Royal Ontario Museum
Ufs Sciences
Miscellaneous Publications

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LIFE SCIENCES MISCELLANEOUS PUBLICATIONS
ROYAL ONTARIO MUSEUM

john w Reynolds The Earthworms

Illustrated hv /T i • • i

daniel l dindal (Lumbncidae

and Sparganophilidae)
of Ontario

Publication date: 15 June 1977

isbn 0-88854-191-0
issn 0082-5093

Suggested citation: Life Sci. Misc. Pub.. R. Ont. Mus.

ROYAL ONTARIO MUSEUM
PUBLICATIONS IN LIFE SCIENCES

The Royal Ontario Museum publishes three series in the Life Sciences:

life sciences contributions, a numbered series of original scientific publications, in-
cluding monographic works.

LIFE sciences occasional papers, a numbered series of original scientific publications,
primarily short and usually of taxonomic significance.

LIFE sciences miscellaneous publications, an unnumbered series of publications of
varied subject matter and format.

All manuscripts considered for publication are subject to the scrutiny and editorial poli-
cies of the Life Sciences Editorial Board, and to review by persons outside the Museum
staff who are authorities in the particular field involved.

LIFE SCIENCES EDITORIAL BOARD

Chairman: a. r. emery
Senior Editor: IAN R. BALL
Editor: allan j baker
Editor: Gordon edmund

john w Reynolds is an Assistant Professor in the Department of Forest Resources, University of
New Brunswick, Fredericton, New Brunswick.

daniel L dindal is a Professor in the Department of Forest Zoology, College of Environmental
Science and Forestry, Syracuse, New York.

^ KOYAL ONTARIO MlLSEUWf

I*

PRICE: $8.00

© The Royal Ontario Museum, 1977
100 Queen's Park, Toronto, Canada M5S 2C6

PRINTED AND BOUND IN CANADA BY THE HUNTER ROSE COMPANY

This book is dedicated to Dr. Gordon E. Gates on the occasion of his 80th
birthday and 51st year of publication on the Oligochaeta.

Contents

Foreword vii
Acknowledgments i\
Introduction 1

General Biology 3

Introductory Remarks 3

General Activity 3

Parasites and Predators 4

Environmental Requirements and the Effects of

Pesticides 4
Rearing and Culturing Earthworms 6

Methods of Study 8

Sampling Techniques 8
Preservation Techniques 10
Ontario Collection Coding 10
Figure Coding 1 1

General Morphology 13

External Structure 13
Internal Structure 14

Glossary 18

Identification of the Earthworms of Ontario 3 1

Key to Sexually Mature Earthworms Found in
Ontario 32

Systematic Section 34

Family LUMBRICIDAE 34
Genus A llolobophora 35

A. chlorotica 36
Genus Aporrectodea 40

Ap. icterica 40
A p. longa 43
Ap. trapezoides 46
Ap. luberculata 50
Ap. turgida 56
Genus Bimastos 61

B. parvus 61

Genus Dendrobaena 64

D. octaedra 65
Genus Dendrodhlus 69

Dd. rubidus 69
Genus Eisenia 74

E. foetida 74
E. rosea 78

Genus Eiseniella 83

£/. tetraedra 84
Genus Lumbricus 88

L. castaneus 89

L.festivus 92

L. rubellus 94

L. terrestris 99
Genus Octolasiun 104

0. cyaneum 105

0. tyrtaeum 108

Family SPARGANOPHILIDAE 112
Genus Sparganophilus 1 12
5. mew 1 13

Distribution and Ecology 1 16

Appendix: Provincial Description 123

Literature Cited 127

v/

Foreword

"I would not enter on tin list of friends,

(Tho' grae'd with polish'd manners and tine sense

Yet wanting sensihilit\ ) the man

Who needlessly sets foot upon a worm".

Wilham Cowper, The Task (1784)

Nobody could needlessly set foot upon one of the giant earthworms of Australia
or Brazil, large specimens of which may attain lengths of 1 1 feet and weigh up
to 1 pound. But to many people earthworms are lowly insignificant creatures
whose main utility is to act as bait for catching larger and more edible animals.
The Canadian earthworms are indeed represented only by the smaller, more
modest, forms and even though to the uninitiated they all seem to be the same
there are in fact several species which are not too difficult to distinguish. In this
book Dr. Reynolds has assembled, for the first time, all pertinent data, both sys-
tematic and biological, on the Canadian earthworm fauna, and with the aid of a
key, and the fine illustrations of Dr. Dan Dindal, any naturalist or fisherman
should be able to name accurately the specimens that he has at hand.

Earthworms are a significant component of the soil fauna and their beneficial
effects on the agricultural properties of soils have been documented since the
time of Darwin. Some idea of the extent of their activity can be obtained by
reflecting on the fact that something apparently so permanent as the monument
of Stonehenge is being buried at the rate of about seven inches per century as a
result of the burrowing activities of earthworms. Because of their effects on the
soil there can be little doubt that a proper understanding of these creatures is
greatly to man's benefit but, as Dr. Reynolds points out, very little is known of
their biology in North America. This book should form a valuable basis for fur-
ther study of these important aspects.

Dr. Faustus, in part I of Goethe's Faust, speaks disparagingly of him who
"finds his happiness unearthing worms". And an old Chinese aphorism warns
"watch the earthworm; miss the eclipse". But anybody who has spent time in-
vestigating and observing the smaller and lesser known animals of this planet
knows that there is much intellectual satisfaction to be gained from such efforts.
The great Victorian naturalist, Thomas Henry Huxley, likened the uninformed
naturalist to a person walking through an art gallery in which nine-tenths of the
pictures have their faces to the walls. With the aid of this book a few more pic-
tures are now on view.

The Royal Ontario Museum is fortunate to have persuaded Drs. Reynolds
and Dindal to collaborate in producing this book, and zoologists, farmers,
fishermen, naturalists, and teachers throughout northern North America should
have cause to appreciate their labours.

Ian R. Ball
Assistant Curator of
Invertebrate Zoology
Royal Ontario Museum

vii

Acknowledgments

[his project was sponsored in pan under a Gerald L. Beadel Research Grant

from the Tall Timbers Research Station, Tallahassee, Florida. The author
wishes to thank Mr. E.V. Komarek, Si. o\ Tail limbers for his support. The au-
thor also expresses his gratitude to Mr. Dennis Clarke of Canadian Motor In-
dustries (Toyota Ltd.) for providing a 4-wheel drive Land Cruiser to conduct
this study. The author is grateful to Mr. and Mrs. C.W. Reynolds o\' Islington,
Mr. D.W. Reynolds o( Mitchell, and Mr. K. Burns of the Biology Department
of Lakefield College for providing laboratory space during the field work. The
contribution of Mr. C.E. Meadows. Oligochaetology Laboratory. Knoxville to
the various laboratory and statistical analyses in this project is appreciated. The
author thanks Mrs. W.M. Reynolds of Tall Timbers for constant encourage-
ment and support during the course of the study, for reviewing the manuscript,
and for her comments, criticisms, and suggestions. The author would also like to
thank Ms. Jennifer Smith of the Royal Ontario Museum for typing the final
drafts of the manuscript and he is especially grateful to Dr. Ian R. Ball of the
Royal Ontario Museum for the considerable time and effort he devoted to ed-
iting the manuscript, which, in the author's opinion, has improved the text. The
author is indebted to the following for providing specimens for examination
from collections in their care: Dr. LR. Ball (ROM). Dr. E.L. Bousfield (CNM),
Mr. S. Fuller (ANSP), Dr. S.B. Peck (Carleton University), Dr. M.B. Pettibone
(USNM), and Mr. D.P. Schwert (University of Waterloo).

IX

The Earthworms
(Lumbricidae and
Sparganophilidae)
of Ontario

Introduction

Earthworms (Annelida. Clitellata, Oligochaeta) are familiar to almost everyone.
In North America, they are one of the most popular forms of live bait for fishing
(Harman. 1955): gardeners hold them in high esteem as nature's ploughmen
(Darwin, 1881); folklore and scientific accounts tell of their medicinal uses (Ste-
phenson. 1930; Reynolds and Reynolds, 1972), and soil inhabiting vertebrates
(moles, voles, etc.) store them as a source of food (Plisko, 1961; Skoczeii, 1970).
The role of some species in organic matter decomposition and mineral cycling
may be important (Bouche, 1972; Edwards and Lofty, 1972), and a great deal
has been written concerning earthworm farming (Myers, 1969; Morgan, 1970;
Shields, 1971). Biology students the world over study their anatomy (mainly
Lumbricus terrestris) in great detail (Whitehouse and Grove, 1943). The great
amount of literature that has been devoted to a group of organisms that are nei-
ther pests nor sources of human nutrition is truly amazing, yet their biology and
distribution are still relatively unknown. Many of the world's hundreds of me-
gadrile ( = terrestrial oligochaetes) species are known only from a limited series
of one or a few specimens.

This text has been designed to introduce the non-specialist to the taxonomy,
nomenclature, morphology, distribution, and general biology of earthworms in
Ontario and neighbouring areas. The identity, distribution, and habitats of these
animals have been surveyed for a variety of habitats in each of the southern
counties and districts of the province. An illustrated glossary is included to-
gether with a new key to the identification of the earthworms of Ontario that
also is applicable to the rest of eastern Canada and to the northern tier of states
of the United States. French and English common names are included for each
species.

The first records of earthworms from Ontario were provided by Eisen (1874).
Recently Reynolds (1972a) reviewed the complete published and verified un-
published records of terrestrial earthworms from this province, and a second re-
port examined those data quantitatively for habitat factors governing megadrile
activity in the Haliburton Highlands (Reynolds and Jordan, 1975). This study is
a continuation of those reports and presents subsequent collections from 50
counties and districts of southern Ontario in detail (Fig. 1). In addition, unpubl-
ished records derived from collections in North American museums and univer-
sities, including records from northern Ontario, are presented for the first time.

Fig. 1 The Counties and Districts of Ontario.

Thirty-eight of the counties and districts have never had any earthworms re-
ported previously.

At present, there are insufficient megadrile data available to utilize fully some
of the habitat information. For example, it would be unwise to try to correlate
in detail megadrile distribution with the distribution of soil types until addi-
tional surveys from other parts of the continent are completed (cf. Jordan et al.,
1976). A preliminary examination of megadrile-vegetation relationships has
been presented recently (Reynolds, 1976b). A provincial description is included
in an appendix to assemble regional habitat information for future use as well as
to familiarize native and foreign readers with Ontario.

The technical terms and conventions necessary for earthworm discussion will
be found in the Glossary (p. 18). For additional information on earthworm ter-
minology Stephenson (1930), Causey (1952), Gerard (1946), Ljungstrom (1970),
or Gates (1972c) may be consulted.

General Biology

Introductory Remarks

There can he little doubt thai earthworms are the best known of all soil animals.
It is common knowledge that the) have a beneficial efTect on the structure and
properties of the soil and that the) influence the decomposition processes in or-
ganic materials. However, it cannot be denied that much work purporting to
demonstrate these aspects has been far from rigorous. In fact, far less is known
than is generalh believed, and most work is applicable only to Europe. The
problem is compounded b\ the fact that many of the hundreds of described
species are known onl\ from a morphological study of a few individuals. Fortu-
natelv, nearlv all of the species present in Ontario and the neighbouring areas
are the widelv distributed European species that have received the greatest at-
tention. Major limitations to the interpretation of the literature have been old
nomenciatural and taxonomic designations (Reynolds, 1973b).

Sources of information on various biological attributes for species found in
Ontario and the surrounding region are Evans and Guild (1948), Bouche (1972),
Edwards and Lofty (1972), Gates (1972c), Reynolds (1973d). and Reynolds et
al. (1974). Recent reviews of earthworm activity will be found in Kevan (1962)
and Wallwork( 1970).

General Activity

The main activities of earthworms that affect the soil involve the ingestion of
soil and the mixing of the main soil ingredients of clay, lime, and humus; the
production of castings of a fine crumb structure which are ejected on the soil
surface by some species; the construction of burrows that enhance aeration,
drainage, and root penetration; and the production of a tilth that makes suita-
ble habitats for the smaller soil fauna and micro-organisms. It should be re-
membered, however, that not all Lumbricidae work in the same manner. Some,
for example, burrow deeply whereas others do not.

The influence of earthworms on the translocation of soil material may be
quite considerable. There have been abundance estimates as high as three mil-
lion worms per acre and their role in soil fertility is very important. Studying
forms that eject casts to the surface, Darwin (1881) estimated that between IVz
and 18 tons of soil per acre per year (about 3 cm per 10 years) can be moved,
and the burial of many Roman ruins in Europe has been attributed to the activ-
ity of earthworms (Atkinson, 1957).

Earthworms are omnivorous and can utilize many materials in the soil as
food, including plant remains, and occasionally animal remains. Lumbricids
can withstand considerable starvation and, in L. terrestris at least, a water loss
of up to 70% of the body weight. Some species can withstand total immersion in
water for many weeks, though normally they avoid waterlogged soils.

The reproductive cycle of many Lumbricidae is quite straightforward. Al-
though hermaphrodite, they possess a mechanism to prevent self-fertilization.
During copulation the two worms lie side by side with their anterior ends over-
lapping. A mucous sheath envelops the worms and holds them tightly together.
Sperm are released from the testes and flow down the seminal groove in the side

of each worm to the spermathecae of its partner. Both worms do this at the
same time. Some time after copulation has taken place, and after the worms
have separated, the egg cocoons are formed. A mucous tube or belt is secreted
around the clitellum. The worm then wriggles out of this belt and as the belt
passes the female apertures the eggs are deposited in it. Spermatozoa to fertilize
the eggs are deposited as it passes the spermathecal openings. On release the
ends of the belt close over to form a cocoon in which the young worms develop.
Cross-fertilization does not occur in all earthworms, however, despite asser-
tions to the contrary in many textbooks. In some species there is parthenogenes-
is, with concomitant reduction of the male apparatus. Pseudogamy, in which
sperm play no part in the development of the egg other than as a stimulant, also
may occur. Thus, even if copulation has been observed, the exchange of sperm
alone is not evidence for amphimixis. The whole question of reproduction in
earthworms has been reviewed by Reynolds (1974c).

Parasites and Predators

Some earthworms {Allolobophora chlorotica and Eisenia rosea) are parasitized
by Pollenia rudis (Fabr.), a calliphorid fly known as the cluster fly, which may
lay its eggs directly in the earthworm or merely on the surface of the soil (Thom-
son and Davies, 1973a, 1973b). Cluster flies are the most common and annoying
of the flies that overwinter in buildings. Other insects such as ants and beetles
are predaceous on earthworms (McLeod, 1954). Furthermore, some earthworms
may act as intermediate hosts of parasitic worms that affect domestic animals
(Kevan, 1962). Reports of mites (Acari) parasitizing earthworm cocoons and
adults {Allolobophora chlorotica and Eiseniella tetraedra) were made by Stone
and Ogles (1953) and Oliver (1962).

Earthworms are also an important component of the diet of many birds and
mammals. In Europe moles may store them as a source of food (Skoczeh, 1970;
Gates, 1972c), usually after biting off four or five of the anterior segments to
prevent the worms from escaping (Evans, 1948b). In North America they are
eaten by many organisms, including some of economic or recreational impor-
tance. According to Liscinsky (1965), for example, the diet of the woodcock
(Philohela minor Gmelin), a favourite game bird in eastern North America, is
primarily earthworms. From my current surveys, and from gut analyses of
woodcock, it appears that in the area bounded by Ontario to Nova Scotia and
Minnesota to Maryland, 90% of the earthworms in the diet of these birds are
Aporrectodea tuberculata, Dendrobaena octaedra, Dendrodrilus rubidus, and
Lumbricus rubellus. Snakes, too, may prey extensively on earthworms. This is
true especially of two of our most common species, the red-bellied snake
(Storeria occipitomaculata occipitomaculata Say) and the eastern garter snake
(Thamnophis sirtalis sirtalis Linnaeus), and perhaps of four or five other species
as well (Logier, 1958). As this book is in proof, the author has examined the gut
contents from Thamnophis butleri Cope collected in Essex and Lambton Coun-
ties, Ontario. The earthworms identified in these snakes' stomachs were
Allolobophora chlorotica, Aporrectodea tuberculata, and Lumbricus terrestris. Ac-
cording to the author and Dr. S.W. Gorham (pers. comm.), this is the first valid
report of earthworm species identified from snake stomachs in North America.
A recent account was presented by Dindal (1970) of a terrestrial turbellarian,

4

Bipalium adventitium Hyman, attacking Demlrodhlus mbidus and Lumbricus
terrestris. This flatworm is currently a major problem in outdoor earthworm
beds in central New York state (Dindal. pers. comm.).

Environmental Requirements and the Effects of Pesticides

Daylight and ultraviolet light are injurious to earthworms unless the intensity is
very low. Temperature relations have been reviewed by Reynolds (1973a), and
Gates (1970) quotes interesting accounts of lumbricids studied from the Arctic
circle; Eisenia foetida, for example, has been found in snow, even though gener-
ally associated with warm habitats such as manure piles, and it remains vigor-
ous below 5° C. In Maine L. lerrestris has been seen copulating while bathed
with melt water, and other individuals crawled from under the ice and remained
active (Gates. 1970).

The pH tolerance (see Glossary) of earthworms varies from species to species
(Reynolds. 1973d). Usually they occur in soil with a pH range of about 4.5 to
8.7 and the earthworm density diminishes as the soil acidity increases. Gener-
ally speaking, the greatest earthworm densities are found in neutral soils.

The type of soil also may influence the distribution and abundance of the var-
ious species. Gates (1961), for example, divides the earthworms of Maine into
three groups depending upon whether or not they are geophagous, in that they
pass much soil through the intestine; limiphagous (mud-eating) or limicolous
(mud-inhabiting); or. finally, litter-feeding, and hence found primarily in or-
ganic matter. From his studies in Sweden. Julin (1949) divided the Lumbricidae
into four ecological groups. These were hemerophiles, species favoured by hu-
man culture; hemerophobes, species averse to culture; hemerodiaphores, spec-
ies indifferent to the influence of culture; and hemerobionts, species entirely de-
pendent on culture. Julin's classification has never been applied to the North
American Lumbricidae with the exception of a preliminary attempt for the
earthworms of Tennessee by Reynolds et al. (1974). Regrettably, there are as yet
insufficient data to permit an attempt for the Ontario earthworms; this is a topic
worthy of further study.

The application of pesticides to control soil pests, or the earthworm parasites
mentioned above, may also kill the earthworms. This devastating effect on
earthworm populations has frequently occurred after the application of orchard
sprays. Fruit growers have long held earthworms in high esteem for their help in
controlling the disease apple scab which is produced by the fungus Ventuha
inequalis (Cke.) Wint. This disease overwinters on the fallen leaves in the or-
chard. One method of cultural control is to burn the fallen leaves and twigs in
the fall of the year. An equally effective and less costly method is to introduce
earthworms (preferably Lumbricus terrestris), which will pull the fallen leaves
into the soil for food and eventual decomposition. According to the findings of
Reynolds and Jordan (1975). for example, earthworms have a distinct prefer-
ence for apple leaves over those of maple. Once the leaves are beneath the soil
surface the conidiospores of the fungus are ineffectual inoculating agents of the
disease. The preventive measure most commonly used for control of apple scab
is frequent spraving of copper sulphate solutions which are toxic to earthworms
(Raw and Lofty, 1959).

Many studies have been conducted to determine the effects of pesticides on

earthworms. There is little effect on earthworms with normal doses of Aldrin
(Edwards and Dennis, 1960; Edwards et al., 1967; Hopkins and Kirk, 1957;
Legg, 1968), or benzene hexachloride (BHC) (Dobson and Lofty, 1956; Lipa,
1958; Morrison, 1950); chlordane is extremely toxic to them (Doane, 1962; Ed-
wards, 1965; Hopkins and Kirk, 1957; Schread, 1952). DDT, of course, has
been studied by many workers. In general, the application of this pesticide at
normal rates does not harm earthworms (Baker, 1946; Doane, 1962; Edwards,
1965; Edwards and Dennis, 1960; Edwards et al., 1967; Hopkins and Kirk,
1957; Thompson, 1971).

Although earthworms are not susceptible to many pesticides at normal dosag-
es, they do concentrate these toxic chemicals in their tissues. Since many of
these chemicals have long-lasting residual periods in the soil, there is ample op-
portunity for earthworms to absorb them from the soil. The importance of this
phenomenon is that these pesticides can become concentrated in the food chain.
Earthworms are eaten by many species of birds and certain species of amphibi-
ans, reptiles, and mammals, which can continue to concentrate these pesticides
in their bodies (Hunt and Sacho, 1969). Additional reports of pesticides and
their effects on earthworms can be found in Edwards and Lofty (1972).

Herbicides, another group of chemicals, also can affect earthworm popula-
tions (Edwards, 1970; Fox, 1964). These chemicals may kill earthworms direct-
ly, or indirectly by killing the vegetation on which they feed.

One last group of potential poisons that could become concentrated in the
food chain are metal residues. Recently, Gish and Christensen (1973) found that
concentrations of certain metals (cadmium, nickel, lead, and zinc) in earth-
worms were many times that of the surrounding soils. This study was the first
report of metal residues in earthworms. Because of the earthworms' position in
the food chain and the current studies in other fields on metal toxicity, this is an
area requiring further investigation.

Rearing and Culturing Earthworms

It may be of interest to some readers to discuss briefly the rearing or culturing of
earthworms. This is not difficult for the species found in Ontario. There are
many books available describing techniques (e.g.. Ball and Curry, 1956; Myers,
1969; Morgan, 1970; Shields, 1971), although their citation here must not be
taken as an endorsement. The location for earthworm containers depends upon
the climate of the region. Outdoor containers or pit-runs (benches) in northern
areas will require insulation during the winter period when the soil is normally
frozen. Smaller wooden pit-runs, or one of the various types of metal tubs, can
be housed in a basement or shed to avoid winter freezing problems. Since the
indoor facilities permit year-round activity, these can be a source of replenish-
ment for outside gardens, compost piles, flower beds or earthworm beds, etc.
The size of the container can vary. A convenient size is a box 50 cm long X 35
cm wide and 15-20 cm deep. Larger containers, when filled with medium and
earthworms, will be extremely hard to move. These boxes should have holes 0.5
cm in diameter drilled in the bottom. Plastic window screening should be placed
on the inside bottom of the box with a burlap lining on top of the screen and
sides of the box before the soil is added. This permits the excess water to drain

and presents (he soil medium from sticking to the box, and also prevents the
earthworms from escaping through the holes.

Various combinations of soil and organic matter can serve as a medium in
which to raise earthworms. A frequently used mixture is 16 soil and % organic-
matter. Sources of suitable organic matter are: decayed sawdust, hay, leaves,
manure, peat moss. sod. or straw. Additional materials which can be added to
the medium to serve as food sources are: chicken starter, cornmeal, and kitchen
scraps and fats. Earthworms are omnivorous and can utilize many materials as
food sources. Some important facts to remember are: 1) the medium should
contain sufficient organic matter so that it will not pack into a dense, soggy
mass. 2) the containers must not be overwatered, and 3) the presence of low-
watt bright white or blue light will prevent the earthworms from crawling on the
surface of the medium and eventually out of the box.

The species most frequently used as fish bait, and therefore the ones most
likely to be cultured, are: Aporrectodea trapezoides, Ap. tuberculata, Ap. turgida,
and Eisenia foetida. Two other species, Lumbricus rubellus and Octolasion
tyrtaeum, have also been sold or reared as fish bait, though not so commonly as
the others mentioned. The night-crawler Lumbricus terrestris is widely used by
fishermen but cannot be commercially cultured economically because of its
long life cycle, low reproductive rate, and large spatial requirements.

Methods of Study

Sampling Techniques

There are numerous methods for sampling earthworm populations. These fall
mainly under the general categories of hand sorting, chemical extraction, elec-
trical extraction, and vibration methods. The effectiveness of these methods de-
pends upon the species and habitat; no one method is equally suitable for all
species and all habitats.

Digging and hand sorting is the most reliable sampling method, and the one
used primarily to obtain the specimens for this study (Low, 1955; Reynoldson,
1955; Satchell, 1955, 1967, 1969; Svendsen, 1955; Nelson and Satchell, 1962;
Zicsi, 1962). Though laborious, digging and hand sorting have been widely used
for sampling earthworms and for assessing the effectiveness of other methods.
Digging to locate earthworms should be done with two factors in mind, mois-
ture and organic matter (cf. Reynolds and Jordan, 1975), and collecting success
will be high if one concentrates on sites where both are present. The digging can
be done with a variety of tools — shovel, trowel, garden fork, soil cores, etc. The
soil can then be pressed and passed through the fingers, or sieves may be em-
ployed. The advantages of this method are two-fold: within a sample area active
individuals, aestivating individuals, and cocoons may be collected, and, in addi-
tion, a well-defined sampling area may be chosen so that quantitative data may
be obtained. There are some disadvantages, however. The method is laborious
and time-consuming, specimens less than 2 cm in length may escape collection,
and, if digging is restricted to the top layers of soil, very large individuals may
escape into the deeper layers. Furthermore, specimens may be damaged and
there is considerable habitat destruction.

Chemical extraction is a method widely used to collect earthworms and was a
second method employed in the present study. Initial studies on chemical ex-
traction were done by Evans and Guild (1947) using potassium permanganate
solution to expel earthworms from the soil. Further experiments with chemical
extraction, notably using formalin, were conducted by Raw (1959) and Waters
(1955). The standardized sampling format that I have employed over the years
of quantitative extraction is based on a 0.25m2 quadrat of soil surface. A solu-
tion of 25 ml of formalin (37% Formaldehyde Solution, U.S. P.) in four and a
half litres of water is sprinkled over each quadrat so that all of it infiltrates the
soil without runoff. The earthworms that surface in the ten minutes following
the application of the expellant are collected. If the collection is to be obtained
for other than scientific purposes (e.g., for bait), the time, strength, and number
of applications can be varied, but it should be noted that solutions stronger than
15 ml formalin per litre of water may kill the grass in lawns, and if specimens
are to be kept alive for more than a few minutes they must be washed in fresh
water immediately upon surfacing because formalin can act as a vermicide.
Other materials such as Mowrah meal have been used to expel earthworms from
the soil (Jefferson, 1955). With a chemical extraction method the sampling time
and labour are reduced, a well-defined sampling area may be chosen, and there
is minimum disturbance of the habitat. The disadvantages of the method are
that only active individuals are collected, and not cocoons and aestivating or hi-
fi

bernating individuals, onl\ shallow dwelling species or species with burrow sys-
tems are collected, there may be poor penetration of the vermicide when certain
soil conditions prevail, and there is a variability of response to the vermicides by
different species. The technique is generally good for Lumbricidae but poor for
the other families.

Electrical extraction, a method described by several authors (Walton, 1933;
Johnstone- Wallace, 1937; Doeksen, 1950; Satchell, 1961). has long been used
bv fishermen to obtain bait. The method requires a generator and one to three
electrodes. The current conducted through the soil acts as an expellant. The ad-
vantage of this method is minimal disturbance to the habitat. The disadvantages
are the excessive time required per sample, the difficulty of defining the exact
limits of the volume of soil treated, and the variability of the physical and chem-
ical properties of the soil (for example, when soil is moist, deep dwelling species
will surface, but if the surface soil is dry the earthworms may go deeper into the
soil). The use of too much current kills the earthworms near the electrodes, and
the response to electricity varies in different species.

Vibration methods, or mechanical extraction, are currently limited to the south-
eastern United States. Various modifications of this technique ("grunting" in
Florida and Georgia, and "fiddling" in Arkansas) are employed by fish bait col-
lectors and yield earthworms in amazing quantities (Vail, 1972; Reynolds,
1972d. 1973d). Mechanical stimulation by vibrations seems to have very little
effect on the Lumbricidae but it is extremely successful for some Acanthodrili-
dae and some Megascolecidae. These two latter families are not found either in
Canada or in Europe, which may account for omission of this technique in Eu-
ropean discussions of earthworm sampling, except for one small note (Edwards,
R., 1967). The advantages of mechanical extraction are the minimal habitat de-
struction and the reduced sampling time required for each sample. The disad-
vantages are the difficulty of defining the exact volume of soil treated, the effects
of the variability of the physical and chemical properties of the soil, and the var-
iable response of the different species.

There are several other sampling methods that may be used. Wet sieving in-
volves washing soil with a jet of water through a series of sieves after the soil
samples have been removed from the field (Morris. 1922; Bouche and Beugnot,
1972a). There are no available data on the efficiency of this method. The disad-
vantages, according to Ladell (1936), are the excessive time required per sample,
the inordinate amount of labour in residue separation, and the damage to the
specimens during separation.

The flotation method employed by Raw (1960) for extracting the microfauna
from soils unsuitable for hand sorting was patterned after a technique designed
by Salt and Hollick (1944). Its advantage is that it can be adapted to extract
earthworm cocoons; thus, all stages of the population can be sampled.

The heat extraction method operates on the principle of the Baermann funnel
(Baermann. 1917) and has been used for extracting small surface dwelling spec-
ies that are difficult to hand sort. Similar designs employing Tullgren funnels
and incandescent lights have been used by this author. Considerable time per
sample is required for this method as the soil samples have to be brought in
from the field and placed on the wire sieves in the funnels for several hours. The
method has limited use for earthworm sampling.

Trapping techniques are unlikely to yield accurate population estimates but
do form a useful method of studying the activity patterns where population den-
sities are low (Svendsen, 1957). A mechanized soil washing method, involving
rotating containers and standing sieves, was described by Edwards et al. (1970).
This method is faster than previous washing techniques and is apparently suita-
ble for most soils.

Several authors have compared and discussed the relative efficiency of ex-
tracting earthworms from soil by two or more of the previous methods (Svend-
sen, 1955; Raw, 1960; Bouche, 1969a; Satchell, 1967, 1969). From my own
observations, the choice of chemical, electrical, or mechanical methods for ex-
traction of earthworms from the soil is greatly dependent on the genus and
species of earthworm to be collected.

Preservation Techniques

The proper preservation of specimens for identification, shipping, and storage
has long been a problem. Few good accounts of preserving techniques are read-
ily available to those who wish to send material to a specialist for examination.
One of the best media for earthworm preservation is 10-15% formalin because it
hardens the specimens to facilitate handling. Weak alcohol solutions leave the
specimens soft and limp while strong alcohol solutions produce an undesirable
brittleness. In both cases, alcohol also causes a condition known as "alcohol
browning". This condition makes the reporting of colour of preserved alcohol
specimens valueless. Generally, formalin does not distort the colour greatly.

A simple and effective technique is to kill the worms by immersing them in
70% ethyl alcohol. When movement stops they are placed on absorbent paper in
a straight position, and allowed to dry for a few minutes. For preservation they
should then be transferred to a container of 10-15% formalin where they will
harden in the position thus placed. They must be straight because curled or
twisted specimens are more difficult to handle when internal examination and
dissection are required. The specimens should be left in this container overnight
and may then be stored in bottles or vials filled with fresh formalin preservative
without much danger of curling. For best results, the preservative should be
changed again in a week, especially for such species as Aporrectodea trapezoides,
Lumbricus rubellus, and L. terrestris. Diffusion of body fluids from these species
to allow replacement with the preservative seems to take a longer period. As a
general rule, the preservative should be changed at weekly intervals until it re-
mains clear.

Ontario Collection Coding

Under the Ontario Distribution for each species (Systematic Section), the col-
lection data have been coded in a consistent manner: location, habitat (when
available), date of collection, collector(s), number of specimens (by age classifi-
cation, explained in the Glossary), and museum number (if any). When an au-
thor and date are given, the collection information can be found in that source.
If the data are given for a literature source, it is because the author has exam-
ined that collection. An asterisk (*) before a location means that the author has
a 35 mm colour slide of the habitat in his photographic collection. The abbrevi-
ations used in the Ontario Distribution records are:

10

ANSP

Academj of Natural Sciences,

m

metre(s)

Philadelphia

mm

millimetre^)

AT

Alexis Troicki

MKG

Matthew K. Graham

BP

h\ pass

n

north

CNM

National Museums of Canada

n.e.

north edge

CO

couim

RC

R. Cain

CWR

Charles W. Reynolds

Rd

road

DIST

district

RGR

Ruth G. Reynolds

DPS

Donald P. Schwert

RLH

R. Landis Hare

DRB

David R. Barton

RNS

R.N. Smythe

DWR

David W. Reynolds

ROM

Royal Ontario Museum

e

east

RVW

R.V. Whelan

e.e.

east edge

s

south

GE

Gustav Eisen

s.e.

south edge

GM

G. Mueller

ST

Steve Tilton

GMN

G. Morley Neale

St

street

GWA

George W. Abbott

TTRS

Tall Timbers Research

H\v\

highway

Station

IMS

Ian M. Smith

TW

Thomas Weir

Jet

Junction

TY

Toshio Yamamoto

JEM

John E. Moore III

w

west

JO

Jack Oughton

w.e.

west edge

JPM

J. Percy Moore

WMR

Wilma M. Reynolds

JRD

John Richard Dymond

USNM

United States National

JWR

John W. Reynolds

Museum (Smithsonian)

km

kilometre(s)

UW

University of Waterloo

LWR

L. Whitney Reynolds

Figure Coding

The figures for each species were drawn with a camera lucida from preserved
specimens in the author's collection. The source of the specimens for each draw-
ing is given in parentheses after each figure caption. The abbreviations used in
all figures are:

a

anus

es

oesophagus

be

buccal cavity

fp

female pore

cag

calciferous gland

g

gizzard

cg

cerebral ganglion

gl

gut lumen

chl

chloragogen cells

GM

genital markings

cl

clitellum

GS

genital setae

elm

coelom

GT

genital tumescence

cm

circular muscle

h

heart

epe

circumpharyngeal connectives

if

intersegmental furrow

cr

crop

int

intestine

cut

cuticle

lm

longitudinal muscle

dp

dorsal pore

lnv

lateral neural vessel

dv

dorsal vessel

m

mouth

epi

epidermis

mf

male funnel

//

mp

male pore

mL

mid-lateral line

n

nephridium

nb

nephridial bladder

np

nephropore

ns

nephrostome

nt

nephridial tube

o

ovary

od

oviduct

OS

ovisac

ph

pharynx

phm

pharyngeal muscle

PP

periproct

Pr

prostomium

ps

peristomium

ptm

peritoneum

s
sep

seta
septum

s§

seminal groove

sm

setal muscle

snv

subneural vessel

sp

spermathecae

spp

spermathecal pore

sv

seminal vesicle

t

testes

TP

tubercula pubertatis

typ

typhlosole

vd

vas deferens

ve

vas efferens

vnc

ventral nerve cord

vv

ventral vessel

12

General Morphology

The Oligochaeta are defined as annelids with internal and external metameric
segmentation throughout the bod\. without parapodia but possessing setae on
all segments except the peristomium and periproct, with a true coelom and
closed vascular system, generally hermaphroditic with gonads few in number in
specific locations, with special ducts for discharge of genital products, with a cli-
tellum that secretes cocoons in which ova and spermatozoa are deposited, and
which are fertilized and develop without a free larval stage.

The following brief discussion refers primarily to the Lumbricidae. which
make up nearly all of the Canadian megadnle fauna. The terms used in this sec-
tion that are not explained in detail in the text will be found in the illustrated
Glossary. For additional information and details of megadrile morphology con-
sult Stephenson (1930) or Edwards and Lofty (1972) for English accounts, and
Avel (1959) or Bouche (1972) for French.

External Structure

Terrestrial oligochaetes vary greatly in size. Bimastos spp. are less than 20 mm
long, the largest tropical species are over 1200 mm (Glossoscolex, Megascolides),
and some Australian forms may reach 3000 mm in length. The largest species in
Canada is Lumbricus terresths (p. 99). which varies from 90 to 300 mm when
mature. The bodv shape is generally cylindrical though usually flattened dorso-
ventrally in the posterior region in the case of burrowing species.

The entire body is divided along the longitudinal axis into segments separated
by intersegmental furrows and septa. This is primary segmentation. There are
also secondary annuli. or furrows, which appear to subdivide some of the indi-
vidual segments, usually in the anterior region. These demarcations are only ex-
ternal. Ljungstrom and Reinecke ( 1969) have suggested using a and B for these
subdivisions and I use y for a third subdivision; the primary segments are num-
bered by roman numerals. There is a loss of uniformity in segmentation at the
anterior end of the earthworm; this condition is referred to as cephalization (cf.
Gates. 1972c). The first body segment, containing the mouth, is known as the
peristomium and may have a tongue-like lobe projecting anteriorly. The pros-
tomium is located above the mouth, and is not a true segment. Its appearance
is often important in species identification. The last, or caudal, body segment is
referred to as the periproct.

Sometimes a swelling may be seen around the body, the clitellum. The lay-
man frequently mistakes this for the scar of a regenerated animal. In fact it is an
epidermal modification of sexually mature specimens where gland cells secrete
material to form the cocoon.

Characteristic of all earthworms are the short bristles or setae, retractile struc-
tures that add to the worm's grip during tunneling and locomotion. The setae
are produced bv cells in the body wall. In the Lumbricidae and Spargnnophil-
idae there are four pairs of setae per segment, except for the peristonv .m and
periproct. which are asetal. The type and position of these setae have been used
as taxonomic characters (see Glossary — setae, setal formula, setal pairings).

The colour of the megadriles is primarily a result of pigment in the body wall.

13

But it may be a secondary result of lack of pigment and the red colour of some
forms is due to haemoglobin in the blood. Some colour is due to the presence of
yellow coelomic corpuscles near the surface, but the presence of chloragogen
cells near the surface is rarely, if ever, an influence on colour. Preliminary re-
sults of current North American studies indicate that the physical and chemical
properties of the soil are a possible influence on earthworm colour.

The body wall, upon which the excretory, genital, and reproductive apertures
all open, comprises six layers. From the outside these are: cuticle, epidermis,
nerve plexus, circular muscle, longitudinal muscle, and peritoneal layer. The
well-developed muscle layers are important in locomotion. The body wall is the
foundation for many glandular swellings such as the clitellum, tubercula puber-
tals, and genital tumescences, all of which have long been employed as taxon-
omic characters.

Internal Structure

The annelids have often been characterized as possessing a "tube-within-a-
tube" body style (Figs. 2 and 3). The outer tube is formed by the body wall and
the inner tube by the alimentary canal. Between these two tubes is the second-
ary body cavity, or coelom, which is divided at each segment by a septum at the
intersegmental furrow. Non-segmental alignment may occur anteriorly in some
species as a result of cephalization. The coelomic cavity is filled with a fluid that
varies in composition interspecifically, and also intraspecifically for those spec-
ies that are euryecious in that they tolerate a wide range of habitat conditions.
Pores in the septa permit the coelomic fluid to pass freely between segments.

The alimentary canal or digestive tract is essentially a tube extending from
mouth to anus. The anteriormost part of the tract consists of a muscular buccal
cavity, followed by a pharynx which has a sucking action during feeding, the oe-
sophagus, the crop, a crushing organ known as the gizzard, and finally the intes-
tine. The intestine may possess a dorsomedian fold, the typhlosole, that serves
to increase the absorptive surface. Many associated structures are connected to
the alimentary system, viz., blood glands, chloragogen cells, calciferous glands,
and salivary glands. An extensive account of the alimentary canal is found in
Gansen(1963).

The circulatory system is closed but there is an extensive sinus between the
intestinal epithelium and the choragogen cells. Extending almost the total
length of the body are three main vessels (Fig. 3): the dorsal vessel, closely asso-
ciated with the alimentary canal for most of its length, and two ventral vessels
(ventral and sub-neural vessels). The ventral vessel is located between the nerve
cord and the alimentary canal, while the sub-neural vessel is located between
the nerve cord and the body wall. These main vessels are connected in each seg-
ment by paired connectives. In several anterior segments these connectives,
termed "hearts", are enlarged and contractile, and possess valves. There are
other trunks and branches which anastomose throughout the body. The circula-
tory or vascular system has not yet achieved its proper position in oligochaete
systematics. Its importance has been discussed by Gates (1972c) and Reynolds
(1973e).

There is no formalized respiratory system in earthworms; exchange of oxygen
and carbon dioxide takes place through the moist cuticle. Respiration normally

14

XXXV

XXX

Fig. 2 Diagrammatic longitudinal section of a lumbncid earthworm showing internal organs.

15

Fig. 3 Diagrammatic cross section of a lumbncid earthworm.

occurs in air but earthworms can exist in water for long periods of time (e.g., six
months) if the water is highly oxygenated (Brown, 1944, Roots, 1956).

The excretory system is composed of a series of coiled tubes called nephridia
(sing, nephridium). These are the main organs for nitrogenous excretion in
earthworms. In the Lumbricidae, they are paired organs in each segment except
the first three and the last. A nephridium occupies part of two successive seg-
ments where the nephrostome, or funnel, is in the anterior segment and the
coiled tube and nephridial bladder are in the posterior segment. The nephridial
bladder passes through the body wall opening to the outside forming the neph-
ropore. The position of the nephropore, as well as the structure and type of ne-
phridium, are used as taxonomic characters. The most complete discussion of
nephridia and their classification was presented by Bahl ( 1947).

The nervous system is concentrated, with a bilobed mass of nervous tissue
(cerebral ganglia) on the dorsal surface of the pharynx which is connected to
subpharyngeal ganglia by a pair of circumpharyngeal connectives. The nerve
cord, a fusion of the circumpharyngeal connectives, extends caudad from the
subpharyngeal ganglia ventrally between the alimentary canal and the body
wall (Fig. 2). In each segment, posterior to iv, a ganglion is formed and three
pairs of nerves (peripheral nervous system), one pair anterior to the ganglion
and two pairs posterior to the ganglion, extend to the motor and sensory areas.
The nervous system is another portion of somatic anatomy that has not yet
achieved its proper position in oligochaete systematics.

The reproductive system has long been used as the main source of taxonomic
characters. In amphimictic species the male gonads are paired testes found in
segments x and xi close to the anterior septa, a condition termed holandric. An-
terior to each testis, in segments ix and x, and also posteriorly in segments xi

16

and \ii. lobed seminal vesicles occur in which the sperm develop. Sperm are
transferred via the sperm tunnel and sperm ducts to a vas deferens that may
traverse several segments before opening to a male gonopore. The female go-
nads are represented b) a pair o\ gonads found in segment xiii. Ripe oocytes
pass through the coelomic fluid into ovisacs which lead via an oviduct to the fe-
male genital pore. In each of segments ix and x there is a pair of sac-like organs,
opening ventrallv. that receive the sperm during copulation. These are the sper-
mathecae.

In recent years the inadequacies and inconsistencies of reproductive charac-
ters in taxonomy have been discussed (Reynolds et al., 1974; Reynolds, 1974c;
Gates, 1974a). Unfortunately, statements such as "Oligochaetes are hermaphro-
dite, and have more complicated genital systems than unisexual animals" (Ed-
wards and Lofty. 1972) are true onlv in the broadest sense (cf. Reynolds, 1974c).
In this study, eight of the 18 lumbricids are parthenogenetic (or unisexual). In
megadriles. only the clitellum. ovaries, oviducts, and possibly ovisacs are essen-
tial to reproduction (cf. Gates, 1974a; Reynolds, 1974c). Therefore, when repro-
duction is parthenogenetic all of the following are no longer required: testes,
seminal vesicles, seminal receptacles, vas deferens, copulatory chambers, copu-
latory penes, prostates and ducts, genital markings, spermathecae, tubercula pu-
bertates. genital and penial setae.

The external position and morphology of the genital apertures, setae and
tumescences, clitella. and tubercula pubertates have been widely used in lumbri-
cid identification. If these characters are constant for a given species, they are
excellent simple characters that non-specialists can use with reliability.

17

Glossary

A, B, C, D These single capital letters
refer to the meridians oflongitude pass-
ing anteroposteriorly along the aper-
tures of the respective setal follicles.
Thus, A represents a line along the a,
the most ventrally located setal folli-
cles.

ss

1 .

| .

1 • /

-!•

c

lj

A

AA, BB, CC, DD See setal formula.

acinus (Fr. acine m.) A sac-like termination of a branched gland.

aclitellate adults (Fr. adultes sans clitellum, anteclitellienne f.) These are prere-
productive individuals without a clitellum but in which genital markings are
obvious. The second number in the age classification formula (q.v.) refers to
such individuals.

adiverticulate (Fr. sans diverticule) Without diverticula, and usually referring to
spermathecae.

aestivation (Fr. estivation f., anhydrobiose) A period of inactivity, or dormancy,
resulting from unfavourable moisture conditions.

age classification formula A series of numbers following a binomen (usually
three or four numbers) separated by dashes indicating the number of:
juveniles — aclitellate adults — clitellate adults — postclitellate adults in a col-
lection. If there are no postclitellate adults in the collection the final zero is
omitted from the formula. See juveniles, aclitellate adults, clitellate adults,
postclitellate adults.

amphigony See amphimixis.

amphimixis (Fr. amphimixie f.) Reproduction involving fertilization of an
ovum by a sperm. In megadriles the same as biparental reproduction. Cf. par-
thenogenesis.

anal segment See periproct.

anastomosis (Fr. anastomose f.) Cross connections of ducts, branches of or-
gans, or, more usually, of blood vessels.

anthropochore (Fr. anthropochore) Transported by man, usually unintentional-
ly. Cf. peregrine.

aortic arch See hearts.

18

asetal (Fr. sans soies) Without setae. Ct\ peristomium, periproct.

atrial gland (Fr. glande atriole f.) Glandular tissue associated with a cleft or
coelomic invagination containing the male pore.

blood glands (Fr. glandes sanguines f.) Follicles clustered in the pharyngeal re-
gion, supposed to function in the production of haemoglobin and blood cor-
puscles.

brain (Fr. cerveau m.) See cerebral ganglion.

buccal cavity (Fr. cavite buccal f.) (be) The first region of the alimentary canal,
between mouth and pharynx (Fig. 2).

C. Abbreviation for circumference (in German publications replaced by U).
See setal formula.

caecum (Fr. caecum m.) A blind diverticulum or pouch from the alimentary
canal.

calciferous gland (Fr. glande de Morren, glande calcifere f.) (cag) Whitish gland
that secretes calcium carbonate and opens into the gut via the oesophageal
pouches. In Lumbricidae. it is generally found in segments x-xiv.

castings (Fr. dejections de surface f., turricules m.) Faeces, the voided earth and
other waste matter that are commonly deposited on the surface of the ground.
Not all species, however, form their casts above the ground.

cephalization (Fr. cephalisation f.) The loss of metameric uniformity at the an-
terior end of the body.

cerebral ganglion (Fr. ganglion cerebral m.) (eg) Concentrated nerve cells
above the alimentary canal that function as a simple brain (Fig. 2).

cf. (confer) Compare.

ehaeta See seta.

chloragogen cells (Fr. cellules chloragogues f.) (chl) Cells surrounding the ali-
mentary canal; their function is uncertain but is attributed to excretion and
regeneration in the literature (Fig. 3).

unguium See clitellum.

circumpharyngeal connective (Fr. connectif circumpharyngien m.) (epe) Nerve
collar, between cerebral ganglion and ventral nerve ganglion (Fig. 2).

clitellate adult (Fr. adulte avec clitellum, clitellienne f.) Those individuals with

19

developed clitellum and genital markings. The third number in the age clas-
sification formula (q.v.) refers to these individuals.

clitellum (Fr. clitellum m.) (cl) A regional epidermal swelling where gland cells
secrete material to form the cocoon. There are two types recognizable. An an-
nular clitellum or cingulum (Fr. anneau m.) encircles the body whereas a cli-
tellum that encompasses only the dorsal and lateral parts of the body is refer-
red to as a saddle (Fr. selle f.). The convention xxvi, xxvii-xxxii, xxxiii means
that the clitellum is generally found on segments xxvii-xxxii, but may in some
individuals overlap onto segments xxvi and/or xxxiii.

SADDLE ANNULAR

In the case of Eisenia rosea the clitellum has been termed flared. This ventral
flared condition is easily recognizable.

FLARED

NON-FLARED

coelom (Fr. cavite coelomique, coelome f.) (elm) The body cavity between the
body wall and the alimentary canal (Fig. 3).

congeneric (Fr. congenere) Belonging to the same genus.

copulation (Fr. accouplement m., copulation f.) Sexual union, mating.

crop (Ft. jabot m.) (cr) A widened portion of the digestive system that lacks the
muscularity of the gizzard, in Lumbricidae anterior to the gizzard and poste-
rior to the oesophagus (Fig. 2).

cuticle (Fr. cuticule f.) (cut) A thin, non-cellular, colourless, transparent outer
layer of the body wall. See iridescence 2.

diapause (Fr. diapause f.) An obligatory resting stage in development.

digitiform (Fr. digit if orme) Finger-shaped.

dorsal pore (Fr. pore dorsal m.) (dp) Small single in-
tersegmental apertures in the mid-dorsal line (mD)
leading to the coelomic cavity (Fig. 3). The con-
vention first dorsal pore 5/6 means that the dorsal
pore is found in the intersegmental furrow between
segments v and vi.

20

dorsal vessel (Fr. vaisseau dorsal m.) (dv) A major blood vessel located above
the dorsal surface of the alimentarx canal (Figs. 2, 3).

ectal Outer, external, toward the body wall.

egg sac See ovisac.

endemic (Fr. endemique) Restricted to a certain region or part of a region, na-
tive. Cf. exotic, indigenous.

ental Inner, internal, away from the body wall.

epidermis (Fr. epiderme m.) (epi) The outer cellular layer of the body wall,
which secretes a protective cuticle (Fig. 3).

epilubic (Fr. epilobique) See prostomium.

eq. Equatorial, see mL.

euryoecious (Fr. eurvoeciques) Having a wide range of habitat tolerance.

exoic (Fr. exoique) Opening to the exterior through the epidermis, referring to
the excretory system.

exotic (Fr. exotique) Introduced, foreign. Cf. endemic, indigenous.

facultative (Fr.facultatiJ) Conditional, having the power to live under different
conditions. Cf. obligatory.

female ducts Gonoducts. See oviducts.

female pores ( Fr. pores femelles
m.) (fp) The external openings
for the oviducts on segment xiv
(Lumbricidae) and ventrad of
the mid-lateral line. They are
usually more difficult to see
than the male pores.

fide On the authority of, or with reference to publication, to a cited published
statement.

flared clitellum ( Fr. clitellum evasse m.) See clitellum.

genital markings (Fr. mamelons antiarrheniques, mamelons periarrheniques m.)
(GM) Glandular swellings, pits or grooves of the epidermis. See genital
tumescences.

genital setae (Fr. soies genitales f.) (GS) See setae.

21

genital tumescences (Fr. papille puberculienne f.) (GT) In Lumbricidae, areas of
modified epidermis (glandular swellings) without distinct boundaries and
through which follicles of genital setae open.

-GT

cl-

girdle See clitellum.

gizzard (Fr. gesier m.) (g) The muscularized portion of the digestive system, in
Lumbricidae, anterior to the intestine and posterior to the crop (Fig. 2).

gonopore (Fr. gonopore m.) See male pores, female pores.

hearts (Fr. coeurs m.) (h) The enlarged, segmental, pulsating connectives of the
blood system between the ventral and one or two other longitudinal trunks
(e.g., dorsal and/or supra-oesophageal) (Fig. 2).

hemerobiont A species dependent on human culture.

hemerodiaphore A species indifferent to the influence of human culture.

hemerophile A species favoured by human culture.

hemerophobe A species averse to the influence of human culture.

hibernation (Fr. hibernation f.) A period of inactivity or dormancy resulting
from unfavourable temperature conditions.

holandric (Fr. holandrique) The condition where the testes are restricted to
segments x and xi, or a homoeotic equivalent.

holoic (Fr. holonephridique) The condition of having a pair of stomate, exoic
nephridia in each segment of the body except the first and last.

homoeotic (Fr. homoeotique) The condition of having glands or organs in a
segment(s) where they do not normally occur. Refers principally to intraspe-
cific variation.

indigenous (Fr. indigene) Belonging to a locality, not imported, native. Cf. en-
demic, exotic.

22

in lift. (//; liiteris) In correspondence.

intersegmental furrow (Fr. sillon intersegmen-

taire m.) (if) The boundary between two con-
secutive segments; the area where the epider-
mis is thinnest and where, in pigmented spec-
ies, colour is lacking.

>

-&.

— >

3

r?

^

>

4

-J

iridescence (Fr. irisation, iridescence f.) In the context of earthworm biology
this refers to 1) the appearance of sperm aggregated on the male funnels
(q.v.), or 2) the appearance of cuticular colour as a result of refracted light.

juveniles (Fr. lanes f.) Those individuals with no recognizable genital mark-
ings such as the clitellum. tubercula pubertatis. tumescences, etc.. i.e., in the
life stage between hatching and the appearance of genital markings. The first
number in the age classification formula (q.v.) refers to these individuals.

lamella (Fr. lamelle f.) Any thin plate- or scale-like structure.

mD (Fr. medio-dorsale) Mid-dorsal line.

mL (Fr. medio-laterale) Mid-lateral line.

raV (Fr. medio-venirale) Mid-ventral line.

male funnel (Fr. entonnir male m.) (mf) The enlargement of the ental end of a
sperm duct with a central aperture through which sperm pass into the lumen
of the duct on their way to the exterior. Sperm may temporarily aggregate on
the funnels, prior to entering the ducts, their presence being indicated by ir-
idescence (q.v.).

male pores (Fr. pores males m.) (mp) The external openings for the male ducts
through which sperm are liberated during copulation. In Lumbricidae they
are usually conspicuous near the mL on segment xv; any variation is noted in
the diagnosis.

23

male sterility (Fr. sterilite male f.) Often cited as evidence for parthenogenesis
(q.v.) and may be indicated by the following: 1) adult retention of juvenile
testes, 2) adults with juvenile seminal vesicles and no evidence of sperm, 3)
the absence at maturity of iridescence on the male funnels, indicating that
there are no mature sperm aggregations, 4) the absence of similar iridescences
in the male ducts and/or spermathecae, and 5) the absence of externally ad-
hesive spermatophores. These criteria will only suggest male sterility in any
given individual and many cases of repeated evidence are required before a
species can be considered male sterile or parthenogenetic.

megadrile (Fr. megadrile m.) Sensu Gates (1972c: 29) and Reynolds and Cook
(1977), this term is synonymous with terrestrial oligochaetes. There is some
morphological basis for the megadrile/microdrile division of the Oligochaeta
(cf. Gates, 1972c). Brinkhurst (in Brinkhurst and Jamieson, 1971: 104) em-
ploys microdrile as a major heading when discussing the aquatic oligochaetes.
In general, these old terms are used to describe terrestrial and aquatic oligo-
chaetes without any systematic judgments.

mesial (Fr. medial) In the middle vertical or longitudinal plane.

metamere (Fr. metamere m.) A segment.

moniliform (Fr. moniliforme) Arranged like a string of beads.

monotypy (Fr. monotypie f.) The situation arising when a genus-group taxon is
established with only one originally included species; or when a family-group
taxon is established with only one originally included genus.

morph (Fr. forme f., morph f.) A group of individuals that share a common
anatomy resulting from degradations, deletions, or other changes from struc-
ture of the ancestral amphimictic population caused by reproductive isola-
tion. Such isolation usually comes about as a result of parthogenesis.

Morren's gland See calciferous gland.

mouth (Fr. bouche f.) (m) The anterior opening to the alimentary canal located
in the peristomium.

mouth cavity See buccal cavity.

muscular tube See nephridial bladder.

nearctie (Fr. nearctique) A zoogeographical region including Canada, the
United States, Greenland, and northern Mexico.

neotype (Fr. neotype m.) A single specimen designated as the type specimen of
a nominal species-group taxon of which the holotype (or lectotype). and all

24

paratypes or all s\ni\pes are lost or destroyed. Neotypification is the act of
selecting a neotype. (For nominal taxon, see taxon.)

nephridial bladder (Fr. vesicule de la nephridie f.) (nb) The extended portion of
the nephridial tube connected to the nephropore (Fig. 3).

nephridial pore See nephropore.

nephridial reservoir See nephridial bladder.

nephridiopore See nephropore.

nephridium (pi. nephridia) (Fr. nephridie f.) (n) The organ for nitrogenous ex-
cretion (Figs. 2. 3).

nephropore (Fr. nephridiopore m.) (np) The external opening of a nephridium
(Fig. 3).

nephrostome (Fr. nephrostome m.) (ns) The ciliated funnel at the ental end of
the nephridium (Fig. 3).

obligator) (Fr. obligatoire) Limited to one mode of life or action. Cf. faculta-
tive.

oesophagus (Fr. oesophage m.) (es) The portion of the gut between the pharynx
(anterior) and crop (posterior), ending in an oesophageal valve (Fig. 2).

omnivorous (Fr. omnivore) Eating both animal and plant tissue.

op. cit. (opere citato) In the work or article previously cited for this writer (no
page cited).

ovary (Fr. ovaire m.) (o) The organ for ova (egg) production (Fig. 2).
oviducal pores See female pores.

oviduct (Fr. oviducte m.) (od) The duct carrying the ova from the coelomic fun-
nel to the exterior (Fig. 2).

ovisac (Fr. ovisac m.) (os) An egg-capsule or receptacle (Fig. 2).

ovum (pi. ova) (Fr. ovule, oeufm.) The female germ cell, matured egg-cell.

palaearctic (Fr. paleoarctique) A zoogeographical region including all of Eu-
rope and the U.S.S.R. to the Pacific Ocean, Africa north of the Sahara Desert,
and Asia north of the Himalaya Mountains.

25

papilla (FT.papille f.) A protruding dermal structure.

parietes ( Fr. parietes m.) Walls or sides of structures.

parthenogenesis (Fr. parthenogenese f.) Uniparental reproduction in which the
ova develop without fertilization by spermatozoa. Cf. amphimictic.

penial setae (Fr. soies de la verge m.) See seta.

peregrine (Ft. peregrin) Widely distributed, not necessarily involving man.

periproct (Fr. pygidium m.) (pp) The terminal (last, caudal) "segment" of the
body, without coelomic cavity, asetal.

peristomium (Fr. peristomium m.) (ps) The first body segment, asetal, and con-
taining the mouth (Fig. 2).

pH (Fr./?// m.) An indication of acidity or alkalinity measured as the negative
logarithm of the hydrogen-ion concentration, and expressed in terms of the
pH scale (0-14) where pH 7 is neutral, less than 7 is acidic, and more than 7 is
alkaline. Previously, North American studies employed an aqueous solution
to make soil pH readings, and these are the figures given in the text, but varia-
tions can occur when the amount of water present in the soil changes as well
as when the amount of dissolved gases in this water, e.g., C02, changes. To
overcome these variations in the pH readings, one of several salt solutions of
differing strengths may be employed instead of water, e.g., KC1 or CaCl:.
(For details, see Peech, 1965.)

pharynx (Ft. pharynx m.) (ph) The portion of the gut between the buccal cavity
(anterior) and the oesophagus (posterior) (Fig. 2).

pinnate (Fr.penne) Divided in a feathery manner.

polymorphism (Fr. polymorphisme m.) Occurrence of different forms of individ-
uals within the same species.

postclitellate adult (Fr. adulte apres clitellum, postclitellienne f.) Postreproduc-
tive individuals without a clitellum but with areas of discolouration in the re-
gions of the clitellum, and with genital markings. If these discolourations

26

disappear (which is not abnormal), differentiation between aclitellate adults
and postclitellate adults may be impossible even after dissection. These indi-
viduals have reverted to an aclitellate state and in the future may become eli-
te 1 1 u t e again and be reproductive. The fourth number in the age classification
formula refers to these individuals, but if such individuals are not present in
the sample then this fourth figure is omitted instead of using a zero.

prostates (Fr. prostates f.) In Lumbricidae. the same as atrial glands, and of
unknown function.

prostatic pores (Fr. pores prostatique m.) See male pores.

prostoniium (Fr. prostomium m.) (pr) The anterior lobe projecting in front of
the peristomium and above the mouth. There are four types as seen in dorsal
view:

EPILOBIC » ■ PROLOBIC * ' TANYLOBIC ' "ZYGOLOBIC1

1) Epilobic: tongue of the prostomium partly divides the peristomium. 2) Prolo-
bic: prostomium demarcated from the peristomium without a tongue. 3) Tany-
lobic: with a tongue that completely divides the peristomium. 4) Zygolobic:
prostomium not demarcated in any way.

pseudogamy (Fr. pseudogamy) The activitation of ova by a sperm without nu-
clear fusion and thus without true fertilization.

pygidium See periproct.

pygomere See periproct.

pyriform ( Fr. pyriforme) Pear-shaped.

quiescence (Fr. quiescence f.) A period of inactivity, or dormancy, resulting
from an unfavourable environment; cf. aestivation and hibernation.

q.v. (quod vide) Which see.

ridge of puberty (Fr. cretes depuberte f.) See tubercula pubertatis.

sacculate (Fr. saccule m.) Provided with sacculi, small sacs or pouches,
saddle See clitellum.

27

secondary annulation (Fr. sillons transversaux m.) (sa) The furrows which occur
between the intersegmental furrows (q.v.). These demarcations are only exter-
nal and are labelled a, /?, or y.

mD

segment (Fr. segment m.) A portion of the
body, along the anteroposterior axis, be-
tween two consecutive intersegmental fur-
rows and the associated septa. Segments
are numbered with lower case roman nu-
merals, i, ii, iii, etc., beginning anteriorly
with the peristomium as i. The older sys-
tem and some microdrile workers used up-
per case numerals, I, II, III, etc.

seminal receptacles See spermathecae.

seminal reservoirs See seminal vesicles.

Trr

seminal vesicles (Fr. vesieules seminales f.) (sv) The storage sacs for an
earthworm's own sperm until copulation.

septum (pi. septa) (Fr. cloison f.) (sep) The internal partition at intersegmental
furrows. Also acts as a supporting membrane for internal organs (Fig. 2).

seta (Fr. soie f.) (s) A solid rod or bristle se-
creted by cells at the ental end of a tubular
epidermal ingrowth, the setal follicle. Setae
are of several types: 1) general: sigmoid
shape with pointed outer tip; 2) genital: asso-
ciated with genital tumescences and/or go-
nopores, and not sigmoid; 3) penial: associ-
ated with the male pores and not sigmoid.
Individual setae are referred to as a, b, c, dy as
shown in the first diagram of this Glossary, a
being the most ventral and d the most lateral
of the setae on a particular segment.

GENERAL

PENIAL

28

setal formula (Fr. des soies f.) The distance between ^DD-^/

the setae, usually measured on segments x and/or \^NA0

xxx, and being an estimate of the space between the / \ \_

A. B, C. and D meridians (q.v.). The data can be ex- I ^_

pressed as a ratio (e.g., AA:AB:BC:CD:DD = 9:3:6: Jj» /* JL

2:30), as groupings (e.g., AA >BC<DC AA=BQ or ^/ \^B

in terms of the circumference. C. (e.g., DD = 1 /2C). /— Ay*-"Y"
See also setal pairings.

setal pairings (Fr. schema de la disposition des soies) Setae may be closely
paired (Fr. soies etroitement geminees), widely paired (Fr. soies distantes), or
separate (Fr. soies ecartees, soies separees).

o

V tf

C ) ( \

CLOSELY WIDELY SEPARATE

somatic (Fr. somatique) Referring to any portion of the anatomy except the re-
productive organs.

sperm (Fr. spermatozoides m., sperme m.) The male germ cells, fertilizing agent.

sperm ducts See vas deferens.

sperm funnel See male funnel.

sperm sacs See seminal vesicles.

spermathecae (Fr. spermatheque f.) (sp) The pouches developed in the septa
which receive sperm from another individual during copulation; the sperm
are stored here until the period of cocoon laying.

spermatophore (Fr. spermatophore m.) A capsule of albuminous matter con-
taining a number of sperm.

spermatozoa See sperm.

spermiducal pores See male pores.

stomate (Fr. stomate) Referring to open nephridia, i.e., with funnel.

tanylobic (Fr. tanylobique) See prostomium.

29

taxon (pi. taxa) (Fr. taxon m.) Any taxonomic unit such as a particular family,
genus, or species. Nominal taxon: The taxon, as objectively defined by its type,
to which any given name whether valid or invalid applies.

testis (pi. testes) (Fr. testicules m.) (t) The organs for sperm production.

testis sac (Fr. sac du testicule m.) Usually a closed off coelomic space contain-
ing one or both testes and male funnels of a segment.

trabeculate (Fr. trabeculaire) Seminal vesicles that develop as connective tissue
proliferations from a septum so as to have numerous irregular spaces that re-
main inconsiderable until spermatogonia (primitive sperm cells) begin to en-
ter.

t liberal la pubertal is (Fr. puberculum m.) (TP) A glandular swelling appearing
near the ventrolateral margins of the clitellum. It is not always present, and it
may be continuous or discontinuous, and of varied size and shape.

typholosole (Fr. typhlosolis m.) (typ) Any longitudinal fold in the gut wall pro-
jecting into the gut lumen, usually at mD or mV (Figs. 2, 3).

vas deferens (Fr. canal deferent m.) (vd) The ducts that carry sperm from the
male funnels to the exterior (Fig. 2).

ventral vessel (Fr. vaisseau ventral m.) (vv) A major blood vessel, located ven-
tral to the alimentary canal and dorsal to the ventral nerve cord (Fig. 2).

vesiculate (Fr. vesiculeux) Having a vesicle or small bladder-like sac.

viz. (videlicet) Namely.

zygolobic See prostomium.

1-1-1-1 See age classification formula.

1/2 See first dorsal pore.

i, ii. iii See segment.

30

Identification of the Earthworms of Ontario

The identification of the local earthworms is not as difficult as most people sus-
pect. The following key. which should be used in conjunction with the Glossary
and diagnoses, has been designed to facilitate identification of the 19 species
recorded in Canada without the necessity of dissecting the specimens. Generally
speaking, mature specimens are essential for a definitive identification by the
non-specialist.

The most useful characters of the key are the nature of the prostomium (zygo-
lobic only in Sparganophilus eiseni. tanylobic in species of the genus Lumbhcus,
and epilobic in all others), the segmental position of the clitellum and tubercu-
lata pubertatis (remembering that the prostomium is not numbered), the ar-
rangements of the setae, and the presence or absence of pigment. These charac-
ters are all readily visible in fresh material.

If only preserved material is available, some difficulty may be experienced in
those parts of the key (couplets 8 and 12) that rely on assessments of colour. In
these circumstances it may be necessary at the appropriate point to refer to the
detailed diagnoses of several species before proceeding further. At couplet 8, for
example, if the colour cannot be assessed confidently it will be necessary to refer
separately to the diagnoses of Dendrobaena octaedra, Dendrodrilus rubidus, and
Eisenia foetida. If the preserved specimen at hand definitely is not one of these
three species then one continues through the key on the assumption that there is
no red pigment present.

The study of the characters used in the key requires no more than a good
hand lens or a low-power binocular microscope. The key itself is strictly dicho-
tomous. The numbers in parentheses after the main couplet numbers indicate
the couplet from which that particular point in the key was reached. They are
inserted to make it easier to retrace one's steps through the key in the event that
an obviously incorrect alternative has been reached.

31

Key to Sexually Mature Earthworms Found in Ontario

1 Prostomium zygolobic; clitellum begins in front of segment xx

Sparganophilus eiseni (p. 1 1 3 )

Prostomium not zygolobic; clitellum begins behind segment xx 2

2 ( 1) Prostomium tanylobic 3

Prostomium epilobic 6

3(2) Clitellum begins in front of segment xxx 4

Clitellum begins behind segment xxx 5

4( 3) Clitellum on segments xxvi, xxvii-xxxi, xxxn; tuberculata pubertatis on segments

xxviii-xxxi Lumbricus rubellus (p. 94)

Clitellum on segments xxvii-xxxiii; tuberculata pubertatis on segments
xxix-xxxii Lumbricus casianeus (p. 89)

5( 3) Clitellum on segments xxxi, xxxii-xxxvii; tuberculata pubertatis on segments

xxxiii-xxxvi Lumbricus terrestris (p. 99)

Clitellum on segments xxxiv-xxxix; tuberculata pubertatis on segments xxxv-xxxvii,
xxxviii Lumbricus feslivus (p. 92)

6(2) Clitellum on segments xxiv-xxxi, tuberculata pubertatis absent Bimastos panus (p. 6 1 )

Clitellum rarely on xxiv-xxxi only; tuberculata pubertatis present 7

7 ( 6) Tuberculata pubertatis small sucker-like discs, usually on segments xxxi, xxxiii, and xxxv

(Fig. 4); clitellum on xxviii, xxix-xxxvii Allolobophora chlorotica (p. 36)

Tuberculata pubertatis not as above 8

8 ( 7) Pigment present, red 9

Pigment absent, or if present not red 11

9 ( 8) Setae closely paired; clitellum on segments xxiv, xxv, xxvi-xxxti; tuberculata pubertatis on

segments xxviii-xxx; sometimes striped (alternate transverse dark and light bands)

Eisenia foetida (p. 74)

Setae widely paired or separate 10

10 ( 9) Clitellum on segments xxvii, xxviii-xxxiii, xxxiv; tuberculata pubertatis on segments

xxxi-xxxiii, usually darkly pigmented Dendrobaena octaedra (p. 65)

Clitellum on segments xxvi, xxvii-xxxi, xxxii; tuberculata pubertatis on segments xxviii,
xxix-xxx Dendrodrilus rubidus* (p. 69)

11(8) Male pores equatorial on xiii ; female pores on xiv ; often yellowish in colour

Eiseniella tetraedra (part) (p. 84)

Male pores not on xiii 12

12(11) Pigmented 13

Not pigmented 15

13 (12) Setae widely paired; clitellum on segments xxii, xxiii-xxvi, xxvii; tuberculata pubertatis on

segments xxiii, xxiv-xxv, xxvi

Eiseniella tetraedra (part) (p. 84)

Setae closely paired 14

14(13) Clitellum on segments xxvi, xxvii, xxviii-xxxiv; tuberculata pubertatis on segments

xxxi-xxxiii Aporrectodea trapezotdes (part) (p. 46)

Clitellum on segments xxvii, xxviii-xxxiv, xxxv; tuberculata pubertatis on segments
xxxii-xxxiv Aporrectodea longa (p. 43)

A rare morph of this species, found in the southern USA. lacks tuberculata pubertatis.

32

15(12) Setae widel) paired (posteriorly at least) 16

Setae closely paired 17

16(15) Clitellum on segments xxix-xxxiv; tuberculata pubertatis on segments

xxx-xxxiii Octolasion cyaneum (p. 105)

Clitellum on segments xxx-xxxv; tuberculata pubertatis on segments
xxxi-xxxiv Octolasion tyrtaeum (p. 1 08)

17(15) Clitellar region often flared ventrallv (bell-shaped in cross section, see glossary, p. 20);

clitellum on segments xxv, xxvi-xxxiii; tuberculata pubertatis on segments xxix-xxxi

Eisenia rosea (p. 78)

Clitellar region not flared ventrally 18

18(17) Clitellum begins in front of segments xxx 19

Clitellum begins behind segment xxx; clitellum on segments xxxiii, xxxiv-xlii,
xhu Aporrectodea icterica (p. 40)

19(18) Genital tumescences often present on segment xxviii, present or absent on segments
xxxiii-xxxiv; clitellum on segments xxvii, xxviii-xxxiv; tuberculata pubertatis on segments

xxxi-xxxiii; male sterile (see glossary, p. 24) Aporrectodea trapezoides (part) (p. 46)

Genital tumescences not present on segment xxviii; male fertile 20

20(19) Clitellum on segments xxvii-xxxiv; tuberculata pubertatis on segments xxxi-xxxiii; genital
tumescences often present on segments xxvi and usually absent on segment

xxxiii Aporrectodea tuberculata (p. 50)

Clitellum on segments xxvii, xxviii. xxix-xxxiv, xxxv, tuberculata pubertatis on segments

xxxi-xxxiii; genital tumescences often present on segments xxvii and xxxiii

Aporrectodea turgida (p. 56)

33

Systematic Section

There are three levels of taxa reported in this section — families, genera, and
species.

For each family (Lumbricidae and Sparganophilidae), a diagnosis and the
designation of the type genus are given. For each of the ten genera occurring in
Ontario there are presented a synonymy, the type species, a diagnosis, and a dis-
cussion.

The information for the 19 species reported from Ontario is presented in the
following order: common name (English and French), synonymy, diagnosis, ex-
ternal anterior illustrations (lateral and ventral views), discussion, biology (habi-
tats, reproduction, etc.), range. North American distribution, Ontario distribu-
tion records, and map. For some species, at the end of the North American
distribution records are one or more regions listed under "new records". This
means that the author has recently received or collected specimens from these
areas. Since there are no published records of the species from these areas, they
are included without citation to give the reader the most complete information
on the distribution of the species in North America. The abbreviations used for
the Ontario distribution records are found in the section Methods of Study (p.
10). Niagara County is the union of the former Lincoln and Welland counties.
For the purposes of this study North American distribution includes Canada,
the United States of America, and Greenland.

It should be noted that unless otherwise stated all biological notes for the
species are compilations from the literature. Very little work has been done on
the biology of earthworms in Canada and most data are from Europe. The most
thorough reviews to which reference may be made for further information are
those of Bouche (1972) and Gates (1972c).

Family LUMBRICIDAE Claus, 1880*

Diagnosis

Digestive system: with an intramural calciferous gland comprising longitudinal
chambers that open at their anterior ends into the oesophageal lumen, a termi-
nal oesophageal valve reaching into xv, an intestine beginning with a "crop" fol-
lowed by a gizzard, a sacculated as well as an unsacculated portion and ending
in an atyphlosolate region, but without intestinal caeca and supra-intestinal
glands. Vascular system: with complete dorsal, ventral, and subneural (and
lateroneural?) trunks, the latter adherent to nerve cord, extraoesophageal
trunks median to the hearts passing to dorsal trunk in region of x-xii, without
supra-oesophageal and lateroparietal trunks. Hearts: lateral, the last pair ante-
rior to segment xii. Nephridia: holoic, vesiculate, ducts passing into parietes in

•Designated by Sims (1973).

34

region of B. Setae, sigmoid and single pointed, eight per segment, in regular
longitudinal ranks, in genital tumescences elongated but slender and longitudi-
nally grooved ectally. Dorsal pores, present. Prostomium epilobic, prolobic, or
tanvlobie. Reproductive s\stem: apertures, all minute, female pores anterior to
the male pores, equatorial and anterior to the multilayered clitellum which is al-
ways behind xvii, Spermathecae, adiverticulate, pores at intersegmental levels.
Ovaries, in xiii. bandlike, each terminating distalh in a single eggstring. Ovisacs,
in \iv. small, lobed. Ova. not volkv. Prostates, none (after Gates, 1972c: 61-62).

Type Genus

Lwnbricus Linnaeus, 1758 (neotypification Sims (1973)).

Genus A llolobophora Eisen, 1873

1873 Allolobophora Eisen, Ofv. Vet.-Akad. Forh. Stockholm 30(8): 46.

1900 Allolobophora-Michddsen, Das Tierreich, Oligochaeta 10: 480.

1910 Allolobophora-Michaeken, Ann. Mus. Zool., St. Petersburg 15: 1.

1930 Allolobophora (part.)-Stephenson, Oligochaeta, p. 905, 906, 907, 908.

1941 Allolobophora (part.)-Pop, Zool. Jb. Syst. 74: 20.

1956 Allolobophora (part.)-Omodeo, Arch. Zool. It. 41 : 180.

1972 Allolobophora (part.)-Gates, Trans. Amer. Philos. Soc. 62(7): 68, 69.

1972 Allolobophora-Bouche, Inst. Natn. Rech. Agron., p. 263, 417.

1975 Allolobophora-Gales. Megadrilogica 2(1): 3.

Type Species

Lumbric us riparius Hoffmeister, 1843 ( = Enterion chloroticum Savigny, 1826)

Diagnosis

Calciferous gland, opening into gut through a pair of vertical sacs posteriorly in
x. Calciferous lamellae continued along lateral walls of sacs. Gizzard, mostly in
xvii. Extraoesophageal vessels, passing to dorsal trunk in xii. Hearts, in vi-xi.
Nephridial bladders, 7-shaped, closed end laterally, ducts passing into parieties
near B. Nephropores, inconspicuous, behind the clitellum irregularly alternating
between levels slightly above B and above D. Setae paired. Prostomium, epilo-
bic. Longitudinal musculature, pinnate. Colour variable, (after Gates, 1972c: 68
and 1975a: 3).

Discussion

Allolobophora was erected by Eisen (1873) without the designation of a type
species and this situation was not corrected by Michaelsen (1900b) in his revi-
sion of the Lumbricidae. Typification of the genus was by Omodeo (1956) who
selected A. chlorotica to be the type. Additional species that Eisen included in
his Allolobophora were: arborea, foetida, mucosa, norvegica, subrubicunda, and
turgida, none of which is now referable to this genus (Gates, 1975b: 7).

35

Allolobophora chlorotica (Savigny, 1826)

Green worm Ver vert
(Fig. 4)

1826 Enterion chloroticum + E. virescens Savigny, Mem. Acad. Sci. Inst. Fr. 5:

183.

1828 Lumbricus anatomicus Duges, Ann. Sci. Nat. 15(1): 289.

1837 Lumbricus chloroticus-Duges, Ann. Sci. Nat., ser. 2, 8: 17, 19.

1843 Lumbricus ripahus Hoffmeister, Arch. Naturg. 9(1): 189.

1845 Lumbricus communis luteus Hoffmeister, Regenwiirmer, p. 29.

1865 Lumbricus viridis Johnston, Cat. British non-paras, worms, p. 60.

1873 Allolobophora riparia-Eisen + A. mucosa Eisen, Ofv. Vet.-Akad. Forh.

Stockholm 30(8): 46, 47.

1882 Allolobophora neglecta Rosa, Atti Ace. Torino 18: 170.

1884 Allolobophora chlorotica-V ejdovsky, Syst. Morph. Oligo., p. 60.

1885 Aporrectodea chlorotica-Orley, Ertek. Term. Magyar Akad. 15(18): 22.
1892 Allolobophora cambrica Friend, Essex Nat. 6: 31.

1896 Allolobophora curiosa Ribaucourt -I- A. Waldensis Ribaucourt + A.

morganensis Ribaucourt + A. cambria (laps.)-Ribaucourt, Rev. Suisse

Zool. 4: 46, 47, 94.
1900 Helodrilus (Allolobophora) chloroticus-Michaehen, Das Tierreich, Oligo-

chaeta 10: 486.

Diagnosis

Length 30-70 mm, diameter 3-5 mm, segment number 80-138, prostomium epi-
lobic, first dorsal pore 4/5. Clitellum xxviii, xxix-xxxvii. Tubercula pubertatis
small, sucker-like discs on xxxi, xxxiii and xxxv. Setae closely paired, AA >
BC, DD = ','2 C anteriorly, and DD < Vi C posteriorly. Setae c and d on x often
on white genital tumescences. Male pores in xv with large elevated glandular
papillae extending over xiv and xvi. Seminal vesicles, four pairs in 9-12. Sper-
mathecae, three pairs opening on level cd in 8/9, 9/10 and 10/1 1. Colour varia-
ble, frequently green but sometimes yellow, pink, or grey. Body cylindrical.

Biology

This species has been found in a wide variety of soil types, with a pH of 4.5-8.0,
including gardens, fields, pastures, forests, clay and peat soils, lake shores and
stream banks, estuarine flats, and among all sorts of organic debris. It has been
found in caves in Europe and North America and also in botanical gardens and
greenhouses in these same continents. Most of the sites where A. chlorotica was
obtained in the present survey were moist, low areas, such as under various
forms of debris and logs in ditches, relatively close to the Great Lakes. Eaton
(1942) reported the habitat preference of this species as "wet and usually highly
organic or polluted soil." In eastern Tennessee almost 85% of the specimens col-
lected were from wet, highly organic habitats (Reynolds et al., 1974).

In appropriate conditions activity, including breeding, possibly occurs all
year. In the northern part of the range there may be a single period of activity in
the summer. There are records of active specimens occurring 300 mm below the
soil surface although the species generally is characterized as shallow burrow-
id

1st dp

xxx

XXV

XXXV

Fig. 4 External longitudinal views of Allolobophora chlorotica showing taxonomic characters, a.
Dorsolateral view. B Ventrolateral view. (ONT: Haldimand Co., cat. no. 734 1)

37

ing. Defecation occurs below the soil surface as does copulation. A. chlorotica is
obligatorily amphimictic (Reynolds, 1974c).

This species has been reported as the secondarily preferred host of the cluster
fly, Pollenia rudis (Fabr.) (Yahnke and George, 1972; Thomson and Davies,
1973b); otherwise it is of minimal economic importance. It seems not to be pre-
ferred by fish, and anglers have found little use for it as bait.

Range

A native of Palaearctis, A. chlorotica is known from Europe, Iran, North Ameri-
ca, South America, North Africa, and New Zealand (Gates, 1972c).

North American Distribution

British Columbia (Smith, 1917). New Brunswick (Reynolds, 1976d), Nova Scotia (Reynolds, 1975a,
1976a), Ontario (Reynolds, 1972a), Quebec (Reynolds, 1975b, d, e, 1976c), Arizona (Gates, 1967),
California (Smith, 1917), Connecticut (Reynolds, 1973c). Delaware (Reynolds, 1973a), District of
Columbia (Smith, 1917), Idaho (Gates, 1967), Illinois (Smith. 1928), Indiana (Smith. 1917), Maine
(Gates, 1961), Maryland (Reynolds, 1974b). Massachusetts (Reynolds. 1977). Michigan (Murchie,
1956), Missouri (Olson, 1936), Montana (Reynolds, 1972c), Nevada (Gates, 1967), New York (Ol-
son, 1940), North Carolina (Smith, 1917), Ohio (Olson, 1928), Oregon (MacNab and McKey-Fend-
er, 1947), Pennsylvania (Eaton, 1942), Tennessee (Reynolds, 1972b), Utah (Gates, 1967), Vermont
(Gates, 1972c), Virginia (Gates, 1949), Washington (MacNab and McKey-Fender, 1947). West Vir-
ginia (Williams, 1942), Wisconsin (Gates, 1972c), Greenland (Levinsen. 1884). New records: Mani-
toba, Alaska, Minnesota.

Ontario Distribution (Fig. 5)

CARLETON CO. Reynolds (1972a); *Ottawa-Carleton Rd 34, Cumberland, under paper, 1 1 May
72, JWR, 0-0-1. DUNDAS CO. *Hwy 2, 5 km w of Iroquois, under logs, 11 May 72. JWR. 0-1-0.
*Hwy 43, 1.29 km e of Chesterville, under rocks in ditch, 1 1 May 72, JWR. 0-0-4. 'Hwy 31. 2.1 km s
of Winchester Springs, under logs, 11 May 72, JWR, 0-0-3. DURHAM CO. 'Hwy 401. .16 km w of
Liberty Rd, Bowmanville, digging under log by railroad tracks, 15 May 72, JWR, 1-0-1. ELGIN
CO. *Hwy 73, 3.06 km s of Harnettsville, under rotten log, 4 May 72, JWR. 0-1-1. *Hwy 73. 6.77 km
n of Aylmer, under logs, 4 May 72, JWR, 3-2-4. *Hwy 3, 6.77 km e of Wallacetown, under pine logs,
4 May 72. JWR, 0-1-0. Hwy 3, 7.9 km w of Frome, under logs. 4 May 72, JWR, 0-0-7. ESSEX CO.
•Hwy 3, 1.45kmeofCottam, under logs, 4 May 72, JWR, 1-1-8. FRONTENAC CO. *Hwy 2. 1.13
km e of Westbrook, under logs and paper, 16 May 72, JWR, 0-0-10. *Hwy 15, 4.03 km n of Hwy
401, under paper in wet ditch, 16 May 72, JWR, 0-0-1. GRENVILLE CO. *Hwy 2. Johnstown, w.e.,
under logs, 1 1 May 72. JWR, 0-1-4. HALDIMAND CO. *Hwy 54, 2.26 km n of Caledonia, digging,
3 May 72. JWR. 2-2-0. H ALTON CO. *Hwy 7, 4.84 km e of Georgetown, under burnt logs in soil. 4
May 73, JWR, 0-0-2. HASTINGS CO. *Hwy Jet 2 and 49. wet ditch, 15 May 72. JWR, 0-2-37. HU-
RON CO. *Hwy 4, 1.29 km s of Wingham, under logs in wet area, 5 May 72, JWR, 0-5-7. KENT
CO. 'River Rd, 6.45 km w of Chatham, under logs, 23 Apr 72, JWR & TW, 0-1-0. Hwy 3, 1.77 km w
of Palmyra, under logs, 4 May 72, JWR, 1-2-15. LAMBTON CO. Hwy 21. 2.1 km n of Wyoming,
under logs in wet ditch, 4 May 72, JWR, 0-0-10. LANARK CO. *Hwy 43, 3.71 km e of Smiths Falls,
under telephone pole in ditch, 1 1 May 72, JWR, 0-0-18. LEEDS CO. *Hwy 15, 2.26 km n of Seeley's
Bay, under logs and concrete blocks in ditch. 16 May 72. JWR, 10-2-15. LENNOX AND AD-
DINGTON CO. *Hwy 2, 9.68 km w of Napanee. under paper in wet ditch. 15 May 72. JWR, 8-2-
17. MIDDLESEX CO. Judd ( 1970). *Hwy 7, 2.26 km s of Parkhill, under posts. 4 May 72. JWR. I-
1-1. NIAGARA CO. 'Niagara Co Rd 12, 3.06 km s of Grimsby, under paper in wet ditch, I May
72, JWR. 0-1-9. NIPISSING DIST. 'Hwy 17. 1.29 km e of Verner, under paper in wet ditch. 13
May 72. JWR & JEM, 1-1-2. NORTHUMBERLAND CO. 'Northumberland-Durham Rd 1. .65
km w of Hwy 33, under logs, 15 May 72, JWR, 3-0-3. ONTARIO CO. 'Hwy 7. 1.61 km e of Green

38

Fig. 5 The known Ontario distribution of A llolobophora chlorotica.

River, under logs. 26 Apr 72. JWR. 2-0-9. PEEL CO. *Hwy 5. .81 km e of Dixie Rd. 29 Apr 72,
JYVR. 0-0-2. »Hwy 7. Brampton, e.e.. under debris. 4 May 73, JWR, 0-0-2. Hwy 401, 1.3 km w of
Dixie Rd. wet ditch. 4 Mav 73. JWR, 0-0-1. PERTH CO. 'Mitchell, next to Collegiate, under logs, 4
Slav 73. JWR & DWR, 2-3-2. PRESCOTT CO. *Hwy 34, 2.58 km n of Vankleek Hill, under logs
and rocks. 11 May 72. JWR. 1-2-6. PRINCE EDWARD CO. *Hwy 33, 1.61 km s of Carrying Place,
under log in wet ditch, 15 May 72, JWR. 3-0-8. *Hwy 33. Consecon, n.e.. dump, 15 May 72, JWR,
0-0-1. 'Hwy 33, 2.58 km e of Hillier. digging. 15 May 72. JWR. 0-3-0. Indian Point, in grab sample,
approximately 1 metre depth in Lake Ontario. 24 Jul 74, DRB, 1-0-2, UW-0001. STORMONT CO.
*Hwy 43. 5.48 km w of Finch, in and under wet straw in wet ditch, 11 May 72, JWR. 0-0-42.
WATERLOO CO. Waterloo. Amos Ave.. 1 Sep 75. DPS, 1-3-0. UW-0003. WENTWORTH CO.
Reynolds (1972a). *Hwy 5, Waterdown, e.e, edge of corn (Zea maize L.) field, 29 Apr 72, JWR, 0-
0-4. YORK CO. 'Edenbrook Park, Islington, under logs and rocks near stream bank, 30 Apr 72,
JWR & DWR. 1-1-2. Scarborough, 21 Kingston Rd.. small wooded valley, 30 Nov 41, JO. 0-0-1.
ROM.

39

Genus Aporrectodea Or ley, 1885

1885 Aporrectodea Orley, Ertek. Term. Magyar Akad. 15(18): 22.

1900 AUolobophora (part.)-Michaelsen, Das Tierreich, Oligochaeta 10: 480.

1930 AUolobophora (part.)-Stephenson, Oligochaeta, p. 905, 906, 907, 908.

1 94 1 A Uolobophora (part.)-Pop, Zool. Jb. Syst. 74: 20.

1956 AUolobophora (part.)-Omodeo, Arch. Zool. It. 41 : 180.

1972 AUolobophora (part.)-Gates, Trans. Amer. Philos. Soc. 62(7): 68, 69.

1972 Allolobophora-Gates, Bull. Tall Timbers Res. Stn. 12:2.

1972 Nicodrilus Bouche, Inst. Natn. Rech. Agron., p. 315.

1975 Aporrectodea-Ga\es, Megadrilogica 2(1): 4.

Type Species

Lumbricus trapezoides Duges, 1828.

Diagnosis

Calciferous gland, opening into gut through a pair of vertical sacs equatorially
in x. Calciferous lamellae continued onto posterior walls of sacs. Gizzard,
mostly in xvii. Extraoesophageal vessels, passing to dorsal trunk in xii. Hearts,
in vi-xi. Nephridial bladders, ^/-shaped, ducts passing into parieties near B.
Nephropores, inconspicuous, irregularly alternating between levels slightly
above B and above D. Setae, paired. Prostomium, epilobic. Longitudinal mus-
culature, pinnate. Pigment, if present, not red (after Gates, 1975a: 4).

Discussion

This forgotten genus originally included Enterion chloroticum Savigny, 1 826 and
Lumbricus trapezoides Duges, 1828. Since Omodeo (1956) designated the former
as the type species of AUolobophora, the latter automatically becomes the type
for Aporrectodea. Bouche (1972) erected a new genus Nicodrilus with Enterion
caliginosum Savigny, 1826 as the type and included Lumbricus trapezoides Dug-
es, 1828 in this new genus. Since Aporrectodea is a valid and available genus,
Nicodrilus must be considered the junior synonym of Aporrectodea.

Aporrectodea icterica (Savigny, 1826)

Mottled worm Ver marbre
(Fig. 6)

1826 Enterion ictericum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 183.

1837 Lumbricus /'c7m'cu.y-Duges, Ann. Sci. Nat., ser. 2, 8: 17.

1886 AUolobophora icterica-Rosa, Atti 1st. Veneto, ser. 6, 4: 685.

1900 Helodrilus /c/en'cw^-Michaelsen, Das Tierreich, Oligochaeta 10: 500.

1926 Bimastus /<vj«/.s-Pickford, Ann. Mag. Nat. Hist., ser. 9, 17: 96.

1938 Eophila icterica-Tetry, Contr. Etude Faune Est France, p. 269.

1972 AUolobophora icterica-Bouche, Inst. Natn. Rech. Agron., p. 273.

1976 Aporrectodea /'c/er/ra-Reynolds, Megadrilogica 2(12): 3.

40

E
o

1st dp

V /

•V

-~sd—

x V-~~-

t*V

r~~ 'j

• ■ •}

i ' »'. i— •>

xv te

. ;J

GT

cl

XX,

XXV

-■ 1 — -li

XXX

XXXVI

xl<

I . — -5J1

xlv

TP.

xx

XXV

XXX

-GT

Fig. 6 External longitudinal views of Aporrectodea icierica showing taxonomic characters, a. Lat-
eral view, b Ventral view. (ONT: Wellington Co., cat. no. ROM 148)

41

Diagnosis

Length 55-135 mm, diameter 3-5 mm, segment number 140-190, prostomium
epilobic, first dorsal pore 4/5. Clitellum xxxiii, xxxiv-xlii, xliii. Tubercula puber-
tals in the form of a band xxxiv, xxxv-xli, xlii, xliii. Setae closely paired, poste-
riorly AA.AB.BCCD = 45:5:25:4; c and d in form of genital tumescences in
ix and a and b in xi-xvii, xxix-xxxiv, and xlii-xlv. Male pores on xv, minute,
about at mid BC, with tumescences small and restricted to xv. Seminal vesicles,
four pairs in 9-12, the anterior two pairs smaller. Spermathecae, three pairs with
ducts opening on level c in 8/9-10/11, sometimes an anterior pair opening in
7/8. Colour, lacking. Body cylindrical.

Biology

In Europe Cernosvitov and Evans (1947), Gerard (1964), and Tetry (1938) re-
ported the species from garden soil, meadows, and orchards. With the exception
of Bouche's (1972) study in France, Ap. icterica has been reported infrequently
and in low numbers in Europe. The species is obligatorily amphimictic (Gates,
1968).
Ap. icterica is not known to have any economic importance.

Range

A native of Palaearctis, Ap. icterica is now known from western Europe and
North America (Reynolds, 1976e; Schwert, 1977).

Fig. 7 The known Ontario distribution of A porreclodea icterica.

42

North American Distribution

Ontario (Schwert. 1977), New York (Reynolds. 1976e).

Ontario Distribution (Fig. 7)

WELLINGTON CO. Schwert~(1977). University of Guelph Arboretum, May-Jul 1976, DPS.

Aporrectodea longa (Ude, 1885)

Black head worm Ver a tete noire
(Fig. 8)

1826 Enterion terrestre (non 1820) Savigny, Mem. Acad. Sci. Inst. Fr. 5: 180.

1837 Lumbricus terrestris-Duges, Ann. Sci. Nat., ser. 2, 8: 17, 18.

1845 Lumbricus agricola (non 1842) (part.) Hoffmeister, Regenwiirmer, p. 5.

1885 A llolobophora longa Ude, Z. Wiss. Zool. 43 : 1 36.

1889 Lumbricus terrestris + L. longus-L. Vaillant, Hist. Nat. Annel. 3(1): 113,

121.
1893 A llolobophora terrestris-Rosa, Mem. Ace. Torino, ser. 2, 43: 424, 444.
1900 Helodrilus (A llolobophora) /<?Mg«5-Michaelsen, Das Tierreich, Oligo-

chaeta 10: 483.
1972 A llolobophora longa-Gates, Bull. Tall Timbers Res. Stn. 12: 114.
1972 Nicodrilus longus longus-Bouche, Inst. Natn. Rech. Agron., p. 322.
1975 Aporrectodea longa-Reynolds, Megadrilogica 2(4): 5.

Diagnosis

Length 90-150 mm. diameter 6-9 mm, segment number 150-222, prostomium,
epilobic, first dorsal pore 12/13. Clitellum xxvii, xxviii-xxxiv, xxxv. Tubercula
pubertatis xxxii-xxxiv. Setae closely paired, posteriorly AA.AB.BCCD = 60:
7:28:5; a and b in form of genital setae on genital tumescences in ix, x, xi, xxxi,
xxxiii, xxxiv, and sometimes xii. Male pores on xv with elevated glandular bor-
ders, sometimes extending to xiv and xvi. Seminal vesicles, four pairs in 9-12,
the anterior pairs smaller. Spermathecae, two pairs with short ducts opening on
level c in 9/10 and 10/11. Colour, grey or brown with slight iridescence dorsally.
Body cylindrical and dorsoventrally flattened posteriorly.

Biology

In Europe Cernosvitov and Evans (1947) and Gerard (1964) reported the spec-
ies from cultivated soil, gardens, pastures, and woodlands, and found it to be
abundant in soils bordering rivers and lakes. According to Gates (1972c) Ap.
longa is found in soils with a pH of 4.5 to 8.0, in greenhouses and botanical gar-
dens, lawns, peat bog, in compost and under manure, including chicken yards
and cow yards, and in many other types of soils. The species is known from
caves in Europe.

In appropriate circumstances year-round activity is possible. Feeding, which
takes place nocturnally on the surface of the soil, seems to be selective and
leaves occasionally are dragged into the burrows. Casts are deposited on the
surface and Ap. longa, together with Ap. nocturna (Evans, 1946), is believed re-

43

1st dp —

xxx

XXXV

XXV

xxx

XXXV

Fig. 8 External longitudinal views of Aporrectodea longa showing taxonomic characters. \ Lateral
view, b Ventral view. (NS: Cumberland Co.. cat. no. 9206)

sponsible for the surface castings in England that Darwin studied so intensively.
This species is obligatorily amphimictic with copulation beneath the soil surface
(Gates, 1972a, 1972c, Reynolds, 1974c).

Ap. longa is not known to have been sold or used for bait in North America
and is of little, if any, economic importance.

Range

A native of Palaearctis, Ap. longa is now known from Europe, North America,
Central America, Africa, and Australasia (Gates, 1972c), and also now from
Iceland (Gates 1972a).

44

North American Distribution

Owing to past confusion in the Aporrectodea trapezoides species group (cf.
Gates, 1972a). many of the records of Ap. longa may have been attributed to
other species such as Lumbricus terrestris or Allolobophora terrestris and various
forms of varieties. There are a few reports of limited collections of this species in
North America (cf. Eaton. 1942; Gates, 1953a; Murchie, 1956; Reynolds,
1973c, 1975a-c. 1976a. c; and Reynolds et al., 1974).

New Brunswick (Reynolds, I976d), Nova Scotia (Reynolds, 1975a, 1976a), Ontario (Smith, 1917),
Prince Edward Island (Reynolds, 1975c). Quebec (Reynolds, 1975d, e, 1976c), Alabama (Gates,
1972a), California (Gates, 1972a), Colorado (Gates. 1967). Connecticut (Reynolds, 1973c), Indiana
(Smith. 1917). Maine (Smith. 1917), Maryland (Reynolds, 1974b). Massachusetts (Reynolds, 1977),
Michigan (Murchie. 1956). New Hampshire (Gates. 1972a), New Jersey (Gates, 1972a), New York
(Eaton, 1942), North Carolina (Gates. 1972a). Ohio (Gates, 1972c), Oregon (MacNab and McKey-
Fender. 1947), Pennsylvania (Bhatti, 1965). Tennessee (Reynolds, 1974a), Vermont (Gates, 1972a).

Ontario Distribution (Fig. 9)

Aporrectodea longa is reported here from Ontario for only the second time. The
first report was made by Smith (1917). Both reports are from the greater Tor-
onto area.

YORK CO. Smith (1917). "Edenbrook Park. Islington, under logs and rocks, 30 Apr 72. JWR and
DWR. 0-1-3.

i i ~7

Fig. 9 The known Ontario distribution of Aporrectodea longa.

45

Aporrectodea trapezoides (Duges, 1828)

Southern worm Ver meridional
(Fig. 10)

1828 Lumbricus trapezoides Duges, Ann. Sci. Nat. 15(1): 289.

1917 Helodrilus (Helodrilus) mariensis Stephenson. Rec. Indian Mus. 13: 414.

1923 Allolobophora (Eophila) mariensis-Stephenson, Fauna British India. Oli-

gochaeta, p. 504.
1931 Allolobophora caliginosa trapezoides-Chen, Cont. Biol. Lab. Sci. Soc.

China (Zool.) 7(3): 168.

1941 Allolobophora caliginosa f. trapezoides-Gatts, Proc. California Acad. Sci.
(4), 23: 452.

1942 Allolobophora caliginosa (part.)-Eaton, J. Wash. Acad. Sci. 32(8): 246.
1948 Allolobophora iowana Evans, Ann. Mag. Nat. Hist., ser. 1 1, 14: 515.
1956 Allolobophora (Microeophila) mariensis-Omodeo, Arch. Zool. It. 41 : 184.
1969 Allolobophora longa (part.)-Reinecke and Ryke, Rev. Ecol. Biol. Sol. 6:

515.
1969 Allolobophora caliginosa (part.)-St0p-Bowitz, Nytt. Mag. Zool. 17(2):

191.
1972 Nicodrilus (Nicodrilus) caliginosus meridionalis Bouche, Inst. Natn. Rech.

Agron., p. 334.

1972 Allolobophora trapezoides-G ales, Bull. Tall Timbers Res. Stn. 12:2.

1973 Allolobophora caliginosa f. trapezoides-Plisko, Fauna Polski, no. 1, p. 108.
1975 Aporrectodea trapezoides-Keynolds, Megadrilogica 2(3): 3.

Diagnosis

Length 80-140 mm, diameter 3-7 mm, segment number 93-169, generally >
130, prostomium epilobic, first dorsal pore 12/13, usually. Clitellum xxvii,
xxviii-xxxiii, xxxiv. Tubercula pubertatis xxxi-xxxiii. Setae closely paired, pos-
teriorly AA > AB, DD < V2C. Genital tumescences including a and b setae
only, in ix-xi, xxxii-xxxiv, often in xxviii and occasionally in the region of
xxvi-xxix. Male pores on xv. Seminal vesicles, four pairs in 9-12. Spermathecae,
two pairs opening in 9/10 and 10/1 1. Colour variable and often lighter behind
the clitellum until near the hind end, then deeper, slate, brown, brownish, red-
dish brown, and occasionally almost reddish, but not purple. Body dorsoven-
trally flattened posteriorly so that a cross section is nearly transversely rectangu-
lar with setal couples at the corners.

Biology

This species is found in a wide variety of habitats, according to the material
examined by Gates (1972a). Similar statements have been made by Smith
(1917), Olson (1928), Eaton (1942), Gates (1967), Reynolds (1973 a-c), and Rey-
nolds et al. (1974). According to Gates (1972a, c) Ap. trapezoides is found in the
earth around the roots of potted plants, in gardens, cultivated fields, forest soils
of various types, on the banks of streams, and sometimes in sandy soil. It has
been recorded from caves in North America and Afghanistan, and in California
and Arizona may occur at elevations of 1525 m or more.
Activity may be year round under suitable conditions but it is not possible yet

46

to make a similar statement concerning breeding. Ap. trapezoids is partheno-
genetic, sometimes with pseudogamy, and male sterility is also common (Gates,
1972c; Reynolds, 1974c).

I his species is often found in earthworm culture beds and is one of the five
species commonl) sold and used for bait in North America (Gates, 1972c).

It should be noted that many literature records of this species must be treated
with caution because oi' taxonomic confusion. For a long time Ap. trapezoides
was considered to be a variety of subspecies of A caliginosa but it is unlikely
that all references to A. caliginosa subspecies, variety, or forma, trapezoides do in
fact refer to Ap. trapezoides (cf. Gates, 1972a: 4).

mp xv

xxxv

XXXVIII

Fig. 10 External longitudinal views of Aporrectodea trapezoides showing taxonomic characters, a.
Lateral view, b Ventral view. (ONT. Waterloo Co., cat. no. 8002)

47

Range

A native of Palaearctis, Ap. trapezoides is now known from Europe, North
America, South America, Africa, Asia, and Australasia (Gates, 1972c), and also
from Iceland (Backlund, 1949).

North American Distribution

Alberta (Gates. 1972a). British Columbia (Gates. 1972a). Manitoba (Gates, 1972a). New Brunswick
(Resnolds, 1976d). Nova Scotia (Reynolds, 1975a. 1976a). Ontario (Reynolds. 1972a). Prince Ed-
ward Island (Reynolds, 1975c), Quebec (Reynolds. 1975b. e. 1976c). Alabama (Gales. 1972a),
Alaska (Gates, 1972a) Arizona (Gates, 1972a). Arkansas (Gates. 1972a). California (Gates. 1967).
Colorado (Gates. 1967). Connecticut (Resnolds. 1973c). Delaware (Reynolds. 1973a). District of
Columbia (Gates. 1966). Florida (Gates, 1972a). Georgia (Gates. 1972a). Idaho (Gates. 1967). Illi-
nois (Gates, 1972a), Indiana (Gates, 1972a). Iowa (Gates. 1967). Kentucky (Gates, 1959). Louisiana
(Harman. 1952). Maine (Gates, 1966). Maryland (Reynolds. 1974b). Massachusetts (Reynolds,
1977). Michigan (Gates, 1972a), Minnesota (Gates. 1972a). Mississippi (Gates, 1972a), Missouri
(Gates. 1967), Montana (Reynolds. 1972c), Nebraska (Michaelsen, 1910). Nevada (Gates. 1967),
New Hampshire (Gates, 1972a), New Jersey (Gates, 1972a), New Mexico (Reynolds et al.. 1974),
New York (Gates, 1972a), North Carolina (Cernosvitov. 1942), Ohio (Gates. 1972a). Oklahoma
(Gates. 1967). Oregon (Gates. 1972a). Pennsylvania (Gates, 1972a), Rhode Island (Reynolds,
1973b), South Carolina (Gates, 1972a), Tennessee (Reynolds, 1974a), Texas (Reddell. 1965). Utah
(Gates. 1967), Virginia (Gates, 1972a), Washington (Gates, 1972c), West Virginia (Gates, 1972a),
Wisconsin (Gates. 1972c). Wyoming (Gates. 1967).

Ontario Distribution (Fig. 1 1)

Aporrectodea trapezoides was first reported from Ontario by Reynolds (1972a).

BRANT CO. *Hwy. 53, 2.74 km e of Mt. Vernon, under logs, 1 May 72. JWR, 0-0-3. »Hwy 54, 5.16
km e of Middleport, in ditch, 3May 72. JWR. 1-0-1. BRUCE CO. *Hwy 4, 6.77 km s of Teeswater.
under logs, 5 May 72, JWR, 0-0-0-1. *Hwy 4, .48 km s of Hwy 9, under logs. 5 May 72, JWR, 2-4-4.
Hwy 9, .81 km w of Bervie, under logs in pasture. 5 May 72. JWR. 0-0-1. *Hwy 21. 6.94 km n of
Kincardine, under logs, 5 May 72, JWR, 0-3-0-1. Hwy 21. 2.76 km n of Port Elgin, under logs, 5
May 72. JWR. 6-0-2-1. CARLETON CO. *Ottawa-Carleton Rd 34. Cumberland, under paper. 1 1
May 72. JWR, 1-1-3. "Ottawa-Carleton Rd 34. .97 mi n of Leonard, under paper in wet ditch. II
May 72, JWR, 2-0-4. DUFFERIN CO. *Hwy 9, 2.1 km w of Hwy 104, digging in road bank, 2 May

72, JWR. 0-0-1. *Hwy 9. 3.71 km w of Orangeville. under logs in ditch, 2 May 72. JWR, 0-0-1.
DUNDAS CO. *Hwy 2. 5.48 km e of Iroquois, under logs, 1 1 May 72. JWR. 0-1-0. *Hwy 31. 2.1 km
s of Winchester Springs, under logs, II May 72, JWR, 1-0-2. *Hwy 43, 1.29 km e of Chesterville. un-
der rocks in ditch, 1 1 May 72, JWR, 3-0-2. *Hwy 43, 1.77 km e of Hallville, under logs. 1 1 May 72.
JWR, 1-5-2. DURHAM CO. *Hwy 401, .16 km w of Liberty Rd. Bowmanville, digging under log by
railroad tracks. 15 May 72, JWR, 6-0-4. ELGIN CO. *Hwy" 73. 3.06 km s of Harrietsville. under rot-
ten log, 4 May 72, JWR, 1-3-1. ESSEX CO. Hwy 3. 3.87 km w of Leamington, under lumber in
dump. 4 May 72, JWR, 1-1-1. *Hwv 3, Ruthven. n.e.. in wet ditch by railroad tracks. 4 May 72,
JWR. 0-1-0. *Hwy 3, 1.45 km e of Cottam. under logs. 4 May 72. JWR. 1-0-11. FRONTENAC CO.
•Hwy 15. 4.68 km s of Seeley's Bay. under logs, 16 May 72, JWR. 0-0-1. GLENGARRY CO. *Hwy
401, 7.42 km w of Summerstown Rd, under log, 1 1 May 72. JWR. 0-2-1. *Hwy 34, 2.58 km n of Lan-
caster, under dung in pasture, 1 1 May 72. JWR, 0-0-1. *Hwy 34. 1 1.13 km n of Lancaster, under log,
1 1 May 72, JWR. 4-1-4. GRENVILLE CO. Hwy 43. Merrickville. under logs. 1 1 May 72. JWR, 1-
1-1. HALIBURTON CO. Reynolds ( 1972a). -Hwy 121, 3.22 km e of Tory Hill, dump. 16 May 72,
JWR. 3-2-7. HALTON CO. *Hwy 7. 4.84 km e of Georgetown, under burnt logs in soil. 4 May 73,
JWR. 0-0-1. Halton Co. Rd, 8.87 km n of llalton Co. Rd 8. wet ditch under logs by digging. 4 May

73. JWR, 0-2-0. HASTINGS CO. "Hwy Jet 62 and 620. under logs. 27 Apr 72. JWR. 0-0-1. HU-
RON CO. *Hwy 4, 3.71 km s of Brucefield, under logs and rocks. 5 May 72. JWR, 0-1-0. Hwy 4,
5.16 km s of Clinton, under logs, 5 May 72, JWR. 0-0-2. *Hwy 4. 2.74 km n of Clinton, under logs. 5
May 72. JWR, 0-0-1. KENT CO. 'Hwy 3. 3.55 km e of Wheatley. digging, 4 May 72. JWR, 5-0-0.
LAMBTON CO. 'Hwy 21. Edy's Mills, s.e.. under railroad ties. 4 May 72. JWR. 0-0-1. LANARK
CO. *Hwy 43. 5.32 km w of Smiths falls, under logs, 1 1 May 72. JWR, 5-1-27. *Hwy 7. 5.32 km e of

48

Fig. 1 1 The known Ontario distribution of Aporreclodea irapezoides.

Maberly, under dung in pasture. 1 1 May 72. JWR. 1-0-7. LEEDS CO. *Hwy 15. 3.22 km s of Mor-
ton, under log. 16 May 72. JWR. 0-0-2. *Hwy 15. 4.84 km n of Elgin, digging. 16 May 72. JWR, 0-
0-7. LENNOX AND ADDINGTON CO. *Hwy 500. .81 km n of Denbigh, under logs, 16 May 72,
JWR. 0-0-1 MANITOULIN DIST. »Hwy 68, 5.97 km n of Birch Island, under rocks, 13 May 72,
JWR & JEM. 0-0-1. *Hwy 68. 3.06 km n of Birch Island, under logs and paper in ditch, 13 May 72,
JWR & JEM. 1-2-1. ♦Hwy 68. Birch Island, s.e., under logs. 13 May 72, JWR & JEM, 1-2-4. *Hwy
68, 2.42 km s of Birch Island, under logs, 13 May 72. JWR & JEM, 0-0-3. MUSKOKA DIST. *Hwy
11. Melissa, s.e.. under paper in ditch. 9 May 72. JWR, 0-0-1. NIAGARA CO. 'Niagara Co. Rd 12,
3.06 km s of Grimsby, under paper in wet ditch, 1 May 72, JWR. 0-3-5. NIPISSING DIST. *Hwy
17, 8.39 km e of Warren, house site— digging, 13 May 72, JWR & JEM, 3-3-4. "Hwy 17, 5.48 km e
of Warren, under logs and dung :n pasture, 13 May 72. JWR & JEM, 0-0-1. ONTARIO CO. *Hwy
7, 1.61 km e of Green Rjver. under logs, 26 Apr 72, JWR, 1-0-1. 'Hwy 47, 10.48 km w of Uxbridge,
digging, 9 May 72, JWR, 0-1-0. Duffenn Creek area, day after use of lampncide. 1 1 May 71, TY, 0-
0-6. ROM-I40. OXFORD CO. *Hwy 59, 1.77 km e of Burgessville, under logs and wood chips, 1
Ma) 72. JWR. 0-1-1. PARRY SOL ND DIST. »Hwy 124, 3.06 km e of McKellar, under logs, 9 May
72. JWR. 0-1-3-1. PEEL CO. Hwy 401. 1.3 km w of Dixie Rd. wet ditch, 4 May 73, JWR, 1-1-6.
PERTH CO. Hwy Jet 7, 8 and 59. Shakespeare, under logs, 3 May 72, JWR, 0-0-1. 'Mitchell, next
to Collegiate, under logs. 4 May 72, JWR & DWR. 0-0-1. PRESCOTT CO. *Hwy 34, 5.48 km s of
Vankleek Hill, under logs. 1 1 May 72, JWR, 1-0-2. Hwy 17, 3.22 km w of Alfred, wet ditch, 1 1 May
72, JWR, 2-2-1. PRINCE EDWARD CO. "Hwy 33, Consecon, n.e., dump, 15 May 72, JWR, 0-1-4.
•Hwy 49. 14.84 km n of Picton. under paper in ditch, 15 May 72. JWR, 0-1-4. RUSSELL CO. *Hwy
17. 4^35 km w of Rockland, under logs, 1 1 May 72, JWR, 1-1-0. SIMCOE CO. 'Hwy 89, 1.94 km e
of Rosemont. under logs in wet ditch, 2 May 72, JWR, 0-0-2. *Hwy 27, 5.97 km s of Cookstown. un-
der logs. 2 May 72. JWR. 9-2-1. Hwy 27, 3.22 km s of Newton Robinson, wet ditch, 2 May 72. JWR,
0-1-1. Barne, 14 Greenfield, under logs and digging in garden. 7 May 72, JWR & MK.G, 4-1-0.
'Barrie. park opposite 14 Greenfield, under rocks and grass clippings, 7 May 72. JWR & GWA, 0-

49

0-1. *Hwy 11, 15.16 km s of Severn Bridge, under logs, 9 May 72, JWR. 0-0-1. STORMONT CO.
•Hwy 43, 5.48 km w of Finch, in and under straw in wet ditch, 1 1 May 72, JWR, 1-0-3. SUDBURY
DIST. *Hwy Jet 64 and 69, under rocks, 13 May 72, JWR & JEM, 1-1-4. *Hwy 69. 1.94 km s of
Hwy 607, under rocks, 13 May 72, JWR & JEM, 3-0-3. VICTORIA CO. 'Hwy 35, 8.71 km s of
Hwy 7, under log, 26 Apr 72, JWR, 0-0-2. *Hwy 7. 7.26 km e of Hwy 35, under log, 26 Apr 72, JWR,
0-1-1. "Hwy 7, 1.94 km w of Hwy 46, under logs and dung, 9 May 72, JWR, 0-1-2. WATERLOO
CO. *Hwy 7 and 8, 2.58 km w of New Hamburg, under paper in wet ditch, 3 May 72, JWR, 4-1-8.
Hwy 7 and 8, 1.77 km w of Petersburg, under log, 3 May 72, JWR, 1-0-4. Hwy 85, 1.45 km n of St.
Jacobs, digging, 3 May 72, JWR, 0-0-3. Hwy 86, West Montrose, e.e., under logs, 3 May 72, JWR, 0-
0-1. Waterloo, Beechwood South, on sidewalk after rain (evening), 15 Jun 75, DPS, 5-1-0-3, UW-
0004. Waterloo, Amos Ave., 1 Sep 75, DPS, 0-0-0-1, UW-0003. WELLINGTON CO. *Hwy 24, 5.16
km e of Eramosa, under logs in cedar (Thuja occidentalis) woodlot, 29 Apr 72, JWR, 0-0-1. Hwy 6,
University of Guelph, on wet driveway behind Soil Science Building, 2 May 72, JWR. 0-0-4. *Hwy
6, 10.64 km s of Arthur, under logs, 2 May 72, JWR, 2-1-0. WENTWORTH CO. *Hwy 5, Water-
down, e.e., edge of corn (Zea maize) field, 29 Apr 72, JWR, 0-0-1. YORK CO. 'Hwy 27. 1.45 km n
of Hwy 7, under logs, 29 Apr 72, JWR, 0-0-1. Edenbrook Park, Islington, quantitative study #2
(formalin), 18 May 72, JWR, 2-1-1. Toronto, 1875, GE, 0-0-4, USNM-4559. Toronto, 1875, GE, 0-
0-2, USNM-19704. Scarborough Bluffs, Brimley Rd, garden, 28 Oct 41, JO, 0-0-1, ROM.

Aporrectodea tuberculata (Eisen, 1874)

Canadian worm Ver canadien
(Fig. 12)

1874 Allolobophora turgida f. tuberculata Eisen, Ofv. Vet-Akad. Forh. Stock-
holm 31(2): 43.

1910 Allolobophora similis Friend, Gardener's Chron. 48: 99.

1911 Aporrectodea similis-Friend, Zoologist, ser. 4, 15: 144.

1927 Helodrilus caliginosus trapezoides-B\ake, 111. Biol. Monogr. 10(4): 63.
1930 Allolobophora turgida (part.) + Allolobophora trapezoides (part.)-Borne-

busch, Forstl. Forstfgs. Danm. 1 1 : 94.
1942 Allolobophora caliginosa (part.)-Eaton, J. Wash. Acad. Sci. 32(8): 246.
1952 Allolobophora arnoldi Gates, Breviora, no. 9: 1.

1962 Allolobophora arnoldi + A. nocturna-Omodeo, Mem. Mus. Civ. Sto. Nat.
Verona 10: 92.

1963 Lumbricus terrestris-Camtron and Fogal, Can. J. Zool. 41 : 753.
1969 Allolobophora caliginosa-va.n Rhee, Pedobiologia 9: 130.

1969 Allolobophora caliginosa (part.)-St0p-Bowitz, Nytt. Mag. Zool. 17(2):

191.
1972 Nicodrilus (Nicodrilus) caliginosus alternisetosus Bouche, Inst. Natn.

Rech. Agron., p. 333.

1972 Allolobophora tubereulata-Gates, Bull. Tall Timbers Res. Stn. 12: 44, 45.

1973 Allolobophora caliginosa f. typica (part.)-Plisko, Fauna Polski, no. 1. p.
107.

1975 Aporrectodea tuberculata-Reyno\ds, Megadrilogica 2(3): 3.

50

Diagnosis

Length 90 150 mm. diameter 4-8 mm. segment number 146-194, prostomium
epilobic. first dorsal pore 11 12 or 12 13. Clitellum xxvii-xxxiv. Tubercula pu-
bertals \\\. Kxxi-xxxiii, xxxiv. Setae closely paired, AB =* CD, AA > BC, DD
~ ':C. Genital tumescences absent in xxxi and xxxiii, present in xxx, xxxii, and
wxiv and frequentl) in xxvi. Male pores in xv between b and c. Seminal vesi-
cles, four pairs in 9 12. Spermathecae, two pairs opening on level c in 9/10 and
10 11. Colour, unpigmented, almost white or greyish or sometimes with light
pigmentation on the dorsum. Body cylindrical.

IstdpH

XXV

xxx

Fig. 12 External longitudinal views of Aporrectodea tuberculoid showing taxonomic characters. A.
Lateral view, b Ventral view. (ONT: Sudbury Dist. cat. no. 8544)

51

Discussion

Aporrectodea tuberculoid was described by Eisen in 1874 with specimens ob-
tained from Niagara County (then Welland County), Ontario. Scandinavian
specimens of Ap. tuberculata seem to have first been identified by Eisen as
Allolobophora cyanea (tide Hoffmeister) (Gates. 1972a). Eisen recorded no Type
Locality for Ap. tuberculata: but subsequently this has been designated as Niag-
ara County. Ontario.

Biology

Gerard (1964) reported this species' habitat as pastureland. In the western
United States Gates ( 1967) found it in wet areas near streams and springs where
there was a large concentration of organic matter. In east Tennessee (Reynolds
et al., 1974) Ap. tuberculata was recorded 75% of the time from ditches, or under
logs or debris such as lumber. Gates (1972c) records it from soils of pH 4.8-7.5
including turf, compost, peat, bogs, and rarely manure. In Ontario this species
was found primarily under logs and rocks in all but four eastern counties (cf.
Fig. 11).

Under favourable conditions activity, including breeding, can be year round.
However, Gates (1972c) states that probably throughout New England, New
York, and Canada aestivation and hibernation are climatically imposed with
breeding restricted to spring and late autumn. Reproduction in this species is
obligatorily amphimictic with copulation beneath the soil surface (Reynolds,
1974c).

Ap. tuberculata has been obtained from culture beds on earthworm farms and
being so common in gardens and fields is the species most likely to be dug for
bait in much of Canada.

Range

A native of Palaearctis, Ap. tuberculata is now known from Europe, North
America, South America, Asia, and Australia (Gates, 1972c), and also from Ice-
land (Backlund. 1949).

North American Distribution

Alberta (Gates. 1972a). British Columbia (Wickett. 1967). Manitoba (Gates, 1972a). New Brunswick
(Reynolds. 1976d). Newfoundland (Gates, 1972a), Nova Scotia (Reynolds. 1975a, 1976a). Ontario
(Eisen, 1874). Prince Edward Island (Reynolds, 1975c). Quebec (Reynolds. 1975b. d. e. 1976c). Sas-
katchewan (Gates, 1972a), Alaska (Gates, 1972a), Arizona (Reynolds et al.. 1974). California
(Gates. 1967), Colorado (Gates, 1967), Connecticut (Reynolds, 1973c), Delaware (Reynolds. 1973a).
Florida (Gates. 1972c), Idaho (dates. 1967). Indiana ((rak-s. 1972a), Iowa (Gates, 1967). Maine
(Gates. 1966). Maryland (Reynolds, 1974b). Massachusetts (Reynolds. 1977). Michigan (Gates,
1972a). Minnesota (Gates, 1972a). Montana (Reynolds. 1972c). Nevada (Gates. 1967). New Hamp-
shire (Gates, 1972a). New Jersey (Gates. 1972a). New Mexico (Gates, 1967). New York (Gates,
1972a). North Carolina (Ciates. 1972a). Ohio (Gates. 1972a). Oregon (Gates. 1972a). IVnnsslvania
(Gates, I972ai. Rhode Island (Reynolds, 1973a), 1 enncssee (Reynolds. 1974a). Utah (Gates. 1967).
Vermont (Reynolds, 1976c). Virginia (Gates, 1972a). West Virginia (Gates. 1972a), Wisconsin
(Gates. 1972a), Wyoming (Gates. 1967). New records: Arkansas. Illinois.

52

Fig. 13 The known Ontario distribution of Aporreclodea tuberculata.

Ontario Distribution (Fig. 13)

\I GOMA DIST. *Hw) 17. 7.58 km e of Spanish, under logs, 13 May 72, JWR & JEM. 4-0-2.
•Hw) 17. .48 km e of Spanish, under logs and paper in ditch, 13 May 72, JWR & JEM, 22-16-13-1.
Gargantua Ba>. 1 1 Jun 75. DRB. 0-1-0-1. UW-0001. BRANT CO. *Hwy 53, 5.48 km w of Cathcart,
under railroad ties. 1 May 72, JWR, 0-1-1. *Hwy 53, 4.35 km e of Cathcart, under logs, 1 May 72,
JWR. 10-8-25. "Hwy 53. 2.74 km e of Mt. Vernon, under logs, 1 May 72, JWR, 1-0-7. *Hwy 54, 5.16
km e of Middleport, ditch. 3 May 72, JWR, 0-1-0. *Hwy 2. 5.45 km e of Paris, under lumber and pa-
per. 3 \la\ 72. J\\ R. 4-0-2. BRUCE CO. *Hwy 4, .48 km s of Hwy 9, under logs, 5 May 72, JWR, 0-
0-2. 'Hwy 21, 6.94 km n of Kincardine, under logs, 5 May 72, JWR, 0-2-6. Hwy 21, 2.76 km n of
Port Elgin, under logs. 5 Ma) 72. JWR. 0-0-4. *Hwy 21, 1.61 km e of Elsinore, under logs (pair in
copula), 5 Mas 72. JWR. 4-4-2-1. COCHRANE DIST. Reynolds (1972a). DUFFERIN CO. Hwy 9,
1.61 km eofHw) 10. under logs. 29 Apr 72. JWR, 17-1-0. *Hwy 9, 2.1 km w of Hwy 104, digging in
road bank, 2 May 72. JWR. 4-1-13. *Hwy 9. 10.16 km w of Orangeville, under junk, 2 May 72,
JWR. 0-0-1. •Hwy 9. 3.71 km w of Orangeville, under logs in ditch, 2 May 72, JWR, 6-4-10. *Hwy
10-24. 7.74 km n of Camilla, under logs. 2 May 72. JWR, 6-1-3. *Hwy 89, 10 km e of Primrose, wet
ditch, 2 May 72. JWR. 0-0-10. DUNDAS CO. *Hwy 31. .81 km n of Hwy 43, under logs. 1 1 May
72. JWR. 0-0-3. DURHAM CO. *Hwy 7A. 1 1.77 km e of Hwy 7-12, digging in ditch. 26 Apr 72,
JWR. 0-0-1. *Hw) 7,\. 2.1 km e of Nestleton Station, under logs, 26 Apr 72, JWR, 2-7-3. *Hwy401,
.16 km w of Libert) Rd. Bowmanville. digging under log next to railroad tracks. 15 May 72. JWR, 0-
0-3. 'Hwy 401. .32 km e of Mill St. Newcastle, under logs, 15 May 72, JWR, 0-0-5. "Hwy 401, 1.61
km e of Newtonville. under logs. 15 May 72. JWR. 2-1-8. Hwy 401, 1.61 km w of Hwy 28, under pa-
per, 15 Maj 72, JW R. 2-0-3 El GIN CO. *Hw) 73. 3.06 km s of Harriettsville, under rotten log, 4
Mas 72. JWR. 0-3-9. Hwy 3. 9.19 km e of St. Thomas, under logs. 4 May 72, JWR. 9-3-0. "Hwy 3,
6.77 km e of Wallacetown, under pine logs, 4 May 72. JWR. 13-2-1. 'Hwy 3, 2.1 km e of Wallace-

53

town, under logs. 4 May 72, JWR. 1-0-4. Hwy 3. 3.87 km w of New Glasgow, under logs in aban-
doned school yard, 4 May 72, JWR, 9-4-3-1. ESSEX CO. *Hwy 3. 4.03 km e of Leamington, under
logs, 4 May 72, JWR, 2-2-3. Hwy 3. 3.87 km w of Leamington, under lumber in dump. 4 May 72,
JWR. 5-4-3. *Hwy 3, Ruthven, n.e., wet ditch by railroad tracks. 4 May 72, JWR, 4-1-0. FRONTE-
NAC CO. Reynolds (1972a). "Hwy 7, 3.71 km e of Hwy 38. under log, 11 May 72, JWR. 4-0-6.
•Hwy 2. .97 km w of Westbrook, under logs, 16 May 72. JWR. 0-0-2. *Hwy 2. 1.13 km e of West-
brook, under logs and paper. 16 May 72. JWR. 0-1-3. *Hwy 15, 4.68 km s of Seeley's Bay. under
logs, 16 May 72, JWR, 5-1-1. *Hwy 7. 15.81 km e of Kaladar, under logs. 16 May 72, JWR. 2-0-3.
GRENVILLE CO. *Hwy 2, 5.16 km w of Prescott, under rocks. 10 May 72. JWR. 3-2-1. Hwy 2,
Cardinal, w.e., under log and paper, 1 1 May 72. JWR. 7-5-0. *Hwy 43, 3.87 km e of Kemptville, un-
der logs, 11 May 72, JWR. 0-0-11. GREY CO. *Hwy 21, 7.26 km w of Spnngmount. under rock, 5
May 72, JWR, 0-0-1. *Hwy 6BP, Rockford, w.e.. under logs, 5 May 72. JWR, 1 1-1-2. *Hwy 6. 3.06
km s of Chatsworth, under logs, 5 May 72, JWR, 9-2-10. *Hwy 6, 6.94 km s of Dornoch, under logs,
5 May 72, JWR, 1-1-2. *Hwy 10, 8.06 km s of Flesherton, under logs, 5 May 72. JWR. 5-1-1. HAL-
DIMAND CO. 'Hwy 3. 6.29 km w of Dunnville, under logs, 1 May 72, JWR. 5-1-2. *Hwy 3. 6.13
km w of Cayuga, under logs, 1 May 72, JWR, 0-1-5. *Hwy 54. 2.26 km n of Caledonia, digging, 3
May 72, JWR, 0-1-2. HALIBURTON CO. *Reynolds (1972a). HALTON CO. *Hwy 5. 9.19 km w
of Hwy 10, under log next to barn, 29 Apr 72. JWR. 0-0-1. 'Halton Co. Rd 3, Esquesing Communi-
ty, under rocks and logs next to Town Hall, 4 May 73, JWR, 7-3-0. *Hwy 7. 4.84 km e of George-
town, under burnt logs in soil, 4 May 73, JWR, 0-0-3. Halton Co. Rd 3, 8.87 1cm n of Halton Co. Rd 8,
wet ditch under logs and digging, 4 May 73, JWR, 7-1-15. Halton Co. Rd 3. .48 km n of Halton Co.
Rd 8, under logs in field, 4 May 73, JWR. 5-5-6-1. Rattlesnake Point, 1 Nov 41. JO, 0-0-2, ROM.
HASTINGS CO. "Hwy 620, 5.16 km e of Coe Hill, under logs. 27 Apr 72, JWR, 1-0-1. *Hwy Jet 62
and 620, under logs, 27 Apr 72, JWR, 5-1-1. Hwy 62, 7.42 km n of Bannockburn, under logs, 27 Apr
72, JWR, 6-0-3. 'Hwy 62, 1.61 km n of Hwy 7, under log in wet ditch, 27 Apr 72, JWR, 0-0-1. HU-
RON CO. *Hwy 4, 3.71 km s of Brucefield, under log, 5 May 72. JWR, 0-0-1. 'Hwy 4. 1.29 km s of
Wingham. under logs in wet area. 5 May 72, JWR, 3-0-1. KENORA DIST. Rushing River Provin-
cial Park, stream trickle in campground, 13-14 Jun 71, ROM Field Party, 0-1-0, ROM-I37. KENT
CO. *River Rd, 6.45 km w of Chatham, under logs, 23 Apr 72. JWR & TW, 0-1-1. "Hwy 3. 9.03 km
e of Port Alma, under log, 4 May 72, JWR, 9-0-4. LAMBTON CO. *Hwy 21, 6.45 km n of Dresden,
under logs, 4 May 72, JWR, 3-6-7. *Hwy 21, Edy's Mills, s.e., under railroad ties, 4 May 72. JWR, 0-
0-6. Hwy 21,2.1 km n of Wyoming, under log in wet ditch. 5 May 72, JWR, 0-0-1. *Hwy 7. 1.61 km
n of Arkona, under log, 4 May 72, JWR, 6-4-1. LANARK CO. *Hwy 43, 3.71 km e of Smiths Falls,
under telephone pole in ditch, 1 1 May 72, JWR, 4-3-4. *Hwy 43, 5.32 km w of Smiths Falls, under
logs, 1 1 May 72, JWR, 0-0-13. *Hwy 7, 4.35 km w of Perth, digging under gate post. 1 1 May 72,
JWR, 3-0-12. LEEDS CO. *Hwy 15. 3.22 km s of Morton, under log. 16 May 72, JWR. 2-0-3. *Hwy
15, 5.64 km n of Morton, wet ditch. 16 May 72, JWR. 0-0-11. *Hwy 15, 4.84 km n of Elgin, digging.
16 May 72, JWR, 0-0-13. *Hwy 15, 4.19 km s of Lombardy, under logs and crawling on the surface
during rain. 16 May 72, JWR. 1 1-14-28. *Hwy 15. 7.1 km n of Lombardy, under paper in ditch. 16
May 72, JWR, 0-0-1. LENNOX AND ADDINGTON CO. *Hwy 2, Napanee, w.e.. under rocks be-
hind EEEE Motel, 15 May 72, JWR, 2-1-2. *Hwy 2, 6.61 km w of Hwy 133. under logs, 16 May 72,
JWR, 1-1-4. *Hwy 500, .81 km n of Denbigh, under logs, 16 May 72, JWR, 6-1-4. Hwy 7. Kaladar.
e.e., under lumber in school yard. 16 May 72, JWR. 0-0-2. MANITOULIN DIST. *Hwy 68, 3.22
km n of Little Current, under logs and lumber. 13 May 72. JWR & JEM, 0-0-8. MIDDLESEX CO.
Judd (1964). *Hwy 73, .97 km n of Mossley, under logs, 4 May 72, JWR, 20-2-8. "Hwy 7. 2.26 km s
of Parkhill, under fence posts, 4 May 72, JWR, 0-0-3. MUSKOKA DIST. 'Hwy 1 1. 1.13 km n of Se-
vern Bridge, under logs, 9 May 72, JWR, 2-0-2. *Hwy 11, 11.45 km n of Severn Bridge, under pine
logs, 9 May 72, JWR, 1-1-4. "Hwy 11. 12.9 km s of Bracebridge, under rocks, 9 May 72, JWR, 3-1-2.
•Hwy 1 1, 9.19 km s of Hwy 516, under logs and rocks in wet ditch, 9 May 72, JWR, 3-1-5. *Hwy 1 1,
Melissa, s.e., under paper in ditch, 9 May 72, JWR. 3-1-1. NIAGARA CO. Niagara, 1873, GE. 0-
2-0, USNM-1156. Niagara, 1873, GE, 0-1-3, USNM-1157. •Queen Elizabeth Hwy. 1.13 km w of
Ontario St, Grimsby, under log. 1 May 72, JWR, 0-0-1. NIPISSING DIST. *Hwy 17. 5.48 km e of
Warren, under logs and dung in pasture, 13 May 72, JWR & JEM, 6-1-7. *Hwy 17. 1.29 km e of
Verner, under paper in wet ditch, 13 May 72, JWR & JEM. 3-1-2. Hwy 17, 7.90 km w of North Bay,
digging under rocks, 13 May 72. JWR & JEM, 12-4-17. Temagami, Bearls (Hudson Bay Co. Post),
30 Jun 37, JO & GMN, 0-0-1. ROM-I27. NORFOLK CO. Hwy 6. 4.84 km s of Jarvis, under log, 1
May 72, JWR, 0-0-1. *Hwy 24, 3.06 km n of Hwy 6, under logs, 1 May 72, JWR, 5-4-0. *Hwy 3,

54

1 I 29 km e of Delhi, under logs. 1 Ma) 72, JWR 8-9-0. *Hw) .let 3 and 59. under junk under pine
trees. | Ma) 72, JWR, 7-1-2. NORTHl MBERLANDCO. *ll\\\ 45, 10.69 km s of Norwood, under
logs around Kirn. 28 \pr "1 JWR. 14-2-0 Hwj 45. 4 35 km n of Roseneath, under logs. 28 Apr 72.
JWR, 4-0-6. »Hw) 45,9.03 km n of Baltimore, under logs, 28 Apr 72. JWR. 0-1-1. Mlwy 45, 4.84 km
n of Baltimore, under logs. 2S Apr "2. JWR, 0-0-2. *Hw) 401, 8.06 km w of Grafton, under log, 15
Ma) 72, JWR. 1-1-2. »Hw) 401, 5.64 km e of Grafton, under logs. 15 Ma) 72, JWR, 4-0-18.
•Northumberland-Durham Rd I. .65 km w of Hwj 33. under log, 15 May 72. JWR. 1-0-0. Lakeport,
on pebhK beach ot 1 .ike Ontario. 25 Jul 74, DRB, 0-0-1. UW-0001. ONTARIO CO. *Hwy 7. 1.61
km e of Green Riser, under logs. 2d Apr 72. JWR. 0-0-4. *Hwy 7-12. .81 km n of Myrtle, under logs.
26 Vpr "2. JWR. 4-3-0. *Hw) 4". 8.22 km e of Uxbndge, under login wet area, 9 May 72, JWR, 3-
1-0. »Hwy 47, 10.48 km w of I xhridge. digging. 9 May 72, JWR, 10-4-1-1. *Hwy 7-12, 3.22 km n of
Blackwater. under log-.. ^ Ma) "2. JWR. 2-3-1. DufTerin Creek area, day after use of Lampricide. 1 1
Ma) 71. TY. 0-0-1. ROM-I41. OXFORD CO. »Hw) 59. 6.54 km n ofHwy 3. under logs. 1 May 72,
JWR, 6-0-2 *Hw) <9. 1.13 km s of Curries, under junk. 1 May 72. JWR, 0-0-4. *Hwy 59, 1.77 km e
of Burgesville, under logs and wood ehips. 1 Ma) ^2. JWR. 2-2-4. *Hwy 97, Washington, w.e., un-
der log, 3 Ma) 72, JWR. 0-0-1. *Hwy 59. .32 km n of Hickson. under logs, 3 May 72, JWR, 0-4-2.
PARRY SOL ND DIST. *Hw) 1 1. 4~03 km s of Scotia Rd, under logs, 9 May 72, JWR. 0-3-1. Hwy
II. 3.87 km s of Katrine, under log, 9 Ma) 72, JWR, 0-0-0-1. *Hwy 1 1, 4.52 km s of Burks Falls, un-
der logs. 9 Ma) 72, JWR. 7- 1 1-0. Hw) 520. 5 km w of Burks Falls, under log, 9 May 72, JWR, 0-1-0.
*Hw) 520. Magnetawan, e.e.. under rocks. 9 May 72. JWR, 3-0-3. *Hwy 124, 3.55 km w of Hwy 69,
under logs. ^ Ma) "2. JWR. 0-0-5. PEEL CO. *Hwy 5. .81 km e of Dixie Rd, under logs. 29 Apr 72,
JW R. 5-0-1. *Hw) 5. 4.35 km w of Hwy 10. under mattress, 29 Apr 72. JWR, 0-0-6. *Hwy 24, 8.87
km n of Erin, under log and dung. 29 Apr 72, JWR, 0-4-0. 'Hwy 9, 4.35 km e of Mono Mills, under
logs. 29 Apr 72. JW R^O-lO. Hwy 401. 1.3 km w of Dixie Rd". wet ditch, 4 May 73, JWR, 0-0-3.
PERTH CO. .81 km n of Fullarton, under rock, 29 Apr 72. DWR & LWR, 0-0-1. *Mitchell. next to
Collegiate, under logs. 4 Ma) 72, JWR & DWR. 8-0-6. PETERBOROUGH CO. *Hwy 28, Lakefi-
eld College, waterfront under logs. 27 Apr 72, JWR & CWR. 4-0-1. "Hwy 504, 1.61 km e of Apsley,
under logs. 27 Apr 72. JWR. 1-1-1. "Hwy 504. .81 km w of Lasswade, old house site under logs, 27
Apr 72, JWR, 10-0-7. PRINCE EDWARD CO. *Hwy 33, 1.61 km s of Carrying Place, under log in
wet ditch. 15 May 72. JWR, 1 1-3-1 1. 'Hwy 33, Consecon. n.e., dump, 15 May 72, JWR, 4-0-1. Hwy
33. Bloomfield. e.e.. under paper. 15 May 72, JWR. 1-0-1. *Hwy 49. Picton, n.e., under leaf pile, 15
Ma) 72. JWR. 4-0-1. RENFREW CO. 'Hwy 500. 12.26 km n of Denbigh, under logs, 16 May 72,
JWR. 3-1-6. RUSSELL CO. *2.42 km s of Limonges, under lumber at old house site, 1 1 May 72,
JWR. 0-0-2 SIMCOE CO. 'Hwy 89. 5.48 km e of Alhston, under logs, 2 May 72. JWR, 12-0-5.
*Hwy 89. 1.94 km e of Rosemont. under logs in wet ditch. 2 May 72, JWR, 0-0-2. Barrie, 14 Greenfi-
eld, under logs and digging in garden. 7 May 72. JWR & MK.G, 35-3-4. *Barrie, park opposite 14
Greenfield, under rocks and in grass clippings, 7 May 72, JWR & GWA, 0-1-9. *Hwy 400, .81 km s
of Hwy 103. under logs. 9 May 72, JWR, 2-1-1. SUDBURY DIST. »Hwy 17, 3.06 km e of Nairne
Centre, under logs in ditch. 13'May 72, JWR & JEM. 0-0-5. VICTORIA CO. *Hwy 7, 7.26 km e of
Hwy 35. under logs. 26 Apr 72. JWR. 5-3-3. *Hwy 7, 1.94 km w of Hwy 46. under logs and dung, 9
Ma) 72, JWrR. 8-6-13. 'Hwj 46. 3.71 km n of Hwy 7. under logs, 9 May 72, JWR, 1-3-1. *Hwy 46,
.81 km n of Argyle. under logs, 9 May 72, JWR, 15-1-2. *Hwy 46, 8.22 km n of Argyle, under logs, 9
Ma) "2. JWR. '3-6-9. WATERLOO CO. *Hwy 24A. 1 1.45 km n of Paris, under log, 3 May 72, JWR,
2-1-5 "Hwy 9". Gait. .81 km w of Hwy 24A, under grass clippings, 3 May 72, JWR. 1-0-3. *Hwy97,
3.71 km w of Hwy 401, under logs, 3 May 72, JWR, 4-1-3. Hwy 85, 1.45 km n of St. Jacobs, digging.
3 Mas 72, JW R. 6-1-4. Waterloo, Laurel Creek Conservation Area, 3 Aug 74, DPS, 2-1-6, UW-0007.
Waterloo. Amos Ave.. 1 Sep 75. DPS. 0-0-0-2, UW-0003. WELLINGTON CO. *Hwy 6, 2.26 km n
of Aberfoyle. under logs, 29 Apr 72, JWR, 3-1-4. *Hwy 24, 4.03 km n of Erin, under logs, 29 Apr 72,
JW R. 4-2-1. *Hwy 24. 5.16 km e of Eramosa, under logs in cedar (Thuja occidentals) woodlot, 29
Apr 72. JWR, 1 1-5-2. Hw) 6, University of Guelph, on wet driveway behind the Soil Science Build-
ing. 2 May 72. JWR, 3-0-1 1. *Hw) 6. 1.13 km s of Ennotville, under paper in wet ditch. 2 May 72,
JWR. 6-0-1 W EM WORTH CO. »Hwy 6. 5.32 km n of Hwy 5. under log, 29 Apr 72, JWR, 0-0-1.
YORK CO. Hw) 27. 8.55 km s of Hwy 9, under logs, 29 Apr 72, JWR, 8-0-1. 'Edenbrook Park, Is-
lington, under rocks and logs near stream bank. 30 Apr 72, JWR & DWR, 9-5-1. Hwy 27, Bell's
Lake, under lumber, 2 May 72. JWR, 5-0-4. Edenbrook Park, Islington, quantitative study 1, forma-
lin. 18 May 72. JWR. 10-0-3. Edenbrook Park. Islington, quantitative study 2, formalin, 18 May 72.
JWR, 2-1-6. Toronto. GE, 0-0-4. USNM-4563. Toronto, GE, 0-4-0, USNM-4564.

55

Aporrectodea turgida (Eisen, 1873)

Pasture worm Ver du paturage
(Fig. 14)

1873 Allolobophora turgida Eisen, Ofv. Vet.-Akad. Forh. Stockholm 30(8): 46.

1874 Allolobophora turgida (part.)-Eisen, Ofv. Vet.-Akad. Forh. Stockholm
31(2): 43.

1946 Allolobophora caliginosa-Evans, Ann. Mag. Nat. Hist., ser. 11, 13: 100,
101.

1947 Allolobophora caliginosa-Cemosvitov and Evans, Linn. Soc. Lond., Syn.
British Fauna, no. 6: 13.

1952 Allolobophora caliginosa + A. molita Gates, Breviora, no. 9: 1,3.
1959 Allolobophora caliginosa-Zicsu Acta Zool. Hung. 5(1-2): 172.
1964 Allolobophora caliginosa (in toto)-Gerard, Linn. Soc. Lond., Syn. British
Fauna, no. 6: 27.

1969 Allolobophora caliginosa (part.)-St0p-Bowitz, Nytt. Mag. Zool. 17(2):
191.

1970 Allolobophora caliginosa-Za]onc, Biol. Prace 16(8): 23.

1972 Allolobophora caliginosa f. typica (part.) + A.c.f. trapezoides (part.)-Ed-

wards and Lofty, Biol, earthworms, p. 217.
1972 Nicodrilus caliginosus caliginosus-Boxiche, Inst. Natn. Rech. Agron.. p.

326.

1972 Allolobophora turgida-Gates, Bull. Tall Timbers Res. Stn. 12: 89.

1973 Allolobophora caliginosa f. typica (part.)-Plisko, Fauna Polski, no. 1, p.
107.

1975 Aporrectodea turgida-Keyno\ds, Megadrilogia 2(3): 3.

Diagnosis

Length 60-85 mm, diameter 3.5-5.0 mm, segment number 130-168, prostomium
epilobic, first dorsal pore 12/13 or 13/14. Clitellum xxvii, xxviii, xxix-xxxiv,
xxxv. Tubercula pubertatis xxxi-xxxiii. Setae closely paired,
AA:AB:BC:CD:DD = 3: 1:2:%: 10. Genital tumescences contain a and b only in
xxx, xxxii-xxxiv, and frequently in xxvii. Male pores on xv between b and c.
Seminal vesicles, four pairs in 9-12. Spermathecae, two pairs, with short ducts
opening at level cd in 9/10 and 10/1 1. Colour, unpigmented with the region an-
terior to the crop flesh pink and the remaining segments pale grey, or occasion-
ally with light pigmentation on the dorsal surface. Body cylindrical.

Discussion

Aporrectodea turgida, along with the previous two species, has given oligochae-
tologists great difficulty until a recent publication by Gates (1972a). This species
was first recorded from Ontario (Niagara County) by Eisen in 1874. Four speci-
mens labelled Ap. turgida from Niagara were sent by Eisen to the United States
National Museum (cat. no. 1 157). One specimen may be an aberrant individual
of Ap. turgida, but the other three appear to be Ap. tuberculata (Gates. 1972a). I
have subsequently examined these specimens and confirm this report.

56

1st dp-

t

V

■■

(•

To
• ; j

X 1

9"

1

XV

"^r

-' ' <

3?

-LO

:IT

XXV

XXX

XXXV

Fig. 14 External longitudinal views of Aporrectodea turgida showing taxonomic characters, a. Lat-
eral view. (ONT: Grey Co., cat. no. 7988) B. Ventral view. (ONT: Parry Sound Dist., cat.
no.. 7891)

Biology

Gates (1972c) records this species from a variety of habitats, including gardens,
fields, turf, forest humus, compost, banks of springs and streams, wasteland and
city dumps, and from a cave in West Virginia. The widespread distribution and
variety of utilized habitats for Ap. turgida were recorded also by Eaton (1942),
Cernosvitov and Evans (1947), Murchie (1956), and Gerard (1964). The amaz-
ing water tolerance of this species was reported by Guild (1951) and Reynolds
et al. (1974). in Scotland and the United States respectively. The variety of habi-
tats utilized by Ap. turgida in Ontario can be seen from the collection data.

1<Y
OYAL ONTARIO MUSEUM

57

Activity may be year round but in most North American areas aestivation
and hibernation are probably climatically imposed (Gates, 1972c). Feeding and
cast deposition occur below the surface. Reproduction is obligatorily amphimic-
tic with copulation beneath the soil surface (Reynolds, 1974c).

Ap. turgida is potentially a useful species for fish bait where available in
sufficient numbers.

Range

Known from Europe, North America. South America, Asia, South Africa, and
Australia (Gates, 1972c). Also from Iceland (Backlund, 1949).

North American Distribution

British Columbia (Gates, 1972a), New Brunswick (Reynolds, 1976d), Newfoundland (Gates. 1972a).
Nova Scotia (Reynolds, 1975a, 1976a), Ontario (Eisen, 1874), Prince Edward Island (Reynolds.
1975c), Quebec (Reynolds, 1975b, d, e, 1976c), Alaska (Gates, 1972a), Arizona (Gates, 1967), Cali-
fornia (Gates, 1967), Colorado (Gates, 1967), Connecticut (Reynolds, 1973c), Delaware (Reynolds,
1973a), Florida (Gates, 1972c), Georgia (Gates, 1972a), Idaho (Gates, 1967), Illinois (Garman,
1888), Indiana (Gates, 1972a), Iowa (Gates, 1972a), Kentucky (Gates, 1972a), Louisiana (Harman,
1952), Maine (Gates, 1972a), Maryland (Reynolds, 1974b), Massachusetts (Reynolds, 1977). Michi-
gan (Gates, 1972a), Minnesota (Gates, 1972a), Montana (Reynolds, 1972c), Nevada (Gates. 1967).
New Hampshire (Gates, 1972a), North Carolina (Garman, 1888), Ohio (Gates, 1972a). Oregon
(Gates, 1972a), Pennsylvania (Gates, 1972a), Rhode Island (Reynolds, 1973b), Vermont (Gates,
1972a), Virginia (Gates, 1972a), West Virginia (Gates, 1959), Wisconsin (Ude, 1885). New records:
Manitoba, Arkansas, Mississippi.

Ontario Distribution (Fig. 15)

ALGOMA DIST. *Hwy 17, .48 km e of Spanish, under log in ditch, 13 May 72, JWR and JEM. 0-
0-1. BRANT CO. *Hwy 54, .64 km w of Onondaga, under logs in wet area, 3 May 72, JWR, 13-4-3.
BRUCE CO. *Hwy 4, 6.77 km s of Teeswater, under logs, 5 May 72, JWR, 4-1-9. *Hwy 4. .48 km s
of Hwy 9, under logs, 5 May 72, JWR, 2-2-14. Hwy 9, .81 km w of Bervie, under logs in pasture, 5
May 72, JWR, 0-0-4. *Hwy 21, 6.94 km n of Kincardine, under logs, 5 May 72, JWR, 7-1-6. Hwy 21,
2.58 km n of Tiverton, under lumber, 5 May 72, JWR, 16-2-1. *Hwy 21, 1.61 km e of Elsinore. under
log, 5 May 72, JWR, 0-0-1. CARLETON CO. *Ottawa-Carleton Rd 35, 2.42 km s of Leonard, un-
der logs, 11 May 72, JWR, 16-8-10. COCHRANE DIST. Reynolds, (1972a). DUFFERIN CO.
♦Hwy 89, 10 km e of Primrose, wet ditch, 2 May 72, JWR, 7-0-9. *Hwy 9, 10.16 km w of Orange-
ville, under junk, 2 May 72, JWR, 3-1-6. *Hwy 10-24, 8.06 km n of Orangeville, under logs. 2 May
72, JWR, 3-2-0. DUNDAS CO. *Hwy 2, 5 km w of Iroquois, under log, 1 1 May 72, JWR, 7-0-1.
*Hwy 2, 5.48 km e of Iroquois, under log, 1 1 May 72, JWR, 0-0-1. *Hwy 31, 3.55 km n of Morris-
burg, under lumber, 1 1 May 72, JWR, 4-1-10. *Hwy 43, 1.77 km e of Hallville. under logs. 1 1 May
72, JWR, 2-2-7. DURHAM CO. "Hwy 401, .16 km w of Liberty Rd, Bowmanville, digging under
log next to railroad track, 15 May 72, JWR, 0-1-4-1. *Hwy 401, .32 km e of Mill St. Newcastle, un-
der logs, 15 May 72, JWR, 1-1-6. ELGIN CO. *Hwy 73, 3.06 km s of Harrietsville, under rotten log,
4 May 72, JWR, 10-4-2. *Hwy 73, 6.77 km n of Aylmer, under logs, 4 May 72. JWR, 5-5-14. Hwy 3,
5.16 km w of Talbotville. under log, 4 May 72, JWR, 2-0-1. Hwy 3, 7.9 km w of Frome, under logs, 4
May 72, JWR, 3-2-5. ESSEX CO. Hwy 3, 4.52 km w of Wheatley, under logs, 4 May 72, JWR. 1-1-2.
•Hwy 2, 8.55 km e of St. Joachim, ditch. 4 May 72, JWR, 0-1-8. *Belle River, s.e.. under logs in
dump, 4 May 72, JWR, 7-2-7. FRONTENAC CO. *Hwy 2. .97 km w of Westbrook. under logs, 16
May 72, JWR, 3-0-9. GLENGARRY CO. *Hwy 401, 7.42 km w of Summerstown Rd, under logs,
11 May 72, JWR, 13-2-13. Hwy 34, 2.58 km n of Lancaster, under dung, 11 May 72, JWR. 1-1-0.
♦Hwy 34, 5.64 km n of Alexandria, under logs, 1 1 May 72, JWR, 1-1-3. GRENVILLE CO. "Hwy 2,

58

Fig. 15 The known Ontario distribution of Aporrectodea lurgida.

Johnstown, w.e.. under logs, 1 1 May 72, JWR, 2-0-3. Hwy 2, Cardinal, w.e.. under log, 1 1 May 72,
JWR. 0-0-1. »Hw) 43, 3.87 km e of Kemptville, under logs, 1 1 May 72. JWR, 0-0-3. GREY CO.
•Hwy 21. 7.26 km w of Spnngmount, under rocks, 5 May 72, JWR, 10-1-4. *Hwy 6, 6.94 km s of
Dornoch, under logs. 5 May 72. JWR, 7-0-4. *Hwy 4. 8.22 km e of Durham, under logs, 5 May 72,
JWR, 4-2-5. HALDIMAND CO. *Hwy 3, 9.03 km e of Dunnville, wet ditch, 1 May 72, JWR, 6-0-4.
•Hwj 3. 6.29 km w of Dunnville, under logs, 1 May 72, JWR, 0-0-2. *Hwy 3, 6.13 km w of Cayuga,
under logs. 1 May 72. JWR. 3-1-4. *Hwy 3, 2.74 km w of Nelles Corners, under logs, 1 May 72,
JWR, 7-2-8. »Hw) 54, 2.26 km n of Caledonia, digging, 3 May 72, JWR, 2-2-4. HALIBURTON
CO. 'Reynolds (1972a). H ALTON CO. Hwy 5, 5.81 km e of Hwy 25, under paper and leaves in
ditch. 29 Apr 72, JWR, 2-4-5. *Hwy 7, 4.84 km e of Georgetown, under burnt logs in soil, 4 May 73,
JWR. 4-0-5. *Hw) 401. 4.51 km e of Trafalger Rd, under logs, 4 May 73, JWR, 1-2-1. HASTINGS
CO. Reynolds (1972a). *Hwy 62, 1.61 km n of Hwy 7, under logs in wet ditch. 27 Apr 72, JWR, 2-
2-3. HURON CO. *Hwy 83. 6.77 km w of Russeldale, under logs, 5 May 72. JWR, 15-2-0. *Hwy 4,
1.61 km n of Exeter, wet ditch. 5 May 72. JWR, 8-0-2. Hwy 4. 1.29 km n of Hensall, under corn
(Zea maize)cobs in field, 5 May 72. JWR, 1-1-4. Hwy 4, 5.16 km s of Clinton, under logs. 5 May 72,
JWR. 5-1-5. *Hwy 4. 2.74 km n of Clinton, under logs, 5 May 72, JWR, 2-3-8-1. KENT CO. *River
Rd. 6.54 km v. of Chatham, under logs, 23 Apr 72, JWR & TW, 1-2-1. Hwy 3, 1.27 km w of Palmy-
ra, under logs. 4 May 72. JWR. 9-9-11. LAMBTON CO. *Hwy 21, 6.45 km n of Dresden, under
logs. 4 May 72. JWR. 0-0-3. Hwj 21. 1 1.29 km n of Dresden, under dung in pasture, 4 May 72.
JWR. 0-1-5'. Hwy 21. .64 km s of Petroha, ditch. 4 May 72, 6-1-5. *Hwy 21, Edy's Mills, s.e., under
rail road ties. 4 Ma\ 72. JWR. 0-0-2. Hwy 21, 2.1 km n of Wyoming, under logs in wet ditch, 4 May
72. JWR. 1-2-7. LAN \KK < O. "Hwj 43, 12.1 km e of Smiths I 'alls, under logs. II May 72. JWR, l-
0-1. 'Hwy 7, 5.32 km e of Maberly.'under dung in pasture, II May 72, JWR, 0-0-1. LEEDS CO.

59

*Hwy 15. 2.26 km n of Seeley's Bay. under logs and concrete blocks in ditch. 16 May 72. JWR, 7-0-
17. ♦Hwy 15. 5.64 km n of Morton', wet ditch, 16 May 72. JWR. 3-1-1. *Hwy 15, 4.84 km n of Elgin,
digging, 16 Maj 72. JWR. 6-1-2. *Hwy 15. .97 km s of Portland, under logs, 16 May 72, JWR. 22-0-
17. *Hwy 15. 4.19 km s of Lombardy, under logs and crawling on the surface during rain. 16 May
72. JWR. 2-2-3. ♦Hwy 15. 7.1 km n of Lombardy. under paper in ditch. 16 May 72. JWR, 0-1-2.
MANITOULIN DIST. *Hwy 68, 5.97 km n of Birch Island, under rock, 13 May 72, JWR & JEM,
0-0-1. *Hwy 68, Birch Island, s.e.. under logs. 13 May 72. JWR & JEM, 7-7-10. 'Hwy 68, 2.42 km s
of Birch Island, under logs. 13 May 72. JWR & JEM. 3-0-6. Hwy 68. 3.22 km n of Little Current,
under logs and lumber. 13 May 72^ JWR & JEM. 1-0-4. MIDDLESEX CO. Judd (1964). *Hws 7.
2.26 km s of Parkhill. under fence posts. 4 May 72, JWR, 3-0-13. MUSKOKA DIST. Hwy' 11,
Huntsville, 620 m s of Ravenscliffe Rd. under logs. 9 May 72. JWR, 4-1-1. NIAGARA CO. Eisen
(1874). *Queen Elizabeth Hwy. 1.13 km w of Ontario St, Grimsby, under logs and lumber. I May
72, JWR, 3-1-2. 'Niagara Co. Rd 12. 3.06 km s of Grimsby, under paper in wet ditch. 1 May 72,
JWR. 0-0-2. ♦Hwy 20. 1.61 km e of Smithville. under logs at old house site. 1 May 72. JWR. 5-6-5.
NIPISSING DIST. *Hwy 17. 8.39 km e of Warren, digging at house site. 13 May 72. JWR & JEM,
9-2-9. NORFOLK CO. "Hwy 6. 4.84 km s of Jarvis, under logs. 1 May 72, JWR, 3-2-1. *Hw\ 24.
3.06 km n of Hwy 6, under logs, 1 May 72, JWR, 0-3-0. NORTHUMBERLAND CO. *Hwy 45,
4.84 km n of Baltimore, under logs, 28 Apr 72, JWR, 6-1-7. ONTARIO CO. 'Hwy 7. 1.61 km e of
Green River, under logs. 26 Apr 72, JWR. 13-3-9. *Hwy 47. 10.48 km w of Uxbridge. digging. 9
May 72, JWR, 0-0-1. OXFORD CO. *Hwy 59, 1.94 km s of Norwich, under log, I May 72, JWR, 6-
3-0."*Hwy 59, 1-1-3 km s of Curries, underjunk, 1 May 72, JWR. 4-2-7. *Hwy 59, 1.77 km e of Bur-
gessville. under logs and wood chips. 1 May 72, JWR, 0-5-9. *Hwy 97, Washington, w.e.. under logs.
3 May 72, JWR, 3-1-4. * Hwy 59, .32 km n of Hickson, under logs', 3 May 72. JWR, 13-6-6. PARRY
SOUND DIST. *Hwy 520, 10.32 km e of Magnetawan, under lumber, 9 May 72, JWR, 2-0-6. *Hwy
124, 2.26 km w of Hwy 520, under log, 9 May 72, JWR, 0-0-1. PEEL CO. 'Hwy 5. 4.35 km w of
Hwy 10, under mattress. 29 Apr 72. JWR, 3-1-1. PERTH CO. .81 km n of Fullarton, under rocks, 29
Apr 72, DWR & LWR. 6-0-5. Hwy Jet 7. 8 and 59, Shakespeare, under logs, 3 May 72. JWR. 0-0-5.
♦Mitchell, next to Collegiate, under logs. 4 May 72. JWR & DWR, 6-4-5. PETERBOROUGH CO.
♦Hwy 28, 5.48 km n of Burleigh Falls, under logs, 27 Apr 72, JWR, 3-0-1. PRESCOTT CO. ♦Hwy
34. 5.48 km s of Vanleek Hill, under logs. 1 1 May 72, JWR, 7-1-4. »Hwy 34, 5.81 km n of Vanleek
Hill, under logs, 1 1 May 72, JWR, 13-0-15. Hwy 17, .64 km e of Plantagenet. under log, 1 1 Ma) 72.
JWR. 0-0-1. PRINCE EDWARD CO. #Hwy 33, Consecon, n.e.. dump. 15 May 72. JWR. 0-0-1.
RUSSELL CO. ♦Hwy 17, 7.74 km e of Rockland, under paper in wet ditch, 11 May 72. JWR. 0-0-
1-1. 4Hwy 17, 4.35 km w of Rockland, under logs, 11 May 72. JWR, 4-0-1. '2.9 km e of Embrun.
under logs, 1 1 May 72, JWR, 2-4-5. SIMCOE CO. 4Hwy 89. 1.94 km e of Rosemont, under logs in
wet ditch, 2 May 72, JWR. 7-10-14. Hwy 27. 3.22 km s of Newton Robinson, wet ditch. 2 May 72,
JWR, 2-3-2. ♦Barne. park opposite 14 Greenfield, under rocks and grass clippings, 7 May 72, JWR
& GWA, 0-0-10. ♦Hwy 1 1. 15.16 km s of Severn Bridge, under logs, 9 May 72. JWR. 4-0-4. ♦Hwy
400, .81 km s of Hwy 103. under logs. 9 May 72. JWR, 3-1-1. STORMONT CO. *Hwy 43. 1.29 km
w of Finch, under logs, 1 1 May 72. JWR. 4-0-16. Hwy 43, 3.71 km w of Monkland. under logs, 1 1
May 72, JWR, 0-1-8. ♦Hwy 138, 3.06 km n of St. Andrews West, under log. 1 1 May 72, JWR. 8-2-4.
SUDBURY DIST. *Hwy'l7. 3.06 km e of Nairne Centre, under logs in ditch. 13 May 72. JWR &
JEM, 11-1-4. Hwy 68, Espanola, n.e.. under log and rock. JWR & JEM, 1-0-1. VICTORIA CO.
♦Hwy 35. 8.71 km s of Hwy 7. under log, 26 Apr 72, JWR. 3-0-4. 4Hwy 7, 1.94 km w of Hwy 46. un-
der log and dung, 9 May 72, JWR, 0-2-1. * Hwy 46, .81 km n of Argyle, under logs, 9 May 72, JWR,
18-1-2. Hwy 48, Bolsover, e.e., under paper in wet ditch, 9 May 72, JWR, 0-0-1. WATERLOO CO.
♦Hwy 97, Gait, .81 km w of Hwy 24A. under grass clippings. 3 May 72, JWR, 5-0-2. Hwy 85, 1.45
km n of St. Jacobs, digging, 3 May 72. JWR. 3-1-1. Hwy 86. West Montrose, under logs. 3 May 73.
JWR. 6-2-3. Waterloo, Amos Ave.. 1 Sep 75, DPS, 0-9-4, UW-0003. WELLINGTON CO. ♦Hwy 6.
2.26 km n of Aberfoyle. under logs. 29 Apr 72. JWR. 7-0-2. ♦Hwy 6. 10.64 km s of Arthur, under
logs, 2 May 72, JWR, 19-2-8. WENTWORTH CO. Reynolds (1972a). 4Hwy 5. Waterdown. e.e..
edge of corn (Zca maize) field. 29 Apr 72. JWR, 21-1-4. YORK CO. 4Hwy 48, .32 km n of Steeles
Avenue, under logs, 26 Apr 72, JWR. 1 1-2-4. ♦Hwy 27. 1.45 km n of Hwy 7. under logs. 29 Apr 72,
JWR. 2-1-3. ♦Edenbrook Park, Islington, under logs and rocks near stream bank. 30 Apr 72. JWR &
DWR, 54-9-4. Edenbrook Park, Islington, quantitative study 1, formalin, 18 May 72, JWR, 2-0-3.
Edenbrook Park. Islington, quantitative study 2, formalin, 18 May 72, JWR, 2-0-2. Kings Township.
20 Sep 41, JRD. 6-0-1. ROM-I25.

60

Genus Bimastos Moore, 1893

1893 Bimastos Moore, Zool. Anz. 16: 333.

1895 Bimastos-Moon, J. Morph. 10(2): 473.

1900 Helodrilus (Bimastus)- Michaelsen, Das Tierreich, Oligochaeta 10: 501.

1930 fi/mav/i/.v Stephenson, Oligochaeta. p. 913.

1969 Binmstos-Gmcs. J. Nat. Hist. Lond. 9: 306.

1972 Bimastos-Gates, Trans. Amer. Philos. Soc. 62(7): 86.

1975 Bimastos-Gaics, Megadrilogica 2(1): 4.

Type Species

Bimastos palustris Moore, 1895.

Diagnosis

Calciferous gland, without marked widening in xi-xii, opening into gut in x
through paired vertical sacs. Calciferous lamellae continued onto posterior walls
of sacs. Gizzard, mainly in xvii (ending at ?). Extraoesophageal trunks, joining
dorsal trunk in xii. Hearts, in vii-xi. Nephridial bladders, (/-shaped, closed ends
laterally, ducts passing into parietes near B. Nephropores, inconspicuous, irreg-
ularly alternating (and with asymmetry), between levels somewhat above B and
well above D. Setae, closely paired. Dorsal pores, present from region of 5/6.
Male pores, equatorial in xv. in atrial chambers invaginated deeply into the coe-
lom and bearing acinous glands. Female pores, equatorial on xiv, shortly above
B. Holandric, seminal vesicles in xi-xii. Spermathecae, tubercula pubertatis and
TP glands, lacking. Prostomium epilobic. Pigment red. (after Gates, 1972c: 86;
1975a: 4).

Discussion

This nearctic genus does not normally occur in Canada. The one collection of
four specimens was from an accidental introduction (cf. B. parvus). Over the
years there has been confusion over the spelling of this and other generic names
(cf. Octolasion. p. 104).

Bimastos parvus (Eisen, 1874)

American bark worm Ver americain de l'ecorce

(Fig. 16)

1874 Allolobophora parva Eisen, Ofv. Vet.-Akad. Forh. Stockholm 31(2): 46.
1889 Lumbricus (A llobophora) parvus-L. Vaillant, Hist. Nat. Annel. 3(1): 142.
1893 Dendrobaena constricta (part.)-Friend, Naturalist, p. 19.

1896 Allolobophora parvus (part.: excl. subsp. A. w<fe/)-Ribaucourt, Rev. Suisse
Zool. 4: 80.

1897 Allolobophora constricta var. germinata (part.) Friend, Zoologist, ser. 4, 1 :
459.

61

1st dp

xxv

XXXIII

Fig. 16 External longitudinal views of Bimastos parvus showing taxonomic characters, a Lateral
view, b Ventrolateral view. (FL: Franklin Co., cat. no. 3320)

1900 Allolobophora (Bimastus) parvus-Michaelsen, Abh. Nat. Verh. Hamburg

16(1): 10, 14.
1948 Eisenia parvus + Bimastus beddardi-Pop, Ann. Acad. Sect. §tiint. Geol.

Geogr. Biol., ser. A, 1(9): 123.
1959 Eisenia parva (part. ?)-Zicsi, Acta Zool. Hung. 5(1-2): 170.
1970 Eisenia parva (part. ?)-Zajonc, Biol. Prace 16(8): 23.
1972 Eisenia parva (part. ?)-Bouche, Inst. Natn. Rech. Agron., p. 386.
1972 Bimastos parvus (part. ?)-Edwards and Lofty, Biol, earthworms, p. 215.
1972 Bimastos parvus-Reynolds, Megadrilogica 1(3): 1.

1972 Bimastos parvus-G ates, Trans. Amer. Philos. Soc. 62(7): 87.

1973 Bimastos parvus (part. ?)-Plisko, Fauna Polski, no. 1, p. 99.

1974 Bimastos parvus-Reynolds, J. Tenn. Acad. Sci. 49(1): 17.

Diagnosis

Length 17 65 mm, diameter 1.5-3.0 mm, segment number 65-97, prostomium
epilobic, first dorsal pore 5/6. Clitellum xxiii, xxiv-xxxi, xxxii. Tubercula puber-

62

tatis absent or in the form o\' indefinite ridges on xxiv. xxv, xxvi-xxx. Setae
closely paired. CD = % A B, AA slightly greater than BC, DD = ViC. Male
pores with small, slightly elevated papillae of a yellowish brown colour on xv.
Seminal vesicles in 11 and 12. Spermathecae absent. Colour, reddish dorsally
and yellowish ventrally.

Biology

B. parvus, like all the members of the nearctic genus Bimastos, is found in close
association with decaying logs and leaves, i.e., habitats high in organic matter.
This association has been reported by Cernosvitov and Evans (1947), Causey
(1952). Murchie (1956). and Reynolds et al. (1974). Gates (1972c) recorded it
from soils with pH of 7.5 that were wetted with effluent from human habita-
tions, and from gardens, fields, humus, manure, dumps, and litter in caves.

Activity seems to be possible all year under favourable conditions (Gates,
1972c). B. parvus is obligatorily, or at least facultatively, parthenogenetic (Rey-
nolds, 1974c). Sperm are rarely present, male sterility is apparent, and sperma-
tophores are unknown (Gates, 1972c).

Range

This species is the only known native North American anthropochore, but is
also now known from Europe. North America, South America, Asia, South Af-
rica, and Australia (Gates. 1972c).

II T

Fig. 17 The known Ontario distribution of Bimasios parvus.

63

North American Distribution

Ontario (Reynolds, 1972a). Arkansas (Causey. 1952). California (Michaelsen. 1900b). Illinois
(Gates, 1972c). Kansas (Gates. 1967). Louisiana (Michaelsen. 1900b). Maryland (Reynolds. 1974b).
Massachusetts (Reynolds, 1977), Michigan (Smith and Green. 1916). Missouri (Olson. 1936). Mon-
tana (Gates, 1972c). Nebraska (Swartz, 1929), Nevada (Gates. 1967). New York (Michaelsen,
1900b), Tennessee (Reynolds, 1974a). Texas (Gates. 1956), Washington (Gates. 1972c). Wyoming
(Gates. 1967). New record: Florida.

Ontario Distribution (Fig. 17)

Bimastos parvus has been reported only once from Ontario, by Reynolds
(1972a), as an accidental introduction into the Canadian Agricultural Arbore-
tum in Ottawa.

CARLETON CO. Reynolds (1972a).

Genus Dendrobaena Eisen, 1873

1873 Dendrobaena Eisen, Ofv. Vet.-Akad. Forh. Stockholm 30(8): 53.

1900 Helodrilus {Dendrobaena) (part.) + Helodrilus (Bimastus) (part. ^Michael-
sen, Das Tierreich, Oligochaeta 10: 488, 501.

1930 Dendrobaena (part.)-Stephenson, Oligochaeta, p. 912.

1972 Dendrobaena (part.)-Gates, Trans. Amer. Philos. Soc. 62(7): 88.

1972 Dendrobaena (part.)-Bouche, Inst. Natn. Rech. Agron., p. 388.

1975 Dendrobaena-Gates, Megadrilogica 2(1): 3.

Type Species

Dendrobaena boeckii Eisen by monotypy in 1873 ( = Enterion octaedrum Savi-
gny, 1826).

Diagnosis

Calciferous gland, without sacs opening into gut at vicinity of 10/11, markedly
moniliform in xi-xii. Calciferous sacs, lacking. Gizzard, mainly in xvii. Extraoe-
sophageal trunks, passing to dorsal trunk in vicinity of 9/10. Hearts, in vii-ix.
Nephridial bladders, ocarina-shaped, with bluntly rounded end laterally and
pointed end mesially, ventral side funnel-shaped and narrowing to pass into
parieties at B. Nephropores, obvious, behind first few segments in one rank on
each side, just above B. Setae, not closely paired. Prostomium epilobic. Longitu-
dinal musculature, pinnate. Pigment, red. (after Gates, 1972c: 88 and 1975a: 3).

Discussion

The above diagnosis, after Gates (1972c and 1975a), is for a genus with D.
octaedra as the type species. But another of Savigny's species, Enterion rubidum,
has been congeneric with D. octaedra for decades almost solely because of simi-
larities in their genitalia. As a result of recent studies based on more conservative
somatic anatomy, Omodeo's subgenus Dendrodrilus ( 1956) has been elevated to
full genus status with Enterion rubidum as the type species (Gates, 1975a: 4).

64

Dendrobaena octaedra (Savigny, 1826)

Octagonal-tail worm Ver a queue octogonale
(Fig. 18)

1826 Enierion octaedrum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 183.

1837 Lumbricus octaedrus + L. vetaedrus (laps.)-Duges, Ann. Sci. Nat., ser. 2,

8: 17.24.35.
1845 Lumbricus riparius (part.) Hoffmeister, Regenwiirmer, p. 30.
1849 Lumbricus fla\ i\ cruris R. Leuckart, Arch. Naturg. 15(1): 159.
1871 Lumbricus puter (part.) Eisen. Ofv. Vet.-Akad. Forh. Stockholm 27(10):

959.
1873 Dendrobaena boeckii Eisen, Ofv. Vet.-Akad. Forh. Stockholm 30(8): 53.
1879 Lumbricus /wcA./7-Tauber, Annul. Danmark, p. 69.
1882 Dendrobaena camerani Rosa, Atti Ace. Torino 18: 172.
1885 Octolasion boeckii-Or\ey, Ertek. Term. Magyar Akad. 15(18): 20.

1887 Allolobophora octaedra-Rosa., Boll. Mus. Zool. Torino 2(31): 2.

1888 Dendrobaena octaedra-\ ejdovsky, Entwickgesch. Unters., p. 41.

1889 Lumbricus (Dendrobaena) camerani + L. (D.) boeckii + L. (D.)
octaedrus-L. Vaillant, Hist. Nat. Annel. 3(1): 1 13, 1 18, 1 19.

1893 Allolobophora (Dendrobaena) octaedra (laps.)-Rosa, Mem. Ace. Torino,

ser. 2,43:424,437.
1896 Allolobophora liliputiana + A. alpinula Ribaucourt, Rev. Suisse Zool. 4:

32.33,37.38.
1900 Helodrilus (Dendrobaena) ocfae^/rws-Michaelsen, Das Tierreich, Oligo-

chaeta 10: 494.
1948 Dendrobaena octaedra f. typica + D. o. var. quadrivesiculata Pop, Ann.

Acad. Sect. §tiint. Geol. Geogr. Biol., ser. A, 1(9): 104, 105, 106.
1964 Dendrobaena octahedra (laps.)-Langmaid, Can. J. Soil Sci. 44:34.
1972 Dendrobaena (Dendrobaena) octaedra-Bouche, Inst. Natn. Rech. Agron.,

p. 388.
1974 Dendrobaena octaedra-Gales. Bull. Tall Timbers Res. Stn. 15: 16.

Diagnosis

Length 17-60 mm. diameter 3-5 mm, segment number 60-100, prostomium epi-
lobic, first dorsal pore 4/5-6/7. Clitellum xxvii, xxviii, xxix-xxxiii, xxxiv. Tuber-
cula pubertatis usually on xxxi-xxxiii. Setae widely spread, AA=AB =CD and
DD is slightly greater. On xvi setae a or b are found on small genital tumescenc-
es. Male pores on xv surrounded by small, often indistinct, glandular papillae.
Seminal vesicles in 9. 11, and 12. Spermathecae, three pairs with long ducts on
level with setae d opening in 9/10-11/12. Body cylindrical, with posterior por-
tion octagonal. Colour, red, dark red to purple.

Biology

According to Gates (1972c), Dendrobaena octaedra is found in soils with a pH of
3.0-7.7, mostly in sites little affected by cultivation. Murchie (1956), from his
studies in Michigan, characterized three types of habitat for the species: in sod
or under moss on stream banks, under logs and leafy debris, or in cool, moist

65

1st dp

XXXV /

XXXV

Fig. 18 External longitudinal views of Dendrobaena octaedra showing taxonomic characters, a.
Lateral view (ONT: York Co., cat. no. 7692) B Ventral view (ONT: Sudbury Dist., cat. no.
7732)

66

ra\ ines and upland seepage areas. Gerard ( 1964) found it often under dung and
in soil high in organic matter. These are the types of habitats in which it has
been found in Ontario. In Europe D. octaedra has been found on mountain tops
and in caves, and it is known from botanical gardens and arboretums in Europe
and North America

Under suitable conditions activity, including breeding, is possible the year
round. In Maine, and therefore probably in Ontario also, there are two breeding
periods. The species is said to be surface living, the upper layers being aban-
doned onl\ for aestivation and hibernation. Feeding is selective and sand and
rock particles are rarely found in the intestine.

Parthenogenetic polymorphism is widespread in D. octaedra, probably more
so than in any other lumbricid. A detailed discussion of the numerous morphs
can be found in a recent paper by Gates (1974b).

In Russia (Kursk) D. octaedra has been regarded as an important converter of
leaf substances and it is believed mainly responsible for the decomposition of
oak leaves (Gates. 1972c). In France, where these worms can be found crawling
on bare mountain rocks, the viscous trails are held responsible for trapping li-
chens and thus initiating a lichen-moss-vascular plant succession (Ribaucourt
andCombault. 1906).

Range

A native of Palaearctis, Dendrobaena octaedra is now known from Europe,
North America. Colombia, Mexico, and Asia. This restriction of a peregrine
lumbricid to the Northern Hemisphere is unusual (Gates, 1972c). Also known
from Iceland (Backlund. 1949).

North American Distribution

Alberta (Gates. 1974b). British Columbia (Gates. 1974b). New Brunswick (Reynolds. 1976d), New-
foundland (Gates. 1974b). Nova Scotia (Reynolds. 1975a. 1976a). Ontario (Reynolds. 1972a), Prince
Edward Island (Reynolds, 1975c), Quebec (Reynolds, 1975b. d. e. 1976c), Alaska (Gates, 1974b).
Arkansas (Causey. 1952). California (Gates. 1972c). Colorado (Smith. 1917). Connecticut (Rey-
nolds. 1973c). Delaware (Re\nolds, 1973a). Georgia (Gates. 1974b), Illinois (Smith, 1928). Indiana
(Jo\ner. 1960). Maine (Gates. 1974b). Maryland (Reynolds, 1974b), Massachusetts (Reynolds,
1977). Michigan (Murchie, 1956). Nebraska (Gates. 1967), New Hampshire (Gates. 1972c), New
Jersev (Davies, 1954), New York (Eaton. 1942). North Carolina (Cernosvitov. 1942), Ohio (Olson,
1932). Oregon (Gates. 1974b). Pennsylvania (Bhatti, 1965), Rhode Island (Reynolds, 1973b). Ten-
nessee (Reynolds, 1972b). Vermont (Gates. 1949), Virginia (Smith. 1928). Washington (Altman,
1936). West Virginia (Reynolds, 1974b), Wisconsin (Gates. 1972c), Greenland (Levinsen. 1884).
New records: Manitoba. Colorado. Minnesota.

Ontario Distribution (Fig. 19)

Dendrobaena octaedra was first reported from Ontario by Reynolds (1972a).
This species is found primarily in the Ottawa Valley, extending westward
through the Lake Nipissing area.

CARLETON CO. 'Ottawa-Carleton Rd 34. Cumberland, under paper. 11 May 72. JWR. 0-0-9.
•Ottawa-Carleton Rd 34, .97 km n of Leonard, under paper in wet ditch. 1 1 May 72, JWR. 1-8-3.
HAI.IBl RIO\ ( O. Reynolds (1972a). HA1 TON CO. *Hwy 401.4.51 km e of Trafalgar Rd. un-
der logs, 4 May 73. JWR' 0-1-1. KENORA DIST. Rushing River Provincial Park, stream trickle in

67

Fig. 19 The known Ontario distribution of Dendrobaena octaedra.

campground, 13-14 Jun 71, ROM Field Party, 1-0-0, ROM-I35. LANARK CO. *Hwy 7, 5.32 km e
of Maberly, under dung in pasture, 1 1 May 72. JWR. 0-3- 1 . MUSKOKA DIST. Island in Go Home
Lake, under moss in rotting log near pond^ 8 Oct 75, RLH. 3-0-0, UW-0001. MANITOULIN DIST.
*Hwy 68, 3.06 km n of Birch Island, under logs and paper in ditch, 13 May 72, JWR & JEM, 2-1-1.
NIPISSING DIST. Hwy 17, 15.81 km e of Sturgeon Falls, dump. 13 May 72. JWR & JEM, 0-0-1.
NORTHUMBERLAND CO. *Hwy 45, Baltimore, under junk. 28 Apr 72, JWR, 1-0-1. PARRY
SOUND DIST. *Hwy 124, 4.35 km e of Dunchurch. under logs and rocks. 9 May 72, JWR. 0-3-1.
RENFREW CO. Reynolds ( 1972a). *Hwy 500, 2.42 km s of Hardwood Lake, under logs and dung
in pasture, 16 May 72, JWR, 2-1-2. *2.42 km s of Palmer Rapids, under paper in ditch. 16 May 72,
JWR. 6-6-15. SUDBURY DIST. *Hwy 17, 3.06 km e of Nairne Centre, under logs in ditch. 13 May
72. JWR & JEM, 2-1-2. YORK CO. Edenbrook Park, Islington, quantitative study 2, formalin. 18
May 72, JWR, 0-0-2.

68

Genus Dendrodrilus Omodeo. 1956

1956 Dendrobaena (Dendrodrilus) Omodeo. Arch. Zool. It. 41: 175.

1969 Dendrobaena (part.) St0p-Bowitz, Nytt. Mag. Zool. 17(2): 214.

1972 Dendrobaena (part.) Gates. Trans. Amer. Philos. Soc. 62(7): 88.
\'>~2 Dendrobaena (part.) Bouche, Inst. Natn. Rech. Agron., p. 388.

1973 Dendrodrilus ; Plisko. Fauna Polski. no. 1, p. 78.
1975 Dendrodrilus-G&tts, Megadrilogica 2(1): 4.

Type Species

Enterion rubidum Savigny. 1 826.

Diagnosis

C'alciferous glands, opening into gut ventrally through a pair of sacs posteriorly
just in front of insertion of 10/11. Calcierous lamellae continued along lateral
walls of sacs. Gizzard, mainly in xvii. Extraoesophageal trunks, passing to dor-
sal trunk in xii. Hearts, in vii-xi. Nephridial bladders, (/-shaped loop. Nephro-
pores. inconspicuous, alternating irregularly and with asymmetry on each side
between a level above B and one above D. Setae, not closely paired. Prosto-
mium epilobic. Longitudinal musculature, pinnate. Pigment, red. (after Gates,
1972c: 88 and 1975a: 4).

Discussion

Species of Dendrodrilus formerly congeneric with species in Dendrobaena be-
cause of similarities in genital anatomy are now separated on the basis of differ-
ences in their more conservative somatic anatomy.

Dendrodrilus rubidus (Savigny, 1826)

European bark worm Ver europeen de Tecorce
(Fig. 20)

1826 Enterion rubidum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 182.

1836 Lumbricus xanthurus R. Templeton, Ann. Mag. Nat. Hist. 9: 235.

1837 Lumbricus rubidus-Duges, Ann. Sci. Nat., ser. 2, 8: 17, 23.
1849 ? Lumbricus valdiviensis E. Blanchard, Hist. Chile 3: 43.

1867 ? Hypogeon havaicus Kinberg, Ofv. Vet.-Akad. Forh. Stockholm 23: 101.

1873 Allolobophora norvegica + A. arborea + A. subrubicunda Eisen, Ofv.
Vet.-Akad. Forh. Stockholm 30(8): 48, 49. 51.

1874 Allolobophora tenuis Eisen, Ofv. Vet.-Akad. Forh. Stockholm 31(2): 44.
188 1 Allolobophora fraissei Orley, Zool. Anz. 4: 285.

1881 ? Dendrobaena puter (part.)-Orley, Math. Term. Kozlem. Magyar Akad.

16: 586.
1884 Allolobophora constricta Rosa, Lumbric. Piemonte, p. 38.

69

1st dp

Fig. 20 External longitudinal views of Dendrodrilus rubidus showing taxonomic characters, a Dor-
solateral view, b Ventrolateral view. (ONT: Manitoulin Dist., cat. no. 7502)

70

1884 Lumbricus subrubicunda (part.)-Levinsen. Vidensk. Meddel. Naturhist.
Forh. Copenhagen, ser. 4. 5: 242.

1885 Octolasion constrictum + (). subrubicundum -Or\ey, Ertek. Term. Magyar
Mead. 15(18): 20.21.

1888 Allolobophora putra (part.)-Vejdovsky, Entwickgesch. Unters., p. 41.

1889 Lumbricus (Allolobophora) constrictus L. Vaillant, Hist. Nat. Annel. 3(1):
113.

1 Ss> 1 Allolobophora nordenskioldii (laps.) Michaelsen. Abh. Ver. Hamburg

11(2): 3.
1891 Allolobophora rubicunda (laps.)-Beddard. Proc. R. Phy. Soc. Edinburgh

10: 273.
1893 Allolobophora putris arborea (part. ?)-Rosa, Mem. Ace. Torino, ser. 2, 43:

433.
1893 Dendrobaena const ricta (part.)-Friend, Naturalist, p. 19.
1896 Allolobophora helvetica + A. darwini Ribaucourt, Rev. Suisse Zool. 4: 18,

82.
1900 Allolobophora (Bimastus) constricta-Michaeteen, Abh. Nat. Ver. Hamburg

16(1): 10.
1900 Helodrilus (Dendrobaena) rubidus-Michaeken, Das Tierreich. Oligo-

chaeta 10: 490.
1908 Helodrilus (Bimastus) cw?.?/ncrw5-Michaelsen, Denskschr. Med.-Naturew.

Ges. Jena 13: 41.
1917 Helodrilus (Bimastus) tenuis-Smith, Proc. U.S. Natn. Mus. 52: 157. 182.
1958 Dendrobaena rivu/icola Chandebois. Bull. Soc. Zool. France 83: 159.

1969 Dendrobaena rubida + D. subrubicunda + D. tenuis-Sl0p-Bowilz, Nytt.
Mag. Zool. 17(2): 220. 224. 227.

1970 Dendrobaena rubida var. typica-Zajonc. Biol. Prace 16(8): 22.

1972 Dendrobaena (Dendrodrilus) rubida rubida + D. (D.) rubida tenuis + D.
(D.) subrubicunda-Bouche. Inst. Natn. Rech. Agron., p. 410, 411, 414.

1972 Dendrobaena subrubicunda + Bimastos tenuis + Dendrobaena
rubida-Edwards and Lofty. Biol, earthworms, p. 215. 216.

1973 Dendrobaena (Dendrodrilus) rubida + D. (D.) r. f. typica + D. (D.) r. f.
subrubicunda + D. (D.) r. f. tenuis-Plisko, Fauna Polski, no. l.p. 79, 84,
85.87.

1975 Dendrodrilus rubidus-Reynolds, Megadrilogica 2(3): 3.

Diagnosis

Length 20-90 mm. diameter 2-5 mm, segment number 50-120. prostomium epi-
lobic. first dorsal pore 5/6. Clitellum xxvi, xxvii-xxxi, xxxii. Tubercula puberta-
ls, if present, xxviii, xxix-xxx. Setae widely paired, AB<^CD, and BC = 2 CD.
Male pores on xv between b and c. Seminal vesicles in 9, 11, and 12. Spermathe-
cae, two pairs with short ducts on level with setae c opening in 9/10 and 10/11.
Body cylindrical. Colour, red and darker dorsally.

Biolog)

Dendrodrilus rubidus has been found in a wide range of habitats including gar-
dens, cultivated fields, stream banks, in moss in running water and wells and
springs, peat, compost, and sometimes in manure. It seems acid tolerant. The

71

species is known from caves in Europe and North America, and in greenhouses,
botanical gardens, and the culture beds of earthworm farms (Gates, 1972c). Dd.
rubidus lives in the upper soil layers though on wet nights the worms have been
seen wandering on the soil surface or climbing trees. Under experimental condi-
tions it can withstand prolonged immersion in water (Roots, 1956). Cernosvitov
and Evans (1947) and Gerard (1964) reported it from under the bark of old
trees, and under moss, leaf mould, or rotten wood in moist areas. In Ontario
Dd. rubidus was most frequently found under logs.

Although activity can be year round it is probable that in much of North
America, including Ontario, a winter rest is imposed by the climate. Copulation
has not been properly studied but one published observation records an unusual
position of ventral apposition, head to head and tail to tail (see Gates, 1972c).
Dd. rubidus is facultatively parthenogenetic with male sterility and absence of
spermathecae common (Gates, 1972c; Reynolds. 1974c).

Range

A native of Palaearctis, Dd. rubidus is now know from Europe, North America,
South America, Asia, Africa, and Australasia (Gates, 1972c). Also known from
Iceland (Backlund, 1949).

North American Distribution

Alberta (Gates, 1972c), British Columbia (Gates. 1972c). Manitoba (Gates, 1972c), New Brunswick
(Reynolds, 1976d). Newfoundland (Eaton. 1942). Nova Scotia (Reynolds, 1975a. 1976a). Ontario
(Eisen, 1874). Prince Edward Island (Reynolds, 1975c). Quebec (Reynolds, 1975d. 1976c). Saskat-
chewan (Gates, 1972c), Alabama (Smith, 1917), Alaska (Michaelsen. 1903a), Arizona (Reynolds et
al.. 1974), Arkansas (Causey, 1953), California (Smith, 1917). Connecticut (Reynolds, 1973c), Dela-
ware (Reynolds, 1973a), Hawaii (Ude, 1905), Idaho (Gates, 1967), Illinois (Smith 1917). Indiana
(Heimburger, 1915), Kentucky (Giavannoh, 1933), Louisiana (Harman, 1952), Maine (Gates. 1949),
Maryland (Reynolds, 1974b), Massachusetts (Reynolds. 1977), Michigan (Smith, 1917). Missouri
(Olson, 1936), Montana (Reynolds, 1972c), Nevada (Gates. 1967). New Hampshire (Southern,
1910), New Jersey (Eaton, 1942), New York (Olson. 1940), Ohio (Olson. 1928). Oregon (Gates,
1972c), Pennsylvania (Gates, 1959), Rhode Island (Reynolds, 1973a), Tennessee (Reynolds. 1974a).
Utah (Gates, 1967), Vermont (Reynolds, 1976c), Virginia (Reynolds, 1974b). Washington (Smith.
1917), West Virginia (Gates, 1972c), Wyoming (Gates, 1967), Greenland (Omodeo, 1955a). New
records: Florida. Georgia. Iowa. Minnesota. New Mexico. North Carolina.

Ontario Distribution (Fig. 21)

Dendrodrilus rubidus was the third species reported for Ontario by Eisen (1874).
Later reports of this species in Ontario were made by Stafford (1902), Gates
(1943), Judd (1964), and Reynolds (1972a).

ALGOM A DIST. *Hwy 17. 7.58 km e of Spanish, under logs. 13 May 72, JWR & JEM, 0-0-1. Al-
ona Bay Mine. 4 Jun 71. GM. 0-1-0. Wawa Mine. Wawa, 5 Jun 71. GM. 0-1-0. BRANT CO. *Hwy
54, .64 km w of Onondaga, under logs in wet area, 3 May 72, JWR. 0-1-3. BRUCE CO. *Hwy 4,
6.77 km s of Teeswater, under logs, 5 May 72, JWR, 0-0- 1. Hwy 21, 2.57 km n of Tiverton, under
lumber, 5 May 72. JWR. 0-1-1. 'Hwy 21. 1.61 km e of Elsinore. under logs, 5 May 72. JWR. 0-1-1.
COCHRANE DIST. Smoky Falls, under lumber, 16 Apr 38. RVW. 0-0-2. ROM-I26. DUFFERIN
CO. *Hwy 10-24. 8.06 km n of Orangeville. under logs, 2 May 72. JWR, 0-1-6. DURHAM CO.
•Hwy 401. .32 km e ol Mill St, Newcastle, under logs. 15 May 72. JWR. 0-0-1. ESSEX CO. *Hwy 3,
Ruthven. n.e.. wet ditch next to railroad tracks. 4 May 72. JWR. 0-1-0. *Hwy 2, 8.55 km e of St. Joa-
chim, ditch. 4 May 72. JWR. 0-1-0. FRONTENAC CO. *Hwy 2. .97 km w of Westbrook. under
logs, 16 May 72. JWR. 0-0-1. 'Hwy 7, 15.81 km e of Kaladar. under logs, 16 May 72. JWR, 0-0-1.
GRENVILLE CO. Hwy 43, Merrickville. in grass clippings, 11 May 72, JWR, 0-0-2. GREY CO.

72

Fig. 2 1 The known Ontario distribution of Dendrodrilus rubidus.

•Hwy 6BP. Rockford. w.e., under logs, 5 May 72, JWR 1-0-1. *Hwy 4, 8.22 km e of Durham, under
logs. 5 May 72, JWR, 0-0-2. HALDIMAND CO. *Hwy 3, 6.29 km w of Dunnville, under logs, 1
Ma) 72. JWR. 0-0-1. HALIBURTON CO. Reynolds (1972a). HASTINGS CO. 'Hwy 62, .64 km w
of Maple Leaf, under log, 16 May 72. JWR, 1-1-3. KENT CO. *River Rd, 6.45 km w of Chatham,
under log. 23 Apr 72, JWR & TW, 0-0-1. *Hwy 3. 9.03 km e of Port Alma, under log. 4 May 72,
JWR. 0-T-0. MANITOULIN DIST. Gates (1943). *Hwy 68, 3.22 km n of little Current, under logs
and lumber, 13 May 72. JWR & JEM, 0-0-2. MIDDLESEX CO. Judd (1964). MUSKOKA DIST.
*Hwv 1 1. 1.13 km n of Severn Bridge, under log. 9 May 72. JWR, 0-0-1. *Hwy 103, 2.26 km n of
Hwj 660. under logs in ditch. 9 May 72. JWR, 0-0-3. NIAGARA CO. Niagara. 1873, GE, 0-0-1,
USNM- 18937. N I PISSING DIST. Hwy 17, 15.81 km e of Sturgeon Falls, dump, 13 May 72. JWR &
JEM. 0-0-2. Lake Temagami. under rotten logs. 2 Jul 37. GMN, 1-0-2, ROM-I24. NORFOLK CO.
St. Williams Forestry Experimental Station, 5 Oct 75, DPS, 0-0-1, UW-0001. ONTARIO CO. Du-
ffenn Creek area, day after use of lampricide. 11 May 71, TY, 0-0-1, ROM-I39. OXFORD CO.
•Hwy 59. 1.94 km s of Norwich, under log, 1 May 72. JWR. 0-0-1. PARRY SOUND DIST. *Hwy
124, 3.06 km eof McKellar. under logs, 9 May 72^ JWR, 1-1-1. PRESCOTTCO. »Hwy 34, 5.81 km
n of Vankleek Hill, under logs, 1 1 May 72, JWR, 0-2-6. Hwy 17. .64 km e of Plantagenet, under log,
II \la\ 72. JWR. 0-0-1. RENFREW CO. Reynolds ( 1972a)' *2.26 km n of Schutt, ditch. 16 May 72,
JWR. 0-0-1. '4.19 km s of Palmer Rapids, under paper in wet ditch, 16 May 72, JWR, 0-0-3. *2.42
km s of Palmer Rapids, under paper in ditch. 16 May 72. JWR, 0-0-3. RUSSELL CO. *2.96 km e of
Embrun. under logs. 1 1 May 72, JWR, 0-0-1. SUDBURY DIST. Webbwood Mine, Webbwood, 1
Jun 71. GM. 1-1-0. THUNDER BAY DIST. Jackfish Mine. Terrace Bay, 6 Jun 71. GM, 0-1-1.
Schneber Mine. 6 June 71. GM. 0-1-2. VICTORIA CO. *Hwy 7. 7.26 km eof Hwy 35, under log, 26
Apr 72, JWR, 0-0-1. WATERLOO CO. 'Hwy 97, Gait, .8l'km w of Hwy 24A, under grass clip-
pings. 3 May 72. JWR. 9-0-9. Waterloo, Laurel Creek Conservation Area, 3 Aug 74, DPS, 0-3-5,
UW-0007. WELLINGTON CO. 'Hwy 6, 1.4 km n of Aberfoyle. under logs, 29 Apr 71, JWR. 0-0-3.
•Hwy 6. 10.64 km s of Arthur, under logs, 2 May 72. JWR. 0-0-2. YORK CO. *Hwy 48, .32 km n of
Steeles Avenue, under logs. 26 Apr 72. JWR. 1-6-8. Hwy 27, 8.55 km s of Hwy 9. under log, 29 Apr
72. JWR. 0-0-1. Edenbrook Park. Islington, quantitative study 2, formalin, 18 May 72, JWR, 0-1-3.

73

Genus Eisenia Malm, 1877

1877 Eisenia Malm, Ofv. Salsk. Hortik. Forh. Goteborg 1: 45.

1900 Eisenia (part.)-Michaelsen. Das Tierreich, Oligochaeta 10: 474.

1969 £«e/i/a-Gates. J. Nat. Hist. Lond. 9: 305.

Type Species

Enterion foetidum Savigny, 1826 by Gates (1969).

Diagnosis

Calciferous gland, without sacs, opening into gut behind insertion of 10/11
through a circumferential circle of small pores. Calciferous sacs, lacking. Giz-
zard, mostly in xvii. Hearts, in vii-xi. Nephridial bladders, sausage-shaped or
digitiform, transversely placed. Nephropores, inconspicuous, in two ranks on
each side, alternating irregularly and with asymmetry between a level just above
B and one above D. Setae, closely paired. Prostomium epolobic. Longitudinal
musculature, pinnate. Pigment, red. (after Gates, 1972c: 96; 1975a: 3).

Discussion

Eisenia was erected for three species, Enterion foetidum Savigny, 1826, and
Allolobophora norvegica and A. subrubicunda Eisen, 1873 by Malm (1877) with-
out the designation of a type species. The two Allolobophora species are now sy-
nonyms of Dendrodrilus rubidus. Gates (1969) redefined Eisenia with Eisenia
foetida as type species, but another of Savigny's species, Enterion roseum, has
been congeneric with Eisenia foetida since Michaelsen (1900b) solely because of
spermathecal pore location. If future revisions are based on the more conserva-
tive somatic anatomy these two species will not remain congeneric. Most Euro-
pean workers have followed Pop (1941) and Omodeo (1956) and have transfer-
red E. rosea to Allolobophora. However, on the basis of somatic anatomy it is not
reasonable to place E. rosea in any genus of which Enterion chloroticum Savigny
is the type species. Until this problem is solved I will continue to use Eisenia
rosea to reduce confusion. Recently Gates (1976) transferred E. rosea to the
genus Aporrectodea.

Eisenia foetida (Savigny, 1826)

Manure worm Ver du fumier
(Fig. 22)

1826 Enterion fetidum {con. foetidum) Savigny, Mem. Acad. Sci. Inst. Fr. 5:
182.

1835 Lumbricus semifasciatus Burmeister, Zool. Hand. Atl. 33: 3.

1836 Lumbricus annularis R. Templeton, Ann. Mag. Nat. Hist. 9: 234.

1837 Lumbricus foetidus-Duges, Ann. Sci. Nat., ser. 2, 8: 17, 21.
1842 Lumbricus olidus Hoffmeister, Verm. Lumbric, p. 25.

1 849 ? Lumbricus luteus Blanchard, Hist. Chile 3 : 42.

74

1st dp

J

Fig. 22 External long.tud.nal views of Eiscnia foetida showing taxonomic characters, a Lateral
view b Ventral view. (ONT: Nipissing Dist.. cat. no. 7604)

1873 ? Lumbricus rubro-fasciatus Baird. Proc. Linn. Soc. Lond. 1 1 : 96.

1873 Allolobophorafoetida-E\sen, Ofv. Vet.-Akad. Forh. Stockholm 30(8): 50.

1877 Lumbricus annulatus Hutton, Trans. N.Z. Inst. 9: 352.

1877 Eisenia foetida-Ma\m. Ofv. Salsk. Hortik. Forh. Goteborg 1 : 45.

1887 Endnlus annulatus W.W. Smith, Trans. N.Z. Inst. 19: 136.

1889 Lumbricus lAllobophora) annulatus + L. (A.) foetidus-L. Vaillant, Hist.

Nat. Annel. 3(1): 147. 149.
1913 Helodrilus (Eisenia) foetidus-Michaeken, Zool. Jb. Syst. 34: 55 1 .
1963 Eisenia foetida var. unicolor Andre, Bull. Biol. Fr. Belg. 81:1.
1972 Eisenia fetida fetida + E. f. andrei Bouche, Inst. Natn. Rech. Agron.. p.

380,381.

75

Diagnosis

Length 35-130 mm (generally < 70 mm), diameter 3-5 mm, segment number
80-110, prostoinium epilobic, first dorsal pore 4/5 (sometimes 5/6). Clitellum
xxiv, xxv, xxvi-xxxii. Tubercula pubertatis on xxviii-xxx. Setae closely paired,
AB = CD, BC < AA, anteriorly DD = ViC but posteriorly DD<xhC. Genital
tumescences may be present around any of the setae on ix-xii. usually around
setae a and b of xxiv-xxxii. Male pores with large glandular papillae on segment
xv. Seminal vesicles, four pairs in 9-12. Spermathecae, two pairs with ducts
opening near mD line in 9/10 and 10/1 1. Colour variable, purple, red, dark red,
brownish red, sometimes alternating bands of red-brown on dorsum with pig-
mentless yellow intersegmental areas. Body cylindrical.

Biology

Olson (1928) found this species in manure and decaying vegetation where mois-
ture concentrations were high. Cernosvitov and Evans (1947) and Gerard ( 1964)
recorded its habitats as manure, compost heaps, and soil high in organic matter,
as well as forests, gardens^ and under stones and leaves. Murchie (1956) re-
ported E. foetida from manure and bait castaways but never from what he con-
sidered "natural" habitats. In Tennessee Reynolds et al. (1974) recorded scat-
tered distribution of this species with it occurring most commonly under logs
and debris and at roadside dumps. According to Gates (1972c) the available
records give a pH range of 6.8-7.6, and while in Scandinavia it has been consid-
ered a species dependent on human culture, E. foetida is known from caves in
Europe and North America, and Russian records report it from taiga, forests,
and steppes. In Ontario E. foetida was found most frequently under logs and
usually not far from human habitation. There are few data available concerning
its natural habitat in North America.

There is little information on rest periods in the life-cycle (Gates, 1972c). One
assumes that under favourable conditions activity can occur throughout the
year. Feeding is selective in that there is minimal ingestion of earth. Copulation
is subterranean and although the species has been thought to be obligatorily
amphimictic, uniparental reproduction is possible, though very rare (Gates,
1972c). Experimental self-fertilizing was demonstrated by Andre (1963). E.
foetida has a maximum life expectancy of 4-5 years, although between 1 and 2
years is more usual.

Eisenia foetida has been reared on earthworm farms and sold in every Cana-
dian province and American state for fish bait. Harman (1955) reported on the
commercial aspects of this species.

Range

A native of Palaearctis, E. foetida is now known from Europe, North America,
South America, Asia, Africa, and Australasia (Gates, 1972c). Also known from
Iceland (Backlund, 1949).

North American Distribution

British Columbia (Gates, 1942), Nova Scotia (Reynolds, 1975a. 1976a), Ontario (Reynolds, 1972a),
Quebec (Reynolds. 1975e, 1976c). Alabama (Gates, 1972c). Arizona (Gates. 1967). Arkansas (Cau-
sey, 1952), California (Gates. 1943), Connecticut (Reynolds. 1973c), District of Columbia (Gates,

76

Fig. 23 The known Ontario distribution of Eiseniafoetida.

1972c). Florida (Harman. 1955), Georgia (Harman. 1955). Hawaii (Michaelsen, 1900b). Illinois
(Smith. 1928). Indiana (Heimburger. 1915), Iowa (Harman. 1955), Kentucky (Giavannoli, 1933),
Louisiana (Harman, 1952). Maine (Gates, 1966), Maryland (Reynolds, 1974b), Massachusetts (Rey-
nolds. 1977). Michigan (Smith and Green. 1916). Minnesota (Mickel, 1925), Mississippi (Harman,
1955), Missouri (Olson, 1936), Nebraska (Swartz, 1929), New Hampshire (Gates, 1972c), New Jer-
se\ (Eaton. 1942). New York (Olson. 1940). North Dakota (Harman, 1955), Ohio (Olson, 1928),
Oklahoma (Harman. 1954). Oregon (MacNab and McKey-Fender, 1947), Pennsylvania (Andrews,
1895), Rhode Island (Reynolds, 1973b), South Carolina (Harman, 1955), South Dakota (Gates,
1967), Tennessee (Reynolds, 1974a), Texas (Harman, 1955), Utah (Gates, 1967), Vermont (Rey-
nolds. 1976c). Virginia (Gates, 1949), Washington (Altman, 1936), West Virginia (Gates, 1967),
Greenland (Michaelsen. 1892). New record: New Mexico.

Ontario Distribution (Fig. 22)

First reported by Stafford (1902), and suspected from Haliburton County (Rey-
nolds. 1972a), Eiseniafoetida is not widespread in Ontario.

ESSEX CO. *Hwy 3, 1.45 km e of Cottam, under logs, 4 May 72, JWR, 0-0-2. HALIBURTON CO.
' Reynolds (1972a). KENT CO. 'River Rd. 6.45 km w of Chatham, compost pile, 23 Apr 72. JWR &
rW, 2-1-2 NIAGARA CO. *Hwy 20. 1.61 km w of Smithville. old house site. 1 May 72. JWR, 0-
0-1 MPISSING DIST. 'Hwy 17, Sturgeon Falls, w.e., under logs. 13 May 72, JWR & JEM. 0-1-5.
PERTH CO. 'Mitchell, next to Collegiate, under log in farmyard, 4 May 72, JWR & DWR, 0-0-1.
PRESCOTT CO. 'Hwy 34, 2.58 km n of Vankleek Hill, under logs and rocks, 1 1 May 72, JWR, 4-
8-1. SUDBURY DIST. 'Hwy 69, 6.61 km s of Hwy 637, under grass clippings, 13 May 72, JWR &
JEM. 0-2-0.

77

Eisenia rosea (Savigny, 1826)

Pink soil worm Ver rose du sol
(Fig. 24)

1826 Enterion roseum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 182.

1837 Lumbricus roseus-Dugbs, Ann. Sci. Nat., ser. 2, 8: 17, 20.

1845 Lumbricus communis anatomicus (part.) HofTmeister, Regenwiirmer, p.

28.
1873 Allolobophora mucosa (part.) Eisen, Ofv. Vet.-Akad. Forh. Stockholm

30(8): 47.
1875 Lumbricus aquatilis Vejdovsky, SB Bohm. Ges., p. 199.
1879 Lumbricus muscosus-Tauber, Annul. Danmark, p. 68.
1882 Lumbricus carneus (err. non Enterion carneum Savigny, 1826)-Vejdovsky,

Brunnenw. Prague, p. 51.

1884 Allolobophora carnea- Vejdovsky, Syst. Morph. Oligochaten, p. 61.

1885 Allolobophora aquatilis + A. aguatilis-Orley, Ertek. Term. Magyar Akad.
15(18): 24, 28.

1893 Allolobophora rosea-Rosa, Mem. Ace. Torino, ser. 2, 43: 424, 427.

1896 Allolobophora danieli rara/'-Ribaucourt, Rev. Suisse Zool. 4: 39.

1900 Eisenia rasea-Michaelsen, Das Tierreich, Oligochaeta 10: 478.

1903 Helodrilus (Bimastus) bimastoides Michaelsen, Mitt. Naturh. Mus. Ham-
burg 19: 13.

1907 Helodrilus (Bimastus) indicus Michaelsen, Mitt. Naturh. Mus. Hamburg
24: 188.

1917 Helodrilus (Eisenia) rasews-Smith, Proc. U.S. Natn. Mus. 52(2174): 165,
166.

1922 Helodrilus (Allolobophora) prashadi Stephenson + A. (Bimastus)
zW/ca-Stephenson, Rec. Indian Mus. 22: 440, 441.

1940 Eisenia rosea f. typica + E. r. f. macedonica + Allolobophora hataii -I- A.
harbinensis + A. dairensis + A. jeholensis Kobayashi, Sci. Rep. Tohoku
Imp. Univ., ser. 4, 15: 285-287, 288-289, 290-291, 291-293, 293-295.

1 949 Eophila kulagini Malevic, Dokl. Akad. Nauk SSSR (Biol.) 47 : 400.

1967 Allolobophora rosea var. alpina Vedovini, Bull. Soc. Zool. Fr. 92: 793.

1970 Allolobophora rosea-Zapnc, Biol. Prace 16(8): 23.

1972 Allolobophora rasea-Edwards and Lofty, Biol, earthworms, p. 217.

1972 Allolobophora rosea rosea-Bouche + A. r. vedovinii Bouche, Inst. Natn.
Rech. Agron., p. 418, 423.

1976 Aporrectodea rosea-Gaies, Megadrilogica 2(12): 4.

Diagnosis

Length 25-85 mm, diameter 3-5 mm, segment number 120-150, prostomium
epilobic, first dorsal pore 4/5. Clitellum, somewhat flared ventrally xxv,
xxvi-xxxii. Tubercula pubertatis usually xxix-xxxi. Setae closely paired,
AA>BC<DD, AByCD, anteriorly DD = ViC, posteriorly DD = ViC. Male
pores with elevated glandular papillae in xv with male tumescences extending
over xiv and xvi. Seminal vesicles, four pairs in 9-12. Spermathecae, two pairs
with short ducts opening near mD line or halfway between mL and d in 9/10
and 10/1 1. Body cylindrical, except in clitellar region. Unpigmented, but colour

78

1st dp — -

Fig. 24 External longitudinal views of Eisenia rosea showing taxonomic characters, a. Dorsolateral
view. (ONI : Perth Co., cat. no. 7664) b Ventrolateral view. (ONT: Nipissing Dist., cat. no.
7535)

79

appears rosy or greyish when alive, and white when preserved.

Biology

Cernosvitov and Evans (1947) and Gerard (1964) recorded E. rosea in soil,
fields, gardens, pastures, and forests, under leaves and stones, and frequently on
river and lake banks. Gates (1972c) mentioned soils of pH 4.9-8.0 and recorded
that it is found often enough in conditions that were thought to justify charac-
terization as "amphibious". Murchie (1956) states, "Eisenia rosea, although
showing considerable adaptability in habitat requirements, is to be regarded pri-
marily as a true soil species". The results of this survey would tend to confirm
Murchie's statement. In soil under logs was the most common habitat of this
species in Ontario. E. rosea is one of the cosmopolitan species that have been in-
troduced by Europeans into all parts of the world. It is known also from caves
in Europe. Asia, and North America, as well as from botanical gardens and
greenhouses. It is the only species widely distributed in the virgin steppes of
Russia and the mountains of the Caucasus (Gates, 1972c).

In suitable conditions activity, including breeding, is possible the year round.
But in northern parts of the range there is a resting stage during winter cold and
summer drought in which both hibernation and aestivation are spent tightly
coiled in a small pink ball (Gates, 1972c). According to Thomson and Davies
(1974), E. rosea produces surface casts, despite some contrary statements in the
literature. The species is parthenogenetic and biparental reproduction of an-
thropochorous morphs is unknown (Gates, 1974c, Reynolds, 1974c). Cernosvi-
tov (1930) reported degeneration, phagocytosis, and reabsorption of sperm and
Tuzet (1946) recorded atypical spermatogenesis. Evans and Guild (1948) reared
isolated individuals to sexual maturity which then produced fertile cocoons.

Eisenia rosea is the primary host of the cluster fly, Pollenia rudis (Fabr.)
(Yahnke and George, 1972; Thomson and Davies, 1973a).

Range

A native of Palaearctis, E. rosea is now known from Europe, North America,
South America, Africa, Asia, and Australasia. It also occurs in Iceland (Back-
lund, 1949). Generally, therefore, it is a cosmopolite although apparently absent
from tropical lowlands (Gates, 1972c).

North American Distribution

Alberta (Gates, 1972c), British Columbia (Gates. 1974c), Labrador (Gates, 1974c). New Brunswick
(Reynolds, 1976d). Nova Scotia (Reynolds, 1975a. 1976a), Ontario (Reynolds. 1972a), Prince Ed-
ward Island (Reynolds, 1975c). Quebec (Reynolds, 1975d, e, 1976c), Arizona (Smith. 1917). Arkan-
sas (Causey, 1952). California (Smith, 1917), Colorado (Gates. 1967). Connecticut (Reynolds,
1973c), Delaware (Reynolds, 1973a). Florida (Gates, 1974c), Georgia (Smith. 1917), Hawaii (Gates,
1972c), Idaho (Gates, 1967), Illinois (Smith. 1917), Indiana (Heimburger. 1915), Iowa (Gates. 1967),
Kansas (Gates, 1974c), Kentucky (Gates. 1959), Louisiana (Smith, 1917). Maine (Smith. 1917).
Maryland (Reynolds. 1974b). Massachusetts (Reynolds. 1977). Michigan (Murchie, 1956), Minne-
sota (Gates. 1974c), Missouri (Olson. 1936), Montana (Reynolds. 1972c). Nevada (Gates. 1967),
New Hampshire (Gates. 1972c), New Jersey (Davies. 1954). New Mexico (Gates, 1967). New York
(Smith. 1917), North Carolina (Pearse, 1946). Ohio (Olson. 1928). Oklahoma (Harman, 1954). Ore-
gon (MacNab and McKey-Fender. 1947), Pennsylvania (Bhatti, 1965), Rhode Island (Reynolds,
1973b), Tennessee (Reynolds, 1974a). Texas (Gates, 1972c), Utah (Gates, 1972c). Vermont (Gates,
1949), Virginia (Gates, 1959), Washington (Gates. 1972c), West Virginia (Reynolds. 1974b). Wyo-
ming (Gates, 1967 1.

80

Fig. 25 The known Ontario distribution of Eisenia rosea.

Ontario Distribution ( Fig. 25)

This species was first recorded from Ontario by Stafford (1902) and then by
Judd (1964) and Reynolds (1972a). In the present survey it was found in all but
four counties in southern Ontario, but additional collecting under favourable
conditions should produce specimens from these counties.

BRANT CO. *Hwy 53. 5.48 km w of Cathcart. under railroad ties, 1 May 72, JWR, 1-1-3. *Hwy 54,
.64 km w of Onondaga, under log in wet area, 3 May 72, JWR, 0-0-1. BRUCE CO. *Hwy 4, 6.77 km
s of Teeswater. under logs. 5 May 72. JWR, 4-1-1. Hwy 21, 2.57 km n of Tiverton, under lumber, 5
Ma> 72, JWR, 0-6-2. *Hwy21, 1.61 km e of Elsinore, under logs, 5 May 72, JWR, 3-0-0-1. CARLE-
TON CO. Reynolds (1972a). *Ottawa-Carleton Rd 35. 2.42 km s of Leonard, under logs, 1 1 May 72,
JWR, 3-1-3. DUFFERIN CO. Hwy 9. 1.61 km e of Hwy 10. under logs, 29 Apr 72, JWR, 0-0-6.
•Hwy 9. 2.1 km w of Hwy 104. digging in road bank, 2 May 72. JWR, 0-0-1. *Hwy 9, 3.71 km w of
Orangeville, under logs in ditch, 2 May 72. JWR. 0-2-1. *Hwy 10-24, 8.06 km n of Orangeville. un-
der logs, 2 Ma> 72, JWR. 3-0-3. DUNDAS CO. *Hwy 2, 5 km w of Iroquois, under log, 1 1 May 72,
JWR. 5-3-5. *Hwj 2. 5.48 km e of Iroquois, under logs, 11 May 72, JWR, 0-0-15. *Hwy 31, 3.55 km
n of Mornsburg. under lumber. 1 1 May 72, JWR. 0-0-2. DURHAM CO. Hwy 7A, 2.1 km e of Nes-
tletown Station, under logs. 26 Apr 72. JWR. 0-2-1. *Hwy 401. .61 km w of Liberty Rd, Bowman-
ville. digging under log next to railroad tracks. 15 May 72. JWR, 2-0-3. *Hwy 402, .32 km e of Mill
St. Newcastle, under logs, 15 May 72, JWR, 0-0-2. *Hwy 402, 1.61 km e of Newtonville, under log,
15 May 72, JWR, 0-0-1. ELGIN CO. *Hwy 73, 3.06 km s of Harrietsville, under rotten log, 4 May
72, JWR, 3-1-6. *Hw> 73. 6.77 km n of Aylmer, under logs, 4 May 72, JWR, 0-0-6. Hwy 3, 9.19 km s
of St. Thomas, under logs. 4 May 72, JWR. 0-2-8. Hwy 3, 5.16 km w of Talbotville, under log. 4 May
72. JWR. 5-0-6. 'Hwv 6.77 km e of Wallacetown, under pine logs, 4 May 72. JWR, 2-2-0. Hwy 3. 7.9
km w of Frome. under logs, 4 May 72, JWR. 1-2-3. ESSEX CO. *Hwy 3, 1.45 km e of Cottam, un-

81

der logs, 4 May 72, JWR, 0-0-4. *Belle River, s.e., under logs in dump. 4 May 72, JWR, 1-0-1.
FRONTENAC CO. *Hwy 15, 4.68 km s of Seeley's Bay. under logs, 16 May 72, JWR, 0-0-3.
GLENGARRY CO. *Hwy 34. 1 1.13 km n of Lancaster, under log, 11 May 72, JWR. 2-1-4. 'Hwy
34, 5.64 km n of Alexandria, under logs, 11 May 72, JWR, 3-0-13. GRENVILLE CO. *Hwy 2,
Johnstown, w.e., under logs, 11 May 72, JWR, 5-0-6. Hwy 43. Merrickville, under log. 11 May 72,
JWR, 0-0-1. GREY CO. *Hwy 6BP. Rockford, w.e.. under log, 5 May 72, JWR, 0-0-1. *Hwy 6, 3.06
km s of Chatsworth, under logs, 5 May 72, JWR. 1-0-3. *Hwy 10, 8.06 km s of Flesherton. under
logs, 5 May 72, JWR, 1-0-1. HALDIMAND CO. *Hwy 3. 6.13 km w of Cayuga, under log. I May
72, JWR, 0-0-1. *Hwy 54, 2.26 km w of Caledonia, digging, 3 May 72, JWR, 0-0-2. HALIBURTON
CO. 'Reynolds (1972a). HALTON CO. *Hwy 5, 9.19 km w of Hwy 10. under logs next to barn. 29
Apr 72, 1-1-0. 'Halton Co. Rd 3, Esquesing Community, under rock, next to Town Hall, 4 May 73,
JWR. 0-1-0. *Hwy 7, 4.84 km e of Georgetown, under burnt log in soil, 4 May 73. JWR. 0-0-1.
HASTINGS CO. *Hwy62, 1.61 km n of Hwy 7. under log in wet ditch, 27 Apr 72. JWR. 0-0-1. HU-
RON CO. *Hwy 83, 6.77 km w of Russeldale, under logs, 5 May 72. JWR, 0-2-5-1. *Hwy 4. 3.71 km
s of Brucefield, under logs and rocks, 5 May 73, JWR. 3-3-7. Hwy 4, 1.29 km n of Hensall. under
corn (Zea maize) cobs in pasture, 5 May 72, JWR, 0-0-1. Hwy 4, 5.16 km s of Clinton, under logs, 5
May 72, JWR, 4-1-3. *Hwy 4, 2.74 km n of Clinton, under log, 5 May 72, JWR, 0-0-1. KENT CO.
Hwy 3, 1.77 km w of Palmyra, under logs, 4 May 72, JWR, 1-1-2. *Hwy 3, 9.03 km e of Port Alma,
under log, 4 May 72, JWR, 0-1-0. *Hwy 3. 3.55 km e of Wheatley. digging, 4 May 72. JWR, 4-1-1.
LAMBTONCO. *Hwy 21, 6.45 km n of Dresden, under log, 4 May 72, JWR, 0-0-1. Hwy 21. 11.29
km n of Dresden, under dung in pasture, 4 May 72, JWR, 0-0-1. Hwy 21, .64 km s of Petrolia, ditch,
4 May 72, JWR, 3-0-1. *Hwy 21, Edy's Mills, s.e., under railroad ties, 4 May 72, JWR, 0-0-1. "Hwy
7, 1.61 km n of Arkona, under log, 4 May 72, JWR, 0-4-2. LANARK CO. *Hwy 43. 12.1 km e of
Smiths Falls, under logs, 1 1 May 72, JWR, 4-0-6. *Hwy 43, 3.71 km e of Smiths Falls, under tele-
phone pole in ditch, 11 May 72. JWR, 0-0-1. *Hwy 43, 5.32 km w of Smiths Falls, under logs, 11
May 72, JWR, 4-0-17. LEEDS CO. *Hwy 15, 2.26 km n of Seeley's Bay, under logs and concrete
blocks in ditch, 16 May 72, JWR, 4-1-9. *Hwy 15, .97 km s of Portland, under logs, 16 May 72,
JWR, 0-0-5. LENNOX AND ADD1NGTON CO. *Hwy 2, Napanee, w.e., under rock behind
EEEE Motel, 15 May 72, JWR, 0-0-1. *Hwy 2, 6.61 km w of Hwy 133, under log, 16 May 72. JWR,
0-0-1. MANITOULIN DIST. *Hwy 68, 5.97 km n of Birch Island, under rock, 13 May 72. JWR &
JEM, 0-0-1. *Hwy 68, Birch Island, s.e., under logs, 13 May 72, JWR & JEM, 4-2-8. MIDDLESEX
CO. Judd (1964). Reynolds (1972a). *Hwy 73, .97 km n of Mossley, under logs, 4 May 72, JWR, 0-
0-2. *Hwy 7, 2.26 km s of Parkhill, under fence posts, 4 May 72, JWR, 0-1-6. NIAGARA CO. *Hwy
20, 1.61 km e of Smithville, old house site, 1 May 72, JWR, 0-1-4-1. NIPISSING DIST. 'Hwy 17,
1.29 km e of Verner, under paper in wet ditch, 13 May 72. JWR & JEM, 0-0-3. NORTHUMBER-
LAND CO. *Hwy 45, 10.65 km s of Norwood, under logs next to barn, 28 Apr 72, JWR, 3-3-3-1.
*Hwy 45, 4.84 km n of Baltimore, under log, 28 Apr 72, JWR. 0-0-1. ONTARIO CO. *Hwy 7, 1.61
km e of Green River, under logs, 26 Apr 72, JWR, 3-0-1. *Hwy 7-12, .81 km n of Myrtle, under logs,
26 Apr 72, JWR, 9-0-2. 'Hwy 7. 5.32 km ne of Sunderland, under logs, 9 May 72, JWR. 0-0-4. OX-
FORD CO. *Hwy 59, 1.77 km e of Burgessville, under logs and wood chips, 1 May 72, JWR, 3-0-7.
*Hwy 59, 1.13 km s of Curries, under junk, 1 May 72, JWR, 5-0-1. *Hwy 59, .32 km n of Hickson,
under logs, 3 May 72, JWR, 0-0-2. PEEL CO. *Hwy 5, .81 km e of Dixie Rd, under logs. 29 Apr 72.
JWR, 0-0-2. *Hwy 5. 4.35 km w of Hwy 10, under mattress, 29 Apr 72, JWR, 0-0-1. *Hwy 24. 8.87
km n of Erin, under log and dung, 29 Apr 72, JWR, 1-1-0. *Hwy 7, Brampton, e.e., under debris, 4
May 73, JWR, 0-1-4. PERTH CO. .81 km n of Fullarton. under rocks, 29 Apr 72, DWR & LWR, 2-
1-3. 'Mitchell, next to Collegiate, under logs, 4 May 72, JWR & DWR, 0-0-2. PETERBOROUGH
CO. *Hwy 28, Lakefield College, under logs near waterfront, 27 Apr 72, JWR & CWR, 14-5-10.
•Hwy 28, 5.48 km n of Burleigh Falls, under logs, 27 Apr 72, JWR, 1-0-6. PRESCOTT CO. *Hwy
34,5.81 km n of Vankleek Hill, under logs, 11 May 72, JWR, 1-1-3. PRINCE EDWARD CO. *Hwy
33, 2.58 km e of Hillier, digging, 15 May 72, JWR, 1-0-0. *Hwy 49, Picton, ne., under leaf pile. 15
May 72, JWR, 5-1-4. RUSSELL CO. *Hwy 17. 4.35 km w of Rockland, under log, 11 May 72. JWR,
2-0-1. SIMCOE CO. *Hwy 89, 5.48 km e of Alliston. under log. 2 May 72, JWR, 0-1-0. *Hwy 27,
5.97 km s of Cookstown, under logs, 2 May 72, JWR, 1-0-1. *Barrie. park opposite 14 Greenfield,
under rocks and grass clippings, 7 May 72, JWR & GWA. 5-1-3. STORMONT CO. 'Hwy 43. 1.29
km w of Finch, under logs, 1 1 May 72, JWR, 4-2-8. Hwy 43, 1.61 km e of Finch, under log. 1 1 May
72, JWR, 3-1-1. Hwy 43, 3.71 km w of Monkland, under logs, 1 1 May 72. JWR, 2-1-2. THUNDER
BAY DIST. Hwy 17, 15.17 km e of Thunder Bay, small stream, 17 Jun 71, ROM Field Party, 9-0-0,

82

ROM -1 34. VICTORIA CO. 'Il«\ 46. 3.71 km nofHwj 7. under log, 9 May 72. J WR. 0-0-1. *Hwy
46. SI km n of Argyle, under logs, 9 Ma> 72, JW R. 1-0-2. WATERLOO CO. *Hw) 97. 3.71 km n of
Hwj 401, under log, 3 \l.i\ 72, JWR.0-0-I. Hwy 7-8. 1.77 km w of Petersburg, under log, 3 May 72.
JWR, 1-2-& Hw) 86, Wesl Montrose, under log, 3 Ma) 72. JWR, 0-0-1. Waterloo. Laurel Creek
Conservation \rea, 3 Any 74, DPS. 0-0-1, UW-0007. Wl I 1 1NGTON CO. *Hwy 24, 5.16 km e of
Eramosa. under logs in cedar (77iu/a occidentalis) woodlot, 29 Apr 72, JWR, 3-0-1. *Hwy 24. 4.03 km
n of Inn. under logs. 29 Apr 72. JWR. 3-0-1. WENTWORTH CO. Reynolds ( 1972a). YORK CO.
*Hu\ Jet 4S and 401, Progress and Bellamy Rdv n« corner under lumber. 26 Apr 72. JWR. 1-1-7.
•Hwj 48, .32 km n ol Steeles \venue, under logs. 26 Apr 72. JWR. 3-5-7. *Hwy 27, 1.45 km n of
Hw) 7. under logs. 29 Apr 72, JWR, 1-4-9. *Edenbrook Park. Islington, under logs and rocks near
Stream bank. 30 Apr 72. JWR & DWR. 25-2-13. Edenbrook Park, Islington, quantitative study 1,
formalin. IS Ma) '1 JWR. 16-3-13. Edenbrook Park. Islington, quantitative study 2. formalin, 18
Ma\ 72, JWR. 36-5-12-2

Genus Eiseniella Michaelsen, 1900

1900 Eiseniella Michaelsen. Das Tierreich, Oligochaeta 10: 471.

Type Species

Enlerion tetraedrum Savigny, 1826.

Diagnosis

Calciferous sacs, in x, digitiform. opening posteriorly into the gut ventrally in re-
gion of insertion of 10/11. Oesophagous of nearly uniform width through
xi-xiv, calciferous channels narrow, lamellae low and continued along the lat-
eral walls of the sacs. Intestinal origin, in xv. Gizzard, in xvii, weak, 17/18 not
fenestrated. Typhlosole, simply lamelliform. Extraoesophageal trunks, joining
dorsal vessel in xii. Hearts, in vii-xi. Nephridial bladder, short, sausage-shaped.
Nephropores, inconspicuous, behind xv alternating irregularly and with asym-
metry between a level just above B and one above D. Setae, not closely paired
behind the clitellum. Prostomium epilobic. Longitudinal musculature, pinnate,
(after Gates. 1972c: 108).

Discussion

Michaelsen (1900b) proposed the new name Eiseniella for a genus erected by Ei-
sen in 1873. Although designated only as a new name for Allurus Eisen, 1873,
Michaelsen included a second genus of Eisen's (Tetragonurus Eisen, 1874) in his
Eiseniella. Both of Eisen's generic names were preoccupied (i.e.. already in use
as generic names in other groups); Allurus Foerster, 1862 had been used as a
genus of hymenopterous insects, while Tetragonurus Risso, 1810 had been em-
ployed as a genus of fish. The type species for both of Eisen's genera are syno-
nyms of Enlerion tetraedrum Savigny, 1826. Insufficient data are available con-
cerning the somatic anatomy of species included at various times in the classical
Eiseniella to determine if they can be congeneric with Eiseniella tetraedra.

83

Eiseniella tetraedra (Savigny, 1826)

Square-tail worm Ver a queue carree
(Fig. 26)

1826 Enlerion tetraedrum Savigny, Mem. Acad. Sei. Inst. Fr. 5: 184.

1826 ? Lumbricus quadrangularis Risso, Hist. Nat. Eur. Merid. 4: 426.

1828 ? Lumbricus amphisbaena Duges, Ann. Sci. Nat. 15: 289.

1837 Lumbricus tetraedrus-Duges, Ann. Sci. Nat., ser. 2, 8: 17, 23.

1843 Lumbricus agilis Hoffmeister, Arch. Naturg. 9(1): 191.

1871 Lumbricus tetraedrus-Eisen + L. t. luteus + L. t. obscurus Eisen, Ofv.
Vet.-Akad. Forh. Stockholm 27(10): 966, 967, 968.

1873 Allurus tetraedrus-Eisen, Ofv. Vet.-Akad. Forh. Stockholm 30(8): 54.

1874 Tetragonurus pupa Eisen, Ofv. Vet.-Akad. Forh. Stockholm 31(2): 47.

1885 Allurus neapolitanus Orley, Ertek. Term. Magyar Akad. 15(18): 12.

1886 Allurus ninnii Rosa, Atti 1st. Veneto, ser. 6, 4: 680.

1889 Lumbricus (Allolobophora) neapolitanus + L. (Allurus) tetraedrus + L.
(Eisenia)pupa-L. Vaillant, Hist. Nat. Annel. 3(1): 1 13, 151, 154.

1889 Allurus hercynius-M'ichadsen + A. dubius Michaelsen + A.
n/wi/7-Michaelsen, Mitt. Mus. Hamburg 7(3): 7, 10.

1890 Eisenia pupa-Benham, Quart. J. Micros. Soc, n.s., 31(2): 266.
1892 Allolobophora tetragonurus-Friend, Sci. Gossip 28: 194.

1892 Allurus tetraedrus + A. amphisbaena + A. flavus + A. tetragonurus-

Friend, Proc. R. Irish Acad., ser. 3, 2: 402.
1896 Allurus tetraedrus-K\bd.\xco\xr\ + A. bernensis + A. novis + A. infinitesi-

malis Ribaucourt, Rev. Suisse Zool. 4: 69, 73, 74.
1900 Eiseniella te?rae<ira-Michaelsen, Das Tierreich, Oligochaeta 10: 471.
1937 Eiseniella tetraedra f. /y/?/ca-Cernosvitov, Rec. Indian Mus. 39: 107.
1974 Eisenia tetraedra (laps.)-Vail, Bull. Tall Timbers Res. Stn. 11:2.

Diagnosis

Length 30-60 mm, diameter 2-4 mm, segment number 60-90, prostomium epi-
lobic, first dorsal pore 4/5-5/6. Clitellum xxii, xxiii-xxvi, xxvii. Tubercula pu-
bertals uniformly broad on xxiii-xxv, xxvi. Setae closely paired,
AA:AB:BC:CD:DD = 3:1:3:1:6-8 posteriorly. Ventral setae on x, or ix and x
modified into genital setae. Male pores on xiii with slightly elevated glandular
papillae in Cernosvitov's "typical form". This is a misnomer because the normal
or "typical" position for other members of the family is on xv. A second form of
the same species, not recorded from Ontario, has male pores on xv. Seminal ves-
icles, four pairs on 9-12. Spermathecae, two pairs opening between d and mD
line in 9/10 andl0/l 1. Body cylindrical in front of clitellum and quadrangular
behind. Colour variable, from dark brown, greenish, ruddy to bright golden yel-
low.

Biology

Eiseniella tetraedra is a limicolous species and shows a marked preference for
damp habitats. It is known from wells, springs, subterranean waters, rivers,

84

1st dp.

QQ

XX

■sg

-cl

sg

-cl

XXV

TP xxv

TP

XXX;

Fig. 26 External longitudinal views of Eiseniella tetraedra showing taxonomic characters, a Lat-
eral view, b Ventral view. (ONT: Huron Co., cat. no. 7549)

85

ponds, lakes, and canals, and may be one of the dominant animals in the dense
moss of swift streams (Gates, 1972c). In Ohio it has been recorded from soils
with a pH range of 6.8 to 8.5, a moisture content of 25-35%, and an organic
matter content of 4-5%. It is the most common megadnle in British caves and is
known from caves in the rest of Europe and in South Africa. Olson (1928, 1936)
and Eaton (1942) reported this species from water-soaked banks of streams,
lakes, and ponds. Murchie (1956) reported it from the bottom deposits of
streams, lakes, or ponds, from wet to very moist stream banks and lake shores,
from bottom lands subject to flooding, or with a high water table, and from
seepage areas around springs at upland sites. The soil type for these sites varied
from peaty organic material to sandy gravel. The sources of specimens for the
present study were all moist to wet habitats.

Under favourable conditions activity can be year round but in Ontario there
are probably summer and winter rest periods. Aestivation involves immobility
and tight coiling in a small mucus-lined cavity; whether hibernation involves
quiescence or diapause is not known. This species is obligatorily parthenogen-
etic (Muldal, 1952; Omodeo, 1955b; Reynolds, 1974c). The first reports of uni-
parental reproduction for megadriles involved experiments with Eiseniella
tetraedra (Gavrilov, 1935, 1939).

Range

A native of Palaearctis, El. tetraedra is now known from Europe, North Ameri-
ca, South America, Asia, Africa, and Australasia (Gates, 1972). It also occurs in
Iceland (Backlund, 1949). It is another cosmopolitan species that has been car-
ried around the world.

North American Distribution

British Columbia (Gates. 1972c), New Brunswick (Reynolds. 1976d), Nova Scotia (Reynolds. 1975a,
1976a), Ontario (Eisen, 1874), Prince Edward Island (Reynolds, 1975c). Quebec (Reynolds. 1975b,
d, e, 1976c), Arizona (Reynolds et al., 1974), Arkansas (Causey. 1952), California (Smith 1917),
Connecticut (Reynolds, 1973c), District of Columbia (Eaton, 1942), Idaho (Gates, 1967), Illinois
(Smith, 1917), Indiana (Heimburger, 1915), Louisiana (Harman, 1952), Maine (Gates, 1966), Mary-
land (Reynolds, 1974b), Massachusetts (Reynolds, 1977). Michigan (Smith 1917), Missouri (Olson,
1936), Montana (Reynolds, 1972c), Nevada (Gates, 1967), New Hampshire (Reynolds. 1976c), New
Jersey (Davies, 1954), New York (Olson, 1940), North Carolina (Pearse, 1946), Ohio (Olson. 1928),
Oregon (MacNab and McKey-Fender, 1947), Pennsylvania (Moore, H.F., 1895), Rhode Island
(Reynolds, 1973b), South Dakota (Gates, 1967), Tennessee (Reynolds, 1974a). Utah (Gates. 1967),
Vermont (Reynolds, 1976c), Virginia (Harman, 1960), Washington (Smith. 1917), West Virginia
(Reynolds, 1974b), Wisconsin (Gates, 1972c), Wyoming (Gates, 1967). New records: Alaska. Geor-
gia, Iowa, Kentucky.

Ontario Distribution (Fig. 27)

Eiseniella tetraedra was the fourth species reported from Ontario by Eisen
(1874). This species has also been recorded in the province by Stafford (1902)
and Judd (1964). The distribution of El. tetraedra in Ontario is generally close to
the Great Lakes and concentrated in southwestern Ontario.

86

Fig. 27 The known Ontario distribution of Eiseniella leiraedra.

BRUCE CO. *Hwy 4. 6.77 km s of Teeswater. under log. 5 May 72, JWR, 0-1-0. "Hwy 21. 6.94 km
n of Kincardine, under log, 5 May 72, JWR. 0-0-1. COCHRANE DIST. Reynolds (1972a). DUR-
HAM CO. Kendal, cold fast stream, 31 May 66, IMS, 4-4-7, ROM-I1. ESSEX CO. *Hwy 3, Ruth-
ven, n.e., wet ditch, next to railroad tracks, 4 May 72, JWR. 0-0-2. FRONTENAC CO. "Hwy 15,
4.03 km n of Hwy 401, under paper in wet ditch, 16 May 72. JWR, 1-0-1. 'Hwy 41, 1.29 km n of
Cloyne, under log in ditch. 16 May 72, JWR, 0-0-1. HALD1MAND CO. *Hwy 3, 6.13 km w of Cay-
uga, under logs. 1 May 72, JWR' 0-2-1. HURON CO. "Hwy 4, 1.61 km n of Exeter, wet ditch. 5
May 72. JWR. 1-1-12. KENT CO. Morpeth, under stones near creek, 21 Apr 75, DRB, 0-0-1, UW-
0001. LAMBTON CO. Hwy 21. 2.1 km n of Wyoming, under logs in wet ditch, 4 May 72. JWR. 0-
0-3. LEEDS CO. *Hwy 15. 2.26 km n of Seeley's Bay. under logs and concrete blocks in ditch. 16
May 72, JWR. 0-0-3. Chaffey's Locks, swamp near Lake Opinicon, 4 Sep 65, IMS, 0-0-1. ROM-II7.
MIDDLESEX CO. Judd(l964). NIAGARA CO. Eisen(1874). NIPISSING DIST. *Hwy 17, 1.29
km e of Verner. under paper in wet ditch. 13 May 72, JWR & JEM, 0-1-1. NORFOLK CO. Rey-
nolds (1972a). NMC-1076. 'Hwy 6, 4.84 km s of Jarvis. under logs, 1 May 72, JWR, 0-2-0. ON-
TARIO CO. Duffenn Creek area, day after use of lampricide. 1 1 May 71, TY, 0-0-5, ROM-I38. OX-
FORD CO. Reynolds (1972a). NMC-1077. PEEL CO. Port Credit, Credit Rjver, 6 May 66, IMS. I-
0-0. ROM. PERTH CO. Hw) Jet 23 and 83. Russeldale, under paper in wet ditch. 5 May 72. JWR,
4-0-3. PRINCE EDWARD CO. 'Hwy 49, 14.84 km n of Picton. under paper in ditch, 15 May 72,
JWR. 2-1-13. RUSSELL CO. *Hwy 17. 7.74 km e of Rockland, under paper in wet ditch, 1 1 May
72. JWR. 1-0-2. STORMON I ( O. *Hwy 43. 5.48 km w of I inch, in and under straw in wet ditch.
1 1 May 72. JWR. 0-0-3. WENTWORTH CO. Reynolds ( 1972a). Strathcona Gardens, Lake Ontar-
io. 7 May 75. DRB. 0-0-2, UW-0001. YORK CO. *Edenbrook Park, Islington, under log near
stream bank. 30 Apr 72. JWR & DWR. 0-0-1.

87

Genus Lumbricus Linnaeus, 1758

1758 Lumbricus (part.) Linnaeus, Syst. Nat. (ed. 10), p. 647.

1774 Lumbricus (part. )-M idler. Verm. Terr. Fluv. 1(2): 24.

1780 Lumbricus (part.)-Fabricius, Fauna Gr0nlandica, p. 277.

1826 Enterion (part.) Savigny, Mem. Acad. Sci. Inst. Fr. 5: 179.

1836 Lumbricus (part.)-Templeton + Omilurus Templeton, Ann. Mag. Nat.

Hist. 9: 235.
1845 Lumbricus (part.)-Hoffmeister, Regenwiirmer, p. 4.
1873 Lumbricus-Eisen, Ofv. Vet.-Akad. Forh. Stockholm 30(8): 45.
1876 Lumbricus (part.)-Claus, Grundziige der Zool. (ed. 3) 1 : 416.

1880 Lumbricus (part.)-Claus, Grundziige der Zool. (ed. 4) 1 : 478.

1881 Lumbricus (part.) + Enterion-Orley, Math. Term. Kozlem. Magyar
Akad. 16: 580, 587.

1894 Allolobophora (part.)-W.W. Smith, Trans. N.Z. Inst. 26: 1 17.

1900 Lumbricus-Michaeken, Das Tierreich, Oligochaeta 10: 508.

1930 Lumbricus-Stephenson, Oligochaeta, p. 914.

1975 Lumbricus-Gates, Megadrilogica 2( 1 ): 3.

Type Species

Lumbricus terrestris Linnaeus, 1758, by Sims (1973).

Diagnosis

Calciferous sacs, in x, digitiform to pyriform, opening into gut posteriorly and
ventrally in region of insertion of 10/11, lamellae continued along lateral walls.
Oesophagus, widened and markedly moniliform in xi-xii, in those segments
with a vertically slitlike lumen which widens as lamellae gradually narrow be-
hind 12/13. Intestinal origin, in xv. Gizzard, mainly in xvii. Typhlosole, high,
rather thick and nearly oblong vertically, grooves not continuous across ventral
face. Extraoesophageal trunks, joining dorsal trunk in region of ix-x. Hearts, in
vii-xi. Nephridial bladder, ./-shaped, closed end laterally, duct passing into par-
ietes near B. Nephropores, obvious, behind xv irregularly alternating, with as-
ymmetry, between levels just above B and well above D. Setae, closely paired.
First dorsal pore, anterior to 1Q/ 11. Prostomium, tanylobic. Body compressed
dorsoventrally behind clitellum and with a more or less trapezoidal transverse
section. Longitudinal musculature, pinnate. Pigment, lacking immediately un-
derneath intersegmental furrows, in circular muscle layer (after Gates, 1972c:
113, 1975a: 3).

Discussion

The genus Lumbricus (Linnaeus, 1758, p. 647) originally contained only two
species, L. terrestris and L. marinus. Since the latter was not a member of the Ol-
igochaeta, the type species was declared to be Lumbricus terrestris (I.C.Z.N.,
Opinion 75). Discussions of what was really meant by L. terrestris Linnaeus,
1758 (and his lengthened definition of Syst. Nat., ed. 12, 1767) have raged ever
since but were officially settled recently (Sims, 1973; Gates, 1973b; and Bouche,
1973). The result of this action was the neotypification of Lumbricus terrestris
Linnaeus by Sims (1973) with an expanded definition, and the deposition of
type material from Sweden in the British Museum (Natural History).

88

Lumbricus castaneus (Savigny, 1826)

Chestnut worm Ver alezan
(Fig. 28)

1826 Enterion castaneus + E. pumilum Savigny, Mem. Acad. Sci. Inst. Fr. 5:

180. 181.
1837 Lumbricus casiancusDuv.es. Ann. Sci. Nat., ser. 2, 8: 17, 22.
1851 Lumbricus iriannularis Grube. In: Middendorff, Reise Sibirien 2(1): 18.
1865 Lumbricus minor Johnston. Cat. British Non-paras, worms, p. 59.
1867 Lumbricus josephinae Kinberg, Ofv. Vet.-Akad. Forh. Stockholm 23: 98.
1871 Lumbricus purpureus Eisen, Ofv. Vet.-Akad. Forh. Stockholm 27(10):

956.
1881 Enterion pupureum + Lumbricus purpureus-Or\ey, Math. Term. Kozlem.

Magyar Akad. 16: 588.590.
1889 Lumbricus (L.) castaneus + L. (L.) purpureus + L. (L.) triannularis-L.

Vaillant, Hist. Nat. Annel. 3(1): 124, 127, 129.

1894 Allolobophora purpureus-'W .W . Smith. Trans. N.Z. Inst. 26: 1 17.

1895 Lumbricus pumilosum (laps.)-Beddard, Monogr. Oligo. (Oxford), p. 722.

1896 Lumbricus castaneus-K\bdL\icom\ + L. morelli Ribaucourt -I- L. perrieri
Ribaucourt. Rev. Suisse Zool. 4: 10, 13, 14.

1900 Lumbricus castaneus-M\chdiz\szn, Das Tierreich, Oligochaeta 10: 510.
1936 Lumbricus castaneus var. disjonctus Tetry, Bull. Soc. Sci. Nancy, n. ser.

1936: 196.
1957 Lumbricus castaneus var. pictus Chandebois, Bull. Soc. Zool. France 82:

417.
1972 Lumbricus castaneus-^ouchk, Inst. Natn. Rech. Agron., p. 362.

Diagnosis

Length, 30-50 mm (generally <35mm), diameter 3-5 mm, segment number
70-100. prostomium tanylobic, first dorsal pore 5/6-8/9. Clitellum xxviii-xxxiii.
Tubercula pubertatis xxix-xxxii. Setae closely paired, AA ^ BC, AB^>CD, DD
^ V2C anteriorly and Z)D<V2C posteriorly. Setae a and b on ix and/or x on pale
genital tumescences fused ventrally. Male pores inconspicuous on xv. Seminal
vesicles, three pairs in 9, 11. and 12+13. Spermathecae, two pairs with short
ducts in 9/10 and 10/11. Colour, deeply pigmented, dark red, chestnut, violet
brown and strongly iridescent. Body cylindrical and dorsoventrally flattened
posteriorly.

Biology

Lumbricus castaneus has been recorded from soils with a pFf range of 4.6-8.0,
from gardens, cultivated fields, pastures, forests, taiga, steppes, among organic
matter such as manure and compost or leaf litter, and in banks by water (Gates,
1972c). It has been found in caves in Europe. Gerard (1964) reported it as ter-
restrial, mostly in soil rich in organic matter, in gardens, parks, pastures, forests,
on river and marsh banks, and under stones, leaves and dung. Apart from this
author's records all North American records fail to give habitat information but
not) findings (Reynolds. 1973a, 1973b, 1973c, 1974b, 1975a-e) are similar to
those of Gerard in England. In Ontario L. castaneus was collected from a stream

89

Fig. 28 External longitudinal views of Lumbricus castaneus showing taxonomic characters, a. Lat-
eral view. (NS: Queens Co., cat. no. 8880) B. Ventrolateral view. (NS: Digby Co.. cat. no.
8889)

bank (Reynolds, 1972a) and Judd (1964) obtained his specimens from the By-
ron Bog near London.

In unfavourable seasons individuals may be found 1 to V/i metres down in the
soil, but little is known of the annual cycle. Copulation, which may not involve
secretion of a slime tube (cf. pp. 3-4), is subterranean. This species is obligatorily
amphimictic (Reynolds, 1974c). Janda and Gavrilov (1939) did not find that iso-
lated individuals laid cocoons.

90

Range

A native of Palaearctis. /.. castaneus is now known from Europe, North America
including Mexico. New Zealand, and St. Helena (dates, 1972c). It also occurs
in Iceland (Backlund, 1949).

North American Distribution

New Brunswick (Reynolds I976d), Newfoundland (Cues. 1942), Nova Scotia (Reynolds. 1975a,
1976a). Ontario (Fisen. 1874). Prince Edward Island (Reynolds, 1975c). Quebec (Reynolds, 1975b.
d. e. 1976c). Delaware (Reynolds, 1973a), Connecticut (Reynolds, 1973c). Idaho (Gates. 1967),
Maine (Gates, 1966), Maryland (Reynolds, 1974b). Massachusetts (Reynolds. 1977). New York
(Smith. 1917). Oregon (Gates. 1972c), Pennsylvania (Bhatti, 1965). Rhode Island (Reynolds, 1973b),
Vermont (Reynolds, 1976c), Virginia (Reynolds, 1974b). West Virginia (Reynolds. 1974b). New rec-
ords: Alaska. Washington

Ontario Distribution (Fig. 29)

Lumbricus castaneus was the fifth species reported from Ontario by Eisen (1874).
It has been recorded only twice since (Judd, 1964 and Reynolds, 1972a). This
species has been recorded from only four counties in the province.

HALIBURTON CO. 'Reynolds (1972a). MIDDLESEX CO. Judd (1964). NIAGARA CO. Eisen
(1874). YORK CO. Scarborough Bluffs. Bnmley Rd. garden. 28 Oct 41, JO. 0-0-1. ROM.

Fig. 29 The known Ontario distribution of Lumbricus castaneus

91

Lumbricus festivus (Savigny, 1826)

Quebec worm Ver quebecois
(Fig. 30)

1826 Enterionfestivum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 180.

1836 Lumbricus omilurus R. Templeton, Ann. Mag. Nat. Hist. 9: 235.

1837 Lumbricus festivus-Duges, Ann. Sci. Nat., ser. 2, 8: 17,21.
1891 Lumbricus rubescens Friend, Nature 44: 273.

1900 Lumbricus festivus-Michaehen, Das Tierreich, Oligochaeta 10: 512.

Diagnosis

Length 48-105 mm, diameter 4-5 mm, segment number 100-143, prostomium
tanylobic, first dorsal pore 5/6. Clitellum xxxiv-xxxix. Tubercula pubertatis
xxxv-xxxvii, xxxviii. Setae closely paired, AA:AB:BC:CD = 34: 12:25:8 anteri-
orly, and 35:10:25:8 posteriorly. Setae on segments v-x are notably enlarged
and more widely paired. Some of the ventral setae on viii-xiv, xviii, xxv-xxxix
are on genital tumescences. Male pores on xv with glandular papillae extending
onto xiv and xvi. Seminal vesicles, three pairs in 9, 1 1, and 12+ 13+ 14. Sper-
mathecae, two pairs with short ducts opening on level c in 9/10 and 10/1 1. Col-
our, ruddy brown, iridescent dorsally and lightly coloured ventrally. Body cylin-
drical and slightly dorsoventrally flattened posteriorly.

Biology

Lumbricus festivus is rare and little is known about its habits. Gerard (1964)
recorded the habitats of this species as pastures, on river banks, in soil beneath
dung, leaves, and under stones. I found the species at ten sites in Ontario, six
times under logs, three times under debris, and once under a leaf pile.

Lumbricus festivus is assumed to be obligatorily amphimictic (Reynolds,
1974c) and copulation occurs beneath the soil surface. In recent surveys (Rey-
nolds, 1975d, 1975e, 1976c) specimens with spermatophores have been found.

Range

Known only from western Europe and northeastern North America, notably
southern Quebec.

North American Distribution

New Brunswick (Langmaid. 1964). Quebec (Reynolds. I975d, e. 1976c), Vermont (Reynolds,
1976c). New record: British Columbia.

Ontario Distribution (Fig. 31)

Lumbricus festivus was first reported from Ontario by Stafford (1902) but his re-
port simply stated "from Ontario, Quebec, New Brunswick and Nova Scotia"
with no habitat or locality information. I have been unable to verify the exist-
ence of this species in New Brunswick or Nova Scotia (Reynolds, 1975a, 1976a,
1976d). The specimens obtained during this study are the first reports of L.
festivus in Ontario since 1902.

92

1stdp-

-sg

XX

<*_

&L

GT

-cl

XXXV

XXXV

Fig 30 External longitudinal views of Lumbricus festivus showing taxonomic characters, a Lateral
view. (ONT: Glengarry Co., cat. nos. 7620 and 762 1) B Ventrolateral view. (ONT: North-
umberland Co., cat. no. 8108)

93

Fig. 3 1 The known Ontario distribution of Lumbncus festivus.

CARLETON CO. *Ottawa-Carleton Rd 35, 2.42 km s of Leonard, under logs, 1 1 May 72. JWR. 1-
1-0. DUNDAS CO. *Hwy 31, 3.55 km n of Morrisburg, under lumber, 1 1 May 72, 0-1-0. GLEN-
GARRY CO. *Hwy 34, 1 1.13 km n of Lancaster, under log, 1 1 May 72, 1-0-0. *Hwy 34. 5.65 km n
of Alexandria, under log, 11 May 72, JWR, 0-0-1. NIPISSING DIST. *Hwy 17. 1.29 km e of Ver-
ner, under paper in wet ditch, 13 May 72, JWR & JEM, 2-1-0. NORTHUMBERLAND CO. *Hwy
45. Baltimore, under junk, 28 Apr 72, JWR, 1-4-1. *Hwy 401, 5.65 km e of Grafton, under logs, 15
May 72, JWR, 3-1-1. 'Northumberland-Durham Rd 1. .65 km w of Hwy 33, under logs, 15 May 72,
JWR, 1-1-1. PEEL CO. *Hwy 5, .81 km e of Dixie Rd, under log, 29 Apr 72, JWR, 0-1-0. PRINCE
EDWARD CO. *Hwy 49, Picton, n.e., under leaf pile, 15 May 72, JWR, 1-0-0.

Lumbricus rubellus Hoffmeister, 1843

Red marsh worm Ver rouge du marecage
(Fig. 32)

1843
1877
1881

1883

Lumbricus rubellus Hoffmeister, Arch. Naturg. 9(1): 187.

Lumbricus campestris (part.) Mutton, Trans. N.Z. Inst. 9: 351.

Enterion rubellum var. parvum + E. r. var. magnum Orley, Math. Term.

Kozlem. Magyar Akad. 16: 588, 589.

Digaster campestris (part.)-Hutton, N.Z. J. Sci. 1 : 586.

94

1887 Endrilus campestns (part.}-W.W. Smith, Trans. N.Z. Inst. 19: 137.

1892 Lumbricus rubellus var. curticaudatiis Friend, J. Linn. Soc. Lond. 24: 312.

1894 Allolobophora rubellus-VJ W . Smith, Trans. N.Z. Inst. 26: 117.

1900 Lumbricus rubellus-Michaehen, Das Tierreich, Oligochaeta 10: 509.

1909 Allolobophora relicta Southern, Proc. R. Irish Acad. 27B(8): 1 19.

1923 Lumbricus rubellus-Slephenson, Fauna British India, Oligochaeta, p.

508.
1972 Lumbricus rubellus rubellus- Bouche + L. r. castaneoides + L. r.

friendoides Bouche, Inst. Natn. Rech. Agron., p. 368, 371, 372.

Diagnosis

Length 50-150 mm (usually >60 mm), diameter 4-6 mm, segment number
70-120, prostomium tanylobic, first dorsal pore 5/6-8/9. Clitellum xxvi,
xxvii-xxxi, xxxii. Tubercula pubertatis on xxviii-xxxi. Setae closely paired, AA
> BC, AB > CD, DD = V2C posteriorly. Genital tumescences in viii-xii (less
frequently on x), xx-xxiii, xxvi-xxxvi. Male pores, inconspicuous, without glan-
dular papillae on xv. Seminal vesicles, three pairs in 9, 11, and 12+ 13. Sper-
mathecae, two pairs with short ducts opening in 9/10 and 10/11. Colour, ruddy
brown or red-violet and iridescent dorsally, pale yellow ventrally. Body cylindri-
cal and sometimes dorsoventrally flattened posteriorly.

Biology

Lumbricus rubellus has been recorded from natural soils of pH 3.8-8.0 and
shows a wide tolerance of habitat factors. Olson (1928, 1936) reported it from
under debris. Eaton (1942) found it in stream banks, under logs, and in woody
peat and stated that it seemed to require a great deal of moisture and organic
matter. Cernosvitov and Evans (1947) recorded the habitats of this species as
places rich in humus, abundant in parks, gardens, pastures, on river banks, un-
der stones, moss, or old leaves. Gerard (1964) also found this species frequently
aggregated beneath dung in pastures as well as the sites mentioned above. In
Ontario, L. rubellus was obtained from a wide variety of habitats. This species
also is known from caves in Europe.

Under suitable conditions activity, including breeding, is year round. L.
rubellus is obligatorily amphimictic (Reynolds, 1974c) and copulation, like defe-
cation, occurs below the soil surface, or in the litter layer, at any time of day. It
seems that copulation does not involve a mucous tube (Gates, 1972c).

This species has been cultured by the fish bait industry (Gates, 1972c). It is
also important in the decomposition of litter.

Range

Lumbricus rubellus is now known from Europe, Iceland, North America, Mexi-
co, Asia, South Africa, and New Zealand (Gates, 1972c).

North American Distribution

British Columbia (Berkeley. 1%8), New Brunswick (Reynolds. 1976d). Newfoundland (Smith.
1917), Nova Scotia (Reynolds, 1975a, 1976a), Ontario (Reynolds, 1972a), Prince Edward Island
(Reynolds, 1975c), Quebec (Reynolds, 1975b. e. 1976c). Alaska (Gates, 1954), Arkansas (Causey,

95

1st dp-

-pr

-GT

-GT

Q

o

■GT

GT

£)

I — TP

Fig. 32 External longitudinal views of Lumbricus rube/lux showing taxonomic characters, a Lateral
view, b Ventral view. (ONT: Parry Sound Dist., cat. no. 7505)

96

1952). California (Smilh 1917). Colorado (dates. 1967). Connecticut (Reynolds. 1973c). Delaware
(Reynolds, 1973a), District of Columbia (Eaton, 1942). Florida (Gates, 1 972c). Idaho (Gates, 1967),
Illinois (Smith. 1928), Indiana (Jo\ner, I960). Mars land (Reynolds, 1974b), Massachusetts (Rey-
nold-. 1977), Michigan (Smith. 1917), Missouri (Olson. 193m. New Hampshire (Reynolds. 1976c),
New Jersej (Davies, 1954), Now York (Olson, 1940), Ohio (Olson, 1928). Oregon (Smith. 1917),
Pennsylvania (Eaton, 1942). Rhode Island (Reynolds, 1973b). Tennessee (Reynolds, 1972b), Utah
(dates. 1967). Vermont (Reynolds, 1976c). Virginia (Reynolds, 1974b), Washington, (Smith, 1917),
West Virginia (Reynolds, 1974b), Greenland (Omodeo, 1955a). New records: Manitoba. Oeorgia,
Minnesota. North Carolina.

Ontario Distribution (Fig. 33)

Lumbricas rubelhis was first reported from Ontario by Stafford (1902) and only
once since then (Reynolds, 1972a). The species is widely distributed across the
province and additional collecting under favourable conditions should fill the
gaps in its distribution.

ALGOMA DIST. "Hwy 17. 7.58 km e of Spanish, under logs, 13 May 72, JWR & JEM, 16-10-8.
*Hwy 17, .48 km e of Spanish, under logs and paper in ditch. 13 May 72. JWR & JEM, 4-1-5. Gar-
gantua Bay. 1 1 Jun 75. DRB. 1-0-0. UW-0001. BRANT CO. »Hwy 53, 5.48 km w of Cathcart. under
railroad ties. 1 May 72. JWR. 0-1-0. *Hwy 54. .64 km w of Onondaga, under log in wet area, 3 May
72. JWR. 0-1-0. BRUCE CO. Hwy 21, 2.57 km n of Tiverton, under lumber. 5 May 72, JWR, 1-0-0.
CARLETON CO. Reynolds (1972a). COCHRANE DIST. Reynolds (1972a). DUFFERIN CO.
•Hwy 9. 10-16 km w of Orangeville. under junk, 2 May 72, JWR, 0-1-0. *Hwy 9, 3.71 km w of Or-
angeville. under log in ditch. 2 May 72, JWR. 0-1-0. DUNDAS CO. *Hwy 34, 1.29 km e of Chester-

Fig. 33 The known Ontario distribution of Lumbricus rubellus.

97

ville, under rock in ditch, 11 May 72, JWR, 0-1-0. *Hwy 31..81 km n of Hwy 43. under logs. 11 May
72, JWR, 1-1-4. DURHAM CO. *Hwy 401, .16 km w of Liberty Rd. Bowmanville. digging under
log next to railroad tracks, 15 May 72. JWR, 1-0-0. *Hwy 401, .32 km e of Mill St. Newcastle, under
log, 15 May 72, JWR, 1-0-0. ELGIN CO. *Hwy 73, 3.06 km s of Harnetsville, under rotten log, 4
May 72, JWR. 2-0-0. *Hwy 73, 6.77 km n of Aylmer, under logs, 4 May 72, JWR. 4-2-3. Hwy 3, 9.19
km e of St. Thomas, under logs, 4 May 72, JWR. 0-2-0. *Hwy 3. 6.77 km e of Wallacetown. under
pine logs, 4 May 72, JWR, 2-0-0. Hwy 3, 3.87 km w of New Glasgow, under lumber in abandoned
school yard, 4 May 72, JWR, 3-0-0. ESSEX CO. Hwy 3, 3.87 km w of Leamington, under lumber in
dump, 4 May 72, JWR, 10-0-0. FRONTENAC CO. *Hwy 2, 1.13 km e of Westbrook, under logs
and paper, 16 May 72, JWR, 5-1-9. *Hwy 15, 4.03 km n of Hwy 401, under paper in wet ditch, 16
May 72, JWR, 0-1-0. *Hwy 41, 1.29 km n of Clovne, under logs in ditch, 16 May 72. JWR. 0-0-8.
GRENVILLE CO. Hwy 2, Cardinal, w.e., under log and paper, 1 1 May 72, JWR, 2-0-0. GREY CO.
*Hwy 6BP, Rockford, w.e., under logs, 5 May 72, JWR, 7-1-0. *Hwy 6, 3.06 km s of Chatsworth,
under logs, 5 May 72, JWR, 2-0-0. HALD1MAND CO. *Hwy 3. 9.03 km e of Dunnville, wet ditch,

I May 72, JWR. 3-0-1. HALIBURTON CO. 'Reynolds (1972a). *Hwy 121. 3.22 km e of Tory Hill,
dump, 16 May 72. JWR, 6-1-21. HURON CO. *Hwy 83, 6.77 km w of Russeldale, under logs. 5
May 72, JWR, 3-1-0. »Hwy 4, 1.61 km n of Exeter, wet ditch, 5 May 72, JWR, 5-1-0. Hwy 4, 1.29 km
n of Hensall, under corn (Zea maize) cobs in pasture, 5 May 72, JWR, 1-0-0. *Hwy 4, 3.71 km s of
Brucefield, under log and rock, 5 May 72, JWR. 2-0-0. *Hwy 4, 2.74 km n of Clinton, under logs, 5
May 72, 2-1-2. *Hwy 4, 1.29 km s of Wingham, under log in wet area, 5 May 72, JWR, 1-0-0. KEN-
ORA D1ST. Rushing River Provincial Park, stream trickle in campground, 13-14 Jun 71. ROM
Field Party, 1-0-0, ROM-I36. LENNOX AND ADDINGTON CO. 'Hwy 500, .81 km n of Den-
bigh, under log, 16 May 72, JWR, 1-0-0. Hwy 7, Kaladar, e.e., under lumber in school yard. 16 May
72, JWR, 2-0-2. MAN1TOUL1N DIST. *Hwy 68, 5.97 km n of Birch Island, under rock. 13 May 72,
JWR & JEM, 0-0-1. *Hwy 68, 2.42 km s of Birch Island, under logs, 13 May 72, JWR & JEM.3-1-
22. MIDDLESEX CO. *Hwy 73, .97 km n of Mossley, under logs, 4 May 72, JWR, 6-1-6. MUS-
KOKA DIST. *Hwy 11, 12.9 km s of Bracebridge, under rock, 9 May 72, JWR, 1-0-0. *Hwy 11,
9.19 km s of Hwy 516, under logs and rocks in wet ditch, 9 May 72, JWR, 3-1-0. NIAGARA CO.
*Queen Elizabeth Hwy, 1.13 km w of Ontario St, Grimsby, under logs and lumber, 1 May 72, JWR,
4-1-0. 'Niagara Co. Rd 12, 3.06 km s of Grimsby, under paper in wet ditch, 1 May 72. JWR, 5-1-6.
*Hwy20. 1.61 km e of Smithville, old house site under lumber, 1 May 72, JWR, 4-1-1. NIPISSING
DIST. *Hwy 17, 5.48 km e of Warren, under log and dung in pasture, 13 May 72. JWR & JEM, 5-
5-10. Hwy 17, 15.81 km e of Sturgeon Falls, dump. 13 May 72, JWR & JEM, 6-4-5. NORFOLK
CO. *Hwy 6, 4.84 km s of Jarvis, under logs, 1 May 72. JWR, 1-1-1. OXFORD CO. "Hwy 97,
Washington, w.e., under log, 3 May 72, JWR, 1-0-0. PARRY SOUND DIST. "Hwy 520, 10.32 km e
of Magnetawan, under lumber, 9 May 72, JWR, 5-2-3. *Hwy 520, Magnetawan. e.e., under rocks, 9
May 72, JWR, 2-0-1. *Hwy 124, 2.26 km w of Hwy 520. under logs, 9 May 72, JWR, 4-3-2. *Hwy
124, 3.06 km e of McKellar, under logs, 9 May 72, JWR, 0-0-3. "Hwy 124, 3.55 km n of Hwy 69, un-
der logs, 9 May 72, JWR, 1-0-3. PETERBOROUGH CO. 'Hwy 28, Lakefield College, under logs
near waterfront, 27 Apr 72, JWR & CWR, 2-0-0. *Hwy 504, 1.61 km e of Apsley, under log. 27 Apr
72, JWR, 1-0-0. RENFREW CO. M.19 km s of Palmer Rapids, under paper in ditch. 16 May 72,
JWR, 2-2-3. RUSSELL CO. *Hwy 17, 7.74 km e of Rockland, under paper in wet ditch, 1 1 May 72,
JWR, 9-1-0. 'Hwy 17, 4.35 km w of Rockland, under logs, 1 1 May 72, JWR, 9-2-4. *2.42 km s of Li-
monges, under lumber at old house site, 1 1 May 72, JWR, 0-1-0. *2.9 km e of Embrun. under logs,

I I May 72, JWR, 3-0-3. SIMCOE CO. Hwy 27, 3.22 km s of Newton Robinson, wet ditch. 2 May
72, JWR, 2-0-0. 'Barrie. park opposite 14 Greenfield, under rock and grass clippings. 7 May 72,
JWR, 6-1-0. STORMONT CO. *Hwy 43, 5.48 km w of Finch, in and under straw in wet ditch, 1 1
May 72, JWR. 3-1-0. SUDBURY DIST. *Hwy 69, 3.39 km n of Estaire, under rocks and grass near
house, 13 May 72, JWR & JEM, 3-0-6. *Hwy 17, 2.74 km e of Whitefish, under paper in ditch, 13
May 72, JWR & JEM, 3-2-8. VICTORIA CO. *Hwy 7, 1.94 km w of Hwy 46, under logs and dung
in pasture, 9 May 72, JWR, 5-0-0. Hwy 48, Bolsover. e.e., under paper in wet ditch. 9 May 72. JWR.
0-1-0. WATERLOO CO. Hwy 86, West Montrose, under log, 3 May 72, JWR. 1-0-0. Waterloo,
Beechwood South, on sidewalk after rain (evening), 15 Jun 75, DPS, 2-0-1, UW-0004. WELLING-
TON CO. *Hwy 24, 5.16 km e of Eramosa, under log in cedar (Thuja ocadentalis) woodlot. 29 Apr
72. JWR, 1-0-0. WENTWORTH CO. Reynolds (1972a). "Hwy 6, 5.32 km n of Hwy 5. under logs,
29 Apr 72, JWR, 2-0-1. YORK CO. Edenbrook Park. Islington, quantitative study I, formalin. 18
May 72, JWR, 34-0-0. Edenbrook Park, Islington, quantitative study 2, formalin, 18 May 72. JWR.
47-0-1.

98

Lumbricus terrestris Linnaeus, 1758

Nightcrawler, Dew-worm Ver nocture rampant

(Fig. 34)

1758 Lumbricus terrestris (part.) Linnaeus. Syst. Nat. (ed. 10), p. 647.

1774 Lumbricus terrestris (part.)-Miiller, Verm. Terr. Fluv. 1(2): 24.

1780 Lumbricus terrestris (part.) + L. norvegicus (part.)-Fabricius, Fauna
Gr0nlandica, p. 277.

1825 Lumbricus terrester (part.)-Blumenbach. Hand. Naturg. (ed. 1 1), p. 365.

1826 Enterion herculeum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 180.
1837 Lumbricus herculeus- Duges, Ann. Sci. Nat., ser. 2, 8: 17, 21.
1842 Lumbricus agricola Hoffmeister. Verm. Lumbric, p. 24.

1867 Lumbricus infelix Kinberg, Ofv. Vet.-Akad. Forh. Stockholm 23: 98.

1872 ? Lumbricus americanus E. Perrier, Nouv. Arch. Mus. Paris 8: 44.

1884 Lumbricus herculeus-Rosa., Lumbric. Piemonte, p. 22.

1896 Lumbricus studeh Ribaucourt, Rev. Suisse Zool. 4: 5.

1900 Lumbricus /w?5/m-Michaelsen, Das Tierreich, Oligochaeta 10: 511.

1937 Lumbricus herculeus-Tetry, Bull. Mus. Hist. Nat. 9: 151.

1953 Lumbricus terrestris-Gx&R, Zool. Anz. 161(1 1/12): 324.

1958 Lumbricus terrestris-Gales, Breviora, Mus. Comp. Zool. 91: 8.

1969 Lumbricus herculeus-Bouche, Pedobiologia 9: 89.

1970 Lumbricus herculeus-Bouche, Rev. Ecol. Biol. Sol 7(4): 541.

1972 Lumbricus herculeus-Bouche and Beugnot, Rev. Ecol. Biol. Sol 9(4): 697.

1972 Lumbricus herculeus-Bouche, Inst. Natn. Rech. Agron., p. 352.

1973 Lumbricus terrestris-Sims. Bull. Zool. Nomencl. 30(1): 27.

NOTE: These are the major synonyms and references for Lumbricus terrestris
Linnaeus. Unfortunately, Lumbricus terrestris has become almost synonymous
with "earthworm" in many parts of the world. This has led to many published
studies and reports about the species which in fact were undertaken on differ-
ent, and frequently distantly related, species (Orley, 1881; Vaillant, 1889; Ste-
phenson, 1930: xi; Causey, 1952; Stebbings, 1962: 905; Cameron and Fogal,
1963; Gates, 1972a: 1 14-1 15; and Gates, 1972c: 120-123).

Diagnosis

Length 90-300 mm, diameter 6-10 mm, segment number 120-160, prostomium
tanylobic. first dorsal pore 7/8. Citellum xxxi, xxxii-xxxvii. Tubercula puberta-
ls xxxiii-xxxvi. Setae enlarged and widely paired in the caudal and cephalic re-
gions (i.e.. AB and CD are greater) but closely paired and smaller in the central
region. AA>BC, AB>CD. and DD = V2C anteriorly, DD<lhC posteriorly.
The ventral setae of x, xxvi, and sometimes xxv are on broad genital tumes-
cences modified into genital setae. Genital tumescences, occasionally in viii-xiv
and xxiv-xxxix. Male pores prominent with large elevated glandular papillae ex-
tending over xiv-xvi. Seminal vesicles, three pairs in 9, 11, and 12+ 13. Sper-
mathecae, two pairs with short ducts opening at 9/10 and 10/11. Body cylindri-
cal and strongly compressed dorsoventrally posteriorly. Heavily pigmented,
brownish-red, or violet colour on dorsum and yellowish-orange on venter.

99

7— mp

W

XXV

GT

Fig. 34 External longitudinal views of Lumbricus lerrestris showing taxonomic characters, a Dor-
solateral view. b. Ventrolateral view. (ONT: Manitoulin Dist., cat. no. 7956)

100

Biology

Lumbricus terrestris has been found in soils of pH 4.0-8.08 and can adapt to a
wide variety of habitats. According to Gerard (1964) it is almost purely terres-
trial and is found in gardens, arable and pasture lands, forests, and river banks.
Gates (1972c) records additional habitats such as streams, mud flats, woody
peat, under logs in a stream bed, under cow pats, and in compost. It has been
found in greenhouses and botanical gardens in Europe and North America and
in European caves (Gates, 1972c). The species does not normally occur in for-
ests in North America (Reynolds, 1976b). After intensive collecting by the au-
thor in eastern North America, it now appears that statements such as
"Lumbricus terrestris is becoming increasingly important in the United States,
following its importation and gradual replacement of endemic populations"
(Edwards et al., 1969) are not true. Many erroneous statements such as this have
been made about L. terrestris without data to support their claims. Data for nat-
urally occurring populations of L. terrestris in North American areas south of
the limits of Quaternary glaciations are lacking (Gates, 1970). A few limited col-
lections of this species were obtained in Tennessee, Maryland, and Delaware,
three states south of the southern limits of Pleistocene glaciation, by Reynolds
et al. (1974), Reynolds (1974b), and Reynolds (1973a), respectively. In Ontario,
L. terrestris was collected from a variety of habitats, but primarily from under
logs. Millions are collected annually from the surfaces of golf courses and lawns
for the bait industry.

Under favourable conditions activity, including copulation, is year round but
a summer and winter rest period may be climatically imposed in certain areas.
The species is obligatorily amphimictic (Reynolds, 1974c) and copulation is
nocturnal and takes place on the surface of the soil. Feeding may be both se-
lective and indiscriminate and certainly during burrowing much soil is swal-
lowed. Casting is usually beneath the soil surface. Unusual activities of this
species are the lining of burrows with pebbles, or faecal earth, and the drawing
into the burrows of leaves. The entrances to burrows may be blocked with
seeds, sticks, straws, and feathers.

Lumbricus terrestris is an important species in litter decomposition. It is also
collected annually at night by the millions as the major species of worm for fish
bait in the northern portion of North America. The relatively long life cycle
makes it unprofitable to rear this species commercially for fish bait (cf.
Aporrectodea tuberculata and Eisenia foetida). It is also gathered annually in
large numbers for biological supply institutions which distribute this species for
use in laboratory studies at high schools and universities, for L. terrestris is
nearly always the textbook example of the oligochaetes. For example, 20,000
specimens per year are shipped to South Africa for biological studies because it
cannot be found locally (Reinecke, in litt.).

Range

A native of Palaearctis, L. terrestris is now known from Europe, Iceland, North
America, South America, Siberia, South Africa, and Australasia (Gates, 1972c).
Reinecke (in litt.), however, cannot confirm its natural occurrence in South Af-
rica.

101

North American Distribution

British Columbia (Berkeley. 1968). New Brunswick (Reynolds. 1976d). Newfoundland (Smith,
1917). Nova Scotia (Reynolds. 1975a, 1976a). Ontario (Reynolds. 1972a), Prince Edward Island
(Re\nolds. 1975c). Quebec (Reynolds, 1975d, e, 1976c). Arkansas (Causey. 1952). California (Smith,
1917), Colorado (Smith. 1917). Connecticut (Reynolds. 1973c). Delaware (Reynolds, 1973a), Dis-
trict of Columbia (Smith, 1917), Hawaii (Reynolds et al., 1974), Idaho (Gates, 1967), Illinois (Smith.
1917), Indiana (Joyner, 1960), Iowa (Gates. 1967), Kansas (Gates, 1967), Maine (Smith. 1917).
Maryland (Reynolds. 1974b), Massachusetts (Reynolds, 1977), Michigan (Smith, 1917). Minnesota
(Gates, 1972c), Missouri (Olson, 1936), New Hampshire (Reynolds. 1976c). New Jersey (Davies,
1954), New York (Olson, 1940). North Carolina (Gates, 1972c), Ohio (Olson. 1928). Oregon (Mac-
Nab and McKey-Fender, 1947), Pennsylvania (Bhatti, 1965), Rhode Island (Reynolds, 1973b). Ten-
nessee (Reynolds, 1974a), Utah (Gates, 1967), Vermont (Gates, 1949), Virginia (Reynolds. 1974b),
Washington (Altman, 1936), West Virginia (Reynolds, 1974b). Wisconsin (Gates. 1972c), Greenland
(Eisen, 1872). New records: Georgia.

Ontario Distribution (Fig. 35)

Lumbhcus terrestris was first reported from Ontario by Stafford (1902), and later
by Judd (1964) and Reynolds (1972a). It is widely distributed over the province,
perhaps, in part, as a result of the fish bait industry.

ALGOMA DIST. *Hwy 17, .48 km e of Spanish, under log and paper in ditch, 13 May 72, JWR &
JEM, 0-2-0. BRANT CO. *Hwy 53, 4.35 km e of Cathcart, under log, 1 May 72, JWR, 1-0-0. *Hwy
54, 5.16 km e of Middleport, ditch, 3 May 72, JWR, 4-0-2. *Hwy 2, 6.45 km e of Paris, under lum-
ber, 3 May 72, JWR, 0-0-1. New Durham, May 30, RC, 1-1-2, ROM. BRUCE CO. *Hwy 4. 6.77 km
s of Teeswater, under logs, 5 May 72, JWR, 0-0-4. *Hwy 4. .48 km s of Hwy 9, Under logs, 5 May
72, JWR, 0-2-0. CARLETON CO. Reynolds (1972a). COCHRANE DIST. Reynolds (1972a).
DUFFERIN CO. *Hwy 10-24, 8.06 km n of Orangeville. under logs. 2 May 72, JWR, 1-1-1. 'Hwy
10-24. 7.74 km n of Camilla, under logs, 2 May 72, JWR, 9-1-0. DURHAM CO. *Hwy 401, 1.61 km
e of Newtonville, under logs, 15 May 72, JWR, 4-0-1. ELGIN CO. *Hwy 3, 2.1 km e of Wallace-
town, under log, 4 May 72, JWR, 0-0-1. FRONTENAC CO. *Hwy 15, 4.68 km s of Seeley's Bay,
under logs, 16 May 72, JWR, 1-1-0. GLENGARRY CO. *Hwy 401. 7.45 km w of Summerstown
Rd, under log, 11 May 72, JWR, 0-0-1. GRENVILLE CO. *Hwy 2, 5.16 km w of Prescott. under
rock, 10 May 72, JWR, 0-0-1. *Hwy 2, Johnstown, w.e., under logs, 1 1 May 72, JWR, 0-1-1. GREY
CO. *Hwy 21, 7.26 km w of Springmount, under rocks, 5 May 72, JWR, 9-0-1. *Hwy 10, 8.06 km s
of Flesherton, under logs, 5 May 72, JWR. 0-0-2. HALDIMAND CO. *Hwy 3, 9.03 km e of Dunn-
ville, wet ditch, 1 May 72, JWR, 0-0-1. *Hwy 3. 6.29 km w of Dunnville. 1 May 72, JWR, 0-0-1.
*Hwy 54, 2.26 km n of Caledonia, digging, 3 May 72, JWR, 1-0-1. HALIBURTON CO. 'Reynolds
(1972a). *Hwy 121, 3.22 km e of Tory Hill, dump, 16 May 72, JWR, 0-1-3. HALTON CO. *Halton
Co. Rd 3, Esquesing Community, under rocks and logs, next to the Town Hall, 4 May 73. JWR. 4-
0-0. *Hwy 7, 4.84 km e of Georgetown, under burnt log, 4 May 73, JWR, 1-0-0. HASTINGS CO.
Reynolds (1972a). *Hwy Jet 2 and 49, wet ditch, 15 May 72, JWR, 1-2-1. HURON CO. *Hwy 4,
2.74 km n of Clinton, under logs, 5 May 72, JWR, 0-1-1. KENT CO. *River Rd, 6.45 km w of Chat-
ham, under logs, 23 Apr 72, JWR & TW, 6-0-3. *Hwy 3, 9.03 km e of Port Alma, under log. 4 May
72, JWR, 0-0-1. *Hwy 3, 3.55 km e of Wheatley. digging, 4 May 72, JWR, 0-0-1. LAMBTON CO.
Hwy 21, .65 km s of Petrolia. ditch. 4 May 72, JWR. 1-1-0. LANARK CO. *Hwy 43, 3.71 km e of
Smiths Falls, under telephone pole in ditch, 1 1 May 72, JWR, 0-1-3. LEEDS CO. *Hwy 15. 2.26 km
n of Seeley's Bay, under log and concrete block in ditch, 16 May 72. JWR. 0-0-2. LENNOX AND
ADDINGTON CO. *Hwy 2. 9.68 km w of Napanee. under paper in ditch. 15 May 72, JWR. 0-0-1.
*Hwy 2, 6.61 km w of Hwy 133, under logs, 16 May 72. JWR. 0-1-1. MANITOULIN DIST. *Hwy
68, Birch Island, s.e., under logs, 13 May 72, JWR & JEM, 2-1-0. *Hwy 68, 3.22 km n of Little Cur-
rent, under log, 13 May 72, JWR & JEM, 0-0-1. MIDDLESEX CO. Judd (1964). 'Hwy 7, 2.26 km s
of Parkhill, under fence posts, 4 May 72, JWR, 2-0-1. MUSKOKA DIST. 'Hwy 11. 11.45 km n of
Severn Bridge, under pine log, 9 May 72, JWR, 0-1-0. NIP1SS1NG DIST. *Hwy 17, 8.39 km e of
Warren, house site under lumber, 13 May 72, JWR & JEM. 1-1-1. NORFOLK CO. *Hwy 6, 4.84
km s of Jarvis, under log, 1 May 72, JWR, 0-0-1. *Hwy 3. 1 1.29 km e of Delhi, under log. 1 May 72.
JWR, 0-0-1. NORTHUMBERLAND CO. *Hwy 401, 5.65 km e of Grafton, under logs. 15 May 72,
JWR. 0-0-2. ONTARIO CO. *Hwy 7. 1.61 km e of Green River, under log, 26 Apr 72, JWR, 1-0-0.
*Hwy 47. 10.48 km w of Uxbridge, digging. 9 May 72. JWR. 0-1-0. OXFORD CO. *Hw\ 59. 1.77

102

Fig. 35 The known Ontario distribution of Lumbricus terrestris.

km e of Burgessville. under log, 1 May 72. JWR. 0-0-1. *Hwy 59, .32 km n of Hickson, under logs, 3
Ma) 72. JWR. 2-1-1. PARRY"sOUND DIST. *Hwy 11, 4.52 km s of Burks Falls, under log, 9 May
72. JWR. 0-0-1. PEEL CO. *Hwy 5. 4.35 km w of Hwy 10. under mattress, 29 May 72, JWR. 0-0-2.
•Hu\ 24. 8.87 km n of Erin, under log in pasture. 29 Apr 72. JWR, 0-0-1. *Hwy 7, Brampton, e.e.,
under debris. 4 May 73. JWR, 1-1-0. Hwy 401, 1.3 km w of Dixie Rd, wet ditch. 4 May 73. JWR, 0-
0-2. PERTH CO. .81 km n of Fullarton. under rocks. 29 Apr 72. DWR & LWR, 4-2-0. 'Mitchell,
next to Collegiate, under logs. 4 May 72. JWR & DWR, 3-0-2. Hwy Jet 23 and 83. Russeldale, under
paper in wet ditch. 5 May 72, JWR, 0-0-1. PETERBOROUGH CO. *Hwy 28, 5.48 km n of Burleigh
Falls, under log. 27 Apr 72. JWR. 0-1-0. PRINCE EDWARD CO. *Hwy 33. 1.61 km s of Carrying
Place, under log in wet ditch, 15 May 72. JWR. 0-0-1. »Hwy 33. 2.58 km e of Hillier. digging, 15
May 72. JWR. 0-0-1. Hwy 33. Bloomfield, e.e.. under paper. 15 May 72. JWR. 0-0-1. RUSSELL CO.
•2.9 km e of Embrun. under logs. 1 1 May 72. JWR, 0-1-1. RENFREW CO. Reynolds (1972a). SIM-
COE CO. "Hwy 27. 5.97 km s of Cookstown. under logs, 2 May 72, JWR, 3-1-0. "Hwy 89, 1.94 km e
of Rosemont. under log in wet ditch. 2 May 72, JWR, 0-0-1. *Hwy 89, 5.48 km e of Alliston. under
log. 2 \la\ 72, JWR. 1-0-0. Barne, 14 Greenfield, digging in garden and under logs, 7 May 72, JWR
&MKG, 6-0-2. *Hw% 400. .81 km s of Hwy 103. under logs, 9 May 72. JWR. 1-1-1. SUDBURY
DIST. 'Hwy Jet 64 and 69. under rocks. 13 May 72, JWR & JEM. 1-2-1. VICTORIA CO. *Hwy 46,
3.71 km n of Hwy 7. under logs. 9 May 72. JWR. 1-0-2. Hwy 48, Bolsover, e.e.. under paper in wet
ditch. 9 May 72. JWR. 1-0-0. WATERLOO CO. *Hwy 24A.' 1 1.45 km n of Paris, under log, 3 May
72. JWR. 1-0-1. 'Hwy 97. 3.71 km w of Hwy 401. under logs. 3 May 72, JWR, 1-0-1. WELLING-
TON CO. *Hwy 24, 4.03 km n of Enn. under log. 29 Apr 72. JWR, 0-1-0. Hwy 6, University of
Guelph, on wet drivewa) behind the Soil Science Building, 2 May 72, JWR, 0-1-3. *Hwy 6, 1.13 km
s of Ennotville. under paper in wet ditch. 2 May 72. 0-0-1. *Hwy 6. 10.64 km s of Arthur, under logs,
2 May 72, JWR. 1-0-2. WENTWOR I H CO. Reynolds < 1972a). *Hwy 5. Waterdown. e.e., edge of
com (Zea maize) field, 29 Apr 72, JWR. 1-0-0. YORK CO. "Hwy 48. .32 km n of Steeles Avenue, un-
der log, 26 Apr 72. JWR. 0-1-0. 'Edenbrook Park. Islington, under logs and rocks near stream bank,
30 Apr 72, JWR & DWR. 34-1-1. Islington. 12 Country Club Drive, crawling on grass. 13 May 72,
RGR & WMR. 0-0-2. Edenbrook Park. Islington, quantitative study 1, formalin, 18 May 72. JWR.
1-0-2. Edenbrook Park. Islington, quantitative study 2, formalin, 18 May 72, JWR, 7-0-3. Scarbor-
ough Bluffs. Bnmle> Rd. garden. 28 Oct 41. JO. 1-0-8. ROM.

103

Genus Oetolasion Orley, 1885

1885 Oetolasion (part.) Orley, Ertek. Term. Magyar Akad. 15( 18): 13.

1889 Lumbricus (Oetolasion) (part.) + L. ( A llobophora) (part.) + Dendrobaena

(part.) + L. (L.j (part.) + Titanus ? (part.)-L. Vaillant, Hist. Nat. Annel.

3(1): 113. 130. 116. 121.93.
1896 Oetolasion + Allolobophora (OctolasionhRibaucourt, Rev. Suisse Zool.

4:95.
1900 Octolasium (part.)-Michaelsen, Das Tierreich, Oligochaeta 10: 504.
1930 Octolasium (part.) Stephenson, Oligochaeta (Oxford), p. 914.
1972 Octolasium-Bouche, Inst. Natn. Rech. Agron., p. 253.
1975 Octolasion-Gates, Megadrilogica 2( 1): 4.

Type Species

Oetolasion lacteum Orley, 1885 (= Enterion tyrtaeum Savigny, 1826).

Diagnosis

Calciferous sacs, in x, large, lateral, communicating vertically and widely with
gut lumen though reaching beyond oesophagus both dorsally and ventrally.
Calciferous lamellae continued onto posterior walls of sacs. Intestinal origin, in
xv. Gizzard, mostly in xvii. Extraoesophageals. passing up to dorsal trunk poste-
riorly in xii. Hearts, vi-xi. Nephridial bladders, ocarina-shaped. Nephropores,
obvious, behind xv in one regular rank on each side, just above B. Setae, behind
the clitellum not closely paired. Prostomium epilobic. Longitudinal muscula-
ture, pinnate, (after Gates, 1972c: 123, 1975a: 4).

Discussion

There has been considerable confusion concerning the spelling of this genus
name since the early 1900s. Michaelsen (1900b) changed many of the Greek
generic endings to Latin endings, i.e., Oetolasion to Octolasium and Bimastos to
Bimastus. According to the International Code of Zoological Nomenclature
(Article 32), the original spelling is the correct spelling. Therefore, Oetolasion
Orley, 1885 and Bimastos Moore, 1893 are the correct spellings and most cur-
rent oligochaetologists are now employing them. The genus Oetolasion Orley,
1885 contains only two species, O. cyaneum and O. tyrtaeum. Species in Europe
and elsewhere often placed in Oetolasion are now considered to belong in the
genus Octodrilus Omodeo, 1956 with Lumbricus complanatus Duges, 1828 as the
type (cf. Bouche, 1972, Gates, 1975a).

104

Octolasion cyaneum (Savigny, 1826)

Woodland blue worm Yer bleu des bois
(Fig. 36)

1774 Lumbricus terrestris (part.) Miiller, Verm. Terr. Fluv. 1(2): 24.

1826 Enterion cyaneum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 181.

1837 Lumbricus cyaneus Duges, Ann. Sci. Nat., ser. 2, 8: 17,21.

1845 Lumbricus stagnalis (part.) HofFmeister, Regenwiirmer, p. 35.

1867 Lumbricus alyattes Kinberg. Ofv. Vet.-Akad. Forh. Stockholm 23: 99.

1889 Lumbricus alyattes + Lumbricus ( Dendrobaena) stagnalis + Lumbricus
cyaneus-L. Vaillant. Hist. Nat. Annel. 3(1): 96, 1 18. 124.

1890 Allolobophora studiosa Michaelsen. Arch. Ver. Mechlenb. 44: 50.

1893 Allolobophora (Octolasion) cvanea (part.)-Rosa, Mem. Ace. Torino, ser. 2,

43:424.455,456.
1896 Allolobophora (Octolasion) cvanea studiosa-Ribaucourl, Rev. Suisse Zool.

4: 95.
1900 Octolasium cvancum- Michaelsen, Das Tierreich, Oligochaeta 10: 506.
1972 Octolasium cyaneum— Boxxche + O. c. var. armoricum Bouche, Inst. Natn.

Rech. Agron.. p. 258, 260.
1972 Octolasium ow?euw-Edwards and Lofty, Biol, earthworms, p. 214.
1972 Octolasion cyaneum-Gates, Bull. Tall Timbers Res. Stn. 14: 31.

Diagnosis

Length 65-180 mm. diameter 7-8 mm, segment number 140-158, prostomium
epilobic, first dorsal pore 11/12 or 12/13. Clitellum xxix-xxxiv. Tubercula pu-
bertals xxx-xxxiii. Setae closely paired anteriorly, CD<^AB<^BC<^AA<^DD,
and widely paired posteriorly, AB^>BC^>CD. Setae of x, xviii, xix, xx, xxi fre-
quently on white genital tumescences. Male pores on xv with well-defined nar-
row papillae. Seminal vesicles, four pairs in 9-12, with the pairs in 11 and 12
larger than the pairs in 9 and 10. Spermathecae, two pairs opening between c
and d in 9 10 and 10/11. Body cylindircal but octagonal posteriorly. Colour,
blue-grey or whitish.

Biology

This species is known from soils of pH 5.2-8.0 and may well be ubiquitous with
respect to this factor. Gates (1972c) records it from under stones in water, in
moss, stream banks, and other limnic habitats. It is known also from ploughed
fields, wet sand, forest soils, and from caves in Europe. Cernosvitov and Evans
( 1947) reported it mostly from under stones and occasionally under moss. Gates
( 1973a) found the species under logs and under rocks near stream beds. Under
logs and rocks was the most common site for O. cyaneum in Ontario.

Activity may be year round but in central Maine summer drought and winter
freezing impose two periods of inactivity (Gates, 1972c), and this is probably the
case in Ontario. In experimental studies casting was below ground but occa-
sional surface casting has been reported (Gates, 1972c). O. cyaneum is obligato-

105

1st dp—;

XXV

XXX

XXXV

mp

Fig. 36 External longitudinal views of Octolasion cvaneum showing taxonomic characters, a Lat-
eral view. b. Ventrolateral view. (ONT: Durham Co., cat. no. 3384)

106

nl\ parthenogenetic (Reynolds. 1974c): copulation has not been recorded and
ma\ never have boon observed (Gates, 1972c). This species is relatively rare in
North America and is of little or no economic importance.

Range

A native of Palaearctis, O. cyaneum is now known from Europe, Iceland. North

America. South America. India. Azores, and Australasia (Gates, 1972c).

North American Distribution

British Columbia (Wickett, 1967), Ontario (Reynolds, 1972a), Nova Scotia (Reynolds. 1976a), Cali-
fornia (Gates. 1966). Colorado (Gates. 1967). Georgia (Gates. 1973a), Indiana (Reynolds et al.,
1974). Iowa (Evans. 1948a). Maine (Gates. 1966). Massachusetts (Reynolds. 1977), New York
(Schwert. 1976). North Carolina (Gates. 1973a). Pennsylvania (Reynolds. 1974b), Tennessee (Rey-
nolds. 1974ai. Virginia (Gates. 1973a), Washington (Gates. 1973a). New records: Quebec. Mississip-
pi-

Ontario Distribution (Fig. 37)

Octolasion cyaneum was first reported from Ontario by Reynolds (1972a) and
the report presented here is only the second account of this species in Ontario.
O. cyaneum has been obtained only from Haliburton and Perth counties.

H ALIBL'RTON CO. 'Reynolds (1972a). PERTH CO. .81 km n of Fullarton. under rocks. 29 Apr
72. DWR& l.WR. 1-1-0.

Fig. 37 The known Ontario distribution of Octolasion cyaneum

107

Octolasion tyrtaeum (Savigny, 1826)

Woodland white worm Ver blanc des bois
(Fig. 38)

1826 Enterion tyrtaeum Savigny, Mem. Acad. Sci. Inst. Fr. 5: 180.

1837 Lumbricus tyrtaeus-Duges, Ann. Sci. Nat., ser. 2, 8: 17, 22.

1845 Lumbricus communis cyaneus + L. stagnalis (part.) Hoffmeister, Regen-

wiirmer, p. 24, 35.
1881 Lumbricus terrestris var. lacteus + L. t. var. rubidus Orley, Math. Term.

Kozlem. Magyar Akad. 16: 584.

1884 Allolobophora profuga Rosa, Lumbric. Piemonte, p. 47.

1885 Octolasion rubidum-Or\ey + O. profugum-Ox\ty + O. gracile Orley +
O. lacteum-6r\ey, Ertek. Term. Magyar Akad. 15(18): 16, 17, 18, 21.

1889 Lumbricus (Allobophora) profugus + L. (O.) gracilis L. Vaillant, Hist. Nat.
Annel. 3(1): 113.

1896 Allolobophora (Octolasion) rubida-R\ba.ucourt + A. (O.) gracilis-
Ribaucourt + A. sylvestris Ribaucourt, Rev. Suisse Zool. 4: 63, 65, 67,
95.

1900 Octolasium lacteum-Michaeteen, Das Tierreich, Oligochaeta 10: 506.

1900 Allolobophora (Octolasion) profuga-MichaeXsen, Abh. Nat. Verh. Ham-
burg 16(1): 16.

1900 Allolobophora profuga-Smhh, Bull. Illinois St. Lab. Nat. Hist. 5: 441.

1917 Octolasium lacteum-Smith, Proc. U.S. Natn. Mus. 52(2174): 178.

1952 Octolosium ladeum (laps.)-Goff, Amer. Midi. Nat. 47: 484.

1971 Octolasium lacteum-Crossley, Reichle and Edwards, Pedobiologia 11:
71.

1972 Octolasium lacteum-Edwards and Lofty, Biol, earthworms, p. 216.

1972 Octolasium lacteum lacteum + O. I. gracile-Bouche, Inst. Natn. Rech.

Agron., p. 253, 257.
1972 Octolasion tyrtaeum-Gales, Bull. Tall Timbers Res. Stn. 14: 35.

Diagnosis

Length 25-130 mm, diameter 3-6 mm, segment number 75-150, prostomium
epilobic, first dorsal pore 9/10-13/14, usually 11/12. Clitellum xxx-xxxv. Tu-
bercula pubertatis xxxi-xxxiv. Setal pairings as in Octolasion cyaneum. Fre-
quently setae a and/or b on xxii and occasionally on ix-xii, xiv, xvii, xix-xxiii,
xxvii, xxxvii, or xxxviii are on genital tumescences and modified into genital se-
tae. Male pores on xv and on large glandular papillae extending over xiv and
xvi, occasionally limited to xv. Seminal vesicles, four pairs in 9-12, with pairs in
11 and 12 larger than pairs in 9 and 10. Spermathecae, two pairs opening on
level C or between c and d in 9/10 and 10/1 1. Body cylindrical but slightly oc-
tagonal posteriorly. Colour variable, milky white, grey, blue, or pink.

Biology

Reported from soils of pH 5.5-8.08, O. tyrtaeum has been found under stones
and logs, in peat, leaf mould, compost, forest litter, gardens, cultivated fields
and pastures, bogs, stream banks, in springs, and around the roots of submerged
vegetation (Gates, 1972c). The species is also known from caves in Europe and

108

1st dp }

— mp

Fig. 38 External longitudinal views of Octolasion tyrtaeum showing taxonomic characters, a. Dor-
solateral view, b Ventrolateral view. (ONT. Bruce Co., cat. no. 7557)

109

North America. Smith (1917) reported this species as commonly found under
logs, leaf mould, and debris of various kinds, in compost heaps, and to some ex-
tent in the soil. Some workers believed this species preferred rich, moist organic
material (Causey, 1952), while others presented data to the contrary (Eaton,
1942). Octolasion tyrtaeum was the most abundant species in Tennessee (Rey-
nolds et al., 1974) and was obtained under logs, debris, and rocks, and by dig-
ging. In Ontario it was most frequently found under logs.

Activity may be year round although summer drought and winter cold may
impose two rest periods. Octolasion tyrtaeum is an obligatorily parthenogenetic
species (Gates, 1973a; Reynolds, 1974c) and copulation occurs below the sur-
face of the soil. The species is of little economic importance although one dealer
in Michigan is reported to have sold it for fish bait (Gates, 1972c).

Range

A native of Palaearctis, O. tyrtaeum is known from Europe, North America,
South America, Asia, Africa, and Australia (Gates, 1972c).

North American Distribution

British Columbia (Wickett, 1967), Manitoba (Gates, 1972c), Nova Scotia (Reynolds, 1975a, 1976a),
Ontario (Reynolds, 1972a), Quebec (Reynolds, 1975d, e, 1976c). Alabama (Gates, 1972c), Arkansas
(Causey, 1952), California (Smith, 1917), Connecticut (Reynolds, 1973c). Florida (Gates. 1972c),
Georgia (Gates, 1972c), Idaho (Gates. 1967), Illinois (Smith, 1917). Indiana (Reynolds, 1972e). Iowa
(Gates, 1967), Kentucky (Allee et al., 1930) Maine (Gates. 1961), Maryland (Reynolds, 1974b),
Massachusetts (Reynolds. 1977). Michigan (Murchie, 1956). Minnesota (Gates. 1973a), Missouri
(Olson, 1936), Nebraska (Gates, 1967), New Jersey (Davies, 1954), New York (Olson. 1940), North
Carolina (Gates, 1973a), Ohio (Smith, 1917), Oregon (MacNab and McKey-Fender, 1947). Pennsyl-
vania (Davies, 1954), South Carolina (Gates, 1973a), Tennessee (Reynolds, 1974a), Utah (Gates.
1967), Virginia (Gates, 1973a) West Virginia (Gates, 1959). New records: Alaska. Mississippi, Wis-
consin.

Ontario Distribution (Fig. 39)

Octolasion tyrtaeum was first reported from Ontario by Reynolds (1972a) and
the present study is only the second report of the species from Ontario. It is
more widely distributed than O. cyaneum and there are two main centres of dis-
tribution, the first around the Kawartha Lakes region in central southern Ontar-
io, and the second in western southern Ontario running along the Niagara Es-
carpment and to the west.

BRANT CO. *Hwy 54, .64 km w of Onondaga, under log in wet area, 3 May 72, JWR, 0-0-1.
BRUCE CO. *Hwy 4, 6.77 km s of Teeswater, under log, 5 May 72. JWR. 0-0-1. *Hwy 4. .48 km s
of Hwy 9, under logs, 5 May 72, JWR, 2-2-4. Hwy 21, 2.57 km n of Tiverton, under lumber. 5 May
72, JWR, 9-0-2. DURHAM CO. *Hwy 7A, 2.1 km e of Nestleton Station, under log, 26 Apr 72,
JWR, 0-0-1. ELGIN CO. Hwy 3, 9.19 km e of St. Thomas, under logs, 4 May 72. JWR. 2-2-8.
GREY CO. *Hwy 6, 6.94 km s of Dornoch, under logs, 5 May 72, JWR. 2-1-2. *Hwy 4, 8.22 km e of
Durham, under logs, 5 May 72, JWR, 1-0-1. HALIBURTON CO. •Reynolds (1972a). HALTON
CO. Hwy 5, 5.81 km e of Hwy 25. under paper and leaves in ditch. 29 Apr 72. JWR, 1-0-1. *Halton
Co. Rd 3, Esquesing Community, under rocks and logs next to the Town Hall. 4 May 73. JWR. 0-
4-3-1. Campbellville, rapids in cold stream, 15 May 66. IMS, ST, & RNS. 0-1-0, ROM-I6. HURON
CO. *Hwy 83, 6.77 km w of Russeldale, under logs, 5 May 72. JWR, 3-0-1. NORTH UMBER-

110

Fig. 39 The known Ontario distribution of Octolasion tyrtaeum.

LAND CO. *Hwy 45. 4.84 km n of Baltimore, under logs, 28 Apr 72, JWR, 1-4-1. OXFORD CO.
•Hwy 97, Washington, w.e., under logs, 3 May 72, JWR, 0-1-3. PEEL CO. *Hwy 5, .81 km e of
Dixie Rd. under logs. 29 Apr 72, JWR. 0-1-1. PETERBOROUGH CO. *Hwy 28, Lakefield College,
under logs near waterfront, 27 Apr 72, JWR & CWR, 4-5-8. VICTORIA CO. *Hwy 7, 7.26 km e of
Hwj 35. under log, 26 Apr 72, JWR, 0-3-1. WATERLOO CO. *Hwy 24A, 1 1.45 km n of Pans, un-
der log, 3 May 72, JWR. 4-0-2. *Hwy 97, 3.71 km w of Hwy 401, under log, 3 May 72, JWR, 0-1-0.
Hwy 7-8. 1.77 km w of Petersburg, under log, 3 May 72, JWR, 1-2-1. Waterloo, Beechwood South,
on sidewalk after rain (evening), 15 Jun 75, 0-0-1, DPS, UW-0004. WELLINGTON CO. *Hwy 6,
2.26 km n of Aberfoyle, under log, 29 Apr 72, JWR, 0-0-1. *Hwy 24, 5.16 km e of Eramosa, under
logs in cedar ( Thuja occidentalis) woodlot, 29 Apr 72, JWR, 0-1-1. *Hwy 6, 10.65 km s of Arthur, un-
der log, 2 May 72, JWR. 0-0-1. WENTWORTH CO. Reynolds (1972a). *Hwy 6, 5.32 km n of Hwy
5, under logs, 29 Apr 72, JWR, 1-0-2.

Ill

Family SPARGANOPHILIDAE Michaelsen, 1921

Diagnosis

Digestive system: without gizzard, calciferous glands, lamellae, caeca, typhlo-
sole, or supra-intestinal glands, with an intestinal origin in front of the testis seg-
ments. Vascular system: with complete dorsal and ventral trunks, two pairs of
anterior lateroparietal trunks, one of which passes to the dorsal vessel and the
other to the ventral vessel in xiv, but without subneural and supra-oesophageal
trunks. Hearts: lateral, moniliform, in vii-xi. Blood glands: protuberances from
capillaries in septal glands. Nephridia: holoic. aborted at maturity in first 12 or
more segments, avesiculate, peritoneal cells investing postseptal portions en-
larged, ducts without muscular thickening passing into parieties in AB. Nephro-
pores: inconspicuous, in AB. Setae: eight per segment. Dorsal pores and pig-
ment, lacking. Prostomium, zygolobous. Anus, dorsal. Reproductive apertures:
all minute and superficial, female pores in xiv, spermathecal pores in front of
testis segments. Clitellum multilayered, including male pore segment. Seminal
vesicles, trabeculate. Ovaries, in xiii, each terminating in a single eggstring. Ova,
not yolky. Ovisacs, in xiv, small and lobed. Spermathecae, adiverticulate (after
Gates, 1972c: 314).

Type Genus

Sparganophilus Benham, 1892 by monotypy (Michaelsen, 1921).

Discussion

This family is still monotypic and contains but a few nearctic species, only one
of which occurs in Canada. Originally, Benham placed the genus in the Rhino-
drilidae, a taxon no longer employed by oligochaetologists. Following Michael-
sen (1900b), most authors have placed Sparganophilus in various subdivisions of
the Glossoscolecidae. In a recent review Brinkhurst and Jamieson (1971) still
considered the genus as belonging to the Glossoscolecidae.

Genus Sparganophilus (Benham, 1892)

1892 Sparganophilus Benham, Quart. J. Micros. Soc. (n.s.), 34: 155.

1895 Sparganophilus-Smilh, Bull. 111. St. Lab. Nat. Hist. 4(5): 142.

1900 Sparganophilus-Michaeken, Das Tierreich, Oligochaeta 10: 463.

1921 Sparganophilus-Michaehen, Arch. Naturg. 86(A): 141.

1930 Sparganophilus-Slephenson, Oligochaeta (Oxford), p. 899.

1971 Sparganophi lus-Brmkhursl and Jamieson, Aquatic Oligochaeta World, p.
810.

1972 Sparganophilus-Gates, Trans. Amer. Philos. Soc. 62(7): 314.

112

Type Species

Sparganophilus tamesis Benham, 1892.

Diagnosis

Calciferous gland and gizzard, absent. Hearts, in vii-xi. Nephridia, absent in ce-
phalic region (segments i-xii). Nephropores, inconspicuous, in AB. Setae,
paired. Prostomium zygolobic. Lateral lines, absent. Colour, unpigmented.

Sparganophilus eiseni Smith, 1895

American mud worm Ver americain de la vase
(Fig. 40)

1895 Sparganophilus eiseni Smith, Bull. 111. St. Lab. Nat. Hist. 4(5): 142.

1906 Sparganophilus eiseni-Moove. Bull. Bur. Fish. 25: 153.

1911 Helodrilus elongatus Friend, Zoologist, ser. 4, 15: 192.

1921 Sparganophilus elongatus-Friend. Ann. Mag. Nat. Hist., ser. 9, 7: 137.

1934 Pelodrilus cuenoti Tetry, C.R. Acad. Sci. Paris 199: 322.

1934 Eiseniella tetrahedra (laps.)-Moon, J. Anim. Ecol. 3: 17.

Diagnosis

Length 150-200 mm, diameter uniformly about 2.5 mm, segment number
165-225, prostomium zygolobic, dorsal pores absent. Clitellum xv-xxv. Tuber-
cula pubertatis xvii-xxii. Setae closely paired, with setae c and d above mL line.
Male pores on xix, female pores on xiv. Seminal vesicles, two pairs, in 1 1 and
12 with a pair of testes in 10 and 1 1. Spermathecae, three pairs, in 7, 8, and 9
with ducts opening just ventral to level C in 6/7-8/9. Gizzard, calciferous
glands, and tvphlosole absent. Prostate-like glands, four pairs, in 13, 14, 15, and
16. Body cylindrical. Colour, unpigmented but sometimes appearing pink with
blue and green iridescence.

Biology

This species is limicolous and confined to the muddy banks of streams, rivers,
ponds, and lakes. Smith (1915) found it to be abundant in the mud of the bot-
tom and margins of many waters east of the Mississippi River. Olson (1928) re-
ported it along the shores of Lake Erie, and similar habitats are recorded for
New York (Lake Ontario) (Olson, 1940), New Jersey (Davies, 1954), Louisiana
(Harman. 1965) and Tennessee (Reynolds, MS).

S. eiseni is relatively rare in North America except for an area bounded by the
Great Lakes, Mississippi River, Atlantic Ocean, and the Gulf States. It is as-
sumed to be amphimictic (Reynolds, 1974c).

113

XXVIII

Fig. 40 External longitudinal views of Sparganophilus eiseni showing taxonomic characters, a.
Dorsolateral view, b Ventrolateral view. (GA: Thomas Co., cat. no. 3285)

114

Range

A native of North America. S. eiseni is known also from Central America and
has been introduced into England and France (Brinkhurst and Jamieson, 1971;
Gates, 1972c).

North American Distribution

Ontario (Moore. 1906), Alabama (Gales. 1967). Connecticut (Reynolds, 1973c), Florida (Smith.
18%). Illinois (Smith. IS95). Indiana (Heimburger, 1915). Iowa (Hague. 1923), Louisiana (Harman.
1965), Maryland (Reynolds, \^-Xb). Massachusetts (Reynolds, 1977). Michigan (Moore, 1906), New
Jersej (Davies, 1954). New, York (Olson. 1940). Ohio (Olson. 1928). Oregon (MacNab and McKey-
Fender. 1947). New records: Arkansas, Georgia, Kentucky, Mississippi, North Carolina, Pennsylva-
nia, Tennessee. Virginia. Wisconsin.

Ontario Distribution (Fig. 41)

Sparganophilus eiseni has been previously recorded from only two sites in On-
tario (Moore. 1906).

BRCCE CO. Howdenvale, 2 metres depth. Lake Huron, airlift sampler, 1 1 Jul 75. DRB, 0-1-0. UW-
0001. HALDIMAND CO. Selkirk Provincial Park, 2 metres depth. Lake Erie, airlift sampler. 5 Aug
75. DRB. 0-1-3. L'W-0001. HALTON CO. Sixteen Mile Creek, intersection of creek and Lower
Baseline Road. 1 Jun 71. AT. 0-0-6. ROM. KENT CO. Rondeau Harbour, 30 Aug 99. JPM. 1-1-0.
\\SP-41Sa i\loore. 1906). MANITOULIN DIST. 2 metres depth. Georgian Bay, airlift sampler
while scuba diving. 17 Aug 74. DRB. 0-0-1. UW-0001. NORFOLK CO. Long Point, 23 Aug 99,
JPM, 4-0-0. ANSP-382 (Moore. 1906). PARRY SOUND DIST. Parry Sound, 3 metres depth, in
sheltered shallow bay, Ekmann dredge. 18 Aug 75. RLH. 0-0-1, UW-box 1.

^r>

Fig. 41 The known Ontario distribution of Sparganophilus eiseni.

115

Distribution and Ecology

Even among oligochaetologists the question of megadrile distribution, particu-
larly in North America, has long been controversial. Except for the southern tip
of Queen Charlotte Island (British Columbia), almost the entire megadrile
fauna in Canada is composed of species of the family Lumbricidae that are pri-
marily European forms. Of the 19 Ontario species, for example, 18 are lumbri-
cids. Two species, Bimastos parvus (Lumbricidae) and Sparganophilus eiseni
(Sparganophilidae), are natives of North America.

There are 16 currently accepted genera in the family Lumbricidae:
Allolobophora, Aporrectodea, Bimastos, Dendrobaena, Dendrodrilus, Eisenia, Ei-
seniella, Eiseniona, Eisenoides, Eophila, Helodrilus, Kritodrilus, Lumbricus, Octo-
lasion, Octodrilus, and Satchellius. Of these, the genera Eiseniona, Eophila, Helo-
drilus, Kritodrilus, Octodrilus, and Satchellius are restricted to Europe and a few
adjacent areas, such as North Africa (in the case of Eophila), except that four
specimens of Satchellius were recorded in New Jersey. Two genera, Bimastos
and Eisenoides, are nearctic but are restricted to the southeastern region of the
United States except for two species, Bimastos beddardi and B. panus, which
have been carried around the world. B. parvus has been reported from Canada
(Reynolds, 1972a), but subsequent investigation (E.L. Bousfield, pers. comm.,
1973) nullified this as the only Canadian natural record because the one collec-
tion of four clitellate adults came from the Arboretum at the Central Experi-
mental Farm, Department of Agriculture, Ottawa.

There are five other nearctic megadrile genera: Argilophilus and Diplocardia
(Acanthodrilidae), Komarekiona (Komarekionidae), Lutodrilus (Lutodrilidae),
and Sparganophilus (Sparganophilidae) (Reynolds, 1976b; Gates, 1977, 1974d,
McMahan, 1976). The first four have not been reported from Canada or from
any areas north of the southern limit of Quaternary glaciation in the United
States. Sparganophilus eiseni has been reported from Canada once (Moore,
1906) but has never been recorded again in Canada until now. This species has
also been introduced, probably by man, into Britain and France.

There is little disagreement among specialists that the centre of origin of the
Lumbricidae is in Europe. Here the family is a very diverse group and the num-
ber and variety of species in the genera are greater than anywhere else in the
world. There is less agreement, however, as to how those lumbricids became es-
tablished in North America, a problem that has been discussed in detail most
recently by Gates (1967, 1970) and Omodeo (1963) and reviewed by Reynolds
et al. (1974) and Reynolds (1975f, 19760-

Assuming that lumbricids have indeed travelled from Europe to North Amer-
ica, either they must have migrated actively by their own power, or they must
have been transported passively by some agent. Since the Atlantic Ocean now
forms an effective barrier to their active migration some workers have hypothes-
ized that long ago they arrived in North America by migration via a land-
bridge, or were perhaps aided by continental drift (e.g., Omodeo, 1963). Such
explanations, however, clearly are inadequate to explain the occurrence of some
of the same species in South Africa, Australasia, and South America. Moreover,
if such hypotheses are accepted then it is necessary to explain why it should be
that of all the North American earthworms only the European lumbricids sur-

116

vived the Quaternary glaciations that otherwise depauperated the North Ameri-
can megadrile fauna.

In fact Gates (1961. 1^70) and Reynolds (1975f, 19760 have assembled a
great deal of data indicating that during the Quaternary all the earthworms of
glaciated North America were exterminated and such extermination extended,
perhaps, almost as far as the area north of the Appalachian Mountains. Early
settlers, for example, found that glaciated portions of the United States and
Canada had no earthworms excepting where they were introduced (see Gates,
1967). Consequently, and contrary to the usual belief, surviving species did not
follow the retreating glaciers. The earthworm species currently found in regions
formerh covered b\ the ice sheets are all introductions, probably by man, either
from other continents or from refugia south of the Appalachians. In recording
the occurrence of some European lumbricids in South Africa, Australasia, and
South America, the efficacy of transportation by man has rarely been ques-
tioned and there is no reason why this method of introduction need be denied
for the large European element in the North American earthworm fauna. Gates
( 1966) therefore suggested that the colonization of North America by European
lumbricids has occurred since 1500 A.D. and was made primarily by European
settlers who brought potted plants and other agricultural material with them to
the New World. Certainly this hypothesis explains both the diversity of the Eu-
ropean element in glaciated North America and the near absence of nearctic en-
demics in the same region. The relative merits of the opposing views of Omodeo
and Gates have recently been assessed from a theoretical viewpoint by Ball
(1975). who concluded that the rival hypotheses were not mutually exclusive,
and that on logical grounds it was not possible to prefer one over the other.

Our detailed knowledge of megadrile distribution within North America is
rather limited and a summary of the North American biogeographical surveys
to date is presented in Table 1 .

Little work has been done on the ecology and distribution of Canadian
earthworms and it is an open field of study. From a study of 14 species from
Maine. 13 of which occur also in Ontario, Gates (1961) concluded that dietary
preferences might be an important factor influencing distribution. He divided
these species into three groups.

The first group comprised those species that pass much soil through the intes-
tine and hence are termed geophagous, viz., A. chlorotica, Ap. longa, Ap. tubercu-
lata, Ap. turgida, E. rosea, L. terrestris, and O. cyaneum. These species are known
from soils of pH as low as 4.5-5.5 and the kinds of soil seem to be of little sig-
nificance. Within this group only L. terrestris normally and regularly copulates
at the surface. Because these species are tolerant of different soil types and oc-
cur in a wide range of habitats they are likely to be widely distributed, aided es-
pecially by the activities of man.

The second group, containing only El. tetraedra, is limiphagous (mud-eating)
or hmicolous (mud-inhabiting). Outside of Maine other species such as
Sparganophdus eiseni and possibly A. chlorotica belong in this group, the mem-
bers of which thrive in mud well under water or in saturated soils. The fact that
on a few occasions El. tetraedra has been found to be geophagous suggests an
adaptability that would prove advantageous if the animal were introduced to a
new area.

The third group consisted of the litter feeders: E.foetida, D. octaedra, Dd. ru-
in

Table 1. Regional earthworm surveys in North America.

Region

Number of

Number of

Total

References

species

counties

number of

obtained

sampled

counties in
state or province

Arkansas

21

22

75

Causey (1952, 1953)

Cape Breton

14

4

4

Reynolds (1975a)

Connecticut

21

8

8

Reynolds (1973c)

Delaware

15

3

3

Reynolds (1973a)

Gaspe

15

8

8

Reynolds (1975e)

Iles-de-la-Madeleine

9

8

10'

Reynolds (1975b)

Indiana

25 +

92

92

Reynolds (MS)

Louisiana

8>

64

64

Tandy (1969)

Maryland

26

23

23

Reynolds (1974b)

Massachusetts

21

13

14

Reynolds (1976e)

Michigan

18

51

83

Murchie(1956)

Missouri

21

31

115

Olson (1936)

New Brunswick

13

15

15

Reynolds (1976d)

New York

18

29

62

Olson (1940). Eaton (1942)

Nova Scotia

15

18

18

Reynolds (1976a)

Ohio

22

55

88

Olson (1928. 1932)

Ontario

18

52

54

Reynolds'

Oregon

25

10

36

MacNab and McKev-Fender
(1947)

Prince Edward Island

12

3

3

Reynolds (1975c)

Quebec (sud cote)

15

37

37

Reynolds (1975d. 1976c)

Tennessee

44 + «

95

95

Reynolds (1974a, MSS)

Washington

16.

9

39

Altman(1936)

'Municipality and iles.

2Tandy was concerned only with the genus Pheretima. Other reports by Gates (1965 and 1967) bring

the state list to 25 species.

'Data included in this book.

"•Includes species currently in manuscript.

bidus, L. castaneus, and L. rubellus. Litter includes all types of accumulation of
organic matter such as leaves, manure, compost, etc. The activities of farmers
and horticulturists greatly affect the distribution of these forms, and also, of
course, of geophagous species that may stray into organic matter. In contrast,
prior to hibernation, litter feeders become geophagous when they burrow down
to the levels where they hibernate.

The extent to which Julin's (1949) ecological classification of the Lumbricidae
(p. 5) can be applied to the Ontario earthworms is unknown. Only Reynolds et
al. (1974) have attempted to apply the system to a North American fauna. Con-
sidering here only those species common to both Tennessee and Ontario, they
placed Allolobophora chlorotica, Dendrobaena octaedra, Dendrodrilus rubidus,
Eisenia foetida, E. rosea, Eisenielta tetraedra, Lumbricus rubellus, and Octolasion
cyaneum among the hemerophiles. The hemerodiaphores included Ap. trape-
zoides, Ap. turgida, and O. (yrtaeum, and information was lacking for Ap. longa,
Bimastos parvus, and Lumbricus terrestris. There were none that appeared to be
true hemerophobes.

118

There is evidence to suggest, how ever, that this classification is not strictly ap-
plicable to Ontario. Thus, in Tennessee Ap. tuberculata, E. rosea, and /..
terrestris are sparseh distributed and probabK will become widespread and
established only with the aid of human culture, whereas in Ontario these species
alread) are widespread in great numbers (Reynolds et al., 1974). For these spec-
ies there is a change from the hemerophilic to the hemerodiaphoric category
from Tennessee to Ontario. The reverse is the case for Ap. trapezoides and
Octolasion tyrtaeum, for these are of restricted distribution in Ontario and wide-
spread in Tennessee (Reynolds et al., op. cit.).

There is no cause to consider any of the local lumbricids as hemerobionts ex-
cept in areas where the natural habitats have been completely destroyed, as in
the Sudbury-Copper Cliff region of the Sudbury District of Ontario.

Ontario lumbricids. like those of Maine (Gates. 1961). must have become ha-
bituated to rather low temperatures. Nothing is known about the state in which
winter is spent but. in Maine at least. E. rosea has been found active under
leaves surrounded by snow and frost and active specimens of Ap. tuberculata, L.
castaneus. D. octaedra and Dd. rubidus, some in breeding condition, have been
found in a ridge surrounded with 75-cm deep snow. L. terrestris has been found
at night on frozen ground and it seems that melt water from snow and ice is not
too cold for many of these lumbricids. Presumably the same holds for the On-
tario species, though it is evident that a study of environmental and climatic fac-
tors affecting the distribution and activity of the Ontario fauna is greatly need-
ed.

In determining the presence and distribution of the earthworms in Ontario,
330 new county and district records were established. Ten of these new distribu-
tion records were found in the ROM unidentified collections by the author, two
came from Carleton University collections, and the remainder were collected by
the author. For a detailed breakdown of these records see Table 2 and the On-
tario Distribution records for each species.

When careful and thorough collection procedures are followed, earthworm
species will be found in associations. The size and number of associations ob-
tained in this study are reported in Table 3. The maximum number of species
found at any one site was eight, while single species, or associations of two or
three, were most frequently encountered.

Associations vary considerably in size depending on habitat and geography.
Reviews of some previously reported habitat associations were presented by
Bouche (1969b) and by Reynolds (1972b, 1973d). Since then, another study
from one of Bouche's (1969b) stations has been reported (Reynolds, 1972d).
This latter report records lumbricid species associations of two, five, and eight
for woodlots in north central France. In the previous Ontario study (Reynolds,
1972a), species associations ranged from four with most sites in the single spec-
ies and two species association categories. The most common association was
Aporrectodea tuberculata — Lumbricus rubellus. Recent lumbricid studies in Ten-
nessee (Reynolds et al., 1974) involving almost 1,700 sites and 18,000 specimens,
recorded associations most frequently of between three and four species per site
with a range of one to eight species. When the remaining families of earthworms
have been analyzed these values should increase.

The most frequent associations found in this study were: 1) Aporrectodea
tuberculata — Aporrectodea turgida — Eisenia rosea, 2) Aporrectodea

119

Table 2. Summary of numbers of new county Distribution Records for Ontario earthworms recorded
herein.

Species

Number of new
county records

First provincial
record

A llolobophora chloroiica
Aporrectodea icierica
Aporrectodea longa
Aporrectodea trapezoides
Aporrectodea tuberculata
Aporrectodea turgida
B i mas l os parvus
Dendrobaena octaedra
Dendrodrilus rubidus
Eisenia foetida
Eisenia rosea
Eiseniella letraedra
Lumbricus castaneus
Lumbricus festivus
Lumbricus rubellus
Lumbricus terrestris
Oclolasion cyaneum
Oclolasion tyrtaeum
Sparganophilus eiseni
Total

25

0

0

34

41

41

0

9

25

7

39

11

1

7

33

37

1

14

5

330

Stafford (1 902)'
Schwert(1977)
Smith (1917)

Reynolds (1972a)
Eisen(1874)
Eisen(1874)
Reynolds (1972a)
Reynolds (1972a)
Eisen(1874)
Stafford (1902)'
Stafford (1902)'
Eisen(1874)
Eisen(1874)
Stafford (1902)'
Stafford (1902)'
Stafford (1 902) '
Reynolds (1972a)
Reynolds (1972a)
Moore (1906)

'First reported by Stafford (1902) but he gave no collection details and no locations other than On-
tario, Quebec, New Brunswick, and Nova Scotia.

tuberculata — Aporrectodea turgida — Lumbricus rubellus, 3) Aporrectodea
tuberculata — Lumbricus terrestris, 4) Aporrectodea tuberculata — Aporrectodea
turgida, and 5) Aporrectodea tuberculata or Aporrectodea turgida — Dendrodrilus
rubidus. Because of the limited diversity of earthworms in Ontario, one should
not be surprised at the repetitive nature of the species associations. Even though
19 species have been recorded from Ontario, this does not represent a particu-
larly diverse fauna when compared with other political regions of North Amer-
ica (Table 1) because the fauna comprises principally exotic European forms;
the nearctic endemic species characteristic of more southern regions are absent.
Tennessee, for example, has over 44 species known to exist in various habitats
within its boundaries (Reynolds et al., 1974). Three of Ontario's 19 species
(Apporectodea icterica, Lumbricus castaneus, and L. festivus) are not found in
Tennessee, the remainder are. The many species in Tennessee not found in On-
tario belong mainly to the nearctic genera Bimastos, Diplocardia, Eisenoides, and
Komarekiona. Recent observations by this author (Reynolds, 1973a, 1973b,
1973c, and 1974b) indicate that the habitats utilized by the nearctic species in
the southern portions of the United States are those which in Ontario are in-
vaded and utilized by exotic European lumbricids. The success of these species
in these habitats no doubt is a result of the lack of competition from endemic
forms.

120

Table 3. Earthworm associations in Ontario b> counties and districts.

Counties and Numb

er of

Frequency of obtaining

he following nurr

iber

districts collec
in cou

or disi

ions

nls

of species per

collection

ricl

1

2

3

4

5

6

7

8

Mgoma

3

0

.33

.33

.33

0

0

0

0

Brant

6

0

.50

.33

0

.17

0

0

0

Bruce

8

.12

.12

.12

.38

.12

0

.12

0

Carieton

5

0

.20

.60

.20

0

0

0

0

Cochrane

3

0

.67

0

0

.33

0

0

0

Dufferin

7

0

.42

.29

0

0

0

0

0

Dundas

7

.14

.57

.29

.29

0

0

0

0

Durham

6

.33

0

.33

0

.17

.17

0

0

1 Igin

8

.12

.25

.12

.25

.12

.12

0

0

\ svC\

7

.29

.29

.14

.29

0

0

0

0

Frontenac

8

.25

,25

.38

.12

0

0

0

0

Glengarr\

4

0

.25

.75

0

0

0

0

0

Grenville

5

0

.40

.40

.20

0

0

0

0

Circs

6

0

0

.83

.17

0

0

0

0

Haldimand

6

.33

0

.17

.33

.17

0

0

0

Haliburton

72

.52

.33

.12

.03

0

0

0

0

Halton

9

.33

.44

0

.11

0

.11

0

0

Hastings

7

.43

.43

.14

0

0

0

0

0

Huron

7

0

0

.57

.29

.14

0

0

0

Kent

6

.33

0

.33

.17

0

.17

0

0

Kenora

1

0

1.00

0

0

0

0

0

0

Lambton

6

.17

.50

.33

0

0

0

0

0

Lanark

5

.20

.20

.40

.20

0

0

0

0

Leeds

7

0

.72

.14

0

.14

0

0

0

Lennox & Addington

5

0

.60

.40

0

0

0

0

0

Manitoulin

6

.17

.17

.17

.50

0

0

0

0

Middlesex

5

.20

.20

.20

.20

0

0

.20

0

\luskoka

9

.56

.44

0

0

0

0

0

0

Niagara

4

0

0

.25

.50

.25

0

0

0

Nipissing

6

.33

0

.50

0

.17

0

0

0

Norfolk

7

.57

.29

0

0

.14

0

0

0

Northumberland

9

.44

.11

.33

.11

0

0

0

0

Ontario

7

.43

.14

0

.29

0

.14

0

0

Oxford

7

.29

.14

.14

.29

.14

0

0

0

Parr\ Sound

11

.45

.45

.10

0

0

0

0

0

Peel'

6

.17

0

.33

.33

.17

0

0

0

Perth

4

0

.50

0

0

.25

0

.25

0

Peterborough

4

.25

25

.25

.25

0

0

0

0

Prescott

5

.20

.60

.20

0

0

0

0

0

Prince Edward

7

.14

.29

.57

0

0

0

0

0

Rentreu

11

.73

.18

.09

0

0

0

0

0

Russell

4

0

.25

.25

.50

0

0

0

0

Simcoe

8

0

.12

.63

.12

.12

0

0

0

Stormont

5

.40

.40

0

20

0

0

0

0

Sudbury

7

72

.14

.14

0

0

0

0

0

Thunder Bay

1

1.00

0

0

0

0

0

0

0

Victoria

7

.14

.14

.43

.29

0

0

0

0

vs.iierloo

10

.10

0

.60

.30

0

0

0

0

Wellington

6

0

.17

.33

.17

.33

0

0

0

Went worth

5

.20

0

.20

.20

0

0

.40

0

York

10

.30

10

.20

,10

.10

0

.10

.10

Total collections

384

Vwerage frequents

.22

.26

.26

.15

.06

.01

.02

.00

of obtaining the

number of species

per collection

Appendix: Provincial Description

It is the author's hope that this text will facilitate and stimulate further studies
of North American earthworms. Certainly we need to know more concerning
their detailed distribution if we are to assess fully their biological importance.
For Ontario readers, therefore, a provincial description now follows. This is in-
tended to form a basis upon which regional surveys can be made. It is to be
hoped that ultimately we will be able to correlate distribution with edaphic and
vegetational factors. The provincial description also will serve as a reference
point for foreign readers unfamiliar with the Canadian environment.

Southern Ontario lies between 42° and 46° N latitude and 75° and 83° W longi-
tude (Fig. 1. p. 2) in the St. Lawrence Drainage Basin and primarily in the Plains
of the lower Great Lakes and St. Lawrence Lowlands, a physiographic region
underlain bv relatively undisturbed Paleozoic sedimentary beds of limestone,
shale, and sandstone. The northern portion of southern Ontario (Algoma, Co-
chrane. Manitoulin, Nipissing, and Sudbury districts) is underlain by Precam-
brian formations in a physiographic region known as the Canadian Shield. All
of Ontario has been glaciated.

Since 1915, soil surveyors have been at work in Canada, and most of the set-
tled area has been mapped on a preliminary scale at least, except for the great
unoccupied areas of northern Canada. One should consult Ehrlich (1968) and
United States Department of Agriculture (1964, 1967 and 1968) for a detailed
explanation of the soil classification schemes used in the following paragraphs
and in other megadrile surveys.

Southern Ontario contains large expanses of three soil orders — Alfisols, Spo-
dosols, and Inceptisols. The Alfisols (Aquic and Ochreptic Hapludalfs), mineral
soils, also known as Grey Brown Podzolic soils, are located in two areas: 1) a
line through Simcoe. Victoria, Peterborough, Northumberland, and Prince Ed-
ward counties and all counties to the west, and 2) a line through Carleton and
Leeds counties and all counties to the east. The parent materials either are gla-
cial in origin or were deposited in the great bodies of water which occupied the
lowlands at the close of the glacial epoch. Under deciduous forests the leaching
is not intense. The surface soil (A horizon) is greyish (10YR 4/2-5/2) (Munsell,
1954; USDA, 1951) and slightly acid, and has a moderate amount of organic
matter. The B horizon is brown (10YR 5/3, 4/3, 3/2, 2/2) and nut-like in struc-
ture because of the accumulation of clay (argillic B horizon). Normally, all lime
is leached from the soil profile which may be slightly acid throughout. Generally
the soils of this suborder (Udalfs) are reasonably fertile and well suited to culti-
vation.

The Spodosols appear to be more weathered than the Alfisols. Spodosols are
mineral soils with a spodic epipedon and a subsurface horizon with an accumu-
lation of organic matter and aluminium oxides + iron oxides. These soils (Spo-
dosols: Boralfic Cryorthods, Typic Cryorthods, and Humic Haplorthods) are
also known as Bisequa and Orthic Podzolic soils and occupy the major region
not included above for the Alfisols. These soils form mostly on coarse-textured,
acid parent material subject to ready leaching occurring in humid climates com-

123

monly where it is cold and temperate. Forests are natural cover vegetation.
Conifers, low in metallic ion content, seem to encourage the development of
Spodosols. As the litter from these low-base species decomposes, a strong acid-
ity develops and percolating water leaches acids down into the profile. Since the
upper horizons are so intensly leached, Spodosols are not naturally fertile. If
properly fertilized, these soils can become quite productive.

The inceptisols are young mineral soils whose profiles contain horizons which
1) have formed quickly, 2) result mostly from alteration of parent materials, 3)
are free of extreme weathering, and 4) lack accumulation of clay, and iron and
aluminium oxides. These Degraded Brown Forest soils (Mollic Alfic Eutro-
chrepts) are located mainly on a line from Bruce to Glengarry counties. These
soils are generally in forests as agricultural productivity may be limited without
considerable expense.

These three soil groups make up nearly all of the soils of southern Ontario.
But there are other minor areas of note, such as the sands or Orthic Regosols
(Entisols: Udipsamments and Udorthents) in portions of Durham, Haliburton,
Norfolk, and Victoria counties. Also there are many areas comprised of Histo-
sols (Fibrists, Hemists, and Saprists), known previously as Bog and/or Half-bog
soils. These organic or bog soils are found covering sizeable areas of Essex,
Kent, Lambton, and Simcoe counties. In both cases, these groups (Entisols and
Histosols) have in recent decades been converted into areas of productive spe-
cialized cropping.

Of the 14 major regions of vegetation in Canada, southern Ontario contains
part of the Great Lakes-St. Lawrence Forest and all of the Niagara Forest
(Rowe, 1972).

The Great Lakes-St. Lawrence Forest extends from Lake-of-the- Woods to
Baie de Chaleur and is essentially a transition between the boreal coniferous
forest and the deciduous forest of eastern North America. The dominant coni-
fers are: white pine (Pinus strobus L.), red pine (P. resinosa Ait.), hemlock
(Tsuga canadensis (L.) Carr.), and white cedar (Thuja occidentalis L.); others, ap-
parently invaders from the north, are: jack pine (Pinus banksiana Lamb.), ta-
marack (Larix laricina (Du Roi) K. Koch), balsam fir (Abies balsamea (L.) Mill.)
and white spruce (Picea glauca (Moench) Voss). The dominant hardwoods are:
sugar maple (Acer saccharum Marsh.), beech (Fagus grandifolia Ehrh.), yellow
birch (Betula alleghaniensis Britt.), red oak (Quercus rubra L.), bur oak (Quercus
macrocarpa Michx.) and white oak (Quercus alba L.) on upland soils; with red
maple (Acer rubrum L.), silver maple (Acer saccharinum L.), elm (Vlmus) and ash
(Fraxinus) in the low ground. This forest region has probably more species and
a greater number of associates than any other in Canada. The vegetation also
includes many smaller plants, shrubs, and herbaceous forms on the forest floor
and cleared lands. Some of the more prominent are: ground ivy (Glecoma
hederacea L.), juniper (Juniperus virginiana L.), witch-hazel (Hamamelis
virginiana L.), sumach (Rhus typhina L.), poison ivy (Rhus radicans L.), service-
berry (Amelanchier oblongifolia T. & G.), wild grape (Vitis labrusca L.), hawt-
horn (Crataegus foetida Ashe), raspberry and blackberry (Rubus spp.), thimble-
berry (Rubus odoratus L.), and honeysuckle (Lonicera (atarica L.). Hawthorns
have taken over many thousands of hectares of pasture land, and large areas are
occupied by tangles of raspberries and brambles. There are many herbaceous

124

plants on the floor of the deciduous forest, such as may apple (Podophyllum
peltatum 1..). herb Robert {Geranium Robertianum L.), Jack-in-the-pulpit
(Arisaema triphyllum (L.) Schott), lily-of-the-valley (Maianthemum canadense

Desf.). trillium (Trillium grandiflorum (Michx.) Salisb.). and sarsaparilla (Alalia
nudicaulis L.). Manv species of aster (Aster spp.) and goldenrod (Solidago spp.),
almost unnoticed in the Forest, dominate large areas of unimproved, low pasture
land. Dry sites are covered bv mullein (Verbascum thapsus L.) and blueweed
(Echium vulgare L). The roadside flora is just as characteristic as that of the for-
est.

The Niagara Forest is a strip along the northern shore of Lake Erie (Essex,
Kent. Lambton. Elgin, Middlesex, Norfolk. Brant. Niagara, Oxford, and Went-
worth counties). Much of what was said concerning the composition and ap-
pearance of the vegetation of the larger Great Lakes-St. Lawrence Forest also
applies to the Niagara Forest. However, there are differences. Except for the
pines on the sand plains there are few conifers while, on the other hand, there
are additional species of deciduous trees such as: blue ash (Fraxinus
quadrangulata Michx.). tulip poplar (Liriodendron tulipifera L.), sassafras
(Sassafras albidum (Nutt.) Nees), magnolia (Magnolia acuminata L.), Kentucky
coffee tree (Gvmnocladus dioica (L.) K. Koch), sycamore (Plantanus occidentalis
L.). black walnut (Juglans nigra L.), pawpaw (Asimina triloba (L.) Dunal), and
others. This area is also the habitat of many small plants not found farther
north.

There are large areas of cultivated land in southern Ontario. Prior to Euro-
pean settlement there were no forages and few natural meadows. The natural
meadows were assigned to settlers on a livestock number basis. The lack of feed
made it necessary to seed and cultivate meadows for wintering cattle. Most of
the forages are of European origin. Recent values for areas of cultivated land in
Ontario are: seeded pasturages 1.35 million hectares, rough pasture >1.30 mil-
lion hectares, and small grains slightly >2 million hectares.

One of the most popular grasses in pasture mixtures is timothy (Phleum
pratense L.), which is used on 75-80% of Ontario farms. Other prominent pas-
ture species are: orchard grass (Dactylis glomerata L.), brome grass (Bromus in-
ermis Leyss.), tall fescue (Festuca elatior L.), reed canary grass (Phalaris
arundinacea L.). Kentucky bluegrass (Poa pratense L.), Canada bluegrass (Poa
compressa L.), red fescue (Festuca rubra L.), perennial ryegrass {Lolium perenne
L.), alfalfa (Medicago sativa L.), red clover (Trifolium pratense L.), white clover
(Trifolium repens L.). and birdsfoot trefoil (Lotus corniculatus L.). Various com-
binations of these species are found in Ontario depending on the soil type, an-
nual precipitation, physiographic position, and topography of the pasture, etc.
For example. Canada bluegrass is well adapted to drier areas while Kentucky
bluegrass can only be sown in moist areas.

The major small grains sown in Ontario are: oats (Avena sativa L.), barley
(Hordcum vulgare L.), wheat (Triticum spp.), rye (Secale cereale L.), sorghum
(Sorghum vulgare Pers.). and very limited areas of rice (Oryza sativa L.).

The climate of Ontario, according to the Koppen system (Pettersen, 1968), is
in the cool snow-forest climatic type (D/b) with warm summers and the absence
of a dry season. The Holdridge system (Sawyer and Lindsay, 1964) places the
province in the cool temperate moist forest bioclimatic formation. Utilizing the

125

Thornthwaite (1948) classification of climate for Ontario, the province is char-
acterized in the following manner:

1. Moisture Regions — Climatic Type (moisture deficiency surplus index): B,
Humid (20-40) — south and west of a line through Huron and Wentworth
counties, southern Kenora and Rainy River districts. B: Humid
(40-60) — the rest of southern Ontario.

2. Seasonal Variation of Effective Moisture: r (little or no water deficiency in
any season) — all of southern Ontario.

3. Average Annual Thermal Efficiency — Thermal Efficiency Type (TE index,
in inches): B1, or Mesothermal (22.44-28.05) — south and west of a line
through Huron and Wentworth counties. C2 or Microthermal
(16.83-22.44) — the rest of southern Ontario.

4. Summer Concentration of Thermal Efficiency — Thermal Efficiency Type
(%): b': (56.3-61.6) — south and west of a line through Huron and Went-
worth counties, b', (61.6-68.0) — the rest of southern Ontario.

The mean annual precipitation for southern Ontario ranges from 711 mm
(Essex and Niagara counties and Manitoulin District) increasing eastward to
916 mm (Prescott County). The mean January air temperature is -1 to -7° C
(maximum) and -12 and -18 °C (minimum) while the mean-July air tempera-
ture is 24 to 27 °C (maximum) and 10 to 16 °C (minimum). For further details,
see Brown et al. (1968) or Thomas (1953).

126

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1977 The first North American record of Aporrectodea icterica (Savigny. 1826) (Oligochaeta,
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SHIELDS. E. B.

1971 Raising earthworms for profit. 9th ed. -Elgin. Shields Publ.

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1973 Lumbricus terrestris Linnaeus, 1758 (Annelida. Oligochaeta): designation of a neotype in
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1970 Gromadzenie zapasow pokarmowych przez neiktore ssaki owadozerne (Insectivora).
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SMITH. F

1895 A preliminary account of two new Oligochaeta from Illinois. -Bull. 111. St. Lab. Nat.
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138

1915 I wo now varieties of earthworms with a ke\ to the described species in Illinois Bull.
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1917 North American earthworms of the famil) Lumbricidae in the collections of the United
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I92S An account of changes in the earthworm fauna of Illinois and a description of one new
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SMITH t and B. R GREEN

1916 The Porifera, Oligochaeta, and certain other groups of invertebrates in the vacinity of
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SMITH « w

1887 Notes on New Zealand earthworms. -Trans. N.Z. Inst. 19: 122-139.

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SOLTIIi RV R

1909 Contribution towards a monograph of the British and Irish Oligochaeta. -Proc. R. Irish
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1902 Notes on worms. -Zool. Anz. 25: 481-483.

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1962 Endemic-exotic earthworm competition in the American midwest. -Nature, Lond.
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STEPHENSON I

1917 On a collection of Oligochaeta from various parts of India and further India. -Rec. In-
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1953 L'ropoda agiians. a mite pest in commercial fishworm beds. -J. Econ. Ent. 46(4): 711.

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1969 A contribution to our knowledge of the systematics and zoogeography of norwegian
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1955 Earthworm population studies: a comparison of sampling methods. -Nature, Lond.

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1969 The earthworm genus Pheretima Kinberg, 1866 in Louisiana. -Ph.D. thesis, Louisiana
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1 M MR. p

1879 Annulata Danica. (1) En kritisk revision af de i Danmark fundene Annulata. Gephyrea,
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1953 Climatological atlas of Canada. -Natn. Res. Coun.. Div. Bldg. Res. 3151(41): 253 pp.

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1964 Soil Classification, a comprehensive system (7th approximation). -Washington, U.S.
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1970 Ecolog) of Soil Animals I ondon, McGraw-Hill. 283 pp.

w m roN * k

1933 The reaction of earthworms to alternating currents of electricity in the soil. -Proc. Ent.
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1955 Numbers and weights of earthworms under a highl) productive pasture. -N.Z.J. Sci.
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WHITEHOUSER H and \ J GROVI

1943 I he dissection ol the earthworm. -London. Univ. Tutorial Press. 72 pp.

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1967 Exotic specimens of earthworms in British Columbia eaten by Coho salmon. -J. Fish.
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1^42 Observations on several species of Metaradiophrya (Protozoa: Giliata). -J. Morph. 70:
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YAHNKE, W. and J A GEORGE

1972 Rearing and immature stages of the cluster fly (Pollenta rudis) (Diptera: Calliphondae)
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/MUM 1

1970 Syniizie dazdoviek (Lumbricidae) na liikach Karpatskej oblasti Ceskoslovenska. -Biol.
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zicsi \

1959 Faunistisch-systematische und okologische Studien Liber die Regenwurmer Ungarns. I.

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Zool. Budapest 4(2-4): 157-161.

141

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