Condor, 80:49-63 @ The Cooper Ornithological Society 1978

VOCALIZATIONS OF SCALED

WALTER L. ANDERSON

The Scaled Quail ( squamta) is ( Wallmo 1956). Sex could often be distinguished in typically a of the desert-grassland and the field during soring and summer bv the color of the head and &at; fall and winter could be Chihuahuan Desert associations. Its range sexed with reasonable certainty only when in the extends from southeastern through hand. much of New , southeastern , With one- and two-entrance Stoddard quail traps western and , and across baited with milo grain, I captured, banded, and Mexico in a broad band to (Johns- back-tagged 72 Scaled Quail at the primary site. The color-coded vinyl backtags were attached by means gard 1973). of elastic loops that I could easily slip over the wings. Several significant studies of Scaled Quail These tags did not interfere with normal running or ecology have been conducted (Wallmo 1957, flight, and allowed recognition of individuals at a Schemnitz 1961, Hoffman 1965, Campbell distance. No losses were caused by capturing and handling, and I saw some tagged birds as long as et al. 1973), but little has been written on eight months after release. the vocal signals and social behavior of the I used a Sony TCSOOB tape recorder and a Dan (Johnsgard 1973, Hatch 1975). De- Gibson electronic parabolic microphone for all field tailed analyses of vocalizations of closely re- recordings (tape speed 7% ips). I also recorded two three-day-old chicks in the lab. I analyzed calls lated species in captivity have been performed on a Kay Electric Company Sonograph, model 6061B, by Williams (1969) for California Quail using the FL-1 circuit and the wide-band filter (Lophortyx californicus), by Ellis and Stokes setting. The audiospectrograms reproduced here were ( 1966) for Gambels’ Quail (L. gamhelii), judged typical by visual inspection. and by Stokes (1967) for Bobwhite (Colinus virginianus). RESULTS This study is based on observations and Scaled Quail are monogamous, with sexes tape-recordings of free-living birds. I intend similar in size and (Wallmo 1956). to describe the repertoire of vocal signals of Both sexes are reproductively mature by the Scaled Quail and discuss each signal in its breeding season following their first winter. selective context. This study may lay the Primary sex ratio approximates 1: 1, but dif- framework for research to identify more com- ferential female mortality results in a male- pletely the interacting forces shaping each biased adult population (Campbell et al. 1973, behavior. Hickey 1955). D uring the early breeding period, of variable length and timing, the METHODS population disperses as pairs and single males; occasionally remnant groups persist when I conducted field work from June 1972 through No- vember 1973 and again on 14-16 May 1974, mainly breeding conditions are unfavorable. in three contiguous square-miles of desert grassland Both parents attend the young. Broods pro- just East of Cochise Stronghold (Dragoon Mountains) vide the nuclei for covey formation, consisting in Cochise County, southeastern Arizona. The 150 of brood mergers or the addition of unsuccess- davs of field work samoled all seasons but emnha- sized summer periods. Within the “Sunsites” devel- ful pairs and unmated males. Essentially all opment of the Horizon Corporation, the site was Scaled Quail were found in coveys of three or subdivided by dirt roads into a uniform grid of 96 more (up to 40) individuals from September blocks, each one-fourth by one-eighth mile. The through March (Anderson 1974). study area lacked buildings, was free from grazing, This brief summary of the annual cycle and received little disturbance. The grid of roads aided both accessibility and observation of the birds suggests the types of interactions that may and permitted me to pinpoint locations of sightings. occur, which in turn influence the evolution Supplemental observations were made at a secondary of social signals. For the call descriptions and site, a waterhole between Oracle and Oracle Junc- interpretations which follow, my data base for tion, Pinal County, Arizona and on four wild quail that I kept as captives from January through August adult vocalizations is shown in Table 1. Each 1973. number shows the separate occasions on which I surveyed quail on systematic transects (10 mi a given call type was used. It does not count per survey) on the Cochise County site. The vehicle served as a blind but I approached on foot if neces- repetitions of a given call; thus the 312 sary to obtain a complete count by flushing a covey. shriek encounters actually represent thousands I used a windmill as an observation tower and of the calls heard. In a very general way, the crawled through the grass to obtain behavioral data frequency distribution indicates the relative on undisturbed birds. I determined sex and age of quail by plumage commonness of a call, but the sample is not

f491 50 WALTER L. ANDERSON

TABLE 1. Minimum sample sizes for behaviors described in this paper. (An encounter with a bird scored one observation for each behavior given by that bird, even if the behavior was repeated.)

Sex Breeding status

Male Unmated 79 10 17 112 Male Mated, no brood 17 3 1 15 13 17 19 Female Mated, no brood 20 6 4 13 Unknown Mated, no brood 4 3 6 2 1 7 Male Mated, with brood : 16 4 11 Female Mated, with brood 7 1 22 Unknown Mated, with brood 12 3 1 12 2 3 1 : Unknown Non-breeding; in covey 114 27 12 117 2+ 2 11 4 11 Unknown Unknown 150+ 10 1 21 18 zoo+ 1

Minimum Totals 416 54 22 232 r 114 312 23 48

* squeals broken down by context: a. another quail or pair of quail moving closer to caller b. another (not Scaled Quail) approaching caller c. context not determined.

unbiased. Loud or conspicuous calls (e.g. second syllable is somewhat longer than the chekar, shriek) tended to be heard more first. easily than soft calls (e.g. CHUPuh, cut cut). A calling quail tipped its head up slightly, My presence often disturbed the birds, prob- opened its bill and pumped its tail on each ably increasing the use of tsing. syllable. Eyes remained open. Quail gave the The table stratifies the calls by the sex and call while standing or walking on the ground breeding status of the caller. Obviously, mated or from an elevated perch. males without broods account for most of Figure 1 is a series of audiospectrograms the tidbitting acts. Males and females showed representing portions of a continuous sequence no differences in the use of chekar or tsing. of chekars that lasted nearly 15 min. The calls Where sex determination was not possible, were made by a hen separated from her mate, some inferences are yet feasible. For example, who flushed when my vehicle stopped. when I watched mated pairs without broods, Initially the hen gave two strong series of I identified males as the callers 24 times when chekars, then reverted to low volume calls the squeal vocalization was given, females (part A), apparently inhibited by the ap- never; for the 10 cases in which I heard pearance of the parabolic microphone. At squeals from one member of a mated pair but low volume, these calls sounded more like did not actually see the caller, it would be safe to assume the caller was the male. In discussing probable functions of each call, I will omit reference to selection for spe- cies-distinctiveness. Four species of quail- Scaled, Gambels,’ California, and Elegant ( Callipepla [ Lophortyx] douglasii)-are con- sidered by some authorities to be congeneric (Johnsgard 1973, Hubbard 1974). It is not known to what extent selection for divergence or convergence has occurred. I will point out homologies and differences where these may be instructive.

THE CHEKAR (SEPARATION) CALL A distinctive two-syllable chekar (Peterson 1961) was often given by individuals of either sex when separated from other quail (Table

1) , Wallmo ( 1957) described it as chin-tang FIGURE 1. Chekar calls given by a female sepa- or chuk-thing, while Johnsgard (1973) in- rated from her mate: A, low intensity calls; B and C, terpreted it as pay-cod. In all cases, the high intensity calls. SCALED QUAIL VOCALIZATIONS 51 repetitions of wip-woo than like chekar. All sequences were initiated by a wip syllable, though either a wip or a woo syllable could have terminated the sequence (Fig. 1A). After 11 sequences of from one to three wip-woo repetitions each, the quail graded into strong chekar series of three to seven repetitions each (Fig. 1B and C). Although spacing of notes was comparable at both volume extremes, the chekars sounded qualita- tively different, because of the increase in overtones. At highest intensity, an uncommon variant appeared (the third, sixth, and ninth syllables in Fig. lB), a sneezy chip that fol- lowed the normal chekar. Shortly after a covey was flushed, one or more quail started the chekar call. The separated birds usually ran or flew toward the caller, even when this meant exposing FIGURE 2. Chekar calls by five different unmated their locations to a potential ground predator. quail. A calling bird often moved toward other birds whom it could see or hear. A quail flying are known to be capable of recognizing their toward a caller frequently overshot or veered mates’ call. away from the actual position of the caller, Unmated males gave series of chekars in- and the birds converged on the ground. Call- terspcrsed among shrieks (described later) ing generally ceased before or shortly after advertising their single status. Five different the covey was fully reassembled. Birds rarely unmated males gave the chekars shown in Fig. called when a flushed covey landed together. 2. These calls were usually given at full in- Sumner (1935) found the tendency to answer tensity. They varied from one to eleven so strong in California that a mechan- chekars per series, averaging six to eight. Each ical imitation of their separation call in the series was separated from the next series or winter often induced return calling by an from a shriek by pauses of 2 to 15 s, and no undisturbed covey. predictable relationship between shrieks and When different birds alternated calls during chekars was evident. reassembly, there were no predictable patterns of exchange. Nor was there adequate pre- CHUPuh cision in the responses of separated mates to suggest antiphonal calling of the form de- Figure 3 demonstrates a low-volume vocaliza- scribed for the Bobwhite (Stokes and Wil- tion given by quail when seeking escape or liams 1968). when re-contacting associates after distur- Chekar calls reassemble covey members bance. It appears to coincide with the “group before dusk if they have scattered while feed- alarm cry” or chink-thank-a’ described by ing. Scaled Quail typically roost on or near Wallmo ( 1957) as being similar to the sepa- the ground in a small compact group, often a ration call but more rapid and with slightly part of the larger covey (Russell 1932). It is different syllabication. It somewhat resembles not known whether or not members of a roost- the wip-woolchekar type of call (Fig. l), but ing party are kin if the covey is comprised of differs from these by lacking the regular in- several families. If separated at night, the tervals of syllables and by the usual crowding quail remain where they land and do not of two syllables (CHUPuh) into the space of assemble until daylight (Wallmo 1957). about 0.2 s. Figure 3C begins with a sequence An incubating female leaving the nest for which sounds like CHUPuh CHUPuh CHU- UP?, with a slightly rising inflection on the either of her daily feeding periods gave the last note. Unlike chekars, in which both chekar call when she was some distance from syllables are equally stressed (though the the nest. The male flew or ran to join his second syllable is longer), the CHUPuh calls mate, accompanying her as she ate, though distinctly emphasize the first syllable. Figure seldom feeding with her. California Quail 3D shows unusual variants of the call. (Williams 1969), Gambels’ Quail (Ellis and Captive birds gave this call while frantically Stokes 1966), and Bobwhite (Stokes 1967) searching for an exit from the pen. If ap- 52 WALTER L. ANDERSON

FIGURE 3. CHUPuh calls by captive quail seeking exit from pen. FIGURE 4. Tsing calls by six different adults in contexts of alarm. proached too closely, a quail stood erect and uttered a tsing note. It then returned to a tsing calls. Each is characterized by a quick, head-low, active search for escape, oriented broad frequency pulse which becomes a high away from the disturbance. frequency descending slur, ending imprecisely. In the field I heard this low-volume call Several show a peculiar splitting of harmonic only 44 times, much less frequently than I bands (Fig. 4A and 4D), suggesting dual heard the chekur or tsing (Table 1) . It control of sound production within the syrinx typically occurred when quail resumed con- (Greenewalt 1968). tact after having been scattered by a distur- The usual stimulus eliciting the tsing call bance or when a female approached her mate appeared to be the sight of potential danger who was giving agonistic squeal calls (dis- approaching on the ground. Frequently cussed later). avoidance followed, particularly if the quail was further approached. Cattle or deer ap- CONTACT NOTES proaching quail were as effective in eliciting the call as were and humans. If the Wallmo (1957) reported “conversation notes,” potential danger did not approach, the calling soft clucks and chirps within an undisturbed quail might resume feeding or drinking, but group or pair. I noted such calls on 22 oc- seemed to keep the intruder in view and re- casions (Table 1) , but since they were audible peated the tsing call occasionally. at just a few meters, the calls may have been Trapped birds gave the call when ap- more common. Gambels’ Quail in my study proached closely and often when handled. area appeared to give similar notes much At all times the calling bird kept its body more frequently. erect, its crest raised, and its eyes oriented Because the calls were soft and given only toward the alarming stimulus. Between calls by undisturbed birds, I was unable to record the bird often flicked its erect crest, which is them. It is possible that they may coincide usually depressed when the bird is relaxed. with wip-woos (low intensity form of Other birds within hearing of the calling chekars) . bird responded with alertness and avoided the stimulus if they could see it. The TSING homologous call in California Quail, psezc, This call (spelled tsinggg by Wallmo 1957) “will often throw a dozen covered cartons full was given by adults and larger juveniles of of banded quail into a pandemonium of both sexes ( Table 1) . The calling bird jerked scuffling and scrambling” (Sumner 19X5:205). its head slightly with each utterance, with the He also found some quail responding to the eyes open. Most calls were given by birds call by approaching in a manner that he in- standing or running, but flushed birds often terpreted as a form of mobbing behavior gave them at the beginning of flight. which might distract the predator. Alterna- Figure 4 displays six audiospectrograms of tively, the responding birds may have been SCALED QUAIL VOCALIZATIONS 53

FIGURE 5. Male Scaled Quail (right) perched as sentinel while the hen and chicks feed below. The erect crests indicate that the quail have been alerted. If approached by a ground predator, the male may give the tsing call and jump to the ground to run or fly with the brood. approaching to learn the predators’ charac- by a hawk. Ellis and Stokes (1966) were teristics ( Kruuk 1976). unable to elicit such a call in captive Gam- Tsing calls occurred most predictably when bels’ Quail, but Williams (1969) found that adults with a young brood were disturbed. California Quail frequently reacted to a sud- Often the adult male, perched above the den overhead stimulus . ground as a sentinel, gave the tsing as he Scaled Quail typically responded to an jumped down to flush with the family (Fig. avian predator by freezing motionless or by 5). In about half the winter-coveys flushed, ducking quickly into heavy cover. On 22 one or more birds gave the call. Even single October 1973, a Coopers’ Hawk (Accipiter males gave the note when alarmed at close cooperii) pursued several quail deep into the range, generally concurrent with escape be- cover of low mesquite, causing the quail to havior. Agonistic encounters with other males burst out in many directions. Some uttered sometimes were followed by tsing notes, per- tsing notes, but two or more gave intense haps accompanying an avoidance tendency. TiCHUNK calls. All flew but a short way, dropping into heavy cover. The hawk re- TiCHUNK mained motionless a few seconds, then tried Although the presence of an “aerial alarm re- unsuccessfully to relocate the scattered quail. sponse” has been noted for many gahiforms (Johnsgard 1973), I found no recent refer- SQUEAL (HEAD-THROW) ences to its existence in Scaled Quail. Bendire The squeal vocalization is a high intensity (1892) offered a second-hand report of a call, initiated as a broad-spectrum of noise gutteral oom-oom-oom given by a quail chased that becomes an ascending complex of har- 54 WALTER L. ANDERSON

gave the call when confronted with another male (or with his own image in a mirror, Johnsgard 1973). If avoidance did not occur immediately, other agonistic gestures (low aggressive rushes or actual fighting) followed, interrupted occasionally by bouts of squealing. Playback of recorded squeals induced more squeals from nearby single males, but mated males generally responded only if another male was in sight. Williams ( 1969) noted but one instance in three years of a squill call by a female California Quail; Ellis and Stokes (1966) detected no meah calls by hen Gambels’ Quail. I heard squealing by wild female Scaled Quail in only 2 of 57 instances in which the caller was seen (Table 1). On 28 June 1973, two pairs arrived at a waterhole simultaneous- FIGURE 6. Squeal calls. A, the same call analyzed ly. One male squealed; the other male at both 8 kHz and 16 kHz scales; B and C, two se- squealed back and moved away, only to en- quences of calls by other males; and D, male squeals and the softer CHUPuh calls of the mate. counter a female Scaled Quail with three small chicks. The male squealed and rushed at the female who squealed back. The male monies (Fig. 6). The quail tosses its head stopped and both birds bobbed their heads upward and backward vigorously, opening the up and down rapidly, but no fighting oc- bill and closing the eyes (Fig. 7). It repeats curred. On 29 June 1973, a pair was flushed the call up to eight times, each repetition 0.4 from chicks still unable to fly. The parents to 0.8 s apart. circled, giving tsing and chekar calls. After A male in breeding condition predictably considerable disturbance, the female gave two

FIGURE 7. Postures of a mated pair of Scaled Q ual1‘ w h en disturbed by another male or a mildly alarm- ing stimulus. The male (right) gives squealsand the female may give CHUPuh calls. SCALED QUAIL VOCALIZATIONS 55

FIGURE 8. Comparison of male shrieks on both the FIGURE 9. Advertisement calls of male quail: A-I, 8 kHz and 16 kHz scales. shrieks by male Scaled Quail; and J, kaa call by male Cambels’ Quail. squeals, then resumed the separation and alarm calls. SHRIEK Figure 6C shows that a male quail may I found the shriek to be the most variable of interject a short syllable between squeals, the major Scaled Quail calIs. This variability particularly in the later parts of a series. These may account for the variety of names applied interjections are given with the head at nor- to it: tree in (Russell 1932), mal level between the violent head-throws. squawk or kzooc1cin Texas ( Wallmo 1957), They appear to be a variant of the CHUPuh and tL;hock in Oklahoma (Schemnitz 1961). call. Figures 8 and 9 illustrate some of the varia- Though primarily a male call, squeals also tion of the call. Figure 8 gives three examples were given outside the breeding season as they appear when analyzed on both the (Table 1) . Occasionally the apparent stimulus 8 kHz and 16 kHz scales. Most whistles last was not another Scaled Quail; I observed 0.2 to 0.3 s but differ greatly in the number birds squeal when approached by humans, and arrangement of harmonics, as well as in cattle, deer, Gambels’ Quail, and even a Log- purity of tone. Figure 9J shows the homol- gerhead Shrike (Lanius Zudovicianus) . Johns- ogous call of a Gambels’ Quail which called gard (1973) reported squeals given by alternately in a long, unsynchronized series retreating quail, with some individuals also with a male Scaled Quail, whose call is giving a chip ’ or chip-eee,’ perhaps variants shown in 91. of the tsing or CHUPuh notes. In some cases Shrieks were given only by males, presum- the quail may be directing the call toward the ably by unmated ones (Table 1). Intervals intruder, demonstrating location and possibly between shrieks in a sample of 77 calls varied aggressiveness so that a large non-predator from 3 to 20 s, averaging 8.2 s. Periodically, a (deer, cattle) may avoid approaching too calling male interjected a series of chekar closely. Squeals from individuals in a covey calls, then resumed the shrieks. If slightly landing close together may be directed toward inhibited, the male gave soft pips before flockmates who have ahghted too near for the resuming shrieks. comfort or safety of the caller. With each shriek, the quail tilted its head Figure 6D presents further evidence of the up, closed its eyes, and pumped its tail alarm context in which the squeal may appear. slightly. On chekars, the bill was tilted just above the horizontal and the eyes remained As cattle approached, the male of a pair gave open. squeals while his mate uttered a series of Single quail typically called from a perch CHUPuhs (Fig. 7). Unlike the uniform (Table 2). Only four of 107 observations in- CHUP interjection occasionally given by volved calling from the ground, and one of males, the females’ calls were not synchronized those quail used a gravel dike to raise its with the pattern of the males’ squeals. effective height to 1.2 m. Choice of perch 56 WALTER L. ANDERSON

TABLE 2. Average height of calling perches chosen quickly after giving chekar calls. At no time by unmated male Scaled Quail. did RWY give the shriek call characteristic of unmated males. On 19 and 20 May, the Type of perch No. cf observations Av. height (m) female was gone and male RWYs’ behavior stump -17 0.8 was totally different from that of the previous Yucca stalk 41 2.1 Mesquite 40 1.5 month. He gave repeated series of shrieks and Sign or fence post 5 1.6 chekars, challenged and fought several un- Ground .4 0.3 mated males, and wandered extensively. Ap-

Total 107 1.5 parently this male (at least two years old, since I banded him as an adult the previous fall) succeeded in acquiring a new mate, for on 25 May and on five more occasions in the usually reflected availability. Sites with good next month, I saw him with a female, and the clearance on all sides were usually chosen, shriek was no longer used. but the quail did not always select the highest A second case of a mated male assuming available perch. the behavior of an unmated bird was set up In 1973 calling began in April, coincident when I trapped a hen but not her mate. The with the breakup of coveys. Both Wallmo male gave many chekars, but the female was (1957) and Schemnitz (1961) found similar inhibited from calling by her fear of the trap initiation in their populations. Frequent call- and me. After one-half hour of separation, ing in both morning and evening periods the male gave a shriek, then more chekars, continued until broods appeared.’ Calling then then several more shrieks. I released the fe- declined noticeably, particularly in the after- male, who flew toward the male, and each noons. By early July, few shrieks were heard. gave soft chekar calls as they ran together. On 18 July, however, I found a significant revival of calling, apparently in response to TIDBITTING the heavy rains the preceding week. These rains may have flooded some nests, forcing During the breeding season I often observed females to re-nest, or may simply have im- a male Scaled Quail attract his mate to him proved the nutritional status of all birds. I when he found a choice food item, particularly heard no shrieks after 8 August 1973 and none an insect. Typically the male faced toward at all in August 1972. the food, pecked incompletely at it, and During the breeding season, playback of uttered a soft cut cut food call. This behavior recorded chekur calls of female Scaled Quail in chickens was called “tidbitting” by Domm stimulated increased calling (both chekars (1927). Stokes and Williams (1971, 1972) and shrieks) by unmated males but elicited described and illustrated the display and call no response from either sex of a mated pair as found in galliforms. (several dozen tests). When I flushed a pair The most common releaser for this behavior with young, the females’ chekars were also on my study area was a small moth. A male likely to induce calling by nearby males. My quail usually pecked it once or twice to sub- findings concur with those of Levy et al. due it, but would not consume it if his mate (1966) that single males tend to approach was near. Only once did I see a male present cheknr calls independent of any visual stimu- the food with his bill’ as is more commonly lus. done by the Bobwhite (Williams et al. 1968). Williams ( 1969) found that administering A female generally responded immediately testosterone proprionate daily to male Cali- to the call by running to the male and con- fornia Quail produced aggressiveness and cow suming the food. The homologous food call calling, behavior characteristic of unmated in Red Jungle Fowl (Gallus gallus) is easy males in the breeding season. Male Scaled to locate, being brief, repetitive, and covering Quail giving shrieks are presumably in repro- a broad frequency range (Stokes 1971) . The ductive condition and capable of breeding. sudden, accurate dashes made by a female The presence of a mate may inhibit the shriek quail out of sight from the male suggest that vocalization in males who may otherwise re- she is able to determine directionality from tain reproductive capabilities. Two examples the sound. When a mated females’ foraging illustrate this. brought her to a moth or other item suitable Marked male RWY was observed with his for the tidbitting display, she ate it immedi- mate for long periods on seven days from 18 ately. April through 18 May 1973. During this time Either parent may tidbit to the young, but I occasionally flushed the pair, who re-united less commonly than for males to females dur- SCALED QUAIL VOCALIZATIONS 57 ing courtship (Table 1; also Williams et al. 1968). Occasionally during the winter, a quail would discover a large or attractive food source and give the cut cut call. Nearby quail would immediately converge upon the calling bird and feed rapidly; often six to ten birds would be in contact at once.

OTHER POSSIBLE SEXUAL CALLS Neither this study nor Johnsgard (1973) found any calling by Scaled Quail during copulation. Both male and female Gambels’ Quail (Ellis and Stokes 1966) and California Quail (Williams 1969) and the female Bob- white (Stokes 1967) occasionally call during copulation. I had no opportunity to observe nest- building behavior in Scaled Quail. Williams FIGURE 10. Calls by juvenile quail: A, cherk calls (1969) described’ a call and associated be- of three-day-old chick; B, peep calls of three-day- havior which accompanied nest-building in old chick; C, low intensity peter calls superimposed California Quail. by tsing of adult; D and E, peter calls of older chicks.

CHICK VOCALIZATIONS Figure 10A illustrates the call of a captive domestic chicks and their hen. Peep peep three-day-old Scaled Quail chick; Wallmo calls stimulated clucking by the hen, which (1957) referred to the call as cherking. It in turn inhibited peeping in the chick, partic- consists of short notes with ascending fre- ularly if they came into contact. quencies, usually terminated by a small The peep peep call of very young chicks descending leg. The homologous call in the soon sounded more like peter peter. The num- domestic chick, “pleasure notes” (see audio- ber of notes per call series decreased from ten spectrogram in Collias and Joos 1953) or or twelve in the first week to series of six (Fig. twitter (Andrew 1964), is extremely similar lOD), then to five (Fig. lOE), and the funda- in pattern. It also bears strong structural mental frequency became lower. These trends similarity to the pipping and twitter calls continued until development of the adult (possibly variants of the same call) of the chekar call with two syllables and many over- young California Quail (Williams 1969). The tones (Figs. 1 and 3). Call configurations cherk was given when a chick was in the suggest that the CHUPuh call is derived from presence of other quail, free from disturbance, the same juvenile call. and not hungry. Figure 1OC shows four peter syllables over When separated from its companions, a which an adult tsing is superimposed. Though young chick emitted lengthy series of peeps. simple in structure, the calls were made by Figure 10B shows the peep peep call of the the older chicks who also gave the calls in same chick that gave the cherks in 10A. Young 10E. Just as occurs in adult chekars, inhibited chicks gave four to five peeps per s, up to chicks call at a lower volume in which twelve peeps per series. Each series was harmonics drop out and overall structure is separated by a pause of three to six s. The simplified. basic form spectrographically is that of a I heard no tsing alarm calls in juvenile quail short inverted “U”. until they were at least half-grown. Wallmo Figure lOC-E shows field recordings of (1957) first heard it in chicks seven weeks chicks separated from their parents. The call old. It was similar to the adult call but lacked is clearly analogous to that already discussed the “metallic ring.” for separation calls (chekurs) of the adults. As such it corresponds to the “lost call” of DISCUSSION Bobwhite chicks (Stoddard 1931) and of Visual signals are little used in Scaled Quail California Quail chicks (Sumner 1935). Call- except for short-range purposes, such as be- ing by both chicks and adults continued until tween mates or rival males. The similarity of reassembly occurred. Collias (1952) found a plumage of both sexes suggests selection for reciprocal relationship between calling of crypticity in Scaled Quail to reduce vulner- 58 WALTER L. ANDERSON

TABLE 3. Synopsis of Scaled Quail vocalizations. ~ .~ ria1ne Sex of call calline Context Probable function Probable information content chekar Both Visual isolation from Reassembly of sepa- Identifies caller, gives one or more regular as- rated birds-pair, location of caller, and sociates; chekar from brood, or covey. shows callers’ motivation another quail Proximity important for for reassembly. If call is reproduction, parental subdued, may indicate investment, and self- a threat is still nearby protection respectively

Unmated bird early in Long-range advertise- Gives callers’ location and breeding season ment for mate willingness to seek contact

CHUPuh Both Approach of another Communicating low in- Identifies caller as non- animal, potentially dan- tensity alarm and pre- threatening flockmate gerous; reassembly after venting associates from alarm avoiding the caller

Female SqueaZ of her mate Preventing male from Identifies caller as mate; attacking or avoiding non-threatening the female contact Both Nearby presence of Maintenance of contact Indicates caller is close notes associates in nonsexual with mate or group by and undisturbed; and non-threatening perhaps encourages reci- situation procity by flockmates, without which the caller will cease calling and be- come disturbed cherk Both Chick in presence of Maintenance of contact Same as adult contact family, unthreatened, with family notes not hungry mwl ’ Both Chick separated from Reassembly of chick Indicates chick is dis- siblings or parents; with usual companions; tressed; same informa- chick cold, wet, or brings parental care tion as adult chekars hungry when needed tsing Both Approach of ground Alerting kin or flock- Indicates caller is predator; capture mates of danger alarmed; posture and avoidance behavior of caller may indicate direction from which danger approaches

TiCHUNK Both (?) Extreme threat, such Predator confusion; re- Perhaps deceives and as hawk attack duce likelihood of cap- startles the attacker ture by increasing by intense reaction reaction time of predator squeal Mostly Male-male encounter Distance-increaser; re- Conveys degree to which male during breeding season duction of actual caller is motivated fighting aggressively; reflects dominance relationships

Chekar calls of hen Inhibition of responsive Identifies caller as aggres- mated to the calling calling by unmated males sive mate of the female male giving chekars

Disturbance by Avoidance of further Alerts intruder to loca- non-predator conflict tion and aggressive motivation of caller shriek Male Lack of mate during Advertisement of sex Identifies caller as breeding season and single status to sexually-motivated, potential mates unmated male

Responsive chekar calls Determination of status Same as above; chal- from unknown bird of other bird lenges bird giving chekars to identify its own sex SCALED QUAIL VOCALIZATIONS 59

TABLE 3. (Continued). _ NEUIE Sex of call callina context Probable function Probable information content cut cut Male Encounter with novel Attraction of female; Shows female a desired food item, female enticement leading to food item; signifies po- nearby copulation; strengthen- tential fitness of male ing of pair bond

Both Novel or concentrated Parental investment as Gives location of caller food, quail nearby functional feeding of and a food source young; attraction of companions around caller as self-protection

ability to diurnal predators hunting by vision. birds. Though the table is nearly complete Short-range signals such as the flicking of in itself, a few comments may be useful. the crest, standing erect with slicked feathers, Both sexes share nearly all calls, consistent and escape movements themselves all readily with the similar roles in parental care and the convey fear motivation to other nearby quail long period of flocking. The persistence with without risking undue exposure to a predator. which separated birds give cheknrs until re- Certain calls are used instead to convey in- assembled suggests strong advantages to flock- formation about intensity of the alarm situa- ing, despite the automatic disadvantages to tion. group living (Alexander 1974). Two other Compared to other forms of signals, sounds calls, the CHUPuh and contact notes, may have several advantages: they (1) are rela- assure other conspecific quails of the non- tively economical to produce, (2) decay threatening motivation of the caller in situa- rapidly, thus avoiding interference with fur- tions of alarm and peacefulness respectively. ther transmissions, (3) travel quickly to CHUPuh notes by a hen when her mate is distant points, (4) are relatively independent giving the agonistic squeal call may prevent of physical obstacles (compared to visual him from mistakenly directing his aggression signals), and (5) provide a wide choice of toward her. available frequencies and intensities for sig- The squeal appears homologous to the nal use (Thorpe 1961:9). Final expressions “caterwaul” of the Bobwhite (Stokes 1967, are influenced by the physical and biological Hatch 1975)) the squill of the California Quail environment. ( Sumner 1935), and the meah of the Gambels’ Otte (1974:406) described six basic types Quail (Ellis and Stokes 1966). In Gambels’ of information that may be conveyed in ani- Quail, the menh may reduce the amount of mal signals: (1) deictic, drawing attention actual fighting (Ellis and Stokes 1966). Wil- to an individual or object, (2) identification, liams (1969) placed two malt California specifying age, sex, relationship, individual, Quail in the presence of a female; the sub- species, and so on, (3) spatial, referring to a ordinate gave only four of the 1s squill series location, a direction, or a distance, (4) re- and those only before the birds had estab- sponse level, reflecting the motivation of the lished a final dominance relation. This sug- signaler or the character of some exploitable gests that the squeal and its homologues may resource, (5) temporal, indicating the timing be used to indicate a males’ willingness to of some event, and (6) event, informing on fight. Fighting can be costly to either com- events taking place. Each signal may impart batant, and a signal showing aggressive several types of information. motivation may spare the caller significant Table 3 presents a concise summary of the expense. If both participants behave more Scaled Quail calls, their usual contexts, their agonistically (“calling the bluff”), actual fights likely functions (as in Williams 1966, Alex- may ensue. ander 1974), and their presumed information In encounters between males, a visual content (in the sense of Otte 1974, not as in signal may accompany the vocal one. In information theory). My interpretations of California and Gambels’ quails, the head- the data correspond with evolutionary predic- throw call exposes the conspicuous black tions, but the logical extension of this study throat. The chestnut throat of the Mountain would be an experimental, hypothesis-testing Quail (Oreortyx pictus) and the black and approach using captive birds of known age, white throat of the Bobwhite are also con- sex, and relationship to other experimental spicuous. The throat of the male Scaled Quail 60 WALTER L. ANDERSON is pale buffy to white, but it remains as one fended his mate in relation to her position, of the few distinctive features separating the without persistent topographic reference. sexes (Wallmo 1956). Critical tests on the Activity space overlap of different pairs was possible functions of the throat patch have considerable. yet to be performed. Unmated males in the breeding season On one occasion I flushed the male of a either wandered or remained near mated pair without also forcing the female away. pairs, a tendency documented for several quail The male landed 75 m away near a single species (Emlen 1939, Genelly 1955, Ellis and male. Both males squealed. The hen began Stokes 1966, Johnsgard 1973, Anderson 1974). a chekar series. The mated male flew toward Unlike songs, which may both at- her but overshot her by 40 m. She continued tract females and repel males, three different to cheknr as the male ran toward her, but he calls by Scaled Quail are employed: the stopped three times to give series of squeals chekar, the usual separation call which brings which overpowered the volume of her calls. two birds together; the shriek, a “qualifier” The context and the behavior which resulted which indicates that the caller is both male suggested incipient antiphonal calling (Stokes and unmated; and the squeal, the agonistic and Williams 1968). In the two Lophortyx call directed toward other males. species they studied, the male superimposed My study supported the assertions by Wall- the agonistic call over the separation call of mo ( 1957), Schemnitz ( 1961), and Johnsgard the female, probably to “inhibit responsive (1973) that the shriek call is characteristic of calling by unmated males.” The headthrow unmated males. A male losing his mate (even call, if my observation was not exceptional, for several hours when she is trapped) may may indicate the response level of the male resume the call, presumably from a change (highly aggressive) and indirectly identify in motivation (reverted to the status of an the female as already mated. unmated male). This agrees with Williams The cut cut food call is clearly in the males’ (1969) on California Quail, Stokes (1967) on reproductive interests if the tidbitting display Bobwhite, and Ellis and Stokes (1966) on increases his chances of mating, and provides Gambels’ Quail. D. E. Brown (pers. comm.) the female with protein which may help her elicited shrieks from one paired male Scaled produce a full clutch of . It is in both Quail by using a tape-recorded female call; parents ’ interest as a functional feeding of he also observed another male call as his mate their offspring. The use of the call after incubated nearby. These unusual observations covey formation is less obvious. Kin selection may represent cases where the pair bond is (Hamilton 1964, Maynard Smith 1964) and weak or where a male is responding as if the protection from predation (Hamilton 1971, recorded calls were those of his own mate. Pulliam 1973) are possible nonexclusive Birds uttering shrieks or chekars, especially hypotheses that could be tested. single males, frequently used elevated perches, The tsing may have evolved to communicate a strategy which minimizes sound interference alarm to kin or frequent associates while mini- by vegetation and increases the effective mizing location cues that might attract range of the calls (Ficken and Ficken 1962, predators. As a high, rather pure whistle Morton 1975). Chekars are repetitive, with which terminates imperceptibly, it is remark- many harmonics and relatively low frequen- ably similar (probably by convergence) to cies, attributes that favor long-distance trans- alarm calls of many bird species (Marler mission and ease of locatability (Marler 1957). Though given most frequently when 1957). The period during which single males flockmates (usually relatives) are near, it is use high and conspicuous perches coincides occasionally given by single males or by birds with the season of lowest pressure from aerial in very large coveys. It is possible that the predators (Anderson 1974). call, at least when given as the bird flushes Apparent sentry behavior by quail is re- immediately before a predator, may startle ported widely in the literature. Typically one the latter and increase its reaction time individual flies up on a post, shrub, or tree ( Humphries and Driver 1967). The tsing and and appears to watch alertly as other individ- its homologues in other quail have been uals feed below. My 48 observations of this labeled “distress calls,” but intensive work is behavior are subdivided by the sex and still necessary to determine how, in the sev- breeding status of the sentinel in Table 1. eral contexts within which such calls are Where sex was known, 30 of 32 sentinels given, the signaler itself is favored. were males. Nineteen observations involved The Scaled Quail in this study did not hold males accompanying their mates, primarily as and defend territories. A paired male de- the incubating females left their nests for SCALED QUAIL VOCALIZATIONS 61 short periods of feeding in mornings and not only economically performs the reassem- evenings. Males tended to be nervous and bly function of the loud chekar calls given easily triggered into giving the squeal call. when a covey is widely separated, but may Rarely did they eat as their mates fed, and also restrict the message to an appropriate rarely did the latter look up from eating. recipient. Again this is best interpreted as male re- A study of vocalizations is a necessary productive effort, whether the watchfulness preliminary to the application of any census served to detect predators, rival males, or technique which depends on song or call both. counts. Biologists extrapolate from spring call Adult males exhibit similar alertness and counts of , quail, and doves to attentiveness when with a brood. Sumner predict fall populations, hence probable (1935) found that male California Quail fed hunter harvests. Yet correlational studies may mostly when the female and young rested in mislead on cause-effect interpretations. Most cover. A male usually perched within one-half quail studies, including this one, describe the meter of the ground as the family fed below male announcement call as characteristic of (Fig. 5), changing his sentinel position to unmated males. A prediction of fall popula- keep in close contact with the feeding brood. tions would thus be based on counts of that At the approach of a “threat,” he typically populations’ unproductive members. Varia- jumped to the ground, with or without giving tions in survivorship may create large devia- a tsing note, and ran or flushed with the tions in numbers of unmated males. When brood. call counts and hunter harvests do correlate Quail which perch as sentinels predictably well, it may be because both factors use lower perches (nearly all below 0.5 m) fortuitously correlate with another factor re- than do calling bachelors (average 1.5 m, lated to productivity. For example, higher Table 2). Sentinels have no demand for long- calling rates may reflect better nutritional range signals, and a higher perch would conditions for all quail, which in turn account simply render them and their associates more for higher productivity. This example merely visible to predators. When approached, bach- emphasizes that we still need detailed life clors tended to flush and fly away directly history studies, upon which we may make from the perch; sentinels usually dropped to intelligent decisions in matters which may the ground and flushed only if the brood affect animal populations. flushed. The latter may reflect a males’ com- mitment to defense of mate and young, SUMMARY though he himself may gain directly by using Vocalizations of Scaled Quail were analyzed the others as cover. Sentinel behavior was observed between 11 audiospectrographically and discussed in re- lation to contexts, functions, and information June and 28 October in 1973. When the social content. Four calls were associated with ag- unit was more than two birds, only three of 28 sightings involved coveys that definitely gregation and contact, three calls with alarm consisted of more than one brood. The ap- or distress, three calls with sexual attraction, parent decline of sentinel activity and the use and a single call with threat/attack encoun- of tsing calls after covey mergers (hence, de- ters. Very young chicks used two vocalizations cline in average relatedness of covey mem- and added others during maturation. Two calls were given primarily by males, but the bers) is consistent with the hypothesis that the behavior evolved because of advantages remainder of the species ’ repertoire was to the inclusive fitness of individuals. Marler shared by both sexes. (1956) similarly found that when family Social adaptations, including vocalizations, groups of the Chaffinch (Fringilla coelebs) can be accounted for satisfactorily in terms of dissolve in the winter so that most groups are benefits to individuals and their kin. Be- composed of unrelated birds, the use of hawk haviors change in frequency seasonally, coin- alarm calls ceases. ciding with changes in reproductive effort, Only the chekar vocalization and its equiv- degree of relatedness of flockmates, and risk alent in chicks were given at strikingly dif- of predation. ferent intensities. The ability to control or grade a vocal display in relation to stimulus ACKNOWLEDGMENTS strength and nearness of conspecific individ- Primary support for this study was provided by the uals increases the potential information con- Arizona Cooperative Wildlife Research Unit and by tent of the call. The use of wip-woos when a National Science Foundation fellowship. I am close to a brood or when danger still threatens grateful for the cooperation of the Arizona Game 62 WALTER L. ANDERSON

and Fish Department, the Willcox office of the U.S. HOFFMAN, D. M. 1965. The Scaled Quail in Colo- Soil Conservation Service, and the Horizon Corpora- rado: range, population status, harvest. Cola. tion, owners of the land where the study was con- Game, Fish, Parks Dep., Tech. Publ. No. 18. ducted. I thank my graduate committee-Lyle K. HUBBARD, J. P. 1974. A&n evolution in the arid- Sowls, C. Roger Hungerford, and Stephen M. Russell lands of North America. Living Bird 12:155- -for their support and encouragement. The follow- 196. ing individuals contributed in some way and have HUMPHRIES, D. A. AND P. M. DRIVER. 1967. my sincere appreciation: David E. Brown, Herbert Erratic displays as a device against predators. Collins, Chris Dunford, Harvey Nessmith, H. Ronald Science 156: 1767-1768. Pulliam, William Radke, and Jerry C. Tash. I am JOHNSGARD, P. A. 1973. and quails of indebted to Allen W. Stokes for his valuable corre- North America. Univ. of Nebraska Press, spondence and to individuals at the University of Lincoln. Michigan who carefully reviewed this manuscript: KRUUK, H. 1976. The biological function of gulls ’ Richard C. Alexander, Mark Bergland, Archibald B. attraction towards predators. Anim. Behav. 24: Cowan, Bobbi S. Low, Dale R. McCullough, Robert 146-153. B. Payne, and Robert W. Storer. My deepest thanks LEVY, S. H., J. J. LEVY, AND R. A. BISHOP. 1966. go to R. M. Anderson whose time, effort, and con- Use of tape recorded female quail calls during cern have strengthened the study from its inception. the breeding season. I. Wildl. Manage. 30: 426-428. - LITERATURE CITED MARLER, P. 1956. Behavior of the Chaffinch, Fringilla coelebs. Behav. Suppl. 5: 1-184. ALEXANDER, R. D. 1974. The evolution of social MARLER, P. 1957. Specific distinctiveness in the behavior. Annu. Rev. Ecol. Syst. 5:325-383. communication signals of birds. Behaviour 11: I ANDERSON, W. L. 1974. 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WILLIAMS, H. W. 1969. Vocal behavior of adult white Quail (Colinus virginianus). Auk 85: California Quail. Auk 86:631-659. 464-476. WILLIAMS, H. W., A. W. STOKES,AND J. C. WALLEN. West Butte Sanctuary Co., 713 Main Street, Cohsa, 1968. The food call and display of the Bob- CA 95932. Accepted for publication 25 April 1977.