Survival and Mobility in a Population of Pacific Coast Song Sparrows (Melospiza Melodia Gouldii)
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SURVIVAL AND MOBILITY IN A POPULATION OF PACIFIC COAST SONG SPARROWS (MELOSPIZA MELODIA GOULDII) RICHARD HALLIBURTON AND L. RICHARD MEWALDT The Song Sparrow (Melospiza melodia) is and Wildlife Service band. At each capture, age, one of the most thoroughly studied passerine sex, location, weight, molt, and reproductive con- dition were recorded in the field. We used plumage, birds in America, and its life history and degree of skull ossification (Norris 1961), and pres- breeding cycle are fairly well known. It is ence of a cloaca1 protuberance (adult males) or also the most polytypic bird in North America, brood patch (adult females) to classify a bird as a with 31 subspecies currently recognized juvenile (in its first calendar year) or adult, if aging was possible. (American Ornithologists ’ Union 1957). Cer- tain nonmigratory populations of Song Spar- RESULTS rows tend to be relatively sedentary, individ- uals rarely moving far from their hatching sites SURVIVAL (Nice 1937, Johnston 1956a). In this report we Survival, as used in this report, means sur- present evidence that the resident population vival from one breeding season to the next of Melospiza melodia gouldii on the Point and is estimated from the proportion recap- Reyes peninsula of California also is sedentary. tured the following year. Females with brood We estimate survival rates, dispersal distance patches were first captured in March, so any of juvenile birds, and home range size. We bird captured after 1 March was considered conclude that low vagility is widespread to have survived to the beginning of the among Song Sparrow populations and discuss breeding season of that year. its consequences. Aduh. Data for all adults of known sex were analyzed together and overall survival MATERIALS AND METHODS rates estimated. Estimates of survival rates ranged from 0.39 to 0.58 over the four years, From May 1966 through December 1970 individuals with a mean of 0.45 (table 1). Survival rates associated with the Avian Biology Laboratory of for males and females were not significantly San Jose State University banded 2522 Song Spar- rows in a capture-recapture study of the granivorous different (P > .05; test for equality of two birds at the southern end of Point Reyes National percentages, Sokal and Rohlf 1969) except in Seashore in Marin County, California. 1966 (P < .Ol) when the difference probably The study area, adjacent to the Pacific Ocean, was due to small sample size. varies in elevation from 60 m at the top of the Survival was independent of age after the cliffs directly above the beach, to 300 m. The study area is approximately 200 hectares, of which birds had lived one year. Over the four years about 130 are suitable habitat for Song Sparrows. 49% of 173 one-year-olds, 43% of 67 two-year- Most of the area is covered by Northern Coastal olds, and 50% of 12 three-year-olds survived Scrub (Munz and Keck 1959). The most common until the next breeding season. These values shrubs are Baccharis pilularis, Artemesia californica, and Rhus dioersiloba. Parts of old marine terraces are not significantly different (P > .05). at 60 m and 180 m elevation were cultivated, and Juveniles. Estimation of juvenile survival is the rest of the area was grazed by livestock for complicated by dispersal. Some juveniles are many years prior to 1966. Since then, these areas not seen the following year even though they have begun to return to their natural state. have survived. Thus, recapture rates may Mean monthly air temperatures range from 4 to 15°C. Daily summer temperatures typically range significantly underestimate the actual survival from 8 to 18°C and winter temperatures from 3 rates if many juveniles disperse from the to 10°C. In winter, air temperature frequently drops study area. to -1°C on slopes facing away from the ocean. Al- most all rain falls from October to April with an- Juveniles recaptured after 1 March were nual totals of 60 to 125 cm during the years of this considered to have survived to the breeding study. Fog is common, especially during the sum- season, though not necessarily to have bred. mer months. Recapture rates (minimum survival) ranged Rirds were mist-netted or captured in grain- baited Glenhaven-Standby Potter traps placed in an from 0.16 to 0.32 (table 2). The higher rate approximate 100-m grid in all suitable habitat. for 1966 is significantly different from the Altogether, 192 capture sites were used during the other years (P < .OOl) and may be due to study, but not all were operated regularly. Every bird was banded with a United States Fish errors in age determination in the late sum- 14991 The Condor 78:499-504, 197G 500 RICHARD HALLIBURTON AND L. RICHARD MEWALDT TABLE 1. Annual survival rates of adult Song TABLE 2. Known minimum survival rates of juve- Sparrows at Point Reyes, 1966b-1969. nile Sung Sparrows at Point Reyes, 1966-1969. Proportion Recaptured Year NU Sb N s N s Year k&k!: Following Year 1966 27 0.74 18 0.33 45 0.58 1966 137 0.32 1967 89 0.39 52 0.42 141 0.40 1967 373 0.21 1968 99 0.55 77 0.48 176 0.52 1968 279 0.18 0.16 1969 112 0.38 65 0.41 177 0.39 1969 509 Total 1298 0.20 Total 327 0.46 212 0.44 539 0.45 a N = Number of banded adults in population. h s = Proportion recaptured the following year. DISPERSAL We define dispersal distance as the distance mer of 1966. Some birds called “juveniles” between a birds hatching site and its point may have been adults. After 1966, age de- of settlement the following breeding season. termination probably was more accurate be- We estimated the hatching site by the first cause field personnel were more experienced, capture site for all juveniles banded before and because degree of skull ossification was 20 June of any year. Restricting the sample estimated. This is reflected by the more con- to birds banded before 20 June limited the sistent survival rates for 1967-69, during sample size to only 36 birds, but birds banded which the minimum for juvenile birds was after 20 June were likely to have moved from 0.18. their hatching site already. Recapture records The number of birds dispersing from the showed that movement nearly always had study area can be estimated by the method ceased by the following March; therefore, of Speirs (1963). He gave the following equa- we used the most frequent (usually the only) tion for estimating the total population, given recapture site after 1 March to estimate the the adult mortality rate: point of settlement. P, = Pi/( l-S,) = P,/M, The distance between banding site and recapture site was measured to the nearest where P, = total adult population, P, = 25 m for each juvenile banded before 20 population of one-year-olds, S, = annual sur- June. The results are compared (fig. 1) vival rate of adults, and M, = annual mor- with similar data for a population of Salt tality rate of adults. Solving for P,, Marsh Song Sparrows (M. m. samuelis) in P, = P&M,. the San Francisco Bay region (Johnston 1956a: fig. 9) and for an Interpoint, Ohio This means that in a stable population, population of M. m. euphonia (Nice 1937). enough juveniles survive to replace the adults To facilitate comparison, the Point Reyes data that die. If we consider P, to be the total were grouped into 100-m increments from population of banded adults and determine the banding site. The distributions were simi- M, from survival estimates (table 1)) we can lar for all three populations; the medians for calculate P,. Calculated, or expected, juvenile the Point Reyes, Salt Marsh, and Ohio popu- survival rates can be compared with the known minimum survival rates (table 3). The expected survival rates range from TABLE 3. Expected survival rates and known min- 0.14 to 0.30. Again, 1966 is inconsistent, prob- imum survival rates of juvenile Song Sparrows at Point Reyes, 1966-1969. ably for the reasons already mentioned. Over the four years the expected juvenile survival YfXI P,” p, % PI, p,r s, s, rate was 0.23. The known minimum survival 1966 137 45 0.42 19 44 0.14 0.32 rate was 0.20. The difference estimates the 1967 373 141 0.60 84 78 0.23 0.21 number of birds that have dispersed from 1968 279 176 0.48 85 51 0.30 0.18 the study area. In other words, 256 of 1298 1969 509 177 0.61 108 83 0.21 0.16 juveniles (20% ) were known to have sur- Total 1298 539 0.55 296 256 0.23 0.20 vived, but 296 (23%) were expected to have RP, = Number of juvenile birds banded; P, = total popu- survived (table 3). The 40 birds unaccounted l,:tion of banded adults; M, = annual mortality rate of adults (from table 1); P,, = expected number of one-year-olds the for are hypothesized to have dispersed be- following year (z P,M,); P,, = actual number of one-year- yond the study area. These represent 3% of olds recaptured the following year; se = exTected survival rate of juveniles (= Pie/PO); s,,, = known minimum survival all juveniles, or 14% of those surviving. rate (P,JP,). SURVIVAL AND MOBILITY IN SONG SPARROWS 501 .-----* POINT REYES (36) x_---________xSAN FRANCISCO BAY (34) +-----o OHIO (34) 200 400 600 800 1000 1200 1400 DISTANCE (171) FIGURE 1. Distance from hatching or banding site of juveniles to recapture site the following spring for three populations of Song Sparrows.