Fishes of Marine Protected Areas Near La Jolla, California
Fishes of Marine Protected Areas Near La Jolla, California
P. A. Hastings,
1
* M. T. Craig,
2
B. E. Erisman,
3
J. R. Hyde,
4
and H. J. Walker
1
1
Marine Biology Research Division, Marine Vertebrate Collection & Center for
Marine Biodiversity and Conservation, Scripps Institution of Oceanography,
University of California San Diego, La Jolla, CA 92093-0208 U.S.A.
2
Department of Environmental and Ocean Sciences, University of San Diego,
San Diego, CA 92110 U.S.A.
3
University of Texas, The University of Texas Marine Science Institute, 750 Channel
View Drive, Port Aransas, TX 78373
4
Southwest Fisheries Science Center, NOAA/NMFS, 8901 La Jolla Shores Dr.,
La Jolla, CA 92037, USA
Abstract.—The marine waters surrounding La Jolla, California have a diverse array
of habitats and include several marine protected areas (MPAs). We compiled a list of
the fish species occurring in the vicinity based on records of specimens archived in
the Marine Vertebrate Collection (MVC) of the Scripps Institution of Oceanography
(SIO). Collection of fishes from La Jolla in the MVC started in 1905, but greatly
accelerated in 1944 when Carl L. Hubbs moved to SIO. By 1964, 90% of the 265
species recorded from the area had been collected and archived in the MVC. The
fishes of La Jolla are dominated by species whose center of distribution is north of
Point Conception (111 species), or between there and Punta Eugenia (96), with fewer
species with southern distributions (57), and one exotic species. Reflecting the
diversity of habitats in the area, soft-substrate species number 135, pelagic species 63,
canyon-dwelling species 123 (including 35 rockfish species of the genus Sebastes),
and hard-bottom species 140. We quantified the abundance of the latter group
between 2002 and 2005 by counting visible fishes in transects along the rocky
coastline of La Jolla, both within and adjacent to one of the region’s MPAs. In 500
transects, we counted over 90,000 fishes representing 51 species. The fish
communities inside and outside of the MPA were similar and, typical of southern
California kelp forests, numerically dominated by Blacksmith, Chromis punctipinnis
(Pomacentridae), and Sen˜orita, Oxyjulis californica (Labridae). Natural history
collections such as the MVC are important resources for conservation biology for
determining the faunal composition of MPAs and surrounding habitats, and
documenting both the disappearance and invasion of species.
The coastal environment in and around La Jolla, San Diego County, California is
notable for its complex and diverse array of habitats within a relatively small area. These
include kelp forests, rocky reefs, rocky intertidal, sandy beaches, sand and mud subtidal
areas, eelgrass and surf-grass stands, pier pilings, and submarine canyons, as well as the
pelagic environment. Because of the proximity of the La Jolla and Scripps submarine
canyons, depths range to over 500 meters within less than 7 km of the coastline. These
diverse habitats support a rich marine community, which has served as the focus of a
* Corresponding author: phastings@ucsd.edu
Bull. Southern California Acad. Sci.
113(3), 2014, pp. 200–231
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variety of scientific investigations (e.g., Limbaugh 1955; Quast 1968; Craig et al. 2004;
Brueggeman 2008).
The nearshore environment of La Jolla is especially important within the context of
marine conservation because it houses a network of marine protected areas (MPAs) of
varying size and age and with varying levels of use restrictions (Fig. 1). Historically, the
first of these was the San Diego Marine Life Refuge established in 1957, which included
the Scripps Coastal Reserve, a part of the University of California Natural Reserve
system (McArdle 1997). Directly to the south is the San Diego-La Jolla Ecological
Reserve (SDLJER), a small no-take reserve on the northern end of the La Jolla rocky
coastline that was established in 1971. The San Diego-La Jolla Ecological Reserve Areas
of Special Biological Significance was established in 1974 and largely overlapped the
SDLJER (McArdle 1997). Recently the SDLJER was included in the Matlahuayl State
Marine Reserve (CDFG 2013). In addition, the area on the southern part of the La Jolla
peninsula was recently designated as the South La Jolla State Marine Reserve, and an
area directly west of that was designated as the South La Jolla State Marine Conservation
Area (CDFG 2013). These MPAs have a variety of use restrictions, but all recognize and
seek to protect the biodiversity of this ecologically important region.
This study documents the fish fauna in and around this series of marine protected
areas. The primary source of information on fishes occurring in the La Jolla area was
the Scripps Institution of Oceanography’s Marine Vertebrate Collection (MVC). The
important roles of natural history collections such as the MVC to conservation biology
have been widely documented (Allman 1994; Pyke and Ehrlich 2010; Drew 2011). These
include compiling biotic inventories, documenting the loss or degradation of habitats and
associated biota, documenting changes in the distribution and occurrence of native
species, and documenting species invasions. In addition to compiling an inventory of fish
species collected in the area and archived in the MVC, we report on diver surveys of the
abundance and diversity of fishes in kelp forests, one of the most prominent habitats in
the area, in and around the SDLJER (now the Matlahuayl State Marine Reserve) from
2002 to 2005.
Brief History of Fish Collecting in the La Jolla Area
The marine fishes of California have been studied for many decades and are well-known
(Miller and Lea 1972, 1976; Hubbs et al. 1979; Love et al. 2005; Allen et al. 2006) with at
least 519 species known from state waters (Horn et al. 2006). In addition to early collections
of fishes from the San Diego area reported by Jordan and Gilbert (1880, 1881), the study of
fishes in the San Diego region of southern California was begun in earnest with the
Albatross surveys (Moring 1999) as reported by C.H. Gilbert (1890, 1896, 1915), as well as
inventories by Eigenmann and Eigenmann (1890) and Eigenmann (1892).
The establishment of the Scripps Institution of Oceanography (SIO) in the San Diego
region in 1903 and its subsequent move to La Jolla in 1905 marked a significant increase
in the study of the region’s biota (Hastings and Rosenblatt 2003). The on-site aquarium
displayed many of the common shallow-water fish species of the area. In 1918, Percy S.
Barnhart (Fig. 2A) was appointed as Collector and Curator of the Aquarium, and in
1926, Barnhart was elevated to the position of Curator of the Biological Collections, a
position he held until 1948. Barnhart studied the local fishes leading to a publication on
the fishes of southern California (Barnhart 1936), and he assembled a small collection of
preserved specimens from the region that ultimately formed the basis of the SIO Marine
Vertebrate Collection (MVC).
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Knowledge of the ichthyofauna of the region greatly increased after 1944 when Carl L.
Hubbs (Fig. 2B) moved to SIO and began actively archiving collections of fishes in the
MVC. In negotiations regarding his forthcoming move from the University of Michigan
to SIO, Hubbs wrote to then SIO Director Harald Sverdrup:
‘‘I would no doubt want to put considerable emphasis on systematic and variational
studies of west coast marine fishes, particularly those in which speciation would be
correlated with oceanographical (sic) conditions…I would no doubt be interested in
exploratory work, for instance with the fauna of the deep basins off the southern
California coast. I will probably be interested too in detailed analyses of the
distribution of fishes along the entire west coast, again as correlated with the
oceanographic conditions.’’ (Shor et al. 1987, pp. 226-227).
Before his arrival in October 1944, Hubbs convinced Sverdrup to invest in facilities to
store his anticipated collection of fishes, leading to the ultimate establishment of the
Fig. 1.
Map of study area with MPAs designated. Kelp forest fishes were surveyed at La Jolla Cove
and Boomers.
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MVC (Shor et al. 1987). Hubbs wasted little time in collecting fishes from the
surroundings of the SIO campus, amassing over 200 collections in his first year and over
500 by the end of the decade (Fig. 3).
Hubbs had another California project in mind when considering the move to La Jolla.
In 1944 he wrote to W. I. Follett in Oakland, with whom he had been corresponding for a
decade:
‘‘I look forward particularly to cooperating with you in making better known the
California fish fauna. I no doubt will have new material published from time to time
on the systematics and biology of the fishes but will definitely hope that you will
Fig. 2.
Photos of A) Percy S. Barnhart, 1935; B) Carl L. Hubbs, 1973; C) Richard H. Rosenblatt, 1979;
D) a view of La Jolla Shores and La Jolla peninsula looking southward from the SIO campus, circa 1910
(arrow indicates small embayment at La Jolla Shores). All images are from the Scripps Institution of
Oceanography Archives, UC San Diego Library.
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maintain your plan to work toward a ‘Fishes of California.’ It will be a pleasure to
make records and other information available for your project.’’ (Shor et al. 1987,
p. 227).
That project was published shortly after Hubb’s death in 1979 (Hubbs et al. 1979).
In 1958, Professor and Curator Richard H. Rosenblatt (Fig. 2C) was hired to oversee
the growing collection of fishes at SIO. He and other researchers and students at SIO
actively collected fishes in and around the La Jolla region. By the end of 1969, over 1,000
collections and well over 3,000 lots of fishes from La Jolla had been archived in the MVC
(Fig. 3). Local collecting of fishes declined after that time due to diverging research
interests, and constraints of space in the MVC for storage of specimens. Since that time,
the MVC has archived specimens of fishes from the La Jolla region primarily when new
or unusual specimens become available, specimens from focused efforts associated with
faunal inventories of the area (e.g., Craig et al., 2004; Craig and Pondella, 2006), and
voucher specimens for the growing MVC tissue collection, established by H.J. Walker in
1993 (Hastings and Burton, 2008).
Materials and Methods
We compiled a list of fishes recorded from La Jolla, California based on specimens
collected and archived in the MVC of the Scripps Institution of Oceanography,
University of California San Diego (Table 1). We included all species collected less than
10 km from shore (east of 117
u20.59 W longitude) and from Torrey Pines State Beach
southward to Tourmaline Beach (32
u549 N - 32u489 N latitude). This area includes the
entire rocky headland of La Jolla, as well as the primary conservation areas in the vicinity
(Fig. 1). These include the San Diego-Scripps Coastal State Marine Conservation Area,
the Matlahuayl State Marine Reserve, the South La Jolla State Marine Reserve and
Marine Conservation Area (CDFG 2013).
Fig. 3.
Number of collections of fishes made within the study area and archived in the SIO Marine
Vertebrate Collection by year (bars) and cumulative number of species recorded from those collections
within the study area (line).
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While some collections date to the early 1900s and a few were made in recent years, most
were made between 1944 and 1969 (Fig. 3). Archived specimens were collected using a wide
variety of sampling methods and were taken from the beach or ocean surface to depths of
over 500 meters. Species are listed in Table 1 arranged in taxonomic order. Common names
of families and species follow those recognized by the American Fisheries Society (Page et
al. 2013). Brief information on habitat or habitats occupied and special occurrences is
provided for each species. Habitat type was divided into three categories as follows: H 5
hard substrates, including rocky reef, rocky intertidal, kelp forest, sides of pier pilings,
boulders or any other type of hard substrate; S 5 soft bottom, including sand, mud, eelgrass
and surfgrass; P 5 pelagic, including species that always swim well above the bottom, as
well as those that periodically or regularly swim several meters above the bottom. Also
indicated in Table 1 are the species collected in the La Jolla and Scripps Canyons from .
30 meters depth (5 Cn), and the species observed in transects conducted in and adjacent to
the SDLJER (5 Tr) between 2002 and 2005 (see below).
Frequency of occurrence is based on the number of lots (occurrences) of each species that
have been archived in the MVC that were collected within the designated area, regardless of
the abundance of the species. Common (C) species are those represented by eleven or more
collections, uncommon (U) species were collected in the study area from three to ten times,
and rare (R) species were collected only once or twice in the La Jolla study area. In a few
cases, species that are known by us or reported by others to be more common in the study
area than indicated by collection records are indicated with an asterisk (*). The
biogeographic distribution of each species was designated based on the mid-point of the
entire known range of the species. Range endpoints are from Horn et al. (2006),
supplemented as needed based on published distribution records (e.g., Love et al. 2005).
Southern species (S) are those whose range midpoint is south of Punta Eugenia on the outer
coast of Baja California (27
u509 N), northern species (N) have range midpoints north of
Point Conception (34
u279 N), and central species (C) have range midpoints between these
well-established biogeographic barriers (Brusca and Wallerstein 1979; Horn et al. 2006).
Between 2002 and 2005, the abundance of fishes associated with kelp forests was visually
surveyed at La Jolla Cove (within the SDLJER, now called the Matlahuayl State Marine
Reserve) and an adjacent site (Boomers) a short distance beyond the reserve boundary
(Fig. 1). Both sites have moderate relief rocky reefs (1-3 m high) scattered throughout the
area and are dominated by red algal turf reefs and kelp forests (Parnell et al. 2005, 2006).
Survey protocols were modeled after established techniques for assessing abundance and
density of conspicuous fishes (McCormick and Choat 1987; Pondella et al. 2005). Randomly
selected, quantitative belt transects were swum by two SCUBA divers for a period of
5 minutes over a distance of 50 m. All fishes (excluding pelagic species) observed within a
two-meter window (i.e., one meter on either side of the diver and one meter above and
below) along the transect were identified to species where possible, and the number of
individuals was counted by each diver. If counts from the divers differed, the average was
recorded. Each transect accounted for 100 m
2
of bottom area surveyed. Three replicate
transects were conducted along rocky reef substrates at each of four depths (3 m, 6 m, 9 m,
and 12 m) for a total of twelve transects at each site per survey period. Surveys were
conducted every two months for a total of six periods per year between January 2002 and
December 2005. At certain periods throughout the study, persistent foul weather prevented
full surveys at each sample site, especially for the shallowest depth contours. However, at
least six transects were conducted at every site during each sample period throughout the
study except for July/August 2004 when continuous high surf precluded surveys at Boomers.
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Ta
ble
1.
List
of
fishes
co
llected
from
La
Jolla,
Calif
ornia,
and
deposited
in
the
SIO
Marine
V
ertebra
te
Colle
ction.
Date
5
yea
r
of
fir
st
MV
C
rec
ord
from
study
area.
Oc
5
freque
ncy
of
oc
currenc
e
in
MVC
coll
ections
from
st
udy
area:
C
5
commo
n,
11
or
mo
re
records;
U
5
uncom
mon,
3
to
10
rec
ords;
R
5
rare
,
1
to
2
records;
*
indicates
spe
cies
known
to
be
more
comm
on
in
the
st
udy
area
than
collec
tion
rec
ords
indica
te
(see
text)
.
Habitat
(Hab)
cate
gories:
H
5
har
d
bo
ttom
(reefs,
etc);
P
5
pelagic;
S
5
soft
bottom
.
‘‘x’’
unde
r
C
n
and
Tr
ind
icate,
respec
tively,
col
lection
rec
ords
for
La
Jo
lla
or
Scripps
Cany
ons
at
.
30
me
ters
de
pth,
and
occurre
nce
in
kelp
forest
trans
ects
in
and
around
the
Mat
lahuay
l
State
Marin
e
Res
erve
(form
erly
San
Diego
-La
Jo
lla
Ec
ological
Res
erve;
see
Table
3).
SE
5
southe
rn
latit
ude
en
dpoint
of
range
(po
sitive
and
negative
valu
es
are
nort
h
and
sou
th
latit
ude,
respec
tively
);
NE
5
northe
rn
latitud
e
endpoi
nt
of
range;
mid
5
middle
latit
ude
of
range.
Distribut
ion
catego
ries:
So
5
southern
,
range
mid
point
sou
th
of
Punta
Euge
nia;
Ce
5
ce
ntral,
ran
ge
midpoin
t
betw
een
Pun
ta
Eugenia
and
Poin
t
Conceptio
n;
No
5
northe
rn,
range
midpoin
t
nort
h
of
Poin
t
C
onceptio
n.
Scien
tific
name
Com
mon
name
Date
Oc
Hab
Cn
Tr
SE
NE
mid
Dist
Myxinif
ormes
M
yxinida
e
–
hagfishe
s
Ep
tatretus
mcconn
augh
eyi
W
isner
&
M
cMillian,
1990
Shor
thead
Hagf
ish
1954
U
S
x
-
23
34
28.5
Ce
Ep
tatretus
stoutii
(Lockington,
1878)
Pacific
Hagf
ish
1948
C
S
x
-
27
60
43.5
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