| Rockfish
Inside and out of the San Juan
Islands Marine Preserves:
Demographics of nearshore rocky reef fish
By Eric Eisenhardt,
School of Aquatic and Fishery Sciences, University of Washington
(from Puget Sound Water Quality
Action Team's
Puget Sound Notes
Number 46, January-2002)
Introduction
Most research on marine reserves to date has taken place in
tropical systems. Studies have shown temporal increases in abundance
and size of fish in marine reserves on tropical reefs (Alcala
and Russ 1990; Roberts and Polunin 1991), and export of adult
biomass outside reserves (Russ and Alcala 1997). A handful of
studies have tested the influence of marine reserves on nearshore
rocky reef fish assemblages in temperate regions (McCormick
and Choat 1987; Palsson 1998; Paddack and Estes 2000; Martel
and others 2000). However, these temperate studies have relied
on comparisons of "fished" and "reserve"
sites (1) with little or no replication of treatments or (2)
across limited time scales.
The San Juan Islands Marine Preserves are fishery reserves
created by the Washington Department of Fish and Wildlife in
1990. They restrict all forms of fishing except for salmon,
herring, and, in certain areas, crab (Murray 1998), and offer
an opportunity to study the effects of harvest restrictions
on rocky reef species. Anglers targeting salmon have little
interaction with rocky reef fish, with the exception of lingcod,
Ophiodon elongatus (K.
Koski, personal communication). There is little or no bycatch
associated with herring fishing. Crabs are fished with pots
and usually in soft-bottom habitats.
Bottomfish angler trips in northern Puget Sound peaked between
1980 to 1983, and by 1994 the annual number of trips was down
to early 1970s levels. A 10-fish bag limit for rockfish was
enacted in 1983, and reduced to five fish in 1994. The current
daily bag limit for rockfish is one fish. In 1992, the lingcod
season was reduced from seven months to six weeks and minimum/maximum
size limits were introduced. Directed commercial fisheries for
rockfish using jig and troll gears were prohibited in the San
Juans in 1984 (Palsson and others 1997). Trawling is allowed
outside reserves, but rarely occurs in San Juan Channel, and
has resulted in total annual rockfish landings of less than
100 pounds since 1994. No lingcod have been caught commercially
during the last few years (W. Palsson, personal communication).
Data presented here were collected from six sites between
July 27 to October 5, 2000. Historical data collected in San
Juan Channel from 1974 to 1976 by Moulton (1977) are presented
for comparison. The three San Juan Islands Marine Preserves
in San Juan Channel that contain nearshore rocky reefs were
selected as study sites. These were paired with non-reserve
sites to provide similar bathymetry, substrate complexity, algal
communities, and exposure to oceanographic processes within
each reserve/non-reserve pair.
The eight target species for which data were collected included
five rockfish: copper (Sebastes
caurinus), quillback (S.
maliger), black (S.
melanops), yellowtail (S.
flavidus) and Puget Sound (S.
emphaeus); lingcod (O.
elongatus), kelp greenling
(Hexagrammos decagrammus), and striped surfperch (Embiotoca
lateralis). These species are distributed over a gradient
of susceptibility and desirability to local angler effort. They
comprise the largest and most conspicuous members of the nearshore
rocky reef fish assemblage in San Juan Channel. Due to space
constraints, this paper will present data for three species:
lingcod, copper rockfish, and Puget Sound rockfish. See Eisenhardt
(2001) for additional information.
It should be noted that this study surveyed 0 to 20 meters
depths. The species studied are also known to inhabit deeper
waters. In addition, the San Juan Islands Marine Preserves encompass
depths greater than the depth range covered by this study. Therefore,
these results should not be extrapolated to all depths.
Methods and materials
Data were collected via 25 meters by two meters visual band
transects by two researchers using SCUBA. Data collected for
each transect included species, length (total length to nearest
centimeter), and depth for all target species sighted, as well
as habitat information, including substrate complexity, slope,
and percent cover of rock, algae, invertebrates, and sediment.
Forty-eight transects were completed at each site. See Eisenhardt
(2001) for detailed methods. Fish densities were computed as
fish per 100 square meters and statistics were computed using
SYSTAT 10.
Results
Length-frequency distributions were used to compare demographics
of fish populations inside and out of reserves in 2000 to the
1970s. Total numbers of fish in reserves and non-reserves provide
abundance estimates, as these data resulted from equal sized
areas surveyed. Area surveyed for historical data was different.
Therefore, 1970s abundance should not be compared to 2000 abundance
using these figures. For results of population densities over
time, see Eisenhardt (2001, in press).
The data are species specific. Statistical anyalyses show a
greater mean length (two-sample t-test: p < 0.001) and greater
density (3 x 2 ANOVA: F = 10.487, p = 0.001) for copper rockfish
in reserve sites compared to non-reserve sites. Large copper
rockfish (larger than 38 centimeters), which have been reported
to reach 35 years in age (Richards and Cass 1986), have virtually
disappeared since the 1970s. Greater mean length (two-sample
t test: p < 0.001) was found for lingcod in reserves compared
to non-reserves, but no statistically significant difference
in density of lingcod was found, due to the high variability
in lingcod densities – despite 35 percent greater mean density
of lingcod in reserves. In addition, 43 percent of lingcod sighted
in reserves were greater in length than 50 centimeters, compared
to only 17 percent in non-reserves. Length at 50 percent maturity
for male lingcod is 51.3 centimeters (Jagielo 1994). Puget Sound
rockfish showed an opposite trend with a greater mean length
(two-sample t: test p = 0.001) in reserves, but greater density
(3 x 2 ANOVA: F = 10.487, p = 0.005) in the non-reserve sites.
Habitat variables were similar between each reserve/non-reserve
pair. Further discussion of results for each of these species
and possible mechanisms follows in the discussion section.
Discussion
Copper rockfish are a commonly targeted bottomfish and are
often caught as bycatch by anglers targeting lingcod. The virtual
disappearance of copper rockfish larger than 38 centimeters
(a 45 centimeters individual was sighted during the 2000 survey)
is important to note, as fish this size were more common in
the 1970s (Moulton 1977). This trend indicates increased mortality
of larger, more fecund individuals since the 1970s, almost certainly
due to fishing pressure. This study shows that to regenerate
an abundance of copper rockfish individuals larger than 38 centimeters,
protection measures will need to be in place for at least 10
years. This is logical for a species that can live to be 35
years old. Given more time, copper rockfish larger than 38 centimeters
will probably become more abundant in reserves.
While there were reproductive adult copper rockfish in both
reserves and non-reserves, densities were greater in reserves – especially
for larger length classes. Given the non-linear increase of
copper rockfish fecundity as a function of length (DeLacy and
others 1964; Washington and others 1978), reserves contain greater
reproductive potential than non-reserves. The term "reproductive
potential" is used to describe the number of eggs produced per
area of habitat by a population (Paddack and Estes 2000).
During the spring of 2000, Sebastes
larvae identified as copper/quillback rockfish complex were
most abundant in the middle of San Juan Channel, and abundance
increased with increasing distance from shore (Chasco and others
2000). This trend indicates that planktonic larvae of the copper/quillback
rockfish complex in San Juan Channel are probably contained
in a common larval pool. Therefore, larvae are probably dispersing
from where they are released to this common pool, and then settling
out as juveniles throughout San Juan Channel. If this is correct
(and since fish in reserves seem to contribute a disproportionately
greater share of larvae to the common pool than the area of
rocky reef habitat encompassed by reserves indicates) reserves
appear to supply a disproportionately large share of copper
rockfish larvae to both reserve and non-reserve areas.
Lingcod are commonly targeted by anglers and are one of the
most highly prized and sought after bottomfish in San Juan Channel.
The low end of the legal size limit for lingcod is 26 inches
(or 66.04 centimeters), the same size at which greater fish
densities in reserves begin to appear. The distribution of lingcod
in reserves is similar to that found in San Juan Channel during
the mid-1970s, while the distribution in non-reserves is lacking
larger individuals. This is strong evidence that removals by
anglers fishing in non-reserve areas structure the demographic
pattern of lingcod in San Juan Channel.
Lingcod inhabiting nearshore rocky reefs seem to be predominantly
males, while females reside in deeper water most of the year
and enter shallower nearshore waters only briefly to spawn (T.
Jagielo, personal communication). Egg nests have been sighted
via SCUBA in the reserves; however, most or all of the broodstock
may not typically be within working SCUBA depths and are possibly
out of the reserve boundaries altogether (except during spawning).
Still, assuming larger males guard larger egg masses during
the nesting season (as seen by this author), larger males should
have a disproportionately large relation (on a per-fish basis)
to the reproductive potential of the population. In this study,
lingcod were significantly larger in reserves. Therefore, the
reserve areas could be contributing disproportionately more
to the reproductive potential of the population than the amount
of area contained in reserves would indicate.
Despite possible disproportionate contribution to lingcod
reproductive potential from reserves, both reserves and non-reserves
have similar densities of subadult recruits, that is densities
of individuals less than 50 centimeters were similar in reserves
compared to non-reserves.
Young-of-the-year lingcod have been found initially in soft-bottom
areas, for example bays and coves, and then disappearing from
this habitat and beginning to appear in nearshore rocky reef
habitat sometime before the end of their first year. Due to
the rocky nature of the San Juan Channel study area, shallow
soft bottom habitat is scarce. Therefore, it is likely that
lingcod found on different rocky reefs (including site pairs
in this study) initially recruited to the same soft-bottom areas.
If reserves are contributing disproportionately more reproductive
potential than a per-area basis would indicate, it would follow
that lingcod in reserves are augmenting recruitment outside
reserves.
Puget Sound rockfish are seldom caught by anglers because
they feed on plankton (Beckman 1995) and their mouths are too
small for a typical bottomfishing hook (B. Miller, personal
communication). They reach a maximum size of 18 centimeters
(Beckman 1995). Fishing is probably not structuring the demographics
of this species. The greater abundance of Puget Sound rockfish
in non-reserves versus reserves is probably due to increased
predation of Puget Sound rockfish inside reserves by the more
abundant and significantly larger lingcod inside reserves. Gut
contents of lingcod revealed Puget Sound rockfish, and lingcod
were often sighted in association with aggregations of Puget
Sound rockfish (W. Palsson, personal communication).
The reserve network seems to play an indirect yet major role
in regulating the density and mean size of Puget Sound rockfish
at the various sites, since reserves appear to regulate the
demographics of lingcod at different sites. Human influences
appear to be structuring ecosystems directly (in the case of
lingcod) and indirectly (in the case of Puget Sound rockfish).
There could be other, unknown, indirect impacts on other species,
for example killer whales (Orcinus
orca) and harbor seals (Phoca
vitulina richardsii).
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