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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.
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.
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
2007 D.A. Aborn 295
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
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
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
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.
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).
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).
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.
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
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
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
Black-throated-blue Dendroica caerulescens Gmelin < 0.01 0.00 < 0.01 < 0.01
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
Ruby-throated Hummingbird* 3
Red-eyed Vireo 3
Summer Tanager 2
Swainson’s Thrush 4
Tennessee Warbler (Vermivora peregrine Wilson) 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
* = 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.
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.
Three anonymous reviewers greatly improved the quality of this manuscript.
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