Condor, 80:49-63 @ The Cooper Ornithological Society 1978 VOCALIZATIONS OF SCALED QUAIL WALTER L. ANDERSON The Scaled Quail (Callipepla squamta) is ( Wallmo 1956). Sex could often be distinguished in typically a bird of the desert-grassland and the field during soring and summer bv the color of the head and &at; fall and winter birds could be Chihuahuan Desert associations. Its range sexed with reasonable certainty only when in the extends from southeastern Arizona through hand. much of New Mexico, southeastern Colorado, With one- and two-entrance Stoddard quail traps western Texas and Oklahoma, and across baited with milo grain, I captured, banded, and Mexico in a broad band to Hidalgo (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 species (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 plumage (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 animal (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.