu in this study. Juveniles were much more
abundant and consequently showed much greater re-
cruitment and disappeararce rates than did those of
either species in this study. Individual seasonal growth
and shrinkage produced a pattern of annual size fluc-
tuation in A . t e n e b r o ~ usimilar to that of both Antho-
pleura species. Ottaway (1980) calculated longevities
(mean 50 yr, maximum 210 yr) for adults, and a growth
rate from recruitment to 4 cm diameter in 8 to 66 yr.
It is difficult to compare the population dynamics of
A. elegantis~itnato that of other species since indi-
vidual "mortality" often occurs via the production of
daughter individuals after fission. Division rates in nat-
ural populations (Sebens 1982b) and rates of disap-
pearance from experimental clones indicate that indi-
viduals have an expected mean lifetime of 2-3 yr,
although some individuals probably live much longer.
Field tneasuretnetit of itidetertninate g r o ~ ~ s t h
Growth rates, and thus reproductive success, de-
creased with increased density of A . .uatitlzogratntnica
in the three control areas. Both values were even
greater in experimental removal areas where density
was lowest. Comparison of growth rate by size class
has been used here for an animal with highly indeter-
minate growth (able to grow and shrink over a wide
range). The method takes into account the fact that a
group of equal-sized individuals will not necessarily
increase together. Some individuals will grow more
than others, and some may even shrink. Calculation
of growth trajectories from such data extrapolates cur-
rent conditions over many decades and assumes that
the mean growth in each size class is representative.
Individuals growing to full adult size and occupying
space for a long period are the successful ones, whose
early growth in particular may have been more rapid
than average. Growth increments for juveniles include
those of individuals subsequently dying (Ricker 1975)
or emigrating. For this reason, the rapid juvenile growth
rates have been used (mean increment plus one stan-
dard deviation).
If no mortality or emigration occurred, mean growth
increments might best describe average growth curves.
Insignificant adult mortality allows the use of mean
increments for adults in this study. Mean increments
could also be used forjuveniles if survival to adulthood
were random and not conditional on rapid early growth.
Some knowledge of differential survivorship at all
stages is necessary to determine the best way to use
growth increments to plot trajectories. Plotting both
mean and most rapid growth (Fig. 14) outlines the re-
gion of most probable average growth trajectories. I t
is less likely that increments lower than the mean of
each size class would describe the average growth in
a given habitat.
Despite the necessary assumptions, this method al-
lows accurate comparison of both growth rate (and its
variability) and expected reproductive success. Growth
430 K E N N E T H P. S E B E N S Ecological Monographs
Vol. 53, No. 4
rate alone does not come close to describing "fitness" Longitudinal fission of Anthopleura elegantissimu
in a given habitat. Only the contribution to the next can be a means of optimizing energy acquisition and
generation, weighted by survivorship and timing of thus fitness (reproductive output). There may be an
offspring production, will suffice. Previous studies that energetically determined optimum size for anemones
have compared individual growth in altered conditions in each habitat type set by the local balance of prey
have used a single size class (Peterson and Andre 1980) availability and metabolic cost (Sebens 1979, 1982~).
or a few individuals over several years (Edwards and Division will occur when anemones exceed this habi-
Huebner 1977). Sea urchin growth on various diets tat-dependent size. Since clonal gonad production de-
(Vadas 1977) has been compared, although original and pends on individual size within the clone, there must
final sizes of individuals may have been at different be some advantage to division at a smaller size in poorer
points on a growth trajectory. (more stressful) habitats. That advantage probably de-
The present method allows a comparison between pends on the ability to continue a high rate of clonal
habitats or between experiments using a large data set growth rather than to reach some larger individual size
and considering all size classes. Both growth rates and without further division.
final adult size can thus be compared statistically (e.g., Gonad volume of A . xanthogrammica increases with
by ANOVA). Annual growth increments were used, individual size in all habitats considered in this study
since there is a known seasonal size fluctuation. Growth (Sebens 1981b). The great differences in cumulative
of annual or short-lived organisms could be compared gonad production (Fig. 16) occur as a result of the
similarly using smaller time intervals, but the method delayed onset of reproductive maturity as well as the
is particularly well suited to long-lived organisms with ultimate maximum gonad production. Since size, rath-
indeterminate growth. er than age, determines reproductive success, and
space near abundant prey is limiting, juvenile A. xan-
Habitat suitability thogrammica may be adapted to search for and rec-
It was possible to estimate the differences in ex- ognize suitable sites and even to space themselves away
pected reproductive success in a variety of habitat types from obstructing adults whenever possible.
using the information on individual and clone growth A. elegantissima occurs over a wide range of phys-
and the data on gonad production (Sebens 1981b). ical conditions, while their prey is generally available
Longitudinal fission of A . elegantissitna does not oc- in all habitat types, and such physical conditions de-
cur more frequently in clones with either larger or termine size at division, which in turn determines mean
smaller mean individual size. However, the gonad pro- individual size in clones. Since reproductive success
duction of large individuals is so much greater than of a clone is directly related to mean individual size,
that of smaller ones that there is a distinct linear re- habitat suitability for A . elegantissima is determined
lationship between annual gonad output and mean largely by physical conditions (including immersion
clonal individual mass. This would be sufficient com- time, which limits feeding). A . xanthogrummicu, on
parison, except that clones that have not reached a the other hand, exists within a rather narrow physical
spatial limitation continue to produce individuals that range, actively moving into more protected microhab-
build additional gonad in subsequent years. The rate itats. Reproductive success is directly related to size,
of increase in gonad production (per original individ- which appears to depend primarily on prey availabil-
ual) is a better comparison of habitat suitability for ity, although it will also be influenced by conditions
growing clones. Since divisions occur in most of the which change energetic cost.
clones, this rate is also proportional to mean individual Habitat suitability can be highly variable. The re-
mass in an approximately linear manner (Fig. 17). sults of this study are similar to those for limpets
Habitats in which clones consist of large individuals (Sutherland 1970), intertidal snails (Paine 1969), mus-
were clearly the more suitable, by at least two orders sels (Paine 1976), and barnacles (Wethey 1979) indi-
of magnitude (dGidt comparison) over habitats with cating that there are certain habitats (usually less
clones of the smallest individual size. Within clones crowded, with more available food, or lower mortali-
of small individuals it is only some of the largest that ty) where individual reproductive success is many times
produce gonad at all. It is interesting to consider why that in other habitats. Future populations will be com-
division occurs at such a small size, producing infertile posed primarily of the offspring of such fortunate in-
individuals. Given the potential individual growth rates, dividuals.
it is very likely that all individuals in such clones could The distribution of anemones over their range of
reach fertile size during years with high prey abun- habitat types is far from the "ideal free distribution"
dance, and thus produce more offspring than if divi- hypothesized for organisms with refined habitat selec-
sion had not occurred. In addition, large size may be tion and assessment abilities, while it may correspond
physiologically disadvantageous or even impossible in more closely to the "ideal-despotic distribution" of
habitats with high stress or low prey availability (Se- Fretwell (1972) because of the advantage afforded es-
bens 1980). tablished individuals. Movement during a phase of ac-
December 1983 SEA ANEMONE HA BITAT SUITABILITY 43 1

tive habitat selection (juvenile) and regulation o f tim- in a rocky intertidal community. Ecological Monographs
ing and division size allow anemones to locate more 41:351-389.
suitable sites and to respond with growth, gonad pro- . 1973. Two cases of resource partitioning in an in-
tertidal community: making the right prediction for the
duction, and division rates appropriate to the physical wrong reason. American Naturalist 107:662-670.
and biological constraints of each habitat. . 1975. Experimental evaluation of ecological dom-
inance in a rocky intertidal algal community. Ecological
Monographs 45: 137-159.
Dunn, D. R. 1975. Reproduction of the externally brooding
I wish to thank the following people for their important sea anemone Epiuctis proliferu Verrill, 1869. Biological
contributions to the formulation and realization of this re- Bulletin 148: 199-218.
search: R. T. Paine, G. H . Orians, A. J. Kohn, T. H . Su- Ebert, T. A. 1980. Estimating parameters in a flexible growth
chanek, J. M. Sebens, M. A. R. Koehl. and L. Francis for equation, the Richards function. Canadian Journal of Fish-
continued encouragement, assistance, and discussion: B. eries and Aquatic Science 37:687-692.
Churchill, D. Duggins, M. A. R. Koehl, S. Libonatti-Barnes, Edmunds, M., G. W. Potts, R. C. Swinfen, and V. L. Waters.
A. Linder, T. Linder, J. Nelson, R. T. Paine, C. Quaife. C. 1974. The feeding preferences of Aeolidiu pupillosa (L.)
Schrank, C. P. Sebens, J. M. Sebens, B. Stedman, C. J. (Mol1usca:Nudibranchia). Journal of the Marine Biological
Sturgis. T. H . Suchanek, S . Waltz, B. Whitney and D. Wood Association of the United Kingdom 54939-947.
for volunteered assistance in field, laboratory, and SCUBA Edwards, D. C., and J. D. Huebner. 1977. Feeding and
diving experiments; P. K. Dayton, R. Fernald, V. Gallucci, growth rates of Polinices duplicutus preying on Myu ur-
P. L. Illg, E. N. Kozloff, C. Nyblade, A. R. Palmer, R. R. encrria at Barnstable Harbor, Massachusetts. Ecology 58:
Strathmann and A. 0 . D. Willows for valuable discussions 1218-1236.
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L. Thorne, D. Denninger, and R. Aronson for reading and invertebrates in shallow water. Limnology and Oceanog-
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