Further Considerations of Competition and the Ecology of the Ants Lasius flavus (F.) and L.

niger (L.)
Author(s): A. J. Pontin
Source: Journal of Animal Ecology, Vol. 32, No. 3 (Oct., 1963), pp. 565-574
Published by: British Ecological Society
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 565

FURTHER CONSIDERATIONS OF COMPETITIONAND THE

ECOLOGY OF THE ANTS LASIUS FLAVUS (F.)
AND L. NIGER (L.)
BY A. J. PONTIN

Zoology Department,Bedford College, Regent's Park, London, N. W.1

In earlier papers (Pontin 1957, 1960, 1961a, b; Odum & Pontin 1961) work on popula-
tions of these ants in 1956-58 was described and the following account is of population
changes observed after a lapse of 5 years and discussion of some aspects not previously
covered.
It is convenient to divide the paper into two parts. The first deals with the history of
the populations from early colonization to eventual extinction, and the second deals
with stabilization of mature populations where plant succession has been checked at a
suitable stage.

Table 1. The density per square metre of colonizingqueens and

mature colonies of Lasius flavus and L. niger
Locality L. flavus L. niger Reference
rNorth Oxford,bare ground 1P6 2-0 Pontin (1960)
Colonizing queens North Oxford,Brassica plot 0 33 044 Pontin (1960)
tTubney, quarry 0-083 0A44

rBowling Alley, Wytham 0-26 0.01

Mature colonies Wytham Park 0-25 -
Silwood, Berks. 0-15 - Richards & Waloff (1954)
ISilwood, Berks. 0-18 - Waloff & Blackith (1962)

POPULATION HISTORY
The populations studied occupied an area of calcareous grassland at Wytham, Berkshire,
and some of their history may be reconstructed. This area, known as the 'Bowling Alley'
(also area'C' in Duffey 1956 and 1962), was used as a pheasant rearing ground up to the
outbreak of the Second World War (Duffey, unpublished). The vegetation is still
markedly different from similar, but undisturbed, areas and contains many herbs
characteristicof dry stony ground. These include: Thymusdrucei, Viola hirta, Hypericum
hirsutum, H. perforatum, Cynoglossum officinale, Verbascumthapsus, Pastinaca sativa,
Hieraciumpilosella, Carlina vulgaris and Brachypodiumpinnatum. There were still many
superficial stones in 1957-far more than used by the ant colonies- and there may well
have been little else left on the area immediately after pheasant rearing ceased. Pheasants
used the hills of Lasius flavus (F.) for dust bathing and destroy them, leaving large
craters, before changing to another. Artificial rearing of pheasants has not been resumed
at Wytham so the present populations date from 1940 when the area must have been
devoid of ants as the result of disturbance of the surface, loss of soil and removal of
plants which are the ultimate source of most of the ants' food.
Queen ants of L. flavus and L. niger (L.) alight upon bare areas after mating and the
lack of shade favours early colony growth (Pontin 1960). Table 1 compares the density
of queens arriving in 1 year with the density of colonies observed in old populations.

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566 Competitionin ants
There is no reason to suppose that the density of colonizing queens was any less 20 years
ago and it may be concluded that the Bowling Alley was rapidly colonized by a large
excess of queens during the years following 1940. In 1956 one L. niger and two L. flavus
queens were found alone under stones in the study area, so colonization was still being
attempted then. Late arrivals become fewer as vegetation increases and are likely to be
killed by well-established colonies (Pontin 1960, 1961b; Waloff & Blackith 1962).
L. niger occupies newly bared areas more readily than L. flavus apparently because
early colony growth in sparse vegetation is quicker with a large surface territoryavailable.
This was observed in a number of abandoned quarries in the Oxford district, and an
area of one at Tubney, Berks., may be cited as a good example. This site was devoid of

138 ~~~~~~~N

101~~~~~~~~~~~~1

82~~~~~~~~~~~~~~~~~~~~8

FIG. 1. Map of the researcharea at Wythamshowing the numbersof alate queensproducedin 1957 by
Lasiusfiavusand L. niger.The circlesare theoreticalterritoriesof L. fiavus and are proportionalin area
to the numberof queens produced. The observedundergroundterritoriesof L. niger are indicatedby
irregularoutlines and the interruptedoutline indicates the territorypreviously occupied by a L. niger
colony which was poisoned in 1956. Increasesin territorysince 1956 are diagonallyhatched. Territory
takenoverfromL. flavusbyL. nigeris cross-hatched. Areasof denseBrachypodium pinnatumaredotted.

vegetation and surface soil in 1955 and has been undisturbed since. Table 1 gives the
results of collecting all the dealated queens which could be found after the mating flight
on 13 Sept. 1956. Most of them had burrowed under stones and many must have remained
undiscovered. In 1961 there were many large L. niger colonies producing alate males and
a few queens, but only small L. flavus colonies, none of which were producing alates.
The vegetation was still sparse with Echium vulgare, Oenothera erythrosepala, Reseda
luteola and Agrostis spp. common and the very dry sandy soil was presumably causing
an unusually slow rate of colonization by plants. Possibly the activities of man have
resulted in the common competition of these two species of ants; Lasius niger being
previously adapted to colonize temporary clearings resulting from treefall, while L. favus
is better adapted to live in more permanent grassland.

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 A. J. PONTIN 567
An estimate of the L. flavus population growth-rate may be made by analysis of the
1956-57 results. In Fig. 1 the theoretical territories of the L. flavus colonies are repre-
sented by concentric circles proportional in area to the number of alate queens produced
(see Pontin 1961a). Increases are shown by cross-hatching and the two areas indicated
by straight outlines are used in the following analysis. Increase of a colony may be
accompanied by decrease in its neighbours and a rather complex treatment of the results
is needed to estimate real increases in density. The radius of a territory is proportional
to the square root of the number of queens produced (Pontin 1961a) and therefore the
distance apart of two neighbouring colonies (d) is proportional to the sum of the square
roots of the numbers of queens produced by them (X/N1+X/N2). For each pair of neigh-
bouring colonies a 'crowding index': (A/N1+A/N2)/d was calculated and all possible

Table 2. Aphidae and Coccidae occurringwith Lasius flavus and L. niger

within the area shown in Figs. 1 and 2 at Wytham,Berks., in 1956-62
Usual position on plant
Host plants (all underground)
APHIDAE
Forda formicaria v. Heyd.* Carex, Brachypodiumpinnatum and Internodes
other grasses
F. marginata (Koch) Festuca, Agrostis and other grasses Internodes
Geoica utricularia (Pass.)* Grasses Roots
G. setulosa (Pass.) Grasses Roots
Baizongia pistaciae (L.) Grasses Roots
Anoecia sp(p). Grasses Nodes and stem bases
Neanoecia krizusi C.B. Grasses Nodes
Paranoecia pskovica (Mordv.) Carex Nodes
Tetraneura ulmi L. (Hal.)* Brachypodium pinnatum Stem bases, roots in early instars
Dysaphis bonomii (H.R.L.)* Pastinaca sativa Root collar and basal petioles
Aphis chloris Koch.* Hypericum perforatum and H. Stem bases
hirsutum
A. poterii (C.B.) Poterium sanguisorba Stem bases
A. jacobaeae Schrk. Senecio jacobaea Stem bases
Trama troglodytes del Gu. Compositae Stem bases and roots
Neotrama caudata (del Gu.) Compositae Roots
Smynthurodes betae Westw. Various dicotyledons Roots
COCCIDAE
Ripersia sp(p). Festuca Internodes
* The commonestand most generallydistributedspecies in this particulararea.

neighbours of each colony were included once in each analysis, but to minimize subjective
bias the four nearest neighbours of each colony were arbitrarily selected and any more
than 4 m away were omitted. This includes a number of pairs which probably did not
have meeting territories, but the maximum available information is used and this mini-
mizes the effect of various unaccounted variables. Plotting crowding indices for 1957
against those calculated for 1956 gives the graphs shown in Figs. 3 and 4. It is reasonable
to assume the same unaccounted variability for the two years and the best fitting straight
lines were calculated as described by Quenouille (1952). Those points which were calcu-
lated using pairs of colonies separated by the two main areas which were filling up fall
along a line passing close to the origin (Fig. 3) indicating that increase is proportional to
the initial density rather than the degree of under-population. The regression coefficient
1 42 (P<O 001) must be squared to obtain an estimate of rate of increase in density,

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568 Competitionin ants
........... . . .
, .. . . . ...
. . . . . . . . ,0.. . ...
METRES -* *..
0 1 3 4 ( ......

0f:::e'$ ~..A..w. ~ 4 :9::::

46 ~ ~ ~ 10......

FIG. 2. Map of researcharea showing the numbers of alate queens produced in 1962 Territoriesof
Lasiusfiavus and L. nigerare shown as in Fig. . Black discs show the position of extinctnests of L. flavus
and the areas occupied by dense stands of Brachypodium pinnatumare dotted. The horizontal dashes
indicate areas of tall herbs other than B. pinnatum.

l957 / xX

Xx XXX

* /mX /

U7XXF X XX

X00 X X

1956
FI.3. Graphs of the crowding indices (<Ni +VN2)/d
for Lasiusflavus colonies in 1957 plotted against those
for 1956. Ni and N2 are numbers of alate queens
produced by pairs of neighbouring colonies and
d is their distance apart in metres. The circles and
squaresreferto colonies filling gaps indicatedin Fig. 1
by the same symbols. Equations for the lines are:
y = 1 05x-0*02 and y = 1 42x?0*26.

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 A. J. PONTIN 569
since the crowding index is a linear concept, and therefore the best estimate of exponential
growth of L. flavus colonies under Bowling Alley conditions in 1956-57 is X 2l0 per
annum. Most of these colonies must have been producing alate queens for at least 5 years
before 1956 according to this estimate and may well be much older, but retarded by
competition. The initial growth-rate is probably larger before alates are emitted each
year and further work is in progress on this. Fig. 2 shows the results of counting numbers
of alate queens produced in the area in 1962 and it may be assumed that the same colonies
still occupy the same sites and except where the vegetation has markedly changed many
of them are still producing similar numbers of alate queens per year. The only case of
movement of a colony is the spread westward of the colony producing 155 queens in
1962 and this may be the result of very asymmetrical crowding.

1957

00

10
0
0. 0

10 1956
FiG. 4. Graphsof the crowdingindices (see Fig. 3) for
Lasiusflavus coloniesin 1956 and 1957before andafter
removal of L. niger (filled circles) and in areas of
increasing L. niger (open circles). Equations for the
lines are: y = 3-143x-6-6 and y = 0-395x+0O67.

There was only one unoccupied mound in the area in 1956 and in north Oxford
colonies have been under observation for 16 years so 22 years is not an unreasonable
life for L.flavus colonies and many of the present colonies could be the original founders.
Nothing seems to be known about queen replacement in L. flavus, but since colonies
normally kill introduced queens and they are capable of living at least 10 years in the
laboratory it may be that the original queens are still alive.
The large colonies of L. flavus and L. niger use different methods to overcome the
effects of shading by short herbs. L. flavus constructs the familiar grass-covered mounds
which are firmly bound together by grasses and resist trampling by grazing animals.
The herbage on these mounds rarely dies except in sandy soils in periods of drought, and
superficial stones are buried by the soil brought up by the ants. One summer may be
enough to cover large stones and on the Bowling Alley a 9 x 9 x lyin. (23 x 23 x II cm)
cement slab was buried 3 in. (7 cm) deep in the five years from 1957 to 1962 by a large
colony. The mound increased in height very little as the cement sank into the space left
by the ants' excavations. There is very little correlation between mound size and the

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570 Competitionin ants
number of ants present on the Bowling Alley since the sparse soil is unevenly distributed
and colonies may become smaller after making a large mound. The colony producing
138 queens in 1957 had no mound, but nested entirely under superficial stones. L. niger
on the other hand, usually nests under superficial stones, or if they are not available
makes friable earth mounds which do not resist grazing animals or heavy rain. Plants
are killed where large aggregations of this ant occur, because it bites through roots
which obstruct its constantly changed galleries. In the summer the nests of L. niger are
often made conspicuous by patches of dead herbs with earthworks at their bases. These
dead plants are not usually associated with aphid damage.
Eventually, unless large surface stones are kept uncovered, all areas of grazed grassland
which were initially colonized by both species become pure L. flavus populations. The
pasture of Wytham Park near the Bowling Alley has reached this stage (Table 1).
The major vegetational change seen on the Bowling Alley after a lapse of 5 years from
1957 to 1962 is the spread of Brachypodiumpinnatumwhich is a grass making very dense
stands completely shading the underlying soil. The other vegetation has also thickened,
apparently as the result of loss of rabbit-grazing, but this has not resulted in dense tall
herbage and has not been accompanied by any striking change in the ant populations.
B. pinnatumspread has, however, been accompanied by extinction of those Lasiusflavus
colonies which are completely surrounded by it. Shading must be the important factor
as those colonies on the edge of Brachypodiumpinnatumstands are unaffectedeven though
most of their territories are in the B. pinnatum, and in other areas where there is more
soil available Lasiusflavus builds exceptionally tall mounds into the sunlight and succeeds
in surviving Brachypodiumpinnatumspread. Dense shading of the soil surface completely
prevents the foundation of new colonies (Pontin 1960).
Ecological succession of ant species may occur by temporary social parasitism (see
Donisthorpe 1927) of Lasius niger by L. umbratus(Nyl.), and then L. umbratusin turn
by L. fuliginosus (Latr.). These latter species are more tolerant of shade and L. fuliginosus
normally tends aphids on trees. On numerous occasions in the spring at Wytham L.
umbratusqueens were found under the same stones as L. flavus nests but separated by a
thin wall of earth and it may well be that temporary social parasitism of L. flavus occurs,
but has been overlooked because of the similarity of the workers L. umbratus and
L. flavus.

STABILITY AND COMPETITION

The definition of competition given in Pontin (1961a) is retained although Milne (1961)
has discussed the subject comprehensively and redefined competition as 'the endeavour
of two (or more) animals to gain the same particular thing, or to gain the measure each
wants from the supply of the thing when that supply is not sufficient for both (or all)'.
This definition is impracticable (apart from requiring subjective assessment of animals'
endeavour) because the only way of knowing if the thing is not sufficient in the case of
interspecific competition is by investigation of the effects of the competition on the
competitors, and for an experimental approach the effect must be specified in the
definition. One may as well define interspecific competition concisely as an interaction
which lowers the densities of the competing species compared with the density each
would reach if present alone, or else drop the term altogether.
The population densities of Lasius flavus and L. niger eventualiy reach a stable level
provided that the vegetation is kept short by grazing for a sufficiently long period. In

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 A. J. PONTIN 571
Fig. 3 the close fit to a straight line through the origin with a slope of unity (regression
coeff. 1.05, P< 0O001, when y = 0, x = 002) indicates stability of the L. flavus popula-
tion and, except for the area of Brachypodiumpinnatum increase, the similarity of the
populations in 1956-57 and 1962 (Figs. 1 and 2) confirms that the two competing
populations are stable. Artificial introduction of Lasiusflavus colonies into the territories
of L. niger colonies in 1956 (Pontin 196la) has resulted in L. flavus occupying territory
previously occupied by L. niger and retaining it until 1962. These introduced L. flavus
produce no queens, but maintain numerous workers just like those which were found
occurring naturally with L. niger.
The L. nigercolony experimentallypoisoned in the autumn of 1956 had either recovered
or else been replaced by another in the interval between 1957, when no L. niger were
observed on this site, and 1962.

- - -- - - - - - - -
r2--

cr1

d2 dA
POPULATION DENSITY
FIG. 5. Hypothetical population growth curve showing equilibrium
density (di) reached when only a stabilizing factor is operating and
the equilibrium density (d2) after an additional non-stabilizing factor
is introduced which removes individuals at a rate ri. A factor removing
individuals at a rate >r2 acts as a distribution limiting factor.

The actualdensityattainedby L. flavus in the densestand most uniformarea of the

Bowling Alley was estimated to be 1130 workers/M2 (Odum & Pontin 1961) and the
maximum density of queens produced under these conditions is 15/M2 (estimated from
the regression analysis: Pontin 1961a, p. 49). The maximum density of queens produced
by L. niger on the BowlingAlley is about 14/M2estimatedfrom the observedterritory
size of the largest colony. It does not follow that this is the maximum density attainable
or that a lower density could not be stable. Fig. 5 is a hypothetical population growth
curve similar to that given by Haldane (1953) where the stabilizing factor operates
increasingly severely with density. If the only factor operating on the population were
intraspecific competition, the population would reach a stable density (di) where the
growth-rate is zero. If, however, a factor killing individuals at a constant rate (ri) is then
introduced, the density will have to be reduced in order to replace the loss in the face of
competition, and a new stable density (d2)will be reached. Any sort of factor could affect
the stable density and one which kills more than r2 or reduces reproduction by this much
will prevent the species occupying the area, i.e. it will limit distribution. Shading is an
exampleof such a factorlimitingL. flavus distribution.
L. niger preyson L. flavus as well as exploitingthe same undergroundfood (Table2
J.A.E.-R

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572 Competitionin ants
and Pontin 196lb), but the effect on L. flavus density is not the result of predation alone
for two reasons: (i) L. niger does not eliminate the L. flavus colonies, even within its own
nest area, by predation, and interspecific competition provides the necessary stabilizing
mechanism (Pontin 1961a); (ii) after removal of L. niger the increase in queen production
by L. flavus was much more rapid than L. flavus filling empty territory (P<0-001, see
Figs. 3 and 4) and there was no time in which new workers could be reared to raise the
larger number of queens. Presumably the L. flavus colonies around the edge of L. niger
colonies manage to maintain large numbers of workers which are not preyed on to any
significant extent, but there is less food available near L. niger for rearing the large alate
queens.
The poisoning of the L. niger colony had far-reachingeffects on the other ant colonies.
Neighbouring L. niger colonies showed a significant increase in queen production
(Pontin 1961a) and this increase was in turn correlated with a decrease in queen produc-
tion by the L. flavus colonies around them (P<0 001, Fig. 4). The populations of the two
species are apparently in such close competition that a wide area is sensitive to a local
disturbance and new stable states are reached as may be seen from comparison of the
1956-57 territories with those of 1962 (Figs. 1 and 2).
It is very difficult to give a quantitative estimate of the effect of either species on the
other because subjective selection of the L. flavus colonies affected must be made and
bias is likely. However, a rough estimate may be made allowing for the difference in
biomass of the alate queens of the two species. The total weight of ten samples of ten
queens each from different colonies was 2-57 g for L. niger and 1-75 g for L. flavus; a
ratio of 1-47:1. The large L. niger colony which produced 687 queens in 1956 lost the
amount of food needed to produce from twenty to eighty L. flavus queens and therefore
could apparently have increased by 2-10 % in the absence of L. flavus. On the other hand
there was theoreticaliy room for 300 more L. flavus queens to be produced in the area
occupied by this L. niger colony in 1956, and 286 more queens were in fact produced by
colonies in and around this area when the L. niger was poisoned. On this basis two-thirds
of the L. niger food came from above the surface and the other third came from under-
ground to the detriment of L. flavus. The 1956 pattern was roughly similar in the case
of the other L. niger colonies, but one of them increased its queen production from
forty-eight to 107 in 1957 and this was accompanied by a reduction of 107 in the queens
produced by the neighbouringL. flavus colonies. Perhaps this large reduction is the result
of the extra food intake of the L. niger colony being used to increase worker number as
weli as queen production. Little information may be gained from the third L. niger colony
in this area as it occupies a site with a lower density of ants than elsewhere for reasons
which are unknown.
These ant populations are rather unusual for animals because the nests are static.
Stable interspecific competition is brought about by L. niger having large epigaeic
territories which overlap the smaller underground territories of L. flavus and result in
intraspecific competition which is more severe in either case than the interspecific
competition (Pontin 1961a). It may be worth while to speculate about stable competition
in other animals in general terms, particularlywith respect to similarity of the competing
species.
The conmmonstatement that species occupy 'different niches' or have 'different
controlling factors' and therefore avoid competition leads to confusion since 'different'
in this context could lead one to conclude that the species are separated ecologically
whereas different, but overlapping, niches must be commonplace.

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 A. J. PONTIN 573
Imagine for the sake of argument the absurdity of two identical species-their survival
when living together would be entirely dependent upon chance (Harper, Clatworthy,
McNaughton & Sagar 1961) and it would be possible only to estimate the probability
of one or the other surviving any given period of time. Differences would presumably
be evolved between such species and, depending on the nature of these differences,either
displacement or stability wili follow. If the species exploit a slightly different range of
resources such that intraspecificcompetition in each case exceeds interspecificcompetition
in severity then an infinitesimal difference will stabilize their cohabitation without the
two populations being stabilized by separate factors. The suggestion that independent
control (e.g. Harper et al. 1961) is necessary for cohabitation is incorrect. If on the other
hand one species evolves mechanisms which interfere with the activities of the other
species more than its own then displacement will follow. Probably the Argentine ant
(Iridomyrmexhumilis (Mayr.)) displaces other species (see Elton 1958) because it lacks
colonial boundaries and fights with other species. The barnacles studied by Connell
(1961) perhaps also compete in this way.
If two different species exploiting an ovierlappingrange of resources come to overlap
in range then stable competition is, a priori, more likely than displacement (Pontin
1961a), but the only possible equilibrium point may be at too low a density for one of
them to maintain itself in the face of other factors, and displacement wili follow. Wide
disparity in efficiency of exploitation by the species is an obvious cause of such displace-
ment and this is more likely if they are distantly related taxonomically. It cannot be
maintained that there is a minimum anatomical difference between sympatric species
which permits their cohabitation as has been suggested (e.g. Slobodkin 1961, p. 172).

ACKNOWLEDGMENTS
I wish to thank Mr H. L. G. Stroyan for his help with aphid identification and also
Mr C. S. Elton, F.R.s., and Dr E. Duffey for information on the history of the area
studied.
SUMMARY
1. The history of the ant populations studied at Wytham is reconstructed as far as
possible,
2. The exponential growth-rate of Lasius flavus colonies is estimated to be x 20 per
annum from their queen production in 1956-57.
3. Three types of factor are considered to be important to populations: (i) density
stabilizing, e.g. competition, (ii) distribution limiting, e.g. shading of L. flavus colonies,
(iii) others which affect density without having the properties of (i) or (ii).

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