2009 SOUTHEASTERN NATURALIST 8(3):537–546 Stopover-site Fidelity at a Near-coastal Banding Site in Georgia Scott G. Somershoe1,*, Don G. Cohrs2, and Doris A. Cohrs2 Abstract - We documented one Seiurus noveboracensis (Northern Waterthrush) exhibiting stopover-site fidelity by returning to a near-coastal stopover site at the Butler Island Auxiliary Station (BIAS) on the Altamaha Waterfowl Management Area in southeast Georgia during banding operations from 1995–2000. The Northern Waterthrush was recaptured in fall 2000 after being banded at BIAS in fall 1997. Although few individual passerine migrants have exhibited stopover-site fidelity, the majority were recaptured at banding stations nearer (<500 km) breeding populations of the respective species. Stopover-site fidelity is rarely documented in the southeast, as migration banding stations are generally located at coastal migrant traps, which primarily capture hatching-year birds that likely migrate to inland locations in subsequent fall migrations and do not return to coastal migrant traps. High percentages of hatchingyear migrants at our near-coastal site suggest that this area is used by migrants in a similar fashion to coastal sites; however, we found possible benefits received by young birds returning to this site in subsequent years. Although rarely documented, passerine migrants may encounter some of the same benefits that wading birds, waterfowl, and shorebirds encounter in using the same stopover locations annually. Introduction Migrant birds of many different groups have been well documented exhibiting fidelity to breeding and non-breeding grounds (Greenwood and Harvey 1982, Holmes and Sherry 1992). Such fidelity provides advantages of increased familiarity with local resources, territorial dominance, and predator avoidance (Brown et al. 2002, Latta and Faaborg 2001). The selection pressures that provide site fidelity to breeding and non-breeding areas may also be expected to be found in en route migrants (Catry et al. 2004). Shorebirds, wading birds, and waterfowl have well-documented fidelity to stopover sites since they tend to be tied to specific habitats that are not generally available across the landscape, i.e., wetlands, streams, mud fl ats, lakes, etc. (Evans and Townshend 1988, Harrington et al. 1988, Robertson and Cooke 1999, Smith and Houghton 1984). We may expect stopover-site fidelity to be less common in passerines, especially long-distance passerine migrants, in relation to waterfowl, shorebirds, and wading birds. Indeed, fidelity to stopover sites along migration routes has rarely been reported in long-distance migrant passerines in North America (Somershoe and Chandler 2000, Winker et al. 1991) and central Iberia (Bermejo and de la Puente 2002, Cantos and Tellería 1994); however Catry et al. (2004) documented up to 12.9% of passage migrants returning to the same sites in Portugal. 1Tennessee Wildlife Resources Agency, PO Box 40747, Nashville, TN 37204. 2PO Box 1908, Darien, GA 31305. *Corresponding author - scott.somershoe@tn.gov. 538 Southeastern Naturalist Vol. 8, No. 3 We find few stopover-site faithful passerines for a couple of reasons. First, suitable stopover sites are generally in greater supply than for wetland- specific species. Passerines are more fl exible in their habitat selection during stopover and can gain weight, or at minimum, maintain weight even in dry forests (Schaub and Jenni 2001). Second, the typical fl ight speed of migrating passerines is lower than for larger-bodied geese and shorebirds; therefore, passerines are more sensitive to wind conditions. A slower fl ying speed would result in passerines experiencing greater wind drift, likely needing many more stopover locations, and migration may be much more dependent on current weather conditions that infl uence whether a bird stops at a previous stopover location or continues with favorable wind conditions (Catry et al. 2004). In North America, stopover-site fidelity has been rarely documented. Nisbet (1969) published the first compilation of records of stopover-sitefaithful transients, which he describes as individuals that return to the site of banding in subsequent years and whose regular breeding and non-breeding areas are at least 161 km from the banding site. Winker et al. (1991) summarized 8 published articles and found a total of approximately 21 individuals of 10 species that exhibited stopover-site fidelity. All of these documented cases were at inland sites (≥40 km from the nearest large water barrier) and were generally located closer to the nearest breeding populations for the species. Somershoe and Chandler (2000) documented the first site-faithful migrants (four Seiurus noveboracensis Gmelin [Northern Waterthrush] and one Dendroica petechia L. [Yellow Warbler]) at a near-coastal stopover site (≤40 km from the nearest large water barrier) in southeast Georgia. The first objective of this paper is to report an additional stopover-sitefaithful migrant at a near-coastal site in Georgia. Second, we assess possible benefits to migrants utilizing the same stopover site annually. Third, we review the literature of stopover-site-faithful passerine migrants in North America and assess species-specific likelihoods of stopover-site fidelity of transients. Finally, we address factors infl uencing the likelihood of migrants to return to stopover sites with respect to distance between the banding station to the nearest breeding population of the respective species, distance between the banding station and the nearest large water barrier, and proportion of hatching-year individuals captured at each site. Methods We banded birds at the Butler Island Auxiliary Station (BIAS) within the Altamaha Waterfowl Management Area, a near-coastal site located approximately 20 km from the Atlantic coastline in southeastern Georgia (McIntosh County; 31°21'N, 81°26'W). Mist nets were located along a series of levees used to control water levels on the management area. The levees and the pond edges adjacent to the net lanes were dominated by Celtis laevigata Willd. (Sugarberry), Prunus serotina Ehrh. (Black Cherry), Diospyros virginiana L. (Persimmon), Morella cerifera (L.) Small (Wax Myrtle), and 2009 S. Somershoe, D.G. Cohrs, and D.A. Cohrs 539 Rubus spp. (Blackberry). Altamaha Waterfowl Management Area has a variety of habitats along the fl oodplain of the Altamaha River, including old diked rice fields, Pinus spp. savannas, tidal creeks, hardwood fl oodplain forests, and old-growth stands of Taxodium distichum L. (Bald Cypress). Although BIAS is not located on the immediate coast nor truly inland, we consider BIAS a near-coastal banding station because of its close proximity (≤40 km) to the Atlantic coast. There are extensive fl oodplain marshes to the east between BIAS, the mouth of the Altamaha River, and Sapelo Island. We also report data from the Jekyll Island Banding Station (JIBS), a nearby coastal banding site, on Jekyll Island, Glynn County, GA (31°00'N, 81°25'W). JIBS is 40 km south of the BIAS site on the immediate Atlantic coast, and nets were placed within early successional habitat at the southern tip of Jekyll Island. The site is dominated by Quercus virginiana P. Mill (Live Oak), Wax Myrtle, Serenoa repens Bartr. (Saw Palmetto), and Persea borbonia L. (Red Bay) (Cohrs and Cohrs 1994). We operated 12–15 mist nets (12 x 2.6 m, 32-mm mesh) at BIAS daily, weather permitting, from 29 August through 5 November 1995–2000. We opened nets at dawn and operated them 6 hours/day (mean of 57 days and 2166 net hrs/year). We operated 9–15 nets at JIBS between 21 September and 25 October 1986–2001 in a similar fashion to our procedures at BIAS, with an average of 22 days and 1374 net hrs/year. Each bird captured was banded with a USGS aluminum band and released at a central banding station under permit to D.G. Cohrs and D.A. Cohrs. For comparison of age ratios at other banding stations in the southeast United States, we assessed banding from two inland banding stations at Wekiva Springs State Park and Rock Springs Run State Reserve (WSSP/RSRSR) near Orlando, Orange County, FL (C.R.D. Brown and R.T. Poole, Orlando, FL, unpubl. data; see Poole et al. [2003] and Somershoe et al. [2009] for site descriptions). Birds were banded in fall migration, i.e., 1 September through 31 October 1997–2001. We combined data because sites were within 5 km and consisted of similar habitats. We assessed age ratios at a coastal site of birds banded at Ft Morgan, AL from 1 September through 30 October 1991–2007, excluding 1998 and 2003 (F.R. Moore, University of Southern Mississippi, Hattiesburg, MS, unpubl. data). We generated chi-square values for contingency table analysis with 95% confidence limits to test the hypothesis that age ratios were not different between sites (Zar 1996). Results One Northern Waterthrush was recaptured on 22 September 2000 after initially being banded as a hatching-year bird at BIAS on 28 September 1997. We did not recapture the individual more than once during either year of encounter. The individual weighed 18.0 g upon original banding and 18.8 g upon recapture in 2000 and had no fat reserves during either capture. With 4 site-faithful Northern Waterthrush from Somershoe and Chandler (2000) and 1 in this paper, 5 of 733 Northern Waterthrush banded at BIAS 540 Southeastern Naturalist Vol. 8, No. 3 from 1995–1999, with returns possible in 1996–2000, exhibited stopoversite fidelity at BIAS (0.7%). We are unable to assess stopover-site fidelity in other species as we did not capture many individuals of most transients species at BIAS during that time span, i.e., only 30 Catharus fuscescens Stephens (Veery), 20 Dendroica caerulescens Gmelin (Black-throated Blue Warbler), and 83 Setophaga ruticilla L. (American Redstart); however, 1 of 166 (0.6%) Yellow Warbler returned to our study site (Somershoe and Chandler 2000). Of the 5 Northern Waterthrush exhibiting stopover-site fidelity at BIAS, 4 of 5 were hatching-year at the time of initial capture. All individuals subsequently returned to BIAS on an ordinal date earlier than that of initial capture, while all but 1 recapture occurred within 10 days of the initial banding date (Table 1). All individuals had the same or greater fat score (range 0–3) upon recapture. Only 1 individual had a lower mass upon recapture (0.4 g lower), while the remaining 4 birds averaged 1.5 g higher mass than at initial capture (range = 0.3–3.2 g). On the other hand, the Yellow Warbler that returned to BIAS was recaptured on an ordinal date 11 days after initial banding and with a reduced fat score and mass (1.5 g lower). We found no stopover-site-faithful individuals of species fitting Nisbet’s (1969) criteria from >4500 individuals of 13 species banded at JIBS from 1986–2000, allowing for recaptures in 2001. We also did not document any stopover-site-faithful individuals from >30,500 individuals of 7 species that do not strictly fit the criteria, but are common during fall migration and may return annually, i.e., Dumetella carolinensis L. (Gray Catbird), Seiurus aurocapillis L. (Ovenbird), Dendroica p. palmarum Gmelin (Western Palm Warbler), Dendroica discolor Vieillot (Prairie Warbler), Geothlypis trichas L. (Common Yellowthroat), Vireo olivaceous L. (Red-eyed Vireo), and Passerina cyanea L. (Indigo Bunting). We did not find stopover-site-faithful birds in either spring or fall at other coastal stopover sites in the southeast United States where large numbers of migrants, including those meeting and not meeting Nisbet’s (1969) criteria are banded annually, i.e., Peveto Woods, LA (>10,000 individuals, 27 species, spring 1982–1991), Johnson Bayou, LA (>19,000 individuals, 25 species, spring 1993–2006), and Ft. Morgan, AL (>31,100 individuals, 31 species, fall 1991–2007) (F.R. Moore, University of Southern Mississippi, Hattiesburg, MS, unpubl. data). In addition, no stopover-site faithful birds were found from over 1500 individuals of 13 transient species fitting Nisbet’s criteria that were banded from 1997–2000 at the inland site WSSP/ RSRSR. We only included species that strictly fit Nisbet’s criteria, as individuals of several migratory species returned to winter grounds annually, e.g., Gray Catbird and Ovenbird (Somershoe et al. 2009). We reviewed and critiqued records of stopover-site-faithful migrants from the literature and consider 25 individuals of 9 species strictly fitting Nisbet’s criteria with the inclusion of the 1 new record in this paper (Table 1). We excluded 3 Dendroica c. coronata L. (Myrtle Warbler) and 2 2009 S. Somershoe, D.G. Cohrs, and D.A. Cohrs 541 Zonotrichia albicollis Gmelin (White-throated Sparrow) possibly accepted by Winker et al. (1991) due to the fact that the banding station was within the winter range of the species (Foy 1975, Johnson and Lee 1974). Twenty-one of the 25 site-faithful migrants (84%) returned to a stopover site within 10 ordinal days of the initial banding date, accounting for leap years. Sixteen of the 25 migrants (64%) were recaptured at banding stations ≤500 km from the nearest respective breeding population, with 7 of these birds (28%) returning to sites ≤250 km of the nearest breeding population (Table 1). Only 7 individuals returned to sites ≥700 km from the nearest breeding population. All documented stopover-site-faithful individuals occurred in fall, with the exception of 1 Northern Waterthrush in spring (Table 1). The 6 individuals that returned to BIAS (Somershoe and Chandler 2000 and this paper) are the only recorded stopover-site-faithful migrants to return to a site ≤40 km from the nearest large water barrier. Table 1. Stopover-site-faithful migrants documented in the literature with capture site, encounter dates with the difference in number of ordinal days (Days) between encounters accounting for leap years, and approximate minimum distance from banding location to nearest breeding population (Dist., in km). Banding Return Species Banding location date date Days Dist. Catharus ustulatusA Shirland, IL 31 Aug 1968 10 Aug 1969 20 550 Catharus ustulatusA Shirland, IL 30 July 1972 26 Aug 1973 26 550 Vermivora peregrinaB Homewood, AL 21 Oct 1967 18 Oct 1968 2 2100 Vermivora peregrinaC Little Falls, MN 24 Aug 1968 29 Aug 1969 4 200 Vermivora peregrinaD Washington Co., MN 23 Aug 1984 19 Aug 1985 6 240 25 Aug 1986 Vermivora ruficapillaC Little Falls, MN 23 Aug 1966 27 Sept 1968 3 160 Vermivora ruficapillaC Little Falls, MN 8 Aug 1968 6 Sept 1969 28 160 Vermivora ruficapillaC Little Falls, MN 21 Aug 1968 12 Aug 1969 8 160 Dendroica petechiaC Tallahassee, FL 18 Sept 1967 7 Sept 1968 10 200 Dendroica petechiaE Butler Island, GA 16 Sept 1995 27 Sept 1996 10 200 Dendroica magnoliaF Nashville, TN 14 Sept 1975 27 Sept 1978 13 500 Dendroica tigrinaG Powdermill, PA 9 Sept 1972 20 Sept 1974 11 300 Dendroica c. coronotaB Powdermill, PA 16 Oct 1961 14 Oct 1962 2 200 Dendroica striataB Raynham, MA 18 Sept 1962 20 Sept 1963 2 200 Dendroica striataB Raynham, MA 25 Sept 1962 24 Sept 1963 1 200 Dendroica striataB Sudbury, MA 9 Sept 1962 24 Sept 1963 15 200 Dendroica striataB Littleton, MA 18 Sept 1967 17 Sept 1968 1 200 Seiurus noveboracensisB Tallahassee, FL 3 Sept 1967 10 Sept 1968 7 900 Seiurus noveboracensisE Shirland, IL 9 Sept 1967 7 Sept 1969 2 200 Seiurus noveboracensisH Hughes Hollow, MD 18 May 1970 15 May 1971 3 200 Seiurus noveboracensisE Butler Island, GA 9 Sept 1995 3 Sept 1997 6 700 Seiurus noveboracensisE Butler Island, GA 10 Sept 1996 6 Sept 1997 3 700 Seiurus noveboracensisE Butler Island, GA 14 Sept 1997 4 Sept 1998 10 700 Seiurus noveboracensisE Butler Island, GA 15 Sept 1996 30 Aug 1999 16 700 Seiurus noveboracensisI Butler Island, GA 28 Sept 1997 22 Sept 2000 6 700 AJohnson and Lee 1974, BNisbet 1969, CRyan 1970, DWinker et al. 1991, ESomershoe and Chandler 2000, FGoodpasture 1979, GLeberman and Clench 1975, HWoodward 1972, Ithis paper. 542 Southeastern Naturalist Vol. 8, No. 3 Age ratios of migrants at coastal, near-coastal, and inland stopover sites were significantly different (Table 2). We found that age ratios at the 2 coastal sites differed for all species. We found that the coastal stopover sites captured a higher percentage of hatching-year birds than the inland and nearcoastal site, with the exception of Yellow Warbler at Ft. Morgan and BIAS. Although nearly all sites had significantly different age ratios, BIAS typically had hatching-year percentages more similar to those from the coastal sites than the inland site in Florida, with the exception of Black-throated Blue Warbler, which had a relatively small sample size at BIAS (Table 2). Discussion The tendency of songbirds to be faithful to stopover sites along the migration route is rare due to a myriad of potential physical and environmental factors. Although this phenomenon is uncommon, we found that 48% of documented site-faithful individuals were either Northern Waterthrush or Dendroica striata Forster (Blackpoll Warbler), suggesting that they may tend to exhibit stopover-site fidelity more often than other species. Benefits of stopover-site fidelity to Northern Waterthrush at BIAS may be evident from the general pattern of higher mass and fat scores upon recapture as adults. Northern Waterthrush are territorial on migration, and there may be benefits of returning to a known stopover site for locating and defending areas of adequate food resources for fat deposition or replenishment (Rappole and Warner 1976). The benefit of stopover-site fidelity to Blackpoll Warbler in eastern Massachusetts may be familiarity of sites for fat deposition, as these areas are concentration areas near departure locations for the long offshore migration undertaken by this species; however, we do not have fat and mass scores for a preliminary assessment (Hunt and Eliason 1999). Birds with more fat reserves during migration may have a greater chance of selecting stopover habitat. In addition, the possible benefit of more rapid deposition of fat and mass at a revisited stopover site and a similar passage window for the majority of stopover-site-faithful migrants may indicate that individuals generally have a relatively small migration window and so efficient use of a familiar stopover site can be important. There may be some physiological benefit of stopover-site fidelity in passerines, but measuring success of returning individuals is difficult, if not impossible. The possible benefits of utilizing a stopover site annually may be important to passerine migrants; however, several factors greatly reduce the likelihood of a passerine returning to a stopover site. One factor is the geographic location and distance of the breeding range to the southern most stopover site. A latitudinal analysis of probability of stopover-site fidelity is difficult to do with each species having a different breeding range and location of breeding populations in the eastern United States and Canada. There appears to be a tendency for stopover-site-faithful migrants to return more often to sites ≤500 km of the nearest breeding population for the respective species. Only 7 individuals returned to sites ≥700 km distant. 2009 S. Somershoe, D.G. Cohrs, and D.A. Cohrs 543 Table 2. Age ratios for new birds captured at 4 fall migration banding locations in the southeast United States. Age ratios represent the number of hatching-year birds for all banded individuals. P-value represents the result of chi-square analysis and contingency table analysis with 95% confidence limits. Inland Near-coastal Coastal Species WSSR/RSRSR Butler Island Jekyll Island Fort Morgan P Catharus fuscescens 422/677 62.3% 27/34 79.4% 58/60 96.7% 158/189 83.6% <0.001 0.622 < P < 0.624 0.771 < P < 0.80 0.951 < P < 0.967 0.832 < P < 0.837 Catharus ustulatus 106/228 46.5% 88/93 94.6% 703/916 76.7% <0.001 0.463 < P < 0.467 0.936 < P < 0.947 0.767 < P < 0.77 Dumetella carolinensis 1728/2561 67.5% 2644/2696 98.1% 10295/11562 89.0% <0.001 0.674 < P < 0.675 0.980 < P < 0.981 0.890 < P < 0.891 Dendroica caerulescens 224/400 56.0% 14/28 50.0% 583/769 75.8% 70/74 94.6% <0.001 0.557 < P < 0.561 0.483 < P < 0.517 0.757 < P < 0.758 0.933 < P < 0.947 Dendroica magnolia 42/71 59.1% 94/108 87.0% 887/1229 72.2% <0.001 0.583 < P < 0.597 0.862 < P < 0.872 0.721 < P < 0.722 Dendroica palmarum 50/66 75.8% 116/121 95.8% 8258/8284 99.7% 1093/1137 96.1% <0.001 0.746 < P < 0.761 0.951 < P < 0.959 0.997 < P < 0.998 0.960 < P < 0.961 Dendroica petechia 139/166 83.7% 153/161 95.0% 383/460 83.3% <0.001 0.832 < P < 0.838 0.944 < P < 0.951 0.831 < P < 0.833 Seiurus aurocapillis 316/599 52.8% 144/153 94.1% 210/286 73.4% <0.001 0.527 < P < 0.528 0.935 < P < 0.942 0.732 < P < 0.735 Seiurus noveboracensis 56/93 60.2% 724/789 91.7% 462/473 97.6% 498/782 63.7% <0.001 0.596 < P < 0.606 0.917 < P < 0.918 0.975 < P < 0.977 0.636 < P < 0.637 Setophaga ruticilla 304/594 51.1% 90/102 88.2% 1328/1470 90.3% 1316/1640 80.2% <0.001 0.511 < P < 0.513 0.874 < P < 0.884 0.903 < P < 0.904 0.802 < P < 0.803 544 Southeastern Naturalist Vol. 8, No. 3 As migrants reach areas more distant from the breeding grounds, the cumulative effects of variable wind conditions are more likely to increase the amount of drift from a possibly predetermined migratory pathway and/ or affect the distance an individual can successfully migrate in a given period of time. The benefit of correcting for such drift errors, albeit slight or significant, may not outweigh the cost of expending energy returning to a familiar patch of forest, whereas the larger benefit of continuing migration when conditions are favorable likely outweigh the smaller benefit of shortstopping a flight for a known location. The distance migrants travel from breeding grounds in Canada and northern United States to southeastern banding stations is generally >500–700 km and may be too far to adjust for drift and utilize familiar locations, thus resulting in a scarcity of stopoversite- faithful records for species that strictly fit Nisbet’s (1969) criteria. The distance of banding stations to the immediate coast (i.e., nearest large water barrier) and the segment of the migratory population of each species using those sites likely affect the probability of stopover-site fidelity. Assessment of age ratios of passage migrants captured at various locations across the southeast support the hypothesis that adult birds appear to use more inland sites during migration, that coastal areas are dominated by inexperienced, and possibly disoriented, hatching-year birds, and that inland sites support more adult birds in fall migration (Moore 1984, Murray 1966, Ralph 1978). Ralph (1981) and Moore (1984) also suggested that adult birds have more experience in migration orientation; thus, we would not expect adult birds to show fidelity to a coastal site since these individuals are not returning to coastal areas in subsequent years. Adult birds may also migrate inland in order to attempt to avoid higher densities of birds along the coastal sites, possibly reduced availability of resources and/or territories on the coast, and reduced risk of fl ight error due to wind drift upon initiating a fl ight from the edge of a water barrier, especially the southeastern Atlantic coast. Many adult birds captured in fall at migrant traps on the coast of the Gulf of Mexico may be captured after returning to land as a correction for over-shooting the coast during a night fl ight and encountering poor migratory conditions (R. Sargent, Hummer Bird Study Group, Clay, AL, pers. comm.). The high percentage of hatching-year birds at BIAS suggest migrants may perceive the site as a coastal location rather than an inland site, although the proximity of the site prevents clear determination of category. Regardless, the 6 returning birds at BIAS represent the first records of sitefaithful transients returning to a site ≤40 km from a large water barrier. The site-faithful birds banded as hatching-year at BIAS in a site dominated by inexperienced, hatching-year birds, returned to utilize the near-coastal location and may have been able to take advantage of familiarity of the site and knowledge of resources in order to increase body fat and mass. The higher percentages of adult birds at inland sites and the lack of site-faithful migrants at banding stations on the immediate southeastern Atlantic coast suggests possible inter-annual variability in migration routes between age classes. We also suggest that stopover-site fidelity may be a small and generally 2009 S. Somershoe, D.G. Cohrs, and D.A. Cohrs 545 unimportant factor to long-distance migrant passerines in North America. Further research comparing stopover length, rates of fat and mass deposition, and age ratios may elucidate differences in migratory behavior of birds between coastal, near-coastal, and inland sites. Acknowledgments We dedicate this paper to the memory of Don G. Cohrs for the education and conservation programs he conducted in south Georgia over 35 years. We thank the staff of the Altamaha Waterfowl Management Area for permission to conduct migration monitoring. 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