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California State University, Northridge Sound CALIFORNIA STATE UNIVERSITY, NORTHRIDGE SOUND COMMUNICATION IN THE •• CALIFORNIA GROUND SQUIRREL A thesis submitted in partial satisfaction of the requirements for the degree of Master of Science in Biology by Nurit !!_evy August, 1977 The thesis of Nurit Levy is approved: California State University, Northridge August, 1977 ii ACKNOWLEIX;MENTS I wish to thank all the people who have helped and encouraged me with this study. Most of all I would like to express my deepest thanks and appreciation to Dr. Jim Dole for the in­ valuable help he gave me throughout my stay at C.S.U.N. I wish to thank him for his enlightening suggestions, ideas, in­ spiring guidance, and for the kindness, patience and innumerable hours he spent in thoughtfully reading the preliminary drafts leading to this manuscript. I also wish to thank him for his most exciting and stimulating lectures which motivated me to do this research and become involved with animal behavior. I am also most grateful to Drs. George Fisler and Charles Weston for their helpful suggestions. Many thanks are extended to Dr. Andrew Starret for providing me with his ultrasonic re­ ceiver; to Mr. Tom Haley who taught me how to use the sonagraph; and to Mr. Tom Nayler who took very special care of it. Sincere thanks to Professor Pedro Durant who was a great help in the early stages of the study, to Mr. Paul Abravaya for designing new traps and to Mr. Jeff Werber for building them. I am especially indebted to ¥~. George Boyle and Mr. Edward Boyle for permitting the use of their ranch for this study and to Mr. Roger Beam for his kindness and cooperation in providing access to the study area whenever needed. My most sincere thanks to Marilyn, for her efficiency and enthusiasm in typing this manuscript. And iii finally, I wish to thank all my fellow graduate students that were of great help and encouragement throughout the study , particularly Betty Rose and Steve Lenchner, for whom my appre­ ciation is boundless. iv TABLE OF CONTENTS ABSTRACT viii INTRODUCTION 1 MA.TERLA.LS AND METHODS 7 Study Area 7 Field Observation Procedure 8 Study of Captive Animals 10 Sound Analysis 11 RESULTS 12 Spectrum of Vocalization 12 1. Chatters 12 2. Squeals 13 3. Whistles 13 4. Grunts 14 5. Screams 14 6, Tooth chatters 14 Ultrasonic Sound 26 FUnctions .of Vocalization 26 a. Response to predators and potential predators 26 Response to aerial predators 27 Response to ground predators 28 Response to conspecifics 29 Vocalizers 30 b. Vocalization used in intraspecific encounters 30 1. Threat 31 2. Blocking 31 3. Supplanting 32 4. Chase 32 5. Fight 33 6. Sand kicking 34 7. Territorial disputes 34 8. Courtship behavior 34 Nature of Sound Used in Agonistic Behavior 35 DISCUSSION 39 LITERATURE CITED 60 v ' ' TABLES Table Page 1. Characteristics of the "chatters" of the California ground squirrel. 16 2. Characteristics of the "squeal" of the California ground squirrel. 17 3. Characteristics of the "whistles" of the California ground squirrel. 18 4. Characteristics of "tooth chatter" of the California ground squirrel. 23 5. Catalog of the vocalizations of the California ground squirrel. 57 vi FIGURES Figure Page 1. Typical chatters of the California ground squirrel. Adult chatters (A,B,C), a chatter emitted by a young animal (D). 20 2. Squeals (A,B) and whistles (C) of the California ground squirrel. 22 J. Tooth chatter (A), pulsed grunt (B), two continuous grunts (C), and scream (D) of the California ground squirrel. 4. A grunt emitted by a male while chasing a female which he shared his burrows with (A), a grunt emitted during male-male fight (B), a section taken at the point of the arrow of the above grunt. 38 vii ABSTRACT SOUND COMMUNICATION IN THE CALIFORNIA GROUND SQUIRREL by Nurit Levy Master of Science in Biology August, 1977 The purpose of this study was to examine the vocal commu­ nication system of the California ground squirrel, Spermophilus beecheyi. Tape recordings of the sounds of marked squirrels were made in the field, while concurrent notes were taken on the be­ havior of the animals; feeding boxes were used for intensifying interactions among individuals, Supplementary data were collected from captive squirrels. The recordings were analyzed with a sonagraph. On the basis of their physical properties, vocalizations fell into six distinct groups: chatters, squeals, grunts, whistles, screams and tooth chatter. By combination and repetition of these sounds, the use of the same sounds in different contexts, and the use of graded signals, the number of messages used for viii communication was increased. Two types of alarm calls were given in response to predators: squeal indicated an immediate alarm; a long call (chatters followed by repeated signals) served as an alerting signal for less immediate danger. Females were found to be more alert than males and gave most of the long alarm calls. A whistle was emitted during sexual chases. During agonistic encounters, grunts and chatters were used in association with domina...'1.ce and a tendency to attack, while a squeal was used to indicate subordination and a tendency to flee. Some sounds (grunts, screams and tooth chatter) were found to be associated with territorial defense. ix INTRODUCTION Communication has been defined as "an action on the part of one organism that alters the probability pattern of behavior in another organism in a fashion adaptive to either one or both of the participants (Wilson, 1975). Its advantage to any species lies in the potential for transmission of adaptive information among the groupmembers. Indeed, communication is an essential part of social organization, for the ability of group members to communicate makes social bonds possible. As might be expected, among vertebrates, a correlation tends to exist between the de- gree of social integration within a species and the complexity of the communication system of that species (Fisler, 1970; Barash, 1977) . The emission of sounds is but one possible means of cornmu- nication, but the rapidity of sound transmission over great distances and through objects and vegetation makes it well adapted for information transfer in many circumstances. Its importance is readily documented by its widespread occurrence among the vertebrates, for the use of sound signals is well known in all groups except the reptiles. For instance, many species of fish use sounds for attraction between sexes, defense against other species, threat and schooling (Moulton, 1963). Amphibian calls are well known to serve in mating, maintaining territories, for warning and for signaling release from clasping in copulation 1 2 (Blair, 1968). Sound emission in reptiles, although minimal, is associated in some groups with reproductive activity and ter­ ritoriality (Pope, 1946; Evans, 1961). Bird vocalizations include complex songs, thought to accomplish functions asso­ ciated with courtship, territorial and aggressive behavior, and simple call notes serving for alarm, distress, bond main­ tenance, flocking, nesting, and attracting other individuals to food sources (Stokes and Williams, 1972). Mammals typically have a well-developed vocal organ and many species have highly developed sound communication systems. ·Primates have been studied extensively in this regard. In one species, the vervet monkey (Cercopithecus aethiops), 21 different situations have been found to cause distinct sounds, including several different agonistic calls, distinct alarm calls for different predators in different proximities, and different infant sounds (Struhsaker, 1970). Vocalizations in rodents are also widespread and in some species extend into the ultrasonic range. Peromyscus nasutus, for instance, vocalizes up to 100KHz (Sebeok, 1968). A very elaborate vocal display known is the "song" of the humpback whale (Megaptera novaegliae) analyzed by Payne and McVay (1962), which is thought to serve in identifying individuals and keeping groups together during migrations. As among vertebrates in general, in some groups of land mamrnals it has been demonstrated that the extent of vocal rep­ ertory correlates with the degree of sociality. Among Canids, for instance, foxes (Vulpes vulpes), characterized by relatively 3 solitary and asocial behavior patterns, show a more limited vocal communicatory repertory than do wolves (Canis lupus) whose social organization is much more complex (Fox, 19?0). Similarly among primates, chimpanzees (Pan troglodytes), the socially most advanced of the nonhuman primates, well-known for their extraordinary cooperation, possess a rich repertory of sounds composed of 25 signals (Busnel, 1963), whereas the less social gorilla (Gorilla gorilla beringei) employs only 16-17 vocal displays. In rodents, a study by Fisler (1970) also shows that the number and complexity of acoustical signals varies from the more socially organized guinea pig (Cavia porcellus)~ to the less social mouse (Reithrodonto~ys megalotis), whereas the least number of sounds is used by the unorganized jird (Meriones unguiculatus). A similar relation occurs among squirrels. The black tailed prairie dog (Cynomys ludovicianus), which lives in a very highly organized social system, employs an exceptionally rich repertory of auditory signals. It emits nine different sounds for different intensities of alarm, intra- and interspecies threat, contact, distress, apprehension and pleasure (King, 1955; Waring, 1967). The Olympic marmots (Marmota olympus merriam), which live in distinct, well-integrated colonies, have an au- ditory communication system composed of seven calls used to indi- cate alarm, distress, departure of a predator, yearling fights and more vigorous adult fights (Barash, 1973). The colonial yellow bellied marmot (Marmota flaviventris) is also knoWn to 4 possess a vocal repertory of seven distinct vocalizations aroused by different stimulus situations (Waring, 1966). As reported by Brand (1976), the chipmunk (Eutamias townsendii), a relatively social squirrel, also uses seven calls for communication. Several are used as alarm calls and the rest in agonistic and courtship situations. On the other extreme of the social scale, fox squirrels (Sciurus niger) live thinly scattered and appear to be rather solitary.
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