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BULLETIN OF MARINE SCIENCE, 73(3): 545-556,2003
EFFECT OF CLIMATIC CHANGE ON THE HARVEST OF TIEE KELP
MACROCYS77SPYRIFERA ON THE MEXCAN PACJFIC COAST
MargaritaCasas Valdez, Elisa Serviere Zaragoza, Daniel Lluch Belda,
Roberto Marcos andRuth Aguila Ramirez
ABSTRACT
The effect of sea temperature, sea level, upwelling, and wind speed on the Macrocystis
pyrifem harvest was evaluated using historical harvest data for the western coast of the
Baja California Peninsula, Mexico. The relationship between the environmental vari-
ables and harvest per unit effort (CPUE) was determined through correlation analysis for
the period from 1956-1998. The effects of temperature, upwelling, and sea level were
inverse, whereas the effect of wind speed was not significant. Multiple regression analy-
sis showed that, of the variables studied, temperature best explained variations in
Macrocystisharvest.
The giant kelp Macrocystispyrifera (Linnaeus) C.Agardh isamember of the large
brown algae (Phaeophyta) that are a conspicuous part of the marine environment. It is a
subtidal species that attaches to solid or soft substrate, produces holdfasts that are either
conical or low mounds. Because of the great size of individual plants, their trunk-like
appearance in the water column, and the extensive surface canopies along the coastline,
these subtidal habitats have been called forests (Foster and Schiel, 1985). Subtidal forests
of M pyrifera occur in many areas of the world, but is most widely distributed in the
southern hemisphere. In the northern hemisphere, M. pynifera commonly occurs from
near Santa Cruz (central California) to Baja California (Foster and Schiel, 1985). Giant
kelp is highly productive.
Macrocystis has been harvested since 1956 along the Pacific coast of Baja California
and exported to the United States for the production of alginates. Recently, it has been
sold in Mexico as meal and f6r abalone aquaculture (McBride, 1998). It is harvested by
specially designed ships that cut the algae at an approximate depth of 1.2 m and transport
it. Two ships participated in the harvest at the area of interest: EL CAPITAN, from 1956-
1966, (storage capacity of 300 t), and EL SARGACERO from 1967 to present, (storage ca-
pacity of 400 t). Ship operations were the same at all locations and did not change over
the study period.
In spite of the importance of kelp, studies of variations in abundance or the size of
harvest are few. Casas et al. (1985) and Hernandez et al. (1989a,b; 1991) reported higher
values of surface canopy and standing crop biomass in spring and summer. Corona (1985)
found a higher harvest with less effort in spring and summer from data on kelp beds from
1978-1984. The first objective of this study is to analyze changes in the size ofMpyiifera
harvest, the effort, and the catch per unit effort (CPUE) along the western coast of the
Baja California Peninsula over the period of large-scale harvest, almost 43 yrs.
There is evidence that climatic changes have altered the productivity of marine ecosys-
tems at several trophic levels (Polovina et al., 1994). The existence of causative relation-
ships between the physical environment and biological productivity has been demon-
strated (Polovina et al., 1995). In the California Current System and in other parts of the
world, changes in climate have been shown to cause fluctuations in the number of small
Bulletin ofMarine Science 545
0 2003 Rosenstiel School of Marine and Atmospheric Science
of the University of Miamni
546 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
pelagic fish (Southar and Isaacs, 1974; Lluch et al., 1989,1991,1992; Baumgartner et al.,
1992; MacCall, 1996). Less isknown about the effect of climate on benthic resources. In
California, there is available information about the effect of environmental variables on
the crab fishery (Wild et al., 1983; Bostford, 1986), abalone (Tegner, 1989, 1991; Davis
et al., 1992), and sea urchin and kelp (North and Zimmerman, 1984; Dean and Jacobsen,
1984, 1986; Tegner and Dayton, 1987; Tegner et al., 1996). Tegner et al. (1996) com-
pared data on the maximum canopy of the kelp forests and the size of the annual harvest
of Macrocystisfor Califomia, and they chose harvest size as the most useful data to relate
to environmental variables. They pointed out that harvest size was a reflection of changes
in consumer demand and harvest productivity, as well as natural disturbances. This data
has the advantage of integrating growth over a long period and has less subjectivity in its
measurement.
The second objective of this study is to show how the abundance of M pyiifera, ex-
pressed as catch per unit effort (CPUE) is related to environmental variables, including
sea surface temperature, sea level, upwelling index, and wind speed on the west coast of
Baja California. It was also determinated which environmental factor best explains the
variability in the abundance of Macrocystis.
METHODS AND MATERIALS
In Mexico, M. pyrifera is distributed along the westem coast of the Baja California Peninsula
from the border with the USA to Punta San Roque, Baja California Sur. However, the alga is
harvested from Islas Coronado (3215rN) to Arrecife Sacramento, Baja California (30'30N) at 15
beds: Islas Coronados, Playas de Tijuana, Punta Mezquite, Salsipuedes, San Miguel and Sauzal,
Bahia de Todos Santos, Punta Banda, Bahia Soledad, Santo Tomais, Punta China, Punta San Jos6,
Punta San Isidro, Punta San Telmo, Isla San Martin, and Bahia del Rosario (Fig. 1). This study
encompasses all of these.
KELP HARVEST DATA.-Daily harvest records from 1956-1998 were provided by Products of the
Pacific, S.A. de C.V. These contained the following information: harvest date, name of the bed,
number of trips made by each ship, and harvest size mt (wet weight). In total, data were extracted
from 3278 daily records.
The volume of the harvest per trip made by each ship, called harvest per unit effort (CPUE),
served as an index of abundance of Macrocystis.To adjust for the difference in the storage capacity
of the two ships, the data on harvest volume trip-' was weighted in the following way:
Period Storage capacity (t) Harves volume/trip (t) Weight factor
1967-98 400 364 1
1956-67 300 222 0.61
PHYSlCAL PARAMETERs.-Sea surface temperatures (SST) were obtained from the data base of
Reynolds and Smith (1994). The database contains monthly averages of SST (°C) from 1956-1992
for 20 x 20 ocean quadrants. Records for latitudes 30°-32° N were selected. For the years 1993-
1998, this information is available for 10 x 10 ocean quadrants. For consistency, we averaged the
monthly values from four quadrants. For years 1956-97, annual averages for the middle level of
the sea (MLS) were obtained from the web page of the Sea Level Center of the University of
Hawaii (http:lluslc.soest.Hlawaii.edu/uhslc/data.html). For years 1956-98, the upwelling index (UI)
for latitudes 28.5°-31.5°N was obtained from the upwelling index database of Bakun (1973), which
maintains records for 3° x 3° ocean quadrants. The upwelling index is expressed as m3s-' mr' of
1
g00
CASAS VALDEZETAL: EFFECTS OFCLIMATIC CHANGEON HARVESFMACROCYSTIS 547
Figure 1.Distribution of the harvest beds of Macrocystispyriferaalong the Baja Califomia Peninsula.
coast. For 1956-89, wind speed data (WS) for latitudes 30°-32°N were obtained from the COADS
(Comprehensive Ocean-Atmosphere Data Set) database, which compiles monthly average wind
speeds in ms-' for 20 x 20 ocean quadrants.
The annual averages for cpue, SST, UII, WS, and MLS were calculated. These average values
were used to calculate the annual anomalies for each of the variables using the following equations:
548 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
Ai =xi - x
n
>xi
x= i-I
n
where, A is the anomaly of the fh year, Xi is the ith annual value, x is the average value for the year
i to n, n is the number of years.
Because the SST and MLS series had a long-term trend, this trend was eliminated through a
linear fit. To eliminate the noise in the interannual variability in the graph representation, the series
were smoothed by using running averages (Statsoft, 1999).
The relationships between abundance of M pynifera and the climatic variables were determined
through correlation analysis using the anomalies of variables and harvest volume/trip values (Ander-
son, 1972). To identify the factor that explains most the variation in harvest volume/trip of
Macrocystis,a multiple linear regression analysis was used (Zar, 1984).
RESULTS
In Baja California, kelp forests of M pyrifera have been harvested since 1956 by the
same company. Annual harvest have ranged from a low of 2980 t in 1983, to 44,250 t in
1989, with an average of 25,900 t (Fig. 2A). The history of the Macrocystis harvest re-
veals two major periods: 1959 to 1977, when the harvest increased to 44,000 t, and 1985-
1997, when the harvest fluctuated around 32,800 t. Three large declines in harvest oc-
curred in 1958,1983, and 1998 (Fig. 2A), declines of 87%, 89%, and 79% with regard to
the average value.
SST from 1956-98 had an average value of 1N7.4C, with a minimum of 16.3°C in 1975
and a maximum of 19.3°C in 1998. There were cool periods in 1956-57, 1960-66 and
1969-76. Warming periods occurred in 1958-59, 1981-84, and a very pronounced one
from 1992-98. In this last year, the SST was the warmest in 43 yrs (Figs. 2B,3A). For the
period 1956-97, MLS varied, on average by 2.03 cm, with a minimum of -67.6 cm in
1964 and a maximum of 107.2 cm in 1984. MSL had negative anomalies in 1960-76 and
1988-90. Positive anomalies occurred in 1958-59,1982-84, and 1990-97 (Figs. 2C,3B).
The average value for the UI from 1956-98 was 122 m3 s-', with a minimum of 67 m
3
3s-' in 1971. There were three periods of negative
s7' in 1966 and a maximum of 169 m
anomalies: 1965-69, 1985-87 and 1992-93. Positive anomalies occurred in 1956-59,
1970-75, 1980-84 and 1989-91 (Figs. 2D,3C). From 1958-90, the average wind speed
was 22.2 m3 s', with a minimum of 17.5 m3 ' in 1967 and a maximum of 29.9 m3 s' in
1987. Clear negative anomalies occurred in 1959 and 1966-70. Positive anomalies oc-
curred in 1971-75 and 1982-89 (Figs. 2E,3D).
Correlation analyses indicated an inverse relationship between the size of CPUE
and SST (r2 = -0.46; P < 0.05; Fig. 3A); MLS (r2 = -0.48; P < 0.05; Fig. 3B), and UI
(r2 = -0.35, P < 0.05; Fig. 3C). The relationship between cpue and WS was not
significant (r2 = 0.006, P > 0.05; Fig. 3D). The results obtained by multiple regres-
sion analysis (SST: -0.45; MLS: -0.02; WS: -0.03 with r= 0.46, P < 0.10) indicate
that changes in SST best explain the variation in harvest volume trip-' of M. pyrifera.
CASAS VALDEZ ETAL: EFFECTS OF CLIMATIC CHANGE ON HARVEST MACROCYS7S 549
50000 a) EINiIIo El Niubo
40000
30000 ElNiio(1958)
20000
10000
0O 11 Illl111HIliillillllllillli
IH lllH ill
1956 1960 1964 1968 1972 1976 1900 1984 1988 1992 1996
20
b)
19
17
16
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995
150
c)
100
O 50
*0
0A
~ 50
.100
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995
Figure 2. Data series of a)Harvest volume of Macrocystispyrifera,b)Sea surface temperature, c)
Middle anomaly level sea, (on followingpage) d) Upwelling index, and e) Wind speed.
550 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
180
d)
160 -
?140-
15~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~3
-E 120
100
80
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995 1998
30
28
26
3- 24
~22
20 -
18
16 ,.. .. .. .. .. .. .L. .. .. .. .. ..........
1956 1960 1964 1968 1972 1976 1980 1984 1988
DISCUSSION
The fishery of M pyrifera does not show signals of overexploitation, since it is at a
stage where effort increases correspond to harvest increases and the CPUE has remained
almost constant from the beginning of the collecting period until the present.
The harvest volume per trip is a reasonable indicator of the abundance of Macrocystis
because approximately 60% of the algal biomass is represented by the surface canopy
and about 95% of the production is in the upper I m of the water column, with a maxi-
mum cutting depth of 1.2 m. Furthermore, harvesting techniques and the ships have re-
mained similar over the period of study, so the volume of each harvest is comparable
throughout this study.
The periods of cooling and warming of the SST found during over 43 yrs are consistent
with those found by Smith (1995) for 1940-1983 at the Scripps Institution of Oceanogra-
phy pier in La Jolla, California.
CASAS VALDEZETAL: EFFECTS OF CLIMATIC CHANGEON HARVEST A CROCYSTIS 551
200 -
150 . 1.5
100 I
1
200
-cpue -.- SST
-150 -IO0
1956 1960 1964 1968 1972 1976 1980
13~~~~~~~~~~~~~~~~~~~0 1984 1988 1992 1996
A-100
200 3
C 150
b)
150
100
100
.50
.,....
., --
15
U-'-200
50
*0
~~~~~~
-50~~~~~~~~~~~~~~~~5
-100
-opuc -- vIS --100
-150
1956 1960 1964 1968 1972 1976 1980 1984 1988 1992 1996
Figure 3.Correlation of harvest volume/trip of Macrocystisppriferawith a)Sea surface temperature
and d) Wind
(SST), b) Middle level sea (MLS), (on following page) c) Upwelling index (UI), represents the
speed (WS). The heavy line represents the harvest volume/trip and the thin line
environmental variable.
Positive anomalies in the CPUE between 1964 and 1975 correspond to a decrease in
the SST. The low temperatures during this period coincide with a Regime Change (Mantua
et al., 1997). Beginning in 1976, the SST increased coincidently with a decrease in the
harvest volume of Macrocystis, with both peaking in the mid-1 980s. In 1988-89, during
mommommommom
552 BULLETIN OF MARINE SCIENCE, VOL 73, NO. 3,2003
250 -
150
,C- 50-
3
a
0
E -50-
eJ
-150
-250
)56 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995 1998
250 6
d)
4
150 -
.2-. 50 -
A -2
ti
s -
:h2LYLWDIL-AO
A-- cE
03
to
A
C, -50 -
-150 -
-ccpue--WS
-250 - , ............ ,., ...... -6
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995 1998
a La Nifia cooling period, the maximum CPUE occurred. Finally, since the beginning of
the 1990s, a strong warming trend is apparent, with a concomitant decrease in the CPUE.
North et al. (1993) related the maximum annual canopy area of the algal bed of Califomia
from 1967-89 to the maximum and minimum quarterly temperatures and to the number
of warm and cool days per year. They found that the former explained 34% of the vari-
ability in canopy area and the latter to 46%.
CASAS VALDEZ ETAL: EFFECTS OF CLIMATIC CIIANGE ON liARVEST MACROCYS77S 553
The effects of high temperatures in the Pacific waters of Baja California Peninsula
during the El Nino years were evident in the harvest and CPUE registered in 1958,1983,
and 1998, amounting to under 5000 t yr'. During El Nifio events, changes in both distri-
bution and abundance of M pyriferahave been described at some localities of California
for 1958 and 1983 (Tegner and Dayton, 1987) and for Baja California Peninsula for 1983
and 1998 (Hemrindez, 1988; Ladah et al., 1999). During El Nifio, high temperatures and
low nutrient concentration results in wide-spread mortality of the giant kelp forest (Gerard,
1984; Tegner and Dayton, 1987, 1991; Tegner et al., 1996).
Increases in sea temperature may reduce the Macrocystis population by limiting the
availability of nutrients. A strong negative correlation was noted between temperature
and nitrate availability (Gerard, 1982; Zimmerman andiKremer, 1986). When nitrate con-
centration falls below 1 pM, as it does off the coast of southern California at approxi-
mately 15.5°C (Zimmerman and Kremer, 1984) and off Bahia Asunci6n, Baja California
Sur at 18.5°C (south of the distribution of this species) (HernAndez et al., 2001) growth
rate of Macrocystisdeclines. Large-scale disappearances may be explained because canopy
growth is supported by transfer ofnitrogen from lower parts of the plant that are bathed in
water with higher nitrate concentrations (Jackson, 1977). If the lower parts of the plants
cannot transfer enough nitrogen to the surface, then the canopy dies back and individuals
succumb, probably because of low nitrogen reserves (Gerard, 1984).
In addition to limiting nitrates, and thereby causing Macrocystisto die off, high tem-
peratures also inhibit successful recruitment of new algal cells to the beds. Thus, the
population cannot recover until the temperature decreases (Jackson, 1977). At the south-
ern limit, Ladah et al. (1999) suggest that microscopic stage of M.pyrifera survived the
stressful conditions during the ENSO, possibly in a dormant state.
The decline in the CPUE when MSL was higher than normal may, in fact be caused by
increased SST. Large positive anomalies in the MLS occur during periods in which the
thermocline is at a greater depth, and at the same time, when SST is higher. The pattern of
response of CPUE to changes in MLS is similar to that previously described for tempera-
ture changes.
Upwelling of deep-sea water normally brings to the surface a rich supply of nutrients
that can benefit marine organisms. The surprisingly inverse correlation found between UI
and Macrocystis abundance may be explained as follows. Some periods (1958-59 and
1980-84) with high upwelling values also had elevated SST. A similar phenomenon was
documented by Hayward (1997) for both the California and Peruvian Currents in the
mid-1970s. Under these circumstances, nutrient enrichment may not occur because the
combination of increased temperature and coastal wave activity cause thickening of the
mixed layer. This prevents Ekman pumping reaching the nutrient-enriched waters, which
under these conditions would be deeper than normal (Shkedy et al., 1995).
Furthermore, the upwelling values used in this study were estimated for large 30 x 30
oceanic.quadrants. Macrocystisbeds would normally occupy only a small portion of such
a quadrant, being restricted to small rocky coastal zones where nutrients are available
from runoff and from benthic sources stirred up by vigorous water movements (Tegner et
al., 1996). Also, there are generally sufficient nitrogen levels in the beds of Macrocystis
in coastal waters off southern California (Dohrman and Palmer, 1981), therefore nutrient
deficiencies likely prevail only during periods of extreme sea warming, such as those
caused by El Nifio (Dean and Jacobsen, 1986). These observations can be extended to the
beds off Baja California.
554 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
From the factors considered in this study, SST is the one variable that explains most of
the changes in the abundance of M. pyrifera. As a result of this species' sensitivity to
temperature (it is temperate affinity), it grows better in cool regimens (Tegner and Day-
ton, 1987). Its low tolerance to variations in temperature were evident during the El Nifio
1982-83 and 1997-98, when this species disappeared from its normal distribution range
in the Baja California Peninsula.
ACKNOWLEDGMENTS
Thanks to Productos del Pacifico, S.A. de C.V. for providing the data of harvest of Macrocystis.
M.C.V., D.L.B. and R.A.R. are fellows of COFAA-IPN. English language text was edited by I.
Fogel at CIBNOR.
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hist6rico de lapesqueriay lavariabilidad del marco ambiental. Pages 191-212 in S.A. Guzmin
del Proo, ed. Mem. Taller Intern. Mexico-Australia sobre reclutamiento de recursos marinos
bent6nicos de Baja California. Secretaria de Pesca - Instituto Politecnico Nacional. 212 p.
Wild, P. W., P. M. W. Law and D. R. MeLain. 1983. Variations in ocean climate and the dungeness
crab fishery in California. Fish. Bull., U.S. 172: 175-188.
Zar, J. H. 1984. Bioestatistical analysis, 2nd ed. Prentice-Hall, Inc. Englewood Clifts, New Jersey.
718 p.
Zimmerman, R. C. and J. N. Kremer. 1984. Episodic nutrients supply to a kelp forest ecosystem in
southern California. J. Mar. Res. 42: 591-604.
_ and __ . 1986. In situ growth and chemical composition of the giant
kelp, Macrocystispyrifera:response to temporal change in ambient nutrient availability. Mar.
Ecol. Prog. Ser. 27: 277-285.
DATE SUBMrrTrED: September 8,2000. DATE ACCEPTED: February 18, 2002.
ADDRSSES: (M.C.V., D.L.B., R.A.R.) Centro Interdisciplinariode Ciencias Marinas- CICIMAR-
IPN.Av. InstitutoPolit6cnicoNacionals/n. Palode SantaRita. Apdo. Postal592.La Paz, B.C.S.,
Col.
Mexico23000. (E.S.Z.) CentrodeInvestigacionesBiol6gicas delNoroeste (CIBNOR), La Paz,B.C.S.,
Mexico. (R.M.) Productosdel Pacifico, S. A. de C V., Ensenada, Baja California,Mexico. CoRRE-
SPONDING AUTHOR: (M.C.V.) CentroInterdisciplinario CienciasMarinas, Apdo. Postal592, La Paz;
de
Baja Califormia Sur, Mexico 23000. Phone: (52) 112-2-53-44; Fax: 112-2-53-22. E-mail:
<mmcasas@ipn.mx> and <mcasasv(hotmaiL,com>.
COPYRIGHT INFORMATION
TITLE: Effect of climatic change on the harvest of the kelp
Macrocystis pyrifera on the Mexican Pacific coast
SOURCE: Bull Mar Sci 73 no3 N 2003
WN: 0330505190002
The magazine publisher is the copyright holder of this article and it
is reproduced with permission. Further reproduction of this article in
violation of the copyright is prohibited.
Copyright 1982-2004 The H.W. Wilson Company. All rights reserved.
EFFECT OF CLIMATIC CHANGE ON THE HARVEST OF TIEE KELP
MACROCYS77SPYRIFERA ON THE MEXCAN PACJFIC COAST
MargaritaCasas Valdez, Elisa Serviere Zaragoza, Daniel Lluch Belda,
Roberto Marcos andRuth Aguila Ramirez
ABSTRACT
The effect of sea temperature, sea level, upwelling, and wind speed on the Macrocystis
pyrifem harvest was evaluated using historical harvest data for the western coast of the
Baja California Peninsula, Mexico. The relationship between the environmental vari-
ables and harvest per unit effort (CPUE) was determined through correlation analysis for
the period from 1956-1998. The effects of temperature, upwelling, and sea level were
inverse, whereas the effect of wind speed was not significant. Multiple regression analy-
sis showed that, of the variables studied, temperature best explained variations in
Macrocystisharvest.
The giant kelp Macrocystispyrifera (Linnaeus) C.Agardh isamember of the large
brown algae (Phaeophyta) that are a conspicuous part of the marine environment. It is a
subtidal species that attaches to solid or soft substrate, produces holdfasts that are either
conical or low mounds. Because of the great size of individual plants, their trunk-like
appearance in the water column, and the extensive surface canopies along the coastline,
these subtidal habitats have been called forests (Foster and Schiel, 1985). Subtidal forests
of M pyrifera occur in many areas of the world, but is most widely distributed in the
southern hemisphere. In the northern hemisphere, M. pynifera commonly occurs from
near Santa Cruz (central California) to Baja California (Foster and Schiel, 1985). Giant
kelp is highly productive.
Macrocystis has been harvested since 1956 along the Pacific coast of Baja California
and exported to the United States for the production of alginates. Recently, it has been
sold in Mexico as meal and f6r abalone aquaculture (McBride, 1998). It is harvested by
specially designed ships that cut the algae at an approximate depth of 1.2 m and transport
it. Two ships participated in the harvest at the area of interest: EL CAPITAN, from 1956-
1966, (storage capacity of 300 t), and EL SARGACERO from 1967 to present, (storage ca-
pacity of 400 t). Ship operations were the same at all locations and did not change over
the study period.
In spite of the importance of kelp, studies of variations in abundance or the size of
harvest are few. Casas et al. (1985) and Hernandez et al. (1989a,b; 1991) reported higher
values of surface canopy and standing crop biomass in spring and summer. Corona (1985)
found a higher harvest with less effort in spring and summer from data on kelp beds from
1978-1984. The first objective of this study is to analyze changes in the size ofMpyiifera
harvest, the effort, and the catch per unit effort (CPUE) along the western coast of the
Baja California Peninsula over the period of large-scale harvest, almost 43 yrs.
There is evidence that climatic changes have altered the productivity of marine ecosys-
tems at several trophic levels (Polovina et al., 1994). The existence of causative relation-
ships between the physical environment and biological productivity has been demon-
strated (Polovina et al., 1995). In the California Current System and in other parts of the
world, changes in climate have been shown to cause fluctuations in the number of small
Bulletin ofMarine Science 545
0 2003 Rosenstiel School of Marine and Atmospheric Science
of the University of Miamni
546 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
pelagic fish (Southar and Isaacs, 1974; Lluch et al., 1989,1991,1992; Baumgartner et al.,
1992; MacCall, 1996). Less isknown about the effect of climate on benthic resources. In
California, there is available information about the effect of environmental variables on
the crab fishery (Wild et al., 1983; Bostford, 1986), abalone (Tegner, 1989, 1991; Davis
et al., 1992), and sea urchin and kelp (North and Zimmerman, 1984; Dean and Jacobsen,
1984, 1986; Tegner and Dayton, 1987; Tegner et al., 1996). Tegner et al. (1996) com-
pared data on the maximum canopy of the kelp forests and the size of the annual harvest
of Macrocystisfor Califomia, and they chose harvest size as the most useful data to relate
to environmental variables. They pointed out that harvest size was a reflection of changes
in consumer demand and harvest productivity, as well as natural disturbances. This data
has the advantage of integrating growth over a long period and has less subjectivity in its
measurement.
The second objective of this study is to show how the abundance of M pyiifera, ex-
pressed as catch per unit effort (CPUE) is related to environmental variables, including
sea surface temperature, sea level, upwelling index, and wind speed on the west coast of
Baja California. It was also determinated which environmental factor best explains the
variability in the abundance of Macrocystis.
METHODS AND MATERIALS
In Mexico, M. pyrifera is distributed along the westem coast of the Baja California Peninsula
from the border with the USA to Punta San Roque, Baja California Sur. However, the alga is
harvested from Islas Coronado (3215rN) to Arrecife Sacramento, Baja California (30'30N) at 15
beds: Islas Coronados, Playas de Tijuana, Punta Mezquite, Salsipuedes, San Miguel and Sauzal,
Bahia de Todos Santos, Punta Banda, Bahia Soledad, Santo Tomais, Punta China, Punta San Jos6,
Punta San Isidro, Punta San Telmo, Isla San Martin, and Bahia del Rosario (Fig. 1). This study
encompasses all of these.
KELP HARVEST DATA.-Daily harvest records from 1956-1998 were provided by Products of the
Pacific, S.A. de C.V. These contained the following information: harvest date, name of the bed,
number of trips made by each ship, and harvest size mt (wet weight). In total, data were extracted
from 3278 daily records.
The volume of the harvest per trip made by each ship, called harvest per unit effort (CPUE),
served as an index of abundance of Macrocystis.To adjust for the difference in the storage capacity
of the two ships, the data on harvest volume trip-' was weighted in the following way:
Period Storage capacity (t) Harves volume/trip (t) Weight factor
1967-98 400 364 1
1956-67 300 222 0.61
PHYSlCAL PARAMETERs.-Sea surface temperatures (SST) were obtained from the data base of
Reynolds and Smith (1994). The database contains monthly averages of SST (°C) from 1956-1992
for 20 x 20 ocean quadrants. Records for latitudes 30°-32° N were selected. For the years 1993-
1998, this information is available for 10 x 10 ocean quadrants. For consistency, we averaged the
monthly values from four quadrants. For years 1956-97, annual averages for the middle level of
the sea (MLS) were obtained from the web page of the Sea Level Center of the University of
Hawaii (http:lluslc.soest.Hlawaii.edu/uhslc/data.html). For years 1956-98, the upwelling index (UI)
for latitudes 28.5°-31.5°N was obtained from the upwelling index database of Bakun (1973), which
maintains records for 3° x 3° ocean quadrants. The upwelling index is expressed as m3s-' mr' of
1
g00
CASAS VALDEZETAL: EFFECTS OFCLIMATIC CHANGEON HARVESFMACROCYSTIS 547
Figure 1.Distribution of the harvest beds of Macrocystispyriferaalong the Baja Califomia Peninsula.
coast. For 1956-89, wind speed data (WS) for latitudes 30°-32°N were obtained from the COADS
(Comprehensive Ocean-Atmosphere Data Set) database, which compiles monthly average wind
speeds in ms-' for 20 x 20 ocean quadrants.
The annual averages for cpue, SST, UII, WS, and MLS were calculated. These average values
were used to calculate the annual anomalies for each of the variables using the following equations:
548 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
Ai =xi - x
n
>xi
x= i-I
n
where, A is the anomaly of the fh year, Xi is the ith annual value, x is the average value for the year
i to n, n is the number of years.
Because the SST and MLS series had a long-term trend, this trend was eliminated through a
linear fit. To eliminate the noise in the interannual variability in the graph representation, the series
were smoothed by using running averages (Statsoft, 1999).
The relationships between abundance of M pynifera and the climatic variables were determined
through correlation analysis using the anomalies of variables and harvest volume/trip values (Ander-
son, 1972). To identify the factor that explains most the variation in harvest volume/trip of
Macrocystis,a multiple linear regression analysis was used (Zar, 1984).
RESULTS
In Baja California, kelp forests of M pyrifera have been harvested since 1956 by the
same company. Annual harvest have ranged from a low of 2980 t in 1983, to 44,250 t in
1989, with an average of 25,900 t (Fig. 2A). The history of the Macrocystis harvest re-
veals two major periods: 1959 to 1977, when the harvest increased to 44,000 t, and 1985-
1997, when the harvest fluctuated around 32,800 t. Three large declines in harvest oc-
curred in 1958,1983, and 1998 (Fig. 2A), declines of 87%, 89%, and 79% with regard to
the average value.
SST from 1956-98 had an average value of 1N7.4C, with a minimum of 16.3°C in 1975
and a maximum of 19.3°C in 1998. There were cool periods in 1956-57, 1960-66 and
1969-76. Warming periods occurred in 1958-59, 1981-84, and a very pronounced one
from 1992-98. In this last year, the SST was the warmest in 43 yrs (Figs. 2B,3A). For the
period 1956-97, MLS varied, on average by 2.03 cm, with a minimum of -67.6 cm in
1964 and a maximum of 107.2 cm in 1984. MSL had negative anomalies in 1960-76 and
1988-90. Positive anomalies occurred in 1958-59,1982-84, and 1990-97 (Figs. 2C,3B).
The average value for the UI from 1956-98 was 122 m3 s-', with a minimum of 67 m
3
3s-' in 1971. There were three periods of negative
s7' in 1966 and a maximum of 169 m
anomalies: 1965-69, 1985-87 and 1992-93. Positive anomalies occurred in 1956-59,
1970-75, 1980-84 and 1989-91 (Figs. 2D,3C). From 1958-90, the average wind speed
was 22.2 m3 s', with a minimum of 17.5 m3 ' in 1967 and a maximum of 29.9 m3 s' in
1987. Clear negative anomalies occurred in 1959 and 1966-70. Positive anomalies oc-
curred in 1971-75 and 1982-89 (Figs. 2E,3D).
Correlation analyses indicated an inverse relationship between the size of CPUE
and SST (r2 = -0.46; P < 0.05; Fig. 3A); MLS (r2 = -0.48; P < 0.05; Fig. 3B), and UI
(r2 = -0.35, P < 0.05; Fig. 3C). The relationship between cpue and WS was not
significant (r2 = 0.006, P > 0.05; Fig. 3D). The results obtained by multiple regres-
sion analysis (SST: -0.45; MLS: -0.02; WS: -0.03 with r= 0.46, P < 0.10) indicate
that changes in SST best explain the variation in harvest volume trip-' of M. pyrifera.
CASAS VALDEZ ETAL: EFFECTS OF CLIMATIC CHANGE ON HARVEST MACROCYS7S 549
50000 a) EINiIIo El Niubo
40000
30000 ElNiio(1958)
20000
10000
0O 11 Illl111HIliillillllllillli
IH lllH ill
1956 1960 1964 1968 1972 1976 1900 1984 1988 1992 1996
20
b)
19
17
16
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995
150
c)
100
O 50
*0
0A
~ 50
.100
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995
Figure 2. Data series of a)Harvest volume of Macrocystispyrifera,b)Sea surface temperature, c)
Middle anomaly level sea, (on followingpage) d) Upwelling index, and e) Wind speed.
550 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
180
d)
160 -
?140-
15~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~3
-E 120
100
80
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995 1998
30
28
26
3- 24
~22
20 -
18
16 ,.. .. .. .. .. .. .L. .. .. .. .. ..........
1956 1960 1964 1968 1972 1976 1980 1984 1988
DISCUSSION
The fishery of M pyrifera does not show signals of overexploitation, since it is at a
stage where effort increases correspond to harvest increases and the CPUE has remained
almost constant from the beginning of the collecting period until the present.
The harvest volume per trip is a reasonable indicator of the abundance of Macrocystis
because approximately 60% of the algal biomass is represented by the surface canopy
and about 95% of the production is in the upper I m of the water column, with a maxi-
mum cutting depth of 1.2 m. Furthermore, harvesting techniques and the ships have re-
mained similar over the period of study, so the volume of each harvest is comparable
throughout this study.
The periods of cooling and warming of the SST found during over 43 yrs are consistent
with those found by Smith (1995) for 1940-1983 at the Scripps Institution of Oceanogra-
phy pier in La Jolla, California.
CASAS VALDEZETAL: EFFECTS OF CLIMATIC CHANGEON HARVEST A CROCYSTIS 551
200 -
150 . 1.5
100 I
1
200
-cpue -.- SST
-150 -IO0
1956 1960 1964 1968 1972 1976 1980
13~~~~~~~~~~~~~~~~~~~0 1984 1988 1992 1996
A-100
200 3
C 150
b)
150
100
100
.50
.,....
., --
15
U-'-200
50
*0
~~~~~~
-50~~~~~~~~~~~~~~~~5
-100
-opuc -- vIS --100
-150
1956 1960 1964 1968 1972 1976 1980 1984 1988 1992 1996
Figure 3.Correlation of harvest volume/trip of Macrocystisppriferawith a)Sea surface temperature
and d) Wind
(SST), b) Middle level sea (MLS), (on following page) c) Upwelling index (UI), represents the
speed (WS). The heavy line represents the harvest volume/trip and the thin line
environmental variable.
Positive anomalies in the CPUE between 1964 and 1975 correspond to a decrease in
the SST. The low temperatures during this period coincide with a Regime Change (Mantua
et al., 1997). Beginning in 1976, the SST increased coincidently with a decrease in the
harvest volume of Macrocystis, with both peaking in the mid-1 980s. In 1988-89, during
mommommommom
552 BULLETIN OF MARINE SCIENCE, VOL 73, NO. 3,2003
250 -
150
,C- 50-
3
a
0
E -50-
eJ
-150
-250
)56 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995 1998
250 6
d)
4
150 -
.2-. 50 -
A -2
ti
s -
:h2LYLWDIL-AO
A-- cE
03
to
A
C, -50 -
-150 -
-ccpue--WS
-250 - , ............ ,., ...... -6
1956 1959 1962 1965 1968 1971 1974 1977 1980 1983 1986 1989 1992 1995 1998
a La Nifia cooling period, the maximum CPUE occurred. Finally, since the beginning of
the 1990s, a strong warming trend is apparent, with a concomitant decrease in the CPUE.
North et al. (1993) related the maximum annual canopy area of the algal bed of Califomia
from 1967-89 to the maximum and minimum quarterly temperatures and to the number
of warm and cool days per year. They found that the former explained 34% of the vari-
ability in canopy area and the latter to 46%.
CASAS VALDEZ ETAL: EFFECTS OF CLIMATIC CIIANGE ON liARVEST MACROCYS77S 553
The effects of high temperatures in the Pacific waters of Baja California Peninsula
during the El Nino years were evident in the harvest and CPUE registered in 1958,1983,
and 1998, amounting to under 5000 t yr'. During El Nifio events, changes in both distri-
bution and abundance of M pyriferahave been described at some localities of California
for 1958 and 1983 (Tegner and Dayton, 1987) and for Baja California Peninsula for 1983
and 1998 (Hemrindez, 1988; Ladah et al., 1999). During El Nifio, high temperatures and
low nutrient concentration results in wide-spread mortality of the giant kelp forest (Gerard,
1984; Tegner and Dayton, 1987, 1991; Tegner et al., 1996).
Increases in sea temperature may reduce the Macrocystis population by limiting the
availability of nutrients. A strong negative correlation was noted between temperature
and nitrate availability (Gerard, 1982; Zimmerman andiKremer, 1986). When nitrate con-
centration falls below 1 pM, as it does off the coast of southern California at approxi-
mately 15.5°C (Zimmerman and Kremer, 1984) and off Bahia Asunci6n, Baja California
Sur at 18.5°C (south of the distribution of this species) (HernAndez et al., 2001) growth
rate of Macrocystisdeclines. Large-scale disappearances may be explained because canopy
growth is supported by transfer ofnitrogen from lower parts of the plant that are bathed in
water with higher nitrate concentrations (Jackson, 1977). If the lower parts of the plants
cannot transfer enough nitrogen to the surface, then the canopy dies back and individuals
succumb, probably because of low nitrogen reserves (Gerard, 1984).
In addition to limiting nitrates, and thereby causing Macrocystisto die off, high tem-
peratures also inhibit successful recruitment of new algal cells to the beds. Thus, the
population cannot recover until the temperature decreases (Jackson, 1977). At the south-
ern limit, Ladah et al. (1999) suggest that microscopic stage of M.pyrifera survived the
stressful conditions during the ENSO, possibly in a dormant state.
The decline in the CPUE when MSL was higher than normal may, in fact be caused by
increased SST. Large positive anomalies in the MLS occur during periods in which the
thermocline is at a greater depth, and at the same time, when SST is higher. The pattern of
response of CPUE to changes in MLS is similar to that previously described for tempera-
ture changes.
Upwelling of deep-sea water normally brings to the surface a rich supply of nutrients
that can benefit marine organisms. The surprisingly inverse correlation found between UI
and Macrocystis abundance may be explained as follows. Some periods (1958-59 and
1980-84) with high upwelling values also had elevated SST. A similar phenomenon was
documented by Hayward (1997) for both the California and Peruvian Currents in the
mid-1970s. Under these circumstances, nutrient enrichment may not occur because the
combination of increased temperature and coastal wave activity cause thickening of the
mixed layer. This prevents Ekman pumping reaching the nutrient-enriched waters, which
under these conditions would be deeper than normal (Shkedy et al., 1995).
Furthermore, the upwelling values used in this study were estimated for large 30 x 30
oceanic.quadrants. Macrocystisbeds would normally occupy only a small portion of such
a quadrant, being restricted to small rocky coastal zones where nutrients are available
from runoff and from benthic sources stirred up by vigorous water movements (Tegner et
al., 1996). Also, there are generally sufficient nitrogen levels in the beds of Macrocystis
in coastal waters off southern California (Dohrman and Palmer, 1981), therefore nutrient
deficiencies likely prevail only during periods of extreme sea warming, such as those
caused by El Nifio (Dean and Jacobsen, 1986). These observations can be extended to the
beds off Baja California.
554 BULLETIN OF MARINE SCIENCE, VOL. 73, NO. 3,2003
From the factors considered in this study, SST is the one variable that explains most of
the changes in the abundance of M. pyrifera. As a result of this species' sensitivity to
temperature (it is temperate affinity), it grows better in cool regimens (Tegner and Day-
ton, 1987). Its low tolerance to variations in temperature were evident during the El Nifio
1982-83 and 1997-98, when this species disappeared from its normal distribution range
in the Baja California Peninsula.
ACKNOWLEDGMENTS
Thanks to Productos del Pacifico, S.A. de C.V. for providing the data of harvest of Macrocystis.
M.C.V., D.L.B. and R.A.R. are fellows of COFAA-IPN. English language text was edited by I.
Fogel at CIBNOR.
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DATE SUBMrrTrED: September 8,2000. DATE ACCEPTED: February 18, 2002.
ADDRSSES: (M.C.V., D.L.B., R.A.R.) Centro Interdisciplinariode Ciencias Marinas- CICIMAR-
IPN.Av. InstitutoPolit6cnicoNacionals/n. Palode SantaRita. Apdo. Postal592.La Paz, B.C.S.,
Col.
Mexico23000. (E.S.Z.) CentrodeInvestigacionesBiol6gicas delNoroeste (CIBNOR), La Paz,B.C.S.,
Mexico. (R.M.) Productosdel Pacifico, S. A. de C V., Ensenada, Baja California,Mexico. CoRRE-
SPONDING AUTHOR: (M.C.V.) CentroInterdisciplinario CienciasMarinas, Apdo. Postal592, La Paz;
de
Baja Califormia Sur, Mexico 23000. Phone: (52) 112-2-53-44; Fax: 112-2-53-22. E-mail:
<mmcasas@ipn.mx> and <mcasasv(hotmaiL,com>.
COPYRIGHT INFORMATION
TITLE: Effect of climatic change on the harvest of the kelp
Macrocystis pyrifera on the Mexican Pacific coast
SOURCE: Bull Mar Sci 73 no3 N 2003
WN: 0330505190002
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