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Hemlock woolly adelgid (Adelges tsugae), or HWA, is member of the Sternorrhyncha suborder of the Order Hemiptera and native to East Asia. It feeds by sucking sap from hemlock and spruce trees (Tsuga spp.; Picea spp.). In eastern North America, it is a destructive pest that gravely threatens the eastern hemlock (Tsuga canadensis) and the Carolina hemlock (Tsuga caroliniana). Though the range of eastern hemlock extends north of the current range of the adelgid, it could spread to infect these northern areas as well. Accidentally introduced to North America from Japan, HWA was first found in the eastern United States near Richmond, Virginia, in the early 1950s. The pest has now been established in eighteen eastern states from Georgia[1] to Massachusetts, causing widespread mortality of hemlock trees. As of 2015, 90% of the geographic range of eastern hemlock in North America has been impacted by HWA.[2]

Hemlock woolly adelgid
Adelges tsugae 3225077.jpg
Evidence of hemlock woolly adelgid on hemlock
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Superfamily: Phylloxeroidea
Family: Adelgidae
Genus: Adelges
Species: A. tsugae
Binomial name
Adelges tsugae
(Annand, 1928)

Contents

CharacteristicsEdit

An adult individual averages a body length of 0.8 mm and is oval in shape.[3] The tiny brown-colored insect has four thread-like stylets that are bundled together and function as a mouthpart. Three times the length of its body, the stylet bundle pierces the host plant's parenchymatic ray tissue to derive nutrition from stored reserves.[4] It may also inject a toxin while feeding.[5] The resulting desiccation causes the tree to lose needles and not produce new growth. Hemlocks stricken by HWA frequently become grayish-green rather than a healthy dark green. In the northern portion of the hemlock's range, death typically occurs four to ten years after infestation. Trees that survive the direct effects of the infection are usually weakened and may die from secondary causes.[6]

The presence of HWA can be identified by its egg sacs, which resemble small tufts of cotton clinging to the underside of hemlock branches. In North America, the hemlock woolly adelgid asexually reproduces and can have two generations per year. Both generations are parthenogenetic and exclusively female.[3] In its native Asian habitat is a third winged generation called Sexupera; although it was previously stated that this generation's sexual reproduction requires a species of spruce that is not found in the Eastern United States and therefore dies, areas of West Virginia as of 2017 are heavily infested with this form.[7] Between 100 and 300 eggs are laid by each individual in the woolly egg sacs beneath the branches. Larvae emerge in spring and can spread on their own or with the assistance of wind, birds and/or mammals. In the nymph stage, the adelgid is immobile and settles on a single tree.[8][9]

Control methodsEdit

Forest levelEdit

The current leading biological control method of hemlock woolly adelgid is Pseudoscymnus tsugae.[10] P. tsugae is a black lady beetle that is relatively host specific, feeding only on three known aldegid species, including HWA. This beetle was discovered in 1992 while feeding on hemlock woolly adelgid in its natural range of Japan. Since 1995, the DCNR's Bureau of Forestry has released hundreds of thousands of adult beetles into affected hemlock forests of the eastern United States to determine its effectiveness at controlling the unchecked spread of the adelgid.[11] From 1995 to 1997, experiments in Connecticut and Virginia found that releasing adult P. tsugae beetles into infested hemlock stands resulted in a 47 to 88% reduction in adelgid densities within 5 months of introduction.[11] The beetle's life cycle is in parallel to the life cycle of the hemlock woolly adelgid. Both lay eggs in the spring and hatching occurs nearly simultaneously. When hatched, P. tsugae larvae are highly mobile and feed on hemlock woolly adelgid eggs and larvae. Each P. tsugae larva can effectively consume approximately 500 adelgid eggs or nearly 100 developing adelgid nymphs.[11]

Laricobius nigrinus is another predatorial beetle used as a biological control in response to hemlock woolly adelgid. Native to the western United States and Canada, L. nigrinus is known to prey exclusively on various woolly adelgids.[10] L. nigrinus adults will lay their eggs on top of wintering adelgid larvae in early spring, and upon hatching the larval beetle feeds on hemlock woolly adlegid.[10]

Also under study is Laricobius osakensis from Japan, a relative of L. nigrinus. They have shown promise in field trials.[12]

Individual treesEdit

The environmentally safest chemical control methods for treating individual trees are non-toxic insecticidal soap and horticultural oil. These are sprayed on the foliage and smother the insects as they dry. Most trees will need to be treated on a yearly basis.[13]

Toxic systemic insecticides may be applied to the foliage and bark of a tree and will persist in killing the adelgid for up to two to four years after application. Caution must be used,[13] and restraint exercised around bodies of water.

Soil drenches/soil injections/bark sprays are used in larger trees that cannot be completely sprayed with insecticidal soaps or foliage insecticides. Tree roots absorb and transport the product into the foliage and kill hemlock woolly adelgid. Soil drenches must be applied when there is adequate soil moisture for the tree roots to absorb the product. These products should not be used in areas that are in close proximity to bodies of water.[13]

Trunk injections are used for large trees that are near water or where soils are too rocky for soil injections or drenches. The chemical is injected directly into the tree and transported to the twigs and needles where the hemlock woolly adelgids are feeding. Adequate soil moisture is also necessary for the tree to take up these products.[13]

SignificanceEdit

Hemlock is a vital component of the New England forest system, and is the third most prevalent tree in Vermont. It provides protection from erosion along stream banks, food for deer and wildlife, and shelter for deer in winter. The tree is also valued both as an ornamental and as an important source of lumber. Unlike the balsam woolly adelgid that attacked only mature balsam fir, HWA infests hemlocks of all ages. Where hemlock occurs in pure stands in that region, the most commonly observed tree species to succeed it is black (sweet) birch.[14] Whereas in the southern extreme of its range, hemlock typically occurs not in pure stands but in linear riparian areas and other moist sites. Succession in these areas is affected by the presence of Rhododendron maximum which often coexists with hemlock, and because of a combination of influences restricts regeneration to shade and otherwise understory-tolerant plant species. Major changes in ecosystem structure and function, including hydrologic processes, are expected with the loss of hemlock.

Loss of the eastern and Carolina hemlock from hemlock woolly adlegid infestation will likely result in many ecological shifts in eastern North America. The understory of hemlock forests is characterized as dark, damp, and cool and is an ideal habitat to various other organisms.[4] The moist environment is preferred by many native amphibian species, particularly newts and salamanders.[15] Some species of birds have close association with the hemlock, especially during mating and nesting periods.[16] Aquatic systems adjunct to hemlock stands are also impacted by the trees' decline. Brook trout is a native fish species to the eastern United States and is known to prefer the cool, shaded streams of hemlock forests during spawning events.[17] Vulnerable animal populations are expected to diminish as a result of loss of hemlock habitat to the invasive hemlock woolly adelgid.

One hopeful factor is that the adelgid does not seem able to survive prolonged or bitter cold.[18] Following the winter of 1999-2000, a considerable dieback of adelgid and subsequent regrowth of infested trees was observed across Connecticut. The same phenomenon was repeated after the prolonged winter of 2013-2014, in time to save numerous nearly-succumbed forests.

A 2009 study conducted by scientists with the U.S. Forest Service Southern Research Station suggests the hemlock woolly adelgid is killing hemlock trees faster than expected in the southern Appalachians, and rapidly altering the carbon cycle of these forests. According to Science Daily, the pest could kill most of the region's hemlock trees within the next decade. According to the study, researchers found "hemlock woolly adelgid infestation is rapidly impacting the carbon cycle in [hemlock] tree stands", and "adelgid-infested hemlock trees in the South are declining much faster than the reported 9-year decline of some infested hemlock trees in the Northeast."[19] In fact, as of 2007 the rate of HWA expansion was recorded as 15.6 km/year south of Pennsylvania and 8.13 km/year (or less) in the northern section of the HWA’s range.[20]

ReferencesEdit

  1. ^ "Helping Hemlocks". Yahoo.com. 
  2. ^ Kok, Loke T.; Salom, Scott M.; et al. "Biological Control of the Hemlock Woolly Adelgid". Virginia Tech College of Agriculture and Life Sciences, Department of Entomology. 
  3. ^ a b Danoff-Burg, Dr. James A. "Invasion Biology Introduced Species Summary Project - Columbia University". www.columbia.edu. Retrieved 2017-04-13. 
  4. ^ a b Vose, J.M.; et al. (2013). "Hemlock woolly adelgid in the southern Appalachians: Control strategies, ecological impacts, and potential management responses" (PDF). Forest Ecology and Management. 291: 209–219. 
  5. ^ McClure M.S. and Cheah, C.A. (2002) "Important Mortality Factors in the Life Cycle of Hemlock Woolly Adelgid, Adelges tsugae Annand (Homoptera: Adelgidae) in the Northeastern United States."
  6. ^ McClure, Mark S. "Hemlock Wooly Adelgid Greenshare Factsheet". University of Rhode Island, University of Maryland Cooperative Extension. Archived from the original on 8 January 2009. 
  7. ^ Personal observation by Harold Kornylak, Virginia Beach, VA on 12 July 2017
  8. ^ "Hemlock Wooly Adelgid". Pennsylvania Department of Conservation and Natural Resources. Retrieved 2 January 2009. 
  9. ^ "Other Exotic Forest Threats - Hemlock Woolly Adelgid". Wisconsin Department of Natural Resources. Archived from the original on 30 May 2009. 
  10. ^ a b c "Forest Health Fact Sheet" (PDF). Pennsylvania Department of Conservation and Natural Resources. Retrieved 2014-02-16. 
  11. ^ a b c Shelton, Anthony. "A Guide to Natural Enemies in North America". Ph.D. Professor of Entomology, Cornell University. Cornell University. Archived from the original on 9 September 2013. Retrieved 16 February 2014. 
  12. ^ Havill, N.P; et al. (2011). Implementation and Status of Biological Control of the Hemlock Woolly Adelgid (PDF). pp. Chapter 21. 
  13. ^ a b c d Sidebottom, Ph.D., Jill. "Recommendations for Hemlock Woolly Adelgid Control in the Landscape". North Carolina Cooperative Extension Service. Retrieved 2014-02-16. 
  14. ^ Orwig, David A.; Foster, David R.; Mausel, David L. (2002-10-01). "Landscape patterns of hemlock decline in New England due to the introduced hemlock woolly adelgid". Journal of Biogeography. 29 (10-11): 1475–1487. ISSN 1365-2699. doi:10.1046/j.1365-2699.2002.00765.x. 
  15. ^ Siddig, A.A.H.; et al. (2016). "Assessing the impacts of the decline of Tsuga canadensis stands on two amphibian species in a New England forest". Ecosphere. 
  16. ^ Tingley, M.W.; et al. (2002). "Avian response to removal of a forest dominant: consequences of hemlock woolly adelgid infestations" (PDF). Journal of Biogeography. 29: 1505–1516. 
  17. ^ A., Siderhurst, Leigh (2010-01-01). Changes in Light Levels with Loss of Eastern Hemlock (Tsuga canadensis) at a Southern Appalachian Headwater Stream: Implications for Brook Trout (Salvelinus fontinalis) (Thesis). James Madison University. 
  18. ^ Talbot Trotter, R.; Shields, Kathleen S. (2009-05-29). "Variation in Winter Survival of the Invasive Hemlock Woolly Adelgid (Hemiptera: Adelgidae) Across the Eastern United States". Environmental Entomology. 38 (3): 577–587. ISSN 0046-225X. doi:10.1603/022.038.0309. 
  19. ^ "Science Daily: Hemlock Trees Dying Rapidly, Affecting Forest Carbon Cycle". University of Toronto. 
  20. ^ Rentch, J.; Fajvan, M.A.; Evans, R.A.; Onken, B. (2008). "Using dendrochronology to model hemlock woolly adelgid effects on eastern hemlock growth and vulnerability" (PDF). Biological Invasions. 11 (3): 551–563. doi:10.1007/s10530-008-9270-x. Archived from the original (PDF) on 6 March 2012. 

External linksEdit