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Abundance, Density, and Diversity of Neotropical Migrants at the Lula Lake Land Trust, GA
David A. Aborn

Southeastern Naturalist, Volume 6, Number 2 (2007): 293–304

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2007 SOUTHEASTERN NATURALIST 6(2):293–304 Abundance, Density, and Diversity of Neotropical Migrants at the Lula Lake Land Trust, GA David A. Aborn* Abstract - I used point counts and mist netting at the Lula Lake Land Trust, GA, a reclaimed mining area, to assess its suitability for migrating and breeding longdistance migratory landbirds. The results suggest that the property is suitable for migrants. The proportion of migrants is similar to other areas of the southeastern United States, as well as other reclaimed forestland. In addition, migrant populations were either stable or increased over the three years of monitoring, and 4% of migrants banded returned in subsequent years. Management of the Trust properties should continue to emphasize maintaining large areas of mature forest, and Trust managers should take measures to ensure that increased human recreational activity on the property does not degrade habitat quality. Introduction In recent years, much interest has been generated regarding population declines of many long-distance neotropical migratory landbirds (hereafter referred to as migrants). While research has shown that the reports of familywide population declines may have been exaggerated (Faaborg 2002), there have nonetheless been population-wide declines for some species (e.g., Dendroica cerulea Wilson [Cerulean Warbler]; Robbins et al. 1992), and regional declines in others (e.g., Coccyzus americanus Linnaeus [Yellowbilled Cuckoo]; Hughes 1999). The causes of the declines are varied, and include habitat fragmentation and loss of both breeding and wintering grounds. Thus, migratory birds still face human-induced threats to their survival, and identifying regions important for migrants and understanding all the factors influencing their life history are important for taking a proactive approach to managing migrant populations. Studies from many areas have demonstrated that migrants are more abundant in large forest tracts than in smaller ones (Ambuel and Temple 1983, Blake and Karr 1984, Gibbs and Faaborg 1990, Hayden et al. 1985, Martin 1988, Robbins et al. 1989). Thus, as result of their area-sensitive nature, neotropical migrants disappear more rapidly than other avian species when forests are cleared or fragmented. In addition to basic loss of area, many species suffer increased nest predation when forests are fragmented because many predators that are normally restricted to forest edges gain access to forest interiors (King et al. 1996, Lovejoy et al. 1986, Wilcove 1985). In addition to predation, recent evidence indicates that forest fragments contain lower food abundance than contiguous forests (Burke and Nol 1998) and also impede male pairing *Department of Biological and Environmental Sciences, University of Tennessee at Chattanooga, Chattanooga, TN 37406-2598; David-Aborn@utc.edu. 294 Southeastern Naturalist Vol. 6, No. 2 success (Bayne and Hobson 2001), both of which may further impact reproductive success for some species. It is clear that bird population trends for a given area are a complex mixture of factors that affect survival and recruitment (Sherry and Holmes 1992). Identifying and protecting large forest tracts have therefore become fundamental parts of many management plans for conserving forest-breeding migrants. Reclaimed mining lands can represent important habitat for many bird species (Allaire 1982, Karr 1968, Laki et al. 2004, Wray et al. 1982). For example, mined areas in Alabama had similar numbers and species of wintering birds as a nearby control area (Terrel and French 1975). In Illinois, mined areas actually had higher avian diversity when compared to similar, undisturbed grasslands (Karr 1968). While there has been much research on the avifauna of reclaimed grasslands, studies of reclaimed forests have received comparatively less attention. Given the importance of forests to many neotropical migrants, such data are valuable for the development of proper management plans. The southern Appalachian region has one the highest densities of breeding neotropical migratory birds in the United States. In some areas, over 80% of breeding-bird communities are composed of neotropical migrants. The Lula Lake Land Trust in Walker County, GA represents a large (􀂧 1800- ha) forest tract in the southern Appalachian region that at one time saw heavy mining activity and is therefore a good location for monitoring the abundance of neotropical migratory birds on reclaimed land. I surveyed the Lula Lake property in order to provide an inventory of the avian species using the area during spring migration and the breeding season. The data will provide valuable information that can be used for management of the property, as well as decisions regarding future land acquisition. Methods Study site The Lula Lake Land Trust (LLLT, 34°56'N, 085°22'W; Fig. 1), established in 1994, seeks to protect and preserve the resources within the Rock Creek watershed. Coal mining, clearcut timber harvests, garbage dumping, and unrestricted public access had left much of the land highly degraded. Once mining ended in the 1950s, and the area was closed to public access and logging in the early 1980s, restoration of the land began by cleaning up dump sites and replanting timbered areas. In addition to the 243-ha core area containing Lula Lake, the Trust has purchased a number of satellite properties, bringing the total holdings to 1780 ha. The condition of the property has been much improved with reforestation, stablization of eroded areas, and better water quality. The habitat varies from relatively xeric upland areas composed of oak-hickory-pine forests, to bottomland areas composed of oak-hickory and laurel-poplar-hemlock forests. The elevation ranges from 250–760 m. 2007 D.A. Aborn 295 Point counts During 2000–2003, ten-minute fixed-radius point counts were conducted weekly at five different locations on the Lula Lake property between late March and mid-June (Fig. 2). Point counts were conducted at the following locations: Bluff Trail: an upland oak-hickory-pine forest in the core property running along the ridge of Lookout Mountain. Ten points were established at this location. Middle Trail: a mixed-deciduous forest on the core property, about half-way between Rock Creek and the mountain ridge. Ten points were established at this location. Long Branch: a 162-ha oak-pine forest along Long Branch Creek. Eight points were established at this location. Five Points: a 202-ha upland section of oak-hickory-pine forest running along the mountain ridge. Ten points were established at this location. Figure 1. Location of the Lula Lake Land Trust, GA. 296 Southeastern Naturalist Vol. 6, No. 2 Ascalon: a 120-ha laurel-poplar forest, with patches of hemlock running along Price Branch Creek, and oak-hickory forest on the surrounding slopes. Eight points were established at this location. Counts began no later than 30 minutes after local sunrise, and the count at the last point started no later than 1000 EST. A single observer stood at each point and recorded all birds seen and heard within a 10- minute period. To estimate density, the observer also recorded whether each bird was inside or outside a 50-m radius centered on the observer. Figure 2. Location of point-count routes. A = Ascalon, B = Bluff Trail, F = Five Points, L = Long Branch, M = Middle Trail. 2007 D.A. Aborn 297 No attempts were made to attract birds during counts. Points were located along trails (with the exception of Ascalon, where a trail line had been flagged but not cut), and were at least 250 m apart to reduce the chances of recounting the same individuals from the previous point. To ensure adequate detection, counts were not conducted if the air temperature was below 0 °C, if there was more than a light drizzle falling, or if the wind speed was greater than 20 km/h (Robbins 1981). Point-count data were analyzed by summing the number of migrant detections at each site. I used the number of detections rather than the number of individuals since the same bird may have been seen or heard at a given point on more than one count. To account for this situation, as well as to compensate for differences in the number of points per site and the number of weeks a site was surveyed, I calculated the average number of individuals per point for each migrant species as: average of the # of detections for a given site/total # of points sampled/# of weeks of sampling for a given site. The number of individuals of each migrant species detected within 50 m was then used to calculate densities (birds/ha). Data were pooled within sites and across years, and I then ran repeated-measures analysis of variance (ANOVA) tests on the average number of detections per point, migrant density, migrant diversity, and number of species between years. Tukey’s multiple comparisons were run if any of the ANOVA tests were significant. I used a Shannon-Weiner diversity index on the pooled data to examine migrant diversity. All these analyses provide information on migrant abundance and richness, as well as community composition (DeSante 1986). A Pearson correlation analysis was run to look for significant population trends. Trends were examined only for species where there were at least 20 detections in all three years analyzed. Data from 2003 were excluded from all point-count analyses for three reasons. First, an unexpected construction project at Long Branch prevented access to the site and also created enough disturbance to potentially influence the data. Second, ownership of Five Points was transferred to the Georgia Department of Natural Resources (DNR). The property was subsequently gated, and I could not receive access. Third, the spring of 2003 was extremely rainy, which greatly reduced the number of counts that were conducted at the remaining sites. Mist netting As another way of inventorying birds at Lula Lake, as well as to assess the suitability of the property for breeding migrants, I mist netted and banded birds in the Middle Trail area. Six nets (12 m x 2.6 m, 30-mm mesh) were set up; 3 in wooded habitat and 3 in more open, shrubby habitat. The nets were opened within 30 minutes of local sunrise and run until 􀂧 1100 h, three days per week. Any birds that were caught were banded with an aluminum US Fish and Wildlife Service numbered leg band, sexed, aged, 298 Southeastern Naturalist Vol. 6, No. 2 and released. Netting data from 2003 were not excluded, as netting effort was not affected by some of the conditions that affected the point-count data (i.e., the lack of access to Long Branch and Five Points). Results Point counts Because of weather conditions, the sites were not surveyed all 12 weeks in each year of the study (Table 1). I tallied 1186 detections of 37 migrant species in 2000, 960 detections of 31 migrant species in 2001, and 1174 detections of 34 migrant species in 2002. Of 91 total species tallied, 40 (44%) were neotropical migrants. As shown in Table 2, Red-eyed Vireo was the most prevalent species in all three years of point counts. Scientific names of the species detected are listed in Table 2. There were no significant population trends between years. There was a significant difference in the number of neotropical migrant species between sites (F = 7.460, p = 0.008). Middle Trail had significantly more migrant species than Ascalon and Long Branch. The sites also differed in the number of detections of migrants (F = 16.467, p < 0.001), with Middle Trail having more detections than all other sites. There were no year-to-year differences in the number of migrant species or detections. There was also no difference in migrant density or diversity between sites or years (Table 3). Mist netting A total of 191 individuals of 41 species were captured during 1488 nethours between 2000 and 2003 (1 net open for 1 hour = 1 net-hour). Of these, 85 individuals (45%) and 23 species (56%) were neotropical migrants. Sixty-one percent of the migrants were in breeding condition, with either a brood patch or cloacal protuberance being present. Out of 15 individuals recaptured in subsequent years, 7 were migrants (Table 4). All the recaptured migrants were individuals that were in breeding condition at the time of their initial capture. Discussion Unfortunately, no data are available on migrant populations prior to reclamation, which makes it difficult to derive definitive conclusions regarding the impact reclamation actions of LLLT have had on long-distance Table 1. The number of times the Lula Lake sites were surveyed in different years. 2000 2001 2002 Five Points 9 6 5 Ascalon 8 8 6 Bluff 8 8 8 Long Branch 10 8 6 Middle 10 10 9 2007 D.A. Aborn 299 Table 3. Average number (standard error) of Neotropical migrant species, individuals, density, and diversity at the different Lula Lake sites. Site Five Points Ascalon Bluff Long Branch Middle Trail # of species 20 (1.33) 18 (0.88) 22 (1.00) 19 (1.53) 25 (0.88) # of detections 157 (46.76) 207 (11.62) 181 (9.68) 177 (27.54) 394 (13.54) Density 0.70 (0.06) 0.52 (0.02) 0.57 (0.08) 0.57 (0.08) 0.73 (0.02) Diversity index 1.41 (0.49) 1.42 (0.49) 1.36 (0.32) 1.44 (0.54) 1.50 (0.50) Table 2. Number of individuals per point at the Lula Lake Land Trust, GA, 2000–2002. Common name Scientific name 2000 2001 2002 Avg Red-eyed Vireo Vireo olivaceus Linnaeus 0.12 0.11 0.16 0.13 Scarlet Tanager Piranga olivacia Gmelin 0.08 0.07 0.10 0.08 Ovenbird Seuirus aurocapillus Linnaeus 0.07 0.05 0.08 0.06 Black-and-white Warbler Mniotilta varia Linnaeus 0.06 0.05 0.06 0.06 Hooded Warbler Wilsonia citrina Boddaert 0.04 0.05 0.07 0.05 Indigo Bunting Passerina cyanea Linnaeus 0.04 0.03 0.05 0.04 Wood Thrush Hylocicla mustelina Gmelin 0.02 0.01 0.02 0.03 Worm-eating Warbler Helmitheros vermivora Gmelin 0.02 0.03 0.04 0.03 Blue-gray Gnatcatcher Polioptila caerulea Linnaeus 0.02 0.03 0.04 0.03 Yellow-throated Vireo Vireo flavifrons Vieillot < 0.01 0.01 0.02 0.01 Acadian Flycatcher Empidonax virescens Vieillot 0.01 < 0.01 0.02 0.01 Louisiana Waterthrush Sieurus motacilla Vieillot 0.02 < 0.01 < 0.01 0.01 Yellow-breasted Chat Icteria virens Linnaeus < 0.01 < 0.01 0.01 < 0.01 Yellow-billed Cuckoo Coccyzus americanus Linnaeus < 0.01 < 0.01 0.01 < 0.01 Swainson’s Thrush Catharus ustulus Nuttall < 0.01 < 0.01 < 0.01 < 0.01 Ruby-throated Archilocus colubris Linnaeus < 0.01 < 0.01 < 0.01 < 0.01 Hummingbird Common Yellowthroat Geothlypis trichas Linnaeus 0.00 0.00 < 0.01 < 0.01 Yellow-throated Warbler Dendroica dominica Linnaeus < 0.01 0.00 < 0.01 < 0.01 Yellow Warbler Dendroica petechia Linnaeus 0.00 < 0.01 0.00 < 0.01 White-eyed Vireo Vireo griseus Boddaert < 0.01 < 0.01 < 0.01 < 0.01 Veery Catharus fuscescens Stephens < 0.01 0.00 0.00 < 0.01 Summer Tanager Piranga rubra Linnaeus < 0.01 < 0.01 < 0.01 < 0.01 Rose-breasted Grosbeak Pheucticus ludovicianus Linnaeus < 0.01 < 0.01 < 0.01 < 0.01 Prairie Warbler Dendroica discolor Vieillot < 0.01 0.01 < 0.01 < 0.01 Northern Rough-winged Stelgidopteryx serripennis < 0.01 < 0.01 < 0.01 < 0.01 Swallow Audubon Northern Parula Parula americana Linnaeus < 0.01 0.00 0.00 < 0.01 Kentucky Warbler Oporornis formosus Wilson < 0.01 < 0.01 0.01 < 0.01 Great-crested Flycatcher Myiarchus crinitus Linnaeus < 0.01 0.00 < 0.01 < 0.01 Eastern Wood-pewee Contopus virens Linnaeus < 0.01 < 0.01 < 0.01 < 0.01 Chimney Swift Chaetura pelagica Linnaeus < 0.01 < 0.01 < 0.01 < 0.01 Canada Warbler Wilsonia canadensis Linnaeus 0.00 0.00 < 0.01 < 0.01 Broad-winged Hawk Buteo platypterus Vieillot < 0.01 0.00 0.00 < 0.01 Blue-headed Vireo Vireo solitarius Coues < 0.01 < 0.01 < 0.01 < 0.01 Blue Grosbeak Passerina caerulea Linnaeus < 0.01 0.00 0.00 < 0.01 Black-throated-green Dendroica virens Gmelin < 0.01 0.01 < 0.01 < 0.01 Warbler Black-throated-blue Dendroica caerulescens Gmelin < 0.01 0.00 < 0.01 < 0.01 Warbler Blackpoll Warbler Dendroica striata Forster < 0.01 < 0.01 < 0.01 < 0.01 Blackburnian Warbler Dendroica fusca Statius Muller < 0.01 0.00 < 0.01 < 0.01 Bay-breasted Warbler Dendroica castanea Wilson < 0.01 0.00 0.00 < 0.01 American Redstart Setophagia reticulla Linnaeus < 0.01 0.00 < 0.01 < 0.01 300 Southeastern Naturalist Vol. 6, No. 2 migrants. There are, however, several indicators that suggest the restoration and management efforts at LLLT have improved the habitat quality for passage and breeding migrants. First, the proportion of migrants on the Trust is comparable to that of reclaimed mining land elsewhere, as well as other locations in the southern Appalachians (Franzreb and Rosenberg 1997). A study of a strip-mined area in east Tennessee found a greater number of species in edge habitats compared with forest habitats (Yahner and Howell 1975). Within both the disturbed habitat and the undisturbed forest, however, five of the seven most commonly detected species were the same as some of the most commonly detected migrants at LLLT (Red-eyed Vireo, Hooded Warbler, Scarlet Tanager, Ovenbird, and Wood Thrush). In a similar study elsewhere in east Tennessee, Curtis et al. (1978) found that breeding- bird diversity on a 13-year-old area reclaimed with a variety of hardwoods, pines, and shrubs was similar to that of nearby undisturbed forests. Migrant diversity increased from 23 species to 54 species just 5 years following reclamation. Most recently, seven years of data on avian communities in reclaimed land, open–canopy forests, and closed-canopy forest found no difference in diversity or abundance among the three habitats (Lacki et al. 2004). Out of 110 species recorded on reclaimed lands, onethird were long-distance neotropical migrants, and 10 species of migrants Table 4. Number of birds mist netted and recaptured at the Lula Lake Land Trust. Scientific names given for species not mentioned in the Table 2. Species Number captured (recaptured) Acadian Flycatcher 3 Black-and-white Warbler 2 Blue-headed Vireo 2 Common Yellowthroat 1 Chestnut-sided Warbler (Dendroica pensylvanica Linnaeus) 1 Gray-cheeked Thrush (Catharus minimus Lafresnaye) 1 Gray Catbird (Dumetella carolinensis Linnaeus) 1 Hooded Warbler 8 (1) Indigo Bunting 9 (1) Magnolia warbler (Dendroica magnolia Wilson) 4 Ovenbird 6 Ruby-throated Hummingbird* 3 Red-eyed Vireo 3 Summer Tanager 2 Swainson’s Thrush 4 Tennessee Warbler (Vermivora peregrine Wilson) 1 Veery 1 White-eyed Vireo 11 (2) Worm-eating Warbler 5 Wood Thrush 5 Yellow-breasted Chat 10 (3) Yellow-throated Vireo 1 Yellow Warbler 1 Total 85 * = captured but not banded. 2007 D.A. Aborn 301 were passage migrants, reflecting the suitability of the habitat as both a stopover site and breeding area (Lacki et al. 2004). A second indicator that reclamation efforts have been successful, at least for the Middle Trail area, is the fact that seven individual migrants returned to the area to breed in subsequent years. Research on some migrant species indicates that prior breeding success is an important factor in breeding-site fidelity (Gavin and Bollinger 1988, Hoover 2002, Payne and Payne 1993, Porneluzi 2003, Sedgwick 2004). For example, in experimental manipulations of reproductive success, Hoover (2002) found that territory fidelity of Protonotaria citrea Boddaert (Prothonotary Warblers) increased where pairs raised two broods, and declined in birds that were unsuccessful in breeding. Similarly, only 9% of Ovenbird males that paired but failed to raise young returned the following year, compared with 54% of successful males (Porneluzi 2003). Female Empidonax traillii Audubon (Willow Flycatchers) also had return rates that increased with breeding performance; however, male fidelity was not correlated with reproductive success (Sedgwick 2004). Finally, while trends were not significant, eight of the 10 most abundant species showed increasing or stable trends over the three years of surveys. If the habitat was unsuitable, then population declines might be expected. It is possible, however, that the habitat is unsuitable, and that territory vacancies became occupied by floaters, thus giving the appearance of a stable population (McShea et al. 1995, Rappole and McDonald 1994). The difference in migrant diversity and abundance among sites is likely a factor of habitat extent and disturbance. Middle Trail is located at the center of the core property and therefore might be considered the largest tract of interior forest. Middle Trail had significantly more detections than the other sites, and had more species than Ascalon and Long Branch. Ascalon and Long Branch are smaller in area, and also suffer encroachment from residential development. Bluff Trail and Five Points are also larger in extent than Ascalon and Long Branch, but since they are located on ridge tops the forests are exposed to greater wind disturbance, and also have a greater slope. These factors may create less favorable habitat characteristics, such as a more open canopy, drier soils, or unsuitable nest locations. Management of the Trust properties should continue to emphasize maintaining large areas of mature forest, as many of the migrants detected are area sensitive (Robbins et al. 1989). To that end, LLLT has partnered with Georgia DNR and other agencies to join separate properties into a single 3300-ha corridor. While much of this corridor will have limited access to the public, there will nonetheless be multi-use trails, picnic areas, and a visitor center. Trust managers should be sensitive to the possible effects of disturbance on bird populations, especially during breeding (see Cooke 1980, Gutzwiller et al. 1998, Riffel et al. 1996). Additionally, measures should be taken to ensure that the increased human activity does not degrade habitat quality. Further monitoring of migratory 302 Southeastern Naturalist Vol. 6, No. 2 bird populations, including examination of return rates, survivorship, and reproductive success, should be conducted as a way of monitoring the health of the LLLT ecosystem. Acknowledgments I thank the following administrators and staff of the Lula Lake Land Trust for their cooperation: Bill Chipley, Adelaide Bratcher, Paul Grant, and Matt Fox. Craig Ely, Melea Langley, and Stephen Hales provided invaluable assistance in the field. 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