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New Asian pheretimoid "jumping earthworm" records (Oligochaeta: Megascolecidae) in Canada.

Authors:
  • Oligochaetology Laboratory, Kitchener, Ontario, Canada

Abstract and Figures

Asian pheretimoid "jumping worms" are found throughout the northeast and midwest United States but had previously only been documented once in Canada, in 2014. In summer 2021, reports were received of "unusual" earthworms in the gardens of private residences in southern Ontario, Canada. Following up on these reports, we provide five new collections of jumping worms including the previously documented Amynthas hilgendorfi (syn. Metaphire hilgendorfi) and Amynthas agrestis in addition to the first records of Amynthas tokioensis (syn. Metaphire levis) and Pithemera bicincta. This brings the total number of known earthworm species in Canada to 35. Jumping worms were collected primarily from household gardens but were also detected in a semi-natural ravine in Toronto, Ontario. Observations by landowners and the extent of invasion in the ravine support suspicions that undocumented jumping worm populations have already been present in Canada and Ontario in particular for some time. Further monitoring and research supported by community-science should be prioritized to determine the distribution and spread of jumping worms in Canada. Key words: Canada, Ontario, Oligochaeta, Megascolecidae, earthworms, Amynthas agrestis, A. hilgendorfi, A. tokioensis, Pithemera bicincta, distribution, new records.
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Printed in Canada ISSN 0380-9633
MEGADRILOGICA
Volume 26, Number 7, October 2021
NEW ASIAN PHERETIMOID "JUMPING EARTHWORM" RECORDS
(CLITELLATA: OLIGOCHAETA: MEGASCOLECIDAE) IN CANADA.
John Warren Reynolds
Oligochaetology Laboratory, 9-1250 Weber Street East, Kitchener, ON Canada N2A 4E1, and
Research Associate, New Brunswick Museum, Saint John, NB Canada E2K 1E5
(e-mail: john.w.reynolds1941@gmail.com ! https://orcid.org/0000-0001-8362-9071)
and
Michael J. McTavish
Mitacs Elevate Postdoctoral Fellow, Smith Forest Health Lab, Institute of Forestry and Conservation,
John H. Daniels Faculty of Architecture, Landscape and Design
University of Toronto, 33 Willcocks St, Toronto, ON Canada M5S 3E8
(e-mail: michael.mctavish@alum.utoronto.ca ! https://orcid.org/0000-0002-3343-8709)
ABSTRACT
Asian pheretimoid "jumping worms" are found throughout the northeast and midwest United States but
had previously only been documented once in Canada, in 2014. In summer 2021, reports were received of
"unusual" earthworms in the gardens of private residences in southern Ontario, Canada. Following up on these
reports, we provide five new collections of jumping worms including the previously documented Amynthas
hilgendorfi (syn. Metaphire hilgendorfi) and Amynthas agrestis in addition to the first records of Amynthas
tokioensis (syn. Metaphire levis) and Pithemera bicincta. This brings the total number of known earthworm species
in Canada to 35. Jumping worms were collected primarily from household gardens but were also detected in a
semi-natural ravine in Toronto, Ontario. Observations by landowners and the extent of invasion in the ravine
support suspicions that undocumented jumping worm populations have already been present in Canada and Ontario
in particular for some time. Further monitoring and research supported by community-science should be prioritized
to determine the distribution and spread of jumping worms in Canada.
Key words: Canada, Ontario, Oligochaeta, Megascolecidae, earthworms, Amynthas agrestis, A. hilgendorfi, A.
tokioensis, Pithemera bicincta, distribution, new records.
RESUME
Les « vers sauteurs » phérétimoïdes asiatiques se retrouvent à travers le nord-est et le centre-ouest des
États-Unis, mais avaient été documentés auparavant qu'une seule fois au Canada, en 2014. À l'été 2021, des rapports
ont été reçus concernant des vers de terre « inhabituels » dans les jardins de résidences privées du sud de l'Ontario,
Canada. Pour faire suite à ces rapports, nous avons répertorié cinq nouvelles collections de vers sauteurs, y compris
les espèces précédemment documentées, Amynthas hilgendorfi (syn. Metaphire hilgendorfi) et Amynthas agrestis,
en plus des premières observations d'Amynthas tokioensis (syn. Metaphire levis) et de Pithemera bicincta. Cela porte
à 35 le nombre total d'espèces de vers de terre connues au Canada. Les vers sauteurs ont été collectés principalement
dans des jardins familiaux, mais ont également été détectés dans un ravin semi-naturel à Toronto, en Ontario. Les
observations des propriétaires fonciers et l'étendue de l'invasion dans le ravin appuient les soupçons selon lesquels
des populations de vers sauteurs non documentées sont déjà présentes au Canada et en Ontario en particulier depuis
un certain temps. D'autres surveillances et recherches appuyées par la science communautaire devraient être
priorisées pour déterminer la répartition et la propagation des vers sauteurs au Canada.
Mots-clé: Canada, Ontario, Oligochaeta, Megascolecidae, vers de terre, Amynthas agrestis, A. hilgendorfi, A.
tokioensis, Pithemera bicincta, répartition, nouvelles observations.
84 New Pheretimoid earthworms in Canada
RESUMEN
Los 'lombrices de tierra saltarinas' feretimoides asiáticas se encuentran en todo el noreste y el medio oeste
de los Estados Unidos, pero anteriormente solo se habían registrado una vez en Canadá, en 2014. En el verano de
2021, se recibieron informes de lombrices de tierra 'no comunes' en los jardines de residencias privadas en el sur de
Ontario, Canadá. Siguiendo esta informacio'n, proporcionamos cinco nuevas colecciones de gusanos saltarines que
incluyen Amynthas hilgendorfi (sin. Metaphire hilgendorfi) y Amynthas agrestis previamente registrados, además
de las primeras citas de Amynthas tokioensis (sin. Metaphire levis) y Pithemera bicincta. Esto eleva el número total
de especies de lombrices de tierra conocidas en Canadá a 35. Las lombrices de tierra saltarinas se recolectaron
principalmente de los huertos familiares, pero también se detectaron en un barranco seminatural en Toronto, Ontario.
Las observaciones de los propietarios de tierras y el alcance de la invasión en el barranco respaldan las sospechas
de que las poblaciones de lombrices de tierra saltarinas no registradas aun ya estuvieron presentes particularmente
en Canadá y Ontario durante algún tiempo. Se debe priorizar el seguimiento y las investigaciónes adicionales
respaldados por la ciencia comunitaria para determinar la distribución y propagación de las lombrices de tierra
saltarinas en Canadá.
Palabras clave: Canadá, Ontario, Oligochaeta, Megascolecidae, lombrices de tierra, Amynthas agrestis, A.
hilgendorfi, A. tokioensis, Pithemera bicincta, distribución, nuevos registros.
ZUSAMMENFASSUNG
Asiatische Pheretimoid-"Springende Würmer" sind im gesamten Nordosten und Mittleren Westen der
Vereinigten Staaten zu finden, wurden in Kanada jedoch einmal im Jahr 2014 dokumentiert. Im Sommer 2021
gingen Berichte über "ungewöhnliche" Regenwürmer in den Gärten von Privathäusern im Süden der kanadischen
Provinz Ontario ein. Im Anschluss an diese Berichte stellen wir fünf neue Sammlungen von Springwürmern zur
Verfügung, darunter die bereits dokumentierten Amynthas hilgendorfi (syn. Metaphire hilgendorfi) und Amynthas
agrestis zusätzlich zu den Erstnachweisen von Amynthas tokioensis (syn. Metaphire levis) und Pithemera bicincta.
Damit steigt die Gesamtzahl der bekannten Regenwurmarten in Kanada auf 35. Springwürmer wurden hauptsächlich
in Gärten gesammelt, wurden aber auch in einer naturnahen Schlucht in Toronto, Ontario, entdeckt. Beobachtungen
von Grundbesitzern und das Ausmaß der Invasion in der Schlucht stützen den Verdacht, dass in Kanada und Ontario
bereits seit einiger Zeit undokumentierte Springwurmpopulationen vorkommen. Weitere Überwachung und
Forschung, die von Community-Science unterstützt wird, sollte Priorität haben, um die Verbreitungvon
Springwürmern in Kanada zu bestimmen.
Schlüsselworte: Kanada, Ontario, Oligochaeta, Megascolecidae, Regenwürmer, Amynthas agrestis, A. hilgendorfi,
A. tokioensis, Pithemera bicincta, geografische Verteilung, neue Datensätze.
INTRODUCTION
The introduction of jumping and thrashing
earthworms, also commonly called 'crazy worms' – a
group of pheretimoid species (Clitellata: Oligochaeta:
Megascolecidae) from China, Japan and Korea, have
increasingly become an ecological, environmental and
conservation issue in forest and urban landscapes in
many countries (Chang et al. 2016, 2021). When I
(JWR) conducted research in the southeastern United
States in the late 1960s and 1970s, these species were
just beginning to be recorded in nature as far north as
Tennessee (Reynolds, 1978). Today they are frequently
found in the American northeast and midwest (Callaham
et al., 2003; Frelich et al., 2006; Nuzzo et al, 2009;
Szlávecz et al., 2011; Görres and Melnichuk, 2012;
Greiner et al., 2012; Gorsuch and Owen, 2014; Görres et
al., 2014a, 2014b; Schult et al., 2016; Qui and Turner,
2017; Moore et al., 2018; O'Keefe and McCulloh, 2021).
The first report of pheretimoid earthworms in
Canada was on the Ojibway Prairie in Essex County,
Ontario just across the Detroit River from Detroit,
Michigan, USA (Reynolds, 2014). In the summer of
2021, reports of "unusual" earthworms in the private
gardens of residents were brought to the attention of
MJM through the Master Gardeners of Ontario Program.
Where possible, site visits were made to verify these
reports and collect earthworm specimens. This is the
first record of collections in Canada, and Ontario in
particular since the original paper, but it has been
expected for a number of years that jumping worms were
already in many other locations in Ontario (Reynolds
and Dobson, 2021).
85
MEGADRILOGICA
COLLECTIONS
Identification of species in collections A-F,
below were undertaken with the aid of keys and
taxonomic information presented in Chang et al. (2016),
Gates and Reynolds (2017), and Reynolds (1978). The
age classification formula (as described in Reynolds,
1977) for juveniles-aclitellate adults-clitellate adults
[as a polyminimal, x-x-x], immediately follows the
scientific names of the species identified in each of the
collections, below.
All earthworms appeared superficially similar
in the field and displayed characteristics consistent with
one or more species of jumping worm, i.e., pale grey
colouration, milky annular clitellum, thrashing behaviour
and autotomy upon being disturbed. Samples were
euthanized in 95% ethanol, refrigerated (for 1–3 days
prior to processing) to limit degradation, transferred to
10% formalin for 24 hours, then stored in 95% ethanol.
Previous collections
A. Essex County, Ontario
1. Ojibway Prairie Provincial Nature Reserve, 5200
Matchette Rd, Windsor, in a large woodchip pile, 28
June 2014. Coll. John W. Reynolds (Reynolds, 2014).
Amynthas agrestis (Goto and Hatai, 1899) 0-0-20+
2. Ojibway Prairie Provincial Nature Reserve, 5200
Matchette Rd, Windsor, under a large piece of plywood
in open grassland, 28 June 2014. Coll. John W.
Reynolds (Reynolds, 2014).
Amynthas hilgendorfi (syn. Metaphire hilgendorfi)
(Michaelsen, 1892) 0-0-5
New collections
B. Wheatley (Kent Co. on Essex Co. Line)
Private rural residence, in a mulched garden, 26 August
2021. High density of casts and earthworms. First
observed by residents in June 2021, source of
introduction unknown. Potential for spread to adjoining
agricultural fields. Coll. Michael J. McTavish.
Amynthas hilgendorfi 0-0-3
Amynthas tokioensis (Beddard, 1892) (syn. Metaphire
levis (Goto and Hatai, 1899)) 0-0-1
C. Dundas (Hamilton) (Wentworth Co.)
Private urban residence, beneath a pile of old
newspapers and along the wood-framed edges of a small
front yard mulched garden, 27 August 2021. Low
density population with scattered casts. First observed by
residents in August 2021, possible introduction through
recent plant and soil additions. Coll. Michael J.
McTavish.
Amynthas agrestis 0-0-1
Amynthas tokioensis 0-0-1
D. East York (Toronto) (York Co.)
Private urban residence, in front, side, and back of house
gardens and backyard greenhouse, 25 August 2021.
Moderate density population with scattered casts.
Unconfirmed observations by residents dating back to
2016, source unknown. Potential for spread to nearby
ravines and through movement of plants to other
properties. Coll. Michael J. McTavish.
Amynthas hilgendorfi 0-1-4
Pithemera bicincta (Perrier, 1875) 0-0-1
E. Toronto (York Co.)
Private urban residence, in back garden, 20 September
2021. Low density population with scattered casts.
Unconfirmed observations by residents dating back to
2017 of unexplained plant decline and behaviourally
atypical earthworms. Possible sources include
contiguous mulched gardens in neighbouring properties
or damp top dressing brought to the property. Potential
for spread to nearby ravines (including Collection F,
below). Coll. Michael J. McTavish.
Amynthas agrestis 0-0-6
F. Blythwood Ravine (Toronto) (York Co.)
Public ravine park surrounded by private homes,
alongside a foot path ~200 m from the ravine access
point, 20 September 2021. High density population with
abundant casts adjacent to foot path and deeper into the
woodlot interior. Possible sources include gardens of
private residences uphill and backing onto the ravine
(including Collection E, above). Depth and extent of
casts suggests the population may have been present for
at least several years. Coll. Michael J. McTavish.
Amynthas hilgendorfi 0-0-3
S. Supplemental. Additional credible, but unconfirmed
observations were submitted from the Niagara Region
and other sites in York County (see So in Fig. 1).
Residents reported a small number of suspicious looking
earthworms from nursery pots of recently purchased
plants but had destroyed and disposed of the individuals
before samples could be collected.
86 New Pheretimoid earthworms in Canada
Fig. 1. Location of pheretimoid collections in Ontario, Canada. A – Ojibway Prairie Provincial Nature Reserve, B
Wheatley, CDundas (Hamilton), DEast York (Toronto), E Toronto, F – Blythwood Ravine (Toronto), S – Fonthill
(Niagara Region) and Toronto (York Region) suspected species, unconfirmed. Species collection sites: M Amynthas
agrestis, M Amynthas hilgendorfi, M Amynthas tokioensis, M Pithemera bicincta.
DISCUSSION
Prior to the collections discussed herein, 33
earthworm species had been recorded in Canada
(Reynolds, 2018, 2021), which included the first record
of Asian earthworms in Canada (Amynthas hilgendorfi,
Amynthas agrestis), collected at the Ojibway Prairie
Provincial Nature Reserve in 2014 (Reynolds, 2014).
This present paper records the second time these
earthworms are reported from Canada and Ontario and
adds the first records of Amynthas tokioensis (syn.
Metaphire levis) and Pithemera bicincta, bringing the
total number of known earthworm species in Canada to
35 (Fig. 1).
Of the newly reported species in Canada,
Amynthas tokioensis has been previously reported from
the United States: Connecticut (Gates, 1966), Delaware,
Illinois, Maryland, Wisconsin (Dávalos et al., 2015),
Florida (Reynolds, 1994b), Georgia (Reynolds, 2009),
New Jersey (Davies, 1954), New York (Gates, 1954),
North Carolina (Reynolds, 1994a), Pennsylvania
(Bhatti, 1954), Tennessee (Reynolds, 1978) and
Vermont (Chang et al., 2016). Although early reports
have been from the eastern United States, the spread of
this species in the southeastern and midwestern U.S.
states is the result of activities in horticulture
(gardening, mulching, potted plantings, contaminated
soil discard, etc.) and bait cast off (Reynolds and
Dobson, 2021).
Chang et al. (2017) found that sympatric
occurrence of at least two, and often all three species
e.g., A. agrestis, A. hilgendorfi and A. tokioensis is
more common than having only one species. In
addition, A. tokioensis was dominant in many of these
earthworm communities.
87
MEGADRILOGICA
Pithemera bicincta has been reported from the
states of Florida (Gates, 1982), Georgia (Gates, 1982)
and Maine (Gates, 1963). The distribution possibilities
were discussed under Amynthas tokioensis above.
Consistent with the association of jumping
worms and horticultural activities (Reynolds and
Dobson, 2021), these new collections were made
primarily from the gardens (often mulched) of private
residences. In addition to being a source of concern for
the aesthetics and productivity of gardens (Johnson et
al. 2021), jumping worms in gardens may also
contribute to spread into nearby natural areas. Several
of the collections were from Toronto, Ontario, and the
surrounding area, where gardens may be a major
distribution pathway into the nearby network of semi-
natural and ecologically sensitive ravines that run
throughout the city, areas that should be prioritized for
future research.
While the only prior collections of jumping
worms in Canada were made in Essex County in 2014
(Reynolds, 2014), private residents provided
unconfirmed reports in 2021 of jumping worm activity
dating back up to five years, supporting the suspicion
that other populations had been present but
undocumented in Canada. This is further substantiated
by the quantity of casts present in the Blythwood
Ravine (Toronto, ON) suggesting the presence of a
large and established population. Given the challenges
of monitoring a large geographic area by a limited
number of professional researchers, there have been
growing calls to recruit gardeners, amateur naturalists,
and others to help submit and track pheretimoid
earthworm observations (Johnson et al., 2021;
Reynolds and Dobson, 2021). These community-
science efforts were key to our new collections and may
play an essential role going forwards in monitoring the
distribution and spread of jumping worms in Canada.
ACKNOWLEDGEMENTS
We are grateful to Wilma M. Reynolds of the
Oligochaetology Laboratory and Mark J. Wetzel of the
Illinois Natural History Survey, Champaign, Illinois for
reviewing the manuscript, their comments and
suggestions. Also, we thank Dr. Josef H. Görres,
University of Vermont for significant suggestions. We
thank Bruce Miller for cartographic assistance and
Cathy Kavassalis of the Master Gardener Program of
Ontario for bringing the observations to our attention.
The authors also acknowledge Dr. Catalina C. de
Mischis, Universidad Nacional de Córdoba (Argentina),
Dr. Jean-Marc Gagnon, Canadian Museum of Nature,
and Dr. Josef H. Görres, University of Vermont for
checking our translations of the abstracts and key
words.
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Reynolds, J.W. 2021. The Earthworms (Lumbricidae,
Megascolecidae and Sparganophilidae) in
Canada. Ottawa: Canada Food Inspection
Agency, ii + 185 pp. (in press)
Reynolds, J.W. and A.M. Dobson. 2021. Earthworms
of Ontario. Ontario Horticultural Association,
Trillium, Summer 2021: 10–13.
Schult, N., K. Pittenger, S. Dávalos and D. McHugh.
2016. Phylogeographic analysis of invasive
Asian earthworms (Amynthas) in the northeast
United States. Invertebr. Biol. 135: 314–327.
doi:10.1111
Szlávecz, K., M. McCormick, L. Xia, J. Saunders, T.
Morcol, D. Whigham, T. Filley and Cs.
Csuzdi. 2011. Ecosystem effects of
non-native earthworms in Mid-Atlantic
deciduous forests. Biological Invasions 13:
1165–1182.
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Web Site for the Journal Megadrilogica:
https://www.inhs.illinois.edu/people/mjwetzel/megadrilogica/
Web Site for Nomenclatura Oligochaetologica – Editio Secunda
A catalogue of names, descriptions, and type specimens of the Oligochaeta:
https://www.inhs.illinois.edu/people/mjwetzel/nomenoligo
Please note:
The website addresses (URLs) for the journal Megadrilogica, its index, and for Nomenclatura Oligochaetologica will
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site addresses.
... The first Canadian report was in Essex County, Ontario at the Ojibway Prairie Bioblitz in June 2014 when I collected Amynthas agrestis and Metaphire hilgendorfi (as Amynthas hilgendorfi) (Reynolds, 2014). It was another seven years before the next report in the scientific literature when Reynolds and McTavish (2021) reported these two previous species and two additional species (Amynthas tokioensis and Pithemera bicincta) (Fig. 1). Reynolds and McTavish, 2021). ...
... It was another seven years before the next report in the scientific literature when Reynolds and McTavish (2021) reported these two previous species and two additional species (Amynthas tokioensis and Pithemera bicincta) (Fig. 1). Reynolds and McTavish, 2021). The next report came from New Brunswick when McAlpine et al. (2022) reported Amynthas minimus, A. tokioensis and Metaphire hilgendorfi from York County (Fig. 2). ...
... In natural outside habitats it has been found in the United States: Florida (Gates, 1937), New Jersey (Davies, 1954), New York (Gates, 1954) and Pennsylvania (Bhatti, 1965), Georgia, North Carolina and Tennessee , Delaware, Illinois and Maryland (Dávalos et al., 2015),Vermont (Chang et al., 2016), Wisconsin (Dávalos et al., 2015). Canada: Ontario (Reynolds and McTavish, 2021), New Brunswick (McAlpine et al., 2022). Diagnosis: (after Sims and Easton, 1972): ...
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This paper discusses the initial spread of pheretimoid earthworms in the USA and recently in Canada. There are now five species recorded from Canada, but that number may eventually increase. A synonomy, description, reproduction, distribution, and photograph is presented for each species, plus a glossary and key to their identification. Key Words: Canada, Oligochaeta, Megascolecidae, distribution, Amynthas agrestis, Amynthas minimus, Amynthas tokioensis, Metaphire hilgendorfi, Pithemera bicincta. RÉSUMÉ Cet article s'intéresse à la propagation initiale des vers de terre phérétimoïdes aux États-Unis et, plus récemment, au Canada, où cinq espèces sont actuellement recensées (ce nombre, à terme, est appelé à augmenter). Pour chaque espèce présentée, en plus d'un glossaire et d'une clé d'identification, sont indiqués les synonymes, la description, la reproduction et la répartition. Mots-clé: Canada, Oligochaeta, Megascolecidae, répartition, Amynthas agrestis, Amynthas minimus, Amynthastokioensis, Metaphire hilgendorfi, Pithemera bicincta.
... This suggests that climate change, underway and pending, may have a significant impact on earthworm diversity and abundance in North America, including that of pheretimoids. Occurences are from Chang et al. (2021), Reynolds (2010Reynolds ( , 2014 and Reynolds and McTavish (2021) and those newly reported here for Atlantic Canada. McAlpine et al. (2020) sampled 34 sites in New Brunswick for earthworms across a variety of habitats in 2014 and 2018 in the 10, 697 ha Grand Lake Protected Natural Area (GL PNA). ...
... Limited data suggests there has been relatively little penetration by European lumbricids into the mixed and conifer forests of New Brunswick, outside of river shorelines, areas adjacent to woods roads, and the disturbed soils associated with residences (McAlpine and Reynolds 2019; McAlpine et al. 2019). Nonetheless, although most of the Ontario sites reported by Reynolds and McTavish (2021) were private gardens, they report evidence of spread into nearby natural woodlands. The establishment of pheretimoids in urban and suburban habitats in Atlantic Canada should therefore not be overlooked as a platform for dispersal into forested sites in the region. ...
... Johnson et al. (2021) found that public reporting allowed them to track the expanding distribution of pheretimoids in the northeastern United States. Likewise, Reynolds and McTavish (2021) relied on reports from gardeners at private residences to document new occurrences of pheretimoids in southern Ontario. Chang et al. (2021) have also promoted the use of coordinated community science projects in tracking the spread of pheretimoids and identifying knowledge gaps and possible management strategies. ...
Article
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First reports of pheretimoid earthworms in Atlantic Canada, from a suburban site in south-central New Brunswick, are documented. Pheretimoids were present at high m-2 densities under hardwood cover, estimated at ~ several hundred in the small area sampled. Preliminary samples collected at the site in October 2021 suggest that Metaphire hilgendorfi, followed closely by Amynthas tokioensis, were dominant, while A. minimus was rare. The latter is the first report of A. minimus from Canada. Associated lumbricids, present in much smaller numbers and in descending order of abundance, included Lumbricus rubellus, Dendrobaena octaedra, and Aporrectodea tuberculata. We suggest that pheretimoid earthworms are probably more widespread in the Atlantic Canadian region than this first report would indicate. Further survey, supported by public education and reporting, will be required to assess the full ecological impact of invasive Asian pheretimoids in eastern Canada. Key words: New Brunswick, distribution, Amynthas minimus, Amynthas tokioensis, Metaphire hilgendorfi
... This number was based on an earlier paper (Reynolds, 2018) where collection specifics were not included. Since the publication of those books, several additional species have been recorded in Canada (Reynolds and McTavish, 2021;. The following chronology reveals the species in Canada, and where and when they were reported. ...
... Graff, 1957; this epigeic species was collected by Virginia F. Zoll on 10 August 2017 during the NB Biota on the Spednic Lake Protected Natural Area, (York Co.), New Brunswick, and first reported without details by Reynolds et al. (20 18) and subsequently with a habitat description in . (Perrier, 1872); this epiendogeic species was collected by Michael McTavish on 25 August 2021 from a house garden in East York (Toronto) (York Co.), Ontario and reported by Reynolds and McTavish (2021). (Beddard, 1892); this epiendogeic species was collected by Michael McTavish on 27 August 2021 from a mulched garden in Dundas (Hamilton) (Wentworth Co.), Ontario and reported by Reynolds and McTavish (2021). ...
... (Perrier, 1872); this epiendogeic species was collected by Michael McTavish on 25 August 2021 from a house garden in East York (Toronto) (York Co.), Ontario and reported by Reynolds and McTavish (2021). (Beddard, 1892); this epiendogeic species was collected by Michael McTavish on 27 August 2021 from a mulched garden in Dundas (Hamilton) (Wentworth Co.), Ontario and reported by Reynolds and McTavish (2021). (Horst, 1893); this epi-endogeic species was collected by Lawrence R. Manzer and Donald F. McAlpine Burton (York Co.), New Brunswick on 25 October 2021 and reported by ; subsequent research has produced this species, collected by Esther Tang (University of Toronto) from Blythwood Ravine/ Sherwood Park (Toronto) (York Co.), Ontario 12 October 2021 (unpublished data). ...
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Recent official documentation of Eisenia andrei Bouché, 1972 in Canada brings the number of known terrestrial earthworm species in Canada to 37. This paper details the chronological record of the 37 species found in Canada by presenting the date collected, the location and collector, where possible. In addition, the name of the species when collected if different than the currently used nomenclature is included. Six of these species have been reported in Canada for the first time in the past five years, and four of them in the past year. Key words: Canada, earthworms, chronological records, nomenclature, Eisenia andrei.
... Earthworm communities ebb and flow with the changing precipitation patterns (Tondoh, 2006;Eisenhauer et al., 2019), but if the moisture is not available to the earthworms, this pattern may not be sustainable. Therefore, soil moisture tension gives a better indication of earthworm's ability to reproduce and feed in a given habitat (Reynolds, 2021). Biological invasions can have strong impacts on biodiversity as well as ecosystem functioning and introduced below ground organisms (e.g., earthworms) may have strong effects. ...
... In the past 50 years, they have spread into the midwestern and the northeastern American soils (Reynolds, 2011c(Reynolds, , 2010c. It was not until recently that I recorded the first populations of these earthworms in Canada (Reynolds, 2014a) and then again in 2021 (Reynolds and McTavish, 2021(McAlpine et al., 2022. I may not live to see it happen, but as the soils in Canada warm, and if precipitation in the form of increased rain and vegetation diversifies, one day earthworms will become a major component of the soil fauna in the Canadian far north. ...
... ; Dymond et al. (1997);Eisenhauer et al. (2007); McAlpine and Reynolds (2018, 2020; McKey-Fender et al. (1994); McLean and Parkinson (1997); Moore and Reynolds (2003); Moore et al. (2009Moore et al. ( , 2018Moore et al. ( , 2022, Panesar et al. (2000); Reynolds (1975aReynolds ( -d, 1976aReynolds ( ,b, 1977a, Reynolds and McTavish (2021), Reynolds and Reynolds (1992), Reynolds et al. (2014Reynolds et al. ( , 2015Reynolds et al. ( , 2019, Welke and Parkinson (2003); Whalen (2004), Wironen and Moore (2006). ...
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Canadian forests cover 43% of its landmass, which represent 10% of the world's overall forested areas and 30% of the world's boreal forests. There are eight forest regions in Canada plus grassland and tundra. Each forest region is represented by a description, map, earthworm diversity and function. Four appendices summarize: the distribution and diversity earthworms in Canadian forests according to dominant tree species and the Canadian and American soil type systems; earthworms recorded from the eight Canadian forest regions; earthworms recorded from the 10 dominant forest soils; and descriptions of the soil type profiles in Canada from which earthworms have been recorded. A map of the soil orders in Canada is presented. A table summarizes the mean values for morpho-anatomical traits characterizing the most frequent ecological categories with which earthworms are associated. The discussion considers arctic earthworm migration and climate change. Key words: Canada, Oligochaeta, Lumbricidae, Megascolecidae, forest regions, earthworms, distribution, soils, Arctic migration, climate change.
... In the introduction section of his paper, Reynolds (2017) also briefly summarized papers discussing the recent invasion of Asian earthworm species (representing the genera Amynthas, Metaphire, Pheretima, Pithemera and Polypheretima, all in the family Megascolecidae) into North America. Moore et al. (2018) and Reynolds and McTavish (2021) provided updated records for species in these and other pheretimoid genera reported from Canada and elsewhere in North America. ...
Article
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In this paper we present the results of a preliminary inventory of the terrestrial oligochaetes, or earthworms, of the Nachusa Grasslands preserve in Lee and Ogle Counties, Illinois, USA, based upon specimens collected from 23 sites in April and October 2015. In total, eight species representing four genera in the family Lumbricidae are reported, including the collection of Bimastos welchi (Smith, 1917) from one site-a new state record for Illinois, and species native to North America. The other seven species collected during this study-Aporrectodea rosea (Savigny, 1826), Aporrectodea trapezoides (Dugès, 1828), Aporrectodea tuberculata (Eisen, 1874), Aporrectodea turgida (Eisen, 1873), Lumbricus rubellus Hoffmeister, 1843, Lumbricus terrestris Linnaeus, 1758, and Octolasion tyrtaeum (Savigny, 1826)-are considered to be introductions, and are relatively common elsewhere in Illinois and in North America.
... In the introduction section of his paper, Reynolds (2017) also briefly summarized papers discussing the recent invasion of Asian earthworm species (representing the genera Amynthas, Metaphire, Pheretima, Pithemera and Polypheretima, all in the family Megascolecidae) into North America. Moore et al. (2018) and Reynolds and McTavish (2021) provided updated records for species in these and other pheretimoid genera reported from Canada and elsewhere in North America. ...
Article
Full-text available
In this paper we present the results of a preliminary inventory of the terrestrial oligochaetes, or earthworms, of the Nachusa Grasslands preserve in Lee and Ogle Counties, Illinois, USA, based upon specimens collected from 23 sites in April and October 2015. In total, eight species representing four genera in the family Lumbricidae are reported, including the collection of Bimastos welchi (Smith, 1917) from one site – a new state record for Illinois, and species native to North America. The other seven species collected during this study – Aporrectodea rosea (Savigny, 1826), Aporrectodea trapezoides (Dugès, 1828), Aporrectodea tuberculata (Eisen, 1874), Aporrectodea turgida (Eisen, 1873), Lumbricus rubellus Hoffmeister, 1843, Lumbricus terrestris Linnaeus, 1758, and Octolasion tyrtaeum (Savigny, 1826) – are considered to be introductions, and are relatively common elsewhere in Illinois and in North America. Key words: Illinois, Oligochaeta, Lumbricidae, earthworms, distribution, checklist, new state records.
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The invasive Asian earthworms, Amynthas tokioensis and A. agrestis , have been successful in entering North American forests in recent decades, with significant damage to both soils and above-ground environments. This success could be driven in part by a polyploid genetic system and parthenogenetic reproduction, often suggested as benefits for invasive species. Therefore, we assessed the genetic population structure, genetic diversity, and reproductive system of both species using morphological traits and panels of microsatellite markers. A total of 216 A. tokioensis and 196 A. agrestis from six sites in Vermont USA were analyzed. Although all worms were morphologically hermaphroditic, all the A. agrestis lacked the male pore (the structure allowing pass of sperm between individuals), and only 19% of the A. tokioensis possessed the male pore. All A. tokioensis earthworms were triploid (scored for three alleles for at least 1 locus, and usually several), and A. agrestis was a mix of triploid and diploid individuals. Notable was the high proportion (80%) of A. agrestis earthworms that were diploid at one site. There was clearly clonal reproduction, with identical seven- locus genotypes observed for earthworms from each site, with as many as 45 individuals with the identical genotype at one site. However, the earthworms were also genetically diverse, with 14 genotypes observed for A. tokioensis and 54 for A. agrestis , and with many singleton genotypes (a single individual). Most genotypes (71% for A. tokioensis and 92% for A. agrestis ) were found at a single site. The greatest number of genotypes was found at a commercial nursery where fully 23/26 A. agrestis earthworms were singleton genotypes. As expected for the pattern of private clone alleles at sites, several measures of geographic genetic differentiation were positive, and as expected for triploid systems, an AMOVA analysis showed high within-individual genetic diversity. The paradox of clear clonal reproduction, but with a great number of genotypes for each species, and the mix of triploid and diploid individuals could be explained if the worms have been sexually reproductive, with the switch to the uniparental system only recently (or even if sexual reproduction is episodic). Last, a large number of microsatellite loci were recovered for each species and there sequence and suggested PCR primers are provided for free use by other researchers.
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Earthworms are considered the best ecological engineers and they are distributed across various ecotypes around the world. The diverse habitats of earthworms make them interesting to explore. In addition, this review discusses the availability of various species of earthworms in relation to the various land use systems in various countries and continents. The structure of the population of earthworms s varies by land use patterns in different agro-climatic regions of the country. Biodiversity is an essential determinant of an ecosystem and earthworms are generally classified into five ecological types, i.e. anecic, corticolous, endogeic, epigeic and limicolous. The seven continents of the world belong to various ecological zones, including various agroecosystems and include a great diversity of earthworms. Researchers examined the paucity of research on the interacting effects of several global changes affecting earthworms in order to understand the various factors related to the diversity of earthworms. Keywords: Diversity, distribution, earthworms, continents, ecosystems.
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In a country where native earthworms are mostly inconspicuous and make up 27% of the species, the vast majority are introduced aliens from Europe (67%) or the Orient (6%). Most of the introduced European species are now ubiquitous over large areas of Canada, although Except for two species, the native earthworms are restricted to British Columbia and the Yukon. Based on fieldwork covering more than 50 years, and the author’s knowledge of the fauna, this book describes and illustrates the 33 species reported in Canada. For each species, genus and family, - synonymy, type species, diagnosis, biology, reproduction, climatic zone, ecological type, origin, range, and North American distribution are described. The book also includes an extensive literature section, illustrated glossary and a key to the 33 species.
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The invasion of jumping worms, a small group of pheretimoid earthworm species from Asia, has increasingly become an ecological, environmental and conservation issue in forest ecosystems and urban-suburban landscapes around the world. Their presence is often noticed due to their high abundance, distinctive “jumping” behavior, and prominent granular casts on the soil surface. Although they are known to affect soil carbon dynamics and nutrient availability, no single paper has summarized their profound impacts on soil biodiversity, plant community, and animals of all trophic groups that rely on soil and the leaf litter layer for habitat, food, and shelter. In this study, we summarize the biology, invasion, and ecological impacts of invasive jumping worms across North America. We highlight potential impacts of this second wave of earthworm invasion, contrast them with the preceding European earthworm invasion in temperate forests in North America, and identify annual life cycle, reproductive and cocoon survival strategies, casting behavior and co-invasion dynamics as the key factors that contribute to their successful invasion and distinct ecological impacts. We then suggest potential management and control strategies for practitioners and policy makers, underscore the importance of coordinated community science projects in tracking the spread, and identify knowledge gaps that need to be addressed to understand and control the invasion.
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Although non-native earthworms are known to drastically change forest soils and understory plant communities, comparatively little is known about how the introduction of different earthworm groups may alter perennial plant functioning (i.e., physiological, morphological, and/or anatomical traits), or how these dynamics may vary between different earthworm groups. Our objective was to assess whether a recent replacement of European earthworms by Amynthas spp. (i.e., Asian “jumping worms”) in a Central Hardwoods forest of the Upper Midwest impacts patterns of water use in dominant overstory Acer saccharum Marshall (sugar maple). We found that (1) soil water content was often greater in Amynthas sites, (2) compared to those growing in sites with only European earthworms, A. saccharum growing with Amynthas had higher midday leaf water potential values (a measure of greater plant water status) early in 2017 but lower midday leaf water potential by the end of 2018, and (3) A. saccharum from Amynthas sites had fewer, but wider xylem conduits than trees from European sites, but this trend was not significant and was not associated with differences in stem hydraulic functioning between the two groups. Overall, these results suggest that Amynthas invasions may not impact the overstory of A. saccharum—dominated forests within the Upper Midwest, at least early during invasions of forests colonized by existing earthworms or during growing seasons with above-average precipitation. Future work should investigate the impacts of different earthworm groups on tree water and nutrient relations over longer periods of colonization and in different tree age classes.
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Exotic species invasions are among the most significant global-scale problems caused by human activities. They can seriously threaten the conservation of biological diversity and of natural resources. Exotic European earthworms have been colonizing forest ecosystems in northeastern United States and southern Canada since the European settlement. By comparison, Asian earthworms began colonizing forests in the northeastern United States more recently. Since Asian species have biological traits compatible with a greater potential for colonization and disturbance than some European species, apprehension is growing about their dispersal into new territories. Here we review the extent of the current northern range of Asian earthworms in northeastern North America, the factors facilitating or limiting their propagation and colonization, and the potential effects of their invasion on forest ecosystems. Data compilation shows that Asian earthworms are present in all northeastern American states. So far, only one mention has been reported in Canada. Data confirm that their distribution has now reached the Canadian border, particularly along the Michigan–Ontario, New York–Ontario, Maine–New Brunswick, and Vermont–Québec frontiers. Studies report that the presence of Asian earthworms is strongly associated with human activities such as horticulture, vermicomposting, and the use of worms as fish bait. Some climatic (temperature, soil moisture) and edaphic (soil pH) factors may also influence their distribution. Controlling their dispersal at the source is essential to limiting their spread, as there is currently no effective way to eradicate established earthworm populations without unacceptable nontarget effects. Proposed management options in the United States include the prohibition of fish bait disposal and better management of the international trade of horticultural goods, commercial nurseries, and vermicomposting industries. We conclude that although regulations and awareness may delay their expansion, Asian earthworms are likely to spread further north into Canada. They are expected to cause important changes to biodiversity and dynamics of the newly invaded forest ecosystems.
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Earthworm invasions are one of the most serious causes of ecological deterioration in the temperate deciduous forests of North America. Non-native earthworms impact understory vegetation, leaf litter layer, carbon dynamics, nutrient availability, and the associated food webs. Here we report a significant status change and confirm expansion of known range of Amynthas agrestis, one of the most serious invasive species in North America, and two of its close relatives, A. tokioensis and Metaphire hilgendorfi. The three species have never been confirmed to co-occur in North American ecosystems. We examined 1760 earthworms collected from 30 sites across northeastern USA, and identified them using a new morphological key. Our data show that sympatric occurrence of at least two, and often all three, species is more common than having only one species. In addition, A. tokioensis was dominant in many of these earthworm communities. The status change in species composition from only one species to two or three and the shift in dominance are most likely caused by previous incorrect species identification. Our results support expansion of known range of A. tokioensis and M. hilgendorfi northward and westward into states with colder winters. This range expansion may have taken place alongside that of A. agrestis in the last 10–20 years, but has long been overlooked. Altogether, results highlight an urgent need for correct species identification. The recognition of an expanding multi-species system represents a unique opportunity to further evaluate complex interactions among co-invading and resident species, and to investigate whether interspecific interactions have unexpected non-additive impacts on ecological processes.
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Phylogeographic studies are useful in reconstructing the history of species invasions, and in some instances can elucidate cryptic diversity of invading taxa. This can help in predicting or managing the spread of invasive species. Among terrestrial invasive species in North America, earthworms can have profound ecological effects. We are familiar with the centuries-old invasions of European earthworms (Lumbricidae) and their impacts on nutrient cycling in soils. More recent invasions by Asian earthworms of the family Megascolecidae are less fully understood. We used data for two mitochondrial gene fragments, cytochrome oxidase I (COI) and 16S rRNA, to examine the relationships among populations of Asian earthworms in the megascolecid genus Amynthas in the northeast United States. Recent reports have indicated that one species in particular, Amynthas agrestis, is having detrimental effects in mixed forest ecosystems, and we were interested in understanding the invasion history for this species. We were surprised to discover three divergent mitochondrial lineages of Amynthas occurring sympatrically in upstate New York. Given the gap between intra- and inter-lineage sequence divergences, we propose that these three lineages represent cryptic species of Amynthas, one of which is A. agrestis. For each of the three lineages of Amynthas, we observed shared haplotypes across broad geographic distances. This may reflect common origins for populations in each lineage, either by direct routes from native ranges or through post-introduction spread by natural dispersal or human-mediated transport within North America. Management efforts focused on horticultural imports from Asia, commercial nurseries within the USA, and on prohibition of bait disposal may help to reduce the further invasion success of Amynthas.
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The invasion of the pheretimoid earthworms in North America, especially the genera Amynthas and Metaphire, has raised increasing concerns among ecologists and land managers, in turn increasing the need for proper identification. However, the commonly used keys to this group are more than 30 years old with outdated taxonomic information and are based primarily on internal morphology. The requirement of significant amount of taxonomic expertise and dissection, even from the first entry of the key, has prevented broader use of these keys. As a result, many publications in the United States have used Amynthas spp. to represent the group without identifying the species. We present here a new key and diagnoses for the 16 pheretimoid earthworm species recorded in North America north of Mexico, including four genera: Amynthas (10), Metaphire (4), Pithemera (1), and Polypheretima (1). The descriptions were based on published records with modifications following inspection of specimens archived at the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Photos of external and internal characters, including male pores, spermathecal pores, genital markings, spermathecae, prostate glands, and intestinal caeca, are presented to help identification. A summary of current knowledge about the ecology and historical context is provided for each species. We also highlight the previously overlooked, and potentially common and widespread co-occurrence of three species-A. agrestis, A. tokioensis, and M. hilgendorfi-and point out that many recent claims of invasion of A. agrestis need to be re-evaluated for potential misidentification.
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Effects of invasive European earthworms in North America have been well documented, but less is known about ecological consequences of exotic Asian earthworm invasion, in particular Asian jumping worms (Amynthas) that are increasingly reported. Most earthworm invasion research has focused on forests; some Amynthas spp. are native to Asian grasslands and may thrive in prairies with unknown effects. We conducted an earthworm-addition mesocosm experiment with before–after control-impact (BACI) design and a complementary field study in southern Wisconsin, USA, in 2014 to investigate effects of a newly discovered invasion of two Asian jumping worms (Amynthas agrestis and Amynthas tokioensis) on forest and prairie litter and soil nutrient pools. In both studies, A. agrestis and A. tokioensis substantially reduced surface litter (84–95 % decline in foliage litter mass) and increased total carbon, total nitrogen, and available phosphorus in the upper 0–5 cm of soils over the 4-month period from July through October. Soil inorganic nitrogen (ammonium– and nitrate–N) concentration increased across soil depths of 0–25 cm, with greater effects on nitrate–N. Dissolved organic carbon concentration also increased, e.g., 71–108 % increase in the mesocosm experiment. Effects were observed in both forest and prairie soils, with stronger effects in forests. Effects were most pronounced late in the growing season when earthworm biomass likely peaked. Depletion of the litter layer and rapid mineralization of nutrients by non-native Asian jumping worms may make ecosystems more susceptible to nutrient losses, and effects may cascade to understory herbs and other soil biota.