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- BULLETIN of the FLORIDA STATE MUSEUM Biological Sciences Volume 22 1977 Number 2 GEOGRAPHIC VARIATION IN THE SONG OF BELDING'S SAVANNAH SPARROW (PASSERCULUS SANDWICHENSIS BELDING.4 RICHARD ALAN BRADLEY e -. , rI. - . .* r S ., 4* f S 2 / a . 1. 1 UNIVERSITY OF FLORIDA GAINESVILLE Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCIENCES, are published at irregular intervals. Volumes contain about 300 pages and are not necessarily completed in any one calendar year. OLIVER L. AUSTIN, JR., Editor RHODA J. RYBAK, Managing Editor Consultants for this issue: Luis F. BAPTISTA JOHN WILLIAM HARDY j Communications concerning purchase or exchange of the publications and all manuscripts should be addressed to: Managing Editor, Bulletin; Florida State Museum; University of Flor- ida; Gainesville, Florida 32611. This public document was promulgated at an annual cost of $1,667.26 or $1.667 per copy. It makes available to libraries, scholars, and all interested persons the results of researches in the natural sciences, emphasizing the circum-Caribbean region. Publication date: June 17, 1977 Price: $1.70 GEOGRAPHIC VARIATION IN THE SONG OF BELDING'S SAVANNAH SPARROW (PASSERCULUS SANDWICHENSIS BELDINGI) RICHARD ALAN BRADLEY' SYNopsis: The song of Belding's Savannah Sparrow, resident from Santa Barbara, California, south to El Rosario, Baja California del Norte, was studied during the spring of 1973. Songs were recorded from individuals at 14 of the 15 remaining breeding localities of this subspecies and analyzed with an audiospectrograph. Recordings of color-banded birds indicated that each male sang a single highly stereotyped song pattern. Detailed analysis of intrapopulational and inter- populational variation was made. The study of interpopulational variati6n yielded a mosaic pattern of song "dialects." The occurrence of such song dialects has been substantiated else- where for a variety of species. The present study involves very small adjacent, but isolated pop- ulations. TABLE OF CONTENTS INTRODUCTION . 58 ACKNOWLEDGEMENTS. 59 METHODS 59 59 Recordings 61 Analysis..... Terminology . 61 REsuLTs ... 61 Call Repertoire ....._ 61 @~SERR~ Features of Song Delivery The Structure of the Primary Song.. Ontogeny of the Song . Individual Variation Intrapopulational Variation Interpopulational Variation Summary of Interpopulational Variation by Location . & DISCUSSION. 65 LITERATURE CITED.. The author is an Associate in Natural Sciences at the Florida State Museum, University of Florida, Gainesville 32611. This paper was submitted in partial fulfillment for the degree of Master of Arts at California State University, Long Beach 90804 Manuscript accepted 18 December 1975. BRADLEY, RtcHARD ALAN. 1977. Geographic variation in the song of Belding's Savannah Sparrow (Passerculus sandwichensis beldingi). Bull. Florida State Mus., Biol. Sci. 22(2):57-100. 58 BULLETIN FLORIDA STATE MUSEUM Vol. 22, No. 2 INTRODUCTION Geographic variation in bird song has been the subject of recent studies and has taken several forms. The early emphasis was on documentation of variation within species having wide distributions. More recently compre- hensive studies have been conducted using recordings from many localities or analyzing variation between local contiguous or isolated populations. Per- haps the best summary of the work on geographic variation in bird song is by Thieleke (1969). Early work on the Chaffinch (Fringilla coelebs) and other species revealed a distinctive mosaic pattern of variation (Thieleke 1969). Song patterns used by most of the birds in one place are similar to their neighbors' songs, while individuals from other populations have different themes. Often an entire system of such variants is found when several different populations are studied. The term "song dialect" refers to this type of variation. Many authors discussed the evolutionary significance of song dialects. Mayr (1963) stated that geographic variation of song is a widespread isolating mechanism in birds. Other workers expressed similar ideas about song as an isolating mech- anism (Thorpe 1961, Marler and Tamura 1964, Lanyon 1969, Nottebohm 1969, Thieleke 1969, Armstrong 1973). Nottebohm (1972) summarized the proposed mechanisms of such isola- tion: "It is necessary to assume that female birds develop a pref- erence for the song dialect of the area where they are born, and that this preference is revealed in their choice of partner. Males in turn must learn to sing the dialect of their birth area. If birds of both sexes migrate or wander between fledgling age and their first breeding season, they must return to breed in the same gen- eral area where they were raised. In this fashion an assortative mating system based on song preferences reinforces a more or less loose philopatry." The Savannah Sparrow (Passerculus sandwichensis) is one of the most widely distributed of North American sparrows. The species breeds from 71°N (Barrow, Alaska) south to 16°N (Hacienda Chancol, Guatemala) and is divided into 18 subspecies that vary considerably in size, proportions, and coloration (American Ornithologists' Union Check-list 1957, supplement 1973). Of the 18 subspecies 8 are restricted to the coastal marshes of California and northwestern Mexico. Van Rossem (1947) presented an excellent review of the seven races that breed in western Mexico. The Savannah Sparrow form that breeds along the coast from Santa Bar- bara south to El Rosario was originally named as a full species, Passerculus beldingi, by Ridgway (1885). This race, now known as Passerculus sand- 1977 BRADLEY: BELDING'S SAVANNAH SPARROW 59 wichensis beldingi (American Ornithologists' Union Check-list 1957) is re- ferred to here as Belding's Sparrow. The name Savannah Sparrow is used when referring to the species in general. Belding's Sparrow is restricted to the tidal salt marshes of southern Cali- fornia and northwestern Baja California. In a preliminary census conducted in conjunction with this study, I estimated a population of fewer than 3,000 singing males (Bradley 1973). The restriction of this subspecies to local patches of salt marsh has produced small isolated populations ideal for the study of geographic variation. The pattern of song variation within the spe- cies, as outlined by Borror (196la) is well suited to the development of song dialects: (1) each bird sings only one song pattern, (2) variation occurs between different individuals, and (3) song patterns used in any particular location show considerable overlap. Because of restrictions of distance and time, recordings could be made of only a limited number of populations. As a result, a somewhat arbitrary de- cision was made to restrict the study to the coastal populations now assigned to P. s. beldingi. Recordings from another coastal subspecies (P. s. alaudinus) in central and northern California indicate a continuum of similar patterns of variation in that subspecies, and perhaps throughout the species. This study was conducted primarily in the spring of 1973, although some additional ob- servations and recordings were made later in 1973 and early in 1974. Tape recordings were made of 280 different individuals at 14 of the 15 known re- maining breeding sites of Belding's Sparrow. No recordings were made at Ensenada Bay, Baja California del Norte, because of time limitations and the small, widely-scattered nature of that population. ACKNOWLEDGEMENTS I thank David R. Bontrager, Marjorie J. Bradley, John William Hardy, and Graeyer Mans- field-Jones for reading all or parts of the manuscript and making many helpful suggestions. I give special thanks to Charles T. Collins and Stuart L. Warter for guiding my research as well as suggesting corrections and improvements of this paper. In addition to the above named per- sons, many others helped me in the field, or obtained permission for my studies in restricted areas. For such help I express my appreciation to Commander Robert Baker, Frank DeVore, L. E. Fellows, Barbara Massey, Larry Pomeroy, Jene St. Germain, Deane Swickard, Margaret Tay- lor, and Bruce Wales. I also thank David Bradley for preparation of Figure 16, assistance in the field, and innumerable hours spent conducting computer analysis of the data. METHODS RECORD]NGS Most of the recordings in this study were made with a Uher 4000 Report S tape recorder and a Uher M 512 microphone mounted on either a 61 cm or a 76 cm fiberglass parabolic reflector. Some additional recordings were made with a Sony TC 800 recorder and a Sony ECM 19 B mi- crophone mounted on the 61 cm reflector. All recordings were made at a tape speed of 19 cm/s. Recordings were made in the field with a hand-held reflector during the peak period of song activity in the morning. Copies of the recordings are on file in the Bioacoustic Archive of the Florida State Museum, Gainesville. 60 BULLETIN FLORIDA STATE MUSEUM Vol. 22, No. 2 Belding's Sparrow is a bird of the coastal salt marshes of southern California and is largely , restricted to Salicorniq-covered tidal flats..Each singing individual, presumed to be a male, de- fends an exclusive territory -in this habitat and regularly sings from one or more exposed perches within his territory. When I surveyed a particular location, I made recordings beginning with an individual at one edge of the marsh and continued in a regular pattern from territory to territory until the entire nesting area had been traversed. Usually I recorded 5 t6 10 songs from each individual. As males of this subspecies defend very small territories (Bradley 1973),each singing bird could be iden- tified by its song perch location. Great care was taken to record songs from distinct individuals. In those cases where a large number of birds were singing from a relatively small area, only a few indivjduals were recorded if any probability of confusion existed. With this method only small samples could be obtained from some sites, but these samples are free from duplication. At some locations the samples were recorded on two or more dates. In each case additional recordings for the sample were made in parts of the marsh where birds had not been previously recofded.