tegneretal97 small scale, low frequency oceanographic effecs on kelp forest succession
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122 Mar Ecol Prog S er 146: 117- 134, 1997
2.5Macrocvs t i s pyr i fera - 8m Central- 5m Central- 2m Central- 8m North
--b 18m South
1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996
l 0 1 18m Central
1.0:
15m Central
1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996
r 2 5--01
22.0
1.5
1 .o 12m Central
0.5
0.0
1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996
1.0-
1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996
18m South
Fig. 3. Macrocy stispyrifera . Density of adult (defin ed as 4 or more stipes, Dayton et al. 1992) giant kelp de termined quarterly at
the long-term study sites. 1983 to 1995. Error bars represent *l SD
s to rm s of 1982-1983 a n d 1988 . Be c a us e bo th e ve n ts conspec if ics . At th e 3 sites w h e r e recruitment
w e r e ca tas trophic in te rm s of the ir des truc tion of occurred immedia te ly after t h e dis turbances (1 8 mgia n t ke lp popu la t ions , t h e first pos t-dis turbance Sou th be c a m e a n urchin ba r re n after th e 1988 storm
cohorts faced little competi t ion from exis t ing a du l t a n d 12 m Ce nt ra l w as initially dom ina te d by Des-
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Tegner et al.: Oceanographic effects on kelp forest succession 125
Fig. 6 . Macrocystis pyrifera. Changesin the mean numbe r of stipes (growth)
per plant and stipe density (carryingcapaci ty) as functions of pla nt densityat (A ) 18 m Central and (B ) 15 m
Central, 1983 to 1995
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12 6 Mar Ecol Prog Ser 146 : 117-134, 1997
7 1 A. P t e r v q o ~ h o r a c a l i f o r n i c a
--C 15mCer.:ral
+ 2m Central
U 8rnNonh
+ 8m South
40 1 B. L a m i n a r i a f a r l o w i i
Fig. 7. Changes In (A ) dens ity of
understory kelp Pterygophora
californica, an d (B ) percent
cover of understory kelp Lamj-
nan d farlorvi!, 1983 to 1995 at
1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 th e 5 permanent sites
permanent sites (except 18 m South du e to sea urchin
grazing in 1986). The very cold La Nina conditionsafter the 1988 storm were associated with extraordi-
nary M. pyrifera growth and declines in L. farlowii
percent cover to zero at all sites between 1989 and
1991. L , farlowili was a bsen t for Interv als of 1 to 3 yr
until th e decline of local M. pyrifera stands allowed
for new recruitment.
Other algae
The other algal groups showed several patterns. The
annual Desmarestia spp. responded to storm and graz-
ing disturbances with short-term bursts in percent
cover (dat a not shown) . Cystoseira osmundacea also
recruited at some sites after each major disturbance,
but only persisted at trace levels in both the 1980s andthe 1990s; his is consistent with the low survivorship of
cohorts of this species obser ved in the 1970s (Dayton et
al. 1984). Coralline alga e (Fi g. lOA), well kno wn for
their de ep distributions (e. g. Lobban et al. 1985),
showed no evidence of response to the different tem-
perature/Macrocystis pyrifera regimes nor to the 1988
storm. Foliose red al gae, although they expen enced
major recruitment peaks at some sites during the
1983-1984 El Nino, were generally similar to coral-
lines. Brown turf (Dictyotaceae) persistence, however,
followed a patte rn very similar to tha t of th e understory
kelps; these recruited strongly after both major distur-
bances at 18 and 15 m Central, persisted at low levels
until the 1988 storm, virtually disappeared from mid
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T e g n e r e t a1 . Oceano graphic e f fec ts on ke lp fores t succession 12 7
0. 5 18m C e n t r a l o 25
0. 4+ acrocystis 0 20- terygophora
0. 3 0 15
0 2 0.10
0.1 0.05
0.0 0.00
1 9 8 3 1 9 8 4 1 9 85 1 9 8 6 1 9 8 7 1 9 8 8 1 9 8 9 1 9 9 0 1 9 9 1 1 9 9 2 1 9 9 3 1 9 9 4 1 9 9 5 1 9 9 6
1 o15m C e n t r a l 6 o
0.8
4.00.6
0 42.0
0. 2
0 .0 00 p
-mX 3 .0 80 L0)z a
c1 6.0
, 2.0 .-m.-mC
4.0D
d 1.0
V)2 0.- 0
.c
0 .0 00 21 9 8 3 1 9 8 4 1 9 8 5 1 9 8 6 1 9 8 7 1 9 8 8 1 9 8 9 1 9 9 0 1 9 9 1 1 9 9 2 1 9 9 3 1 9 9 4 1 9 9 5 1 9 9 6 ?
90
18 m Northc
I C
Flg 8 Macrocys t i spynfe ra an d Pterygophora californica. Cha nges in ke lp dens i t i e s a t each of t he pe rm anen t s i te s , 1983 to 1995.
Error ba rs repres ent * l SD. Note sca l e changes a mon g s pec ies and s i t e s
1989 through 1992, an d then reapp eared as pos t-1988- sen ce of s tress , an ecosys tem achiev es a higher leve l of
s torm M. pyrifera cohorts dec lined (Fig. 1 0 B ) . orga n iz a tion , whe re a s in the p re s e nc e of s e v e re s t r e ss ,
the process of organiza tion is like ly to be s lowed an d
the u l t im a te s tea dy s ta te tha t m igh t be r e a c he d would
DISCUSSION be a t a lower leve l of orga n iz a tion . T urk ing ton e t a l .
(1993) , es t in g the hypo thes is tha t the e ffec ts of inte r-
Odu m (1981) comp ared the diffe rent tra jec tor ies of spec if ic competi t ion would dec lin e a long gradient s ofs uc c es s ion in e nv i ronm e nts w i th a nd w i thou t s e ve re de c re a s ing nu t r i e n t s a nd inc re a s ing d i s tu rba nc e
e nv i ronm e nta l s t r e s s. He s ugge s te d tha t , in the a b - a m on g pa s tu re p la n t s , de m ons t r a te d tha t nu t r i e n t s
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12 8 Mar Ecol Prog Ser 146: 117-134, 1997
0.518m Centra l
30
0.4- E Macrocystis- aminaria
0.320
0.210
0
0.0 0
1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996
U mm 'EL 3.0a 12m Centra l 6 .$a- E2 2.0.-U) ;C
m
10
.-2 g- U
2
0 0.0 0 :
1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 l994 1995 1996 gI" a
18m North
0.518m South
4
0.43
0.3
2
0.2
01
0.0 0
Fig. 9 . Changes In Macrocystispyrdera dens i ty an d Lamjnarja farlowii percen t cover at each of the perm anen t s i tes , 1 .983 o 1995.E rro r bars rep resen t t SD Note sca le changes amon g spec les and s i t es
and disturbance interact to determine community
structure. Here we compared succession after 2 cata-
strophic disturbances under contrasting oceano-
graphic regimes, the nutrient-stressed El Nino condi-
tions of 1983-1984 and , on the opposite end of the
spectrum, the very cold, nutrient-rich La Nina condi-tions of 1988-1989. These different oceanographic
regimes had different effects on succession, notably on
the performance of the competitive dominant itself and
on its competitors. In both cases the unusual oceano-
graphic conditions lasted for about 2 yr, but the appar-
ent effects on community structure persisted for the life
of the post-disturbance Macrocystis pyrifera cohorts,
despite relatively normal oceanographic conditionsfrom 1985 to 1987 and relatively warm waters in recen t
years.
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Tegner et al.. Oceano graphic effects on kelp forest succession 129
A. Articulated Coral l ines
I 18mCenlral
1 15mCenlral- 2m Cenlral
5 2 0l8m Nonh
LQ)
5 - 8m South0
0
ad
C
a,
2 1 0a
a
0
B. Brown Turf (Dictyotaceae)
Fig. 10. Chan ges in pe rcen t cover of (A ) articulated corallines an d (B ) brown turf algae (Dictyotaceae) at the 5 permanen t sites,
1983 to 1995
MacrocystispyrLfera performance varied considerably
over the study period. Intense recruitment followed bothmajor disturbances, but there the similarities ended.
Giant kelp plants that survived the 1982-1983 storms
formed a sparse canopy in late spring, but the canopy be-
gan to deteriora te in summer, and by October, the tallest
plants were 6 to 8 m below the surface; there was also
subst antia l mortality of plan ts that survived the storms
(Tegner&Dayton 1987).New cohorts of adu lts recruited
in summer and fall 1983, but these had poor growth
(Dayton & Tegner 1984) and survival relative to post-
disturbance cohorts in 1988 (Fig. 5) . Canopy area was
very low in 1983-1985 before it began to recover; 1986
and 1987 were relatively normal years in terms ofcanopy coverage and oceanography (Fig. 2). Excellent
growth conditions in 1988 were reflected in the more
rapid post-storm recovery of the canopy an d the better
survival of initial post-disturbance cohorts (Fig.5) .SomeM.pyrifera plants that had lost all their stipes in the 1988
storm produced new stipes and lived several months
longer, whereas the nutrient-stressed plants damaged
by the 1982-1983 storms generally had very poor sur-
vival and no recovery was observed of plants tha t had
lost all their sti pes (Dayton et al. 1992). Two years of
anomalously cold water resulted in peak s in both canopy
area and kelp harvest in 1990 (Fig. 2). The post-1988-
storm M.pyrifera cohorts showed their sensitivity to the
relatively warm water, inte rmittent El Niiio conditions of
1991-1993 with reductions in harvest and canopy area ,
poor growth in stipe number, and reduced carryingcapacity, but appa rently not throu gh mortality (Figs. 2 , 3
& 6).
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134 Mar Ecol Prog Ser 146: 117-134, 1997
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Manuscript first received: May 28, 1996
Revised version accepted: October 14, 1996