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A geographically variable model of hemlock woolly adelgid spread

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Abstract

The hemlock woolly adelgid (HWA) (Adelges tsugae) has spread through the range of eastern hemlock (Tsuga canadensis) leaving dead hemlock trees in its wake. We combined county and previously unavailable township level infestation records with geographic variables including plant hardiness zone, elevation, forest cover type, urbanization, precipitation, temperature, and hemlock range in a geographic information system to build models linking HWA spread to the structure and characteristics of the environment. We explored the connections between site variables and spread rate using quantile regression, Tobit regression, a parametric model for heterogeneous error variance, and classification and regression tree modeling. The results show that since 1990 HWA has spread at a rate closer to 12.5 km/yr rather than the 20–30 km/yr previously estimated, however spread rates vary significantly with environmental variables. County-based data show a faster rate of range expansion in the south, 15.6 km/yr, and very slow spread in the northern part of the current range of HWA. The rate of spread based on township records for Pennsylvania and north is 8.13 km/yr. Our study suggests that HWA spreads more slowly in colder areas, with a mean minimum temperature of  − 26.1°C (plant hardiness zone 5B) delineating areas of slower range expansion. HWA also spreads more slowly during cold years, those with lower than average mean January temperatures. Our analysis demonstrates the value of quantile regression, Tobit regression, and classification and regression tree models for understanding the spread of invasive, exotic insects.

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References

  • Amemiya T (1984) Tobit models: a survey. J Econom 24:3–61

    Article  Google Scholar 

  • Beatty SW (1984) Influence of microtopography and canopy species on spatial patterns of forest understory plants. Ecology 65:1406–1419

    Article  Google Scholar 

  • Beers TW, Dress PE, Wensel LC (1966) Aspect transformation in site productivity research. J For 64:691–692

    Google Scholar 

  • Breiman L, Friedman JH, Olshen RA, Stone CJ (1984) Classification and regression trees. Chapman and Hall/CRC, Boca Raton, FL

    Google Scholar 

  • Cade B, Noon B (2003) A gentle introduction to quantile regression for ecologists. Front Ecol Environ 1:412–420

    Article  Google Scholar 

  • Cade BS, Terrell JW, Schroeder RL (1999) Estimating effects of limiting factors with regression quantiles. Ecology 80:311–323

    Article  Google Scholar 

  • Cheah CAS-J, McClure MS (2002) Pseudoscymnus tsugae in Connecticut forests: the first five years. In: Onken B, Reardon R, Lashomb J (eds) Hemlock woolly adelgid symposium. USDA Forest Service and Rutgers University, East Brunswick, NJ, pp 150–165

    Google Scholar 

  • Daly C, Taylor G (2000) United States average annual precipitation, 1961–90. Spatial Climate Analysis Service at Oregon State University (SCAS/OSU), Corvallis, Oregon

    Google Scholar 

  • Daly C, Gibson W, Taylor G (2002) Average daily minimum temperature: annual 1971–2000. 103-year high-resolution temperature climate data set for the conterminous United States. Spatial Climate Analysis Service, Oregon, Corvallis, OR

    Google Scholar 

  • DeGraaf RM, Yamasaki M, Leak WB, Lanier JW (1992) New England wildlife: management forested habitats. General Technical Report NE-144, USDA Forest Service, Northeastern Forest Experiment Station, Radnor, PA, p 271

  • ESRI (2004) ArcGIS. version 9.1. Environmental Systems Research Institute (ESRI), Redlands CA

    Google Scholar 

  • Evans A (2005) A randomized branch sampling method for hemlock woolly adelgid. In: Onken B, Reardon R (eds) Proceedings of the 3rd symposium on hemlock woolly adelgid. USDA Forest Service, Ashville, NC, USA, pp␣63–72

    Google Scholar 

  • Gannoun A, Saracco J, Yuan A, Bonney G (2005) Non-parametric quantile regression with censored data. Scand J Stat 32:527–550

    Article  Google Scholar 

  • Godman RM, Lancaster K (1990) Eastern hemlock. In: Burns RM, Honkala BH (eds) Silvics of North America. Agriculture Handbook 654, USDA, Forest Service, Washington, DC, p 877

    Google Scholar 

  • Gregoire TG, Dyer ME (1989) Model fitting under patterned heterogeneity of variance. For Sci 35:105–125

    Google Scholar 

  • Grosholz ED (1996) Contrasting rates of spread for introduced species in terrestrial and marine systems. Ecology 77:1680–1686

    Article  Google Scholar 

  • Havill N (2005) Using mitochondrial DNA to determine the native range of the hemlock woolly adelgid. In: Onken B, Reardon R (eds) Proceedings of the 3rd symposium on hemlock woolly adelgid. USDA Forest Service, Ashville, NC, USA, p 211

    Google Scholar 

  • Heger T, Trepl L (2003) Predicting biological invasions. Biol Invasions 5:313–321

    Article  Google Scholar 

  • Kelty MJ (1989) Productivity of New England hemlock/hardwood stands as affected by species composition and canopy structure. For Ecol Manage 28:237–257

    Article  Google Scholar 

  • Kittredge DB, Ashton PMS (1995) Impacts of deer browse on regeneration in mixed-species stands in southern New England. North J Appl For 12:141–145

    Google Scholar 

  • Kizlinski ML, Orwig DA, Cobb RC, Foster DR (2002) Direct and indirect ecosystem consequences of an invasive pest on forests dominated by eastern hemlock. J Biogeogr 29:1489–1503

    Article  Google Scholar 

  • Koenker R, Hallock KF (2001) Quantile regression. J Econ Perspect 15:143–156

    Article  Google Scholar 

  • Liebhold A, Luzader E, Halverson J, Elmes G (1992) The spatial dynamics of invasions by exotic forest pests. General Technical Report NE-175, USDA Forest Service, Northeastern Forest Experiment Station, Radnor, PA, pp 125–132

  • Liebhold AM, MacDonald WL, Bergdahl D, Mastro VC (1995) Invasion by exotic forest pests: a threat to forest ecosystems. For Sci Monogr 30:1–38

    Google Scholar 

  • Little EL (1999) Digital representation of the atlas of United States trees. Digital Edition. US Geological Survey, Reston, VA

  • Mack RN, Simberloff D, Lonsdale WM, Evans H, Clout M, Bazzaz FA (2000) Biotic invasions: causes, epidemiology, global consequences, and control. Ecol Appl 10:689–710

    Article  Google Scholar 

  • McClure MS (1990) Role of winds, birds, deer, and humans in the dispersal of hemlock woolly adelgid (Homoptera: Adelgidae). Environ Entomol 19:36–43

    Google Scholar 

  • McClure MS (1991) Density-dependent feedback and population cycles in Adelges tsugae (Homoptera: Adelgidae) on Tsuga canadensis. Environ Entomol 20:258–264

    Google Scholar 

  • Monserud RA, Leemans R (1992) Comparing global vegetation maps with the Kappa statistic. Ecol Model 62:275–293

    Article  Google Scholar 

  • National Arboretum (1990) USDA plant hardiness zone map. No. 1475. USDA National Arboretum, Washington, DC

    Google Scholar 

  • National Climate Data Center (2005) Area-weighted state, regional, and national monthly and seasonal temperature and precipitation. Asheville, NC

  • National Imagery and Mapping Agency (1996) Digital terrain elevation data level 0. National Imagery and Mapping Agency, Fairfax, Virginia

  • Onken B, Reardon R (2005) Proceedings of the 3rd symposium on hemlock woolly adelgid. FHTET-2005-01, USDA Forest Service, Ashville, NC, p 363

  • Orwig DA, Foster DR, Mausel DL (2002) Landscape patterns of hemlock decline in New England due to the introduced hemlock woolly adelgid. J Biogeogr 29:1475–1488

    Article  Google Scholar 

  • Parker BI, Skinner M, Gouli S, Ashikaga T, Teillon HB (1998) Survival of hemlock woolly adelgid (Homoptera: Adelgidae) at low temperatures. For Sci 44:414–420

    Google Scholar 

  • Pimentel D, Zuniga R, Monison D (2005) Update on the environmental and economic costs associated with alien-invasive species in the United States. Ecol Econ 52:273–288

    Article  Google Scholar 

  • Reardon R, Onken B (2004) Biological control of hemlock woolly adelgid in the eastern United States. FHTET-2004-04, USDA, Forest Service, Forest Health Technology Enterprise Team, Morgantown, WV, p 24

  • Ross R, Redell L, Bennett R (2002) Mesohabitat use of threatened hemlock forest by breeding birds of the Delaware Water Gap National Recreation Area. In: Onken B, Reardon R, Lashomb J (eds) Hemlock woolly adelgid symposium. USDA Forest Service and Rutgers University, East Brunswick, NJ, pp 353– 354

    Google Scholar 

  • SAS Institute Inc. (2002) SAS OnlineDoc. version 9. SAS Institute Inc., Cary, NC

    Google Scholar 

  • Sharov AA, Liebhold AM (1998) Model of slowing the spread of gypsy moth (Lepidoptera: Lymantriidae) with a barrier zone. Ecol Appl 8:1170–1179

    Article  Google Scholar 

  • Sharov AA, Pijanowski BC, Liebhold AM, Gage SH (1999) What affects the rate of gypsy moth (Lepidoptera: Lymantriidae) spread: winter temperature or forest susceptibility? Agric For Entomol 1:37–45

    Article  Google Scholar 

  • Shields K, Cheah CAS-J (2005) Winter mortality in Adeles tsugae populations in 2003 and 2004. In: Onken B, Reardon R (eds) Proceedings of the 3rd symposium on hemlock woolly adelgid. USDA Forest Service, Ashville, NC, USA, pp 354–356

    Google Scholar 

  • Shigesada N, Kawasaki K, Takeda Y (1995) Modeling stratified diffusion in biological invasions. Am Nat 146:229–251

    Article  Google Scholar 

  • Simberloff D (2000) Global climate change and introduced species in United States forests. Sci Total Environ 262:253–261

    Article  PubMed  CAS  Google Scholar 

  • Simberloff D, Von Holle B (1999) Positive interactions of nonindigenous species: invasional meltdown? Biol Invasions 1:21–32

    Article  Google Scholar 

  • Skellam JG (1951) Random dispersal in theoretical populations. Biometrika 38:196–218

    PubMed  CAS  Google Scholar 

  • Skinner M, Parker B, Gouli S, Ashikaga T (2003) Regional responses of hemlock woolly adelgid (Homoptera: Adelgidae) to low temperatures. Environ Entomol 32:523–528

    Google Scholar 

  • Snyder CD, Young JA, Lemarie DP, Smith DR (2002) Influence of eastern hemlock (Tsuga canadensis) forests on aquatic invertebrate assemblages in headwater streams. Can J Fish Aquat Sci 59:262–275

    Article  Google Scholar 

  • Therneau TM, Atkinson EJ (1997) An introduction to recursive partitioning using the RPART routine. Technical Report 61, Mayo Clinic, Section of Statistics, p 52

  • Tingley MW, Orwig DA, Field R, Motzkin G (2002) Avian response to removal of a forest dominant: consequences of hemlock woolly adelgid infestations. J Biogeogr 29:1505–1516

    Article  Google Scholar 

  • US Census Bureau (1990) Census 1990, Summary File 3

  • US Geological Survey (2000) Urbanization. National Atlas web edition. US Geological Survey, Reston, VA

    Google Scholar 

  • US Geological Survey and US Forest Service (2000) Forest cover types. US Geological Survey and USDA Forest Service, Reston, VA

  • Wallace MS, Hain FP (2000) Field surveys and evaluation of native and established predators of the hemlock woolly adelgid (Homoptera: Adelgidae) in the southeastern United States. Environ Entomol 29:638–644

    Article  Google Scholar 

  • Ward JS, Montgomery ME, Cheah CAS-J, Onken BP, Cowles RS (2004) Eastern hemlock forests: guidelines to minimize the impacts of hemlock woolly adelgid. NA-TP-03–04, USDA Forest Service, Northeastern Area State and Private Forestry, Morgantown, WV, p 32

  • Waring KM, O’Hara KL (2005) Silvicultural strategies in forest ecosystems affected by introduced pests. For Ecol Manage 209:27–41

    Article  Google Scholar 

  • White MA, Mladenoff DJ (1994) Old-growth forest landscape transitions from pre-European settlement to present. Landsc Ecol 9:191–205

    Article  Google Scholar 

  • Williamson MH (1996) Biological invasions. Chapman and Hall, London

    Google Scholar 

  • Williamson M, Fitter A (1996) The character of successful invaders. Biol Conserv 78:163–170

    Google Scholar 

  • Work TT, McCullough DG, Cavey JF, Komsa R (2005) Arrival rate of non-indigenous insect species into the United States through foreign trade. Biol Invasions 7:323–332

    Article  Google Scholar 

  • Young RF, Shields KS, Berlyn GP (1995) Hemlock woolly adelgid (Homoptera: Adelgidae): stylet bundle insertion and feeding sites. Ann Entomol Soc Am 88:827–835

    Google Scholar 

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Acknowledgements

We are grateful to Jennifer Bofinger, Charlie Burnham, Jason Denham, Don Ouellette, Alan Sior, and Brent Teillon for providing township records of HWA infestation as well as the John F. Enders Fund and the Connecticut Forest and Parks Association for financial support. R. Talbot Trotter III, and Brian Cade provided insightful comments on the manuscript.

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Correspondence to Alexander M. Evans.

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Evans, A.M., Gregoire, T.G. A geographically variable model of hemlock woolly adelgid spread. Biol Invasions 9, 369–382 (2007). https://doi.org/10.1007/s10530-006-9039-z

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