THE ACCLIMATION PROCESS BETWEEN FISH ANT
ANEMONES IN MONTEREY BAY
STEPHEN EMILLER
Stanford University at Hopkins Marine Station
Biology 175H, 1989
ABSTRACT
Although there have been many studies on the relationships
between certain fish and sea anemones in the tropics, very little
is known about the fish-anemone interactions in the colder waters
of the northeastern Pacific Ocean. Äfter a seven week
investigation in Monterey Bay, I found that none of the fish
observed will acclimate to the tentacles of either of the intertidal
sea anemones Anthopleura xanthogrammica or Anthopleura
elegantissima, although the wooly Sculpin (Clinocottus analis)
will rest on the base or column of either.
also found that the Painted Greenling (Oxylebius pictus), as
well as other rock dwelling fish, will acclimate to the Strawberry
Anemone (Corynactis californica). The acclimation process takes
approximately 3-5 days, depending on the amount of contact
between the acclimating fish and the anemones, and the amount of
previous contacts with the anemones by other fish. There are two
mechanisms involved in the acclimation process. First of all,
through repeated contact, the fish picks up mucus from the
anemones and adds it to its own mucus layer. This causes each
anemone to recognize the acclimated fish as a part of itself and
prevents the fish from stimulating the anemone's chemoreceptors
for nematocyst discharge. Secondly, the anemones become
habituated to contact by the fish.
INTRODUCTION
There are two fish that were concentrated on during this
study. The first were the wooly Sculpins, Clinocottus analis.
These fish inhabit the intertidal and shallow subtidal zones. They
are found from Baja, California to Cape Medocino, California and
are extremely abundant in the tidepools and shallows at Hopkins
Marine Station in Monterey
The other fish studied were Painted Greenlings (also called
the convict fish), Oxylebius pictus. These fish live at depths
ranging from just below the intertidal to 50 meters along the
northeastern Pacific coast from Baja California to British
Columbia. They live under the canopy forming brown kelp
Macrocystis pyrifera Adult greenlings are residential, usually
occupying areas less than 10 square meters (DeMartini), and feed
from the rocks on a variety of invertebrates, especially
crustaceans and worms (Burgess and Axelrod).
Two intertidal species from the order Actiniaria were used
in this study: Anthopleura xanthogrammica and Anthopleura
elegantissima They each have strongly adherent bases and have
columns covered with verrucae, which are evaginations with
modified ectoderm and no nematocysts which are able to cling to
such things as rocks and shells (Carlgren). They have large, strong
tentacles and can therefore catch fairly large prey. Both practice
dioecious reproduction, although A. elegantissima also has the
ability to reproduce by longitudinal fission, forming aggregations
of a single clone (Sebens). They are part ially supported by the
symbiotic algae Zooxanthellea (Friese).
The other anemone studied was the Strawberry Anemone,
Corynactis californica. This anemone-like species is in the order
Corallimorpharia It is morphologically similar to corals, but it
lacks any skeleton. It has a smooth column and thin tentacles with
acrospheres at the distal ends. Acrospheres are globular ends of
tentacles containing numerous potent nematocysts (Carlgren).
This anemone is very abundant in Monterey Bay and occurs in
California from San Diego to Sonoma County. It is found in shaded
areas from the low intertidal to 30 meters. Corynactis californica
reproduces asexually by longitudinal fission and is found in clonal
groups (Hand).
In a recent publication, The Biology of Nematocysts
(Thorington and Hessinger). Thorington and Hessinger described
their study of nematocyst discharge and listed some of the factors
affecting this discharge. Like most researchers of nematocysts,
they found that concomitant mechanical and chemical stimulation
of the cnidocytes is required to initiate or trigger the discharge
of the cnidae. The cnidocytes are specialized secretory cells on
the tentacles that are also considered independent effectors and
directly perceive stimuli for discharge.
The mechanism for discharge involves chemosensitization of
the cnidocytes by the active chemicals, which then lower the
mechanical threshold of cnida discharge. Chemical stimulation
alone has occasionally been found to cause discharge, but only
under more stringent conditions (using very harsh chemicals) than
are found in the natural environments of the anemones. In some
anemone tentacles mechanical stimulation alone can cause some
discharge, but in every case adding appropriate chemical
stimulation to mechanical stimulation causes much greater
discharge. Some of the active chemicals have been isolated and
include n-acetylated sugars and various amino compounds.
N-acetylated sugars are common in the mucus layers of many
organisms that are prey for anemones. These sugars cause
immediate discharge on contact with the tentacles. Then, after
nematocysts have pierced the prey, body fluids containing
numerous amino compounds are released and stimulate further
discharge.
For years cnidocytes were considered independent effectors
with direct control over the cnidae discharge and no intervention
of the nervous system. Certain observations, however, show that
the nervous system seems to have either a modulatory or an
inhibitory influence on cnidae discharge. For example, a
swimming anemone, Stomphia coccinea, will not capture any prey
while its pedal disc is detached from the substratum. It was also
found, using coverslips coated with food extract and the anemone
Anemonia sargassensis, that tactile stimulation of the body
column increases the amount of discharge. Even the ambient light
intensity, though cnidocytes are not considered to be
photosensitive, affects the responsivness of cnidocytes and the
ability to capture prey. Other factors influencing the
physiological state of the anemones, such as ions in the ambient
medium and the amount of recent feeding, also seem to affect the
amount of cnidae discharge (Thorington and Hessinger).
The acclimation process between anemonefish and their
hosts has been studied for years, but most of these studies have
taken place in the tropics where the fish actually lives within the
tentacles of a single host. There are a few different theories on
how the acclimation takes place.
The first set of theories assumes that the anemone makes
certain changes and is responsible for the acclimation (Gohar).
They feel that the anemone changes or inhibits its pattern of
nematocyst discharge due to the behavioral, physical, or chemical
stimuli of the acclimating fish.
Contrary to these are the theories in which the fish is
responsible for its protection from the anemones by in some way
changing its mucus layer. Some feel that the mucus layer after
acclimation contains no active agents or other factors which are
known to stimulate the chemoreceptors, causing nematocyst
discharge upon contact (Lubbock). Öthers feel that certain
chemicals in the mucus layer inhibit nematocyst discharge
(Davennport & Norris). This inhibition can occur in one of two
ways. There is either direct inhibition of the
cnidocyte-nematocyst complex on the tentacles, or this inhibitior
is accomplished through the organism's nervous system.
The final theories assume that the mucus of the anemone is
transferred to the acclimating fish. One German researcher,
Schlichter concluded in 1976 that substances which inhibit
nematocyst discharge are produced by the anemones themselves
(Mariscal). Through repeated contact, the acclimating fish coats
its body with this inhibitor as to no longer induce nematocyst
discharge. Another possible explaination is that the anemone
identif ies the acclimated fish as a part of itself by recognizing its
own mucus on the fish.
MATERIALS AND METHODS
All of the Clinocottus analis used were collected from
tidepools during low tide at Hopkins Marine Station. All other fish
tested (Oxylebius pictus, Coryphopterus nicholsii, and Orthonopias
triacis) were captured with anet from depths of 5-10 meters at
Hopkins Marine Station, the breakwater, or Lovers Point, all of
which are located in Monterey Bay.
For Tests With Anthopleura;
In each experiment l used a tank with two A. xanthogrammica
and one A. elegantissima, with approximate diameters of 15, 10.
and 10 centimeters respectively. All three anemones were
carefully removed from rocks in the intertidal zone of Hopkins
Marine Station. They were left in isolation for one week before
any experimentation was started, in which time they settled on
rocks in the observation tank. During observational experiments
(watching fish in tanks with anemones), all contacts and reactions
were noted, described and timed. In the experiment with forced
contact between anemones and Wooly Sculpins (Clinocottus
analis), the sculpins were held by the jaw with forceps and placed
tail first on one side of the ring of tentacles. Care was taken not
to touch the fish with my hands
For Tests With Corynactis californica:
One rock of C californica was collected from a depth of 8
meters at the breakwater of Monterey Bay. All other rocks were
collected from the Del Monte shalebeds of Monterey Bay at depths
of 13-16 meters. For all observational experiments, tanks would
be watched for a couple hours and then left for a few hours
repeatedly over several days. All rocks with C californica were
mapped such that individual anemones could be accounted for (See
Maps 1-3).
During a given observation period, all contacts were recorded
by anemone number and type of contact. Three types were used: hit
hard, hit, and brushed. One of four types of reactions by the
anemone was then recorded: closed, closed 1/2, slight retraction,
or none. When an anemone closed, the amount of time before
opening was also marked for some experiments. For any
observation period, a "closure value" could be calculated. Each of
the reaction types was assigned a number: closed=3, closed
1/2=2, slight retraction=1, and none=O. For all contacts in a
period, these numbers were added up and divided by the total
number of contacts. This would result in a closure value between
O and 3.
One experiment using various parts of Painted Greenlings
was also done. A fish was captured in a clean glass bowl to
prevent contamination. The fish was then placed on its side in a
clean pan, keeping its tail up with forceps. The spinal cord just
behind the head was quickly severed and the tail and side facing up
were left uncontaminated. The pectoral fin, ventral fin, and tail
were then removed and tested on anemones. The anemones were
put in just enough water to cover them such that the body parts
could be touched to anemones without letting the areas cut touch
the water. This prevented internal body fluids from entering the
water and possibly triggering the chemoreceptors.
The final experiment done with C californica involved an
attempt to acclimate my finger to a clonal group of anemones from
the shalebeds. Irubbed the end of my index finger against 172 of
the anemones (which were closed) and then tested the reactions of
open anemones to my touch.
RESULTS: FIELD OBSERVATIONS
Both Anthopleura xanthogrammica and Anthopleura
elegantissimawere studied in the tide pools of Hopkins Marine
Station. The only contact ever seen involved a wooly Sculpin and
an A. Xanthogrammica The sculpin was sitting on the column of
the anemone, approximately 1/2 way between the oral and pedal
discs. Another observation involved an A. xanthogrammica, four
Wooly Sculpins, and a Monkey Faced Eel. The anemone was
attached to a vertical rock surface just above the bottom of the
pool such that its tentacles reached the bottom. This formed a
small shelter between the sand, rock, and tentacles (Diagram 1)
saw a fish go behind the tentacles, but it was too dark to see
where it went. I could see that nothing was touching the
anemone's tentacles. By putting a net on one side of the anemone
and sticking my hand under on the other side, I captured the four
sculpins and the eel. Finally, a fellow student found a bead that
she had used to mark a small Clinocottus analis near the mouth of
an anemone in one of her study tide pools.
All field observations with C californica were made using
SCUBA. Painted Greenlings were almost always found on the same
rocks as C californica, though contact was only observed once. A
fairly large greenling was found at 8 meters on a field of
anemones covering approximately one square meter. Using my
hand, I caused him to move three times before he left. Each time
he landed on another part of the anemone cluster, and none of the
anemones closed. The only reactions observed were occasional
slight retractions.
Another interesting observation was the abundance of
Snubnose Sculpins (Orthonopias triacis) and Blackeyed Gobies
(Coryphopterus nicholsiD on the same rocks as the greenlings and
C californica A Spubnose Sculpin was once seen with its pectoral
fin on an anemone that was still open. Finally, a Northern
Spearnose Poacher (Agonopsis emmelane) was found resting on
four anemones. I pulled him off by the tail and released him. He
immediately returned to the anemones, none of which showed any
reaction.
RESULTS: TANK OBSERVATIONS
With Anthopleura:
A Painted Greenling with bubble disease was placed in a
small tank with the three Anthopleura In the first hour, the fish
made contact with only the tentacles 5 times. In all 5 cases, the
anemones responded with a feeding motion; the tentacles rolled in
toward the mouth and then immediately back out. Then, as the fish
died an hour later, it landed in one of the A. xanthogrammica
twice. Both times the anemone attempted to eat the fish (exp. 1).
The other greenling tested with Anthopleura was in a much larger
tank. It stayed in the far left 1/3 of the tank, away from the
anemones, for the entire week of observation (Map *4).
Similar feeding responses were observed both times a
Clinocottus analis touched an Anthopleura (exp. *4). But unlike
the Painted Greenlings, Wooly Sculpins were often seen resting on
the pedal discs and columns of these anemones.
With Corynactis californica:
All the Painted Greenlings that acclimated to C californica
did so in approximately 3-5 days. For example, a greenling from
Hopkins Marine Station was put in a tank with two different clones
of C californica, one orange and one pink, neither of which had
been previously in a tank with fish. As time passed, the closure
values for the orange clone decreased, showing acclimation. These
values are listed in Table *1 and graphed in Figure +1. In the first
75 hours, there were only 9 observed contacts with the pink
anemones, so closure values were not calculated for individual
observation periods. Of these 9 hits, 4 caused complete closure
while 5 caused anemones to close 1/2 (a combined closure value of
2.4). However, in the 80 hour and 89 hour observation periods,
there were more hits and closure values of 0.2 and 1.0 were found
This demonstrates the greenling's ability to acclimate to more
than one clone at a time.
Interesting results were found when a greenling was
acclimated to the orange anemones in one tank, moved to a new
tank with pink anemones, and then returned to the original tank 11
days later. The original acclimation closure values with the
orange anemones can be found in Table *2. Figure *2 shows the
closure values when the fish is put in and taken out of the first
tank, when the fish is put in the second tank, and when it is
returned to the original tank. Note that after 11 days, the fish
was still fairly acclimated (closure values for hit hard, hit, and
brushed were 1.0, 0.81, and 0.6 respectively).
It is also interesting to follow different situations for a
particular group of C californica One at a time, four fish were
introduced to the clonal groups of anemones in one of the tanks
(exp. 6, 7 (Table *2), 8, & 10). Figure +3 shows the closure values
calculated from the first observation périod for each of these four
fish (2 greenlings, a Snubnose Sculpin, and then another greenling)
Each greenling had been previously isolated and all were
approximately the same size, yet the closure values upon
introduction to the tank go down with each successive greenling.
This implies some type of mechanical habituation by the anemones
to repeated contact by fish.
Another example of this mechanical habituation is the
acclimation periods of the second and third greenlings mentioned
above. The third greenling was entirely acclimated after 72 hours.
The second greenling, however, was only partially acclimated:
after 72 hours and was not fully acclimated until over 100 hours
after introduction to the tank. This is also shown in Figure r3.
Finally, I looked at all the "hit hard" contacts throughout my
observations. For each contact, I counted how many times that
particular anemone had been hit before. For anemones that closed,
they had previously been hit an average of 1.6 times. Anemones
that closed 172 had been hit an average of 4.0 times, while slight
retraction anemones had been hit an average of 5.5 times. This
shows the mechanical habituation by individual anemones.
RESULTS: FORCED CONTACT EXPERIMENTS
Using forceps to hold wooly Sculpins by the jaw, contact
with the tentacles of Anthopleura was forced. One small A.
xanthogrammica had its mouth already partly open. A small (3.
cm.) sculpin was touched to the tentacles, and as soon as there
was contact, the fish was engulfed. Within 1 second I could not
even see the fish, let alone try to save it. Next I tested a similar
fish (3.3 cm.) with A. elegantissima It was caught and
disappeared as quickly as the fish above. Both anemones were
fully opened around 8 minutes after catching the fish. Obviously,
Anthopleura are willing to eat Clinocottus analis.
Body parts from two acclimated greenlings were tested with
both acclimated and non acclimated anemones. The results can be
found in Table *3. For the pectoral fins of both fish and the
ventral fin of the second fish, the acclimated anemones responded
C
less than the previously isolated anemones. But for the tails of
both fish and the ventral fin of the first fish, there is little or no
difference, possibly due to contamination by blood when the fish
had its spinal cord cut.
As a final experiment, l acclimated my index finger to a
clone of C californica After ten minutes of rubbing my finger on
closed anemones, I was able to lightly touch the tentacles of an
opened anemone on the same rock without causing a reaction.f
then touched this anemone with a different finger, the anemone
would close. The acclimated finger was covered by a slimy
substance that must have been anemone mucus. It is this mucus
that protected my finger from nematocyst discharge.
DISCUSSION AND CONCLUSIONS
Observations with Anthopleura show that fish did not
acclimate to the tentacles of either A. elegantissima or A.
xanthogrammica. Painted Greenlings avoid these anemones
completely. Wooly Sculpins, on the other hand, merely avoid the
tentacles. They will rest on both the pedal disc and column of an
anemone, and were even seen using an anemone for protection in
the field. If contact with Anthopleura tentacles does occur,
however, the anemone will attempt to eat the fish.
It is also clear that Painted Greenlings (and probably
Northern Spearnose Poachers, Snubnose Sculpins, and maybe
Blackeyed Gobies) do acclimate to the Strawberry Anemone,
Corynactis californica This acclimation period takes
approximately 3-5 days.
The acclimation process seems to occur through two main
mechanisms. The first involves a change in the mucus layer of the
fish. Though there have been claims that the fish secretes the
protective substances, it seems more likely that the fish picks up
anemone mucus through repeated contact. It is this mucus layer
that prevents the fish from stimulating the chemoreceptors to
cause nematocyst discharge. This is demonstrated by the
acclimation of my finger to the anemones. Another observation
that supports this theory was that even after acclimation, on the
rare occasion that the face of a fish hit an anemone, the anemone
always closed. If the fish was producing a protective mucus layer
itself, the face would also be protected. Since the face rarely
touched anemones, it did not accumulate enough anemone mucus
for protection.
The Painted Greenling seems able to maintain this
protection through its mucus layer for a long time. One fish was
removed from its acclimated anemones for 11 days and was still
fairly acclimated upon return to the tank. This is quite different
from the inability of Amphiprion bicinctus to remain acclimated
to Radianthus koseirensis (Graefe, 1964). Graefe observed that if
this fish was seperated from its host anemone for 1 hour or more,
it would be stung upon its return.
One main theory behind mucus protection is that the mucus
does not contain any of the active chemicals needed to trigger the
chemoreceptors. Another claims that certain chemicals inhibit
nematocyst discharge, possibly by raising the threshold to
mechanical stimuli. Finally, the anemone may recognize its own
mucus on the fish and identify the fish as part of itself.
It seems very unlikely that the mucus contains inhibitors to
nematocyst discharge. If inhibitors were present in the anemone's
mucus layer, they would interfere in the anemone's normal feeding
and protective reactions.
One experiment that may support the lack of excitatory
factors was the testing of greenling body parts with anemones.
While some fins caused a notable difference in reactions between
acclimated and nonacclimated anemones, others did not. Those
that did not may have been contaminated by blood when the fish
was killed. If the fins were covered with something that inhibited
nematocyst discharge, the contamination should not have caused
all anemones to react the same. Even if the contamination
contained chemicals that were active enough to override the
inhibition (lower the threshold to mechanical stimult more than
the inhibitors raised it), the reactions of acclimated anemones
should still have been somewhat less than those that were not
affected by the inhibitors at all. Inhibition must not be the
mechanism. On the other hand, if the mucus layer merely
contained no active chemicals, contamination would be expected to
cause similar reactions by both acclimated and nonacclimated
anemones. It is still possible, however, that there was merely a
lack of mucus on the areas touched against the anemones. This
would also cause all anemones, acclimated or not, to react the
same.
The most likely mechanism for protection is that the
anemone recongnizes the fish as part of itself. All observations
agree with this expanation. For example, consider that any
particular greenling can acclimate to more than one clone of C;
californica at a time. An acclimated fish put in a tank with a
different clone of anemones will be entirely unacclimated. So
even if there are no active chemicals to cause discharge of the one
clone, another clone will react. A fish in a tank with two clones
would pick up mucus from both clones. Assuming that the lack of
active chemicals causes protection, this combined mucus should
still contain chemicals from each clone that are active enough to
cause discharge on the other clone. If, however, each clone gives
the fish mucus that it can recognize, the fish will be recongnized
by both and will not cause nematocyst discharge on either. This is
what was observed, causing me to believe that this is the more
probable mechanism.
The other important aspect of acclimation involves
habituation by the anemones to repeated contact by the fish. This
was first observed in a three clone tank of C californica in which
successive greenlings showed lower initial closure values and
shorter acclimation periods. It was also seen in individual
anemones in that stronger reactions were observed for anemones
that had been previously hit less times.
The mechanism for this habituation is unknown, though it
probably involves modulation by the nervous system. Thorington
and Hessinger (Thorington and Hessinger) showed that various
types of modulation do occur for certain species of anemones. For
example, one experiment showed that tactile stimulation of the
body column of Anemonta sargassensis will increase the amount
of discharge by the cnidae on the tentacles. No tests were done on
habituation to repeated contact.
Besides the mechanism of habituation to contact for the
anemones, there are other aspects of acclimation that need to be
studied further. A complete examination of anemone mucus would
help to determine how the anemone recognizes its own mucus if
this indeed is the mechanism behind the protection from
nematocyst discharge. Possibly one could determine the chemicals
detected by the chemoreceptors. It would also be interesting to
see if an active chemical present on the fish could cause discharge
despite recognition by the anemone of its own mucus.
e
LITERATURE CITED
Burgess, W.E. and Axelrod, H.R.,1984, Fishes of California and Western
Mexico, TFH Pub. Inc, New Jersey.
Carlgren, O., 1949, A Survey of the Plychodactiacea, Corallimorpharia
and Actiniaria, London.
Davenport, D. and Norris, K., 1958, Observations on the Symbiosis of
the Sea Anemone Stoichactis and the Pomacentrid Fish,
Amphiprion Percula, University of Califonia.
DeMartini, E.E., 1946, The Adaptive Significance of Territoriality and
Egg Cannibalism In the Painted Greenling, Doctoral
dissertation, University of Washington.
Friese, U. Erich, 1972, Sea Anemones, TFH Publications, Hong Kong.
Gohar, H.A.F., 1948, Commensalism Between Fish and Anemones, Pub.
Marine Biology, Stn Ghardaga 6, pp 35-44.
Hand, C., 1955, Sea Anemones of Central California, Wasmann journal
of Biology.
Lubbock, R., 1979, Why are Clownfish not Stung by Sea Anemones,
University of Cambridge, G.B..
Mariscal, R.N., 1966, The Symbiosis Between Tropical Sea Anemones
and Fishes: a Review. pp 157-171 in Bowman, R.I. ed., The
Galapagos: Proceedings of the Symposia of the Galapagos
International Scientific Project, University of CA press,
Berkeley.
Sebens, K., 1977, Habitat Suitability, Reproductive Ecology, and the
Plasticity of Body Size in two Sea Anemone Populations,
Doctoral Dissertation, University of Washington
Thorington and Hessinger, 1988, Control of Discharge: Factors
Affecting Discharge of Cnidae. pp 233-250 in Hessinger and
Lenhoff ed.s, The Biology of Nematocysts, Academic Press, San
Diego.
APPENDIX
Exp. *1: During the dying process (the last four minutes), the fish
would shoot to the surface and then sink to the bottom. He
landed with his tall in the small A. xanthogrammica and his
tail was held by the tentacles. He swam out after 10 seconds
and again went to the surface and sank. Again he landed with
his tail in the small A. xanthogrammica and appeared to be
dead. The anemone pulled the greenling in towards its mouth
and after 90 seconds he was nearly 1/2 way in. Suddenly, the
greenling started thrashing violently and after seven seconds
was free. He landed on the sand and died.
Exp. *4: A Wooly Sculpin was dropped into the tank. It was still
somewhat dulled from CO» and just sank when 1 dropped it in.
On the way down, it touched three tentacles of the large A
xanthogrammica On contact, the fish immediately swam hard
away from the anemone. The three touched tentacles rolled in
and then immediately back out. The only other contact between
anemone tentacles and fish occurred when a piece of squid was
dropped to an anemone. A sculpin saw the squid just before it
reached the anemone and swam for it. But before it got to the
squid, the fish touched some tentacles and immediately
changed direction and swam away quickly. The anemone got the
squid and closed around it.
A Painted Greenling was also in this tank for over a week.
After the first hour, in which he sat on the right side of the
tank where I had dropped him in, I never saw him as far right as
the most left anemone (Map 4). He completely avoided them all.
Exp. 46: A Painted Greenling was put in a tank with at least three
different clones of C californica (Map 1). During observations
in the first 90 minutes, closure values for hit hard, hit and
brushed were 3, 3, and 2 respectively. The fish died of bubble
disease before further observations could be made.
Exp. *8: A Snubnose Sculpin was put in the same tank as (exp. 16).
In the first 90 minutes, closure values for hit hard and hit
were 3.0 and 1.13 respectively. For the next three days, the
fish touched no anemones, and was then removed.
Exp. +10: Another Painted Greenling was placed in the same tank
(exp. *6) and observed for the first 1 hour. The closure value
for hit during this period was 2.2. Observations were not made
again until 72 hours after introduction to the tank. Closure
values at this point were 1.0, 0.0, and 0.0 for hit hard, hit, and
brushed. After 80 hours, hit and brushed still had closure
values of 0.0.
c
TABLE *1 (PAINTED GREENLING WITH CORVNACTIS CALIFORNICA)
CLOSURE VALUES
THREE TYPES OF CONTACT
HIT
TIME (In Hours)
HIT HARD
BRUSHED
2.79
O-
1.33
—
1.5-3
1.0
Z.12
18-20
2.30
2.0
24-26
2.57
1.0
32-33
1.80
39-41
1.83
54-55
2.54
64-65
1.60
70-71
3.0
1.0
0.8
75-76
1.0
0.0
80-81
3.0
1.29
88-89
1.06
106-107
0.55
TABLE *2 (PAINTED GREENLING WITH CORWACTIS CALIFORNICA)
CLOSURE VALUES
THREE TYPES OF CONTACT
TIME (In Hours)
HIT
HIT HARD
BRUSHED
O-
2.63
1.0
65-66
3.0
0.0
72-73
1.0
1.17
0.0
95-96
1.0
0.56
0.0
117-118
1.0
1.50
0
TABLE F3 (PAINTED GREENLING FINS TOUCHED TO BOTH ACCLIMATED
AND NONACCLIMATED CORWACTIS CALIFORNICA)
FISH FIN TYPE ANEMONE
REACTION TIME BEFORE FULLY OPENED
TAIL
CLOSED 1/2
2.50
ACCL
410
TAIL
ACCL
CLOSED 1/2
420
TAIL
NON
CLOSED
5:15
TAIL
NON
CLOSED
410
VENTRAL ACCL
CLOSED
VENTRAL ACCL
5.30
CLOSED
VENTRAL NON
5:15
CLOSED
5:10
VENTRAL NON
CLOSED
PECTORAL ACCL
2:40
CLOSED
3:40
PECTORAL ACCL
CLOSED
PECTORAL NON
8:50
CLOSED
PECTORAL NON
9.30
CLOSED
CLOSED
5:10
TAIL
ACCL
6:40
TAIL
ACCL
CLOSED
6.30
TAIL
CLOSED
NON
VENTRAL ACCL
0.30
CLOSED 1/2
450
VENTRAL ACCL
CLOSED 1/2
VENTRAL NON
9.30
CLOSED
4.15
PECTORAL ACCL
CLOSED 17/2
PECTORAL ACCL
3.10
CLOSED 1/2
PECTORAL ACCL
3.50
CLOSED 1/2
PECTORAL ACCL
6.20
CLOSED 173
PECTORAL NON
CLOSED
12:00
PECTORAL NON
CLOSED
8:10
ACCL= Anemone clone fish was acclimated to
NON= isolated clont
FIGURE LEGENDS
FIGURE 1: This graph shows a Painted Greenling becoming
acclimated to the C californica shown in Map 3. The line
represents closure values for hits, the dark spots on top
represent hit hard, and the plus signs represent brushes. The
one high brush mark and the highest hit mark are from
observations in which the greenling's face hit and closed
anemones.
Figure 2: This shows closure values for a greenling in a variety of
different situations. First it is introduced to orange C
californica and then is taken out after acclimation (140 hours).
It is put in with a new clone (pink) and again is completely
unacclimated. Finally, after 11 days, the fish is put back in
with the orange anemones and is still fairly acclimated, but not
completely.
Figure 3: The first four bars represent the introduction of different
fish to the same clone of C californica Notice the initial
closure values go down for each successive greenling. The final
two bars show that the second greenling introduced took longer
to acclimate than the third greenling. This is due to
mechanical habituation.
8
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ACKNOWLEDGEMENT
would like to thank the professors and staff (especially in the
library) at Hopkins Marine Station for all their help. I would also like
to thank the many graduate students who were willing to take the
time to help when asked. Also, I thank Maggie Harrison for being my
relfable dive partner. Finally, I want to give special thanks to Chuck
Baxter for his time, insights, and guidance.