r. Insect Physiol., 1967, Vol. 13, pp. 699 to 716. Pergmam Press Ltd.

Printed in Great Britain

POPULATION DENSITY, SURVIVAL, GROWTH, AND

DEVELOPMENT OF THE AMERICAN COCKROACH

DENIS R. A. WHARTON, JOHN E. LOLA, and MARTHA L. WHARTON

Pioneering Research Division, U.S. Army Natick Laboratories, Natick,
Massachusetts 01760
(Received 7 November 1966)

Abstract-The mortality rate of larvae of Pwipkzneta americana increases with

increasing population density. The mortality curve is not linear but cascading
and, like growth, mortality alternates with ecdysis. Three major phases of
mortality are recognized: the first, with a high mortality rate during the first to
the sixth week; the second, of brief duration and low mortality, during the sixth
to the ninth week; and the third, of moderately high mortality which persists
until development is completed. The mortality phases are dependent respectively
on (1) a maximal increase in the instantaneous percentage rate of growth and a
high frequency of ecdysis, which expose the larvae to injury and high mortality
when in groups, (2) a slowing down of the growth rate, and (3) sexual develop-
ment and continuing growth in the last instar associated with the metamorphosis
of the larva to the adult. The data support the concept of a homology between the
last larval instar of the hemimetabolous insect and the pupal stage of the holo-
metabolous species.
Isolation retards development of larvae and exerts its retarding influence at all
stages of growth. In the present experiments the optimal density for growth of
larvae is about 25.
The average weight of adult insects tends to decrease with an increase in the
population density of the larvae. On the other hand, the weight-time curve of
emergent adults varies with density of the group.
The results indicate the presence of a factor(s) which affects the rate .of
maturation of the insect throughout its development, which is a function of group
size and is sex-dependent in that males are more sensitive to its influence.
Cannibalism among larvae is observed chiefly with the young and is con-
sidered to be sanitary rather than predatory in character.

INTRODUCTION
THE effects of population density on the survival, growth, and development of many
species of animals vary greatly depending upon species and developmental stage
(ALLEE, 1934). Conditions most suitable for survival may not be those required for
optimal growth. Some species develop most advantageously in isolation, whereas
others respond more favourably when in groups. Food, available space, sanitation,
cannibalism, and certain other factors become increasingly important as the
population density increases. Among insects, population growth has been shown
to be influenced by the counteracting influences of mating and cannibalism
(especially of the eggs). Mating is optimal at moderate densities which permit
opportunity to copulate and result in a favourable rate of egg fertility (PEARL and
PARKER, 1922; PEARL et al., 1927; PEARL, 1932), whereas cannibalism is more

699
700 DENISR. A. WHARTON,
JOHNE. LOLAANDMARTHAL. WHARTON

prevalent the denser the population (PARK, 1932; CHAPMAN 1928; CHAPMANand
WHANG, 1934; MACLAGAN,1932; BOYCE, 1946). PARKet al. (1965) have elaborated
on Park’s studies on cannibalism in Tribolium and find it inherently complex and
involved in every aspect of population growth.
While crowding is unfavourable to the growth of many insects, isolation causes
Tenebrio larvae to grow more slowly than those that are grouped (MICHAL, 1931).
LANDOWSKI(1937) noted that larvae of the cockroach BZatta orientalis L. kept under
uniform conditions grow more slowly when isolated than when grouped, and that
the emergent adults weigh more than adults grown from groups of 2 to 16 larvae.
PETTIT (1940) showed that isolated larvae of BZatteZZagermalzica (L.) also grew
more slowly than groups of larvae and this observation was confirmed by WILLIS
et al. (1958) with several species of cockroaches. CHAUVIN (1946), on the other
hand, stated that isolated larvae of B. germanica grow more rapidly than those in
groups. However, when an optimal ‘vital space’ of 7 ml/larva is provided, groups
of 2 and 5 surpass the isolates in their rate of growth. PIQUETTand FALES(1952)
state that 2 in2 is optimal for the adult Periplaneta americana (L.), however, our
data will show that this area is not optimal for either growth or survival of the larvae
of this species.
The damaging effects of crowding on survival, reproduction, fecundity, and
growth rate whether in insects, tadpoles (ADOLPH, 1931), fish (CERNAJEV,1931), or
mice (CREW and MIRSKAIA, 1931; VETULANI, 1931) have been attributed pre-
dominantly to physical rather than chemical factors (BILSKI, 1921; ALLEE, 1934;
CHAUVIN, 1946), and by some authors to psychological influences (PEARL and
SURFACE,1909; ADOLPH, 1931; VETULANI, 1931). More recently, the work of
BUTLER (1954), NORRIS (1954), L%CHER (1956), and LOHER (1961) in various
species of insects, and of LEE and BOOT (1955), WHITTEN(1956), BRUCE(1959), and
PARKESand BRUCE(1961) in mice, indicates that growth processes may be controlled
to a considerable and often decisive extent by specific pheromones.
Although the biology of the American cockroach, P. americana, has been
studied from many points of view, and the insect is being increasingly used for
various experimental purposes, little has been published about its maintenance in
large numbers. Various types of cages or containers for housing cockroaches have
been devised to control space or labour (PIQUETTand FALES, 1952), but not with-
standing the concern sometimes expressed for overcrowding (GIER, 1947), there is
a general lack of information pertaining to the effect of population density on its
survival and growth from the larval state to maturity.
Preliminary observations with P. americana, which differed from expectations
based on Landowski’s and Chauvin’s findings, together with the growing recogni-
tion of exocrine or pheromone influences, have led to the present study of the effects
of population density in the cockroach.

MATERIALS AND METHODS

The P. amerz’canaused in these experiments were of known age from our own
large colony and maintained on Purina Laboratory Chow pellets (PLC) and water.
 EFFECTS OF POPULATION DENSITY dN AMERICAN COCKROACH 701

The isolated insects were generally kept in 250 ml beakers, and the grouped ones
in 6 x 8 in. or 10 x 12 in. round museum jars which were smeared lightly on the
inside with petrolatum to a depth of 2-4 in. Except where stated to the contrary,
the grouped insects were provided with an inverted cylindrical paper container for
shelter, while the isolates in beakers were without shelter. The larvae were main-
tained either at 29 + 1°C in the dark and exposed to artificial light only when
actually worked with; or at room temperature in the dark, except for about 9 hr
of artificial light during 5 days a week. The larvae were otherwise undisturbed
except once a week when water and food were replenished. Usually the larvae
were counted and weighed every 3 weeks. This interval of handling was used to
minimize injury to the young and teneral larvae, and was adequate for most
purposes. Except where noted, adults were removed from the jars upon emergence
and weighed 1 week later. They were always counted in survival data.

RESULTS

The effect of density on survival and ecdysis

The mortality of newly hatched larvae increased with increasing population
densities up to 1000 (Fig. 1). It is evident from the slopes of the survival curves
that a most severe mortality took place during the first 3 weeks and that this period
constituted the greatest test to the survival of larvae throughout their development.
To determine more precisely the form of the survival curve, daily counts were

80
pooo

$? 70
(75,65,50)

3 60
>-: (225,200,175)

"0 3 6 9 12 I5 I8 21 24

Time, wk

FIG. 1. Survival of different population densities of larvae. Data for groups 15,100,
and 500 are the average of duplicates, and the data for groups 50 and 200 the
average of triplicates. Groups 50, 65, and 75 were approximately alike in their
survival and are combined for economy of graphic presentation, as are groups 175,
200, and 225.
702 DENIS R. A. WHARTON,JOHN E. LOLA ANDMARTHAL. WHARTON

made during the first 3 weeks after hatching. To avoid the risk of injuring the
larvae by frequent handling and counting, 27 groups of 100 each of newly hatched
larvae were placed in 6 x 8 in. battery jars. A different group was counted and
weighed on successive days to obtain a correlation of survival and early growth
(Fig. 2). Following the first ecdysis (at hatching) growth expressed as weight takes

l *-*
.l

..
..**’ . . 5
30,
*5 , I I I I , 1 1 1 I I 1 1 I I I I I I I I I I 1 I 0
o I 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 I8 192021 22 232425
Time, days

FIG. 2. Survival and weight curves obtained from data from a different group of
100 larvae on each successive day. Each group was counted for survival and
weighed once, and then discarded. Each circle on curve represents the percentage
survival of the particular group of 100 larvae examined on the indicated day; each
closed circle represents the average weight of the individuals in that group. The
ecdysial times and percentage ecdysed were as follows:

2nd Ecdysis 3rd Ecdysis 4th Ecdysis

Day % Day % Day %

7 20 14 3 21 3
8 90 15 22 75
9 97 16 ;i 23 83
10 100 18 99 24 94
25 100

place for several days and then levels off for a few days during which the second
ecdysis occurs. Growth proceeds again following the second ecdysis and tapers
off again after several days during the period of the third ecdysis, after which it is
resumed. The moulting period becomes more protracted as development proceeds
and hence the lag period of growth becomes less distinct. The mortality curve is not
linear but rather forms steps which coincide with the moulting cycle of the insect:
survival decreases following the ecdysis and becomes stable immediately prior to
and during the succeeding ecdysis.
TABLE ~-MORTALITY AND GROWTH OF LARVAE AT 3 WEEK PERIODS THROUGHOUT THEIR DEVELOPMENT IN GROUPS OF DIFFERENT
DENSITIES, BASED ON THE NUMBERS AND WEIGHTS AT THEBEGINNINGOFEACHPERIOD

Group
---
25 50 75 17.5 250

% No. % NO. % No. % No. % No.

3-week % weight adults % weight adults % weight adults % weight adults % weight adults
period mortality increase emerged mortality increase emerged mortality increase emerged mortality increase emerged mortality increase emerged

o-3 8 900 26 672 9.5 657 30 555 26 580

3-6 0 363 8.1 422 I.5 515 12.3 548 14.6 433
6-9 0 210 0 0 199 0 178 0 0 117 1.9 173
9-12 73.9
12-15 0
0 57.8
3.5 126 2.9
6.1 96.5
20.9 25 6.0
2.1 69.8
22.2 15
42 9.3
7.2 106
242 0
24 2:; 21.1
15-18 0 3”:
18-21 20 15.8 2 0 0 21 0 49.8 41 0 17.3
0.4 56
6 8 23.3 4
21-24 0 24.6 1

Group

500 650 750 1000

o-3 28.8 600 23.8 552 19.2 374 23.5 500 27.5 306
3-6 16.1 447 15.4 416 40 385 12.7 395 45.8 371
6-9 153 0 4.0 146 2.2 215 9.4 12s 4.8 256
9-12 ;:; 67.6 10 8.7 60.8 0 11.3 71.7 0 11.5 21.4 0 IO.4 59.4 0
12-15 12.6 16.8 77 17.0 15.9 48 14.6 13.3 68 28.1 48.6 22 18.2 47.3
15-18 1.3 9.7 67 11.3 20.0 95 7.9 24.3 67 19.9 17.5 44 11.0 12.8
18-21 0 16.2 9 4.3 17.1 45 0 9.9 13.5 15.3 63 5.1 22.4 65
21-24 4.5 10.1 17 Il.4 13.6 24.8 46 22.4
24-28 223 22 19.7
704 DENISR. A. WHARTON,
JOHNE. LOLAand MARTHAL. WHARTON

Mortality and growth rate

Ecdysis is therefore one factor involved in mortality, and the consequent
exposure of teneral larvae to the hazards that go with large numbers accounts for a
large number of deaths with increasingly high densities. There are, however, other
intrinsic factors such as growth that modify the death rate, as shown by an examina-
tion of the mortality and growth rates of groups of different densities (Table I).
The percentage mortality, based on the number of surviving larvae at the
beginning of successive 3 week periods from the time of hatching, is highest during
the first 3 weeks and continues high during the second 3 week period regardless of
the population density. A sharp drop occurs in all groups during the 6 to 9 week
period, and this respite is followed by a rise in mortality during the ninth to the
twelfth and following weeks. Thus, we recognize three major phases of mortality
in a population of larvae: (1) the early phase of a high mortality rate during the
first to the sixth week, (2) a phase of low mortality during the sixth to the ninth
week, and (3) a mortality of moderate proportions and of long duration which
reaches its height during the fifteenth to the eighteenth week and gradually
subsides. The instantaneous rate of growth at 3 week intervals (percentage weight
increase) is maximal during the first phase, drops sharply during the second phase,
and continues at a lower level for the rest of larval development. The relatively
short intervals of ecdysis during their rapid rate of growth expose the young larvae
in a teneral state more frequently in a given period of time than is the case with
older larvae; so that it is young larvae in groups that are most likely to be injured
and killed. Isolated larvae grow slowly and their mortality rate is very low. It is
only when larvae are exposed to the stresses of the group that they suffer a high
mortality.
The second or middle phase of mortality occupies the brief period of the sixth
to the ninth week and is characterized by a declining growth rate and low mortality.
It would appear that the larva is entering a resting period in which physiological
inequalities incident to rapid growth have come to a balance. The larva at this stage
may be considered as having completed larval development before entering upon
the new metabolic processes and hormonal balances of the third phase which will
culminate in the sexual adult, and which, as the rising mortality rate shows, makes
it vulnerable to stress.
During the third phase the growth rate is minimal, but this phase is marked by
the onset of a moderate mortality, which coincides with the differentiation of the
larva and its metamorphosis to the adult. The fact that this phase should be so
much more lethal than the second period, indicates that the larva undergoes a
severe physiological disturbance in its final metamorphosis.
Growth as weight gain may be expressed in different ways which convey
entirely different information (BRODY, 1945 ; THOMPSON, 1948). In Fig. 3 two
modes are used: the first plots average weight per larva at 3 week intervals and
generally yields sigmoid curves ; and the second represents the instantaneous
percentage of rate of growth. Whereas the slopes of the sigmoid curves give growth
rates that gradually increase to a maximum between the sixth and ninth week
 EFFECTS OF POPULATION DENSITY ON AMERICAN COCKROACH 705

800

1200-

IIOO- 700

(25,50,75)

1000 - (750,1000) P 600

900-

---x-m-m I I 1
‘3 6 9 12 15 18 21 24 27 30 33’

Time, wk

FIG. 3. Growth(weight) curves for different densities of larvae in jars. The retarded
growth in the l’s is exaggerated because of their being grown in beakers (cf. Fig. 4).
The curves with solid lines present the average weight at 3 week intervals. The
broken-line curve is that for the representative group of (500, 650) and shows the
instantaneous rate of growth (percentage weight increase of Table 1) at 3 week
intervals. It emphasizes the relative rate of growth during these intervals.

and then decline, the instantaneous percentage rate of growth shows that the
percentage weight increase is maximal during the early stages of growth and
decreases rapidly. The first curve shows the increase of total weight, whereas the
second shows the percentage increase, or the rate of increase at the stated time
intervals. It is evident from the second curve, which illustrates the data of
group (500, 650) of Table 1, that the young animal is expending its energy in
growth at a more intense rate than the older, and it is this aspect of growth that
is relevant to the mortality data that we are now considering.

The effect of density on optimal growth and development

The effect of population density on the growth and development of cockroach
larvae presents a very different picture from that of survival. When the percentage
of the survivors emerging as adults in the different density groups is plotted against
time, a series of generally sigmoid curves is obtained which shows a difference
between the insects grown in isolation or in small groups and those grown in large
groups (Fig. 4). Except for the group of (750, lOOO), the isolated insects took
706 DENIS R. A. WHARTON, JOHN E. LOLA AND MARTHA L. WHARTON

(500,,650)
100
90
80
70
60
50
40
30
20
10
0

Time, wk

FIG. 4. Development curves for different population densities of larvae grown in

6 x 8 in. battery jars (except for 1B and 2B, which were grown in 250 ml beakers),
expressed as percentages of all surviving insects, both larvae and adults. Vertical
lines crossing curves indicate limits of averages for the group of (500,650) and
(750,1000).

longest to mature, while larvae in groups of from 2 to 25 matured increasingly early

as the numbers were increased. Beyond this point there is an abrupt reversal, so
that the time of emergence increases according to the density. The reversal is so
consistent and far reaching that the group of 200 (not shown on chart) emerges at
the rate of the 2’s, and the group of (750, 1000) actually emerges more slowly than
the isolates.
The data are also plotted in terms of the time it takes a certain percentage of the
survivors of the different density groups to develop (Fig. 5). The curve is drawn
for the emergence of 50 per cent of the survivors of the different density groups in

IS0
0

160
0
0

60c

1 I II, I I 1 I
IO 100 1000

Populotion density

FIG. 5. Development data for 50,25,and 90 per cent emergence in days vs. log of
population density. Curve drawn for 50 per cent emergence.
 EFFECTS OFPOPULATIONDENSI~ONAMERICAN COCKROACH 707

days vs. the log of population density. It shows a rapid shortening of the develop-
ment time resulting from increasing the group numbers from 1 to about 25 ; then
the curve breaks so that the time for development now increases gradually with
increasing initial density. When the data are plotted arithmetically the break in the
curve is remarkably sharp, and thereafter the time for development is approximately
linear with density. Since the same type of sharply breaking curve is obtained
regardless of whether the data are plotted for 25, 50, or 90 per cent emergence, we
conclude that a factor exists which affects the rate of development of the grouped
insects throughout their development. This factor acts with increasing intensity
as the size (density) of the group is increased to an optimum of about 25 (under
these conditions), and persists with diminishing effect as the size of the group is
further increased.
This effect predominates in the males, whereas except for the l’s there is little
variance in development time from group to group of females (Table 2). However,
unless large numbers or several groups are observed, this finding could easily go
unnoticed because of the sex differences in development rates, and the not infrequent
chance in small groups of the larvae, being preponderantly of one sex.

TABLE %--MEAN TIMES IN DAYS FOR DEVELOPMENT OF ISOLATED AND GROUPED LARVAE, AND
WEIGHTS OFEMERGENTADCZTS

Popula- Mean time for development + S.D. Mean weights k S.D.

tion
density No. Males No. Females No. Males No. Females

1 3 143 * 12 2 140 +_21 3 905+ 50 2 1097* 68

2 5 125 zk15 4 100 + 10 5 801 zk 55 4 1005* 79
5 7 121 -t_21 8 95f. 9 7 864+ 86 8 964+ 43
10 30 lOSj,ll 36 100 + 11 30 783 f 84 36 IOllf 80
15 14 107kl2 15 102+17 14 777* 73 1s 96Ort 99
20 33 106klO 17 lOI& 9 33 798+114 16 960 f 126
25 20 102+11 24 100+11 20 744+ 50 23 923 rf 79
30 28 Ill+12 23 99+13 27 782+106 23 937 XL144
60 53 1074 8 44 98~~ 8 53 704+ 70 44 886+ 87
200 43 113+112 44 103 + 12 43 624+ 55 43 8605 85

These findings contrast with those of LANDOWSKI (1937) in which isolated B.

orient& larvae took longer to mature than larvae in groups up to his maximum of
16, which showed no reduction in development time as the size of the group
increased.
The promotion of growth and development by grouping is effective at any larval
stage; however, its influence ceases when groups are disbanded. In Fig. 3, for
example, isolated larvae are seen to have grown very slowly up to the twenty-
fourth week of age, but their rate of growth was immediately accelerated when
they were grouped in twenties. Conversely, newly hatched larvae were distributed
in groups of 10 and at each ecdysis the individuals of a different one of the groups
708 DENIS R. A. WHARTON, JOHN E. LOLA AND MARTHA L. WHARTON

were isolated, and kept so, until development was completed. The groups thus
formed a series in which the larvae were isolated for decreasing lengths of time,
separated by their ecdysial intervals (Table 3). The results show that isolation
retarded development at all stages, and that the earlier the larvae were isolated the
more they were retarded in development.

TABLE ~---EFFECT OF ISOLATIONOF LARVAEAT DIFFERENTSTAGESOF GROWTH ON WEIGHT AND

TIME FOR DEVELOPMENT

Day Moult No.” Weight of adults: mean + S.D.

isolated
after 2345 6 7 8 9 10 Males Females
Lot hatching Days after hatching? No. Weight No. Weight

0 10 31 60 84 141 185 231 275 317 771* 92 2 829klll

7th 7 23 45 68 102 145 190 243 282 762 + 70 5’ 995 + 132
16th 14 29 76 108 151 201 257 287 748 + 75 2 926zk 32
23rd 22 44 72 121 193 217 255 756 + 38 6 93Ok 88
35th 32 71 117 182 231 266 659 169 5 925 zk 97
43rd 41 94 142 199 251 701 f 58 3 926 f 161
57th 55 124 1.59 184 703 4 36 7 945 * 140
71st 147:. 709 +_61 5 1005* 88
71st 147$ 713 + 69 4 946?1 69

* Eclosion of nymph is considered as first moult.

t Mean of days from hatching.
$ The eighth and ninth moults became intermingled and therefore could not be assigned
a definite time.

As contrasted with their growth in 6 x 8 in. jars, the growth of isslated larvae
in 250 ml beakers is extraordinarily slow, taking 305 days to adulthood as against
142 days in jars. This slowness is not due to the smaller amount of space alone but
also to the lack of shelter, for larvae grown in 6 x 8 in. jars with shelter grow at a
faster rate than those without shelter whether they are isolated or grouped (Table 4).

TABLE ~-RATE OF DEVELOPMENTOF LARVALIN JARPWITH AND WITHOUT SHELTERS

With shelter Without shelter

Time Weight Weight

(wk) Condition (mg) Condition (mg> Significance

5 3 groups of 10 89.1 + 5.2 3 groups of 10 67.4k 6 P-CO*01

5 8 isolates 66.0 f 19 9 isolates 44.0 f 14 P<O*Ol

Retardation of development can be affected as severely by fluctuations in tempera-

ture, as seen in the development of larvae isolated in beakers at room temperature
with a range of about 20 to 32”C, and at an incubator temperature of 29 + 1°C.
 EFFECTS OF POPULATIONDENSITY ON AMERICAN COCKROACH ,709

As shown by Table 5, both the male and female larvae took very much longer to
mature at room temperature than at the uniform incubator temperature. There was
a significant difference in the weights of the emergent females, the P value being
<O-02. The difference in temperature did not affect -the relative development
times significantly, although at room temperature there was deviation from the
rule that female larvae complete their development earlier than males.

TABLE 5--EFFECTOFVARIABLRTEMPFBATURR ON GROWTH OFISOLATED LARVAE

(IN BRARRRS)

Days to maturity Weight of emergent adults

-
No. Males No. Females No. Males No. Females

Incubator,
29+1”C 24 316+66 23 3102 79 24 754+102 23 886 z!z103’
Room
temperature 68 550 + 156 70 512+144 68 761* 85 68 955 f 118”

* The difference in weight is statistically significant (P<O*O2).

The e@ect of density on the weight of larvae and of the emergent adults
The influence of larval population density on the weights of larvae and of the
emergent adults is complex. Isolated larvae, which take longest to develop, yield
heavier adults than those from larvae grown in groups (Table Z), and the denser
the group the less will the larvae weigh at any given time or at their ultimate
development (Fig. 3). However, within groups of up to 1000 it is characteristic for
the later larvae that develop to weigh as much as or more than the heaviest in that
group, and therefore the typically sigmoid growth curve acquires a secondary rise
towards completion of development in large groups in which a comparatively high
percentage of the larvae develop slowly. For example in Fig. 3 the (750, 1000)
group shows a distinct secondary rise after the twentieth week, at which stage
survival is about 35 per cent in the 1000 group. Although the weights of this group
of larvae and of the resultant adults average the lowest of the series because of the
intensely suppressive effect of numbers during the first weeks, the later larvae and
the resultant adults weigh as much as those from small groups. It appears from
this relationship that as the numbers, that is density, of groups of larvae become
reduced, a more favourable condition for growth obtains, and this occurs early
enough in the development of the insect to be significant. Furthermore, this
indicates that originally large numbers have caused no permanent depression or
depletion of the growth factors in the larvae as measured by weight, in contrast to
the permanent injury which affects their survival as demonstrated in the mortality
curves.
Consideration of the various groups shows that the highest mean weight of the
adults is found in the isolates of both sexes, and the lowest mean weight is found
in the densest group. Within groups of all densities there is a definite tendency for
46
710 DENISR. A. WHARTON, JOHN E: LOLA AND MARTHA L. WHARTON

the later adults that emerge to weigh most. Time of development is not, however,
the only determiner of adult weight. A plot of the mean development time for
males of different densities forms a breaking curve, with development time
increasing after an optimal density is reached; nevertheless, the mean weights
decrease consistently as the density of the groups increases. As the group densities
increase, the weights of the females decline ‘even though there is no significant
variance in development time between the groups.

1000

jr ~~~~_

5 .**/
2
a 1 I I I1 11 11 1 11 ‘1 ”
65OlI 12. 13 14 15 16 17 18 19 20 21 22 23 24 25 26

Development time, wk

FIG. 6. cumulative average weight8 in mg corresponding with time of emergence

for adults from groups of 2, 5, 10, 15, 20, 25, 30, 60, and 200 larvae grown in
6 x 8 in. battery jars in the incubator. The middle curves show the percentage
emerged against the same time scale.

7801-
I Oriainol oooulation:500 -

Females

5*i2 I t xx 20
I6 I 24
1 28
1 32
’ 36
’ 40
’ 44
’ 4s
’ 52
’ 56
’

Time, wk

FIG. 7. Cumulative average weights in mg corresponding with time of emergence

for adults from a group of 500 larvae grown in a 6 x 8 in. battery jar at room
temperature.
 EFFECTS OF POPULATION DENSITY ON AMERICAN COCKROACH 711

Although between groups of different density there are significant differences

in the times for larval development and the weights of emergent adults, a trace of
the cumulative average weights of the emergent adults from. groups of 2 to 200
larvae (Fig. 6) shows that the females develop earlier and weigh more than the
males, and that in both sexes the emergent adults weigh more the longer the stage
of larval development. The steepest rise is in the first 25 per cent to emerge, and
is due to the slight but steady increase in the average weight, whereas the continued
terminal rise is due to the much higher weights characteristic of the relatively few
late-emerging adults. This type of curve is also given by larger groups of larvae
kept at room temperature (Fig. 7) even though they grow at only half the rate of
those kept in the incubator and may ultimately weigh as much (Table 5). However,
isolated larvae kept at room temperature in 250 ml beakers without shelter have
yielded adults whose weights have shown a decline with time of emergence.

The in.uence of cannibalism on growth and survival

Larvae tend to form dense clusters regardless of the total available space, and
this gregariousness of the insect is responsible for much of the injury and resulting
cannibalism of the young larvae. A good deal of milling about with resulting
collisions occurs, so that it is the defenceless ecdysing and tender larva that is the
most susceptible to injury. Crippled or dead, it is removed by cannibalism, which
acts more as a sanitary or scavenging measure, we believe, than as .a form of
predation. Thus, when 10,000 newly hatched larvae are put in a large cage, 23 x
14 x 9 in., with seemingly ample room and supplied with several inverted con-
tainers as shelters, they will occupy the shelters very unevenly, so that some are
crowded with jostling larvae. When inspected during the day, the cages are free
from dead bodies, leaving no reason to suspect that a heavy mortality may be
occurring. Nevertheless, a count made 1 to 2 weeks later will show that only about
50 per cent of the larvae have survived. On the other hand, when only 2500 larvae
occupy the same area, the probability of injury from crowding and the resultant
mortality are much reduced. Also, when 500 to 1000 newly emerged larvae were
provided with ample food and water in 6 x 8 in. and 10 x 12 in. battery jars which
contained a single shelter, and provided available areas of approximately 150 in2
and 300 in2, respectively, the mortality rates in the small and large jars were 72 and
48 per cent for the 500’s, and 77 and 56 per cent for the 1000’s.
Since optimal growth and development occur in the smaller groups with few
deaths and little or no cannibalism, but are retarded as population density and
cannibalism increase, it is interesting to consider the possible benefit of cannibalism
to the predator. In Tribolium it is said to satisfy the water needs of larvae in con-
ditions of low humidity (HOLDAWAY, 1932), but there seem to be no data regarding
its possible nutritional effect on animals provided with an adequate diet. There-
fore, we set up two experiments with larvae in their fifth instar: in the first, larvae
which were maintained on the standard PLC and water diet were fed in addition
freshly killed larvae of their own size three times weekly; and in the second,
starved larvae maintained on water were similarly fed killed larvae three times
712 DENISR. A. WHARTON,JOHNE. LOLAANDMARTHAL. WHARTON

weekly. The original mean weights of the 10 test animals and their controls in the
first experiment were 189 it: 38 mg and 176 + 29.5 mg, respectively. After 7 weeks’
feeding, the weights were 320 f 110 mg and 301 t_ 111 mg, respectively. The
results of the first experiment, therefore, showed no significant difference in
weights between the cannibals and the controls, and there were no deaths. In the
second experiment, the wholly starved larvae lost weight, as would be expected,
and one died on the thirty-second day. After 7 weeks, the fed larvae weighed
226 + 41 mg and the starved 112 + 31 mg. Although the larvae feeding on their
fellows gained weight and had all survived when the experiment was ended at 7
weeks, their weight failed to compare with the controls fed PLC, which weighed
281+ 31 mg. Thus, it is apparent that cannibalism served the purpose of survival
in the absence of other food, but did not promote growth when an adequate diet
was available.

DISCUSSION
It has been our unvarying observation (Fig. 1; Table 1) that the greatest
mortality is suffered by young larvae. This contrasts with LANDOWSKI’S (1937)
finding with B. orientalis that mortality increases with age and with reduction of
the space available per head to the growing larva. Thus, it appears that the increas-
ing mortality which occurs with increasing population density is due to the greater
incidence of injury from collisions consequent to the teneral state at ecdysis,
especially of the young larvae. Such ‘passive’ cannibalism as occurs with the eating
of crippled or dead larvae is beneficial to the colony as a scavenging, sanitary
measure, but aggressive cannibalism is considered to be damaging by creating a
state of alertness and ‘anxiety’ which interferes with appetite and growth. As
contrasted with its sanitary role, cannibalism in the form of predation among the
larvae may also be a lethal instrument to counteract the tendency of the insects to
cluster, and so establishes more tolerable living conditions. Most cannibalism
among larvae appears to be caused by the commotion and resultant injury induced
by crowding, which depends partly on the ambient conditions of light, temperature,
humidity, and so forth. Cannibalism has been observed to be most intense among
young larvae and far outweighs that which occurs among older larvae or adults,
except possibly with respect to the cannibalism of eggs.
Nevertheless, in the current series of experiments in which an adequate diet
was fed, there is no evidence of cannibalism having benefited the insects. The
most heavily populated groups, in which cannibalism was greatest, were not the
ones to grow most rapidly or to produce the largest adults. However, we have
observed with adult cockroaches that where the metabolism is imbalanced, as by
feeding antibiotics, as the result of which protozoa1 growth is suppressed and the
character of the microflora greatly altered, the insects become ravenous and
pugnacious with little regard for the vigour of their opponents (WHARTON et aZ.,
1965), and we assume this to be a way of satisfying nutritional needs. Although
this might also be a means of satisfying thirst, induced by a possible but not evident
distaste for the antibiotic solution, there is no obvious reason to suppose that
 RFFRCTS OF POPULATION DENSITY ON AJVIERIC4N COCKROACH 713

cannibalism among the cockroach larvae serves such a purpose in these experiments.
It is initiated by the presence of injured larvae and is not primarily an instrument
of growth regulation or population control among larvae.
Since cannibalism appears to provide no food advantage to larvae already
supplied with an adequate diet, it may be considered to have originated from
dietary need. Its persistence in abundantly supplied colonies therefore probably
serves more importantly as a sanitary function than as a means of controlling
population density.
Of the several remarkable effects of population density on growth and matura-
tion, perhaps the most striking is the difference that the addition of adults or of
only one larva makes to the growth rate of an isolate; and the relative size of the
paired larvae seems to make no difference, as shown by the fact that one larva grew
to above average size in the presence of a persistent runt. Increased growth rate
has also been reported for the house cricket, Gryllulus dome&us, reared in twos
and threes CHAWIN (1958) or tens (MCFARLANE, 1962) rather than singly; and
VETULANI (1931) has shown the same for mice. KUNKEL (1966) found that increas-
ing the population density of B. germanica larvae had the general effect of reducing
the average weight gain. This conforms generally with our results with P. americana
even though the minimal number of 0.2 larvae per cm2 which Kunkel used would
be equivalent to about 50 larvae per battery jar as compared with one in our
experiments with P. americana. This leap from the constraint of isolation stands
in sharp contrast to the conclusion of CHRISTIAN and DAVIS (1964) that ‘since
social and behavioural features are density dependent, they become evident only
at high population levels’.
In view of the demonstration of a maturity-promoting effect on immature
adults of the desert locust by a pheromone which acts through absorption on
contact, or through the olfactory sense (NORRIS, 1954; LOHER, 1961), and of the
fact that the stimulus to growth of the larva of P. americana is removed when the
larva is isolated, it is tempting to postulate that a pheromone is the factor that
operates to promote, or, by its absence, depress, growth of the cockroach larva
under the influence of numbers. But it would be unrealistic to attribute accelerated
growth and development to pheromones solely; for although the isolated larva is
free from the often lethal harassment of the crowd and has high survival, isolation
deprives the insect of benefits that may possibly accrue from feeding on the
secretions of others, such as the mucous-like substance on the tergites and cerci
(ROTH and STAHL, 1956); it deprives them of the stimulus of disturbance which,
with the associated competition, increases the metabolism and appetite, as grouped
cockroaches are more active than single ones and do, in fact, eat more (WHARTON
and WHARTON, 1959); and it deprives them of the sense of security and well-being
that their natural gregariousness presumably imparts by physical and possibly
psychological means. Whatever the individual contribution of these various
factors, it is evident from the rather precise consistency of the data that a quantita-
tive relationship exists, and that while association is beneficial, a point is reached
at which its benefits are overcome by the stress of concentration. Whatever the
714 DENIS R. A. WHARTON,JOHN E. LOLA AND MARTHAL. WHARTON

operative factors may be, it is evident that they are different from those that affect
mortality.
The view of mortality as incidental to the physiological state of the insect raises
the question of precisely what the imbalances are that are more lethal or less so.
This paper does not propose definitive answers. However, the suggestion seems
valid that the rapid growth and relatively frequent ecdysis of the young larvae,
which exposes them to injury in the presence-of crowds, are accountable for the first
phase of mortality. The second phase is certainly one of rest, as indicated by the
decreased growth rate, and one in which a new balance of hormones is being
established whose effects become manifest in the third phase. The third phase is
that in which the sexual apparatus and secondary appurtenances of the adult are
being formed and which occurs when growth is still progressing, though at, a
reduced rate. The data prompt the suggestion that this phase of development of
the cockroach, a hemimetabolous insect, is equivalent to the pupal stage of
differentiation of a holometabolan, and that its sensitiveness to stress derives from
the fact that it metamorphoses directly from the active larval state to the adult
while still growing. This state of stress resolves itself in the evolution of the more
orderly segregation of the differentiating phases which is achieved in the pupa. If
these hypotheses are correct, our results add substantial support to HINTON’S
(1963) theory of the function of the pupal stage, but emphasize the physiological
efficiency of dividing the labour of metamorphosis more equably as an evolutionary
force in the origin of the pupa.

REFERENCES
ADOLPHE. F. (1931) The size of the body and the siie of the environment in the growth of
tadpoles. Biol. Bull., Woods Hole 61, 350-375.
ALLEEW. C. (1934) Recent studies in mass physiology. Biol. Rev. 9, 1-48.
BILSKI F. (1921) See L~DOW~KI (1937) Zoc. n’t.
BOYCEJ. M. (1946) The influence of fecundity and egg mortality on the population growth
of Triboliuna confusum Duval. Ecology 27, 290-302.
BRODY S. (1945) Time Relations of Gowth of Individuals and Populations. Chapter 16.
Reinhold, New York.
BRUCEH. M. (1959) An exteroceptive block to pregnancy in the mouse. Nature, Land. 184,
105.
BUTLERC. G. (1954) The method and importance of the recognition by a colony of honey
bees (A. mellifea) of the presence of its queen. Trans. R. ent. See. Land. lOS, ll-29.
CBRXAJEVW. P. (1931) Ein Beitrag zum Studium von Wachstumfaktoren an Korpfesbrut
mit besonderer Beriicksichtigung des Einfluss der Gr&senverhiiltnisse des zur Verfii-
gung stehenden Lebensraumes. 2. Fisch. 29, 217-282.
CHAPMAN R. N. (1928) The quantitative analysis of environmental factors. Ecology 9,
698-711.
CHAPMANR. N. and WHANG W. Y. (1934) An experimental analysis of the cause of popula-
tion fluctuations. Science, N. Y. 80, 297-298.
CHAUVINR. (19%) Notes sur la physiologie comparee des Orthopteres--V. L’effet de groupe
et la croissance larvaire des Blattes, des Grillons et du Phaneroptere. BUZZ.SOC. r001. Fr.
71, 39-48.
CHAUVIN R. (1958) L’action du groupement sur la croissance des grillons (GryZZ~lus
dome&us). J. Insect Physiol. 2, 235248.
 EFFECTS
OF POPULATION
DENSITYON AMERICAN
COCKROACH 715

CHRISTIANJ. J. and DAVISD. E. (1964) Endocrines, behavior, and population. Science, N. Y.

146, ISSO-1560,
CREWF. A. E. and MIR~KAIAL. (1931) The effects of density on an adult mouse population.
Biologia gen. 7, 239-250.
GIER H. T. (1947) Growth rate in the cockroach Periplaneta americana (Linn.). Ann. ent.
Sot. Am. 40,303-317.
HINTONH. E. (1963) The origin and function of the pupal stage. Proc. R. ent. Sot. Lond. (A)
38, 77-85.
HOLDAWAYF. G. (1932) An experimental study of the growth of populations of the ‘flour
beetle’ Tribolium confusum Duval, as affe.cted by atmospheric moisture. Ecol. &lonogr. 2,
261-304.
KUNKELJ_ G. (1966) Development and the availability of food in the German cockroach,
Blattella germanica (L.). J. Insect Physiol. 12, 227-235.
LANDOWSKIJ. (1937) Influence of isolation and cohabitation on the development and growth
of the nymphs of Periplaneta orientalis L. C. R. Sot. Sci. Lett. Warsaw, 4, 190-203.
(Translated from the Polish.)
LEE S. V. and BOOT L. M. (1955) Spontaneous pseudopregnancy in mice. Acta physiol.
pharmac. n&rl. 4,442-444.
LOHER W. (1961) The chemical acceleration of the maturation process and its hormonal
control in the male of the desert locust. Proc. R. Sot. (B) 153, 380-397.
L~SCHERM. (1956) Hemmende und fijrdemde Faktoren bei der Entstehung von Ersatz-
geschlechtstieren bei der Termite Kalotermes flavicollis Fabr. Rev. suisse Zool. 63,
261-267.
MCFARLANEJ. E. (1962) A comparison of the growth of the house crkket (Orthoptera:
Gryllidae) reared singly and in groups. Can. J. Zool. 40, 559-560.
MACLAGAND. S. (1932) An ecological study of the ‘Lucerne flea’ (Smynthurus viridis). Bull.
ent. Res. 23, 101-145.
MICHAL K. (1931) Die Beziehung der Populationsdichte zurn Lebendsoptimum aus der
ZahlenverhSiltniss der Geschlechter bei Mehlwurm und Stubentliege. Biologia gen. 7,
631-646.
NORRIS M. J. (1954) Sexual maturation in the desert locust (Schistocerca gregaria Forskgl)
with special reference to the effects of grouping. Anti Locust Bull. 18, l-44.
PARKT. (1932) Studies in population physiology. The relation of numbers to initial popula-
tion growth in the flour beetle Tribolium confusum Duval. Ecology 13, 172-l 81.
PARK T. MERTZ D. B., GRODZINSKIW., and PRUS T. (1965) Cannibalistic predation in
populations of flour beetles. Physiol. Ziiol. 38, 289-321.
PARKESA. S. and BRUCE H. M. (1961) Olfactory stimuli in mammalian reproduction.
Science, N. Y. 134, 1049-I 054.
PEARLR. (1932) The influence of density of population upon egg production in Drosophila
naelanogaster. J. exp. Zool. 63, 57-84.
PEARLR., MINER J. R., and PARKERS. L. (1927) Experimental studies on the duration of life
-XI. Density of population and life duration in Drosophila. Am. Nat. 61, 289-318.
PEARLR. and PARKERS. L. (1922) On the influence of density of population upon the rate of
reproduction in Drosophila. Proc. natn Acad. Sci. U.S.A. 8, 212-219.
PEARLR. and SURFACEF. M. (1909) A biometrical study of egg production in the domestic
fowl. Bull. Bur. Anim. Ind. U.S. Dep. Agric. 110,I-80.
PETTITL. C. (1940) The effect of isolation on growth in the cockroach Blattella germanica.
Ent. News 51, 253.
PIQUETTP. G. and FALES J. H. (1952) Rearing cockroaches for experimental purposes.
U.S. Dept. Agric. E.T. 301, l-12.
ROTH L. M. and STAHL W. H. (1956) Tergal and cereal secretion of Blatta orientalis L.
Science, N.Y. 123,798-799.
THOMPSOND’ARCY W. (1948) On Growth and Form. Macmillan, New York.
716 DENIS R. A. WHARTON, JOHN E. LOLA ANDMARTHAL. WHARTON

VETULANIT. (1931) Untersuchungen iiber das Wachstum der Siiugetiere in Abhiingigkeit

von der Anzahl zusammengehaltener Tiere. Biologiu gen. 7, 71-98.
WHARTOND. R. A. and WHARTONM. L. (1959) The effect of radiation on the longevity of
the cockroach, Periplaneta americana as affected by dose, age, sex and food intake.
Radiat.Res. 11,600-615.
WHARTOND. R. A,, WHARTONM. L., and LOLA J. (1965) Cellulase in the cockroach, with
special reference to Peri$laneta americana L. y. Insect Physiol. 11,947-959.
WHITTEN W. K. (1956) Modification of the oestrus cycle of the mouse by external stimuli
associated with the male. r. Emiocr. 13,399-404.
WILLIS E. R., RISER G. R., and ROTH L. M. (1958) Observations on reproduction and
development in cockroaches. Ann. ent. Sac. Am. 51,53-69.