BIOACOUSTICS, MATERNAL INVESTMENT,
AND DEVELOPMENTAL STRATEGIES
IN THE MOLE CRICKETS,
SCAPTERISCUS ACLETUS AND VICINUS
TIMOTHY G. FORREST
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
I believe in fires at midnight...
Silken mist outside the window-
Frogs and newts slip in the dark
Too much hurry ruins a'body
I'll sit easy, fan the spark
thank you for everything.
Now is the solstice of the year.
Winter is the glad song that you hear.
Ring out, ring Solstice Bells.
I am most appreciative of the guidance that Tom Walker and Jim Lloyd have
given me. Their high standards of scholarship and science have been truly inspiring.
I thank them both for what I have received in terms of knowledge, opportunity, and
friendship. My work will always be a product of their teachings. I wish to thank
Marty Crump for adding a new dimension to my interest in animal sound
production. Her knowledge of anuran biology has been most rewarding and has
made me look at insects in a different light. All three gave helpful suggestions that
much improved this dissertation.
Henry Bennet-Clark, Dave Green, Les Bernstein, and Ginny Richards have
all been helpful in transmitting their knowledge of sound and acoustics, without
which I would never have completed Chapter I. Ken Prestwich provided
unpublished data and his help with the energetic of sound production was
invaluable. Frank Slansky was extremely helpful with comments on Chapters II and
III that much improved their content. Bill Oldacre provided the schematics for the
portable multiplexing unit and was always willing to give his time. John Allen
furnished a computer program that made the mentally impossible visually possible.
Sue Wineriter gave many hours of work on many drafts of many figures. Her
expertise in art and illustration is much appreciated. I would also like to thank my
friends Will Hudson and Steve Wing for making this time and degree an enjoyable
and enlightening one.
I also wish to acknowledge my parents for their help when help was needed
most. And finally I thank my family. Sue, I thank for her devotion and dedication
that have given me confidence in myself and the drive to finish. Her many hours of
assistance in the field will always be remembered. Justin, I thank you for making
Sue and me very happy.
TABLE OF CONTENTS
LIST OF FIGURES. vi
I BIOACOUSTICS 3
Introduction . 3
Materials and Methods 21
II OVIPOSITION AND MATERNAL INVESTMENT 36
Introduction . 36
Materials and Methods 37
III SIZE TACTICS AND DEVELOPMENTAL STRATEGIES 55
Size Tactics in Mole Crickets 62
Materials and Methods 63
REFERENCES CITED 78
BIOGRAPHICAL SKETCH 84
LIST OF FIGURES
1. Pressure variation in a 2 Hz sine wave. 6
2. Phase relationship between velocity
and displacement of a pendulum. 10
3. Acoustic interference and the
summation of sinusoids. 14
4. Sound produced by a vibrating disc.. 19
5. Relationship between total power output,
male size and soil moisture in Scapteriscus. 26
6. Total power output as a function of
sound pressure above the calling burrows of
Scaptcriscus acltus. 29
7. Sound field of Scapteriscus acletus. 31
8. Scapteriscus egg laying seasons. 41
9. Relationship between female size and
total offspring produced in acletus.. 45
10. Relationship between female size and
average number of eggs per clutch in acletus. 47
11. Effect of female age on mass of eggs produced in
12. Costs and benefits associated with maturing. 58
13. Hypothetical cost/benefit curve associated
with changes in an individual's size. 60
14. Relationship between adult pronotal length
and mass in Scapteriscus. .66
15. Seasonal size distributions in mole crickets. 69
16. Seasonal change in size distribution in
mole crickets.. 72
Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
BIOACOUSTICS, MATERNAL INVESTMENT,
AND DEVELOPMENTAL STRATEGIES
IN THE MOLE CRICKETS,
SCAPTERJSCUS ACLETUS AND VICINUS
Timothy G. Forrest
Chairman: Thomas J. Walker
Major Department: Entomology and Nematology
Three aspects of the reproductive ecology of the southern and tawny mole
crickets, Scapteriscus acletus and vicinus, were investigated.
Twenty-two sound fields of calling males were measured under field
conditions. Total power output ranged from 1.9 to 22.4 uWatt and was positively
correlated with male size and soil moisture. Efficiency of sound production was
only 0.02 to 0.16 percent. The shape of the sound field approximates a hemisphere.
Lifetime oviposition by caged, individual female mole crickets was
monitored outdoors. Oviposition is highly seasonal, and females that mature in the
fall wait until the following spring to oviposit. Most eggs are laid in May and June
and percent hatch of clutches decreases later in the season. Dry weight of eggs,
number of eggs per clutch, and percent hatch decrease with successive clutches from
individual females. Average investment in eggs per clutch ranged from 10 to 24
percent of female body weight for acletus and from 5 to 16 percent for vicinus
females. Lifetime investment (dr weight of eggs) was greater than 100 percent of
female body weight (wet weight) for 6 of 12 aclctus females, but only 1 of 8 vicinus
females had a lifetime investment in eggs equaling its body weight. In acletus, but
not vicinus, large females produced more offspring and the average size of their egg
clutches was larger than those of smaller females.
Insect size tactics and developmental strategies are discussed in relation to
decisions individuals make about when to mature. Such decisions carry with them
costs and benefits in terms of an individual's fitness. Whenever size affects
reproductive success, selection should act such that individuals evaluate the costs
and benefits due to changes in size and mature when the cost/benefit ratio is
maximized. Predictions about seasonal changes in adult sizes within a natural
population of mole crickets are tested. The changes in seasonal distributions of
adult sizes of mole crickets support the predictions and suggest that individuals
make decisions about when to mature based on costs and benefits associated with
changes in size.
The southern and tawny mole crickets, Scapteriscus aeletus and vicinus, were
accidentally introduced into the United States around the turn of the century
(Walker and Nickle 1981). Because they lack a specialized predator/parasite
complex their populations have gone unchecked, their numbers have increased, and
they are one of the major pests of turf grasses and pasturelands in the southeast.
Their economic importance has prompted large amounts of funding for research
aimed at controlling mole crickets and lessening the damage caused by them. Only
recently has there been progress toward this goal. The primary stumbling block has
been an insufficient knowledge of the basic biology of these pest species. During
the past decade numerous researchers have contributed to our understanding of the
animals and their elusive subterranean lifestyle.
Mole cricket biology. Scapteriscus spp. are usually associated with habitats
that are temporary and early successional. In the southeastern United States both
acletus and vicinus are common in pastures, cultivated fields, lawns, and other turf.
All adults have long hindwings and are capable of flight. Dispersal flights occur
during the spring and fall, consist mostly of females, and are important to the
crickets' ecology in that they allow individuals to colonize newly opened areas or to
leave unsuitable or deteriorating ones (Ulagaraj 1975, Walker et al. 1983). Flying
crickets land in response to their species-specific calling song (Ulagaraj 1975,
Ulagaraj and Walker 1973, 1975). Adults are easily collected at traps broadcasting
such songs. Both males and females respond to the calls and preferentially land
near males whose calls are louder than those of neighboring males (Forrest 1980,
The two species differ markedly in their food habits with acletus being
primarily carnivorous on soil arthropods, earthworms, etc., whereas vicinus is
mainly an herbivore, feeding on roots as well as grass blades (Matheny 1981).
In Gainesville both species are univoltine; however, they differ in that about
85 percent of the vicinus population overwinter as adults compared with only 25
percent of the acletus population (Hayslip 1943). The rest of the populations
overwinter as late-instar juveniles that mature the following spring when eggs are
laid. Egg clutches are laid during the spring in underground ovoid chambers (4 by 3
cm) constructed by females. Once the eggs are laid, the chamber is sealed and no
parental guarding of eggs occurs. Eggs within a clutch hatch synchronously (within a
24-h period), and after a few days the young tunnel to the surface.
The topics that follow are part of the ongoing effort to understand mole
cricket populations and biology. The subjects add to the understanding of the
reproductive ecology of mole crickets and should be useful in monitoring changes in
the population and in designing models concerning mole cricket population
dynamics. Chapter I covers the bioacoustics of mole crickets and is an investigation
in the efficiency of acoustic communication by the two crickets. Chapter II deals
with female oviposition and the maternal investment by females and how these are
influenced by season, female size and female age. The third chapter is a study of
how individuals evaluate costs and benefits associated with size and how these
relate to the decisions of whether or not to mature.
Let me bring you songs from the wood:
To make you feel much better than you
Since man's beginnings the sounds of animals have frightened, amused, and
intrigued him. With the advent of the tape recorder and audiospectrograph, he
could accurately quantify the sounds of animals, and in the last two decades the
interest and research in the areas of animal communication and bioacoustics have
increased dramatically. A new generation of digital recording and measuring
devices has made endeavors into bioacoustics even more reliable and from an
empirical standpoint more profitable. It is from the world of insects that such
research has really profited. Here there is a wealth of species with a wide variety of
sound producing and sound receiving mechanisms, many hundreds of which have
evolved independently, each one having been shaped by natural selection for
efficient communication purposes. A thorough background of the physical nature
of vibration and sound will be given and this understanding will be the basis for a
discussion of mole cricket bioacoustics and sound production.
Vibration and Sound
In simple terms sound is a disturbance or vibration in some elastic medium. This
disturbance need not be uniform or repetitive. Such a disturbance transmits energy
to the particles of the medium; the particles move from their resting position,
collide with neighboring particles and transmit the energy to them. These neighbors
transmit the energy to other neighbors and so on. Because of the elasticity of the
medium, the particles tend to return to their resting position after being displaced.
Thus the energy is transferred in waves away from the source of the disturbance.
Waves can be of two types depending on the axis the particles move relative to the
movement of the wavefront. When the particle motion is perpendicular to the
movement of the wave it is said to be transverse, whereas particle motion parallel to
the motion of the wave is called longitudinal. Sound waves in air are longitudinal;
waves on the surface of water are transverse.
The characteristics of a sound will depend upon the type of disturbance as
well as the physical characteristics of the medium. In most instances the medium is
air, although sound can be transmitted through liquids and solids and many insects
use these media for communication. For simplicity I will restrict my discussion to
sound in air, although the principles generally apply to all sound.
Pressure, power and intensity. A disturbance in air causes a change in the
density of molecules and in the pressure of the gas. These fluctuations in the
pressure of a sound wave are easily detected and measured with instruments, such
as microphone, that are sensitive to such pressure changes. Because of their ease of
measurement, changes in instantaneous pressures are commonly used to describe
sound. Perhaps the most well known sounds are sinusoids in which the pressure
fluctuates in a sinusoidal manner and can be described by the equation
where pressure (p) is a function of time (t), A is the amplitude of the pressure
change, pi is the constant 3.14159..., f is the frequency of the sine wave and theta is
the pase angle. Since the instantaneous pressure changes sinusoidally with respect
to both time and space, p(x), the pressure at some distance (x) from the source, can
Figure 1. Pressure variation in a 2 Hz sine wave. Pressure, p, is a sinusoidal
function of time, t, and the wave has an amplitude of A. Positive
values of p(t) arc compressions and negative values are rarefactions.
also be described by substituting x/c for t in the above equation, where c is the
velocity of sound in the medium. In air, c is about 330 meters per second. Note
that the plus and minus signs in the space formula are used to describe sound
travelling in opposite directions from the source.
Figure 1 shows the instantaneous pressure changes for a 2 Hz (cycles per
second) wave with initial phase angle theta=0. At t=0 and x=0 the pressure is 0,
although the pressure may have been any value between -A and A depending on the
initial phase angle. With an initial phase angle of 900 the pressure at t=0 is equal to
A. When pressure, p(t), is positive, the particles of the medium are compressed.
This part of the sound wave is called the compression. When the pressure is
negative, a vacuum occurs and this part of the wave is the rarefaction. The sinusoid
in Fig. 1 is repetitive every 1/2 second because the sin(0)=sin(2*pi); i.e. the period is
1/2 second and is equal to the reciprocal of the frequency for any sinusoid. The
wavelength (lambda) of the sinusoid is the distance the wave travels in one period
and equals period*c or c/f. The 2 Hz wave has a period of 1/2 second and a
wavelength of 115 meters. Insects typically use much higher frequencies with
shorter wavelengths. For instance the calling song of the southern mole cricket,
Scapteriscus acletus, has a carrier frequency of 2.7 kHz with a wavelength of 12 cm.
In dealing with sound one often wishes to know the power or intensity (I) of
the sound. Intensity differs from the scalar quantity of pressure, in that intensity is a
vector quantity having both magnitude and direction and its units are given in
energy per time per unit area or Watts per square meter (W/m2) whereas units of
pressure are in force times area (Newton-meter2: Nm2). Sound power, the total
power produced by a source, is measured in the same units as electrical power, the
Watt. Since pressure is an easily measured aspect of sound it is important to
understand the relationship between the scalar pressure and the vector intensity.
Intensity is the product of the instantaneous pressure (p) and particle velocity (u)
I=p(t)*u(t) or p*u. 2
This equation is analogous to the power equation used in electrical systems where
power is voltage times current.
Particle velocity describes the fluid movement of the medium. Near the
source particle velocity and pressure are 900 out of phase. This relationship is
particularly important and has relevance to measurement of pressure, and therefore
intensity. To illustrate this phase difference consider a pendulum (Fig. 2). At its
resting position the pendulum hangs vertically and there is no movement, no kinetic
energy and no potential energy. However, suppose the pendulum is displaced some
distance, x, from the resting position. If it is held at this position, the pendulum has
obtained some potential energy but has no kinetic energy. Releasing the pendulum
will cause it to swing toward its resting position and there is an exchange between
potential and kinetic energy. When the pendulum has reached its resting position
its velocity is maximum, its displacement a minimum. The pendulum then swings
past the resting position, and because of gravity the kinetic energy begins to change
back to potential energy. At the end of the swing the pendulum reaches its
maximum displacement (now -x) and its velocity has slowed to a minimum of zero.
It can be seen that the phases of displacement (=pressure) and velocity (=particle
velocity) are shifted by 900. The phase relationship changes as the sound moves
away from the source and can be described by
Tangent theta=lambda/r. 3
Lambda is the wavelength of sound, r is the distance from the source, and theta is
the phase angle (in radians) between particle velocity and pressure (Bennet-Clark
1971 and refs). It can be seen that the phase shift approaches 900 as r becomes zero
and the phase difference between particle velocity and pressure decreases to zero as
the distance r increases. Though the boundaries are not well defined, the near field
is where pressure and particle velocity are not in phase, whereas in the far field
particle velocity and pressure are in phase. For practical purposes the far field
begins at distances greater than about 1 wavelength of the lowest frequency sound
produced by the source. In the far field the sound field is said to be purely active
(Gade 1982). This is useful because in the active field particle velocity can be
calculated from pressure using the acoustical equivalent to Ohm's law, current
equals the ratio of voltage to resistance:
where poc is the characteristic impedance of the medium, about 40 dyne*sec/cm3
for air (Gade 1982). Substituting p/poc for particle velocity in the intensity equation
(2) it can be seen that intensity is proportional to the square of the pressure
and can thus be calculated using measurements of pressure-- provided these
measurements are taken in the far field.
Decibels. Pressure, power and intensity levels are often given in decibel
units. These units are logarithmic expressions of the ratio of any two sound
pressures or powers. Normally the levels are described relative to some standard
reference. For instance, the commonly used reference for power level is 10-12 W,
that for intensity is 10-12 W/m2, and that for pressure is 2x10-5 Nm2 = 2x10-5 Pa =
.0002 dyne/cm2 = .0002 ubar. All of these standards represent 0 decibels (dB).
Sound intensity, I in dB, can be calculated using the equation
I=10 log W/Wr 6
where W is the intensity and Wr is the reference or 10-12 W/m2. An intensity ratio
of 2 (i.e. W/Wr = 2) is 3 dB relative to the reference. A power level, PWL, can be
calculated using the same equation except that the reference is 10-12 W. Often the
terms power and intensity are interchanged although each has a specific meaning.
Similarly sound pressure level, SPL in dB, can be calculated by
SPL= 20 log P/Pr. 7
In this equation P is the pressure and Pr is the reference. A pressure ratio of 2
equals 6 dB relative to the reference. Note from the equations that for any given
level in dB the power ratio and intensity ratio, W/Wr, equals the square of the
pressure ratio, P/Pr. Since 0 dB corresponds to both references, a sound pressure
level can be converted into an intensity level directly, provided, of course, the
pressure is measured in the far field.
Acoustic interference. Consider two sound waves travelling in opposite
directions. One logical question to ask is: What happens when the two waves
meet? The resultant pressures are simply the sum of the pressures of the two waves.
This is graphically shown for a pair of equal-amplitude sinusoids in Figure 3. In
Fig. 3a the waves are approaching each other from opposite directions. In 3b the
waves are positioned in space so that a portion of the waves occupying the same
space are in phase (i.e. the compressions of one wave occurs with the compression
of the other) and the pressures add to produce pressure that is greater than the
pressure of each individual wave. This is called constructive interference. In 3c the
pressures sum to zero since the phases of the individual waves are opposite and they
exactly cancel. This condition is destructive interference. As each wave continues
on its way they are unchanged even though there were great pressure changes when
the two waves met. When two sounds interfere destructively the energy of the
sound is not lost. When the pressure decreases, the particle velocity increases and
Figure 3. Acoustic interference and the summation of sinusoids.
a. Two equal-amplitude sine waves of the same frequency travelling
in opposite directions toward each other.
b. During this instant in time the two occupy the same space and the
pressure of the resultant sinusoid (solid line) is the sum of the two
individual waves. The resultant has an amplitude greater than either
of the individual waves; this condition is termed constructive
c. At this instant the individual waves are opposite in phase and their
pressures sum to zero (solid line). This is destructive interference.
-- -- -- - .. .. .. .. .
:I: : ::
:: :: -:I
: II '
I :; :: :: ::
I : : : i : : :
: : : :
i : : : :
there is a conservation of energy. The sum of any two sinusoids that have the same
frequency is another sinusoid of the same frequency but with a different amplitude.
Propagation and the inverse square law. In an ideal environment sound
waves radiate at the same rate in all directions from the source, and thus, the
wavefront is spherical in nature. Because the same acoustic energy must occupy a
greater surface area as the distance from the source increases, sound intensity
decreases with distance from the source, since intensity is measured in Watts per
unit area. The surface area of a sphere increases proportionately with the square of
its radius (4*pi*r2), and thus the intensity of a sound is inversely proportional to the
square of the distance from the source--the inverse square law.
Suppose the intensity is measured at some distance from the source. The
intensity at twice that distance will be one-fourth or about 6 dB less [10*log(0.25)].
If the power ratio is 0.25, the pressure ratio is 0.5. For the same doubling of
distance the pressure will also decrease by 6 dB [20*log(0.5)]. Pressure, the scalar
quantity, is proportional to the distance from the source. For every doubling of
distance the pressure is halved and it is this proportionality that will be used to
calculate sound fields of crickets.
The study of insect bioacoustics, the special problems that insects have in
using acoustic signals to communicate, and the biophysical aspects of sound
production began more than 15 years ago and have been reviewed by
Bennet-Clark 1971, 1975, Michelsen and Nocke 1974, Michelsen 1983 and Elsner
and Popov 1978. The interest in the field has continued to grow primarily because
the animals are easily maintained, they are easily manipulated for experimental
purposes, and they offer a host of sound producing mechanisms to study.
Many insect groups use acoustic signals to communicate: ants do it, cicadas
do it, lacewings do it, and moths and beetles do it. The complexity of the
communication varies from very simple sounds of hitting a part of the body on the
substrate to sounds that are extremely complex with distinct signals for special
contexts.- For instance crickets have different types of calls classified as calling,
courtship, or aggressive (Alexander 1961). The different life stages of passalid
beetles use acoustic signals to communicate, adult to young and young to adult.
These beetles have a repertoire of as many as seven different sounds used in 13
different behavioral contexts (Schuster 1975). Some species of insects may use
acoustic signals to communicate to other species of animals--for example, many of
the tiger moths that use sounds to confuse and deceive bat predators (Fullard et al.
However, insects are very inefficient at producing sound because the
efficiency of sound production is dependent on the relationship between the size of
the oscillator or sound source, the frequency of the sound produced by the oscillator
and the immediate surroundings of the oscillator. To be an efficient radiator, the
diameter of the sound source must be about the size of, or larger than, the
wavelength of sound produced by the source. Because insects are small, and thus
their sound producing organs are small, they must produce a sound with very small
wavelengths to be efficient. That is, they must use high frequency sounds. However,
there is a disadvantage. High frequencies are attenuated by air at a faster rate than
low frequencies (Wiley and Richards 1978, Michelsen 1978). Therefore, given the
same power output for a high and low frequency signal, the intensity at the same
distance from the source will be greater for the low frequency than the high
frequency sound. Because of this, low frequency sounds are better for long distance
communication and insects must make a trade-off between the effective distance of
communication and the efficiency of signal production.
Insect muscles cannot contract at the high rate needed to produce
frequencies in the kHz range. Therefore, insects usually use some mechanism for
frequency multiplication. Most often this is accomplished by stridulation (Elsner
1983). During stridulation one part of the insect body is rubbed against another.
One part usually has a row of teeth, and during each stroke a number of teeth are
struck, each adding to the sound. The frequency of the muscle contraction is
multiplied by the number of teeth hit.
One of the most familiar groups of singing insects are the Orthoptera. This
group contains the grasshoppers, which produce song in at least three different ways
(Otte 1970). It includes cockroaches, some of which produce a hissing sound by the
expulsion of air from the spiracles (Nelson and Fraser 1980). But above all, the
crickets and katydids are the most noted and familiar singers of the group. Few
people have not heard a cricket.
Sound production in the Ensifera. The Ensifera are an ancient insect group
and include the Tettigoniidae (katydids), the Gryllidae (crickets), the Gryllotalpidae
(mole crickets) and the Haglidac. The forewings are similarly modified for sound
production in the four families. While calling the wings are raised above the
abdomen and moved back and forth one on top of the other. Located on the
underside of the Cu2 vein of each wing is a row of teeth called the pars stridens or
file. On the posterior margin of each wing is the plectrum or scraper. Stridulation
occurs during the closing stroke of the wings when the plectrum is pulled against the
grain across the teeth of the file. In katydids the left wing is usually over the right,
in crickets the right is usually over the left, while in haglids and mole crickets each
configuration occurs with equal frequency (Masaki et al. in press, Forrest in press).
Figure 4. Sound produced by a vibrating disc.
a. As the disc moves to one side it compresses the particles of air and
at the same time produces a vacuum on the other side of the disc.
The shading of the circles represents the degree of displacement of
the particles (ie. black circles are compressions and no circles are
b. Sound output and phase relationships of a dipole source.
c. Sound field of the dipole source. If the vibrating disc is small
relative to the wavelength of sound produced, the outputs from each
side of the disc will destructively interfere along the edges of the
disc. The result is a dumbbell shaped sound field.
-,-/I I IIII
In some species the wings have become specialized for their tasks and there is an
asymmetry between the left and right wings.
The file and scraper are closely associated with specific membranous areas of
the wings. The cell or cells are caused to vibrate at the frequency of the
tooth-impact rate of the scraper on the file. The cells are resonant near the
impact frequency, and the vibrations of the cells produce the sound.
The vibration of the wing membrane can be likened to the movement of a
piston or disc. As the disc moves back and forth in air, it produces a compression
and a rarefaction simultaneously on opposite sides of the disc. That is, a sound
wave is generated from each side of the disc, each being equal in amplitude and
frequency, but opposite in phase (Fig. 4). When the diameter of this doublet or
dipole source is small relative to the wavelength of sound produced, the sounds
radiating from the two sides of the disc destructively interfere along the edges of the
disc (Fig. 4b). The result is a dumbbell shaped sound field with a maximum
pressure perpendicular to the disc surface and a minimum pressure along its edges
(Fig. 4c). Of course the particle velocity at the edges is high and the intensity is
equal in all directions.
The membranes of crickets are less than one-tenth the wavelength of sound
produced, and they are inefficient primarily because of their size. Energetically,
producing sound may cost a male ten times as much as resting (Prestwich and
Walker 1981). However, because calling contributes importantly to a male's
reproductive success, selection should be strong on males to increase signaling
efficiency and effectiveness. Crickets have evolved a number of behaviors that use
baffles to increase calling efficiency (see Forrest 1982 and refs.). The study of these
baffles and their effect on sound production is one important aspect of cricket
mating systems and reproductive biology.
One such baffle system is used by the southern and tawny mole crickets,
Scapteriscus acletus and vicinus. Both species construct a calling burrow shortly
after sunset, prior to the female flight period. The burrow is shaped by the cricket
and expands exponentially from the throat to the opening at the soil surface
(Nickerson et al. 1979).
Because the males call from within a burrow, their sound fields are very
suitable for measuring. The position of the male in the burrow is constant, as is the
position of the burrow opening. The calling songs are continuous trills that change
less than 1 dB in sound pressure during the 1 to 1.5 hours of calling (Forrest 1981,
1983), and an observer can move around outside the burrow with little disturbance
to the calling male. If the animal is disturbed, he runs down into the burrow system
but returns in a short period of time to the same calling position and resumes
advertising for females.
I measured 22 sound fields of Scapteriscus males (17 acletus, 5 vicinus) under
field conditions to determine the power output of the crickets, the efficiency of
sound production and the directional properties of the sound field.
Materials and Methods
Sound pressure level readings were taken using nine Bruel & Kjaer, model
4125 condenser microphones and type 2642 preamplifier. The typical frequency
response for these microphones is flat (1 dB) from 200 Hz to 10 kHz for a free
field response and an angle of incidence of 00. Directional characteristics show less
than 1 dB loss for +300 deviations from a 00 incidence angle for signal frequencies
at 2, 4, and 8 kHz (Bruel & Kjaer specifications).
The output of each microphone could be adjusted independently, and each
microphone was calibrated using a model 4230 B&K pistonphone calibrator.
Initially, each microphone was calibrated after each sound field was measured.
Later the calibration was done every two weeks. The stability of the microphones
was such that later they were calibrated only every month. Never was the level of
any of the microphones more than 0.5 dB from the calibration level of 94 dB.
The outputs of the microphones were multiplexed to a single input of a
model 2219 Brucl & Kjaer sound level meter that measured the sound pressure re
0.0002 dyne/cm2 (dB A). The entire system was battery operated and field portable.
Cables from the microphones were long enough so that the observer and equipment
were more than a meter from the sound source and nearest microphone. This
avoided reflection of sound waves off the observer's body, and interference in the
pressure readings was reduced to a minimum.
The nine microphones were equally spaced around a semicircle of tubular
aluminum (every 22.50, +20) such that the diaphragm of each microphone was 25
cm from the center of the semicircle and the angle of incidence was zero (+2). An
angle of incidence of 00 is recommended when the direction of the sound source is
known (Brucl 1983). At 25 cm the microphones were always more than 1
wavelength of the carrier frequency of the calling song of the mole crickets (acletus
2.7 Khz or 12 cm wavelength, vicinus 3.3 Khz or 10 cm wavelength). Thus sound
pressure measurements were taken in the far field (Michelscn 1978).
By rotating the semicircle in 22.50 steps along one of the axes, sound
pressure levels could be taken at every 22.50 latitude and longitude of a hemisphere
surrounding the sound source located on the ground in the center of the semicircle.
A stand anchored one end of the hemisphere to the ground and a handle that
moved the semicircle locked every 22.50. Axis of rotation was always with the
longitudinal axis of the cricket so that on every 22.50 rotation the two microphones
at the ends of the semicircle remained at the same anterior and posterior position
relative to the cricket burrow. This arrangement provided a check for changes in
the crickets' output and added reliability to the measurements. Altogether, pressure
measurements were taken at 65 different positions surrounding the opening of the
cricket's burrow with the anterior and posterior positions each having 9 separate
measurements. Sound fields with more than +0.5 dB variation in these 9
measurements were not used in analysis. These measurements represent the most
detailed measurements of cricket sound fields.
All sound fields were measured outdoors at the Horticultural Farm of the
University of Florida. Sound fields of crickets were measured while they called in
the field or while they called from soil-filled, 12-liter buckets buried so that the
bucket lip and the soil in the bucket were at ground level. Only crickets calling on
level ground were measured and all obstacles that might cause interference or
reflections were removed from the area.
To determine the influence of male size on acoustic output, the males were
captured and the lengths of their pronota along the midline were measured to the
nearest 0.1 mm using vernier calipers. The mass of each male was measured to the
nearest 1.0 mg with a model AC100 Mettlcr balance. A soil sample (2 cm dia X 4.5
cm deep) was taken within 5 cm of the burrow opening. This sample was dried and
moisture content determined. Soil temperature at a depth of 2.5 cm was also taken.
Calculation of sound fields and power output. Each of the 65 sound pressure
levels measured at 25 cm was converted to a radial distance from the cricket at
which the pressure would be 70 dB. For instance a pressure that read 73 dB at 25
cm would be 70 dB at about 35 cm. The points were converted to polar coordinates
and the areas of all triangles defined by adjacent triads of points were summed to
obtain the surface area of the sound field. A test of this procedure revealed that it
underestimates the surface areas of 24-200 cm dia hemispheres by only 2-3%.
Table 1. Power input, output and efficiency of calling male Scaptcriscus.
PNL MASS TEMP PULSEa SOIL POWER PWL POWERb EFF.
RATE MOIST OUT IN
mm mg 0C p/sec uW dB mW %
7.7 574 26.8 56.2
7.7 574 25.2 54.0
7.7 574 25.4 54.3
7.7 574 30.7 61.7
7.7 574 27.2 56.8
8.1 614 29.1 59.4
8.1 614 25.7 54.7
8.6 737 25.8 54.8
8.6 737 30.2 61.0
8.6 737 30.8 61.8
8.8 776 20.4 47.3
8.9 676 28.8 59.0
8.9 676 27.7 57.5
9.4 772 22.5 50.2
9.5 875 22.4 50.1
9.9 1172 19.1 45.4
10.0 1162 20.9 48.0
8.3 890 N.A.
8.3 890 25.4 139.0
8.4 889 26.6 145.3
8.4 834 21.4 118.0
8.4 N.A. 21.2 116.9
a Based on relationship to soil temperature (Ulagaraj 1976).
b Based on data from Prestwich and Walker 1981.
N.A.- Not available or data not taken.
Figure 5. Relationship between total power output and male size and total power
and soil moisture in Scapteriscus.
a. Total power (dB re 10-12 W) as a function of male size (pronotal
length mm). Power (in Watts) of males with more than one sound
field measurement were averaged for regression analysis. Vertical
lines at these points show the range in power for individual qales.
Regression equation is y = 2.4x + 46.4 (acletus only p<0.05, r =0.45).
Open circles are acletus males, filled circles are vicinus males.
b. Total power as a function of soil moisture (percent of dry weight of
soil sample). Connected points are for sound fields by the same
male at different soil moistures. Percent soil moisture was log
transformed for regression analysis y = 6.2log(x) + 61.5 (acletus only
p<0.05, r =0.40).
8.5 9.0 9.5
Pronotal Length (mm)
5.0 10.0 15.0 20.0
= = * * * * *
Since the measurements were taken in the far field, the 70 dB isobar
represents a 0.01 mW/m2 intensity level and the total power, in mW, is given by the
intensity level times the surface area.
The acoustic power produced by calling male mole crickets ranged from 1.9 to 22.4
uW (62.8 73.5 dB re 10-12 W) for the 17 acletus sound fields and from 2.3 to 9.6
uW (63.5 69.8 dB) for the 5 vicinus sound fields (Table 1). Part of this variation in
the power output was due to the size of the male and the moisture in the soil
surrounding the male's burrow. There was a significant relationship between male
size and power level (p<0.05, r2=0.45; Fig. 5a, Note that regression was done only on
sound fields from acletus males and that the acoustic powers for individuals for
which more than one sound field was measured were averaged and used for
computing the regression). There was also a significant relationship between the
soil moisture and power level (acletus only p<0.05, r2=0.40; Fig. 5b, Percent soil
moisture was log transformed for regression analysis). The data from individual
males also showed the same trend. That is, their power output increased with
increasing soil moisture (5 out of 7 acletus and 1 of 1 vicinus). However, a check to
see if soil moisture and male size were correlated revealed no significant
relationship and large males did not produce more power simply because they
called from moister soil.
The power output of a male can be estimated by a single measurement of the
sound pressure level directly above the burrow. There was a direct relationship
between the sound pressure directly above the burrow and the power level of the
sound field produced (p<0.05, r2=0.62; Fig. 6).
Using soil temperature, the pulse rate of each cricket (wing stroke rate) was
estimated (Ulagaraj 1976). Since the power used by the cricket is directly related to
Figure 6. Total power outpuA (dB re 10-12 W) as a function of sound pressure (dB
re 2x10l- Nm ) directly above the calling burrow. Sound pressures
are measured at 25 cm from the burrow opening. The regression is
significant (p<0.05, r-=.62) y = 0.58x + 26.0.
S 0 0
0 y .58x + 26.0
I I I
55 70 75
Sound Pressure (dB)
Figure 7. Sound field of Scapteriscus acletus. Sound field is base on an average of
17 radial distances at each of 65 radii. Standard errors around the 65
points are always less than 4 cm. Anterior is designated by ant, posterior
by pst, right by rit and left by Ift.
a. View from above the sound field.
b. Looking from the crickets' left.
c. A perspective view.
.p si. . . . ...
the wing stroke rate while calling (Prestwich and Walker 1981), I estimated the
percent efficiency of sound production for each sound field. Subtracting an
estimated resting metabolism jf a)out 370 uliter O2/g h (Prestwich unpublished,
personal communication 1986) from the metabolic rates while calling revealed that
the mole crickets were very inefficient. Percent efficiency ranged from 0.02 to 0.16
percent (Table 1). The data for estimated power used while calling are similar to
actual measurements of calling metabolism of male acletus (Prestwich, personal
Because the individual sound fields were variable, an average sound field
based on the average radial distances along each of the 65 radii was computed for
the 17 sound fields from acletus males. The standard errors of the 65 means were
about 2.5 and always less than 4 cm. The shape of the sound field most closely
resembled a hemisphere and the mean distances did not differ significantly from the
radius of a 26 cm hemisphere. At ground level (X-Y plane, Fig. 7) the transverse
axis of the sound field was 48.4 cm and the longitudinal axis was 53.7 cm across.
Height of the sound field was 26.6 cm. Unlike the sound fields of other crickets, the
burrow opening acts as a point source and radiates equally in all directions. The
total power of the averaged sound field was 4.33 uW or 66.4 dB Using 26 cm as the
radius for a hemisperical sound field, the power should be 4.25 uW or 66.3 dB.
The acoustic power produced by male Scapteriscus is less than one-
hundredth of the power produced by the French mole cricket, Gryllotalpa vineae.
Bennet-Clark (1970) calculated a mean power output of 1.2 mW. Assuming that
this was produced by a total muscle power of 3.5 mW, Bennet-Clark calculated an
efficiency of 28-35 percent. However, the power used was probably underestimated
since he estimated power used from data on grasshopper flight muscles. Using data
from Prestwich and Walker (1981) Gryllotalpa vineac should use about 70 mW of
power for an average 3.3 g cricket calling with a pulse rate of 66 pulses/sec. The
efficiency then becomes 1.7 percent, a little less efficient than a typical loudspeaker.
This is still more than ten times more efficient than Scapteriscus. Since female
Scapteriscus preferentially land at louder males (Forrest 1980, 1983), males are
expected to call at their maximum output, but because Scapteriscus are one-sixth
to one-third the size of Gryllotalpa it seems likely that they are unable to produce
as much acoustical power because of their much smaller muscle mass. Note that the
power produced by Scapteriscus acletus was directly related to the size of the male
(Fig. 5a). Soil moisture also influenced output probably because it enables the
animal to pack the sandy soil better, and thus, it absorbs less of the acoustic energy.
Bennet-Clark (1970) also noted that moist soil increased output in Gryllotalpa
The efficiency of sound production is somewhat lower in Scapteriscus spp.
than that reported for other animals that use acoustic signals to communicate.
Hylid frogs are about 1 to 3 percent efficient (Prestwich and Bruger, unpublished).
The frogs are probably more efficient because the calls are produced by vibration of
a vocal sac that acts much like a pulsating sphere. Such a source does not have the
acoustic short circuit a dipole source has, and if the vocal sacs of the frogs are the
proper dimensions relative to the wavelength of their calling songs they may have
resonant properties that would also increase the efficiency. Male bladder cicadas,
Cystosoma saundersii, produce about 0.35 mW of acoustic power and are about 0.82
percent efficient. Like the frogs, the abdomen of the cicada acts as a baffle and
resonating system (MacNally and Young 1981). Other crickets have only slightly
higher calling efficiencies than those of Scaptcriscus. Anurogryllus arboreus are
about 0.2 percent efficient and Oecanthus celerinictus are from 0.3 to 1.3 percent
efficient (Prcstwich unpublished).
The sound fields of individual males were often irregular although they were
similar in overall shape to the average distribution pattern (Fig. 7). Since a new
burrow is constructed each evening, the openings may be very different from cricket
to cricket and from night to night for the same cricket and this probably causes the
irregularities. The distribution pattern of sound from a mole cricket burrow differs
considerably from the directional patterns of other crickets. The burrow acts as an
infinite baffle around the vibrating membranes raised in the throat of the horn
(Bennet-Clark 1970, Nickerson et al. 1979). The burrow opening then produces a
point source of sound and the waves radiate equally in all directions. The result is a
spherical shaped distribution pattern.
The sound fields of other crickets usually have some directional properties.
Anurogryllus arboreus (Paul and Walker 1979) and Gryllus campestris (Nocke
1971) have directional characteristics due to the destructive interference along the
edges of the wings. Probably the most noticeable sound fields for their directional
quality are those of tree crickets (Williams 1945). The sound pressure may be more
than 20 dB less at the sides of the cricket compared to anterior and posterior
pressures, and males often turn while calling seemingly to direct sound in different
directions (Toms 1984, Forrest unpublished). Male Gryllotalpa vineae also produce
a directional pattern because the two burrow openings act like dual speakers, and
there is a characteristic radiation pattern dependent upon the relationship of
distance between the two openings and the wavelength of the sound produced. This
radiation pattern of Gryllotalpa vineae is such that it should increase the probability
that a female will intercept the sound fields (see Bennet-Clark 1975 and Forrest
1982 for a discussion of selection on directional sound fields).
In summary, the power output of male Scapteriscus is relatively small
compared to other species of acoustically signaling animals, and male mole crickets
are very inefficient at producing sound (Table 1). Large males produce greater
sound power than smaller males, and moist calling sites increase the total output
(Fig. 5). The radiation pattern of sound from the calling burrow is hemispherical in
shape (Fig. 7), and the total power can be estimated from a single measurement of
sound pressure above the burrow (Fig. 6).
OVIPOSITION AND MATERNAL INVESTMENT
It's only the giving that makes you what you are.
Females of some insects may lay as many as 600,000 eggs during their
lifetime although the norm is usually a few hundred (Hinton 1981). These nutrient-
filled eggs are a major investment in offspring, and parental investment, especially
the differences in investment between the sexes, is a predominant force in selection
on reproductive strategies (Trivers 1972), life history tactics, and resource allocation
(Boggs 1981). In most instances, investment by males ends at copulation or
fertilization (see Thornhill  on paternal investment in insects), and females
generally do not invest after oviposition. Unlike paternal investment, which is
frequently difficult to assess, maternal investment can often be measured simply by
determining the mass of eggs produced.
In crickets oviposition and oviposition sites are diverse and eggs are
extremely variable in size, number laid, and whether or not they are diapausing.
While a great deal of attention has been given to nuptial feeding and paternal
investment in katydids (Gwynne 1981, 1983), crickets (Sakaluk 1984), and
cockroaches (Mullins and Keil 1980), few studies have involved orthopteran egg
deposition and measurement of maternal investment in eggs. This study reports the
oviposition of southern and tawny mole crickets. I followed the reproductive output
of females caged under field conditions and studied the influence of size, season,
and aging on female investment in eggs and female fitness as measured by the
number of hatchlings produced. Egg laying in these species was earlier described by
Hayslip (1943), but his study concerned mole cricket life histories and lacked
long-term monitoring of individual females. Walker and Nation (1982) studied
fall mating, egg maturation, and sperm storage of individuals but did not examine
maternal investment in offspring.
Materials and Methods
All studies of oviposition were done in Gainesville, Florida, and involved
females that were collected after they were attracted to the conspecific male calling
song: either an artificial song (Walker 1982) or the natural song of a male calling
from a soil-filled bucket surrounded by a trapping device (Forrest 1983). Females
were placed in buckets of soil (19-liter) and were fed 10 cm3 of ground, dry dog
food each week, enough to ensure that food was not a limiting factor in egg
production. Dog food was chosen because it had proven to be an adequate food in
maintaining other crickets in the laboratory. The buckets were kept outdoors under
two conditions. 1) The buckets were on the ground and screen lids prevented the
crickets' escape. 2) Buckets were buried flush with the soil surface, surrounded by a
doughnut-shaped pitfall, and covered with a cylindrical, hardware cloth cage.
Females had to fly to leave the buckets and in so doing they hit the cage, fell, and
were trapped. Trapped females were then placed in new buckets and the old
buckets were checked for eggs. If females could not leave because of screen lids, or
if they did not fly from a bucket, their buckets were examined for oviposition and
mortality about every 21 days (13 to 24 days depending on temperature).
Examination of buckets consisted of carefully scraping thin layers of soil (0.5-
1 cm) from the bucket using a spatulalike piece of sheet metal. This allowed the egg
cells to be excavated without damaging the eggs. The depth and the number of eggs
of each egg cell were noted and the eggs were placed on moist soil in numbered,
aluminum containers. To estimate investment in each clutch, a sample of eggs was
dried (at 30-370C) and weighed to the nearest 1.0 mg using a model AC 100 Mettler
balance. Remaining eggs were kept in the aluminum containers at room
temperature and monitored daily until hatched.
Females of each species (n= 11 vicinus, 9 acletus) were collected before
oviposition during the 1982 fall flight season and placed singly in buckets with a
male (condition 1 above). During the following spring, surviving females and spring
females collected before oviposition were kept (condition 2) throughout the flight
and reproductive seasons. Because females were found to lay no eggs in the fall,
data from spring 1982 were included in analysis of reproductive season.
Oviposition Cycles and Flight
During the springs of 1982 and 1983, females were kept in buckets (condition
2) and their flights monitored to determine the relationship between oviposition
and flight activity. The duration between hatching dates of successive egg clutches
from individual females was used as an estimate of time between egg clutches.
Effects of Size and Age on Fecundity
Females varying in size were kept singly in buckets (condition 2) with a male
and monitored for oviposition throughout their reproductive lifetime. Only females
collected prior to any reproductive activities or that were known virgins were used.
The wet weight (to the nearest 1.0 mg) of females was measured only once prior to
their use in the experiment with a model AC100 Mettler balance. Pronotal length
measured to the nearest 0.1 mm with vernier calipers was used as an indication of
female size. In most cases, pronotal length was used because it has a direct
relationship to female weight (r2=0.72 aclctus, 0.66 vicinus, p < 0.001 see Chapter
III), is less variable than weight, and is easily measured in the field.
Insect eggs of many species absorb water during development, thus
increasing both volume and weight (Hinton 1981). Because of differences in water
content among eggs, dry weights were used as a measure of investment. Investment
in individual eggs (mean dry weight per egg of samples), investment in clutches
(mean dry weight of egg times number of eggs), and number of offspring produced
(percent hatch times number of eggs) were measured for successive clutches of each
Mortality. Longevity, and Seasonal Oviposition
Of 9 acletus and 11 vicinus monitored during the fall, only 3 (33%) and 4
(36%) survived the winter. Since mortality was concentrated in colder months, this
high mortality was probably due to buckets being placed above ground, increasing
their exposure to cold temperatures during the winter. There was no apparent
influence of size on mortality. No eggs were laid by any of the fall females and
when females were found while examining buckets for eggs they were motionless or
extremely sluggish. Of all females monitored during 1982 and 1983, 54% (n = 24) of
vicinus and 41% (n = 22) of acletus females died before reproducing (including one
vicinus and one acletus that were killed during examination of their buckets).
Longevity of females in buckets was highly variable and ranged from 16 to 278 days
(mean 91) for acletus and from 7 to 254 days (mean 89) for vicinus females.
Egg-laying season was similar for the two species during the springs of 1982
and 1983 (Fig. 8). Oviposition started in late April and stopped in late July or early
August, with most egg clutches produced in May and June. The proportion of
Figure 8. Oviposition season in mole crickets (a) S aclctus and (b) S. vicinus.
Graphs show the monthly proportion of egg clutches laid. For each
month, bar on the left represents data for spring 1982 and the bar on
right data for spring 1983. Note greater proportion of unsuccessful
clutches (0-50% hatch) late in the season.
S> 50% hatch
APR MAY JUN JUL AUG
Table 2. Correlation matrix of paired comparisons for S. acletus (top half of table)
and S. vicinus (bottom half, underlined).
Clutch % Hatch Egg No. of eggs
no. wt per clutch
Clutch -0.40 -0.72 -0.32
% Hatch -0.61 0.38 0.32
wt -0.66 0.49 0.30
No. of eggs
per clutch -0.61 0.65 0.57
a Spearman's rank correlation (rho). All correlations are significant (p < 0.01).
unsuccessful clutches (<50% hatch) increased as the reproductive season progressed
(Fig. 8). This was probably the effect of decreased hatching in later clutches of
females (Table 2) rather than an influence by season alone.
Oviposition Cycles and Flight
The average interval between clutches, as estimated by the hatching dates of
successive clutches, was 10 days for acletus (n = 38, 95% CL 8-11 days) and 9 days
for vicinus (n = 32, 95% CL 7-12 days).
Females rarely flew between egg clutches. In only 1 of 68 instances did a
vicious female (n = 9) take flight between the production of egg clutches. Of all
females producing eggs, seven vicinus never left buckets, three flew prior to egg
production but not after, and one flew after egg production but not before. Female
acletus (n = 13) were more likely to take flight between clutches (13 of 43
occasions). For acletus females producing eggs, six never flew from their buckets,
two flew after they began laying eggs, and five flew both before and after they
Reproduction and Investment by Individuals
Females produced variable numbers of eggs and clutches; some died before
ever reproducing, while others produced more than 450 eggs and as many as 10
clutches. Part of the variation in fecundity was due to size differences among the
females. Excluding females that produced fewer than three clutches, there was a
significant positive relationship between pronotal length of acletus females and the
total number of offspring produced (sum of the number of eggs per clutch times
percent hatch of the clutch, Fig. 9, p < 0.05) and between pronotal length and
average number of eggs per clutch (Fig. 10, p < 0.01). These relationships were not
significant for vicinus females. There was no relationship between female size and
the number of clutches or the size of eggs produced for either species.
Figure 9. Relationship of size and total number of offspring produced for 12
S. acletus females eaged outdoors. Line is calculated using the
regression equation (r = 0.33, p < 0.05).
y 4.5(x) 264.3
7.0 7.4 7.8 8.2 6 9.0
Pronotal Length (mm)
Figure 10. Mean number of eggs per clutch for 12 S. acletus females varying in
size caged outdoors. Line is calculated using the regression
equation (r = 0.50, p <0.01).
7.0 7.4 7.8 8.2
Pronotal Length (mm)
y 7.9 W -24.6
Figure 11. Plot of ranked egg weight for successive clutches of female mole
crickets. Egg weight is the mean dry weight per egg of a sample
from each clutch. Clutches with the same mean weight were given
averaged ranks. Points connected by lines represent points at the
same position on the graph. The number of females producing a
specific number of clutches can be found by counting the points for
each clutch number. Note that females invest less (smaller weight =
higher rank) in later clutches (higher clutch numbers). (a) S. acletus,
1 2 3 4 5 6
S S vicinus
1 2 3 4 5 6 7 8 9 10
7 8 9
Average investment per clutch was from 10 to 24 percent of female body
weight for aclctus and ranged from 5 to 16 percent of female body for vicinus
females. Total lifetime investment was greater than 100 percent of female body
weight for 6 of 12 acletus females and for 1 of 8 vicinus females. Since dry weight of
eggs was measured and female body weight was measured while the crickets were
alive (i.e., they do not represent dry weight of females), the investments in eggs are
actually substantially larger.
In both species, investment per egg and per clutch decreased as females
produced successive clutches. There was a significant inverse correlation between
weight of eggs and clutch number (Fig. 11) as well as an inverse correlation between
the number of eggs per clutch and clutch number (Table 2). Also, the percent hatch
for clutches and mass per egg were significantly and negatively correlated with
clutch number (Table 2). Later clutches of a female had fewer eggs, smaller eggs,
and lower percent hatch.
My observations on oviposition did not differ from Hayslip's (1943) Plant
City, Florida, observations with regard to depth and average number of eggs found
for egg cells of acletus or vicinus. Oviposition season was also similar for vicinus;
however, acletus in Plant City extended oviposition into September. This may be
the result of warmer temperatures and a longer growing season farther south (ca.
190 km), conditions that result in a bivoltine life cycle for acletus in South Florida
(Walker et al. 1983). I did not find fall oviposition in buckets by females of either
species, but flush samples taken during November in Gainesville have contained
hatchling acletus suggesting that fall oviposition may occur, although young
juveniles apparently do not survive the winter (Stackhouse W., E. L. Matheny and
W. G. Hudson unpublished data).
Even though oviposition seasons of acletus and vicinus are similar, flight
seasons are not. Most of the flight season of vicinus is over by the start of
oviposition, while acletus flights are concurrent with oviposition (compare Fig. 2 of
Walker et al.  with Fig. 8). This agrees with differences in flight from buckets
in the two species. That acletus females are more likely to fly between clutches may
have been one factor contributing to the more rapid spread of this species after its
introduction into the United States (Walker and Nickle 1981).
Clutch cycles run between 7 and 12 days as evidenced by the average time
between hatching of successive egg clutches of females. This agrees with earlier
work that estimated a 9 day period for a female to mature an egg clutch based upon
the measurement of primary oocytes of females (Forrest 1981). That flying females
have small oocytes suggests that they recently matured or laid a clutch and are
starting a new clutch cycle (Forrest 1981). Ngo and Beck (1982), working on
dispersal flights of acletus, noticed a significant 10-day peak in recaptures of
marked and released females. Oviposition cycles may have accounted for this peak.
They also found 12-day cycles in overall flight activity and attributed it to female
oviposition cycles. This assumes that females are synchronous in their egg laying. I
found no such synchrony. Because acletus females in my tests flew between only
30% of the egg clutches produced, it seems unlikely that peaks in flight activity are
due to oviposition cycles. Such peaks are more likely caused by environmental
factors making flights favorable or synchronizing maturation.
Larger eggs generally produce higher quality, more vigorous progeny that
may be more mobile, may suffer less mortality during early stages, may develop
faster, or may require fewer instars (Capinera 1979). But given finite resources,
producing larger eggs may not always increase the fitness of the female and a trade-
off should exist between number of eggs produced and their size (Smith and
Frctwell 1974, Parker and Begon 1986). Producing small eggs is apparently
disadvantageous for mole cricket females; because egg size and hatching success
were positively correlated in both species. Producing fewer eggs may also hamper
escape from the egg cell if siblings must cooperate while digging to the surface.
Large clutches of large eggs then should maximize a female's fitness. However, as
females aged they produced smaller clutches with smaller eggs. If clutch size is
important and resources are limited as females get older, they may divert some of
their resources away from egg size, at the cost of hatching success, to increase the
clutch size. That later clutches are smaller with smaller eggs could have also been
the effect of longer lifespans than would normally occur under natural conditions.
Organisms rarely survive their full potential life span and theories on the evolution
of senescence suggest that selection should act to increase reproductive output early
in life at its expense later (Hamilton 1966). Females kept in buckets were protected
from hazards, and, thus, their becoming senescent may have influenced the size of
their later clutches. Another orthopteroid, the stick insect Diapheromera vclii,
shows a similar decline in the rate of egg production and a decrease in investment in
eggs later in life (Sivinski 1977).
Gwynne (1983) related sexual differences in mating behaviors to differences
in parental investment in Tettigoniidae and other Orthoptcra. As sexual selection
theory predicted (Trivers 1972), when males invest more they are more selective of
mates than females, whereas females are more selective and males more
competitive when maternal investment is greater than paternal investment.
Investment in progeny by female mole crickets is far greater than that for males.
Males provide no paternal investment except the possible energy a female receives
from consuming the small spermatophore (<1 mm diam, estimated 0.1% of male
body weight) a male passes to the female during mating. On the other hand, a
female's average investment per clutch may be as much as 24% of her body weight.
Behavioral differences between the sexes support predictions based on differences
in parental investment. Male mole crickets display acoustically, are aggressive, and
often fight near calling burrows, whereas females are discriminating in their choice
of mates (Forrest 1983).
In a model of optimum egg size and clutch size in insects, Parker and Begon
(1986) showed that if larval success was determined solely by the size of the egg,
then selection would act such that females all produced eggs that were the same
size. A consequence of this is that large females can produce larger clutches than
smaller females. Data from mole crickets fit the model. There was a positive
correlation between egg size (investment) and hatching success of clutches for mole
crickets. Mean clutch-specific egg size did not differ between large and small
females nor did the rate of decline of investment in eggs or egg clutches. Also,
larger females produced larger clutches.
Although a direct influence of size on fecundity is common in insects, few
studies have been conducted under natural conditions and few data are available for
Orthoptcra. I found a significant relationship between female size and mean clutch
size (Fig. 10) and total offspring produced (Fig. 9) for acletus but not vicinus
females. That only one of eight vicinus females invested more than her body weight
in lifetime egg output compared with six of twelve acletus females suggests that this
difference may be due to the inability of vicinus, primarily a herbivore, to assimilate
the dog food provided as food. Female vicinus would then have to rely on juvenile
reserves for egg production, and although a larger female may carry greater reserves
these would probably not be enough to produce more than a few clutches. Dog
food, however, was able to keep the females alive for an average of 90 days and in
some cases for more than 250 days, indicating that this diet was reasonably adequate
for the crickets. Female vicinus might simply consume more of the poor quality
food, and, thus, expend more energy to compensate for its lack of nutritional value
(see Slansky  for a review of insect nutritional ecology).
Large acletus females can produce almost 3 times as many offspring and
more than 1.5 times as many eggs per clutch as small females. Gwynne (1984)
found that female size and nuptial feeding by males directly affected fecundity in the
bushcricket, Requena verticalis. Females feeding on seven spermatophylaxes more
than doubled egg production, and the size of eggs produced also increased. There is
also a relationship between the number of ovariolcs (and presumably reproductive
output) and size in betwecn-species comparisons of melanopline grasshoppers
(Bellinger and Pienkowski 1985).
Female investment and fecundity and their relationship with size tactics and
developmental strategies in insects have received almost no attention, though such
selective forces shape population size structure and influence population dynamics.
Assuming a direct relationship between fecundity and size, Masaki (1967) explained
the latitudinal size variation in the Emma field cricket, Telcogryllus emma, on the
basis of a balance between the opposing selection pressures of increasing fecundity
with increasing size and decreasing size with shortened growing season of more
northern latitudes. I have used similar explanations for within-population size
variation and changes in the seasonal distributions of sizes of both male and female
mole crickets (see Chapter III). These types of data should improve models of
population dynamics and help explain the life-history strategies in insects from an
individual, evolutionary perspective.
SIZE TACTICS AND DEVELOPMENTAL STRATEGIES
From early days of infancy,
through trembling years of youth
Long murky middle age
and final hours long in the tooth.
Size is one of the most important attributes of any organism, and within any
species there is often a great deal of variation in size among individuals. An
animal's size will profoundly affect the prey it can cat, what predators it has, its
physiology and energetic, and its reproductive success (Peters 1983). Thus an
organism's size and life history are intimately related. However, research on life
history strategies has virtually ignored the relationship between size and
development, and has focused primarily on tactics of diapause, migration and
dispersal (Blakley 1981; Denno and Dingle 1981).
Animals with complex life cycles are interesting with regard to size tactics
because certain life stages are specifically evolved for growth and increasing body
size, while other stages are adapted for dispersal and/or reproduction. Amphibians
and insects are perhaps the best known groups with complex life cycles. Of the two,
insects offer a simpler system in which to study size tactics because growth occurs
only in juvenile and non-reproductive stages (see Wilbur and Collins 1973; Smith-
Gill and Berven 1979; Werner 1986 for a discussion of amphibian metamorphosis).
Unlike amphibians, there is usually no growth once an insect becomes adult (i.e., no
increase in size although abdominal stretching may occur especially in females), and
adult size is determined by what happens during juvenile stages. Individuals can
make decisions about their size by using certain cues to determine when to molt to
Costs and Benefits
Regarding decisions, I am not implying that the animals make a conscious
decision but rather these are evolved responses to cues the individual has available.
The decisions an individual makes about changes in size carry with them costs and
benefits. Although difficult to assess, these costs and benefits are ultimately
measured in terms of fitness. Accordingly, selection should act such that individuals
evaluate the costs and benefits and make those decisions that will maximize fitness.
For insects the decision an individual makes about its size can be simplified to a
decision of whether or not to become an adult. Figure 12 shows some of the costs
and benefits associated with a yes or no decision to mature, that is to molt to pupa
or adult. For instance, at a point in the development of an individual the decision
to continue as a juvenile (no-decision Fig. 12) has associated with it costs due to
delayed reproduction with the risk of dying before maturing and reproducing.
There is also the cost of limited mobility since juveniles cannot fly. Benefits to an
individual remaining a juvenile are due to an increase in fitness because of a longer
period of growth and thus a further increase in size, assuming there is a direct effect
of adult size on fitness (see below).
Graphically the cost/benefit curves might look like Figure 13. Early in its
development an individual might benefit a great deal because of small increases in
size while the costs of the same size change increase only slightly. Later on small
changes in size increase fitness (benefits) very little while costs for the same amount
of change become increasingly more expensive. The individual is expected to adjust
its development such that it maximizes the difference between benefits and costs
Figure 12. Costs and benefits to an individual associated with yes or no
decisions of whether or not to become an adult.
BENEFITS attain larger size
COSTS delayed reproduction
Figure 13. Hypothetical cost/benefit curves for changes in an individual's size.
Selection is expected to act such that individuals weigh costs and
benefits and become adult when the difference between the curves is
maximized (dashed line).
(i.e. the individual is expected to make the decision to molt to adult at the size
indicated by the vertical line). Cues such as population density, availability and
suitability of food and mates, size of competitors, sex ratio, length of the remaining
growing season, and "expected" temperature (particularly important for
poikilotherms) should be used in making the decision of when to molt.
Although some authors have focused on an optimal body size within a
species (sce Roff 1981), it is the variation in sizes among individuals and the
deviation from the optimal that needs to be investigated. Previous work on size
variation in insects dealt with variation between populations. Primarily latitudinal or
altitudinal differences among the populations were used to explain the variation
(Masaki 1967, 1978; Dearn 1977), and Roff (1980,1983) has proposed models that
use season length, = generation length, which varies with latitude and altitude, as a
tool in predicting insect size and life histories. Proximate causes of size variation in
insect populations include temperature gradients (Sweeney and Vannote 1978),
larval diet (Slansky 1982; Andersen and Nilssen 1983; Palmer 1984) and population
density (Peters and Barbosa 1977). Borgia (1979) discussed variation in male size
and the decisions males should make in weighing costs and benefits of changes in
size. Borgia was primarily interested in male size and its importance in a male's
ability to control specific resources, but such tactics and decisions should be
applicable whenever size influences an individual's fitness.
Size and fitness in insects
For poikilotherms, reproductive success or output is almost always
influenced by size (Peters 1983). It is well known that for many insects there are
positive relationships between female size and the number of eggs, size of eggs, and
size of clutches produced (Hinton 1981). Only recently have there been attempts to
correlate male reproductive success with size. In species where males compete for
females (female defense polygyny) or for resources used by females (resource
defense polygyny) large males often have a reproductive advantage over smaller
males that may have to adopt alternative mating strategies to acquire mates. Where
female choice is involved, females often choose males on the basis of cues
correlated with male size (Thornhill and Alcock 1983 and refs.) Thus it is well
documented among insects that large size confers some advantage in fitness for
both sexes. For this reason insects are appropriate for investigating the occurrences
of size tactics and developmental strategies.
Size Tactics in Mole Crickets
Costs and benefits associated with size are almost impossible to measure
directly since they may differ from one individual to the next and may change at any
time during an individual's life. However, developmental strategies should be
reflected in adult sizes, and certain predictions about changes in size distribution for
a species can often be tested. To examine whether individuals weigh costs and
benefits associated with changes in size I tested predictions about seasonal
differences and within-season changes in size distributions for two species of mole
Size is an important component in the sexual success of an individual. For
males, the intensity of the calling song is correlated with male size--larger males
produce louder calls (Forrest 1980,1983 and Chapter I). Flying females selectively
respond to and land at louder calls, and all things being equal, larger males will
attract and mate with more females than smaller males (Forrest 1983). Larger
females also have a reproductive advantage over smaller females. Large females
may produce as many as 3 times as many offspring and more than 1.5 times as many
eggs per clutch than smaller females (see Chapter II).
Since reproduction occurs only during a brief period the following spring,
during the fall the cost to an individual due to delayed reproduction associated with
remaining a juvenile is zero (see Fig. 12). One would predict that individuals that
are smaller than average would overwinter as juveniles to increase their size, while
those that are larger than average would gain relatively little by increasing in size,
but more by becoming adult, dispersing to favorable overwintering sites, and
preparing for reproduction early the following spring. There may be a cost of
delayed maturation due to limited movement. For instance, if an individual found
itself in an unfavorable site where the likelihood of surviving the winter is very low
(i.e., the costs are very high), that individual should mature and disperse to a more
favorable site. However, this cost would be borne by large individuals as well as
small, while the additional cost of maturing at a small size is borne only by small
individuals. Therefore I expected a greater proportion of larger than average
individuals in fall flights compared to spring flights.
The second prediction involves an expected change in the size distribution
during the spring reproductive season. Because the egg laying (reproductive)
season is brief, individuals that remain juveniles incur greater and greater costs as
the season progresses. Because the chance of surviving to the following
reproductive season (the next spring) is very small, the costs of delaying
reproduction increase dramatically and soon outweigh the benefits of increasing in
size. I predicted that as the spring season progressed smaller and smaller
individuals should mature.
Materials and Methods
Quarter-monthly samples (up to 50 individuals for 1981, 25 for 1982) of adult
male and female mole crickets, acletus and vicinus, were obtained from a routine
sound trapping station (Walker 1982) on the Agronomy Farm of the University of
Florida. Samples were taken during the spring and fall flight seasons of both years.
Since fall adults are of the same generation as adults of the following spring,
measurements spanned three generations, and samples from spring 1981 and fall
1982 represented only part of the 80-81 and 82-83 generations respectively.
To prevent desiccation, adults were placed in 13 dram snap cap vials
containing moist soil but no food. Measurements of pronotal length (to nearest
0.1mm measured along the midline using vernier calipers) and mass (to nearest 1.0
mg using a model AC100 Mettler balance) were taken for each individual within 24
hr of capture.
There was a significant positive relationship between mass and pronotal length (Fig.
14). A linear model is shown because this model explained as much of the variation
in the data as did the more usual model of mass equals the cube of a linear
dimension. The regressions differed between species, whereas within a species the
sexes had similar slopes that did not differ significantly. In both species the
intercept was smaller for males than females (significant for acletus but not vicinus),
thus for a given pronotal length males weighed less even though males on average
are the larger sex (Fig. 14).
Figure 15 shows the size distributions for each sex during spring and fall
seasons (combined for both years). Distributions were approximately normal and
male and female sizes were broadly overlapping. On average males were larger
than females (Fig. 15, Table 3). For acletus the distributions changed seasonally.
Fall adults were larger than spring adults (Fig. 15a & b). The distributions for
vicinus females suggest the opposite trend (though not significant Table 3), but the
exaggeration of the fall proportions is the result of a small sample of fall vicinus
Figure 14. Linear relationship between pronotal length and mass for male and
female S. acletus (circles) and vicinus (squares). Lines are plotted
using regression equations and the endpoints represent the extreme
values of pronotal length for each group. All regressions are
significant p<.001. N=1222 male and 1431 female acletus; 284 male
and 714 female vicinus.
* acletus d y=168x -848 r2=.85
O acletus9 y=163x -760 r2=.72
* vicinus dr y238x -1147 r2=.78
O vicinus 9 y=249x -1173 r2= .66
6.0 7.0 8.0 9.0 10.0 11.0 12.
Pronotal Length (mm)
Table 3. Mean pronotal length (mm), standard deviation, and number of adult
mole crickets trapped during spring and fall flight of 1981 and 1982.
SPECIES SPRING FALL
sex 1981 1982 1981 1982
males 8.8, 7a 8.8, .7a 9.5, .6b 9.3, .5b
(n=594) (369) (32) (227)
females 7.9, .5a 7.9, .5a 8.4, .4c 8.2, .4b
(709) (371) (102) (249)
males 8.4, .5b 8.2, .5a ---- 8.3, .4ab
(162) (119) (0) (3)
females 7.9, .4b 7.8, .4a 8.0, .6ab 7.8, .4ab
(379) (273) (3) (59)
aMeans in the same row followed by different letters are significantly different
Figure 15. Frequency distributions (expressed as a proportion of season total)
of adult mole crickets captured during 1981 and 1982 flights at
S aletus s0
or Srn n=l60a
n 'all n.35;
~.miIIflmfl LA Eli ______________
65 70 75 80 85 90 95
Pronotal Length (mm)
100 105 110 115
S ademus d
. sano n=259
65 70 75 80 85 9.0 95 100 105 110 115
Pronotal Length [mm)
| B* sg n-652
0 fa n=62
65 70 75 80 85 90 95 100 105 110 115
Pronotal Length (mm)
S -cmnus dlf
* sprng n-281
65 70 75 80 85 90 95 100 105 110 115
Pronolal Length [mm)
(n=62). Only three vicinus males were captured during the two fall seasons and
were not included in the figure.
Mean pronotal lengths were significantly larger for fall aclctus than for
spring individuals, whereas vicinus did not differ in average pronotal lengths
between the two seasons. -Pronotal lengths were similar for seasons compared
between years (Table 3). Note also that fall individuals typically showed less
variation in size than those in the spring.
Mean pronotal lengths of quarter-monthly samples decreased during the
spring flight season and reached a minimum toward the end of the reproductive
season (Fig. 16). During the spring there was a significant relationship between
pronotal length and the week of the season, but this relationship was not significant
for fall individuals. This relationship explained less than 10 percent of the variation
in the data when individual measurements were used in the regression analysis (Fig.
16). A regression using sample means (for samples with more than 10 individuals)
and date of sample explained 37-89% of the variation in the mean pronotal length
(Fig. 16). Data for vicinus males are not shown because few samples contained
more than 10 males (1981 n=4, 1982 n=5). The relationship was significant for
individual measurements of male vicinus for spring of 1982 (p<.05) and both years
combined (p=.05); however no significant relationships were found between the
nine sample means and date of capture.
The data on seasonal and distributional shifts in mole cricket sizes support
the predictions and suggest that individuals weigh costs and benefits associated with
changes in size (Fig. 12). There was one major difference between the two species
that may indicate different overwintering strategies. There was a significant
difference between fall and spring adult sizes for aclctus but not vicinus. According
Figure 16. Plot of mean pronotal length of quarter-monthly samples (where
N>10) for adult mole crickets captured during 1981 and 1982 flights.
For all means shown standard errors are about 0.1mm. Trend lines
plotted from regression equations for individual measurements for
1981 and 1982 combined (solid) or separately in the case of vicinus
(dashed). Spring declines in sample means were significant--
A) 1981:r2=.38, p<.03. 1982: .89, <.001. 1981+82: .70, <.001.
B) 1981:r2=.89, p<.001. 1982: .72, <.001. 1981+82: .79, <.001.
C) 1981:r2=.45, p<.03. 1982: .62, <.004. 1981+82- NS.
A A A
A A sA A A
Sep Oct I Nov I Dec 1 Feb Mar Apr I May Jun Jul
A A A a a
A A A-A
A A a
Sep I Oct I Nov Dec Feb I Mar Apr I May I Jun Jul
- *-- -
Sep I Oct I Nov I Dec 1 1 Feb I Mar I Apr I May I Jun I Jul
to the predictions made, individuals that are smaller than average in the fall should
wait until the following spring to mature. That adult vicinus arc less variable in size
than acletus (Fig. 15) may have made the detection of seasonal differences more
difficult. It is interesting that a greater percentage (85%) of the vicinus population
overwinter as adults than in the acletus population (25%). Forage quantity and
quality are greatly reduced during the winter months and, since vicinus is mainly
herbivorous, this may limit the potential for growth during that part of the year.
Benefits due to overwintering as a juvenile would be lessened and it may be more
important to locate overwintering sites that are favorable for survival and allow
early reproduction the following spring.
I found that as the reproductive season progressed there was a decline in the
pronotal lengths of individuals for successive samples. That is, a significant
relationship was found between pronotal length and capture date. Although the
model did not explain much of the variation in measurements of individuals the
overall trend in the data (sample means Fig. 16) was highly significant. This might
be expected since all sizes of individuals should be maturing throughout the season,
but the proportion of small individuals increases dramatically toward the end of the
season. Small individuals have probably found themselves in areas where growing
conditions are poor and they mature at the smaller size in order to makes the 'best
of a bad situation' (Dawkins 1980).
Rogers (1942), working with a number of tipulid species, noticed a decline
(as much as 20%) in the size of late season specimens and attributed the decline to
lack of food and moisture. Similar seasonal changes were also found in the
milkweed leaf beetle, Labidomera clivicollis (Palmer 1984). There was a significant
gradual decline in male and female sizes during the growing season caused by a
decline in the amount of available food. Food quality was not discussed and may be
even more important. Slansky (1982) pointed to the growing interest in insect
nutritional ecology and discussed the decisions individuals must make relative to
nutrition and their final adult size.
For mole crickets, the similarity from year to year in the time at which the
smallest sample means occur suggests that photoperiod may be one proximate cue
used in the molting decision. However, cold weather during the spring of 1981
caused a delay in the vicinus flights (Walker et al. 1983) and probably delayed
maturation and caused the shift seen in the seasonal decline in size for 1981
(dashed lines Fig. 16c). Since acletus flights occur later in the spring, cold weather
had no affect on delaying maturation or the decline in size during the spring
months. Other proximate causes for the observed decline, such as food and
temperature, seem unlikely. Food quantity and quality as well as temperature are
increasing during the months of May, June, and July.
Insect size variation, though environmentally induced, may still be under the
influence of natural selection. The mechanisms can be simple. It is well known that
in many insects different numbers of molts precede adulthood. For instance, mole
crickets may have from 6 to 10 molts before becoming an adult. Molting is brought
about by changes in hormone liters of the individual. These hormone levels, while
under genetic control, are known to be influenced by many environmental cues.
Therefore even though variation in size can be caused environmentally, tle
decisions to molt have a genetic basis that can be acted on by natural selection.
One pervasive point in most studies of development is that an individual
must reach a 'critical size' before becoming adult (Blakley and Goodner 1978,
Blakley 1981, Palmer 1984). Starvation of juveniles before they attain this size
results in death. On the other hand, some cockroaches apparently determine their
ultimate size at a specific instar and adjust the number of instars to reach this size at
adulthood (Tanaka 1981). Slansky (1982) suggested that this minimum size is
adjusted by selection and the balance between costs of prolonged development and
the costs of reduced size and fitness.
The understanding of size variation as it relates to developmental strategies
should broaden our understanding of an organism's life history. An individual's size
is the direct result of its development. Size has been implicated as the factor
affecting a number of alternative reproductive behaviors in insects (Thornhill and
Alcock 1983) and is a major determinant in the sex change of sequential
hermaphrodites (Charnov 1979). Size tactics may even be one generation removed.
For instance females that provision broods and female parasites may adjust the
amount of investment in their offspring, thus influencing their offspring's adult size
and reproductive behaviors (Alcock 1979). Once size tactics and the developmental
strategies of an organism are explained from an individual's standpoint, new light
will be shed onto other aspects of that organism's life history.
Male mole crickets produce sound. Like most crickets these acoustic signals
function to attract females for mating purposes. Even though males use their
burrows as a baffle and they call from within an acoustic horn, the production of
sound is an inefficient process. Males are less than 0.2 percent efficient in
producing from 2 to 22 uWatts of acoustic power. The power produced is
dependent on male size and the moisture of the soil surrounding the calling burrow.
The sound field is hemispherical, and thus, has no directional characteristics. By
preferentially landing at louder males, females may use the calling songs to locate
large mates or moist soil for oviposition.
Oviposition occurs during the spring, primarily in May and June. The
investment in eggs, the numbers of eggs, and the percent hatch of eggs decline as
females age. Large females produce more hatchlings and have larger egg clutches
than smaller females.
Because size is such an important aspect of the reproductive success of both
sexes, males and females should make decisions about when to mature that will
maximize fitness. Each decision about whether or not to mature has certain costs
and benefits that must be evaluated. During the fall, individuals that are smaller
than average overwinter as juveniles in order to increase in size before spring
reproduction. As the reproductive season progresses the cost of delaying
maturation increases and smaller individuals mature.
It is important to note that the observations are made from the standpoint of
the individual. As such, they can be used to predict aspects of mole cricket life
history and will be useful in the models concerning the population dynamics of these
Alcock, J. 1979. The evolution of intraspecific diversity in male reproductive
strategies in some bees and wasps. Pages 381-402 in Blum, M.S. and N.A.
Blum, eds. Sexual selection and reproductive competition in insects.
Academic Press, New York.
Alexander, R.D. 1961. Aggressiveness, territoriality, and sexual behavior in field
crickets (Orthoptera: Gryllidae). Behavior 17:130-221.
Andersen, J. and A.C. Nilssen. 1983. Intrapopulation size variation of free-living
and tree-boring Coleoptera. Can. Entomol. 115:1453-1464.
Bellinger, R.G. and R.L. Pienkowski. 1985. Interspecific variation in ovariole
number in Melanopline grasshoppers (Orthoptera: Acrididae) Ann.
Entomol. Soc. Am. 78:127-130.
Bcnnet-Clark, H.C. 1970. The mechanism and efficiency of sound production in
mole crickets. J. Exp. Biol. 52:619-652.
Bennet-Clark, H.C. 1971. Acoustics of insect song. Nature 234:255-259.
Bennet-Clark, H.C. 1975. Sound production in insects. Sci. Prog. (Oxford) 62:263-
Blakley, N. 1981. Life history significance of size-triggered metamorphosis in
milkweed bugs (Oncopeltus). Ecology 62:57-64.
Blakley, N. and S.R. Goodner. 1978. Size-dependent timing of metamorphosis in
milkweed bugs (Oncopeltus) and its life history implications. Biol. Bull.
Boggs, C.L. 1981. Nutritional and life-history determinants of resource allocation in
holometabolous insects. Am. Nat. 117:692-709.
Borgia, G. 1979. Sexual selection and the evolution of mating systems. Pages 19-80
in Blum, M.S. and N.A. Blum, eds. Sexual selection and reproductive
competition in insects. Academic Press, New York.
Bruel, P.V. 1983. Sound level meters--the Atlantic divide. Brucl & Kjaer Technical
Capinera, J.L. 1979. Qualitative variation in plants and insects: effect of propagule
size on ecological plasticity. Am. Nat. 114:350-361.
Charnov, E.L. 1979. Natural selection and sex change in pandalid shrimp: test of a
life-history theory. Am. Nat. 113:715-734.
Dawkins, R. 1980. Good strategy or evolutionary stable strategy? Pages 331-367 in
Barlow, G.W. and J. Silverberg, eds. Sociobiology: beyond nature/nurture?
Westview Press, Boulder, Colo.
Dearn, J.M. 1977. Variable life history characteristics along an altitudinal gradient
in three species of Australian grasshopper. Oecologia 28:67-85.
Denno, R.F. and H. Dingle, eds. 1981. Insect life history patterns: habitat and
geographical variation. Springer-Verlag, New York.
Elsner, N. 1983. Insect stridulation and its neurophysiological basis. Pages 69-92 in
Lewis, B. ed. Bioacoustics a comparative approach. Academic, London.
Elsner, N. and A.V. Popov. 1978. Neuroethology of acoustic communication. Adv.
Insect Physiol. 13:229-355.
Forrcst, T.G. 1980. Phonotaxis in mole crickets: its reproductive significance. Fla.
Forrest, T.G. 1981. Acoustic behavior, phonotaxis, and mate choice in two species
of mole crickets (Gryllotalpidac: Scapteriscus). M.S. thesis, Univ. of Florida,
Forrest, T.G. 1982. Acoustic communication and baffling behaviors of crickets. Fla.
Forrest, T.G. 1983. Calling songs and mate choice in mole crickets. Pages 185-204 in
Gwynne, D.T. and G. K. Morris, eds. Orthopteran mating systems: sexual
competition in a diverse group of insects. Westview Press, Boulder, Colo.
Forrest, T.G. (in press). Sinistrality in the southern and tawny mole crickets
(Gryllotalpidae: Scapteriscus). Fla. Entomol.
Fullard, J.H., M.B. Fenton and J.A. Simmons. 1979. Jamming bat ccholocation: the
clicks of arctiid moths. Can. J. Zool. 57:647-649.
Gade, S. 1982. Sound intensity (Part I. theory). Bruel & Kjaer Technical Review
Gwynne, D.T. 1981. Sexual difference theory: Mormon crickets show role reversal
in mate choice. Science 213:779-780.
Gwynne, D.T. 1983. Male nutritional investment and the evolution of sexual
differences in Tettigoniidae and other Orthoptera. Pages 337-366 in Gwynne,
D.T. and G. K. Morris, eds. Orthopteran mating systems: sexual competition
in a diverse group of insects. Westview Press, Boulder, Colo.
Gwynne, D.T. 1984. Courtship feeding increases female reproductive success in
bushcrickets. Nature (London) 307:361-363.
Hamilton, W.D. 1966. The moulding of senescence by natural selection. J. Theor.
Hayslip, N.C. 1943. Notes on biological studies of mole crickets at Plant City,
Florida. Fla. Entomol. 26:33-46.
Hinton, H.E. 1981. Biology of insect eggs. Pergamon, Oxford.
MacNally, R. and D. Young. 1981. Song energetic of the bladder cicada,
Cystosoma saundersii. J. Exp. Biol. 90:185-196.
Masaki, S. 1967. Geographical variation and climatic adaptation in a field cricket
(Orthoptcra: Gryllidae). Evolution 21:725-741.
Masaki, S. 1978. Climatic adaptation and species status in the lawn ground cricket.
11 body size. Occologia 35:343-356.
Masaki, S., M. Kataoka, K. Shirato and M. Nakagahara. (in press). Evolutionary
differentiation of right and left tegmina in crickets. Bull. Ital. Entomol. Soc.
Mathcny, E.L. 1981. Contrasting feeding habits of pest mole cricket species. J. Econ.
Michelsen, A. 1978. Sound reception in different environments. Pages 345-373 in
Ali, M.A. ed. Sensory ecology. Plenum, New York.
Michelsen, A. 1983. Biophysical basis of sound communication. Pages 3-38 in Lewis,
B. ed. Bioacoustics a comparative approach. Academic, London.
Michelsen, A. and H. Nocke. 1974. Biophysical aspects of sound production in
insects. Adv. Insect Physiol. 10:247-296.
Mullins, D.E. and C.B. Keil. 1980. Paternal investment of rates in cockroaches.
Nelson, M.C. and J. Fraser. 1980. Sound production in the cockroach,
Gromphadorhina portentosa: evidence for communication by hissing. Behav.
Ecol. Sociobiol. 6:305-314.
Ngo, D. and H.W. Beck. 1982. Mark-release of sound-attracted mole crickets: flight
behavior and implications for control. Fla. Entomol. 65:531-538.
Nickerson, J.C., D.E. Snydcr and C.C. Oliver. 1979. Acoustical burrows constructed
by mole crickets. Ann. Entomol. Soc. Am. 72:438-440.
Nocke, H. 1971. Biophysik der Schallerzeugung durch die Vorderfliigel der Grillen.
Z. Vcrgl. Physiol. 74:272-314.
Otte, D. 1970. A comparative study of communicative behavior in grasshoppers.
Misc. Pub. Mus. Zool. Univ. Mich. #141.
Palmer, J.O. 1984. Environmental determinants of seasonal body size variation in
the milkweed leaf beetle, Labidomcra clivicollis (Kirby) (Coleoptera:
Chrysomclidae). Ann. Entomol. Soc. Am. 77:188-192
Parker, G.A. and M. Begon. 1986. Optimal egg size and clutch size: effects of
environment and maternal phenotype. Am. Nat. 128:573-592.
Paul, R.C. and T.J. Walker. 1979. Arboreal singing in a burrowing cricket,
Anurogryllus arboreus. J. Comp. Physiol. 132:217-223.
Peters, R.H. 1983. Ecological implications of body size. Cambridge University
Peters, T.M. and P. Barbosa. 1977. Influence of population density on size,
fecundity, and development rate of insects in culture. Ann. Rev. Entomol.
Prestwich, K.N. and T.J. Walker. 1981. Energetics of singing crickets: effects of
temperature in three trilling species (Orthoptera: Gryllidae). J. Comp.
Roff, D. 1980. Optimizing development time in a seasonal environment: the "ups
and downs" of clinal variation. Oecologia 45:202-208.
Roff, D. 1981. On being the right size. Am. Nat. 118:405-422.
Roff, D. 1983. Phenological adaptation in a seasonal environment: a theoretical
perspective. Pages 253-270 in Brown, V.K. and I. Hodek, eds. Diapause and
life cycle strategies in insects. Dr. W. Junk Publishers, Boston.
Rogers, J.S. 1942. The crane flies (Tipulidac) of the George Preserve, Michigan.
Misc. Publ. Mus. Zool. Univ. Mich. #53.
Sakaluk, S.K. 1984. Male crickets feed females to ensure complete sperm transfer.
Schuster, J.C. 1975. Comparative behavior, acoustic signals, and ecology of the new
world Passalidac (Coleoptera). Ph.D. dissertation, Univ. of Florida,
Sivinski, J. 1977. Factors affecting mating duration in the walkingstick
Diapheromera velii (Walsh) (Phasmatodea: Heteronemiidae). M.S. thesis,
Univ. of New Mexico, Albuquerque.
Slansky, F. 1982. Insect nutrition: an adaptationist's perspective. Fla. Entomol.
Smith, C.C. and S.D. Fretwell. 1974. The optimal balance between size and number
of offspring. Am. Nat. 108: 499-506.
Smith-Gill, S.J. and K.A. Berven. 1979. Predicting amphibian metamorphosis. Am.
Sweeney, B.W. and R.L. Vannote. 1978. Size variation and the distribution of
hemimetabolous aquatic insects: two thermal equilibrium hypotheses.
Tanaka, A. 1981. Regulation of body size during larval development in the German
cockroach, Blatella germanica. J. Insect Physiol. 27:587-592.
Thornhill, R. 1976. Sexual selection and paternal investment in insects. Am. Nat.
Thornhill, R. and J. Alcock. 1983. The evolution of insect mating systems. Harvard
University Press, Cambridge.
Toms, R.B. 1984. Directional calls and effects of turning behavior in crickets. J.
Entomol. Soc. Sth. Afr. 47:309-312.
Trivers, R.L. 1972. Parental investment and sexual selection. Pages 136-179 in
Cambell, B. ed. Sexual selection and the descent of man, 1871-1971. Aldine-
Ulagaraj, S.M. 1975. Mole crickets: ecology, behavior, and dispersal flight
(Orthoptera: Gryllotalpidae: Scapteriscus). Env. Entomol. 4:265-273.
Ulagaraj, S.M. 1976. Sound production in mole crickets (Orthoptera:
Gryllotalpidae: Scapteriscus). Ann. Entomol. Soc. Am. 69:299-300.
Ulagaraj, S.M. and T.J. Walker. 1973. Phonotaxis of crickets in flight: attraction of
male and female crickets to male calling songs. Science 182:1278-1279.
Ulagaraj, S.M. and T.J. Walker. 1975. Response of flying mole crickets to three
parameters of synthetic songs broadcast outdoors. Nature 253:530-532.
Walker, T.J. 1982. Sound traps for sampling mole cricket flights (Orthoptera:
Gryllotalpidae: Scapteriscus). Fla. Entomol. 65:105-110.
Walker, T.J. and D.A. Nickle. 1981. Introduction and spread of pest mole crickets:
Scapteriscus vicinus and S. acletus reexamined. Ann. Entomol. Soc. Am.
Walker, T.J. and J.L. Nation. 1982. Sperm storage in mole crickets: fall matings
fertilize spring eggs in Scapteriscus aclctus. Fla. Entomol. 65:283-285.
Walker, T.J., J.A. Reinert and D.J. Schuster. 1983. Geographical variation in flights
of the mole cricket, Scapteriscus spp. (Orthoptera: Gryllotalpidae). Ann.
Entomol. Soc. Am. 76:507-517.
Werner, E.E. 1986. Amphibian metamorphosis: growth rate, predation risk, and the
optimal size at transformation. Am. Nat. 128:319-341.
Wilbur, H.M. and J.P. Collins. 1973. Ecological aspects of amphibian
metamorphosis. Science 182:1305-1314.
Wiley, R.H. and D.G. Richards. 1978. Physical constraints on acoustic
communication in the atmosphere: implications for the evolution of animal
vocalization. Behav. Ecol. Sociobiol. 3:67-94.
Williams, M. 1945. The directional sound waves of Oecanthus nigricornis argentinus
or a violinist listens to an insect. Entomol. News 54:1-4.
Timothy G. Forrest was born Friday, the 13th of April, 1956, in St. John's,
Newfoundland, Canada. He received his Bachelor of Science degree in 1978 from
Florida Southern College where he majored in biology and mathematics. He
received his Master of Science degree in entomology from the University of Florida
during the spring of 1981. Since that time he has been working toward his
doctorate. In January of 1982 he married Susan Lynn Mashke. They became
parents on June 25, 1986 when their son, Justin Broc, was born.
Along the coast road, by the headland
the early lights of winter glow
I'll pour a cup to you....
Raise it up- say Cheerio.
I certify that I have read this study and that in my opinion it conforms to
acceptable standards of scholarly presentation and is fully adequate, in scope and
quality, as a dissertation for the degree of Doctor of Philosophy.
Thomas J. Walker, Chairman
Professor of Ehtomology and Nematology
I certify that I have read this study and that in my opinion it conforms to
acceptable standards of scholarly presentation and is fully adequate, in scope and
quality, as a dissertation for the degree of Doctor of Philosophy.
James E. Lloyd
Professor of Entomology and Nematology
I certify that I have read this study and that in my opinion it conforms to
acceptable standards of scholarly presentation and is fully adequate, in scope and
quality, as a dissertation for the degree of Doctor of Philosophy.
Martha L. Crump
Associate Professor of Zoology
This dissertation was submitted to the Graduate Faculty of the College of
Agriculture and to the Graduate School and was accepted as partial fulfillment of
the requirements for the degree of Doctor of Philosophy.
December 1986 ---
Dean, College of Agriculture
Dean, Graduate School