224 Florida Entomologist (95)1 March 2012

PSOCOPTERAN SPECIES ASSOCIATED WITH EASTERN HEMLOCK IN THE SOUTHERN APPALACHIANS

Carla Coots1,2, Paris Lambdin1, Jerome Grant1, Rusty Rhea3, and Edward Mockford4 1Department of Entomology and Plant Pathology, The University of Tennessee, Knoxville, TN 37996

2Corresponding author, e-mail: [email protected]

3USDA Forest Service, Forest Health Protection, 200 Weaver Boulevard, Asheville, NC 28804.

4School of Biological Sciences, Illinois State University, Normal, IL 61790-4120.

Eastern hemlock, Tsuga canadensis (L.) Car- collected using direct sampling (beat sheet, hand- rière, comprises a vital component of biological picking, and trunk vacuuming) with collected diversity (Jordan & Sharp 1967; Lapin 1994; Tin- specimens placed into labeled (date, site and tree gley et al. 2002; Buck 2004; Buck et al. 2005; Dill- location, sampling type) alcohol-filled vials and ing et al. 2007; Dilling et al. 2009) and economi- taken to the laboratory for identification. Beat- cal (Travel Industry Association 2006; Woodsen sheet samples (4 per tree) were taken at each 2001) and environmental stability (Evans 2002; direction by striking each branch 5 times with Snyder et al. 2004) within its geographical range. a one-meter rod, while visual observations were Hemlock woolly adelgid, Adelges tsugae Annand conducted on each tree for 5 min per stratum, and (Hemiptera: Adelgidae), is an exotic species direct trunk vacuuming occurred on 61 cm of the capable of rapidly reducing populations of eastern circumference of the tree’s trunk. hemlock throughout the eastern United States During this study, 3,740 adult and nymph pso- (McClure & Fergione 1977; Buck et al. 2005; copteran specimens were collected and identified, Ellison et al. 2005). As such, documenting the representing 44 species, 23 genera, and 11 fami- biodiversity of associated with this host lies (Table 1). These species represent 14.3% of tree is imperative. More than 400 insect species the species (307) and 41% of the families of Pso- have been recorded to be associated with eastern coptera documented in North America (Mockford hemlock in the southern Appalachians (Wallace 1993; Lienhard & Smithers 2002). In addition, 29 & Hain 2000; Buck et al. 2005; Lynch et al. 2006; species represent new records for Tennessee and Dilling et al. 2007; Dilling et al. 2009). 33 species represent new county records for the Psocopterans comprise a major component of state. The number of psocopteran species on east- diversity on eastern hemlock, occupy- ern hemlock comprises almost 10% of the known ing an important ecological role as major consum- insect species associated with this host tree. ers of microepiphytes as well as an integral food The greatest number of species was recorded source for predators (Thornton 1985; Mockford for followed by and Perip- 1993). Population densities of psocopterans can socidae. Species within these 3 families represent be extremely high. For example, densities of Pso- 37%, 23%, and 11%, respectively, of the total spe- coptera on the bark of larch, i.e., Larix decidua cies identified on eastern hemlock. The families Mill., have been reported at > 4,000/m2, which , , , Thornton (1985) equates to > 6,000/m2 of land , and were each rep- surface. In this study, we document the psocop- resented by only 1 species. However, the least teran richness within the canopy of eastern hem- number of specimens for a species was collected lock in the southern Appalachians. for Lachesilla andra Sommerman (n = 16) (La- Eastern hemlocks (n = 6 ) were selected on 5 chesillidae) followed by L. anna Sommerman (n Nov 2005 at Indian Boundary within the Chero- = 22), Peripsocus stagnivagus Chapman (n = 27) kee National Forest located in Monroe County (), and Lachesilla corona Chapman in southeast Tennessee (35°23.787 N, 84°06.662 (n = 29) (Lachesillidae). The greatest number of W). All trees (14-18 m in height) were located in specimens was collected for Cerastipsocus veno- a shortleaf pine–oak (type 76) forest (Eyre 1980). sus (Burmeister) (n = 246) (Psocidae) followed by Because of the diversity of habitat preferences in flavidus (Stevens) (n = 201) (Caecili- trees and mobility among species of , usidae). Species identified from eastern hemlock 3 sampling methods were selected to capture a constitute the highest number of psocopteran spe- wide range of psocopteran species and were ex- cies recorded from a single tree species. ecuted monthly from Nov 2005 through Sep 2008. The dominant distribution pattern of the spe- Also, a platform lift was used to sample the bot- cies identified is the eastern deciduous forest tom, middle and top stratum on each tree in Aug pattern as defined and described by Mockford 2006, Sep 2007, and Jun 2008. Specimens were (1993) (i.e., corresponding with delimited east- Scientific Notes 225 . habitats

and , Habitat*** distribution , Foliage and branches of conifers, larch, spruce, also oak and su - pine, juniper, mac (1), hemlock (2) Hemlock, spruce, larch, and other co - nifers (1) Shaded tree trunks, outcrops, and stone and cement structures (1) Variety of broad-leaf and needle leaf Variety trees Bark of tree trunks and branches, Bark of tree trunks and branches, stone outcrops (1) Dead leaves (1) Conifers, dead grasses, drying grain drying grasses, dead Conifers, crops (1) Dead vegetation and grasses (1) Dead leaves of broad-leaf plants and oc - casionally conifers (1) Foliage of junipers and occasionally junipers of Foliage pine (1) Dead leaves on a variety of woody and woody of variety a on leaves Dead herbaceous plants (1) Foliage of pines, junipers, and conifers (1) Dead leaves and shrubs trees (1) Dead standing grass stems and leaves (1) Hanging dead leaves, occasionally foli - age of conifers (1) Dead leaves (1) Hanging dead leaves (1) Foliage of broad-leaf and conifer trees, ground litter (1) Broad-leaf plants (1), hemlock (2) Evergreen trees and shrubs, live oak, and wax myrtle (1) American holy, Foliage of pines and juniper, noted on Foliage of pines and juniper, spruce and Douglas fir (1) Broad-leaf trees, rarely conifers (1), hemlock (2) methods

No No No No Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes No No No No Yes No Tennessee Tennessee New Record c olle tion

NA COS NA NA NA EUS NA NA EUS NA NA NA NA NA NA COS COS NA EUS & WUS NA NA COS Distribution** c orres p ondin g

and

6 49 38 94 76 46 67 74 48 36 29 55 22 66 35 67 87 59 No. 110 122 123 201 Specimens ala c hians pp A

H, T H, T H, T B,H,T T B, H B,T,H B, T B, T B, H,T B B, H B, T B B, H, T B, H B, H B, T B, H B, H B, H B, H, T Method* Collecting southern

the

in

Author Mockford (Dalman) (Hagen) (Müller) (Packard) (Walsh) (L.) (Aaron) Chapman Chapman Chapman Chapman Sommerman Sommerman (Walsh) Ribaga (Enderlein) (Walsh) (Mockford) (Chapman) (Banks) (Stevens) hemlo c k

eastern

Species with

maculosus alboguttatus lugens unipunctatus hageni rufa pedicularia nubilis major forcepeta corona contraforcepeta anna andra amabilis meridio nalis cryptomeriae conterminus sommermanae quillayute pinicola flavidus asso c iated

Genus Peripsocus Peripsocus Lichenomima Mesopsocus Echmepteryx Lachesilla Lachesilla Lachesilla Lachesilla Lachesilla Lachesilla Lachesilla Lachesilla Lachesilla Anomopsocus Ectopsocus Ectopsocopsis Teliapsocus Xanthocaecilius Valenzuela Valenzuela so c o p tera P of

p e c ies 1. S able *B, Beatsheet; H, Handpick; and T, Trunk Vacuum Trunk *B, Beatsheet; H, Handpick; and T, United States; and CUS, Central States America; COS, Cosmopolitan; EUS, Eastern United States; WUS, Western **NA, North ***Habitats recorded by (1) Mockford (1993) and (2) Dilling et al. (2009) Peripsocidae Peripsocidae Myopsocidae Mespsocidae Lepidopsocidae Lachesillidae Lachesillidae Lachesillidae Lacheillidae Lachesillidae Lacheillidae Lachesillidae Lachesillidae Lachesillidae Lachesillidae Ectopsocidae Dasydemellidae Caeciliusidae Caeciliusidae Caeciliusidae Caeciliusidae T Family 226 Florida Entomologist (95)1 March 2012 . habitats

and , , Habitat*** distribution , , methods

Unknown Branches of broad-leaf and conifer trees (1) Branches of broad-leaf trees, hemlocks, trees, broad-leaf of Branches and bald cypress (1) Trunks and branches of broad-leaf Trunks conifer trees (1) Trunks and branches of broad-leaf Trunks conifer trees (1) Bark o f trees and shaded stone outcrops (1) Branches of broad-leaf and conifer trees and shrubs (1) Trunks and branches of broad-leaf Trunks trees and pines, occasionally stone out - crops (1) Trunks and branches of broad-leaf Trunks trees and conifers (1) Branches of broad-leaf trees, shrubs, and conifers (1), hemlock (2) Trunks and branches of broad-leaf Trunks trees and conifers (1) Dead branches o f broad-leaf tress (1) Trunks and branches of broad-leaf Trunks trees and pines (1) Trunks and branches of broad-leaf Trunks conifer trees (1) Trunks and branches of broad-leaf Trunks conifer trees (1) Branches o f broad-leaf and conifer trees (1) Branches of broad-leaf and conifer trees (1), hemlock (2) Bottomland forest trees (1), hemlock(2) Trees and shrubs, stone outcrops (1), stone outcrops and shrubs, Trees hemlock (2) Needle-like and broad-leaf trees (1), hemlock (2) Trunks and branches of trees, spanish Trunks moss , and occasionally stone outcrops (1) Hemlock, juniper, spruce, pine, and Hemlock, juniper, broad-leaf trees (1) tion c olle c

Yes Yes Yes Yes Yes Yes Yes No Yes Yes No Yes Yes Yes Yes Yes Yes No No No No Yes Yes g Tennessee Tennessee New Record ondin p orres c

and

EUS & WUS NA NA EUS & CUS COS CUS EUS EUS NA NA EUS EUS & CUS NA NA NA NA NA EUS EUS NA EUS NA Distribution** hians c ala 5 pp 43 66 57 41 62 59 78 63 37 27 55 No. A 114

113 139 169 186 165 246 147 154 148 Specimens southern

the

in B, H, T B, H, T B, H, T B,T B, T B, H, T B, H, T B, H, T B, T H, T B, H, T B, H B, H, T B, H, T B, H, T H, T B, H, T B, H, T H, T H, T B, T H, T

Method* Collecting k c hemlo

Author eastern

(L.) (Banks) (Chapman) Betz Betz Sommerman (Walsh) Aaron (Walsh) (Walker) (Hagen) (Banks) (Banks) (Burmeister) (Chapman) (Hagen) (Banks) (Aaron) (Danks) (Rambur) Chapman (Hagen) with

iated c Species asso

cruciatus slossonae quaesitum merum castum alexanderae polluta leidyi purus novaescotiae moesta infumatus floridanus venosus lithinus quieta opposita maculosa badonneli subfasciatus stagnivagus madidus tera p o c so P of

Genus ies c e p S Graphopsocus Trichadenotecnum Trichadenotecnum Trichadenotecnum Trichadenotecnum Trichadenotecnum Ptycta Psocus Metylophorus Metylophorus Loensia Indiopsocus Hyalopsocus Cerastipsocus Blastopsocus Blaste Blaste Aaroniella Aaroniella Peripsocus Peripsocus Peripsocus ) ) ontinued 1. (C 1. able *B, Beatsheet; H, Handpick; and T, Trunk Vacuum Trunk *B, Beatsheet; H, Handpick; and T, United States; and CUS, Central States America; COS, Cosmopolitan; EUS, Eastern United States; WUS, Western **NA, North ***Habitats recorded by (1) Mockford (1993) and (2) Dilling et al. (2009) Stenopsocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Psocidae Philotarsidae Peripsocidae Peripsocidae Peripsocidae T Family Scientific Notes 227

ern deciduous forest (Braun 1972) in the eastern Dilling, C., Lambdin, P., Grant, J., and Buck, L. 2007. United States). This eastern deciduous forest Insect guild structure associated with eastern hem- pattern is noted as having poorly documented lock in the southern Appalachians. Environ. Ento- northern delineation due to limited collecting mol. 36: 1408-1414. in this area, but generally is found from central Dilling, C., Lambdin, P., Grant, J., and Rhea, R. 2009. Community response of insects associated with east- New Brunswick to central Ontario and south- ern hemlock to imidacloprid and horticultural oil ward to peninsular Florida. Lachesilla pedicu- treatments. Environ. Entomol. 38: 53-66. laria (L.) and Teliapsocus conterminus (Walsh) Ellison, A. M., Bank, M. S., and Clinton, B. D. 2005. were noted as having other distribution patterns Loss of foundation species: consequences for the (Mockford 1993). In the eastern United States, T. structure and dynamics of forested ecosystems. conterminus (Walsh) generally follows the east- Front. Ecol. 3: 479 -486. ern deciduous forest pattern but has a southern Evans, R. A. 2002. An ecosystem unraveling? In Proc. limit that extends farther into Florida (Mockford Hemlock Woolly Adelgid in Eastern United States 1993). is noted as having Symp. 5-7 Feb 2002. East Brunswick, New Jersey. Lachesilla pedicularia 6 pp. a large outdoor range, and as such does not fit Eyre, F. 1980. Forest cover types of the United States into a specific distribution pattern. Distribution and Canada. Soc. Am. Foresters. Washington, DC. of male Ptycta polluta (Walsh) was documented Jordan, J. S., and Sharp, W. M. 1967. Seeding and plant- to be the southern Appalachians in Tennessee ing hemlock for ruffed grouse cover. Res. Paper NE- (Mockford 1993) with bisexual populations of P. 83. USDA For. Serv., Upper Darby, Pennsylvania, 7 polluta and Peripsocus subfasciatus (Rambur) pp. appearing to be restricted to hemlock stands. Lapin, B. 1994. The impact of hemlock woolly adelgid on Graphopsocus cruciatus (L.) has been classified resource in the Lower Connecticut River Valley. Re- as an introduced species (Mockford 1993), and port for the Northeastern Center for Forest Health Research. USDA For. Serv., Hamden, CT, 10 pp. has a varied distribution, including Tennessee, Lienhard, C., and Smithers, C. 2002. Psocoptera (Insec- in the eastern and western United States, where ta) World Catalogue and Bibliography. Instrumenta it has been documented most often from areas Biodiversitatis V. Museum d’histoire naturelle, Ge- around coastal waterways and inlets. The major- nève, 745 pp. ity of the species identified are noted as having a Lynch, C., Lambdin, P., Grant, J., Reardon, R., and Rhea, variety of habitat preferences and broad distri- R. 2006. Survey for established predators of elongate bution range (Table 1). hemlock scale in Tennessee. Florida. Entomol. 89: 527-528. Mcclure, M. S., and Fergione, M. B. 1977. Fiorinia Summary externa and Tsugaspidiotus tsugae (Homoptera: Di- aspididae); distribution, abundance, and new hosts Psocopterans collected (3,740) represented 44 of two destructive scale insects of eastern hemlock in species, 23 genera, and 11 families. Most species Connecticut. Environ. Entomol. 6: 807-811. of Psocoptera found in association with eastern Mockford, E. L. 1993. North American Psocoptera hemlock in the southern Appalachians exist on a (Insecta). Flora and Fauna Handbook 10. Sandhill variety of vegetative hosts, and as such their popu- Crane Press. Gainesville, Florida, 455 pp. lation numbers should be minimally impacted by Snyder, C., Young, J., Smith, D., Lemarie, D., Ross, any decline in eastern hemlock populations. How- R., and Bennett, R. 2004. Stream ecology linked to eastern hemlock decline in Delaware Water Gap ever, loss of eastern hemlock populations could be National Recreation Area. U. S. Geological Survey. detrimental to a few species (i.e., P. polluta and P. Kearneysville, WV (www.lsc.usgs.gov/aeb/2048-03/ subfasciatus) that are dependent upon this host for dewa.asp). survival. Of those species identified, 29 represent Thornton, I. W. B. 1985. The geographical and ecologi- new records for Tennessee and 33 species repre- cal distribution of arboreal Psocoptera. Annu. Rev. sent new county records for the state. Entomol. 30: 175-196. Tingley, M. W., Orwig, D., Field, R., and Motzkin, G. 2002. Avian response to removal of a forest domi- References Cited nant: consequences of hemlock woolly adelgid infes- tations. J. Biogeogr. 29: 1505-1516. Braun, E. L. 1972. Deciduous forests of eastern North Travel Industry Association. 2006. The economic im- America. Hafner Co., New York. XIV, 596 pp. pact of travel on Tennessee counties. Res. Dept. Buck, S. L. 2004. Insect fauna associated with eastern Travel Ind. Association Amer. Washington, DC. 82 hemlock, Tsuga canadensis (L.), in the Great Smoky pp. Mountains National Park. MS Thesis, The Univer- Wallace, M. S. and Hain, F. P. 2000. Field survey and sity of Tennessee, Knoxville, TN. evaluation of native and established predators of Buck, S. L., Lambdin, P., Paulsen, D., Grant, J., and Sax- the hemlock woolly adelgid (Homoptera: Adelgidae) ton, A. 2005. Checklist of insect species associated in the southeastern United States. J. Environ. Ento- with eastern hemlock in the Great Smoky Moun- mol. 29: 638-644. tains National Park and environs. Tenn. Acad. Sci. Woodsen, M. M. 2001. Forest invaders-insects that in- 80: 1-10. vade hemlock trees. Am. Forests 4: 12-13.