Biota of Freshwater Ecosystems
Identification Manual No. 8
FRESHWATER LEECHES (ANNELIDA:HIRUDINEA) OF NORTH AMERICA
by
Donald J. Klemm
Museum of Zoology, Mollusk Division
The University of Michigan
Ann Arbor, Michigan 48104
for the
ENVIRONMENTAL PROTECTION AGENCY
Project # 18050 ELD
Contract # 14-12-894
May 1972
For sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, D.C. 20402 - Price $2.50
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JSYIRCKMLSTAL Pi.Gi'i'CrlGU AGEHGX
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FOREWORD
"Freshwater Leeches (Annelida:Hirudinea) of North America"
is the eighth of a series of identification manuals for se-
lected taxa of invertebrates occurring in freshwater systems.
These documents, prepared by the Oceanography and Limnology
Program, Smithsonian Institution for the Environmental Pro-
tection Agency will contribute toward improving the quality
of the data upon which environmental decisions are based.
Additional manuals will include, but not necessarily be lim-
ited to, freshwater representatives of the following groups:
amphipod crustaceans (Gammaridae), branchiuran crustaceans
(Argulus), isopod crustaceans (Asellidae), decapod crayfish
crustaceans (Astacidae), polychaete worms (Polychaeta),
freshwater planarians (Turbellaria), aquatic dryopoid beetles
(Dryopoidea), and freshwater clams (Sphaeriacea).
111
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ABSTRACT
An illustrated key to 63 species of North
American freshwater leeches (Annelida:Hirudinea)
is given with notes on their collection, preser-
vation, important diagnostic features and distri-
bution .
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CONTENTS
Section Page
I Introduction 1
Collection 3
Preservation 3
Identification 4
II Species List and Ranges 9
III Key to the Freshwater Leeches (Hirudinea) of North America 17
IV Acknowledgements 45
V References 47
VI Index to Scientific Names 51
VII
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FIGURES
Page
1 Anterior suckers 17
2 General body shape of Glossiphoniidae 18
3 General body shape of Piscicolidae 19
4 Arrangement of eyes 20
5 Arrangement of eyes 20
6 Arrangement of eyes 21
7 Posterior sucker of Aotinobdella ccnneotens 21
8 Anterior end of P. montifera 22
9 Posterior sucker of P. pediculata 23
10 Ventral view of P. parasitica 23
11 Dorsal view of H. stagnates 24
12 Posterior suckers of Piscicola spp. 26
13 Arrangement of copulatory gland pores 28
14 Dissection of mouth and buccal cavity 30
15 Annulation of D. (Mooreobdella)3 Nephelopsis and Erpobdella 31
16 Male genital atrium of N. obseura 31
17 Dissection of male genital atrium and ejaculatory duct 33
18 Dorsal view of Marvinmeyepia luoida, Oligobdella biannulata3
Aotinobdella anneotens3 A. triannulata* A. inequiannulata3
Plaoobdella montifera. 34
19 Dorsal view of Plaoobdella pedioulata3 P. hollensis3 P.
parasitica^ P. ornata, P. rmltil-ineata, P. pap-Lllifera. 35
20 Dorsal view of Batracobdella phalera, B. paludosa, B. picta,
Eelobdella stagnalis, H. elongata, H. punatatolineata. 36
21 Dorsal view of Eelobdella papillata3 H. lineata* H. fusoa3
Glossiphonia heteroalita, G. eomplanata, G. aomplanata
mollissima. 37
22 Dorsal view of Theromyzon maouloswn, T. rude, T. tessulatum^
Pisoioola punotata, P. salmositiea, P. m-Llneri. 38
23 Dorsal view of Pisoicola geometra, P. zebva, Cystobvandhus
vividusj C. vervilli, C. virginieus, Myzobdella moore-i. 39
24 Dorsal view of Illinobdella riahardsoni. I. elongatat I. alba3
Pisoioolavia reduotaf Macrobdella ditetra, M. decora. 40
25 Dorsal view of Maorobdella sestertia, Hirudo medioinalis3
Peraymoovensis kingi3 P. Iateralis3 Philobdella flori-dana3
P. gracilis. 41
26 Dorsal view of Bdellarogatis plumbea, Mollibdella gvandis.
Peroymoorensis Iateromaculata3 P. marmoratis3 Nephelopsis
obscura, Erpobdella punatata. 42
27 Dorsal view of Erpobdella punctata annulata3 E. triannulata3
Dina anoculata3 D. (Mooveobdella) fervida3 D. (M.) bucera3
D. (M.) miorostoma. 43
28 Dorsal view of Dina latevalis, D. parva3 D. dubia. 44
Vlll
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SECTION I
INTRODUCTION
Historically, leeches are derived from an ancient protostome stock which
gave rise to the annelids, thus establishing the metameric line. The ear-
liest fossil annelids of the middle Cambrian indicate that they were al-
ready differentiated into well-established groups (such as Polychaeta,
Oligochaeta, Hirudinea). The group (Annelida: Hirudinea) is relatively
homogeneous, and the highly specialized annelids are thought to have
arisen from an oligochaete progenitor.
Leeches are predominantly freshwater invertebrates, but there are many
marine forms, as well as numerous terrestrial species, occurring mainly
in the tropics. Unfortunately to the layman all leeches are popularly
considered to be "bloodsuckers". However, their food habits are far more
diverse than most people realize; many are not parasitic. These animals
are predaceous, ectoparasitic, or scavengers with anterior and posterior
suckers that serve as organs of attachment, feeding, and locomotion.
They are morphologically adapted for obtaining and digesting food consist-
ing chiefly of the blood of fishes, turtles, frogs, salamanders, birds
and mammals; they may also consume carrion, blood fluids, tissues, and
whole invertebrates, such as annelids (including leeches), insect larvae,
and mo Husks.
As a scientifically neglected group of invertebrates, they may, in the
near future be more important in ecosystems, in invertebrate pathology,
and as hosts to other parasitic forms, or even as Vectors in the life
cycles of parasites of other animals.
A number of keys to leeches have been published. Some of them are, as
follows: Meyer (1946a) for fish leeches (Piscicolidae) in North America;
Pennak (1953) for the United States; Moore, J. P. (1959) for North America;
Mann (1962) for Central Europe, the British Isles, and North America; Soos
(1963 to 1969b) for genera of the world with a catalogue of the species,
and Davies (1971) for Canada. Other geographically restricted keys have
been written: Moore, J.P. (1906, 1912, 1922) for the Great Lakes Region,
Minnesota, and southern Canada; Bere (1929) for Jasper Park Lakes
(Alberta, Canada); Miller (1929, 1937) for Ohio and Michigan; Eddy and
Hodson (1950) for the north central United States; Moore, J.E. (1964,
1966) for Alberta; and Keith (1960) for Minnesota. All of these keys,
however, have limited use because they are based on sparse collections;
they also are in need of some of the recent important taxonomic changes
made by Richardson (1969) and Soos (1963, 1965, 1966a, 1969a and b).
Since published leech accounts have been regrettably meager and 'widely
scattered in various journals throughout the world, an expanded compila-
tion has been prepared from published sources as well as from the late J.
P. Moore's unpublished records and Soos (1970) records of zoogeographic
distribution of the leeches of North America. It should be stressed
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that previous keys have a more restricted number of species and have
proven somewhat difficult to interpret and utilize. Therefore, the cri-
tical examination of leech collections and type specimens at the Academy
of Natural Sciences in Philadelphia, especially those in the U. S. Nation-
al Museum, Washington, D. C., and an extensive survey of the leeches of
Michigan (Klemm, 1972) have provided materials which aided considerably
in the preparation of this illustrated key. Notes have also been provid-
ed on important diagnostic characters, and they are intended to alleviate
some of the problems inherent in earlier keys. This key is to be used
primarily as a guide for determining the species of freshwater leeches of
North America, north of Mexico, and might not serve effectively outside of
that area.
By way of clarification of systematic difficulties Davies (1971) included
Erpobdella octoculata (Linnaeus, 1758) in his key on the basis of an iden-
tification made by J. P. Moore of Erpobdella atomaria (Cavena, 1820), a
new record for North America in Rawson (1953), but Moore, J. P. (1959)
omitted them from his key. Soos (1966a) considered E. atomaria a variety
of E. octoculata and recorded its distribution only in Europe and Japan.
Mann (1961) and Soos (1968) stated that E. octoculata was restricted only
to the Palaearctic Region.
There is still doubt concerning the correct generic rank for Vina fervida
(Verrill, 1874), D. microstoma (Moore, J. P., 1901), and D. bucera (Moore,
J. P., 1949). Moore, J. P. (1959), Mann (1961), and Meyer (1968) recog-
nized Mooreobdella (Pawlowski, 1955) as a valid genus containing these
three species. However, Soos (1966a and 1968), whose opinion is followed
here, regarded Mooreobdella as a subgenus of Dina. For the purposes of
this key E. atomaria and E. octoculata are ommitted and the name
Mooreobdella is retained only in subgeneric rank.
Soos (1969b) lumped Eelobdella fusca (Castle, 1900) and H. lineata
(Verrill, 1874) into one species, H. triseralis (E. Blanchard, 1849);
whereas, Moore J. P. (1952 and 1959) separated H. fusca and H. lineata
taxonomically into two species. Because there remains some doubt con-
cerning the correct species rank, H. fusca and H. lineata have been re-
tained and H. triseralis omitted.
Finally, for each species recorded in the Species List and Ranges section,
references are given to works containing more complete descriptions.
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COLLECTION
Leeches usually avoid light and are best collected by hand or gently with
forceps. They are usually found attached to the undersides of stones,
boards, logs, other objects littering the lake or stream bottom, submer-
gent aquatic vegetation or masses of organic debris. Blood-sucking ecto-
parasites are sometimes collected with their hosts, but most freshwater
species drop off after taking a meal. Some of the active swimming
Hirudidae and Erpobdellidae can also be collected at night with a dipnet.
PRESERVATION
In all species the identification is extremely difficult and often im-
possible because of unsuitable preservation. Therefore, if the leeches
are dropped alive into preservatives such as 70% alcohol, 4% formalde-
hyde, etc., they contract strongly and such features as the eyes and the
genital pores are difficult to discern. To get full advantage of this
key, live leeches or well-preserved specimens should be used. For best
results, live leeches should be narcotized in soda water (carbonated
water), shreds of cigarette tobacco to give the water a faint tint, or
in weak solutions of alcohol, chloroform, chlorotone, sodium nembutal,
or propylene phenoxetol added very gradually over a 30 to 60 minute
period. The leeches, depending on size, should be completely narcotized
and relaxed in 30 to 60 minutes. Excessive relaxation in chloroform,
chlorotone, etc., causes the furrows between the annuli to disappear.
When the leeches no longer respond to probing or pinching with a probe
or forceps, they are then rapidly drawn between the fingers and washed
so that the mucus secreted during the preceeding process is washed off.
Seventy-percent alcohol is injected into the body cavity of Erpobdellidae
to insure preservation of the reproductive organs which might be neces-
sary for positive identification. Next the leeches are straightened and
placed between two glass plates or back to back Petri dishes with only
slight pressure so they are held in normal position and not permanently
flattened. Thus the leeches are kept in and extended position without
displacing or distorting internal organs. This preparation is then im-
mersed in a fixative (2% formaldehyde) to fix the leeches in this posi-
tion. After the fluid has thoroughly penetrated and the leeches have
fully stiffened, they are removed, washed and placed in vials or jars
containing 80% alcohol.
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IDENTIFICATION
The diagnostic features which are very important in leech identification
are: size of mouth, eye number and position, relative size of suckers,
presence or absence of papillae, tubercles, and pulsatile vesicles, number
of annuli between the male and female gonopore, the male reproductive
system in some species, and annulation. Color differences occur within
some species. In the key color refers to living leeches and those colors
that are known to persist for several years with proper preservation. In
some specimens that have been preserved for a period of time, color
changes occur by the action of the preservative, and the deeper pigments
may be completely obscured by the opacity of the surface tissue. In some
cases it is important that the living color be noted on the label. In
the key, reference is made to internal anatomical features of some species.
Dissection is, therefore, necessary. If reference is made in a couplet to
an internal feature after reference to an external one, the key can be used
without resorting to the dissection provided the external features are
clearly seen. Internal dissections are sometimes necessary in the examin-
ation of the genital atrium and atrial cornua to distinguish the Dina-
Mooreobdella complex, Nephelopsis obscura, and in the examination of the
jaws and teeth for the identification of some Hirudidae.
The body of the leech consists of 34 somites or segments designated I
through XXXIV; each somite is represented by a ganglion in the central
nervous system. Characteristically, each neuromeric somite is divided ex-
ternally by superficial furrows in 2-16 rings or annuli. Somites that
have the full number of annuli (termed complete or perfect somites) are
found in the middle of the body and this number is generally characteris-
tic of the genus or species. Incomplete or abbreviated somites occur at
both ends of the body. The annuli features are best seen after careful
narcotization. The specimen should be blotted dry and examined in air
under oblique illumination, and the annuli features can be most easily
seen in the lateral margins of the ventral surface.
Moore (1898) recognized that the nerve cord ganglia are placed in the
middle annulus of the somite. The recognition of the triannulate somite
is basic, and also that more complex divisions may be derived by repeated
binary division of the annuli. Counting from the head end, those of the
three primary annuli are numbered al, a2, a3. Annulus a2 (the neural or
sensory annulus) contains the ganglion and is marked externally by trans-
verse rows of minute, cutaneous sensillae (segmental receptors). Repeated
bisection of the three primary annuli give more complex annulation, bl, to
b6; repeated subdivision give tertiary annuli cl to c!2, and quaternary
annuli dl to d24. The annular composition of complete somites is usually
characteristic of genera and the composition of incomplete somites is fre-
quently characteristic of species.
In addition to the sensillae, which are confined to the sensory annuli,
other visible surface organs are the eyes, papillae, and tubercles.
When present on the caudal sucker the simplest ocelli, which are imbedded
in pigment, are termed oculiform spots (Fig. 3a). They are usually pres-
ent in the Piscicolidae but are absent in the Glossiphoniidae. The number
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and position of the eyes are important taxonomic features. The Glossi-
phoniidae have one to four pairs of eyes, the Piscicolidae zero to two
pairs, Erpobdellidae have three or four pairs (except Dina anoculata
which has none), and the Hirudidae five pairs of eyes. In Batraaobdella
and sometimes in Glossiphonia coalescence of the eyes may occur (Fig. 6h-
j). The lobed nature of the eyes usually indicates the original condition.
The relative distance between eyes is another important diagnostic fea-
ture in identificiation (Fig. 6a-d). If the distance between them is
equal to or greater than the eyes, they are termed well separated; if
they touch they are termed fused, and if the distance between a pair of
eyes is less than the diameter of a single eye, they are termed close to-
gether. In the Erpobdellidae the eyes are arranged in separate labial
and buccal groups (Fig. 4b-d), and in the Hirudidae the eyes are arrang-
ed in a submarginal arch (Fig. 4a). If the leech has been fixed and the
eyes cannot be seen, the head can be flattened between two glass slides.
If the eyes are hidden by pigment, decolorize the head by immersion in
5% caustic potash (Mann, 1961). Papillae are limited to the small, minute
protrusible sense organs that are often scattered in small or great
numbers over the dorsal surface. Tubercles are the large, smooth, coni-
cal, or rounded cone projections that include some of the dermal tissues
and muscles, and they often themselves covered with papillae. The arrange-
ment of the papillae and tubercles is an important characteristic in dis-
tinguishing certain of the Helobdella and Plaaobdella. In H. punatatolin-
eata tubercles are absent; in H. lineata they are small, smooth, and
conical; in H. fusoa they are absent or limited to the middorsal line of
the posterior somites; and in H. papillata, the tubercles are prominent
in five to nine longitudinal rows. In P. omata numerous tubercles are
present; the surface of the tubercles is covered with papillae which re-
sults in a very rough or warty appearance. In P. pcccas-iti-oa the tubercles
are inconspicuous or absent; in P. multilineata they are small and more
uniform; and in P. papi-flifera they are small, white but conspicuous in
five longitudinal rows.
The anterior suckers of leeches are diagnostic and may be very prominent
or simply consisting of the expanded lips of the mouth. In the Glossi-
phoniidae the anterior sucker is small and only slightly distinct from
the body, but in the Piscicolidae the anterior sicker is always expanded
and usually distinctly marked off from the body. The posterior sucker
is generally directed ventrally. In all the Piscicolidae (except
Uyzobdella, Illinobdelta, and Pisciaolayia), the body is divided into two
distinct regions (Fig. 3a): a narrow anterior (trachelsome) region, and
a longer and wider posterior (urosome) region. In the genera Pisoicola
and CystobTanchus only, the neural annuli of the urosome bear 11 pairs of
pulsatile vesicles. In Pisoicola the vesicles are small and sometimes
invisible in preserved specimens but in CystobTanohus the pulsatile vesi-
cles are large and clearly visible in both living and preserved individu-
als.
The alimentary canal is a tube from mouth to anus and is divided into the
buccal chamber, pharynx, esophagus, stomach or crop, intestine, and
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rectum. In the Rhynchobdellida the mouth is a small pore within the an-
terior sucker, on the rim of it, or in the center of the sucker, (Fig.
la-c). The pharynx of the Rhynchobdellida is muscular and protrusible
through the mouth as a proboscis. In the Gnathobdellida and Pharyngob-
dellida the mouth is medium to large (Fig. Id-f) and occupies the entire
cavity of the anterior sucker. In the Hirudidae, the buccal cavity, which
may or may not contain j aws, is separated from the cavity of the sucker by
a flap of skin called the velum (Fig. 14a,b). The presence or absence of
papillae on the velum in the key can also be used as a diagnostic fea-
ture between Bdellarogat-Ls plimbea (papillate) and Mollibdella grand-is
(smooth). All other Hirudidae have three muscular jaws (two ventrolat-
eral and one dorsomedial). The free edge of each bears teeth arranged
in either one (monstichodont) or in two (distichodont) rows. To examine
the velum and jaws the specimen should be pinned out; a median incision
should be made from the lower lip of the anterior sucker back far enough
for the margins to be pinned out to expose the inner surface of the
pharynx. Details of the teeth can only be seen by removal of a jaw and
making a temporary or permanent mount on a microscope slide.
The anus of most species opens on the dorsal surface a few annuli anter-
ior to the posterior sucker (except in Plaaobdella pediaulata and
Illinobdella viehcacdsoni'). In P. pediaulata it opens between somites
XIII and XXIV with the posterior annuli forming a slender stalk (pedun-
cle) bearing the posterior sucker and in I. ridhardsoni the anus opens
15 annuli anterior to the posterior sucker.
Leeches are hermaphroditic. The male and female gonopores are visible
on the middle of the ventral surface of somites XI and XII and are gen-
erally separated by two to five annuli. The male gonopore is large,
more readily visible, and anterior to the female gonopore. The female
pore is often difficult to see. It is seen most easily immediately
after narcotization; its position often being revealed by some color
difference which is sometimes lost during fixation.
The female reproductive system comprises a pair of ovisacs, terminating
in ducts which join to form a common duct or vagina. The male reproduc-
tive system consists of testisacs in metameric patterns (five to six in
Rhynchobdellida and nine to ten in Hirudidae) but in the Erpobdellidae
they are small, numerous, and arranged in grape-like clusters. The vasa
efferentia connect the testisacs to the vasa deferens on each side.
These ducts lead into the seminal vesicle and ejaculatory duct which
opens into the atrium. The atrium is a median chamber and consists of
three parts: a thin-walled eversible bursa, a thickened-walled glandu-
lar chamber, and a muscular median one, as well as a pair of lateral
horns (atrial cornua), which receive the ejaculatory duct. In
Nephelopsis obscura the atrial cornua is spirally arranged like a ram's
horn (Fig. 16), but in the Dina-Mooreobdella complex the cornua are
simply curved, globular, rounded, ellipsoidal, or short and curved,
(Fig. 17a-e). Some species also have ejaculatory ducts with or without
long preatrial loops (Fig. 17a-e).
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To confirm diagnostic features of the internal reproductive system,
dissection is necessary. The best way to accomplish this is to pin the
preserved specimen out with the ventral surface up. A transverse in-
cision should then be made across the body four or five annuli posterior
to the male gonopore. Cuts should then be made anteriorly up the later-
al margins of the body for about 26 annuli. The posterior edge of the
flap thus made can now be lifted forward to expose the inner tissue
which can be cleared away to fully expose the atrium and preatrial loops.
External copulatory glands with pores are present in a linear pattern of
either four, eight or 28 (Fig. 13 a-cl on adult Maarobdella deaoir>a3
M. ditetra.1 and M. sestertla. These copulatory glands are located five
annuli posterior to the female gonopore.
Glossiphonia heteroalita and Marvinmeyeria lucida are two unusual species
in that the male and female ducts open into a single gonopore.
In the Pharyngobdellida, the genera are separated on the pattern of an-
nulation. While Erpobdella has all its annuli of similar size, Dina-
Mooreobdella complex and N. obscura have every b6 annulus larger than
the others and subdivided or partially divided by a faint transverse
furrow (Fig. 15a, b).
Names in brackets [=] are synonyms used in older taxonomic keys. The
average size of each species is given in the key but many variations
appear in the species.
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SECTION II
SPECIES LIST AND RANGES
Phylum Annelida
Class Hirudinea Lamarck, 1818
Order Rhynchobdellida Blanchard, 1887
Family Glossiphoniidae
Genus: Aotinobdella Moore, 1901
Aotinobdella anneotens Moore, 1906 (Refs 1,20,29)
Continental Distribution: North America
North American Distribution: Canada (Ontario)
Aotinobdella inequiannulata Moore, 1901 (Refs 1,20,32)
Continental Distribution: North America
North American Distribution: USA (Illinois, Michigan, Minnesota, Ohio)
Aotinobdella tviannulata Moore, 1924 (Refs 1,20,28)
Continental Distribution: North America
North American Distribution: Canada (Ontario), USA (Michigan, New
Hampshire, Virginia)
Genus: Batvaoobdella Viguier, 1879
Batracobdella miohiganensis Sawyer, 1972 (Ref. 46)
Continental Distribution: North America
North American Distribution: USA (Michigan)
Batraoobdella paludosa (Carena, 1824) (Refs 13,51)
Continental Distribution: North America, Palearctic Region
North American Distribution: Canada (Nova Scotia)
Batraoobdella phalera Graf, 1899 (Refs 1,7,58,59)
[= Plaeobdella phalera (Graf)]
Continental Distribution: North America
North American Distribution: Canada (Manitoba, Nova Scotia, Ontario,
Quebec), USA (Colorado, Connecticut, Florida, Iowa, Michigan, New
York, Ohio, Wisconsin)
Batracobdella piota Verrill, 1872 (Refs 1,2,29,51,54,58,59)
[= Plaoobdella piota (Verrill)]
Continental Distribution: North America
North American Distribution: Canada (Ontario, Quebec), USA (Arkansas,
Colorado, Connecticut, Louisiana, Michigan, Missouri, New York, Ohio,
South Dakota, Utah, Virginia, Wisconsin)
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Genus: Glossiphonia Johnson, 1816
Glossiphonia complanata (Linnaeus, 1758) (Refs 2,8,30)
Continental Distribution: North America, Europe, Asia (Kashmir, China,
India, Japan), Africa (Belgian Congo), South America (Argentina ?)
North American Distribution: Canada (generally distributed), USA
(Alaska, California, Colorado, Connecticut, District of Columbia,
Florida, Idaho, Indiana, Illinois, Iowa, Massachusetts, Michigan,
Minnesota, Missouri, Mississippi, New York, North Dakota, Ohio,
Oregon, South Dakota, Utah, Washington, Wisconsin, Wyoming)
Glossiphonia aomplanata mollissima Moore, 1898 (Refs 26,40)
Continental Distribution: North America
North American Distribution: USA (Alaska, Bering and Kodiak Islands)
Glossiphonia heteroclita (Linnaeus, 1761) (Refs 2,8)
Continental Distribution: North America, Europe, Asia (India), Africa
(Ethiopia, Belgian Congo)
North American Distribution: Canada (British Columbia, Newfoundland,
Quebec, Ontario, Manitoba, Alberta, Saskatchewan), USA (Connecticut,
Indiana, Iowa, Michigan, Mississippi, Ohio, Pennsylvania, Wisconsin)
Genus: Helobdella Blanchard, 1896
Helobdella elongata (Castle, 1900) (Refs 4,7)
[= Glossiphonia nepheloidea Graf, 1899 (nomen nudum)]
Continental Distribution: North America
North American Distribution: Canada (Ontario, Quebec). USA (Florida,
Georgia, Indiana, Iowa, Louisiana, Massachusetts, Michigan, Minnesota,
Mississippi, Missouri, North Dakota, Ohio, Wisconsin)
Helobdella fusoa Castle, 1900 (Refs 4,30)
Continental Distribution: North America
North American Distribution: Canada (Alberta, Manitoba, Northwest
Territories, Ontario, Quebec, Saskatchewan), Mexico (Federal District,
Yucatan), USA (Alaska, California, Colorado, Connecticut, Florida,
Indiana, Illinois, Iowa, Michigan, Minnesota, Mississippi, Missouri,
Nebraska, Ohio, Oklahoma, Pennsylvania, Vermont, Wisconsin)
Helobdella lineata (Verrill, 1874) (Refs 37,57)
[= papillifera var. d Verrill, 1874]
Continental Distribution: North America
North American Distribution: Canada (Ontario), USA (Florida, Michigan,
Nebraska, Ohio, Wisconsin)
Helobdella punetatolineata Moore, 1939 (Ref. 35)
Continental Distribution: North America
North American Distribution: USA (Puerto Rico, Louisiana, Wisconsin)
Helobdella papillata (Moore, 1952) (Refs 20,37)
[= papillifera var. b Verrill, 1872]
Continental Distribution: North America
North American Distribution: Canada (Ontario), USA (Connecticut,
Michigan, Ohio, Wisconsin)
10
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Eelobdella stagnalis (Linnaeus, 1758) (Re£s 2,30)
Continental Distribution: Cosmopolitan
North American Distribution: Canada (generally distributed), Mexico
(Guanajuato), Costa Rico, USA (Alaska, Arizona, California, Colorado,
Connecticut, Florida, Illinois, Indiana, Iowa, Massachusetts,
Michigan, Minnesota, Mississippi, Maine, Montana, Missouri, Nebraska,
New York, North Dakota, Ohio, Oregon, Pennsylvania, Utah, Vermont,
Wisconsin, Wyoming, New Jersey)
Helobdella transversa Sawyer, 1972 (Ref. 46)
Continental Distribution: North America
North American Distribution: USA (Michigan)
Genus: Mawinmeyeria Soos, 1969
Marvinmeyepia luoida Moore, 1954 (Refs 21,54)
[= Oaulobdella luoida (Moore)]
Continental Distribution: North America
North American Distribution: Canada (Alberta, British Columbia,
Manitoba), USA (Michigan)
Genus:. Oligobdella Moore, 1918
Oligobdella biannulata (Moore, 1900) (Ref. 27)
Continental Distribution: North America
North American Distribution: USA (North Carolina, South Carolina)
Genus: Plaaobdella Blanchard, 1893 (emend. Autrum, 1936)
Plaoobdella hollensis (Whitman, 1892) (Refs 2,30)
Continental Distribution: North America
North American Distribution: Canada (Manitoba, Ontario), USA (Florida,
Iowa, Massachusetts, Michigan, Minnesota, Wisconsin)
Plaaobdella montifera Moore, 1906 (Refs 1,29,30)
Continental Distribution: North America
North American Distribution: Canada (British Columbia, Ontario,
Saskatchewan), USA (Connecticut, District of Columbia, Indiana,
Illinois, Iowa, Kansas, Michigan, Minnesota, Missouri, New York, North
Carolina, Ohio, Tennessee, Washington, Wisconsin)
Plaaobdella rmltil-ineata Moore, 1953 (Ref. 38)
Continental Distribution: North America
North American Distribution: USA (Florida, District of Columbia,
Illinois, Iowa, Michigan, North Carolina, Oklahoma, Texas, Utah)
Plaaobdella ornata (Verrill, 1872) (Refs 30,57)
[= P. Tugosa Verrill, 1872)]
Continental Distribution: North America
North American Distribution: Canada (Alberta, British Columbia,
Manitoba, Nova Scotia, Ontario, Quebec, Saskatchewan), Mexico
(scattered questionable reports), USA (California, Colorado,
Connecticut, District of Columbia, Florida, Idaho, Illinois, Indiana,
Iowa, Louisiana, Maryland, Massachusetts, Michigan, Minnesota,
Missouri, Nebraska, New Jersey, New Mexico, New York, North Carolina,
North Dakota, Ohio, Oklahoma, Oregon, Pennsylvania, Virginia,
Washington, Wisconsin)
11
-------
Plaoobdella papillifera (Verrill, 1872) (Re£s 21,37,56)
Continental Distribution: North America
North American Distribution: Canada (Alberta, Manitoba, Northwest
Territories), USA (Michigan)
Plaoobdella parasitica (Say, 1824) (Refs 2,30)
Continental Distribution: North America
North American Distribution: Canada (Alberta, Manitoba, Nova Scotia,
Saskatchewan, Ontario), Mexico (scattered reports), USA (Arizona,
Colorado, Connecticut, Delaware, District of Columbia, Florida,
Illinois, Georgia, Indiana, Iowa, Louisiana, Maine, Maryland,
Massachusetts, Michigan, Minnesota, Mississippi, Missouri, Nevada,
New Jersey, New York, North Dakota, Ohio, Pennsylvania, South
Dakota, Tennessee, Virginia, West Virginia, Wisconsin)
Plaoobdella pedioulata Hemingway, 1908 (Refs 1,9,30)
Continental Distribution: North America
North American Distribution: USA (Illinois, Iowa, Maine, Minnesota,
Missouri, Oklahoma, Wisconsin)
Genus: Theromyzon Philippi, 1867
Thevomyzon maoulosim (Rathke, 1862) (Refs 2,24,54)
[= T. meyevi (Livanow, 1902]
Continental Distribution: Holarctic Region, North America
North American Distribution: Canada (Alberta), USA (California, Iowa,
Michigan, Minnesota, Montana, North Dakota, Oregon, Pennsylvania)
Theromyzon rude (Baird, 1869) (Refs 2,40,54)
[= T. oooidentalis Verrill, 1874]
Continental Distribution: North America
North American Distribution: Canada (Alberta, British Columbia,
Northwest Territories, Saskatchewan), USA (Alaska, Iowa, Colorado,
Michigan, Montana, Oregon, South Dakota)
Theromyzon tessulatum (0. F. Muller, 1774) (Refs 2,30)
Continental Distribution: Holarctic, Neotropic, North America, Europe,
Asia, South America
North American Distribution: Canada (Nova Scotia, Saskatchewan), USA
(Alaska, Colorado)
Family Piscicolidae Johnston, 1865
Genus: Cystobvanohus Diesing, 1859
Cystobranahus verri-lli Meyer, 1940 (Refs 2,17)
Continental Distribution: North America
North American Distribution: Canada (Ontario, Saskatchewan), USA
(Arkansas, Illinois, Iowa, West Virginia)
CystobTanohus vividus Verrill, 1872 (Ref. 26)
Continental Distribution: North America
North American Distribution: USA (Massachusetts, North Carolina,
Virginia)
Cystobranahus virginieus Hoffman, 1964 (Ref. 10)
Continental Distribution: North America
North American Distribution: USA (Virginia)
12
-------
Genus: Il'iinobdella Meyer, 1940
Illindbdella alba Meyer, 1940 (Refs 17,18)
Continental Distribution: North America
North American Distribution: Canada (Ontario), USA (Connecticut,
Illinois, Michigan, Minnesota, New York, Tennessee)
Illinobdella elongata Meyer, 1940 (Refs 17,18)
Continental Distribution: North America
North American Distribution: Canada (Ontario), USA (Illinois, Michigan,
Minnesota)
Illindbdella riohardsoni Meyer, 1940 (Refs 17,18)
Continental Distribution: North America
North American Distribution: USA (Illinois, California, Kansas,
Michigan, Minnesota, Ohio, New York, Tennessee)
Genus: Myzobdella Leidy, 1851
Myzobdella moovei (Meyer, 1940) (Refs 17,43)
[= Illinobdella moorei (Meyer)]
Continental Distribution: North America
North American Distribution: Canada (Alberta, Ontario, Saskatchewan),
USA (Alabama, Arkansas, California, Colorado, Connecticut, Florida,
Illinois, Iowa, Louisiana, Michigan, Minnesota, New Jersey, New York,
Ohio, Oklahoma, Tennessee, Texas)
Genus: Pisoioola De Blainville, 1818
Pisoioola geometra (Linnaeus, 1758) (Refs 2,8)
Continental Distribution: Holarctic, Europe, North America, South
America
North American Distribution: Canada (Saskatchewan), USA (District of
Columbia, Michigan, Minnesota, Wisconsin)
Piscicola milneri (Verrill, 1874) (Refs 2,17)
Continental Distribution: North America
North American Distribution: Canada (Alberta, Ontario, British
Columbia, Quebec, Saskatchewan), USA (Alaska, California, Maine,
Michigan, Minnesota, New York, Wisconsin)
Pisoioola punctata (Verrill, 1871) (Refs 2,17)
Continental Distribution: North America
North American Distribution: Canada (British Columbia, Ontario, Nova
Scotia, Quebec, Saskatchewan), USA (District of Columbia, Illinois,
Louisiana, Michigan, Minnesota, Mississippi, Missouri, Montana, New
Jersey, New York, North Carolina, Virginia, Wisconsin)
Pisoioola salmositica Meyer, 1946 (Ref. 19)
Continental Distribution: North America
North American Distribution: Canada (British Columbia), USA
(Washington)
Pisoioola zebra Moore 1898 (Refs 17,26)
Continental Distribution: North America
North American Distribution: Canada (Nova Scotia)
13
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Genus: Piseieolaria Whitman, 1889
Piscicolaria reducta Meyer, 1940 (Refs 17,18)
Continental Distribution: North America
North American Distribution: Canada (Ontario), USA (Connecticut,
Kansas, Illinois, Maine, Michigan, New Jersey)
Order Gnathobdellida Vaillant, 1890
Family Hirudinidae [=Hirudidae]
Genus: Bdellarogatis Richardson, 1969
Bdellarogatis plumbea (Moore, 1912) (Refs 15,30,45)
[= Haemopis plumbea (Moore)]
Continental Distribution: North America
North American Distribution: USA (Iowa, Michigan, Minnesota, Ohio)
Genus: Mollibdella Richardson, 1969
Mollibdella grandis (Verrill, 1874) (Refs 15,30,45,57)
[= Haemopis grandis (Verrill)]
Continental Distribution: North America
North American Distribution: Canada (Alberta, Ontario, Manitoba, New
Brunswick, Newfoundland, Prince Edward Island, Quebec, Saskatchewan),
USA (Connecticut, Maine, Michigan, Minnesota, New York, Ohio, Virginia,
Wisconsin)
Genus: Pereymoorensis Richardson, 1969
Pereymoorensis kingi (Mathers, 1954) (Refs 14,45)
[= Haemopis kingi (Mathers)]
Continental Distribution: North America
North American Distribution: USA (Iowa, Colorado)
Percymoorensis lateralis (Say, 1824) (Refs 2,15,30,45,46)
[= Haemopis latevdlis (Say)] [= H. terrestris Forbes, 1890]
Continental Distribution: North America
North American Distribution: USA (Florida, Illinois, Iowa, Colorado,
Kansas, Maine, Michigan, Minnesota, Missouri, North Carolina, Tenne-
see)
Peroymoovensis lateromaculaba (Mathers, 1963) (Refs 16,45)
[= Haemopis lateromaaulata (Mathers)]
Continental Distribution: North America
North American Distribution: USA (Iowa, Minnesota)
Percymoovensis marmoratis (Say, 1824) (Refs 2,15,30,45)
[= Haemopis marmoratis (Say); Moore, 1901]
Continental Distribution: North America
North American Distribution: Canada (generally distributed), USA (Alas-
ka, Colorado, Connecticut, District of Columbia, Illinois, Iowa, Indiana,
Kansas, Maryland, Massachusetts, Michigan, Minnesota, Missouri, Montana,
New Jersey, New Mexico, New York, Ohio, South Carolina, Utah, Virginia,
Wisconsin)
14
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Genus: Hirudo Linnaeus, 1758
Hirudo medioinalis Linnaeus 1758 (Ref. 13)
Continental Distribution: Europe, West Asia, North America (introduced)
North American Distribution: USA (New Jersey?, Pennsylvania?)
Genus: Maorobdella Verrill, 1872
Maorobdella deoora (Say, 1824) (Refs 2,30,45,53)
Continental Distribution: North America
North American Distribution: Canada (Nova Scotia, Alberta, Manitoba,
New Brunswick, Ontario, Prince Edward Island, Quebec, Saskatchewan),
Mexico (Nuevo Leon), USA (Colorado, Connecticut, District, of Columbia,
Illinois, Iowa, Kansas, Massachusetts, Maryland, Michigan, Minnesota,
Maine, New Hampshire, New Jersey, New Mexico, New York, North Carolina,
North Dakota, Ohio, Pennsylvania, Vermont, Virginia, Wisconsin)
MaoTobdella ditetra Moore, 1953 (Refs 34,38,53)
Continental Distribution: North America
North American Distribution: USA (Alabama, District of Columbia,
Florida, Louisiana, Mississippi, Missouri, North Carolina, South
Carolina, Texas)
Maorobdella sestertia Whitman, 1886 (Refs 2,53,60)
Continental Distribution: North America
North American Distribution: USA (Massachusetts, Louisiana)
Genus: Philobdella Verrill, 1874
Philobdella floridana (Verrill, 1874) (Refs 37,56)
Continental Distribution: North America
North American Distribution: USA (Florida, Louisiana, North Carolina)
Philobdella gvaailis Moore, 1901 (Refs 2,28)
Continental Distribution: North America
North American Distribution: USA (Illinois, Louisiana, Michigan,
Missouri, Texas)
Order Pharyngobdellida Johnson, 1913
Family Erpobdellidae
Genus: Dina R. Blanchard, 1892
Dina anoculata Moore, 1898 (Refs 26,47)
Continental Distribution: North America
North American Distribution: Canada (British Columbia), USA (Califor-
nia, Oregon)
Dina dubia Moore § Meyer, 1951 (Refs 2,40,47)
Continental Distribution: North America
North American Distribution: Canada (Alberta, Northwest Territories,
Saskatchewan), USA (Alaska, Colorado, Iowa, Michigan, Utah)
15
-------
Dina lateralis (Verrill, 1871) (Refs 2,37,49)
Continental Distribution: North America
North American Distribution: Canada (British Columbia, Ontario,
Manitoba, Quebec, Saskatchewan), USA (Colorado, Connecticut, Florida,
Maine, Michigan, Minnesota, Wisconsin)
Dina parva Moore, 1912 (Refs 30,47)
Continental Distribution: North America
North American Distribution: Canada (Alberta, Ontario, Quebec,
Saskatchewan), USA (Colorado, California, Indiana, Iowa, Michigan,
Minnesota, New York, Virginia, Wisconsin, Wyoming)
Dina (Mooreobdella) buoera Moore, 1949 (Refs 36,38,41,56)
Continental Distribution: North America
North American Distribution: USA (Michigan)
Dina (Mooreobdella) fewida (Verrill, 1871) (Refs 41,47,56)
Continental Distribution: North America
North American Distribution: Canada (Alberta, Ontario, New Brunswick,
Nova Scotia, Prince Edward Island, Quebec, Saskatchewan), USA (Colo-
rado, Illinois, Indiana, Iowa, Michigan, Minnesota, Missouri, North
Dakota, Ohio, Pennsylvania)
Dina (Mooreobdella) microstoma Moore, 1901 (Refs 2,41,47)
Continental Distribution: North America
North American Distribution: Canada (Ontario), Mexico, USA (Califor-
nia, Colorado, Illinois, Indiana, Iowa, Louisiana, Michigan, Missouri,
New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Texas,
Wisconsin)
Genus: Erpobdella De Blainville, 1818
Erpobdella punotata (Leidy, 1870) (Refs 2,30,52)
Continental Distribution: North America
North American Distribution: Canada (generally distributed), Mexico
(Federal District, Jalisco, Hidalgo, Yucatan), USA (Alaska, Arizona,
California, Colorado, Connecticut, District of Columbia, Georgia,
Idaho, Illinois, Indiana, Iowa, Louisiana, Maine, Massachusetts,
Michigan, Minnesota, Missouri, Montana, Nebraska, New Hampshire, New
Jersey, New York, North Dakota, Ohio, Oregon, Pennsylvania, Utah,
Vermont, Virginia, Washington, Wisconsin, Wyoming)
Erpobdella punotata annulata Moore, 1922 (Refs 31,52)
Continental Distribution: North America
North American Distribution: Canada (Ontario), USA (Michigan, Oregon,
Washington)
Eyypobdella tviannulata Moore, 1908 (Refs 33,52)
Continental Distribution: North America
North American Distribution: Central America (Guatemala, Yucatan),
USA (California, North Dakota)
Genus: Nephelopsis Verrill, 1872
Nephelopsis obsoura Verrill, 1872 (Refs 2,30)
Continental Distribution: North America
North American Distribution: Canada (generally distributed), USA (Alas-
ka, Colorado, Idaho, Iowa, Michigan, Minnesota, Mississippi, Montana,
New York, North Dakota, Utah, Washington, Wisconsin, Wyoming)
16
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SECTION III
KEY TO THE FRESHWATER LEECHES (HIRUDINEA) OF NORTH AMERICA
Mouth in form of small pore on anterior sucker from which muscu-
lar pharyngeal proboscis may be protruded (Fig. la-c); no
jaws or denticles. Order Rhynchobdellida
Mouth medium to large, occupying entire anterior sucker
forming rounded lips (Fig. Id-f); no protrusible proboscis;
teeth either present or absent
e f
Fig. 1. Ventral views of the anterior suckers to show
mouth differences: a- pore on rim of sucker; b- pore within
sucker; c- pore near center of sucker; d-f- mouth occupying
entire sucker.
17
-------
2(1) Body flattened, much wider than head (Fig. 2), never cylindri-
cal (except Relobdella. elongata which is subcylindrical),
not divided into distinct anterior (trachelsome) and post-
erior (urosome) regions; anterior sucker not freely expanded
but more or less fused to head; somites 3-annulate; eggs
and young attached to ventral surface of adults during their
development. Family Glossiphoniidae 4
Body at rest cylindrical, may or may not be divided into dis-
tinct narrow (trachelsome) and wider posterior (urosome)
regions (especially in contraction) (Fig. 3a-e); anterior
sucker expanded, distinctly separate from body (Fig. 3a-e
and Fig. 3f, g); 7 or more annuli per somite (except
P-iso-ioolapla. veduata which is 3-annulate); oculiform ocelli
or spots sometimes present on posterior sucker (Fig. 3a);
eyes 0, 1, or 2 pairs (Fig. 3f, g); pulsatile vesicles pre-
sent along lateral margins of urosome in Pisaicola sp. and
Cystobranehus sp. (Fig. 3a-c); eggs and young never attached
to ventral surface of adults during their development.
Family Piscicolidae 30
Fig. 2. a-d- General body shape of Glossiphoniidae.
3(1) Eyes 5 pairs, arranged in arch on somites II to IV (Fig. 4a);
jaws either present or absent; pharynx short, not extending
to clitellum; testes large, arranged in metameric pairs.
Order Gnathobdellida. Family Hirudinidae 42
Eyes 3 or 4 pairs in separate labial and buccal groups (Fig. 4b-
d) or absent (Dina anoculatd); no jaws; pharynx extending to
XIII, about 1/3 of body length; testes small and numerous in
grape-bunch arrangement. Order Pharyngobdellida. Family
Erpobdellidae 53
18
-------
trachelosome
pulsatile
vesicles
urosome
oculiform
spot
ray
eyes
Fig. 3. a-g- General body shape of Piscicolidae in dorsal view.
19
-------
t
Fig. 4. Dorsal views of arrangement of eyes: a- Hirudinidae;
b-d- Erpobdellidae.
4(2) 3 or 4 pairs of eyes (Fig. 5a-c) 10
1 or 2 pairs of eyes (except Plaoobdella hollensis which also
has accessory eyes) (Fig. 6a-j) 5
Fig. 5. Dorsal views showing positions of eyes: a- Theromyzon
sp.; b- Glossiphonia oomplanata; c- G. heteroolita.
5(4) Mouth within anterior sucker cavity, clearly not on rim (Fig.
lb)
Mouth apical or subapical on rim of anterior sucker (Fig. la)
9
6
6(5) Posterior sucker conspicuous with marginal circle of about 30
or 60 glands and retractile papillae, their positions being
indicated dorsally by faint radiating ridges (Fig. 7).
Genus Aat-inobdella 11
Posterior sucker without marginal circle of glands and retrac-
tile papillae 7
20
-------
a
A
accessory
eyes
Fig. 6. Dorsal views of eyes: a- well separated; b- fused; c,d-
close together; e- fused with accessory eyes; f-j- eyes showing
variation (coalescence) in the eye position.
7(6) 1 pair of eyes well separated; male
and female gonopores united in
common pore; body smooth, not tu-
berculate, dark streaks between
eyes in living and well preserved
specimens; dorsum dusky due to
nearly uniform distribution of
minute blackish chromatophores ,
dusky diffusion becoming banded
towards margins; length 15-22 mm
(Fig. 18a) : Marvinmeyeria luoida
1 pair of eyes fused (Fig. 6b) (ex-
cept P. hollensis which also has
several pairs of minute accessory
eyes (Fig. 6e) or close together
as in P. monttfera which has
eyes almost separated by their
diameter (Fig. 6c)); dorsum papil-
late, tuberculate or smooth ..... 8
marginal
papulae
dorsal
ridges
Fig. 7. Dorsal view of
posterior sucker of
Actinobdella anneetens
showing the marginal pa-
pillae and dorsal ridges,
21
-------
8(7) 1 pair of eyes confluent (fused) or close together; body
usually papillate and/or tuberculate. Genus Placobdella .. 13
Dorsum smooth, without conspicuous tubercles; translucent,
speckled with scattered green and brown pigment cells which
give pale olive-green color; large posterior sucker; 1 pair
of fused eyes; length 6.3-? mm (Fig. 18b):
Oligobdella biannulata
9(5) 1 or 2 pairs of eyes (Fig. 6b or 6f,g); if only 1 pair of eyes
present, these may be close together or coalesced (Fig. 6h-
j); gonopores separated by 2 annuli. Genus Batraoobdella .. 19
[see Sawyer (1972) for a new species not in this key]
1 pair of eyes, well separated (Fig. 6a); gonopores separated
by 1 annulus. Genus Helobdella 21
[see Sawyer (1972) for a new species not in this key]
10(4) 3 pairs of eyes in 2 paramedian rows or in roughly triangular
pattern (Fig. 5b,c); body firm. Genus Glossiphonia 26
4 pairs of eyes in 2 paramedian lines on somites II-V (Fig.
5a); body after egg laying translucent and soft. Genus
Thevomyzon 28
11(6) Posterior sucker with about 60 dorsal ridges (Fig. 7); dorsal
tubercles in 5 longitudinal rows; 6 equal annuli per segment;
length 7-11 mm (Fig. 18c): Actinobdella anneotens
Posterior sucker with about 30 dorsal ridges 12
12(11) Dorsal tubercles prominent in 5 longitudinal rows; posterior
sucker with 30 dorsal ridges; 3 annuli per somite; length
7-22 mm (Fig. 18d): Actinobdella triannulata
Tubercles confined to mid-dorsal row; about 30 dorsal ridges
on posterior sucker; 6 unequal annuli per somite; length
9-21 mm (Fig. 18e) : Aotinobdella inequ-Lannulata
13(8) Anterior somites distinctly widened to form discoid head
(Fig. 8); 3 dorsal, prominent, tuberculate keels or ridges;
color dull greenish-grey or pale olive-brown; length 9-16
mm (Fig. 18f) : Placobdella. montifeva
Without distinct, discoid head 14
Fig. 8- Dorsal view of anter-
ior end of P. montifera,
showing the discoid head.
22
-------
14(13) Anus separated from posterior sucker by slender stalk (pe-
duncle) (Fig. 9); body surface smooth; high dorsally and
contractile; 1 pair of eyes close together, hard to detect
in adults (small pigment masses at II-IV); color brownish;
length 20-35 mm (Fig. 19a): Plaodbdella pediculata
Anus close to posterior sucker, no slender stalk (peduncle).. 15
Fig. 9. Posterior sucker
and its slender stalk (pe-
duncle) of P. pedioulaba:
a- ventral view; b- lateral
view.
15(14)
16(15)
17(16)
Accessory (supplementary) eyes present (Fig. 6e); color light
olive-green variegated with brown, pale yellow and color-
less areas; length 38-51 mm (Fig. 19b): Plaodbdella hollensis
No accessory eyes 16
Ventrally striped blue, brown or green (Fig. 10); dorsal tu-
bercles inconspicuous or absent; color variable, usually
dark greenish-brown; few papillae, body depressed; length
38-64 mm (Fig. 19c): Plauaobdella parasitiaa
Not ventrally striped; dorsal tubercles prominent 17
Fig. 10- Ventral view of P.
parasitica showing stripes.
Median longitudinal rows of tubercles, especially conspicuous,
all tubercles bearing several papillae which gives rough,
warty appearance; color pattern of dorsum fine mixture of
brown, greens, and yellows with or without median dorsal
light or dark stripe interrupted by short brown lines; venter
unstriped; length 38-35 mm (Fig. 19d): Plaoobdella ovnata
With small more uniform tubercles and papillae 18
23
-------
18(17) Nearly uniform dark van dyke brown above, slightly paler at
head, with much darker brown median dorsal stripe contin-
uous for entire length but fading at head; about 30 dark
brown lines beneath surface which remain after preservation;
length 48-51 mm (Fig. 19e): Plaaobdella multilineata
5 prominent longitudinal rows of conspicuous, conical, tuber-
cles on dorsum, appearing as whitish spots on coffee brown
stripes; ground color, pale or dark brown with faint white
or yellow spots; length 15-45 mm (Fig. 19f):
Plaeobdella papillifera
19(9) A dense white or yellowish bar on somite VI; dorsal tubercles
usually large, acute, pale yellow or brown, deeply pigmented
and arranged in median longitudinal dorsal row and 2 other
rows halfway to margins; color greenish-brown, flecked with
dark spots; length 10-25 mm (Fig. 20a): Batvacobdella •ghaleva
No dense white or yellowish bar 20
20(19) 2 pairs of eyes (arrangement frequently modified by coalesc-
ence of eyes in various ways (Fig. 6f-j)); tubercles ab-
sent; color green or brownish-green; length 7-20 mm (Fig.
20b): Batvaodbdella paludosa
1 pair of eyes fused (Fig. 6b); dorsum smooth with 4 rows of
white spots; color dark greenish-brown, finely variegated
with orange; not ventrally striped; usually thin median
dorsal stripe; may or may not have semi-circular orange
spots along lateral margins; length 13-25 mm (Fig. 20c) :
Batraaobdella picta
21(9) Dorsal nuchal gland (or scute) conspicuous on somite VIII
(Fig. 11); no distinct dorsal tubercles; color dusky brown,
green or pink; length 9-14 mm (Fig. 20d): Helobdella stagnalis
No dorsal nuchal gland (or scute) present 22
Fig. 11- Dorsal view of H.
stagnalis showing position
of the nuchal gland. nuchai
gland
24
-------
22(21) Body subcylindrical, slender, smoothly rounded; trans-
lucent white or colorless; eyes 1 pair, widely separate
on somite IV; posterior sucker small and terminal in posi-
tion; length 9-25 mm (Fig. 20e): Helobdella elongata
Body moderately flattened; wider posteriorly (Fig. 2a, b);
anterior sucker confluent with body; posterior sucker
ventral in position 23
23(22) Dorsum with distinct transverse rows of white spots and prom-
inent longitudinal brown stripes; tubercles absent or
nearly so; length 10-20 mm (Fig. 20f):
Helobdella punatatolineata
Dorsum without distinct transverse rows of white spots 24
24(23) With 5 to 9 longitudinal rows of prominent and numerous tu-
bercles on dorsum, smooth, conical (black or dark brown);
general color brown; length 9-14 mm (Fig. 21a) :
Helobdella papillaba
Without 5 to 9 rows of prominent and numerous tubercles on
dorsum 25
25(24) Tubercles small, smooth and conical; deeply pigmented and
often double, many fine longitudinal light and dark brown
lines; length 9-14 mm (Fig. 21b): Helobdella lineata
Tubercles absent or limited to mid-dorsal line of posterior
body somites; color coffee brown or gray with 6 or 7
white spots on every 3rd annulus; length 10-14 mm (Fig.
21c): Helobdella fusea
26(10) Body entirely smooth, without papillae, 3 pairs of eyes in
rough triangular pattern of groups of 2 each (Fig. 5c);
with or without median dorsal brown stripe, but no paired
lines; color, uniform amber or whitish; length 6-9 mm
(Fig. 21d): Glossiphonia heteroolita
Body with papillae and/or tubercles; eyes 3 pairs in para-
median rows (Fig. 5b) but may have coalescence of eyes ... 27
27(26) Body with papillae in 6 longitudinal rows; 1 pair of dark
brown stripes broken by pale spots on dorsum and venter;
ground color brown, green, or gray; length 14-25 mm (Fig.
21e): Glossiphonia oomplanata
Body with large paramedian knob-like tubercles; dorsum with
large pale spots that reduce the dark brown ground color
almost to reticulum; paramedian dark stripes, heavy and
uninterrupted; ventrum bears no white spots; length 15-25
mm (Fig. 21f): Glossiphonia oomplanata mollissima
28(10) Gonopores separated by 2 annuli; olive-green or brown, nearly
transparent, flecked with black spots or conspicuously
spotted on dorsum; length 20-26 mm (Fig. 22a):
Theromyzon maculosim
Gonopores separated by 3 or 4 annuli 29
25
-------
29(28) Gonopores separated by 3 annul!; color variable with rounded
spots of yellow, orange or brown; length 20-30 nun (Fig.
22b): Theromyzon rude
Gonopores separated by 4 annuli; color transparent amber or
greenish with rounded spots; length 15-30 mm (Fig. 22c) :
Theromyzon tessulatwn
30(2) Caudal sucker flattened, as wide or wider than widest part of
body; pulsatile vesicles (11 pairs) on margin of body;
suckers distinctly set off from body (Fig. 3a-c) 31
Caudal sucker concave, weakly developed and narrower than wid-
est part of body (Fig. 3d, e); pulsatile vesicles absent .. 38
31(30) Pulsatile vesicles small, difficult to see on preserved spe-
cimens; body not clearly divided into anterior and posterior
regions (Fig. 3a) 32
Pulsatile vesicles large, clearly seen after preservation;
body distinctly divided into anterior and posterior regions
(Fig. 3b, c) 36
32(31) 8 to 14 ocelli spots on posterior sucker (Fig. 12a-c) 33
Oculiform spots absent from posterior sucker (Fig. 12d);
suckers clearly marked off from body; pulsatile vesicles
small (11 pairs) on sides of body; no dark rays on caudal
sucker; cephalic eyes 2 (or 1) pairs; color translucent,
greenish; length 14-16 mm (Fig. 22d): Piscioola punotata
Fig. 12. Dorsal view of posterior suckers: a- oculiform spots
only; b- oculiform spots and rays; c- crescent-shaped spots and
rays; d- oculiform spots absent.
33(32) Oculiform spots (10-14) on posterior sucker (Fig. 12a-c) 34
Crescent-shaped spots (8-10) on posterior sucker (Fig. 12c);
2 pairs of eyes; body and suckers may or may not be covered
with more or less disorderly arrangement of thickly flecked,
stellate spots; length 10-31 mm (Fig. 22e):
Pisoioola salmositiea
26
-------
34(33) Posterior sucker with dark rays and 12-14 oculiform spots .... 35
No dark rays, but 10-12 oculiform ocelli on posterior sucker
(Fig. 12a); color in mid-region of body yellowish with
brownish stellate flecks, disposed roughly in 5 longitu-
dinal rows; dorsally these flecks from slightly acute
(triangular) angle and spread out laterally; length 16-24
mm (Fig. 22f): Pisaiaola rrrllneri
35(34) 12-14 oculiform ocelli on posterior sucker separated by dark
rays (Fig. 12b); color greenish, yellowish, or brownish,
usually finely sprinkled with minute black or brown cells,
disposed more or less regularly in longitudinal rows;
length 20-30 mm (Fig. 23a): Pisaioola geometra
Posterior sucker with 14 dark, irregular brown rays each
terminated by an oculiform spot; 2 pairs of eyes, anterior
eyes larger; length 19-? mm (Fig. 23b): Piscieola zebra
36(31) Segments in middle of body with 6 annuli; 2 regions of body
sharply distinguished; lateral vesicles large; color of
dorsum dusky brown or purplish-brown, finely speckled with
stellate points of darker brown, and with irregular rows
of conspicuous, small, rounded, opaque, white spots along
upper surface of dorsum; length 15-? mm (Fig. 23c):
Cystobvanehus vividus
Segments in middle of body with 7 annuli 37
37(36) Gonopores separated by 2 annuli; pulsatile vesicles large;
void of pigment cells or with brownish-black spots pro-
fusely distributed over entire surface of body and
suckers; caudal sucker very large; length 10-30 mm (Fig.
23d): Cystobranehus vencilli
Gonopores separated 'by 1 or 2 annuli; pulsatile vesicles
large; posterior sucker with about 10 oculiform spots and
only slightly greater in diameter than anterior sucker;
length 9-15 mm (Fig. 23e) : Cystobrandhus virg-inious
38(30) Body divided into narrow anterior region and wider poste-
rior region (Fig. 3d); eyes 1 pair; no color or with
brownish-black stellate pigment cells; gonopores separated
by 8 annuli; length 24-26 mm (Fig. 23f): Myzobdella moorei
Body not divided into 2 regions (Fig. 3e) 39
39(38) Anterior sucker spatulate or bell-shaped (Fig. 3g) 40
Anterior sucker weakly developed, not spatulate or bell-
shaped 41
27
-------
40(39) Eyes present or absent, if present located on posterior half
of spatulate oral sucker; anus 15 annuli from posterior
sucker; color greenish, body wall usually devoid of pig-
ment; length 15-20 mm (Fig. 24a): Illinobdella riehardsoni
Eyes 1 pair, present on posterior half of somewhat spatulate
or bell-shaped sucker; sides of body nearly parallel; only
slight decrease in width anteriorly, practically none
posteriorly (this species is readily recognised by its
proportions); anus 10 annuli or less from posterior
sucker; length 25-30 mm (Fig. 24b): Illinobdella elongata
41(39) Body thin, devoid of pigment; eyes 1 pair on posterior half
of weakly developed anterior sucker; gonopores separated
by 8 annuli; length 9-10 mm (Fig. 24c): Illinobdella alba
Color consists of brown-black pigments arranged in 6 longitu-
dinal rows, especially 2 most dorsal rows; eyes 1 pair
in posterior region of oral sucker; somite 3-annulate;
length 6-8 mm (Fig. 24d): Piscioolaria reduata
42(3) Copulatory glands present behind gonopores on ventral surface
(Fig. 13) 43
Copulatory glands absent 45
O
'
^
O
o
3=;
o
XIII
i
'I
XIV
I
Fig. 13. Diagrams of the arrangement of copulatory gland pores:
a- MaoTobdella deoora;b- M. sestertiaj c- M. ditetra. (After
Moore).
43(42) About 21 bright red or orange spots in mid-dorsal line 44
Red spots absent from mid-dorsal line; 8 copulatory gland
pores (Fig. 13c); 2 annuli between gonopores; about 50
teeth per jaw; color of dorsum drab brown with darker
median field 1/3 body width, venter yellowish with some
or no black blotches; length 100-150 mm (Fig. 24e):
Maarobdella ditetva
28
-------
44(43)
45(42)
46(45)
47(46)
48(47)
49(47)
50(45)
4 copulatory gland pores (Fig. 13a); 5 annul! between gono-
pores; color of dorsum green with median row of red or
orange spots, lateral black spots; ventral surface red or
orange; about 65 teeth per jaw; length 110-150 mm (Fig.
24f): MaoTobdella decora
24 copulatory gland pores (Fig. 13b); 2 1/2 annuli between
gonopores; color of ventral side bright reddish brown with
few scattered flecks of black, dorsum olive-green with
median row of orange or red spots, lateral black flecks;
length 50-100 mm (Fig. 25a): MacTobdella sestert-ia
Dorsum with 1 or more colored stripes 46
Dorsum without colored stripes 50
Dorsal surface with single median stripe 47
Dorsum with pair of longitudinal red stripes; color pattern
variable, greenish background and pattern of irregular
black markings near lateral margins, ventral surface
black with white and gray markings; jaws with 35-100 teeth;
length 30-150 mm (Fig. 25b) : Hirudo medioinal-ls
Dorsal surface with median black stripe
Dorsal surface with another colored median stripe
48
49
With distinct median dorsal black stripe; ground color
uniform brownish- green to olive, with scattered black and
yellowish-orange blotches (usually more black than yellow-
orange); margins conspicuously mottled with yellowish-
orange blotches forming broken longitudinal lines; ventrally
darker, plumbeous and uniform, occasional yellowish-orange
blotches; teeth 9-12 pairs; length 60-90 mm; young with
metameric bands (Fig. 25c) : Pereymoorensis kingi
Dorsal surface with median black stripe and orange marginal
stripes, uniformly gray, ventrally lighter, few or no
dark blotches; jaws with 20-25 pairs of teeth; length 150-
200 mm (Fig. 25d) : Percymoorensis lateral-is
With dark brown median stripe, lateral margins with irregu-
lar black stripes sometimes broken but no discrete spots;
20 teeth per jaw; length 40-85 mm (Fig. 25e) :
Ph-ilobdella floridana
With light yellow median dorsal stripe and dorsolateral
brownish-black irregular spots; about 40 (35-48) teeth
per jaw; length 40-85 mm (Fig. 25f) : fhllobdella
Jaws absent or vestigial (Fig . 14a) .......................... 51
Jaws very small and retractable into narrow-mouthed tubular
pits, 10-16 coarse pairs of teeth per jaw (Fig. 14b, c) ... 52
29
-------
Fig. 14. Ventral view of the dis-
section of the mouth and buccal cavi-
ty and cross section of teeth: a-
Mollibdella grandis showing velum and
absence of jaws; b- Percymoorensis
maxmorat-Ls showing velum, relative
size of jaws; c- distichrodont ar-
rangement of teeth.
51(50) Dorsum grayish in color with few or no blotches, with red-
dish or orange band along margins; lower surface of velum
finely and closely papillate; gonopores in middle of
annuli separated by 5 annuli; pharynx with 15 internal
ridges; length 150-200 mm (Fig. 26a) : Bdellax-ogatis plwnbea
Color highly variable, but usually shades of dull green, gray,
or plain, always more or less blotched with black; lower
surface of velum smooth (Fig. 14a); pharynx with 12 inter-
nal ridges; gonopores in furrows, separated by 5 annuli;
length 175-300 mm (Fig. 26b): Mollibdella grandis
52(50) Jaws with 10-14 pairs of teeth (commonly 11-12); color olive-
green dorsally, heavily blotched with few scattered
yellow blotches, ventrally darker, uniform gray, few in-
distinct black or yellowish blotches; caudal sucker large,
about 3/4 width of body, discoid, broadly attached by very
short pedicel which tapers to direct attachment to somite
XXVII; length 50-85 mm (Fig. 26c):
Peraymoorens'is lateromaculata
Jaws with 12-16 pairs of teeth (Fig. 14b, c); color usually
blotched and more or less thickly flecked with black-brown
and yellow-gray; posterior sucker about 1/2 width of body;
length 75-100 mm (Fig. 26b) : Percymoorensis marmorat'is
53(3) Somites 5-annulate (bl, b2, a2, b5, and b6, with b6 never
distinctly subdivided all annuli being equal in length)
(Fig. 15b); dorsum usually with 2 or 4 longitudinal stripes
of brown of black spots but may be heavily black barred
form or pigmentless. Genus Erpobdelta 55
Somites 6- or 7-annulate, annuli differing in length with b6
either subdivided or longer than others, in any group
of 6 consecutive annuli at least 1 is either narrower or
wider than rest, narrower or wider annulus occurring
regularly along body (Fig. 15a); eyes 3 or 4 pairs 54
30
-------
sensillae,
Fig. 15. Annulation: a-
Dina (MooTeobdella)-flep'hel-
opsis showing the subdivi-
sion of b6; b- ErpobdeZla
showing the undivided
nature of annulus b6.
bl
55
b6
bl
bl
b2
a2
b5
b6
bl
54(53) Anterior and posterior 2 pairs of eyes arranged in parallel
(Fig. 4d); atrial cornua spirally coiled like rain's
horn (Fig. 16a, b); coloration gray, spotted with black
blotches or plain; length 26-40 mm (Fig. 26e):
Nephelopsis obseura
Atrial cornua simply curved (Fig. 17a-e); 3 or 4 pairs of
eyes (except D. anooulatd); dorsum with spots, longitudin-
al stripes, or none. Genus Dina: Subgenus Mooreobdella .... 57
55(53) 2 annuli between gonopores 56
3 annuli between gonopores; color, olive or dull green, mark-
ed with 4 longitudinal stripes composed of numerous small
black spots with pale centers in which sensory papillae are
situated, ventral surface and lateral margins plain gray or
ashy, unpigmented; 3 pairs of eyes, 1st pair much larger on
somite II; atrial cornua simply curved, ejaculatory duct
with long preatrial loop (Fig. 17a); length 25-30 mm (Fig.
27b): Erpobdella triannulata
XII ganglion
preatrial
loop
cornua
(paired horns)
genital
atrium
Fig. 16. Male genital atrium of N. dbsowca: a- dorsal view;
b- lateral view. (After Moore).
31
-------
56(55) 2 or 4 rows of black spots on dorsum or plain with few black
areas; atrial cornua simply curved, ejaculatory duct with
long preatrial loop (Fig. 17a); 3 pairs of eyes, 1st pair
smallest; length 80-100 mm (Fig. 26f): Erpobdella punctata
Same as above except heavily black barred form (Fig. 27a) :
Evpobdella punotata annulata
57(54) Eyes 3 or 4 pairs (Fig. 4b, c) 58
Eyes absent; gonopores in furrows separated by 2 annuli;
longitudinal stripes down body; ejaculatory duct with long
preatrial loop; length 10-15 mm (Fig. 27c): Vina anooulata
58(57) 3 pairs of eyes (except sometimes 4 pairs in D. (M.) fervidd)
(Fig. 4c) 59
4 pairs of eyes (Fig. 4b) 62
59(58) Gonopores separated by 2 annuli 60
Gonopores separated by 2 1/2 to 4 annuli 61
60(59) Atrium globular with prominent horns longer than its diameter
(Fig. 17b); eyes (3 or 4 pairs); gonopores separated by 2
annuli normally on rings; posterior sucker large; ejacu-
latory duct without long preatrial loop; color varying,
either lacking pigment entirely, or pale red with darker
clouding or with 2 narrow or broad, dark longitudinal
stripes extending over body, including always 1 light median
stripe; length 25-51 mm (Fig. 27d):Dina (Mooveobdella) fervida
Atrium globoid as above (Fig. 17c); ejaculatory duct without
long preatrial loop; gonopores separated by 2 annuli on
rings or in furrows; length 25-32 mm (Fi'g. 27e) :
Vina (M.) buoera
61(59) Gonopores separated by 3 annuli, usually in furrows; atrium
ellipsoidal, wider than long, with horns shorter than diam-
eter of median atrium (Fig. 17d); ejaculatory duct without
long preatrial loop; eyes 3 pairs; color reddish from blood
showing through or light yellowish or light grayish or unpig-
mented; length 30-50 mm (Fig. 27f): Dina (M.) microstoma
Gonopores separated by 3 1/2 annuli; 2 rows of black spots on
dorsum; atrial cornua short, merely curved (Fig. 17e); ejacu-
latory duct with preatrial loop; eyes 3 pairs; color liver
or reddish in life; length 25-50 mm (Fig. 28a): Dina lateralis
62(58) With few dark spots or no pigment; gonopores usually separated
by 3 1/2 (sometimes 2 1/2 or 3) annuli, male on annuli,
rarely in furrow, female in furrow; ejaculatory duact with
preatrial loop; length 25-29 mm (Fig. 28b): Dina parva
With dark brown or blackish median dorsal stripe visible in
cephalic half, fading out posteriorly, usually heavily
blotched; gonopores usually separated by 3 1/2 (sometimes
4) annuli, male on annuli, female in furrow; ejaculatory
duct with preatrial loop; length 25-41 mm (Fig. 28c):
Dina dubia
32
-------
Fig 17. Dorsal view of dissected male genital atrium and ejaculatory
duct with or without long preatrial loop: a- E. punatata; b- D. (M.)
fewida; c- D. (M.) buoera; d- D. (M.) miorostoma; e- D. lateralis.
(After Moore)
c- cornua (paired horns); g- XII, twelfth ganglion; g.a.- genital
atrium; p.l. preatrial loop; w- without preatrial loop.
33
-------
]fffe*
\Z3Kfi:.
•\sA'^/'\
\i-
f
Fig. 18. a- Marvinmeyeria lucida; b- Oligobdella biannulata; c- Aotinob-
della anneotens; d- A. triannulata; e- A. inequiannulata; f- Plaoobdella
montifera.
34
-------
Fig. 19. a- Plaeobdella pediculata; b- P. hollens-is; c- P. pafasitioa;
d- P. ornata; e- P. multilineata; f- P. papillifera.
35
-------
i6T*-Q';»:o UdJ
j'^'tJ'!-''VP ''';?
•tj •'•'-.'/.. f ;:i ;>v
550 d
Fig. 20. a- Batraaobdella phalera; b- S. paludosa; c- B. piota;
d- Eelobdella stagnalis; e- H. elongata; f- ff. punetatolineata.
36
-------
Fig. 21. a- Helobdella papillata; b- ff. lineata; c- 5. fusea; d-
Gloss-Lphonia 'heteicoolita; e- ff. complanata; f- G. eomplanata molliss-ima.
37
-------
*&£££&
Fig. 22.
a- Thevomyzon maculostm; b- y. rwcZe; c- T. tessulatim; d-
punotatai e- P. salmositiaa; £- P. nrilneri.
38
-------
Fig. 23. a- Pisoioola geometra; b- P. zebra; c- Cystobranohus vividus;
d- C1. verrilli; e- C. virginicus; £- Mysobdella moorei.
39
-------
f
Fig. 24. a- Illinobdella T-ichacdsan-i; b- J. elongata, c- J. a£i>a; d-
Pisoioo'Lar'La reduota; e- MaerobdeHa ditetra; f- M. decora.
40
-------
f
Fig. 25. a- Maevobdella sestertia; b- Hirudo medicinalis; c- Pepcymooren-
sis kingi; d- P. lateralis; e- Philobdella floridana; f- P. graeilis.
41
-------
f
Fig. 26. a- Bdellarogatis plumbea; b- Mollibdella grandis; c- Pevaymoor-
ensis lateromaaulata; d- P. mormoratis; e- Nephelops-is obsaura; f-
Erpobdella punatata.
42
-------
im
It
f
Fig. 27. a- Evpobdella punotata annulata; b- E. triannulata; c-
anooulata; d- D. (Mooreobdella) fervida; e- D. CM.J bucera; f- P.
microstoma.
43
-------
b
Fig. 28. a- Dina lateralis; b- P. porya; c- P. dub-La.
44
-------
SECTION IV
ACKNOWLEDGEMENTS
I wish to express sincere thanks to Dr. Bert M. Johnson, Biology Depart-
ment, Eastern Michigan University, for his initiating me into leech
research; to Dr. Henry van der Schalie, Museum of Zoology, Mollusk
Division, The University of Michigan, for his continued help in leech-
snail studies and particularly for his influence in making this key
possible; and to the Smithsonian Institution, The United States National
Museum, Department of Invertebrate Zoology, Division of Worms, in making
their leech collection and type specimens available for examination.
My studies have been supported (in part) by PHS Training Grant No. 5 Tl
Al 41 from the National Institute of Allergy and Infectious Diseases,
U. S. Public Health Service.
45
-------
SECTION V
REFERENCES
1 Autrum, H. 1936. Hirudineen. Bronn's Klassen und Ordnungen des Tier-
reichs, Bd. 4, Abt. 3, Buch 4, Teil 1, pp. 1-96, 60 Text-figs.
2 1939. Hirudineen. Bronn's Klassen und Ordnungen des Tierreichs,
Bd. 4, Abt. 3, Buch 4, Teil 2, pp. 1-662, 8 Text-figs.
[literature to 1938]
3 Bere, R. 1929. Reports of the Jasper Park Lakes Investigations 1925-
1926. III. Leeches. Contributions to Canadian Biology and Fish-
eries, 4:117-183.
4 Castle, W. E. 1900. Some North American freshwater Rhynchobdellidae
and their parasites. Bulletin of the Museum of Comparative
Zoology at Harvard College,, 36(2):17-64.
5 Davis, R. W. 1971. A key to the freshwater Hirudinoidea of Canada.
Journal of the Fisheries Research Board of Canada, 28(4):543-
552.
6 Eddy, S. and W. C. Hodson. 1950. "Taxonomic Keys to the Common
Animals of Minnesota Exclusive of the Peroschi Worms, Insects
and Birds." Burgess Publishing Company, Minneapolis, Minneso-
ta. 123 pp.
7 Graf, Arnold. 1899. Hirudinienstudien. Abhandlungen der Kaiser
Leopoldinisch-Carolinischen Deutschen Akademie der Naturforscher,
72:217-404.
8 Harding, W. A. 1910. A revision of British leeches. Parasitology, 3:
130-201.
9 Hemingway, E. E. 1908. 'Plaoobdella pediculata n. sp. American
Naturalist, 42:527-532.
10 Hoffman, R. L. 1964. A new species of Cystobranchus from southeastern
Virginia (Hirudinea:Piscicolidae). American Midland Naturalist,
72(2):390-395.
11 Keith, M. M. 1960. A simplified key to the leeches of Minnesota.
Proceedings of the Minnesota Academy of Science, 27:190-199.
12 Klemm, D. J. 1972. The leeches (Annelida:Hirudinea) of Michigan.
Michigan Academician, 4 (4): in press.
13 Mann, K. H. 1962. The systematics of freshwater and terrestrial
leeches, pp. 147-168 in "Leeches (Hirudinea). Their Structure,
Physiology, Ecology and Embryology." Pergammon Press, Oxford.
201 pp.
14 Mathers, C. K. 1954. Haemopis kingi n. sp. American Midland
Naturalist, 52(2):460-468.
15 1961. A study of North American species of the genus Haemopis
(Annelida:Hirudinea) with a description of a new species
Haemopis lateromaculatum. University Microfilms. Ann Arbor,
Michigan, pp. 1-48. Order No. 61-5593.
16 1963. Haemopis lateromaculatum n. sp. (Annelida:Hirudinea)..
American Midland Naturalist, 70:168-174.
17 Meyer, M. C. 1940. A revision of the leeches (Piscicolidae) living
on freshwater fishes of North America. Transactions of the
American Microscopical Society, 59:354-376.
47
-------
18 Meyer, M. C. 1946a. Further notes on the leeches (Piscicolidae)
living on freshwater fishes of North America. Transactions of
the American Microscopical Society, 65:237-249.
19 1946b. A new leech Pisciaola salmositica. Journal of
Parasitology, 32(5):467-476.
20 1968. Moore on Hirudinea with emphasis on his type specimens.
Proceedings of the United States National Museum, 125(3664),
32 pp.
21 Meyer, M. C. and J. P. Moore. 1954. Notes on Canadian leeches.
Wasmann Journal of Biology, 12:63-96. [Oculobdella lucida]
22 Miller, J. A. 1929. The leeches of Ohio. Contributions. Franz
Theodore Stone Laboratory, 2:1-38.
23 1937. A study of the leeches of Michigan with a key to orders,
suborders and species. Ohio Journal of Science, 37(2):85-90.
24 Moore, J. E. 1964. Notes on the leeches (Hirudinea) of Alberta.
National Museum of Canada Natural History Papers, 27:1-15.
25 1966. Further notes on Alberta leeches (Hirudinea). National
Museum of Canada Natural History Papers, 32:1-11.
26 Moore, J. P. 1898. The leeches of the U. S. National Museum.
Proceedings of the United States National Museum, 21:543-563.
27 1900. A description of Microbdella biannulata with especial re-
gard to the constitution of the leech somite. Proceedings of
the Academy of Natural Science of Philadelphia, 5(2):50-73.
28 1901. The Hirudinea of Illinois. Bulletin of the Illinois State
Laboratory of Natural History, 5:479-547.
29 1906. Hirudinea and Oligochaeta collected in the Great Lakes
region. United States Bureau of fisheries. Bulletin, 25:153-172.
[Actinobdella annectens, Placobdella montifera]
30 1912. The leeches of Minnesota. Minnesota. Geological and
natural history survey. Zoological series, No.5, Pt. 3:63-150.
[Haemopis plumbea, Dina parva]
31 1922. The freshwater leeches (Hirudinea) of southern Canada.
Canadian Field Naturalist, 36:6-11, 37-39. [Erpobdella punctata
annulata}
32 1924. The leeches (Hirudinea) of Lake Nipigon. Publications of
the Ontario Fisheries Research Laboratory, No. 23:17-31, 1 Fig.
[Actinobdella triannulata}
33 1908. The leeches of Lake Amatitlan. in Meek. The zoology of
Lakes Amatitlan and Atitlan, Guatemala, with special reference
to ichthyology. Publications. Field museum of Natural History.
Zoological series, 7:199-201. [Erpobdella triannulata}
34 1936. in Brandt. Parasites of certain North Carolina Salientia.
Ecological monographs, 6:491-532. [Macrobdella ditetra}
35 1939, Helobdella punctato-lineata, a new leech from Puerto Rico.
Puerto Rico Journal of Public Health and Tropical Medicine, 14:
422-429, 2 Figs.
36 1949. Hirudinea. pp. 38-39, in R. Kenk. The animal life of
temporary and permanent ponds in southern Michigan. Miscella-
neous Publications. Museum of Zoology, University of Michigan,
No. 71:1-66. [Dina bucera]
48
-------
37 Moore, J. P. 1952. Professor A. E. Verrill's freshwater leeches. A
tribute and critique. Notulae Naturae (Philadelphia), No. 245:
1-15, 7 Figs. [Helobdella papillata]
38 1953. Three undescribed North American leeches (Hirudinea).
Notulae Naturae (Philadelphia), No. 250:1-13, PI. 1, 2 Figs.
[Placobdella multilineata]
39 1959. Hirudinea, pp. 542-557. in Ward, H. B. and G. C. Whipple
(Eds). "Freshwater Biology." 2nd Ed. John Wiley and Sons, Inc.,
New York, N. Y. 1248 pp.
40 Moore, J. P. and M. C. Meyer. 1951. Leeches (Hirudinea) from Alaskan
and adjacent waters. Wasmann Journal of Biology, 9:11-77, Pis
1-11. [Dina dubia]
41 Pawlowski, L. K. 1955. Revision des genus Erpobdella de Blainville
et Dina R. Blanchard (Hirudinea). Bulletin de la Soaiete des
Sciences et des Lettres de Lodz. Class III, 6:1-15.
42 Pennak, R. W. 1953. Ch. 13, pp. 302-319. "Freshwater Invertebrates
of the United States." Ronald Press Co., New York, N. Y. 769 pp.
43 Raj, P. J. S. 1962. Morphological and biological studies of two
species of piscicolid leeches. University Microfilms, Ann Arbor,
Michigan, Order No. 62-4393. [Myzobdella moorei]
44 Rawson, D. S. 1953. The bottom fauna of Great Slave Lake. Journal of
the Fisheries Research Board of Canada, 10:486-520.
45 Richardson, L. R. 1969. A contribution to the systematics of the
hirudinid leeches with description of new families, genera and
species. Acta zoologica Academiae scientiarwn hungaricae, 15:
97-149.
46 Sawyer, R. J. 1972. North American freshwater leeches, exclusive of
the Piscicolidae with a key to all species. Illinois Biological
Monographs, 46:1-154.
47 Soos, A. 1963. Identification key to the species of the genus Dina
R. Blanchard, 1892 (Emend. Mann, 1952) (Hirudinea:Erpobdellidae)
Acta Universitatis Ssegediensis Acta Biologica, 9:253-261.
48 1965. Identification key to the leech (Hirudinoidea) genera of
the world with a catalogue of the species: I Family: Piscicoli-
dae. Acta zoologica Academiae scientiarum hungaricae, 2:417-463.
49 1966a. Identification key to the leech (Hirudinoidea) genera
of the world with a catalogue of the species: III Family:
Erpobdellidae, Acta zoologiea Academicae scientiarum hungaricae,
12:371-407.
50 1966b. On the genus Glossiphonia Johnson, 1816, with a key and
catalogue to the species Hirudinoidea:Glossiphoniidae. Annales
Historico-Naturales Musei Nationales Hungarici, 58:271-279.
51 1967. On the genus Batracobdella Viguier, 1879, with a key and
catalogue to the species Hirudinoidea:Glossiphoniidae. Annales
Historico-Naturales Musei Nationales Hungarici, 59:243-257.
52 1968. Identification key to the species of the genus Erpobdella
de Blainville, 1818 (Hirudinoidea:Erpobdellidae). Annales
Historico-Naturales Musei Nationales Hungarici, 60:141-145.
53 1969a. Identification key to the leech (Hirudinoidea) genera
of the world with a catalogue of the species. V Family:
Hirudinidae. Acta zoologica Academiae scientiarwn hungaricae,
15:151-201.
49
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54 Soos, A. 1969b. Identification key to the leech (Hirudinoidea) genera
of the world with a catalogue of the species. VI Family:
Glossiphoniidae. Aota zoologica Academiae scientiarum hungaricae3
15:397-454.
55 1970. A zoogeographical sketch of the freshwater and terrestrial
leeches (Hirudinoidea). Opusaula Zoologioa (Budapest), 10(2):
313-324.
56 Verrill, A. E. 1872. Description of North American freshwater
leeches. American Journal of Soienoe and Arts, 103:126-139.
57 1874. Synopsis of the North American freshwater leeches. Report
of the United States Commissioner of Fisheries 1872-73, Pt II,
pp. 666-689.
58 Viguier, C. 1879a. Anatomie comparee des Hirudinees. Organisation
de la Batracobdelle (Batraaobdella latasti, C. Vig.). Compte
rendu hebdomadaires des seances de I'Aoademie des sciences.
Paris, 89:110-112.
59 1879b. On the organisation of Batraohobdella (B. latasti, C.
Vig.). Annals and Magazine of Natural History, 4:250-251.
60 Whitman, C. 0. 1886. The leeches of Japan. Quarterly Journal of
Microscopical Science, n.s., 26:378-382, PI. XX, Figs 57-59.
50
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SECTION VI
INDEX TO SCIENTIFIC NAMES
Aotinobdella, 9,20
annectens, 9,22
inequiannulata, 9,22
triannulata, 9,22
alba (Illinobdella), 13,28
annectens (Aatinobdella), 9,22
anooulata (Vina), 5,15,18,32
atomaria (Erpobde1ta)3 2
BatraoobdeI la, 5,9,21
miahiganensis, 9
paludosa, 9,24
pnalera, 9,24
piota, 9,24
Bdellarogatis, 14
plumbea, 6,14,30
biannulata (Oligobdella), 11,22
buaera (Dina (Mooreobdella)), 16,32
oomplanata (Glossiphonia), 10,25
mollissima (Glossiphonia), 10,25
Cystobranehus, 5,12,18
verrilli, 12,27
virginious, 12,27
vivLdus, 12,27
decora (Maorobdella)3 7,15,29
Dina, 2,4,15,31
anooulata, 5,15,18,32
dubia, 15,32
fervida, 2
latefalis, 16,32
(Mooreobdella)3 4,6,7,16,31
buoera, 16,32
fervida* 16,32
miorostomaf 2,16,32
parva, 16,32
ditetra (Maerobdella)3 7,15,28
dubia (Dina)3 15,32
elongata (Helobdella)3 10,18,25
(Illinobdella), 13,28
Erpobdella3 7,16,30
atomariaj 2
ootoculata, 2
punotata3 16,32
annulata, 16,32
tviannulata, 16,31
Erpobdellidae, 15
fervida (Dina (Mooreobdella))3 2,16,
32
floridana (Philobdella)3 15,29
fusaa (Helobdella)3 2,5,10,25
geometra (Piscicola)3 13,27
Glossip'honia, 5,10,22
Gomplanata3 10,25
moll-iss-Cma, 10,25
heteroalita3 7,10,25
nepheloidea3 10
Glossiphoniidae, 9
graeilis (Philobdella)3 15,29
grandis (Haemopis)3 14
(Mollibdella)3 6,14,30
Eaemopis grandis3 14
kingi3 14
Iateralis3 14
lateromaoulata3 14
marmoratis3 14
plumbea3 14
terrestris3 14
Helobdella, 5,10,21
elongata, 10,18,25
fusaa, 2,5,10,25
Hneata, 2,5,10,25
pap-illata, 5,10,25
punetatolineata, 5,10,25
stagnalis, 11,24
transversa, 11
triseralis, 2
heteroclita (Glossiphonia), 7,10,25
Hirudidae, 14
H-irudo, 15
med-ioinalis, 15,29
hollensis (Plaoobdella), 11,20,21,
23
Illinobdella, 5,13
alba, 13
elongata, 13,28
moorel, 13
richardsoni, 6,13,28
inequiannulata (Aotinobdella), 9,22
king-i (Haemopis), 14
(Peroymoorensis), 14,29
lateralis (Dina), 16,32
(Haemopis), 14
(Peraymoorensis), 14,29
lateromaaulata (Haemopis), 14
51
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lateromaeulata (Peroymoorensis),
14, 30
lineata (Helobdella)3 2,5,10,25
luoida (Marvinmeyeria)3 7,11,21
(Ooulobdella), 11
Maorobdella, 15
decora, 7,15,29
ditetra, 7,15,28
sestertia, 7,15,29
maaulosum (Theromyzon), 12,25
marmor,atis (Haemopis), 14
(Peroymoorensis)3 14,30
Marvinmeyeria, 11
luoida, 7,11,21
medioina Us (Hirudo) 3 15,29
meyeri (Theromyzon), 12
miohiganensis (Batraoobdella), 9
miorostoma (Dina (Mooreobdella))3 2
16,32
mi Ineri (Pisoioo la) 3 13,27
Mollibdella, 14
grandis, 6,14,30
montifera (Plaeobdella)3 11,21,22
moorei (Illinobdella)3 3
(Myzobdella)3 13,27
Moopeobdella3 2
multilineata (Plaoobdella)3 5,11,24
Myzobde11a3 5,13
moorei3 13,27
nepheloidea (Glossiphonia)3 10
Nephelopsis3 16
obsoura3 4,6,7,16,31
obseura (Nephelopsis)3 4,6,7,16,31
ooaidentaUs (Theimmyzon) 3 12
oatoeulata (Erpobdella)3 2
Oculobdella luo-lda, 11
Oligobdella, 11
bisamulatat 11,22
ornata (Plaoobdella), 11,23
paludosa (Batraoobdella)3 9,24
papillata (Helobdella), 5,10,25
papillifera (Plaoobdella)3 5,12,24
parasitioa (Plaoobdella)3 5,12,23
parva (Dina)3 16,32
pedioulata (Plaoobdella), 6,12,23
Peraymoorensis, 14
kingi3 14,29
lateralis, 14,29
mcamoYatis, 14,30
lateromaaulata, 14,30
phalera (Batraoobdella), 9,24
(Plaoobdella), 9
Philobdella, 15
floridana, 15,29
15,29
piota (Batraoobdella), 9,24
(Plaoobdella), 9
Pisoioola, 5,13,18
geometra, 13,27
milneri, 13,27
punotata, 13,26
salmositioa, 13,26
zebra, 13,27
Pisoioolaria, 5,14
reduota, 14,18,28
Piscicolidae, 12
Plaoobdella, 5,11,22
> hollensis, 11,20,21,23
montifera, 11,21,22
multilineata, 5,11,24
ornata, 11,23
papiIlifera, 5,12,24
parasitioa, 5,12,23
pedioulata, 6,12,23
phalera, 9,
piota, 9
rugosa, 11
plumbea (Bdellarogatis)3 6,14,30
(Haemopis), 14
punotata (Erpobdella), 16,32
annulata (Erpobdella), 16,32
(Pisoioola), 13,26
punotatolineata (Helobdella), 5,10,
25
reduota (Pisoioolaria), 4,18,28
riohardsoni (Illinobdella), 6,13
rude (Theromyzon), 12,26
rugosa (Plaoobdella), 11
salmositioa (Pisoioola), 13,26
sestertia (Maorobdella), 7,15,29
stagnalis (Helobdella), 11,24
terrestris (Haemopis), 14
tessulatum (Theromyzon), 12,26
Theromyzon, 12,22
maaulosum, 12,25
meyeri, 12
oooidentalis, 12
rude, 12,26
tessulatum, 12,26
(Helobdella), 11
52
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triannulata (Aatinobdella)3 9,22 vivginieus (Cystobi>ana1ws)3 12, 27
(Erpobdella)> 16,31 wividus (Cystobranehus), 12, 27
triseralis (Helobdella)3 2 zebra (Pisoioola)y 13,27
verriZZ-i (Cystobranohus)3 12,27
53
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SELECTED WATER
RESOURCES ABSTRACTS
INPUT TRANSACTION FORM
1. Report No-
3. Accession No.
w
4. Title BIOTA op FRESHWATER ECOSYSTEMS IDENTIFICATION MANUAL "i/'ft^Jjbfi?,;., ,
NO. 8. Freshwater Leeches (Annelida:Hirudinea) of North f. ' •"" ;'*! ";,>,-'' *
America, t
Report K#.
7. Author(s)
Klemm, D. J.
9. OrganizationThe University of Michigan,
Ann Arbor,
Michigan,
12. Sponsoring Organization
15. Supplementary Notes
10. Project No.
18050 ELD
. Contract/Grant No.
14-12-894
3. Type of Report and
Period Covered
16. Abstract
An illustrated key to 63 species of North American freshwater
leeches (Annelida:Hirudinea) is given with notes on their
diagnostic features and distribution.
17a. Descriptors
*Aquatic fauna, *Annelida, Distribution, Preservation,
176.identifiers identification Manual, illustrated Key, *Hirudinea, *Leeches,
North America, Collection, Species List,
17c. COWRR Field & Group
10A
18. Availability
19. Security Class.
(Report)
20. Security Class.
(P*ge)
2t. ffo.of
Pages
22. Prict
Send To:
WATER RESOURCES SCIENTIFIC INFORMATION CENTER
US DEPARTMENT OF THE INTERIOR
WASHINGTON. D C 20240
Abstractor Donald J. Klemm
I institution The University of Michigan
WRSIC 102 (REV JUNE 1971)
ft U. S. GOVERNMENT PRINTING OFFICE : 1972 O - 468-268
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