Amynthas agrestis (crazy worm)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Environmental Impact
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Amynthas agrestis (Goto and Hatai, 1899)
Preferred Common Name
- crazy worm
Local Common Names
- USA: jumpers; snake worm; wigglers; wood eel
Summary of InvasivenessTop of page
Amynthas agrestis is an epigeic (litter-dwelling) Asian earthworm. It is native from Japan and the Korean Peninsula and has been introduced to the eastern United States, where it has spread widely, predominantly in forests. It has also been recorded from one location in Canada, near the USA border. Introduction and long-distance spread are thought to be associated with either transport of horticultural materials (for greenhouses, nurseries, and planting locations) or release of earthworms used as fishing bait.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Annelida
- Class: Oligochaeta
- Order: Haplotaxida
- Genus: Amynthas
- Species: Amynthas agrestis
Notes on Taxonomy and NomenclatureTop of page
The taxonomy of A. agrestis is and has been relatively stable. The species was originally described in the genus Perichaeta (Goto and Hatai, 1899), moved to Pheretima, and finally to Amynthas (Sims and Easton, 1972). Blakemore (2003) briefly and incorrectly listed the species in Metaphire, correcting this in subsequent publications (e.g. Blakemore, 2009).
DescriptionTop of page
The following description is based on the most recent summary of A. agrestis by Chang et al. (2016):
A. agrestis is a moderately sized, surface-dwelling (epigeic) earthworm of adult size 70-160 mm length and 5-8 mm width. Live specimens are red, but often appear somewhat brownish or purplish, with some iridescence. A. agrestis moves in a serpentine motion and will thrash/jump when disturbed, and it will often (but not always) autotomize a posterior body region in defence. As with other Amynthas, the clitellum is pale and surrounds the entire body in segments 14-16, each segment has a ring of many (40 or more) setae (perichaetine) in the middle of the segment. This group of earthworms is identified by using the number and location of sexual pores and swellings (genital markings) on the body and the shape of sexual (spermathecae) and somatic (intestinal caecae) organs. A. agrestis usually lacks male pores and genital markings posterior to the clitellum (both found ventrally in segment 18); when present, the pores are small and slit-like, and the markings are large, circular, slightly concave pads, just anterior and median to the pores. A. agrestis typically has three pairs of spermathecal pores ventrally at the intersegmental furrow between segments 5 & 6, 6 & 7, and 7 & 8, with corresponding spermathecae inside segments 6, 7 and 8. Spermathecal number may be reduced. Genital markings anterior to the clitellum are often absent; when present, these appear as slightly discoloured and wrinkled areas located ventrally in segments 7 and/or 8. These may be single/median or paired. Internally, spermathecae have a duct shorter than the ampulla, with a thin, cylindrical diverticulum that is longer than the combined duct and ampulla (illustrated in Chang et al., 2016). Prostate glands are usually absent but, when present, are found in segment 23, and can extend through segment 16. Paired intestinal caecae have 5-7 finger-shaped divisions (manicate). They branch from the intestine in segment 27 and extend anteriorly for a few segments.
A. agrestis and other species in this group can be challenging to identify due to variability in several characteristics. In case of preserved specimens, identification can be even harder due to poor visibility of specific characteristics because of low quality of preservation. Positive identification of the species requires dissection of an adult specimen for confirmation of location and shape of spermathecae, intestinal caecae, and other structures. As of 2017, for the known species of earthworms in North America north of Mexico, an adult (clitellate) earthworm with all of the following characteristics should be identified as A. agrestis: perichaetine setae, size >50 mm, male pores absent, three pairs of spermathecae. As described above, other combinations of characteristics are also possible, but confirmation of the species would be needed.
Using morphological characteristics, juveniles can only be identified as pheretimoid (Amynthas or related genera). Internal organs grow gradually, thus moderately sized juveniles can sometimes be positively identified with dissection.
DistributionTop of page
The native range of A. agrestis includes the four main islands of Japan and the Korean Peninsula. Following introduction, it is now widespread in eastern North America, from Alabama and Georgia to New Hampshire and Vermont. Additional locations include northern Louisiana, eastern Texas, southeastern Oklahoma (Tandy, 1969; Gates, 1982; Damoff and Reynolds, 2009), one location in Wisconsin (Qiu and Turner, 2017), one in Ontario (Reynolds, 2014) and a greenhouse in Maine (Gates, 1966). While many of these localities consist of natural or managed habitats, such as arboreta (e.g., Gates, 1953), A. agrestis has been recorded in many human-created environments (e.g., culture beds, nurseries, compost heaps; Gates, 1954; 1958; 1963; Reynolds, 1977).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Japan||Present||Native||Present based on regional distribution|
|-Honshu||Present||Native||Aomori Prefecture, Iawati Prefecture, Takahashi, Tokorozawa, Oarai, Sendai, Oshima Island|
|Canada||Present||Introduced||Invasive||Present based on regional distribution|
|United States||Present||Introduced||Invasive||Present based on regional distribution|
|-Alabama||Present||Introduced||Invasive||Dallas, Montgomery, and Tuscaloosa Cos.|
|-Kentucky||Present||Introduced||Invasive||Daniel Boone National Forest - single record in a remote area, needs confirmation|
|-Louisiana||Present||Introduced||Invasive||Widespread north of the Red River|
|-Maryland||Present||Introduced||Invasive||First North American record|
|-New Hampshire||Present||Introduced||Invasive||Belknap Co.|
|-Ohio||Present||Introduced||Invasive||Clinton, Lake, and Summit Cos; Original citation: Reynolds (2015a)|
|-Pennsylvania||Present||Introduced||Invasive||Original citation: Reynolds (2008c)|
|-South Carolina||Present||Introduced||Invasive||Darlington and Jasper Cos.|
|-Texas||Present||Introduced||Invasive||Widespread in east Texas|
|-Vermont||Present||Introduced||Invasive||Present in multiple sites throughout the state|
|-Virginia||Present||Introduced||Invasive||Henry Co; Original citation: Reynolds (2015b)|
|-West Virginia||Present||Introduced||Invasive||Summers Co.|
|-Wisconsin||Present||Introduced||Invasive||Madison, University of Wisconsin - Madison Arboretum|
History of Introduction and SpreadTop of page
Details of the introduction of A. agrestis into the USA are unknown. Presumably, as is the case for other invasive earthworms, the species was introduced multiple times, with the first introductions occurring before the first A. agrestis record in 1939 (Howell, 1939). Asian earthworm introductions were ongoing at least half a century prior - the first pheretimoid record in North America was from a greenhouse (hot-house) in Illinois, in 1888 (Garman, 1888). Recognising this fact, Howell (1939) speculated about the species route of introduction: “introduced into this country with botanical products imported probably 15 to 20 years ago.” Details of the introduction of A. agrestis to Canada are unknown. The species may have been transported directly from its native range or from the USA.
Early records of the species are all associated with exotic plants. It is thus likely that, as Howell (1939) surmised, the original introduction(s) came with soil or other organic material accompanying plants from Asia. These were then planted in arboreta (Gates, 1953; Qiu and Turner, 2017) or grown in nurseries/culture beds/greenhouses (e.g., Gates, 1958; Gates, 1963). Plants and the earthworms associated with them were then transported and planted at various locations, which contributed to widely dispersing the species (Gates, 1958; Tandy, 1969).
An additional route of dispersal has been through the use of the species as bait for anglers (Gates, 1958). The species has been observed for sale in bait shops (Callaham et al., 2003) and there is evidence that populations have been started intentionally to provide a local source of bait (Tandy, 1969). Many of the invaded natural habitats are near actively fished waters (Tandy, 1969; Callaham et al., 2003; Snyder et al., 2011).
Risk of IntroductionTop of page
The routes of introduction/spread mentioned in the 'History of Introduction and Spread' section (horticultural materials and bait) continue to be relevant today and may unknowingly result in new invasions in remote or protected areas (e.g., Callaham et al., 2003; Snyder et al., 2011). The risk of introduction from horticultural materials may extend to any organic matter product associated with horticulture or landscaping, including soil, leaf litter, compost and mulch. Mulch (pieces of shredded wood up to 5 cm) is not an ideal habitat for this species, but survival in/under mulch has been documented and is linked to a high risk for transport and establishment of this species in gardens (Bellitürk et al., 2015).
In addition to local purchase for bait or composting, earthworms purported to be A. agrestis can also be ordered online (Ziemba et al., 2016). Earth-moving activities (e.g., road building) have been shown to act as pathways of introduction and spread for other invasive earthworms, being a potential route of transport for A. agrestis.
Habitat ListTop of page
|Terrestrial-managed/Protected agriculture (e.g. glasshouse production)||Present, no further details||Harmful (pest or invasive)|
|Terrestrial-managed/Managed forests, plantations and orchards||Present, no further details||Harmful (pest or invasive)|
|Terrestrial-managed/Disturbed areas||Present, no further details||Harmful (pest or invasive)|
|Terrestrial-managed/Rail / roadsides||Present, no further details||Harmful (pest or invasive)|
|Terrestrial-managed/Urban / peri-urban areas||Present, no further details||Harmful (pest or invasive)|
|Terrestrial-natural/semi-natural/Natural forests||Present, no further details||Harmful (pest or invasive)|
|Terrestrial-natural/semi-natural/Riverbanks||Present, no further details||Harmful (pest or invasive)|
|Other/Soil||Present, no further details||Harmful (pest or invasive)|
Biology and EcologyTop of page
A. agrestis genetics has been little studied, although interest in using molecular techniques to aid identification has been increasing. Schult et al. (2016) used two loci to examine the genetic divergence of Amynthas spp. and suggest that there may be cryptic speciation. However, morphological identification of some individuals by an expert was able to refer specimens in each lineage to a valid species, only by using species morphology.
All earthworms are hermaphrodites and many, including A. agrestis, have developed parthenogenesis (Gates, 1958; Chang et al., 2016). Thus, most individuals encountered in the native or introduced range lack male pores and/or other parts of the male sexual system. Unlike many earthworms, A. agrestis has an annual life cycle (Tandy, 1969; Reynolds, 1978; Callaham et al., 2003; Chang et al., 2016). They overwinter as cocoons, hatch in spring, reach adulthood in summer, lay cocoons and all adults die by winter. A. agrestis lays spherical or subspherical cocoons of slightly under 2 mm diameter (Snyder et al., 2013). Cocoon colour is variable, from yellow to brown to red. Most cocoons contain one individual, but ~2% contain two individuals (Ikeda et al., 2015). Cocoon viability after 4 months has been shown to be between 60 and 70% (Ikeda et al., 2015). External sexual structures do not develop until near sexual maturity.
Physiology and Phenology
In Japan (Kanagawa Prefecture), A. agrestis juveniles were found in April-May and adults beginning in June (Uchida and Kaneko, 2004). In Louisiana, USA, juveniles were found April-June and adults were present in May-September (Tandy, 1969). In Tennessee, USA, juveniles or aclitellate adults were found May-June and adults were collected in June-December (Reynolds, 1978). Adults collected in September in Tennessee and North Carolina laid cocoons between September and November (Blackmon, 2009). At a high elevation site in north Georgia, USA (~1450m above sea level), juveniles were collected July-September, while adults were collected August-November (Callaham et al., 2003). Similarly, in Vermont, USA adults were observed July-November (Görres et al., 2014).
A. agrestis is known to bioaccumulate heavy metals, particularly Se, Zn, Cd and Hg (Richardson et al., 2015).
A. agrestis has a life cycle lasting ~1 year. Little research has monitored the cocoon stage. Development from hatchling to adult is estimated to be a minimum of 90 days, based on frost-free periods of invaded locations (Görres and Melnichuk, 2012). Although, in the field, adults die in late fall or early winter, they have been kept alive in the laboratory through February (Snyder et al., 2013; Ikeda et al., 2015).
Soil and litter both play a large role in the biology of A. agrestis. Soil is a requirement for survival, weight maintenance and reproduction: in mesocosms with no soil (leaf litter only), A. agrestis lost weight and died, producing no cocoons (Ikeda et al., 2015). In this same study, A. agrestis with only soil survived, lost more weight than treatments with both litter and soil, and produced a limited number of cocoons. A. agrestis performs better on partially-decomposed leaf litter over fresh leaves, and appears to derive some nutrition from soil (Snyder et al., 2013; Ikeda et al., 2015). A combination of all three substrates provided the best nutrition (Ikeda et al., 2015). Zhang et al. (2010) found that A. agrestis feeds more on soil than Lumbricus rubellus (an epigeic European species also introduced in the USA) and, when needed, can shift its feeding strategy to more available soil or litter microbes.
A. agrestis can also survive in soil with mulch on top. Bellitürk et al. (2015) found that, in addition to survival and maturation over a 7 week incubation period with spruce, cedar, or pine mulch, A. agrestis increased lignin-digesting enzyme activity (phenoloxidase and peroxidase) in these soils. Mortality rates in this experiment were 10-30%, but there was no control against which to measure the effect of mulch on this variable.
In the laboratory, adult A. agrestis did not survive at soil temperatures of -5, 5 or 35°C, but did survive at 12 and 25°C; moisture played a role, with no survival at 8% moisture and 25°C, but higher survival in wetter soils at the same temperature (Richardson et al., 2009). Field studies in Vermont indicate that 5°C may not be a minimum threshold temperature and that specimen size may play a role (Görres et al., 2014). Görres and Melnichuk (2012) suggest that the frost-free period (air temperature) may be an important predictor of A. agrestis tolerance of a site, because of time required to mature.
Temperature may play an important role for the development and hatching of cocoons. Blackmon (2009) examined whether cocoons would hatch at temperatures ranging from 5 to 30°C. Hatching rate was highest at 10°C and varied between 33, 60, and 100%, depending on the site where earthworms were collected. No hatching occurred at 5°C, and hatching occurred sooner at higher temperatures. Blackmon (2009) also tested whether changes in temperature (e.g., a winter cooling followed by spring warming) would affect hatching. Cocoons were lowered to 5, 10 or 15°C and incubated for 15, 30 or 60 days. Hatching occurred at all incubation lengths, but only at 10ºC, with the exception of one cocoon at 5°C. Cocoons also predominantly hatched 5-6 weeks after warming, regardless of incubation length.
ClimateTop of page
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Dw - Continental climate with dry winter||Tolerated||Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)|
|Df - Continental climate, wet all year||Preferred||Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
Notes on Natural EnemiesTop of page
The centipede Scolopocryptops sexspinosus and centipedes from the family Cryptopidae, rusty crayfish (Faxonius rusticus), wandering broadhead planarian (Bipalium adventitium), ribbon leech (Nephelopsis obscura) and salamanders Desmognathus monticola, Plethodon metcalfi and P. teyahalee have been documented to prey on A. agrestis (Craft, 2009; Gorsuch and Owen, 2014; Gao et al., 2017).
Means of Movement and DispersalTop of page
Natural dispersal of A. agrestis can be as fast as 12 m yr-1, but appears to be highly variable and weather dependent (Snyder et al., 2011).
A. agrestis has been - and continues to be - introduced across eastern North America due to two accidental means: the movement of horticultural/landscaping materials (Howell, 1939; Gates, 1958; Tandy, 1969; Bellitürk et al., 2015) and use as fishing bait (Gates, 1958; Callaham et al., 2003). Earth-moving activities, such as road building, may also have introduced the species to new locations, but this has not been formally documented.
A. agrestis has been intentionally introduced to some environments, in order to start populations of bait near fishing spots (Tandy, 1969). This may have occurred on a broader scale, but it has not been further documented.
Pathway CausesTop of page
|Botanical gardens and zoos||Raised to feed platypuses||Yes||Gates (1954)|
|Escape from confinement or garden escape||Movement with soil or other organic material accompanying plants||Yes||Gates (1958)|
|Horticulture||Movement with soil or other organic material accompanying plants||Yes||Yes||Gates (1958; 1966); Tandy (1969)|
|Hunting, angling, sport or racing||Intentional and accidental introduction for bait||Yes||Yes||Gates (1958); Tandy (1969); Callaham et al. (2003)|
|Internet sales||Can be ordered online||Yes||Yes||Ziemba et al. (2016)|
|Landscape improvement||Movement with soil or other organic material accompanying plants||Yes||Yes||Görres and Melnichuk (2012); Bellitürk et al. (2015)|
|Nursery trade||Movement with soil or other organic material accompanying plants||Yes||Yes||Gates (1954; 1958); Görres and Melnichuk (2012)|
Pathway VectorsTop of page
|Mulch, straw, baskets and sod||Likely dispersed to many new locations via mulch||Yes||Yes||Bellitürk et al. (2015)|
|Plants or parts of plants||All stages possible in soil or other organic material accompanying plants||Yes||Yes||Tandy (1969)|
|Soil, sand and gravel||All stages possible in soil||Yes||Yes||Gates (1953; 1954; 1958)|
|Can be ordered online||Yes||Yes||Ziemba et al. (2016)|
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Growing medium accompanying plants||Yes||Pest or symptoms usually visible to the naked eye|
Impact SummaryTop of page
Environmental ImpactTop of page
Impact on Habitats
Snyder et al. (2011) found that, in the Great Smoky Mountains National Park, USA, A. agrestis was effective at removing partially-decomposed litter layers and changing the surface soil structure. Increased presence of A. agrestis was associated with increased amounts of large (> 2 mm) water-stable soil aggregates and a decrease in the thickness of the partially-decomposed (F and H) litter layer. However, A. agrestis presence was not found to change C:N or microbial biomass. Snyder et al. (2013) attributed litter decrease to direct consumption by A. agrestis.
Qiu and Turner (2017) found, both in mesocosms and in the field, decreased amounts of leaf litter due to co-invasion of A. agrestis and A. tokioensis. In mesocosms, surface soils (0-5 cm) showed increased organic matter, total carbon, total nitrogen, C:N and available phosphorus; in the field, these changes occurred, but not at a consistent rate over time. Deeper soils had an increase in inorganic nitrogen and dissolved organic carbon (Qiu and Turner, 2017).
A. agrestis presence also increases activity of lignin-digesting enzymes (Bellitürk et al., 2015).
Impact on Biodiversity
A. agrestis changes the chemistry and physical structure of the litter layer and upper soil, and thus is expected to impact any organisms that use these horizons. Removal of litter, in particular, will affect many detritivorous fauna and litter-dwellers. Snyder et al. (2011; 2013) found that millipede diversity and abundance declined in the field due to A. agrestis invasion, and that there was evidence for direct competition between millipedes (Sigmoria lyrea [Falloria lyrea]) and A. agrestis. Effects on other species are virtually unknown.
Gorsuch and Owen (2014) found that A. agrestis was less likely to be predated than lumbricid earthworms by wandering broadhead planarian (Bipalium adventitium), ribbon leech (Nephelopsis obscura) and seal salamander (Desmognathus monticola). However, this is the only record of A. agrestis from Kentucky and some behaviours described do not fit what is typical for A. agrestis. Given the recent discoveries of multiple Amynthas species living in the same habitat (Chang et al., 2016), it is possible that multiple species were unknowingly used in this experiment.
Several authors have raised concerns about the interaction between A. agrestis and salamanders. Craft (2009) conducted a mesocosm study with salamanders (Plethodon metcalfi and P. teyahalee) using soil from an A. agrestis invasion front. This study showed no difference in salamander body or faeces mass, but the total organic carbon, total nitrogen, and C:N differed significantly, suggesting that prey quantity and/or quality were diminished in the invaded area. A. agrestis was also not a preferred food item in feeding trials.
Ziemba et al. (2015; 2016) found that red-backed salamanders (Plethodon cinereus) and pheretimoid earthworms co-occurred less than expected. Leaf litter layers were qualitatively thinner and lower quality in invaded sites, but there was no effect of invasion on salamander body condition, number of eggs, or tail breakage. In a laboratory study, Ziemba et al. (2015; 2016) also found that P. cinereus used lower quality habitats and consumed less food in the presence of pheretimoid earthworms.
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Abundant in its native range
- Highly mobile locally
- Fast growing
- Reproduces asexually
- Ecosystem change/ habitat alteration
- Reduced native biodiversity
- Threat to/ loss of native species
- Competition (unspecified)
- Rapid growth
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
UsesTop of page
Uses ListTop of page
Animal feed, fodder, forage
- Sport (hunting, shooting, fishing, racing)
Detection and InspectionTop of page
A. agrestis is surface-dwelling and is best found by hand-searching through the leaf litter, at the soil surface, and under logs/rocks (Snyder et al., 2011). Decreased levels of partially-decomposed leaf litter and a granular surface soil may indicate an earthworm invasion, but not necessarily of this species. Wetter areas (e.g., along stream banks, valleys, deeper pockets of leaf litter) and areas impacted by human activity (e.g., roadsides, developed edges of forests) are more likely to support populations of A. agrestis.
Similarities to Other Species/ConditionsTop of page
A. agrestis can be easily confused with other pheretimoid earthworms. Amynthas alone includes hundreds of species. Sixteen species in Amynthas and closely related genera have been found in North America; Chang et al. (2016) provide a detailed key for these species. Most species can be easily differentiated from A. agrestis by size, colouration in life, and location of easily observable pores and markings. A. tokioensis and Metaphire hilgendorfi are quite similar to A. agrestis and the three species have been shown to coexist in some locations (Chang et al., 2016). Chang et al. (2016) provide a comparison table to help differentiate these species.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
Blackmon (2009) and Ikeda et al. (2015) examined the role of fire in the control of A. agrestis invasion. Neither study found direct mortality, but both appear to have found indirect impacts: indirect/delayed mortality and weight loss, potentially due to starvation (Blackmon, 2009), and decreased cocoon viability (Ikeda et al., 2015).
Gaps in Knowledge/Research NeedsTop of page
Despite its wide dispersal across eastern North America over the last century, the biology and ecology of A. agrestis are very poorly known. The same is true for other invasive pheretimoid species, for which often no information exists beyond their taxonomic description (Burtelow et al., 1998). Understanding traits which aid A. agrestis’ dispersal, establishment and spread (e.g., life history traits, Callaham et al., 2003) is necessary, in order to fully understand and combat these invasions. Tracing pathways of invasion and quantifying their contribution to A. agrestis spread would also be very useful in slowing or stopping the spread of this species.
Earthworm invasions may be nearly impossible to eliminate without wholesale ecosystem destruction, but prescribed fire is one avenue that has shown promise and which needs further exploration.
More research is needed on the diversity of parthenogenetic morphs within the pheretimoids and the distribution of each invading species, as well as effective techniques to accurately identify species.
ReferencesTop of page
BellItürk, K., Görres, J. H., Kunkle, J., 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, 27-31. http://www.sciencedirect.com/science/journal/09291393 doi: 10.1016/j.apsoil.2014.11.007
Bernard, M. J., Neatrour, M. A., McCay, T. S., 2009. Influence of soil buffering capacity on earthworm growth, survival, and community composition in the western Adirondacks and central New York. Northeastern Naturalist, 16(2), 269-284. doi: 10.1656/045.016.0208
Bhatti HK, 1965. Earthworms of Swarthmore, Pennsylvania, and vicinity. Proceedings of the Pennsylvania Academy of Science, 39(2), 8-24.
Blackmon JH, 2009. The use of fire in the control of invasive, epigeic earthworm species in the southeastern United States. MPhil Thesis. Athens, GA, USA: University of Georgia.
Blakemore RJ, 2003. Japanese earthworms (Annelida:Oligochaeta): a review and checklist of species. Organisms, Diversity, and Evolution, Electronic Supplement, 11, 1-43. http://www.senckenberg.uni-frankfurt.de/odes/03-11.pdf
Blakemore RJ, 2009. Cosmopolitan earthworms - a global and historical perspective. In: Annelids as model systems in the biological sciences, [ed. by Shain DH]. New York, USA: John Wiley & Sons, Inc. 257-283.
Burtelow, A. E., Bohlen, P. J., 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. In: Applied Soil Ecology [Soil organisms and soil resource management. Proceedings of the XII International Colloquium on soil zoology, Dublin, 22-26 July, 1996], 9(1/3) . 197-202.
Callaham, M. A., Jr., Hendrix, P. F., Phillips, R. J., 2003. Occurrence of an exotic earthworm (Amynthas agrestis) in undisturbed soils of the southern Appalachian Mountains, USA. Pedobiologia, 47(5/6), 466-470. doi: 10.1078/0031-4056-00214
Chang ChihHan, Snyder, B. A., 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), 495-529. http://www.mapress.com/j/zt/article/view/zootaxa.4179.3.7 doi: 10.11646/zootaxa.4179.3.7
Craft JJ, 2009. Effects of an invasive earthworm on plethodontid salamanders in Great Smoky Mountains National Park. MPhil Thesis. Cullowhee, NC, USA: Western Carolina University.
Damoff GA, Reynolds JW, 2009. The earthworms (Oligochaeta: Acanthodrilidae, Eudrilidae, Lumbricidae, Megascolecidae, Ocnerodrilidae, and Sparganophilidae) of east Texas, USA. Megadrilogica, 13(8), 113-140.
Davies H, 1954. A preliminary list of the earthworms of northern New Jersey with notes. Breviora, 26, 1-13.
Gao MeiXiang, Taylor, M. K., Callaham, M. A., Jr., 2017. Trophic dynamics in a simple experimental ecosystem: interactions among centipedes, Collembola and introduced earthworms. Soil Biology & Biochemistry, 115, 66-72. http://www.sciencedirect.com/science/journal/00380717 doi: 10.1016/j.soilbio.2017.08.001
Garman H, 1888. On the anatomy and histology of a new earthworm (Diplocardia communis, gen. et sp. nov.). Bulletin of the Illinois State Laboratory of Natural History, 3, 47-77.
Gates GE, 1953. Further notes on the earthworms of the Arnold Arboretum, Boston. Breviora, 15, 1-9.
Gates GE, 1954. Exotic earthworms of the United States. Bulletin of the Museum of Comparative Zoology, 111(6), 219-238.
Gates GE, 1958. On some species of the oriental earthworm genus Pheretima Kinberg, 1867, with key to species reported from the Americas. American Museum Novitates, 1888, 1-33.
Gates GE, 1963. Miscellanea Megadrilogica. VII. Greenhouse earthworms. Proceedings of the Biological Society of Washington, 76, 9-18.
Gates GE, 1966. Requiem - for megadrile utopias. A contribution toward the understanding of the earthworm fauna of North America. Proceedings of the Biological Society of Washington, 79, 239-254.
Gates GE, 1982. Farewell to North American megadriles. Megadrilogica, 4(1-2), 12-77.
Gorsuch, J. P., Owen, P. C., 2014. Potential edaphic and aquatic predators of a nonindigenous Asian earthworm (Amynthas agrestis) in the Eastern United States. Northeastern Naturalist, 21(4), 652-661. http://www.eaglehill.us/NENAonline/articles/NENA-21-4/21-Gorsuch.shtml doi: 10.1656/045.021.0415
Goto S, Hatai S, 1899. New or imperfectly known species of earthworms. No. 2. Annotationes Zoologicae Japonenses, 3, 13-24.
Görres JH, Melnichuk RDS, Bellitürk K, 2014. Mortality pattern relative to size variation within Amynthas agrestis (Goto & Hatai, 1899) (Oligochaeta: Megascoledcidae) populations in the Champlain Valley of Vermont, USA. Megadrilogica, 16(2), 9-14.
Görres, J. H., Melnichuk, R. D. S., 2012. Asian invasive earthworms of the genus Amynthas Kinberg in Vermont. Northeastern Naturalist, 19(2), 313-322. http://www.eaglehill.us/nena doi: 10.1656/045.019.0212
Hatai S, 1930. Note on Pheretima agrestis (Goto and Hatai), together with the description of four new species of the genus Pheretima. The Science Reports of the Tôhoku Imperial University Fourth Series (Biology), 5(4), 651-667.
Howell CD, 1939. The responses to light in the earthworm, Pheretima agrestis Goto and Hatai, with special reference to the function of the nervous system. Journal of Experimental Zoology, 81(2), 231-259.
Ikeda, H., Callaham, M. A., Jr., O'Brien, J. J., Hornsby, B. S., Wenk, E. S., 2015. Can the invasive earthworm, Amynthas agrestis, be controlled with prescribed fire?. Soil Biology & Biochemistry, 82, 21-27. http://www.sciencedirect.com/science/journal/00380717 doi: 10.1016/j.soilbio.2014.12.011
Kobayashi S, 1938. Earthworms of Korea I. The Science Reports of the Tôhoku Imperial University Fourth Series (Biology), 13(2), 89-170.
Qiu JiangXiao, 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), 73-88. http://link.springer.com/article/10.1007/s10530-016-1264-5 doi: 10.1007/s10530-016-1264-5
Reynolds JW, 1978. The earthworms of Tennessee (Oligochaeta) IV. Megascolecidae, with notes on the distribution, biology, and a key to the species in the state. Megadrilogica, 3(7), 117-129.
Reynolds JW, 2008a. The earthworms (Oligochaeta, Lumbricidae) of Maine, USA. Megadrilogica, 12(4), 61-72.
Reynolds JW, 2008b. The earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae and Sparganophilidae) of Missouri, USA. Megadrilogica, 12(6), 89-104.
Reynolds JW, 2008c. The earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae and Sparganophilidae) of Pennsylvania, USA. Megadrilogica, 11(12), 132-146.
Reynolds JW, 2014. A checklist by counties of earthworms (Oligochaeta: Lumbricidae, Megascolecidae and Sparganophilidae) in Ontario, Canada. Megadrilogica, 16(10), 111-135.
Reynolds JW, 2015a. A checklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae, and Sparganophilidae) in Ohio, USA. Megadrilogica, 18(1), 1-20.
Reynolds JW, 2015b. Earthworms of the families Lumbricidae, Megascolecidae and Sparganophilidae in the collections of the Virginia Museum of Natural History. Megadrilogica, 19(2), 11-22.
Reynolds JW, Görres JH, Knowles ME, 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), 125-140.
Reynolds, J. W., 1994. Earthworms of North Carolina (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae, Meascolecidae, Ocnerodrilidae and Sparganophilidae). In: Megadrilogica , 5(6) . Lindsay, Canada: Oligichaetology Laboratory, Sir Sandford Fleming College.18 pp.
Richardson, D. R., Snyder, B. A., 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), 325-334. doi: 10.1656/058.008.0211
Richardson, J. B., Görres, J. H., Jackson, B. P., Friedland, A. J., 2015. Trace metals and metalloids in forest soils and exotic earthworms in northern New England, USA. Soil Biology & Biochemistry, 85, 190-198. http://www.sciencedirect.com/science/journal/00380717 doi: 10.1016/j.soilbio.2015.03.001
Schermaier AF, 2013. Influence of earthworms on plant and soil invertebrate communities of the Cleveland Metroparks. MPhil Thesis. Cleveland, OH, USA: Cleveland State University.
Schult N, Pittenger K, Davalos S, McHugh D, 2016. Phylogeographic analysis of invasive Asian earthworms (Amynthas) in the northeast United States. Invertebrate Biology, 135(4), 314-327.
Sims, R. W., Easton, E. G., 1972. A numerical revision of the earthworm genus Pheretima auct. Megascolecidae: Oligochaeta) with the recognition of new genera and an appendix on the earthworms collected by the Royal Society North Borneo Expedition. Biological Journal of the Linnean Society, 4(No.3), 169-269. doi: 10.1111/j.1095-8312.1972.tb00694.x
Snyder, B. A., Callaham, M. A., 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), 349-358. http://www.springerlink.com/content/m422k472t0w4p574/ doi: 10.1007/s10530-010-9826-4
Snyder, B. A., Callaham, M. A., Jr., Lowe, C. N., Hendrix, P. F., 2013. Earthworm invasion in North America: food resource competition affects native millipede survival and invasive earthworm reproduction. Soil Biology & Biochemistry, 57, 212-216. http://www.sciencedirect.com/science/journal/00380717 doi: 10.1016/j.soilbio.2012.08.022
Stebbings JH, 1962. Additions to the Illinois and Missouri earthworm fauna. American Midland Naturalist, 67(2), 504-505.
Tandy RE, 1969. The earthworm genus Pheretima, Kinberg, 1866, in Louisiana (Oligochaeta: Megascolecidae). DPhil Thesis. Baton Rouge, LA, USA: Louisiana State University and Agricultural and Mechanical College.
Uchida T, Kaneko N, 2004. Life history of Megascolecidae earthworms in forest soils at Kanagawa, Japan. Edaphologia, 74, 35-45.
Zhang WeiXin, Hendrix, P. F., Snyder, B. A., Molina, M., Li JianXiong, Rao XingQuan, Siemann, E., Fu ShengLei, 2010. Dietary flexibility aids Asian earthworm invasion in North American forests. Ecology, 91(7), 2070-2079. http://www.esajournals.org/doi/abs/10.1890/09-0979.1 doi: 10.1890/09-0979.1
Ziemba, J. L., Cameron, A. C., Peterson, K., Hickerson, C. A. M., Anthony, C. D., 2015. Invasive Asian earthworms of the genus Amynthas alter microhabitat use by terrestrial salamanders. Canadian Journal of Zoology, 93(10), 805-811. http://www.nrcresearchpress.com/doi/full/10.1139/cjz-2015-0056#.ViYOe1LvodU
Ziemba, J. L., Hickerson, C. A. M., Anthony, C. D., 2016. Invasive Asian earthworms negatively impact keystone terrestrial salamanders. PLoS ONE, 11(5), e0151591. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0151591 doi: 10.1371/journal.pone.0151591
Bernard M J, Neatrour M A, McCay T S, 2009. Influence of soil buffering capacity on earthworm growth, survival, and community composition in the western Adirondacks and central New York. Northeastern Naturalist. 16 (2), 269-284. DOI:10.1656/045.016.0208
Bhatti HK, 1965. Earthworms of Swarthmore, Pennsylvania, and vicinity. In: Proceedings of the Pennsylvania Academy of Science, 39 (2) 8-24.
Blackmon JH, 2009. The use of fire in the control of invasive, epigeic earthworm species in the southeastern United States. MPhil Thesis., Athens, GA, USA: University of Georgia.
CABI, Undated. Compendium record. Wallingford, UK: CABI
CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Callaham M A Jr, Hendrix P F, Phillips R J, 2003. Occurrence of an exotic earthworm (Amynthas agrestis) in undisturbed soils of the southern Appalachian Mountains, USA. Pedobiologia. 47 (5/6), 466-470. DOI:10.1078/0031-4056-00214
Damoff GA, Reynolds JW, 2009. The earthworms (Oligochaeta: Acanthodrilidae, Eudrilidae, Lumbricidae, Megascolecidae, Ocnerodrilidae, and Sparganophilidae) of east Texas, USA. In: Megadrilogica, 13 (8) 113-140.
Davies H, 1954. A preliminary list of the earthworms of northern New Jersey with notes. In: Breviora, 26 1-13.
Gates GE, 1953. Further notes on the earthworms of the Arnold Arboretum, Boston. In: Breviora, 15 1-9.
Gates GE, 1954. Exotic earthworms of the United States. In: Bulletin of the Museum of Comparative Zoology, 111 (6) 219-238.
Gates GE, 1958. On some species of the oriental earthworm genus Pheretima Kinberg, 1867, with key to species reported from the Americas. In: American Museum Novitates, 1888 1-33.
Gates GE, 1963. Miscellanea Megadrilogica. VII. Greenhouse earthworms. In: Proceedings of the Biological Society of Washington, 76 9-18.
Gates GE, 1966. Requiem - for megadrile utopias. A contribution toward the understanding of the earthworm fauna of North America. In: Proceedings of the Biological Society of Washington, 79 239-254.
Gates GE, 1982. Farewell to North American Megadriles. In: Megadrilogica, 4 12-77.
Gorsuch J P, Owen P C, 2014. Potential edaphic and aquatic predators of a nonindigenous Asian earthworm (Amynthas agrestis) in the Eastern United States. Northeastern Naturalist. 21 (4), 652-661. http://www.eaglehill.us/NENAonline/articles/NENA-21-4/21-Gorsuch.shtml DOI:10.1656/045.021.0415
Goto S, Hatai S, 1899. New or imperfectly known species of earthworms. No. 2. In: Annotationes Zoologicae Japonenses, 3 13-24.
Hatai S, 1930. Note on Pheretima agrestis (Goto and Hatai), together with the description of four new species of the genus Pheretima. In: The Science Reports of the Tôhoku Imperial University Fourth Series (Biology), 5 (4) 651-667.
Howell CD, 1939. The responses to light in the earthworm, Pheretima agrestis Goto and Hatai, with special reference to the function of the nervous system. In: Journal of Experimental Zoology, 81 (2) 231-259.
Kobayashi S, 1938. Earthworms of Korea I. In: The Science Reports of the Tôhoku Imperial University Fourth Series (Biology), 13 (2) 89-170.
Qiu JiangXiao, 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), 73-88. http://link.springer.com/article/10.1007/s10530-016-1264-5 DOI:10.1007/s10530-016-1264-5
Reynolds J W, 1994. Earthworms of North Carolina (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae, Meascolecidae, Ocnerodrilidae and Sparganophilidae). In: Megadrilogica, 5 (6) Lindsay, Canada: Oligichaetology Laboratory, Sir Sandford Fleming College. 18 pp.
Reynolds JW, 1978. The earthworms of Tennessee (Oligochaeta) IV. Megascolecidae, with notes on the distribution, biology, and a key to the species in the state. In: Megadrilogica, 3 (7) 117-129.
Reynolds JW, 2014. A checklist by counties of earthworms (Oligochaeta: Lumbricidae, Megascolecidae and Sparganophilidae) in Ontario, Canada. In: Megadrilogica, 16 (10) 111-135.
Reynolds JW, Görres JH, Knowles ME, 2015. A checklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae and Megascolecidae) in the states of Maine, New Hampshire and Vermont, USA. In: Megadrilogica, 17 (9) 125-140.
Schermaier AF, 2013. Influence of earthworms on plant and soil invertebrate communities of the Cleveland Metroparks. MPhil Thesis., Cleveland, OH, USA: Cleveland State University.
Snyder B A, Callaham M A, 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), 349-358. http://www.springerlink.com/content/m422k472t0w4p574/ DOI:10.1007/s10530-010-9826-4
Stebbings JH, 1962. Additions to the Illinois and Missouri earthworm fauna. In: American Midland Naturalist, 67 (2) 504-505.
Tandy RE, 1969. The earthworm genus Pheretima, Kinberg, 1866, in Louisiana (Oligochaeta: Megascolecidae). DPhil Thesis., Baton Rouge, LA, USA: Louisiana State University and Agricultural and Mechanical College.
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11/07/17 Original text by:
Bruce A. Snyder, Georgia College & State University, Milledgeville, GA, USA
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