Intraspecific Competition in Petella cochlear Born

Author(s): G. M. Branch
Source: Journal of Animal Ecology , Feb., 1975, Vol. 44, No. 1 (Feb., 1975), pp. 263-281
Published by: British Ecological Society

Stable URL: https://www.jstor.org/stable/3862

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 263

INTRASPECIFIC COMPElvllvlON IN PATELLA COCHLEAR

BORN

BY G. M. BRANCH

Department of Zoology, University of Cape Town

INTRODUCTION

Competition results from the requirement by two or more individuals of a common

resource which is, or may become, limiting to these individuals. The limitations which
may result can affect a variety of functions such as growth, size or mortality, but the
most important limit is that placed on reproductive potential. Closely related species
usually have the greatest overlap in the resources they require, and are more likely to
compete if they are coexistent. This reaches its highest intensity in intraspecific relation-
ships where the requirements of the individuals may be identical. In limpets, food and
space
be are the major resources competed for. Obviously the intensity of competition will
density-dependent.

The present work is concerned with the quantitative effects of competition on Patella
cochlear Born populations. Of all the South African limpets, P. cochlear is the most
obvious subject for such an analysis, because the populations vary in density from 90 to
1700 per square metre: no other limpets experience densities of this order. Furthermore,
populations of P. cochlear are stable in time, thus enabling mortality to be estimated
from the composition of the age groups. This stability also increases the accuracy of
predictions made from the results. A comparable analysis was also completed on P.
granularis
barnacle L., but will be considered in a separate paper in relation to competition with
populations.

MATERIALS AND METHODS

Most of the work was done at Dalebrook and Kalk Bay in the Cape Peninsula. Zonation
and densities
measured were assessed
with a turbulometer by line transects and random quadrats. Wave action was
(Field 1968a).

Growth rates were obtained by measurement of labelled animals, and the details of this
technique have previously been described (Branch 1974b). Reproductive output was
determined for subsamples of random populations by calculating gonad indices (between
somatic and gonad weights) just prior to and just after spawning. (See Branch 1974a for
details
related of the method.) From the results, the output of eggs per square metre could be
to density.

Deductions about mortality are based on the composition of age-groups in populations

isof known
based growth
on that of rate.
The technique has previously been described (Branch 1974b) and
Deevey (1947).

COMPETITION

Densities of Patella cochlear may vary considerably, but numbers up to 1700 per m2 have
been recorded. The numbers are closely related to wave action, highest densities being

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264
//
Intraspecific competition in Patella cochlear Born

found in regions of fairly strong wave action. Ebling et al. (1948) have shown that Patina
pellucida (L.) also occurs where water movement is fairly strong, and Atapattua (1972)
remarks that in Ceylon Cellana occurs only on wave-washed shores. Patella argenvillei Kr.
also occurs predominantly where wave action is fairly strong.
As wave action decreases, so do the densities of P. cochlear, and in sheltered waters it is
totally lacking. Here, the lower shore may become covered by an algal turf which is
normally excluded by limpet browsing. The converse may also be true: that dense algal
growth in sheltered areas excludes the limpets. The competition between diffierent algal
species under different conditions of wave action may also be important in limiting P.
cochlear. The main foodplant of P. cochlear, lithothamnion, is overgrown in sheltered
areas and disappears as a result. Experiments will be described below which suggest the
limpets are responsible for maintaining the lithothamnion and preventing its overgrowth.
In the absence of P. cochlear, P. Iongicosta Lam. may penetrate into these sheltered areas.
Under extreme wave action, P. cochlear is replaced by mussel beds, particularly those of
Aulacomya ater (Molina) or by Octomeris angulosus Sowerby.
As Ballantine (1961) has pointed out, the prevailing wave action of a shore is often most
easily and accurately defined in terms of the biological components present, and in this
respect Patella cochlear is a particularly good indicator species.
Wave action is difiicult to quantify, but Field (1968b) has shown a convincing numeri-
cal correlation between wave action and numbers of P. cochlear. In the present work a
turbulometer (Field 1968a) was used to measure wave action on a short term basis, and its
relationship to P. cochlear densities is shown in Fig. 1. Although the data are scattered, a
significant linear relationship exists over the central range of wave action. Extreme wave
action could not be measured.

600 -

400- /
/

,% _ / @
c /
8 / @

200- //

s / @

s i/ ss
/
O 8 I O _< 1 1. 1 1 . I
3 4 5 6 7 8 9
Wave action

(arbitrury turbulorneter units)

FIG. 1. Relationship between wave action and density of Patella cochlear.

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 G. M. BRANCH 265

The intensity of intraspecific competition must obviously be related to density. In P.

cochlear an unusual condition exists in that increasing numbers of juveniles are found on
the shells of adults, as the density rises. Frequently, smaller juveniles live on the larger
juveniles in a multi-tiered arrangement. As lithothamnion covers the shells, these provide
both substrate and-food for the juveniles. However, the juveniles do not restrict their
feeding to the shell surface, and crawl down on to the rocky substrate to feed. The result is
that competition is almost 'two-layered': juveniles compete on the shells, while all size
groups compete on the rocky substrate. As the size groups all compete to some extent, it is
not practical to separate their interactions. Furthermore, the space occupied by each
animal will be devoid of food, so that food available on the shell will merely compensate
for that which is lost because of the physical presence of the animal. Thus the amount of

O I I I I 1 1
20 40 60
Length mm

FIG. 2. Accumul
from left to right, * * 1225; * ff ff 355; *--- * 345; o- - - - o - - - -
365;o o-200;ff- 100;t A 125.

food remains constant, but will be shared by incr

increases. Consequently competition for food has
its effect on all limpets present in a given sam
eliminates competition for space, but it can only
food, and not eliminate it. Its effect can be me
size, growth rate, standing crop, mortality rates
As density increases, the mean size of P. coch
accumulative percentage size frequency shows thi
ponderance of smaller animals in high-density po
ties have approximately equal numbers of juvenil
contain a majority of larger, older animals. In

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266 Intraspecific competition in Patella cochlear Born

juveniles is greatest in the high-density populations. The maximum size (taken as a mean
of the five largest animals) is also negatively correlated with density (Fig. 3), and the
regression of the relationship is:

y = -0 017x+70 1

(where y = maximum size and x = density).

The y intercept (701 mm) should give an estimate of the maximum size attained by
P. cochlear at Dalebrook, and does in fact correspond closely to the largest recorded
animal (72a3 mm).
70- 0

o\ o

E w
E 9_
c C o

a) \

250-

40-
X- r 125
x x x

/>
, cq

x/ -oo X

I cs

X I _ 75 E
I M

/ x

/ -50
/ I l l l l I
0 200 400 600 800 1000 1200

Density/m2

FIG. 3. Maximum length (o) and biomass ( x ) of Patella cochlear relative to density.

More important than size is the standing crop which can be supported in relation to
density. Fig. 3 shows the relationship between biomass (dry flesh weight per m2) and
density. As density increases, so biomass must increase. If competition is lacking the two
will be related in a linear fashion and at densities below 250/m2 this is the case.
At densities of above 450/m2, a limit is reached to the standing crop which can be
supported by available resources. The biomass then remains static at about 125gm/m2
despite further increases in density. Competition must become progressively more and
more intense as the resources are distributed among more and more animals. Between
densities of 250 to 450/m2 competition must exist, but at a relatively lower intensity.
The most important aspect of competition is its effect on output of gametes, for this has
a direct influence on recruitment, and hence the future abundance of the species. This
effect may be direct in that the relative size of the gonad is reduced; alternatively (or
additionally), it may be indirect and simply reduce the mean size of the sexual individuals.
As has previously been shown (Branch 1974a), larger animals produce proportionally
more gametes, so that any reduction in size will effectively reduce gamete output.
An analysis of five sites at Dalebrook was undertaken just before and just after
spawning (in April and October respectively) to assess gamete output relative to density.

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 G. M. BRANCH 267

At one site the data were too scattered to give significant results and this site is omitted.
Results from the remaining sites (A, B, C and E) are summarized in Figs 4a and b, and
show that density has little direct effect on the output: gamete output relative to body
weight was constant in all four sites. A slightly greater output occurred at Site A (density
125/m2) but was not statistically higher than the other sites. Regressions for these data
and statistical comparisons of slopes and intercepts are given in Appendix I. As no
significant differences existed, the data were pooled and total regressions calculated for
mature ovaries (April) and spent ovaries (October).

- (a)

A
Totol

0,6-

3
0,4-
o

0,2-

t l l
A _ 6

0,2- (b)

C A Totol

.. w
3
0,1 -
o

l . l l l

2 4 6

Somotic wt gm

FIG. 4. Reproductive output of Patella cochlear at four sites (A, B, C, E). (a) mature gonad;
(b), spent gonad. Density: A, 125/m2; B, 220/m2; C, 320/m2; E, 720/m2.

The relationships were as follows:

Mature ovaries: y = 0 1600x-0 2147

Spent ovaries: y = 0 0444x-0 061 5

(y = gonad weight, x = somatic weight).

The assumption was then made that these relationships are true for all the populations
subsequently analysed and that the difference between the two regressions is a reflection

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 268
1I
1,
1. I I I
IntraspeciJ5ic competition in Patella cochlear Born

of average gamete release in relation to somatic weight. Shell length (1) is related to body
weight (w) by the exponential function:

W = Clb = 0*0001537 1 51

From this, shell length was related to gamete output (Fig. 5). This relationship allows
analysis of numerous large samples, because shell length is the quickest dimension to
measure to assess the composition of a population. In addition, the output which was
measured directly at sites A, B, C and E has been included and agrees well with the calcu-
lated output (Fig. 6).

0,8- /

0,6 /
I

zs I
O 0X4 /
I
I

0,2- /

0 20 40 60
Length mm

FIG. 5. Patella cochlear shell leng

Calculations for nine sites of vary

weight) decreases almost linearly a
average size of the sexual indivi
percentage of juveniles in the popu
total output (per unit area) initi
density
density.
of about 430/m2 and then

Thus increased competiton at higher densities not only establishes a maximum stand-
ing crop, but sharply decreases the output of eggs.
The effect of competition on growth rates was directly measured at three sites, ranging
in density from 90 to 620/m2. Figs 7a, b and c show the increment per six months,
relative to initial length. Regressions were calculated for each site to give the mean growth
(Appendix II). These values could then be used to obtain the shell growth and body
growth throughout life (Figs 8a and b). This presumes that the densities remain constant
from year to year, as has already been demonstrated for other P. cochlear populations
(Branch 1974b).

Quite clearly there is a dramatic reduction of growth rate as density increases. This
will, of course, reflect back on the reproductive potential.
From the size of different age groups (Fig. 9) estimates can be made of mortality

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 G. M. BRANCH 269

between one age group and the next. These estimates are only reliable if the population is
stable in time (Deevey 1947). Stability has been demonstrated in medium-density popula-
tions (Branch 1974b), but it is possible that low-density populations are more subject to
the vagaries of random settlement. For this reason, the value of survivorship curves must
be comparative and not absolute.
Several features are apparent from the survivorship curves (Fig. 10). Firstly, as density
increases, there is a proportionally greater mortality of juveniles. If mortality in the firs
20% of the life span is ignored, the curves become remarkably similar. Density then

012
-66

55- 010

44- 008 2

33- 006 8
o

oF
0F
22- 004

ll 002

I i I I I I
220 440 660 880 I 100 1320

Density/m2

FIG. 6. Gonad output in Patella cochlear, relative to density. Gamete output per m2: ,
measured; o, calculated. Gamete output per unit biomass; , measured; , calculated.

effects survival of juveniles rather than adults. This is principally because as density
increases, a large percentage of the juveniles are found on the shells of adults (Fig. 11).
Previously it has been shown that massive mortality occurs during transition from shell to
rock (Branch 1974b), so that the population of juveniles settling on shells will radically
affect the mortality pattern. In this way, the mortality of juveniles is density-dependent,
while, once a scar has been established on a rock, mortality ceases to be a function of
density.
The second trend that is important follows logically from this. High-density popula-
tions will have proportionately larger numbers of juveniles. Both the population structure
(Fig. 9) and the accumulative size composition (Fig. 2) demonstrate this.
At densities of (about) 450 per m2, the standing crop reaches a plateau. As densities
increase above this value, the supply of foodplants remains relatively constant, because it
coats the limpet shells. A constant biomass is therefore supported by a constant food

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270 IntraspeciJ5ic competition i

supply, so that competition for fo

further. Competition for rock s
resultant increase in juvenile mort
mean size, and in reproductive pot

(D

6B

660/m2

4 - \

2
-\S\
\.\ *\

l 'm s\ l l
20 40 60 80
Initial length mm

FIG. 7. The effect of density on the growth of Patella cochlear.

DISCUSSION

In the concentrated populations of Patella cochlear, considerable in

ha$ been demonstrated. Crowding results in reduced growth, lowe
sizesg higher mortality of juveniles, and more importantly, a red

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 G. M. BRANCH 271

unit area). In populations of less than 250/m2 competition is slight, although growth
rates may be moderated by density. Competition is most intense in populations of over
450/m2, because at this density biomass reaches a ceiling. Above this value, gonad output
drops dramatically. Some aspects of these results parallel those of Sutherland (1970),
working on Acmaea scabra Gould. Analysing high and low level populations, he showed
that high on the shore (at the 'edge' of the range), growth is greater, mortality lower and
biomass higher.

(a)
6-

,/,o,/--'i 1 1 1

(b)

60-
4 _ _--

q / / P

l l l l
5 lo 15 20 25
years

FIG. 8. Growth of Patella cochlear at various densities: s 9°/m2; ---, lsO/m2;

- - - -, 660/m2. (a) total wet flesh weight, (b) shell length.

Density is related directly to wave action. Quantitative values for settling are impossible
to obtain, but the size of the first-year group is a reasonable indication of success. Table 1
summarizes this, and shows that high density is always related to high rates of settling,
and that this is consistent and not due to random settling in a given year. High density is

Table 1. Size of first-year groups in two successive years, in relation to total

density

Total density Density of first- Density of first-

per m2 year group year group
per m2 (1972) per m2 (1973)
90 19 7
99 18 29
120 29 31
- 198 26 36
341 95 90
- 352 109 120
363 139 131
1210 448 461

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 nqo
L lD
_
r 1 _ m . . FT s
Intraspecific competition in Patella c

in other words, due to a consistently high rat

settlements are due solely to the wave action
response as welI, is not known.

td) + .s + A+++++XA
10- ::

(b) + + t + f + + t + + i +X+s

_ kC'tAS't
| l l I , , I

(d)

+ + ttii itAiXli4w

30 40 50 60 70
t ength rnm

FIG. 9. Size frequency distribution in Patella cochlear, in relation to density per square
metre: (a) 90lm2; (b) 150lm2; (c) 6601m2; (d) 12101m2. Arrows indicate calculated year
groups, determined from measured growth rates.

Thus in Patella cochlear there exists a feedback mechanism controlling overall popula-
tion levels. Gonad output and survivaI are highest in low-density populations, but settling
is greatest in areas of wave action, resulting in high-density populations in which low
growth, high mortality (particularly of juveniles), small $ize, and low gamete output are all

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 G. M. BRANCH 273

features. The problems of density cannot be ameliorated by emigration, because P.

cochlear occupies such a narrow zone on the shore, and because they 'home' rigidly to a
fixed scar.
The numbers of P. cochlear are clearly self-regulated, provided one can assume the

1000-x :,*s

I m.ss * ..
500- !si Xx xvx x x... (a)
a, 'xs *

axh x"x *X

Q\ %
\ \ xs

5- V x.

a \ "<\.
g I I I 1-g-2
2g. 20 40 60 80 100
a Age as % of maximum age

500 >'x . (b)

\; 'X t x

100- wQ \ 'X.X

50- tl\ @ 'x

% :
's
\
*: *. x

10- *E *. \

\ \ * \

1 OY' \
*' *L%& x I
5 iO 15 20 25

Age (yeors)

FIG. 10. Survivorship curves for Patella cochlear at various densities: * , 90/m2;
x- - - x, 150/m2; o- - - - o, 660/m2; * *, 1210/m2. (a) Age expressed as a percentage
of maximum age, (b) age in years.

quantity of gametes released is related to the number of larvae which will subsequently
settle. It would be naive to assume a directly proportional relationship, when other
factors, such as intraspecific larval competition, are likely to influence numbers, but there
are reasons for suggesting that gamete output and larval settling rate are correlated. Firstly,
L. Hutchings (personal communication) has shown that at least on the west coast of
the Cape Peninsula, zooplankton are not responsible for 'grazing out' the phytoplankton,

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274
-
Intraspecific competition in Patella cochlear Born

so that larval competition for food is unlikely to be a limiting factor. Secondly, nearly all
the Patella spp. spawn at the same time as P. cochlear (Branch 1974a), so that if competi-
tion is occurring, it will be most acute on an interspecific rather than an intraspecific level.
Reduced gamete output would thus mean fewer P. cochlear larvae to compete with the
remaining species. Thirdly, there is evidence from an area of oil pollution, where mortality
resulted in reduced gamete output (per m2) about four months after the oil spill: larval
settlement was correspondingly reduced. Subsequently the sparsely distributed survivors
grew rapidly, reached a larger maximum size, and achieved a higher gamete output,
associated with a three-fold increase in larval settling (Branch 1973). Finally, selection is
likely to favour an 'optimal' release of gametes which on a long-term basis allows for the
wastage associated with broadcast fertilizers, but not for 'excessive' gamete production
over and above this. If this is so, then again a reduction in the amount of gametes liber-
ated will influence the numbers of larvae subsequently settling.

70 -
tn t

8 60- */

g 50- * /
*
qO 40- +
S 30- S
20- *
I I . I
500 1000 1500

Density m2

FIG. 11. Patella cochlear: the relationship between density and the percentage of juveniles
living on the shells of older animals.

Fig. 12 represents the balance which exists between sparse and dense populations. In
this way a dynamic equilibrium is maintained, and the total population remains stable.
Variations in density are local and related to prevailing wave action.
In many ways this is analogous to Chitty's (1967) hypothesis, put forward to explain
the 'crashes' which occur in vole populations. He argued that as density rises, mutual
interference between individuals also increases, resulting in a reduced rate of increase and
selection for aggressive animals which are more likely to survive the mutual interference.
Obviously this is the start of a vicious circle which intensifies until the population
crashes. In the subsequent low-density populations, interference is lacking, selection
occurs against aggressiveness, and the population again builds up to repeat the cycle.
Chitty's hypothesis is an explanation of temporal variations in populations, while in
P. cochlear the variations are spacial and overall constancy is maintained in time. Some
limpets react aggressively when they encounter other members of the same species
(G. M. Branch, unpublished), pushing against them until they retreat. In P. cochlear it is
difficult to conceive of short-term selection for or against aggressiveness, because with
external fertilization and larval dispersal, larvae will probably settle under different
conditions to those experienced by the parents. Selective pressures are likely to change
from generation to generation. On a long-term basis, aggressiveness is likely to be selected
against because it would lead to waste of energy in crowded populations. This does appear
to have happened, for intraspecific reactions are notably mild in P. cochlear (as will be

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 G. M. BRANCH 275

described in a later paper). In denser populations, interference must occur between

individuals; for when feeding they will encounter and react to an increasing number of
other animals.
One of the interesting features in P. cochlear is that increasing density does not reduce
the gonad size (relative to body size) (Fig. 4), while growth is much reduced (Figs 7 and 8).
This suggests that gonad growth takes precedence over somatic growth. It has previously
been suggested that male limpets grow more slowly because of their proportionately
larger gonads (Branch 1974a). Evidence is also available that in P. glSanularis somatic
growth is curtailed if food is short, while gonad development is relatively unaffected.
Fig. 14 is a semidiagrammatic three-dimensional view of part of the shore at Dalebrook,
indicating the zonation and numbers of P. cochlear. The shore shelves gradually to just

Wesk wave action Strong \:vave action

Settlement low Settlement high
i <
DENSITY LOW DENSITY HIGH

Juvs on rock | \eSpacing Spacing > ; Juvs on shells

/ rundom A uniform X
/ Resources /\ Absdute food \
/ adequate , shortuge \
# gompetition ow Larvae Competition high < <
Juv. mortality ; , , Juv. mortality
low \ Rapid growth A Low (lrowth high
\ Large size Small size /

<;
High
'' + /
gonodial Low gonadial
oj< Soutput

FIG. 12. Gonad

above low water

The outer edge
shelf above is w
cochlear are den
shore is protect
indicate a med
samples are in
ceiling, and gonad output is suboptimal.
At first sight it appears strange that much of the total population exists under 'self-
imposed' competitive stress, and that the self-regulatory mechanism has not been selec-
tively refined until densities are optimal with respect to intraspecific competition.
There are probably three reasons why this has not occurred. Firstly, recruitment occurs
from low-density populations to offset low gamete release in high-density populations,
thus enabling the situation to continue.
Secondly, lithothamnion is partly maintained by the activities of the limpets because
they graze away other algae and settling spores preventing overgrowth by other algae.

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276 Intraspecific competition in Patella cochlear Born

Algal spores are frequently found in the gut contents. At limpet densities of less than
300/m2, algae begin to develop on the rocks in the P. cochlear zone and competitively
displace the lithothamnion (Fig. 13a). Higher densil;ies prevent this, aided by the greater
wave action which tears away the less robust erect algae.

- >

o-
.s
n
-

c c
C5

Q:
o

LL

>t

>s
*fi5

a)

A,

O 0,5 1 2 3 4 5 6 7 8 10 15 20
DenSity Of Ecoch/ew xD2/m2

FIG. 13 (a) Histogram of the frequency of Patella cochlear dens

between P. cochlear density and the percentage algal cover
which is able to develop despite limpet grazing. (b) Hypothet
petition between members of the P. cochlear population. A: Lim
too dispersed; B: co-operation increases relative to density; C: c
mum; D: intraspecific competition limited or absent; E: compet
reaches a maximum and competition (particularly in terms of
intense.

This raises the question of physico-biological interaction:

do erect algae flourish and therefore prevent P. cochlear s
thamnion, the foodplant of P. cochlear), or conversely do
lished, because of reduced wave action, thus permitting alg

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 G. M. BRANCH 277

possibilities are probably linked in a dynamic balance, as the unpublished work of M.

Wood (1935) suggests. She cleared areas in the cochlear zone to study recolonization, and
showed how a dense covering of algae (especially Ulva sp.) developed in the absence of
Patella cochlear.: :n the subsequent two years, larval settlement was prevented by the algal

Prevailing wave action

l-::::::::l
|-.iiii-.iil >500/m2

250- 500/ m2 Pcoch/ear density

< 250/m2

m
1,0

L.W.S O

-170

I 100/ m2 u 20 °/O cover

I tOO-300/m2 S 20-50 % cover

I -700/m2
FIG. 14 (a) Semidiagrammatic three-dimensional view of Dalebrook shoreline, showing the
relative densities and zonation of Patella cochlear. (b) Transect at Dalebrook, showing
relative distribution of P. cochlear, P. Iongicosta and algal cover.

mat, and lithothamnion disappeared except where stray invading limpets cleared the
algal turf away.
Finally, competition for space occurs between P. cochlear and P. Iongicosta. The details
of this interaction will be described in a subsequent paper, but briefly, P. Iongicosta can
only establish itself in the cochlear zone if P. cochlear densities are less than about 200/m2.

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278 Intraspecific xmpetition in Patella cochlear Born

The transect in Fig. 14b indicates the relationship between these two limpets and algal
cover. Clearly both algae and P. Iongicosta are displaced by P. cochlear, but compete with
the latter when its densities are low enough. Although all the algae have been lumped
together, it must be remembered that different species (with different tolerances) may
occur above and below P. cochlear. Nevertheless, in the absence of P. cochlear (or after its
experimental removal), algae encroach from both above and below into the cochlear
zone: there can be no doubt that the activity of the limpets excludes the algae, and not
merely the tolerances of these algae.
P. cochlear also prevents barnacles settling. Cyprid larvae and small barnacles have
been found in the gut, and clearing experiments show that large numbers of Balanus
algicola Pisbry establish themselves in the cochlear zone in the absence of this limpet
(unpublished notes of M. Wood, 1935). The latter eXect has been strikingly demonstrated
by P. White (1973, unpublished), from whose work Plate 1 is taken.
The density of Patella cochlear must therefore be high to prevent competition or inva-
sion of the zone. High. densities are maintained by recruitment from low-density popula-
tions, the latter existing on the fringe of the habitat at the lower limits of wave action
that can be tolerated. At the centre of the range (high wave action, high density) the
maintenance of the cochlear community is assured unless catastrophic events (such as oil
pollution) modify the environment. At the periphery of the range, interspecific reactions
are stronger and more varied, and the numbers of P. cochlear are likely to oscillate,
depending on whether local conditions favour one or other ofthe species in the community.
Almost certainly, all other limpets will experience similar, if less intense, intraspecific
competition. This is certainly true of P. granularis, P. vulgata L. (W. Ballantine, unpub-
lished), Acmaea scabra (Sutherland 1970) and A. digitalis Esch. (Breen 1973). Competition
is also indicated by the interesting remark Forster (1967, p. 294) makes about Haliotis
tuberculata L.: 'in fished populations the rates of growth are possibly rather higher'
(than in unfished areas).
W. Ballantine (personal communication) has suggested that limpets may simultaneously
compete for food while co-operating to maintain their habitat. Groups of Patella vulgata
collectively prevent overgrowth by Fucus although individually incapable of this but
they may also compete for food.
A comparable situation is true in Patella cochlear. Densities of about 200 to 300/m2 are
required to exclude P. Iongicosta, and of about 350/m2 to prevent algal growth covering
lithothamnion. Thus at levels less than 350/m2, co-operation will increase with density.
while in excess of this, no further advantage is gained by increasing the density. It has also
been shown that intraspecific competition is limited if densities are less than 250/m2, and
most intense when these exceed 450/m2. This relationship is summed up in Figs 13a and
b. Here it is apparent that the median density of P. cochlear falls in the range where co-
operation and competition are balanced. Many populations however exceed this value.
The stimulating work of Paine and others (Paine 1966, 1969, 1971; Dayton 1971) has
shown that experimental removal of top predators from food 'subwebs' results in a reduc-
tion of diversity and often a monopolization of space by a single species. Predation keeps
dominants at a level where other species can still share the habitat. In the case of P.
cochlear, its complete domination of the cochlear zone may be due to relative immunity
from predation, or its potential population growth may be much higher than in other
species, resulting in densities so high that predation is insufficient to prevent monopoliza-
tion. The latter possibility might also explain the 'unnecessarily' high densities in P.
cochlear, which allow the monopoly of space this species enjoys.

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Journal of Animal Ecology G. M. Branch, Plate 1

I Cleared - Control I

(Top) Patella cochlear were experimentally removed from the left-hand side of the plot: extensive settle-
ment of the barnacle Balanus algicola Pilsbry has occurred in this area. On the right-hand side Patella
cochlear were left undisturbed (their shells are covered by coralline algae), and have excluded the
barnacles by their grazing. Reduction 1/5. Photograph by G. M. Branch.

(Bottom) Small Patella cochlear which invaded the experimentally cleared plot (arrowed), grazed away
or forced off the established barnacles. Beyond their grazing range in the lower part of the picture,
barnacles are still present. Reduction 2/3. Photograph by G. M. Branch.

(Facing p. 278)

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 G. M. BRANCH 279

In all the limpets mentioned above, density in the centre of the range is such that intra-
specific competition is considerable. This in itself raises interesting possibilities-both
ecological and commercial. How general is the phenomenon that intraspecific competition
is greater than 'desired' ? And is this effect a necessary corollary to maintaining the species
in the face of interspecific and predatory pressures ?
These questions have basic ecological implications for population control mechanisms,
but their formulation into general principles must await data from a far wider range of
animals.
In his excellent review on competition, Miller (1967, p. 4) succinctlyraises the problem
that 'there is still no satisfactory explanation for the tremendous diversity of species. . .
Observed in natural systems'. The answer is likely to be complex, but one facetis suggested
from the present work. For one species totally to exclude another it must saturate the
habitat completely. This has been observed in the centre of the cochlear zone: but here
intraspecific competition reduces gamete output to the point where the population
would crash if recruitment from elsewhere did not occur. If this were true throughout the
limpet's range, the population could not be maintained, and saturation of the total
habitat is thereby prevented. If intraspecific competition cannot rise beyond certain levels
without self-regulation occurring then it may be impossible totally to exclude other species.

ACKNOWLEDGMENTS

Working under my supervision, Miss P. White showed the effects of rem

cochlear from selected sites and Plate 1 was taken at her experimental si
J. H. Day kindly criticized the first draft of this paper.

SUMMARY

(1) The density of Patella cochlear varies from 90 to 1700/m2, and is cor
wave action. Highest densities occur with moderate to strong wave action.
(2) As density increases, mean and maximum sizes decrease, growth rate is lo
mortality of juveniles is greater. The latter is partly because a greater pr
juveniles occur on shells of other limpets, and mortality is high during their tr
rocky substrate.
(3) Standing crop increases with density until the latter reaches about 450/m
this value, biomass remains static at about 125 gm/m2 despite further
density.
(4) This suggests that competition is most intense above this density.
(5) Density has no direct effect on gonad size, but at above 430/m2 gonad output
m2 decreases relative to density because mean size is reduced.
(6) Low densities are associated with high gonad output, high densities with low outpu
This results in a negative feedback controlling overall numbers.
(7) High-density populations remain high because of recruitment, settlement be
highest where wave action is strong.
(8) Intraspecific competition is slight at densities of less than 250/m2 and most inten
over 450/m2. The median density at Dalebrook is 300-400/m2, but many populatio
have higher densities.
(9) Densities of about 300/m2 are required to exclude other limpets and to prevent th

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280 Intraspecif c competition in Patella cochlear Born

foodplant lithothamnion from being overgrown by other algae. To maintain monopoly of

the cochlear zone, high densities of P. cochlear are required.
(10) Concomitantly intraspecific competition is high, but it is suggested that this is
balanced against co-operation promoting exclusion of other limpets from the community
and maintenance of the foodplant.

REFERENCES

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shores. Field Studies, 1(3), 1-19.
Branch, G. M. (1973). The biology of Patella cochlear with reference to oil pollution. S. Afr. natn.
oceanogr. Symp. Abstr., Aug. 1973: 18.
Branch, G. M. (1974a). The ecology of Patella Linnaeus from the Cape Peninsula, South Africa. II.
Gonadial cycles. Trans. roy. soc. S. Afr. 41 (2), 111-60.
Branch, G. M. (1974b). The ecology of Patella Linnaeus from the Cape Peninsula, South Africa. III.
Growth rates. Trans. roy. soc. S. Afr. 41 (2), 161-93.
Breen, P. A. (1972). Seasonal migration and population regulation in the limpet Acmaea (Collisella)
digitalis. Veliger, 15, 133X1.
Chitty, D. (1967). The natural selection of self-regulatory behaviour in animal populations. Proc.
Ecol. Soc. Australia, 2, 51-78.
Dayton, P. K. (1971). Competition, disturbance, and community organisation: the provision and subse-
quent utilisation of space in a rocky intertidal community. Ecol. Monogr. 41, 351-89.
Deevey, E. S. (1947). Life tables for natural populations of animals. Q. rev. Biol. 22, 283-314.
Ebling, F. J., Kitching, J. A., Purchon, R. D. & Bassindale, R. (1948). The ecology of the Lough Ine
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Ecol. 17, 223-44.
Field, J. (1968a). The 'turbulometer' an apparatus for measuring relative exposure to wave action on
shores. Zool. Afr. 3, 115-18.
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shore samples in False Bay, South Africa. Zool. Afr. 3, 119-37.
Forster, G. R. (1967). The growth of Haliotis tuberculata: results of tagging experiments in Guernsey
1963-65. J. mar. biol. Ass. U.K. 47, 287-300.
Miller, R. S. (1967). Pattern and process in competition. Adv. ecol. Res. 4, 1-74.
Paine, R. T. (1966). Food web complexity and species diversity. Amer. Nat. 100, 65-75.
Paine, R. T. (1969). The Pisaster-Tegula interaction: prey patches, predator food preference and inter-
tidal community structure. Ecology, 50, 950-61.
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(Received 16 May 1974)

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 A >0 05 >0 05 <0e02 >0 05
G. M. BRANCH 281

Appendix I. Relationship between somatic weight (x) and gonad weight (y)
forfive sites at Dalebrook from mature and spawned samples

Site Regression Correl. P

coeff. r

Mature A y = 0 173x-0-204 0 94 <0-001

(April 1972) B y = 0 143x-0-196 0-76 <0-02
C y = 0 190x-0-308 0 97 <0-001
D y= 0070x+0030 050 >005
E y = 0 107x-0-144 0-91 <0-02
Total y = 0 160x-0-214 0-87 <0-001

Spent A y = 0045x-0-067 091 <0-001

(October 1972) B y = 0050x-0094 077 >005
C y= 0053x-0-068 0-76 <0-02
E y = 0038x-0a052 0a66 >005
Total y = 0044x-0a061 0a81 <0a001

Significance of difference in slopes (above diagonal) and intercepts

(below diagonal)

Mature A B C D E

B >005 \ >005 >005 >005

C >005 >005 \ >005 >005

D >005 >005 >0 05 \ >005

E >005 >005 >005 >005 \

Spent: A B C E

A \ >005 >005 >005

B >005 \ >005 >005

C >005 >005 \ >005

E >005 >005 >005 \

Appendix II. Regressions of increment in shell length (per six months) (y),
against initial shell length (x)

Density Regression r P
A 150/m2 y=-0080x+485 079 <0001
B 9o/m2 y =-0 113x+7-708 0-88 <0 001
C 660/m2 y=-0070x+321 0-82 <0-001

Tests of difference in slopes (above diagonal) and intercepts (below)

A B C

A \ <005 <005

B <0-01 \ >0 05

C < 0-01 < 0 01 \

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