AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2756, pp. 1-15, figs. 1, 2 April 5, 1983

Vocal Behavior, Morphology- and Hybridization of Australian Spotted and Yellow-rumped Pardalotes (Aves, Pardalotus)

LESTER L. SHORT,' JENNIFER F. M. HORNE,2 AND RICHARD SCHODDE3

ABSTRACT In the course of varied Australian field studies between the two taxa in details of structure, pitch, during 1979 and 1980 we were able to devote some and temporal arrangement of elements. Some effort to investigating the vocalizations and other vocalizations intermediate in form, pitch and tim- behavior of the closely related, largely allopatric ing, together with morphological data, indicate (Pardalotus punctatus) in convergence oftheir characters in areas ofcontact Queensland, New South Wales, and Victoria; and in Victoria. Three of four specimens collected in to a lesser extent the Yellow-rumped Pardalote (P. Victoria are hybrids or likely hybrids as deter- xanthopygus) in Victoria. A few specimens were mined by their morphology and vocalizations. The collected of whose voices were recorded on vocal repertory as presented for the two taxa may tape. Morphological studies of these and other be complete, but more data are needed, especially specimens demonstrate the similarity of the two from P. xanthopygus. The extent of their hybrid- taxa. Their vocal repertory includes seven major ization, and hence their taxonomic status remain vocalizations. Of these, six are calls, five in punc- to be established fully, since they meet in three tatus and three in xanthopygus. Probably all six different regions (southwestern Australia, South calls are found in both forms. Differences in those Australia, and southeastern Australia), and the only of the calls found in both are nil or slight. Males detailed studies, reported here, cover but one re- of both species sing songs and abbreviated songs gion, and that only partly. similar generally in quality and tone, but differing

INTRODUCTION During the course of investigating hybrid- ior of the closely similar Spotted Pardalote ization in various Australian birds we (Pardalotus punctatus) and Yellow-rumped attempted to gather some data on the behav- Pardalote (P. xanthopygus). Our field studies ' Chairman and Curator, Department of Omithology. American Museum of Natural History; Adjunct Professor, City University of New York. 2 Research Associate, National Museums of Kenya, Nairobi, Kenya. I Research Scientist, CSIRO Division of Wildlife Research, A.C.T., Australia.

Copyright © American Museum of Natural History 1983 ISSN 0003-0082 / Price $1.50 2 AMERICAN MUSEUM NOVITATES NO. 2756

took place sporadically, as we could take time National University's Research School of from our major projects, in Queensland, New Biological Sciences; funding for his fieldwork South Wales, and Victoria during August came from the Australian National Univer- 1979 to January 1980. Previous studies of sity and the Leonard C. Sanford Fund of the the behavior of these pardalotes have been American Museum of Natural History. sparse and lacking in many details (Chandler, Throughout the text kHz is used to indicate 1961; Brunt, 1962; Richards, 1972) and fur- kilohertz, AMNH, the American Museum of ther investigations are still needed. We pre- Natural History, and mm., millimeters. sent our data on their vocalizations and mor- phology in the hope that they will provide an RESULTS appropriate framework for new investiga- tions of these two morphologically similar VOCAL REPERTORY OF pardalotes, which geographically replace each PARDALOTUS PUNCTATUS AND other generally but meet in southwestern PARDALOTUS XANTHOPYGUS Australia, South Australia, and southeastern We describe the vocalizations that we Australia (see map in Salomonsen, 1961, p. recorded from Pardalotus punctatus, P. xan- 11). Pardalotes (genus Pardalotus) form a thopygus, and possible hybrids between them, small group with no close relatives, including for comparison some of P. xan- and the relationships of which are contro- thopygus (from South Australia, western Vic- versial (Morony, Bock, and Farrand, 1975; toria, and New South Wales) kindly provided Schodde, 1975; Sibley, in litt.). by Swaby. The repertory is not considered Horne made tape-recordings using a Stel- complete, although that of major vocaliza- lavox Sp-7 recorder, a Schoeps CMC-45 con- tions of P. punctatus may be so. Our discus- denser microphone, and a 76-cm. parabolic sion is in the framework of meaning, func- reflector. Audiospectrographic analysis was tion, and motivation used in studies of conducted with a Kay Elemetrics Sonagraph Winkler and Short (1978). The arrangement 6061-B. Playback was by means of a Sony is from structurally simpler, single-note calls cassette recorder using copies of our Stella- to more complex calls and songs. Some com- vox-recorded material. Dr. R. J. Swaby made parisons are made with the available to us for audiospectrographic anal- complex (Pardalotus striatus, Short et al., in ysis a cassette copy tape of various of his prep.), congeners that are broadly sympatric recordings of Pardalotus xanthopygus and P. with both P. punctatus and P. xanthopygus punctatus, obtained by use of diverse tape- (note the lack of such comparisons of songs, recorders and microphones. The use ofcopy- which are quite different in these two major tape material inevitably affects the analytical pardalote groups). results, by reducing clarity and overtones, for BEGGING CALL: We include this call for the example. However, the timing of the ele- sake of completeness, although our only ex- ments and aspects oftheir form and pitch are amples come from three young calling con- evident in those we have figured. We made stantly while being fed by an apparent Yel- specimen comparisons at the American low-rumped Pardalote in Bat Ridge Nature Museum of Natural History. Reserve, 14 km. west of Portland, Victoria, We are especially indebted to Dr. Swaby on November 22, 1979. The only pardalote for letting us use his recordings for analyses, songs heard there were "weee-eee" songs typ- and to Mr. Norman Robinson for his sug- ical of the Yellow-rumped Pardalote (see be- gestions and aid. Mr. Richard Weatherly par- low). Begging calls tend to be conservative, ticipated in the fieldwork in Victoria. Various that is, very similar or identical in closely other individuals too numerous to mention related species of diverse taxonomic groups individually assisted us in the field, and Mr. (e.g., woodpeckers, see Winkler and Short, Robert Stolberg aided us in our laboratories. 1978). Hence the Spotted Pardalote Begging We are grateful to wildlife authorities in Vic- Calls are apt to prove similar or identical with toria for permission to obtain specimens. those we describe. The call (fig. IA) is vari- Short did his studies in Australia while he able in the emphasized pitch and composed was a Visiting Fellow in the Australian of short, fast notes 0.02 to 0.025 second in 1 983 SHORT, HORNE, AND SCHODDE: PARDALOTES 3

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(. Xt- v 0;5 1.0 1S 2.0 0;5 1.0 1.5 TIME IN SECONDS FIG. 1. Wide-band sonagrams of calls and songs of Australian Pardalotus punctatus and P. xantho- pygus. A. Begging Call, young of xanthopygus, Bat's Ridge, Victoria, November 22, 1979. B. Fast Chip Call, male punctatus, Orphan Gully, Queensland, September 21, 1979. C. Short Weep Call, notes of punctatus, Orphan Gully, Queensland, September 21, 1979, then two of hybrid male xanthopygus X punctatus, Mitre, Victoria, November 6, 1979. D. Long Weep Call, double note of male punctatus, near Heywood, Victoria, November 24, 1979. E. Long Weep Call, triple note, of hybrid male xanthopygus X punctatus, Mitre, Victoria, November 6, 1979. F. Whee Call, male punctatus, Orphan Gully, Queensland, September 21, 1979. G. Weet Call, near Heywood, Victoria, November 24, 1979, probable punctatus (possibly same as fig. 2H). H. Weet Call, longer form, punctatus, Terang, Victoria, November 28, 1979. I. Sa-weet Abbreviated Song, male punctatus, Terang, Victoria, November 28, 1979. J. Sa-weet Ba-bee Song, male punctatus, Orphan Gully, Queensland, September 21, 1979. K. Sa-weet Ba-bee Song, male punctatus, near Heathmere, Victoria, November 25, 1979. L. Same song as K, but read twice the stated frequency and twice the stated time (sonagram at 0-16 kHz, rather than 0-8 kHz). duration. Essentially each note contains fast- quality. These variations likely reflect prox- rising, partially falling, then rising, peaking imity ofthe feeding adult and relative hunger and dropping elements. Emphasis is on the of the young. Notes are uttered at 15 to 17 partial drop and following rise to a peak at per second and may be continuous or in 3.8 to 5 kHz. Some notes are simply rapid, bursts. Localization of the calling young and peaked, and inverted V-shaped; others are the eliciting of feeding by the adults are ob- noisy and fast with structure obscured son- vious functions of the call. agraphically. Harmonic tones are evident, as FAST CHIP CALL: Our only examples ofthis in most begging calls, yielding its high piping call are from a male Spotted Pardalote 4 AMERICAN MUSEUM NOVITATES NO. 2756 strongly stimulated by song playback at its call (fig. 1C) from a backcross hybrid male nest near the North Branch of Dalrymple Yellow-rumped Pardalote (see below) west of Creek, 26 km. north-northeast of Warwick, Mitre, Victoria, on November 6, 1979, hence southern Queensland on September 21, 1979. it likely occurs in both taxa. These soft notes As he perched, hopped, and stamped his feet often mark interactions of a pair, especially at the entrance to the nest he displayed toward near the nest, and apparently are the "Pe he" the "singing intruder" represented by the notes of Richards (1972, p. 155). A female recordist, erecting the red rump and spotted may give Short Weep Calls as the male sings crown feathers (similar in part to the display nearby. Thus, pair contact may be a function described by Richards, 1972, p. 155), and ofthis call. It also may be uttered by the male chip-calling in bursts and in long series of up interspersed among his songs without the to one minute continuously, or with but short immediate presence of the female, perhaps pauses over a 10-minute period. He in an attempt to establish contact with her. approached the recordist several times, right LONG WEEP CALL: This vocalization to her feet, turning about, calling and dis- resembles the Short Weep Call, often being playing. The notes (fig. 1 B) are sharp, oflow volume, but it is longer, it usually rises mechanical chips but 0.01 to 0.02 second in rather than falls in pitch, and it frequently is duration, at a tempo of 8 to 9 per second. uttered rapidly in a two-note or three-note The note sonagraphically resembles an excla- series. The few recorded examples preceded mation point without the basal dot, but struc- songs and could have been uttered by either turally is a very fast, soft drop from 7 or even the singing male or a nearby mate, or an inter- 8 kHz to about 1.9 kHz, followed instantly active second male. Five examples ofSpotted by a rise to the loud peak, continuing to the Pardalotes show (fig. 1 D) rising pitch in each end ofthe note at 5.1 to 5.5 kHz. Most sound of two notes, the notes being 0.05 to 0.13 is in the rise and peak, and in fact the initial second in duration, giving 0.20 to 0.30 sec- drop is lacking in some (less well recorded) ond for their total duration. The frequency recordings. Richards (1972, p. 155) noted rises from 3.1 or 3.2 kHz to 3.25 or 3.35 kHz. "zit" or "tik" calls from displaying Spotted Four samples, all from the backcross hybrid Pardalotes, that may represent the Fast Chip male Yellow-rumped Pardalote west ofMitre, Call just described. Preceding the male's Fast Victoria, on November 6, 1979, contain one Chip Call he uttered several Whee Calls (see to three notes (fig. 1E) at a pitch of 3.35 to below). He also sang intermittently. The 3.5 kHz, slightly higher than those of the female came and went during the male's dis- Spotted, and differing by their even, rather play, several times adding her Short Weep than rising pitch. The notes are 0.10 to 0. 125 Calls to his utterances. We cannot determine second in duration, and the three-note series the functions and meaning of the Fast Chip are rendered in 0.48 and 0.51 second. Five Call on the basis ofthis one observation, but double notes of Long Weep Calls obtained it clearly is associated with visual displays of by Swaby at Bunn's Bore near Bordertown, the male at the nest. Fast Chip Call notes are South Australia, on May 19, 1974, appear unlike any notes of the Striated Pardalotes similar to those described from the Mitre, that we studied much more intensively. Victoria, Yellow-rumped Pardalote; songs SHORT WEEP CALL: Notes given in diverse recorded by Swaby that day at Bunn's Bore situations and rendered "weep" are often of are of the high-pitched "wee-eee" type (see very low volume. Usually uttered by Spotted below) ascribed to the Yellow-rumped Par- Pardalotes in series (fig. 1C) of two to 35 or dalote. These Long Weep Calls from Bunn's more, they consist ofrelatively short, usually Bore are 0.07 to 0.13 second in duration, the down-dropping sound at 2.75 to 3.8 kHz, second note of each pair being shorter than with a weak to moderate overtone at 7 kHz the first. Their pitch is at 3.3 to 3.4 kHz, near or more. Each is 0.035 to 0.085 second (mean that of those described for the Spotted Par- of 32, 0.05 5 second) in duration. In series the dalote, but there is no rise in pitch. Until tempo varies, but approximates 2 per second there are more data it seems prudent to draw with an interval of 0.37 to 0.66 second no conclusions from the apparent differences between notes. We have recorded the same just noted. The function of this call in regard 1 983 SHORT, HORNE, AND SCHODDE: PARDALOTES 5 to the Short Weep Call particularly needs song. Individuals stimulated by playback study. often show greater variation in their songs WHEE CALL: A single, double, or occasion- than prior to playback. ally a series call, a whistled "wheee," was SA-WEET ABBREVIATED SONG: The Sa-weet associated with periods of excitement such is a double-noted introduction to the full song as human intrusion near the nest, and play- of Pardalotus punctatus that may be uttered back of songs at a male. The 10 notes studied by itself, usually interspersed among full songs (fig. IF), all of punctatus, are 0.245 to 0.36 or full songs and Weet Calls (the Weet Call second at an even pitch or dropping slightly also could be considered an Abbreviated between 3.55 and 3.7 kHz initially and 3.45 Song, or rather a replacement Abbreviated and 3.55 kHz terminally, and show a weak Song, as it is not part of the typical song). harmonic tone at 6.75 to 6.9 kHz, or more The 29 examples of Sa-weet Abbreviated usually, none. In series they are uttered at a Songs (fig. 11) studied sonagraphically, when variable tempo with intervals of 0. 12 to 1.27 compared with 43 examples ofSa-weets from second. The pitch is higher than that of the full songs show that these Sa-weets are equiv- full song (see below), and its even pitch lack- alent except that the Abbreviated version ing modulation renders it unlike any other varies a bit more overall, and tends to be calls of the Spotted, Yellow-rumped or longer, note for note, than when preceding Striated pardalotes. This call preceded sev- the "Ba-bee" ofthe full song. The 29 Sa-weet eral of the Fast Chip Calls of the male dis- Abbreviated Songs are 0.22 to 0.265 second playing at the Dalrymple Creek site men- in duration, with the "sa" taking 0.015 to tioned under those calls. 0.07 second and the "weet," 0.105 to 0.16 WEET CALL: This loud note is associated second. The "sa" is at 2.75 to 2.85 kHz, and closely with either abbreviated (eight cases, the "weet" at 2.65 to 2.8 kHz. Overtones "sa-weet," see below) or full (nine cases, "sa- usually are absent or at most weak on the weet ba-bee," see below) songs of Spotted "sa"l at 5.5 and 7.8 kHz, and weak to mod- Pardalotes. It is the first of the songlike calls erate at 5.2 to 5.3 and moderate to strong at to be considered. A "weeet" or "weee-eet" 7.6 to 7.8 kHz on the "weet." The shorter describes the call (fig. 1G) which often has a "sa"1 often is weaker than the "weet," and terminal ("-eet") element (fig. 1 H). The dura- may be inaudible in distantly recorded calls; tion ofthe call is 0.09 to 0.20 second, or, with it tends to rise slightly in pitch. The "weet" the "eet," 0.25 to 0.29 second. Its frequency is strong and either level or wavering some- is between 4.25 and 4.65 kHz, mainly 4.3 to what in pitch with occasional signs of mod- 4.5 kHz, with overtones above 8 kHz (weak ulation. There always is a gap between the to strong), at 12.5 to 13.5 kHz (stronger), and two elements exceeding 0.05 second in du- sometimes at 16 kHz. In form the note shows ration. a distinctive, more or less pronounced, rise SA-WEET BA-BEE SONG: This is the typical initially then an almost imperceptible fall; the song (fig. 1J, K) of the Spotted Pardalote. terminal "-eet," when present (fig. 1 H), begins Brunt (1962) was correct in declaring that below the tail ofthe longer initial "weee" and only one bird, not two (see Chandler, 1961) rises slightly before ending. Consecutive calls utters the full Song, and in our experience it have not been recorded, rather the Weet Call is always given by the male. At a distance the precedes or follows other forms of song by weakest element, the "sa," tends to drop out; 1.0 to 1.8 second. No Weet Calls have been the "ba-bee" often runs together such that a found in the material of xanthopygus avail- drawn out single note is heard, but overtones able to us. A single Weet Call note comprises always reflect the dual nature of that portion the initial element in some variant full songs of the song. Overall the full song is but 0.67 described below. We have been unable to find to 0.80 second (mean 0.747 second) long in behavioral differences associated with vari- 43 examples from Queensland to Victoria, ation in the song form of Spotted Pardalotes; and it shows relatively little variation (com- some individuals in a locality may sing only pare fig. IJ and 1K). The Sa-weet Abbrevi- typical songs, while others may switch back ated Song has been analyzed above, and the and forth among two or even three types of "sa-weet" of the full song is similar but 6 AMERICAN MUSEUM NOVITATES NO. 2756 shorter, the "sa" lasting from 0.01 to 0.06 the vicinity of the male, whether singing second and "weet," from 0.065 to 0.12 sec- without playback or in response to playback. ond in duration. The break or gap between However, the loud, penetrating song and its the "sa" and "weet" is 0.05 to 0.12 second, constant repetition provide a clear indication with a mean of 0.09 second. The Sa-weet is of the singer's location to its mate, and loca- followed in 0.17 to 0.24 second (mean 0.209 tion and perhaps stimulation of the female, second) by the loud, pulsed, modulated Ba- pair-bond maintenance and other functions bee ("baa-a-bee-ee-ee," or "baa-a-ee-eee-ee") may be served by this song. of 0.24 to 0.33 second (mean 0.29 second) WIT WEE-EEE SONG: This is the Yellow- duration. The Ba-bee may be connected rumped Pardalote's equivalent (fig. 2A) ofthe throughout by sound, but usually the "Ba" Spotted Pardalote's Sa-weet Ba-bee Song. ends in several pulses ofsound at 0.07 to 0.19 When uttered in full it is a much longer, if second, after which there may be a slight break structurally simpler song than that of punc- before the "bee." tatus, with a duration of 1.12 to 2.28 seconds The frequency of the "sa" is at 2.75 to 2.9 (mean 1.63 seconds). In 30 full songs the short, kHz (mean 2.83 kHz), the "weet" at 2.6 to weak "wit" (it may be lacking, see below) is 2.85 kHz (mean 2.72 kHz), the "ba" at 3.1 0.02 to 0.07 second (mean 0.054 second) in to 3.3 kHz (mean 3.19 kHz), and the "bee" duration, and the "wee-eee" is 0.21 to 0.32 at 3.0 to 3.3 kHz (mean 3.17 kHz). The first second (mean 0.276 second), the "wee" last- two notes thus are at a distinctly lower pitch, ing 0.10 to 0.21 second and the "eee" 0.01 usually with a slight drop from the "sa" to to 0.115 second. Some temporal variation is the "weet." The weaker "sa" lacks harmonic due to the weakness of the "wit" and "eee" tones (in 31 of 43 cases), or they are weak, elements, only part of which may be loud whereas the "weet" shows overtones at 5.15 enough to reproduce well sonagraphically. to 5.4 kHz (mean 5.25 kHz), usually a stronger There also is considerable variation in the one at 7.5 to 7.85 kHz (mean 7.75 kHz), and length ofthe song (fig. 2A, B) due to the vari- often others at about 10.4, 13.4, and 15.1 able interval between the "wit" and the "wee,"9 kHz. The "ba" and "bee" essentially are at that being 0.82 to 1.92 second (average 1.31 one pitch (equal in 18 cases, the "ba" is barely second). Thus the two main parts ofthe song higher in 13, a trifle slower in 2 of 33 checked are separated by at least several times the for this feature). Overtones are usually weak interval between the two main parts ("sa- to strong but they delimit the "ba" and "bee" weet" and "ba-bee") of the Spotted Parda- more clearly than does the fundamental tone. lote's song. It can be seen clearly from figure 1L, which In pitch the initial "wit" is variable, provides analysis up to 16 rather than 8 kHz, between 2.4 and 3.1 kHz, but averaging 2.91 that the alternate overtones are emphasized kHz, about at the maximum for the "sa" note (at about 9 and 15 kHz) and contribute sub- of the Sa-weet Ba-bee Song. The "wee-eee" stantially to the sound characterizing this rather consistently shows a drop in pitch (in song. 27 of 30 songs), averaging 3.74 kHz for the The functions of the song undoubtedly are "wee" and 3.58 kHz for the "eee" elements; complex, as in the alternating of this form of the low extreme of both "wee"" and "eee" the song with the previous two forms. Reac- barely reaches the upper frequency limit of tion to playback of song is intense and often the "ba-bee" in the Spotted Pardalote's song, long-lasting (a relatively infrequently singing the range being 3.15 to 3.9 kHz for the "wee" male, upon playback, may sing much more and 3.0 to 3.8 kHz for the "eee." Harmonic frequently not only that day, but the next as tones are lacking in sonagrams from copies well, with no further playback attempted), of Swaby's recordings, but they are clearly inferring a territorial function, and, with cor- kHz related visual displays, a definite aggressive marked and strong (at 5.8 and over 8 motivation (Winkler and Short, 1978). The for the "wit," and, especially, at 7.5 to 7.25 singing male usually actively seeks the source kHz for the "wee-eee") in the hybrid Mitre, ofthe playback, approaching and circling the Victoria, male discussed below. recordist at a range of 1 meter or even less. The "Wit" note is sometimes uttered apart The female may or may not be attracted to from the full song (examples by Swaby from 1983 SHORT, HORNE, AND SCHODDE: PARDALOTES 7

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FIG. 2. Wide-band sonagrams of more calls and songs of Australian Pardalotus punctatus and P. xanthopygus. A. Wit Wee-eee Song, xanthopygus, Sedan, South Australia, August 31, 1975 (Swaby). B. Longer Wit Wee-eee Song, xanthopygus, Swan Reach, South Australia, May 1977 (Swaby). C. Wee-eee Song, xanthopygus, Tailem Bend, South Australia, June 1, 1975 (Swaby). D. Variant song ("eep-beeee"), male punctatus, Orphan Gully, Queensland, September 21, 1979. E. Variant song ("wee-eee-eee"), xanthopygus, Kangaroo Island, South Australia, March 31, 1974 (Swaby). F. Possibly hybrid punctatus X xanthopygus song, Nelson, Victoria, September 12, 1967 (Swaby). G. Possibly hybrid punctatus X xan- thopygus song, Overland Corner, South Australia, October 8, 1973 (Swaby). H. Weet Call, likely hybrid male punctatus x xanthopygus, Heywood, Victoria, November 24, 1979. 1. Sa-weet Beee Song, likely hybrid male punctatus X xanthopygus, Inglewood, Victoria, November 5, 1979. J. Weet-beee Song variant, some male as I. K. Wit Wee-eee Song, male hybrid xanthopygus X punctatus, Mitre, Victoria, November 6, 1979. L. Another song, same type as K, of same male. M. Wee-eee Song, same hybrid male as in K.

Hattah Lakes, Victoria, August 30, 1978), lacking the Wit introductory note. This is in although not to the same extent as in the Sa- contrast to the Ba-bee of the Sa-weet Ba-bee weet of the Sa-weet Ba-bee Song. In contrast Song, which appears not to be given apart the Wee-eee is uttered, apparently frequently, from the Sa-weet elements. without the Wit; the lack of a Wit note may It should be noted that certain Wee-eee be ascribed to lack offocus ofthe microphone Songs (lacking the "wit," see fig. 2C) closely or distance from the bird, as this note is rel- resemble the "weet-it" version (nearly dou- atively weak, but our Mitre recordings were ble-noted version) of the Spotted Pardalote's at close range and we could hear songs uttered Weet Call. The two are of the same duration 8 AMERICAN MUSEUM NOVITATES NO. 2756 and general form, the Wee-eee differing from between punctatus and xanthopygus makes the "weet-it" only in its slightly higher pitch any convergence (i.e., the higher pitch tend- and distinct downward rather than upward ing toward that of xanthopygus) noteworthy. inflection at the end. VARIANT YELLOW-RUMPED PARDALOTE VARIANT SPOTTED PARDALOTE SONGS: Sev- SONG: Among the vocalizations on a tape eral songs do not match the descriptions just provided by Swaby is a three-noted, loud provided in all oftheir characteristics. A male song, a "wee-eee-eee" (fig. 2E), uttered March from Orphan Gully, Queensland, just prior 31, 1974, on Kangaroo Island, South Aus- to giving the typical song shown in figure 1J, tralia, by a presumed Yellow-rumped Par- uttered an "eep-beee" (fig. 2D). This latter dalote. This song, 0.59 second in duration, vocalization is loud and somewhat resembles contains notes consecutively 0.18, 0.14, and the full Sa-weet Ba-bee Song in overall form, 0.12 second in duration, and separated by but it lacks the "sa" note, the "eep" is not gaps of0.05 and 0.10 second. The three notes like the second "weet" note of the full song, show only a very slight drop in pitch, from and the drawn out "beeee" is continuous, 3.4 to 3.35, 3.35 to 3.3, and then 3.3 to 3.25 with no indication of a break. The initial kHz. They could be considered an extended "eep" rises peculiarly (somewhat like a Weet Wee-eee Song (lacking the introductory Wit Call, but that call is longer and usually higher note), but, although their pitch is in the lower pitched), and its frequency (3.35 kHz) is well range ofthat song, the notes are farther apart above that of the "Sa-weet" introductory (the first two together are 0.37 second in dura- notes of the full song. The "beee" is, at 0.32 tion, the maximum for the Wee-eee Songs at second, within the range of the full "Ba-bee" hand being 0.32 second), the second note is of a typical full song, and it is at the high too long, and there is a third note. There is extreme of pitch (3.3 kHz) of the "Ba-bee." resemblance to a three-noted Long Weep Call There are several pulses indicated, as in the of punctatus (fig. 1E), but the Wee-eee-eee full song. One significant aspect of this vari- notes lack the regular form and shape ofthose ant song is the slightly dropping pitch from notes sonagraphically, and show a suggestion the first to the second note, which, with the of the frequency modulation that marks the high pitch of the first note imparts a bit of Ba-bee portion of the full song of punctatus, the quality of a Yellow-rumped Pardalote's and to a lesser extent the Wee-eee portion of song. Another point is its occurrence in the that of xanthopygus. We tentatively consider same individual singing typical punctatus full this vocalization a variant form of the song songs, showing that punctatus is capable of of the Yellow-rumped Pardalote; Spotted giving songs tending toward those ofxantho- Pardalotes do not occur on Kangaroo Island. pygus in at least some aspects. POSSIBLY HYBRID SONGS: Two series of We note that songs recorded September 3, songs provided us by Swaby are of interest 1973, near Karoonda, South Australia, by in that they exhibit peculiarities of structure Swaby are at the extreme high pitch of those suggesting intermediacy, or at any rate a ten- of P. punctatus from elsewhere, although the dency in songs presumed to be ofone species, five examples studied are in other respects toward the song of the other species. One typical of Sa-weet Ba-bee Songs. They are array consists of songs from Nelson, south- rather weak, without overtones, in our copy western Victoria, recorded by Swaby on Sep- tape, hence we do not show the sonagrams, tember 12, 1967. In form and sound the song but the pitch of the "sa" was at 2.8, 2.85, (fig. 2F) appears to be a distantly recorded 2.85, 2.9, and 2.95 kHz in the five songs, the Sa-weet Ba-bee Song, i.e., ofthe Spotted Par- "weet" is at 2.85 in one and 2.9 kHz in four dalote, but lacking the "sa" note. The prox- examples, the "ba" is at (3.3 kHz), or just imity of the initial "weet" to the "ba-bee" above (3.35 kHz) the extreme high of3.3 kHz (interval 0.17 second in all three examples) for punctatus, and the "bee" is within the is that ofthe Sa-weet Ba-bee Song. The initial range ofpunctatus. This variation may prove "weet" is at 3.25 to 3.35 kHz in the three to be within that for punctatus in South Aus- examples, and this frequency is considerably tralia, but geographical proximity there above that of the Sa-weet in a typical punc- 1 983 SHORT, HORNE, AND SCHODDE: PARDALOTES 9 tatus song. There is a rise in pitch from the be of hybrids, and that studies of the status Weet to the Ba-bee, which is virtually at an of P. xanthopygus and P. punctatus in the even pitch, at 3.65, 3.7 and 3.85 kHz, in these vicinity of Nelson and Overland Corner songs, the pitch being well above that of the would be productive. Ba-bee of punctatus songs and within that of the Wit Wee-eee Song ofxanthopygus. Thus, in effect, these songs are like that ofpunctatus VOCALIZATIONS OF THREE but pitched like that of xanthopygus. COLLECTED MALES Among songs recorded by Swaby at Over- We recorded vocalizations of three males land Corner on the Murray River in October that we later collected, their behavior or mor- 1973 are a group from October 8 that are phology or ecology causing us to question typical in every way of those we have their identity. Their vocalizations are described for P. xanthopygus, namely drawn described here, and their morphology dis- out Wit Wee-eee Songs. However, on both cussed below. October 5, and 8, 1973, he tape-recorded the Male, 5 km. south-southeast of Heywood, voice ofa pardalote uttering a distinctive song Victoria (AMNH 824921). Within but near (fig. 2G) with attributes of both the Sa-weet the edge of second growth moist eucalypt Ba-bee and Wit Wee-eee songs. This song woodland we found numbers of Spotted Par- may be rendered Sa-weet Wee-eee, and five dalotes, as determined by song, on Novem- examples show it to be variably long (1.7 to ber 24, 1979. One of the males in this area 2.13 seconds, in the range of xanthopygus) attracted our attention by its pale rump that with an interval of 1.12 to 1.53 second appeared neither yellow nor rusty. We sought between the "weet" and the "wee" (interval this individual, tape-recording as we did so. that of xanthopygus). The first, "sa" note is A number ofvocalizations were recorded, but 0.02 to 0.08 second (overlaps time of both are not definitely ascribable to the bird in "sa" and "wit," to longer than either), and question (e.g., the Weet Call, fig. 1G). We the "weet," coming 0.07 to 0.1 1 second after finally obtained a clear view of the bird and the "sa," is 0.125 to 0.16 second in duration recorded it while in view up to the time it (the "weet" exceeds the duration ofboth "wit" was secured. Its vocalizations include a Weet and "weet" notes, respectively of the song of Call (fig. 2H), uttered before a Sa-weet Abbre- xanthopygus and ofpunctatus). The Wee-eee, viated Song; the Weet is a bit low in pitch at 0.27 to 0.285 second, with the "wee" at (4.1 kHz) for typical punctatus, as even other 0.105 to 0.15 second and the "eee" at 0.075 calls from the same locality are pitched above to 0.1 second, is entirely within the range of 4.2 kHz. Not much can be made of this, for xanthopygus in these parameters. In fre- we have no Weet Calls of xanthopygus. We quency all notes exceed that ofpunctatus, and recorded eight Abbreviated Sa-weet Songs fall within the range ofxanthopygus, with the and three full songs from the male we sought. "sa" at 3.05 to 3.15 kHz, the "weet" at 3.1 The Sa-weet Songs are typical of those to 3.2 kHz and descending slightly by up to described for punctatus in all respects, 0.1 kHz, the "wee" at 3.8 to 3.9 kHz descend- although the pitch, particularly of the weet ing by 0.05 to 0.1 kHz, and the "eee" at 3.7 element, is at the low end of the spectrum to 3.8 kHz and descending very slightly. (some at 2.6 kHz) in the sample ofpunctatus Effectively the second "weet" note has been Sa-weets available to us. The full songs are added to a song otherwise like that of xan- punctatus-like also, although they too tend to thopygus; the overall effect is that of a long- be low-pitched (two of three "sa" elements interval, high-pitched punctatus song with at 2.65 kHz). We conclude from the available xanthopygus-like elements. Certainly the vocal data for this male that its repertory was "weet" element otherwise is lacking in the punctatus-like, ascribing no cause for the ten- repertory ofxanthopygus as currently known, dency ofits vocalizations toward a low pitch. and is equivalent in structure to the "weet" Other vocalizations from this site include two note of a punctatus Sa-weet Ba-bee Song. Long Weep Calls, three Weet Calls, two Sa- We suggest that these peculiar songs may weet Abbreviated Songs, seven Sa-weet Ba- 10 AMERICAN MUSEUM NOVITATES NO. 2756 bee Songs, and two odd songs, not well Male, 3 km. west of Mitre (41 km. west of recorded, rendered "weep-bee." All are typ- Horsham), AMNH 824922. This male, ical of punctatus except the last, odd songs. apparently alone, sang and called in dry scle- These are unusual in the even pitch, at 3.25 rophyll woodland dominated by kHz, of the weaker, short (0.04 to 0.06 sec- leucoxylon and E. baxteri. We analyzed a ond) initial note and of the longer, pulsating series ofShort Weep Calls (fig. I C), four Long "bee" note, which closely resembles the "ba" Weep Calls of one to three notes (fig. 1E), element of Sa-weet Ba-bee Songs. We regard four Wee-eee Songs (fig. 2M), and nine Wit these odd songs as variant punctatus songs Wee-eee Songs (fig. 2K, L). The Short Weep (both unfortunately show too much back- and Long Weep calls resemble those ofpunc- ground noise to be figured). tatus, as discussed above, comparable calls Male, 4 km. northwest of Inglewood (50 of xanthopygus being unavailable for analy- km. northwest ofBendigo), National Museum sis. In form the songs clearly resemble those ofVictoria collection. On November 5, 1979 of P. xanthopygus, with a short preliminary this bird and its mate attracted our attention note followed after a pause by a double note because ofthe habitat in which we saw them that drops. The pitch of all elements is that which was mallee scrub and partly second of xanthopygus (Wit, 2.85-3.05 kHz; Wee- growth. The male called and sang vigorously, eee, dropping 3.9 to 3.65 kHz). However, in and since we saw no other individual than their temporal facets and in some aspects of its mate, we attribute all the vocalizations we structure the songs are not quite like those of describe to the two birds. Both appeared xanthopygus. The Wee-eee Songs and ele- punctatus-like in plumage. We recorded Short ment of the full song are longer than in xan- Weep Calls (probably of the female), eight thopygus in six cases. The Wit note ofthe Wit Weet Calls, five Abbreviated Sa-weet Songs, Wee-eee Song in seven of nine cases (see fig. eight full (Sa-weet Ba-bee) songs, and five 2K) is separated from the Wee-eee by less variant Weet-bee Songs. The songs and Weet than the 0.82 second minimum in xantho- Calls were uttered by the male of the pair. pygus songs (the interval is 0.66 to 0.94 sec- The Weet Calls and Short Weep Calls differ ond in the bird from Mitre, 0.82 to 1.92 sec- not at all from those described above for ond with a mean of 1.31 second in 30 punctatus (and for xanthopygus in the case xanthopygus songs). Well recorded songs of of the latter call). The Sa-weet and Sa-weet both types from the male from Mitre show a Ba-bee songs also fall within the range ofvari- separate terminal element following the Wee- ation of punctatus (a Sa-weet Ba-bee Song is eee (fig. 2L, M), and, preceding this terminal shown in fig. 21), although the Ba-bee section element, an even-pitched element at a fre- of the full song is rather continuous, more a quency above the dropping "-eee" segment "beeeeee" with no or almost no break. The (fig. 2K, L, M). The Wee-eee portion also is variant Weet-beee Songs (fig. 2J) resemble a well connected in most songs from the Mitre variant punctatus song from Queensland (fig. bird, giving a "weeeeee" sound rendering it 2D), and even more closely, the Weep-bee difficult to discern a break. While all these variant song described above for the Hey- variations are not in the direction of punc- wood male. The rising first note at 3.1 to 3.5 tatus songs, the temporal shortening of the kHz, the slightly lower but very evenly song is. That shortening and the observed pitched, continuous beee element, and tem- temporal variation noted above in recordings poral relations of the elements are notewor- by Swaby from Nelson and Overland Corner, thy. We see no strong indication ofwide spac- lead us to suspect that there is punctatus influ- ing of the notes, increased pitch of the song, ence shown in the songs of the male from or a broken beee element, dropping in pitch, Mitre. any of which might represent a tendency toward xanthopygus. Rather the Weet-beee Songs are considered as a variant punctatus MORPHOLOGY AND HYBRIDIZATION song, pending further elucidation of the rep- Pardalotus xanthopygus is a trifle larger ertory of xanthopygus. (longer winged, longer tailed) than is P. punc- 1983 SHORT, HORNE, AND SCHODDE: PARDALOTES I1I

tatus; its bill is the same length but tends to length, 28.91 vs. 30.53 mm.; bill length (ex- be slightly less broad, and less deep. The two posed culmen) 6.89 vs. 7.23 mm.; bill width taxa differ in a number of features, but these across nostrils, 4.29 vs. 4.11 mm.; and bill are such that they well may be genetically depth at center ofnostrils, 3.91 vs. 3.67 mm. closely tied. Basically, xanthopygus is paler Overlap in all cases is very extensive (even and punctatus is washed with buffy cinnamon complete in bill length and bill depth). Even to chestnut; reduction ofthis single color array in the case of the character showing greatest in punctatus would render it into "xantho- difference, tail length, 10 of 16 punctatus fall pygus." Dorsally, the spotting is similar in within the range of xanthopygus and 10 of the two, but the spots are washed yellowish 15 of the latter fall within the range of punc- buff to cinnamon in punctatus. The lower- tatus. What we see then are tendencies, that most back and rump are yellow in xantho- is, for xanthopygus to attain slightly greater pygus and cinnamon to chestnut cinnamon tail, wing and bill lengths, with punctatus in punctatus (but these areas may be under- having a slightly wider, deeper bill. The men- lain with yellow in punctatus, which has yel- sural differences, ifindeed larger samples were low undertail coverts). Both are yellow on the to prove them valid, are trivial, not even ap- undertail coverts, but these are cinnamon- proaching those found between subspecies of or buff-tinged in punctatus. The underparts one species. These data clearly offer no help of punctatus are variably washed in cinna- in determining hybridity of any individual mon buff or creamy buff, whereas xantho- bird. pygus has such color extremely reduced, Four male specimens were obtained in the mainly appearing whitish or buffy whitish course of our studies in Victoria during below. Finally, the yellow throat of xantho- November 1979. They were compared with pygus is less golden, that of punctatus, more more than 50 males of each of P. punctatus golden yellow. All ofthese traits involve rich- and P. xanthopygus representing all seasons ness of (mainly cinnamon) color, and it may of the year. The morphology of the newly be that they are affected by very few genes collected birds is treated in order of the date or gene combinations. of their collection. The colors are affected by wear, chestnut Male, 4 km. northwest of Inglewood (50 cinnamon or buff cinnamon fading to buff km. northwest of Bendigo), National Mu- throughout; the back spotting of very worn seum of Victoria collection. This bird was punctatus may show little buff, closely resem- studied and its voice recorded November 5, bling the white-spotted condition of xantho- 1979, in mallee scrub, where it was singing pygus. Some worn punctatus, e.g., male (see above), accompanied by a female, ap- AMNH 698754 from Selby, Victoria, parently its mate. It weighed 8.4 grams and approach xanthopygus not only in color of its testes measured 2 by 1 mm. Its bill is the back spots, but in yellowish appearing in longer than that ofeither form (from samples the (pale cinnamon) rump and in the paleness used in comparison); at 7.8 mm. the bill length of the underparts. These factors, taken with might be taken as an indication of xantho- the very close resemblance of the two taxa in pygus influence, but bill lengths overlap in all other traits (some ofthem involving com- our samples (though averaging 0.34 mm. plex patterns and strong color markings), longer in xanthopygus). The yellow of the make the determination of hybrids difficult. throat has a golden tone and some cinnamon Measurements also disclose very little dif- traces posteriorly. Ventrally it is pale buffy ference between these two pardalotes. Com- cinnamon, less richly colored than punctatus paring 16 adult males of P. punctatus from The New South Wales and Victoria taken Sep- and within the range of xanthopygus. tember to January with 15 males of P. xan- undertail is richly colored, golden yellow with thopygus from Victoria obtained in Septem- strong cinnamon edging at the base. Its rump ber to November gave these mean results, is rich cinnamon rufous tending to chestnut, punctatus preceding xanthopygus in each case: well within the range of punctatus. The back wing length (chord) 57.19 vs. 57.87 mm.; tail spotting shows but slight buffy edging, espe- 12 AMERICAN MUSEUM NOVITATES NO. 2756 cially posteriorly and in the scapular areas. this may reflect fading; it is too richly cin- Thus, possible xanthopygus influence is evi- namon for xanthopygus, and within the range dent only in ventral and dorsal color, and in ofvariation exhibited by punctatus. The yel- bill length. The specimen does not appear low undertail coverts have richly cinnamon exceptionally worn (which would result in bases, as in punctatus. Its dorsal spotting is fading and an "approach" to xanthopygus), peculiarly variable, and as the bird is not and we consider this bird a likely hybrid of moulting, this variation is important. A few punctatus X xanthopygus or a backcross feathers bear quite rusty spots, most are very product of such a hybrid crossed with punc- faintly buff-tinted, and a few anterior back tatus. feathers bear almost white spots. Some faded, Male, 3 km. west of Mitre (41 km. west of worn punctatus come close to matching this Horsham), AMNH 824922. Observed, voice specimen in dorsal coloring, but none is so recorded, apparently alone in open dry scle- variable, and none has such pale anterior rophyll woodland on November 6, 1979. It spotting. The rump color also is mixed, nearly weighed 8.6 grams and its testes (damaged) chestnut at the tips ofsome feathers and paler measured maximally 5 mm. In measure- cinnamon elsewhere with a hint of yellow. ments this specimen falls within the range of The rump feathers are worn and partly faded. overlap between punctatus and xanthopygus, The uppertail coverts barely show tipping of except for tail length (31.3 mm.), in which it red-orange, not red, and the combination of slightly exceeds the maximum measurement this lack ofred and the mixed coloring of the for punctatus. In its coloration this specimen rump was what probably attracted our atten- falls completely within the range of variation tion to it. Thus, the bird may be a hybrid or ofxanthopygus except for its undertail coverts a backcross product (hybrid X punctatus), as and rump. The undertail has yellow feathers indicated by its back and rump coloring (its slightly cinnamon toward their bases; this long bill may be considered suggestive ofsuch cinnamon is lacking or barely hinted at in the a cross, though not significant in itself). xanthopygus specimens at hand. The rump Male, 4.3 km. southwest of Heathmere (12 feathers, basically yellow, show cinnamon at km. north-northwest of Portland), National their bases, edges, and tips, definitely exceed- Museum of Victoria collection. On Novem- ing the variation within xanthopygus, but ber 25, 1979 we obtained this bird, weighing closer to that form than to punctatus. The 7.5 grams and with testes 2 X 1.5 mm. (right white-spotting of the lower back seems unaf- testis smaller, 1 by 1 mm.), in heathy, moist fected by the cinnamon evident in nearby sclerophyll forest. Its vocalizations were tape- rump feathers. We conclude that this speci- recorded (and proved identical in every way men represents a backcross product of a with those of punctatus). All its measure- hybrid between xanthopygus and punctatus ments are within the range ofoverlap between with xanthopygus. punctatus and xanthopygus. The specimen is Male, 5 km. south-southeast of Heywood quite richly colored, with golden yellow throat (18.5 km. north-northeast of Portland), tinged cinnamon, richly cinnamon under- AMNH 824921. The voice of this male was parts, broadly cinnamon-based undertail recorded, and the bird taken at the edge of coverts, buffy cinnamon back spotting (quite moist sclerophyll forest on November 24, pale anteriorly, but matched by some punc- 1979. It weighed 7.8 grams, and its testes tatus), and rusty chestnut uppertail coverts. measured 2 by 1 mm. Other pardalotes in the We have no hesitation in considering this area included rusty-rumped and yellowish- bird to represent P. punctatus; others with rumped birds. Mensurally this specimen is chestnut rumps were seen in the surrounding within the range of overlap of both forms, forest. except that it has a long bill (8.0 mm.), which a may or may not reflect tendency toward SUMMARY OF MORPHOLOGICAL AND xanthopygus. The throat is golden yellow in THESE SPECIMENS this bird, less rich than in many punctatus, VOCAL DATA FOR but within the range ofvariation ofthat form. Combining results of the- morphological Its underparts are a trifle pale cinnamon, but comparisons and vocal data, we find: 1) the 1 983 SHORT, HORNE, AND SCHODDE: PARDALOTES 13

Inglewood male morphologically is a likely ofterritory, both species being highly aggres- hybrid with punctatus-like vocalizations that sive toward conspecific birds and toward other show some peculiarities, and we conclude that species as well (both are aggressively inter- it is a likely hybrid or backcross product; 2) active with, and dominant over the sympatric the Mitre male is mainly xanthopygus-like in Pardalotus striatus, despite the latter being morphology and in voice, but shows definite larger than either punctatus or xanthopygus). tendencies toward punctatus in both, hence The songs function probably in synchrony of we consider it a backcross product ofa hybrid reproduction and pair-bond maintenance with xanthopygus; 3) the Heywood male is between mated birds, judging from the great mainly punctatus-like in morphology and in frequency ofsongs and "excited" behavior of vocalizations, but with peculiarities in both singing males in the presence of their mates. cases-we conclude that it represents a hy- Finally, the very often repeated songs clearly brid or the result of the crossing of a hybrid indicate the location ofthe singers to the mate, with punctatus; and, 4) the Heathmere male and to adjacent conspecific pairs. morphologically and vocally is punctatus-like The results we have presented concerning (with some vocal-peculiarities), and we assign interbreeding of P. punctatus and P. xantho- it to punctatus. pygus require amplification and further study, especially in those areas in which these two pardalotes meet. Our vocal data are sugges- DISCUSSION AND CONCLUSIONS tive of at least limited hybridization between the Spotted and Yellow-rumped pardalotes. Of six calls described for these pardalotes, We emphasize the need for further study of three (Fast Chip, Whee, Weet calls) are so far the variation in vocalizations of both forms. known only from P. punctatus, although Also, the ontogeny of the vocalizations and approaches to the Whee and Weet calls have inheritance ofvarious calls and songs require been noted above in P. xanthopygus. The investigation. Nonetheless certain individual special circumstances under which the Fast birds show vocal convergence, and they come Chip Calls were recorded in punctatus suggest from areas of contact or overlap between that they are uttered under similar circum- punctatus and xanthopygus. Most impor- stances in xanthopygus. Another call, the tantly the intermediacy we have described in Begging Call, was recorded only from xan- vocalizations ofsome individuals is matched thopygus, but such calls are generally similar by their morphological intermediacy. This in congeneric species (e.g., of woodpeckers; concordance oftwo diverse character systems Winkler and Short, 1978), and are apt to be and their occurrence of course strengthens similar in punctatus and xanthopygus and our bases for believing that these pardalotes certainly to occur in punctatus. The Short interbreed, although we must rely mainly on Weep and Long Weep calls described for both the morphological data until the vocaliza- species are alike in the case ofthe former call, tions of the two are subjected to more inten- and their Long Weep Calls differ only slightly. sive studies. This does not detract from the It thus appears likely that the six calls occur importance ofthe vocal data, which in them- in both species, and that they are very similar selves represent a significant advance in our if not in every case identical in the two. Data knowledge of the behavior of pardalotes. do not yet permit us to state that the calls are Sample sizes from areas of contact so far functionally identical, but we have no evi- are insufficient to establish the incidence of dence to the contrary. hybridization between punctatus and xan- The songs ofmales ofthe two species differ thopygus. In particular, human modification in the form oftheir elements, their pitch, and ofhabitat may have limited or enhanced con- their temporal features, although the general tact between the two in recent times, as Sal- structure of the songs and their quality are omonsen (1961, p. 14) has suggested. Hence, similar. Both sing abbreviated versions of not only are appropriate samples needed from their respective songs, the context and func- central and western Victoria, southern South tion ofwhich are not clear in relation to those Australia and southwestern (Western) Aus- ofthe full songs. The songs function in defense tralia, but the ecological situation in the areas 14 AMERICAN MUSEUM NOVITATES NO. 2756 of sampling needs to be investigated concur- Keast, J. Allen rently. 1961. Bird speciation on the Australian con- The Spotted Pardalote has been assumed tinent. Bull. Mus. Comp. Zool., vol. 123, (e.g., Salomonsen, 196 1; see also Keast, 196 1) pp.305-495. to be a Bassian (wet, southern Australian) Morony, John J., Jr., Walter J. Bock, and John Farrand, Jr. derivative, and the Yellow-rumped Parda- 1975. Reference list of the birds of the world. lote, an Eyrean (arid, interior Australian) Amer. Mus. Nat. Hist., x + 207 pp. derivative of their common ancestor, with Richards, B. spreading of both in a complicated manner 1972. The nesting of the Spotted Pardalote. to the west and to the east subsequent to spe- Austral. Bird-Watcher, vol. 4, pp. 150- ciation such that both taxa now have popu- 156. lations isolated in southwestern, south-cen- Salomonsen, Finn tral and southeastern Australia. Until we 1961. Notes on (Aves, Dicaei- know more about the interactions of the two dae) 5. The genera Oreocharis, Para- forms in all their areas of contact it seems mythia, and Pardalotus (except the superspecies Pardalotus striatus). Amer. premature to speculate on their history and Mus. Novitates, no. 2067, 24 pp. we defer this until a later time. Schodde, Richard LITERATURE CITED 1975. Interim list of Australian . . Royal Austral. Orn. Union, Brunt, B. Melbourne, vi + 46 pp. 1962. The call of the Spotted Pardalote. Aus- Winkler, Hans, and Lester L. Short tral. Bird-Watcher, vol. 1, p. 204. 1978. A comparative analysis of acoustical Chandler, L. G. signals in pied woodpeckers (Aves, 1961. Victorian pardalotes. Austral. Bird- Picoides). Bull. Amer. Mus. Nat. Hist., Watcher, vol. 1, pp. 153-156. vol. 160, pp. 1-109.

ADDENDUM After completion of the manuscript hybrids (in Australian National Wildlife Col- Schodde, in his review of Australian parda- lection, Canberra) of xanthopygus and punc- lotes for the passerine portion of the Royal tatus taken at Peebinga, South Australia, in Australasian Ornithologists' Union Check- May 1976; prior to 1950 only xanthopygus list, was able to study specimens at the var- had been collected around Peebinga. The ious Australian museums. This study re- Spotted Pardalote frequents wetter, generally vealed evidence of some introgression, and coastal eucalypt forests and woodlands, and hybridization between P. punctatus and P. the Yellow-rumped occurs in drier "mallee" xanthopygus from the Mt. Lofty and south- woodland from the coast to farther inland. ern Flinders Ranges southeast through the The areas where these habitats meet, inter- lower Murray "Mallee" and the Ninety-mile digitate, or intergrade are precisely where the Desert to western Victoria. For example, two pardalotes meet, and it appears that hu- among 73 specimens from the South Austra- man factors (cutting of wetter forests and lian Museum representing the area just men- woodlands, and of mallee, forcing the birds tioned no fewer than 49 blend the morpho- to move elsewhere) are facilitating their con- logical features of both forms to varying tact. degrees and in diverse combinations. This Study of larger series shows a tendency for strongly supports the likelihood of hybrid- disparate sexual dimorphism in these par- ization as a causal factor for the vocal vari- dalotes, for, whereas male and female punc- ation discussed herein. tatus are identical in their buffy back-spot- That shifts in habitat and human-induced ting, female xanthopygus are more coarsely, modifications ofthe environment account for more buffy and less white spotted than males. some ofthe interbreeding is indicated by two This fact and the duller, buffier rump of fe- 1983 SHORT, HORNE, AND SCHODDE: PARDALOTES 15

male xanthopygus suggest that buff coloring from the same transcontinental stock in the is the ancestral condition, which may imply west and spreading east. the origin of xanthopygus from a more punc- Finally, very recent observations ofsinging tatus-like ancestor (following Gloger's Eco- male punctatus make it obvious that the song geographic Rule). is indeed primarily territorial in function. Schodde notes that the Karoonda record- Singing males use an uncluttered vantage ings of Swaby may have been from a hybrid point such as a bare twig just beneath the punctatus x xanthopygus or been influenced sheltering canopy of a small tree. There, they by gene flow from xanthopygus, in view of raise the body to an almost upright, stretching the introgression he has only recently found posture, crane the neck and tilt the head and to be widespread in the Murray Mallee. bill slightly downward in voicing each song- The possibility must be considered phrase, twisting this way then that way as if (Schodde, in recent publications) that both to aim the signal in several directions, or to species discussed herein are indeed Bassian peer in different directions for a prospective in origin, with punctatus arising in the east challenger. and spreading west, and xanthopygus arising