The University of Notre Dame

Sound Communications of Black-tailed, White-tailed, and Gunnison's Prairie Dogs

Author(s): George H. Waring
Source: The American Midland Naturalist, Vol. 83, No. 1 (Jan., 1970), pp. 167-185
Published by: The University of Notre Dame
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American Midland Naturalist

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Sound Communications of Black-tailed, White-tailed,
and Gunnison's Prairie Dogs
GEORGE H. WARING,
Department of Zoology, Colorado State University, Fort Collins 80521

ABSTRACT: The sounds of black-tailed, white-tailed, and Gunnison's

prairie dogs were studied in Colorado and Wyoming from February
1964 to June 1966. Observations, photographs, and tape recordings
were made in the field and were supplemented by data collected from
captive prairie dogs.
The sounds of black-tailed prairie dogs were named and had the
usual function as follows: ( 1 ) "repetitious barks" alert; (2)
"chuckle" alert; (3) "chatter barks" - threat; (4) "wee-oo" -
contact; (5) "snarl" - threat; (6) "growl" threat; (7) "scream" -
distress; (8) "raspy purr" - pleasure; and (9) "tooth chatter" -
threat.
The sounds of white-tailed prairie dogs were: (1) "repetitious
barks" alert; (2) "chuckle" alert; (3) "laughing barks" -
contact; (4) "snarl" - threat; (5) "growl" threat; and (6)
"scream" - distress.
The sounds of Gunnison's prairie dogs were: (1 ) "repetitious
barks" - alert; (2) "rapid barks" - apprehension; (3) "chuckle" -
alert; (4) "raspy chatter" - contact; (5) "snarl" threat; (6)
"growl" - threat; (7) "scream" distress; and (8) "tooth chatter"
threat.
INTRODUCTION

The present research is a study of the sounds emitted by three

species of vociferous, social rodents: the black-tailed prairie dog
(Cynomys ludovicianus (Ord) ), the white-tailed prairie dog (C. leu-
curus Merriam) and the Gunnison's prairie dog (C. gunnisoni Baird).
Previous systematic study of prairie dog sounds has been done by King
(1955), Tembrock (1963), Tileston and Lechleitner (1966) and
Longhurst (1944).
The purpose of this study was to describe quantitatively and pic-
torially the sounds produced by these three species.

METHODS AND MATERIALS

From February 1964 to June 1966, field observations and tape
recordings were made of prairie dogs in Colorado and Wyoming.
Representative observations at all daylight hours during the seasons
each species was active aboveground were made from blinds. Addi-
tional data were obtained from captive prairie dogs.
Black-tailed prairie dogs were studied in two locations, both in
Larimer Co., Colorado. One area is at an elevation of 1615 m at the
west edge of the Great Plains, lat 40?34'N; long 105?09'W. The
other area of study is at an elevation of 1490 m, lat 40033'N; long
1t0458'W.
One area where white-tailed prairie dogs were studied is at an

1 Present address: Department of Animal Industries and Department of

Zoology, Southern Illinois University, Carbondale, Illinois 62901.

167

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168 THE AMERICAN MIDLAND NATURALIST 83(1)

elevation of 2420 m in the Larimer River Valley, Larimer Co.,

Colorado (lat 40055'N; long 105?56'W). Another area studied was
at 2375 m elevation near the Colorado-Wyoming border, Larimer Co.,
Colorado (lat 40059'N; long 105?49'W). The third area was in
Albany Co., Wyoming, at an elevation of 2360 m (lat 41006'N;
long 105030'W).
Gunnison's prairie dogs were studied in three locations, also. One
location was near the type locality for the species in Saguache Co.,
Colorado, at 2865 m elevation (lat 38008'N; long 106?43'W). The
second was at 3030 m at the base of Black Mountain, Park Co.,
Colorado (lat 39006'N; long 106002'W). The third was beside the
South Platte River in Douglas Co., Colorado, at an elevation of
1935 m (lat 39t17'N; long 105?12'W).
The sounds of the prairie dogs were recorded in the field and
laboratory on a professional model W-610-EV Amplifier Corporation
of America TransMagnemite battery-operated, spring-motor tape re-
corder, driven at 15 in/sec. At this speed the recorder had a calibrated
frequency response of 100 to 12,000 Hz ? 2 db. The speed was
extremely accurate. For example, pure tones of 100, 1000, and 4001
Hz were replayed at 100, 999, and 4001 Hz. The microphone was an
Electro-Voice model 644 Sound Spot unidirectional microphone.
In the laboratory, the tape recordings were analyzed on a sound
spectrograph (Kay Electric Company model 661-A Sona-Graph).
Type "B" Sona-Graph paper was used. Wide-band, HS (high-frequency
pre-emphasis), and original speed analysis were used first. Narrow-
band and half-speed spectrograms were made for further interpretation
when needed. The original recordings were transferred to the Sona-
Graph from an Ampex PR- 10 tape recorder. A Hewlett-Packard
model 350D attenuator was used in conjunction with the PR-10
recorder. Frequency, duration, and interval measurements were taken
from the sound spectrograms.
Photographs (16mm motion pictures and 35mm transparencies)
were taken to supplement my field observations of postures and
movements.
I live-trapped many of the prairie dogs, determined their sex,
marked them with either Nyanzol "A" or "D" fur dye, then released
them while I was still at the capture site.
Pups were studied after they first came aboveground at approxi-
mately 1.5 months of age.
RESULTS
It is necessary to rename some of the sounds already described by
other workers. My approach will be consistently to name the sounds
by terms which refer to their physical, rather than their functional,
characteristics. Naming the sounds by their possible function (e.g.,
territorial call) limits the sound to one meaning, whereas sometimes
a sound appears to have different functions under different ecological
conditions as recognized by Collias (1960) and Waring (1966).
The "fundamental" of a sound will be used to refer to the first

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 169

harmonic or lowest frequency component consistently visible on the

sound spectrograms of multiovertone sounds. The "dominant" fre-
quencies or harmonics are those with the greatest amplitude on the
spectrograms (cf. Watkins, 1967). By the "function" of a sound,
I refer to the purpose or evolutionary advantage for the sound being
emitted under certain circumstances (e.g., a sound functioning to
warn others of danger).
I detected no difference in the sound repertoire of the prairie
dogs between different study areas of the same species.

I. BARKS

Repetitious Barks.-The sounds commonly heard by any visitor to

a prairie dog town are the "repetitious barks." Pups after a few days
aboveground and the older male and female prairie dogs emitted
the "repetitious barks" characteristic of their species whenever they
were alarmed. If there was sudden or imminent danger, more than
one prairie dog in the neighborhood sometimes gave the barks, al-
though commonly only the first individual to see the intruder gave
the barks and continued until the intruder was gone or until it no
longer caused a fear response. In every instance observed, the vocaliz-
ing animal was alert and remained so while producing the call.
Nearby prairie dogs became alert as soon as the "repetitious barks"
began, but commonly they awaited visual cues from the vocalizer,
other prairie dogs, or their own observations before taking further
action. If one prairie dog ran for a mound, others did likewise. If no
further acts of alarm occurred, the nonvocalizing prairie dogs looked
about and then returned to their previous activities, even when the
vocalizer continued to bark. Prairie dogs that had wandered far from
their burrows often ran back to their mounds whenever the "repeti-
tious barks" began. Prairie dogs occasionally emitted these sounds for
prolonged periods when I could see no reason for alarm.
The vocalizing posture was sitting up on the haunches, standing
on all four legs, or standing up on the hind legs. The black-tailed
prairie dog was also seen to sit, crouch and even lie prone while
vocalizing. The head was held with the nose raised to a horizontal
level with the eyes. Unless the intruder was quite distant, the prairie
dogs presented one side to the intruder and watched with monocular
vision. The mouth of the black-tailed prairie dog was opened only
partially during vocalization, whereas the mouth of the other two
species opened completely with each sound, then partially closed
before again opening for the next sound emission, the tongue of these
two species was obvious from a side view and bulged outward at each
sound. The abdominal muscles of all three species contracted notice-
ably with each emission of the sounds.
The "repetitious barks" of the black-tailed prairie dog can be of
one or two syllables, or a rather indefinite syllable transitional between
the two (Table 1; Figs. tA, iB, tC). There seems to be no uniform
difference in the function or purpose between the two extremes; how-

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170 THE AMERICAN MIDLAND NATURALIST 83(1)

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 17 1

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172 THE AMERICAN MIDLAND NATURALIST 83(1)

ever, under utmost alarm the prairie dogs frequently gave the more
obvious two-syllable sounds. The frequency of the first syllable was
always equal to or higher than that of the second. Intervals between
barks are shortest during the sudden or close approach of danger.
The animal sometimes continued the barks with only a few short pauses
for nearly an hour, although the call was normally given for less than
30 minutes. The intensity and intersound intervals became variable
during the long barking sequences. On a few occasions single barks
were heard which alerted nearby prairie dogs. These sounded iden-
tical to the components of the "repetitious barks."
In the black-tailed species, vertical tail flicking was done frequently
by vocalizing and nonvocalizing individuals when alert or uneasy.
The tail of the vocalizer was flicked with each bark. As an intruder
approached, the prairie dogs moved into the craters of their mounds
and peered over the rim with only their heads and flicking tails

I I

TIME (0.1 seC intervals.)

Fig. 1.-Sound spectrograms of sounds of black-tailed prairie dogs. A

Monosyllable Repetitious Bark. B. Bisyllable Repetitious Bark. C. Indefinite-
syllable Repetitious Bark. D. Chuckle. E. Chatter Barks. F. Wee-oo. The
analyzing filter bandwidth is 300 Hz; high-frequency pre-emphasis is utilized.

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 173

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174 THE AMERICAN MIDLAND NATURALIST 83(1)

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 175

showing as they continued to bark. In the other two species, the tail
was motionless and down during alarm behavior.
White-tailed prairie dogs produced very short "repetitious barks"
which were repeated so rapidly they became a vocal chatter (Table 2;
Fig. 3A). Unlike the continuous barks of the black-tailed prairie dog,
these sounds were emitted in sets of about 30 sounds each. After one
set, a pause of 1-15 seconds preceded another set of barks. Within a
set the intersound intervals were rather constant. The first barks of
each set were composed of the purest tones; later barks became noisy.
The "repetitious barks" of the Gunnison's prairie dog (Table 3;
Fig. 4A) consisted of barks with a rather long duration compared to
those of the other two species. The sounds were repeated at approxi-
mately three per second and were grouped into short sets of 2-25
barks per set with a pause of 3-15 seconds between each set. Inter-
sound intervals were shortest upon extreme alarm. The first sets
commonly had the most sounds per set.
Chatter Barks.-Sometimes black-tailed prairie dogs emitted barks
so rapidly that the sound was like a chatter; hence, I shall call them
"chatter barks" (Fig. IE). These sounds were often given by an adult
female which had retreated from a male during the reproductive

Ic

. r | - ; ; ;-- -- -A'- r ---

~~~~~~~~~~~~~~~~~A i . !

IOOJ

W~~~~~~~~~~~~~

TIME (0.1 sec intervals)

Fig. 2.-Sound spectrogram of sounds of black-tailed prairie dogs. A.

Raspy Purr. B. Snarl. C. Scream. D. Growl. E. Tooth Chatter. The ana-
lyzing filter bandwidth is 300 Hz; high-frequency pre-emphasis is utilized.

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 176 THE AMERICAN MIDLAND NATURALIST 83(1)

season. The sounds were directed to the male if he continued to

approach her. Prairie dogs before and after chasing another prairie
dog sometimes made these sounds. One August, some pups made the
"chatter barks" when they appeared fearful of an adult male nearby.
Rapid Barks.-Gunnison's prairie dogs sometimes gave a long
series of "rapid barks" when seemingly alarmed or apprehensive.
The intersound interval and sound duration were less than for the
"repetitious barks" of this species. There commonly was an obvious,
low-pitched gular or some type of inhalation sound between barks
(Fig. 4B). These sounds were heard when male and female prairie
dogs were transported in a car or were held captive in bive-traps at
camp. A captured female pup made these sounds for several minutes
at a time when strong winds preceded an approaching storm. She
crouched in the live-trap with her head held low. A male pup in
another live-trap nearby made a weak series of somewhat similar
barks during the strong winds. However, each sound was only the

I.e . , .,,
I A . i A:1

Ah
S. ~ ~ ~ ~ ~~~~.

TIME (0.1 s'ec intervals)

Fig. 3.-Sound spectrogramns of sounds of white-tailed prairie dogs. A.
Repetitious Barks. B. Chuckle. C. Laughing Barks. D. Snarl. E. Scream.
F. Growl. The axnalyzing filter bandwidth is 300 Hz; high-fre-quency pre-
emphasis is utilized.

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 177

last half of the usual "rapid bark." The geometric mean duration of
12 sounds was 20 msec and that of 11 intersound intervals was 150
msec. Modified in this way, the sounds seemed to be a repetitious
whimper (Fig. 4C). The following day, this animal was dying from
sylvatic plague. I never heard the whimper sounds again. Without
further evidence, I consider them as a type of "rapid barks."
Another Gunnison's prairie dog produced the "rapid barks" upon
seeing my microphone beside its mound as it came out of its burrow.
A nearby prairie dog became alert, then went down its burrow.

II. CHUCKLES

The barks emitted by prairie dogs as they ran down, or deep

within, their burrows sounded like a "chuckle." Both sexes and indi-
viduals older than two months often produced these sounds as they
were released from live-traps and ran into their burrows.
The "chuckles" are different for each species. The sounds emitted

C:<~~~~~~~~~~~~l

w D

701
I 3 I

TIME (0.1 sec intervals)

Fig. 4.-Sound spectrograms of the sounds of Gunnison's prairie dogs. A.
Repetitious Barks. B. Typical Rapid Barks. C. Modified Rapid Barks. D.
Chuckle. E. Raspy Chatter. F. Growl. The analyzing filter bandwidth is
300 Hz; high-frequency pre-emphasis is utilized.

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178 THE AMERICAN MIDLAND NATURALIST 83(1)

by black-tailed prairie dogs were a brief series of their "repetitious

barks"; however, the barks sounded different coming from within
the burrow because the higher frequencies were lost in the burrow
system and the fundamental became the more dominant audible fre-
quency (Fig. ID). White-tailed prairie dogs produced their "repeti-
tious barks" and Gunnison's prairie dogs gave their "rapid barks,"
but again the higher frequencies were lost in the burrow system. (Figs.
3B and 4D).
III. CONTAGIOUS VOCALIZATIONS

Each species had one vocalization that was repeated by neighbor-

ing individuals once the call was initiated. The allelomimetic call of
the black-tailed prairie dog was highly contagious and was repeated
immediately by several neighbors. Among the other two species, the
calls were repeated after a delay of several seconds.
Wee-oo.-The two-syllable call (Fig. 1F) which sounded like
"wee-oo" (w&eoo) was the most unusual of all the vocalizations of the
black-tailed prairie dog because of the associated behavior. The first
syllable was emitted as they extended their heads up and back and
raised onto their hind legs with their backs arched. The forelegs were
extended out and upward above the horizontal. Finally, the animals
were in a quasi-opisthotonous position for a fraction of a second
before they relaxed and came down to stand on all four feet. On the
way down, they emitted the second syllable. The whole act occurred
in less than one second. Frequently neighboring prairie dogs began
to repeat the call before the originator had given the second syllable.
The situations where the "wee-oo" was given varied greatly. Some
black-tailed prairie dogs periodically gave it as they paused during
feeding or from their resting place on the mounds. Two individuals
sometimes suddenly stopped and gave the call after one had been
chasing the other. Often they gave the sounds soon after first peeking
out of their burrow entrances early in the morning or immediately
after a dive by an aerial predator. Captive prairie dogs once habitu-
ated to their new surroundings gave the "wee-oo" after hearing a
human cough. My pet prairie dog gave these sounds whenever our
domestic dog shook itself and rattled the metal tags attached to its
collar. Once a male pup made five of these vocalizations within its
burrow immediately after being released from several days in cap-
tivity. On a few occasions, a single sound which seemed to be the
first syllable of the "wee-oo" was heard. It was produced especially
by pups during peaceful situations.
Laughing Barks.-When white-tailed prairie dogs were undisturbed
and the environment seemed peaceful, both sexes periodically emitted
a rather short set of vocalizations which sounded like "laughing
barks" (Fig. 3C). The call was given in sets of 2-15 sounds per set.
The first one or two sounds of these musical complex tones were as
much as 40% longer in duration than the following sounds in the set.
The animals stood, sat upi hunchbacked, or were slowly running when
they emitted the sounds. The head was outstretched causing the mouth

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 179

to be extended forward and up. Others continued their activities

for several seconds then made the sounds themselves.
Raspy Chatter.-The allelomimetic call of the Gunnison's prairie
dog in peaceful surroundings was a "raspy chatter," consisting of low
intensity, low-pitched, rather noisy sounds repeated with irregular
intersound intervals 2-15 times in a short sequence (Fig. 4E). The
vocalizing postures were the same as for the "laughing barks" of the
white-tailed prairie dogs.
IV. SNARLS

Free-ranging black-tailed prairie dogs were heard giving a brief

"snarl" when threatening another prairie dog that persisted in attempt-
ing to mount or paw the unreceptive individual. The vocalizer often
feigned an attack. The sound was characterized by being a broad band
noise with resonance bands and often some undulation in pitch
(Fig. 2B).
A black-tailed prairie dog held in the talons of a hawk repeatedly
"snarled" as it wiggled and fought to free itself. Individuals in live-
traps often "snarled" when a hand was repeatedly waved past the side
of the trap. Adults, especially, gave these sounds and resumed a pos-
ture of sitting or lying partially on one side, mouth opened ready
to bite, and often raised one foreleg.
Though snarls were not heard among free-ranging white-tailed
prairie dogs, individuals of both sexes (adult males especially) in live-
traps often gave "snarls" with their mouths opened as if preparing to
bite when a hand threatened them. They often raised one or both
forelegs as they faced the threatening object. The sound was a loud,
moderately high-pitched noise with two or more resonance bands
above 1500 Hz (Fig. 3D).
The threatening sound heard between free-ranging Gunnison's
prairie dogs had a snarl-like characteristic with undulation of pitch.
It sounded similar to the "snarl" of the other two species and of higher
pitch than the "growl." A female caught in a trap with a male snarled
at the male whenever he crowded or crawled over her. Another
female snarled briefly while attempting to free herself from the grasp
of a male during copulation. I was unsuccessful in tape-recording
these sounds.
V. GROWLS

Captive males and females of all three species emitted a "growl."

In the black-tailed prairie dog this sound was given when the menace
was not too close or too annoying. For example, when a hand was
passed directly over a captive prairie dog, "snarls" were given, but
when the hand was moved away or was not yet close to the captive,
growls" were commonly heard. The sounds were pulsed and had no
undulation of the resonance bands (Fig. 2D). Free-ranging black-
tailed prairie dogs occasionally emitted "growls" when annoyed by
the presence of another nearby. Quiet "chatter barks" sometimes
followed.
White-tailed prairie dogs made the "growls" when threatening my

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 180 THE AMERICAN MIDLAND NATURALIST 83(1)

hand or a probing stick, and often during occasions when they had
also emitted "snarls." Their "growl" was a low-pitched, low-intensity
sound with resonance below 1500 Hz (Fig. 3F).
The "growl" of Gunnison's prairie dogs was similar to the "growl"
of the other two species (Fig. 4F). The "growling" prairie dog was
usually sitting, ready to bite, and often raised one or both forelegs.
Compared to the other two species, Gunnison's prairie dogs were
generally silent when live-trapped and handled.

VI. SCREAMS

On a few occasions, captive black-tailed prairie dogs made a low-

amplitude, musical, abbreviated "scream" immediately following a
"snarl." The sound was primarily of low frequencies and had har-
monics (Fig. 2C).
One September morning a young black-tailed prairie dog ap-
peared with one or both hind legs lame. I watched it groom its left
hind foot. Then it wandered on, feeding and attempting to keep its
weight off the left foot. Suddenly it shrieked, seemingly in pain, as it
darted a meter away. The vocalization was a three-syllable, musical,
high-pitched sound lasting one second. Possibly it reinjured itself, for
it groomed both hind legs and then limped away. The sounds seemed
like prolonged, though higher pitched, forms of the "scream."
Adult female white-tailed prairie dogs and pups of either sex often
produced a "scream" when held captive by my hand. Occasionally
the sounds were made within the live-traps. Their "scream" was a
short sound undulating in pitch and had clear harmonic structure
(Fig. 3E).
A screaming sound was made by a Gunnison's prairie dog as a
digit was removed for identification purposes. The rather musical
sound was like a slightly prolonged bark of high and variable pitch.

VII. P URR

A captive female black-tailed prairie dog began making "raspy

purring" sounds at one year of age when it was scratched by a human
being. Pet prairie dogs seem to appreciate having their bodies rubbed
and scratched. The sounds were rather short and low in amplitude and
pitch (Fig. 2A). Often the "wee-oo" followed the "purr."
Social epimeletic grooming was observed only twice in my study
areas; I could not detect any sounds on either occasion. Therefore,
I am unable to say whether the "raspy purr" occurs in the wild.

VIII. TOOTH CHATTER

The most audible nonvocal sound black-tailed prairie dogs nmade

was the "tooth chatter." By anteroposterior and posteroanterior move-
ments of the lower jaw, the prairie dogs clicked the tips of the lower
incisors against the upper incisors. The result was a rapid clicking
or chattering sound. Both sexes produced these sounds; however,
during the reproductive season captive males especially produced the

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 181

"tooth chatter" toward their usual human handlers and were hostile
toward them. Occasionally a pet prairie dog being scratched emitted
the "tooth chatters," although the chatters were barely audible.
Each clicking sound was an extremely brief, wide band noise, and,
as shown in Fig. 2E, there is resonance in the lower frequencies pos-
sibly due to the oral cavity.
These sounds could only be heard when the animal was close to
the blind. Sometimes the subject was sitting alone on the mound;
other times the sound was given as two prairie dogs were alert and
close together. I never was certain that I heard it during disputes
between two prairie dogs. Dr. R. R. Lechleitner (personal communica-
tion) has heard "tooth chatter" from black-tailed prairie dog males
caged side by side during the period the testes were in the scrotum.
A male Gunnison's prairie dog in a live-trap when threatened with
my hand made a number of "tooth chatters" which sounded identical
to those of the black-tailed prairie dog; however, the intensity was
so low the sounds were almost inaudible.

IX. OTHER SOUNDS

Black-tailed prairie dogs of both sexes held around the back of the
neck sometimes produced a grunting or quiet coughing noise. It was
of low intensity and varied from one individual to the next in pitch and
duration. In general, it was low-pitched and brief (less than 1 sec).
The captive seemed in discomfort due to the constriction of my hand.
Each species had vocalizations that functioned as scolding sounds.
Black-tailed prairie dogs annoyed by others of their species, but not
directing an attack toward them, sometimes made one to three squeak-
ing or chirping sounds. My pet black-tailed prairie dog emitted several
of these sounds in a low but varied intensity if we annoyed her. These
sounds may have been a short version of the "chatter barks." I once
heard a white-tailed prairie dog emit a squeaky bark when seemingly
scolding another prairie dog after they had greeted each other. They
leaped apart as the one emitted the sound. G-unnison's prairie dogs
sometimes produce one or several squeaky barks when seemingly
trying to scold another prairie dog. The sounds were of low intensity
and short duration. They may have been modifications of the "rapid
barks."
DISCUSSION
The sounds of the prairie dogs are a useful tool for identification
of species in the field. Yet, most of the sounds of each species, though
physically different, are functionally similar to sounds of the other two
species. Each species has sounds functioning for threat, alert, or group
cohesion (Tables 1, 2, 3). Some sounds are similar in certain physical
properties between species but have different functions. For example,
the individual sounds of the "laughing barks," the contact call of
white-tailed prairie dogs, appear somewhat similar in sound structure
to the alerting "repetitious barks" of the Gunnison's prairie dog. The
threatening "chatter barks" of the black-tailed species and the "rapid

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182 THE AMERICAN; MIDLAND NATURALIST 83(1)

barks" of apprehensive Gunnison's prairie dogs are also somewhat

similar in sound structure.
Since earlier observers of prairie dogs have not published sound
spectrograms nor have they given a detailed description of the sounds,
I have had difficulty knowing whether I have heard the same sounds.
In Table 4, I have attempted to list the names of the sounds given by
previous authors which seem synonymous with those described by me.
King (1955) and Smith (1958) were able to distinguish both

TABLE 4.-Synonymous names of the sounds of prairie dogs

Name used in Previous name with

this paper original author

Black-tailed Prairie Dog:

Repetitious Barks Alarm note (Merriam, 1902)
Squit-tuck (Seton, 1926)
Warning Bark (King, 1955)
Hawk Warning (King, 1955)
Alarm Bark (Smith, 1958)

Chuckle Chuckle (King, 1955)

Chatter Barks ? Muffled Barks (King, 1955)

? Churr (King, 1955)
? Chirr (Anthony, 1955)

Wee-oo Yelp (Jillson, 1871)

Cry (Wilder, 1872)
Song (Seton, 1926)
Territorial Call (King, 1955)
All Clear (Anthony, 1955)

Snarl Snarl (King, 1955)

Scream Scream (King, 1955)

Growl Skr-skirr (Seton, 1926)

? Churr (King, 1955)
Growl (Tileston, 1961)

Tooth Chatter Tooth Chatter (Squire, 1925)

White-tailed Prairie Dog:

Repetitious Barks Alarm Notes (Cary, 1911)
? Warning Bark (Tileston, 1961)

Chuckle Chuckle (Tileston, 1961)

Laughing Barks Querulous Cry (Cary, 1911)

All Clear (Tileston, 1961)

Growl Growl (Tileston, 1961)

Gunnison's Prairie Dog:

Repetitious Barks Warning Bark (Longhurst, 1944)

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1970 WARING: PRAIRIE DOG SOUND COMMUNICATIONS 183

''warning (alarm) bark" and a "hawk warning" of the black-tailed

prairie dog. The sounds I heard during alarm due to aerial predators
were the same on the sound spectrograph as those emitted for ground
predators. Since more than one individual would often vocalize as a
hawk or an eagle made a dive and since the call was made only
during the rapid maneuver of the bird, the call seemed very rapid and
consisted of only a few barks. Nevertheless, I found no reason to
separate their "repetitious barks" into two or more distinct types of
calls. All my observations suggested to me that the prairie dogs, unless
far from their burrows, seek the protection of their burrows because
of visual cues and not because they heard the "repetitious barks." Often
alarm was communicated without the use of sounds.
The "chatter barks" seem not to have been described by previous
observers, unless they are what King (1955) terms the "muffled barks"
or possibly the "little barks" (p. 76) which are given among disputing
prairie dogs. They may be the sounds Anthony (1955) described as
a "low 'chirr'."
Several authors have discussed the "wee-oo" of the black-tailed
prairie dog. I adopted the name from Seton's (1926) description.
It is a difficult call to label with a single function; for as King (1955)
aptly points out, it is. given in a variety of circumstances. I conclude
that it and the corresponding call of the other two species are contact
or group-cohesion calls-sounds of social familiarization serving to
maintain the social bonds among the group. Other mammals have
sounds of similar function (Tembrock, 1963), although rarely is there
such a complex action pattern as seen during the "wee-oo" of the
black-tailed prairie dog.
The "screams" I heard of the black-tailed species might be similar
to or a slightly modified version of what King (1955) and Smith
(1958) call a "scream." I would expect such a sound to vary greatly
from one situation to another depending on the state of fear or distress
of the animal.
What Tileston (1961) and I name the "growl" of the black-tailed
species might be a portion of the sounds King (1955) labels as "dis-
puting churr" or what Seton (1926) described as a "skr-skirr" sound.
No distinct sounds of the black-tailed prairie dog were heard that
could be identified as those King (1955) named "defense barks."
The "warning barks" Tileston (1961) described for the white-
tailed prairie dog seems to be the same as the "repetitious barks" for
this species except for the intersound interval. His description of
intervals of 0.5 second is far longer than the data I collected. The
"laughing barks" of this species are probably made up of what Cary
(1911) called the "querulous cry" and seem also to be the "all clear"
mentioned by Tileston (1961).
Burnett and McCampbell (1 926), Longhurst (1 944), and Scheffer
(1947) all seem to have been referring to the "repetitious barks" when
discussing the sounds of the Gunnison's or Zuni prairie dogs. None of
the other sounds has been mentioned previously in the literature.

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 184 THE AMERICAN MIDLAND NATURALIST 83(1)

Within the genus Cynomys there appear to be affinities between

C. leucurus and C. gunnisoni when sound production characteristics
are used; yet, the evidence suggests that the species are distinct. In
both species, alerting sounds are grouped into sets of barks, mouths are
opened to extreme at each emission, and tails are motionless during
these sounds. Contact calls of both species are given as a series of
sounds using similar postures. In C. ludovicianus the alerting sounds
are a continuous series of barks accompanied by partially opened
mouths and flicking tails; the contact call is a two-syllable call given
with a specific complex action pattern. When C. ludovicianus is com-
pared to the other two species, there is similarity to C. gunnisoni.
Only in these two species did tooth chatters and continuous (first
order) sound sequences seem to exist. Similarity is especially evident
between the "chatter barks" of black-tailed prairie dogs and the
"rapid barks" of Gunnison's prairie dogs.
There are no obvious affinities linking Cynomys to Spermophilus or
Marmota using only the sound emissions. Alarm and threat sounds
in the three genera are distinct aside from the tooth chatter (cf.
Waring, 1966; Balph and Balph, 1966). At present among the three
genera, contact calls appear to be limited to Cynomys.

Acknowledgments.-Laboratory work was niade possible by the cooperation

of the Colorado State University Speech and Hearing Clinic. Partial support
came from the F. C. Jean Research Award of the Colorado-Wyoming Academy
of Science and from U. S. National Science Foundation (Summer, 1965) and
U. S. Public Health Service (1-F1-GM-30,852-01) fellowships.
This research, under the direction of Dr. R. R. Lechleitner, represents part
of the requirements for the Ph.D. degree at Colorado State University. Drs.
P. H. Baldwin, N. H. Booth, J. A. Modrick, 0. W. Olsen, Mr. R. E. McFar-
lane, my wife, and numerous landowners graciously gave valuable assistance.
To all the above I give my sincere thanks.

REFERENCES

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Cynomys ludovicianus. J. Mammal., 36:69-78.
BALPH, D. M. AND D. F. BALPH. 1966. Sound communication of Uinta
ground squirrels. Ibid., 47:440-450.
BURNETT, W. L. AND S. 'C. MCCAMPBELL. 1926. The Zuni prairie dog in
Montezuma County, Colorado. Colorado State Entomol. Circ., 49:3-15.
CARY, M. 1911. A biological survey of Colorado. U.S. Dep. Agr., N. Amer.
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COLLIAS, N. E. 1960. An ecological and functional classification of animal
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SUBMITTED 23 DECEMBER 1968 ACCEPTED 20 JANUARY 1969

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