GEOGRAPHIC VARIATION IN SONG AND DIALECTS OF THE PUGET SOUND WHITE-CROWNED SPARROW

Transcription

1 GEOGRAPHIC VARIATION IN SONG AND DIALECTS OF THE PUGET SOUND WHITECROWNED SPARROW LUIS F. BAPTISTA Because of thei simplicity of stuctue and eizines, and the evolutionay and ecological geogaphic vaiation, songs of cowned spa significance of these findings ae discussed. ows (Zonotichia spp. ) have een popula sujects of ioacoustical studies. DeWolfe et METHODS AND MATERIALS al. (974) descied geogaphic vaiation in Songs wee sampled fom localities in Oegon, song of the migatoy suspecies gamelii of Washington and Bitish Columia (Tale, Fig. ), the Whitecowned Spaow (Zonotichia etween 7 June and 4 July 970. Field ecodings leucophys) ut found no dialects. Howeve, wee made on a Uhe 4000 RepotS tape ecode with a Uhe micophone mounted on a 4in. paasong dialects have een descied fo the olic eflecto. Tane sueed was 7. ins. Sons wee sedentay suspecies nutialli ( Male and Ta spectogaphed on a Kay Elemetic Sonogaih Mamua 9, 94, Bake 974, 97, Baptista chine (Model 709A) set at high shape and wide 97) and the migatoy, montane eeding and settings. In the laoatoy songs wee played suspecies oiantha (Bake 97, Oejuela on Ampex and Uhe 4000 Repot C tape ecodes. Some 8 songs fom 70 individuals elonging and Moton 97). Although a few audioto the suspecies pugetensis wee ecoded. Notaspectogams of songs of the migatoy Puget tions wee made of the songs of individuals that Sound Whitecowned Spaow (Z.. pugeten could not e ecoded (Tale ). As in Bantista sis) have een illustated (Baptista 974)) a ( 97), quantitative vaiation in the following chaacteistics wee teated: () duation of each song in detailed study of song in this suspecies of seconds, () nume of elements (syllales plus Whitecown has not een undetaken pevi phases), () nume of syllales in the teminal ously. till, (4) nume of kinds of elements, ( ) epetition Taxonomists distinguish two acial com index, i.e., nume of elements divided y the nume plexes of Whitecowned Spaows, the dake of kinds of elements, () highest fequency in khz, and (7) lowest fequency in khz. nuttallipugetensis goup of the Pacific coast Each ecoded song was assigned a nume. By and the pale gameliiom anthaucophys efeence to a andom nume tale, a single song complex of the mountains and inteio (Banks fom each id was selected fo statistical teatment. 94, Cotopassi and Mewaldt 9). Song If the audiospectogam thus selected poved unclea may e a chaacte sepaating these two su due to some ecoding atifact, I used the next cleanest audiospectogam fom which the aove quantifispecies complexes ( Baptista 97). Howeve, ale chaacteistics wee taken. Individuals wee until songs of all the foms ae studied in de gouped accoding to dialect aea, the latte ased tail, the use of vocalizations as a taxonomic on visual inspection of the teminal tills in each song. tool cannot e evaluated popely. Six dialect ateas wee thus ecognized (Fig. ). Based on osevation of adults attending fled&es. Song dialects of migatoy Whitecowned feeding nestlings, o diecting high intensity ala& Spaows may function as a valuale means calls at the investigato, I concluded that all populafo students of id migation to identify the tions sampled wee eeding. winteing gounds of each eeding population ( DeWolfe et al. 974). Regional dialects TERMINOLOGY of ids may also e used as makes in the Tems and symols used to descie pugetensis song study of inteactions etween populations o ae those of Bantista ( 974) and ae illustated in supopulations (Banks 94: 8, Payne 97, Figue. These* ae defined elow with the symols appeaing in ackets. Baptista 97). Song dialects may e used to ) Note: any continuous tace on the audio spectotace the oigins of invading populations (Bapgam. tista 97, Mundinge 97). Additionally, ) Syllale [S]: the unit of epetition in a till. King (97:44) pointed out that, Geogaph In this pape TRILL efes specifically to the ical vaiation of avian song is of inteest as an teminal tills (Fig. A, B ) symolized y SS. ) Complex syllales [CS]: goups of fou o moe index of speciation. notes foming moe o less coheent units (Figs. 4, This pape teats geogaphical vaiation in F). They follow the intoductoy whistle (song B, song of the migatoy suspecies pugetensis Fig. ) o the intoductoy uzz (song A, Fig. ) in of the Whitecowned Spaow, descies e pugetensis themes. gional dialects fo this taxon, and discusses 4) Whistle [W] : a continuous tace on the audiospectogam geneally exceeding 0. set and constant the consequences of inteactions etween in pitch. This intoduces evey pugetensis song. ovewinteing pugetenzis and esident nutialli. ) Buzz o Viato [B] (of Male and Tamua Data ae compaed with those fo othe em 9) : a fequencymodulated whistle containing 0 The Condo 79:70, 977

2 VARIATION IN WHITECROWNED SPARROW SONG 7 o moe pulses/set, following the citeion of Boo (90). ) Buzzwhistle [BW]: a uzz attached to a whistle, occuing as the second phase in some songs (e.g., H in Fig. 7). 7) Phase: a whistle, uzz, o uzzwhistle. This tem does not include syllales. 8) Song type: used intechangealy with theme and motif to denote all the elements listed aove in vaious pemutations and cominations. Using symols, song A (Fig. ) may e descied as WBCSCSBSS, and song B as WCSBSS. Bitish HABITAT.. pugetensis in my study aeas occupied thee diffeent types of haitat (Tale ) : RESIDENTIAL AREAS As in sedentay nuttalli (Blanchad 94, Banks 99, Male and Tamua 9, Baptista 97), pugetensis often was found in paks and gadens. Shuey was used fo nest sites, lawns povided foaging aeas, and tall tees and oof tops often seved as singing peches. FOREST CLEARINGS AND EDGES Lewis (97) studied pugetensis on Camano Island, Washington and pointed out that logging activity poaly inceased haitat availale to Whitecowned Spaows. Recentlylogged aeas, in ealy successional stages ecause of gazing, povide eeding haitat fo the species. Banks (94:4) epoted on populations of pugetensis nea the Noth Santiam Rive, Oegon, that had invaded ecentlylogged aeas. I found these spaows at the ecotone etween conifeous foests and the gassy aeas on the Pacific coastal sand dunes (e.g., at Fot Stevens, Tale ). On San Juan and Vancouve islands they often occupied foest cleaings such as those descied y Lewis and Banks. SAND DUNES Lage aeas of the Oegon and Washington coastal dunes ae coveed y thick mats of gasses (e.g., Ammophila aenaia, Festuca ua, Poa macantha) and sedges (Caex sp.). These gasslands may e dotted with small conifes (Pinus contota, Picea sitchensis), shus such as Scottish oom ( Cytisus sp.), lupines ( Lupinus littoalis, L. aoius), and coyote ush (Bacchais pilulais). At some localities (e.g., Ocean City, Washington) small tees and ushes ae ae, so that Whitecowned Spaows sang fom the gound o used pieces of diftwood as singing _.. Ocedn City.iwin Haos.,. Leadette+paint,...;:,...I..,: F<Lewis s......i Oegon miles, ::,:,:..,.,.; 0 0 IO....Newpot ;.. ; l ::::: 0 dialect I._. _..... (. :.::._ :: l dialect... A dialect Adialect 4 A dialect A dialect FIGURE. Map of localities sampled. Round symols signify populations singing southen themes. Tiangula symols designate populations singing nothen themes. Diffeent pattens epesent diffeent dialects. posts. Nests wee poaly uilt on the gound. At Fot Cany and Leadette Point, Washington, willows (SaZix sp.) ae a vey conspicuous pat of the floa. Haitat at these localities is eminiscent of that occupied y montane Whitecowned spaows,.. oiantha (DeWolfe and DeWolfe 9, Moton et al. 97). Floa and topogaphy of the Oegon coastal dunes have een teated in detail y Wiedemann et al. ( 99). At some localities sand

3 8 LUIS F. BAPTISTA < Till.+ kh B F Whistle \ Complex + Syllale j CBun + h Till > seconds FIGURE. A nothen theme (A) and southen theme (B) illustating teminology used. TABLE. Numes of ids singing nothen (N) and/o southen (S) themes. Localities N Both S Hyids Dialect No. Bdg. Noted Hait& Bandon Beach Bullad Beach Coos Elay Newpot Tillamook Oceanside Cape Meaes Manzanita Tolovana Cannon Beach Ecola Seaside Geahat Astoia Fot Stevens Fot Cany Long Beach Leadette Pt. Tokeland Twin Haos Ocean City Pacific Beach Dosewallips Rainow Lodge Eldon Fot Lewis Tacoma Seattle Anacotes San Juan Island Butchad Gadens Nanaimo June. IHighway 4 Comox Cleveland Dam Stanley Pak () ;;, 4 4 () 4(l) (l) 4 &) 4 (l) (4) l!() :() (l) ) 7, t 4 4, t: i ;, f ; d I if d d : :: f, f, ; ; f Totals W8) (l) 80( ) (l) = attending nestlings o fledglings. cl z dunes; f = foest cleaings o foest edge; = esidential aeas. A numes in paentheses epesent sutotal of ids head ut not ecoded.

4 VARIATION IN WHITECROWNED SPARROW SONG 9 4 c FIGURE. Fou dialects within nothen _. _ themes: a: two motifs ecoded at Fot Cany, Washing ~~~ ton; 4a4: two songs ecoded at Tokeland, Washington; a: a song ecoded at Ocean City, Washington; : a theme fom Seattle. Washington: ad: fou motifs fom San luan Island, Washington. Note the vaiation in the mophology of theteminal syllales in the fou songs. dunes occu etween the Pacific Ocean and the gassy stetches occupied y the spaows. At othe localities (e.g., Fot Cany), gassy aeas and the attendant spaows wee found almost to the wate s edge. DeWolfe and DeWolfe (9) descied and compaed haitats fo thee taxa of Whitecowned Spaow, including the suspecies pugetensis. All haitats encounteed in my study oe the chaacteistics listed y these authos, namely, gassland, ae gound, and shuey. The dunes occupied y pugetensis wee the most open of all the haitats encounteed as ushes and small tees wee patchy and vitually asent in some aeas. THEMES OF PUGETENSIS The song of Z.. pugetensis is aout two seconds long and etween two and eight khz in fequency. Based on diffeences in the sequencing of whistles, uzzes, complex and simple syllales, and distinct complex syllale types, I distinguished two geogaphical goups of songs; these will e efeed to as Nothen and Southen themes. THE NORTHERN THEME Typically, the nothen theme egins with a whistle and is followed y a uzz, two complex syllales, anothe uzz, and a teminal till ( WBCSCSBSS; Fig. A). Some ids sang songs containing only one complex syllale (WBCSBSS; Fig., song ). TWO ids sang songs (not illustated) with thee complex syllales ( WBCSCSCSB). A few individuals sang songs lacking the fist uzz which nomally constitutes the second phase (WCSCSBSS; Fig., song ). The complex syllales and teminal tills vay geogaphically and will e teated in detail. THE SOUTHERN THEME In contast to the aove, the typical southen theme egins with a whistle and is followed y a complex syllale, a uzz, and finally a till ( WCSBSS; Fig. B). The complex syllales in southen themes ae quite unlike those in nothen themes (compae Figs. 4 and ) and will e discussed late. The teminal tills in southen themes ae moe o less population specific.

5 0 LUIS F. BAPTISTA DISTRIBUTION OF NORTHERN AND SOUTHERN THEMES Figue illustates the distiution of the two song types. With a few exceptions ids eeding in the sampling localities fom Bandon to Ecola, Oegon, sang the southen theme (dialects and ). Along the Pacific coast, ids eeding at Fot Stevens and Astoia, Oegon, and ids eeding in Washington fom Fot Cany to Pacific Beach sang nothen themes (dialects, 4 and ). Bids sampled at localities in Puget Sound sang nothen themes (dialects and ). The latte included populations on the mainland of Washington and Bitish Columia as well as ids on San Juan and Vancouve islands. The southen theme is thus distiuted in almost pue fom fo at least 00 mi of the Pacific coastline (Fig. ). The nothen theme anges ove 40 mi of this same coast. Bids singing nothen themes fom mainland localities on Puget Sound ange at least 70 mi. On Vancouve Island, the same song theme occus fom Butchad Gadens to Comox, a distance of at least 0 mi. CONTACT AREAS BETWEEN NORTHERN AND SOUTHERN THEMES At Geahat and Seaside, Oegon, ids singing nothen o southen themes occued togethe (Tale ). Thee ids sang oth themes. The two song types appea to occu sympatically along at least 0 mi of coastline. One ilingual id ecoded at Seaside, Oegon on June 970 was oseved matching nothen and southen themes sung y fou neighos. I tied to get it to match songs played fom my tape ecode ut was unsuccessful. In contast, a id (pug&e&s) ecoded at Cescent City, Califonia, and seveal ids (nuti&) ecoded in the San Fancisco Bay aea, Califonia, sang two song types each and would match themes sung y neighos as well as playack of ecoded song (Baptista 97). At Ecola, Oegon, I could find only five singing Whitecowned Spaows. Fou ids sang typical southen themes, and one id sang two atypical nothen themes (Fig. 7) descied late. REPERTOIRE SIZE AND VARIATION WITHIN THEMES Five ids out of 9 sang two themes, each theme containing diffeent syllale types. The est sang one theme each. This is simila to data fo othe taxa of Zonotichia (King 97, DeWolfe et al. 974, Oejuela and Moton 97) and fo anothe emeizine, the Otolan Bunting, (Emeiza hotulana; Conads and Conads 97:9). Although I did not ecod any ids who sang thee themes, Banks (94) epoted such an individual of pugetensis. A id ecoded at Tokeland, Washington, sang a theme with a whistle and a uzz in the intoduction ( WBCSCSBSS) and a vaiant of the latte with the intoductoy uzz asent ( WCSCSBSS). Five individuals each sang a song with two complex syllales (WBCSCSBSS) and vaied it y sometimes dopping one complex syllale (WB CSBSS). One id ecoded at Seaside, Oegon, would haitually dop one of its complex syllales when eplying to playack of its own song. A nuttdi in Bekeley, Califonia, misimpinted with a pugetensis theme, ehaved likewise ( Baptista 974). Except as noted aove, vaiation within each nothen o southen theme consisted mostly of lengthening o shotening the intoductoy whistle, lengthening o shotening the viati, o vaying the nume of syllales in the teminal tills. This degee of steeotypy has een epoted fo othe aces of Whitecowned Spaow, namely, nuttdi (Male and Tamua 9, Konishi 9, Bake 974, Baptista 97), gamelii (DeWolfe et al. 974) and oiantha (Oejuela and Moton 97). As an example of individual vaiation, audiospectogams of songs ecoded fom one id at Anacotes, Washington, yielded the following statistics: duation in seconds :.8.99 (X.9 f 0.47 s.d.); no. of elements: 4 (ii.4 * 0.79); no. of teminal syllales: 9 (ii ); no. of kinds of elements: 4 in all audiospectogams; epetition index:.7.0 (ii.0 C 0.0) ; highest fequency (khz) :.7.00 (.8 0.); lowest fequency:.00 khz in all audiospectogams. SONG DIALECTS NORTHERN THEMES Geogaphic vaiation in teminal tills. Data on the geogaphic distiution of teminal tills ae summaized in Tale and illustated in Figue. Teminal till a (song a, Fig. ) occued in pue fom fom Astoia, Oegon to Long Beach, Washington (Tale, Fig. ). At Seaside and Geahat, Oegon, these songs wee found sympatically with southen themes (Tale ). Till 4a (Fig. ) was ecoded in Washington fom Leadette Point to Twin Haos.

6 VARIATION IN WHITECROWNED SPARROW SONG TABLE. Distiution of eight teminal till types. Localities & 9 4a a a c d Seaside Geahat Astoia Fot Stevens Fot Cany Long Beach Leadette Pt. Tokeland Twin Haos Ocean City Pacific Beach Dosewallips Binnon Eldon Fot Lewis Tacoma Seattle Anacotes San Juan Island4 Butchad Gadens Nanaimo4 June. Highway 44 Comet Cleveland Dam Stanley Pak 4 4 i Total ltemina tills of southen themes as in song Za, Fig.. a to d ae the teminal tills of the nothen themes in Fig.. Dosewallips is the state pak on the outskits of Binnon. Vancouve Island. Vancouve City. At Ocean City two ids used this teminal till wheeas used till nume (song a, Fig. ). Songs 4a and 4 (Fig. ) illustate individual vaiation in the mophology of the syllales making up the teminal tills in a population; the inveted V shaped note is lage in song 4 than in song 4a. Teminal till a (Fig. ) was ecoded at two localities along the Pacific coast and six localities on Puget Sound (Tale ) fom Dosewallips to Seattle. On San Juan Island in Puget Sound, fou vaiants of the teminal till wee pesent (Fig., songs a to d). The two vetical lines in the teminal syllales of song a ae appoximately the same length. In song the second vetical component in the teminal syllale is noticealy longe than the fist. In song c, the second longe vetical am appeas thickened nea the ottom extemity. In song d the thickened potion is twisted so that each syllale teminates with a small inveted U. I found no evidence of clumped distiution ( sudialects ) of these teminal syllales on San Juan Island. Howeve, on Vancouve Island, till was found to the south (Butchad Gadens, Tale ) and tills c and d in moe nothely localities (Nanaimo to Comox). The single id ecoded at Anacotes used till a, and seven of eight ids ecoded at Vancouve City (Cleveland Dam and Stanley Pak) used till d (Tale ). One id in Stanley Pak, Vancouve City, used till. Tills a to d, theefoe, seemed to eappea in geogaphically distant localities on San Juan and Vancouve islands as well as Anacotes and Vancouve City on the mainland nea Puget Sound. Thus, wheeas tills a to a ae quite unifom in mophology within populations, teminal syllales epesented in songs fom the nothen Puget Sound populations ae moe vaiale, fou vaiants having een ecoded (tills a to d). Geogaphic vaiation in complex syllales. Two complex syllales nomally follow the two intoductoy phases in the typical nothen theme (WBCSCSBSS), Usually oth complex syllales ae mophologically simila, ut in a few cases they wee unlike (Fig., song ~). Eight complex syllale types wee epesented in the songs sampled (Fig. 4, to 8). Syllale types 9 to (Fig. 4) ae simple in stuctue and ae meely the fist susyllales of syllales one, thee and fou, espectively. Most complex syllales ae distiuted in mosaic fashion, i.e., the same syllale type may e epeated in geogaphically distant loca

7 LUIS F. BAPTISTA IA A 4 FIGURE 4. 8: Complex syllales and thei vaiants fom nothen themes. Syllale A is a vaiant of syllale, diffeing slightly in the mophology of the second susyllale. Syllale A is a vaiant of syllale, again diffeing in the mophology of the second susyllale. The latte is an inveted V in syllale and a steak in syllale A. 9: Susyllales of complex syllales used y some individuals. They epesent fagments of complex syllales, and 4, espectively, which appeaed in place of complex syllales in some songs (e.g., song c in Fig. ). of sample). Data fo dialect ae, theefoe, simila to those found in nuttalli in which vaiaility within populations was also low (Male and Tamua 9, 94, Bake 97, Baptista 97). Complex syllale fist makes its appeaance at Tokeland, Washington (Tale ) and continues to occu until Comox, Vancouve Island. At Ocean City, Washington, syllale occued in the songs of of 4 ids (.7% of sample). On San Juan Island, Washington, seven of the eight complex syllales illustated in Figue occued (9% of sample). Type was the dominant complex syllale on the island ut was found in only ids (40% of the sample). This syllale type was found at no othe locality. Syllale type 8 occued only once (Fig., a), in a song fom San Juan Island. Howeve, Bake (974, Fig. ) illustated songs of nuttalli fom San Mateo Co., Califonia containing syllales simila to type 8 (Fig. 4, this study). SOUTHERN THEMES tions (Tale ). Nonetheless, a geogaphical Geogaphic vaiation in teminal til.s. In tend is evident. Syllale type (Tale ) song la (Fig. ) all the syllales in the temiis the dominant fom in the populations fom nal till ae mophologically simila. In songs Seaside, Oegon to Long Beach, Washington. l to Id, the fist syllale in the teminal till Most ids singing dialect also use syllale is moe gently sloping (i.e., slued diffetype (4 out of 48 ids ecoded o 9.7% ently) fom the est of the syllales in the TABLE. Fequency of occuence of syllales in nothen themes. Syllale numes coespond to those in Figue 4. Localities Othe Dialect# Seaside Geahat Astoia Fot Stevens Fot Cany Long Beach Leadette Pt. Tokeland Twin Haos Ocean City Pacific Beach Dosewallips Binnon Eldon Fot Lewis Tacoma Seattle Anacotes San Juan Island Butchad Gadens Nanaimo June. Highway 4 Comox Cleveland Dam Stanley Pak z * z Total 9 * Theme Hl in Fig. 7.

8 VARIATION IN WHITECROWNED SPARROW SONG Id I FIGURE. la to Id: fou vesions of dialect. Songs la to lc wee ecoded on Bandon Beach, Oegon. Song Id was ecoded on Bullad Beach, Oegon. a to d: fou vaiations of dialect. Song a was ecoded at Tillamook, Oegon. Song was ecoded at Cape Meaes, Oegon. Song c is fom Manzanita, Oegon. Song d is fom Newpot, Oegon. till. All the syllales, howeve, ae similaly shaped. In songs a to d, (Fig. ) the fist two syllales following the uzz ae moe elaoately stuctued than the est of the syllales in the teminal tills. These fist syllales may e notched (songs a and, Fig. ) o may consist of paied notes (songs c and d, Fig. ). The est of the syllales in these tills ae simple in stuctue, and mophologically simila to each othe and to the syllales in the tills of songs la to Id. Although the teminal simple syllales ae simila in all southen themes sampled fom Bandon Beach to Geahat, Oegon (Fig. l), ased on the pesence o asence of the two moe elaoate paied o notched syllales immediately following the uzz (Fig., songs a to d), two dialect egions may e distinguished. Dialect anges fom Bandon Beach to Coos Bay, Oegon and dialect fom Newpot to Geahat, Oegon (Tale 4, Fig. ). In addition to diffeences in syllaic stuctue, the caie fequencies in the viati found in song dialect (Fig., songs a to d) vay little, wheeas the uzzes in dialect ae noticealy inflected downwad (Fig., songs la to Id). Song dialects in Z. cape&s (Notteohm 99, 97, King 97) and Z.. nuttalli (Male and Tamua 9, Baptista 97) ae ased mainly on the stuctue of the syllales in the teminal potions of the till. Dialects in southen pugetensis themes (this study), howeve, ae distinguished on the asis of elements in the middle potions of the song (complex syllales, uzzes, paied syllales). In this espect southen pugetensis dialects ae eminiscent of some song dialects of the Shottoed TeeCeepe ( Cethia hachyductyla; Thielcke 9). Geogaphic vaiation in complex syllales. I distinguish six complex syllale types in southen themes ( Fig. ). Type diffes fom type in the shape (fequency span) of each note, e.g., the fist note is longe in, ut the last note is longe in type. Although the diffeences ae small, I teat and as sepaate categoies ecause each occued 8 o moe times and ecause thei elative fequencies vaied acoss populations. Types A, B, and C ae clealy vaiants of type. Complex syllale B occued only once, in the song of a id ecoded at Seaside, Oegon; complex syllale C also was found only once, in the song of a id ecoded at Cape Meaes, Oegon. Numes of ids using syllale types, A, B, and C wee pooled in Tale 4. Syllales 4 and 4A ae also vey simila. Syllale 4A was found in only two ids, ecoded at Newpot, Oegon. Numes of ids using 4 and 4A wee also pooled in Tale 4. Complex syllale types to 4A (Fig. ) ae distiuted geogaphically in mosaic fashion (Tale 4), occuing in diffeent fequen

9 4 LUIS F. BAPTISTA TABLE 4. Fequency of occuence of complex syllale in southen themes. Syllale numes coespond to those in Fig.. A c Localities 4 7 Othe Dialect # Bandon 0 l Bullad s Beach Coos Bay Newpot 7 4 Tillamook Oceanside I Cape Meaes Manzanita sconds Tolovana Cannon Beach Ecola Seaside 4 Geahat Fot Cany ld Total 8 4 a One hyid theme (Fig. 7, H) in this sample. Hyid theme (Fig. 7, H). c Bids epesented y unclea spectogams identifiale only as southen themes. d Hyid theme (Fig. 7, Hl). ties in widely sepaated populations. With only one exception (song Hl in Fig. 7), these complex syllales wee used only y ids singing dialect (Tale 4). Complex syllales and (Fig. ) wee used solely y ids in dialect aea one (Tale 4). Although stuctually simila to complex syllales and, complex syllale 7 was found only in the songs ecoded fom two ids at Newpot, Oegon (Tale 4), i.e., in dialect aea two. It is conceivale, theefoe, that with moe samples, especially fom localities intemediate etween dialect aeas one and two (i.e., etween Coos Bay and Newpot, Fig. l), all the complex syllales in Figue may e shown to occu with teminal tills of oth dialects and. It is nonetheless notewothy that on the asis of pesent data the two dialect aeas have no complex syllales in common (Tale 4, Fig. ). Geogaphic vaiation in the intoductoy phases. DeWolfe et al. (974, Fig. ) distinguished thee types of intoductoy whistles (Phase A of Male and Tamua 9) in the songs of.. gamelii. These wee designated Continuous, Segmented, o Boken. Phase A in pugetensis themes is emakaly unifom geogaphically (Figs.,, and 7). These whistles wee all segmented due to amplitude modulation, appeaing in audiospectogams as thee moe o less staight dak lines joined y two lighte aeas. Fequency modulation was slight in these whistles. Audiospectogams in Oejuela and Moton (97) and Bake (97) indicate that FIGURE. Complex syllales and thei vaiants fom southen themes. intoductoy whistles in songs of oiantha ae simila to those in pugetensis. In contast, Phase A in nuttalli in the San Fancisco Bay aea may show geat individual vaiaility in oth amplitude modulation and fequency modulation (Baptista 97, 97). This was especially evident in the songs fom Teasue Island, San Fancisco Bay. The second intoductoy phase (Phase B of Male and Tamua 9) is also sometimes quite vaiale in nuttalli themes. These ae most commonly a pue whistle o a uzz, and sometimes uzzwhistles, whistleuzzes, o shot tills (staccato phases of Baptista 97). Phase B is asent in southen themes of pugetensis (Fig. ). Howeve, within the nothen themes, with only thee exceptions descied late, Phase B when pesent was a viato in pugetensis motifs. Whistles and uzzwhistles as second phases wee quite ae in the populations sampled. EXCEPTIONAL SONGS Although most individuals in a song population sang one of the local themes teminating with tills chaacteistic of goups of populations, a few exceptions wee encounteed. These ae descied elow, accoding to locality. VARIANT THEMES Seaside, Oegon. Eleven ids in this population sang typical southen themes (WCS BSS). One othe individual sang a song sustituting the uzz with a uzzwhistle ( WCSBWSS; song VI, Fig. 7). This uzzwhistle was the only one of its kind encounteed. Newpot, Oegon. Eleven of 4 ids ecoded at this locality sang the local southen theme ( WCSBSS). These songs typically

10 VARIATION IN WHITECROWNED SPARROW SONG I seconds : FIGURE 7. Vl to V ae vaiant themes. Vl ecoded at Seaside, Oegon. V and V ecoded at Newpot, Oegon. Hl to H4 epesent some hyid themes. Hl ecoded at Fot Cany, Washington. H and H ecoded at Bandon, Oegon. H4 ecoded at Ecola, Washington. ended with a till of simple syllales, each consisting of one downwad inflected note (Fig., a to d). Howeve, thee ids sang songs ending with syllales consisting of paied notes; one id sang songs with teminal syllales illustated in V (Fig. 7), and two ids sang theme V. A typical teminal till is illustated in Vl (Fig. 7) fo compaison. Astoia, Oegon. One id sang a song (not illustated) with two whistles in the intoduction (WWCSCSBSS). This was the only song of this kind ecoded (see also section on geogaphic vaiation in phases). Although this individual sang an unusual theme, it was mated and was oseved feeding a fledgling. HYBRID THEMES Fot Cany, Washington. Thiteen of 4 ids sampled sang nothen themes. One id (Hl in Fig. 7) sang a theme which appeaed to e a typical southen theme with egad to sequencing of elements in the intoductoy potion (WCSBSS). Howeve, the last thee teminal syllales wee mophologically simila to those in the local nothen themes (compae with song a, Fig. ). The long downwadinflected uzz was poaly also oowed fom a nothen theme (compae with second uzz in song a, Fig. ). Bandon, Oegon. Twelve of 4 ids ecoded at this locality sang one of the local southen themes (Fig., songs la and l). Two individuals sang what appeaed to e hyid themes consisting of elements fom nothen and southen themes comined. One of these (H in Fig. 7) sang a song which egan with a whistle, followed y a type (Fig. ) complex syllale, a uzz, and then thee type (Fig. 4) complex syllales. The last thee (type ) complex syllales poaly wee oowed fom a nothen theme, and the intoductoy (WCSB) potion fom a southen theme. The second id (H in Fig. 7) sang a theme which appeaed to e a typical nothen theme with egad to sequencing of elements, i.e., WBWCSCSBSS. The two complex syllales ae typical of nothen themes (type, Fig. 4). Howeve, the last two simple syllales ae typical of the local southen themes (see song la, Fig. ). The uzzwhistle compising the second phase in H s song is atypical of pugetensis. It was the only one of its kind encounteed. Ocean&&, Oegon. One id that sang infequently and could not e located fo ecoding, sang a theme simila to H (Fig. 7). Ecola, Oegon. Fou ids sang typical southen themes; one id, howeve, sang theme H4 (Fig. 7). This egins with a whistle, followed y a uzz, a complex syllale, and then a till of simple syllales. The complex syllale is typical of nothen themes (type, Fig. 4). The teminal syllales, howeve, ae typical of the dominant local southen theme (compae with teminal syllales in theme C, Fig. ). This id sang a single second theme (not illustated) which ap

11 LUIS F. BAPTISTA peaed to e an incomplete nothen theme ( WBCSCS ). This individual appeaed to e mated. In pugetensis, as in nuttalli (Baptista 974, 97), then, most ids in a eeding population sing a song identifiale as one of the local themes. Howeve, a few ids sing vaiant themes modifying some of the phases and syllales. Still othes impovise y comining elements fom nothen and southen themes and constucting hyid themes. Among emeizines, impovisation of this kind is ae in Whitecowned Spaows (Baptista 97) and Otolan Buntings (Eme&z hotulanu; Conads and Conads 97:9). Howeve, syllale oowing appeas to e the ule in song development in some populations of Song Spaows (Melospiza mlodia; Hais and Lemon 97, Eehadt and Baptista 977). In nuttalli, hyid themes wee found only in zones of seconday contact whee two paental themes wee found sympatically (Baptista 97). Howeve, hyid pugetensis themes descied aove wee found some distance away fom seconday contact zones. How then wee these themes acquied? Fist, these ids may have hatched in aeas of seconday contact etween two dialects whee they developed thei hyid themes and then dispesed to the places whee they wee ecoded (Tale ). Second, they could have picked up elements of a second theme fom migants in passage. Thid, individuals in the sampled localities could e singing othe dialects not encounteed y the investigato. These second themes could have influenced song development in some individuals, giving ise to hyid themes. Fouth, if some leaning in some individuals takes place eyond a shot citical peiod (typical of nuttalli, Male 970), these individuals could oow syllales fom flock mates duing migation o on thei winteing gounds. QUANTITATIVE ASPECTS OF SONG VARIATION Milligan and Vene (97) demonstated that Whitecowned Spaows may distinguish etween song dialects. Baptista (97) found that individuals can distinguish themes within dialects, individuals with ivalent epetoies matching specific themes duing playack expeiments. Fequency and/o tempoal chaacteistics of avian song often seve fo ecognition of species o individuals. (Falls 9, Emlen 97, Books and Falls 97). Tempoal and fequency paametes of song fu

12 VARIATION IN WHITECROWNED SPARROW SONG 7 nish hythms and pitch diffeences which could function along with syllale mophology in ecognizing dialects o themes. To quantitatively examine song vaiation, I summaized data fom seven paametes fo each of the six dialects (Tale ). If song dialects act as ethological aies to pai fomation etween dialect aeas (Notteohm 99), it would e meaningful to test diffeences etween dialects that ae actually geogaphically juxtaposed in natue. Thus, as in Baptista (97) I conducted univaiate tests etween neighoing pais of dialects fo the seven paametes measued (Tale ). Based on ttests, dialect pais diffeed in fou o five of the seven quantified chaacteistics. Although dialects and seemed on aveage to e iefe than dialects to, ttests demonstated thei diffeences to e insignificant (Tale ). Song length, theefoe, is simila in all population pais of pugetfmsis tested. Although ttests showed the diffeences in mean nume of elements etween dialects and to e significant (Tale ), I do not elieve this to e iologically eal. Having spent many hous in the field with Whitecowned Spaows, I elieve that seveal ids in populations and wee not singing complete songs, i.e., when the ids wee fully motivated thei songs would have contained moe teminal syllales, thus moe elements, and consequently would have lasted longe. Significant diffeences etween populations tested wee found in the () nume of elements and kinds of elements, () epetition index, () highest, and (4) lowest fequencies. These data diffe fom those in viduine finches (Vidua spp.) in which Payne ( 97) found no diffeences in the same paametes tested acoss dialect populations. DISCUSSION COMPARISONS WITH NUTTALLI Song dialects of sedentay nuttalli studied y Male and Tamua (9, 94), Bake ( 97), and Baptista ( 97) ae vey local in distiution. Song dialects of migatoy pugetemis ae moe widespead (Fig. ). In this espect the latte ae eminiscent of o~iantha, studied y Bake (97) in Coloado. Channels of wate 0. to. mi wide appea to e effective aies to the dispesal of nuttalli in the San Fancisco Bay aea of Califonia, so that distinct song dialects occu on islands in the Bay (Baptista 97). Pugetensis ecoded on San Juan and Vancouve islands in Puget Sound sang the same themes as ids taped at Anacotes and Vancouve City on the Bitish Columia mainland (Fig., Tale ), suggesting some dispesal etween island and mainland populations. Contact zones etween neighoing nuttalli dialects ae often naow (Bake 97, Baptista 97). In contast, the one contact aea etween nothen and southen themes of pugetensis, studied in detail heein, is at least 0 mi wide (Tale, Fig. ). Whee nothen and southen pugetensis themes occu sympatically, a few ids sing oth song types. In nuttalli a few ids also sang two dialects in zones of seconday contact (Bake 97, Baptista 97). Bake ( 97) found ids singing two dialects ealy in the eeding season ut not late. He could not detemine whethe ilingual individuals had moved o had emained and failed to use one song type. One possile explanation fo Bake s osevations is that ecause of countesinging and song matching, the moe common theme in the aea is used moe and moe often y individuals with ivalent epetoies, and the ae song is soon left unused (Baptista 97). The nume of ilingual individuals in oth pugetensis and nuttalli may thus e highe than that epoted y field investigatos. PUGETENSIS AS SINGING TUTORS OF NUTTALLI Pugetensis aive at the San Fancisco Bay aea aout midsepteme and do not leave until ealy o midapil (Blanchad 94, 94). Fall aival is often announced with song (Blanchad 94, Davis 98:4). In late Feuay, aout fou weeks pio to migation, song egins to e moe fequent and loude ( Blanchad 94, DeWolfe 98). Bids elonging to the sedentay suspecies nuttalli, hatched ealy o late in the eeding season in the San Fancisco Bay aea would thus hea the songs of winteing pugetensis o ids in passage. As a esult, eight of nine esident teitoial individuals ecoded in the San Fancisco Bay aea singing pugetensis themes wee identified as nuttalli misimpinted with songs sung y migants (Tale ). One of these (P) could have een a pugetemis that emained in its winteing gounds though the following sping. Seven of the nine individuals sang nothen themes and two sang southen themes (Tale ). Because complex syllale types,, and wee the most often encounteed in nothen themes of pugetensis (Tale ), it is not supising that the complex syllales used y misim

13 8 LUIS F. BAPTISTA TABLE. Summay of data on songmisimpinted nuttdi in Califonia. Individual COUllty l Complex Syllale Type Yeas 7c Till@ ecoded Pl Main P* Conta Costa P Alameda P4 Alameda P Conta Costa P Alameda P7 Alameda P8g Alameda Pgh San Fancisco Total + T a 99 a 99 a 9870 none 97 + none 97 + a 97 none 97 + a 974 a 97 :* Data fom Baptista 974 unless othewise noted. See Fig. 4 fo illustations of these syllales. c See Fig. fo illustation of this syllale. d Teminal tills as in Figs. and. e Vaiant of type. f Suspecies unknown. g Data fom Baptista and Wells 97. h Data fom Baptista, unpul. pinted nuttalli singing nothen themes wee also of these widespead foms (Tale ). Thee ids sang pugetensk themes with no teminal tills (Tale ). Pehaps as a esult of low andogen levels, visitant pugetewis fequently sang patial songs. The thee nuttalli in question poaly had incomplete pugetensis songs as models fo mimesis (Baptista 974). Pl sang a nothen pugetensis theme ending with potions of a southen pugetensis till. Pl and P4 occasionally tacked on the local nuttalli tills to thei pugetensis motifs. The othe six ids sang complete songs typical of eeding populations on the Pacific coasts of Oegon, Washington and Puget Sound. Banding studies have shown that Pacific coastal and Puget Sound populations of pugetensis do indeed winte in the San Fancisco Bay aea (Blanchad 94, Cotopassi and Mewaldt 9). Themes sung y winte visitants matched those sung y the misimpinted nuttalli (Baptista 974). Because occasional pug&en& may emain on thei winteing gounds the following eeding season (Blanchad 94), and ecause song is not necessaily a aie to pai fomation, I suggested that pugetensis may occasionally mate with female nuttalu encounteed on thei winteing gounds (Baptista 97, 974). SONG DIALECTS AND THE STUDY OF MIGRATION Out of, Whitecowned Spaows anded ove 4 yeas, only I98 wee ecoveed (Cotopassi and Mewaldt 9). Only six song dialects wee distinguishale in the many hundeds of miles of pugetewis haitat. Vaiation within populations was low as in nuttalli. It would seem possile, theefoe, to identify the oigins of populations of pugetewk on thei winteing gounds y analyzing thei songs. Peequisite to this would e () demonstating the staility of these song dialects fom yea to yea, () ecoding songs fom the isolated inteio populations, e.g., in the Willamette Valley, () otaining ecodings of ids in the gaps in my tansect, e.g., etween Coos Bay and Newpot o etween Newpot and Tillamook, Oegon. GENETIC STRUCTURE OF POPULATIONS Individual vaiation in the song leaning pocess is well documented and has een intepeted y some authos as impefect copying (Male 9:, Lemon 9, Payne 97, Koodsma 974, Baptista 97), and y othes as a positive pocess temed dift (Hais and Lemon 97, Lemon 97). Thee is aundant evidence of individual vaiation in theme copying in the songs of pugetends, giving ise, fo example, to vaiants of the complex syllale types (Figs. 4, )) o to occasional individuals singing songs missing a second phase (Fig., ). Given geogaphic isolation, oal tadition and the passage of time, these mino deviations may accumulate, esulting in lage diffeences and new distinct dialects. Because the diffeences etween dialects (teminal tills) ae smalle than the diffeences etween the nothen and southen themes (mophology of complex and simple syllales and sequencing of elements), I postulate that dialect popula

14 VARIATION IN WHITECROWNED SPARROW SONG 9 tions wee sepaated fom each othe fo a shote time than the populations singing nothen vesus southen themes. It is tempting to speculate, theefoe, that the genetic diffeences etween neighoing dialect populations ae smalle than the genotypic diffeences etween populations singing nothen vesus southen themes. This hypothesis may pehaps e tested in a study of potein polymophisms such as Notteohm and Selande (97) and Bake (974, 97) have employed fo seveal taxa of Zonotichia. SONG DIALECTS IN RELATION TO HABITAT Notteohm (99, 97) and King (97) found that each song dialect in Zonotichia cape&s coesponded to a diffeent haitat. Bake (97) and Baptista (97) found no coelation etween haitat and song dialects in sedentay Z.. nuttalli. Thee appeas to e no coespondence etween song dialects and haitat in migatoy pugetensis. GENETIC RELATIONSHIPS OF THE PUGET SOUND POPULATIONS Banks (94:4) discussed these populations and stated that, The Puget Sound populations poaly do not epesent ecent invasions fom moe stictly coastal goups. Geogaphically and mophologically they ae oviously closely elated to the Vancouve Island ids. Based on the mophology of syllales constituting the teminal tills, I distinguished two song dialects on Puget Sound. Songs ecoded fom Vancouve City and Anacotes wee simila to themes fom San Juan and Vancouve islands. Howeve, motifs ecoded fom Dosewallips to Seattle on the southen shoes of the Sound wee simila to songs fom Ocean City and Pacific Beach on the Pacific coast. This suggests that the Puget Sound populations may epesent at least two distinct gene pools. The nothen mainland populations (Anacotes, Vancouve) may e genetically simila to those on Vancouve Island as poposed y Banks (94). Howeve, the southen mainland populations (Dosewallips to Seattle) may e genetically close to Pacific coast populations. SUMMARY Geogaphic vaiation in song, and song dialects wee studied in eeding populations of the migatoy suspecies pugetensis of the Whitecowned Spaow. Using mophology and sequencing of elements, two widely distiuted song themes wee distinguishale. At zones of seconday contact etween these two song types a few ids wee ilingual. Syllales in the medial potions of the songs tended to eoccu in mosaic fashion ove hundeds of miles. With few exceptions, syllales in the teminal tills wee chaacteistic of goups of adjacent populations thus pemitting the distinction of six dialect aeas. Vaiation within populations in five of these dialects was low, eminiscent of the dialects of nuttalli. Song dialects of nuttalli ae moe local in distiution than those in pugetensis. A few nuttalli may occasionally lean patial o complete songs fom ovewinteing pugetensis. Based on song mophology, I suggest that dispesal acoss wate aies poaly has een moe fequent in pugetensis than in nuttalli. In contast to Zonotichiu cape&s, I could find no coespondence etween song dialect and specific haitat types in pugetensis. The possile use of song as a tool in the study of migation in pugetensis was teated. Finally, I comment on the possile elevance of song vaiation to the genetic stuctue of populations of pug&e&s along the Pacific coast and in Puget Sound. ACKNOWLEDGMENTS I thank Baaa DeWolfe, Ned K. Johnson, Matin L. Moton, Paul Mundinge, and Lewis Petinovich who ead the manuscipt and offeed valuale comments. Field wok was made possile v a gant fom the Fank M. Chapman Memoial Fund OF the Ameican Museum of Natual Histoy. Audiospectogaphic wok was conducted in the Mooe Laoatoyof Zoology. Caol Beuchat and Eic Dawson assisted in the nenaation of the figues. Gene M. Chistman p&&d the map. This study was conducted while I was a student at the Museum of Veteate Zoology, Univesity of Califonia, Bekeley. LITERATURE CITED BARR, M. C Genetic stuctue of two populations of Whitecowned Spaows with diffeent song dialects. Condo 7:. BAKER, M. C. 97. Song dialects and genetic diffeences in Whitecowned Spaows (Zonotichia leucophys). Evolution 9#:4. BANKS, R. C. 99. Development of nestling Whitecowned Spaows in cental coastal Califonia. Condo :909. BANKS, R. C. 94. Geogaphic vaiation in the Whitecowned Spaow Zonotichia leucophys. Univ. Califonia Pul. Zool. 70:. BAPTISTA, L. F. 97. Wild House Finch sines Whitecowned Spaow song. Z. Tiepsychol. 0:70. BAPTISTA, L. F. 97. De Einfluss de Geslnge de Zugvogelasse pugetensis auf die Gesangsentwicklung de Standvogelasse des Weisskopfammefinken ( Zonotichia leucophys nuttalli). In Dtsch. Onithol. Ges. Jahesvesammlung (97), Saaiicken. J, Omithol. 4:7980.

15 70 LUIS F. BAPTISTA I I BAPTISTA, L. F The effects of songs of winteing Whitecowned Spaows on song development in sedentay populations of the species. Z. Tiepsychol. 4: 477. BAPTISTA, L. F. 97. Song dialects and demes in sedentay populations of the Whitecowned Spaow (Zonotichia leucophys nuttalli ). Univ. Califonia Pul. Zool. 0: l. BAPTISTA, L. F., AND H. WELLS. 97. Additional evidence of songmisimpinting in the Whitecowned Spaow. BidBanding 4:97. BLANCHARD, B. D. 94. The Whitecowned Spaows (Zonotichia Zeucophys) of the Pacific seaoad: envionment and annual cycle. Univ. Califonia Pul. Zool. 4:78. BLANCHARD, B. D. 94. Migation in Pacific Coast Whitecowned Spaows. Auk 9:47. BORROR, D. J. 90. The analysis of animal sounds, p. 7. In W. E. Lanyon, and W. N. Tavolga [eds.], Animal sounds and communication. Am. Inst. Biol. Sci., Washington, D.C. BROOKS, R. J., AND J. B. FALLS. 97. Individual ecognition y song in Whitethoated Spaows. III. Song featues used in individual ecognition. Can. T. Zool. :7497. CONRADS, K., AND W. CONRADS. 97. Regionaldialekte des Otolans (Eme&a hotulana) in Deutschland. Vogelwelt 9:800. CORTOPASSI, A. J., AND L. R. MEWALDT. 9. The cicumannual distiution of Whitecowned Spaows. BidBanding : 49. DAVIS, J. 98. Singing ehavio and the gonad cycle of the Rufoussided Towhee. Condo 0: 08. DEWOLFE, B. B. 98. Puget Sound Whitecowned Spaow. In A. C. Bent, and Collaoatos. Life histoies of Noth Ameican cadinals. goseaks, untings, towhees, finches, spaows; and allies. Pt.. U.S. Natl. Mus. Bull. 7. DEWOLFE, B. B., AND R. H. DEWOLFE. 9. Mountain Whitecowned Spaows in Califonia. Condo 4:8789. DEWOLFE, B. B., D. D. KASKA, AND L. J. PEYTON Pominent vaiations in the songs of Gamel s Whitecowned Suaows. _ BidBanding 4: 4. EBENHARDT, C., AND L. F. BAPTISTA Intaspecific and intespecific song mimesis in Califonia Song Spaows. BidBanding, in pess. EMLEN, S. T. 97. An expeimental analysis of the paametes of id song eliciting species ecognition. Behaviou 4:&7. _ FALLS.. B. 9. Poneties of id song eliciting esponses fom teitoial males. Poc. th Int. Onithol. Cong., p. 97. HARRIS, M. A., AND R. E. LEMON. 97. Songs of Song Spaows ( Melospiza melodia) ; individual vaiation and dialects. Can. J. Zool. 0:009. KING, J. R. 97. Vaiation in the song of the Rufouscollaed Spaow, Zonotichiu capensis, in nothwesten Agentina. Z. Tiepsychol. 0: KONISHI, M. 9. The ole of auditoy feedack in the contol of vocalization in the Whitecowned Spaow. Z. Tiepsychol. : KROODSMA, D. E Song leaning, dialects and dispesal in the Bewick s Wen. Z. Tiepsychol. : 80. LEMON, R. E. 9. The song epetoies of cadinals ( Richmondena cadinalis) at London. Ontaio. Can. J. Zool. 4:99: LEMON, R. E. 97. How ids develop song dialects. Condo 77:840. LEWIS, R. A. 97. Repoductive iology of the Whitecowned Spaow (Zonotichia leucophys pugetensis Ginnell ). I. Tempoal oganization of epoductive and associated cycles. Condo 77:49. MARLER, P. 90. Bid songs and mate selection, p In W. E. Lanyon, and W. N. Ta Volga [eds.], Animal sounds and communication. Am. Inst. Biol. Sci., Washington, D.C. MARLER, P A compaative appoach to vocal leaning: song development in Whitecowned Spaows. J. Comp. Physiol. Psychol. 7: l. MARLER, P., AND M. TAMURA. 9. Song dialects in thee populations of Whitecowned Spaows. Condo 4:877. MARLER, P., AND M. TAMURA. 94. Cultually tansmitted pattens of vocal ehavio in spaows. Science 4: MILLIGAN, M. M., AND J, VERNER. 97. Intepopulational song dialect discimination in the Whitecowned Spaow. Condo 7:08. MORTON, M. L., J. L. HORSTMANN, AND J. M. OSBORN. 97. Repoductive cycle and nesting success of the Mountain Whitecowned Spaow (ZOnotichia leucophys oiantha) in the cental Siea Nevada. Condo 74:. MUNDINGER, P. 97. Song dialects and colonization in the House Finch, Capodacus mexicanus, on the east coast. Condo 77 :4074. NOTTEBOHM, F. 99. The song of the chingolo, Zonotichia capensis, in Agentina: desciption and evaluation of a system of dialects. Condo 7:99. NOTTEBOHM, F. 97. Continental pattens of song vaiaility in Zonotichia cape&s: some po ssile ecological coelates. Am. Nat. 09:04. NOTTEBOHM, F., AND R. K. SELANDER. 97. Vocal dialects and gene fequencies in the chingolo spaow (Zonotichia capensis). Condo 74: I7 4. OREJUELA, J. E., AND M. L. MORTON. 97. Song dialects in seveal populations of Mountain Whitecowned Spaows (Zonotichia leucophys wiztha) in the Siea Nevada. Condo 77:4 PAYNE, R. B. 97. Behavio, mimetic song and song dialects, and elationships of the paasitic indigo ids (Vidua) of Afica. Onithol. Monog.. THIELCKE, G. 9. Stammesgeschichte und geogaphische Vaiation des Gesanges unsee Baumlaiife (Cethia familiais L. und C. achydactyla Behm). Z. Tiepsychol. 8: WIEDEMANN, A. M., L. J. DENNIS, AND F. H. SMITH. 99. Plants of the Oegon coastal dunes. Oegon State Univ. Bookstoe. Mooe Laoatoy of Zoology, Occidental College, Los Angeles, CA Accepted fo pulication Noveme 97.