Marine Biology 34, 137-142 (1976)

© by Springer-Verlag 1976

Changing Lobster Abundance and the Destruction of Kelp Beds

by Sea Urchins
P. A. Breen and K. H. Mann

Department of Biology, Dalhousie University; Halifax, Nova Scotia, Canada

Abstract

In a study area in Nova Scotia , Canada , abundance of the lobster Homarus americanus
decreased by nearly 50% in 14 years. The lobster is a major predator of sea
urchi ns, and during the past 6 years the sea urchin Strongylocentrotus droebachiensis
has destroyed 70% of the beds of Laminaria spp . in the area . Implications for
management are discussed.

Introduction Canada , sea urchins (Strongylocentrotus

droebachiensis) are usually controlled by
Large brown seaweeds (kelps) are e x- lobsters (Homarus americanus) ; and thus ,
tremely produc t ive , a n d thus play a v i- that reduced lobster a b undance h as al -
tal role in the coastal food chains of l owed sea urchi ns to increase, r esulti n g
many parts of the world (Mann , 1973) . in destruction of beds of kelp (L aminaria
Local distributions of seaweeds , however , spp . ) . This paper presents the results
are often restricted by sea urchins from tests of this hypothesis .
(Forster , 1959; Kitching and Ebling , Testing the hypothes i s as a whole
1961 ; Jones and Kain, 1967 ; Paine and would have involved manipulating lobster
Vadas , 1969 ; Ormond and Campbell , 1971; population density in large areas , then
Bernard and Miller 1973 ; Shepherd, 1973) , examining the effect on sea urchins and
and sea urchins have destroyed beds of kelp. In pi l ot experiments , attempts to
kelp (North , 1965; Leighton et al ., 1966) increase lobste r dens i ty fai l ed , because
and seagrasses (Camp et al . , 1973 ; Ogden lobsters rapidly dispersed and disap-
et al . , 1 97 3) . The resulting loss of peared . We therefore tested each sepa-
pr i mary productivity and c hanges in com- r ate component of the hypothesis , using
munity structure are matters f or concern. St. Margaret 's Bay, Nova Sco tia, as the
The abundance of sea urchins may nor- study area . The questions asked were :
mally be limited by predation . In Ca l i - whether kelp had in fact become less
fornia , predati on by the sea otter Enhy- abundant , as casual observation sug-
dra lutris has depleted local sea - u r chin gested ; whether sea urchins a re the
populations, allowing kelp beds to ex- cause of kelp disappeara nce ; wheth e r
pand (McLean , 1962; North , 1965; Ebert , l obsters could be a key predator on sea
1968 ; Lowry and Pearse, 1973) . Differing urchins; and whether lobster abundance
leve l s of sea- otter predation are thought had decreased i n the study area .
to account for diffe r ences i n sea urchin
and kelp abundances in the Aleutian Is-
lands (Estes and Palmisano , 1974). Paine Reduction of L a mina ria (Kelp) Beds
and Vadas (1969) suggested that preda-
tion by a large sea star (Pycnopodia sp . ) Reduction in ke l p beds a l ong a 15-km
accounted for the mosaic of sea urchin length of the shore was observed be-
and algal distributions they observed tween 1968 and 1973. Six transects were
in Washington (USA). Crabs restrict the made through the kelp beds at right
distribution of sea urchins in Britain , ang l es to the shore , from high water to
and thus influ ence algal distrib uti o n the outer limi t of k elp (Mann , 1972 ).
(Muntz et al . , 1965). The total length of transect occupied by
We earlier put forth the hypothesis kelp beds at each survey is given in
(Mann and Breen , 1972) that , in easte rn Table 1. By 1973 , kelp b eds were abse nt
138 P.A. Breen and K.H. Mann : Lobsters , Sea Urchins and Kelp l
Table 1. Laminaria spp. Total length (m) oc-
20 At kelp edge l. 71
cupied by kelp on 6 transects in 1968 and 1973

Date Transect
A c D E F G

1968 11 51 73 43 124 183 ~ 20

1973 0 47 0 0 10 150 u
"'
N
·;;;;
c
0. 55
2.0 ~---~,---~,---~

"'rn II
I
- 5 years 0. 15
I 40
I
I
I
I
I
.-
I 20
I
I
I
-
I
I
~ I I 20 40 60
0 5 10 15 Sea urchin size class (mm)
2
Sea urchi n biomass (kg m- 1
Fig. 1. Strongylocentrotus droebachiensis and Fig . 2. Strongylocentrotus droebachiensis. Size
Liminaria longicruris. Relation between sea- composition of sea- urchin populations at various
urchin biomass at kelp edge and rate of kelp- bed times after kelp disappearance . Values on the
recession. Dotted line indicates the postulated right show biomass in kg.m- 2
value o f sea-urchin biomass below which kelp
does not re cede

or greatly reduced on 4 transects, and to persist, no macrophytes ever appeared

were slightly reduced on the remainder. during the experiment.
Kelp continued to decline in 1974, and When sea urchins were placed at
we estimate that nearly 70% of the kelp ma r ked sites inside a ke lp bed , no de-
on this shore has disappeared since struction of kelp took place. Sea ur-
1968. No newly formed kelp beds were chins crawled onto the blades of kelp
ever found in the study area. There is plants , and were then dislodged and dis -
evidence that kelp beds have been de- persed when the blades we r e swayed by
stroyed in other parts of eas t ern Canada wave action. By contrast, sea urchins at
(Neish, 1973). the edge of a receding kelp bed were
abundant enough to weight down the blades
of kelp and prevent the ir movement. We
postulate a t hre sho ld abundance of sea
Are Sea Urchins the Cause of Kelp Disappearance? urchins which is necessary for them to
graze destructively upon kelp.
If sea urchins are the immediate cause Sea- urchin biomass and the rate of
of kelp disappearance, then kelp should kelp disappearance were both measured
grow on barren rock after sea urchins during 1973 at various points along the
are removed, kelp should be destroyed edge of a rapidly receding Laminaria longi -
when sea urchins are placed in kelp beds , cruris bed. There was a si gnificant posi -
and the rate of kelp disappearance tive correlation (Spearman rank r =
should be positively related to sea- 0.606; P <0 . 0 1) between sea- urchin b io-
urchin abundance. In February, 1972, all mass and kelp recession rate (Fig. 1).
sea urchins (Strongylocentrotus droebachiensis) As the previous experiment suggested,
were removed from a barren area where there appeared to be a threshold biomass,
they were common, and the area was kept near 2 kg/m2, below which sea urchins
free of sea urchins for the next 18 did not cause the kelp to r ecede .
months. Some algae became visible within From these experiments it can be con-
a month, and Laminaria longicruris began to cluded that sea urchins are the immedi-
appear at the end of April. By June 1973, ate cause of kelp disappearance, and
L. longicruris was the dominant algal spe- that sea urch i ns prevent algae from re-
cies, with a biomass equal to that in a colonizing bare rock.
nearby, old-established bed. In a con- After all kelp in an area has been
trol area where sea urchins were allowed destroyed, both the biomass and average
P.A. Breen and K.H. Mann: Lobsters, Sea Urchins and Kelp 139

u
Are Lobsters a Key Predator on Sea Urchins?
~

=> ~
l§ 113'
0. 15 The possibility that the l obster is a
~
b•O. 16
~
key predator on sea urchins was examined
=
·;:: ~

in two steps:
g' ~
] '!5 0. 10 First, prey-species selection and
'!5 "'
~
feeding rate of lobsters was measured in
~ ~

underwater cages that simulated natural

~ :;:c b·O. 0238 • 0. 017 x sea urchin abundance lkgl
conditions as far as possible. Of 6 prey
2' ~
·- => 0. 05
0 species offered to the lobsters Homarus
~ ~ americanus, only the green crab Carcinides
~ 'O
E' maenas was selected at a higher rate than
12 16 sea urchins , while the rock crab cancer
Sea urchin abundance lkg per cage! irroratus and sea urchins were selected
Fig. 3 . Interaction of Strongylocentrotus droe - about equally. Of these species, sea
bachiensis biomass and temperature in deter- urchins are the most abundant in kelp
mining changes in feeding rate of the lobster beds (Miller et al . , 1971), and so are
Homarus americanus probably the major item in the lobster
d i et. Using sea urchins as prey, lobster
feeding- rate was measured as a funct i on
of both temperature and prey biomass
(Fig. 3).
N
' E 10000 Second, th i s information was in te-
_go 300 400 grated in a simulation model with ex-
~
~ isting information on lobster and sea-
E 8000
0 urchin population dynamics (Wilder, 1953;
'5
c Scarratt, 1972 , 1973a ; Miller and Mann,
~=> 6000 1 973). The model was used to discover
the conditions under which lobsters
~ might be a key predator on sea urchins.
E 4000
:::J
·;: Represented i n the model were: numbers
.-e of lobsters and sea urchins in a series
:::J
3
2000 of age classes; urchin recruitment ,
growth and natural mortality; and month-
ly , size-selective feeding by lobsters
20 40 60 80
on sea urchins. The lobster population-
2 size was chosen so as to represent a
Lobster biomass lg. m· 1
Fig. 4. Model results . Change in equilibrium natural, unfished population . After the
Strongylocentrotus droebachiensis biomass at model was built, l obster abundance and
various Homarus americanus biomass values . Each
sea- urchin recruitment- rate were the
line represents a different annual sea-urchin
only parameters that varied. Details of
settlement rate, per m2, indicated at top of
the model are given in Breen (1974).
Results are shown in Fig . 4. At a
curve . Horizontal dashed line represents density
g i ven value of sea- urchin recruitment-
above which kelp beds are destroyed
rate , the equilibrium sea- urchin biomass
- - - - - - - - - - - - - - - - - - - - - - - - - d e c r e a s e d as lobster biomass increased.
At a fixed value of l obster b i omass ,
sea- urchin biomass increased as recruit-
ment-rate increased.
From the fie l d observations shown in
Fig. 1, it was inferred that Strongylocen -
size of sea urchi ns decrease rapidly trotus droebachiensis do not destroy Lami -
(Fig. 2). This may i ndicate that starva- naria spp. at a biomass less than 2 kg.m2 ,
tion occurs. Because of continuing set- and are thus "under control". This value
tlement from the plankton, however, a was, therefore, used to describe the
residual population remains, composed point at which the model sea-urchi n popu-
mainly of small individuals. These ap- l ation was "controll ed " ; the results are
parently exist by grazing microscopic summarized in Fig. 5. When lobsters were
plants as well as detritus from the in- abundant and sea-urchin recruitment- rate
tertidal zone. Despite the low biomass low, sea urchins were controlled. As
of these populations, they prevent a l- lobster abundance decreased and sea- ur-
gae from re- colonizing. Thus , areas chin recruitment-rate inc r eased , sea
which have been cleared of kelp by sea urchins were controll ed l ess often.
u rchi ns r emain barren for at least 6 The model was expanded to include the
years and probably longer . rock crab cancer irroratus and the At lant i c
140 P.A. Breen and K.H. Mann: Lobsters, Sea Urch ins and Kelp

wolf fish Anarhi chas lupus, both of which

!;:_ are also sea- urch in predators (Hirnrnelman
')'
E 400 , - - - - - - - - - - - -- - - - - - -1
and Steele, 1971), and which appeared to
;g J Sea urchins ! be abundant in t he study area. The struc-
.5 : not 'control I ed' J ture of the model remained the same, but
300
~
I
I
I I predatory inter-actions became more com-
I I
c I
I
I
I
plex: lobsters and wolffish ate both
E 200 I
I
I
I crabs and sea urchins, and crabs ate sea
"' 1 Sea urchins 1
~ I
J_ _ _ _ _ _ _ ___ _~oo~~led '
_ _ _______ J
I
:
urchins (Scarratt and Lowe, 1972).
c
JOO Results from this model are shown in
~::> Fig. 6. It was discovered that the addi-
</)
"'"' tion of wolf fish had nearly no effect ,
20 40 60 and that changes in the resu lts were
caused almost entirely by the crabs . Sea
-z,
Lobster biomass lg. m urchins were controlled except when lob-
Fig. 5. Summary of model results. Broken lines ster abundance was very low. Results of
indicate expected natural limits of Homarus each model thus indicate that lobsters
americanus biomass and Strongylocentrotus droe- could be a key predator on sea urchins
bachiensis recruitment rate . Solid line connects under conditions likely to be found in
points at which sea-urchin equilibrium biomass nature.
equals 2000 g.m-2

500
Sea urchin biomass Changes in Lobster Abundance
' !;:_ not'controlled'
"'' E 400 -------------,
The final stage in testing the hypothesis
II
.,;
.5
0 was examination of changes in Homarus
I
I
O> 300 I
I americanus biomass in the study area. Lob-
"' I
I ster abundance was inferred from catch
o>
c I
I per unit effor t , derived from Canadian
~ 200 Sea urcl1ins I
I
government fishery statistics. The best
c 1 'controlled'
L____________________J
! measure of effort available was the num-
~::> JOO ber of licensed lobster fishermen. As
:ii
v'l
shown in Fig. 7, catch per unit effort
declined by 50% from 1959 to 1973 . It
0 20 40 60
2
would be incorrect to cite lobster fi sh-
Lobster biomass (g. m- 1 ing alone as the cause of this decline.
Fig . 6 . Summary of model results, when Cancer The re is evidence that environmental
irroratus and Anarhichas lupus are included as variation causes fluctuations in lobster
well as Homarus americanus. Broken lines indicate abundance (Wilder , 1965; Dow , 1 969 ;
expec ted natural limits of lobster biomass and Flowers and Saila , 1972). For instance,
Strongylocentrotus droebachiensis recruitment there appears to be a positive relation
rate. Solid line connects points at which se a - between river runoff and larval lobster
urchin equilibrium biomass equals 2000 g.m- 2 abundance (Scarratt , 1973b) , and hence
between river runoff and the subsequent
catch (Sutcliffe, 1973). Environmental
c factors , as well as fi shing, may have
e contributed to the decline in lobster
~ 800
-............. stocks seen in the study area.
§"'
------- The hypothesis is thus s upported by
0
i;
600
--------- the evidence: lobsters appear likely as
a key predator on sea urchins, lobsters
·c:::>
have declined in abundance, sea urchins
li; 400
Q. have increased in number and have de-
,.
.t:::
'-'
'-'
stroyed kelp beds at a high rate.
... 200
::>
c
c
<
Discussion and Conclusions
J960 1965 1970
Fig. 7. Homarus americanus. Lobster landings When kelp beds are destroyed , their high
(continuous line) in St . Margaret's Bay, Nova primary productivity is lost. Although
Scotia, 1959-1 973, expressed as lbs per man. the role of algal production in coastal
Crosses indicate values based on estimated num- systems is poorly understood, it is prob-
ber of men. Continuous line joins points, broken able that changes will occur at higher
line is a fitted regression trophic levels. Kelp beds of Laminaria

J
P.A. Breen and K.H. Mann: Lobsters , Sea Urchins and Kelp
spp. also host populations of rock crabs,
the cunner Tautogolabrus adspersus, and
other small fishes and invertebrates,
which virtually disappear when kelp is
destroyed. The direct relations between
lobsters and kelp are not known, but it
is possible that small lobsters depend
on kelp beds for shelter. Scarratt (1972)
found large numbers of small lobsters
(Homarus americanus ) in beds of Irish moss
(Chondrus crispus) . It is also probable that
lobster productivity is direct l y re l ated
to kelp productivity . Th us , one may ex-
pect l obster production to decrease
further as kelp disappears . If so, then
all the changes that occur as kelp de-
clines are destabilizing, in that natural
controls on sea urchins are removed as
kelp beds are removed.
The barr en- rock community that re-
places the kelp bed community appears
persistent. Although sea urchins found
there are small and not abundant , they
are able to graze algae as they settle
and thus the substrate r emains clear of
seaweed cover . If sea urchins continue
to settle at the present rate, it is
difficult to visualize a mechanism that
would a l low kelp beds to re- appear . A
possibility is that the suppl y of larval
sea u rchins could be great l y reduced
after all kel p disappears from a large
area, since most of the breeding popula-
tion is found at the kelp e dge. However ,
in Newfoundland (Himmelman, 1969), kelp
is confined to the zone of strong wave
action just below low- tide level, and
all other rocky areas are inhabited by
sea urchins apparently living on detri -
tus from t h e narrow k elp refuge. Such an
arrangement might persist indefinitely.
If the ideas presented above are cor-
rect , we have described a one-way change
in the structure of rocky subtidal com-
munities in eastern Canada, which will
have a great effect in other communities
and on the production in commercia l
coastal fisheries. A broad survey is
needed to determine precisely how wide-
spread is kelp-bed destruction. Shou l d
it prove as widespread as casual reports
suggest, measures to incease the abun-
dance of lobsters may be the only way to
protect the remaining kelp-bed communi-
ties.
Acknowledgements. The work was supported by
National Research Counci l of Canada grants to
K. H. Mann and I.A . McLaren. The Marine Ecology
Laboratory, Bedford Institute of Oeceanography ,
Environment Canada, provided laboratory space,
equipment and ship time . P . A. Breen received
scho l arship support from Dalhousie University
and the National Research Council of Canada .
141
Literature Cited
Bernard, F . C . and D. C . Miller: Preliminary
investigation on the red sea urchin resources
of Briti sh Columbia (Strongylocentrotus fran -
ciscanus (Agassiz)). Tech. Rep. Fish . Res . Bd
Can . 400, 1- 38 ( 1973)
Breen , P.A. : Relations among lobsters , sea ur-
chins and kelp in Nova Scotia , Ph . D. thesis,
190 pp . Halifax, Nova Scotia , Dalhousie
University 1974
Camp , D. K., S.P . Cobb and J.F . van Breedveld :
Overgrazing of seagrasses by a regular urchin,
Lytechinus vari egat us . Bioscience 2 3, 37-38
(1973)
Dow, R. L . : Cyclic and geographic trends in sea-
water temperature and abundance of American
lobster . Science , N.Y. 164, 1060-1063 (1969)
Ebert , E . E .: A food habits study of the southern
sea otter , Enhydra lutris nereis . Calif. Fish
Game 54 , 33- 42 ( 1968)
Estes, J.A . and J.F. Palmisano : Sea otters : their
role in structuring nearshore communities .
Science , N.Y. 185, 1058-1060 (1974)
Flowers, J . M. and S . B. Saila : An analysis of
temperature effects on the inshore lobster
fishery . J . Fish. Res . Bd Can. 29, 1221-1225
(1972)
Fors ter, G. R.: The ecology of Echinus esculentus
L. quantitative distribution and rate of
feedi ng. J . mar . biol . Ass . U. K . 38 , 361-367
(1959)
Himmelman, J .H.: Some aspects of the ecology of
Strongylocentrotus droebachiensis in eastern
Newfoundland, 125 pp . M. Sc. Thesis, St .
John ' s , Newfoundland : Memorial University
1969
and D. H. Steele : Foods and predators of the
green sea urchin Strongylocentrotus droe-
bachiensis in Newfoundland waters . Mar . Biol .
9, 315- 322 ( 1971)
Jones, N. S . and J.M . Kain: Subtidal algal colo-
nization following removal of Echinus. Helgo-
l anaer wiss. Meeresunters . 15, 460-466 (1967)
Kitching , J . A. and F.J. Ebling : The ecology of
Lough In e . XI . The control of algae by Para-
centrotus lividus (Echinoidea) . J . Anim .
Eco l. 30, 373-383 ( 1961)
Lei ghton , D.L., L . G. Jones and W.J. North:
Ecological relationships between giant kelp
and sea urchins in Southern California. In:
Proc . int. Seaweed Symp . ( 5 , Halifax)
141-1 53 (1966) . (Ed. by E . G. Young and J . L .
McLachlan . Oxford: Pergamon Press 1966)
Lowry , L.F. and J.S. Pearse : Abalon es and sea
urchins in an area inhabited by sea ot t ers .
Mar. Biol. 23, 213-2 19 (1973)
Mann , K. H. : Ecological energetics of the seaweed
zone in a marine bay on the Atlantic coast
of Canada . I. Zonation and biomass of sea-
weeds . Mar. Biol. 12 , 1-10 (1972)
Seaweeds: their productivity and strategy
for growth . Science , N.Y. 182, 975- 981 (1973)
and P . A. Breen: The relation between lobster
abundance , sea ur chins , and kelp beds . J .
Fish. Res . Bd Can . 29, 603- 609 (1972)
142 P . A. Breen and K.H. Mann : Lobsters, Sea Urchins and Kelp
McLean, J.H. : Sublittoral ecology of kelp beds
of the open coast near Carmel, California .
Bi o l . Bull. mar . biol . Lab., Woods Hole 122,
95-11 4 (1962)
Miller , R . J . and K. H. Mann : Ecological ene r get-
ics of the seaweed zone in a marine bay on
the Atlantic coast of Canada . III. Energy
transformations by sea urchins. Mar. Biol.
1 8 I 99- 114 (19 7 3 )
--and D. J . Scarratt : Production potential of a
seaweed- lobster community in eastern Canada.
J . F ish . Res . Bd Can . 28 , 1733-1738 (19 7 1)
Muntz , L ., F . J . Ebling a n d J. A. Kitching : The
e co l ogy of Lough Ine . XIV. Predatory acti vi-
ties of large crabs . J. Anim. Ecol. 34,
315- 329 (1965)
Neish, I.C . : The distribution of sea urchins in
Charlotte County, New Brunswick. Report to
the New Brunswick Department of Fisherie s
and Environment, 22 pp. Halifax: Applied
Mar i ne Research Ltd. 1973
North , W. J .: Urchin predation . A. Prog . Rep .
Calif . I nst . Technol. 1964- 1965 , 57-59 ( 1965) .
(See also annual progress reports since this
date)
Ogden, J.C . , R.A. Brown and N. Salesky : Grazing
by the echinoid Diadema antillarum Phillipi:
formation of halos around West Indian patch
reefs. Science, N.Y . 182, 715-717 (1973)
Ormond , R. F . G. and A.C. Campbell : Observations
on Acanthaster planci and other coral reef
ech i noderms in the Sudanese Red Sea. Sy mp .
zool . Soc. Lond. 28, 433- 454 (1971)
Paine, R.T . and R. L . Vadas : The effects of
grazing by sea urchins, Strongylocentrotus
spp . , on benthic algal populations. Limnol .
Oceanogr . 14, 710-719 (1969)
Date of f inal manuscrip t acceptance : Octobe r 6 , 1975 . Communicated by T . R . Parsons, Vancouver
Scarratt, D.J.: Investigations into the effects
on lobsters of raking Irish moss, 1970-1971 .
Tech . Rep . Fish Res . Bd Can . 329, 1-20
( 1972)
The effect of raking Iris h moss (Chondrus
crispus) on lobsters in Prince Edward Island .
Helgolander wiss. Meeresunters. 24, 415 - 424
(1973a)
Abundance, survival and vertical and diurnal
distribution of lobster larvae in Northumber-
land S t rait, 1962-63, and their relationships
with commercial stocks . J. Fish . Res . Bd Can .
30 , 1819 -1 824 ( 1973b)
and R . Lowe: Biology of rock crab (Cance r
irroratus ) in Northumberland Strait . J . Fish.
Res. Bd Can. 29, 161- 166 ( 1972)
Shepherd, S . A . : Competition between sea urchins
and abalones. Aust . Fish . 32 (6) , 4- 7 (1973)
Sutcliffe, W. H . , Jr .: Correlations between
seasonal river discharge and local landings
of American lobster (Homarus americanus ) and
Atlantic halibut (Hippoglossus hippoglossus)
in the Gulf of St . Lawrance . J . Fish. Res .
Bd Can . 30, 856 - 859 (1973)
Wilder, D. G.: The growth rate of the American
lobster (Homarus americanus) . J . Fish. Res .
Bd. Can. 10, 371- 412 (1953)
Lobster conservation in Canada . Rapp. P. - v .
Reun. Cons . perm . int . Explor Mer . 156, 21 - 29
(1965)
Dr. P . A . Breen
Pacific Biological Station
P.O . Box 100
Nanaimo V9R 5K6 , B . C .
Canada