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52 Mar. Ecol. Prog. Ser. 77:49-63, 1991
A . Average Drift By Site
30 1
18m 18m 18m 1% l 2 r
Nonh S o u l h Central Central Central
Site
B. Average Drift By Season
Table 1. Species composition of all drift collections (n = 131)
from all 5 sites from 1983 through 1986. Macrocystispyrifera is
also broken dow n by parts of the pla nt
SpeciesOO
Macrocystis pyrifera 85.37
Blades 38.88
Stipes 25.71
Fronds 12.42
Haptera 6.82
Sporophylls 1.54
Other laminarians
Pterygophora californicaPelagophyc us porra
Laminaria farlowii
Egregia rnenziesiiEisenia arborea
Agarurn firn briatumOther Phaeophyta
Desmares t ia l~yu la t a
Dictyopteris undulata
Cystoseira osmundace aPach ydictyon corn'aceurn
Dictyota flabellata
Rhodophyta
Foliose red algae
Articulated corallines
Angiosperms
Phyllospadiu spp.
Total weight 657 065 g
A. Central Forest Sites
wlnler sprlng summer fall
Se a so n
B. Outer Edge Sitesduring 1983 o 1985, ncludi ng cases of zero valu es for
both species, follocved by increases in 1986 an d 1987. lo o
These curves sugge st that recruitment was consistently 90
r
better at e dge of the forest sites than inside the bed. To 80 -
test this hypothesis, recruitment rates were ranked for 70 -
eac h of the 9 determinations at the 5 sites and analyzed60 -
w t h the Friedman 2-way ANOVA . The rankng s were l ~ m ~ ~ l t h
significant for both species (p< 0.01). posteriori tests 2 - am South
40 - - am Central(Nemenyi 1963)were conducted to determine which
30 Isites were significantly different. The mean percent 1983 1984 1985 1986
r e c ru i tmen t (lga3o fo r each site in th e Orde r ofFig 3. Macrocys t i spyn fera . Variance in the percentage of the
its ranking is given in 2 . Thus, there may drift constituted by M.pyrifera, spring 1983 through the end of
have been minor differences between the 2 species, 1986, by year. (A) Central forest sites. (B ) Outer edge sites
Fig. 2. (A) Average drift by site, all samples combined, spring E 60 -1983 through the end of 1986 (B )Average drift by season, all
2 50 -sites combined, spring 1983 through the end of 1986. Sites, 2and especially seasons, were significantly different (see text) 40 -
--f la m Central- sm Central- 2m Central
l
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Mar Ecol. Prog. Ser. 77: 49-63, 1991
18m Centra l
15m Central
Description of events at 18m S- hanciscanus- purpuralus Until the fall of 1984, ther e was no indication th at18 m S was different from other forest edge sites. At
that time plants were stressed throughout the forest
12m Centra l
Fig. 10. Densities were determin ed once in 1975 on
100 X 2 m transects at the 12 and 18 m C sites running
through the same areas a s the present 25 m transects;
the habi tats are similar but not identical. ( The densities
at 12 m C were reported incorrectly in Tegner&
Dayton1981. The values should b e 0.8 and 2.1 m-2 for Strongy-
locentrotus franciscanus an d S. purpuratus, respec-
tively.) Purple urchin densities do not appear to have
change d since 1975,bu t red urchin densities have been
reduced by 1 to 2 orders of magn itude. Th e purple to
red urchin ratio has increased from 1.6 to 14.6 at 1 8 m C
and from 2.8 to 291 at 12 m C.
Fig. 5. Strongylocentrotus purpuratus and S. franciscanus.Recruitment rates during the mid 1970s (Tegner & Dayton
1981) compare d with 1983 to 1987 as computed on a n annu albasis for 3 sites in the central forest. Data were computed from
m2 samples of urchin aggr egation s
through 1987 at 4 of the 5 st udy sites (Fig. 7; ANOVA, p
> 0.05). The densit ies of both species increased a t 18 mS (p < 0.001 in each case) . At this site, an examination
of the density changes line by line illustrates that some
of th e increase can be att ributed to migration from an
A. 18m Central
B. 12m Central
Z Density < 35 mm- enslly > l00mm
urchin-dom inated area south of line D, which increased Fig. 6 .Strongylocentrotus franciscanus. Densities of young-of-
faster than line C, etc. (Fia.8). However, increases in the-year and large adult red urchins from the mid 1970s\ d ,
recruitment also preceded increases in density , espe- (Tegner& ~ a ~ t o n 1 9 8 1 )nd 1983 to 1987 at 18m C nd 12 m
C sites computed from the mZ amples of urchin aggregations.cially for Strongylocentrotus franciscanus (Fig. 9). The decline in adult density between the mid 1910s and theDensities from the mid 1980s are compared with 1980s was signif icant at both site s and juvenile recruitment
values obtained in 1975 (Tegner & Dayton 1981) in was significantly lower at 12 m C in the 1980s (see text)
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Tegner & Dayton: Sea urchins, El Ninos, and kelp forests 63
Sokal, R. R., Rohlf, F. J. (1969). Biometry. W. H. Freeman and
Co., San Francisco
Tegner, M . J. (1980). Multispecies conside rations of r esource
managem ent in Southern California kelp beds. Can. Tech.
Rep. Fish. Aquat. Sci. 945: 125-143
Tegner, M. J. (1989). The feasibility of en hanc ing red se a
urchin (S trongylocentrotus francis canus) stocks in Califor-
nia; a n analysis of the options. Mar. Fish. Rev. 51(2): 1-22
Tegner, M. J. , Dayton, P. K. (1977). Sea urchin recruitment
patter ns and implications of commercial fishing. Science
196: 324-326
Tegner, M. J. , Dayton, P. K. (1981). Population str ucture ,
recru itment , and mortality of two sea urchins (Str ongy-
locentrotus franciscanus and S. purpurat us) in a kelp
forest. Mar. Ecol. Prog. Ser . 5: 255-268
This article was presented by N. Holland, La Jolla, California,
USA
Tegner, M. J., Dayton, P. K. (1987). El NiAo effects on South-
ern California kelp forest communities. Adv. ecol. Res. 17:
243-279
Tegner, M. J., Levin. L. A. (1983). Spiny lobsters and s ea
urchins: analysis of a predator- prey interac tion. J. exp.
mar Biol. Ecol. 73: 125-150
Tutschulte, T C. , Connell, J. H . (1988). Feeding behavior and
algal food of thr ee specie s of aba lones ( Hahoti s) in South-
ern Calif ornia. Mar Ecol. Prog. Ser . 49: 57-64
Vadas. R. L. (1977). Preferential feeding: an optimization
strategy in sea urchins. Ecol. Monogr. 47: 337-371
Wilson, K. C., Haaker, P. L., Hanan, D. A. (1977). Kelp restora-
tion in Southern California. In: Krauss, R. (ed.) The marine
plant biomass of the Pacific Northwest coast. Ore gon State
University Press, Corvallis, p. 183-202
Manuscript first received: Nove mber 6, 1989
Revised version accepted: A ugust 21, 1991