Journal of lnsect Behavior, Vol. 9, No.

1, 1996
Factors Regulating Predation by First-Instar Spined
Assassin Bugs [Sinea diadema (Fabricius)]
(Hemiptera: Reduviidae)
J. R. Tayl or I and J. M. Schmi dt 1'2
Accepted July 11, 1994; revised September 12, 1995
Selected nutritional and developmental factors regulating the predatory behavior
of first-instar spined assassin bugs Sinea di adema (Fabricius) (Hemiptera: Redu-
viidae) were investigated. The longevity of unfed nymphs provided with free
water, bean pod sections, or glucose solutions was not significantly greater than
that of unfed nymphs which were not provided with a source of water. First-
instar S. di adema that were provided with larvae of Ephestia kuehniella Zeller
(Lepidoptera: Pyralidae) as prey began feeding 1.9 (+0.9) days after hatching.
In contrast, first-instar S. di adema that were provided with conspecifics of the
same age did not begin feeding until they were 3. 9 (+0.9) days old. These
results suggest that the potential nutritional benefits to be gained f rom feeding
equal or outweigh the risk of attacking prey capable of effective self-defense
only when nymphs have not f ed f or 4 days after hatching. Providing the nymphs
with water or glucose solutions significantly delayed the onset of conspecific
predation. Additional data are presented which demonstrate that first-instar S.
di adema are not restrained f rom preying on siblings by kin recognition:
KEY WORDS: Reduviidae; interspecific predation; intraspecific predation; nutrition; kin recog-
nition.
I NT RODUCT I ON
According to opt i mal foraging t heory, a predat or should select prey that maxi -
mi ze its net energy gain (see Pyke et al., 1977; Krebs, 1978). The handling'
' Department of Environmental Biology, University ofGuelph, Guelph, Ontario, Canada N 1G 2W 1.
2To whom correspondence should be addressed.
23
0892-755319610100-0023509.5010 9 1996 Plenum Publishing Corporation
24 Taylor and Schmidt
time required for a specific prey species is an important factor in prey selection
because handling time can be costly in terms of energy expenditure by the
predator. The reaction of the prey when it is attacked may be a deciding factor
in prey selection, especially for a generalist predator that attacks many different
types of prey. Handling time is greatly affected by the time it takes a predator
to subdue the prey (Molles and Pietruszka, 1983) and prey with vigorous defen-
sive reactions are often rejected by satiated predators (Marston et al. , 1978;
Evans 1982). The physiological state of the predator may also influence prey
selection. Hunger can often prompt a predator to increase the range of prey
species that it will accept (Pastorok, 1980; Molles and Pietruszka, 1987). In
addition, hunger can alter an animal' s response to the risk of predation (Croy
and Hughes, 1991). Conversely, animals that have recently fed, or have nutrient
reserves, may be less likely to attack prey that are capable of counterattack.
In this study, we investigated the effect that different foods have on intra-
specific predation by first instars of a predatory bug. Sinea diadema (Fabricius)
(Hemiptera: Reduviidae) is a common North American predatory bug that feeds
on a wide variety of prey species (Readio, 1924). It employs a sit-and-wait
strategy for capturing prey with its raptorial forelegs. Prey encountered in the
field by S. diadema nymphs include various soft-bodied insects such as lepi-
dopteran and coleopteran larvae. In addition, the nymphs are likely to encount er
a variety of predatory arthropods that are both potential prey and predators,
including spiders, nabid bugs and ot her reduviid nymphs. In a series of four
experiments we determined some of the factors that regulate the response of S.
diadema nymphs to potentially hazardous prey. The hazardous prey used in
these experiments were conspecific nymphs of the same age and similar nutri-
tional status.
In our first experiment we tested the effect of different diets on nymphal
survival in order to establish the relative value of water, glucose solutions and
conspecific prey. In the second experiment first-instar S. diadema were offered
two types of prey: the soft-bodied larvae of Ephestia kuehniella Zel l er (Lepi-
doptera: Pyralidae) and conspecific nymphs. The larvae of E. kuehniella are the
usual food fed to S. diadema in our laboratory. Conspecifics of the same age
and size were chosen as the second prey species because they are representative
of prey with similar aggressive qualities and defensive capabilities. Furthermore,
cannibalism occurs frequently under laboratory conditions (Readio, 1924). I f a
predator does alter its behavior according to its nutritional status, particularly
the likelihood of continuing an attack on an aggressive prey, then it must use
physiological cues associated with recent feeding activity to regulate its behav-
ior. In our third experiment we evaluated whet her the availability of wat er or
glucose solution delayed cannibalism among first instars.
Since female S. diadema lay their eggs in clutches, cannibalism among
siblings possibly occurs in the field. To control for any affect this may have on
Predation by Spined Assassin Bugs 25
predatory behavior observed in the experiment, we conducted an additional test
to determine whether individuals cannibalize kin less readily than they canni-
balize nonkin.
MATERI ALS AND METHODS
Experiment 1: Development and Life Span of First Instars Fed
Different Diets
Sinea diadema egg clutches were collected from captive adults every 2 to
3 days. The eggs were kept in petri dishes until the time of hatching. In all
tests, nymphs less than 16 h old were placed individually in 5-cm-diameter petri
dishes. Every 24 h, nymphal mortality and devel opment were recorded. Obser-
vations ceased when a nymph either had died or had developed to the second
stadium. There were 20-25 replicates of each treatment. Three diets were tested.
Test 1. Bean vs. No Bean. (a) Each nymph was provided with a section of
green bean pod (Phaseolus vulgaris) which was replaced with a fresh section
every 2 days. Nothing else was provided. (2) Each nymph was provided with
neither food nor water.
Test 2. Water vs. No Water. (1) Each nymph was provided with a 1-cm 2
piece of absorbent 3M Supersorb material moistened with water. More water
was added as needed every 2 days. (2) Each nymph was provided with a dry
1-cm 2 piece of absorbent 3M Supersorb material.
Test 3. Variable Amount s of Prey. In this test, nymphs were allowed to
consume a fixed number of prey. Nymphs were provided with a 1-cm 2 piece of
absorbent 3M Supersorb material moistened with water, Prey consisted of con-
specifics that had been killed by freezing. On the day of hatching, each nymph
was provided with one prey item. Twent y-four hours later, each prey was checked
for indications that it had been fed upon (i . e. , sucked out). I f the prey had been
fed upon, the nymph was recorded as consuming a prey item. Each day prey
were replaced with freshly killed nymphs regardless of whet her they had been
fed upon. This procedure was repeated every day until the nymph had consumed
its allotment of prey. The nymphs were fed a total of either 2, 3, 4, 5, or 6
conspecifics.
In addition, 20 newly emerged nymphs were divided into t wo groups of
10 nymphs each. One group was provi ded with water, while the ot her group
was not provided with water. When the nymphs were 3 days old they were
weighed and the average weight of a nymph was calculated.
For tests 1 and 2, the Mann- Whi t ney U test was performed to test the
significance of the observed difference between the lifespans of treatment nymphs
and control nymphs.
26 Taylor and Schmidt
Experiment 2: Occurrence o f Predation and Cannibalism
All nymphs were less than 16 h old and unfed at the beginning of the
experiment. One or three nymphs were placed in a 5-cm-diameter petri dish
with a section of green bean pod. There were three treatments.
(1) A single nymph was provided with four 2-week-old E. kuehniella lar-
vae that were replaced every 48 h with fresh larvae.
(2) A group of three nymphs was not provided with any prey.
(3) A single nymph was not provided with any prey.
Treatment 3 was used to determine the age at which the nymphs died of
hunger.
Each dish was checked every 24 h. In treatment 1, nymphs were checked
for distended abdomens, which indicated that a nymph had fed. Observations
of each nymph continued until the nymph had fed. In treatment 2, occurrences
of cannibalism were recorded. Cannibalism was assumed to have occurred i f
there was at least one dead nymph in the dish and one, or both, of the remaining
nymphs had distended abdomens. Observations of each group of nymphs con-
tinued until the first incidence of cannibalism. In treatment 3, nymphal mortality
was recorded. Observations of each nymph continued until the nymph died.
There were 20- 25 replicates of each treatment.
The Mann-Whi t ney U test was performed to test the significance of the
observed difference between the age at which nymphs began feeding on E.
kuehnieUa larvae in treatment 1, and the age at which cannibalism began in
treatment 2. The Mann-Whi t ney U test was also performed to test the signifi-
cance of differences between the mortality distribution of nymphs maintained in
groups in treatment 2 and the mortality distribution of single nymphs in treat-
ment 3.
Experiment 3: The Effect of the Availability of Water and Glucose on the
Life Span and on the Age at Which First Instars Begin to Canni bal i ze
On the day that nymphs hatched, one or three nymphs were placed in a
5-cm-diameter petri dish. Liquids were presented by moistening t wo 0. 5-cm 2
pieces of absorbent 3M Supersorb material with wat er or with a 20 % glucose
solution. Diets were refreshed every 48 h. The following six treatments were
tested for their effects on either the life span of nymphs or the age at which
nymphs began to cannibalize.
(1) A single nymph was provided with wat er only.
(2) A single nymph was provided with wat er and a 20 % glucose solution.
Predation by Spirted Assassin Bugs 27
(3) A single nymph was provided with neither wat er nor glucose (control).
(4) A group of three nymphs was provided with wat er only.
(5) A group of three nymphs was provided with wat er and a 20% glucose
solution.
(6) A group of three nymphs was provided with neither water nor glucose
(control).
Every 24 h, nymphal mortality and devel opment in treatments 1, 2 and 3
and occurrences of cannibalism in treatments 4, 5 and 6, were recorded. Can-
nibalism was assumed to have occurred i f there was at least one dead nymph
in the dish and one, or both, of the remaining nymphs had distended abdomens.
In treatments 1, 2 and 3, observations ceased when a nymph had either died,
or had developed to the second stadium. In treatments 4, 5 and .6, observations
ceased when one of the three nymphs was dead. . Ther e were 34 replicates of
each treatment.
A Kruskal-Wallis test was performed to test for significant differences in
the age at which nymphs begin to cannibalize among treatments with grouped
nymphs. It was also used to test for differences in the lifespan of nymphs among
treatments with single nymphs. Pairwise comparisons wer e performed using the
Mann-Whi t ney U test.
Experiment 4: Selective Cannibalism of Unrelated Nymphs
Animals collected from the same area may be closely related. To obtain
animals that were not related, animals were collected from three different pop-
ulations at sites at least 10 km apart.
To ensure that offspring were full siblings, virgin females were paired with
a single male. Each pair was kept in a 5 x 7-cm plastic bottle from which egg
clutches were collected every two days. The parentage of each clutch was noted.
Three nymphs were placed in each petri dish. The three nymphs consisted
of a pair of full sibling nymphs (kin) and one nymph from an egg clutch produced
by adults collected from a different area (nonkin). Nymphs were marked with
a dab of acrylic paint on the back of the head for identification. Nymphs were
checked daily f or cannibalism. Cannibalism was assumed to have occurred i f
one nymph was dead and one, or both, of the remaining nymphs had distended
abdomens. There was a total of 35 replicates. Replicates in which t wo nymphs
were dead were omitted from the final analysis.
I f nymph s do not discriminate bet ween kin and nonkin, then the nonkin
nymph should be the first nymph killed in one-third of the replicates. A one-
tailed G test f or goodness of fit (Sokal and Rohlf, 1981) was performed to test
the null hypothesis that nymphs do not discriminate between kin and nonkin.
28 Taylor and Schmidt
RE S UL T S
Experiment 1: Development and Life Span of First Instars Fed
Different Diets
I n t he first test, none o f t he first-instar S. di adema pr ovi ded wi t h onl y gr een
bean pod devel oped into second instars. The avai l abi l i t y o f gr een bean pod al so
had no significant effect on t he mean life span o f t he first instars ( P > 0. 10;
Ma nn- Whi t ne y U test) (Tabl e I). I n t he second test, t he avai l abi l i t y o f free
wat er al so had no si gni fi cant effect on t he life span o f first instars (Tabl e I) ( P
> 0. 10; Ma nn- Whi t ne y U test). I n t he t hi rd test, t he pr opor t i on o f nymphs
devel opi ng i nt o second instars i ncr eased wi t h t he quant i t y o f conspeci fi cs con-
sumed (Tabl e II).
Nymphs t hat had been pr ovi ded wi t h wat er wei ghed 0. 32 mg on aver age
when t hey wer e 3 days ol d, whi l e nymphs t hat had been depr i ved o f wat er
wei ghed onl y 0. 24 mg on average.
Experiment 2: Occurrence of Predation and Cannibalism
Nymphs began feedi ng on E. kuehniella l arvae 1.9 ( + 0 . 9 ) days aft er hat ch-
ing. I n cont rast , nymphs began feedi ng on conspeci f i cs 3. 9 ( + 0 . 9 ) days aft er
hat chi ng ( P < 0. 05; Ma nn- Whi t ne y U test) (Tabl e III). The life span o f nymphs
t hat wer e kept i ndi vi dual l y and wer e not pr ovi ded wi t h pr ey was 6. 0 ( + 1 . 0 )
days.
Experiment 3: The Effect of the Availability of Water and of Glucose on
the Life Span and on the Age at Which First Instars Begin to Cannibalize
First instars that were kept i ndi vi dual l y and t hat wer e pr ovi ded wi t h wat er,
and wi t h wat er and gl ucose sol ut i on, sur vi ved 5. 2 ( + 0 . 9 ) and 5. 6 ( + 1. 6) days,
Table I. Mean Life Span (+SD) of First Instars Provided Either with a Section of Green Bean
Pod or with No Food and Water; Also, the Mean Life Span (+SD) ot First Instars Provided with
Either Water or with No Food and Water"
Bean h No bean Wate/' No water
Life span (days) 6.0 + 1.0 5.5 -I- 1.0 6.7 + 1.4 6.2 + 1.4
(n) (24) (25) (25) (22)
"Nymphal mortality was recorded approximately every 24 h.
bTreatment mean was not significantly different from control mean (P > 0.10; Mann-Whitney U
test).
Predation by Spined Assassin Bugs 29
Table H. The Number of First Instars, Fed 2, 3, 4, 5, or 6 Dead Conspeeifics, to Complete
Development to Second Instar"
No. of dead conspeeifics provided t o
first instars 2 3 4 5 6
No. of first instars to develop to
second instar 0 6 10 14 16
Percentage of first instals to develop to
second instar 0 30 50 70 80
I l l l
~ = 20 for each column. Data are also presented as the percentage of nymphs to complete
development.
r es pect i vel y. The s e nymphs di d not l i ve s i gni f i cant l y l onge r t han n y mp h s t hat
wer e not pr ovi de d wi t h wa t e r ( 4. 9 -t- 1.1 days ) ( P > 0. 10; Kr u s k a l - Wa l l i s t est )
( Tabl e I V) . Ho we v e r , n y mp h s pr ovi de d wi t h wa t e r o r wi t h a gl uc os e s ol ut i on
began t o c a nni ba l i z e when t hey wer e s i gni f i cant l y ol de r ( 4. 0 ___ 1. 0 and 4. 5 +
1.3 da ys , r es pect i vel y) t han nymphs t hat wer e not pr ovi de d wi t h wa t e r ( 3. 2 +
0. 7 days ) ( P < 0. 05; Ma n n - Wh i t n e y U t est s) ( Tabl e I V) . The age at whi ch
canni bal i s m be ga n a mong nymphs pr ovi de d wi t h wa t e r was not s i gni f i cant l y
di f f er ent f r om t hos e pr ovi de d wi t h a gl uc os e s ol ut i on ( P > 0. 10; Ma n n - Wh i t n e y
U t est ) ( Tabl e I V) .
Experiment 4: Se l e c t i ve Ca n n i b a l i s m o f Un r e l a t e d Nymphs
Twe nt y- ni ne r epl i cat es wer e us ed in t he anal ys i s . I n ni ne, or 31%, o f t he
obs er vat i ons , t he nonki n nymph was t he fi rst nymph t o be ki l l ed. The r e is no
evi dence t hat f i r st - i nst ar S. di adema pr e f e r t o c a nni ba l i z e nonki n r at her t han ki n
( P > 0. 10; G t est f or goodne s s o f fit).
Table i l l Mean Age (+SD) at Which Nymphs Began Feeding on Nonconspecific
and Conspecific Prey and the Lifespan of Nymphs Kept Individually
Age at which nymphs began feeding (days)
Single nymphs provided with Grouped nymphs provided with Life span of single nymphs
E. kuehniella larvae no prey" provided with no prey
1.9 + 0.9 3.9 + 0.9 6.0 + 1.0
(n = 25) 01 = 20) (n = 24)
"Age at which nymphs began feeding on larvae was significantly different from the life span of
single nymphs provided with no prey (P < 0.05; Mann-Whitney U tests).
30 Taylor and Sc hmi dt
Table I V. Mean Li fe Span ( 4 - SD) o f Nymphs that Were Maintained Individually and the Mean
Age ( +SD) at Whi ch First Instars Kept in Groups o f Three Began to Cannibalize (n = 34 for
Each Treatment): Three Di et s Were Testf, d ~
Diet
Wat er Wat er & 20% glucose No wat er
provi ded solution provi ded provi ded
Single nymphs- - l i f espan b (days) 5. 2 + 0. 9 5. 6 + 1.6 4. 9 + 1.1
Three nymphs - - age at first
incidence of cannibalism c
(days) 4. 0 + 1.0 b 4. 5 4- 1.3 b 3.2 4- 0. 7 c
"Nymphal mortality was recorded approximately every 24 h.
~'Life span of single nymph and age at whi ch first inslars began to cannibalize were significantly
different for each diet ( P < 0. 05; Mann- Whi t ney U tests).
CMeans within roW followed by the same letter (superscript) were not significantly different ( P <
0. 05; Mann- Whi t ney U tests).
DISCUSSION
Nutritional Factors Regulating the Predatory Behavior of First-lnstar
S. diadema
As shown in Experiments 1 and 3, first-instar S. diadema did not develop
into second instars when provided with only water, bean pod, or a glucose
solution. The life spans of nymphs provided With moisture sources were not
si'gnificantly greater than t hose of nymphs reared without a source of water.
Therefore, first-instar S. diadema cannot obtain sufficient nutrients from these
substances to complete development or 'prolong survival. Comparable results
have been obtained with Sinea confusa, Zelus renardii Kolenati, and Z. tetra-
canthus (Stoner et al., 1975). Even the availability of high-protein plant food,
such as dandelion pollen, did not sustain the development of first-instar S.
complexa, S. confusa, Z. renardii, and Z. tetracanthus, although their life spans
were five to nine times greater than the life spans of nymphs provided only with
water (Stoner et al. , 1975).
In contrast, several predatory hemipterans have been found to develop to
second instar on a diet of only plant food and water. Podisus maculiventris (Say)
(Hemiptera: Pentatomidae) developed into second instars when fed potato foli-
age and water (Ruberson et al., 1986); Geocoris punctipes (Say) (Hemiptera:
Lygaediae) developed into second instars when fed bean pods (Naranjo and
Stimac, 1985; Stoner, 1970); and Orius insidiosus (Say) (Hemiptera: Antho-
coridae) (Kiman and Yeargan, 1985) and O. tristicolor White (Salas-Aguilar
and Ehler, 1977) developed into second instars when fed bean pod and water.
Predation by Spined Assassin Bugs 31
Podisus maculiventris and G. punctipes are predatory members of primarily
phytophagous families (Cobben, 1979). In contrast, the Reduviidae are com-
prised solely of predators and arc probably descendants of a much older carniv-
orous offshoot (Sweet, 1979). The different evolutionary origins oftbese families
may.account for their differing abilities to use plant food. Podisus'maculiventris
and G. punctipes may. still possess physiological adaptations that allow them to
fully use plant nutrients. For instance, amylase activity was detected in the saliva
of P. maculiventris, 'but not in the saliva of Zelus renardii, Sinea confusa, or
G. punctipes (Cohen, 1990).
Reduviids appear to require animal protein for development. The only
nymphs of S. diadema to complete development among Tests 1, 2, and 3 in
Experiment 1 were those that fed on dead conspecifics. Parker (1965) noted that
the occurrence of molting in first instars of the reduviid, Psilius tipuliformis,
was closely linked with the supply of insect food. Edwards (1962a) found that
the reduviid, Rhinocoris carmelita, required at test one meal of prey for moulting
to occur. These results demonstrate that prey is essential for development. Fur-
thermore, as shown in Experiment 1, Test 3, the probability of successfully
developing to the next stadium increases with the quantity of prey ingested.
Experiments 1 and 3 also raise questions about the purpose of drinking in
reduviids. First-instar S. diadema drank readily when either free water or bean
was provided (personal observation). Edwards (1962b) studied the drinking
behavior of Rhinocoris carmelita, Platymeris rhaidamanthus, and Reduvius per-
sonatus (Linnaeus) and found that drinking was stimulated by contact between
water and the tibial pads of the forelegs and midl.egs, the tips of the antennae,
or the beak. Moreover,. when drinking, the reduviids could discriminate between
ionic solutions of different concentrations. These observations suggest that drink-
ing is a common behavior, and therefore nymphs must derive some benefit from
it. ,However, in Experiment 1 water availability did not increase the survival of
S. diadema nymphs. Moreover, S. diadema nymphs did not need to drink water
prior to killing and feeding on prey (personal observation). Thus, the signifi-
cance of drinking behavior in reduviids is not clear. Possibly, it facilitates
feeding by decreasing the time required to paralyze and ingest the prey. Large
amounts of fluid may increase the production of saliva. Saliva contains the toxins
and proteolytic enzymes needed for the paralysis and preoral digestion of the
prey (Smith, 1985). Moreover, large amounts of dilute saliva are neded to form
a temporary suspension of the digested tissue which: is then ingested (Smith,
1985). A comparison of the time required for the onset.of paralysis of prey and
ingestion of prey by nymphs that are provided with water and nymphs that are
not provided with water, is neded to test this hypothesis.
Nymphs that were provided with E. kuehniella larvae did not begin feeding
until they were 2 days old. Edwards (1966) noted that when emerged Zelus
exsanguis Stal (Hemiptera: Reduviidae) nymphs are "maintained at high humid-
32 Tayl or and Schmi dt
ity they will not take prey for three to four days . " Sinea diadema nymphs and
Z. exsanguis nymphs must be sustaining themselves for the first few days after
emergence on nutrient reserves persisting from the egg stage.
When conspecifics were the only prey available to first-instar S. diadema,
feeding did not begin until nymphs were nearly 4 days old. It is possible that
E. kuehniella larvae are more active and therefore encounter nymphs more
frequently. However, differences in encounter rates between nymphs and larvae
and between nymphs and conspecifics were considered negligible because of
the simplicity and small size of the experimental arenas and relatively long time
period over which the experiment was conducted.
The reaction of the prey during encounters is probably the major factor in
prey selection. In response to being attacked by a nymph, the E. kuehniella
larvae thrashed their bodies for about 30 s, until the injected salivary toxin
paralyzed them. In contrast, S. di adema nymphs reacted to being attacked by
posturing and striking with their raptorial forelegs. Sinea di adema nymphs had
a more vigorous defensive behavior than E. kuehniella larvae and were thus
more successful at repelling attacks. The defensive behavior of the prey may
also communicate to the predator the degree of risk associated with attacking.
There is little risk of injury to a S. di adema nymph attacking an E. kuehniella
larva. However, a nymph that is attacking a conspecific of the same age and
size is at significant risk of being injured and even killed. Thus, it appears that
conspecifics are attacked and killed only when a nymph is compelled by hunger
to persist in the attack, such as after a prolonged period of food deprivation.
This suggests that when nymphs are four days old and have not fed, the nutri-
tional benefits to be gained from feeding on a conspecific equal or outweigh the
handling costs and risk of attacking that prey.
Interactions between recently hatched nymphs and between older, starved
nymphs do not differ significantly in duration, however, the behaviors displayed
do change markedly with age. During interactions between 2-day-old, unfed
nymphs the predominant behaviors are posturing and striking with the forelegs.
The nymphs rarely hold or grapple with each other with their forelegs. In con-
trast, 5-day-old, starved nymphs interact at close quarters, often grappling with
each other while attempting to insert their stylets (Talyor and Schmidt, 1994).
Grappling was observed in only 18% of the interactions between 2-day-old
nymphs. In contrast, 67% of the interactions between 5-day-old nymphs con-
tained grappling. Cannibalism occurs only in interactions during which grap-
pling occurs (Taylor and Schmidt, 1994).
The observation that the consumption of conspecifics is delayed compared
to consumption of E. kuehniella larvae suggests that S. d i a d e m nymphs may
have information concerning their own nutritional status. This information may
regulate the likelihood that a nymph will grapple with, and attempt to immobilize
an aggressive prey. Physiological cues that may be involved include distension
Predation by Spined Assassin Bugs 33
of the digestive tract, tissue dehydration, hemol ymph volume, and the concen-
tration of specific hemol ymph components such as trehalose. In Experiment 3,
nymphs that were not supplied with water began to cannibalize at a significantly
younger age than nymphs that were provided with either water or a glucose
solution. Again, this result is surprising as water availability did not increase
the survival of first instars in Experiment 1. Assuming that the age at which
nymphs begin to cannibalize is an indication of their motivation to feed, and
hence their hunger level, then nymphs that were not provided with wat er were
"hungr i er " than nymphs of the same age that were provided with water. When
water is available, nymphs can consume substantial amounts. First instars that
had water available since hatching weighed 0. 32 mg on average when they were
3 days old. In comparison, nymphs that had been deprived of water weighed
only 0. 24 mg on average when they were 3 days old. Possibly, distension of
the gut stimulated stretch receptors of the abdominal wall, indicating to the
nymph that its gut was partially full. Distension of the gut has been shown to
stimulate stretch receptors which control intake of food in flies (Dethier, 1976)
and the reduviid Rhodnius prolixus (Wigglesworth, 1965). Still, ingesting so
much water that it lowers a nymph' s motivation to feed appears counteradaptive.
In fact, nymphs that do not drink appear to have an advantage since they are
more aggressive and are more likely to capture prey than nymphs that do drink.
Further research is needed to understand the behavioral and physiological impli-
cations of drinking.
Kin Recognition and Canni bal i sm
There was no evidence that S. diadema nymphs behave differently toward
kin and nonkin. This is not surprising since kin recognition among conspecifics
engaged in cannibalism is relatively rare outside of the eusocial insects (Wilson,
1987). There is no evi dence of kin recognition in land snail larvae (Baur, 1987),
acarid mites (Radwan, 1993), several species of waterstriders (Nummelin, 1989;
Carcamo and Spence, 1993), and lace bugs (Tallamy and TaUamy, 1993).
Cannibalism has been cited as a significant obstacle to mass-rearing pred-
atory insects for biological control (DeBach and Rosen, 1991). However, our
results indicate that the risks and possibility of effective count er attack associated
with preyi ng on conspecifics may del ay or reduce the incidence of cannibalism.
I f predators such as S. diadema are provided with adequate supplies of l ower
risk prey, losses due to cannibalism can be reduced.
ACKNOWLEDGMENTS
This research was supported by the Natural Sciences and Engineering
Research Council of Canada. The authors thank H. Nadel for helpful discussion.
34 Taylor and Schmidt
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