Maine Invasive Species Network

Crazy Worm Resources from the University of Vermont

(Bolded articles are ‘must-read.’)

Classification and Identification of Pheretimoids

• Blakemore, R.J., 2012. Japanese earthworms revisited a decade on: (Oligochaeta: Megadrilacea), Zoology in the Middle East, 58(sup4), January 2012, pp.15-22.
  • Japanese earthworms revisited a decade on (Oligochaeta: Megadrilacea) (PDF)
• Chang, C.h., Snyder, B.A. and Szlavecz, K., 2016. Asian pheretimoid earthworms in North America north of Mexico: An illustrated key to the genera Amynthas, Metaphire, Pithemera, and Polypheretima (Clitellata: Megascolecidae). Zootaxa, 4179(3), pp. 495-529.
  • Asian pheretimoid earthworms in North America north of Mexico: An illustrated key to the genera Amynthas (PDF)
• Gates, E.G. and J.W. Reynolds. 2017. Preliminary Key to North American Megadriles (Annelida, Oligochaeta), based on external characteristics, insofar as possible. Megadrilogica 22.

Distribution of Pheretimoids in the USA

• Callaham Jr, M.A., Hendrix, P.F. and Phillips, R.J., 2003. Occurrence of an exotic earthworm (Amynthas agrestis) in undisturbed soils of the southern Appalachian Mountains, USA: The 7th international symposium on earthworm ecology. Cardiff, Wales, 2002. Pedobiologia, 47(5-6), pp.466-470.
  • Occurrence of an exotic earthworm (Amynthas agresfis) in undisturbed soils of the southern Appalachian Mountains, USA (PDF)
• Chang, C.H., Johnston, M.R., Gorres, J.H., Davalos, A., McHugh, D. and Szlavecz, K., 2017. Co-invasion of three Asian earthworms, Metaphire hilgendorfi, Amynthas agrestis and Amynthas tokioensis in the USA. Biological Invasions, pp.1-6.
• Hendrix, P.F., Callaham Jr, M.A., Drake, J.M., Huang, C.Y., James, S.W., Snyder, B.A. and Zhang, W., 2008. Pandora’s box contained bait: the global problem of introduced earthworms. Annual Review of Ecology, Evolution, and Systematics, 39, pp.593-613.
• Keller, R.P., Cox, A.N., Van Loon, C., Lodge, D.M., Herborg, L.M. and Rothlisberger, J., 2007. From bait shops to the forest floor: earthworm use and disposal by anglers. The American Midland Naturalist, 158(2), pp.321-328.
• Moore, J.D., Gorres, J. and Reynolds, J.W., 2017. Exotic Asian pheretimoid earthworms (Amynthas spp., Metaphire spp.): Potential for colonization of south-eastern Canada and effects on forest ecosystems. Environmental Reviews.
• Reynolds, J.W., Gorres, J.H. and Knowles, M.E., 2015. A checklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae and Megascolecidae) in the states of Maine, New Hampshire and Vermont, USA. Megadrilogica, 17(9), pp.125-140.
  • A checklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae and Megascolecidae) in the states of Maine, New Hampshire and Vermont, USA (PDF)

Ecology of Pheretimoids

• Belliturk, K., Gorres, J.H., Kunkle, J. and Melnichuk, R.D.S., 2015. Can commercial mulches be reservoirs of invasive earthworms? Promotion of ligninolytic enzyme activity and survival of Amynthas agrestis (Goto and Hatai, 1899). Applied Soil Ecology, 87, pp.27-31.
• Burtelow, A.E., Bohlen, P.J. and Groffman, P.M., 1998. Influence of exotic earthworm invasion on soil organic matter, microbial biomass and denitrification potential in forest soils of the northeastern United States. Applied Soil Ecology, 9(1-3), pp.197-202.
• Chang, C.H., Szlavecz, K. and Buyer, J.S., 2017. Amynthas agrestis invasion increases microbial biomass in Mid-Atlantic deciduous forests. Soil Biology and Biochemistry, 114, pp.189-199.
• Gates, G.E., 1956. Reproductive organ polymorphism in earthworms of the oriental Megascolecine genus Pheretima Kinberg 1867. Evolution, 10(2), pp.213-227.
• Gorres, J.H. and Melnichuk, R.D., 2012. Asian invasive earthworms of the genus Amynthas Kinberg in Vermont. Northeastern Naturalist, 19(2), pp.313-322.
• Gorres, J.H., Connolly, S.T., Chang, C.H., Carpenter, N.R., Keller, E.L., Nouri-Aiin, M. and Schall, J.J., Winter hatching in New England populations of invasive pheretimoid earthworms Amynthas agrestis and Amynthas tokioensis: a limit on population growth, or aid in peripheral expansion?. Biological Invasions, pp.1-5.
• Greiner, H.G., Kashian, D.R. and Tiegs, S.D., 2012. Impacts of invasive Asian (Amynthas hilgendorfi) and European (Lumbricus rubellus) earthworms in a North American temperate deciduous forest. Biological Invasions, 14(10), pp.2017-2027.
• Hiller, B.J. and Barclay, J.S., 2011. Concentrations of heavy metals in American Woodcock harvested in Connecticut. Archives of environmental contamination and toxicology, 60(1), pp.156-164. Not directly related to Amynthas, but compare with Richardson et al., 2015.
• Loss, S.R. and Blair, R.B., 2011. Reduced Density and Nest Survival of Ground-Nesting Songbirds Relative to Earthworm Invasions in Northern Hardwood Forests. Conservation Biology, 25(5), pp.983-992. This is not a pheretimoid paper but these wildlife effects are likely to occur with Amynthas too.
• Qiu, J. and Turner, M.G., 2017. Effects of non-native Asian earthworm invasion on temperate forest and prairie soils in the Midwestern US. Biological Invasions, 19(1), pp.73-88.
• Reynolds, J.W., 1974. Are oligochaetes really hermaphroditic amphimictic organisms. Biologist, 56(2), pp.90-99.
  • Are oligochaetes really hermaphroditic amphimictic organisms? (PDF)
• Richardson, D.R., Snyder, B.A. and Hendrix, P.F., 2009. Soil moisture and temperature: tolerances and optima for a non-native earthworm species, Amynthas agrestis (Oligochaeta: Opisthopora: Megascolecidae). Southeastern Naturalist, 8(2), pp.325-334.
• Richardson, J.B., Gorres, J.H., Jackson, B.P. and Friedland, A.J., 2015. Trace metals and metalloids in forest soils and exotic earthworms in northern New England, USA. Soil Biology and Biochemistry, 85, pp.190-198.
• Snyder, B.A., Boots, B. and Hendrix, P.F., 2009. Competition between invasive earthworms (Amynthas corticis, Megascolecidae) and native North American millipedes (Pseudopolydesmus erasus, Polydesmidae): effects on carbon cycling and soil structure. Soil Biology and Biochemistry, 41(7), pp.1442-1449.
• Snyder, B.A., Callaham Jr, M.A., Lowe, C.N. and Hendrix, P.F., 2013. Earthworm invasion in North America: food resource competition affects native millipede survival and invasive earthworm reproduction. Soil Biology and Biochemistry, 57, pp.212-216.
  • Earthworm invasion in North America: food resource competition affects native millipede survival and invasive earthworm reproduction (PDF)
• Snyder, B.A., Callaham, M.A. and Hendrix, P.F., 2011. Spatial variability of an invasive earthworm (Amynthas agrestis) population and potential impacts on soil characteristics and millipedes in the Great Smoky Mountains National Park, USA. Biological Invasions, 13(2), pp.349-358.
• Zhang, W., Hendrix, P.F., Snyder, B.A., Molina, M., Li, J., Rao, X., Siemann, E. and Fu, S., 2010. Dietary flexibility aids Asian earthworm invasion in North American forests. Ecology, 91(7), pp.2070-2079.
• Ziemba, J.L., Hickerson, C.A.M. and Anthony, C.D., 2016. Invasive Asian Earthworms Negatively Impact Keystone Terrestrial Salamanders. PloS one, 11(5), p.e0151591.

Molecular Ecology

• Chang, C.H., Rougerie, R. and Chen, J.H., 2009. Identifying earthworms through DNA barcodes: Pitfalls and promise. Pedobiologia, 52(3), pp.171-180.
  • Identifying earthworms through DNA barcodes: Pitfalls and promise (PDF)
• Keller, E., J. Schall, J. Görres. 2017. Genetic Structure of Two Invasive Earthworms, Amynthas agrestis and Amynthas tokioensis (Oligochaeta, Megascolecidae) and a molecular method for species identification. Megadrilogica 22: 140-149
  • Genetic Structure of Two Invasive Earthworms, Amynthas agrestis and Amynthas tokioensis (Oligochaeta, Megascolecidae) and a molecular method for species identification (PDF)
• Schult, N., Pittenger, K., Davalos, S. and McHugh, D., 2016. Phylogeographic analysis of invasive Asian earthworms (Amynthas) in the northeast United States. Invertebrate Biology, 135(4), pp.314-327.

Pheretimoid Control

• Ikeda, H., Callaham Jr, M.A., O’Brien, J.J., Hornsby, B.S. and Wenk, E.S., 2015. Can the invasive earthworm, Amynthas agrestis, be controlled with prescribed fire?. Soil Biology and Biochemistry, 82, pp.21-27. Can the invasive earthworm, Amynthas agrestis, be controlled with prescribed fire? (PDF)
• Compiled by Josef Gorres, jgorres@uvm.edu, Plant and Soil Science Department, Jeffords Hall, University of Vermont, Burlington, Vermont 05405.
• Maryam Nouri-Aiin, Maryam.Nouri-Aiin@uvm.edu, Plant and Soil Science Department, Jeffords Hall, University of Vermont, Burlington, Vermont 05405.