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Insect species associated with eastern hemlock in the Great Smoky Mountains National Park and environs.

 ABSTRACT -- Eastern hemlock, Tsuga canadensis (L.), Carr is an
 integral part of forested systems in eastern North America now
 threatened by invasions of exotic pests. Due to the threat of
 exotic pests, information on the species associated with eastern
 hemlocks was collected to compile a species listing. The insect
 fauna associated with eastern hemlock was assessed at four sites
 representing new and old growth trees and ten alternate sites in the
 Great Smoky Mountains National Park and adjacent areas in eastern
 Tennessee. Sites were sampled using malaise/pan traps, pitfall traps
 and direct collection. Species diversity was assessed using the
 Shannon index and species richness estimates were made using the
 program estimates. The rich insect fauna of eastern hemlocks yielded
 2,832 specimens representing 292 species in 101 families and ten
 orders. Species richness and abundance were highest at Chimney Tops
 old growth site with 107 species and 801 specimens. Species richness
 estimators projected between 415 and 550 species associated with
 eastern hemlock. Several pests of eastern hemlock, including the
 hemlock woolly adelgid, Adelges tsugae (Annand), elongate hemlock
 scale, Fiorinia externa Ferris, hemlock scale, Abgrallaspis ithacae
 (Ferris), and hemlock looper, Lambdina fiscellaria (Guenee), as well
 as natural enemies of these pests, were collected.


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Eastern hemlock, Tsuga canadensis (L.), Carr is an important component of forests throughout the eastern United States and possesses both intrinsic and unique values. Eastern hemlocks are ecologically important representing an integral component of many old growth communities providing distinct microclimates for an array of wildlife. The areas dominated by eastern hemlocks are cool and shaded making them attractive to both wildlife and people. This tree is often found near streams where it reduces the water temperature making the stream favorable to native brook trout. Within forests where eastern hemlocks are threatened can be found a myriad of outdoor recreational opportunities, a rich and diverse assemblage of regional flora and fauna, scenic waterfalls and historic structures, as well as a variety of environmental and outdoor educational opportunities. Also, eastern hemlock is among the most widely grown evergreens in ornamental landscapes. Because of these scenic mountains in the Southern Appalachians, more than 14 million visitors annually contribute over five billion dollars to the local economy of eastern Tennessee.

Forest decline is an issue confronting the southern Appalachian region with significant damage caused by several invasive insect pests. Introduced species such as Adelges tsugae (Annand), hemlock woolly adelgid, and Fiorinia externa Ferris, elongate hemlock scale, represent pernicious pests with the potential to cause widespread destruction to eastern hemlocks in the eastern United States (Danoff-Burg and Bird, 2002; McClure and Fergione, 1977; Stimmel, 1980, 2000). Wallace and Hain (2002) concluded that none of the four established predators collected from eastern hemlock had a significant impact on populations of hemlock woolly adelgid. The Great Smoky Mountains National Park (GSMNP) includes some of the largest remnants of eastern hemlock in the world, which are presently confined to about 35,399 ha at various elevations. Because information on the status of the insect fauna on eastern hemlock within the region is lacking, this study was initiated to identify and assess those insect species associated with this important tree.

MATERIALS AND METHODS

Four primary test sites (each 20 m by 40 m), each consisting of three trees, were established representing mature and new growth hemlocks at high and low elevation gradients within the GSMNP in eastern Tennessee. Two lower sites (760 m) at Elkmont representing new (35[degrees]39'56.388"N, 83[degrees]35'04.915"W) and old (35[degrees]39'47.733"N, 83[degrees]35'10.036"W) growth trees and two higher sites (1,149 m) at the Chimney Tops representing new (35[degrees]38'1.74"N, 83[degrees]28'11.4"W) and old (35[degrees] 37'49.44"N, 83[degrees]28'3.18"W) growth trees were selected for study. These sites were located in areas not infested by the hemlock woolly adelgid. Each site consisted of three old growth (diameter at breast height > 20 cm) or new growth (diameter at breast height; dbh < 20 cm) trees. The Elkmont new growth site is located in a xeric oak forest (type 7) and the Elkmont old growth site is part of a pine forest (type 9). Chimney Tops old growth is located in a tulip poplar forest (type 6), while the Chimney Tops new growth site is located in cove hardwoods (type 3). Specimens were obtained from these four sites using Malaise/pan traps, pitfall traps, and direct (beat-sheet, sweep-net and handpicking) sampling from 1 June 2002 through 30 November 2002 and from 5 June 2003 through 2 September 2003.

Modified malaise/pan traps were placed in the tree canopy of each of three trees at the four primary test sites in the GSMNP to sample the insect fauna. Trap frames were constructed using PVC pipe (60 cm by 60 cm by 60 cm) and covered with polyester netting (#156). The collecting unit consisted of a plastic cup (60 mm wide by 65 mm deep, 120 mL volume) that contained 30-60 mL of 50% propylene glycol (Sierra[R]) and tap water. The pan (15 cm wide by 65 cm wide by 12 cm deep) was hung under the frame and contained 900-1000 mL of 50% propylene glycol and water. Samples were obtained from all collection units and pans biweekly, labeled, and taken to the laboratory for processing. Pitfall traps were used to sample ground-dwelling species at two trees per site. Four shallow holes (8 cm deep; one in each cardinal direction at the canopy's peripheral edge) per tree were dug into the ground for placement of traps. Each trap consisted of two plastic cups (60 mm wide by 65 mm deep/120 mL volume) with a plastic cover. One cup was placed inside the other to aid in sample collection and reduce flooding. The outer cup had a drainage hole, while the inner cup was filled with a 50% mixture of propylene glycol and tap water. Plastic covers with 90[degrees] directional fans were placed on the surface of the ground above the pitfall traps to prevent flooding and direct insects into the trap. Two pitfall traps at each tree were retrieved every 14 days/site and taken to the laboratory for processing and identification. On each sampling date, the collection cup with preservative was removed and replaced with fresh preservative. Visual observations and direct sampling of insects were conducted every 14 days (15-20 min/tree) within each study site using beat sheet, sweep-net and handpicking to obtain specimens from the branches and foliage. Sweep-net (a canvas net bag 38 cm diameter and 82 cm deep) and beat-sheet (1 [m.sup.2]) samples were collected, placed into 2 d vials or zip-lock bags, labeled (date, site number, host number), and taken to the laboratory for processing and identification.

Specimens also were obtained during this time period using direct sampling (beat-sheet, sweep-net, handpicking) from 12-13 trees at eight additional sites in the GSMNP that had been found to be recently infested with the hemlock woolly adelgid (2002). These included: Anthony Creek (83[degrees]44'32.99"W, 35[degrees]34'47.45"N), Cataloochee Cove (83[degrees]5'38.86"W, 35[degrees]36'8.65"N), Gregory Ridge (83[degrees]50'1.47"W, 35[degrees]32'53.16"N), Laurel Falls (83[degrees]33'57.26"W, 35[degrees]40'47.75"N), Lynn Camp (83[degrees]38'8.78"W, 35[degrees]36'2.42"N), Meigs Creek (83[degrees]36'33.98"W, 35[degrees]38'51.92"N), Panther Creek (83[degrees]58'58.96"W, 35[degrees]33'50.52"N) and Stoney Branch (83[degrees] 50'53.04"W, 35[degrees]37'15.38"N). Also, collections at Lynnhurst Cemetery (83[degrees]55'46.843"W, 36[degrees]01'24.177"N), Knoxville, Tennessee and Biltmore Estates (82[degrees]32'53.142"W, 35[degrees]32'22.481"N), Asheville, North Carolina were made from June through December 2004. Species identified from these collection sites were included in the species listing for eastern hemlock.

Some specimens were sent to specialists for identification [L. Davis (USDA-ARS-CMAVE, Gainesville, Florida 32608); R. Gordon (Northern Plains Entomology, Willow City, South Dakota 58384); A. Mayor (Great Smoky Mountains National Park, Gatlinburg, Tennessee 37738); M. Peterson (Iowa State University, Ames, Iowa 50011); K. Vail and J. Skinner (University of Tennessee, Knoxville, Tennessee 37996)]. Other specimens were identified using standard keys, and voucher specimens were placed in the University of Tennessee Insect Museum. All identified species were systematically arranged into Cornell drawers for incorporation into the GSMNP and University of Tennessee insect museum.

Data Analysis -- A species list was developed from specimens obtained from all collection methods at all sites. Data entered into a computer database (Excel[R]) consisted of species, family, order, site, number of specimens, collection type and collection date. To determine species richness for each new or old growth site, the database was sorted by site and the species for each site were counted. Species richness data for uncommon or rarely encountered species was determined by dividing those species represented by a single specimen by the total number of species at the sites to obtain a collection ratio. All species were compared to the Tennessee Natural Heritage Program "Rare Invertebrates List" to determine their threatened/endangered status at the state level (Withers, 1997). The malaise/pan trap sampling method data were used to assess the insect fauna in relationship to their association with the host tree. Insect diversity, basic composition, and evenness were determined for insects at the sites using the Shannon index (Vandermeer, 1981). Significant differences were determined by using chi-square analysis, and output values were considered significant at P [less than or equal to] 0.05. To determine how many species may potentially be present at a given primary study site, species richness estimators (ACE, ICE, Chao 1, and Jackknife 1) were obtained from the program EstimateS (Colwell, 2000).

RESULTS AND DISCUSSION

During this study, 2,832 specimens representing 292 species in 101 families and ten orders (Table 1) were collected. Five dominant orders [Coleoptera (127 species), Diptera (79 species), Lepidoptera (27 species), Hymenoptera (23 species) and Hemiptera (19 species)] represented 94.2% of all species collected. Species richness was more varied at the Chimney Tops new growth and old growth sites with 85 and 107 species, respectively, compared to the Elkmont new growth (106 species) and old growth (103) sites. The number of species collected only at a specific site was highest (41) at the Chimney Tops old growth site and lowest (26) at the Elkmont new growth site. More site-specific species were collected at the old growth sites (73 species) than at the new growth sites (53 species). Although the lowest number of species was recorded at the Chimney Tops new growth site, the number of species (n = 27) unique to this site was similar to that at the Elkmont new growth site. As a result, 43.1% of the species recorded were obtained at one of the four primary sites. When all sites are included, 61.9% of all species collected were from a specific site. The number of specimens collected also varied among the four primary sites (ranging from 486 at the Chimney Tops new growth site to 801 at Chimney Tops old growth site). However, specimen abundance differed significantly only for Chimney Tops old growth site ([[lambda].sup.2] = 245, d.f. = 3, P = 0.05).

Significant differences (F = 103.30, d.f. = 5, 3, P < 0.05) were noted in the monthly abundance for 2002 and 2003 with the highest numbers occurring in the spring and early fall as expected. When the same months are compared across both years, significantly more insects were captured in 2002 ([[lambda].sup.2] = 63.177, d.f. = 1, [alpha] = 0.05). Specimen abundance for the months of June and July was quite variable (range 88-456) when compared among years (2002 and 2003). Some 44 species represented by 15 or more specimens were collected constituting 72.9% of the total abundance and 15.1% of the total species richness. Species (127) represented by a single individual collected during this study denoted 43.5% of the total species richness at the study sites.

No significant differences ([[lambda].sup.2] = 3.339, d.f. = 3, [alpha] = 0.05) were detected for diversity and evenness among the four primary sites (Table 2). Malaise/pan traps were the most successful single collection method used to sample the insect fauna on eastern hemlock. Malaise/pan traps accounted for 176 species or 60.3% of species richness and also had the highest Shannon diversity value at 4.14 (Table 3). Pitfall traps, considered both an ecologically sensitive and cost-effective collection method (Work et al., 2002), captured only 90 species resulting in the lowest Shannon diversity value (2.91). However, 53 species were collected only from pitfall traps. Direct collection (handpicking, beat sheet, and sweep netting) accounted for 66 species from all sites with 59 species obtained specifically from this collection method. Direct collection had the highest Shannon evenness value of any single collection method at 0.83 (Table 3). Some 40 species were collected by multiple collection methods (any combination of the trap types) that accounted for 13% of the total species richness. These data demonstrate that various collection methods are essential to comprehensively sample the diverse fauna on eastern hemlock.

Species richness estimators varied for the number of insect species collected at the four primary sites. The estimators ACE and Jack 1 resulted in the most conservative estimates at every site including the estimates of overall species richness. The ICE estimator provided the most liberal estimates at every site, except Chimney Tops new growth site. The range for Elkmont new growth site was from 175 to 225 species with 104 species observed (Sob). Elkmont old growth site estimates ranged from 175 to 245 species with 102 species observed (Sob), Chimney Tops old growth site estimates ranged from 185 to 270 species with 107 species observed (Sob), and species estimates for Chimney Tops new growth site was 145 to 230 species. The species estimates for all of the sites combined ranged from 415 (Chao 1) to 550 (Sob) species. The [r.sup.2] values for all of these estimates are strong (above 0.950) suggesting a high level of confidence in each estimate. Except for those species that specifically feed on mature trees, it can be inferred from these data that the insect community is rather stable.

Species of interest that could have a significant impact on survival of eastern hemlock and the existing habitats are the two exotic pests, hemlock woolly adelgid and elongate hemlock scale. The hemlock woolly adelgid has become a significant pest of eastern hemlock in several areas throughout the eastern United States. The hemlock woolly adelgid was discovered at eight alternate sites within the Park and continues to increase its range and damage to the host trees. The exotic elongate hemlock scale is a bisexual species with two generations annually in the southern Appalachians often co-existing with the hemlock woolly adelgid (Kosztarab, 1996; McClure, 2002). We recently discovered this species infesting eastern hemlocks in the GSMNP and in Knox County, Tennessee (Buck, 2004). Another hemipteran that is commonly found infesting eastern hemlock within the region is the hemlock scale, Abgrallaspis ithacae (Ferris), a native species that feeds on the underside of the needles of eastern hemlock. This species occurs throughout the eastern hemlock range but rarely reaches damaging levels in forests due to presence of natural enemies (Stimmel, 2000).

The carabid beetle, Sphaeroderus stenostomus Weber, the most abundant species (199 specimens) captured, feeds exclusively on snails often located on the forest floor (Downie and Arnett, 1996). Sphaeroderus stenostomus is reported to be established in several southeastern states and overwinters as both larvae and adults (Downie and Arnett, 1996). Due to these qualities, the cool moist nature of the hemlock forest floor makes an excellent hunting ground for this predator, which was represented at all four primary sites. Another carabid, Scaphinotus andrewsii Harris, is a generalist predator collected in the unique microclimates produced by eastern hemlock (Ball and Bousquet, 2001). This beetle was represented by 13 specimens at the two Chimney Tops sites. The family Agyrtidae is represented by 11 species in six genera in North America north of Mexico. However, only one species, Necrophilus pettiti Horn, is found in eastern North America (Peck, 2001). Members of this family are adapted to cool climates often near mountainous regions, cool streams, or high elevation snowfields. Although not commonly collected (Peck, 2001), 24 specimens of this species were collected from all four sites. The buprestid beetle, Dicerca tuberculata (Laporte and Gory), was collected at the Anthony Creek site and is known to feed on Pinus spp., Picea spp., Abies spp., Larix spp., Thuja spp., and Tsuga spp. (Downie and Arnett, 1996). The cerambycid beetle, Leptura subhamata Randall, is known to feed on decaying hemlock and pine (Yanega, 1996). Five specimens of this species were found at Elkmont new and old growth sites. The larvae of these beetles bore into roots and wood. In all, 123 cerambycid beetles representing 20 species were collected.

A geotrupid beetle, Geotrupes hornii Blanchard, was represented by 118 specimens from both Elkmont sites and the Chimney Tops new growth site constituting the third most abundantly collected species. This beetle species is common throughout the eastern United States and feeds on fungi (Arnett, 2000; Downie and Arnett, 1996). Another commonly collected beetle (84 specimens) was Glischrochilus sanguinolenta (Olivier). This nitidulid was collected at all four GSMNP sites. Five species of nitidulids (112 specimens) were collected during this study (Table 1). The family Nitidulidae, commonly known as sap beetles, is represented by 2,800 species in 172 genera worldwide of which 165 species and 30 genera are found in the United States. Members of this family are primarily sapro-phagous or mycetophagus except a few species that live in flowers, decaying fruit, or fungi (Habeck, 2002).

Another insect species collected that feeds on snails was a sciomyzid dipteran, Euthycera arcuata (Loew). About 200 dipteran species worldwide feed on terrestrial or freshwater snails, their eggs, and larvae (Berg and Knutson, 1978). This insect lays its eggs on the backs of snails and feeds on the snail larvae. The second most commonly occurring species of Diptera was Monoclona rufilatera Walker, a mycetophilid represented by 122 specimens collected from the Elkmont new growth site and both the mature and new growth Chimney Tops sites. Mycetophilids are commonly found in shady, damp places near fungi or decaying vegetation. Although a few species are predaceous as larvae, most feed on fungi and few are considered pests (Borror et al., 1989). Five specimens of the rarely collected fly, Dryomyza simplex Loew (Dryomyzidae), were collected from the Chimney Tops old growth site. These insects are found as larvae in decaying organic matter similar to that found in moist forest situations (Borror et al., 1989).

The most abundant Hymenoptera collected was the ant, Aphaenogaster fulva Emery. All 102 specimens of A. fulva collected were found at the two Elkmont sites except for one specimen at the Chimney Tops old growth site. Abundance was concentrated at these two sites because of several colonies of A. fulva located around and between the Elkmont sites. These ants are indigenous to the southern Appalachian Highlands, New England, and Nova Scotia (Creighton, 1950).

CONCLUSIONS

In assessing the insect biodiversity associated with eastern hemlock, 2,832 specimens representing 292 species were collected and identified. The number of species identified from eastern hemlock falls within the expected range for the number of species recorded on other trees (LaForest, 2000; Trieff, 2002) within the region. From these data, we conclude that a rich, stable insect fauna associated with eastern hemlocks exists within the test sites at this time. Species richness and abundance were highest at the two old growth sites, and species richness ranged from 85 to 107 species at the Chimney Tops new and old growth sites, respectively. Species estimates for all sites combined ranged from 415-550 species.

Some 101 families in ten orders were represented in this study with species of Coleoptera comprising 43.5% of all insects collected. Because of the importance of bio-diversity to the well being of forest systems and the intricate nature of insect communities, information on the status and function of eastern hemlock forest systems is imperative for the development of management decisions. Eastern hemlocks are valuable for their aesthetics, contribution to ecological stability of the species-rich forests in eastern North America and economic benefit to the region. Unfortunately, the health of these trees and the habitat structure of the forest systems are severely threatened by the invasion of exotic insect species.</p> <pre> TABLE 1. Insects associated with eastern hemlock in the Great Smoky Mountains National Park. Order

Family Genus Species Orthoptera Acrididae

Arphia sulphureus Gryllacrididae Camptonotus carolinensis Gryllidae Gryllus

assimilis Gryllidae Allonemobius

fasciatus Gryllidae Oecanthus exclamationis

Rhaphidophoridae Ceuthophilus brevipes

Rhaphidophoridae Ceuthophilus maculatus Blattodea Blattellidae

Ischnoptera deropeltiformis Blattidae

Periplaneta americana Hemiptera Acanthosomatidae Elasmucha

lateralis Adelgidae Adelges tsugae (c) Cicadellidae Gyponana sp.

Cicadellidae Osbornellus limosus Cicadellidae Scaphoideus sp. Cicadidae

Tibicen canicularis Coreidae Acnthocephala

terminalis Coreidae Leptoglossus oppositus

Diaspididae Abgrallaspis ithacae

Diaspididae Chionaspis pinifolia Diaspididae

Fiorinia externa Lygaeidae Kleidocerys

resedae Membracidae Glossonotus sp.

Membracidae Platycotis vittatus

Pentatomidae Banasa calva Pentatomidae

Mormidea lugens Scutelleridae Tetyra

bipunctata Thyreocoridae Corimelaena

pulicaria Tingidae Corythuca pruni Psocoptera Caeciliidae sp. 1 Ectopscocidae sp. 2 Lachesillidae

sp. 3 Peripsocidae

sp. 4 Coleoptera Agyrtidae Necrophilus pettiti

Alleculidae Capnochroa fuliginosa

Alleculidae Isomira quadristriata

Alleculidae Isomira sericea Bruchidae

Cryptocephalus quadruplex Buprestidae Dicerca tuberculata Cantharidae Podabrus

tomentosus Cantharidae Silis

bidentatus Cantharidae Trypherus latipennis

Carabidae Agonum melanarium

Carabidae Agonum tenuis Carabidae

Calosoma externus Carabidae Calosoma

marginalis Carabidae Carabus

sylvosus Carabidae Cyclotrachelus conviva

Carabidae Dicaelus politus

Carabidae Dicaelus teter Carabidae

Harpalus pensylvanicus Carabidae

Lebia analis Carabidae Scaphinotus

debilis Carabidae Scaphinotus andrewstii

Carabidae Scaphinotus guyotii

Carabidae Scarites subterraneus Carabidae

Sphaeroderus stenostomus Cerambycidae

Analeptura lineola Cerambycidae Anthophylax

cyaneus Cerambycidae Bellamira scalaris

Cerambycidae Brachyleptura circumdata

Cerambycidae Brachysomida bivittata Cerambycidae

Clytus ruricola Cerambycidae Cyrtophorus

verrucosum Cerambycidae Idiopidonia pedalis

Cerambycidae Leptorhabdium pictum

Cerambycidae Leptura emarginata Cerambycidae

Leptura subhamata Cerambycidae Microgoes

oculatus Cerambycidae Pidonia aurata

Cerambycidae Pidonia densicollis

Cerambycidae Pidonia ruficollis Cerambycidae Prionus imbricornis Cerambycidae

Prionus laticollis Cerambycidae Strangalepta

abbreviata Cerambycidae Typocerus velutina

Cerambycidae Urgleptes faceta

Ceratocanthidae Germarostes globosus Chrysomelidae

Altica viridana Chrysomelidae Diabrotica

undecimpunctata

howardi Cleridae Cymatodera bicolor

Cleridae Placopterus thoracicus

Coccinellidae Anatis labiculata Coccinellidae

Chilocorus stigma Coccinellidae Cycloneda

munda Coccinellidae Harmonia axyridis

Coccinellidae Hyperaspis signata

Coccinellidae Psyllobora vigintimaculata Coccinellidae Rhyzobius lophanthae Coccinellidae

Sasajiscymnus tsugae (d) Coccinellidae Scymnillus

horni Coccinellidae Scymnus loewii

Curculionidae Curculio caryae

Curculionidae Cyrtepistomis castaneus Curculionidae

Hypera punctata Curculionidae Myrmex

myrmex Curculionidae Odontopus calceatus Curculionidae Panscopus erinaceus

Elateridae Agriotes oblongicollis

Elateridae Athous brightwelli

Elateridae Athous posticus Elateridae

Athous rufifrons Elateridae Athous

scapularis Elateridae Conoderus

lividus Elateridae Ctenicera signaticollis

Elateridae Hemicrepidius memnonius

Elateridae Danosoma obtectus Elateridae

Lacon discoidea Elateridae Limonius

griseus Elateridae Melanactes piceus

Elateridae Melanotus americanus

Elateridae Melanotus decumanus Elateridae

Melanotus hyslopi Elateridae Malanotus

pertinax Erotylidae Megalodacne heros

Eucnemidae Isorhipis ruficornis

Geotrupidae Bolboceras simi Geotrupidae

Geotrupes blackburnii Geotrupidae Geotrupes hornii Geotrupidae Geotrupes

semiopacus Geotrupidae Geotrupes splendidus Lampyridae Ellychnia corrusca

Lampyridae Pyropyga decipiens

Lycidae Plateros centralis Melandryidae

Dircaea liturata Meloidae Meloe

americanus Mordellidae Mordellistena

arida Mordellidae Mordellistena limbalis

Mordellidae Mordellistena ornata

Mordellidae Tomoxia serval Nemonychidae

Cimberis pilosus Nitidulidae Cryptarcha

ampla Nitidulidae Glischrochilus fasciatus

Nitidulidae Glischrochilus quadrisignatus

Nitidulidae Glischrochilus sanguinolenta

Nitidulidae Stelidota octomaculata Pyrochroidae Dendroides concolor Scarabaeidae

Copris minutus Scarabaeidae Dichelonyx

albicollis Scarabaeidae Dichelonyx linearis Scarabaeidae Dichelonyx subvittata

Scarabaeidae Onthophagus hecate

Scarabaeidae Onthophagus striatulus Scarabaeidae

Serica atracapilla Scarabaeidae Serica

georgiana Scolytidae Dendroctonus

terebrans Scolytidae Pityogenes plagiatus

Silphidae Nicrophorus defodiens

Silphidae Nicrophorus marginatus Silphidae

Nicrophorus orbicollis Silphidae Nicrophorus pustulatus Silphidae Nicrophorus

sayi Staphylinidae Bisnius blandus

Staphylinidae Philonthus cyanipennis

Staphylinidae Tachinus fimbriatus Tenebrionidae Arthromacra aenea Tenebrionidae

Helops aereus Tenebrionidae Meracantha

contracta Tenebrionidae Tarpela micans

Tenebrionidae Tarpela undulatus Neuroptera

Chrysopidae Chrysopa sp. Coniopterygidae

Coniopteryx sp. Hemerobiidae Hemerobius

stigma Lepidoptera Arctiidae Halysidota tesselaris

Geometridae Lambdina fiscellaria

Geometridae Melanolophia canadaria Geometridae

Nematocampa limbata Geometridae Prochoerodes

transversata Geometridae Stamnodes gibbiocostata Hesperiidae Epargyreus clarus

Lymantriidae Orgyia leucostigma

Noctuidae Catocala cerogama Noctuidae

Catocala epione Noctuidae Cucullia

intermedia Noctuidae Feralia

comstocki Noctuidae Hypena baltimoralis

Noctuidae Hypena madefactalis

Noctuidae Hyppa xylinoides Noctuidae Lithophane baileyi Noctuidae

Lithophane petulca Noctuidae Orthodes

cynica Noctuidae Parallelia bistriaris

Noctuidae Pseudorthodes vecors

Noctuidae Sunira bicolorago Nymphalidae

Speyeria diana Papilionidae Papilio

glaucus Pyralidae Herpetogramma thestealis Pyralidae Pantographa limata

Thyatiridae Pseudothyatira cymatophoroides

Zygaenidae Pyromorpha dimidiata Mecoptera

Panorpidae Panorpa appalachia Diptera Acroceridae

Eulonchus marialiciae Anthomyiidae Anthomyia pluvialis Anthomyiidae Emmesomyia

socialis Anthomyiidae Hydrophoria sp.

Anthomyiidae Hylemya alcathoe

Anthomyiidae Pegomya sp. Asilidae

Efferia aestuans Bibionidae Penthetria

heteropterus Calliphoridae Calliphora

vomitoria Calliphoridae Lucilia coeruleiviridis

Calliphoridae Lucilia pallescens

Calliphoridae Pollenia rudis Ceratopogonidae

Atrichopogon sp. Ceratopogonidae Culicoides

sanguisuga Chironomidae Chasmatonotus bicolor

Chironomidae Parametriocnemus lundbeckii

Drosophilidae Amiota sp. Drosophilidae

Drosophila sp. Dryomyzidae Dryomyza

simplex Empididae Rhamphomyia sp.

Heleomyzidae Allophyla atricornis

Heleomyzidae Amoebaleria sp. Heleomyzidae

Suillia sp. Lauxaniidae Camptoprosopella sp. Lonchaeidae Lonchaea caerulea

Lonchaeidae Lonchaea sp. Micropezidae

Rainieria antennaepes Muscidae Helina

sp. Muscidae Mesembrina latreillii

Muscidae Mydaea sp. Muscidae

Phaonia sp. Muscidae Potamia

sp. Muscidae Thricops rufisquama

Mycetophilidae Boletina sp. Mycetophilidae

Brevicornu sp. Mycetophilidae Dynatosoma

fulvidum Mycetophilidae Dynatosoma placidum

Mycetophilidae Leptomorphus subcaerulea

Mycetophilidae Monoclona rufilatera Mycetophilidae Mycetophila sp. Mycetophilidae

Mycomya sp. Mycetophilidae Orfelia

sp. Mycetophilidae Phronia sp.

Mycetophilidae Saigusaia cincta Mycetophilidae

Synapha tibialis Mycetophilidae Zygomyia

ornata Periscelididae Periscelis annulata

Phoridae Dohrniphora cornuta

Phoridae Megaselia sp. Sarcophagidae

Blaesoxipha atlanis Sarcophagidae Boettcheria

cimbicis Sarcophagidae Boettcheria sp.

Sarcophagidae Fletcherimyia sp. Sarcophagidae

Sarcophaga sp. Sarcophagidae Tripanurga

sp. Sarcophagidae Udamopyga niagarana

Scathophagidae Scathophaga nigrolimbata

Sciaridae Bradysia sp. Sciaridae

Phytosciara flavipes Sciomyzidae Euthycera

arcuata Simuliidae Prosimilium mixturn

Syrphidae Eristalis sp. Syrphidae

Ferdinandea buccata Syrphidae Ferdinandea

dives Syrphidae Mallota bautias

Syrphidae Spilomyia sp. Syrphidae

Syrphus rectus Syrphidae Syrphus

sp. Syrphidae Xylota sp.

Tabanidae Chrysops geminatus

Tachinidae Trigonospila pallipes Tipulidae

Austrolimnophila toxoneura Tipulidae Epiphragma fasciapennis Tipulidae Limonia

indigena Tipulidae Limnophila politissima Tipulidae Elephantomyia westwoodi

Tipulidae Metalimnobia cinctipes

Tipulidae Tipula duplex Xylophagidae

Dialysis sp. Hymenoptera Aphelinidae Encarsia

citrina Apidae Bombus bimaculatus

Apidae Bombus fervidus

Apidae Bombus impatiens Apidae

Bombus perplexius Formicidae

Acanthomyops claviger Formicidae Acanthomyops

interjectus Formicidae Camponotus pensylvanica

Formicidae Camponotus americanus

Formicidae Aphaenogaster fulva Formicidae

Prenolepis impairs Halictidae Augochloropsis

metallica Halictidae Augochlora purus

Halictidae Augochlorella striata

Halictidae Lasioglossum bruneri Ichneumonidae

sp. 1 Ichneumonidae

sp. 2 Ichneumonidae

sp. 3 Sphecidae Cerceris sp.

Tenthredinidae Tenthredo carolina

Vespidae Dolichovespula maculata Vespidae

Vespula vulgaris Vespidae Vespula sp. Order Author Site (a)

Method (b) n Orthoptera (F.) 1 PF 1 Gershacker 1,2,3 PF

18 (F.) 1 PF

2 (DeGeer) 1,2 PF

10 Davis A1 DI

1 Scudder 1,2,3,4 PF

25 Harris 1,2,3,4 MA,PF 80 Blattodea Brunner 1,2 MA,PF 3

(L.) 1,2 MA 2 Hemiptera

(Say) 3,A2,A3 PF,DI 3 (Annand) A1-A8 DI 100 DeLong

2,3 MA 2 DeLong

1 MA 1 Delong & Beery

1,2 MA 2 (Harris)

2 MA 1 (Dallas)

1 MA 1 (Say) A1 DI 1 (Ferris) A1

DI 8 (Fitch) A1,A4

DI 3 Ferris A4

DI 6 (Panzer) A1

DI 1 A2

DI 1 (F.) 1,2,3,4 MA 9 (Say) A1,A2 DI

2 (F.) A1 DI

1 (Herrich-Schaeffer) 1 MA

1 (Germar) A1 DI

1 Osborn & Drake A1 DI

1 Psocoptera A9 DI 2 A9 DI 8

A10 DI 4

A9 DI 3 Coleoptera

Horn 1,2,3,4 MA,PF 24 (Melsheimer) 1,2 PF 10 Couper

A5 DI 1 (Say)

1,2,3, A4 MA 26 Newman

2 MA 1 (LaPorte & Gory)

A2 DI 1 (Say)

A1,A2 DI 2 (Say) A5 DI 1 (Germar) A5

DI 3 (DeJean) 3

PF 1 (LeConte) 4

PF 1 (Say) 1,2,3

PF 21 Casey 1,2,3,4

PF 32 Say 1,2,3,4

PF 8 LeConte 1,2 PF 19 DeJean 1,2,3,4 PF

21 Bonelli 1,2,4 PF

26 DeGeer 3 PF

1 DeJean A1 DI

1 (LeConte) 3,4 PF

28 Harris 3,4 PF 13 (LeConte) 4 PF 1

F. 1,2,3 PF 35

Weber 1,2,3,4 PF 199

(Say) 1,3,4 MA,PF 11

(Haldeman) A6 DI 1

(Say) 3 MA 2

(Olivier) 3 MA 1 (Say) 2 PF 1 (Olivier)

1,2,3,4 MA 15 (Olivier)

4 MA 1 (LeConte)

3,4 MA 5 (Haldeman)

4 MA 1 F.

2 MA 1 Randall 1,2 MA 5 (LeConte) 2

MA 1 (Horn) 3,4

MA 36 (Casey) 3,4

MA,PF 16 (Say) 3

MA 1 (L) 1

DI 1 (Drury) A8

DI 2 (Germar) 1,2,4 MA 20 (Olivier) 2 MA

1 (Say) 2 DI

1 (Say) 2 PF

1 Schaeffer A3 DI

2 Barber 2,A2 PF,DI

2 (Say) 2,3,4 MA,PF 7 (Olivier) 4 MA 1

(Say) A3,A4 DI 4

(Say) A4 DI 8

(Say) A2 DI 1

(Palles) A1,A2 DI 17

Olivier 1 MA 1

(Say) A2,A3 DI 29 Blaisdell A9,A10 DI 34 (Sasaji & McClure) A1,A3 DI 4 Gordon

A9,A10 DI 11 Mulsant

A9,A10 DI 6 (Horn)

A3 DI 1 (Roelofs)

1,2,A3 MA,PF 11 (F.)

1,2,A5 PF,MA 4 (Herbst) 1 MA 1 (Say) 1,A2,A3,A4,A5 MA,DI 28 (Say) A8 DI

1 (Melsheimer) 1,2,4 MA,PF

10 (Kirby) 1,2,A5 PF,DI

9 (Melsheimer) 2 MA

1 (Randall) 3 MA

1 (Say) 3 MA 1 (DeGeer) 4 PF 1

(Melsheimer) 1,2,4 MA 7

(Herbst) 1,2 MA 2

(Say) 1 PF 1

(Weber) 3 PF 1

Beauvois 2 MA 1

(DeGeer) 4 MA 1 (Herbst) 1,2 MA 8 (Erichson)

3 MA 1 Zwaluwenburg

1,2 MA 14 (Say)

1,2 MA 5 (Say)

2,3,4 PF,DI 7 (Say)

1,2,3,4 MA,PF 51 (Wallis) 1,2 MA,PF 2 (F.) 1

PF 1 Blanchard 1,2,4

MA,PF 118 Jekel 1,2,3

PF 11 (F.) 1,2

PF 7 (L.) 1,A1

PF 2 (Harris) 2 MA 1 Green 1,2 PF

2 LeConte 2,3 MA,PF

2 Leach 4 PF

8 LeConte 1 MA

8 (Melsheimer) 1 MA

1 (Melsheimer) 1 MA 1 (Say) 4 MA 3

(LeConte) A2 DI 1

Erichson 1,2 MA 2

(Olivier) 1,2,3,4 MA,PF 19

(Say) 4 MA 1

(Olivier) 1,2,3,4 MA,PF 84

(Say) 1,2,4 MA,PF 6 (Newman) 3,4 MA 6 (Drury)

2 MA 2 Burmeister

1,2 MA 5 (Gyllenhal)

3 MA 1 LeConte

1,2,3,4 MA 11 (Panzer)

1 PF 1 (Beauvois)

1,2 PF 2 (Kirby) 1,2,3 MA 13 Leng 1,2,3,4

MA,PF 23 (Olivier) A1

DI 1 (LeConte) A2

DI 1 Mannerheim 2,3,4 MA,PF 29 (F.) 1 MA

1 Say 2,3,4 PF

68 Herschel 2 PF

2 Laporte 3 PF

1 (Gravenhorst) 3 PF

1 (F.) 1,2 PF 5 Gravenhorst 1,2 PF 17

Say 2,3 MA 3

Germar 1 MA 1

(Beauvois) 1,2 MA 7

(F.) 4 PF 1

(LeConte) 1 MA 1 Neuroptera

A9 DI 2

A9 DI 4 Stephens

A10 DI,MA 12 Lepidoptera (J.E. Smith)

1 MA 1 (Guenee)

3,4,A1, MA, DI 9

A2,A3,A5 (Guenee) A4 DI

4 (Haworth) 2 MA

1 (Drury) 1,3,4 MA,PF

5 (Walker) 3 MA

1 Cramer 3,4 MA 2 (Smith) A1 DI 1

(Guenee) 3 MA 7

Drury 2 MA 1

(Speyer) 4 MA 1

(Grote) A4 DI 4

(Guenee) 4 MA 1

(Guenee) 4 MA 1 (Guenee) 3 MA 3 Grote

3 MA 1 (Grote)

3 MA 1 Guenee

3 MA 1 Hubner

3 MA 1 (Guenee)

4 MA 1 (Guenee)

3 MA 1 (Cramer)

1 MA 1 L. 3 DI 1 (Walker) 2

MA 7 (Grote & Robinson) 3

MA 1 (Guenee) 4

MA,PF 11 Herrich-Schaffer A4

DI 2 Mecoptera Byers 1,3,4

MA,PF 13 Diptera Brimley 4

MA 1 (L.) 2 MA 1 (Stein) 3 MA

5 3, 4 MA

8 (Walker) 3 MA

2 2,3,4 MA

15 (L.) 1 PF

1 (Say) A1 DI 1 (L.) 2,3,4 MA 7

(Macquart) 4 MA 1

(Shannon) 3,4 MA,PF 30

(F.) 1,2 MA 3

A2 DI 1

(Coquillett) A6 DI 1

Rempel 4 MA 1 Johannsen A1 DI 1

4 MA 6

2,4 MA 6 Loew

3 MA 5

A3 DI 1 (Meigen)

1,2,3 MA,PF 3

3,4 MA 3

3,4 MA 7

4,A1 MA 2 Walker

A2 DI 1

3,4 MA 5 (Say) 1 PF 1 1,3

MA,PF 9 Robineau-Desvoidy 3,4

MA,PF 25 2,3,4

MA,PF 7 3

MA 3 2

MA 1 (Schnabl) 3,4 MA 33 1 MA

3 4 MA

1 Coquillett 3 PF

5 Johannsen 3 MA

1 (Coquillett) 2 PF

1 Walker 1,3,4 MA 122

1,2,3,4 MA 32

3 MA 2

4 MA 1

2,3 MA 54

(Johannsen) A1 DI 1

(Coquillett) 3 MA 1 Loew 1 MA 1 (Fallen)

2 MA 1 (Bigot)

1,2 MA 2

4 MA 1 Aldrich

1,2,4 MA,PF 6 (Townsend)

1 MA 1

1,2,3 MA,PF 5 2 PF 1 1,2

MA,PF 8 1

PF 1 (Parker) 3

MA 1 Cresson 1

MA 1 3

MA 1 (Meigen) 3 MA 6 (Loew) 3,4 MA,PF

4 Syme & Davies A1 DI

2 3 MA

1 (Loew) 1,2 MA

2 (Osten Sacken) 1,2 MA 2 (Walker) 1 MA 4

1,3 MA 3

Osten Sacken 3 MA 7

1,3,4 MA 4

3,4 MA 2

Wiedemann 2 MA 1

(Reinhard) 3 MA 1 (Osten Sacken) 4 MA 2 (Say)

3 MA 1 (Osten Sacken)

2 MA,PF 5 (Alexander)

A2 DI 1 Osten Sacken

4 MA 1 Say 3 MA 1 Walker 4

PF 1 1

PF 1 Hymenoptera Craw A9,A10

DI 100 Cresson 2,3,4

MA 30 (F.) 2,3,4

MA 34 Cresson 2,3,4

MA 33 Cresson 2,3,4 MA 51 (Roger) 2 PF

2 Mayr 1 MA

2 (DeGeer) 2 PF

7 Mayr 1 DI

1 Emery 1,2,3 MA,PF 102 (Say) 1 PF 45 (F.) 4 MA 1

(Say) 2,3,4 MA 15

(Provancher) 1,2,3,4 MA 14

(Crawford) 4 MA 1

3 MA 12

3 MA 1

3 MA 1

2 MA 1 (Rohwer)

1 MA 1 (L.)

1,2,4 PF 3 (L.)

1,3,4 MA,PF 86

2 PF 2 (a) Primary Sites: 1 = Elkmont new growth, 2 = Elkmont old growth, 3 = Chimney Tops old growth, and 4 = Chimney Tops new growth. Alternate Sites: A1 = Anthony Creek, A2 = Cataloochee Cove, A3 = Gregory Ridge, A4 = Laurel Falls, A5 = Lynn Camp, A6 = Meigs Creek, A7 = Panther Creek, A8 = Stoney Branch, A9 = Lynnhurst Cemetery, and A10 = Biltmore Estates. (b) Methods of sampling: DI = direct sampling (beat-sheet, handpicking, sweep-net), MA = Malaise/pan trap, PF = pitfall traps. (c) Adelges tsugae Annand (Adelgidae), a recent invasive species, was collected at sites A1-A8 by direct sampling. (d) Sasajiscymnus tsugae, a biocontrol agent released in 2002 against Adelges tsugae was observed at sites A1 and A3. </pre> <pre> TABLE 2. Shannon diversity (H') and evenness (E) values for insect fauna at new and old growth eastern hemlock sites, Great Smoky Mountains National Park, 2002-2003. Site Shannon's H' Shannon's E Elkmont New 3.94 * 0.84 Growth Elkmont Old

3.77 0.80 Growth Chimney Tops Old 3.62 0.76 Growth Chimney Tops New 3.64 0.81 Growth All Sites 4.51 0.79 * No significant differences ([[lambda].sup.2] = 3.339, d.f. = 3, [alpha] < 0.05, n = 282 species) noted among sites for species diversity or evenness. </pre> <pre> TABLE 3. Shannon diversity (H') and evenness (E) values for collection methods used to sample the insect fauna associated with eastern hemlock, Great Smoky Mountains National Park, 2002-2003. Collection Method Shannon's H' Shannon's E Malaise/pan traps 4.14 0.79 Pitfall traps

2.91 0.72 Direct sampling 3.18 0.83 Multiple traps (a) 3.31 0.85 (a) Multiple traps include samples obtained from all three collection methods at the four primary sites evaluated in the Great Smoky Mountains National Park. </pre> <p>ACKNOWLEDGMENTS

We are grateful to K. Johnson (Great Smoky Mountains National Park, Gatlinburg, Tennessee 37738), B. Hascher (Biltmore Estates, Asheville, North Carolina), R. Rhea (USDA Forest Service, Asheville, North Carolina) and C. Limebarger (Lynnhurst Cemetery, Knoxville, Tennessee) for providing the study sites. We also thank L. Davis (USDA-ARS-CMAVE, Gainesville, Florida 32608), R. Gordon (Northern Plains Entomology, Willow City, South Dakota 58384), A. Mayor (Great Smoky Mountains National Park, Gatlinburg, Tennessee 37738), M. Peterson (Iowa State University, Ames, Iowa 50011), K. Vail and J. Skinner (University of Tennessee, Knoxville, Tennessee 37996) for identifying several of the species collected. We are also grateful to the Great Smoky Mountains National Park (DLIA/ATBI) and the USDA Forest Service for their partial support of this project.

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L. BUCK, P. LAMBDIN, D. PAULSEN, J. GRANT, AND A. SAXTON

The University of Tennessee, Department of Entomology and Plant Pathology, Knoxville, TN 37996 (LB, PL, DP, JG)

The University of Tennessee, Department of Animal Science, Knoxville, TN 37996 (AS)
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