The Great Smoky Mountains National Park All Taxa Biodiversity Inventory:
A Search for Species in Our Own Backyard
2007 Southeastern Naturalist Special Issue 1:215–228
1Department of Ecology and Evolutionary Biology, 569 Dabney Hall, University of
Tennessee, Knoxville, TN 37996. 2Department of Zoology, 381 David Clark Labs,
North Carolina State University, Raleigh, NC 27695. 3USGS Biological Resources
Discipline, 1314 Cherokee Orchard Road, Gatlinburg, TN, 37738. *Corresponding
author - firstname.lastname@example.org.
Rarity and Diversity in Forest Ant Assemblages of
Great Smoky Mountains National Park
Jean-Philippe Lessard1,2,*, Robert R. Dunn2, Charles R. Parker3,
and Nathan J. Sanders1
Abstract - We report on a systematic survey of the ant fauna occurring in hardwood
forests in the Great Smoky Mountains National Park. At 22-mixed hardwood
sites, we collected leaf-litter ant species using Winkler samplers. At eight of those
sites, we also collected ants using pitfall and Malaise traps. In total, we collected 53
ant species. As shown in other studies, ant species richness tended to decline with
increasing elevation. Leaf-litter ant assemblages were also highly nested. Several
common species were both locally abundant and had broad distributions, while many
other species were rarely detected. Winkler samplers, pitfall traps, and Malaise traps
yielded samples that differed in composition, but not richness, from one another.
Taken together, our work begins to illuminate the factors that govern the diversity,
distribution, abundance, and perhaps rarity of ants of forested ecosystems in the
Great Smoky Mountains National Park.
Robert Whittaker's (1952, 1956) classic work showed that the elevational
gradient in the Great Smoky Mountains National Park (GSMNP) strongly
infl uences plant and insect communities. Since Whittaker’s work, few studies
(see Stiles and Coyle 2001, Van Pelt 1963, Watson et al. 1994) have
explored elevational gradients in diversity in the southern Appalachians.
As for most taxa, ant diversity often varies systematically along elevational
gradients ( Brühl et al. 1998; Fisher 1996, 1998; Olson 1994; Sanders 2002;
Sanders et al. 2003), but no studies to date have explicitly examined elevational
gradients in ant diversity in the southern Appalachians.
At least two early investigators focused on the ant fauna of the southern
Appalachians. First, Cole (1940) performed one of the earliest ant surveys of
any National Park and one of the first systematic surveys of any taxon in the
GSMNP (though he focused only on the Tennessee side of GSMNP). Cole
(1940) thoroughly inventoried the ant fauna of the GSMNP and provided
notes on the distribution and autecology of the species he observed in his
forays. Second, Van Pelt (1963) studied the ant communities of the southern
Blue Ridge Mountains with a particular interest in the variation in regional
216 Southeastern Naturalist Special Issue 1
patterns of ant diversity and community composition at low and high-elevation
sites. These early studies by Cole and Van Pelt provide contemporary
ecologists with a unique knowledge of the history of ant assemblages in the
southern Appalachian Mountains.
Because of the ecological importance and near ubiquity of ants in most
terrestrial ecosystems (Hölldobler and Wilson 1990), understanding the
causes and consequences of ant diversity is critical to preserving both ecosystem
functions and services that ants provide (Folgarait 1998). As part
of the All Taxa Biodiversity Inventory (ATBI; [Sharkey 2001]), we have
sought to understand the biotic and abiotic controls on ant diversity. Our
goals in this paper are to: (i) estimate litter ant species richness at 22 forest
sites in the GSMNP; (ii) document the major spatial patterns in the diversity,
distribution, and abundance of ants in the GSMNP; (iii) assess whether
ant assemblages are nested, where nestedness is a measure of the extent to
which species-poor assemblages are subsets of species-rich assemblages;
and (iv) examine how different sampling techniques yield different components
of ant assemblages. To accomplish the first three goals, we used data
from leaf-litter ant assemblages collected at 22 sites during 2004 and 2005.
To accomplish the fourth goal, we used a combination of different sampling
techniques (pitfall traps, Malaise traps, and Winkler extractors) to inventory
ant workers and alates at eight sites in the GSMNP.
We collected ants at 22 forested sites using Winkler extractors in 2004
and 2005. At eight of those 22 sites, we also collected ants using pitfall traps
and Malaise traps as part of C. Parker’s “ATBI Pilot Study.” The 22 sites
were chosen to cover nearly the entire range of elevation in the GSMNP
(260–2021 m). Our sampling design ensured that about 80% of the elevational
range in the GSMNP was sampled. Sites were chosen on both the NC and TN
side of the GSMNP, and from the southern to the northern boundaries of the
park (Fig. 1). All sites were located in mixed-hardwood forests, which is the
main forest type found throughout the park (White 1983), and all sites were
located away from roads, heavily visited trails, or other human disturbances.
Leaf-litter ant sampling (Winkler Extraction). At each of the 22 sites
described above, we randomly placed a 50-m x 50-m plot. Within the corners
of this plot, we placed a 10-m x 10-m sub-plot, and within the corners
of each 10-m x 10-m sub-plot, we sampled ants in four 1-m2 plots. Thus, at
each site, there were 16 1-m2 plots. We collected the leaf litter inside each
1-m2 plot and sifted it through a coarse mesh screen of 1-cm grid size to
remove the largest fragments and concentrate the fine litter (see Longino
and Colwell 1997, Longino et al. 2002). The litter fragments that did not
fit through the mesh, twigs, and sticks in each 1-m2 plot were inspected for
colonies. The concentrated fine litter from each sample was then suspended
2007 J.-P. Lessard, R.R. Dunn, C.R. Parker, and N.J. Sanders 217
in mini-Winkler sacks for two days in the lab. All worker ants that were
extracted from the 1-m2 plots were identified and enumerated and are stored
in Sander's ant collection at the University of Tennessee.
Pitfall trapping. We used pitfall traps to sample ants at eight forested
sites (Table 1) within the GSMNP every two weeks from October 2000
through April 2003. At each of the eight sites, ten pitfall traps were placed
at least 3 m apart along an approximately 30-m transect. Traps were 6-cm
diameter plastic cups buried fl ush with the soil surface and partially filled
with propylene glycol. Pitfall traps effectively capture ground-foraging ants
(Bestelmeyer et al. 2000), while not always capturing those that are exclusively
Malaise traps. Two Malaise traps (1.6 m × 1.8 m × 1.0 m) were placed
on the ground 75–100 m from one another at each of the eight sites at which
pitfall traps were placed (Longino and Colwell 1997, Longino et al. 2002).
The contents of the alcohol-filled traps were collected every two weeks from
January 1999 through January 2002.
For each site and sampling technique, the observed number of species is
simply the tally of species collected at the site. We examined how species
richness (the total number of ant species occurring at a site) varied with
elevation. To estimate species richness, the number of species that would be
collected if sampling were to go to completion, we used the Chao2 estimator
(Chao 1987, Colwell and Coddington 1994) as:
SChao2 = SObs + Q1
2 / 2Q2,
where SObs is the number of species that occurred in the sample, Q1
is the number of species that occur in only one sample (uniques), and Q2 is
Figure 1. Map of the Great Smokies National Park (GSMNP) showing the 22 sites
(white dots) where leaf-litter ants were sampled. The shadings represent the repartition
and proportion of the main habitat-cover types in the GSMNP.
218 Southeastern Naturalist Special Issue 1
Table 1. Minimum and maximum elevation and number of sites occupied for forest ant species
sampled in Great Smoky Mountains National Park. The elevation records are based on 8 sites
for which data for the three different sampling techniques (malaise, pitfall, and Winkler sacks)
were available. The number of occurrences are shown for each of the sampling techniques.
Species Min Max Pitfall Winkler Malaise
Amblyopone pallipes (Haldeman) 594 1530 5 5 6
Aphaenogaster fulva Roger 594 1530 3 1 0
A. lamellidens Mayr 1530 1530 1 0 0
A. rudis Enzmann 594 1673 6 7 6
A. sp. 594 1380 0 0 2
Brachymyrmex depilis Emery 594 896 2 0 0
Camponotus americanus Mayr 594 594 0 0 1
C. chromaiodes Bolton 594 1033 4 0 4
C. mississippiensis Smith 594 594 0 0 1
C. nearcticus Emery 594 1380 1 0 3
C. pennsylvanicus (De Geer) 594 1530 3 1 5
C. snellingi Bolton 594 594 0 0 1
C. subbarbatus Emery 594 594 0 0 1
Crematogaster ashmeadi Mayr 594 594 0 0 1
C. pilosa Emery 594 1673 1 0 1
C. sp. 594 594 0 0 1
Cryptopone gilva (Roger) 594 896 2 0 0
Formica subsericea Say 594 994 0 0 2
Lasius alienus (Foerster) 594 1673 3 5 4
L. latipes (Walsh) 594 1530 0 0 2
L. nearcticus Wheeler 594 594 1 0 0
Monomorium minimum (Buckley) 594 594 0 0 1
Myrmecina americana Emery 594 1530 6 5 5
Myrmica latifrons Cole 594 896 0 0 2
M. pinetorum Wheeler 594 896 2 0 2
M. punctiventris Roger 594 1673 2 2 2
M. sp. 594 594 0 0 1
Paratrechina sp. 594 594 0 0 1
P. sp1. 594 594 0 0 1
Ponera pennsylvanica Buckley 594 1673 3 2 4
Prenolepis imparis Emery 594 1530 4 1 2
Proceratium croceum (Roger) 594 594 0 0 1
P. pergandei (Emery) 594 594 0 0 1
P. sp. 594 594 0 0 1
P. sp.1 594 594 0 0 1
Pyramica ohioensis (Kennedy & Schramm) 594 594 0 1 0
P. ornata (Mayr) 594 594 1 0 0
P. rostrata (Emery) 594 594 0 1 0
P. sp. 594 594 0 0 1
Solenopsis molesta (Say) 594 1033 2 0 1
Stenamma brevicorne (Mayr) 594 1530 5 3 0
S. diecki Emery 594 1673 7 6 1
S. impar Forel 1033 1530 1 1 0
S. meridionale Smith 594 1673 6 5 0
S. schmittii Wheeler 594 1673 5 3 0
S. sp. 594 594 0 0 1
S. sp.1 994 1530 0 0 3
S. sp.2 594 1673 0 0 7
Tapinoma sessile (Say) 594 1530 0 0 2
Temnothorax curvispinosus Mayr 594 1033 0 0 3
T. longispinosus Roger 594 1380 2 1 4
T. sp. 594 1033 0 0 2
T. sp.1 1380 1380 0 0 1
2007 J.-P. Lessard, R.R. Dunn, C.R. Parker, and N.J. Sanders 219
the number of species that occur in two samples (Colwell and Coddington
1994). The Chao2 index uses data on the rare species collected in the samples
(Q1 and Q2) to estimate the number of additional species that are present
at the site, but were not recorded in the samples. As in other biodiversity
studies of this kind, (Colwell et al. 2004, Kaspari et al. 2000, Longino et al.
2002, Ratchford et al. 2005), we treated each sampling unit (a 1-m2 quadrat
for the Winkler sampling or a pitfall trap for the pitfall trapping) as a sample.
Because the Chao2 estimator is sensitive to sample size (Colwell and Coddington
1994), we used Colwell’s EstimateS (Colwell 2004) to construct
50 randomized accumulation curves for each site to calculate the standard
deviation of the estimated species richness. Across all sites, the asymptotic
richness estimator was very similar to the observed total species richness
(r2 = 0.64, p < 0.001), and sampled diversity reached a plateau at all but two
sites. Therefore, for ease of interpretation, we report only the observed richness
for each site and sampling technique.
Elevational gradient. To examine whether leaf-litter ant diversity varied
systematically with elevation, we plotted observed species richness from the
Winkler samples from 22 sites against elevation. We assessed the relationship
using linear regression.
Nestedness and rarity. We conducted two analyses to understand the
distribution of rare ant species in the GSMNP hardwood forests: a nestedness
analysis and a simple correlation between the diversity of rare ants and
overall diversity. Nestedness analyses provide a measure of the extent to
which the species found at species-poor sites are exclusive of or a subset of
those found at species-rich sites. If sites are nested, those species at speciespoor
sites are a subset of those at species-rich sites. If sites are not nested,
then species at species-poor sites are not necessarily found in species-rich
sites. We performed an analysis of community nestedness using a presence/
absence matrix comprising 22 sites and 38 species detected in the leaf-litter
ant sampling. Nestedness can be assessed using the “nestedness temperature
calculator” (NTC) implemented by Atmar and Patterson (1995). The NTC
provides a T value between 0 and 100 describing the degree of nestedness
of a given set of communities. A T value close to 0 is highly nested (where
species at less diverse sites are strict subsets of those at more diverse sites),
whereas a T value near 100 describes a random assemblage. The NTC further
allows testing for statistical significance by generating 50 random matrices
based on the original data set. The mean T value produced in the process is
then compared to the observed T value and used to calculate a confidence
interval. As an additional test of the distribution of rare ant species in the
GSMNP forests, we looked at the correlation between the diversity of rare
species (defined as those species found at no more than 4 sites) and overall
ant diversity. In many regions, the diversity of rare species and overall diversity
do not co-vary (Jetz et al. 2004), and as a consequence, conservation
of diversity per se will not necessarily conserve those species most at risk.
Abundance and distribution. We examined the shape of the abundance220
Southeastern Naturalist Special Issue 1
distribution curve by ordering species’ frequencies of occurrence (the
number of times they were detected) in litter samples, from the most rare to
the most common species. This allowed us to illustrate the relative proportion
of rare and common species in our litter samples. Then, to test whether
abundant ants also tend to be widespread within the GSMNP forests, we
regressed the number of occurrences (the number of 1-m2 plots in which a
species was detected) against the number of sites (out of 22) at which it was
detected. If many species were uniques and/or singletons, it would be an
indication that sampling at the scale of the park was relatively incomplete
even though sampling in individual sites seemed relatively complete (see
Comparing sampling techniques. To compare the three sampling techniques
(Winkler samples, pitfall traps, and Malaise traps), we first asked
whether the number of species collected by one technique was correlated
with the number of species collected by the other two techniques at the eight
sites at which each of the three sampling techniques were employed. Then,
to assess the similarity in composition of the assemblages sampled with
each technique, we used the Jaccard’s similarity index. The comparison was
limited to the 8 sites where all three sampling techniques had been used.
Leaf-litter ant assemblages. In total, we detected 38 leaf-litter ant species
at the 22 sites. The number of species per m2 ranged from 0–10, and the number
of species per site varied from 2–22. In 20 of the 22 sites, the estimators
reached an asymptote, indicating that further sampling would have added
no new species. Leaf-litter ant species richness declined significantly with
increasing elevation (Fig. 2, r2 = 0.63, p < 0.001).
Nestedness and rarity. The 22 litter-ant assemblages sampled were
nested (T =18.37°). The NTC randomization process generated 50 matrices
that had an average T value of 62.2 ± 4.11°. The original matrix had a significantly lower temperature than the mean T for the simulated matrices (p =
2.64e-26), indicating that species-poor assemblages were composed of a subset
of species-rich assemblages. The core species of most assemblages were
Aphaenogaster rudis, Myrmecina americana, Stenamma diecki, S. meridionale,
Lasius alienus, Amblyopone pallipes and Ponera pennsylvanica (Fig.
3). The diversity of rare species was well correlated with overall diversity,
such that the most diverse sites had the most rare species (Pearson r = 0.87,
p = 0.001 ).
Abundance and distribution. The species abundance distribution was approximately
log normal, with a few common species and a tail of rare species
(Fig. 4a). The abundance of individual species (the number of 1-m2 plots a
species was detected in) increased with the number of sites they occupied
(Fig. 4b, r2 = 0.74, p < 0.0001). One “species,” A. rudis, represented a large
percentage of all occurrences. Aphaenogaster rudis is likely a group of species
rather than a single species (Umphrey 1996), but distinguishing species
2007 J.-P. Lessard, R.R. Dunn, C.R. Parker, and N.J. Sanders 221
within the group is possible only by examining their karyotypes. Regardless,
the A. rudis species group appears to be tremendously successful in the
southern Appalachia, a success that warrants further examination.
Pitfall and Malaise samples. Our inventory of the eight ATBI sites yielded
a total of 15,340 ant individuals and 53 species, all sampling techniques
(Winkler + pitfall + Malaise traps) combined (Table 1). Malaise traps at the
same eight sites captured 30 species and over 10,000 alates ; pitfall traps
captured 25 species and 2796 workers; and litter samples yielded 17 species
and 2242 workers. Due to the limited availability of taxonomic work on ant
alates, we identified them to morphospecies and tried to be as conservative
as possible in splitting species to avoid infl ating the count of the number of
species. We expect this conservatism to cause only a slight, if any, increase
in the number of species recorded, and thus no significant effect on the relative
species richness yielded by the different sampling techniques.
Comparing sampling techniques. To assess whether different sampling
techniques provide equivalent estimates of the number of species
sampled at our sites and the overall pattern of diversity, we correlated
the different measures of species richness with one another. Measures of
observed species richness were positively correlated to each other (Table
Figure 2. Elevational gradient in leaf-litter ant species richness for 22 sites surveyed
during summer 2004–2005 in the Great Smokies National Park.
222 Southeastern Naturalist Special Issue 1
2). The myrmecofauna collected with pitfall traps and Winkler extractors
were similar to one another, but the fauna detected in the Malaise traps
differed markedly from the faunas collected in Winkler extractors and
pitfall traps (Table 3).
To our knowledge, our work is the first systematic sampling of the ant
fauna in the GSMNP. As with other taxa in the GSMNP (e.g., Whittaker
Figure 3. Presence/absence matrix of forest ant species illustrating the nestedness
pattern of the 22 leaf-litter ant assemblages. Species are sorted from top to bottom
by the number of sites they occcupy.
Table 2. Correlation coefficients showing relationship between richness detected by three
First sample Second sample Correlation coefficient P
Pitfall traps Winkler samples 0.835 0.010
Pitfall traps Malaise traps 0.878 0.004
Winkler samples Malaise traps 0.847 0.008
Table 3. The similarity in species composition among assemblages collected with three sampling
techniques. Jaccard’s values near 1 indicate more similarity, and values near 0 indicate
First sample Second sample Jaccard similarity index
Pitfall traps Winkler samples 0.703
Pitfall traps Malaise traps 0.480
Winkler samples Malaise traps 0.392
2007 J.-P. Lessard, R.R. Dunn, C.R. Parker, and N.J. Sanders 223
Figure 4. a) Species abundance distribution in 22 sites (black lines) and 288 onem2
plots of leaf-litter samples, and b) correlation between the relative abundance
of leaf-litter ant species detected and the number of sites at which they occur.
224 Southeastern Naturalist Special Issue 1
1952), elevation strongly infl uenced leaf-litter ant diversity. Ant species
richness decreased monotonically with elevation, a common pattern for
many insect elevational diversity gradients (Rahbek 2005). Our results support
the observations made by Van Pelt (1963), who found a greater number
of ant “forms” and “nests” at lower elevations in the Blue Ridge mountains
than at higher elevations. Similarly, Wang et al. (2001) found that ant species
richness decreased with elevation in an oak forest of the central Appalachians.
A regional faunistic survey of the ants of Georgia (Ipser et al. 2004)
also found that ant species richness generally declined with elevation. Of
course, the ants do not really respond to elevation per se. Instead, they respond
to some biotic (e.g., productivity) or abiotic (temperature, geometric
constraints) variable that covaries with elevation. The next step in our work
in the GSMNP is to understand the biotic and abiotic factors that shape both
spatial and temporal gradients in diversity (Dunn et al. 2007).
Leaf-litter ant assemblages in the GSMNP are highly nested. Speciespoor
assemblages (generally at the highest elevations) are made up of a
subset of those species that occur at the most species-rich assemblages
(generally at the lowest elevations). As a consequence, the diversity of
rare species tracked the overall pattern of diversity. Ellison et al. (2002)
calculated a T value of 15.1 for the bog ant assemblages in New England,
comparable to the value we observed here. The ant fauna of the New England
bogs is characterized by a few “bog specialists;” here we find a few
In our study, ant species collected at the high-elevation sites were widely
distributed across the elevational gradient surveyed, whereas the distributions
of low-elevation species were often restricted to a few low-elevation
sites. Aphaenogaster rudis, M. americana, P. pennsylvanica, and S. diecki
were widely distributed along the elevational range covered by our study.
The most frequently collected ant species, A. rudis, was abundant within
plots, common across multiple plots within sites and found at most sites in
the GSMNP, while others species were represented by just a few individuals
in one or a few plots (e.g., Cryptopone gilva, Pyramica ornata, Proceratium
pergandei). The causes of rarity and abundance in ants remain poorly
explored. Some authors (e.g., Davidson et al. 2003) have suggested that
the most abundant ants (in tropical forest canopies) tend to be homopteratenders
that can monopolize large territories and pools of sugar resources.
In contrast to ants in tropical systems, A. rudis, a behaviorally subordinate
ant (Fellers 1987, Smallwood 1982), is not known to rely extensively on
homoptera exudates, and does not maintain exclusive foraging territories
(Lessard, pers. observ.; Smallwood 1982). Instead, Aphaenogaster rudis,
like other species in the genus Aphaenogaster (ants in this genus are referred
to as the gypsy ants; T.G. McGlynn, California State University, Dominguez
HillsCarsn, California, pers. comm.), migrates from nest to nest frequently
and feeds on a wide variety of food resources. Interestingly, all of these
behaviors are shared by Aphaenogaster araneoides Emery in Costa Rica,
2007 J.-P. Lessard, R.R. Dunn, C.R. Parker, and N.J. Sanders 225
where A. araneoides is extremely abundant in an even more diverse ant
community (McGlynn 2006, McGlynn et al. 2004).
The rare ants, those species found at only a few sites, represented a mix
of different life histories. Rarity may be real or only apparent. Apparent
rarity means that the low abundance of certain species in a set of samples
is strictly a sampling artifact. In the current study, the combination of sampling
techniques and the high correlation between observed and predicted
richness suggest that the species that we describe as being rare actually are
rare. Furthermore, species that were rare in space (litter samples) were also
rare in time (two years of pitfall trapping), suggesting that they have both
low local abundance and low frequency of occurrence at the meso-scale.
Most of the rare species may be at the edge of their climate envelope within
the GSMNP and hence found predominately at the lowest, warmest, and
most diverse sites within the park.
Perhaps not surprisingly, different sampling techniques detected different
ant species (Longino et al. 2002, Martelli et al. 2004). The composition
of assemblages collected solely by litter extraction did not differ dramatically
from the assemblages detected by pitfall traps. However, it is worth
noting that there were two Pyramica spp. that were detected in the leaf-litter
samples that were not collected in the pitfall traps. Similarly the reproductive
ant fauna collected in the Malaise traps was different from either
the pitfall or leaf-litter samples. Taken together, these results suggest that
documenting ant diversity in the GSMNP will require multiple sampling
techniques, as is the case in other systems (Delabie et al. 2000, Longino et
al. 2002). However, it is worth noting that most (68%) of the identifiable ant
species collected here were collected by litter extraction and that the overall
patterns of diversity detected by the different methods were highly concordant
such that all sampling methods showed similar declines in diversity
with increasing elevation.
The ant species that occur at these high-elevation sites generally have
broad elevational ranges, whereas many of the species that occur at low
elevations are found almost exclusively at low-elevation sites. Similarly,
many of the species that occur at high elevations, with broad elevational
ranges, also occur at high latitudes (e.g., Lessard and Buddle 2005) and
have broad latitudinal ranges. Ant assemblages in GSMNP are highly
nested, and most species-poor sites in GSMNP occur at high elevations.
This suggests that these broad-ranged species are able to tolerate climatic
extremes that are frequent at high elevations and latitudes. Unlike examples
of elevational gradients in the southwestern US (e.g., Fleishman et
al. 2000), we find few high-elevation endemics and a number of relatively
rare species at low elevation. More extensive sampling, especially of other
habitat types in GSMNP, could further illuminate the causes and consequences
of ant diversity.
226 Southeastern Naturalist Special Issue 1
Thanks to Jaime Ratchford, Melissa Geraghty, Raynelle Rino, Kerri Crawford,
Donny Mai, and Kristin Lane for help in the field and Matt Fitzpatrick for help with
Figure 1. We are grateful to Noa Davidai, James Trager, and two reviewers who
provided comments on the manuscript. This research was funded by grants to N.J.
Sanders and R.R. Dunn by Discover Life in America.
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