J. comp. Physiol.

80, 255--266 (1972)

9 by Springer-Verlag 1972

Communication by Surface Waves

M a t i n g B e h a v i o r of a W a t e r Strider (Gerridae)

R. Stimson Wilcox
Department of Zoology, School of General Studies, Australian National University,
Canberra

Received May 8, 1972

Summary. 1. During mating behavior males and females of an Australian

species of Rhagadotarsus (Gerridae) communicate with patterned sequences of
surface waves (signals) produced by leg movements (Figs. 1, 3). Individuals
produce signals while free on the surface or while grasping floating or fixed
objects; these objects then become copulation and oviposition sites (Fig. 2).
Wave patterns are produced during precopulation, copulation, postcopulation,
and aggression (Fig. 3). In the laboratory, mating behavior occurs diurnally, noctur-
nally and in total darkness.
2. Females were attracted and stimulated to oviposit by artificial precopula-
tory signals. Visual cues were shown not to be necessary for precopulatory
behavior and oviposition to occur (Table 1).
3. Female response to artificial calling signals of different frequencies corre-
sponded to the frequency range (approximately 17-29 waves/sec) present in male
calling signals; and females discriminated a difference in frequency of 1.5-2.0 waves/
sec (Table 2).

Introduction
Studies on surface-predating aquatic Hemiptera have demonstrated
these insects' ability to sense and orient relative to surface waves, b u t
have not examined whether this ability is involved in intraspecific
communication (Baerends, 1939; Rabe, 1953; Rensing, 1961; Wolda,
1961; Markl and Wiese, 1969; Wiese, 1969; Meyer, 1971; Murphey,
1971a, h). This s t u d y describes mating behavior in a species of the
water strider genus Rhagadotarsus Breddin (Gerridae, Rhagadotarsinae)
found near Cairns, Queensland, Australia. I n this species mating behavior
involves production and reception of patterned sequences of surface waves
(signals) b y males and females. The observations and experiments repor-
ted herein indicate t h a t this mode of communication functions in attrac-
tion of females to males, in stimulation leading to copulation and
oviposition, and in communication of aggression. To m y knowledge,
communication b y patterned surface waves during mating behavior has
not previously been described.
18 g. corn!0.Physiol., Vol. 80
256 R . S . Wilcox:

Fig. 1. Calling signal of patterned sequence of surface waves being produced b y

a male of Rhagadotarsus on a pond near Cairns, Queensland. Body of male is the
oval white spot at center of pattern. Male is grasping a small piece of wood
with his forelegs while producing signal. Air 28.8 ~ C, W a t e r 26.9 ~ C

Materials and Methods

Rhagadotarsus has not previously been reported in Australia. The small species I
studied (body length: males about 3.9 cm; females a b o u t 4.6 cm) close!y resembles
the description of R. kraepdini, a species known from India across to the Philip-
pines (Hungerford and Matsuda, 1960; Matsuda, 1960; Cheng and Fernando,
1969). Specimens of the species I studied are deposited in The Australian Museum.
The populations I studied inhabited quiet parts of the Barron River above
Redlynch, about 8 miles from Cairns. I n 1970-71, mating behavior occurred year-
round there. The description of laboratory mating behavior closely resembles
observations and motion picture films made in the field a t the Cairns locality
during 21-25 J a n u a r y 1971. Laboratery observations a n d experiments were made
in a 60 X 90 X 8 cm tray containing about 4 cm of water. The insects were main-
tained on vestigial-winged Drosophila.
 Communication by Surface Waves 257

Fig. 2. A Male of Rhagadotarsus producing signals while grasping a floating

styrafoam block. B Dead male model (foreground) held at styrafoam signal-gene-
rating float by glass stylus (angling from right). Live female, attracted by arti-
ficial signals, grasping right midleg of male with her foretarsi. C Copulation;
male above female. D Female (left) ovipositing in styrafoam; male remains nearby

Males of Rhagadotarsus produce signals when stationary, while (a) free on the
surface or (b) grasping floating or fixed objects on the surface with their forelegs
(Figs. 1 and 2A). In the laboratory, I glued a styrafoam float to a glass stylus
and attached the stylus to the balance arm of a Weston D. C. galvanometer posi-
tioned above the water surface in the tray. Males grasped and oscillated the float,
the output from the galvanometer being fed into a Grass P15 amplifier and a
Brush Mark 280 pen recorder (Fig. 3). Recordings of signals generated by indivi-
duals near the float closely resembled (except in amplitude) recordings from the
same individuals grasping the float. All photographs (except Fig. 1), recordings and
experiments were made at air temperatures between 20.0 ~ and 21.9 ~ C and water
temperatures between 18.5 ~ and 20.2 ~ C.
I conducted seven experiments in the laboratory. Experiment 1 examined
whether Rhagadotarsus would undergo mating behavior nocturnally and in total
18"
258 R . S . Wilcox:

darkness. Total darkness was achieved by taping an opaque sheet of plastic over
the entire tray. Mating behavior was recorded by connecting the galvanometer
and float system to a Rika Denkl Model B-1 activity recorder, and to the Brush
pen recorder.
Experiments 2-6 were designed to test (a) whether precopulatory signals (see
description in Mating Behavior) actually function in precopulatory communi-
cation, by observation of female response to artificial precopulatory signals;
and (b) whether visual cues presented by males are necessary for precopulatory
behavior and oviposition to occur, by observation of female response in the
presence or absence of a model (and its associated visual cues) during the gene-
ration of artificial precopulatory signals.
In experiments 2, 3 and 4 a dead male was positioned as a model on the
float by means of a horizontally-sliding jig which held a glass stylus to which
the model was glued (Fig. 2 B). The model could be moved back and forth on the
surface with the jig, in a manner similar to that of courting males. In experiments
5 and 6 the model and stylus were removed.
Artificial calling signals (Fig. 3C), courtship-calling signals (Fig. 3D) and court-
ship signals were generated by driving the galvanometer, stylus and float system,
with a Tektronix Type 161 pulse generator gated by a Type 162 waveform
generator which I triggered manually while watching the response of individual
females. Female response was scored (in increasing levels of response) according
to whether females (a) approached within 5-10 cm of the float; (b) approached
within 2-3 cm; (c) produced courtship signals while within 2-3 cm; (d) touched,
spanned or pulled a leg of the model with their foretarsi (after which the model
was moved back from the float about 1 cm); (e) grasped the float; (f) oviposited
in the float. This sequence of response levels corresponds to the normal sequence
exhibited by females in response to the signals of live males. Table 1 shows the
signals used in each experiment. Artificial signals were recorded by attaching a
second galvanometer and stylus system to the float.
New individuals were used for each experiment except in three cases, wherein
a female from one experiment was used in at most one other experiment. During
pilot trials, different positioning of the apparatus in the tank had no apparent
effect on results; and use of models made of dead females and varnish-coated dead
males gave results similar to use of dead males.

Fig. 3. A Calling signals of male of Rhagadotarsus. Central portion of each

signal approximately 18.5 waves/see. One signal expanded ten times (right);
similar expansions in B, C and D. B Courtship-calling signals. C Artificial calling
signals (17.5waves/sec). D Artificial courtship-calling signals. E At 1, male
begins signaling with courtship-calling signals, increasing the number of waves
to calling signals at 2; at 3 a female approaches within 5-10 cm of male, who
changes to courtship (e.g., three arrows-barely discernible blips) and courtship-
calling signals; between 3 and 4 female approaches within 2-3 cm; and at 4 female
grasps and pulls leg of male, causing movements of float. F Male copulatory
signals. See text for explanation. G At 1 a male produces courtship-calling signals;
at 2 another male approaches closely; at 3 first male produces aggressive signal,
but second male grasps float also, and dispute follows, with both males grasping
float; 4, lower frequency (9-13 waves/sec) signal; 5, higher frequency (23-30 waves/
sec) s i g n a l . - All time markers at 1-second intervals. A, B, C, D (right) recorded
at a chart speed of 20 cm/sec; all other signals recorded at 2 cm/sec
 Communication by Surface Waves 259

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 260 R . S . Wilcox:

Table 1. Results of experiments using artificial signals played to individual female

Rhagadotarsus (three females per experiment, 10 trials per female), with and
without a model on the signal-generating float. Each n u m b e r indicates positive
response, expressed as the average percent of all trials conducted per experiment.
A = calling signals (22.0 waves/see); B = courtship-calling signals, C = courtship
signals

Ex- Artifi- Model Increasing level of response

peri- cial
ment signals on re- approached within Produced Pulled/ Grasped Ovi-
float moved courtship spanned float potsied
5-10 cm 2-3 cm signals leg a

A B Cb X 100 83 40 23 0 0
B C X 100 80 35 13 0 0
Ac X 100 I00 100 100 10 0
Ac X 100 100 100 -- 80 20
B C X 55 20 0 -- 0 0

a Model moved back about 1 cm from float if female spanned and/or pulled leg
of model with her foretarsi.
b Calling signals switched to a mixture of courtship-calling and courtship signals
when female approached within 5-10 em.
c Signals decreased in intensity when female approached within 5-10 cm.

Experiment 7 examined the extent to which females were attuned to the

frequency range of calling signals. I tested the frequency response of three females
b y subjecting t h e m individually to calling signals produced (with no model present)
a t 10 different frequencies ranging from 12.5 to 32.0 waves/see (see Table 2). These
frequencies bracketed the normal calling signal frequency range (approximately
17-29 waves/see). Each female was subjected first to a test frequency, with a
m a x i m u m of 40 calls, and scored according to categories a, b and c of the score
response levels used in experiments 2-6, optimal response thus being defined as
production of courtship signals. If the response was suboptimal I waited (if
necessary) until the female h a d returned to 20-35 em from the float, then produced
a m a x i m u m of 40 calling signals of 22.0 waves/see as a control, discarding the
trial if the control response was suboptimal. The control frequency of 22.0 waves/
sec approximated the average frequency of the central portion of calling signals
(see Calling Signal Analysis).
A pilot experiment on female frequency response which involved test frequency
versus control frequency, produced alternately b y two galvanometers placed 20 cm
apart, gave results similar to b u t less consistent t h a n the design utilized herein.
The binary design had the i m p o r t a n t drawback t h a t females stimulated b y a n
optimal frequency (as defined herein) showed a higher response level toward
suboptimal frequencies immediately after being stimulated b y the optimal frequency
t h a n they did without prior optimal stimulation, thus biasing the data.
I n experiments 2-7, prior to trials each female was observed to be attracted
to a live male's signals, then isolated for at least i day. I watched the behavior of
a female for a t least 1 minute prior to a trial, began a trial when she was relati-
vely quiescent and 20-35 cm from the float, and (except where noted) used
3 females per experiment, with 10 trials per female. Consecutive trials were spaced
 Communication by Surface Waves 261

Table 2. Averaged response of three (or two where noted) females, tested indivi-
dually, to artificial calling signals of 10 different frequencies (waves/sec). Each
number indicates positive response, expressed as the average percent of all trials
conducted at the pertinent frequency

Waves/sec Increasing level of response

Approached within Produced No.
courtship trials
5-10 cm 2-3 cm signals

12.5 35 25 0 11 (2 ~ 2)
14.0 70 60 0 15 (2 ? 9)
16.0 83 63 23 30
17.5 83 67 67 30
20.0 100 100 70 30
22.0 100 100 100 98 (control)
25.0 100 100 100 30
27.5 97 97 90 30
29.0 93 93 67 30
32.0 47 43 7 30

at least 15 minutes apart. As the response of females was very similar, I have
pooled data within experiments except where noted in the discussion.
In Fig. 3, each upward peak corresponds to the crest of a wave. Signal frequen-
cies (waves/see) were c~Iculated from measurements between peaks, made with a
graticule (graduated in 1/10 ram) which was laid onto the graph paper and viewed
through a binocular microscope. Measurements were taken from signals recorded
at a chart speed of (a) 2 em/sec, where peaks were measured in groups of 5,
or (b) 20 era/see, where measurements were made between adjacent peaks. Using
artificial calling signals of 10 seconds' duration, recorded at both chart speeds,
calculations made from gi'aticule measurements were accurate to i 0.2-0.3 waves/
see in comparison to the average waves/see of the 10-second signals.

Results and Discussion

Mating Behavior
D u r i n g p r e c o p u l a t o r y behavior, males p r o d u c e r e l a t i v e l y large-ampli-
t u d e waves in groups either of 7-15 (designated calling s i g n a l s ; F i g s . 1
a n d 3A) or of 2 - 3 (designated courtship-calling signals; Fig. 3B), b y
in-unison p o s t e r o v e n t r a l a n d a n t e r o d o r s a l strokes of their mid]egs a n d
possibly also hindlegs. T h e y also p r o d u c e low a m p l i t u d e waves singly
or in groups of u p to a b o u t 30, b y vertical in-unison foreleg move-
m e n t s (designated courtship signals; Fig. 3 E , arrows). F e m a l e s a p p a r -
e n t l y p r o d u c e o n l y courtship signals, also b y v e r t i c a l foreleg m o v e m e n t s .
Males which are grasping objects b e g i n a n o r m a l p r e c o p u l a t o r y
sequence with calling signals (Fig. 3 E , 2), sometimes preceding these
w i t h several courtship-calling signals a n d t h e n progressively increasing
262 R.S. Wilcox:

the wave number into calling signals (Fig. 3 E, 1 to 2). Females respond
by moving more or less directly toward the signal source. When a
female approaches within 5-10 cm a male usually switches abruptly
from calling signals to courtship signals interspaced with courtship-
calling signals, diminishing the intensity of the courtship-calling
signals as the female approaches closer, and producing almost entirely
courtship signals when she is within 2-3 em (Fig. 3E, 3 to 4). When
within 2-3 cm, the female responds with courtship signals, and usually
also (a) touches or (b) spans and moves laterally or (c) grasps and briefly
pulls, a mid- or hindleg of the male with her foretarsi (Fig. 2B and
3E, 4). The male then releases the object and backs away about 1 cm; the
female moves forward, grasps the object and usually produces courtship
signals; and the male mounts and copulates (Fig. 2C).
In 7 recorded copulations, males copulated for 60-69 sec, producing
3-5 copulatory signals while copulating. Each signal began with
2-6 strokes of one midleg (Fig. 3F, duration of 1) then continued with
waggling of both midlegs with 4-27 waggles (Fig. 3F, duration of 2).
Whether these are strictly tactile or also surface wave signals is ob-
viously debatable.
After dismounting, the male backs away about 1 cm from the female,
remains stationary facing her, and produces sporadic posteopnlatory
signals, apparently identical to courtship signals, while the female releases
the object, turns around and maneuvers her hindlegs onto the object,
excavates a hole, and oviposits (Fig. 2D). After ovipositing the female
moves away, and the male usually returns to the object and begins
signaling again.
Males often attract (and subsequently copnlate with) females with
only courtship-calling and courtship signals if the females are nearby,
but soon begin calling signals if no females approach. Occasionally
males produce only calling signals before females approach within
2-3 cm.
After a female grasps an object she may not copulate but instead
turn around and oviposit, the male remaining nearby as after copulation;
or a female may leave without copulating or ovipositing. Both sexes
will copulate repeatedly (one male copulated 5 times in 1 hour, 3 of these
times with 1 female) and females will oviposit repeatedly without copu-
lating meanwhile.
Males free on the surface may produce calling signals which attract
females, then respond to a female's approach by approaching the
female. When within 2-3 cm of each other, both sexes may produce
courtship signals and they tend to remain near each other after such an
encounter, usually continuing to produce precopulatory signals; but
 Communication by Surface Waves 263

copulation and/or oviposition do not occur unless the male subsequently

grasps an object.
I saw females oviposit only in objects at which males had just
been signaling. Oviposition sites were therefore determined by males.
Females are probably not too stereotyped in choosing oviposition sites,
however, since in nature, solitary colonizing winged females would have
to oviposit in the absence of males.
During mating behavior males may produce aggressive signals with
their midlegs, apparently with regard to the possession or defense of
signaling sites. Aggression occurs mainly towards males, occasionally
toward females. Fig. 3 G shows the distinctive lower frequency (9-13 wa-
ves/see ; Fig. 3 G, 4) and higher frequency (23-30 waves/see ; Fig. 3 G, 5)
signals produced during an encounter between two males both grasping
the float. Males also produce these signals when free on the surface,
where fights often occur, usually as a consequence of the approach of a
male to within about 8-10 cm of a site at which another male is signaling,
at which point the signaler leaves the site, approaches the intruder,
produces aggressive signals and, if the intruder does not retreat, engages
in physical attack, returning to the site if successful in repelling the
intruder. Fights may last up to several minutes.

Experiments
Experiment 1. Use of the activity recorder showed that mating beha-
vior occurs nocturnally and in total darkness in the laboratory.
I pen-recorded one complete mating sequence, including oviposition acti-
vity, under total darkness conditions.
Experiments 2-6. Table 1 shows the results of experiments 2-6.
Experiments 2-4. With the model present at the float, generation of
calling signals (22.0 waves/sec) switched to a mixture of courtship-
calling and courtship signals when the female approached within 5-10 cm
(experiment 2); courtship-calling and courtship signals only (experi-
ment 3); and calling signals only (experiment 4), attracted females
within 5-10 em in all trials and within 2-3 cm in 80-100% of (all)
trials.
In experiments 2 and 3, females responded with courtship signals
in 35-40% of trials and spanned and/or pulled the model's leg in
13-23% of trials; but when the model was moved back about 1 cm
from the float after a leg was spanned and/or pulled, the females never
moved forward to grasp the float. However, calling signals only
(experiment 4) always elicited courtship signals and spanning and/or
pulling of the model's leg; and after the model was moved back from
the float, two of the three females grasped the float, in three trials
264 R.S. Wilcox:

(10% of trials). Because in normal conditions females are more often

exposed to, and respond optimally to, conditions imitated by experi-
ments 2 and 3 rather than 4, it seems likely that the courtship-calling
and courtship signals in experiments 2 and 3 were relatively less natural
than the calling signals in experiment 4, with less effective results.
Experiments 5 and 6. With the model absent from the float, genera-
tion of calling signals only (experiment 5) elicited as good a response
as when the model was present (in experiment 4). Moreover, females
oviposited in 20% of trials--perhaps because up to 20 calling signals
were generated after a female was within 2-3 em of the float, thus
maintaining wave stimulation more constantly than when, in experi-
ment 4, calling signals were stopped just before moving the model back.
(In support of this idea, during pilot trials, females attracted by calling
signals only, often would oviposit if calling signals were continued after
the model was moved back.)
On the other hand, generation of courtship-calling and courtship
signals only (experiment 6) elicited no response or, at most, attraction to
within 2-3 cm, a notable contrast to the success of experiments 2 and 3.
I repeated experiment 6 with one of the three females, with similar
results; then repeated it again with this female but with the stylus
(minus the model) resting on the float, thus testing whether (a) absence
of the model's visual characters or (b) alteration of wave signal character-
istics was responsible for the low response in the experiment. The female
then produced courtship signals in 7 of the 10 trials. I t thus seems likely
that the presence of a model or stylus on the float influenced charac-
teristics of the courtship-calling and/or courtship signals, and that the
absence of the model was of lesser importance to the females than the
alteration in signal characteristics.
The results of experiments 2-6 confirm laboratory and field obser-
vations that females are (a) attracted by and (b) stimulated to reply
with courtship signals and to grasp an object at the signaling site by (a)
calling signals and/or (b) courtship-calling and courtship signals; and
show thatcalling signals alone are sufficient to stimulate females to
oviposit in the laboratory. These results, plus the fact that females will
copulate and undergo oviposition activity in total darkness, indicate that
visual cues presented by males are not necessary for precopulatory beha-
vior and oviposition to occur.

Calling Signal Analysis

Analysis of calling signals showed that each signal comprises a stable
sequence of frequencies ranging between approximately 17-29 waves/see,
with low variability within and between males. For example, calling
signals of three males were recorded within 1 hour. Analysis of three
 Communication by Surface Waves 265

signals per male (ehart speed 20 em/see) showed that each signal began
with higher frequencies in the first two or three waves (23.3-29.4 waves/
see), stabilized at intermediate frequencies during the central portion
of the signal (18.7-24.5 waves/see) and ended with one or two low
frequency waves (about 10.0-17.4 waves/see). Mean frequencies for central
portions of signals for each respective male were 21.5 waves/see, standard
deviation • 1.06; 21.5 4- 0.72 ; and 21.1 4- 0.53, mean frequency between
males being 2 1 . 3 i 0 . 3 6 . By inspection, variability within males is not
significantly different from variability between males.
Males collected in August 1971 (winter) tended to produce calling
signals with lower frequencies, at the same laboratory temperatures,
than males collected in November (spring) 1971. For example, the
calling signals with central portions of 18.5 waves/see in Fig. 3A were
produced by a male collected on 3 August 1971; and the three males
whose signal analysis is reported above with calling signals centering
at about 21.5 waves/see--were collected on 16 November 1971. Also, evi-
dence from pilot trials indicated that both calling signal frequency
and female frequency response alter up or down with increase or de-
crease of laboratory temperature. The range of temperature in which
laboratory experiments were conducted was probably not large enough
to influence the results significantly. The three females used in
experiment 7 (below) were eolleeted on 16 November 1971.
Experiment 7. The results of experiment 7 (female frequency response)
are presented in Table 2, and show (a) that females were attracted to
within 5-10 em by all 10 frequencies, in 35% or more of trials, (b)
that when females were attracted within 5-10 em they normally con-
tinued to approeh to within 2-3 em and (e) that courtship signals (opti-
mal response) were produced in 67 % or more of trials at frequencies
between 17.5-29.0 waves/see, optimal response being elicited in at most
23 % of trials at frequencies above and below this range.
Individually, the three females responded optimally in 60 % or more
of trials to frequencies between 22.0-29.0 (female A); 16.0-29.0 (female
B); and 17.5-27.5 (female C) waves/see, each responding optimally to
frequencies above and below these ranges in at most 20 % of their respec-
tive trials. Thus the response of females t3 and C especially corresponded
to the range of frequencies present in male calling signals. In addition,
female A distinguished a difference in frequency of 2.0 waves/see (test
frequency of 20.0 waves/see versus control frequency of 22.0 waves/see)
in 9 of 9 trials; and female C distinguished a difference in frequency
of 1.5 waves/see in 9 of 9 trials where I altered the experimental design
and used 17.5 rather than 22.0 waves/see as the control frequency versus
the 16.0 waves/see test frequency. These frequency discriminations were
tested at the lower limit of each female's optimal response range.
266 R. S. Wilcox: Communication by Surface Waves

Evidence from pilot trials on other females indicated discrimination of

similar a c u i t y at the higher limit.

This investigation was supported by an Australian-American Education Foun-

dation grant and by funds from the Department of Zoology, The Australian
National University.

References
Baerends, G.P.: Waarnemingen en proeven aan de ruggezwemmer Notonecta
gIauca. De levende natuur 42, 11-17, 45-51 (1939).
Cheng, L., Fernando, C. H. : A taxonomic study of the Malayan Gerridae (Hemi-
ptera:Heteroptera) with notes on their biology and distribution. Oriental
Insects 8, 97-160 (1969).
Hungerford, H. B., Matsuda, R. : Keys to subfamilies, tribes, genera and subgenera
of the Gerridae of the world. Univ. Kans. Sci. Bull. 41, 3-24 (1960).
Markl, H., Wiese, K.: Die Empfindlichkeit des Riiekensehwimmers Notonecta
glauca L. ffir Oberfl~chenwellen des Wassers. Z. vergl. Physiol. 62, 413-420
(1969).
Matsuda, R.: Morphology, evolution and a classification of the Gerridae
(Hemiptera--Heteroptera). Univ. Kans. Sci. Bull. 41, 25-632 (1960).
Meyer, H.W.: Visuelle Sehliisselreize fiir die AuslSsung der Beutefanghandlung
beim Bachwasserl~ufer Velia caprai (Hemiptera, Heterop~era). Z. vergl. Physiol.
72, 260-297 (1971).
Mm~hey, R . K . : Motor control of orientation of prey by the waterstrider,
Gerris remiqis. Z. vergl. Physiol. 72, 150-167 (1971).
Murphey, R . K . : Sensory aspects of the control of orientation to prey by the
waterstrider, Gerris remiqis. Z. vergl. Physiol. 72, 168-185 (1971).
Rabe, W. : Beitr~ge zum 0rientierungsproblemder Wasserwanzen. Z. vergl. Physiol.
85, 300-325 (1953).
Rensing, L. : Beitr~ge zur vergleichenden Morphologie, Physiologie und Ethologie
der Wasserl~ufer. Zool. Beitr. Berlin (N.F.) 7, 447485 (1961).
Wiese, K.: Wahrnehmung yon Oberfl~chenwellen geringer Amplitude durch den
Wasserl~ufer. Naturwissenschaften 56, 575 (1969).
Wolda, H. : Response decrement in the prey catching activity of Notonecta glauca
L. (Hemiptera). Arch. neerl. Zool. 14, 61-89 (1961).

Dr. R. Stimson Wilcox

Department of Zoology
School of General Studies
Australian National University
Canberra, Australia

Present address:
Department of Biological Sciences
Purdue University
Lafayette, Indiana 47907, U.S.A.