VOWLUZATION BEHAVWR OF THE RING-NECKED PHEASANT Thesis far its Begree of M. S. PMCFEIGAR STATE UNEVERSETY GARY HEINZ 18$? Hawaii w m I'll/WU / Ill/III/l/Illllll j Michiganstm 3 1295fl/l/Il/l/clzll1/[Lo 1023 University / ATERlAL TO ‘ MENTARY M i RE mififw THE. BOOK PLEASE "0‘" AC m Ammo LmRARY _.___———————-—--—' VWZN’ .. - __ 1-3.; I ~—' ABSTRACT VOCALIZATION BEHAVIOR OF THE RING—NECKED PHEASANT by Gary Heinz Only a few of the many ring-necked pheasant vocali- zations are commonly heard in the field; yet these vocali- zations are important in understanding the activities of this bird. This study is an attempt to record for a perma- nent collection all ring—necked pheasant calls, to describe the behavioral significance of these calls, to analyze the structure of each call, to investigate the ontogeny of calls, and to prOpose possible uses of sound techniques for studying ring—necked pheasants. Typical sound spec— trographs and a #5 rpm record of calls accompany verbal descriptions of vocalizations. The study was carried out from June to September of 1966, and both wild and captive pheasants were observed. The following calls are described for males: crow, subcrow,* alarm, flight, hiss, antagonistic,* and pecked*, for females: pecked,* precopulatory,* flight, brood gathering, and brood caution; for chicks: content, caution, and flock. *These calls were previously undescribed in the liter— ature for the ring-necked pheasant. Gary Heinz Intra-individual variation in call structure exists but is not as great as inter-individual variation, and it is often possible to identify a particular bird by the constancy of its call structure. The inter—relationships among calls and the meaning of calls in total pheasant behavior are complex problems to study, and the study of calling behavior is complicated by inter—individual varia— tion in reSponse to Specific stimuli. The use of calls to census, trap, and study pheasants could be put to better use as an important tool for the wildlife biologist and behavioral scientist alike. Some calls elicit fairly stereotyped responses from pheasants, and playback of tape recorded calls into the field can often attract wild ringnecks and evoke calling from them. Considerably more research is needed to increase the understanding of some less well understood calls, to ex- plain the ontogeny of calls, and to find practical appli- cation of calls in the field to study ring-necked pheasants. VOCALIZATION BEHAVIOR OF THE RING-NECKED PHEASANT By Gary Heinz A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Fisheries and Wildlife 1967 ACKNOWLEDGMENTS I sincerely thank: The Michigan State University Agricultural Experi- mental Station and the Department of Fisheries and Wild- life for financing the study. My major professor, Dr. Leslie W. Gysel, for the con- ception of an animal vocalization study, for suggestions concerning eXperimental design, and for encouragement throughout the study. Committee members, Dr. Rollin H. Baker and Dr. Martin Balaban, for counseling on the design of the study and for helpful criticisms on the text. Dr. William L. Thompson of Wayne State University for use of facilities in his laboratory of biological acoustics and for training in the use of the sound Spectro- graph and interpretation of spectrographic results. Mr. and Mrs. Alfred Mayer who made available their pheaanfi;farm for sound recording and observation of pheasant behavior, supplied pheasants for pen studies, and offered valuable information on pheasant calls and behavior. Mr. Donald Riggs for advice on the use of the sound Spectrograph and for supplying equipment. ii The Rose Lake Wildlife Experimental Research Station Staff who provided pen facilities. Dr. P. P. Kellogg who supplied ring-necked pheasant calls from the Cornell University Library of Natural Sounds. The professors, fellow students, and others who offered sincerely appreciated advice and encouragement. iii TABLE OF CONTENTS ACCNOWLEDGMENTS LIST OF TABLES LIST OF FICUFES . . . INTRODUCTION MATERIALS AND METHODS CATALOG OF CALLS. Adult Males Crow. Subcrow. Alarm Flight Hiss. . Antagonistic Pecked Adult Females. Pecked . Precopulatory. Flight . . . Brood Gathering Brood Caution. Chicks Content. Caution. Flock . . . . . Calls of Uncertain Meaning Males Females. . . . . Ontogeny of Vocalizations. Early Development Content. . . Flock Caution. Crow. Alarm Flight Hiss. iv Page ii vi vii [UMP—W4 CDEKOQUWU‘! W U) MUGUJUOLUUO ”KO CID'N] Gwen) 41’ t. +_ \ \ \DDI‘: C)O\O O\ 0“. (Tx Ox (h (‘1‘. (DUI m \le U1 U1. U1 m U Ix} m H i—J H-fl—SJ (“to m e—J Page DISCUSSION. . . . . . . . . . . . . . . 6H Catalog of Calls. . . . . . . . . . 6A Complexity of Vocalizations . . . . . . . . 6A Call Variation . . . . . . . . 66 Application of Sound Techniques. . . . . . . 68 SUMMARY AND CONCLUSIONS . . . . . . . . . . 71 Study Methods. . . . . . . . . . . . 71 Pheasant Vocalizations. . . . . . . . . 72 Application of Sound Techniques. . . . . . . 7A Areas of Possible Research . . . . . . . . 74 LITERATURE CITED. . . . . . . . . . . . . 75 Table 1 LIST OF TABLES Summary of ring-necked pheasant vocalizations vi Page 73 lo. 12. 13. 14. LIST OF FIGURES Maximum variation by day and time of day found in the crow call of a male . . . . Intra-individual and inter—individual variations in the crow call of eight males Inter-individual variation in the subcrow call in four groups of males . . . . . . Intra—individual and inter—individual variations in the alarm call of ten males . . Intra-individual and inter~individual variations in the flight call of three males . Intra-individual variation in the hiss call of a male . . . . . . . . . . . Intra-individual variation in the antagonistic call of a male . . . . . . . . . . Inter-individual variation in the pecked call of three males . . . . . . . . . Inter—individual variation in the peeked call of three females . . . . . . . Intra—individual and inter—individual variations in the precopulatory call of three females . . . . . . . . lntra-individual variation in the flight call of a female . . . . . . . . Intra-individual variation in the brood gathering calls of a female . . . . . . Intra-individual variation in the brood caution calls of a female . Intra-individual and inter—individual variations in the content call of two chicks . vii 1A 18 22 27 BO 30 35 35 Al Al Al 49 Figure 15. 16. 17. 18. Intra-individual variation in the caution call of a chick Intra-individual and inter-individual variations in the flock call of five chicks Calls of uncertain meaning The ontogeny of vocalizations viii Page 149 “9 55 6O INTRODUCTION The ring-necked pheasant, Phasianus colchicus, has a great repertoire of vocalizations, but only a few of these calls are commonly heard in the field. In an attempt to establish the importance of vocal communication in the behavior of this bird, a study was carried out from June to September of 1966 with the following objectives: (1) to identify and tape record for a permanent collection as many distinctly different calls as possible, (2) to illustrate the structure of and establish the behavioral significance of each call, (3) to describe, in part, the ontogeny of calls, and (4) to gain insights into the use of the knowledge about calls and the understanding of their effects on pheasants when played back to census, trap, and study the behavior of ring—necked pheasants. A 45 rpm record is included in the Jacket on the front cover of this thesis. Calls representative of those discussed in the thesis are found on this record. MATERIALS AND METHODS Both penned and wild pheasants were studied; obser— vation of penned birds was carried out largely at the Mayer Pheasant Farm in Brighton, Michigan. Additional studies of captive pheasants were made at the Rose Lake Wildlife Research Center in Clinton County, Michigan, and at the Fisheries and Wildlife Department pens at Michigan State University. Field studies were carried out at the Rose Lake Wildlife Research Center; the Ingham County, Michigan, crowing survey route; Michigan State University farmlands; and in other areas in Michigan. All calls were recorded at seven and one—half inches per second on a Uher 4000 Report-L portable tape recorder equipped with a Uher M514 microphone and a twenty-four inch parabola. Structural analysis of sounds was made with a Kay Electric 66lA Sonograph on wide band analysis. Descriptions of the behavioral significance of calls were arrived at both by observing the calling behavior of unmolested pheasants and by manipulating the stimulus situ— ations to which pheasants were exposed. This latter method included playback of tape recorded calls to captive and wild birds. A portable Ampli-Vox 8-200 Diplomat amplifier and loudspeaker was used to playback recorded calls. The vocal response of males to tape recorded crowing was studied on the Ingham County crowing route on eight days in mid—July. Twenty fixed stops were visited in the morning and evening of four survey days during the peak of crowing (from one—half hour before to one and one—half hours after sunrise and from one and one-half hours before to one-half hour after sunset), and all runs were made in the same direction over the survey route. Each stop was visited for two minutes, the first of which was a silent period; during the second minute a tape containing six dif- ferent pheasant crows, spaced ten seconds apart, was played over loudSpeakers directed toward opposite sides of the road. The number of crows heard from the field was recorded for each minute. 0n the remaining four survey days the (procedure was the same, but only the odd numbered stops were visited, this time with two minutes of silent listen- ing and two minutes of taped crowing. 0n the last five days the time lapse between a tape recorded call and the crow of a wild male was also estimated and recorded. Other observations of pheasants were made when birds were presented with mounted pheasants and when penned and wild pheasants were approached and intimidated. Observa— tions were also made of chicks which were deprived of food, warmth, and the companionship of other chicks. The ontogeny of calls was studied by observing vocal behavior of various aged chicks and by following two particular chicks, a male and a female, in their vocal devel- opment for several weeks. Ideas for possible use of calls to census, trap, and study pheasant behavior evolved from an understanding of the meaning of the various calls, from observations of pheasant response to tape recorded playback of calls, and from study of the structural characteristics of pheasant calls. To induce crowing, two adult males were injected with testosterone propionate U.S.P. Some crows which were ana— Ibmad sound Spectrographically were provided on tape by P. P. Kellogg from the Library of Natural Sounds at Cornell University. Nomenclature used in the description of sono- grams follows, inlbuge part, definitions set down by Davis (1964). CATALOG OF CALLS Adult Males Adult males have a remarkable range of vocalizations. The calls of males are generally more raucous, lower pitched, and louder than those of females. In the field, the male's crow call, alarm call, and flight call are the most fre- quently heard of all pheasant calls. As with all other ring-necked pheasant calls, there are apparently no male calls which extend beyond the range of human hearing. Only minor portions of any ring-necked pheasant calls have been noted to fall outside the 250 to 10,500 cycles per second range of hearing (Stewart, 1955) of this bird. In the cata— log which follows, the behavioral significance of, the structure of, the variations found in, and the literature concerning each male vocalization are discussed. 2:911 This harsh two syllable sound, described by Delacour (1951:231) as "korrk-kok" and by Leffingwell (1928) as ”a w k - kack," can be heard under favorable conditions for about nine-tenths of a mile (Kimball,l949), making it the loudest of pheasant calls. Wind, topography, plant cover, man-made obstructions, and background noises generally reduce the distance crowing carries. Ball (1950) listed 5 one-fourth of a mile as frequently being a more reasonable estimate of the carrying distance of the crow. Occasional crowing is heard all year but reaches a peak during the Spring when males are defending territories and are breeding. Dawn and dusk are favored times for crowing, but some crows are heard throughout the day and even on moonlit nights (Kimball, 1949). Growing frequency apparently is not affected by temperature, dew, nebulosity (Kozicky, 1952), relative humidity, fog, or mild rain (Kimball, 1949). The behavioral significance of crowing is not defi- nitely known. This call probably warns other cocks not to treSpass into territory defended by another male (Allen, 1956 and Leffingwell, 1928). From June 19 to July 24 field studies were carried out to determine the reSponse of wild cocks to playback of tape recorded crowing. The location of crowing males was determined, and these males were cautiously approached to within about fifty yards. From a concealed spot a prerecorded crow was played over a loud— speaker every six and one-half seconds on aloop of tape. In earlier tests it was determined that the sound equip— ment which was used could broadcast such a call easily one- half mile for human ears and probably further for a ring- necked pheasant. Of ten trials run in the above manner in late June and early July, eight were successful in attracting wild males up to the loudspeaker. Cocks always approached silently to within ten to fifty feet from the loudSpeaker and then began giving the alarm call. The time required to attract a male varied from two to six minutes; in no case was a tape played for more than twenty minutes if it failed to attract a male. The nature of this alarm call response is discussed further under the alarm call. Later in July, as crowing of wild males greatly decreased, it became considerably more difficult to attract cocks to a tape recorded call. Apparently the crowing of a male with- in the territory of another male acts as a challenge call and provokes the approach of the resident bird. Only rarely does a male respond to taped crowing within its territory by crowing itself. Males were attracted during all daylight hours, but no attempts to attract males were made at night. It has been suggested (Allen, 1956) that crowing may also function to attract the attention of hens, but Ruffing (1952) never saw hens attracted to crowing males during his intensive field study of crowing behavior, and through- out my study I never observed hens approaching tap re— corded crowing; however, trials were made well past the peak of the breeding season. The most acceptable explana- tion is that crowing alerts hens to the presence of a cock in their vicinity; although contact between the sexes may result, quite independently of crowing, through the fre- quenting of the same feeding, dusting, or roosting areas. The stimulus for crowing has never been adequately investigated. Ruffing (1952:68) commented, ”The evidence, though fragmentary, indicates that crowing in pheasants is a neurophysical response to certain hormonal stimuli, coupled with external sensory releasors." There is a direct relationship between gonad weight and crowing fre- quency (Kimball, 1949) which indicates that testosterone may play a role in establishing the urge to crow. In an attempt to induce crowing, a male, not heard to crow during several early morning observation periods, was intra- muscularly injected on August 17 with one dose of 100 mg testosterone propionate, suspended in corn oil. A great increase in the size of and a deepening of the red color of the face patch took place within a few days, and the first crowing from this bird was heard on August 29. Growing continued for several days but diminished in fre- quency after the first four or five days, when the cock could be heard crowing about once every ten minutes in early morning and somewhat less at dusk. Such evidence does not indicate, however, that testosterone is the sole stimulus for crowing. Various sounds and vibrations such as thunder, car doors Slamming, other males crowing (Kimball, 1949); explosions (McClure, 1944); earthquakes (Leffingwell, 1928); and "any unusual noise" (Bent, 1932:319) can evoke crowing. Gates (1966) demonstrated that the intensity of crowing is not indepen- dent of population density, indicating a greater competi— tion for territory and/or a mutual stimulation of crowing; yet Ruffing (1952:70) in intensive direct observation of males never observed one cock responding to the crow of another. An experiment, described under Materials and Methods, was designed to compare the natural rate of crowing in wild males with the rate of crowing during playback of tape recorded crowing as a stimulus. Since so little crowing was heard, the data were considered insufficient to analyze for the effects of different stops, time of day, or length of the listening period. A paired t test failed to show a significant differ— ence between the natural rate of crowing and the rate of crowing with tape recorded crows as a stimulus: P(-l.4 :,t 1 1.4) > .05 It is concluded then that, with the techniques used, there was no significant effect of tape recorded calls on the number of crow calls heard. Although wild males seldom appeared to respond to tape recorded crowing, I noted that calls from males in the field closely followed taped calls. I therefore estimated, for five crowing survey days, the time lapse between a taped crow and a crow from a wild male. If taped calls did not act as a crowing stimulus, the time lapses between taped crows and crows from the field should have varied randomly from zero to ten seconds (the time interval between taped crows), and the expected mean time lapse should have been five seconds. A t test was used to test whether time lapses departed significantly from five seconds. The observed mean time lapse for fourteen IO recorded calls was 2.857 seconds with a variance of 2.593. Random sampling and a normal population must be assumed. The null hypothesis that the observed mean time lapse did not differ significantly from the eXpected value of five seconds is rejected with a one tailed test: P (t _>_ 4.984) < .005 This test indicates that wild males did reSpond to taped crowing. The paired t test may have failed to show the same result for two reasons. Sampling was probably inadequate, and males close to the loudSpeaker seldom crow, but instead give the alarm call response. It is possible that only males which had not recently crowed, and were therefore predisposed to crow, answered taped crowing; other males which had crowed recently were perhaps not inclined to crow and thus could not be induced to crow with taped calls. During summer months when crowing is seldom heard, few males would be predisposed to crow at the moment of taped stimulus; this would account for the fact that, although few males responded, those that did crow answered taped calls quickly. Sound spectrographic analysis (Figures 1 and 2) reveals that the crow is distinctly disyllabic and that the first syllable is usually longer than the second syllable. Both syllables contain the same set of harmonics with the fundamental at about 800 to 1000 cycles per second (cps) and with overtones at each 800 to 1000 cps 11 Figure 1. Maximum variation by day and time of day found in the crow call of a male. FREQUENCYHILOCYCLES PEI! SE00") I. Iaiiieii August 31 0:10 All q," ww- - 'v "wee-nu. M 12345678 September I 1:05 AI Jr“! 'fll f-l. "fill-"v I" n} ler’v 1!! I ' “."W. 'l'" J‘ a, Magnet 30 721.5 M! 0- ‘11? n .w 7 A ' g) ’HWM 7 e 6" . H, ' 6- 5‘ '1 H| 1:5," .0 5‘ 44 . a 4‘ 3" '- '.' ' - 3‘ 2‘ Wm» r, e 24 l‘ . ememm. w l- 1 2 a 4 5 3 '7 e .. lens: 31 5:29 II '1 74 ‘ 7, be 61 sJ 5' 41 4‘ 3+ 3‘ 2‘ 2. h ‘ '1 I 2 3 4 5 6 7 a '1 (Must 31 6:43 PM 8. 7‘ 7. 6‘ 61 5‘ ‘ 51 “ 4‘ 3‘ 3, 2« _.:',:w~.- = ~ ~ , 2. l' um I. h I 2 3 4 5 6 7 8 .; smegma ezoem 8‘ . 7‘ 61 51 44 3-1 2-1 I TIIEUENTHS OF A SECOND) a '2 5 3 3 3 } 5 September 5 5:50 AM I e M 0 rt «mum w:— ."b.l" ' 125555§i 13 Figure 2. Intra-individual and inter—individual variations in the crow call of eight males. FIEIIEICY (IllICVClES PEI SECIII) .' W l . I. ‘ ‘ 7 WA! A V ’ ,, «~33 ‘ 3:“ 'l‘e F A” a in.) II- 64 ' (‘2 W ‘4 ' “i" 5‘ im‘ "& “WWW : 43': ‘- HI! 0h M A 3‘ . 21 ‘1 oeaeese 6123456 .. 2 I I .. e 7‘ 6‘ n Hui" " w 1w»: m w; “ been an M ' 4 1‘ e I Ii 5555356 éiiiisi (viiiiii l: 3 .1 ,. ‘ .}1 .' 5 7« 1+ 4’“ ,.: 1- I . .I 1 ‘l ‘ I.“ w . ‘ v - “ me:- me 5‘ ’2; 5' " h “ M h ‘: " H p 3‘ e. 3‘ ...~ 3. .. 2* 2: 2‘ '4 I: " 5155533 oTiiiiZ 01233? .' 1 .' 9'7"!” 1"” 7‘ 7‘ '. Q! '4' 6- 61 5‘ “um! I. 5‘ Hg; 1. a. m u 34 2' 3‘ I 1‘ 0123455 61i55§3 012345 "I! (IEIIIS If A SECIII) l5 interval above the fundamental. The harsh quality of the pheasant crow is caused by the frequency modulation and amplitude modulation found in this call. Harmonic (fre- quency) bands often bow up and down, particularly at the beginning and end of a syllable. Pheasant crows may differ in pitch, timing, and loudness. In an individual cock, differences in these three factors are slight. Eight calls, representative of maximum variations found in over forty calls of the same male, illustrate intra—individual variation of the crow (Figure 1). Small changes in syllable lengths and differ- ences in the distribution of sound in the eight harmonic bands comprise the greatest variation. Some apparent variation is caused by distortion and artifacts produced by the tape recorder equipment and by the sound Spectro— graph. The typical consistency of crow structure within a male is better shown by cocks l and 2 (Figure 2). Although intra—individual variation is slight over different days and at different times of the day (Figure l), more study is needed to follow males over longer periods of time to detect possible seasonal differences in call structure. Inter-individual variation in the structure of the crow call is greater than intra-individual variation. Variations in timing and harmonic structure are easily seen on sonograms (Figure 2). The eight calls shown give a good idea of differences in pitch and timing of calls; variation in loudness was not investigated in this study. 16 Calls with emphasis upon the higher harmonics (cocks 2 and 3) sound higher pitched than a crow (cock 6) which has primary emphasis upon the lower harmonics. Differences in call length and pause between call syllablesene easily seen; cock 4 shows a rare case in which the first syllable is longer than the second syllable. Cock 1 has a crow with a fairly high fundamental of about 1000 cps (the frequency band actually covers 300 or 400 cps and is only approxi- mately centered at 1000 cps), whereaS'Uwacrow of cock 7 has its fundamental frequency band centered at about 800 cps. These differences in crow structure have been reported in the field (Ball, 1950; Delacour, 1951:236; and Ruffing, l952:9) as differences in the quality of the crow. Sound Spectrographic analysis also enables one to distinguish between crows which could not easily be told apart by the human ear. Subcrow Leffingwell (1928) noted that when males crow in early Spring, the vocal performance is not accompanied by wingbeating, and also that in August and September some males of that year's hatch attempt to crow, but without the wingbeat. In my study during the summer, males also voiced a crow-like call, but with no wing flapping. It was common to hear penned cocks give this call in chorus, apparently stimulating each other to call. Three or four birds may call within a two-second interval (Figure 3), u H A. igure 3. Inter—individual variation in the subcrow call in four groups of males. EIEOIEICV (IIlOCYClES PEI SECOND) eeoezeeee "IE (IEIIIS If I SECIII) l9 and often the reSponse of one bird to another's call is so quick that calls may overlap. Only occasionally is any wingbeating seen. In the field this call would be difficult to distinguish from crowing unless the observer were close enough to the male to notice the absence of wingbeating. The function of the subcrow is presently unclear. It may serve as a subcall to the fully developed territorial crow, much as subsongs function in the vocal repertoires of songbirds. There is no explanation available for the chorus calling frequently noted with the subcrow. The structure of the subcrow is similar to that of the crow (Figure 3). The two syllables of the subcrow are more nearly alike in length than they are for the crow, but the frequency and amplitide modulations and the bowing of frequency bands is about the same for the subcrow as for the crow. Presently, nothing can be said of intra- individual variation in this call. Inter-individual dif- ferences in harmonic structure seem greater than in the crow, while differences in timing of the subcrow appear less than in the crow. Alain The alarm call, perhaps the most regularly heard of all pheasant calls, is frequently given when a strange or menacing Sight or sound confronts captive or wild males. The alarm call of one bird often elicits the same call from other nearby males, and occasionally with penned or wild 20 birds a chorus of alarm calls will arise, usually at dawn or dusk hours, with no apparent stimulus. The meaning of the alarm call is complex for there is a broad spectrum of stimulus stiuations which elicits it. Captive males usually can be induced to make the alarm call simply by entering their cage or otherwise alarming them, Mwneasin the field wild cocks may give this call as one approaches them or works in an area close to them. The alarm call also can sometimes be evoked by playing tape recordings of the crowing call within the territory of a male. The alarm call response to crowing is fairly stereo- typed. The calls at first follow each other quickly, often less than one-half second apart, but gradually dimin- ish over a period of minutes to about one call every two or three seconds. Birds may call for more than ten minutes this way or until frightened off; provoked captive males have sometimes continued calling for more than one—half hour. The ability to induce this alarm call response to crowing apparently decreases as the breeding season passes, Beebe (1931, 11:45) stated that the alarm call, "a ' may function to attract hens or act as a lower cackle,’ guiding call to the feeding grounds._ As yet there is no evidence to support the first claim, and only once were males heard giving the alarm call during what was presumed to be undisturbed feeding activity. Since the alarm call is given under so many circumstances it may be wrong to assign to it any single meaning. 21 The structure of the alarm call is similar to that of the crow and subcrow, but the harmonic structure is far more variable, and sharp changes in frequency often occur throughout the duration of the call (Figure 4). The funda— mental generally lies a little below 1000 cps as with the crow and subcrow, and bowing of the harmonics is very common. Leffingwell (1928) described the call as "a di- or tri—syllabic call which may be given as cucket, tucket, or tucketuck.” Actually the alarm call exists in mono, di, and trisyllabic forms, but the mono and disyllabic calls are by far the most common. Calls sometimes seem to gain one syllable due to a change of inflection within a syllable, and ”tucket" and "tucketuck" described by Leffingwell are probably really mono and disyllabic calls. The three syllable call does exist but was heard only once in the field and never among the thousands of alarm calls heard from penned males. No tape recording was made of a tri— syllabic alarm call, but the repetitions by the wild male were identical to the ear and can be described as the simple addition of one short, less accented syllable before a typical two syllable alarm call. There was no obvious dif- ference in behavioral Significance of the three forms of the call, and the same male has been heard to give both mono and disyllabic calls, although each bird has a particu- lar form of the call which it generally gives. Intra—individual variation is greater in the alarm call than in the crow, particularly if a male has been alarm 22 Figure 4. Intra—individual and inter—individual variations in the alarm call of ten males. PIEIIEICYHILINCLES PEI SE00") a“? 1‘! em: cell let-e nieyllelic cell vatietiee emu eecke: e e s leeeeyllebic cell neietiee eleu cocks: .1 Q .' .14" "IE (IEIIIS IE I SECIII) 24 calling for some time. For this reason it is more diffi— cult to distinguish males by the alarm call than by the crow call. Inter-individual variation is based more on whether a bird gives a one or two syllable call and on peculiarities of frequency bands than on timing of the call. One syllable calls are usually about one decisecond long, and two syllable calls are generally composed of two one—decisecond syllables. The pause between syllables in disyllabic calls varies from essentially no time (some calls are neither distinctly mono nor disyllabic) to slightly more than one decisecond. Flight When flushed from hiding, males generally give the flight call as they take to wing. No evidence is available to indicate whether some males never give this call or if all males use it irregularly, but the latter explanation seems most likely. The flight call sounds very much like the initial few moments of the alarm call of some males on the ground. Birds begin calling immediately upon taking flight and con— tinue calling for as short a time as two seconds or for Ilonger than seven seconds. Calls are spaced fairly evenly, aldout one to two deciseconds apart, until the few firal Céalls which may be more irregularly Spaced. Leffingwell (1928) described this call as "invariably tIfiisyllabic,” however during my study no trisyllabic flight 25 calls were recorded. As with the alarm call, two syllable calls often seem trisyllabic because of changes within the first syllable. Flight calls which are technically monosyllabic have been recorded, but they are borderline cases between one and two syllable calls. The flight call is most often given when males are forced to flight as they are approached to within a few feet in the field. On one occasion in my study a male gave the flight call as it flew toward a loudSpeaker broad- casting crow calls within his territory. When rising fromea roosting area in early morning, males sometimes utter a flight call. These latter examples indicate that the flight call may have a broader meaning than merely escape from danger. The flight call is quite similar to the alarm call in sound spectrographic structure; and as with the alarm call, the harmonic structure of the flight call is much more irregular than that of the crow call. Marked changes may occur in the frequency bands of the call (Figure 5). Fundamental frequencies of flight calls are generally lOOO cps or less, and the timing of calls is about the same as for the alarm call. Intra-individual variation is greater, however, for the flight call than for the alarm call. There are no recordings available of more than one entire flight call sequence for any male, so variations in the “pattern of calls given by a male as he is flushed are not known. Within one sequence of calls, however, 26 Figure 5. Intra-individual and inter-individual variations in the flight call of three males. -..e-- u I 6 .9 fl 0;. n 2.. ‘ Imu‘v EWN 2 .. .. m - BAP! 2 Iae .‘ e. N 3 I. 9. o ‘3‘“ 6 376AMMH¢quh 2 ‘1 . 4 2 *2 I”. l ‘ “It, 2 I l 2 . “70 err I‘ll”. .4 o n” J E J 2 PW . fl ., rt 3 . p I- ‘u j.“ I v6 0" .In. 3 L «4‘9, 0 .. I I .r v-3 . $3». - M I . I .2 o.4 - . v.53 3 , _QPHV 1 u - r .m 1...... ' 4.. .8.‘ .4. 2 rnmvhmme mrmrmmmu 07654321 32.3» a: 3333.58.33: M I. II I! HIE (PENIS If I SEIIII) 28 intra-individual variation is noticeable. Occasionally a cock will use both one and two syllable calls in the same flight sequence, and variation may exist in the structure of the calls and also in the pause between syllables in two syllable calls. Inter-individual variation in harmonic structure and call timing is much like that found in the alarm call; but two syllable flight calls rarely exceed a total length of two deciseconds and usually are only about one and one—half deciseconds long. 112: This call is given under different circumstances and may convey more than one meaning. During courting, males make a hissing sound after a display pose is held (Leffing- well, 1928) and also after c0pulation (Delacour, 1951:237). In my study males also sometimes hissed when approached or handled. In one case a male hissed as a microphone was extended to within inches of him as he roosted in a pen late at night. Following several hisses the bird flew about making the alarm call. Other males placed in bags for tranSport hissed as they struggled to get out. Females have also been heard hissing when ap— proached, but no recording is available of this sound. The hiss is audible only at close range, and no reSponse to this sound was observed in nearby pheasants. Sonograms of hisses show a very wide band of fre- quencies involved in this sound (Figure 6). The fundamental Figure 6. Figure 7. 29 Intra-individual variation in the hiss call of a male. Intra-individual variation in the antagonistic call of a male. flEflIElflflllOCYClES PEI SECOND) NM “'1". 'W' n,“ g“ ,"v '4 I. II 20 22 "I! "II": If I SEC...) 32')“, ummm O 2 4 31 begins near baseline and extends upward to nearly 1,500 cps with faint overtones sometimes visible. Intra-individual variation is almost entirely in call length, which may be little more than one decisecond or as long as nine or more deciseconds. The interval between hisses varies from a fraction of a second to several seconds. No recordings are available to show inter-individual variation, but to the ear the hisses of different males sound the same. Antagonistic In my study only penned males were observed giving this call. Calling males generally ran along the fence separating them from the cock in the adjacent pen and pecked at the sheet metal which extended from the ground to about two feet up the fence. The male in the pen adjacent to the calling bird often reSponded with the same call and behavior. Such incidents often lasted several minutes with both males becoming quite vocal. This same call was evoked from males by placing within their pens a mounted male ring-necked pheasant. Live males cautiously approached the mount, slowly circled it, and then pecked at it. It is possible that the antagonistic call, given during the breeding season, would provoke attack from another male. This call was also heard from at least one female, believed to be either an intersex or sex-invert pheasant (see Morejohn and Genelly, 1961, for a description of sex deviations in ring-necked pheasants). One hen repeatedly 32 gave this call as she attacked and once mounted another hen in the pen. Structurally this call is fairly complex. Although the most intense sound occurs in a wide frequency band at about 1,000 cps, parts of the call extend throughout the 8,000 cps frequency range of the sonograph (Figure 7). There can be a noticeable variation in the call of one male. Two very low syllables at about 500 cps often initiate the antagonistic call sequence. These syllables are about two deciseconds long and are Spaced about one decisecond apart. After another one decisecond pause, ten or more one-half decisecond syllables, spaced about one decisecond apart, follow; these syllables are loudest at about 1,000 CpS. Then follow several pulses of calls varying in length from one decisecond to over five deci- seconds. Pulses are generally spaced about three to five deciseconds apart. Call syllables vary from one to five centiseconds in length, and often the last syllable of a pulse of calls is about twice as long as the others. Intra—individual variation in call syllable length rarely exceeds two centiseconds. The interval between syllables is constant for an individual but ranges from less than one centisecond to a few centiseconds in different males. There also exists both intra- and inter-individual varia— tion in pitch and volume of the antagonistic call. 33 Pecked When pecked by another pheasant of either sex, males emit a dissonant sound which is easily recognized as the pecked call. The pecked male momentarily retreats, but shows no signs of antagonism, and other nearby pheasants show no response to the call. Sound spectrographic structure is apparently harmonic with emphasized frequency bands from about 2,000 to M,000 ops (Figure 8). This call is harsh, and sometimes rapid changes in frequency occur. No recordings are available for more than one call from any male, but comparisons among different cocks show more variation than is found in most calls. Most variation is found in the harmonic structure and not in the length of the call, which varies little from an average duration of about one decisecond. Adult Females Hen pheasants also have a considerable range of vocal- izations, but their calls are not as loud as those of males and are less frequently heard. Females apparently have no territorial calls and did not voice calls of an antagonistic nature. Hens produce sounds extending over the same fre- quency range as males, but generally female calls are highs: pitched. A listing of female calls follows. 34 Figure 8. Inter—individual Figure 9. Inter— variation in the pecked call individual variation of three males. in the pecked call of three females. Figure 10. Intra-individual and inter—individual variations in the precopulatory call of three females. "Elli“! (IIlICVClES PEI SE00") I! W 3 0’ .1 7‘ W. 7 O1 .. “ . ‘ ’1 1 . q. ‘ 4‘ ‘* V . N 3* " u w " 24 -' fr 0 2‘ 1 " --—6-I-r--—-u—u—-v- O I o I I O 2 4 6 O '0 I! I4 |6 II 30 12 O 71 . ’ ‘4 :1 Im at “'3. M 3‘ . 2‘ ‘1 t A 4 ~L-"W 74332333333353740414440 "I! ('fl'l‘ .f I “3"” 36 Pecked This call seems identical to the male pecked call in meaning and in the stimulus. Also like the male's call, it is given only once when the hen is pecked, and there is very little subsequent reaction save for a momentary re- treat. Other pheasants nearby show no reSponse to the call. The female pecked call is a sharp, high pitched sound. It is characterized by a rapid slurring of the frequency upward, a bowed frequency band in the middle of the call, and a rapid slurring of the frequency downward again (Fig— ure 9). Generally a relatively constant frequency band exists at 2,000 to 4,000 cps. Since no more than one call was recorded for any hen, there was no opportunity to determine the amount of intra— individual variation. Like the male pecked call, consid- erable variation exists among calls of different birds. The length of this call varies from slightly less than one decisecond to over three deciseconds. The same degree of 'variation is found in the frequency of greatest intensity. Usually this band occurs at about u,000 to 6,000 cps, but it may be located as high as 7,500 cps and perhaps higher. This variation found in both male and female pecked calls is greater than that found in most other pheasant calls. The reason seems to reside in the fact that the pecked call is an unplanned cry which carries, perhaps, little meaning other than momentary pain. Such a sponta- neous call might not be expected to possess any constant 37 features other than those controlled by the structure of the bird's syrinx and by the rapid expulsion of air through this organ. Other calls, more Specific in meaning (e.g. mating, territorial, or antagonistic) could be expected to evolve in time into fairly concrete forms, understandable to all members of the species and conveying important infor- mation for survival. Precopulatory This call has been reported for the bluegrouse, Dendrogapus obscurus, (Stirling and Bendell, 1966); but not for the ring-necked pheasant. In my study, evidence for the precopulatory call comes only from observations of penned pheasants. This call is given under quite specific circumstances. Hens crouch into the submitting posture and begin giving the precopulatory call, sometimes calling even after being mounted by the male. Males respond to the hen‘s calling by quickly approaching and mounting her. Calling may last several seconds; and hens call during all daylight hours but most often in early morning. By midsummer this call was seldom heard. Like the pecked call, the precopulatory call is com— posed of an upward slur, a bowed frequency band, and a downward slur of frequency (Figure 10). There appears to mean harmonic structure with the fundamental at about 2,500 to 3,000 cps and the first overtone, always more emphasized, 38 at 5,000 to 6,000 cps. Other overtones extend off the sono- graph scale. Intra-individual variation is easily noticeable; usually most differences occur in the harmonic structure of the call with variations of no more than a few centi— seconds in length and one or two deciseconds in duration of the pause between calls. The harmonic variation can be as great as 500 cps in the frequency of the fundamental and overtones, although some hens are much more consistent in their calls. Inter-individual variation is considerable. Call lengths vary from a little less than one decisecond to greater than three deciseconds, and the interval between calls ranges from as short as two deciseconds to as long as seven deciseconds. Differences in the complex frequency pattern of this call are also easily seen. Flight In contrast to the loud raucous call uttered by a flushed cock, the hen's flight call is a soft, high-pitched squeak. This call was generally given by a hen with a brood, although hens with young did not always make this sound. Kozlowa (1947) said of this call: If the intrusion happens to occur at the moment of rest, the hen takes wing silently and the chicks at once crouch on the ground under_available cover. If, on the contrary, the family is surprised when feeding and the chicks are scattered about in the grass, the female, when taking wing, gives a soft cry. At this signal, evidently meaning alarm, 39 the young immediately disperse, running or flying away. Those hens which do call begin as soon as they are flushed and continue calling for a few seconds. Since the young have either taken flight with the hen or have run to cover even before the call is given, the function of the call in the hen-brood relationship cannot be pinpointed; but it likely functions as a warning call. The female flight call is a slurred syllable. The frequency rises sharply from zero to a fundamental at about 2,500 to 5,000 cps, then declines rapidly back to zero (Figure 11). Intra-individual variation is similar to that described for the pecked and precopulatory calls but is not as great; a hen is farily constant in pitch and timing of her flight calls. Inter-individual variation is also reduced in this call. Calls vary from four to six centiseconds in length with greater variation in the intervals between calls, which range from less than one decisecond to over eight deciseconds apart. Calls generally become more irregularly Spaced as the hen continues to call. Differences exist in harmonic structure but are difficult to interpret pre- cisely from available Spectrograms. Brood Gathering There are two calls which seem to be associated with a hen's attempts to gather together her chicks after being Figure 11. Figure 12. Figure 13. 40 Intra-individual variation in the flight call of a female. Intra-individual variation in the brood gathering calls of a female. Intra-individual variation in the brood caution calls of a female. fllllllflfllllflflls PEI SE00") I1 74 .4 N M 31 14 Black | U. ‘ Bark "I! (mm If I SECIII) 4 4 . 42 frightened from them. One call is a soft low—pitched Cluck and the other is a harsh and fairly loud barking sound. Both calls are used under the same circumstances and are usually heard together in a sequence of gathering calls. When a hen and her brood are surprised, quite often the hen will fly thirty to sixty yards distant while her young run to cover or also fly to safety. From ten minutes to an hour or more later the hen is usually heard calling from about ten or twenty yards away as she returns to the spot from which she was flushed. Chicks, which normally become active before the hen's return and cheep loudly, generally become quiet when the hen calls but may resume cheeping if the hen pauses very long in her calling. Hens wander about searching for scattered chicks, and the calls of chicks probably serve to attract the attention of the hen. One hen was observed waiting on a roadside until she hear the call of one of her chicks, at which time she quickly ran toward the young bird. Hens generally cease calling after ten or fifteen minutes, presumably having gathered together the brood. If frightened away in her first attempt to call together her chicks, a hen will return later and resume calling. The response of hens to situations dangerous to the brood varies greatly. Some hens must be approached to within a few feet before they will flush, whereas others take flight immediately upon sighting possible danger. One hen whose chicks crouched motionless in low grass led me slowly away from the brood by hopping just fifteen to twenty feet ahead until she was far from the young, at which time she flew to safety. She returned within a few minutes to call together her brood. Holcomb (1964) reported even bolder behavior of a hen which flew to within four feet of a man holding one of her loudly cheeping chicks. As she approached she made ”clucking and squawking sounds,” presumably to alarm a predator into temporarily releasing the chick. A "low cluck" (Beebe; 1931, 11:46) and a ”kee kee kee” (Bent, 1932:315) have been reported as gathering calls for chicks, and these descriptions fit fairly well the brood gathering calls recorded in my study. Structurally the two brood gathering calls are quite dissimilar (Figure 12). The cluck is a wide band of sound from baseline to about 1,500 cps with some streaks ex- tending higher. Extreme frequency and amplitude mtdulaticn is present. Calls average fourteen centiseconds in lergth with about eight to fourteen decisecond intervals between calls. Intra-individual variation lies almost entirely in small changes in frequency and amplitude modulation, and inter—individual variation of the same sort is also small. The bark is higher pitched; the most intense fre- quencies are at about 1,800 to 2,300 cps with lower ard higher pitched streaks. There is also a faint bowed fre— quency band at about 4,000 to 4,500 cps. Calls are 44 consistently Six to seven centiseconds long and are Spaced every two or three deciseconds apart. Intra-individual variation is small, and inter-individual variation seems to lie mainly in pitch. Brood Caution The brood caution call is an abrupt, low—pitched sound which is usually uttered softly and in rapid pulses. It does not sound unlike the ground predator warning call of the domestic hen, Gallus gallus (Collias and Joos, 1953) and it serves a similar function of cautioning the chicks against apparent danger. This call in the ring—necked pheasant has been mentioned in the literature (Beebe, 1931, 31:45; Delacour, 1951:2393 and Holcomb, 1964), but, like most other calls, it has not been carefully described. It is not yet entirely known why the caution call is sometimes heard with gathering calls in the same se— quence of calls. On one occasion a hen in the process of gathering her chicks was approached and flushed. She gave a few fairly loud caution calls at the first strange noise and continued to caution call more softly until she was frightened away. In a few minutes she returned, this time usirg the caution call exclusively. A second time she was stalked, and, although considerable noise was made when a microphone was held within three feet of her, she continued to voice a soft caution call from her hiding place in deep weeds. An attempt to approach even closer caused her 45 to flush, and after two hours of additional waiting she was not heard again. If or how she ever gathered together her chicks is not known. Scattered chicks generally cease calling when the hen gives the caution call, but this is not an invariant rule. In one case a hen called steadily from one side of a road while only twenty feet across the road her chicks continued to frantically cheep and mill about in Spite of her caution calls. Eventually she flew across giving a peculiar squeaking call, and when she landed twenty feet from the young (in the direction opposite the danger) every one of her ten or so chicks instantly bolted for her, and they all hastily took to cover. The structure of this call is unique for it consists of a set of very low pitched harmonics with the fundamental beginning as low as 100 cps and extending up for about 400 ops (Figure 13). Frequently there is approximately an equal emphasis upon the fundamental and the first over— tone, and there are seldom more than four plainly visible harmonics. Calls average about four centiseconds in length with one to three decisecond intervals between calls. There can be very noticeable intra-individual vari- ation, but this is usually only in the harmonic structure. To the ear there appears to be a transition of calls from the gathering cluck to the caution call with a decrease ir length and increase in harmonic structure of the call. Often the first call of a series is different and is 46 actually composed of two typical caution calls with no pause between them. Variation among hens is generally in harmonic structure with little change in the tempo of calling or in the length of calls. Chicks Leffingwell (1928) reported five calls of ring-necked pheasant young up to seven weeks of age. Only one, the alarm call, was not distinctly recognized in my study. It is arbitrary at what age one describes the calls peculiar to chicks and claims further vocalizations to be imperfectly developed adult calls. Obviously the ontogeny f calls is a continuum, and the complete Spectrum of adult calls is acquired gradually. The following calls are those which were noted in young within the first seven weeks of life, and the nomenclature follows that established by Leffingwell (1928). Content This call, described as ”ter-rit” or ”ter-wit” with the accent on the last syllable (Leffingwell, 1928), is usually heard when chicks are warm, feeding, settling down for the night, or resting with other chicks. Evidence for this call comes only from the observation of captive chicks, isolated from an adult hen. The content call, like other chick vocalizations, is more musical than the usually harsh calls of the adults. Also like the remaining chick calls, the content call is 47 difficult to describe structurally since it changes con— siderably as the young mature. A better discussion of this development of the calls is found under the Ontogeny of Calls. Only the general structure of chick calls will be discussed here. The content call consists of a set of harmonics with the fundamental at 3,000 cps or greater (Figure 14). Gen- erally only the fundamental is seen on sonograms since the overtones often extend beyond the 8,000 cps range. Calls show mild frequency modulation, but almost no amplitude modulation. Various patterns of rising and falling fre- quencies exist in different chicks. Calls of young chicks sometimes exceed five deciseconds, while calls of older chicks decrease to about one decisecond in length. Intra-individual variation is remarkably reduced in older chicks and is only somewhat greater in the recently hatched. The volume of the call, which varies most, may rise from nearly inaudible when a chick is resting to a volume which can be heard over twenty feet away when a chick is busily feeding. Inter-individual variation is greater than intra- individual differences, particularly in the ascending and descending form of the harmonics. Within any age group the call length varies little, and intra— and inter- individual variations in the pause between calls are about the same with a range from about one to greater than five deciseconds. 48 Figure 14. Intra-individual and inter~individual variations in the content of two chicks. Figure 15. Intra-individual variation in the caution call of a chick. Figure 16. Intra-individual and inter-individual variations in the flock call of five chicks. mumvmmvms PEI SEN") l' " '\ 7i ‘1 ’1 .. an M 4:.» H N 3i 21 h ------------------------------- LI—v---—i-i-l— 0| 2 3 4 5 6 7 I 9 '0" I2 0 1 O ' g a l c 7‘ ). o. 5" t I, in 4- , 3‘ g“. M ['4 2‘ 7 "3 I" l I: I ------- -W --------- w 0 I 2 3 4 5 6 7 8 O I 2 3 4 5 6 O I a 1 , 2 7‘ ‘fi ‘0 7‘ ‘ 51 4.5- b b h- 11 o ‘ '1 5 i 4 i i no (a u Illutmu It I scan) 50 Caution When a strange object is presented to a chick, the young bird cautiously approaches the object calling ”terreep” or ”turreep" (Leffingwell, 1928) and pecks at it, assuring itself of the nature of this strange thing. The same call is voiced when chicks become antagonistic and peck at each other. This call is composed of a rapid succession of like syllables and is louder than the content call. Chicks cease calling when they become familiar with a new sur- rounding or stop antagonistic behavior toward each other. Leffingwell (1928) considered the caution call a modified content call, and in fact, both calls are given together sometimes. Structurally the caution call bears only slight resemblance to the content call, although ap- parently its harmonic structure is pitched the same (Figure 15). Call syllables are about two centiseconds long, and each successive syllable is often slightly higher pitched than the one before it. Pauses between the three syllables are about two centiseconds long, making the total call about one decisecond long. Calls are usually Spaced two to four deciseconds apart. Intra-individual variation is slight and is mostly in the frequency changes in call syllables and also in the 4. intervals between calls. Recordings are insufficient to provide much data on inter—individual variation, but presently differences appear slight. 51 Flock The flock call was frequently heard in the field when chicks were scattered from the hen. Leffingwell (1928) described the call as ”tee-erp" or "pre-erp,” repeated at short intervals. Chicks frightened to cover usually remain silent for several minutes, but then begin moving about and occasionally give the flock call. Calling gradually increases until a chick gives a group of five or more calls. The calls of one bird stimulate others hidden elsewhere to call,1wereasaaforeign sound may cause all chicks to tempo- rarily stop calling. Penned chicks which are isolated also give the flock call, but cease calling as soon as they are returned to other young. Hunger, cold, or other stressful situations also elicit the flock call, which indicates that this call may function as a general distress sound and not exclusively as an aid in reuniting a brood with the hen. Leffingwell (1928) described separately a fright call which sounded like the flock call but was louder and more vehement. The call was reported given when a chick was held captive by an enemy, and served to frighten away nearby pheasants. In my study there was no reason to believe a separate fright call exists; however, chicks were never seen held by predators. Hand-held young gave vehement flock calls. The possible existence of a fright call cannot yet be ruled out. 52 Flock calls, like content calls, vary considerably among individuals but are in general longer than content calls and are much louder. Flock calls are easily heard one hundred feet away, particularly after chicks have been calling for some time and have increased the intensity of the call. The fundamental of this call often covers 2,000 cps, and generally calls have fundamentals somewhere between 2,000 and 4,000 cps (Figure 16). Frequency bands sometimes slur upward and downward rapidly, but in some chicks the call is nearly of constant pitch. Small differences in harmonic structure is the most noticeable form of intra-individual variation. Inter- individual variation, by contrast,is great; frequency bands in various chicks slur downward, upward, once upward and once downward, or upward and downward more than once. The pitch varies 2,000 cps or more, a difference easily heard in the field. Calls are one to three deciseconds long with two or three decisecond intervals between calls in one group. Groups of calls are usually separated by sev- eral seconds. Calls of Uncertain Meaning There were many calls heard during the study for which no precise meaning could be given. Some of these calls which were often heard or which were considered important, although seldom heard, are discussed below. 53 knees Since this study was carried out during the summer, some of the male's Spring breeding calls were undoubtedly missed. Some calls were heard, however, which may have been related to courting activities. One call, accompanied by very stereotyped behavior, was observed in a few penned males. Cocks would call with a fairly high pitched, but soft, sound while they stood in place and pecked the ground. Such behavior might continue for several minutes, after which the bird would resume its normal activities. Hens did not reSpond to this call, perhaps because the male's diSplay was incomplete or because hens were not very inter— ested in males at this time of the year. Kozlowa (1947) reported a "kutj-kutj-kutj" call for males which pecked at the ground around some food as an invitation to a hen to come to feed. This call was easily recognized in different males, but noticeable intra-individual as well as inter—individual variation exists (Figure 17). Call structure is harmonic with a fundamental at about 3,000 to 4,000 cps, and calls, which are Spaced every few deciseconds, are seldom longer than one decisecond. Occasionally other soft calls could be heard during these diSplays, and low clucking calls of one cock, injected with 100 mg testosterone propionate, attracted hens. Males also voiced a low chicken-like call when ap- proached by man and in some cases with no apparent stimulus. kW 4:. Figure 17. Calls of uncertain meaning. "EWING! (KILOCYCLES PEI SECOND) 'lalo “nu" all r Cull Cut 2 \ ’\ '\ 210. b '0» '5 14 0246.810120246810 film “stun call 5‘ j I, ' ii \ "r". ’5‘” .~"' “w. N != ‘ . O 2 4 O I l0 l2 l4 I. I. 20 22 24 "I! "II": If I Slflll) This call is low pitched with frequencies extending from baseline to about 1,000 cps (Figure 17). Females is a high pitched squeak which hens sometimes voice insist- ently while standing very erect and alert. In penned females there does not seem to be any Special stimulus, but the one wild .hen from which this call was recorded had just been frightened from her brood and stood on one side of a road calling, as her chicks cheeped loudly on the other side. As She flew to join the brood she emitted the same squeak call. Like the pecked call, this squeak is a slurring upward and downward of frequencies with major emphasis upon the high frequencies, here as high as 7,000 ops with some sound extending off the sonagram (Figure 17). Call lengths are about one decisecond with irregular pauses between calls. Given under similar circumstances is another call, a high pitched syllable which is usually one of many such calls in a rapid series. Hens using this call seem to be alert and may call for more than a minute. Considerable inter—individual variation exists, but calls are always harmonically structured. Call lengths vary from five tetw er ((1 (D centiseconds to over two deciseconds, and pause 57 calls are usually about two or three deciseconds long (Figure 17). One hand-held hen gave a distress call, but other hens failed to use this call when held captive. A similar call has been reported for Gambel quail, Lophortyx gambeli (Ellis and Stokes, 1966). Other pheasants became alarmed upon hearing the distress call of the captive hen, and some males began giving the alarm call. Structurally the call is characterized by marked changes in frequency throughout its three and one—half decisecond duration (Figure 17). Calls are spaced about four deciseconds apart. Ontogeny of Vocalizations The study of the development of calls is an essen- tially unexplored area of pheasant vocalizations. Only small fragments of the total pattern of development were discovered in this study. To gain a good understanding of vocal ontogeny would require a separate study. Some gen— eral observations about call development follow. Early Development As chicks begin pecking through the shell, faint calls are first heard, although vocal sounds before hatching may occur, as in the domestic chicken (Balaban, personal com— munication). These early chick vocalizations are charac— terized by a rapid rise in the frequency of the sound and a subsequent decline in frequency, sometimes with numerous frequency changes in between. These frequency changes may 58 sometimes exceed 3,000 cps. Calls of a chick which had just begun breaking its beak through the shell show a variety of call structures, some of which are quite simi- lar (Figure 18). Most of these calls show severe frequency modulation, generally not found to such a degree in older birds. As with other initial vocalizations, their meaning and behavioral significance are difficult to understand. At the moment when a chick completely breaks free of its shell, a different type of call is often heard. The structure of this call is very unstable with instan— taneous changes in frequency and peculiar distributions of sound at various frequencies (Figure 18). Just after hatching, chick calls show less frequency modulation and fewer sharp changes in frequency level (Figure 18). Call length is variable and frequencies still range over 2,000 to 3,000 cps. After a few hours, chick calls (nowpresumably con- tent calls) become very long (Figure 18). Some calls are nearly five deciseconds long and continue to exhibit the upward and downward pattern of frequencies. By this age greater stability has been achieved in call structure and relatively consistent patterns are evident. Within a few days, call lengths have shortened to one or two deciseconds, and calls are quite constant in structure (Figure 18). Thereis little knowledge of when young acquire the full spectrum of pheasant calls. Some The ontogeny of vocalizations. "HIE"! (IllICYClES PEI SECOII) Cutout .. rm: mum all: It "I." 0' WWII 7‘ \ OT A 1' I.“ m ». 9 3i 2' 21 o 2 4 4 4 l0 ------- o 2 4 4 4 to m last site: mum m hm III 5 an m a, . 4 4' 1i I1 w--W- , , t Ti 0 I 2 3 4 5 o 2 4 6 I I0 12 o I 2 4 . Sum: III M". 3 III. {light 14 um 71 71 .. a « "{ 2 .1 Ilsa 14 cut: old 71 7. 6‘ 64 5‘ 5‘ 4. 4‘ ‘ thMfl/‘V ‘ 4 ' ' ‘ 7» 4. fl 3 U ‘ \.,‘ :4 " I w ")~ M ‘9 a “ ‘“ ~ “ . -.».t" q i vb 1': 4'4 {4 {4 :3 {a o a 4 4 "I! "II!” If I 8“...) 6l chronology has been discovered for the few calls which follow. Content This call appears moments after hatching. Chicks which are still wet give the content call when placed in an incubator with other young. It is difficult, however, to say whether the initial calls of the hatching chick are content calls or are merely the first efforts to exercise the vocal apparatus. 512% Day old chicks will voice a flock call if isolated from other chicks or if not kept warm. This call probably evolves early, perhaps along with the content call, al- though there are no recordings of flock calls this early. The content call and flock call seem to be the first calls developed in the chick and logically are derived from the initial hatching calls. Caution This call was first heard at about four weeks in chicks which were kept under fairly close observation. The first instances of use were in antagonistic situations between chicks. Later the same call was used when strange objects were introduced into the chicks' pen. Leffingwell (1928) reported that at about seven or eight weeks of age males often try to crow. This fact has also been reported by pheasant farm owners, but the actual crow was never heard from young males during my study. Juvenile males do make a variety of harsh sounds, but these seem more like alarm calls from which crowing may possibly develop. When captured, a five week old male gave an alarm call with a fairly typical structure (Figure 18). Easily recognizable alarm calls were voiced by a seven week old male; and more fully developed alarm calls were recorded from a fifteen week old male, although even at this age the structure of the calls was not as constant as it is in adults (Figure 18). Flight ,_.——._ Fourteen week old male captives gave flight calls when they were released into the field. These calls sounded very similar to those of adult cocks, but were more irregu— larly spaced than in adults and often the juvenile flight call was of short duration (Figure 18). v . L—T ‘ I," 1“ A a A; K7,“ k) Yourg of both sexes have hissed as early as five oi six weeks. As with adultsjthe call was given when the 63 young were intimidated by man or attacked by other chicks. Fifteen week old male young hissed when handled and had essentially the same call structure as adults (Figure 18). DISCUSSION Catalog of Calls The catalog contains many gaps in the understanding of pheasant vocalizations but does give evidence that calls play an important role in pheasant behavior. The importance of calls might be expected since the ring—necked pheasant has poor olfactory reception, has its vision often blocked by dense cover, but has remarkably acute hearing, thus auditory stimuli should predominate as behavioral cues. It is not surprising that a great number of calls have evolved to solve the problems of avoiding danger, breeding, caring for and educating young, and ordering social con— tacts. Complexity of Vocalizations Too often in the literature the reader finds an ap— parent simplicity of the reported animal calls. It is doubtful that such simple vocal systems exist in these animals, and such a system is certainly not found in the ring-necked pheasant. Inter—individual variability in call structure is easily recognized, but also significant is the behavioral variability related to these vocalizations. There is a great latitude of responses among pheasants to the same 614 65 stimuli, and with this understanding in mind, one is not trapped into looking for perfect norms of behavior or for exact correlations of calls to Specific stimulus situations. It should not be puzzling that not all pheasants respond in the same way and with the same calls to a given stimulus. For example, some males are aggressive, whereas others are timid and seldom give the antagonistic call. Hens in the field use different patterns of calls to gather together their broods. One of two captive chicks gave the flock call immediately upon separation from its penmate, Mrneasiflwaother, treated identically throughout raising, seldom gave this call when isolated and was considerably tamer. One of two hens held in large pens would calmly walk around you if approached,vdmmeasthe other hen in the same pen would invariably crouch fearfully and at the last moment Spring into frantic flight to escape. The behavior of these individuals could be predicted, but it would be a mistake to either classify all pheasants of the same age and sex by this behavior or to consider pheasant behavior as entirely unfathomable when presented with obvious variability. Certainly this variability im— poses problems in interpreting the meaning of calls, but the general patterns of behavior, which are the most impor- tant to study, can still be recognized after sufficient observation has been made. Some variability in pheasant behavior might be attrib- utable to the genetic variability of the so—called 66 ri g—necked pheasant. Classification of pheasants varies, but deinster (l95hzl) states that three species--Phas aiu ve rsi color, Phasianus torquatus, and Phasianus colchicus U) everal subspecieS)-—have been periodically introduced into the United States. MacMullan (1960) says of Michigav asant stock: (Ii Ph Michigan's pheasants are a mixture of a number of subspecies of the genus Phasianus. Examination oi the plumage of coc k pheasants picked at random from Michigan' sgmeas .ut range would show this mixed ancestry; rarely would one find a pheasant typical of any one subspecies. Taxonomists and historians are not completely agreed on the derivation and taxonomic status of the species and subspecies of Phasian us. Intro- ductions of several subSpecieS, freely interbreeding where their ranges overlap, from severl wideSpread areas in Asia, and long confinement and artificial mixing and selection in game farms have thoroughly confused the genetic composition of the game farm birds, from which our wild birds come. further: m 5 Michigan pieasants most nearly resemble colouatu:: commonly, cocks show coloration irdicat’ng no xcllCHi ancestry. Rarely one sees a bird that shows charac- teristics oi vers icolcr. Call Variation Ac pavyirg physical differences are probably be— havioral differences and call variation. Inter-individual Variation in all structure is likely the result of d:if- (3 frience in the structure and control of the vocal appara- tiugiwedeas intra-individual variation probably re ults from muscular control of organs associated with voiaiizctltrn and perhaps from seasonal changw in vyiin eal structure. in 6'7 A certain amount of variation in some calls probably serves no function at all, but some inter—individual vari— ation might function to identify a calling pheasant to other nearby pheasants. Edminster (1954:11) remarked that males guard their harem's broods and respond to alarm notes of these hens. It has also been reported (Delacour, 195lz237) that females answer the ”nuptial call" of only the cock to which they are mated. Such examples indicate that pheasants recognize and reSpond apprOpriately to calls of other individual pheasants. As discussed in the Catalog of Calls, hens gathering up their broods do not always use just one call. The great variability in the use of chick gathering calls leads one to believe that possibly chicks merely recog- nize the voice of their mother and reSpond, by hiding or approaching, to various tones of the hen's calls, rather than to specific forms of calls. On several occasions I made attempts to attract chick: with tape recorded calls of a hen which was not their mother. These calls were broadcast toward cover known to be concealing scattered chicks (their flock calls could be heard), but young were never attracted in close enough to be seen. This cannot be considered conclusive evidence that the chicks recognized that it was not their mother calling because many other factors could have been operating to keep the young from approaching; and in fact in cases 68 when a hen's brood calls were recorded and then played back to her brood in the field, chicks were also never seen approaching. Tape recorder buzz, other strange noises, imprOper volume control, and other factors could nullify the natural effect of a call. Application of Sound Techniques Methods for censusing pheasants are inadequate, and sound techniques offer promise here. The results of the Engham County crowing survey study, although not conclusive during the summer, indicated that it may be possible to induce crowing by broadcasting into the field tape recorded crowing calls. This method has proved successful in elic- iting calls from crows, Corvus brachyrhynchos, (Frirgs and Frings, 1957); songbirds (Oech and Oech, 1960); and chukar partridges, Alectoris graeca (Bohl, l965). Tape recorded crowing might induce more calling from wild males and might also result in more consistent crowing. The precopulatory call discovered in this study also has possibilities for attracting wild males. Playback of irlirtg arid C+ the precOpulatory call of the blue grouse (S Bendell, l966) caused wild males to call and/or attracted them. The definite attraction this precopulatory call has for male pheasants should encourage further work with it in the field. The most promising call for censusing males could be the alarm call. Males defending territories seem /‘ 09 urable to resist arswering a tape recorded crow with the alarm call reSponse already described in the Catalog cf Calls. If, in fact, males invariably approach crowing calls which are heard within their territory, then this reSponse could give good population indices by the number of responding males out of a given number of randomly selected sites for playback of taped crowing. Censusing hens offers greater problems, but Stokes (lgfl) has reported a tidbitting call of the male chukar partridge which attracts the female. The use of the flock call of chicks to attract hen pheasants also warrants an attempt. One technique which has not yet been tried for any game Species is the use of sound spectrographic methods for censusing birds. The differences which exist in call structure of crowing males is sufficient to confidently identify individuals. Tape recording of crows heard ir the field and subsequent spectrographic analysis could reveal the number of crowing males within recordirg dis- >< tance. Further approaches might be in Lircoln lrde methods adapted to sound spectrographic analysis. instead of marked animals, one would have pheasarts whose calls had been recorded in a given area. Subsequent recordings would reveal the proportion of new calls recorded to repeats (recorded during the initial effort). The assump— tions concerning rates of crowing in different males and .2, O the presence of no non-crowing males would, of course, have to be tested. Playback of calls offers good methods for new capture techniques. Tape recordings of singing male woodcocks, ‘T1 T nilohela minor, lured males into a collapsable ret trap in one study (Sheldon, l955). During the breeding season male ringnecks could probably be captured by taking advan- tage of the alarm call response to crowing. Wild males, attracted by tape recorded crowing within their territory, might enter a trap to attack a mounted cock. The female precopulatory call might also be used to attract males. The use of mechanical calls and taped calls to attract and elicit vocalizations has been reported for many groups of animals. The eastern crow (Frings and Frings, 1957), the mule deer, Odocoileus hemionus (Diem, 1954); the coyote, CaniS latrans (Alcorn, 19M6); the bullfrog, Eana C+ 0 —T1 ”D ca 9. f .;_iana (Capranica, 1965), and many other species, representing diverse groups of animals, have been studied. The success of the use of calls to study these animals should encourage more research with this technique for understanding the behavior of the gallinaceous birds. Fri:- ciples and techniques used with unrelated species are ofter applicable for use with the ring-necked pheasant. SUMMARY AND CONCLUS 0N8 Study Methods The combination of field and pen studies is a neces- sity in a project of this type since some calls are very seldom observed in the wild and can be heard in reasonable numbers only with birds held captive ard easily studied. Unfortunately,with both field and pen research the very presence of an observer often modifies pheasant behavior and makes difficult the interpretation of calls. Pen studies also suffer from the lack of completely natural stimuli. The ease and efficiency of pen studies must be taken into account, though, eSpecially since field work is invariably made difficult because of dense cover, pheasant mobility, and uncontrolled variables such as weather, human or animal interference, and background noises. The transport of cumbersome recording equipment is also a field problem, but to accurately d scribe the mearirg of a call it must eventually be observed in the field under ratural conditions. Since only late spring and summer observations were made in this study, probably not all calls were discoverei, and those calls recorded are still not entirely understood. |Fl Pheasant Vocalizations Ring—necked pheasants possess a surprising range of calls to meet their social and individual needs. No attempt should be made to oversimplify vocalizations since there exists considerable complexity in calling behavior. However, the great variability in pheasant calling behavior should not discourage attempts to draw general conclusions from available data. Genetic variation, which is great, helps to explain some of the behavioral differences observed in pheasants. Table 1 summarizes the important facts about the best understood pheasant calls, but this list must not be considered complete since other calls were heard but were not thought well enough understood to include in the table. Application of Sound Techniques Since pheasants often reSpond in fairly characteristic ways to certain calls, playback of tape recorded calls into the field may prove helpful in censusing, in trapping, and in studying the behavior of ring-necked pheasants. Taped calls can evoke calling in wild birds and can often attract them to the sound source. The breeding calls seem most promising for attracting pheasants and for elicitirg calls since the behavioral reSponses to these calls are strong and are least variable. The crow call, precopulatory call, and male courting calls stimulate Specific responses from other pheasants, and studies of other gallinaceous '73 .TABLE l.--Summary of ring-necked vocalizations. \ Possible Function and Call Age Sex 'Probable Stimulus Behavioral Response Crow Hormonal (?), loud Establish territory, noises, crowing attract hens (?) Subcrow . Hormonal (?), same call Subcall to crowing ' by others Alarm Approach of a man or Warning signal, contact animal, alarm call of call, escape lothers £4) Flight m Approach to within a Warning signal, escape 2 few feet by man or animal by flight Hiss Courting situation, Stimulate courtship in copulation, intimidation female, alarm Antagonistic w Antagonistic, confron— Challenge signal, fl tation with another male attack other male :3 Peeked 3 Pecked Pain, momentary retreat Pecked Pecked Pain, momentary retreat Precopulatory Hormonal (?), Receptivity toward male, courting activity attract male Flight 3 Approach to within a Escape by flight, alert 3 few feet by man or animal young to possible danger . Q) m Brood Gathering Separation from brood Attract scattered chicks 'Brood Caution Possibly dangerous Alert chicks to danger sound or sight [Content Warmth, food, companion- Contact call ship with other chicks ' Caution m Confrontation with Approach to object, fi fi strange object, aggressive behavior E o antagonism toward toward object or o m another chick another chick Flock Isolation, hunger, Regroup brood, cold attract hen birds reveal that such calls are practical as aids to field study. Areas of Possible Research The ontogeny of calls is not well understood. There is no information on the variation in the process of call acquisition or on the innate and learned aspects of vocal— izations. Studies following particular birds from the egg to the adult are necessary here. Additional work is needed to complete the catalog of calls. Many calls have not been reported, while others, which are rarely observed, lack an adequate understanding of their behavioral significance in the vocabulary of the pheasant. There is also not enough known about the seasonal variations in calling behavior. The true significance of intra-individual and inter- individual variations in calls is uncertain. The possitle functions such differences may serve in the activities of pheasants deserves further investigation. Little is krown about the extent to which pheasants recognize and respond to individual calls of other pheasants. Essentially nothing is known of the physiology in- volved as stimuli for calls. Hormonal factors certainly play some role in the establishment of the breeding calls and perhaps others. Much work is needed to develop practical uses of calls to census, trap, and study pheasant behavior in the field. Sound techniques could help solve problems in these areas. LITERATURE CITED Alcorn, J. R. 1946. On the decoying of coyotes. J. Mamm. 27(2):122—126. Allen, D. L. 1956. Pheasants in North America. The Stackpole Company, Harrisburg; and Wildlife Manage- ment Institute, Washington. xv + 490. Ball, K. E. 1950. Breeding behavior of the ring-necked pheasant on Pelee Island, Ontario. Canadian Field— Nat. 64(6):201-207. Beebe, W. 1926. Pheasants, their lives and homes. Doubleday, Page and Company, Garden City. xv + 309. Bent, A. C. 1932. Life histories of North American gallinaceous birds. Dover Publications, Inc., New York. xi + 490. Bohl, W. H. 1965. Experiments in locating wild chukar partridges by use of recorded calls. J. Wildl. Mgmt. 20(1):83-85. Capranica, R. R. 1965. The evoked vocal reSponse of the bullfrog: a study in communication by sound. Research Monograph No. 33. The M.I.T. Press, Cam- bridge. x + 110. Collias, N. and M. Joos. 1953. The spectrographic analysis of sound signals of the domestic fowl. Behavior 5(3): 175—188. Davis, L. I. 1964. Biological acoustics and the use of the sound spectrograph. The Southwestern Nat. 9(3):118-145. Delacour, J. 1951. The pheasants of the world. Country Life Limited, London; Charles Scribner's Sons, New York. 347 pp. Diem, K. L. 1954. Use of a deer call as a means of locating deer fawns. J. Wildl. Mgmt. 18(4):537-538. Edminster, F. C. 1954. American game birds. Charles Scribner's Sons, New York. xviii + 490. 75 76 Ellis, C. R. and A. W. Stokes. 1966. Vocalizations and behavior in captive Gambel Quail. Condor 68(1):72-80. Frings, H. and M. Frings. 1957. Recorded calls of the eastern crow as attractants and repellants. J. Wildl. Mgmt. 21(1):9l. Gates, J. M. 1966. Crowing counts as indices to cock pheasant po ulations in Wisconsin. J. Wildl. Mgmt. 30(4):735-7 u. Holcomb, L. C. 1964. Aggressive behavior of hen pheasant while protecting chicks. Wilson Bull. 76(4):380. Kimball, J. W. 1949. The crowing count pheasant census. ‘ J. Wildl. Mgmt. 13(1):lOl-l20. Kozicky, E. L. 1952. Variations in two spring indices of male ring-necked pheasant populations. J. Wildl. Mgmt. 16(4):429—437. Kozlowa, E. V. 1947. On the spring life and breeding habits of the pheasant (Phasianus colchicus) in Tadjikistan. Ibis 89:423:428. Leffingwell, D. J. 1928. The ring—neck pheasant-—its history and habits. Occasional Papers of the Charles R. Conner Museum. The State College of Washington. No. l, 36 pp. MacMullan, R. A. 1960. Michigan pheasant populations. Game Division Report 2277, Michigan Dept. Cons., Lansing. xiii + 169. McClure, H. E. 1944. Censusing pheasants by detonations. J. Wildl. Mgmt. 8(1):6l—65. Morejohn, G. V. and R. E. Genelly. 1961. Plumage dif- ferentiation of normal and sex-anomalous ring—necked pheasants in response to synthetic hormone implants. Condor 63(2):lOl-110. Oech, V. and L. Oech. 1960. The use of recorded bird songs as a bird—censusing technique. Flicker 32(2): 46-47. Ruffing, E. J. 1952. An intensive study of the crowing behavior of the ring-necked pheasant. M.S., Ohio State University. xi + 103. Sheldon, W. G. 1955. Methods of trapping woodcocks on their breeding grounds. J. Wildl. Mgmt. l9(l):lO9- 115. *3 *J Stewart, P. A. 1955. An audibility curve for two ring— necked pheasants. Ohio J. Sci. 55:122—125. Stirling, 1., and J. F. Bendell. 1966. Census of the blue grouse with recorded calls of a female. J. Wildl. Mgmt. 30(1):184-187. Stokes, A. w. 1961. Voice and behavior of the chukar partridge. Condor 63(2):1ll—l27. FOR SUPPLEMENTARY MATERIAL TO ACCOMPANY THIS BOOK PLEASE iNQUIRE |N AUDIO LIBRARY HICHIGRN STRTE UNIV. LIBRQRIES 31293101601023