INTRODUCTION
A question of fundamental importance to the biologists that study
the intertidal zone concerns the impact of the resident and visiting
fishes on the community. Most intertidal studies are carried out while
the tide is out and thus predatory or browsing activities of fishes,
especially the nearshore forms that move in with the tide, is not generally
appreciated. Dietary studies done on tidepool and coastal species yield
some information but a focused community approach is really required to
resolve the problem.
A number of studies on the diet and abundance of nearshore fishes
of California can be found throughout the literature. Limbaugh (1955)
described the habits of the fishes found in Southern California kelp
forests. Quast (1968a-c) examined the diets and population sizes of the
kelp associated fishes found at two sites in Southern California and one
site in Baja California. Miller and Geibel (1973) researched the various
fish populations in the kelp forest off the Hopkins Marine Station, the
site of this study. Finally of importance, Feder, Turner, and Limbaugh
(1974) studied the relationship between the kelp plant Macrocystis pyrifera
and the fishes inhabiting its forest in Southern California. In
addition, there have been studies carried out on the tidepool fishes
specifically. Mitchell (1955) analyzed gut contents of tidepool fishes
on the Palos Verdes Penninsula in Southern California.
This study focused on the species in and adjacent to the intertidal.
examining their diets, behavior patterns and abundance to see what impact
these predators have upon this zone. The intertidal algal turf is a
region only periodically available to open water fishes and it thus seems
possible that a less-exploited food supply can be found there. Observations
show a large number of fish moving into the intertidal at high tide
despite often surgey, churning conditions which would seem to make it an
inhospitable environment. In addition, the large population of resident
tidepool fishes must exert a substantial predatory effect on the intertidal
algae and resident invertebrates. This study takes a habitat approach
investigating both the permanent residents and the larger invaders of
the intertidal zone to get an over-all picture of the impact of fish on
this rich algal dominated community.
MATERIALS AND METHODS
Collection of Specimens
The period of collection for specimens to be used in gut analyses
lasted from April 23 to May 28, 1979. All fish were taken from very
shallow water (eight feet or less) between the Monterey Cannery and
the northwest corner of the Hopkins Marine Reserve (see Figure 1 for
collection sites). Three methods of collection were employed. The larger
fish, those not left in tidepools at low tide were speared while skin
diving using a pole spear. Spearing was an ideal method of collection
because fish could be taken directly from either the intertidal zone
or from water very near shore at low tide. Care was applied in not
depleting any population from a particular area.
The larger tidepool fishes were taken by poke-pole fishing. A ten-
inch monofilament leader with a number 6 or 8 size hook was attached to
the end of a fishing rod, mussel was used for bait. By dangling the
baited hook into crevices and between rocks reasonably large fish were
captured from pools at low tide. Smaller tidepool fish were collected
with a twelve-inch diameter dipnet. Catching the quick-moving sculpins
was often difficult unless they were in a shallow pool.
Following collections, the fish identifications were made with
references to Miller and Lea (1972), Gotshall (1977), Bolin (1975).
Fitch and Lavenberg (1975). Bane (1971), and Roedel (1953). Then the
entire intestinal tract of each fish was removed and preserved in 85%
isopropyl alcohol.
Gut Analyses
Within a week after its collection, each preserved intestinal
tract was sliced open and the partially digested food material was
examined under a dissecting microscope. Estimates were made of stomach
fullness. The food material was then identified as precisely as possible
to major group and when appropriate to genus and species, using Light
(1975) and Abbott and Hollenberg (1976) as references. Make note that
the entire gut was examined, not just the stomach, because in most cases
there was identifiable material found beyond the stomach.
Transect Fish Observation Study
A twenty meter long transect line perpendicular to the shore, was
chosen in a gently sloping area of the intertidal zone in the Hopkins
Marine Reserve. This became the site for a study on intertidal densities
and feeding habits of several fish species. The designated region included
ten meters to either side of the transect line making a total transect
area of 400 square meters stretching along twenty meters of coastline.
Twelve observational high tide dives were made in the area each one
lasting for approximately twenty minutes. A clipboard with plastic-
coated paper was used to record numbers of each species of fish observed
within the area. Notes were also taken on the foraging and feeding
behavior of the fish sighted.
RESULTS
A total of 129 fish were collected, eighty-eight were speared,
twenty-one were caught by poke-pole fishing, and twenty were captured
with the dipnet. The species collected and the number of each taken is
listed below:
Clinocottus analis
28
Wooly Sculpin
Embiotoca jacksoni
18
Black Surfperch
Striped Surfperch Embiotoca lateralis
Oxyjulis californica
Senorita
Damalichthys vacca
Pile Surfperch
Sebastes atrovirens
Kelp Rockfish
Scorpaenichthys marmoratus
Cabezon
Cebidichthys violaceus
Monkeyface-eel
Rainbow Surfperch Hypsurus caryi
Hexagrammos decagrammus
Kelp Greenling
Paralabrax clathratus
Kelp Bass
Sebastes rastrelliger
Grass Rockfish
Black Prickleback Xiphister atropurpureus
Striped Kelpfish
Gibbonsia metzi
The results are presented seperately for each of the majjor species.
The composite results are presented for the diets in Table 1, intertidal
food items are treated in Figure 2, and Tables 2 and 3. Comparisons of the
diets of juvenile and adult wooly sculpins are given in Figure 3.
The abundance of fish in the study quadrat are reported in Figure 4
and Table 4. In addition, complete stomach contents data is in the appendix.
Black Surfperch
(Embiotoca jacksoni)
All eighteen specimens were collected by spear, the standard
length size range was 21.0 cm to 27.5 cm. This was one of the most
frequent visitors to the intertidal, present in 100% of the transect
observation periods. An average of three specimens were present during
each period, the adults being three or four times more common than the
juveniles
Its principal foods were polychaetes, amphipods, isopods, gastropods,
crabs, and algae, however the algae consumed was most likely incidental,
taken for the invertebrates found on its fronds. This fish definitely
feeds a great deal intertidally because 70% of all specimens had at least
three seperate intertidal species in their guts. In addition, between
sixty and seventy percent of the food items found came from the intertidal.
A great deal of its foraging seems to have occurred among Phyllospadix
blades since 75% of the specimens examined had this plant in their guts.
In the field, the black surfperch was observed nibbling at low
intertidal Phyllospadix that was encrusted with bryozoa, mouthfuls were
taken approximately every thirty seconds. Another common feeding
technique noted several times begins with the fish taking a mouthful
from the sand bottom or algal turf. It chews for a few seconds then
expells sand and bits of algae from its mouth, presumably retaining
invertebrates to swallow. In spite of the expulsion of sand from their
mouths, most of these fish had large quantities of sand in their guts.
sometimes comprising up to 90% of their entire gut material.
(Embiotoca lateralis)
Striped Surfperch
All sixteen specimens were taken by spear, the standard length
range was 21.0 cm to 29.0 cm. The striped surfperch was observed in
83% of the transect observations, averaging 2.5 fish per time period,
juveniles were rarely encountered.
Polychaetes, amphipods, isopods, gastropods, crabs, and algae
dominated their diet as in the black surfperch. Between seventy and eighty
percent of their food came from the intertidal. Furthermore, 57% of
the specimens had three or more intertidal species of invertebrates
or algae in their guts. The mouth structure is very much like that of
the black surfperch with short, stubby teeth.
The sane feeding behavior employed by the black surfperch of spitting
out sand was noted in the striped surfperch, except that the quantity
of sand expelled seemed to be noticeably less. Often the striped surfperch
was observed foraging side by side with the black surfperch and the
pile surfperch in the intertidal.
Pile Surfperch
(Damalichthys vacca)
The eleven specimens were collected by spear, their standard
length size range was from 12.5 cm to 29.5 cm. The pile surfperch
was always found in the intertidal, appearing 100% of the days of the
transect study in an average number of 3.3 fish per time period,
juveniles composed one-fourth of this number. It was however, the
shyest member of this family and usually left the area immediately as I
entered the site, so this figure may actually be substantially larger.
The diet was composed primarily of gastropods and hermit crabs
or (Paguridae), this significantly differs from the foods of the black
and striped surfperches. Approximately sixty percent of their food
came from the intertidal and ten of eleven specimens had at least one
intertidal food item. This species also has the typical perch mouth
but with teeth slightly sharper than the black or striped surfperches.
This fish was never observed nibbling from the algal turf, instead
it often hovered over sandy areas. A favorite hideaway while not in the
intertidal is underneath the kelp canopy where schools often aggregate.
Kelp Rockfish
(Sebastes atrovirens)
Nine specimens were collected by spear, the size range of standard
lengths was 20.5 cm to 30.0 cm. The kelp rockfish was never observed
in the intertidal transect area, however it commonly was found in very
shallow water of nearby locations.
Interestingly, nearly 60% of the fish taken had empty stomachs, and
none had over 20% of their guts filled. The principal identifiable
foods were tiny fish, gastropods, and shrimps. Quast (1968c) also noted
this high percentage of empty stomachs and speculated that perhaps
they were suffering from malnutrition. Larson (1972) instead proposed
that their diet of primarily soft-bodied organisms passes through the
gut very quickly so that there is little accumulation.
This species was most often seen swimming very close to the bottom.
On occasions they were found hovering vertically head-down near a shallow
water kelp plant, appearing much like a kelp blade.
(Oxyjulis californica)
Sennorita
Sixteen specimens were collected by spear with a standard length
size range of 14.0 cm to 19.6 cm. The senorita is an extremely abundant
fish in shallow waters and commonly enters the intertidal. It was observed
in 75% of the transect observation periods, averaging 2.9 fish at a time
within the 20 x 20 meter intertidal transect area.
Its diet consisted of bryozoa, algae, fish and in smaller quantities
gastropods, isopods, and shrimps. Roughly one-half of the food found in
the gut was found intertidally, there was also a very high sand content.
The teeth on this fish are tiny but very sharp and the mouth projects
forward from their pointed snout.
These fish were often seen picking at algae and the sand bottom,
a well known habit of the senorita. Limbaugh (1955) says they "feed on
most any animal protein, continually picking at small objects on the
bottom, on plants, other fishes, or from the water itself." Once in the
field, a senorita was seen that picked up a Tegula funebralis snail,
dropped it, picked up another one and swam off with it.
(Scorpaenichthys marmoratus)
Cabezon
Five specimens were collected by spear and three by hook and line,
standard lengths ranged from 14.2 cm to 30.0 cm. The cabezon was found
both in tidepools and subtidally, entering the intertidal at high tides.
It was observed in the 20 x 20 meter quadrat only 42% of the time.
averaging 0.5 individuals per time period. Juveniles were far more common
than adults, however, several large fish up to 40 cm in length were found
in very shallow intertidal water.
10
Eighty percent of all food items were from the intertidal and
seventy-five percent of the specimens had four or more different intertidal
species present in the gut. The most common foods were the crabs, found
in 100% of the stomachs, also tiny fish and shrimps were important.
The many species of intertidal algae found show their extensive intertidal
feeding. Endocladia muricata, one of the highest growing marine algae
was found in the gut of one larger specimen.
Actual feeding was never observed, but the fish was most often
seen lying motionless against the bottom presumably waiting for prey to
appear. Their mottled coloration blends extremely well with the algae-
covered rocks, an unwary potential prey invertebrate could easily oversee
this cryptic fish. During surgey periods the cabezon wedges itself
into rock crevices for protection where it can withstand the turbulance
and remain in the intertidal to feed.
(Clinocottus analis)
Wooly Sculpin
Twenty-eight specimens were collected, sixteen with a dipnet and
twelve using hook and line, all were taken from tidepools. The standard
lengths ranged from 3.7 cm to 14.0 cm. This is the most abundant
tidepool fish in the study area, usually several can be found living
under any tidepool rock. The transect abundance figures are extremely
deflated because of the sculpins' small sizes, inconspicuous coloration.
and lack of movement. However, a trained eye should be able to spot
several sculpins within a square meter of sandy intertidal substrate.
Approximately 90 of their food came from the intertidal, the rest
is most likely taken as part of the drift that washes inshore. Overall,
they consumed a wide variety of foods, dominated by crabs, gastropods,
and polychaetes. However, there was significant difference between the
foods of the small and large members of the species. The small fish
ate far more polychaetes and isopods than the larger fish. The large
fish in turn, consumed many more crabs and limpets. In fact, 80% of
the large specimens had one or more of the limpets Collisella scabra
and Collisella digitalis in their guts while not a single specimen
under 9.9 cm standard length had recently eaten a limpet. Their rows
of fine, sharp teeth and strong fleshy jaws are apparently well designed
to remove these limpets from rocks.
Wooly sculpins are speedy swimmers, and use this to their advantage
in their technique of feeding. They too lie motionless, camoflauged
against the bottom waiting for prey to appear, once they spot their
victim they dart out and grab it before the crab or worm knows what has
happened, this procedure was noted on several occasions.
(Cebidichthys violaceus
Monkeyface-ee
Seven specimens were collected by hook and line, one was captured
using a dipnet, all were found in tidepools, under rocks, or in caves.
The standard lengths covered the widest range of any fish studied, from
12.0 cm to 37.0 cm. Quantitative transect abundance studies were very
difficult to perform since this fish is rarely visible in the open
water, at most a head may be seen hanging beneath a rock. The frequency
of capture by hook and line however, suggest its great abundance in pools,
The monkeyface-eel has a very unique diet, consisting almost
entirely of algae. The only animals present were bryozoan colonies
which were located on the algal fronds. This bryozoa was present in
nearly 60% of the specimens yet always in very small ammounts. In all
seven species of intertidal algae were found in the guts. Not a single
subtidal species was found implying that all of the monkeyface-eels
feeding occurs in the intertidal.
Although this species has a primarilyherbiverous diet, the specimens
were caught using mussel for bait. As the meat was dangled at the
mouth of a cave the monkeyface-eel inhabitant stuck its head out, grabbed
it, and quickly retreated to its cave
Other Fishes
Several other species were dealt with but to a much less extent
than the preceding fishes. They will be dealt with briefly.
Four specimens of the kelp greenling (Hexagrammos decagrammus),
all females, were speared in the intertidal or close by in very shallow
water. They were occasionally observed in the transect area, present
17% of the time, with an abundance of O.24 individuals.
Crabs composed a good part of their diets and in each case several
small eels and fishes had also been consumed. There were in addition.
a few intertidal species, though the sample number is very small, and
percentages could not be determined. Kelp greenlings were usually
observed swimming very close to the bottom searching for food.
Four rainbow surfperch (Hypsurus caryi) specimens were speared in
shallow water. This species was never actually observed in the intertidal.
it usually inhabits deeper water. They are also very shy fish, not
comfortable around divers, much like the pile surfperch. Their diet
consisted mostly of amphipods and isopods.
Two specimens of the kelp bass (Paralabrax clathratus) were taken
very near the intertidal zone. Their guts were entirely filled with
fish remains. These fish are very prominent in the deeper kelp beds
and most likely do little feeding in the intertidal.
Two grass rockfish (Sebastes rastrelliger) were speared in the low
intertidal. Only some crab parts and polychaetes were identified from
their highly digested gut contents. This fish too is a common inhabitant
of the kelp beds and most likely has little to do with the intertidal.
One black prickleback (Kiphister atropurpureus) was caught on
hook and line from a tidepool. Its gut contained several species of
intertidal algae, some bryozoa and a Tricolia pulloides snail on an
algal frond. The diet and habitat of this fish seems very similar
to that of the monkeyface-eel, although it does not appear to be as
abundant in the study area.
One striped kelpfish (Gibbonsia metzi) was also caught by hook
and line from a tidepool crevice. Its gut contained an isopod and a
Pachycheles rudis crab. These fish were often seen hiding among the
bases of Phyllospadix and algae in very shallow water and feeding on the
invertebrates in these low intertidal forests.
DISCUSSION
The three surfperches examined appear to have an important impact
on the mid to low intertidal. The black and striped occupy a very
similar feeding niche while the pile surfperch differs from these two
species significantly. The bulk of the diets of the black and striped
surfperches comes from the algal turf community. By pulling off chunks,
they obtain a great deal of polychaetes, sipunculids, small gastropods,
amphipods, isopods, and tiny crabs. This procedure results in an intake
of unwanted algae and sand. It is assumed that algae is not the desired
foodstuff since most algal pieces are expelled after a mouthful is taken
and becomes part of the drift.
It is interesting to speculate on the impact this feeding method
has on the intertidal turf. Assume thirty mouthfuls are taken a day per
fish, each consisting of approximately 2 cm of turf area. Consequently,
about 60 cm" is pulled off each day by one fish. Intertidal fish density
estimates for fishes closely associated with the intertidal show
approximately one black or striped surfperch every 3-6 meters of shore
length. Imagine the extent of turf removal over a few months or even
weeks. Perhaps these fish are the gardeners limiting the growth of the
algae, their exclusion from an area may result in lush algal growth.
Invertebrate grazing upon algal fronds may have only minor impact on
the algal forest when compared to the fishes' activities.
Although the pile surfperch is often seen foraging with black and
striped surfperches within the intertidal, this species is exploiting
a quite different resource. The low presence of algae and associated
15
turf invertebrates indicates the pile surfperch is not foraging on the
algal turf. Instead it appears to be a selective visual feeder on larger
snails and hermit crabs. Sandy and bare rocky substrates are favorite
feeding locations. Hermit crabs were found both in and out of shells
along with an abundance of broken shell pieces in the gut which indicates
their ability to crack shells in their mouths and throats, Quast (1968c).
The impact the pile surfperch has upon the intertidal zone differs
significantly from that of the black and striped surfperches, since its
feeding does not involve incidental algal pruning. The pile surfperch
is definitely an important predator of the tidepool-inhabiting hermit
crabs. Larger hermit crabs in tidepools have little to fear because
of their size and protective shell. However, once the tide comes in these
fish predators appear, forcing the hermits to take cover. This may be
a primary reason why the hermit crabs often come up with the tide to
inhabit the shallowest, most protected water possible, Prohaska (1979).
The senorita does not directly affect the intertidal as much as
the surfperches do, however its scavenging carries over into this
zone at high tides. This species avoids fast-moving, large prey items,
and is described as a picker on small, slow, or immobile forms. Bryozoa
occurred most frequently in its diet in this study. The high occurence
of fish remains may come from scavenging activity or feeding on the smaller
juvenile rockfish. The most practical method of feeding for this species
with its tiny mouth is to pick at their food material working over
bottom and algal substrate. This fish is even known to feed on zooplankton.
when abundant in the water. Because of the wandering and picking
16
habits of the senorita it seems to have no preference for feeding
intertidally since its nearshore abundance could provide a much higher
intertidal density than was actually observed.
The cabezon primarily comes to the intertidal to feed on crabs.
Crabs were often removed from the gut with their legs and claws all
present and neatly folded indicating that they were captured by surprise,
probably because they did not perceive the cryptic cabezon. This fish
exerts a similar effect on the intertidal crabs such as Pachygrapsus,
by invading their environment only at high tides like the pile surfperch
does to Pagurus.
The tidepool fishes inevitably occupy extremely important niches
in the food chain of the intertidal. They are the top predators for at
least half the tidal cycle. Without doubt, if the wooly sculpin was
excluded from the intertidal the entire community would change drastically
The difference in diet between large and small wooly sculpins
can be simply explained by their different mouth and body sizes. A long
slender worm would be a far more practical prey item for a young sculpir
than would a bulky crab or limpet, which are preferred by the adults.
The general small size of wooly sculpins allows them to remove invertebrates
from algae without taking large quantities of it into their stomachs.
The most important algal feeders seem to be the monkeyface-eels
and the pricklebacks. By observing the fullness of the guts it appears
that these are herbiverous organisms which require large quantities of
plant material to obtain sufficient nutrition. Although bryozoa was
frequently encountered it never seemed to be abundant enough to provide
adequate food energy to fuel these large fishes.
Large monkeyface-eel specimens often had entire blades of Iridaea
cordata and good-sized clumps of Gigartina papillata in their gut.
These large algal pieces must have been pulled off the intertidal
turf to eat. If this fish was scavenging from the drift the guts would
probably contain substantial quantities of subtidal algae species as well.
These species may be important trimmers of the intertidal algae.
Larger blades and fronds of algae are abundant and easier to obtain
than the short "well-pruned" plants.
A 30 cm monkeyface-eel could easily consume 225 cm of mature
Gigartina papillata a day. These fish are known to be territorial, often
living in a particular cave for several years. At this rate of feeding.
one of these fish would very soon deplete the nearby algae surrounding
its home. Since no unusually bare patches were observed near monkeyface-
eel caves, it is assumed that most feeding occurs away from their home,
This may be an excellent species to test for dietary sufficiency of
algae on aquarium specimens along with enzymatic studies to see if
herbivory is truly the mode of feeding in this species. The previous
suspected herbiverous species (opaleye, zebraperch, and halfmoon)
have been placed in doubt (Quast 1968b).
In closing, this study has demonstrated the significant roles
that tidepool and nearshore fishes play in intertidal ecology and has thus
established the need for further study in the field. Different environments,
seasonal variations, and diurnal cycles are among the essential elements
for follow-up investigations on the fishes of the diverse intertidal
community.
18
SUMMARY
1. Studies were done on exploitation of intertidal food resources by
resident intertidal species and inshore movement of the nearshore
fishes. Presence of intertidal food items in the gut contents of
several nearshore fishes suggest extensive feeding occurs in the
intertidal during high tides.
2. Gut content analyses of tidepool fishes implies that most if not all
of their feeding occurs intertidally. Wooly sculpins were the most
common tidepool species and consumed a wide variety of food items.
The monkeyface-eel appears to be herbiverous and consumes large
quantities of intertidal algae.
Observations and gut content examinations of the black and striped
surfperches show they obtain tiny invertebrate food items by picking
off the algal turf.
The pile surfperch concentrates its diet on large visible food items,
primarily hermit crabs and gastropods.
The feeding activities of the black and striped surfperches and the
monkeyface-eel result in tearing and trimming of the algal turf and
thus have substantial impact on the intertidal community.
ACKNOWLEDGEMENTS
I would like to thank Bill Magruder for his help in algae
identification, Dr. Don Abbott for his energy and inspiration, and
especially to my advisor Chuck Baxter, a long time fish-lover, for
his wisdom and his Green Death.
19
20
LITERATURE CITED
Abbott, Isabella A. and George J. Hollenberg. 1976. Marine algae of
California. Stanford University Press, Stanford, Calif. 827 p.
Gilbert W. and Anneka W. 1971. Bay fishes of Northern California.
Bane
Mariscos Publications, Southampton, N. Y. 137 p.
Bolin, Rolf L. 1975. Key to intertidal fishes, in S. F. Light, Intertidal
invertebrates of the Central California coast, rev. by Ralph I.
Smith, Frank A. Pitelka, Donald P. Abbott, Francis H. Weasner and
others. Univ. of Calif. Press, Berkeley. p. 656-688.
Howard M., Charles H. Turner, and Conrad Limbaugh. 1974.
Feder
Observations
on fishes associated with kelp beds in Southern California. Calif.
Dept. of Fish and Game Fish Bull. 160, 131 p.
Fitch, John E. and Robert J. Lavenberg. 1975.
Tidepool and nearshore
fishes of California. Univ. of Calif. Press, Berkeley. 146 p.
Gotshall, Daniel W. 1977.
Fishwatchers' guide to the inshore fishes
of the Pacific coast. Sea Challengers, Monterey. 105 p.
Keen, A. Myra and Eugene Coan. 1974. Marine molluscan genera of western
North America. Stanford Univ. Press. Stanford, Calif. 258 p.
Larson, Ralph John. 1972.
The food habits of four kelp-bed rockfishes
(Scorpaenidae, Sebastes) off Santa Barbara, California. MS thesis
Univ. of Calif., Santa Barbara. 57 p.
Light
S. F. 1975. (Eds.) Ralph I. Smith and James T. Carlton. Intertidal
invertebrates of the Central California coast. Univ. of Calif.
Press, Berkeley. 716 p.
Limbaugh, Conrad. 1955. Fish life in the kelp beds and the effects of
kelp harvesting. Univ. of Calif. Inst. Mar. Res., IMR Ref. 55-9: 234 p
Miller, Daniel J. and Robert N. Lea. 1972. Guide to the coastal marine
fishes of California. Calif. Dept. of Fish and Game Fish Bull.
157. 213 p.
Miller, Daniel J. and John J. Geibel. 1973. Summary of blue rockfish and
lingcod life histories; a reef ecology study; and giant kelp.
Macrocystis pyrifera, experiments in Monterey Bay, California.
Calif. Dept. of Fish and Game Fish Bull. 158. 137 p.
Mitchell, D. F. 1953.
An analysis of California tidepool fishes. Amer.
Midland Naturalist 49-3: 862-871.
Prohaska, J. R. 1979. The effects of mid-intertidal tidepools on the
adjacent algal community. Unpublished manuscript on file at the
Hopkins Marine Station Library, Pacific Grove, California.
J. C. 1968a. Estimates of the populations and standing crop of
Quast
kelp bed fishes. In North, W. J. (Ed.) The biology of giant kelr
beds (Macrocystis) in California. 1971. J. Cramer Publisher,
Lehre. p.509-539.
J. C. 1968b. Fish fauna of the rocky inshore zone. Ibid., p.
Quast
481-507.
Quast, J. C. 1968c. Observations on the food of kelp-bed fishes. Ibid.,
p. 541-579.
Phil M. 1953. Common ocean fishes of the California coast.
Roedel
Calif. Dept. of Fish and Game Fish Bull. 91, 168 p.
22
FIGURE LEGENDS
Figure 1. Map of study area showing collection sites and modes of collection.
Figure 2. Percent of food items in guts found intertidally for each
species. Compiled both by species and number of organisms
percentages. Species considered intertidal are those listed
in Tables 2 and 3.
Figure 3.
Comparisons of diets between large (standard length299 mm, n-9)
and small (standard length84 mm, n-14) wooly sculpins, shown
in percent of stomachs in which present.
Figure 4. Intertidal non-cryptic fish densities per 20 x 20 meter area.
data from Table 4.
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PERCENT OF INTERTIDAL FOODS IN GUT
CONTENTS BY SPECIES AND NUMBERS
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SIANIT
JVGINVTISouOd
vankhovua
Jvalunova
Vidluvo
VaodiHaNv
Vaodosi
VIZVHOKTOd
HVInONnaIS
Sharpnose
Surfperch
Shiner
Surfperch
Juvenille
Rockfish
Surfperch
Black
Surfperch
Sennorita
Stripec
Surfperch
Calico
Surfperch
92
NUMBER OF NON-CRYPTIC FISH PER
TRANSECT (20x 20m)
O
27
TABLE LEGENDS
Table 1. Major food items in the diets of important intertidal fishes,
compiled as percent of stomachs in which present.
Table 2. Common intertidal animal food species in the guts of several
intertidal fishes. Vertical ranges from Light (1975) and
Keen and Coan (1974). Ranges represented by (high intertidal,
middle intertidal, low intertidal, subtidal), where X denotes
presence in a zone, O denotes absence from a zone. Compiled
as percent of stomachs in which present.
Table 3.
Common intertidal algae food species in the guts of several
intertidal fishes. Vertical ranges from Abbott and Hollenberg
(1976). Ranges represented by (high intertidal, middle
intertidal, low intertidal, subtidal), where X denotes presence
in a zone, O denotes absence from a zone. Compiled as percent
of stomachs in which present.
Results of transect fish count study, with mean numbers of
Table 4.
adults and juveniles present. Also percent of observation
periods present in the intertidal. Total of twelve observation
periods.
5.
saa.
-

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.
a
9
a
-
O
a
5


5

-
a
5

a
28
PERCENT OCCURENCE
Collisella
digitalis (X000)
Collisella
scabra (X000)
Pachygrapsus
crassipes (XX00)
Petrolisthes
cinctipes (XX00)
Lassea
cistula (XXXX)
Hemigrapsus
oregonesis (OXX0)
Margarites
salmonens (OXXo)
Pagurus
hemphilli (OXXO)
Pagurus
samuelis (OXX0)
Tegula
funebralis (OXXO
Barleeia
(OXXX,
haliotiphila
Collisella
limatula (OXXX)
Tricolia
pulloides (OXXX)
Mimulus
foliatus (O0X0)
Pachycheles
rudis (O0X0)
Pagurus
(ooxo)
granosimanus
Cancer
(OOXX)
antennarius
Pugettia
producta (OOXX)





.
PERCENT OCCURENCE
Endocladia
muricata (XX00)
Gigartina
agardhii (XX00)
Gigartina
papillata (XX00)
Hildenbrandia
occidentalis (XX00
Gigartina (OXX0)
canaliculata
Gigartina (OXX0)
leptorhynchos
Pterocladia (OXXO
caloglossoides
Rhodoglossum
affine (OXXO)
Microcladia
coulteri (OXXX)
Pterosiphonia
baileyi (OXXX)
Cryptopleura
corallinara (00X0
Laurencia
pacifica (O0X0)
Gastoclonium
coulteri (OOXX)
Gellidium
arborescens (OOXX
Iridaea
cordata (OOXX)
Neoptilota
hypnoides (OOXX)
Phyllospadix (O0XX
(vascular plant)





S
Table 3
SPECIES
Black Surfperch
Striped Surfperch
Pile Surfperch
Shiner Surfperch
Sharpnose
Sufperch
Calico Surfperch
Senorita
Cabezon
Kelp Greenling
Wooly Sculpin
Juvenile Rockfish
ADULTS
2.33
2.25
2.50
5.25
6.75
0.83
2.91
0.08
0.16
1.08
JUVENILES
0.67
0.25
0.83
0.42
0.08
5.00
Table 4
7 OF
DAYS IN
LNTERTIDA
1007
837
1007
502
677
427
757
427
177
422
337
REMARKS
Oftenseen feeding in
intertidal, alone or in
groups of 2-4 fish.
Forages in intertidal,
alone or in groups of
2-A other fish.
A shy surfperch, always
present in the intertida
When present, in large
schools.
Present in groups of 4
ranging to large schools
of 30 or more.
Occasionally a few
isolated individuals.
Sometimes in schools,
often just solitary
individuals.
Cryptic fish, undoubtedl
more abundant than the
figures suggest.
An occasional visitor to
the intertidal, found
ear the bottom.
Small, cryptic fish.
Actually, tremendously
ore common
Frequent schools of
unidentified young.
32
APPENDIX LEGEND
S: Sipuncula
P: Polychaeta
I: Isopoda
A: Amphipoda
C: Caridea
Pa: Paguridae
Po: Porcellanidae
B: Brachyura
Pol: Polyplacophora
G: Gastropoda
Bi: Bivalvia
E: Ectoprocta
F: Fish
P: Protozoa
A: Algae
Numbers represent number of individuals of that particular group,
an X refers to uncoutable organism remains or simply presence of a group.
The numbers given for the algae are the number of species.
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