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114(3) 105-163 (2015)

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FEB 1 7 70 18

December 2015

Southern California Academy of Sciences

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Bull. Southern California Acad. Sci.

114(3), 2015, pp. 105-122
© Southern California Academy of Sciences, 2015

The Physical Characteristics of Nearshore Rocky Reefs in
The Southern California Bight

Daniel J. Pondella, II,1* Jonathan Williams,1 Jeremy Claisse,1,2 Becky Schaffner3
Kerry Ritter3 and Ken Schiff 3

lVantuna Research Group, Occidental College, 1600 Campus Rd., Los Angeles, CA 90041
2 current address: Biological Sciences Department, California State Polytechnic
University, Pomona, CA 91786

3SCCWRP, 3535 Harbor Blvd., Suite 110, Costa Mesa, CA 92626

Abstract. — We present a GIS method for mapping and characterizing nearshore reef
habitats. Utilizing this technique, we were able to successfully map all nearshore
(<30 m depth) rocky reefs in the Southern California Bight and then quickly assess and
characterize these data layers with expert opinion. The southern California coastline is
1198 km in length, with the eight Channel Islands and mainland comprising 503 km and
695 km of coastline, respectively. This is approximately the same amount of coastline as
the rest of California. Within this region, we identified and characterized 122 natural
reefs comprising 49,055 hectares, which is 26.6% of the 184,439 ha of nearshore habitat
in the bight, the remainder comprised of soft bottom. Reefs varied appreciably in size
ranging from 6 - 2498 ha. We sampled a subset of these reefs using a generalized
random tessellation stratified design and quantified their physical characteristics as
measured by scuba surveys. The reefs also varied with respect to habitat type and five
distinct sub-habitat types varying from sheer oceanic pinnacle reefs to low-lying cobble
were observed. The distribution of reef types varied between the mainland and islands.
Island reefs were, in general, higher relief and had a greater percentage of rocky
substrate. Mainland reefs generally had lower relief and a higher percentage of sand and
cobble substrates.

The Southern California Bight (SCB) is a unique and increasingly critical stretch of the Cali-
fornia coastline. The physical constitution of the coastline along the mainland SCB is primarily
picturesque sandy beaches, broken up by rocky-headlands. In contrast, the remainder of the
state is dominated by iconic palisades associated with the coastal uplift from the shearing of
the right-lateral strike slip-transform San Andreas fault system (Zoback et al. 1987). Similar
and associated strike-slip faults and resulting uplift are the origins of the major coastal head-
lands in the SCB that are broken up by sandy beaches (Emery 1960). The San Andreas fault sys-
tem that runs along the coast in central and northern California moves inland proximate to Point
Conception the upper limit of the bight. The SCB is floored by a ~300 km wide region of exten-
sively faulted and extended continental cmst comprising Mesozoic metamorphic and intmsive
igneous rock as well as Neogene sedimentary and volcanic units (Crouch and Suppe 1993).
This region of submerged continental cmst is referred to in the geological literature as the Cali-
fornia Continental Borderlands (CCB). It differs markedly from the continental shelf north of
Point Conception, which is typically less than 100 km wide. The northern end of the CCB is
formed by the east-west oriented Transverse Ranges, a large fault-bounded cmstal block that

* Corresponding author: pondella@oxy.edu

105

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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

underwent 90° of clockwise rotation between 15 MYA and 5 MYA, the age of the SCB (Luyen-
dyk 1991). The unique east-west transverse ranges of southern California extend through the
CCB as the Northern Channel Islands (San Miguel, Santa Rosa, Santa Cruz and Anacapa)
and, as such, these islands are comprised of metamorphic and igneous rock (Atwater 1998). Dif-
ferential subsidence along the many faults that cut the CCB has produced the distinctive topog-
raphy of islands separated by ~ 1 km deep basins. Rotation of the transverse range block and
the submergence of extended continental crust in its wake created the SCB from a preexisting
coastline that had relatively straight, NW-SE trend and continuous with central and northern
California (Atwater 1998).

The Northern Channel Islands currently appear as extensions of the Santa Monica Mountains
and were all connected at the last glacial maximum (Graham et al. 2003). The orientation of
these islands is an indication of the torsion caused by the shear of the North American and Pa-
cific Plates forming the SCB during the Miocene (Atwater 1989). The subsequent uplift of
metamorphic rock from the Catalina Schist formed Catalina Island. Meanwhile, San Nicolas
Island is primarily an eroding anticline comprised of sandstone and shale marked by character-
istic marine terraces with some igneous rock (Kemnitzer 1933). In contrast, Santa Barbara
Island juts imposingly out of the sea with vertical cliffs up to 150 m in height and is comprised
of brittle igneous rock, which exhibits less pronounced marine terracing. Kemnitzer (1933) also
noted that a rock sample he received from Begg Rock, an exposed pinnacle reef 8 km off the
west end of San Nicolas Island was also volcanic. San Clemente Island lies on the San Clemente
fault, is formed of volcanic rock and has an anticlinal structure and prominent marine terracing
(Olmsted 1958). The origins of the various rocky reef habitats in the SCB are diverse and com-
plex, with considerable spatial variability.

It was previously estimated that the amount of nearshore reef habitat (< 30 m depth) was pro-
portional to the rocky intertidal habitat, approximately 15% of the mainland (Stephens et al.
2006). The southern California islands, however, support a greater proportion of coastal reefs
versus soft substrate in the nearshore environment (Ebeling 1980, Pondella and Allen 2000).
Due to accessibility and increasing stress by a growing population, these reefs are under a vari-
ety of anthropogenic stressors (e.g., turbidity, river plumes, sedimentation, overfishing and pol-
lution) and subject to harmful algal blooms (Stull et al. 1987, Homer et al. 1997, Dojiri et al.
2003, Schiff 2003, Love 2006, Pondella 2009, Foster and Schiel 2010, Sikich and James
2010, Erisman et al. 2011), which in many instances are not well understood and in all cases
necessitate a bight-wide perspective and coordination to contextualize and manage. These reefs
have been impacted by sewage, habitat loss, mnoff and climate change and, as such, can serve
as a model for dealing with these complex anthropogenic interactions (North 1964, Steneck
2002, Ford and Meux 2010). It has been demonstrated that large-scale management actions
can have significant positive effects on this complex ecosystem (Pondella and Allen 2008). In
2012, a network of Marine Protected Areas (MPAs) was created throughout the Bight (CDFG
2012). These MPAs were generally placed on rocky headlands, as this habitat is limited in
the region. This limited reef habitat was the most contentious issue during the implementation
process even though the amount of this reef habitat and the relative spatial distribution and char-
acterization remained unknown making current and future evaluations difficult (Pondella 2009).
Marine Protected Areas in California limit catch of extractable resources within their boundaries
(CDFG 2012). The establishment of these MPAs, while not specifically designated as fishery
management tools during implementation, was in part due to the decline of commercial and
recreational fisheries. Fisheries associated with rocky reefs in the region have been particularly
impacted. Examples include rockfishes (Love et al. 1998), abalone (CDFG 2005) and most
recently the kelp and sand basses (Erisman et al. 2011), and these serial depletions have caused

NEARSHORE REEF CHARACTERISTICS OF THE SOUTHERN CALIFORNIA BIGHT

107

significant socioeconomic damage. A critical task for advancing various research, restoration,
assessment and resource management programs is the quantification and characterization of
this nearshore habitat.

While general biogeographic patterns have been discerned for this ecosystem (Murray and
Littler 1981, Pondella et al. 2005), the gap in our knowledge of the quantity, structure and habi-
tat quality of shallow nearshore reefs in the SCB is surprising. These gaps in knowledge are
similar in other ecosystems where management actions need to be implemented and managers
are challenged by a paucity of quantitative data (Mumby and Harbome 1999, Pittman et al.
2011). Further complicating our understanding of this nearshore ecosystem is the necessity of
modeling processes on both small and large spatial scales (1 01— 1 05 m) (Garcia-Charton 2004)
as physical forcing and associated oceanographic conditions will be critical for contextualizing
reef performance into the future. Similarly, easily expressible metrics of ecosystem health are
needed for managers and non-scientific audiences. While the declines in fishery species are
well documented, the effects of pollution on rocky reefs in this area are not well understood.
Whether analyzing pollution, fishing practices, or ecological performance (including MPA
effectiveness), these processes are all couched within the extent, characteristics and variation
in the underlying habitat. Here we report on a novel method to determine the spatial scale of
reefs in the SCB. Then, we contextualize this system by describing the underlying substructure
and amount of nearshore rocky reefs in the region establishing a template for future research.

Materials and Methods

The methods in this study were composed of three sections. First, we assembled and mapped
all the available GIS layers for the region. The remote sensing techniques used in this study did
not characterize reef types. Thus, these hard bottom layers were then reviewed by experts in the
region to determine accuracy and characterize habitat types. Following this mapping exercise,
we conducted a stratified random draw to determine sites for a field-sampling program based
upon biogeographic region (which were based upon fish assemblages) insuring statistically
equal representation of reefs throughout the SCB. The field-sampling program then character-
ized a subset of reefs allowing inferences to the reef types of the SCB as a whole.

Mapping. — The best available compilations of mapped rocky reef habitat in the SCB were
assembled using GIS. These included maps of hard bottom habitats and kelp canopy (Kelner
2005). GIS spatial analysis techniques were used to integrate existing spatial data that charac-
terizes bottom type, kelp cover, and bathymetry to create a preliminary habitat map. Using these
data in GIS, we met with experts who have conducted multiple subtidal scuba research projects
on various geographic areas of the SCB. These working groups delineated reefs (< 30 m depth)
(Figure 1) and categorized each as either a major reef complex, patchy reef complex, cobble
reef, offshore or pinnacle reef, or manmade. Reef dimensions were made based upon the avail-
able GIS layers, while reef types were based upon expert opinion. The size of each reef was cal-
culated in GIS and categorized as large, medium or small based upon the distribution of reef
sizes. All other nearshore (< 30 m depth) substrate was classified as soft bottom. In better-
studied regions (e.g., Palos Verdes, Santa Catalina Island) investigators identified reefs on a
finer scale (continuous reef tracts were identified as multiple smaller reefs). Therefore, so as
not to bias the sampling draw by these regions for the survey portion of this study (see below),
reefs in these regions were grouped into larger reef areas. Similarly, to not deemphasize large
reef tracks, reef designations were adjusted to be as consistent as possible in size and distribu-
tion throughout the bight while mindful of natural habitat gaps. At Horseshoe Kelp in Los
Angeles County and Point Loma, the large reef areas were broken into two and three reefs,
respectively, for the sampling draw so as to not underestimate their impact.

108

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 1. Nearshore rocky reefs of the SCB. Reefs are color coded by biogeographic province (cold vs. warm)
and numbers correspond to the table used for the sampling draw (Appendix 1).

Station Draw. — Reefs were coded as island or mainland within each biogeographic province,
San Diegan (warm temperate) or Oregonian (cold temperate). Biogeographic province (Appen-
dix 1) was determined for the eight Channel Islands by biogeographic assessment of benthic fish
assemblages studied during the 2003-04 CRANE survey (Tenera 2006). In this biogeographic
analysis young-of-year (YOY) fishes whose density is seasonal, and highly abundant pelagic
species ( Engraulis mordax and Sardinops sagax ) present at only two sites, were excluded
from the data set. All statistics were run using PRIMER (version 6). The number of fishes
observed by station was Log (x+1) transformed. A Bray-Curtis similarity matrix was then cal-
culated and a hierarchical cluster analysis was performed. Using the similarity matrix, non-
metric multi-dimensional scaling was performed and using 45% similarity ellipses calculated
from the Bray-Curtis cluster the biogeographic regions were determined (Figure 1, Appendix
1). Mainland reefs were divided along previous described biogeographic assemblages utilizing
Santa Monica Bay as the faunal break between Oregonian and San Diegan faunas (Horn and
Allen 1978, Horn et al. 2006). Manmade reefs (i.e., breakwaters and jetties) were not included
in this mapping effort because they are well mapped and not part of the random station draw.
For the spatial scale aspect of this program, 60 natural rocky reefs (Figure 1; Appendix 1)
from this map were selected using a nested random draw (Stevens and Olsen 2004), a probabil-
ity-based design developed for monitoring aquatic resources, through EPA’s Environmental
Monitoring and Assessment Program (EMAP) (Stevens 1999). The advantage of the general-
ized random tessellation stratified design (GRTS) is that it allows for random sampling in a
way that provides good spatial coverage (without the clumping of sites often seen with simple

NEARSHORE REEF CHARACTERISTICS OF THE SOUTHERN CALIFORNIA BIGHT

109

random sampling). In addition, various strata or subpopulations can be defined and weighted
proportionally to a host of subpopulation characteristics (e.g., the size of the resource, the
size of the reef, variability of subpopulation estimates) so as to maximize efficiency when esti-
mating population totals or comparing among subpopulations. Two additional reefs (Escondido
and Big Rock) were not included in the random draw but were sampled as a part of this study to
fill in a gap in Santa Monica Bay.

Field Sampling. — teams of SCUBA divers that accessed sample sites from a research vessel
collected data visually. A single site consisted of at least 250 m of reef habitat. Within each site
four depth strata (if present) were sampled and geo-referenced. These strata were the inner (~5
m), middle (—10 m), outer (—15 m) and deep (—25 m) portions of a natural reef. Within each
depth stratum Uniform Point Contact (UPC) sampling protocol was completed. Therefore, the
maximum sampling effort for a site includes 8 UPC transects - 2 transects per each of the 4
depth strata. All transects were 30 m in length. Substrate type and relief were recorded at
each meter mark along the 30 m transect tape to estimate percent cover. Substrate type was
defined as: bedrock (>1 m), boulder (1 m-10 cm), cobble (<10 cm), or sand. Substrate relief
was defined as the maximum relief (0-0.1 m, 0.1-1 m, 1-2 m or >2 m) within a rectangle cen-
tered on the point that is 0.5 m along the tape and 1 m wide. The percentage of each type of
substrate category (bedrock, boulder, cobble or sand) was determined by pooling the number
of contact points for all replicates at each site by category, and dividing the sum of each category
by the total number of contact points at that site. Percentage of reef relief category (0-0. 1 m,
0.1-1 m, l-2m or >2m) was calculated in the same manner. Reef structure categories (% relief
and substrate) were square root transformed and normalized prior to being clustered using Eu-
clidean distances.

Results

In our calculations the southern California coastline is 1198 km in length. The islands com-
prise 503 km of coastline while the mainland coast has a length of 695 km. On the mainland,
rocky reefs are offshore (within 500 m) of 176 km (25.4%) of the coastline. At the islands, reefs
are offshore of 377 km (75.1%) of the coastline. For the islands the faunal break was in the mid-
dle of Santa Cruz Island, on the mainland it fell in the middle of Santa Monica Bay (Figure 1).
In the cold temperate region reefs span offshore of 290 km of the coast and in the warm tem-
perate region they span 263 km of coastline. We identified 122 natural reefs (< 30 m depth)
comprising 49,055 hectares in the SCB (Figure 1, Table 1). A greater fraction (60.8%) of reefs
were found in the cold temperate. This was in part due to the Santa Rosa and San Nicolas
Islands where the greatest expanse of reefs were identified (9088 and 5250, respectively).
A priori, eighty-nine reefs were classified as major reef complexes, seventeen as patchy reef
areas, two cobble reefs, and twelve pinnacle/offshore deep reefs (Appendix I). 10,164 ha of
the reefs identified in this study were previously described as soft bottom habitat. Demarcated
by the 30-m isobath, there are 184,439 ha of nearshore habitat in the bight, of which reefs com-
prised approximately a quarter (26.6%) while the remainder was sand.

Natural reefs (< 30 m depth) ranged in size from 6.2 (Begg Rock) to 2498 ha (Cojo) followed
by Talcott at Santa Rosa Island (2493 ha) (Appendix 1). The total for three Point Loma reef des-
ignations, which are continuous, is 2296 ha. Santa Rosa and San Nicolas Islands contained the
largest contiguous reef tracks and kelp beds in the SCB. The lee of Santa Rosa Island (Rodes,
Talcott and Carrington Point) comprised 5284 ha and the four reefs at west end of San Nicolas
Island comprised 4663 ha. On the mainland, Cojo Anchorage was the largest reef (2498 ha) fol-
lowed by three Point Loma reefs. The mean size of a natural reef was 409 hectares (sd + 497).
The distribution of reef areas was plotted and reefs were classified into three size classes. Sixty-

110

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 1 . The following metrics for the Southern California Bight are summarized below for the islands, main-
land, the cold temperate (Oregonian) and warm temperate (San Diegan) provinces: the length of the Southern Cali-
fornia coastline (Mexico to Point Conception); reef coastline length in km (reefs which are within 500 m of the
coast); and the area of natural reef habitat. The total amount of nearshore habitat in SCB is 184,439 and the
non-reef habitat is primarily sand.

SCB coastline length (km)

Mainland

Island

Total

694.5

502.7

1197.2

Reef coastline length (km)

Mainland

176.2

Cold

290.7

Island

377.4

Warm

262.9

Total:

553.6

Total:

553.6

Reef habitat (ha)

Cold

Warm

Total

Mainland

8213.8

10823.6

19037.4

Island

21587.4

8430.1

30017.4

Anacapa

545.1

545.1

Cortez Bank

1359.6

1359.6

San Clemente

3593.2

3593.2

San Miguel

3461.8

3461.8

San Nicolas

5249.9

5249.9

Santa Barbara Island

888.5

888.5

Santa Catalina

931.1

931.1

Santa Cruz

2365.4

2472.3

4837.7

Santa Rosa

9087.5

9087.5

Tanner Bank

63.2

63.2

Grand totals:

29801.2

19253.7

49054.9

seven reefs were classified in the small category (6-293 ha), with 40 as medium (325-932 ha)
and 13 as large (1086-2498 ha). Reef size categories had a mean of 95 ha (sd + 69) for small
reefs, 558 ha (sd + 183) for medium reefs and 1567 (sd + 484) ha for large reefs.

To begin to assess the range in physical habitat characteristics of the nearshore rocky reefs in
the SCB, we began with a physical characterization of the reef habitat including substrate type
and relief (Appendix II). Island reefs were primarily composed of bedrock or boulders (85.9%)
while mainland reefs had a more even mix of substrate types (Figure 2). Nearly half (47.8%) of
mainland reefs had a 0-0.1 m relief - more than double the fraction at the islands (23.3%). The
amount of 1-2 m and >2 m relief reef habitat at the islands were 2 and 6 times the fraction
found on the mainland, respectively. For relief, breakwaters were generally more similar to
island reefs. Reef structure, classified by relief and substrate through cluster analysis and over-
laid on a nMDS plot (Figure 3; Appendix II), varied from an oceanic pinnacle (Begg Rock) that
was a sheer vertical structure composed of bedrock and an intertidal component to mainland
reefs such as Carp Reef with large fractions of sand with little relief. Five reefs were not classi-
fied into a reef type (Figure 3) since they did not form distinct clusters in the cluster analysis (i.
e., had relatively high distance from the other reefs). Five reef types were found. Type 1
included a pinnacle reef (Begg Rock) and breakwaters comprised almost completely of bedrock
or large boulders. The second grouping (Type 2) was low relief and cobble reefs (Carp Reef and
La Jolla) that had significant fractions of sand. Type 3 reefs were predominantly island reefs
with some exceptions (Big Rock, Cabrillo Breakwater, Point Loma North, Point Vicente and
Little Corona). These reefs were almost completely composed of high relief (1-2 m) bedrock.

NEARSHORE REEF CHARACTERISTICS OF THE SOUTHERN CALIFORNIA BIGHT

Islands

Mainland Breakwaters

51.1%

■ Bedrock
Boulder
Cobble
Sand

■ >2m
l-2m

.Mm

O-.lm

Fig. 2. Substrate type and relief categories for island reefs, mainland reefs and breakwaters.

Alternatively, Type 4 reefs were predominantly mainland reefs with three island reefs (East
Quarry, SCAI, Lil Flower, SCLI, and Lion’s Head, SCAI). These reefs were comprised of bed-
rock and boulders with large fractions of lower relief (0-1 m) components. Type 5 reefs were
bedrock reefs that were primarily flat (0-0.1 m relief). Thus, reefs can be grouped into five
major reef categories: low relief and cobble (Type 2), flat reefs (Type 5), moderate relief
(Type 4), high relief (Type 3), and pinnacles (Type 1). Three of these reefs (Banana Rock,
Southeast Rock and Point Dume), found on the perimeter of the nMDS plot, were pinnacle reefs
(similar to Type 1) that jut abruptly up from a sandy substrate. These types of habitats can be
particularly difficult to sample with a 30 m tape, as portions of the transect may wind up on
the sand, obfuscating the results.

Discussion and Conclusions

While the 122 natural reefs that were identified in the SCB spanned three orders of magnitude
in size (6 to 2498 hectares), most were relatively large major reef complexes and they were dis-
tributed throughout the San Diegan (warm temperate) and Oregonian (cold temperate) biogeo-
graphic regions. Island reefs tended to be higher relief, primarily bedrock. In general Mainland
reefs were lower relief and had more variable substrate composition. Mainland reefs typically
were associated with littoral cells and longshore sediment transport and have larger fractions
of sand (Figure 2)(Inman and Frautschy 1966). We report that approximately a quarter of the
nearshore (<30 m) habitat of the bight is comprised of rocky reef habitat. This is a greater per-
centage than would be expected from just analyzing the GIS layers available in 2008 (Kelner
2005) or from an extrapolation based upon rocky intertidal habitat (Stephens et al. 2006).
This technique was successful at elucidating some generally unexpected patterns. The largest
reefs in the SCB and the western coast of North America were at Santa Rosa and San Nicolas
Islands. The kelp forest on west end of San Nicolas Island, while not the longest in terms of lin-
ear coastline, illustrated the utility of this study. The potential contribution of large reef islands

112

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Point Dume

o

2D Stress: 0.14

SCAI - Banana Rock

o

X KH02

Little Corona

^ SCAI - Iron Bound Cove
Al - West Isle V

V

Cabrillo Breakwater

SCRI - Scorpion SCAI - Little Harbor V SCRI - Pelican

V V V

SCAI - Ripper's Cove HK - Reference

A Reef aSCAI - East Quarry

o >

A Cabrillo Jetty

Water Habitat^ Ridges

Resort Point A SCLI ’ Lil^ower

POLA-Pier 400

POLA - #2 Angels Gate ■ f
■ KH03

Point Loma North

B|9 R^k SCRI -Valley

V

POLA - Angels Gate East

Reef Type

■ 1

X 2

V 3

A 4

• 5

O Unclassified

V

SCRI - Gull Isle

SNI^utch Harbor West SCRI - Yellow Banks
V SBL^util

SMI - Cuyler V SNI - Unnamed Reef

SCAI - Lion's Head
PointFerminAA3pa|ms

Whites PointA

A

Rocky Point

Barn Kelp

X

Nicholas Canyon v

X x

Escondido

X Little Dume

Naples

Leo Carillo Laguna

V ^R^Rodes SBI- Cat Canyon

SRI -Johnson's Lee North bK - Kodes Q

V Point Loma south

SCLI - China Point a Deep Hole

o »

V Lead Better Beach

SCLI - Pyramid Cove A

SRI - Jolla Vieja” SNI - Sand Spit
Cojo

• 0C

jBI - Arch SN| _ Dutch Harbor 0

Crystal Cove

o

HK - Southeast Rock

X Carp Reef

Fig. 3. Reef structure nMDS plot based on Euclidean distances using UPC substrate and relief measures. Reef
Type determined by cluster analysis. Colors refer to biogeographic provinces: blue = cold temperate islands,
orange = warm temperate islands, light blue = cold temperate mainland, red = warm temperate mainland.

habitats to the ecology of this region is important. This study also identified a substantial
amount of the previously described soft bottom habitat as hard bottom by experts and over flight
data of giant kelp canopy. Part of this difficulty is that side scan surveys are limited to the peri-
meter of kelp beds and the nearshore environment changes over time, but utilizing multiple data
layers increases the detection of reefs. More fine-grained reef mapping approaches have been
and continue to be developed since this program (e.g., Claisse et al. 2012, Parnell 2015). Incor-
porating more data layers in the future will increase the accuracy of this reef layer.

What is evident is that the nearshore rocky reefs in the SCB are highly variable in terms areal
extent and physical habitat structure. Based upon relief and substrate characteristics alone, there
are five major reef types in the SCB. Efforts need to be made to understand the influence of reef
habitat characteristics (substrate type, rugosity and relief) on the associated biota (e.g., Parnell
2015). Nearshore reefs in the SCB are typically comprised of igneous, metamorphic or mud-
stone rock (Emery 1960). These rock types may be the cause of additional habitat variation in
terms of the biota they support and the rates at which they erode. Further, the geological pro-
cesses that created the reefs in the Miocene are manifested in the composition and amount of
habitat. The geology of our islands and mainland, while quite variable, mirrors the composition
of the proximate reefs. Where volcanic processes (Santa Barbara Island, Begg Rock) and the
uplift of the Catalina Schist result in dramatic palisades, the resulting fringing nearshore reefs
are also sheer and tight to the shoreline. The Northern Channel Islands are essentially a relo-
cated mountain range and have proportionally large nearshore reefs. The eroding marine
benches observed on San Nicolas and San Clemente Islands produced kelp beds. As an exam-
ple, the entire offshore side of San Clemente Island is a continuous reef and the island is —34
km in length. The geological processes observed on these islands (eroding anticlines, marine
benches, sheer palisades, etc.) are mirrored in the nearshore subtidal habitat.

NEARSHORE REEF CHARACTERISTICS OF THE SOUTHERN CALIFORNIA BIGHT

113

While macroscale processes vary considerably, individual reefs are significantly diverse as
well. This habitat heterogeneity impacts the ecology of the region. In the SCB, rocky reef ver-
tical relief was correlated with increased fish density and production with high relief reef signif-
icantly outperforming low relief reefs (Ambrose and Swarbrick 1989, Anderson 1989, Pondella
et al. 2002, Pondella et al. 2006). Depth has also been shown to be a useful characteristic in
modeling reef habitats (Claudet et al. 2006, Claisse et al. 2012, Parnell 2015); we did not
include depth in our analyses here, but note that depth components may be a significant factor
in reef performance. For instance, Horseshoe Kelp (in Los Angeles County) was only distrib-
uted in the deepest strata while many others lacked a deep stratum and some did not have a
shallow stratum. A finer-scaled approach evaluating the influence of depth strata on reef perfor-
mance would be beneficial. The structure, amount and distribution of reefs in the SCB vary
appreciably and are important to consider in the potential performance of this system.

Approximately 122 natural rocky reefs/reef complexes comprise approximately one-quarter
(26%) of the subtidal habitat in the nearshore (<30 m depth) SCB. Prior to this study, estimates
of nearshore subtidal (<30 m) rocky reef habitat were inferred from the linear distribution of
intertidal rock and these estimates significantly underestimated the amount of shallow subtidal
reef habitat in the SCB. The mapping exercise undertaken in this region was the most exhaus-
tive to date and is the best estimate of reef area for the region. We were able to accomplish this
effort relatively quickly and inexpensively using previously collected data sets and expert inter-
views. Data from multiple sources including side-scan sonar, aerial overflights, satellite imag-
ery, subtidal visual surveys and professional judgments were combined to create our estimates
of habitat extent. As more spatial data sets become available, they should be integrated into
more fine-scaled reef maps.

Acknowledgements

We would like to thank Donna Schroeder of the Bureau of Ocean Energy Management,
David Kushner of the National Park Service and Bill Power of the Los Angeles Counties Sani-
tation Districts for their assistance in the mapping process. Scott Bogue from Occidental Col-
lege assisted in the review of the geological literature. This is a product of SCCWRPs Bight
’08 Rocky Reef Program and in addition was supported by the following: California State Uni-
versity, Long Beach; Channel Islands National Marine Sanctuary; Heal the Bay; Los Angeles
Regional Water Quality Control Board; Marine Science Institute, UCSB; Los Angeles Bay-
keeper; MBC Applied Environmental Sciences; Merkel and Associates, Inc.; Montrose Settle-
ments Restoration Program; National Marine Fisheries Service; Ocean Science Trust;
Partnership for the Interdisciplinary Study of Coastal Oceans; Port of Los Angeles; Reef Check
California; San Diego Coastkeeper; San Diego State University; Santa Monica Bay Restoration
Commission; Scripps Institution of Oceanography; Southern California Edison; United States
Geological Survey; US Navy.

Literature Cited

Ambrose R.F., and S.L. Swarbrick. 1989. Comparison of fish assemblages on artificial and natural reefs off the
coast of southern California. Bulletin of Marine Science, 44:718-733.

Anderson T.W., E.E. DeMartini and D.A. Roberts. 1989. The relationship between habitat structure, body size and
distribution of fishes at a temperate artificial reef. Bulletin of Marine Science, 44:681-697.

Atwater T.M. 1989. Plate tectonic history of the northeast Pacific and western North America. Pp. 21-79 in The
Geology of North America, Vol. N, The Eastern Pacific Ocean and Hawaii (Winterer, E.L., D.M. Hussong
and R.W. Decker, eds.). The Geological Society of America.

114

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Atwater T.M. 1998. Plate tectonic history of southern California with emphasis on the western transverse ranges
and northern channel islands. Pp. 1-8 in Contributions to the geology of the northern Channel Islands,
southern California (Weigand P.W. ed.). American Association of Petroleum Geologists, Pacific Section.

Bohannon R.G., and E. Geist. 1998. Upper crustal structure and neogene tectonic development of the California
continental borderland. Geological Society of America Bulletin, 110:779-800.

CDFG. 2005. Abalone recovery and management plan, final. California Department of Fish and Game. The
Resources Agency, Sacramento.

CDFG. 2012. Guide to the southern California marine protected areas: Point Conception to Califomia-Mexico
border.

Claisse J.T., D.J. Pondella, II, J.P. Williams, and J. Sadd. 2012. Using GIS mapping of the extent of nearshore rocky
reefs to estimate the abundance and reproductive output of important fishery species. PLoS ONE, 7:e30290.

Claudet J., D. Pelletier, J. Jouvenel, F. Bachet, and R. Galzin. 2006. Assessing the effects of marine protected area
(MPA) on a reef fish assemblage in a northwestern Mediterranean marine reserve: Identifiying community -
based indicators. Biological Conservation, 130:349-369.

Crouch J.K., and J. Suppe. 1993. Late cenozoic tectonic evolution of the Los Angeles basin and inner California
borderland: A model for core complex-like crustal extension. Geological Society of America Bulletin,
105:1415-1434.

Dojiri M., M. Yamaguchi, S.B. Weisberg, and H.J. Lee. 2003. Changing anthropogenic influence on the Santa
Monica Bay watershed. Marine Environmental Research, 56:1-14.

Ebeling A.W., R. J. Larson, and W. S. Alevizon. 1980. Habitat groups and island-mainland distribution of kelp-
bed fishes off Santa Barbara, California. Pp. 403— 431 in The California Islands, Proceedings of a Multidis-
ciplinary Symposium (D.M. Powers, ed.). Santa Barabara Museum of Natural History.

Emery K.O. 1960. The sea off southern California. John Wiley & Sons, Inc., New York. 365 pp.

Erisman B.E., L.G. Allen, J.T. Claisse, D.J. Pondella, E.F. Miller, J.H. Murray, and C. Walters. 2011. The illusion
of plenty: Hyperstability masks collapses in two recreational fisheries that target fish spawning aggrega-
tions. Canadian Journal of Fisheries and Aquatic Sciences, 68:1705-1716.

Ford T., and B. Meux. 2010. Giant kelp community restoration in Santa Monica Bay. Urban Coast, 2:43-46.

Foster M.S., and D.R. Schiel. 2010. Loss of predators and the collapse of southern California kelp forests (?): Alter-
natives, explanations and generalizations. Journal of Experimental Marine Biology and Ecology, 393:59-70.

Garcia-Charton J.A., A. Perez-Ruzafa, P. Sanchez-Jerez, J.T. Bayle-Sempere, O. Renones, and D. Moreno. 2004.
Multi-scale spatial heterogeneity, habitat structure, and the effect of marine reserves on western mediterra-
nean rocky reef fish assemblages. Marine Biology, 144:161-182.

Graham M.H., P.K. Dayton, and J.M. Erlandson. 2003. Ice ages and ecological tansitions on temperate coasts.
Trends in Ecology and Evolution, 18:33-40.

Hickey B.M. 1993. Physical oceanography. Pp. 19-70 in Ecology of the Southern California Bight (Reisch, D.J, J.
W. Anderson, and M.D. Dailey, eds.). University of California Press, Berkeley

Horn M.H., and L.G. Allen. 1978. A distributional analysis of California coastal marine fishes. Journal of Biogeo-
graphy, 5:23-42.

Horn M.H., L.G. Allen, and R.N. Lea. 2006. Biogeography. Pp. 3-25 in The Ecology of Marine Fishes: California
and Adjacent Waters (Allen, L.G., D.J. Pondella II, and M. Horn, eds.). University of California Press, Los
Angeles.

Homer R.A., D.L. Garrison, and F.G. Plumley. 1997. Harmful algal blooms and red tide problems on the U.S.
West coast. Limnology and Oceanography, 42:1076-1088.

Hubbs C.L. 1960. The marine vertebrates of the outer coast. Systematic Zoology, 9:134-147.

Inman D.L., and J.D. Frautschy. 1966. Littoral processes and the development of shorelines. Proceedings of the
Coastal Engineering Specialty Conference, ASCE, Santa Barabar California. Pp. 511-536.

Kelner J.J., R. Christensen, R. Clark, C. Caldow and M. Coyne. 2005. Chapter 2, in: A Biogeographic Assessment
of the Channel Islands National Marine Sanctuary (Clark, R., J. Christensen, C. Caldow, J. Allen, M. Mur-
ray and S. Macwilliams eds.). NOAA Technical Memorandum NOS NCCOS 21: 89-134.

Kemnitzer L.E. 1933. Geology of San Nicolas and Santa Barbara Islands southern California. Masters Thesis,
California Institute of Technology, Pasadena California.

Lekic V., S.W. French, and K.M. Fischer. 2011. Lithosperic thinning beneath rifted regions of southern California.
Science, 334:783-787.

Love M. 2006. Subsistence, commercial, and recreational fisheries. Pp. 567-594 in The Ecology of Marine
Fishes: California and Adjacent Waters (Allen, L.G., D.J. Pondella II, and M. Horn, eds.). University of
California Press, Los Angeles.

Love M.S., J.E. Caselle, and W. Van Buskirk. 1998. A severe decline in the commercial passenger fishing vessel
rockfish ( Sebastes spp.) catch in the southern California bight, 1980-1996. CalCOFI Rep, 39:180-195.

NEARSHORE REEF CHARACTERISTICS OF THE SOUTHERN CALIFORNIA BIGHT

115

Luyendyk B.R 1991. A model for neogene crustal rotations, transtension, and transpression in southern California.
Geological Society of America Bulletin, 103:1528-1536.

Mumby P.J., and A.R. Harbome. 1999. Development of a systematic classification scheme of marine habitats to
facilitate regional management and mapping of Caribbean coral reefs. Biological Conservation,
88:155-163.

Murray S.N., and M.M. Littler. 1981. Biogeographical analysis of intertidal macrophyte floras of southern Cali-
fornia. Journal of Biogeography, 8:339-351.

North W.J. 1964. Ecology of the rocky nearshore environment in southern California and possible influences of
discharged wastes. International Conference on Water Pollution Research, London, September, 1962, Per-
gamon Press, Oxford.

Olmsted F.H. 1958. Geologic reconnaissance of San Clemente Island California. United States Department of the
Interior. Geological Survey Bulletin 1071-B. United States Government Printing Office, Washington.

Parnell P.E. 2015. The effects of seascape pattern on algal patch structure, sea urchin barrens, and ecological pro-
cesses. Journal of Experimental Marine Biology and Ecology, 465:64-76.

Pittman S.J., R.T. Kneib, and C.A. Simenstad. 2011. Practicing coastal seascape ecology. Marine Ecology Pro-
gress Series, 427:187-190.

Pondella D., II. 2009. Science based regulation: California’s marine protected areas. Urban Coast, 1 :33-36.

Pondella D.J., and L.G. Allen. 2008. The decline and recovery of four predatory fishes from the southern Califor-
nia bight. Marine Biology, 154:307-313.

Pondella D.J., L.G. Allen, M.T. Craig, and B. Gintert. 2006. Evaluation of eelgrass mitigation and fishery
enhancement structures in San Diego Bay, California. Bulletin of Marine Science, 78:115-131.

Pondella D.J., B.E. Gintert, J.R. Cobb, and L.G. Allen. 2005. Biogeography of the nearshore rocky-reef fishes at
the southern and Baja California islands. Journal of Biogeography, 32:187-201.

Pondella D.J., II, and L.G. Allen. 2000. The nearshore fish assemblage of Santa Catalina Island. Pp. 394-400 in
The Proceedings of the Fifth California Islands Symposium. D.R. Browne, K.L. K.L. Mitchell, and H.W.
Chaney, eds.) Santa Barbara Museum of Natural History, Santa Barbara, California.

Pondella D.J., J.S. Stephens, and M.T. Craig. 2002. Fish production of a temperate artificial reef based on the den-
sity of embiotocids (teleostei: Perciformes). ICES Journal of Marine Science, 59:S88-S93.

Schiff K. 2003. Impacts of stormwater discharges on the nearshore benthic environment of Santa Monica Bay.
Marine Environmental Research, 56:225-243.

Sikich S., and K. James. 2010. Averting the scourge of the seas: Local and state efforts to prevent plastic marine
pollution. Urban Coast, 1:35-39.

Steneck R.S., M.H. Graham, B.J. Bourque, D. Corbett, J.M. Erlandson, J.A. Estes and M.J. Tegner. 2002. Kelp
forest ecosystems: biodiversity, stability, resilience and future. Environmental Conservation, 29:436—459.

Stephens J.S., Jr., R. Larson, and I. D. J. Pondella. 2006. Rocky reefs and kelp beds. Pp. 227-252 in The Ecology
of Marine Fishes: California and Adjacent Waters (Allen, L.G., D.J. Pondella II, and M. Horn, eds.). Uni-
versity of California Press, Los Angeles.

Stevens D.L.J., and A.R. Olsen. 2004. Spatially balanced sampling of natural resources. Journal of the American
Statistical Association, 99:262-278.

Stevens J., D. L. and A. R. Anthony. 1999. Spatially restricted surveys over time for aquatic resources. Journal of
Agricultural, Biological, and Environmental Statistics, 4:415-428.

Stull J.K., K.A. Dryden, and P.A. Gregory. 1987. A historical review of fisheries statistics and environmental and
societal influences off the palos verdes peninsula, California. CalCOFI Rep., 28:135-154.

Tenera Environmental. 2006. Compilation and analysis of CIAP nearshore survey data. California Department of
Fish and Game: 80 p.

Zoback M.D., M.L. Zoback, V.S. Mount, J. Suppe, J.P. Eaton, J.H. Healy, D. Oppenheimer, P. Reasenberg, L.
Jones, C.B. Raleigh, I.G. Wong, O. Scotti, and C. Wentworth. 1987. Evidence of the state of stress of
the San Andreas fault system. Science, 238:1105-1111.

Appendix I. Station numbers corresponding to Figure 1, sampled reefs, biogeographic region, reef category.

116

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Bull. Southern California Acad. Sci.

114(3), 2015, pp. 123-128
© Southern California Academy of Sciences, 2015

Comparison of the Marine Wood Borer Populations in Los
Angeles Harbor in 1950-1951 with the Populations in 2013-2014

Donald J. Reish,1’* Thomas V. Gerlinger,2 and Robert R. Ware2

1 Department of Biological Sciences, California State University, Long Beach,

California 90840

2Coastal Resources Management, Inc., Long Beach, California 90804

Abstract. — A 14-month study was conducted of the marine wood borers present on
suspended wooden blocks replaced monthly at nine stations in Los Angeles Harbor in
2013-2014, and compared to the results of a marine borer study conducted in 1950-
1951. Many environmental changes have occurred in the harbor over the past 63 years.
The harbor land mass of the outer harbor has been expanded towards the breakwater,
channels deepened, and the water quality improved as a result of pollution abatement.
Existing pilings removed, replaced or covered with two layers of polyethylene. The
isopod Limnoria tripunctata and the bivalve Lyrodus pedicellatus were the principal
species. The station located in the Consolidated Slip area of the inner harbor was the site
of 87 and 58 percent of the Limnoria and Lyrodus , respectively, counted during the
study. Neither of these species present at this station in 1950-1951. The dissolved
oxygen concentration at this station improved from a mean of 0.1 ppm in 1950-1951 to
a mean of 6.7 in 2013-2014 as a result of improved environmental conditions. Larval
settlement at Cerritos Channel numbered in the thousands in 1950-1951 but only 22
were counted during the 2013-2014 survey. This difference was attributed to the effect
of piling covering or removal. It is recommended that existing creosoted pilings be
covered or removed in the inner harbor area since this was the region of greatest
occurrence of wood borers in 2013-2014.

The purpose of this study is to determine if there have been any changes in the marine wood
boring animal population in Los Angeles Harbor since the study conducted in 1950-1951. The
principal marine wood boring animals in Los Angeles Harbor belong to the isopod genus Lim-
noria and to the bivalve ship worm family Teredinidae. Four species have been reported from
Los Angeles harbor: Limnoria tripunctata, L. quadripunctata, Lyrodus diegensis [ =Teredo die-
gensis ] and Bankia setacea (Menzies, et al., 1963). Two additional species of cmstaceans are
known to ingest wood but unknown to cause damage to wood pilings. The amphipod Chelura
terbrans, which lives within burrows made by Limnoria, and the copepod Tisbe gracilis, which
occurs on wood blocks, ingests wood under laboratory conditions. (Barnard, 1955; Barnard and
Reish, 1957).

The earlier study was initiated by Robert J. Menzies who proposed to study the occurrence
and biology of the wood boring isopod Limnoria in Los Angeles Harbor as the subject for
his doctoral dissertation at the University of Southern California. He contacted Carrol Wake-
man, chief testing engineer of the Port of Los Angeles, in 1949 for assistance in his proposal.
Wakeman was interested and offered the assistance of a boat. Menzies’ major professor, Dr. John
L. Mohr, suggested that the study be expanded to include the bivalve wood borers and fouling

* Corresponding author: Donald.Reish@csulb.edu

123

124

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

organisms. Mohr then established the Southern California Marine Wood Borer Council. The
members included Wakeman, Harold Schiller of Baxter Creosoted Co., and other graduate stu-
dents of Mohr. The group met in the home of Mohr to plan for the study. The one-year study, of
whom the author of this paper (DJR) participated, was initiated in March 1950 and terminated in
March 1951. Wood blocks were suspended at three depths for a 28-day period at 15 stations in
Los Angeles — Long Beach Harbors. The wood blocks were removed, fouling organisms
scrapped off and preserved, replaced with clean wood blocks and the wood borers identified
and counted. Additional blocks were suspended for four months in order for the pallets, the
identifying systematic character, of the bivalve Family Teredinidae, to mature. Publications
on the occurrence of wood borers and fouling organisms as result of this study included: Men-
zies (1951), Menzies, et al. 1963), Barnard (1950, 1955, 1958), Reish, (1954, 1971a). The
Council ceased to exist after the completion of the study. No survey of the occurrence of
wood borers in the Los Angeles — Long Beach Harbors has been conducted since 1950-1951.
Therefore, the objectives of this study were to compare the occurrence of wood borers in Los
Angeles Harbor and their relationship to the environmental conditions in 1950-1951 with those
in 2013-2014.

Los Angeles Harbor has undergone many changes in the ensuing 63 years since the previous
wood borer study and only those which influence wood borer activity are enumerated herein.
The Main Channel and West Basin have been dredged to accommodate larger ships. The harbor
land mass has been extended towards the breakwater by construction of Piers 300 and 400
(compare Figure la with Figure lb). Pollution abatement was initiated in the late 1950s and
extended into the late 1960s which resulted in an improvement of water quality (Reish,
1971b). Pilings in the harbor were primarily wood timbers which had been creosoted to prevent
the infestation of wood borers. However, the isopod Limnoria tripunctata is capable of penetrat-
ing creosoted wood pilings (Lee and Miller, 1980). The inner harbor area was characterized in
the 1950-1951 period with little or no dissolved oxygen (Menzies, et al., 1963). With the
improvement of water quality (Reish, 1971b) L. tripunctata and to a lesser extent Lyrodus
pedicellatus was able to infect pilings and other wood structures in the inner harbor. As a means
of combating the invasion of wood borers, the harbor department covered the wood pilings with
6 mm inner wrap of polyethylene and 20-30 mm of polyvinylchloride outer wrap which killed
any existing borers and fouling organisms by suffocation and excluded new infestations.

Materials and Methods

Station designations coincided, in so far as possible, with those in 1950-1951. The stations
are indicated in Figure la for the 1950-1951 survey, Figure lb for those sampled in 2013-
2014 and described in Table 1 . The study began on August 18, 2013 and terminated on October
1, 2014. Douglas fir wood was purchased from a building supply company as in 1950. The
wooden blocks were smooth and without knots. A 1.5 cm diameter hole was drilled in a
wood block measuring 3.5 x 15 cm. The block was attached to the rope and suspended at
mid-water depth for usually a 28-day period. A rope was weighted at the end and attached to
a harbor structure. The block was removed at the end of the time period and replaced with a
new block. A second block was attached to the rope at selected stations and removed in approxi-
mately four months to examine the pallets to distinguish between the two bivalve teredinid spe-
cies. The water temperature and dissolved oxygen concentration were measured electronically
each time with a YSI 556 multi-parameter meter at a depth of three feet from the surface (Table
1). The fouling organisms were removed from the block with a paint scraper and preserved in
formalin for later identification of polychaetes. Each wood block was examined on the day of
recovery and the wood borers were identified under a dissecting microscope. Limnoria

WOOD BORER POPULATIONS IN LOS ANGELES HARBOR

125

Fig. la. Map of Los Angeles — Long Beach Harbors showing the station locations in the Los Angeles Harbor
survey conducted in 1950-1951. lb. Map of Los Angeles Harbor showing the station locations in the 2013-2014
survey.

tripunctata is distinguished from the other species of the genus by the possession of three
bumps on the telson which are visible under a dissecting microscope.

Results

Water temperatures three feet below the surface ranged from a low of 13.2°C (55.8°F) at Sta-
tion K located near the harbor entrance in April 2014 to a high of 22.1°C (71.8°F) at Station C
located in the Consolidated Slip area in September 2014 (Table 1). The mean temperature in Los
Angeles Harbor during the 14-month study period was 17.7°C (63.8°F). The dissolved oxygen
concentration ranged from a low of 5.97 ppm at Station D in Cerritos Channel to a high of 10.6
ppm at Station A located in Slip 1 in the inner harbor in December 2013. The mean dissolved oxy-
gen concentration in Los Angeles Harbor for the 14-month study period was 7.6 ppm. Three spe-
cies of wood borers were present on the test blocks: the isopod Limnoria tripunctata , the bivalve
Lyrodus pedicellatus and the amphipod Chelura terebrans. The two colder water species L. quad-
ripunctata and Bankia setacea present in the 1950-1951 study did not occur in the 2013-1014
study. Limnoria was present at all stations but Station K located near the harbor entrance. Station
C located in Consolidated Slip accounted for over 87% of the Limnoria counted during the 14-
month period (Table 2). It was present throughout the survey period at Station C with peaks in
occurrence in April through July 2014 on both the monthly and quarterly exposed wood blocks.
There were only 19 Limnoria present at nearby Station B, the second most in occurrence on the

126

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 1. Environmental description, location of stations sampled in 2014-2014.

Station letter

Location

Depth* (in feet)

Water temperature °C
range; mean

Dissolved oxygen ppm
range; mean

A

Slip 1, Berth 160

11.3

15.3-21.2 (18.1)

6. 1-8.5 (7.5)

B

Harbor Basin 5

19.9

14.2-20.4 (17.4)

6. 7-8. 5 (7.5)

C

Consolidated Slip

39.2

14.9-22.1 (17.7)

6.0-7. 8 (6.7)

D

Cerritos Channel

32.4

15.2-20.9 (18.7)

6. 6-9.0 (7.6)

J

Fish Harbor

17.9

13.6-22.6 (18.2)

6. 7-9. 9 (7.6)

K

Shoaling Marker A

20

13.2-21.1 (16.8)

7.3-9. 1 (7.5)

L

Entrance W. Channel

31.6

11.3-21.6 (16.9)

6. 6-8. 6 (7.6)

N

Port Pilot station

20.3

13.4-21.2 (16.9)

7. 1-8.8 (7.9)

Q

Entrance W. Basin

47

15.2-21.8 (17.8)

6. 5-9. 9 (8.1)

* Water depth relative to mean low water in outer Long Beach Harbor.

wood blocks. Only 1 1 Limnoria were counted from the four outer stations J, K, L and N during
the 14-month study. Lyrodus pedicellatus was present at least once at all stations. Station C in the
Consolidated Slip area accounted for 58% with a peak in September through October 2014.
Nearby stations Q and B accounted for 19 and 13%, respectively. The bivalve occurred at all
four outer station with largest numbers (29) observed at Station L. The amphipod Chelura
terebrans was only found at Station C with one taken from monthly exposed wood block on
October 1 and two from the longterm exposure ending in August in 2014.

Discussion

The mean water temperature at the nine stations in Los Angeles Harbor was similar in both
studies with a median of 17.7°C (63.8°F) in 2013-2014 compared to 17.3°C (63.1°F) in
1950-1951. The lowest temperatures were measured in both studies at Station K; the highest
temperature, 26.1°C (79°F), occurred at Station Q in the earlier study which was probably the
result of shallow water at that time. The highest temperature of 22.6°C (72.7°F) was measured
at Station J in Fish Harbor in 2013-2014. The dissolved oxygen concentrations were different
with the range and mean of 5.97-10.6 (7.6) ppm in 2013-2014 compared to 0.0-7. 7 (2.9) ppm
in 1950-1951. The lack of dissolved oxygen occurred several times in the inner harbor and dur-
ing fish canning season at Station J in Fish Harbor in 1950-1951. Three species of wood borers
were present on the test blocks in 2013-2014: Limnoria tripunctata, Lyrodus pedicellatus, and

Table 2. Total number of wood borers present on wood blocks 2013-2014.

Limnoria Lyrodus

Station

letter

Monthly

Quarterly

Monthly

Quarterly

A

1

*

14

*

B

16

*

65

*

C

244 1

2641

266

D

3

1

15

7

J

2

*

6

*

K

0

*

3

*

L

5

1

25

9

N

5

22

1

8

Q

1

*

63

*

* Quarterly wood blocks not exposed at these stations.

1 One Chelura terebrans taken on October 2104 monthly block and two on the long term block August 2014.

WOOD BORER POPULATIONS IN LOS ANGELES HARBOR

127

Table 3. Comparison of the monthly total number of wood borers in 1950-1951 with 2013-2014.

Station

1950-1 95 C

2013-2014

Limnoria

Lyrodus

Limnoria

Lyrodus

A

2

1

1

14

B

rare2

rare2

16

65

C

0

0

244

26

D

10

28,800

3

15

J

present2

present2

2

6

K

43

rare4

0

3

L

14

18

5

25

N

328

18

5

1

Q

1

1

1

63

1 Numbers not always given in Menzies, et al., 1963.

2 L. tripunctata, L. quadripunctata, and teredinids present when dissolved oxygen was high.

3 All L. quadripunctata.

4 L. pedicellatus rare in warm periods; 13 Bankia setacea in cold periods.

Chelura terebrans. The two colder water species L. quadripunctata and Bankia setocea were
not present in 2013-2014 (Table 3). While the water temperatures were similar, the absence
of these two species may be attributed to the lack of floating wood which would bring the colder
water species from the north into the harbor by the prevailing California Current. Picking up
floating pieces of wood was a convenient source of Limnoria which were used in laboratory
experiments (Anderson and Reish, 1967). It was not until recent times that floating wood and
debris were picked up by harbor personnel. There were three notable differences in comparing
the wood borer data: (1) The near absence or lack of Limnoria and Lyrodus at the inner most
harbor Stations A, B, C and Q in 1950-1951 was attributed to little or no dissolved oxygen in
the water in 1950-1951, but the majority of these species occurred at these stations in 2013-
2014 because of the improvement of water quality (Table 3). (2) Thousands of newly settled lar-
vae of Lyrodus were noted at Station D in Cerritos Channel in 1950-1951 (Menzies, et al., 1963)
but only a few were present in 2013-2014. This species has a pelagic larval stage and is capable
of swimming some distance, but probably the majority came from local untreated or damaged
pilings in the earlier study. These pilings had since either been removed or wrapped with poly-
vinylchloride to prevent infestation. (3) Station N in 1950-1951 was the primary site of
Limnoria in 1950-1951 but not in 2013-2014 (Table 3) because all wood pilings had been
wrapped. Wood borer infestation on test wooden blocks was low in the outer harbor stations
and is probably now a minor problem to existing structures.

Conclusions

On the basis of the two studies conducted 63 years apart, the primary area of wood borer
activity is now located in the inner harbor area especially at Station C in the Consolidated
Slip and to a lesser extent in Basin 5. Three wood boring species were present in 2013-2014:
Limnoria tripunctata , Lyrodus pedicellatus and Chelura terebans. The colder water species
Limnoria quadripunctata and Bankia setacea were not taken in 2013-2014. The covering of
existing pilings with polyvinylchloride killed any existing populations by suffocation and pre-
vented new infestations. It is recommended that either the existing pilings in Cerritos Channel
be covered or removed. As a result, wood boring activity and potential damage will become
minimal since new harbor construction predominately uses concrete pilings and any creosoted
pilings that are added are wrapped.

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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Acknowledgements

This study was supported by a contract from the Port of Los Angeles through the Ocean
Studies Institute. We wish to thank Kathyn Curtis and Andrew Jirik of the Port of Los Angeles
for their interest in this study. Nicholas DeSilva (CRN) assisted in field collection. Special
thanks are due to Dr. Geraldine Knatz, former director of the Port of Los Angeles, for her inter-
est and encouragement.

Literature Cited

Anderson, J.W. and D.J. Reish, 1967. The effects of varying dissolved oxygen concentrations and temperature on
the wood-boring isopod genus Limnoria. Marine Biology, 1:56-59.

Barnard, J.L., 1950. The occurrence of Chelura terebrans Philippi in Los Angeles and San Francisco Harbors.
Bull. So. Calif. Acad. Sci. 49:90-97.

Barnard, J.L., 1955. The wood-boring habits of Chelura terebrans in Los Angeles Harbor. In Essays in the natural
sciences in honor of Captain Alan Hancock. Univ. So. Calif., Los Angeles, p. 87-98.

Barnard, J.L., 1958. Amphipod crustaceans as fouling organisms in the Los Angeles-Long Harbors, with reference
to the influence of seawater turbidity. Calif. Fish Game, 43:161-170.

Barnard, J.L., and D.J. Reish., 1957. First discovery of marine wood-boring copepods. Science, 125:236.

Johnson, M.W. and R.J. Menzies, 1956. The migratory habits of the marine gribble Limnoria tripunctata Menzies
in San Diego Harbor, California. Biol. Bull., 110:54-68.

Lee, W.L. and M.A. Miller, 1980. Isopoda and Tanaidacea: The isopods and allies. Pp. 536-558 in Intertidal Inver-
tebrates of California (Morris, R. H., D. P. Abbott and E. C. Harderlie, eds.). Stanford Univ. Press, Stan-
ford, CA. 690 pp.

Menzies, R.J., 1951. A new species of Limnoria (Crustacea: Isopoda) from Southern California. Bull. So. Calif.
Acad. Sci., 50:86-88.

Menzies, R.J., J. L. Mohr, and C. M. Wakeman, 1963. The seasonal settlement of wood borers in Los Angeles —
Long Beach Harbors. Wasmann J. Biol., 21:97-121.

Menzies, R.J., 1971a. Seasonal settlement of polychaetous annelids on test panels in Los Angeles-Long Beach
Harbors 1950-1951. J. Fish. Res. Bd. Canada, 28:1459-1467.

Menzies, R.J., 1971b. Effect of pollution abatement in Los Angeles Harbors. Marine Pollution Bull., 2:71-74.

Reish, D.J. 1954. Polychaetous annelids as associates and predators of the wood borer Limnoria. Wasmann
J. Biol., 12:223-226.

Bull. Southern California Acad. Sci.

114(3), 2015, pp. 129-140

© Southern California Academy of Sciences, 2015

Evidence for Negative Effects of Drought on Baetis sp. (Small
Minnow Mayfly) Abundance in a Southern California Stream

Elizabeth Montgomery,1* Rosi Dagit,1 Crystal Garcia,1 Jenna Krug,1 Krista Adamek,1
Sandra Albers,1 and Katherine Pease2

1 Resource Conservation District of the Santa Monica Mountains 30000 Mulholland Hwy,

Agoura Hills, CA 91301

2Heal the Bay, 1444 9th Street, Santa Monica, CA 90401

Abstract. — Benthic macro invertebrate (BMI) sampling was conducted at two sites in
Topanga Creek from 2003-2014. During this period, Southern California experienced
extreme drought conditions (US Drought Monitor 2014). Examining trends in species
composition over this period allows for a relatively long-term analysis of potential
effects of drought on BMI communities. The Southern California Coastal Index of
Biotic Integrity (SCC-IBI; Ode 2007) was applied to BMI samples from Topanga Creek
to measure the effects of drought on quantitative biotic integrity. The following trends
regarding the BMI community of Topanga Creek emerged during the course of this
study: 1) Wet year rainfall in Topanga Creek Watershed positively correlated to relative
and per sq. ft. springtime abundance of Baetis sp ., relative abundance of Simulium sp. up
to 78.7 cm (31") rain, and negatively correlated to relative abundance of Chironomidae n.
d., 2) percent algae cover in April and May positvely correlated to abundance per sq. ft.
Baetis sp. and Simulium sp ., and 3) multiple regression analysis revealed a negative
relationship between Chironomid n.d. and Baetis sp. abundance. BMI are an important
food source for endangered steelhead trout and other native aquatic and terrestrial
insectivorous species of special concern; significant changes to the BMI community could
have trophic reprecussions for these and other wildlife. Long-term monitoring is important
for tracking the influence of changes in climatic conditions on BMI community.

Introduction

Benthic macroinvertebrate (BMI) communities, made up of snails, worms, insect larvae and
nymphs, freshwater cmstaceans, and other bottom-dwelling organisms of a freshwater stream,
are a vital indicator of riparian ecosystem health (Ode et al. 2005). BMI sampling adds a biotic
element to standard water quality testing procedures and is an invaluable tool for ecologists,
resource management professionals, and anyone interested in investigating and maintaining
healthy rivers (Fetscher et al. 2009). As primary consumers and decomposers of allochthonous
and autochthonous detritus, diatoms, and macrophytes, benthic macroinvertebrates are the most
basic link between aquatic and riparian vegetation and the rest of the stream community (Merritt
et al. 2008, Covich et al. 1999). In Topanga Creek, BMI are an important food item for aquatic
reptiles and amphibians, Arroyo chub, and federally endangered southern California steelhead
trout. The adult imago of many aquatic insects are an important food source for birds, bats,
and other terrestrial insectivores, especially within riparian zones.

Some BMI taxa, such as Ephemeroptera (mayflies), may appear and disappear from the
benthos within a matter of weeks, while others, like some Odonata (dragon/damselflies),

* Corresponding author: montgomerylizzy@gmail.com

129

130

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

develop under water over the course of a year or more (Voshell 2002). Shifts in species compo-
sition may be the result of a current or past disturbance to stream integrity (Boulton et al. 1992).
Habitat preferences, water quality thresholds, and additional life-history traits have been
described for nearly 4,000 North American lotic macroinvertebrate species making it possible
to key in on the nature of ecological disturbance such as sedimentation, pollution, or increasing
temperatures (Vieira et al. 2006). Indices of Biotic Integrity (IBI) have been developed using
this information to assign numeric and descriptive scores of ecological health to freshwater sys-
tems, such as the Southern California Coastal Index of Biotic Integrity (SCC-IBI) applied here
(Fetscher et al. 2009, Ode 2007, Ode et al. 2005).

Drought can negatively affect stream function and aquatic biota by inducing low flows,
pool isolation or drying, increased water temperature, sedimentation, and/or reduced dis-
solved oxygen (Lake 2011). Freshwater habitat and refuges suitable or preferred for particular
BMI species can be severely reduced by prolonged drought (Lake 2003). Griswold et al.
(2008) found that BMI community composition was altered by drought periods, favoring
smaller-bodied species with shorter life cycles. Pool stagnation and fine sediment accumula-
tion associated with drought have been shown to reduce Baetis sp. abundance (Iversen et al.
1978, Kaller and Hartman 2004). Less-tolerant or rheophilic families such as Baetidae and
Simuliidae can face decline or expiration in prolonged drought conditions, while more toler-
ant species such as Chironomidae and Ceratopogonidae can become more abundant (Wright
and Symes 1999).

The goal of this study was to examine the response of benthic macroinvertebrates to drought
in Topanga Creek from 2003 to 2014. Four additional creeks were included to see if any regio-
nal patterns emerged within the Santa Monica Mountains (Fig.l). Topanga Creek, Arroyo
Sequit, Cold Creek, and Solstice Creek are considered regional reference streams, while a large
dam and wastewater treatment plant impact Malibu Creek.

Materials and Methods

Study Location

Topanga Creek, a small perennial coastal mountain stream, drains a 47-km2 watershed into
the Santa Monica Bay, Los Angeles County. The watershed is part of the California Floristic
Province biodiversity hotspot and provides vital habitat for federally endangered southern steel-
head trout ( Oncorhynchus mykiss), 22 species of reptiles and amphibians, and numerous other
plants and animals. Approximately 70% of the Topanga Creek watershed is protected parkland
managed by California Department of Parks and Recreation. The remaining 30% is developed
within the village of Topanga (pop. 8,289, US Census 2010). Both sampling sites are below the
village of Topanga, and share a narrow canyon with a two-lane highway that runs alongside the
creek in some areas.

Annual BMI sampling in Topanga Creek was carried out from 2003-2014. Drought intensity
in this period varied by season and year throughout the decade between periods of normal win-
ter rainfall and extreme drought conditions (US Drought Monitor 2014). The drought intensified
as winter rains in 2012 and 2013 were insufficient to alleviate dry conditions, and by 2014 it
was declared the worst drought on record since NO A A record-keeping began a century ago
(NOAA 2014). Rain data for Topanga, Arroyo Sequit, Malibu, and Solstice watersheds were
acquired from the Los Angeles County Department of Public Works. Algae percent cover
was recorded during monthly foot surveys throughout the sample reaches 3 200-3 700m and
4000-4500m.

EFFECTS OF DROUGHT ON BAETIS SP. MAYFLIES

131

Created 8/1 2/14

Fig. 1. Topanga Creek and Santa Monica Bay sampling sites 2000-2014. *Lower Topanga at 3200m, Upper
Topanga at 4500m.

Sample Collection

BMI samples were collected in spring (Apr 22-May 27) in 2003-2014 (except for 2008 and
2009) at two main stem locations: Upper Topanga (UT; 4500m) and Lower Topanga
(LT; 3200m). LT is located within the low gradient (<3%) reach of the creek and UT is located
above the Topanga Canyon Bridge (mile marker 2.02) in the higher gradient (3-6%) reach. Sam-
pling protocol in years 2003-2012 followed the California Stream Bioassessment Protocol
(CSBP; CDFG 2003). Standard 1 -ft. wide D-shape kick nets were deployed left, center, and
right of three consecutive riffle transects, for a composite sample of nine kicks. In 2013 and
2014, the SWAMP Bioassessment Procedure (Ode 2007) was employed. A 1-ft D-shape kick
net was used to collect samples every 15m along a 150m transect, alternating along the way
between 25%, 50% and 75% from right bank, for a composite sample of 1 1 kicks. Samples col-
lected from UT in 2004 and 2007 were not viable for processing and were not included in the
analysis. All Topanga Creek samples were preserved in s=90% ethanol or by freezing.

Additional samples were collected from Topanga Creek from five additional sites in May,
July, September, November, and December of 2013, and February, April, and June of 2014 fol-
lowing California Stream Bioassessment Method (CDFG 2003) to compare results between
sampling methods and examine phenology.

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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Regional stream survey samples were collected in June or July from 2003-2014 by Heal the
Bay (Santa Monica, CA) at four additional upper Santa Monica Bay creeks: Arroyo Sequit
(AS 19), Cold Creek (CC2, CC3, CC11), Malibu Creek (MCI, MC15), and Solstice Creek
(SC 14). In 2003 BMI samples were collected using the California Stream Bioassessment Proto-
col (CDFG 2003); from 2005-2007, sampling was conducted using the US EPA targeted riffle
composite (TRC) procedure (Peck et al. 2004; Ode 2007), and starting in 2008, sampling was
conducted according to SWAMP Bioassessment Procedure (Ode 2007).

Sample Processing

Using a dissecting microscope, organisms were picked from the sample and then sorted and
identified according to standard taxonomic effort or to the lowest practical taxon based on speci-
men condition and size. Identifications and regional distribution were confirmed using the Cali-
fornia DFW Aquatic Bioassessment Laboratory (ABL) Digital Reference Library (CDFW 2014)
and Merritt et al. (2008). When identification was not possible, photographs were sent to the
CDFW Aquatic Bioassessment Laboratory or identified to the lowest taxonomic level possible
and recorded as non-distinct within that taxon. Heal the Bay regional comparison samples were
processed by SLSII bioassessment lab.

BMI Analysis

The relative abundance (%) and per sq. ft. abundance of each taxa present was calculated per
sample. Processed samples were assessed according to SCC-IBI (Ode et al. 2005). For this
index, seven metrics are used to assess ecosystem health: combined taxa richness of Ephemer-
optera, Plecoptera, and Trichoptera taxa (EPT), Coleoptera taxa richness, predator taxa richness,
% non-insect taxa, % tolerant taxa (TV>7), % intolerant individuals (TV<3), and % collector-
gatherer + collector-filterer (CG+CF) individuals. Information regarding tolerance values and
functional feeding groups was obtained on CAMLnet (Ode 2003). As SCC-IBI is designed
for samples of 500 individuals, samples >500 were subsampled by randomizing sets, generat-
ing random numbers and selecting individuals 1-500.

Bray-Curtis Index of Dissimilarity was applied to whole samples, not sub-samples of 500, as
it accounts for both sample size and shape (Bray and Curtis 1957).

c ElY - x,-t|

E(xi/ - x,*)

Bray-Curtis calculations were based on the abundance per sq. ft. of eight taxa categories: Baetis sp., Chir-
onomidae n.d., Simulium sp., Ostracod, Elmidae, Acari, Planarian, and Hy dropsy chidae.

Multiple regression analysis was performed applying a generalized linear model (Poisson
regression) run in R software with Baetis sp. abundance per sq. ft. as the dependent variable
and as a covariate. Other co variate factors input were wet year rainfall, percent algae cover,
and Chironomidae n.d. abundance per sq. ft.. Simulium sp. was not included because Simulium
sp. abundance and algae were highly correlated.

Results

BMI Community Composition

A total of 15,303 macro invertebrates were collected from UT and LT between 2003 and 2014.
The number of individuals per sample ranged from 104 to 3,514. Six phyla, 21 orders, and a
total of 76 taxa were represented. The majority of individuals (Relative Abundance ‘RA’ =
89%) fell within the class Insecta. Three genera and one family from class Insecta accounted

EFFECTS OF DROUGHT ON BAETIS SP. MAYFLIES

133

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Fig. 2. Relative Abundance of 6 Major Taxon Categories: Upper and Lower Reaches Topanga Creek 2003-2014.

for 76% of total abundance; Baetis sp. (small minnow mayflies nymphs, 44% RA), Chironomi-
dae n.d. (non-biting midge fly larva, 23% RA), and Simulium sp. (black fly larva, 9% RA;
Fig. 2). Other families present throughout the sample period included: Planarian n.d. (flat-
worms, 3% RA), Elmidae n.d. (riffle beetle larva, 3% RA), Acari (water mites, 2% RA), Hydro-
psychidae n.d., (net spinning caddisfly larva, 2% RA) and Ostracoda n.d. (seed shrimp,
2% RA).

In samples from 2003-2012, Baetis sp. comprised 51% RA (7.6<n<214.2 per sq. ft.) and
was the first or second most abundant taxa in all samples. In 2013 and 2014, Baetis sp. declined
to 3% RA (abundance 0.8<n<1.3 per sq. ft.) of all four samples. Simulium sp. abundance
decreased from 11% RA 2003-2012, to less than 1% in 2013-2014. Between 2003-2012 and
2013-2014, Chironomidae RA increased from 15% to 66% RA. In samples collected on five
occasions from five additional sites in Topanga Creek between May 2013 and June 2014, Baetis
sp. and Simulium sp. each comprised less than 2% of all samples, and never more than 8% of
any sample, confirming low abundance across sampling protocols.

Bray-Curtis dissimilarity coefficients for abundance per sq. ft. of eight dominant taxa calcu-
lated between all years ranged from 0.06 (LT 2010 to 201 1, highly similar) to 0.95 (three pairs,
highly dissimilar). Dissimilarity coefficients for 2013 and 2014 were each significantly higher
than 2004, 2006, 2007, 2010, 2011, and 2012, t{ 14,26) = 2.6<4.3,/? < .05 (two-tailed), upper
and lower scores combined. The year 2005 had above average rainfall (61.2 inches) and also
a high species composition dissimilarity coefficient. Wet year rainfall explained variation
in Bray-Curtis dissimilarity coefficients, which were significantly higher in years with
lower rainfall where above average rainfall years 2005 and 2011 were excluded (R“=0.85,
F i i3=30.5, p <0.01).

Functional feeding group composition differed less than taxa composition, and was domi-
nated by collector-gatherers in all years 2003-2014 (range: 63% to 95% RA; Fig. 3). Collec-
tor-filterers were the second most abundant feeding group overall, and they declined
significantly from 2003-2012 to 2013-2014, ^( 16) =2.9, p>.05 (two-tailed).

134

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

BCG ■ FC SC P &SH ■ Non-Distinct

Fig. 3. Relative Abundance of 6 Major Feeding Group Categories: Upper and Lower Reaches Topanga Creek
2003-2014.

Rainfall Correlations

Baetis sp. relative abundance and abundance per sq. ft. significantly and positively correlated
to wet year rainfall (WYR; R2 = 0.41, F1>16= 11.36, p<0.01, R2 = 0.80, F1j16=11.36,
p <0.00000 1; Fig. 4a.). Correlation to relative abundance was stronger when above average rain-
fall in 2005 (61.22”) was removed; however Baetis sp. abundance measured per sq. ft. was high-
est this year. Simulium sp. RA also positively correlated to wet year rainfall (WYR), only with
2005 data removed (R2 = 0.25, F114=4.79 ,p<0.05. Chironomidae n.d. RA negatively corre-
lated to WYR with no upper limit to rainfall (R2=0.26, F1>16=5.49, p<0.05; Fig. 5). Neither
Chironomidae n.d. nor Simulium sp. abundance per sq. ft. correlated to WYR, nor did

Fig. 4. (a) Relative abundance of Baetis sp. and wet year rainfall in Topanga Creek (b) Abundance per sq ft.

of Baetis sp. and observed algae in Topanga Creek. Black triangles depict Lower Topanga samples and gray
circles depict Upper Topanga

EFFECTS OF DROUGHT ON BAETIS SP. MAYFLIES

135

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of Chironomidae n.d. and wet year rainfall in Topanga Creek. Black triangles depict Lower Topanga samples and
gray circles depict Upper Topanga.

any FFG group. In regional samples, only Solstice Creek also showed a significant and
positive correlation between Baetis sp. RA and WYR (R2=0.60, F15 = 7.38, p<0.05 ;
Fig. 6).

Abundance per sq. ft. for eight dominant taxa was compared to observed algae cover in both
reaches from 2007-2014. Baetis sp. and Simulium sp. were found to positively and significantly
correlated to percent algae cover (R2 = 0.77, F1>9=29.9, p<0.0005, R2=0.69, F1?9=19.6,
p <0.005; Fig. 4b.). Multiple regression analysis was run to examine the relationship between
Baetis sp. abundance per sq. ft. and WYR, algae, and Chironomidae n.d.. Rainfall and algae
remained significant predictors of Baetis sp. (p<.0005, p<.05). When controlling for rain
and algae, Chironomid n.d. abundance per sq. ft. was also a significant predictor of Baetis sp.
abundance (p<.0005).

SCC-IBI Analysis

The Southern California Coastal Index of Biotic Integrity (SCC-IBI; Ode et al. 2005) was
applied to determine if the marked change in taxonomic composition resulted in decreased
scores of biotic integrity in 2013-2014. Three out of ten samples from Lower Topanga and
five out of eight samples from Upper Topanga had >500 individuals and were sub-sampled
to 500 individuals appropriate for SCC-IBI scoring. LT04 (n=464) was also included in
SCC-IBI analysis. Overall, there was no significant correlation between total SCC-IBI scores
and wet year rainfall or other creek conditions (Table 1). SCC-IBI scores ranged from 22-57,
categorized as ‘poor’ to ‘fair.’ The lowest metric score across all samples was % intolerant indi-
viduals, which never surpassed a score of 1 . Conversely, the highest metric on average was %
tolerant taxa (average score of 6).

Discussion

Drought is a defining feature of many ecosystems including those within Mediterranean cli-
mates, and it is predicted that droughts will become more frequent and intense over the next few
decades (Houghten et al. 1995). Severe drought is known to affect BMI assemblages and result
in alterations to community composition (Resh et al. 2012). The current drought in Southern
California is the most severe on record (NOAA 2014). While these conditions cause concern
for the integrity of freshwater ecosystems, they also provide an opportunity to study potential
implications of climatic patterns on freshwater benthic macroinvertebrate communities. In

136

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

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Fig. 6. Relative abundance of Baetis sp. compared to wet year rainfall in Solstice Creek.

2013, the Topanga Creek BMI community experienced a sharp decline in Baetis sp. and Simu-
lium sp. abundance, and increase in Chironomidae n.d.. Each of these taxa abundance correlated
to WYR, suggesting a strong relationship between annual precipitation and BMI community
composition in Topanga Creek. This correlation also held true for another Santa Monica Moun-
tains stream, Solstice Creek.

For Simulium sp. in Topanga Creek, the correlation to rainfall did not hold up in the case of
above average rainfall (155.5 cm), and in fact all taxa appeared to diverge from the linear trend
under these high rainfall conditions. This might be due to discharge rates high enough to dis-
place certain BMI taxa. Both Baetis sp. and Simulium sp. also positively correlated to algae
cover in April and May. When available, algae can be a preferred food source for Ephemerop-
tera, resulting in more robust growth and gill size (Mayer and Likens 1987, Gupta et al. 1994).

Chironomidae n.d. relative abundance was favored in low rainfall conditions. Multiple
regression analysis showed that Baetis sp. and Chironomidae n.d. abundance per sq. ft. was sig-
nificantly related when rainfall and algae were held constant, however this was not evident in a
direct comparison. This indicates that there is a distinct relationship between Baetis sp. and
Chironomidae n.d. that is mediated by abiotic and biotic factors such as precipitation and algal
productivity. This relationship might hinge on competition for food or habitat, or sedimentation
dynamics. Angradi et al. (1999) found that Ephemeroptera taxa richness is reduced when fine
sediments accumulate, while some genus of Chironomidae, particularly burrowers or sediment
case-makers, increase. Frost et al. (1995) affirms that changes in the abundance of one species
can lead to disproportionate response from other species. Krug et al. (2012) found that both Dip-
tera and Ephemeroptera larva are important food sources for endangered southern steelhead
trout ( Oncorhynchus mykiss ), occurring in 69% and 92% of 13 stomach samples collected in
Topanga Creek in March 2011. Continued monitoring is recommended to measure the resi-
liency of Baetis sp. and Simulium sp. under future rain conditions.

Both Baetis sp. and Chironomidae n.d. are classified as collector-gatherers with a tolerance
value of 6 (Ode 2003). Despite changes in community demographics, trophic structure can

EFFECTS OF DROUGHT ON BAETIS SP. MAYFLIES

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remain stable (Beche et al. 2009; Vannuchi et al. 2013). This occurred to some extent in
Topanga Creek, as functional feeding group distribution was less variable than species compo-
sition over time. Collector-gatherers made up the far majority, which is typical in riparian lower
reaches (Hawkins et al 1981). The shift in community was accompanied by a simplification of
FFG array. The decline of collector- filterer abundance correlated to Simulium sp. declines, indi-
cating Simulium sp. was a significant collector-filterer in both the Upper and Lower Reach of
Topanga Creek.

Although Topanga Creek is facing impacts of drought, it remains an important reference
stream and continues to provide crucial habitat to wildlife. Catastrophic shifts in BMI commu-
nity compositions have been observed to occur from stable to alternate states, sometimes irre-
versibly (Scheffer et al. 2001; Beche et al. 2009). In other cases, communities have
rebounded within 1-3 years post-drought (Boulton 2003). How the aquatic and riparian zone
insectivorous wildlife of Topanga Creek respond to the replacement of Baetis sp. with Chirono-
mid n.d. and other changes among the BMI community are currently unreported. Continued
long-term monitoring of BMI will support regional advances in understanding species compo-
sition trends and how they are influenced by climatic disturbances such as drought.

Conclusions

Long-term BMI monitoring is important in Topanga Creek, and other drought-affected fresh-
water ecosystems, to determine the influence of climatic events on taxonomic composition. In
this study, drought intensification aligned with declines in Baetis sp. and Simulium sp. relative
abundance. Algae presence also influenced the abundance of these two taxa, although there was
no notable trend between rainfall and algae found in this study. Long-term data sets will enable
scientists to track what components of BMI composition are constantly in flux, responding to
biotic and abiotic processes and events, and which are perennially resilient up to the point at
which catastrophic events may induce long-term disturbance within the established community.
Future work in this watershed should focus on the potential recovery response of Baetis sp.
post-drought.

Acknowledgements

This study was made possible by funding from the office of LA County Supervisor Zev Yar-
oslavsky, dedicated work by the Resource Conservation District of the Santa Monica Mountains
Stream Team, AmeriCorps Watershed Stewards Project members Kayti Christianson, Carrie
Fong, David Gottesman, Diana LaRiva, Tessa Reeder, and volunteers Uriel Cobain, Allie Irwin,
Monica Jarquin, Ariane Jong, Matt Kirby, Gabby Njm, Robert Ruzicka, Vanessa Thulsiraj,
Karen Vu, Sylvia Zamudo, Amy Zimmer-Faust, Mark Ziman. Dan Pickard, CDFW and Ethan
Bell, Stillwater Sciences provided additional help. Finally, thanks to CA Department of Parks
and Recreation for their efforts to preserve and protect Topanga Creek.

Literature Cited

Angradi, T.R. 1999. Fine sediment and macroinvertebrate assemblages in Appalachian streams: a field experiment
with biomonitoring applications. Journal of North American Benthological Society, 16(1 ):49— 66.

Beche, L.A., RG. Connors, V.H. Resh, and A.M. Merenlender. 2009. Resilience of fishes and invertebrates to pro-
longed drought in two California streams. Ecography, 32:778-788.

Boutlon, A.J., C.G. Peterson, N.B. Grimm, and S.G. Fisher. 1992. Stability of an Aquatic Macroinvertebrate Com-
munity in a Multiyear Hydrologic Disturbance Regime. Ecology, 73:2192-2207.

Boulton, A.J. 2003. Parallels and contrasts in the effects of drought on stream macroinvertebrate assemblages.
Freshwater Biology, 48:1173-1185.

EFFECTS OF DROUGHT ON BAETIS SP. MAYFLIES

139

Bray, R.J., and J.T. Curtis. 1957. An ordination of the upland forest communities of Southern Wisconsin. Ecolo-
gical Monographs, 27:325-349.

CDFG. 2003. California Stream Bioassessment Procedure: Protocol Brief for Biological and Physical Habitat
Assessment in Wadeable Streams. California Department of Fish and Game Aquatic Bioassessment
Laboratory, Water Pollution Control Laboratory. Retrieved September 1, 2014 from www.epa.gov/
region9/qa/pdfs/csbp_2003 .pdf.

CDFW. 2014. Aquatic Bioassessment Laboratory (ABL) Digital Reference Collection. Retrieved August 20, 2014
from www.dfg.ca.gov/abl/Lab/califomia_referencecollection.asp.

Covich, A.P., M.A. Palmer, and T.A. Crowl. 1999. The Role of Benthic Invertebrate Species in Freshwater Eco-
systems. BioScience, 49:119-127.

Fetscher, A. E., L. Busse, and P.R. Ode. 2009. Standard Operating Procedures for Collecting Stream Algae Sam-
ples and Associated Physical Habitat and Chemical Data for Ambient Bioassessments in California:
Retrieved September 9, 2014 from www.waterboards.ca.gov/water_issues/programs/swamp

Frost, T.M., S.R. Carpenter, A.R. Ives, and T.K. Kratz. 1995. Species compensation and complementarity in eco-
system function. Pp. 224-39 in Linking species and ecosystems (Jones, C.G., and J.H. Lawton, eds.). New
York: Chapman and Hall.

Griswold, M. W., R.W. Berzinis, T.L. Crisman, and S.W. Golladay. 2008. Impacts of climatic stability on the
structural and functional aspects of macroinvertebrate communities after severe drought. Freshwater Biol-
ogy, 53:2465-2483.

Gupta, A., S. Gupta, and R.G. Michael. 1994. Seasonal abundance and diet of Cloeon sp. (Ephemeroptera: Bae-
tidae) in a northeast Indian Lake. Archaeological Hydrobiology, 130(3):349-357.

Harrington, J. 1996. California Stream Bioassessment Procedures. Rancho Cordova, CA: California Department
of Fish and Game, Water Pollution Control Laboratory.

Hawkins, C.P., and J.R. Sedell, 1981. Longitudinal and seasonal changes in the functional organization of macro-
invertebrate communities in four Oregon streams. Ecology, 62:387-397.

Houghton, J.T., L.G.M. Filho, B.A. Callander, N. Harris, A. Kattenberg, and K. Maskell. 1995. The Science of
Climate Change, Intergovernmental Panel on Climate Change, Cambridge University Press, Cambridge,
572 pp.

Iverson, T.M., P.W. Wiberg-Larsen, S. Hansen, and F.S. Hansen. 1978. The effect of partial and total drought on
the macro invertebrate communities of three small Danish streams. Hydrobiologia, 60(3):235-242.

Kaller, M.D., and K.J. Hartman. 2004. Evidence of a threshold level of fine sediment accumulation for altering
benthic macroinvertebrate communities. Hydrobiologia, 518:95-104.

Krug, M.J., E. Bell, and R. Dagit. 2012. Growing up fast in a small creek: diet and growth of a population of
Oncorhynchus mykiss in Topanga Creek, California. California Fish and Game, 98(1 ):3 8 — 46.

Lake, P.S. 2003. Ecological effects of perturbation by drought in flowing waters. Freshwater Biology,
48:1161-1172.

Lake, P. S. 2011. Drought and Aquatic Ecosystems: Effects and Responses. John Wiley & Sons, Ltd, Chiche-
ster, UK.

Mayer, M.S., and G.E. Likens. 1987. The importance of algae in a shaded headwater stream as food for an abun-
dant caddisfly (Trichop tera). Journal of the North American Benthological Society, 6:262-269.

Merritt, R.W., K.W. Cummings, Berg, M.B. eds. 2008. An Introduction to the Aquatic Insects of North America.
Kendall Hunt. Dubuque, Iowa.

NOAA. 2014. California Facing Worst Drought on Record. Retrieved January 29, 2014 from www.climate.gov/
news-features/event-tracker/califomia-facing-worst-drought-record

Ode, P.R. 2003. CAMLnet: list of California macro invertebrate taxa and standard taxonomic effort. Aquatic
Bioassessment Laboratory, Rancho Cordova. Retrieved September 10, 2014 from http://www.safit.org/
ste.html.

Ode, P.R., A.C. Rehn, and J.T. May. 2005. A quantitative tool for assessing the integrity of southern California
coastal streams. Environmental Management, 35:493-504.

Ode, P.R. 2007. Standard Operating Procedures for collecting Macroinvertebrate Samples and Associated Physical
and Chemical data for Ambient Bioassessments in California. SWAMP Bioassessment Procedures.
Retrieved September 8, 2014 from www.waterboards.ca.gov/water_issues/swamp.

Peck, D. V., J. M. Lazorchak, and D. J. Klemm, eds. 2004. Environmental Monitoring and Assessment Program-
Surface Waters: Field Operations and Methods for Measuring the Ecological Condition of Wadeable
Streams. EPA/620/R-94/004F. US Environmental Protection Agency, Washington, DC.

Resh, V.H., L.A. Beche, J.E. Lawrence, R.D. Mazor, E.P McElravy, A.P. O’Dowd, D. Rudnick, and S.M. Carlson.
2012. Long-term population and community patterns of benthic macroinvertebrates and fishes Northern
California Mediterranean-climate streams. Hydrobiologia. DOI 10.1007/sl0750-012-1373-9.

140

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Scheffer, M., D. Straile, E. H. Van Nes and H. Hosper, 2001. Climate warming causes regime shifts in lake food
webs. Limnology and Oceanography, 46:1780-1783.

US Census Bureau. 2010. State & County QuickFacts. Retrieved September 8, 2014 from http://quickfacts.census.
gov/qfd/states/06/0678960.html

U.S. Drought Monitor Map Archive. The National Drought Mitigation Center. Lincoln, Nebraska. Retrieved
August 1, 2014 from http://droughtmonitor.unl.edu/

Vannuchi, R, M. Lopez-Rodriguez, J. Tiemo de Figueroa and E. Gaino. 2013. Structure and Dynamics of a
Benthic Trophic Web in a Mediterranean Seasonal Stream. Journal of Limnology, 72:606-615

Vieira, N.K.M., N.L. Poff, D.M. Carlisle, S.R. Moulton, M.L. Koski, and B.C. Kondratieff. 2006. A database of
lotic invertebrate traits for North America: U.S. Geological Survey Data Series 187, http://pubs.water.usgs.
gov/dsl87.

Voshell, J.R. 2002. A Guide to Common Freshwater Invertebrates of North America. McDonald and Woodward
Publishing Company.

Wright, J.F., and K.L. Symes. 1999. A nine-year study of the macroinvertebrate fauna of a chalk stream. Hydro-
logical Processes, 13:371-385.

Bull. Southern California Acad. Sci.

114(3), 2015, pp. 141-148

© Southern California Academy of Sciences, 2015

The Bigeye Scad, Selar crumenophthalmus (Bloch, 1793) (Family
Carangidae), New to the California Marine Fauna, with a List to
and Keys for All California Carangids

Milton S. Love,1* Julianne Kalman Passarelli,2 Chris Okamoto,2 and Dario W. Diehl3

1 Marine Science Institute, University of California, Santa Barbara, CA 93106
2Cabrillo Beach Marine Aquarium, 3720 Stephen M White Dr, San Pedro, CA 90731
3 Southern California Coastal Water Research Project, 3535 Harbor Blvd., Suite 110,

Costa Mesa, CA 92626

The anomalously warm waters of the northeast Pacific 2014-2015 brought with it a variety of
subtropical and tropical fish species previously unusual or absent from California waters (Bond
et al., 2015; pers. comm. M. McCrea, pers. comm.; J. Shepherd). On 1 February 2015, Mr. Kei-
chi Yamamoto speared a fish that we have identified as Selar crumenophthalmus (Bloch, 1793),
the bigeye scad (Fig. 1). The fish was captured in the midwaters of a kelp bed (bottom depth 8
m) off Rancho Palos Verdes (33°48'N, 118°24'W), southern California. This is the first time
this species has been reported from off California. The fish he speared was one of approximately
10 conspecifics that were swimming with a school of juvenile jack mackerel, Trachurus symme-
tries (Ayres, 1855). This specimen is housed in the fish collection at the Natural History
Museum of Los Angeles County, LACM 58288-1.

We identified this fish (22.4 cm fork length) through the following diagnostic characters: Two
papillae on the bone margin at the rear of the gill chamber (the lower one larger than the upper);
the presence of a large eye (eye width greater than snout length) with adipose eyelid covers; and
dorsal, anal, and pectoral fin counts and gill raker counts (Table 1 ). Meristics from our specimen
fall within the range reported by other studies. Hardened scutes were present in the posterior lat-
eral line. No fmlets were present. The fish color was olive on dorsum, with a golden stripe along
flanks, and a silvery belly. There was a diffuse golden ring around the eye.

Bigeye scad are normally circumtropical. In the western Pacific, they have been found as far
northward as the Pacific coast of southern Japan (Nakabo, 2002) and Sea of Japan (Parin, 2003).
Previous to this capture, the eastern Pacific range was from Lagunas Ojo de Liebre-Guerrero
Negro, central Baja California (Galvan-Magana et al., 2000) to Cabo Blanco, Peru (Chirichigno,
1974), including the Gulf of California (Smith- Vaniz in Fischer et al., 1995), Islas Galapagos
(Grove and Lavenberg, 1997), Isla de Malpelo, Isla de Coco, Isla Clipperton, and Islas Revilla-
gigedo (Robertson and Allen, 2008). This is a pelagic species found from surface waters to
depths of 170 m (558 ft) (Randall et al., 1990, Allen and Robertson, 1994). It reaches a maxi-
mum length of 70 cm (Kuiter and Tonozuka, 2001).

Of particular interest is that Mr. Yamamoto first observed a similar number of what were
likely the same species in the same location in November 2014, also mixed in with juvenile
jack mackerel. In addition, Mr. Yamamoto continued to observe bigeye scad at this location
to as late as early March 2015, at this time swimming with a school of juvenile Pacific barracuda
(Sphyraena argentea Girard, 1852). These sightings suggest that at least some bigeye scad may
remain within a relatively circumscribed area for lengthy periods.

* Corresponding author: love@lifesci.ucsb.edu

141

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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 1. A bigeye scad, Selar crumenophthalmus (Bloch, 1793), taken on 1 February 2015, off Rancho Palos
Verdes (33°48'N, 118°24'W), southern California. Catalogued in the Los Angeles County Museum of Natural
History as LACM 58288-1.

The addition of the bigeye scad to the California marine fauna brings the total number of jack
species taken from these waters to 16 (Table 2). The last key to the California species (Miller
and Lea, 1972) was based on 13 species and we include a new key to all known California spe-
cies, using the key to eastern Pacific species created by W. F. Smith-Vaniz in Fischer et al.
(1995) as a starting point. We note that this new key was tested on individuals of all carangids
found in California waters. In addition to the California species, we have also included in the
key the Japanese amberjack, Seriola quinqueradiata Timminck & Schlegel, 1845. In 2015,
this species was found inside the wreckage of a Japanese vessel destroyed in the Fukishima tsu-
nami that had floated to the Oregon coast (J. Burke, pers. comm.) raising the possibility of
future captures off California. In addition, this species is often sold in California fish markets
and in our experience this has led enforcement officers to confuse it with the eastern Pacific spe-

cies, the yellowtail, Seriola dorsalis Gill (1863).

Key to the California Carangidae

la Head profile strongly oblique; very small, embedded scales, leading to apparently naked

body; very compressed body in adults 2

lb Head profile not strongly oblique; no embedded scales; moderately compressed body in

adults 3

2a Oblique head profile has a slight concavity in front of eyes; small juveniles with an oval

black spot above the straight portion of the lateral line: Selene peruviana

2b Oblique head profile lines nearly straight in front of eyes; small juveniles with 4 or 5

interrupted dark vertical stripes on the body: Selene brevoortii

3a From lb: Head profile not strongly oblique...

Posterior straight part of lateral line with hardened scutes or bony shields, scutes can be

poorly developed 4

3b Posterior straight part of lateral line without scutes (only pored scales, not enlarged) 12

Table 1. Meristics of bigeye scad, Selar crumenophthalmus (Bloch, 1793) from this study, Moser et al.
(1996), Nakabo (2002), and Robertson and Allen (2015).

Source

Dorsal fin

Anal fin

Pectoral fin

Gill rakers 1st arch

This study

VIII+1,27

11+1,23

23

11+271

Moser et al. (1996)

VIII+1, 24-27

11+1,20-23

19-23

9-12+27-37

Nakabo (2002)

VIII+1, 23-28

11+1,21-23

9-12+24-31

Robertson and Allen (2015)

VIII+24-27

11+1,21-23

9-12+27-37

1 Counts from right arch as left gill arches were lost during capture.

BIGEYE SCAD, NEW TO CALIFORNIA

143

Table 2. Members of the family Carangidae that are known from California waters, with notes on their max-
imum sizes, and geographic and depth ranges.

Caranx caballus Gunther, 1868. Green Jack. To at least 70 cm TL (Allen and Robertson 1994). Monterey Bay,
central California (Lea and Walker 1995) to Chile (Pequeno 1989), including Gulf of California (Robert-
son and Allen 2002), Islas Galapagos (Miller and Lea 1972), and Hawaii (Randall and Carlson 1999).
Surf zone and to 100 m (min.: Carlisle et al. 1960; max.: De La Cruz-Aguero et al. 1997). D VIII +
1,21-25; A II + 1,17-24; Pect. 20; LL scutes 45-56; GR 13-16 + 25-32; Vert. 25. Recently as Caran-
goides caballus (Randall 2007).

Caranx caninus Gunther, 1867. Pacific Crevalle Jack. To 100 cm TL (Smith- Vaniz in Fischer et al. 1995).
Warm waters of eastern Pacific; Huntington Beach, southern California (Miller and Curtis 2008) to Iqui-
que, northern Chile (Sielfeld et al. 20 1 0), including Gulf of California (Robertson and Allen 2002), Islas
Galapagos (Grove and Lavenberg 1997), and other offshore islands (Robertson and Allen 2002). Surface
to 350 m (Smith- Vaniz in Fischer et al. 1995). D VII-VIII + 1,18-23; A 11+1,15-18; Pect. 1,18-21; LL
shields 25-42; GR 15-19 on lower limb; Vert. 24. Some previous records report this as Caranx hippos',
however that species is now considered a separate Atlantic and Caribbean taxa.

Caranx sexfasciatus Quoy & Gaimard, 1825. Bigeye Trevally. To 120 cm TL (Sadovy and Cornish 2000).
Pacific and Indian oceans; southern Japan (Nakabo 2002); San Diego Bay, southern California (Lea
and Walker 1995) to northern Peru (Robertson and Allen 2015), including lower Gulf of California
(Robertson and Allen 2002) and Islas Galapagos (Grove and Lavenberg 1997). Shallow, near reefs
(Moser 1996), 1-96 m (min.: Gonzalez-Acosta et al. 1999; max.: Myers 1999). D VII-VIII + 1,19-22;
A II + 1,14-17; Pect. 18-21; Pelvic 1,5; GR 4-8 + 15-19; Vert. 25.

Caranx vinctus Jordan & Gilbert, 1882. Cocinero. To 38 cm TL (Amezcua Linares 1996). San Diego Bay,
southern California (Lea and Rosenblatt 2000) to Tumbes, Peru (Chirichigno and Velez 1998), including
central and southern Gulf of California (Robertson and Allen 2002). Surface to 50 m (De La Cruz-
Aguero et al. 1997). D VII-VIII + 1,22-24; A 11+1,18-21; Pect. 19; GR 11-12 + 28-30; Vert. 25. Also
recently as Carangoides vinctus (Angulo et al. 2013).

Chloroscombrus orqueta Jordan & Gilbert, 1883. Pacific Bumper. To 31 cm SL [about 38.8 cm TL) (Amezcua
Linares 1996). San Pedro, southern California (Miller and Lea 1972) to Chilca, Peru (Beltran-Leon and
Rios Herrera 2000), including Gulf of California (Smith- Vaniz in Fischer et al. 1995) and Isla Malpelo
(Robertson and Allen 2002). Shallow coastal waters and estuaries (Moser 1996) to 53 m (Zeballos
1998). D VII-VIII + 1,26-30; A II + 1,25-30; Pect. 19-21; Pelvic 1,5; LL shields nearly obsolete, less
than 20; GR 8-10 + 30-37; Vert. 24.

Decapterus muroadsi (Temminck & Schlegel, 1844). Amberstripe Scad. To about 55 cm TL (Robertson and
Allen 2002). Warm waters of Pacific; Japan to East China Sea (Masuda et al. 1984); Pacific Grove, cen-
tral California (Miller and Lea 1972) to Peru (Moser 1996), Easter Island (Pequeno 1989), and Islas
Galapagos (Grove and Lavenberg 1997); apparently not in Gulf of California (Robertson and Allen
2002). Surface to 320 m (Mundy 2005). D VII+VIII + 1,29-33 + finlet; A II + I, 25-29 + 1 finlet;
Pect. 22-23; LL enlarged shields about 30; GR 13-16 + 37-42; Vert. 24. Includes Decapterus hypodus
Gill, 1862, and Decapterus scombrinus (Valenciennes, 1846) as junior synonyms.

Naucrates ductor (Linnaeus, 1758). Pilotfish. To 70 cm TL (Smith- Vaniz in Fischer et al. 1995). Circumglo-
bal; in western Pacific as far north as Japan (Nakabo 2002) and southern Kuril Islands (Savinykh 1998);
Vancouver Island, British Columbia (Eschmeyer and Herald 1983) to Chile (Pequeno 1989), including
Gulf of California (Smith- Vaniz in Fischer et al. 1995) and such offshore islands as Islas Galapagos
(Miller and Lea 1972). Surface to 150 m (Robertson and Allen 2002). D III-VI + 1,24-29; A I-II +
1,15-18; Pect. 18-20; Pelvic 1,5; GR 5-8 + 12-19 = 18-26; Vert. 25.

Selar crumenophthalmus (Bloch, 1793). Bigeye Scad. To 70 cm (Kuiter and Tonozuka. 2001). Circumglobal;
Pacific coast of southern Japan (Nakabo 2002) and Sea of Japan (Parin 2003); Palos Verdes, southern
California (this paper) to Cabo Blanco, Peru (Chirichigno 1974), including Gulf of California (Smith-
Vaniz in Fischer et al. 1995) and Islas Galapagos (Grove and Lavenberg, 1997), Isla de Malpelo, Isla
de Coco, Isla Clipperton, and Islas Revillagigedo (Robertson and Allen, 2008). Surface to 170 m
(min.: Allen and Robertson 1994; max.: Randall et al. 1990). D VIII + 1,23-28; A II + I, 20-23; Pect.
19-23; GR 9-12 + 24-37; Vert. 24.

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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 2. Continued.

Selene brevoortii (Gill, 1 863). Mexican Lookdown. To 42 cm TL (Robertson and Allen 2002). Seal Beach,
southern California (Jarvis et al. 2009) to northern Chile (Sielfeld et al. 2010), including Gulf of California
(Smith- Vaniz in Fischer et al. 1995). Coastal waters, including bays and estuaries (Moser 1996), to 50 m
(Robertson and Allen 2002). D VII-VIII + 1,20-24; A II + 1, 1 7-1 9; Pect. 18-19; GR 5-9 + 28-34; Vert. 24.

Selene peruviana (Guichenot, 1866). Pacific Moonfish. To variously between 35 cm TL (Velasco and Thiel
2002) and 85 cm TL (Franke and Acero 1993). Long Beach, southern California (Miller and Lea
1972) to Chile (Pequeno 1989), including Gulf of California (Smith- Vaniz in Fischer et al. 1995) and
Islas Galapagos (Grove and Lavenberg 1997). Inshore (Grove and Lavenberg 1997) and shallow coastal
waters (Moser 1996) to 450 m (Franke and Acero 1993). D VIII + 1,20-24; A II + 1,16-19; LL shields
nearly obsolete; GR 8 + 28-32; Vert. 24.

Seriola dorsalis Gill (1863). Yellowtail. To at least 1.5 m (5 ft) TL (Miller and Lea 1972). In eastern Pacific
from northern British Columbia (54°35'N, 31°00'W; Nagtegaal and Farlinger 1981) to central Mexico
(Robertson and Allen 2015), including Gulf of California (Miller and Lea 1972), Islas Galapagos (Grove
and Lavenberg 1997), and Isla Malpelo (Robertson and Allen 2002). Unverified reports from Gulf of
Alaska off Kodiak Island and Cordova (Mecklenburg et al. 2002). Primarily epipelagic, recorded from
surface to at least 91 m (300 ft) (min.: Miller and Lea 1972; max.: N. Ben-Aderet, pers. comm.). Indivi-
duals from the eastern Pacific have recently been referred to as the widely distributed Seriola lalandi
Valenciennes, 1833. However, we follow Martinez-Takeshita et al. (2015) and limit the distribution of
S. lalandi to the Southern Hemisphere, resurrecting S. aureovittata Temminck & Schlegel (1845) in
the northeast Pacific, and S. dorsalis in the eastern Pacific. D IV-VII + 1,29-39; A O-II + 1,19-25;
Pect. 19-20; Pelvic 1,5; GR 5-10 + 15-22; Vert. 25.

Seriola rivoliana Valenciennes, 1833. Almaco Jack. To 160 cm FL [about 176 cm TL] (IGFA). Circumglobal;
Korea (Kim et al. 1997) and southern Japan (Nakabo 2002); Oceanside, southern California (Eschmeyer
and Herald 1983) to Cabo Blanco, Peru (Chirichigno and Velez 1998), including Gulf of California
(Smith- Vaniz in Fischer et al. 1995) and Islas Galapagos (Eschmeyer and Herald 1983). Pelagic, at
depths of 1-250 m (min.: Kuiter and Tonozuka 2001; max.: Robertson and Allen 2002). D VIII +
1,26-33; A II + 1,18-21; Pect. 19-22; LLs 167; GR 6-9 + 16-19 = 22-28; Vert. 24.

Trachinotus paitensis Cuvier, 1832. Paloma Pompano. To 50.8 cm TL (Miller and Lea 1972). Redondo Beach,
southern California (Miller and Lea 1972) to Chile (Pequeno 1989), including Gulf of California and
Islas Galapagos (Miller and Lea 1972). Shallow inshore areas (Miller and Lea 1972) and to 100 m
(Amezcua Linares 1996). D VI- VIII, 20-27; A II+I (occasionally III), 20-24; Pect. 16-18; Pelvic 1,5;
GR 9-11 + 11-17; Vert. 24.

Trachinotus rhodopus Gill, 1863. Gafftopsail Pompano. To 61 cm TL (Miller and Lea 1972). Zuma Beach,
southern California (Miller and Lea 1972) to Callao, Peru (Chirichigno and Velez 1998), including
southern and central Gulf of California (Robertson and Allen 2002) and Islas Galapagos (Miller and
Lea 1972). Surface to 30 m (min.: Robertson and Allen 2002; max.: Amezcua Linares 1996). D V-
VI + 1,19-21; A II + 1,17-21; Pect. 18-19; Pelvic 1,5; GR 8-10 + 13-16 = 21-26; Vert. 24.

Trachurus symmetricus (Ayres, 1855). Jack Mackerel. To 81.3 cm TL (Miller and Lea 1972). Pacific Ocean
south of Aleutian Islands (Mecklenburg et al. 2002 [UW 1 5469]) and in Gulf of Alaska to Gulf of Cali-
fornia (Smith- Vaniz in Fischer et al. 1995) and to Acapulco, Mexico (Palacios-Salgado et al. 2014). Pri-
marily pelagic, surface (Miller and Lea 1972) from surf zone (Carlisle et al. 1960) and offshore to 403 m
(Hart 1973). Although Trachurus murphyi Nichols, 1920, found off South America, is considered by
some authors (e.g., Grove and Lavenberg 1997) to be a subspecies of T. symmetricus, DNA evidence
indicates it is a separate species (Poulin et al. 2004). D VIII + 1,28-38; A II + 1,22-33; Pect. 21-24; Pel-
vic 1,5; LLs 87-111, the later 40-55 as enlarged shields; GR 7-15 + 25^12 = 32-57; Vert. 23-25.

Uraspis helvola (Forster, 1801). Whitemouth Jack. To 58 cm TL (Jimenez Prado and Bearez 2004). Indo-Paci-
fic; perhaps southern Kuril Islands (Savinykh and Shevtsov 2001) and southern Japan (Nakabo 2002);
Santa Catalina Island, southern California (Miller and Lea 1972), southernmost Gulf of California (J.
Snow, pers. comm.), and at a number of more southerly locations (Robertson and Allen 2015) to Ecua-
dor (Bearez 1996), including Islas Galapagos (Grove and Lavenberg 1997). 10-300 m (Robertson and
Allen 2002). D V-VIII + 1,25-30; A II + 1,19-22; Pect. 1,22; LL with 32-38 keeled shields; GR 6 + 16
= 22. The Santa Catalina Island specimen was originally identified as Uraspis secunda (Fitch 1972).
However, U. secunda appears to be an Atlantic and Indo-Central Pacific species (W. Smith- Vaniz and

R. Robertson, pers. comms. to M. L.).

BIGEYE SCAD, NEW TO CALIFORNIA

145

4a Lateral line with reduced (very stunted), barely perceptible scutes; a black spot on the

saddle of the upper region of the caudal peduncle; strongly compressed body:

Chloroscombrus orqueta

4b Well-developed lateral line scutes; caudal peduncle without black spot; body not strongly

compressed 5

5 a Dorsal accessory branch of the lateral line extends at least to below origin of 2nd dorsal fm,

usually farther posteriorly: Trachurus symmetricus

5b Dorsal accessory branch of the lateral line terminating before origin of dorsal fin 6

6a Independent 2-rayed finlet at caudle peduncle (dorsal and ventral): Decapterus muroadsi

6b Second dorsal and anal fins without finlets 7

7a Cleithrum, under the operculum, with a furrow (groove) on the ventral side, a large papilla-
like structure immediately above the groove; eye nearly covered by an adipose eyelid,

forming a vertical slit on the center of the pupil: Selar crumenophthalmus

7b Smooth-edge cleithrum; adipose eyelid marginally covering eye but not pupil 8

8a Tongue, roof, and floor of mouth white, the rest black; no teeth on vomer or palatines: . . .

Uraspis helvola

8b Lining of mouth not distinctly black and white; teeth present on vomer and palatines 9

9a Breast without scales except for a small patch of scales at base of pelvic fins (prepelvic):

Caranx caninus

9b Breast completely scaled 10

10a Number of scutes in the lateral line 26-42; total number of gill rakers on the lower limb

(including rudiments) 15-22; lobe of second dorsal fm with a white tip and its height into

fork length 5.0 to 6.6 times in adults: Caranx sexfasciatus

10b Number of scutes in the lateral line 42-56; total number of gill rakers on the lower limb

(including rudiments) 27-30; lobe of second dorsal fm without white tip and its height into

fork length 6.0 to 8.0 times in adults 11

11a In adults, body without dark vertical stripes; scaly basal sheath along the lobes of the dorsal
and anal fins with relatively narrow scales, above which the fins are almost entirely covered
with small scales; back of the eye covered by a weak adipose eyelid extending to the rear

edge of the pupil: Caranx caballus

lib In adults, body with 8-9 dark incomplete vertical stripes; scaly basal sheath along the lobes
of the dorsal and anal fins with relatively wide scales above which the fins are devoid of
small scales; back of the eye covered by a weak adipose eyelid that does not reach the
posterior edge of the pupil: Caranx vinctus

12a From 3b: Posterior straight part of lateral line without scutes. . .

Caudal-peduncle grooves (or shallow notch) present dorsally and ventrally; base of soft
dorsal and anal fins unequal in length with anal-fin base shorter (only about 45-70%) than

dorsal-fin base length 13

12b Caudal-peduncle grooves absent; base of soft anal fm as long as, or only slightly shorter

than, base of dorsal fm 15

13a Fleshy keel on caudal peduncle well developed; soft anal-fin rays 15-17; body with 5-6

dark vertical bars, extending to bases of dorsal and anal fins: Naucrates ductor

13b Fleshy keel on caudal peduncle absent to moderately developed; soft anal-fin rays 18-22;

body lacking obvious wide vertical bars 14

14a Longest dorsal soft-ray about 1/2 length of head; relatively slender supramaxilla; caudal fin
yellowish; yellow stripe along mid-body; head longer than body depth at origin of dorsal
fm: Seriola dorsalis

Note: Seriola quinqueradiata can be differentiated from S. dorsalis by 1) the latter’s more
rounded dorsoposterior comer of upper jaw (Figure 2) and 2) a pectoral fin shorter than
pelvic fin versus 1) more angular and 2) almost equal in length.

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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

/

Seriola quinquerodiato

Fig. 2. An illustration comparing the upper jaw shapes of Seriola dorsalis and S. quinqueradiata. Note the
more rounded dorsoposterior comer of the upper jaw of S. dorsalis.

14b Longest dorsal soft-ray about 2/3 length of head in fish over 18 cm fork length; relatively
broad supramaxilla; caudal fin dark to dusky; black stripe radiating from mandibles, through eye,

to 1st dorsal spines; head shorter than body depth at origin of dorsal fin: Seriola rivoliana

15a From 12b : Caudal-peduncle grooves absent...

First 4-5 dorsal and anal soft-rays elongated, longer than head length in fish over 8 cm fork
length; in adults, 4-6 incomplete vertical bars extending beyond the lateral line; 22-26

gill rakers (8-10 on upper limb): Trachinotus rhodopus

1 5b First 4-5 dorsal and anal soft-rays equal to or shorter than head length; in adults, no vertical bars as
described above; 15-18 gill rakers (5-7 on upper limb): Trachinotus paitensis

Acknowledgments

We thank Keichi Yamamoto for bringing this specimen to our attention. Richard Feeney, Nat-
ural History Museum of Los Angeles County, provided samples of California jacks for testing
the key. The following researchers reviewed the Carangidae key: Don Buth, Craig Campbell,
Dario Diehl, Rick Feeney, Robin Gartman, Pete Major, Jim Mann, Mike Mengel, Julianne Pas-
sarelli, Terra Petry, Bill Power, Jim Rounds, and Fred Stem.

Literature Cited

Allen, G.R. and D.R. Robertson. 1994. Fishes of the tropical eastern Pacific. University of Hawaii Press,
Honolulu.

Amezcua Linares, F. 1996. Peces demersales de la platforma continental del Pacifico Central de Mexico. Institute
de Ciencias del Mar y Limnologica, Universidad Nacional Autonoma de Mexico.

Angulo, A., C.A. Garita-Alvarado, W.A. Bussing, and M.I. Lopez. 2013. Annotated checklist of the freshwater
fishes of continental and insular Costa Rica: additions and nomenclatural revisions. Check List
9:987-1019.

Bearez, P. 1996. Lista de los peces marinos del Ecuador continental. Rev. Biol. Trop., 44:731-741.
Beltran-Leon, B.S. and R. Rios Herrera. 2000. Estadios tempranos de peces del Pacific Colombiano. Republica de
Colombia. Institute Nacional de Pesca y Acuicultura INPA.

BIGEYE SCAD, NEW TO CALIFORNIA

147

Bond, N.A., M.F. Cronin, H. Freeland, and N. Mantua. 2015. Causes of impacts of the 2014 warm anomaly in the
NE Pacific. Geophys. Res. Letters 42, doi:10.1002/2015GL063306.

Carlisle, J.G. Jr., J.W. Schott, and N.J. Abramson. 1960. The barred surfperch ( Amphistichus argenteus Agassiz)
in southern California. Calif. Dep. Fish Game Fish Bull. 109.

Carpenter, K.E. and V.H. Niem (eds.). 1999. The living marine resources of the Western Central Pacific. Vol. 3.
Batoid fishes, chimaeras and bony fishes part 1 (Elopidae to Linophrynidae). Vol. 4. Bony fishes part 2
(Mugilidae to Carangidae). FAO, Rome.

Chirichigno, F.N. 1974. Clave para identificar los peces marinos del Peru. Instituto del Mar del Peru Informe,
No. 44.

Chirichigno, F.N. and J. Velez D. 1998. Clave para identificaticar los peces marinos del Peru (segunda edicion,
revisada y actualizada). Instituto de Mar de Peru. Publicacion Especial.

De La Cruz-Agiiero, J., M.A. Martinez, V.M. Cota-Gomez, and G. De La Cruz-Aguero. 1997. Catalogo de los
peces marinos de Baja California sur. Centro Interdiscipinario de Ciencias Marinas.

Eschmeyer, W.N. and E.S. Herald. 1983. A field guide to Pacific Coast fishes of North America from the Gulf of
Alaska to Baja California. Houghton Mifflin, Boston.

Fitch, J.E. 1972. The cottonmouth jack, Uraspis secunda, added to the marine fauna of California. Calif. Fish
Game, 58:245-246.

Franke, R. and A. Acero P. 1993. Peces carangoideos del Parque Gorgona, Pacifico Colombiano (Osteichthys:
Carangidae, Nematistiidae y Coryphaenidae). Rev. Biol. Mar., Valparaiso, 28:51-73

Galvan-Magana, F., F. Gutierrez-Sanchez, L.A. Abitia-Cardenas, and J. Rodriquez-Romero. 2000. The distribu-
tion and affinities of the shorefishes of the Baja California Sur lagoons. Pp. 383-398 in Aquatic ecosys-
tems of Mexico: status and scope. Ecovision World Monograph Series (M. Munawar, S.G. Lawrence, I.
F. Munawar, and Malle, D.F. eds.). Backhuys Publisher, Leiden, The Netherlands.

Gonzalez-Acosta, A.F., G. De La Cruz-Aguero, J. De La Cruz-Aguero, and G. Ruiz-Campos. 1999. Ictiofauna
asociada al manglar del Estero el Conchalito, Ensenada de la Paz, Baja California Sur, Mexico. Oceanides,
14(2): 121— 1 3 1 .

Grove, J.S. and R.J. Lavenberg. 1997. The Fishes of the Galapagos Islands. Stanford University Press, Stanford,
California.

Hart, J.L. 1973. Pacific Fishes of Canada. Fish. Res. Board Can., Bull. 180.

Jarvis, E.T., H.L. Gliniak, O. Homing, and C. Linardich. 2009. Occurrence of juvenile Mexican lookdown, Selene
brevoortii (Gill, 1863), in Seal Beach, California. California Fish and Game, 95:188-192.

Jimenez Prado, P. and P. Bearez. 2004. Peces marinos del Ecuador continental/marine fishes of continental Ecua-
dor. SIMBIOE/NAZCA/IFEA TOMO II. Quito.

Kim, Y.S., Y.U. Kim, and G. Ahn. 1997. First record of the carangid fish, Seriola rivoliana from Korea. Korean J.
Ichthyol., 9:244-247.

Kuiter, R.H. and T. Tonozuka. 2001. Pictorial Guide to Indonesian Reef Fishes. Part 1. Eels-Snappers, Muraeni-
dae-Lutjanidae. Zoonetics, Australia. 302 pp.

Lea, R.N. and R.H. Rosenblatt. 2000. Observations on fishes associated with the 1997-98 El Nino off California.
CalCOFI Rep., 41:117-129.

Lea, R.N. and H.J. Walker Jr. 1995. Record of the bigeye trevally, Caranx sexfasciatus , and Mexican lookdown,
Selene brevoorti , with notes on other carangids from California. Calif. Fish Game, 81:89-95.

Martinez-Takeshita, N., C.M. Purcell, C.L. Chabot, M.T. Craig, C.N. Paterson, J.R. Hyde, and L.G. Allen. 2015.
A tale of three tails: cryptic speciation in a globally distributed marine fish of the genus Seriola. Copeia,
103(l):357-368.

Masuda, H., K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino. 1984. The Fishes of the Japanese Archipelago.
Tokai Univ. Press.

Mecklenburg, C.W., T.A. Mecklenburg, and L.K. Thorsteinson. 2002. Fishes of Alaska. American Fisheries
Society, Bethesda, Maryland.

Miller, D.J. and R.N. Lea. 1972. Guide to the coastal marine fishes of California. Calif. Dep. Fish Game Fish
Bull. 157.

Miller, E.F. and M.D. Curtis. 2008. First occurrence of a Pacific crevalle jack, Caranx caninus, north of San
Diego, California. Bull. S. Calif. Acad. Sci., 107:41-13.

Moser, H.G. (ed.). 1996. The early stages of fishes in the California Current region. CALCOFI (California Coop-
erative Oceanic Fisheries Investigations) Atlas No. 33. U.S. Department of Commerce, NOAA, NMFS,
Southwest Fisheries Science Center, La Jolla, California.

Mundy, B.C. 2005. Checklist of the Fishes of the Hawaiian Archipelago. Bishop Museum Press, Honolulu,
Hawaii.

Myers, R. F. 1999. Micronesian Reef Fishes. Coral Graphics, Barrigada, Guam.

148

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Nagtegaal, D.A. and S.R Farlinger. 1981. First record of two fishes, Seriola dorsalis and Medialuna californien-
sis, from waters off British Columbia. Syesis 13:206-207. [Journal dated 1980 but not issued until 1981.]

Nakabo, T. (ed.). 2002. Fishes of Japan with Pictorial Keys to the Species, English edition. Tokai University Press,
Tokyo.

Palacios-Salgado, D. S., A. Ramirez- Valdez, A. A. Rojas-Herrera, J. G. Amores, and M. A. Melo-Garcia. 2014.
Marine fishes of Acapulco, Mexico (eastern Pacific Ocean). Mar. Biodiv. 44:471-490.

Parin, N.V. 2003. An annotated catalogue of fish-like vertebrates and fishes of the seas of Russia and adjacent
countries: Part 3. Orders Perciformes (excluding suborders Gobioidei, Zoarcoidei and Stichaeoidei) and
Tetraodontiformes. J. Ichthyol. 43:S1-S40.

Pequeno R.G. 1989. Peces de Chile lista sistematica revisada y comentada. Rev. Biol. Mar. (Valparaiso) 24:1-132.

Poulin, E., L. Cardenas, C.E. Hernandez, I. Komfield, and F.P. Ojeda. 2004. Resolution of the taxonomic status of
Chilean and California jack mackerels using mitochondrial DNA sequence. J. Fish Biol. 65:1160-1164.

Randall, J. 2007. Reef and Shore Fishes of the Hawaiian Islands. Sea Grant College Program, University of
Hawai’i, Honolulu.

Randall, J. and B.A. Carlson. 1999. Caranx caballus, a new immigrant carangid fish to the Hawaiian Islands from
the tropical eastern Pacific. Pac. Sci. 53:357-360.

Randall, J., G.R. Allen, and R.C. Steene. 1990. Fishes of the Great Barrier Reef and Coral Sea. University of
Hawaii Press, Honolulu.

Robertson, D.R. and G.R. Allen. 2002. Shorefishes of the Tropical Eastern Pacific: an Information System. CD-
ROM. Smithsonian Tropical Research Institute, Balboa, Panama.

Robertson, D.R. and G.R. Allen. 2008. Shorefishes of the tropical eastern Pacific: an information system. Version
1.0 (2008). Smithsonian Tropical Research Institute, Balboa, Panama, www.neotropicalfish.org/sftep.

Robertson, D.R. and G.R. Allen. 2015. Shorefishes of the Tropical Eastern Pacific: an Information System. Ver-
sion 2.0. Smithsonian Tropical Research Institute, Balboa, Panama, http://biogeodb.stri.si.edu/sftep/en/
pages. Accessed 2 June 2015.

Sadovy, Y. and A.S. Cornish. 2000. Reef fishes of Hong Kong. Hong Kong University Press, Hong Kong.

Savinykh, V.F. 1998. Nekton composition of near-surface waters of the subarctic front zone in the northwest part
of the Pacific Ocean according to the data of drift-net catches. J. Ichthyol. 38:18-27.

Savinykh, V.F. and G.A. Shevtsov. 2001. Two new species from Russian fauna from the waters of the southern
Kuril Islands. J. Ichthyol. 41:122-124.

Sielfeld, W., J. Laudien, M. Vargas, and M. Villegas. 2010. El Nino induced changes of the coastal fish fauna off
northern Chile and implications for ichthyogeography. Rev. Biol. Mar. Oceanog. 45, S 1:705-722.

Smith- Vaniz, W. F. 1995. Carangidae. Pp. 940-986 in Guia FAO para la identification para los fines de la pesca.
Pacifico centro-oriental (Fischer, W., F. Krupp, W. Schneider, C. Sommer, K. E. Carpenter, and V. H. Niem
eds.). Vol. II, Vertebrados, Parte 1. Vol. Ill, Vertebrados, Parte 2. FAO, Rome.

Velasco, A. and R. Thiel. 2002. A bilingual field manual for the identification of juvenile fish over soft bottom off
the Pacific Coast of Colombia a review from literature. Arch. Fish. Mar. Res. 50:55-118.

Zeballos, J., M. Samame, and M. Robero. 1998. Estructura especiologica demersal observada durante el Crucero
de Evaluation de la Merluza entre Puerto Pizarro y Huarmey. BIC Jose Olaya Balandra 9806-07. Inf. Inst.
Mar Peru 138:87-100.

Bull. Southern California Acad. Sci.

114(3), 2015, pp. 149-163
© Southern California Academy of Sciences, 2015

A Flora of the Ballona Wetlands and Environs

E.A. Read

E Read and Associates, Inc. Orange, CA 92866, marshmistress@msn.com

A list of vascular plant taxa is provided for a study area that encompasses the current Ballona
Wetlands Ecological Reserve, Ballona Freshwater Wetlands, and areas that were part of the his-
toric Ballona Wetlands and adjacent bluffs. The flora for this area has been documented in
unpublished reports and voucher collections but not compiled into a single list. A total of 407
taxa comprise this flora, of which 40% are not indigenous to California. This percentage is
higher than the percentage of non-indigenous species in California as a whole (17%). Out of
242 native species reported for the study area, 26 percent (62 species) have not been observed
or collected since major hydrological alterations of the watershed began in the 1930s. Restora-
tion planned for the Ballona Wetlands Ecological Reserve offers an opportunity to re-introduce
some of the historically native flora, including rare and endangered species.

The study area is located about two miles north of Los Angeles International Airport (Fig. 1 ). Limits
of the study area were defined based on maps of the historical extent of the wetlands and adjacent
uplands. Land use planning in this region has been controversial for at least 40 years, with various pro-
posals ranging from development to restoration. With the exception of occasional botanical collec-
tions, systematic floristic surveys were not documented until Clark (1979), when a single season of
field work was conducted in March and April of 1978. Subsequent floristic surveys in 1980-1981
by Gustafson (1981) and in 1990 by Henrickson (1991) covered more than one season. While these
surveys were supported in part by voucher collections deposited with the Natural History Museum of
Los Angeles County, results of these and subsequent surveys (Read 1995; MEC Analytical Systems
2005; Johnston 2011) were not published. This paper combines plant lists and historical collection
records for the Ballona region into a single list and updates nomenclature. Historical conditions, occur-
rence of rare species, and occurrence of non-native invasive taxa are also discussed.

Latin nomenclature was updated based on current taxonomy (Baldwin et al. 2012). In cases
of species subject to extensive taxonomic revisions, older taxonomies were used to verify iden-
tification of historical collections (Abrams 1902, 1917; Munz 1974). Historical maps were
examined to locate landmarks cited in voucher collections but not present on current
maps. These historical maps consisted of a United States Coast and Geodetic Survey in 1876
(Grossinger et al. 2011) and a vegetation mapping survey of the Redondo 15-minute quadrangle
by Jensen (1933). These maps were overlaid on current topography using Geographic Informa-
tion System software to verify locations of historical landmarks relative to the study area.
Reports of archaeological surveys (Altschul et al. 1992, 2003) were reviewed for historical
land alterations potentially affecting the flora.

All plant taxa reported from the study area are listed in Table 1 . This table is necessarily abbre-
viated for presentation here. The entire database, including details such as accession numbers and
references for each observation or collection, can be provided upon request. Only taxa indigenous
or naturalized in the study area were included in this table in order to make the list more mean-
ingful for analysis, restoration and management. In other words, non-invasive, non-native orna-
mental taxa known to be confined to existing or abandoned landscapes were omitted. A total of
407 taxa are shown in Table 1, of which about 40 percent (165 taxa) are not native to California.
Changes in taxonomy over time, especially below the species level, have a minor effect on these
statistics.

149

150

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Out of the 242 native taxa listed, about 26 percent (62 species) have not been observed or col-
lected since the 1930s when there were major hydrological alterations of the area for flood con-
trol and farming. One of these historical species, Ballona cinquefoil ( Potentilla multijuga), is
believed to have been endemic to Ballona but extirpated sometime after its last collection in
1890 by H.E. Hasse from a “brackish area in coastal sage scrub”.

Human impacts in the study area over the past two centuries have been documented exten-
sively by others (Dark et al. 2011; Grossinger et al. 2011), so it is important to recognize that
historical botanical collections from the study area do not represent pristine (pre-human distur-
bance) conditions or a complete flora of the region. A survey map of the wetlands was not pub-
lished until 1876, which is 57 years after cattle herds were pastured in the Ballona Creek
drainage beginning around 1819 (Altschul et al., 1992 p. 56). After 1819, the earliest known
botanical collections from Ballona were by S.B. Parish 63 years later, in 1882.

FLORA OF BALLONA WETLANDS

151

Thus, given at least two centuries of land alteration in the region, it is not surprising that a
substantial portion of the flora is not native. The proportion of non-native flora is high when
compared to the proportion of 17% non-native taxa in California as a whole1, and especially
high considering the small size of the study area (1,824 ha) relative to the geographic extent
of California (42,476 ha).

Most of the non-native flora has established in the absence of systematic efforts at restoring
conditions suitable for sustaining native species in a heavily urbanized watershed, but not all of
the invaders have been equally successful. One of the first records of non-native species is of
fennel ( Foeniculum vulgare ), collected by Abrams in 1902, yet this species has not invaded
the wetlands as extensively as a relative newcomer, pampas grass ( Cortaderia selloana ). There
are no historic collections of Cortaderia from the study area and its occurrence at Ballona was
unremarked until it was reported by Clark (1979).

Coastal estuarine and saltmarsh taxa are noticeably rare in the historical collections. Most of
the historic flora is composed of species associated with upland, freshwater or brackish habitats.
While there could be various reasons for this, the high proportion of non-estuarine taxa in his-
torical collections is consistent with documentation by Dark et al. (2011) of springs and seasonal
ponds that were prevalent historically but no longer exist. One example of this freshwater flora
is giant horsetail ( Equisetum telmateia ), collected by Abrams in 1901 from Ballona Creek while
it was still a meandering, natural stream. Another example is Oregon ash {Fraxinus latifolia)
collected by Ewan in 1932 along Ballona Creek in the area of what is now Culver City. This
is the only native tree represented in the historic flora.

Examination of the plant list also reveals interesting inconsistencies in the taxonomic records.
For example, Senecio califomicus was collected by Abrams in 1901, but later collectors (Gustafson
1981; Henrickson 1991) identified only Senecio vulgaris , a non-native species. This may be a sim-
ple misidentification, but alternatively could be an example of niche displacement of a native Sene-
cio by an exotic species.

Lastly, determination of which species are truly “native” to the study area is complicated by
changes in taxonomy (e.g., “lumping” vs. “splitting” varieties and subspecies), misidentifications,
and intentional introduction of taxa for “restoration” without documentation of variety or seed ori-
gin. A species of pincushion ( Chaenactis glabriuscula ), an annual herb associated with sandy
soils, has been reported several times since it was first collected in 1901. This taxon persists
into the current period, but tracing its history of occurrence is complicated by taxonomic re-clas-
sifications of some historical collections, and “restoration” efforts that distributed C. glabriuscula
seeds of unspecified variety and unknown origin. Currently, a variety considered rare by the Cali-
fornia Native Plant Society and identified as C. g. var. orcuttiana persists in dunes that are subject
to ongoing restoration efforts, but it is not clear whether this variety is indigenous, intentionally
planted, or some combination of both. Some voucher specimens from the study area are classified
as the more common variety, C. g. var. glabriuscula , and deserve further study.

Historical maps show “marsh” but are silent on the issue of what proportion of the wetlands
consisted of freshwater vs. saltwater or brackish marsh. However, botanical records show that
the study area was occupied by a highly diverse ecosystem. The area encompassed not just wet-
lands, but a wide variety of other habitat types including dunes, freshwater marshes, seasonal
ponds, and riparian communities. Taxonomic inconsistencies in the botanical record introduce
some margin of error in calculating species richness, but this is largely of academic interest.
Habitat restoration in the study area offers a unique opportunity to reverse a decades-old trend

1 Calculated from Baldwin et al. (2012), p. 1521 .

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SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

of exotic invasion at the expense of native diversity, while recognizing that significant altera-
tions of the watershed render it impossible to fully restore the historic flora. This study provides
a starting point for enhancing function and diversity of the Ballona Wetlands.

Table 1. Checklist of the Flora of the Ballona Wetlands and Environs.

Earliest

FERNS

Reference

Collection

Equisetaceae

Horsetail Family

Equisetum telmateia Ehrh. ssp. braunii (J.Milde) Hauke

giant horsetail

DS, POM

1901

MAGNOLIDS

Lauraceae

Laurel Family

Umbellularia californica (Hook. & Am.) Nutt.

California bay

FWM

R

Saururaceae

Lizard's-Tail Family

Anemopsis californica (Nutt.) Hook. & Am.

yerba mansa

POM, RSA,UC

1902

EUDICOTS

Adoxaceae

Muskroot Family

Henrickson 1991,

Sambucus nigra L. ssp. caerulea (Raf.) Bolli

blue elderberry

FWM

R

Aizoaceae

Iceplant Family

Aptenia cordifolia (L.f.) Schwantes*

baby sun rose

RSA

1981

Carpobrotus edulis (L.) N.E. Br.*

freeway iceplant

RSA

1980

Delosperma litorale (Kensit) L. Bolus*

ice plant
coppery

RSA

1981

Malephora crocea (Jacq.) Schwantes*

mesembryanthemum

RSA

1981

Mesembryanthemum crystallinum L.*

common iceplant

Henrickson 1991

R

Mesembryanthemum nodiflorum L.*

slender-leaved iceplant

RSA

1980

Sesuvium verrucosum Raf.

western sea-purslane

UC

1902

Tetragonia tetragonioides (Pall.) Kuntze*

New Zealand spinach

RSA

1981

Amaranthaceae

Amaranth Family

Amaranthus albus L.*

tumbleweed

RSA

1930

Amaranthus blitoides S. Watson

procumbent pigweed

RSA

1980

Amaranthus californicus (Moq.) S. Watson

Californian amaranth

Gustafson 1981

R

Amaranthus deflexus L.*

Amaranthus tuberculatus (Moq.) J.D. Sauer var. rudis (J.D.

large fruited amaranth

RSA

1981

Sauer) Costea & Tardif*

common waterhemp

Henrickson 1991

R

Anacardeaceae

Sumac Family

Malosma laurina (Nutt.) Abrams

laurel sumac

POM

1980

Rhus integrifolia (Nutt.) Rothr.

lemonade berry

Gustafson 1981

R

Rhus ovata S. Watson

sugar bush

RSA

1981

Schinus molle L.*

pepper tree

RSA

1980

Schinus terebinthifolius Raddi*

Brazilian pepper tree

Henrickson 1991

R

Apiaceae

Carrot Family

Apium graveolens L.*

celery

RSA

1902

Conium maculatum L.*

poison hemlock

RSA

1981

Cyclospermum leptophyllum (Pers.) Britton & P. Wilson*

marsh parsley

RSA

1981

Foeniculum vulgare Mill.*

fennel

POM,RSA,UC

1902

Oenanthe sarmentosa DC.

water parsley

POM, UC

1902

Asterisk (*) indicates taxon not native to California. Grey background indicates a native taxon known only from
historical (pre-1940) collections or reports. References: For herbarium specimens, the following acronyms from the
Consortium of California Herbaria are used: CAS (California Academy of Sciences); CDA (California Department of
Food and Agriculture); DS (Dudley Herbarium in CAS); JEPS (Jepson Herbarium, UC Berkeley); NY (New York
Botanical Garden); POM (Pomona Herbarium in the Rancho Santa Ana Botanic Garden); RSA (Rancho Santa Ana
Botanic Garden); SFV (California State University Northridge); UC (University Herbarium, UC Berkeley); UCD
(UC Davis); UCR (UC Riverside). Where an author and year are cited, the taxon was reported but no voucher
specimens could be located. “FWM” refers to a taxon that was observed by this author as having been planted at the
constructed Ballona Freshwater Marsh but no records or reports of historical occurrence at Ballona were found.
Earliest Collection: year of oldest voucher specimen is given. “R” means reported with no record of collection.

FLORA OF BALLONA WETLANDS

153

Table 1 . Continued.

Apocynaceae

Asclepias fascicularis Decne.

Asteraceae

Acroptilon repens (L.) DC.*

Amblyopappus pusillus Hook. & Am.

Ambrosia acanthicarpa Hook.

Ambrosia chamissonis (Less.) Greene
Ambrosia psilostachya DC.

Anthemis cotula L.*

Artemisia californica Less.

Artemisia douglasiana Besser
Artemisia dracunculus L.

Baccharis glutinosa Pers.

Baccharis pilularis DC.

Baccharis salicifolia (Ruiz & Pav.) Pers.

Bidens laevis (L.) Britton et al.

Brickellia californica (Torr. & A. Gray) A. Gray
Centaurea melitensis L.*

Centromadia parryi (Greene) Greene ssp. australis (D.D.
Keck) B.G. Baldwin

Chaenactis glabriuscula DC. var. orcuttiana (Greene) H.M.
Hall

Chondrilla juncea L.*

Cichorium intybus L.*

Cirsium vulgare (Savi) Ten.*

Coreothrogyne filaginifolia (Hook. & Am.) Nutt.

Cotula australis (Spreng.) Hook, f.*

Cotula coronopifolia L.*

Deinandra fasciculata (DC.) Greene

Deinandra paniculata (A. Gray) Davidson & Moxley

Dimorphotheca fruticosa (L.) DC.*

Encelia californica Nutt.

Ericameria ericoides (Less.) Jeps.

Ericameria pinifolia (A. Gray) H.M. Hall

Erigeron bonariensis L.*

Erigeron canadensis L.

Euryops pectinatus (L.) Cass.*

Euthamia occidentalis Nutt.

Gazania linearis (Thunb.) Druce*

Glebionis coronaria (L.) Spach*

Grindelia camporum Greene
Hedypnois cretica (L.) Dum.-Cours.*

Helianthus annuus L.

Helminthotheca echioides (L.) Holub*

Heterotheca grandiflora Nutt.

Dogbane Family

narrow-leaf milkweed

RSA

1930

Sunflower Family

Russian knapweed

POM

1980

dwarf coastweed

DS

1901

annual bursage

RSA

1980

beach bur-sage

DS,POM,RSA

1899

western ragweed

DS,POM,UC,RSA

1902

mayweed

DS

1901

California sagebrush

DS,RSA,UC

1902

mugwort

DS, POM, RSA

1901

tarragon

DS,RSA,UC

1902

marsh baccharis

DS,POM

1903

coyote brush

POM

1980

mulefat

POM,UC

1907

bur-marigold

DS,UC

1902

California brickellbush

Henrickson 1991

R

tocalote

POM

1980

southern tarplant

JEPS, POM, RSA, U
C

1890

Orcutt's pincushion

POM, RSA

1980

skeleton weed

POM

1981

chicory

POM

1980

bull thistle

Gustafson 1981,
Henrickson 1991

R

Califomia-aster

DS,POM

1899

Australian cotula

Gustafson 1981,
Henrickson 1991

R

brass-buttons

Gustafson 1981,
Henrickson 1991

R

clustered tarplant

POM

1980

paniculate tarplant

Henrickson 1991

R

trailing African daisy

Gustafson 1981,
Henrickson 1991

R

California brittlebush

DS

1899

California goldenbush

POM

1981

pine-bush

Henrickson 1991

R

flax-leaved horseweed

Gustafson 1981,
Henrickson 1991

R

horseweed

POM

1980

gray-leaf euryops

Henrickson 1991

R

western goldenrod

RSA,UC

1902

treasure flower, African
daisy

Henrickson 1991

R

garland or crown daisy

UCR

1996

common gumplant

POM

1980

Crete weed

POM

1981

common sunflower

POM

1980

bristly ox -tongue

RSA

1980

telegraph weed

POM

1980

154

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 1.

Hypochaeris glabra L.*

Isocoma menziesii (Hook. & Am.)

Iva axillaris Pursh

Jaumea carnosa (Less.) A. Gray
Lactuca serriola L.*

Lactuca virosa L.*

Laennecia coulteri (A. Gray) G.L. Nesom
Lasthenia californica Lindl.

Lasthenia coronaria (Nutt.) Omduff

Lasthenia glabrata Lindl. ssp. coulteri (A. Gray) Omduff

Logfia filaginoides (Hook. & Am.) Morefield

Malacothrix saxatilis (Nutt.) Torrey & A. Gray
Matricaria discoidea DC. *

Pluchea odorata (L.) Cass. var. odorata

Pseudognaphalium bioletti Anderb.

Pseudognaphalium califomicum (DC.) Anderb.
Pseudognaphalium canescens (DC.) Anderb.
Pseudognaphalium microcephalum (Nutt.) Anderb.
Pseudognaphalium ramosissimum (Nutt.) Anderb.
Pseudognaphalium stramineum (Kunth) Anderb.

Senecio californicus DC.

Senecio vulgaris L.*

Silybum marianum (L.) Gaertn.*

Solidago velutina DC. ssp. californica (Nutt.) Semple
Sonchus asper (L.) Hill*

Sonchus oleraceus L.*

Stephanomeria diegensis Gottlieb
Stephanomeria exigua Nutt.

Stephanomeria virgata Benth.

Symphyotrichum subulatum (Michx.) G.L. Nesom
Taraxacum officinale F.H. Wigg.*

Tragopogon porrifolius L.*

Xanthium spinosum L.

Xanthium strumarium L.

Betulaceae

Alnus rhombifolia Nutt.

Boraginaceae

Amsinkia intermedia Fisch. & C.A. Mey
Amsinkia spectabilis Fisch. & C.A. Mey
Cryptantha clevelandii Greene

Cryptantha clevelandii Greene var. florosa I.M. Johnst.
Cryptantha intermedia (A. Gray) Greene
Cryptantha microstachys (A.Gray) Greene
Heliotropium curassavicum L. var. oculatum (A. Heller)
Tidestr.

Phacelia parry i Torr.

Phacelia ramosissima Lehm.

Continued.

smooth cat's ear

Henrickson 1991

R

coastal goldenbush

FWM

R

poverty weed

FWM

R

fleshy jaumea, marsh
jaumea

POM,UC

1899

prickly lettuce

POM

1902

bitter lettuce

Henrickson 1991

R

Coulter's horseweed

POM

1980

California goldfields

POM,RSA

1901

crowned or royal
goldfields

JEPS,POM

1901

Coulter's goldfields

POM

1903

California cottonrose

POM

1901

cliff aster

Gustafson 1981,
Henrickson 1991

R

pineapple weed

POM

1981

saltmarsh fleabane

UC

1902

two-color rabbit-
tobacco

POM

1901

ladies' tobacco

POM,UCR

1981

cudweed

POM

1981

Wright's cudweed

POM,UC

1902

pink cudweed

POM

1981

cottonbatting plant

POM

1981

California ragwort

POM

1901

common groundsel

RSA

1930

milk thistle

RSA

1981

California goldenrod

Henrickson 1991

R

prickly sow thistle

RSA

1981

common sow thistle

RSA

1981

wreathplant

RSA

1980

small wirelettuce

Gustafson 1981

R

twiggy wreath plant

RSA

1981

annual saltmarsh aster

POM,UC

1901

common dandelion

Henrickson 1991

R

purple salsify

UC

1897

spiny cocklebur

RSA

1981

cocklebur

UC

1902

Birch Family

white alder

FWM

R

Borage Family

common fiddleneck

POM

1901

seaside amsinkia

RSA

1951

cryptantha

UCR

2011

coastal cryptantha

POM

1901

common cryptantha

POM

1901

Tejon cryptantha

POM

1901

seaside heliotrope

POM

1930

Parry's phacelia

Henrickson 1991

R

branching phacelia

POM, RSA

1901

FLORA OF BALLONA WETLANDS

155

Table 1.

Brassicaceae

Brassica nigra (L.) W.D.J. Koch*

Brassica rapa L.*

Cakile maritima Scop.*

Caulanthus lasiophyllus (Flook. & Am.) Payson
Descurainia pinnata (Walter) Britton ssp. brachycarpa
(Richardson) Detling

Descurainia pinnata (Walter) Britton ssp. menziesii (D.C.)
Detling

Erysimum sujfrutescens (Abrams) Rossbach

Hirschfeldia incana (L.) Lagr.-Fossat*

Lepidium didymum L.*

Lepidium lasiocarpum Nutt.

Lepidium latipes Hook.

Lepidium latifolium L.*

Lepidium nitidum Nutt.

Lepidium pinnatifidum Ledeb.*

Lepidium virginicum L. ssp. menziesii (DC.) Thell.

Lobularia maritima (L.) Desv. *

Raphanus sativus L.*

Sinapis alba L.*

Sisymbrium altissimum L.*

Sisymbrium irio L.*

Tropidocarpum gracile Hook.

Cactaceae

Opuntia littoralis (Engelm.) Cockerell

Caryophyllaceae

Polycarpon depressum Nutt.

Polycarpon tetraphyllum (L.) L. var. tetraphyllum*

Silene gallica L.*

Spergula arvenis L. *

Spergularia bocconi (Scheele) Graebn. *

Spergularia macrotheca (Cham. & Schltdl.) Heynh.
Spergularia marina (L.) Besser
Spergularia villosa (Pers.) Cambess.*

Chenopodiaceae

Arthrocnemum subterminale (Parish) Standi

Atriplex argentea Nutt. var. expansa (S. Watson) S.L. Welsh
& Reveal

Atriplex californica Moq.

Atriplex lentiformis (Torr.) S. Watson
Atriplex lentiformis (Torr.) S. Watson subsp. lentiformis
Atriplex leucophylla (Moq.) D. Dietr
Atriplex nummularia Lindl.*

Continued.

Mustard Family

black mustard

Gustafson 1981,
Henrickson 1991

R

field mustard

RSA

1930

sea-rocket

Gustafson 1981,
Henrickson 1991

R

California mustard

DS

1899

tansy mustard

DS

1901

Menzies' tansy mustard

POM

1901

suffrutescent

wallflower

DS,POM,RSA

1901

summer mustard

Gustafson 1981,
Henrickson 1991

R

lesser swine cress

Gustafson 1981,
Henrickson 1991

R

shaggyfruit pepperweed

DS,POM

1901

dwarf peppergrass

DS

1904

perennial pepperweed

Henrickson 1991

R

peppergrass

RSA

1954

featherleaf pepperweed

RSA

1981

wild peppergrass

RSA

1930

sweet alyssum

Gustafson 1981,
Henrickson 1991

R

radish

UCR

1996

white mustard

Gustafson 1981,
Henrickson 1991

R

tumble mustard

Henrickson 1991

R

London rocket

Gustafson 1981,
Henrickson 1991

R

slender keel fruit

UC

1897

Cactus Family

coast prickly-pear

Henrickson 1991

R

Pink Family

California allseed

POM

1901

four-leaved all seed

RSA, UCR

1981

small-flower catchfly

RSA, UCR

1980

stickwort, startwort

RSA

1981

Boccone's sand-spurrey

Henrickson 1991

R

sticky sand-spurrey

POM, RSA

1901

saltmarsh sand-spurrey

CDA,POM

1901

hairy sand-spurrey

Henrickson 1991

R

Goosefoot Family

pickleweed

POM, RSA, UC

1902

silverscale

UC

1902

California orach

RSA

1981

big saltbush

RSA

1981

big saltbush

UC

1902

beach saltbush

RSA,UC

1899

bluegreen saltbush

Henrickson 1991

R

156

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 1.

Atriplex patula L.

A triplex prostrata DC. *

Atriplex rosea L.*

Atriplex semibaccata R. Br.*

Atriplex suberecta I. Verd.*

Bassia hyssopifolia (Pall.) Kuntze*

Beta vulgaris L.*

Chenopodium album L.*

Chenopodium berlandieri Moq.

Chenopodium murale L.*

Dysphania ambrosioides (L.) Mosyakin & Clemants*
Dysphania pumilio (R.Br.) Mosyakin & Clemants*
Salicornia pacifica Standi
Salsola tragus L.*

Suaeda calceoliformis (Hook.) Moq
Suaeda calif ornica S. Watson
Suaeda taxifolia (Standi.) Standi.

Cleomaceae

Peritoma arborea (Nutt.) H.H. litis
Convolvulaceae

Calystegia macrostegia (Greene) Brummit ssp. cyclostegia

(House) Brummit

Calystegia soldanella (L.) R.Br.

Convolvulus arvensis L.*

Cressa truxillensis Kunth
Cuscuta californica Hook. & Am.

Cuscuta indecora Choisy var. indecora
Cuscuta campestris Yunck
Cuscuta salina Engelm
Dichondra occidentalis House
Crassulaceae

Crassula connata (Ruiz & Pav.) A. Berger
Dudleya lanceolata (Nutt.) Britton & Rose

Cucurbitaceae

Citrullus lanatus (Thunb.) Matsum. & Nakai var. citroides
(L.H. Bailey) Mansf.*

Cucurbita foetidissima Kunth.

Euphorbiaceae

Chamaesyce albomarginata (Torr. & A. Gray) Small
Chamaesyce maculata (L.) Small*

Chamaesyce polycarpa (Benth.) Millsp.

Chamaesyce serpens (Kunth) Small*

Croton calif ornicus Mull. Arg.

Croton setigerus Hook.

Euphorbia peplus L.*

Euphorbia terracina L.*

Euphorbia virgata Waldst. & Kit.*

Ricinus communis L.*

Continued.

spear orach

RSA

1980

fat-hen

RSA

1980

tumbling orach

RSA,UC

1902

Australian saltbush

RSA,UC

1902

sprawling saltbush

RSA

1981

five hom bassia

RSA

1980

CDA,POM,RSA,U

common beet

C

1902

lamb's quarters

RSA

1981

pitseed goosefoot

RSA

1980

nettle leaf goosefoot

CDA,RSA

1930

Mexican tea

RSA,UC

1902

Tasmanian goosefoot

Henrickson 1991

R

pickleweed

RSA,UC

1902

Russian thistle

RSA

1980

Gustafson 1981,

homed seablite

Henrickson 1991

R

California seablite

Gustafson 1981

R

woolly sea-blite

Spiderflower Family

POM, RSA

1901

bladderpod

Morning-Glory

Family

DS,POM

1899

coast morning glory

RSA

1981

beach morning glory

SFV

1994

orchard morning glory

RSA

1930

alkali weed

POM,RSA,UCR

1901

chaparral dodder

RSA

1981

large-seeded dodder

Henrickson 1991

R

field dodder

RSA

1981

salt dodder

RSA,UC

1902

western dichondra

Henrickson 1991

R

Stonecrop Family

pygmy-weed

POM, RSA

1901

lance leaf liveforever

Gourd Family

POM

1901

wild watermelon

RSA

1981

calabazilla, buffalo
gourd

RSA

1980

Spurge Family

rattlesnake weed

RSA

1981

spotted spurge

RSA

1981

small seeded spurge

RSA

1981

creeping spurge

RSA

1980

California croton

POM, RSA

1899

turkey-mullein

RSA

1930

petty spurge

RSA

1981

carnation weed

RSA,UCR

1996

leafy spurge

Henrickson 1991

R

castor bean

RSA,UCR

1981

FLORA OF BALLONA WETLANDS

157

Table 1 . Continued.

Fabaceae

Acacia baileyana F. Muell.*

Acacia cyclops G. Don*

Acacia dealbata Link*

Acacia longifolia (Andrews) Willd.*

Acacia neriifolia A. Cunn. ex Benth*

Acmispon americanus (Nutt.) Rydb. var. americanus

Acmispon glaber (Vogel) Brouillet
Acmispon heermannii (Durand & Hilg.) Brouillet
Acmispon strigosus (Nutt.) Brouillet

Albizia distachya J.F. Macbr.*

Albizia lophantha (Willd.) Benth.*

Astragalus pycnostachyus A. Gray var. lanosissimus (Rydb.)
Munz & McBumey

Astragalus tener A. Gray var. titi (Eastw.) Bameby

Astragalus trichopodus (Nutt.) A. Gray

Astragalus trichopodus (Nutt.) A. Gray var. lonchus (M.E.

Jones) Bameby

Hoffmanseggia glauca (Ortega) Eifert
Lupinus bicolor Lindl.

Lupinus chamissonis Eschsch.

Lupinus excubitus M.E. Jones var. hallii
Lupinus longifolius (S. Watson) Abrams
Lupinus succulentus K. Koch
Lupinus truncatus Nutt.

Medicago polymorpha L.*

Melilotus albus Medik.*

Melilotus indicus (L.) All.*

Phaseolus limensis Macfad.*

Spartium junceum L.*

Frankeniaceae

Frankenia salina (Molina) I.M. Johnst.

Geraniaceae

Erodium botrys (Cav.) Bertol.*

Erodium cicutarium (L.) Aiton*

Erodium moschatum (L.) Aiton*

Grossulariaceae
Ribes malvaceum Sm.

Hamamelidaceae
Liquidambar styraciflua L.*

Juglandaceae

Juglans californica S. Watson
Juglans regia L.*

Legume Family

Cootamundra wattle

RSA

1981

western coastal wattle

RSA,UCR

1996

silver wattle

Gustafson 1981,
Henrickson 1991

R

Sydney golden wattle

Henrickson 1991

R

black wattle

Henrickson 1991

R

American bird's foot
trefoil

RSA

1980

deerweed, California
broom

RSA

1980

Heermann's lotus

UC

1902

strigos lotus

RSA

1981

Australian albizia

Gustafson 1981,
Henrickson 1991

R

plume acacia

CDA,RSA

1981

Ventura marsh milk
vetch

NY,POM,RSA,UC

1882

coastal dunes milk
vetch

DS

1891

Santa Barbara
milkvetch

Henrickson 1991

R

Santa Barbara
milkvetch

DS

1901

pig-nut, hog potato

Henrickson 1991

R

miniature lupine

POM

1981

beach blue lupine, dune

POM, RSA

1917

lupine

grape soda lupine

RSA,UCR

1980

longleaf bush lupine

POM

1901

arroyo lupine

RSA

1981

blunt leaved lupine

POM,RSA

1899

California burclover

POM

1981

white sweetclover

POM

1980

sourclover

POM,UCD,UCR

1978

lima bean

POM

1981

Spanish broom

Henrickson 1991

R

Frankenia Family

alkali heath

POM,RSA

1917

Geranium Family

broad leaf filaree

RSA,UCR

1981

redstem filaree

Gustafson 1981,
Henrickson 1991

R

greenstem filaree

UCR

1996

Gooseberry Family

chaparral currant

RSA

1981

Witch-Hazel Family

sweet-gum

Henrickson 1991

R

Walnut Family

southern California
black walnut

FWM

R

English walnut

Henrickson 1991

R

158

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 1.

Lamiaceae

Lycopus americanus W.P.C. Barton
Marrubium vulgar e L.*

Mentha x piperita L.*

Mentha x smithiana R.A. Graham*

Salvia apiana Jeps.

Salvia carduacea Benth.

Salvia columbariae Benth.

Salvia leucophylla Greene
Stachys ajugoides Benth.

Stachys albens A. Gray

Stachys bergii G.A. Mulligan & D.B.Munro

Lythraceae

Lythrum californicum Torr. & A. Gray
Ly thrum hyssopifolia L.*

Malvaceae

Malacothamnus fasciculatus (Torr. & A. Gray) Greene
Malva nicaeensis All.*

Malva parviflora L.*

Malva pseudolavatera Webb & Berthel.*

Malvella leprosa (Ortega) Krapov.

Montiaceae

Calandrinia ciliata (Ruiz & Pav.) DC.

Myrsinaceae

Anagallis arvensis L.*

Myrtaceae

Eucalyptus camaldulensis Dehnh.*

Eucalyptus globulus Labill.*

Eucalyptus tereticornis Sm.*

Eucalyptus viminalis Labill.*

Nyctaginaceae

Abronia maritima S. Watson

Abronia umbellata Lam.

Oleaceae

Fraxinus latifolia Benth.

Fraxinus velutina Torr.

Onagraceae

Camissoniopsis bistorta (Torr. & A. Gray) W.L. Wagner &
Hoch

Camissoniopsis cheiranthifolia (Spreng.) W.L. Wagner &
Hoch ssp. suffruticosa

Camissoniopsis lewisii (P.H. Raven) W.L. Wagner & Hoch
Camissoniopsis micrantha (Spreng.) W.L. Wagner & Hoch
Epilobium campestre (Jeps.) Hoch & W.L. Wagner
Epilobium ciliatum Raf.

Oenothera elata Kunth.

Continued.

Mint Family

bugleweed

UC

1902

horehound

RSA

1981

peppermint

UC

1902

redstem mint

POM

1902

white sage

FWM

R

thistle sage

POM,UCR

1901

chia

POM,UCR

1901

purple sage

FWM

R

hedge-nettle

POM,UC

1917

white hedge-nettle

POM, RSA, UC

1902

Berg's hedge-nettle

UC

1932

Loosestrife Family

California loosestrife

POM,RSA,UC

1902

hyssop loosestrife

RSA,UCR

1980

Mallow Family

chaparral mallow

RSA

1981

bull mallow

POM

1902

cheeseweed

RSA,UC

1932

Cretan mallow

RSA,UCR

1981

alkali-mallow

POM,RSA,UCR

1917

Miner's Lettuce

Family

red maids

UC

1897

Myrsine Family

scarlet pimpernel

RSA

19J0

Myrtle Family

red gum, river red gum

RSA

1981

blue gum

Henrickson 1991

R

forest red gum

Gustafson 1981

R

manna gum

RSA

1981

Four O' Clock Family

red sand- verbena

POM

1901

beach sand-verbena

POM, RSA

1901

Olive Family

Oregon ash

UC

1932

velvet ash

RSA

1981

Evening-Primrose

Family

California sun cup

POM, RSA, UC

1897

beach evening-primrose

POM,RSA,UCD

1901

Lewis' evening

primrose

POM

1899

miniature suncup

RSA

1981

smooth boisduvalia

POM

1901

slender willow herb

RSA

1930

evening primrose

RSA

1981

FLORA OF BALLONA WETLANDS

Table 1.

Orobanchaeae

Castilleja exserta (A. Heller) T.I. Chuang & Heckard ssp.
exserta

Chloropyron maritimum (Benth.) A. Heller ssp. maritimum'

Oxalidaceae

Oxalis pilosa Nutt.

Oxalis pes-caprae L.*

Papaveraceae

Eschscholzia californica Cham.

Papaver heterophyllum (Benth.) Greene

Plantaginaceae

Antirrhinum coulterianum A. DC.

Antirrhinum nuttallianum A. DC. ssp. nuttalianum
Plantago erecta E. Morris
Plantago lanceolata L.*

Plantago major L.*

Platanaceae
Platanus racemosa Nutt.

Polygonaceae

Chorizanthe parryi Wats, var . fernandina (Wats.) Jeps.
Eriogonum fasciculatum Benth.

Eriogonum fasciculatum Benth. var. fasciculatum
Eriogonum gracile Benth.

Eriogonum parvifolium Sm.

Lastarriaea coriacea (Goodman) Hoover
Mucronea californica Benth.

Persicaria hydropiperoides (Michx.) Small
Persicaria lapathifolia (L.) Gray
Persicaria maculosa Gray*

Polygonum aviculare L.*

Rumex conglomeratus Murray*

Rumex crispus L.*

Rumex fueginus Phil.

Rumex salicifolius Weinm.

Ranunculaceae
Clematis ligusticifolia Nutt.

Delphinium parryi A. Gray

Delphinium parryi A. Gray ssp. maritimum (Davidson) M. J.
Wamock

Rosaceae

Heteromeles arbutifolia (Lindl.) M. Roem.

Horkelia cuneata Lindl. var. sericea A. Gray
Potentilla anserina L. ssp. pacifica (Howell) Rousi
Potentilla multijuga Lehm.

Pyracantha cf. crenulata (D. Don) M. Roem.*

Rosa californica Cham. & Schltdl.

Rubus ursinus Cham. & Schltdl.

Continued.

Broomrape Family

purple owl's-clover

POM,RSA

1901

salt marsh bird's beak

DS,POM

1901

Oxalis Family

hairy wood sorrel

Henrickson 1991

R

Bermuda buttercup

Gustafson 1981,
Henrickson 1991

R

Poppy Family

California poppy

Henrickson 1991

R

wind poppy

RSA

1899

Plantain Family

Coulter’s snapdragon

POM

1911

Nuttall's snapdragon

UC

1902

California plantain

POM

1899

English plantain

POM, RSA

1902

common plantain

RSA

1980

Sycamore Family

western sycamore

FWM

R

Buckwheat Family

San Fernando Valley
spineflower

DS

1901

California buckwheat

RSA

1981

coastal California

Gustafson 1981,

buckwheat

Henrickson 1991

R

slender buckwheat

POM,RSA

1901

seacliff wild buckwheat

RSA,UC

1902

leather-spineflower

POM

1901

California spineflower

DS,POM

1899

false waterpepper

POM,UC

1901

willow weed

RSA

1930

lady's thumb

RSA

1980

knotweed, knotgrass

CDA,RSA

1930

clustered dock

RSA

1930

curly dock

RSA

1930

golden dock

POM,RSA

1902

willow dock

RSA

1981

Buttercup Family

western virgin's bower

RSA

1981

larkspur

DS

1901

maritime larkspur

DS

1901

Rose Family

toyon

Henrickson 1991

R

Kellogg's horkelia

Abrams 1902

R

Pacific silverweed

POM,UC,UCR

1900

Ballona cinquefoil

POM

1890

Himalayan firethom

Henrickson 1991

R

California rose

Henrickson 1991

R

California blackberry

FWM

R

160

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 1 . Continued.

Rubiaceae

Galium angustifolium A. Gray

Salicaceae

Populus fremontii S. Watson ssp .fremontii
Salix exigua Nutt. var. exigua
Salix goodingii C.R. Ball
Salix laevigata Bebb
Salix lasiolepis Benth.

Scrophulariaceae
Myoporum laetum G. Forst.*

Scrophularia californica Cham. & Schltdl.

Verb as cum virgatum Stokes*

Solanaceae

Datura wrightii Regel

Lycium ferocissimum Miers*

Lycopersicum esculentum Mill.*

Nicotiana clevelandii A. Gray
Nicotiana glauca Graham*

Petunia parviflora Juss.

Solanum americanum Miller
Solanum douglasii Dunal

Solanum nigrum L.*

Solanum physalifolium Rusby var. nitidibaccatum (Bitter)
Edmonds*

Solanum xanti A. Gray

Tamariscaceae

Tamarix ramosissima Ledeb.*

Ulmaceae

Ulmus parvifolia Jacq.*

Urticaceae

Urtica dioica L. ssp. holosericea (Nutt.) Thome
Urtica urens L.*

Verbenaceae

Phyla lanceolata (Michx.) Greene
Verbena bracteata Lag. & Rodr.

Verbena lasiostachys Link

Verbena lasiostachys Link var. lasiostachys

Zygophyllaceae

Tribulus terrestris L.*

MONOCOTS

Agavaceae

Hesperoyucca whipplei (Torr.) Trel.

Alismataceae

Sagittaria montevidensis Cham. & Schltdl ssp. calycina
(Engelm.)

Amaryllidaceae
Narcissus tazetta L.*

Madder Family
narrowly leaved
bedstraw

Willow Family

POM

1980

Fremont cottonwood

POM

1981

narrow-leaved willow

FWM

R

Gooding's black willow

FWM

R

red willow

DS,POM

1901

arroyo willow

Figwort Family

DS,POM,UC

1932

myopomm

RSA

1981

California figwort

RSA

1930

wand mullein

RSA

1981

Nightshade Family

Jimson weed

RSA

1980

African boxthom

CD A, RSA

1978

tomato

RSA

1981

Cleveland's tobacco

POM, RSA

1901

tree tobacco

RSA

1980

wild petunia
American black

RSA

1981

nightshade

RSA

1981

Douglas' nightshade

POM,RSA,UCR
Gustafson 1981,

1901

black nightshade

Henrickson 1991

R

nightshade

RSA

1981

chaparral nightshade

Tamarisk Family

Henrickson 1991

R

saltcedar

Henrickson 1991

R

Elm Family

Chinese elm, lacebark
elm

Henrickson 1991

R

Nettle Family

hoary nettle

POM, RSA

1902

dwarf nettle

RSA

1981

Vervain Family
lance leaf lippia

POM,UC

1902

bracted verbena

POM

1902

western verbena

RSA

1981

western vervain

RSA

1930

Caltrop Family

puncture vine

RSA

1980

Century Plant Family
chaparral yucca
Water-Plantain
Family

FWM

R

arrowhead

RSA

1981

Amaryllus Family

paper white

RSA

1981

FLORA OF BALLONA WETLANDS

161

Table 1 . Continued.

Araceae

Lemna minuta Kunth
Arecaeae

Phoenix canariensis Chabaud*

Phoenix dactylifera L.*

Washingtonia robusta H. Wendl.*

Cyperaceae

Bolboschoenus maritimus (L.) Palla
Bolboschoenus maritimus (L.) Palla ssp. paludosus (A.
Nelson) T. Koyama

Bolboschoenus robustus (Pursh) Sojak

Carex praegracilis W. Boott
Cyperus eragrostis Lam.

Cyperus esculentus L.

Cyperus involucratus Rottb.*

Eleocharis macrostachya Britton
Eleocharis montevidensis Kunth

Schoenoplectus americanus (Pers.) Schinz & R. Keller
Schoenoplectus californicus (C.A. Mey.) Sojak

Iridaceae

Iris pseudacorus L.*

Sisyrinchium bellum S. Watson

Juncaceae

Juncus acutus L.

Juncus balticus Willd. subsp. ater (Rydb.) Snogerup
Juncus bufonius L.

Juncus mexicanus Willd.

Juncus textilis Buchenau

Poaceae

Agrostis stolonifera L.*

Arundo donax L.*

Avena barbata Link*

Avena fatua L.*

Brachypodium distachyon (L.) P. Beauv.*

Bromus carinatus Hook. & Am. var. marginatus (Steud.)

Barkworth & Anderton

Bromus catharticus Vahl var. catharticus*

Bromus diandrus Roth*

Bromus hordeaceus L. *

Bromus madritensis L. ssp. rubens (L.) Husn.*

Cortaderia jubata (Lem.) StapP

Cortaderia selloana (Schult. & Schult. f.) Asch. & Graebn. *
Cynodon dactylon (L.) Pers.*

Digitaria sanguinalis (L.) Scop.*

Distichlis littoralis (Engelm.) H.L. Bell & Columbus
Distichlis spicata (L.) Greene

Arum Family

duckweed

RSA

1930

Palm Family

Canary Island palm

RSA

1981

date palm

MEC 2005

R

fan palm

Henrickson 1991

R

Sedge Family
saltmarsh bulmsh

DS,RSA

1904

saltmarsh bulmsh

RSA,UCR

1981

seacoast bulmsh

Gustafson 1981,
Henrickson 1991

R

black creeper, freeway
sedge

CAS, DS, POM

1889

tall flatsedge

RSA,SD,UCR

1980

yellow nutgrass

RSA

1981

umbrella sedge

Gustafson 1981,
Henrickson 1991

R

creeping spike rush

RSA

1930

sand spikerush

DS,RSA

1904

Olney's three-square
bulmsh

DS,RSA

1900

southern bulmsh

DS,RSA

1901

Iris Family

horticultural iris

RSA

1981

western blue-eyed-
grass

FWM

R

Rush Family

spiny rush

DS

1901

Baltic rush

Gustafson 1981,
Henrickson 1991

R

toad msh

RSA

1981

Mexican msh

RSA

1981

mat or basket msh

RSA

1930

Grass Family

creeping bent

Gustafson 1981,
Henrickson 1991

R

giant reed

RSA

1981

slender wild oat

Henrickson 1991

R

wild oat

RSA

1981

false brome

Henrickson 1991

R

mountain brome

Gustafson 1981

R

rescue grass

POM

1981

ripgut grass

POM

1930

soft chess

POM

1981

red brome

POM

1901

purple pampas grass

Gustafson 1981,
Henrickson 1991

R

Uruguayan pampas
grass

RSA

1980

Bermuda grass

RSA

1930

hairy crab grass

POM

1981

shore grass

Henrickson 1991

R

salt grass

UC

1902

162

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 1 . Continued.

Echinochloa crus-galli (L.) P. Beauv.*

Ehrharta erecta Lam.*

Elymus condensatus J. Presl.

Elymus glaucus Buckley

Elymus trachycaulus (Link) Shinn, ssp. trachycaulus
Elymus triticoides Buckley
Festuca arundinacea Schreb.*

Festuca myuros L.*

Festuca perennis (L.) Columbus & J.P. Sm.*

Festuca tementula (L.) Columbus & J.P. Sm.*

Hainardia cylindrica (Willd.) Greuter*

Hordeum brachyantherum Nevski

Hordeum murinum L. ssp. leporinum (Link) Arcang.*

Hordeum intercedens Nevski

Hordeum vulgar e L.*

Koelaria macrantha (Ledeb.) Schult.

Lamarckia aurea (L.) Moench*

Leptochloa fusca (L.) Kunth ssp. uninervia (J. Presl) N.

Snow

Melica imperfecta Trin.

Parapholis incurva (L.) C.E. Hubb.*

Paspalum dilatatum Poir.*

Pennisetum setaceum (Forssk.) Chiov.*

Phalaris minor Retz.*

Phalaris paradoxa L.*

Poa annua L.*

Polypogon monspeliensis (L.) Desf.*

Polypogon viridis (Gouan) Breistr.*

Schismus barbatus (L.) Thell.*

Setaria parviflora (Poir.) Kerguelen
Sorghum halepense (L.) Pers.*

Stenotaphrum secundatum (Walter) Kuntze*

Stipa cernua Stebbins & Love
Stipa lepida Hitchc.

Stipa miliacea (L.) Hoover var. miliacea *

Stipa pulchra Hitchc.

Ruppiaceae

Ruppia maritima L.

Themidaceae

Dichelostemma capitatum (Benth.) Alph. Wood
Typhaceae

Sparganium erectum L. ssp. stoloniferum (Grabner) C. Cook

& M.S. Nicholls

Typha domingensis Pers.

Typha latifolia L.

barnyard grass

POM

1981

panic veldt grass

POM,UCR

1981

giant wild-rye

Henrickson 1991

R

western wild-rye

POM

1901

slender wheatgrass

Abrams 1902

R

beardless wild rye

POM

1901

tall fescue

Henrickson 1991

R

rattail sixweeks grass

POM

1913

rye grass

POM,RSA,UCR

1902

Darnel ryegrass

POM

1901

barb grass

POM,UCD

1901

meadow barley

POM

1930

hare barley

POM

1981

bobtail barley

POM

Gustafson 1981,

1901

barley

Henrickson 1991

R

June grass

POM

1901

goldentop

Henrickson 1991

R

Mexican sprangletop

POM

1981

little California melica

POM

1901

sickle grass

POM

1981

dallis grass

POM

1980

crimson fountain grass
Mediterranean canary

Henrickson 1991

R

grass

UC

1932

Hood canary grass

POM

1981

annual blue grass

RSA

Gustafson 1981,

1981

annual beard grass

Henrickson 1991

R

water beard grass

RSA

Gustafson 1981,

1930

Mediterranean grass

Henrickson 1991

R

knotroot bristle grass

RSA

1981

Johnson grass

UC

1902

Saint Augustine grass

RSA

1981

nodding needle grass

Henrickson 1991

R

foothill needle grass

FWM

R

smilo grass

POM

1981

purple needle grass

Ditch-Grass Family

FWM

R

ditch-grass

Brodiaea Family

RSA

1980

blue dicks

Cattail Family

Henrickson 1991

R

bur reed

RSA,UC

1902

southern cattail

RSA

1981

broad-leaved cattail

RSA

1980

Literature Cited

Abrams, L.A. 1902. Additions to the flora of Los Angeles County, please change to Bull. S. Calif. Acad. Sci., 1
(7): 87.

Abrams, L.A. 1917. Flora of Los Angeles and Vicinity. New Era Printing Company, Lancaster, Pennsylvania.
Altschul, J.H., J.A. Homburg, and R.S. Ciolek-Torrello. 1992. Life in the Ballona: Archaeological Investigations
at the Admiralty Site (CA-LAN-47) and the Channel Gateway Site (CA-LAN-1596-H). Statistical
Research, Inc. Technical Series No. 33.

FLORA OF BALLONA WETLANDS

163

Altschul, J.H., A.Q. Stoll, D.R. Grenda, and R. Ciolek-Torrello. 2003. At the Base of the Bluff: Archaeological
Inventory and Evaluation along Lower Centinela Creek, Marina del Rey, California. Statistical Research,
Inc. Playa Vista Monograph Series Test Excavation Report 4.

Baldwin, B.G., D.H. Goldman, D.J. Keil, R. Patterson, T.J. Rosatti, and D.H. Wilken, editors. 2012. The Jepson
manual: vascular plants of California, second edition. University of California Press, Berkeley.

Clark, J. 1979. Ballona Vegetation Survey. Report dated May 10, 1979, prepared for The Conservation
Foundation.

Dark, S., E.D. Stein, D. Bram, J. Monteferante, T. Longcore, R. Grossinger, and E. Beller. 2011. Historical Ecol-
ogy of the Ballona Creek Watershed. Southern California Coastal Water Research Project Technical
Report #67 1 .

Grossinger, R.M., E.D. Stein, K.N. Cayce, R.A. Askevold, S. Dark, and A.A. Whipple. 2011. Historical Wetlands
of the Southern California Coast: An Atlas of US Coast Survey T-Sheets, 1851-1889. San Francisco Estu-
ary Institute Contribution # 586, Southern California Coastal Water Research Project Technical Report #
589. data download for T-Sheet #T 1432b from caltsheets.org/socal/index.html.

Gustafson, R.J. 1981. The Vegetation of Ballona. Undated report prepared for Playa Vista.

Henrickson, J. 1991. Botanical Resources of Playa Vista. Report dated May 12, 1991, prepared for Playa Vista.

Jensen, S.A. 1933. Vegetation Map of the Redondo 15-Minute Quadrangle. Downloaded from http://vtm.
berkeley.edu.

Johnston, K., 2011. The Ballona Wetlands Ecological Reserve Baseline Assessment Program 2009-2010 Final
Report. Prepared for the California State Coastal Conservancy and California Department of Fish and
Game. Issued November, 2011.

MEC Analytical Systems, 2005. 2004 Biological Study of Ballona Wetlands, Los Angeles County, California.
Final Report, prepared for US Army Corps of Engineers Los Angeles District.

Munz, P.A. 1974. A Flora of Southern California. University of California Press, Berkeley, California.

Read, E. 1995. Sensitive Plant Surveys and Vegetation Update for Playa Vista. Report prepared for Impact
Sciences, Inc.

CONTENTS

The Physical Characteristics of Nearshore Rocky Reefs in The Southern California
Bight. Daniel J. Pondella, II, Jonathan Williams, Jeremy Claisse, Becky
Schaffner, Kerry Ritter, and Ken Schiff _ .r„ 1 05

Comparison of the Marine Wood Borer Populations in Los Angeles Harbor in
1950-1951 with the Populations in 2013-2014. Donald J. Reish, Thomas V
Gerlinger, and Robert R. Ware 123

Evidence for Negative Effects of Drought on Baetis sp. (Small Minnow Mayfly)
Abundance in a Southern California Stream. Elizabeth Montgomery, Rosi
Dagit, Crystal Garcia, Jenna Krug, Krista Adamek, Sandra Albers, and
Katherine Pease . ..... 1 29

The Bigeye Scad, Selar crumenophthalmus (Bloch, 1793) (Family Carangidae),

New to the California Marine Fauna, with a List to and Keys for All California
Carangids. Milton S. Love, Julianne Kalman Passarelli, Chris Okamoto, and

Dario W. Diehl . .. ... 1 4 1

A Flora of the Ballona Wetlands and Environs. E.A. Read 149

Cover: Ballona Wetlands, photo credit to Lisa Fimiani.