&EPA
United States
Environmental Protection
Agency
Environmental Monitoring and _
Support Laboratory
Cincinnati OH 45268
600382025
April 1982
Research and Development
Leaches
(Annelida: Hirudinea)
of North America
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EPA-600/3-82-025
April 1982
LEECHES (ANNELIDA: HIRUDINEA) OF NORTH AMERICA
by
Donald J. Klemrn
Aquatic Biology Section
Environmental Monitoring and Support Laboratory
U.S. Environmental Protection Agency
Cincinnati, Ohio 45268
ENVIRONMENTAL MONITORING AND SUPPORT LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U.S. ENVIRONMENTAL PROTECTION AGENCY
CINCINNATI, OHIO 45268
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DISCLAIMER
This report has been reviewed by the Environmental Monitoring and
Support Laboratory, U.S. Environmental Protection Agency, and approved for
publication. Mention of trade names or commercial products does not
constitute endorsement or recommendation for use.
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FOREWORD
Environmental measurements are required to determine the quality of
ambient water, the character of effluents, and the effects of pollutants on
aquatic life. The Environmental Monitoring and Support Laboratory -
Cincinnati conducts an Agency-wide quality assurance program to assure
standardization and quality control of systems for monitoring water and
wastewater and carries out research to develop, evaluate, standardize, and
promulgate methods to:
* Measure the presence and concentration of physical, chemical, and
radiological pollutants in water, wastewater, bottom sediments, and
solid waste.
* Concentrate, recover, and identify enteric viruses, bacteria, and
other microorganisms in water.
* Measure the effects of pollution on freshwater, estuarine, and marine
organisms, including the phytoplankton, zooplankton, periphyton,
macrophyton, macroinvertebrates, and fish.
* Automate the measurement of physical, chemical, and biological quality
of water.
The effectiveness of measures taken to maintain and restore the biological
integrity of the Nation's surface waters is dependent upon our knowledge of
the changes in the taxonomic composition of the aquatic life caused by
discharges of toxic substances and other pollutants and upon the level of our
understanding of the complex relationships that prevail in aquatic
ecosystems. Leeches compose a significant portion of the fauna in a variety
of freshwater and marine habitats. On a world-wide basis some species are
important economically as parasites of mammals, fishes, and birds.
Occasionally, in North America some species are blood sucking pests to humans
in aquatic recreational areas. Other species are important as parasites and
predators of an assortment of animals. Also, leeches act as hosts of
parasites or as vectors for parasites of vertebrates. Several of the more
common species are associated with polluted waters, while others are
associated with water of high quality. This manual was developed to assist
biologists in identifying specimens of leeches to the species level and in
evaluating data collected during studies of the effects of toxic substances
and other pollutants on indigenous communities of aquatic organisms.
Robert L. Booth
Director
Environmental Monitoring and Support
Laboratoary - Cincinnati
iii
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PREFACE
The previous manual, "Freshwater Leeches (Annelida: Hirudinea) of
North America," Identification Manual No. 8, Biota of Freshwater
Ecosystems, published in 1972, was prepared under contract with the
Oceanography and Limnology Program, Smithsonian Institution, with funds
provided by the United States Environmental Protection Agency.
In the ten years that have elapsed since publication of the above
manual, additional work has been done in North America on the ecology and
systematics of the leeches. This publication serves to correct several
errors found in the above manual, to make several desirable taxonomic
changes, to add new sections and various relevant information to the text
with the intent of making the manual more useful, and to add new and
improved quality illustrations. The new materials include expanded
methods for collecting, narcotizing, fixing, and preserving leeches, notes
on systematics, distribution maps, a glossary, synonymies, and an expanded
bibliography. This is not a monographic revision, but offers some new
insights into the systematics of the North American freshwater leeches.
Hopefully, this publication will make identification easier and will
stimulate much needed research on the ecology of leeches, especially on
the water quality requirements and pollution tolerances of individual
species.
If errors are found in the manual, they should be transmitted to the
author. Also, constructive criticism regarding the manual will be
gratefully appreciated.
1v
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ABSTRACT
Leeches are represented in North America by four orders, five
families, 22 genera, and 63 species. The primitive family
Acanthobdellidae is represented by one genus and species. The families
Glossiphoniidae are represented by 10 genera and 29 species, the
Piscicolidae by four genera and 10 species, the Hirudinidae by three
genera and 13 species, and the Erpobdellidae by four genera, nine species
and two subspecies. The systematics are still incompletely worked out
for some of the groups, which makes a definitive listing of species
somewhat arbitrary at this time. Taxonomic problems are discussed, and
the present key in most instances reflects a conservative approach to the
lower taxa and represents all the known species. External characters are
used mostly in the key to separate the taxa, but some dissection of
internal anatomy may be required for the identification of a few species
in the families Hirudinidae and Erpobdellidae.
The main features of this publication are an introduction, methods of
collection, narcotization, fixation and preservation, important notes on
systematics and identification, an illustrated key for species level
identification, a glossary, species distribution, a partial synonymy, and
a selected bibliography which includes the references cited in the text
and other publications which provide additional information on taxonomy
and ecology of the Hirudinea.
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CONTENTS
Foreword ill
Preface 1v
Abstract * v
Figures viii
Tables xiii
Distribution Maps xiv
Acknowledgments xvi
Section 1. Introduction 1
Section 2. Methods 6
A. Collection 6
B. Narcotization, Fixation, and Preservation 7
C. Depository for Leeches 10
Section 3. List of Species 11
Section 4. Notes on Systematics 15
Section 5. Notes on Identification 21
Section 6. Key to Leeches (Hirudinea) of North America, North of
Mexico 31
Section 7. Glossary 95
Section 8. Species Distribution 101
Section 9. Partial Synonymy 137
Selected Bibliography 151
Index to Scientific Names 174
vii
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FIGURES
Number Page
1. Some activities of leeches that may have a direct
or indirect effect upon the ecology of fishes 3
2. Diagrams of methods to expose the male reproductive system
of Erpobdellidae 29
3. Acanthobdella pel e din a 31
4. Anterior (oral) suckers 32
5. General body shapes of Glossiphoniidae 33
6. General body shapes of Glossiphoniidae 34
7. Arrangement of eyes, Glossiphoniidae 34
8. Arrangement of eyes, Glossiphoniidae 35
9. Arrangement of eyes, Glossiphoniidae 36
10. General bod|y shapes and external characters of Piscicolidae . . 37
11. General bocjy shapes of tyyzobdella and Piscicolaria 38
12. Attachment sites of fish leeches 39
13. Arrangement of eyes, Hirudinidae and Erpobdellidae 40
14. General body shapes of Hirudinidae or Erpobdellidae 41
15. Placobdella montifera, dorsal view 42
16. Placobdella nuchalis, dorsal view 42
17. Placobdella hollensis, dorsal view 44
18. Harvinmeyeria lucida, dorsal view 44
19. Proboscis and scute of Helobdella stagnalis, dorsal view. ... 44
20. Helobdella stagnalis, dorsal view 44
VTM
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FIGURES - continued
Number
21. Helobdella elongate, dorsal view 46
22. Helobdella transversa, dorsal view 46
23. Helobdella fusca, dorsal view 46
24. Helobdella papillata, dorsal view 47
25. Helobdella triserialis, dorsal view 47
26. Helobdella triserial is, dorsal view 47
27. Helobdella triserialis, dorsal view 48
28. Batracobdella phalera, dorsal view 49
29. Batracobdella michiganensis, dorsal view 50
30. Placobdella translucens, dorsal view 50
31. Caudal sucker of Placobdella pediculata 51
32. Caudal sucker of BatracobdelTa cryptobranchii 51
33. Caudal sucker of Actinobdella inequiannulata 51
34. Caudal sucker of Actinobdella annectens 51
35. Actinobdella inequiannulata, dorsal view 52
36. Actinobdella inequiannulata, ventral view 52
37. Actinobdella inequiannulata, dorsal view 52
38. Actinobdella annectens, dorsal view 52
39. Placobdella pediculata, juvenile form 53
40. Placobdella pediculata, lateral view 53
41- Placobdella pediculata, dorsal view 53
42. Batracobdella cryptobranchii, dorsal view 53
43. Placobdella parasitica, ventral view 54
44. Placobdella parasitica, dorsal view 54
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FIGURES - continued
Number Page
45. Placobdella parasitica, dorsal view 55
46. Bactracobdella pieta, dorsal view 56
47. Oligobdella biannulata, dorsal view 56
48. Placobdella ornata, ventral view 57
49. Placobdella ornata, dorsal view 57
50. Placobdella papillifera, ventral view 58
51. Placobdella papillifera, dorsal view 58
52. Placobdella paplllifera, dorsal view 58
53. Placobdella multilineata, dorsal view 58
54. Batracobdella paludosa, dorsal view 59
55. Alboglossiphom'a heteroclita, dorsal view 59
56. Alboglossiphom'a heteroclita, dorsal view 60
57. Tri annul ate somite of Glossiphom'a complanata and
Boreobdel 1 a verrucata showing papillae arrangement 61
58. Glossiphom'a complanata, dorsal view 61
59. Boreobdella verrucata, dorsal view 61
60. Theromyzon biannulatum, Theromyzon rude, dorsal view 62
61. Theromyzon tessulatum, dorsal view 63
62. Piscicolaria reducta, dorsal view 64
63. Hyzobdella lugubris, dorsal view 64
64. Hyzobdella lugubris, dorsal view 65
65. Caudal suckers of Piscicolidae 66
66. Cystobranchus meyeri, lateral and dorsal view 67
67. Cystobranchus virginicus, dorsal view 67
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FIGURES - continued
Number Page
68. Cystobranchus mammillatus, dorsal view 68
69. Cystobranchus vern'Th', dorsal view 68
70- Piscicola punctata, dorsal view 68
71* Piscicola salmositica, dorsal view 69
72. Piscicola milneri, dorsal view 69
73. Piscicola geometra, dorsal view 69
74. External reproductive structures of Hirudinidae 70
75- Macrobdella dttetra, dorsal view 72
76- Macrobdella sestertia, dorsal view 72
77. Hacrobdella decora, dorsal view 73
78- Macrobdella diplotertia, dorsal view 73
79- Pnilobdella floridana, dorsal view 75
80. Philobdella gracilis, dorsal view 75
81• HIrado medicinalis, dorsal view 75
82. Haemopis septagon, dorsal view 76
83. Haemopis terrestris, dorsal view 78
84. Haemopis kingi, dorsal view 78
85. Mouth and buccal cavity of Hirudinidae 79
86* Haemopis lateromaculata, dorsal view 80
87« Haemopis marmorata, dorsal view 80
88. Mouth of Haemopis plumbea, ventral view 81
89. Mouth of Haemopis grandis, ventral view 81
90. Haemopis plumbea, dorsal view 82
91. Haemopis grandis, dorsal view 82
xi
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FIGURES - continued
Number Page
92. Male and female gonopores of Erpobdellidae 83
93. Dissected male reproductive system of Erpobdellidae 84
94. Mooreobdella melanostoma, dorsal view 86
95. Erpobdella punctata coastal is, dorsal view 86
96. Erpobdella punctata punctata, dorsal view 86
97 • Erpobdella punctata punctata, dorsal view 87
98. Dina anoculata, dorsal view 87
99. Mooreobdella bucera, dorsal view 87
100. Mooreobdella fervida, dorsal view 89
101. Mooreobdella microstoma, dorsal view 89
102. Mooreobdella tetragon, dorsal view 89
103. Dissected male reproductive system of Erpobdellidae 91
104. Nephelopsis obscura, dorsal view 92
105. Dina dubia, dorsal view 93
106. Dina parva, dorsal view 93
xii
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TABLES
Number
1. The theoretical table of the annul 1 (after Moore, 1898) . . .
22
X111
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DISTRIBUTION MAPS
Number
Pages
1. Acanthobdella peledlna 105
2. Actlnobdella annectens, A. inequlannulata. 106
3. A1bogloss1phon1a heteroclita 107
4. Batracobdella cryptobranchll, ft. michiganensis, B. phalera ... 108
5. Batracobdella paludosa, £. plcta 109
6. Boreobdella verrucata, Glosslphonia complanata 110
7. Helobdella elongata, H_. fusca Ill
8. Helobdella papHlata, Marvlnmeyeria lucida 112
9* Helobdella stagnalls, ]U transversa 113
10. Helobdella triserialls 114
11. Ollgobdella blannulata, Placobdella hollensls 115
12. Placobdella montlfera, P_. tnultillneata 116
13. Placobdella nuchal is, £. ornata 117
14. Placobdella papllUfera, £. pedlculata 118
15. Placobdella parasltlca, £. trans!ucens 119
16. Theromyzon blannulatum, J. rude, J. tessulatum 120
17. Cystobranchus mammlllatus, C. meyerl 121
18. Cystobranchus verri 11 i a £. vlrglnlcus 122
19. Myzobdella lugubrls, Plsclcola geometra 123
20. Plsclcola punctata, P^. salmosltica 124
21. Plsclcola nrflnerl, P1sc1colar1a reducta 125
x1v
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DISTRIBUTION MAPS - continued
Number Pages
22. Haemopis grandis, H. kingi 126
23. Haemopis lateromaculata, H.. marmorata 127
24. Haemopis plumbea, H. septagon, H. terrestris 128
25. Macrobdella decora, M. diplotertia 129
26. Macrobdella ditetra, fl. sestertia 130
27. Philobdella floridana, P. gracilis 131
28 • Dina anoculata, J). dubia, £. parva 132
29* Erpobdella punctata coastalis, _E. £. punctata 133
30. Hooreobdella bucera, Q. fervida, PI. melanostoma 134
31. Mooreobdella microstoma, fl. tetragon 135
32. Nephelopsis obscura 136
xv
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ACKNOWLEDGMENTS
I wish to extend my thanks to Roy T. Sawyer, Bruce Daniels, and Thomas
P. Poe for their reviews of the technical contents of the manuscript. I
also thank Cornelius I. Weber for reading the manuscript and making
valuable suggestions. I especially thank Marvin C. Meyer and Roy T. Sawyer
for sharing their thoughts and opinions on Hirudinea ecology and taxonomy.
I am grateful to the following persons and institutions: Marian H.
Pettibone and Meredith L. Jones, Division of Worms, Department of
Invertebrate Zoology, U.S. National Museum of Natural History, Smithsonian
Institution, Washington, D. C.; John D. Unzicher, Illinois Natural History
Survey, Urbana, Illinois; Richard Franz, Florida State Museum, The
University of Florida, Gainesville, Floirda; H. W. Levi, Museum of
Comparative Zoology, Harvard University, Cambridge, Massachusetts; Donald
G. Huggins, State Biological Survey of Kansas, The Unversity of Kansas,
Lawrence, Kansas; James Reddell, Texas Memorial Museum, The University of
Texas, Austin, Texas; Peter Frank, Invertebrate Zoology, National Museums
of Canada, Ottawa, Canada; Philip Lambert, British Columbia Provincial
Museum, Department of the Provincial Secretary, Victoria, British Columbia,
Canada, and The Academy of National Sciences of Philadelphia, Department of
Invertebrate Zoology, Philadelphia, Pennsylvania for making the leech
collections of these museums available for my examination.
I am particularly indebted to George Ford and Frances 0. Paulson of the
Worm Division, U.S. National Museum of Natural History, Smithsonian
Institution, Washington, D.C. for their kind assistance with the leech
collections and library references.
The following persons have helped to supply specimens for either
identification or verification and for distribution records and should
receive special thanks: Omar Amin, Michael T. Barbour, L. Todd Beck, C.
Dale Becker, David C. Beckett, Michael D. Bilger, R. Bland, Edward Bolton,
Eugene M. Burreson, Stephen W. Carney, William H. Clark, Mary G. Curry,
Bruce A. Daniels, Charles R. Demas, Scott R. Derrickson, William G.
Deutsch, Graig W. Dye, John M. Epler, Tom M. Freitag, Dean Furbish, Joseph
W. Gorsuch, Mike Gross, George L. Harp, Charles 0. Hatcher, A. K. Hauck,
Ethel Helsel, Jarl K. Hiltunen, Glenn L. Hoffman, Peter Hovingh, John R.
Howland, Richard Howmiller, James L. Hulbert, Guy M. Johnson, Michael W.
Jones, Jerry Kaiser, Martin L. Kopenski, Robert E. Kuntz, David R. Lenat,
Philip A. Lewis, Gerry Mackle, Anthony F. Maciorowski, Jacqueline Madill,
Willis E. McConnaha, Mike Mclntosh, Charles N. Merckel, Lewis Giles Miller,
Robert L. Newell, Preston Newman, Jr., Gene K. Okamoto, Jeff Osborn, John
A. Osborne, Peter M. Nolan, Lee 0. Pendergraft, Wendell L. Pennington,
Thomas" P. Poe, Janet L. Polk, Bayliss Prater, Buford C. Pruitt, Jr., Fred
W. Robe, Henry W. Robison, Brenda J. Rogers, Harvey D. Rudolph, Robert P.
xvi
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ACKNOWLEDGMENTS - continued
Rutter, Gerard M. Sala, Harry M. Savage, Steward C. Schell, Gerald
Schnytema, Donald Schultz, Douglas G. Smith, Gerry Smrchek, August A.
Staats, Charles S. Steiner, Kurt S. Stimpson, Daniel Stoneburner, Donald
Tarter, George Te, William M. Turner, Bruno Vincent, Robert E. Watson, Mark
J. Wetzel, Steve White, Wilbur J. Widmer, Nixon A. Wilson.
I would also like to acknowledge the use of the following
illustrations: Thomas P. Poe (Fig. 12b-e);- Roy T. Sawyer and Rowland M.
Shelley (Fig. 52).
The secretarial assistance of Cordelia Nowell and Diane Schirmann is
gratefully acknowledged.
xvii
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SECTION 1
INTRODUCTION
The leeches are evolutionary derived from an ancient protostome
stock which gave rise to the segmented worms (Phylum Annelida), thus
establishing the metameric line. The earliest fossil record of leeches
comes from the Jurassic Period of Bavaria (Kozur, 1970). Other fossil
annelid records of the middle Cambrian Period indicate that they were
already differentiated into well-established groups (such as Polychaeta,
Oligochaeta, and Hirudinea). The leeches are considered the most highly
specialized annelids and are thought to have arisen from an oligochaete
progenitor.
The Class Hirudinea is divided into four Orders: Acanthobdellida,
Rhynchobdellida, Gnathobdellida, and Pharyngobdellida. On a world-wide
distribution Soos (1969b,1970) indicated that the Class Hirudinea contains
over 500 species, approximately 140 genera, and ten families. Currently,
in North America the leech fauna is composed of five families, 22 genera,
and 63 species.
leeches are predominantly freshwater invertebrates, but there are many
marine forms, as well as numerous terrestrial species which occur mainly
in the tropics. Unfortunately, all leeches are often disliked by the
layman and popularly considered to be "bloodsuckers" because of the
bloodsucking habits of a few species. As a result, it is usually this
activity that attracts the attention of anyone who encounters them.
However,- their food habits are far more diverse than most people realize;
many are not sanguivorous. These animals are predatory or parasitic with
anterior and posterior suckers that serve as organs of attachment,
feeding, and locomotion. Some species are morphologically adapted for
obtaining and digesting food consisting chiefly of the blood of fishes,
turtles, crocodilians, frogs, salamanders, birds, and mammals; other
species can also consume blood fluids, tissues, and whole live or dead
invertebrates, such as annelids (including leeches), crustaceans, insect
larvae, and mollusks.
Members of the Class Hirudinea compose a significant part of the North
American invertebrate fauna in both lentic and lotic waters. However, we
know very little about the ecology, natural history, and water quality
requirements of individual species. Herrmann (1970), Klemrn (1972a), and
Kopenski (1969) are the only broad ecological studies for the group in
North America. Sawyer (1974) has also reviewed the pertinent literatuare
for the ecology of freshwater leeches especially in relationship to
various kinds of pollution. The mention of other important papers with
specific information on natural history and ecology can be found in the
Selected Bibliography.
1
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Leeches are important components of food webs (Fig. 1), as predators,
parasites, vectors of parasites, and as food of aquatic animals.
Economically, the leeches of North America are not nearly as serious a
pest as tropical leeches, but they occasionally become numerous and
bothersome to humans in aquatic recreational areas. More importantly,
their activities may have a direct or indirect effect upon the life
histories of fishes (Hoffman, 1979; Khaibulaeu, 1970; Paperna and Zwerner,
1974; Poe, 1972; Shuster, Smith, and McDermott, 1951; Thompson, 1927; and
others listed in the Selected Bibliography. Leeches are also hosts
(Corkum and Beckerdite, 1975; Fish and Vande Vusse, 1976; Vande Vusse,
1980) or intermediate hosts (Becker, 1965-1977; Becker and Katz, 1975;
Khan, 1980; Klemn, 1975; Mann, 1962; Putz, 1972) in the life cycles of
potentially disease-causing blood protozoans, trematodes, and cestodes of
fishes. In addition, certain leeches infest waterfowl, but the incidence
and significance of these infestations are poorly understood or documented
(Meyer and Moore, 1954; Roberts, 1955; Keymer, 1969; Bartonek and Trauger,
1975; Trauger and Bartonek, 1977; McKinney and Derrickson, 1979; Davles
and Wllkialis, 1981).
Leeches are sometimes ignored in macrolnvertebrate analysis or
misidentified by investigators not familiar with their morphology or
taxonomy. Frequently authors of environmental studies have recorded the
group by class, family, genus or merely as "leeches." The improper or
Inadequate treatment of the leeches is attributable at least in part, to
the lack of a practical key to all species, unfamiliarity with the current
literature, and the difficulty associated with identifying some preserved
specimens. To determine the water quality requirements and pollution
tolerances of aquatic organisms, the animals must be identified to the
species level (Resh and Unzicker, 1975; Carricker, 1977).
This publication excludes the leeches of the West Indies, an
archipelago 1n the Caribbean Sea between North and South America, which
are discussed in Sawyer and Kinard (1980); the leeches of Central and
South America which are discussed to genera 1n Ringuelet (1976) and the
exclusively marine leeches. An identification manual to the marine fauna
of North America is not currently available, but is badly needed.
However, the following publications will help and should be consulted for
identifying most marine taxa: Burreson (1976a,b; 1977a-d), Burreson and
Allen (1978), Davies (1978), Knight-Jones (1962), Khan and Meyer (1976),
Meyer and Khan (1979), Moore (1946), Moore and Meyer (1951), Sawyer, et
al. (1975), Sawyer and Kinard (1980), Soos (1965), and Appy and Dadswell
(1981).
Earlier taxonomic keys to the leeches include: Moore, J.P. (1918,
1959), Klemm (1972b), and Sawyer (1972) for North America; Mann (1962) for
the British Isles, Central Europe, and North America; Pennak (1953, 1978)
for the United States; Meyer (1946a) and Hoffman (1976) for the fish
leeches of North America; Hayunga and Grey (1976) for fish leeches of the
genus Cystobranchus; Soos (1962-1969b) for genera or species of the world
with catalogues of the species; Davies (1971) for Canada; Klemm (1976) for
leeches found in North American mollusks; Ringuelet (1976) for families
and genera of MesoAmerica and South America; Mann (1964) and Elliot and
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FIG. 1. Some feeding and behavioral activities of leeches that may have
a direct or indirect effect upon the ecology of fishes: (a) leech
feeding upon the eggs of fish (and other aquatic animals); (b) fish
feeding on leech; (c) leeches feeding on the blood of fishes; (d) leech
infesting an aquatic reptile (birds, etc.); (e) leeches feeding on snails
(and other macrolnvertebrates). (Modified from Shuster, Smith, and
McDermott, 1951).
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Mann (1979) for the British Isles. Keys for specific geographic regions
of North America include: Moore, J.P. (1906, 1912, 1922) for the Great
Lakes Region, Minnesota, and Southern Canada; Bere (1929) for Jasper Park
(Alberta, Canada); Miller (1929; 1937) for Ohio and Michigan; Ed(ly and
Hodson (1950) for the North Central United States; Keith (1960) for
Minnesota; Moore, J.E. (1962; 1965) for Alberta, Canada, and Sawyer and
Shelley (1976) for North and South Carolina.
All the above keys have limited use today because they are either
outdated, are based only on sparce collections, or are limited in the
number of species for which they are written. Furthermore, some of the
authors neglected consideration of the overall morphological variation
found in some species, and thus these keys have been deficient for use in
the identification of specimens collected from throughout the geographic
ranges of the species. Finally, they are in need of the taxonomic changes
proposed by Meyer (1968), Sawyer (1972), Daniels and Freedman (1976),
Klemm (1976, 1977), Sawyer, et al. (1975), and Sawyer and Shelley (1976).
They lack the several new species recently described by Meyer (1975),
Hayunga and Grey (1976), Johnson and Klemm (1977), Sawyer and Shelley
(1976), the redescription of a species by Smith (1977), and new and
important geographical distribution records for many species.
Therefore, I have attempted in the present manual to analyse
critically the leech species of the families Glossiphoniidae,
Piscicolidae, Hirudinidae, and Erpobdellidae in North America. This
entailed the examination of over 15,000 live and preserved specimens
collected or received by the author for identification or verification by
the many individuals mentioned in the Acknowledgment and also the
examination of leech collections from museums also mentioned in the
Acknowledgment.
The manual includes a methods section, a species list, notes on
systematics and identification, an illustrated key, a glossary, a species
distribution section, a section on synonymy for each species in North
America, and an extensive bibliography. The section on Methods contains
information on collecting and processing specimens. The forms with
uncertain taxonomic status are discussed in the section on Notes on
Systematics. In the section on Notes on Identification, emphasis has been
placed on external characters. Where internal characters are used,
dissection methods are given. The dichotomous key comprises as many
external characters for identification as necessary, and the drawings were
made from living and preserved specimens. All the illustrations with the
exception of Figures 12b-e and 52 were drawn by the author. To facilitate
identification, each species and the most common variety have been
illustrated. The morphological terms for the diagnostic characters used
to differentiate the species are defined in the Glossary. The section on
Species Distribution consists of a discussion and maps for each species
found in North America. The bibliography of the primary literature on
leeches of North America is presented to encourage and aid future
investigations. Additional works are found in the bibliographies
of Sawyer (1972), Klemm (1972a, 1977), and Windsor (1972).
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This manual was prepared to provide USEPA and other biologists in
Federal, state, and private water monitoring agencies with an updated and
improved key of the species, to consolidate available information on
distribution so that specimens of leeches can be identified to species,
and to evaluate data collected during water quality studies concerning the
effects of toxic substances and other pollutants.
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SECTION 2
METHODS
COLLECTION
In surveying any body of water for its leech population it is
important to remember that they usually avoid strong sunlight. Free-
living species are found attached to the sides, in cracks, and under
surfaces of a variety of substrates such as rocks, boards and logs, or
almost any inanimate object littering both lentic and lotic environments.
This includes submergent vegetation, leaf packs, and masses of other
organic debris.
Many species of Glossiphoniidae and Piscicolidae are well adapted as
ectoparasites in that they pierce and suck the blood and tissue contents
of their hosts. The collection and examination of vertebrates
(amphibians, aquatic reptiles, fishes, and waterfowl) and macroinverte-
brates (snails, mussels, and so on) are, therefore, occasionally required
when collecting certain species of these leeches. The parasitic forms
remain attached to the host and cling there for a period of time, and then
drop off after feeding and creep away to seek the shelter of a suitable
substrate where they may remain while digesting their meal. Most
parasitic forms are generally found free-living during part of their
development or reproductive cycle (deposition of cocoons, or eggs, and
brooding young) and require a substrate for attachment.
Most species of Hirudinidae have jaws and teeth and are either
sanguiorous (blood sucking), or macrophagous (swallowing their
invertebrate prey whole), but the few species lacking jaws and teeth are
strictly macrophagous. Species of Erpobdellidae also lack jaws and are
macrophagous. Hirudinids and erpobdellids are usually collected
free-living.
Some species of Haemopis (Hirudinidae) are amphibious, while H.
septagon and some populations of H. terrestris, are terrestrial. The
terrestrial species are collectecTfrom moist places under rocks, boards,
logs, or almost any type of inanimate object, usually near water.
Sometimes, however, terrestrial populations are found a great distance
from water. On occasion, these leeches can even be collected when they
are foraging for food along the shores of bodies of water or on land when
moist atmospheric conditions exist, especially at night. Leeches are
collected either by hand, with forceps, or any other instrument which can
be used to gently dislodge them. Some benthic forms may be collected with
bottom samplers (grabs or dredges) while some of the actively swimming
species can be collected with a dipnet. In addition, dipnets are useful
in collecting debris, vegetation, and organisms which may be examined for
leeches.
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NARCOTIZATION, FIXATION, AND PRESERVATION
In some cases, it may be extremely difficult or impossible to identify
some leech specimens to the species level because of faulty preparation
and unsuitable preservation. Leeches are very sensitive, and respond to
irritants and react differently to different substances and concen-
trations. They have a soft, highly contractile body. Therefore, if
leeches are dropped alive into preservatives such as 70% alcohol
(ethanol), 5-10% formalin solution (commercial formaldehyde), for
instance, they contract strongly and sometimes such features as the eyes,
general boc|y shape, and the genital pores become distorted and difficult
for the non-specialist and occasionally even for the expert to discern.
When leeches are properly anesthetized prior to fixation, there is usually
less muscular contraction. The body shape remains more uniform, and there
is less variabilty of the anatomical structures. Many specimens of
leeches which are preserved rapidly without anesthetization or fixation
can still be identified to the species level, but this is not the ideal
procedure because it can delay or prevent the identification of some
specimens. Experience will help alleviate this problem. The preserved
leech should be straight, moderately extended and undistorted. For best
results they should be fixed and preserved in fluids strong enough to
prevent maceration or softening of the tissues but not so strong that they
are rendered overly hard and brittle. Leeches are ruined completely when
they dry out.
Leeches should never be flattened between glass plates and the like
except for special histological purposes. Flattening causes distortion of
the body and internal organs, and the important external organs of the
integument can become altered, thus causing difficulity in determining the
diagnostic characters.
The characteristic color patterns on the dorsal surface of some
leeches, especially the blue, green, red, and yellow pigments (see
chromatophore), can be dissolved or altered by preserving in ethanol, and
it is sometimes very desirable that the color of the living leeches be
noted on the label of the specimen jar and used in the identification of
some specimens. However, some of these colors seem to persist for longer
periods of time if the specimens are first fixed in formalin and then
preserved in alcohol. Brown and black chromotophores, usually remain
after using either preservative. Some people prefer to use alcohol
because formalin is odoriferous, irritating to the skin and eyes, and
causes the specimens to become hard and brittle over time.
The addition of ecological data to the record can help in the
identification and increases the value of the specimen. Sometimes, to
ensure proper identification and to make a thorough anatomical study of a
species, several specimens are needed. A single specimen, especially when
poorly preserved, may be either impossible or at least difficult to
identify to the species level and may, therefore, not make a good specimen
for morphological studies.
If large specimens of the eropbdellids or hirudinids are placed
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directly in 70% alcohol (ethanol) rather than first fixed in formalin, the
body cavities of the specimens should be injected with 70% alcohol to
insure preservation of the internal organs, which may be necessary for
identification of some specimens.
The following methods for preparing specimens for taxonomic studies
and a reference colection are recommended by the author, and all will
generally give good results:
Narcotizing
Specimens can be narcotized through direct placement into carbonated
water, in 70% alcohol {ethanol), or in a 5-10% solution of Chloretone
added slowly to the container of water containing the leeches. These
should be added gradually, increasing the concentration, until all
movement stops or the specimens no longer respond to probing. Depending
on size and number being nacotized, the leeches should be completely
relaxed in a 15 to 30 minute period.
Another method requires adding a few drops of chloroform to the water
containing the leeches (The chloroform will sink to the bottom if
introduced beneath the surface of the water with a pipette or eye
dropper), and covering the container with a glass plate until the leeches
are anesthetized.
One per cent solution of propylene phenoxetol or sodium nembutal may
also be used for narcotizing leeches. Excessive relaxation of specimens
in these nacotizing agents can cause the furrows between the annuli to
diasppear. Therefore, the specimens must be fixed in 10% formalin as soon
as possible after nacotization.
Specimens can be placed directly in a 4% ether solution (4 ml ether:
96 ml water) in a stoppered bottle. The volume of ether solution should
be approximately 10 times the volume of the leeches. This procedure
(Richardson, 1975) will result in the narcotization and death of the
leeches in two to five hours, depending on the size of the specimens and
the temperature (e.g. small specimens and at warmer temperatures). This
solution is bacteriostatic and if specimens for systematics and general
morphological studies, are left in it for 15 to 20 hours at high room
temperature, 35 to 406C, the leeches do not deteriorate appreciably.
However, for best results, do not leave in the stock solution for long
periods of time because leeches having a very weak or thin muscular bo
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Fixing
After narcotization, large specimens should be drawn between the
fingers and washed so that the mucus is removed if secreted. Small
specimens can be wiped free of mucus with a small brush or piece of paper
toweling. Then, straighten the specimen out and place between damp paper
toweling or filter paper in a tray, such as a dissecting tray, and gently
flood with 10% formalin for 12 hours to fix and prevent softening of the
tissue. Small specimens can be fixed in a shorter period of time. After
the tissue is hard, the leeches are placed in the final preservative.
Preserving
Wash specimen in water to remove the formalin of fixation and preserve
in 70%-80% ethyl alcohol (ethanol), or preserve specimens in 5% buffered
formali n.
If whole mount slides of some specimens are prepared for anatomical
details, the modified technique of Palmeieri, et at. (1973) is excellent.
The most critical step in preparing a quality specimen for slides or
display is anaesthetizing. Place specimens in culture dish with clean
water and refrigerated at 35 to 40°F (2 to 4.5°C) for 40 minutes.
Refrigeration slows movement and makes the leech easier to handle. After
refrigeration, place the leech between two microscope slides, one of which
is covered with paper toweling, and apply slight pressure. The
"sandwiched" leech is then placed under running hot tap water
(approximately 180°F or 82°C) for about one minute. This has a
narcotizing effect and yields straight, flat leeches without excess
mucus. Care must be taken to use only hard paper toweling, as the fibers
of soft papers tend to adhere to the leech. Elastic bands or spring clips
(Meyer, 1957) should be placed around the slides, and the entire
leech-paper towel-slide complex placed into a 10% neutral formalin, (FAA,
Bouin's, or Flemming's fixatives) fixing solution for about 24 hours.
After fixation, the leeches should be removed from between the slides,
cleaned and placed in a Petri dish filled with distilled water.
Leeches should be stained with Mayer's paracarmine stain (Gray, 1954)
or Harris1 hematoxylin. The specimens are stained for 12 to 24 hours,
then destained in a "\% HCL-70% ethanol solution until the leech epidermis
is free of stain. After being destained, leeches should be neutralized in
1% MH40H-70% ethanol solution.
With large leeches, counterstaining may not be desirable, but to show
organs of the integument of small leeches, fast green or eosin may be
used. Stained specimens should be dehydrated by passing them through a
series of ethanol solutions, and remain at least 1-4 hours at each ethanol
concentration (35% -50% -70% -80% -95% -100% -100%). Leeches should be
cleared for at least 20 minutes in methyl salicylate (xylol tends to
harden leeches) and mounted in a neutral pH mounting medium. When
mounting large leeches, cover slips can be supported with chips of
microscope slides. See Meyer and Olsen (1971) for methods for whole
mounting and histological sectioning of leech material.
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DEPOSITORY FOR LEECHES
The value of a Hirudinea collection is that it will provide permanent
preservation for type-specimens, other voucher specimens, and records of
collection data, and that it may serve biologists both here and abroad as
a reference for further systematic and ecological studies.
Leech material which is no longer needed should be deposited in The
United States National Museum of Natural History or some other museum that
is concerned with the safety of the specimens, has a reputation for
properly maintaining the collection, and will allow accessibility of the
voucher specimens for scientific research.
Specimens with proper collection data can be sent to the Division of
Worms, Department of Invertebrate Zoology, U.S. National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 40560. The specimens
should be correctly preserved and labelled. The following information
should be included for each specimen: *(1) the locality, including state,
county, and distance from nearest town, if known, *(2) date collected, (3)
scientific name (if known), (4) habitat, *(5) collector, (6) name of
person who identified the specimen, and (7) other ecological information
if available, such as found free-living, or parasitic, and host; the
scientific name of the host (if known), and so on. Specimens sent to
museums without the minimum (*) accompanying data are worthless.
10
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SECTION 3
LIST OF SPECIES
Phylum Annelida
Class Hirudinea Lamarck, 1818
Order Acanthobdellida Livanow 1905
Family Acanthobdellidae Livanow, 1905
Genus: Acanthobdella Grube, 1851
Acanthobdella peledina Grube, 1851
Order Rhynchobdellida Blanchard, 1887
Family Glossiphoniidae
Genus: Actinobdella Moore, 1901
Actinobdella annectens Moore, 1906
Actinobdella inequiannulata Moore, 1901
Genus: Alboglossiphonia Lukin, 1976
Alboglossiphonia heteroclita (Linnaeus, 1761)
Genus: Batracobdella Viquier, 1879
Batracobdella cryptobranchii Johnson & Klemm, 1977
Batracobdella michiganensis Sawyer, 1972
Batracobdella paludosa (Carena, 1824)1
Batracobdella phalera (Graf, 1899)
Batracobdella picta (Verrill, 1872)
Genus: Boreobdella Johansson, 1929
Boreobdella verrucata (F.R. Muller, 1844)
Genus: Glossiphonia Johnson, 1817
Glossiphonia complanata (Linnaeus, 1758)
Notes On Distribution.
11
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Genus: Helobdella R. Blanchard. 1896
Helobdella elongata (Castle, 1900)
Helobdella fusca (Castle, 1900)
Helobdella papillata (Moore, 1906)
Helobdella stagnalis (Linnaeus, 1758)
Helobdella transversa Sawyer, 1972
Helobdella triserialis (E. Blanchard, 1849)
Genus: Marvinmeyeria Soos, 1969
Marvinmeyeria lucida (Moore, 1954)
Genus: Oligobdella Moore, 1918
Oligobdella biannulata (Moore, 1900)
Genus: Placobdella blanchard, 1893
Placobdella hollensis (Whitman, 1892)
Placobdella montifera Moore, 1906
Placobdella multilineata Moore, 1953
Placobdella nuchalis Sawyer & Shelley, 1976
Placobdella ornata (Verrill, 1872}
Placobdella papillifera (Verrill, 1872)
Placobdella parasitica (Say, 1824)
Placobdella pediculata Hemingway, 1908
Placobdella translucens Sawyer and Shelley, 1976
Genus: Theromyzon Phillippi, 1867
Theromyzon biannulatum Klemm, 1977
Theromyzon rude (Baird. 1869)
Theromyzon tessulatum (O.F. Muller, 1774)
Family Piscicolidae Johnston, 1865
Genus: Cystobranchus Diesing, 1859
Cystobranchus mamm///atus(Ma\m. 1863)
Cystobranchus meyeri Hayunga & Grey, 1976
Cystobranchus verrilli Meyer 1940
Cystobranchus virginicus Hoffman, 1964
Genus: Myzobdella Leidy, 1851
Myzobdella lugubris Leidy, 1851
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Genus Piscicola de Blainville, 1818
Piscicola geometra (Linnaeus, 1 758)
Piscicola milneri (Verrill, 1 874}
Piscicola punctata (Verrill, 1871)
Piscicola salmositica Meyer, 1946
Genus: Piscicolaria Whitman, 1889
Piscicolaria reducta Meyer, 1 940
Order Gnathobdellida Vaillant, 1890
Family Hirudinidae Whitman, 1886
Genus: Haemopis Savigny, 1822
Haemopis grandis (Verrill, 1 874)
Haemopis King! Mathers, 1 954
Haemopis lateromaculata Mathers, 1963
Haemopis m armor at a (Say, 1824)
Haemopis plumbea Moore, 1912
Haemopis septagon Sawyer & Shelley, 1976
Haemopis terrestris (Forbes, 1890)
Genus: Hirudo Linnaeus, 1758
Hirudo medicinalis Linnaeus, 1758 2
Genus: Macrobdella Verrill, 1 872
Macrobdella decora (Say, 1 824)
Macrobdella diplotertia Meyer, 1 975
Macrobdella ditetra Moore, 1953
Macrobdella sestertia Whitman, 1886
Genus: Philobdella Verrill, 1874
Philobdella floridana (Verrill, 1874)
Philobdella gracilis Moore, 1901
..
Order Pharyngobdellida Johnson, 1913
\
Family Erpobdellidae
Genus: Dina R. Blanchard, 1892
Dina anoculata Moore, 1 898
Dina dub/a Moore & Meyer, 1951
Dina parva Moore, 1912
2See Notes On Distribution.
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Genus: Erpobdella de Blainville, 1818
Erpobdella punctata coastalis Sawyer & Shelley, 1976
Erpobdella punctata punctata (Leidy, 1870}
Genus: Mooreobdella Pawlowski, 1955
Mooreobdella bucera Moore, 1949
Mooreobdella fervida Verrill, 1871
Mooreobdella melanostoma Sawyer & Shelley, 1976
Mooreobdella microstoma (Moore, 1901)
Mooreobdella tetragon Sawyer & Shelley, 1976
Genus: Nephelopsis Verrill, 1872
Nephelopsis obscura Verrill, 1872
14
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SECTION 4
NOTES ON SYSTEMATICS
The taxonomy of Hirudinea has suffered greatly in the past because of
the brief and ambiguous descriptions of species which have been based only
on a single or few specimens, and on the use of a few, mainly external
characters, such as eyes and color. In fact, the lack of distinct
taxonomic characters even occurs in some genera. Furthermore, authors
have usually neglected consideration of the full range of variation of
forms throughout geographic ranges. Consequently, the critical
examination and comparison of museum specimens with additional leech
material and the descriptions of these earlier species has led to an
excessive number of synonyms (cf. Partial Synonym Section).
The taxonomy employed in this manual is basically that proposed by
Moore (1959) and Klemm (1972b, 1976, 1977), and the variations proposed by
Sawyer (1972), Sawyer et a!., (1975), as well as Sawyer and Shelley
(1976). In preparing this manual, however, a clarification and refinement
of the nomenclature for the leeches of North America has been attempted.
A discussion of specific and generic changes and major existing problems
fol 1ows.
FAMILY GLOSSIPHONIIDAE
Under the genus Glossiphom'a Johnson, 1817, some hirudinologists
recognize anatomical differences between £. complanata (Linnaeus, 1758)
and G. heteroclita (Linnaeus, 1761) at greater than species level. For
example, Lukin (1976) recognized the strong difference between the two
species and, therefore, erected the subgenus Alboglossiphom'a to
accommodate heteroclita. Sawyer (personal communication) and I believe
the distinction between the two species to be at the generic level. Thus,
I have elevated Alboglossiphonia to generic rank, including under it
heteroclita, but retaining compTanata under Glossiphonia.
Glossiphom'a swampina Bosc (1802), redescribed by Sawyer (1973), is
reported from the Carolinas (Sawyer and Shelley, 1976). They suggest that
G< swampina represents a disjunct derivative of the northern species, A.
Feterocjita. However, Moore (1952) indicated that Gi. swampina is A.
heteroclita or a very closely related species. In an earlier publication,
I (Klemm, 1976) discussed the two species and suggested that (a. swampina
might be a color variant of A. heteroclita. The new material from Quebec
and Maryland was examined ancT the appearance of the specimens agrees with
that of £. swampina, thus extending its distribution. After examining
specimens of both color forms from the Carolinas and the Great Lakes
region, and comparing their anatomy, I am convinced that these two forms
15
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represent color differences within one species. I consider £. swampina a
synonym under A_. heteroclita.
Variations of taxonomic characters due to polymorphism are discussed
in Sawyer (1972) and Klemm (1976, 1977) for Helobdella fusca (Castle,
1900), H. punctatolineata Moore (1939), and H. triserial is (E. Blanchard,
1849). After examining and comparing the anatomy of many North American
specimens that resemble H. punctatolineata with specimens from the West
Indies, I have concluded that there are no distinguishing characters to
separate the two species, H. punctatolineata and H. triseri all's. Thus, H_.
punctatolineata has been pTaced in synonymy under~H. trlseri'alTs.
In addition to the typical color form of H_. fusca (Fig. 23b), four
other unpapillated forms were encountered during this study. Three forms
(Fig. 23b-d) resembled specimens discussed by Sawyer (1972). The fourth
form is new (Fig. 23e). It has a whitish dorsal surface with irregularly
scattered black stellate pigment cells (chromatophores). During this
study, specimens of this form were received only from Quebec. Until the
degree of polymorphism of H. fusca is determined, it should remain a
distinct species.
Several color variations (Figs. 25a-b, 26a-b, 27) of H. triseri all's
have been reported (Ringulet, 1943, 1944, 1945; Sawyer, VS72, Klemm, 1976,
1977, and this study). A new distinct form (Fig. 26c), not previously
discussed, was also encountered during this study. Its dorsal surface
contains the typical three rows of black (with some pale white) papillae
but also had irregular shaped clusters of metameric white spots which fuse
together to form three prominent longitudinal white stripes and two
smaller white stripes in the anal region. The dorsal ground color is gray
with some longitudinal gray striping. Specimens of this form were
examined from the Detroit River, near Fighting Island, and from Quebec.
There is still considerable confusion in the systematics of the genus
Theromyzon (Klemm, 1977), and the various nominal species have undergone
considerable synonymy (Soos, 1969; cf. Partial Synonymy Section). In
North America, the identification of J_. biannulatum Klemm (1977), T. rude
(Baird, 1869), and J. tessulatum (O.F. Muller, 1774) is based mainTy on
color and the number of annuli (two, three, or four) between the male and
female gonopores. Soos (1969) recognizes T. maculosum (Rathke, 1862), T.
rude, and T^ tessulatum as the species witF the two, three or four annuTi
between gonopores. Eavies (1971, 1973) and I (Klemm, 1972a,b) later
accepted the synonymy of Soos for both J. maculosum and T. rude. However,
in my review of Theromyzon of North America (Klemm, 197/T, Sawyer
(personal communication) and I are still not convinced that the
establishment of J. maculosum bearing the two annul 1 between gonopores and
unique pigmentation pattern, occurs in North America. Furthermore, Sawyer
(personal communication) and I (Klemm, 1977) believe that Moore's (1912)
description of T. occidental is (cf. Partial Synonymy) from Minnesota is
neither T. rude nor T. maculosum. Therefore, I (Klemm, 1977) proposed the
new name~T.~5Tjinnulatuni for Moore's (1912) occidental is, the Theromyzon
form from North America with the two annul 1 between the gonopores. Based
on Meyer and Moore's (1951) contention that the number of annuli between
16
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gonopores of T. rude varies, Sawyer (1972) and I (Klemm, 1977) postulated
that the validity of separating these species on the basis of only the
number of annuli between gonopores is questionable. To resolve this
problem, a detailed examination of the internal anatomy of the various
species of Theromyzon and a comparison of the annuli between gonopores of
specimens in different geographical ranges (cf. Species Distribution
Section) is needed to determine whether only one or two species exist in
North America. Until these problems are critically examined, I recognize
the following three American Theromyzon taxa: T. biannula turn (with two
annuli separating the gonopores), T. rude (with three annuli between the
gonopores), and T. tessulatum (gonopores separated by four annuli). A
fourth species, T. propinquus Ringuelet (1947), known from Argentina,
closely resembles T. rude b~having three annuli between the gonopores and
might be synonymous.
FAMILY PISCICOLIDAE
In North America, the taxonomic status of several species of
Piscicolidae is uncertain. A description of the external morphology of
Piscicola zebra is given by Moore (1898), but there is no information
concerning the internal morphology to ascertain that its species status is
sound. P. zebra was, taken from the lip of the sea lamprey, Petromyzon
marinus Tan anadromous parasitic fish), at Arichat, Cape Breton, Nova
Scotia, in 1890, but has not been reported since. Soos (1965), Davies
(1971), and I (Klemm, 1972b) considered it a legitimate species based on
Moore's earlier description. However, the only available syntypes (USNM
4818) are poorly preserved and difficult to assess. Sawyer et al. (1975)
could not determine the validity of the species and concluded that it
should be considered as species inquirendae. I have reconsidered the
status of this taxon, and am now of the opinion that until new material is
collected and a detailed study of the anatomy is published, this species
should not be recognized as valid.
Some doubt also exists as to whether Piscicola geometra (Linnaeus,
1758) and ?_. nrilneri (Verrill, 1874) are synonymous.Current knowledge on
the differences between the two species shows that P. milneri has 10-12
(ususally 10) punctiform eyespots on the caudal suc¥er and lacks dark
pigmented rays, while £. geometra has 12-14 punctiform eyespots and the
presence of dark pigmented rays. According to Mann (1962, p. 161),
another difference lies in the annuli between the gonopores. There are
two in P. mi 1neri and three in P_. geometra. Moore, (1959) and Mann (1962)
state tfiat in both species, sperm ducts are simply looped. Meyer (1946)
and Moore and Meyer (1951) indicate that the general body forms of the two
are closely similar; Soos (1964) stated that especially the size and color
of P. geometra, are also highly variable. Holmquist (1975) and I (Klemm,
1977) maintain that some variability exists in the number of punctiform
eyespots in the caudal sucker and in the presnece or absence of the
pigmented rays as a result of the age of the specimens. A critical
examination of the internal anatomy of mature specimens of both species is
needed to determine species clarification. However, Holmquist (1975), as
well as this author, suspect that the two species are identical.
17
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I included Cystobranchus vividus Verrill (1872) in my earlier key to
freshwater leeches (Klemm, 1972b) on the bases of the descriptions by
Verrill (1872) and Moore (1898). Later, Sawyer, et al. (1975) reassigned
Verrill's species vividus to the genus Calllobdella van Beneden and Hess
(1863), as C. vivida (Verrill, 1872), because of the lack of no type
material amT because of the vague and inadequate descriptions of both
Verrill and Moore which caused confusion in the specific name vividus.
Since £. vivida is a marine and estuarine species, I have not included it
in this manual of freshwater leeches.
In regard to another piscicolid problem, Sawyer (1972) and Sawyer and
Shelley (1976) state that Cystobranchus virgi'nicus Hoffman, 1964, known
from Virginia and West Virginia, may be transferable to another genus.
FAMILY HIRUDINIDAE
The genus Haemopis Savigny (1822), in the family Hirudinidae, was
recently subdivided by Richardson (1969) into three genera,
Percymoorensis, Mollibdella, and Bdellarogatis. This revision was
followed by Soos (1969b), Da vies U971, 19/Z, 1973), and me (Klemm,
1972a,b). However, Sawyer (1972), in his morphological investigation of
the genus Haemopis, rejected Richardson's revision of Haemopis into the
new genus Percymoorensis for the North American species H. marmorata (Say,
1824), H. terrestrisTForbes, 1890), H. laterpmaculata Mathers (1963), and
11- kingT Mathers (1954) and the placement of H. cjrandis (Verrill, 1874)
and H. plumbea Moore (1912) into the monotypic genera Mollibdella and
BdelTarogatis respecitvely. Later, Sawyer's view was strengthened with
additional anatomical information and the description of a new species, H_.
septagon (Sawyer and Shelley, 1976), which does not belong in any of
Richardson's narrowly defined genera. They indicated that if the revision
is accepted, a new monotypic genus must be erected to accummodate H_.
septagon. Consequently, they placed the new species in the genus
haemopis. This manual and previously I (Klemm, 1977) follow the
recommendations of Sawyer (1972) and Sawyer and Shelley (1976) for the
genus Haemopis in North America.
Verrill (1874) established Philobdella as a subgenus of Hacrobdel1 a on
the basis of the external genital region, which is characterized by having
either copulatory glands containing pores behind the gonopores, or
copulatory depressions and pits around the gonopores (Fig. 73a-e). By
1898, J.P. Moore had elevated Philobdella to full generic rank. Soos
(1969b) grouped Philobdella with Macrobdella, but others have accepted
Moore's treatment (Pennak, 1953, 1978; Sawyer, 1972; Klemm, 1972b, 1977),
and I have retained it in this manual.
Moore (1952) and I also doubt whether there are two species of
Philobdella Verrill, 1874 (?_. floridana Verrill, 1874 and P. qracilis
Moore, 1901). The type specimen, P. floridana, is from LaFe Okeechobee,
Florida, and is known only from the original description. Moore (1901),
in a morphological study of specimens of Philobdella from Illinois,
described a new species, P. gracilis. Most authors continue to
18
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distinguish P_. floridana and P_. gracilis as separate species, primarily on
the basis of pigmentation and number of teeth. Nothing is known of the
internal anatomy of P. floridana, except the dentition of the jaws. Moore
(1959) stated that tfie denticles, as orgim'ally described for P.
floridana, are 20 teeth per jaw, and in P. gracilis are 40 (3^48) per
jaw. He indicated the dentition as partTy distichodont. Richardson
(1972), in an anatomical study of P_. gracilis from S. Carolina, indicated
only a single row (monostichodont) of 20 acute teeth, with no indication
of distichodonty. The study of Richardson (1972) also showed that the
number of teeth of £_. gracilis differed greatly from the number reported
in Moore's (1901) orginial description, 40 (35-48), but was similar to the
number of teeth in P_. floridana. Therefore, the variability in the number
of teeth suggests it is a poor diagnostic character for the two species.
During this study, specimens which resemble £. floridana were sent to me
from Jacksonville, Florida, and were found to have 24 monostichodont teeth
per jaw. However, the internal anatomy of these specimens has not been
studied. The variability of teeth and color of these forms indicate that
there is considerable doubt that Philobdella is represented in the
southern states as two species. A comparison of the internal anatomy of
the two forms would resolve the question.
FAMILY ERPOBDELLIDAE
The taxonomic rank of certain erpobdellid groups, Dina R. Blanchard,
1892, Erpobdella de Blainville, 1818, and Mooreobdella Pawlowski (1955),
sustained a number of changes over the yeaFs"! The Dina-Erppbdella complex
still remains unresolved. Sawyer (1972) and I (Klemm, 1977) have
discussed the taxonomic characters that have caused the confusion. In
Sawyer's opinion (personal communication) the genus Dina does not actually
occur in North America. Moore (1959), Meyer (1968), Sawyer (1972), Sawyer
and Shelley (1976), and I (Klemm, 1977) consider Mooreobdella a distinct
genus, endemic to North America, and consists of five species: bucera
Moore, 1949; fervida, Verrlll, 1871; melanostoma, Sawyer and Shelley,
1976; microstoma, (Moore, 1901); and tetragon, Sawyer and Shelley, 1976.
A subspecies, Erpobdella punctata annulate, of E. punctata (Leidy,
1870) was described by Moore (1922) from British CoTumbia .Tt differs
from £. D. punctata only by the conspicuously barred or completely black
annulT of its dorsum. Meyer and Moore (1954) reported finding both
subspecies throughout Canada, and I (Klemm, 1972a,b) have reported these
forms from Michigan, Washington, and Oregon. Moore (1959) Soos (1966a),
Davies (1971), I (Klemm, 1972a,b) retained this color form at the
subspecies level. Sawyer (1972), however, synonymized £. p annulata under
E. D. punctata. During this study, I found no evidence of anatomical
Difference between the two forms, other than color and size, and recognize
the synonymy of Sawyer (1972).
Dina lateralis Verrill (1871), a poorly described species, was
relegated to synonym under Erpobdella punctata punctata by Sawyer (1972).
Soos (1966a) and Klemm (1972b) retain the species, based on Moore's
interpretation (1952, 1959) of the species, as D. lateralais or £.
lateralis (Verrill). However, no type materal Ts available, no new
19
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material has been collected, and because of the vague and inadequate
description by VerHll, I concur with the synonymy of Sawyer.
Dina anoculata Moore (1898), described from San Diego County,
California, is poorly known. Moore's vague description of the
morphological type states that the species has no eyes, the gonopores are
separated by two annuli, and the body is colored with longitudinal
stripes. However, nothing is known of its internal reproductive system.
Sawyer (personal communication) stated he had collected specimens from San
Diego with three pairs of eyes. I have examined several syntypes (USNM
4844) from San Diego County, California, and have added to Moore's
description the internal anatomy of the reproductive system. Based on
information from Moore (1898); Soos(1963), Sawyer (personal
communication), as well as an examination of preserved material, the
following is a more complete description for j). anoculata:
Eyes absent or if present, 3 pairs, in separate labial
and buccal groups (Fig. 13b); genital pores separated by
two annuli (Fig. 92d); male pore in furrow XIb6/XIIb1,
female pore also in furrow XIIb2/a2. Annulation:
Somites I-IV:1, V-VI:3, VII-XXIV:5, XXV:3, XXVI-XXVII:! (2);
three annuli behind anus. Atrium (Fig. 93d) is globoid with
prominent horns (cornua) projecting laterad and with long
preatrial loops of vas deferens extending to ganglion XI.
I have not seen fresh material, but the ground color of specimens
preserved in alcohol is dull yellowish gray, immaculate below and on the
margins, but largely replaced above by four longitudinal stripes (vague in
some specimens) of grayish or dull black. The outer pair of stripes is
submarginal, duller in color, and narrower than the more distinct inner
pair, which is separated by a median stripe of ground color.
20
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SECTION 5
NOTES ON IDENTIFICATION
The taxononric key in this manual, which has been constructed mainly of
external anatomical and biological characters, is sufficient for the
identification of most North American leeches to the species level.
However, the identification of a few species of Haemopis and some
Erpobdellidae, especially species of the genera Mooreobdella and Dina, may
require dissection. If reference is made in a couplet to an internal
feature after reference to an external one, the key can be used first for
the identification of some specimens without resorting to dissection,
provided the external characters are clearly discernible. Also, some
collections will contain specimens that can not be identified to the
species level for various reasons, but they should be identified to the
lowest taxonomic category (taxon) possible, based only on characters of
the specimen used in the key. A glossary of diagnostic terms has been
provided.
The external diagnostic features which are important for identifying
the leeches are: presence or absence of chaetae in the cephalic region,
size of mouth, general shape of body, form of suckers, form of cephalic
region, number and arrangement of eyes, presence or absence of jaws and
number of teeth (denticles), eyespots (ocelli), papillae, pulsatile
vesicles, sensillae, digitate processes on rim of caudal (posterior)
sucker, caudal sucker separated from body on narrow pedicle (peduncle),
copulatory gland pores, the number of annuli per somite (segments) and
between gonopores, and the pigmentation patterns.
Only seven internal characters are used in the key, four of which are
used to determine certain species of Haemopis: the presence or absence of
jaws, the number of teeth (denticles) per jaw, the velum, and the internal
ridges of the pharynx. The shape of atrium and atrial horns, and the
length of the ejaculatory ducts are three other internal anatomical parts
used in the identification of some species of erpobdellids. Biological
characters useful for the identification of specimens include hosts,
methods of feeding, the manner of locomotion, and caring for eggs and
young, and ecological and geographical variations.
The body of the leech consists of 34 somites (segments), designated by
Roman numerals (I through XXXIV). Each somite contains a ganglion in the
central nervous system. Each neuromeric somite is divided externally by
superficial furrows into 2-16 annuli or rings . 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 characteristic of
21
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the genera or species. Incomplete or abbreviated somites occur at both
ends of the body. The annul i are best seen after careful narcotization
or proper preservation. Reference to annulation in the key is always to
complete somites, and with the aide of a dissecting stereomicroscope the
outline of the annuli can be seen most easily in the lateral margins of
the ventral surface. Moore (1898) and Castle (1900) independently
recognized that the nerve cord ganglia are located in the middle annulus
of the somite, and that the triannulate somite is the basic form. In the
3-or 5-annulate somite for example, the middle annulus is aligned with
the ganglion and is known as the neural or sensory annulus. The
annulation suggested by Moore (1898) (Table 1) is used throughout the
key. Counting from the cephalic end, the triannulate somite consist of
three primary annuli (best seen in middle of body), numbered al, a2, and
a3 (or sometimes written al-3). Annulus a2 (the neural annulus) contains
the ganglion and is marked externally by transverse rows of minute,
cutaneous sensillae (segmental receptors), which are difficult to see in
some specimens. Repeated bissection of the three primary annuli give
rise to more complex annulation.
Table 1 . Theoretical table of the annuli produced
by repeated bisection of the three primary annuli
(after Moore, 1 898).
Primary Secondary Tertiary
61
c2
Quaternary
I rfl
' * *1 W»
j
t/4
d5
dG
tf7
48
c5
rflO
d16
rf18
I d2-\
I
-------
External Features
Chaetae—
The genus Acanthobdella consists of only two species, A. peledina and
A. livanowi, which are considered primitive leeches. In addition to
"their hirudinean characters, these species show their peculiarity by
possessing some oligochaete-like evolutionary traits. Externally, the
main anatomical character that separates the Acanthobdella from all other
leeches is the presence of chaetae (setae) in the first five anterior
somites (Fig. 3a).
The important diagnostic characters for £. peledina (Fig. 3) are the
irregularly spindle-shaped body, narrower anteriorly than posteriorly
(Fig. 3b,c), lacking an oral sucker and consisting of 30 somites
(segments). The last four somites are united to form a small caudal
sucker which has its concavity directed posteriorly. In the cephalic
region, the three pairs of eyes are small and located {one pair each) on
the 3rd, the 6th, and 10th annuli. The chaetae bundles, consisting of
two pairs (10 pairs in all) (Fig. 3a) in approximately equal distance are
located on the ventral surface (on the posterior margins) of the 1st, the
2nd, the 3rd, the 6th, and 10th annuli (counting posteriorly). The color
of live A^. peledina, as described in the literature, varies in specimens
from gray to red in appearance or dark greenish with yellowish spots
anteriorly. In some individuals the body can have 6-7 dark transverse
bands, and sometimes these bands occur over the entire body. Preserved
specimens often lose their color and become whitish in appearance. The
main diagnostic characters which separate A^. livanowi from A^. peledina
are the enlarged, expanded (orbicular) anterior somites of the cephalic
region, and the oral sucker like depression. The chaetae bundles in the
anterior somites of the ventral surface of /V. livanowi are also of
unequal distance in arrangement.
In addition to the sensillae, which are confined to the sensory
annuli, other visible organs of the integument are the cephalic eyes
(ocelli) and papillae (tubercles). Eyespots when present on the caudal
sucker and lateral margins (Fig. 10), which are usually embedded in
pigment, are also termed ocelli. Varying in number and shape
(punctiform, crescentiform), eyespots are used in the identification of
some species of piscicolids. They are absent in the Glossiphoniidae.
The number and position of the eyes of the cephalic (anterior) region are
important taxonomic features.
Cephalic Eyes—
The glossiphoniids have one to four pairs of eyes (Figs. 7-9), the
piscicolids, none to two pairs (Fig. 10,11), the erpobdellids> three or
four pairs (Fig. 13b-d), except Dina anoculata, which can have none or
three pairs (Sawyer, personal communication), and the hirudinids, five
pairs (Fig. 13a). As reported only by European authors, coalescence of
the eyes occurs occasionally in Gossiphom'a complanata, Alboglossiphonia
heteroclita, Batracobdella paludosa, and Boreobdella verrucata occurs
23
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(Fig. 9j-m). The relative distance between eyes is another important
diagnostic character in identification. If the distance between them is
equal to or greater then the diameter of the eyes, they are termed "well
separated" (Fig. 7). If they touch, they are termed "fused" or
"confluent" (Fig. 8g-n). If the distance between a pair of eyes is less
then the diameter of a single eye, they are termed "close together" (Fig.
8a-f). However, some variation in eye location can occur ocasionally, as
a result of rapid preservation.
The eyes are arranged in separate labial and buccal groups (Fig.
13b-d) in the erpobdellids, and the eyes are arranged in a submarginal
(parabolic) arch (Fig. 13a) in the hirudinids. If the leech has been
fixed and the eyes cannot be seen, the head of small specimens can be
flattened between two glass slides, which will generally make the eyes
visible. If the eyes are hidden by dark pigmentation, decolorize the
head of the preserved specimen by immersion in 5% caustic potash or
Amman's lactophenol (specimens in formalin work best) for 10 to 30
minutes, but the time will vary with the specimen. Amman's lactophenal
is prepared as follows: lOOg phenol, 100 ml lactic acid, 200 ml
glycerine, 100 mL water.
Papillae—
The papillae of the dorsal surface vary in size, type, and arrange-
ment. They may be limited to small, minute protrusible sense organs that
are often scattered in small and great numbers over the dorsal surface,
or are large (tubercle-like) smooth, conical, or rounded cone projections
that include some of the dermal tissue and muscles, and which are often
themselves covered with minute papillae. The segmental arrangement of
the papillae is an important characteristic used in distinguishing
between certain species of Helobdella and Placpbdella, and also
separating Glossiphom'a complanata from Boreobdella verrcuata. In £.
complanata, if the papillae are present, they lie on the 2nd (middle
ring) of the somite only (Fig. 57a). A reliable characteristic for IK
verrcuata is the large and distinct papillae on the 2nd and 3rd rings of
the somite (Fig. 57b). In Helobdella triserialis, the papillae are
small, smooth, and conical in three or less longitudinal rows. In H_.
papillate, the papillae are prominent and arranged in five to nine
longitudinal rows. Numerous large and scattered papillae are present on
the dorsum of Placobdella ornata, and the surface of these papillae is
covered with minute papillae which results in a very rough or warty
appearance. In £. multilineata the papillae are small, less numerous, on
the dorsum, but with the larger ones tending to be segmentally arranged
in five distinct, longitudinal rows. In P. papillifera, the dorsal
papillae occasionally vary from small to Tnconspicuous, or large and
pointed, and are arranged in five to seven distinct longitudinal rows.
However, in specimens of the above species, variation occurs in these
chararacters, and they must be used with other features such as shape and
pigmentation, as indicated in the key.
24
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Cephalic Region--
The form of the cephalic region {anterior somites) of leeches is a
diagnostic character. In the Glossiphoniidae, the cephalic region of all
the species (except P_. montifera and P. nuchalis) is not, or is only
slightly, differentiated (Figs. 7a-f,B,9T^The mouth pore is located on
the rim or within the oral sucker (Figs. 4a,b,d,}. Both P_. montifera and
P. nuchalis have the cephalic regions in the form of a distinct "discoid"
Fead (Fig. 7g,h). The anterior sucker of the piscicolids is always
expanded and usually distinctly marked off from the bocly, and the mouth
pore is in the middle of the sucker (Figs. 4c,10,ll). In Erpobdellidae
and Hirudinidae, the mouth is medium to large in size and the anterior
sucker is little more than the expanded lips of the mouth opening (Fig.
4e-h).
Caudal Region--
The form of the caudal (posterior) sucker is characteristic of some
leech species, and is included in the key as a diagnostic feature.
Species of Actinobdella possess digitate processes (glands and papillae)
on the rim of the caudal sucker (Figs. 33,34), and the hemispherical
sucker is separated from the body on a short, narrow pedicel (peduncle).
In preserved specimens, the digitate processes are usually retracted.
They are finger-like when everted (Fig. 33a,b). The position of the
digitate processes is indicated on the dorsal surface of the caudal
sucker by faint radiating ridges (Figs. 33,34). In Batracobdella
cryptobranchii and Placobdella pediculata, the digitate processes are
absent, but the posterior sucker is separated from the body on a narrow
pedicel (Figs. 31,32). Other minor characters not discussed here will be
found appropriately in the key.
Body Regions—
In all the Piscicolidae, with the exception of some specimens of
Pisci'colaria reducta and Myzobdella lugubris, the body is
characteristically divided into two distinct regions (Figs. 10,11); a
narrow anterior trachelosome region and a larger and wider posterior
urosome region. However, the separation of these two regions is less
conspicuous and not always observible in preserved specimens of
Piscicola, but they are always present in Cystobranchus. Some preserved
specimens of Myzobdella and Piscicol aria also show a distinct separation
of these two regions due to contraction (Fig. llc-e).
Pusatile Vesicles—
In the genera Piscicola and Cystobranchus only, the neural annuli of
the urosome bear 11 pairs of pulsatile vesicle (Fig. 10). In Piscicol a
the vesicles are small or usually undifferentiated in preserved specimens
(Fig. 10f,g,h,i), but in Cystobranchus, the pulsatile vesicles are large
and clearly visible in both living and preserved individuals (Fig.
10a,c,d,e).
25
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Pigmentation--
Pigmentation patterns and color are important diagnostic features for
some species of leeches. Coloration features, such as blotches, dots,
spots, stripes, bands, lines are easy to determine in living as well as
freshly preserved specimens, and usually persist for sometime. They may
fade away in some specimens which have been preserved in alcohol or
preserved for a considerable length of time. Green and blue pigments are
lost almost immediately in alcohol, although they usually persist longer
in formalin. The red color of blood is lost almost as soon as the green
and blue, and the other reds and yellows are lost more slowly. The
browns and blacks usually remain for years. It should be noted, however,
that in some species the colors of the dorsal surface change with the
development, age, and environmental conditions. Also, the pigmentation
pattern of the dorsal surface in several species is a poor diagnoistic
character because several color forms exist.
Size--
The average size of each species is given in the key, but many
variations appear in the species due to the age and development of the
specimen collected and also due to contraction during processing.
Finally, it should be noted again that most specimens of leeches can
be identified to the species level. The key will, however, enable the
user to identify these difficult forms to the lowest taxonomic level of
which diagnostic characters of the specimen are available to a particular
taxa.
Internal Features
Digestive Tract--
The alimentary canal is a tube from mouth to anus and Is divided into
the buccal chamber, pharynx, esophagus, stomach to crop, intestine with
or without diverticula, and rectum. In the Glossiphoniidae and
Piscicolidae, the mouth is a small pore on the rim or in the center of
the anterior sucker (Fig. 4a-d). The pharynx of the Rhynchobdellida is
muscular and protrusible through the mouth as a proboscis (Fig. 4d).
The proboscis has a crown-like tip, and is adapted to penetrate both
invertebrate and some vertebrate tissues. In Hirudinidae and
Erpobdellidae, the mouth is medium to large in size (Fig. 4e-h) and
occupies the entire cavity of the anterior sucker. In the hirudinids,
the buccal cavity, which may or may not contain jaws with denticles, is
separated from the cavity of the sucker by a flap of skin called the
velum (Fig. 85). The shape and form of the anterior sucker, the absence
of teeth, the number of internal ridges (fleshy pods) of the pharynx, and
the presence or absence of papillae on the velum are used as diagnostic
features to separate Haemppfs pi umbea and Haemppi's grandis. The velum is
finely papillated in H. piumbea and smooth in H. grandis. In H. piumbea,
the lip of the oral sucker is broad, thick, flat, and rounded, and the
aperature is transverse and lower margin anteriorly convexed (Fig. 88).
26
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The lip of the anterior sucker of H_. grand is is narrow, thin and arched,
and the aperature is elongated (Fig. 89).All other species of
Hirudinidae have three muscular jaws, two ventrolateral and one
dorsomedial (Fig. 85a-d). The free edge of each bears teeth arranged in
either one (monostichodont) or in two (distichodont) rows. Moore (1952)
indicated that some or all of the teeth in old or poorly preserved
specimens of H_. marmorata can fall off when the cuticle of the preserved
leeches separate"?!Therefore, separating H_. marmorata from H_. grandis
only on the basis of the presence or absence of teeth can be difficult,
perhaps impossible in some specimens. To examine the velum and jaws
(Fig. 85), the specimen should be positioned ventrally or pinned out in
such a manner. A medial incission 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 buccal cavity and pharynx. Details of
the denticles (Fig. 85a,b,d) can only be seen by first removing a jaw and
making a whole mount, using CMCS, Hydramount or other mounting media on a
microscope slide, and examining with a compound microscope to determine
the number of teeth.
Reproductive System
External Reproductive Features--
The male and female gonopores are visible on the middle of the
ventral surface of somites XI and XII and are generally separated by two
to seven annuli. Alboglossiphonia heteroclita and Marvinmeyeria lucida
are two unusual species in tnat the male and female ducts open into a
single gonopore. The male gonopore is large, more readily visible in the
mature specimen, and anterior to the female gonopore. In some specimens
the female pore can be difficult to find due to its small size. It is
seen most easily immediately after narcotization, and its position often
being revealed by a small opening or some color difference which may be
lost if the specimen is not properly preserved.
Another important characteristic used in determining species of
erpobdellids is the number of annuli between the male and female
gonopores (Fig. 92). If it is necessary to make evident the female
opening, the following procedure has sometimes proven successful. The
leeches are narcotized, fixed in 10% formalin, and preserved in formalin
or alcohol. The leech is then placed in creosote-beechwood or Amman's
lactophenol (clearing agents) for 15 to 30 minutes, depending on the size
of the leech (the larger the leech the longer the time), until the female
gonopore begins to stand out. Amman's lactophenol works best with
specimens preserved in formalin. Gentle application of heat will speed
up the process. The area of the gonopores should periodically be
observed under a stereomicroscope, for if the specimens are left too long
in the clearing agent, desiccation and wrinkling ensues. Sawyer (1972)
indicated that slight variations in the location and number of annuli
between gonopores exist in some specimens of Hirudinidae and
Erpobdellidae. Meyer and Moore (1954), Sawyer (1972), and Klemm (1977)
stated that some variation occurs in species of Theromyzon of
Glossiphoniidae. Therefore, some specimens of these taxa can only be
27
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determined to the generic level, (cf. Notes on Systematics).
External copulatory glands with pores are present in a linear or
transverse pattern of either 4, 6, 8, or 28 (Fig. 74a-d) on adult
Macrobdella decora, fl. diplotertia, M. ditetra, and fl. sestertia.
Immature forms may not have a set number of copulatory pores (Sawyer and
Pass, 1972), but other characters in the key can be used for
identification. The copulatory glands are located on the ventral
surface, four to five annuli posterior to the female gonopore. In some
immature specimens the copulatory glands and pores are absent and other
diagonstic characters are indicated whenever possible in this key for
identification. In Philobdella, the gonopores and copulatory pits are
surrounded by an undifferiated glandular area (Fig. 74e).
Internal Reproductive Features--
Leeches are hermaphroditic. The female reproductive system is
comprised of ovisacs, terminating in ducts which join to form a common
duct or vagina. The male reproductive system consists of testisacs in
metameric patterns, five to ten in the Glossiphoniidae and Piscicolidiae
and nine to ten in the American Hirudinidae. But in the Erpobdellidae,
they are small, numerous and arranged in grape-like clusters. The vasa
deferentia connect the testisacs to the vasa deferens on each side.
These ducts lead into the seminal vesicles and ejaculatory ducts, which
open into the atrium. The shape of the atrium is diagonistic for some
species of Erpobdellidae (Figs. 93,103). This organ is a medium chamber
and consists of three parts: a thin-walled eversible bursa, a thick
walled glandular chamber, and a muscular medium chamber, as well as a
pair of lateral horns (atrial cornua), which receive the ejaculatory
ducts. In Nephelopsis obscura, the atrial cornua is spirally arranged
like a ram's horn (Fig. 103a,b). In the Mooreobdella and Dina-Erpobdella
complex, the atrium is simply curved, globular, rounded, ellipsoidal, or
short and curved (Figs. 93,103c,d). Some species also have ejaculatory
ducts with or without long preatrial loops (Figs. 93,103).
To examine the male reproductive structures (Fig. 2a-c), which may
be necessary in the identification of some species of Erpobdellidae, the
following procedures should be followed: (1) specimens fixed in formalin
should be used when dissection is necessary, (2) position or pin out the
preserved specimen with the dorsal surface up, (3) make a transverse
incision across the boc(y 4 or 5 annuli posterior to the male gonopore,
(4) cuts anteriorly up the lateral margins of the body for about 15
annuli, and (5) the posterior edge of the flap thus made can now be
lifted forward or removed to expose the inner tissue which can be
carefully cleared away to fully expose the atrium and ejaculatory ducts.
Another method (Fig. 2d) involves positioning or pinning the
specimen ventral side up and make a rectangular cut to remove a portion
of the body wall, beginning about 4 to 5 annuli in front of the male pore
and including 3 annuli behind the female pore. Once this portion is
removed, the connective tissue surrounding the male reproductive
structures should be carefully teased away, thus exposing the genital
28
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atrium and ejaculatory ducts.
To accurately Identify small forms of the family Erpobdellidae, a
technique which sometimes works is the following: (1) fix the leech in
formalin, (2) stain and destain, (3) run through an alcohol series, (4)
clear and mount as described earlier, to determine the size and shape of
the atrium, atria! horns, and length of the vasa deferentia (ejaculatory
ducts}.
FIG, 2. Diagrams of methods to show male reproductive system of
Erpobdellidae: (a) dash lines indicate cut on dorsal surface, (b,c)
dorsal portion of annuli removed and connective tissure teased away
displaying the reproductive structures; (d) dash lines indicate cut on
ventral surface.
29
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-------
SECTION 6
KEY TO THE LEECHES (HIRUDINEA) OF NORTH AMERICA, NORTH OF MEXICO
Chaetae present in cephalic region of anterior somites (Fig. 3a);
body shape, cylindrical, spindle-like (Fig. 3b,c); length 10-22
mm. Order Acanthobdellida, Family Acanthobdellidae
Acanthobdella peledina Grube, 1851
Chaetae absent from cephalic region of anterior somites; body
shape various 2
chaetae
FIG. 3. Acanthobdellidae: (a) cephalic region, ventral view; (bsc)
general bocjy shape (cf. Notes on Identification).
2(1) Mouth a small pore on rim or within oral sucker cavity from which
a pharyngeal proboscis can be protruded (Fig. 4a-d); jaws
absent; rarely swims. Order Rhynchobdellida 3
Mouth opening medium to large, occupying entire oral sucker
cavity, forming rounded lips, not pore-like (Fig. 4e-h);
lacking a protrusible proboscis; jaws and teeth either present
or absent; good swimmers 4
31
-------
- mouth pore
oral sucker
mouth
oral sucker
proboscis
mouth pore
oral sucker
mouth
- mouth
mouth
f
FIG. 4. Ventral views of anterior (oral) suckers to show mouth and
sucker differences: (a) pore on rim of sucker; (b) pore within sucker;
(c) pore near center of sucker; (d) SEM of protruding muscular proboscis
cavity of Helobdella triserial is (X 250); (e-h) mouth occupying entire
sucker cavity.
32
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3(2) Body at rest flattened dorsoventrally, posterior half usually much
wider than tapering cephalic end (Figs. 5,6), never cylindrical
(except Helobdella elongata which is very sub-cylindrical,
terete, Fig. 5c); not differentiated into two bocjy regions;
oral sucker ventral and more or less confluent with cephalic
region (except Placobdella montifera, P. nuchal is, (Figs.6d,e;
7g,h); eyes 1, 2, 3, or 4 pairs (Figs.~7-9); eyespots (ocelli)
never on caudal sucker or lateral margins of body; 3 annuli per
complete somite (except Oligobdella biannulata which is
2-annulate); free-living or predaceous, parasitic on
invertebrates (insects, oligochaetes, snails, and so forth) and
vertebrates (crocodilians, turtles, fishes, or waterfowl);
young always attached to ventral surface of parent; eggs in
membraneous sacs either attached to ventral surface of parent
or to substrates and covered by parent's body. Family
Glossiphoniidae 5
Body at rest cylindrical, narrow, (Figs. 10,11), posterior half
can be slightly flattened; may be divided (especially in con-
traction) into a narrow neck (trachelosome) and wider body
(urosome) regions (Fig. 10,11); oral sucker expanded, distinct
from neck (Figs. 10,11) eyes 0, 1, or 2 pairs (Figs. 10,11); 7
or more annuli per complete somite (except Piscicolaria reducta
which is 3-annulate); with or without eyespots (ocelli) on
caudal sucker and lateral margins of urosome (Fig. 10); with or
without lateral pulsatile vesicles (Figs. 10,11); young never
attached to ventral surface of parents; cocoons attached to
substrates, young never brooded; rarely found free-living,
usually parasitic on fishes (Fig. 12). Family Piscicolidae. .
34
FIG. 5. General body shapes of Glossiphoniidae in dorsal view: (a)
ovate-lanceolate; (b) lanceolate; (c) subcylindrical (terete); (d,e)
various shapes.
33
-------
FIG. 6. Glossiphonbiidae, general body shapes
ventral view.
(a) dorsal view; (b-e)
9 *
f
discoid
head \^
FIG. 7. Glossiphoniidae, dorsal views of eyes: (a-f) eyes well
separated, cepahlic region undifferentiated; (g,h) eyes well separated,
cepahlic region differentiated "discoid head."
34
-------
g
I
m
n
FIG. 8 Glossiphonildae, dorsal view of eyes: (a-f) close together, less
than diameter of one eye; (g-n) eyes touching, confluent (fused).
35
-------
„, eyes
accessory
eyes
L \ ±
1—
f
S
•
m
FIG. 9. Glossiphoniidae, dorsal view of eyes: (a) fused eyes with
accessory eyes, Placobdella hollensls; (b) Batracobdella paludosa; (c,d)
A1bog1ossiphpm'a~heteroclfta; te-g) Glosslp'npm'a complanata, Boreobdeila
yerrcuata; (h,i) TherOinyzon~sp. (j-m) eyes showing variation
(coalescence) in eye position.
36
-------
pulsatile
pi-gmented
ray
sucker
proboscis
eyes
S
FIG. 10. General body shapes of Piscicolidae: (a-k) dorsal view showing
body regions, ocelli arrangements, and external structures.
37
-------
f
FIG. 11. General body shapes of PiscicoHdae: (a-d) Myzobdella
lugubris, dorsal view; (e,f) Pisclcolan'a reducta, dorsal view.
38
-------
Cystobranchus
\
Placobdella pediculata
a
Myzobdella lugubris
Myzobdella lugubr-is
P-isc-icolapia reducta
FIG. 12. Some attachment sites of fish leeches: (a) Placobdella
pediculata attached under the operculum of the freshwater drum
(Aplodlnotus grunniens); (b) Cystobranchus verrllll attached to the gill
arch and body of a flathead catfish (Pylodictis o]iva|"js); (c) Myzobdella
lugubris attached to the chin of the channel catfish tlctalurus
unctatus); (d) Myzobdella lugubris attached to the pectoral fin of a
lue gill (Lepomjs macrochirusj; (e) Piscicolaria reducta attached to the
caudal fin of a logperch (Percina caprodes).Part of the operculum and
gill (a) was removed to show attachment location. Illustrations (b-e)
after Poe, 1972.
39
-------
4(2) Eyes, 5 pairs, forming a parabolic arch, somites II-VI (Fig. 13a);
body large, linear, elongate (Fig. 14); jaws with teeth (except
none in Haemopis grandis, Haemopis plumbea); muscular pharynx,
short, not extending to clitellum, testisacs large, arranged in
metameric pairs; free-living or predaceous and blood-suckfng.
Order Gnathobdellida, Family Hirudinidae 43
Eyes, 0, 3, or 4 pairs in separate labial and buccal groups
(Fig. 13b-d ); body moderate size, linear, elongate (Fig. 14);
lacking jaws or teeth; mouth armed with muscular pharyngeal
ridges; pharynx extending to somite XIII, about 1/3 bocty
length; testisacs small and numerous in grape-like clusters;
free-living or predaceous. Order Pharyngobdellida, Family
Erpobdellidae. . . 1 56
FIG. 13. Arrangement of eyes, dorsal view: (a) Hirudinldae: (b-d)
Erpobdellidae.
40
-------
s
w
d
FIG. 14. General body shapes: (a-e) H1rud1nidae; (f-j) small
hirudinlds; (f-j) Erpobdellldae.
41
-------
5(3} One pair of eyes {Figs. 7,8), except Placobdella hollensis, which
also has a series of paired accessory eyes (Fig. 9a). . ... .6
More than one pair of eyes (F1g. 9b-i) 28
6(5} Anterior somites (oral sucker) of cephalic region distinctly
expanded to form a discoid head, set off from body by a narrow
neck constriction (Figs. 6d,e;7g,h) 7
Anterior somites (oral sucker) of cephalic region
undifferentiated, more or less continuous with body (Figs.
5,6a-c»7a-f,8,9), not distinctly expanded to form a discoid
head, no narrow neck constriction 8
7(6) Dorsum with 3 prominent pointed (tuberculate) keels or ridges (may
not be decernible in live specimens); eyes one pair, separated
by their diameter; color greenish-gray or pale olive-brown;
free-living, parasitic on fish; length 9-16 mm (Fig. 15). . . .
Placobdella montifera Moore, 1906
Dorsum smooth; eyes one pair, separated by their diameter; color
greenish-gray; free-living, parasitic on fish; length 15-25 mm
(Fig. 16) Placobdella nuchal is Sawyer & Shelley, 1976
15
d-iscoi-d head
16
FIG. 15. Placobdella montifera: (a) normal resting shape; (b)
contracted shape. FIG. 16. PTacobdel1 a nuchalis.
42
-------
8(6)
9(8)
With a series of accessory (supplementary) eyes behind single pair
of functional eyes in cephalic region (Fig. 9a); color light
olive-green variegated with brown, pale yellow or colorless
areas; adults often swim; feeding habits unknown; length 20-30
mm (Fig. 17) Placobdella hollensis (Whitman, 1892)
Without accessory eyes 9
Eyes distinctly separated, usually by the diameter, or more of
one eye (Fig. 7a-f)
10
Eyes close together, separated by less than the diameter of one
eye, or confluent (fused), touching, (Fig. 8)
,16
10(9) Dorsum unpigmented, uniformly pigmented, or pigmented with
longitudinal or transverse bands, lines or stripes and with or
without metameric whitish spots (sensillae) on every 3rd
annulus in middle of body region, or dorsum with scattered
chromatophores (Figs. 21-27); heavily to sparsely or not
papillated, or with a chitinous scute (nuchal plate) in
anterior region; crop caeca 1-6 pairs; gonopores separated by
at least 1 annulus; free-living, parasitic, predaceous on
invertebrates. Helobdella 11
Dorsal and ventral surfaces heavily pigmented with uniform,
minute, blackish chromatophores, with thin dark paramedial
lines extending into anterior region; dorsal surface smooth, no
papillae or scute; crop caeca 6 pairs, gonopores united; length
15-22 mm (Fig. 18) Marvinmeyeria lucida (Moore, 1954)
11(10) With a dark brown chitinous scute, can be faint in juveniles (or
may fall off in individuals preserved for a long time) on
somite VIII of dorsum in anterior region (Fig. 19); color dusky
brown, gray, green, or pink, length 9-14 mm (Fig. 20)
Helobdella stagnalis (Linnaeus, 1758)
Without a chitinous scute in the anterior region of dorsum. . . .12
12(11) Dorsum without papillae (tubercles), smooth 13
Dorsum papitlated (few, scattered, or arranged in 3 to 9
longitudinal rows) 15
43
-------
17
eyes
accessory
eyes
19
20
scute (nuchal
plate)
FIG. 17. Placobdella hollensls. FIG. 18. Marylnmeyerla luclda. FIG.
19. He!obdel1 a stagnalis, photomicrograph showing protruding proboscis
and scute (X TOO). FIG. 20. Helobdella stagnalIs.
44
-------
13(12) Body plgmented, with or without longitudinal or transverse bands,
lines and/or stripes or uniform; body flat, posterior wider
than tapering cephalic end; usually 6 pairs of crop caeca. . .
14
Body unpigmented, elongate and subcylindrical (terete) (Fig. 5c);
lateral margins of body almost parallel, body smoothly rounded;
posterior sucker small, terminal, translucent, sometimes opaque
white or gray; 1 pair of crop caeca; length 9-25 mm (Fig. 21).
Helobdella elongate. (Castle, 1900)
14(13) Dorsum with transverse rusty-brown, interrupted bands alter-
nating with irregular whitish bands, the latter consisting of 8
to 10 confluent white metameric spots on each neural annul us;
length 10 mm (Fig. 22). . . . Helobdella transversa Sawyer, 1972
Dorsum without .transverse pigmentation; color uniform coffee-
brown or with longitudinal whitish stripes alternating with
coffee-brown stripes and/or lines; with or without small
whitish or pale spots in anal region, or dorsum with scattered
chromatophores; occasionally preserved individuals show minute
sensillae on annuli of dorsum; a variable species; length 10-14
mm (Fig. 23a-e) Helobdella fusca (Castle, 1900)
15(12) Dorsum rough, with 5 to 7 or 9 longitudinal rows of large,
whitish, rounded conspicuous papillae, on each neural annulus;
dorsum yellowish-brown or unpigmented; length 9-14 mm (Fig.
24) Helobdella papillata (Moore, 1906)
Dorsum with 3 rows (typically) or fewer of small, black-tipped or
uniformly pale white papillae; pigmentation uniform, or
arranged in numerous longitudinal light and dark brown, gray,
or black lines and/or stripes, or dorsum with 3 broad,
longitudinal white stripes, one median row and one on each side
submarginal, also at posterior end 2 short white stripes, few
or many whitish spots on neural annuli, unaligned or if
aligned, then confined to areas lateral to papillae, or color
uniform with papillae few, scattered, or in 3 rows; whitish
spots on neural annuli unaligned or aligned; a variable
species; length 10-29 mm (Figs. 25,26,27)
Helobdella trisen'alls (E. Blanchard, 1849)
45
-------
21
FIG. 21. Helobdella elongata. FIG. 22. Helobdella transversa. FIG.
23. Helobdella fusca"(a) typical pigmented form; (b) non-pigmented
form; (c) scattered pigmented form; (d,e) white blotched pigmented form.
46
-------
FIG. 24. Helpbdella papillata. FIG. 25. Helobdella tnsenalis: (a,b)
showing variability of [Dapmae and pigmentation. FIG. 26. Helobdella
triserial is: (a-c) showing variability of papillae and pigmentation.
47
-------
FIG. 27. Helobdella triserial is: (a,b) showing variability of papillae
and pigmentation.
16(9) Dorsum with conspicuous white genital and anal patches, one or
more medial white patches and a white ring (bar) in neck region
(Figs. 28,29,30) 17
Dorsum without white patches and white ring (bar) in neck region
19
48
-------
FIG. 28. BatracobdeTIa phalera: (a-d) variable forms; (e) Immature form,
49
-------
17(16) Dorsum smooth, no papillae (tubercles) 18
Dorsum with three series of usually dark-tipped papillae or white
pointed papillae, sometimes with a thick ban encomposing
middorsal series of papillae and marginal dots; with or without
white patches approximately between genital and anal patches;
body convex, thick, solidly or lightly pigmented, opaque;
length 6-10 mm (Fig. 28) free-living or parasitic on fish. . .
Batracobdella phalera (Graf, 1899)
18(17) Body ovate-lanceolate, excessively flattened, thin, with 5
(sometimes raised) longitudinal rows of white prominences
surrounded by yellowish dots, equidistant longitudinally and
transversely; length 6-10 mm (Fig. 29)
Batracobdella michiganensis Sawyer, 1972
Body lanceolate, excessively flattened, thin, lightly pigmented
with small brown, cutaneous chromatophores; or pigmented with a
broken longitudinal median stripe and two rows of submarginal
spots; length 8-11 mm (Fig. 30)
Placobdella trans!ucens Sawyer & Shelley, 1976
30
FIG. 29. Batracobdella michiganensis. FIG. 30.
trans!ucensl(a,b) variable pigmented forms.
Placobdella
50
-------
19(16) Caudal sucker well developed, rim moderately thick, bulbous,
separated from body on a distinct pedicel (peduncle), (Figs.
31-34) 20
Caudal sucker well developed, rim thin, continuous with body, not
separated on a pedicel 23
20(19) With marginal circle of 30-60 retractile digitate processes on
rim of caudal sucker, when everted, projecting (finger!ike)
along inner margin of caudal sucker cavity, usually retracted
in preserved specimens, position marked dorsally on outer rim
by faint radiating ridges, may appear as whitish radiating
bands (Figs. 33,34). Actinobdella 21
Without digitate processes on caudal sucker 22
31
32
pedicel-
digitate
processes
pedicel
33
retracted
digitate
processes
c
v
digitate
processes
FIG. 31. Caudal sucker and slender stalk (pedicel) of Placobdella
pediculata: (a) lataeral view; (b) ventral view. FIG. 32.Caudal
sucker, short pedicel, of Batracobdel1 a cryptobranchii, lateral view.
FIG. 33. (a-c) Actinobdella I'nequiannulata, various views of caudal
sucker showing digitate processes.FIG. 34. Actianobdella annectens,
dorsal view of caudal sucker showing digitate processes.
51
-------
21(20) Caudal sucker with about 30 digitate processes on rim, short
pedicel (Fig. 33); bocjy shape slender, round, and elongate to
thick, and strongly convex dorsally In engorged adults; dorsal
papillae, 1-5 series, acute, median, variable, or absent (In
some Individuals the papillae are replaced by a longitudinal
mlddorsal ridge (Fig. 35) or groove; usually parasitic on fish;
body somites 3-annulate, often with Incipient secondary
annulation; free-living, parasitic on fish; length 7-22 mm
(Figs. 36,37) Actinobdella inequiannulata Moore, 1901
Caudal sucker with about 60 digitate processes on rim, short
pedicel (Fig. 34); otherwise similar to £. inequiannulata (has
not been found since original description), may be a synonym of
A. inequiannulata); length 7-11 mm (Fig. 38)
Actinobdella annectens Moore, 1906
35
36
37
38
FIG. 35. Actinobdella inequiannulata, dorsal view showing middorsal
ridge. FIG. 36.Actinobdella InequTannulata, ventral view of engorged
adult. FIG. 37. Actlnobdella 1nequiannulata, dorsal view with minute
papillae. FIG. 381Actinobdella annectens, dorsal view with papillae.
52
-------
22(20) Body high dorsally and contractile; dorsum smooth; caudal sucker
set off from body by a slender stalk (pedicel) (Fig. 31),
absent or short in juveniles smaller than 1 cm (Fig. 39); bocjy
color brownish or grayish, opaque; usually encountered on
gills, attached to the isthmus or inside of the opercula (gill
chamber) of freshwater drum (Aplodinotus grunniens); length
20-35 mm (Figs. 40,41)
Placobdella pediculata Hemingway, 1908
Body moderately flattened; dorsum smooth; caudal sucker set off
from boc(y by a short pedicel (Fig. 32); bocty color of preserved
specimens cream to golden, live individuals reddish brown due
to blood in crop, usually translucent; (some preserved
specimens have 8 rows of inconspicuous sensillae on annuli of
the dorsum); known only from the Ozark Hellbender
(Cryptobranchus allegam'ensis bishopi); length, up to 17 mm
(Fig. 42). . ,
Batracobdella cryptobranchii Johnson & Klemm, 1977
39
FIG. 39. Placobdella pediculata, dorsal view of juvenile without slender
stalk (pedicel).FIE. 40.Placobdella pediculata. lateral view. FIG.
41. Placobdella pediculata, dorsal view of contracted adult. FIG. 42.
Batracobdella cryptobranchTi, dorsal view.
53
-------
23(19) Dorsum not roughly papillated, usually smooth, with low smooth
domes, often suppressed papillae, or without papillae
(tubercles) 24
Dorsum roughly papillated 26
24(23) Ventral surface not striped 25
Ventral surface with 8-12 bluish, greenish, or brownish longi-
tudinal stripes or lines (Fig. 43); dorsal pigment pattern
variable and intricate, usually dark greenish-brown, with a
middorsal cream colored stripe or band of variable width and
with irregular lateral patches; body somites 3-annulate;
usually parasitic on turtles, often found free-living; length
38-64 mm (Figs. 44,45)
Placobdella parasitica (Say, 1824)
FIG. 43. Placpbdella parasitica, ventral view showing striping. FIG,
44. Placobdella parasitica:Ta,b) dorsal view showing variable
pigmentation patterns.
54
-------
IG. 45. Placobdella parasitica: (a-c) dorsal view showing variable
pigmentation patterns.
25(24) Color of dorsum dark greenish-brown, finely variegated with
orange, with a thin dark median line (can be absent in
preserved individuals), with 4-5 rows of yellowish or whitish
metameric dots; boc(y somites 3-annulate; caudal sucker small;
commonly encountered on mating frogs, toads, salamanders, and
their larvae; often found free-living; length 13-25 mm (Fig.
46) Batracobdella picta (Verrill, 1872)
Color of dorsum pale olive-green, brown, or gray; boc(y somites,
2-annulate; caudal sucker large, conspicuous; known from the
mountains of North and South Carolinas; parasitic on
desmongnathine salamanders; length 2-7 mm (Fig. 47)
Oligobdella biannulata (Moore, 1900)
55
-------
S-annuiate
2-annulate
FIG. 46, Batracobdella pi'cta: (a) form with dark median line; (b) form
lacking median line.FIG. 47. Oligobdella biannulata.
26(23) Dorsum warty, entirely covered with rounded papillae, varying in
size, randomly arranged, or with the larger papillae in 5
longitudinal metameric rows and numerous other papillae
randomly arranged of varying size; all papillae bearing
clusters of secondary, minute papillae at the apex; usually
with a brown band down middorsal line, interrupted 4-5 times,
sometimes with a light colored band; ventral surface unstriped,
with scattered dark chromatophores or plain (Fig. 48); body
usually flat, lanceolate; color of dorsum a mixture of brown,
green, and yellow, but variable; free-living or parasitic on
turtles; length, up to 40 mm (Fig. 49)
Placobdella ornata (Verrill, 1872)
Dorsum less rough, not entirely covered with papillae, with large
and/or small more uniform simple pointed papillae, the larger
ones in 5-7 longitudinal metameric rows, some bearing only one
or a few secondary, minute papillae at the apex 27
56
-------
FIG. 48. Placobdella ornata, ventral view. FIG. 49. Placobdella
ornata: (a,b) dorsal view snowing variability of papillae and
pigmentation.
27(26) Body usually ovate-lanceolate, strongly convex; dorsum with 5-7
longitudinal rows of large white pointed papillae; some bearing
only one or a few secondary minute papillae at the apex; other
small papillae varying in size to inconspicuous, not forming
distinct rows; papillae on caudal sucker; dorsum with a vague
narrow, continuous (sometimes interrupted) median longitudinal
stripe, contained in a wider bluish stripe, encompassing a
middorsal row of papillae; light colored stripes on each side
of the middorsal row of papillae joining in neck region; color
of dorsum greenish-blue with longitudinal striping or color
above obscure yellowish brown, produced by alternating narrow
lines of flesh-color and olive-green (giving a checkered
appearance); base color usually brown in preserved individuals;
ventral surface usually with 2-8 bluish longitudinal stripes,
lines (Fig. 50); without scattered dark chromatophores;
free-living, parasitic on turtles; length 15mm or larger (Figs.
51,52) Placobdella papillifera (Verrill, 1872)
Body usually lanceolate, flat; dorsum with few or numerous,
uniform, small papillae, with the larger ones in 5 longitudinal
rows of varying degree; middorsal stripe narrow, dark, usually
continuous; color of dorsum usually with brownish longitudinal
lines, stripes; ventral surface plain or with dark colorerd
chromatophores in vague longitudinal lines; free-living or
parasitic on turtles and alligators; length 40-50 mm (Fig. 53).
Placobdella multilineata Moore, 1953
57
-------
FIG. 50. Placobdella papillifera: (a,b) ventral view showing striping,
FIG. 51. PIacobdelTa papi 11 iTera'. FIG. 52. PlacobdelJa papillifera
(From Sawyer and Shelley, 1976). FIG. 53. Placobdella multi1ineata:
(a,b) dorsal view showing variability in papillae and pigmentation.
58
-------
28(5)
Eyes 2 pairs (Fig. 9b), arrangement modified sometimes by
coalescence of eyes in various ways (Fig. 9j-m); dorsum
smooth, usually with two longitudinal interrupted lines, but no
large pigment spots; color variable, green or brownish; one
dubious record, may not be established in North America, length
7-20 mm (Fig. 54) Batracobdella paludosa (Carena, 1924)
Eyes 3 or 4 pairs, may be coalesced (Fig. 9c-i)
29(28) With 3 pairs of eyes, may be coalesced of eyes in various ways
(Fig. 9c-g). Alboglossiphom'a, Glossiphonia, and Boreobdella.
With 4 pairs of eyes (Fig. 9h-i ). Theromyzon. .
30(29) Eyes equidistant in 2 paramedian rows (Fig. 9e-g)
29
30
32
31
Eyes in an approximately triangular pattern, the 1st pair always
closer than the posterior pairs (Fig. 9c,d); body smooth,
pigmentation slight, brownish-black chromatophores in sparse
clusters, and often with a dark median, longitudinal stripe
(sometimes interrupted) on dorsum but without paired stripes;
or with small 4 to 7 middorsal irregular, transverse bars,
composed of brownish-black pigmentation; body translucent;
free-living, parasitic on mollusks, invertebrates; length 6-9
mm (Fig. 55,56)
Alboglossiphom'a heteroclita (Linneaus, 1761
55
FIG. 54. Batracobdella paludosa. FIG. 55. Alboglossiphom'a
heteroclita": (a,b) variable pigmented forms.
59
-------
FIG. 56. Alboglossiphonia heteroclita: (a-c) variable plgmented forms.
31(30) Dorsum rarely with papillae on 2nd (middle) annulus of the
triannulate somite, usually smooth (Fig. 57a); with a pair of
dark paramedlal stripes, dorsally and ventrally (can be absent
in young), interrupted by a pair of dorsal metameric white,
yellow spots (which may be slightly raised in preserved
specimens), paramedially and marginally; body opaque, color
brown, green, or gray; 6 or 7 pairs of crop caeca; free-living,
predaceous on invertebrates; length, up to 25 mm (Fig. 58). . .
Glossiphonia complanata (Linnaeus, 1758)
Dorsum always with large and distinct papillae at median of the
2nd and 3rd annuli of the triannulate somite (Fig. 57b), with
several smaller papillae towards the margins; with large size
and varying number of pale whitish spots (some irregularly
shaped) which reduce the dark brown-gray ground color almost to
a reticulum; on the dorsal side, on a level with the genital
pores of the ventral side, two large, irregular blotches; with
a pair of heavy dark paramedian stripes, dorsally and
ventrally, interrupted by papillae dorsally; bo(|y opaque, color
brown, green or gray, spotted with yellow; 7 pair of crop
caeca; length, up to 25 mm (Fig. 59)
Boreobdella verrucata (Fr. Muller, 1884)
60
-------
57
a.2 somite
O o) a2 somite
aS /
somite
o o) a.2 somite
FIG. 57. Schematic view of two nrfddorsal segments: (a) Glosslphom'a
complanata; (b) Boreobdella yerrucata. FIG. 58. Glossiphom'a
comjjlanafa'. FIG, 59. Boreobdella verrucata.
32(29} Gonopores separated by 2 annuli; body shape variable, gelatious,
nearly translucent; color olive-green, flecked with black
chromatophores, but variable, or with longitudinal rows of
cream yellow spots on dorsum; distributed in central and
eastern United States and Canada; free-living or parasitic
(esp. within mucosa of nasal chamber and the conjunctiva of the
eyes) of waterfowl; length 20-26 mm (Fig. 60)
Theromyzon biannulatum Klemm, 1977
Gonopores separated by 3 or 4 annuli
61
33
-------
FIG. 60. Theromyzon biannulatum or Theromyzon rude:
pigmentation.
(a,b) variations in
33(32) With 3 annuli between gonopores; body shape variable, gel-
atinous, nearly translucent; color variable, with 2 paramedial
pairs and marginal pairs of yellow, orange, or brown spots on
dorsum; distributed in central and western United States and
Canada; free-living or parasitic (esp. within mucosa of nasal
chamber and conjunctiva of the eyes) of waterfowl; length 20-30
mm (Fig. 60) Theromyzon rude (Baird, 1863)
With 4 annuli between gonopores; body shape variable,
gelatinous, nearly translucent, color amber or greenish, with 2
thin paramedian black lines on dorsum or with longtudinal rows
of cream colored, yellow spots, but variable; distributed in
Europe but has been reported from western United States and
Canada; free-living or parasitic (esp. within mucosa of nasal
chamber and conjunctiva of the eyes) of waterfowl; length 15-30
mm (Fig. 61) Theromyzon tessulatum (O.F. Muller, 1774)
62
-------
FIG. 61. Theromyzon tessulatum: (a,b) variations in pigmentation.
34(3) With 0 or 1 pair of eyes located in posterior half of oral sucker
(Fig. 10j); without pulsatile vesicles along lateral margins of
body region (Fig. lla-f); caudal sucker concave, weakly
developed, smaller than boc(y region; body usually not
differentiated into distinct neck (trachelosome) and body
(urosome) regions (except in some preserved individuals of
Myzobdel 1 a and Pisctcolaria, Fig. 11) 35
With 0 or 2 pairs of eyes on oral sucker (Fig.
pairs of pulsatile vesicles (may or may not
Piscicola) along lateral margins of urosome
caudal sucker well developed, as wide as or
width; body usually separated into distinct
10k); with 11
be conspicuous in
(Fig. 10); oral and
wider than body
neck (trachelosome)
and body (urosome) regions (except in some preserved
individual, Fig. 10)
36
63
-------
35(34) Dorsurr? with a series of 6 brownish-black longitudinal stripes,
medial pair most conspicuous, extending from eyes to anal
region; other pairs submarginal; preserved individuals can be
contracted into 2 body regions; small oral and caudal suckers;
body stout flattened centrally and convexed dorsally; mid-body
somites 3-annulate; length 6-8 mm (Fig. 62}
Piscicolaria reducta Meyer, 1940
Dorsum without longitudinal stripes, boc(y elongate, narrow,
and cylindrical or separated into a distinct narrow neck
(trachelosome) and a wide boc(y (urosome) regions; morphological
variance of body shape of individuals known from fresh,
brackish, and marine waters; colorless or with scattered
stellate pigment cells; small oral and caudal suckers; mid-boc|y
somites 12- or 14-annulate; parasitic on a variety of fishes;
length 9-30 mm (Figs. 63,64}
Myzobdella lugubris Leidy, 1851
62
63
FIG. 62. Piscicolaria reducta: (a-c) variable bocty shapes.
Myzpbdsella lugubris, variable body shape.
FIG. 63.
64
-------
FIG. 64. Hyzobdella Tugubris: (a-d) variable body shapes.
36(34) Pulsatile vesicles large, each covering 4 annul 1, and
conspicuous even after preservation (Fig. 10a,c,d,e); body
sharply separated Into 2 regions (Fig. 10c,d,e,j, a small
narrow trachelosome and a*wide urosome, boa> cyclindrical, may
be flattened dorsoventrally; 7 annul 1 per segment.
Cystobranchus 37
Pulsatile vesicles usually small, each covering 2 annull, obscure,
difficult to see after preservation of specimens (Figs. 10f,
g,h,i); body not clearly divided into trachelosome and urosome
regions (Fig. 10a,f,g,h,i); body cylindrical or slightly
flattened; 14 annul 1 per segment. Piscicola 40
37(36) With eyespots (ocelli) on caudal sucker (Fig. 65a-d) 38
Without eyespots (ocelli) on caudal sucker (Fig. 65e-g) 39
65
-------
g
FIG. 65. Dorsal view of caudal sucker of Piscicolidae: (a,b) punctiform
eyespots only; (c) crescentiform eyespots and pigmented rays; (d)
punctiform eyespots and pigmented rays; (e-g) eyespots (ocelli) absent.
38(37) With 8 punctiform eyespots on caudal sucker (Fig. 65a);
2 rows of 12 lateral ocelli on each side of body (Fig. lOb,
66); caudal sucker approximately the same size as the body at
its widest point; cephalic eyes 2 pairs; body cylindrical,
slightly flattened; color, numerous dusky brown-black stellate
pigment cells all over body, segmentation clearly marked by
white bands or void of pigment; length 4-7 mm (Fig. 66). . . .
Cystobranchus meyeri Hayunga & Grey, 1976
With 10 punctiform eyespots on caudal sucker (Fig, 65b); no
lateral ocelli absent; caudal sucker approximately the same
size as the body at it widest point; cephalic eyes 2 pairs;
body cylindrical, slightly flattened, void of pigment, length
9-15 mm (Fig. 67)
Cystrobranchus virgim'cus Hoffman, 1964
66
-------
66
lateral
ocelli
FIG. 66. Cystobranchus meyeri: (a) lateral view showing eyespots on
body region and caudal sucker; (b) dorsal view. FIG. 67. Cystobranchus
virginicus.
39(37) Eyes absent; caudal sucker large, as wide or wider than
urosome (Fig. 65f); oral sucker small, distinct from neck
region; devoid of pigment, or body tinged with brownish-gray
transverse bands, and six more or less distinct bands on the
neck or body tinged with brownish-gray, sprinkled with stellate
flecks; body especially flattened; length up to 30 mm (Fig.
68) Cystobranchus mammillatus (Malm, 1863)
Eyes 2 pairs; caudal sucker large, wider than urosome (Fig. 65g);
oral sucker small, distinct from neck region; devoid of
pigment, or with brownish-black stellate flecks profusely
distributed over entire body; body not especially flattened;
length 10-30 mm (Fig. 69)
Cystobranchus verrilli Meyer, 1940
40(36) Caudal sucker with 8-14 eyespots (ocelli) (Fig. 65b-d) 41
Caudal sucker without eyespots (ocelli) (Fig. 65e); cephalic eyes
2 pairs (rarely 1 pair); body sucker large, clearly marked off
from body; gonopores separated by 4 tertiary annuli; length
14-16 mm (Fig. 70)
Piscicola punctata (Verrill, 1871)
67
-------
69
70
FIG. 68. Cystobranchus mamminatus. FIG. 69. Cystobranchus verrillt:
(a) unpigmented fonn; (b) plgmented form. FIG. 7v.Plsclcola punctata,
41(40) With 10-14 punctiform eyespots on caudal sucker (Fig. 65b,d). . .42
With 8-10 strongly crescentiform eyespots on caudal sucker
(Fig. 65c); gonopores separated by 2 annuli; sperm duct much
convoluted; known from western U.S. and Canada; parasite of
salmonid fishes; length 10-31 mm (Fig. 71)
Piscicola salmositica Meyer, 1946
68
-------
71
72
73
FIG. 71. Pisclcola salmositica. FIG. 72. Piscicola milnerl. FIG. 73.
Piscicola geometra.
42(41) With 10-12 (usually 10) punctlform eyespots on caudal sucker;
dark rays absent (Fig. 65b), but variable; gonopores separated
by 2 annuli; sperm duct simply looped; color in mid-body region
yellowish with brownish-black stellate flecks disposed roughly
in 5 longitudinal rows; dorsally these flecks form a slightly
acute triangle and spread out laterally; length 16-24 mm (Fig.
72) Piscicol a milneri (Verrilli, 1874)
With 12-14 punctiform eyespots on caudal sucker, separated by
dark pigmented rays (Fig. 65d); gonopores separated by 3
annuli; sperm duct simply looped; color greenish, yellowish, or
brownish, usually finely sprinkled with minute brownish-black
stellate flecks, disposed more or less regularly in
longitudinal rows; length 20-30 mm (Fig. 73)
Piscicola geometra (Linnaeus, 1758)
69
-------
43(4) With copulatory gland pores on the ventral surface, at about
10-11 annuli posterior to the male gonopore (Fig. 74a-d).
Macrobdella 44
Without copulatory gland pores on ventral surface (Figs. 74e-h)
47
•e.g.
f
s
FIG. 74. Hirudinldae, external reproductive structures, ventral view:
(a) Hacrobdella ditetra; (b) Hacrobdella decora; (c) Hacrobdella
diplotertia; (d)~Hacrobdena sestertia; (e} Philobdella gracilis; (f)
Haemppis "mannerata: tg? Haemppis terrestris; (h) Haemopis septagon. c.g,
copulatory glands; c.p. copulatory pits; ?rf gonopores (male and female).
70
-------
44(43) Dorsum with about 21 red or orange dots in median line 45
Dorsum without red or orange dots; ventral surface with 8
copulatory gland pores (2 rows of 4) (Fig. 74a); 2 annuli
between gonopores; color of dorsum drab brown or gray, usually
with a darker median field, 1/3 body width, usually with
lateral black irregular dots; ventrum yellowish with some or no
black blotches; length 100 to 150 mm (Fig. 75)
Macrobdella ditetra Moore, 1953
45(44) With 4 or 6 copulatory gland pores on ventral surface
(Fig. 74b,c) .- 46
With 24 copulatory gland pores (with rows of 2 groups, containing
6 gland pores each) situated on a raised pad in adults, (Fig.
74d); 2-2 1/2 annuli between gonopores, color of dorsum olive
green with a median row of red or orange dots, with faint black
irregular striping along midline, lateral margins with row of
black spots; ventral surface reddish or orange, with some black
spots; length 100-150 mm (Fig. 76)
Macrobdella sestertia Whitman, 1886
46(45) Four copulatory gland pores (2 rows of 2) on ventral surface (Fig.
74b); 5-5 1/2 annuli between gonopores; color of dorsum green
with median of red or orange dots, lateral black spots; ventral
surface red or orange with some black spots; length 110-150 mm
(Fig. 77) Macrobdella decora (Say, 1824)
Six copulatory gland pores (3 transverse rows of 2 each)
(Fig. 74c); 4 1/2-5 annuli between gonopores; color of dorsum
light gray with median row of red or orange dots, lateral
margins with row of black spots; ventral surface light yellow
or gray, lateral margins of dorsum same color as ventrum;
length 100-150 mm (Fig. 78)
Macrobdella diplotertia Meyer, 1975
47(43) Glandular area around gonopores; gonopores separated by 3-4
annuli, obscured by deep copulatory depressions and pits (Fig.
74e); dorsum usually with a yellow or brown median stripe.
Philobdella 48
Lacking glandular area around gonopores; gonopores separated by
5-7 annuli (Fig. 74f-h); dorsum with a black median stripe
(sometimes faint), several longitudinal stripes, or no stripes
either mottled, blotched, or plain 49
71
-------
a
FIG. 75. Macrpbdella dltetra; (a,b) variable pigmented forms. FIG
76. Macrobdella sestertla.
72
-------
77
78
FIG. 77. Macrobdella decora,
FIG. 78. Macrobdella djplotgertia.
73
-------
48(47) Middorsal stripe dark brown, if present, with two faint
reddish-brown bands along each side toward margins, separated
by a narrow black stripe, lateral margins with irregular black
stripes, sometimes broken but no discrete spots; 20-26 teeth
per jaw; length 40-85 mm {Fig. 79)
Philobdella floridana (Verrill, 1874)
Middorsal stripe light yellow, with dorsolateral brownish-black
irregular flecks or spots; ventral surface light yellow with
some irregular black flecks or spots near margins; about 35-48
teeth per jaw; length 40-85 mm (Fig. 80)
Philobdella gracilis (Moore, 1901)
49(47)
50(49)
Dorsum uniformily gray with a median longitudinal black stripe;
uniformly dark-olive green with faint longitudinal dark stripes
along midline and numerous small irregular scattered black
flecks; or with no middorsal stripes but with few, moderately
to heavily blotched, spotted, mottled with olive, yellow, dark
gray or black; sometimes uniform color. Haemopls
50
Dorsum with 4 or 6 longitudinal reddish-yellow stripes
forming an ornate pattern on dorsum; color pattern variable,
greenish with irregular black margins; ventral surface black
with white and gray markings; probably not established in North
America, sometimes purchased in drug stores; length, to 100mm
(Fig. 81) Hirudo medicinalis Linnaeus, 1758
Gonophores separated by 5-5 1/2 annuli, female gonopore small
(Fig. 74f,g)
51
Gonopores separated by 6 1/2-7 annuli (Fig. 74h); female gonopore
large, conical (nipple-like) in adults, flattened in immatures;
15 pair of teeth per jaw; dorsum uniformly dark olive-green
with faint longitudinal dark stripes along midline and numerous
small irregularly scattered black flecks; dorsal portion of
caudal sucker with black flecks; parts of certain annuli darker
than others dorsally; ventrum, lighter olive-green without
flecks; occasionally with yellow marginal band; body firm;
length, to 200 mm (Fig. 82)
Haemopis septagon Sawyer & Shelley, 1976
51(50) Dorsal surface with a median black stripe 52
Dorsal surface with few, moderately to heavily blotched, spotted,
or irregular scattered black flecks, mottled with olive,
yellow, dark gray or black; sometimes uniform color; no
middorsal black stripes 53
74
-------
•rf.fi ffii
79
80
81
FIG. 79. Philobdella floridana.
81. Hirudo medicinal!s.
FIG. 80. Philobdella gracills. FIG
75
-------
FIG. 82. Haemopis septagon: (a-d) variable plgmented forms,
76
-------
52(51) Color of dorsum uniformly black or slate gray, with median
longitudinal black stripe and reddish-orange or brownish-yellow
band along margins; ventrally lighter, uniform, few or no dark
blotches or flecks; posterior sucker smaller than body width;
jaws with 20-25 pairs of teeth; body firm; length 150-200 mm
(Fig. 83) Haemopis terrestris (Forbes, 1890)
Color of dorsum brownish-green to olive, with scattered black
and yellowish-orange blotches (usually more black than yellow-
orange), middorsal black stripe and sometimes paired lateral
longitudinal dark stripes; margins conspicuously mottled with
yellowish-orange blotches forming broken longitudinal lines,
ventrally darker, slate gray, uniform, occasional yellowish-
orange blotches; posterior sucker as large as body width; young
with metameric black transverse bands (Fig. 84a); jaws with
9-12 pairs of teeth; body firm; length 60-110 mm (Fig. 84b). .
Haemopis kingi Mathers, 1954
53(51) With jaws and teeth (Fig. 85a-d) 54
Without jaws and teeth (Fig. 85e-g) 55
54(53) Jaws with 10-12 pairs of teeth; gonopores separated by 5 annuli,
color olive-green with moderately to heavy black blotching
dorsally with few scattered yellow blotches; ventrally darker,
uniform gray, few indistinct 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. 86). . . .
Haemopis lateromaculata Mathers, 1963
Jaws with 12-16 pairs of teeth; gonopores separated by 5 annuli;
color variable, usually olive-green, yellowish-gray with
moderate to heavy black mottling or blotched dorsally and
ventrally; or uniform slate gray with few irregular black
blotches, resembling H. grandis; caudal sucker about 1/2 width
of body; length 75-10TT mm (Fig. 87)
Haemopis marmorata (Say, 1824)
77
-------
FIG. 83. Haemopis terrestris: (a) adult; (b) juvenile. FIG. 84.
Haemppis kingi: (a) dorsal view of tw
checkerboard color pattern; (b) adult.
78
-------
velum
eeth
pharynx—
folds
pharynx
folds
velum
pharynx
folds
pharynx fold
g
FIG. 85. Dissected mouth and buccal cavity: (a,b) Haemopi's marmorata,
ventral view; (c) Haemopis marmorata, pharynx and associated structures;
(d) Haemopi's marmorata, teeth and jaw, lateral view; (e) location of
velum and pharynx, without teeth and jaws, ventral view; (f) Haemopis
plumbea, dissected pharynx: (g) Haemopis grandis, dissected pharynx. The
pharynx (f,g) is shown as opened along the midventral line, centered on
the middorsal line.
79
-------
86
\&.<&3%st&jmj':-jp&
m JS5£*Kfti;3P£TO
m
£$
87
FIG. 86. Haemopls lateromaculata. FIG. 87. Haemopls marmorata,
variable forms: (a) dark colored, mottled phase; (b) light colored,
mottled phase; (c) dark colored, immaculate phase.
80
-------
55(53) Margin of oral sucker thick, rounded, aperture transverse, the
lower margin anteriorly convex (Fig. 88); lower surface of
velum (Fig. 85e), closely and finely papillate; pharynx with 15
folds (Fig. 85f); color of dorsum grayish with few, or no black
blotches, with reddish or yellowish-orange band along margins;
gonopores in middle of annuli, separated by 5 1/2 annuli;
length 140-200 mm (Fig. 90)
Haemopis piumbea Moore, 1912
Margin of oral sucker thin, aperture elongate (arched), the
lower margin sharply concave (Fig. 89); lower surface of velum
(Fig. 85e), smooth; pharynx with 12 folds (Fig. 85g); color
variable usually shades of dull green, gray, or plain, always
more or less blotched with black or none; gonopores in furrows,
separated by 5 annuli; length 150-300 mm (Fig. 91)
Haemopis grand 1_s (Verrill, 1874)
89
FIG. 88. Haemopis piumbea, oral sucker. FIG. 89.
oral sucker.
Haemopis grandis,
81
-------
91
FIG. 90. Haemopis plumbea. FIG. 91. Haemppis grandis, variable forms:
(a) dark colored, heavy spotted, blotched phase; (b) light colored,
irregularly spotted, blotched phase; (c) dark colored immaculate phase.
56(4) With 0-3 pairs of eyes (Fig. 13b) 57
With 4 pairs of eyes (Fig. 13c,d) 64
82
-------
-------
f
g
FIG. 93. Dorsal view of dissected male genital atarium and ejaculatory
ducts with or without long preatrial loops: (a) Hooreobdel1 a
melanostoma; (b) Erpobdella punctata punctata or ErpobdeTja punctata
coastal isinc) inmature, Erppbdella punctata punctata; td) Dina
anoculata"; (e) Mooreobdella bucera; (f) Mooreobdella fervida; tg,h)
c. cornua (paired horns); g. XII, twelfth
Mooreobdella microstoma.
ganglion; g.a. genital atrium; p.l. preatrial loop; w. without preatrial
loop.
84
-------
58(57) With 2-2 1/2 annuli between gonopores in furrows or on the rings
(Fig. 92a-g) 59
With 3-4 1/2 annuli between gonopores in furrows or on rings
(Fig. 92h-l) 63
59(58) Gonopores separated by 2 annuli (Fig. 92c-e,g), except
Mooreobdella bucera (sometimes 2 1/2 annuli, Fig. 92f).
60(59)
61(60)
60
Gonopores separated by 2 1/2 annuli (Fig. 92b), male gonopore on
ring, female gonopore in furrow; without paired paramedial
black strips and sensillae; color uniform gray; atrium longer
than wide, atrial cornua (horns) simply curved, sperm ducts
with preatrial loops, extending anteriorly to ganglion XI (Fig.
93b); length, to 100 mm (Fig. 95)
Erpobdella punctata coastal Is Sawyer & Shelley, 1976
Dorsum pigmented with longitudinal black flecks, stripes,
spots, heavily barred, almost black, or lightly pigmented.
Dorsum either lacking pigment, uniform gray (pale red or darker
clouding, sometimes with minute black pigment) or pigmented
with two narrow or broad, dark longitudinal stripes extending
over entire body, including always lighter median stripe. . .
61
62
With 2 or 4 rows of black pigment concentrations on dorsal
surface, forming longitudinal stripes or irregular spots,
median 2 pronounced, submarginal ones wanting in immature
forms; some individuals with black bars or heavily pigmented,
almost black on dorsum; occasionaly a white form .with median
scattered minute black chromatophores; eyes 3 pairs; sensillae
on annuli may be conspicuous; gonopores in furrows, separated
by 2 annuli (Fig. 92c), male gonopore in adults very large;
atrium longer than wide, atrial cornua (horns) simply curved,
sperm ducts with preatrial loops, extending anteriorly to
ganglion XI (Fig. 93b,c); length to 100 mm (Fig. 96,97). . . .
Erpobdella punctata punctata (Leidy, 1870)
With 4 longitudinal stripes on dorsal surface of grayish or dull
black, of which outer pairs, submarginal, duller in color, and
narrower than more distinct inner pair, well separated by a
median stripe of ground color or lacking pigment, uniform gray;
eyes lacking or 3 pairs; no sensillae on annuli; gonopores
separated by 2 annuli in furrows (Fig. 92d); male gonopore
surrounded by circle of papillae; atrium longer than wide,
atrial (cornua) horns simply curved, sperm ducts with preatrial
loops, extending anteriorly to ganglion XI (Fig. 93d); length
10-15 mm (Fig, 98) Pina anoculata Moore, 1898
85
-------
96
FIG. 94. Hooreobdella melanostoma. PIS. 95. Erpobdella punctata
coastalls. FIG. 96. Erppbdella punctata punctata": raT~2 rows of black
pigment concentrations; (b) 4 rows of black pigment concentrations.
86
-------
97
98
99
•-
ma
FIG. 97, Erpobdella punctate punctgta: (a) black barred phase; (b) dark
pigment phase; (c) striped phase. FIG. 98. Dina anoculata. FIG. 99.
Mooreobdella bucera.
87
-------
62(60) Gonopores separated by 2 (sometimes 2 1/2) annuli in furrows
or on rings (Fig. 92e,f); color uniform smokey gray without
black pigment; live specimens reddish from cutaneous blood
vessels, eyes 3 pairs; atrium globoid, with cornua (horns)
projecting laterally, with sperm ducts lacking preatrial loops,
ending abruptly at atrium (Fig. 93e); length 10-30 mm (Fig. 99)
,,.,.... Hooreobdella bucera Moore, 1949
Gonopores separated by 2 annuli, usually in furrows (Fig. 92d);
color uniform smokey gray, lacking pigment entirely, or with
darker clouding, sometimes with minute black pigment, or with
two, narrow or broad, dark longitudinal stripes extending over
dorsum, including always a lighter median stripe, eyes usually
3 (sometimes 4) pairs; atrium globoid with prominent cornua
(horns) longer than its diameter projecting anteriorly, sperm
ducts lacking preatrial loops, ending abruptly at atrium (Fig.
93f); length 20-50 mm (Fig. 100). ...
... Hooreobdella fervida Verrill, 1871
63(58) Gonopores separated by 3 annuli, usually in furrows (Fig. 92h);
color reddish from blood showing through or in preserved
specimens, light yellowish-gray or smokey-gray, no black
pigment; atrium ellipsoidal, wider than 1ong» with cornua
(horns) shorter than diameter of median atrium, projecting
anteriorly, with sperm ducts lacking preatrial loops, ending
abruptly at atrium (Fig. 93g,h); eyes 3 pairs, length 30-50 mm
(Fig. 101) Mooreobdella microstoma (Moore, 1901)
Gonopores separated by 4-4 1/2 annuli, usually on rings
(Fig. 92faj); color of dorsal surface uniformly smokey-gray
without black pigmentation; atrium wider than long, atria!
cornua (horns) projecting anterolaterally to anteriorly; with
sperm ducts lacking preatrial loops, ending abruptly at atrium;
eyes 3 pairs; length to 40 mm {Fig. 102)
Mooreobdella tetragon Sawyer & Shelley, 1976
64(56) Gonopores separated by 2 annuli (Fig, 92d,g) 65
Gonopores separated by 31/2 (sometimes 2 1/2-4) annuli
(Fig. 92k,1) 66
68
-------
100
101
IK
FIG. 100, Hooreobdel1 a ferylda: (a-c) variable forms. FIG. 101,
Mooreobdella microstore: I a) without developed clitellum; (b) with
cfevel oped" ell tell urn." FIG. 102. Hooreobdella tetragon.
89
-------
65(64} Eyes 4 pairs, anterior 2 pairs and posterior 2 pairs arranged
almost in parallel (Fig. 13d); color variable, dorsum
greenish-brown, covered with sparce scattered black or light
colored blotches, interlacing or irregular spots, or plain
(uniform), no striping; most of mid-body annuli partially
subdivided once or twice (Fig. 92g); atrial cornua (horns)
spirally coiled like a ram's horn with sperm ducts forming
preatrial loops, extending anteriorly to ganglion XI (Figs.
103a,b); gonopores separated by 2 annuli in furrows (Fig. 93g);
length up to 100 mm (Fig. 104)
Mephelopsis obscura Vem'll, 1872
Eyes 4 (usually 3) pairs, not arranged in parallel (Fig. 13c);
color uniform smokey gray, lacking pigment entirely or with
darker clouding, sometimes with minute black pigment, or with
two, narrow or broad, dark longitudinal stripes extending over
dorsum, including always a lighter median stripe; atrium
globoid with prominent cornua (horns) longer than its diameter
projecting anteriorly, sperm ducts lacking preatrial loops,
ending abruptly at atrium (Fig. 93f); gonopores separated by 2
annuli, usually in furrows (Fig. 92d); length 20-50 mm (Fig.
100) Mooreobdella fervida Verrill, 1871
66(64) Dorsum with a variable dark brown or black middorsal stripe,
obvious in anterior (cephalic) half, sometimes fading
posteriorly; dorsum color greenish, heavily mottled, minutely
spotted with pale yellow or white irregular transverse rows;
gonopores separated by 3 1/2 annuli (sometimes 4), usually on
rings, male gonopore on ring, female gonopore in furrow (Fig.
91k); 4 pairs of eyes, second pair of labial eyes behind 1st
pair (Fig. 13c); atrium with meaian chamber wider than long,
large cornua (horns) directed anteriorly and curved ventrad,
with sperm ducts forming preatrial loops, extending anteriorly
to ganglion XI (Fig. 103c); length 20-60mm (Fig. 105)
Dina dubla Moore & Meyer, 1951
Dorsum lacking a middorsal stripe; dorsum unpigmented, or
color uniformly smokey gray with varigated dark and light
pigment; gonopores usually separated by 3 1/2 (sometimes 2 1/2
or 3) annuli, male on ring, rarely -fn furrow, female in furrow
(Fig. 921); 4 pairs of eyes, second pair of labial eyes behind
1st pair (Fig. 13c), atrium with simply curved cornua (horns),
with sperm ducts forming preatrial loops, extending anteriorly
to ganglion XI (Fig. 103d); length 25-30 mm (Fig. 106)
Dina parva Moore, 1912
90
-------
p. I.
-p. Z.
_. Z.
FIG. 103. Dorsal view of dissected male genital atrium and ejaculatory
il loops: (a,b) Nj
dubia; (d) Dina parva. c. cornua (paired" horns); g. XII, twelfth"
ducts with long preatrial loops: (a,b) Nephelopsis obscura; (c) Dina
na parva. c. cornua (paired m
ganglion; g.a. genital atrium; p.l. preatrial loop.
-------
_ (a) light colored, irregularly spotted
colored,i rregularly spotted form; (c) imnaculate form.
FIG. 104. Nephelopsls obscura:
form; (b) dart
92
-------
105
106
FIG. 105. Dlna dubla. FIG. 106. 01 na parva
clitell urn; TFTwlW developed cliteTTum.
(a) without developed
93
-------
-------
SECTION 7
GLOSSARY
Acanthobdellid (pi. s): A primitive leech in the family Acanthobdellidae.
Accessory Eyes: A series of supplementary eyes (ocelli) behind the single
pair of functional eyes in the cephalic region of Placobdella hollensis.
Annuli (sing.
Annul us): External body rings or superficial transverse
Basically there are 3 primary annuli (tri-
furrows of the somites,
annulate) per somite, labelled by convention as al, a2, a3 (or sometimes
written al-3), (except in Ollgobdella biannulata which has biannulate
somites). In some genera of leeches, each annul us may be subdivided (see
Table 1) into secondary annuli, bl, b2, b3 ... b6 (or bl-6), and still
further into tertiary annuli cl, c2, c3 ... (or cl-12), and rarely still
further into quarternary annuli dl, d2, d3 ...d24 (or dl-24). Each somite
corresponds with one ganglion in the central nervous system. The ventral
nerve cord ganglia are placed in the middle annulus of the somite.
Annulation can be most easily seen in the lateral margins of the middle
region of the body.
Anterior sucker: Oral sucker.
a
Atrial cornua: Horns or hornlike prolongations of the atrium.
Atrium: A variously shaped, male reproductive organ, consisting of three
parts: a thin-walled eversible bursa, a thick walled glandular and
muscular medium chamber, and a pair of atrial cornua openings into the
latter of similar structures of the male reproductive system. The atrium
opens externally through the male gonopore.
Buccal cavity: Mouth cavity.
Bulbous: Bulb shape (Figs. 31, 32, and 33).
Caudal ocelli: Dot-like or crescent-shaped eyespots on the caudal sucker
of certain piscicolids. See punctiform and crescentiform ocelli.
Caudal sucker: Posterior sucker.
Cephalic Region: Head region.
Chaetae (sing. Chaeta): Hair or bristle-like structures of some classes
of Annelida. The term setae, however, seems to dominate recent English
language publications.
95
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Chromatophore: A cutaneous pigment cell or group of pigment cells which
under control of the nervous system can be altered in shape and size to
produce a color change in some rhynchobdellids.
CliteTliim: A regional epidermal saddle or swollen glandular portion of
the integument in the area of the gonopores of certain leeches, especially
the Hirudinidae and Erpobdellidae, visible in adults during the breeding
period. It contains gland cells that secrete material to form cocoons.
Complete somites: Segments having the full number of annuli or body rings
characteristic of the genus.
Copulatory depressions: A glandular area around the gonopores and
copulatory pits on the ventral surface in Philobdella.
Copulatory gland pores: Pores present in a linear or transverse pattern
of either 4, 6, 8 or 24. They are located on the ventral surface, 4 or 5
annuli, posterior to the female gonopore in Hacrobdella.
Copulatory pits: Thick depressions and prominences of the glandular area
around the gonopores of Philobdella. See copulatory depressions.
Crescentiform ocelli: Crescent-shaped or crescentric eyespots on the
caudal sucker of Piscicola salmositica.
Crop (Stomach): Sac-like dilatation or largest part of the alimentary
canal adapted for storage of blood, and so on; may be a single, straight
tube, or contain several paired diverticula.
Crop caeca: Segmental pouches or diverticula of the crop (stomach).
Denticles: Small teeth-like processes; two rows of teeth (distichodont)
wholly or partly in Phi'lobdella and some species of Haemopis or one row of
teeth (monostichodont) in Hirudo and Hacrobdella.
Digitate processes: A marginal circle of 30-60 retractable processes,
finger!ike when everted on the inside rim of the sucker cavity in
Actinobdella. The digitate processs are usually retracted in preserved
specimens but their position is evident by faint radiating ridges or bands
visible on the outer rim of the caudal sucker.
Discoid head: The oral sucker and cephalic region widely expanded,
circular and demarcated from the body by a constriction; in Placobdella
montifera and Placobdella nuchal is.
Erpobdellid (pi. s): A leech in the family Erpobdellidae.
Eyes (Ocelli): Photoreceptors confined to the cephalic "head" region,
formed from a number of light sensitive cells (eyespots) backed by a
pigmented cup. The eyes are located on the dorsal or lateral surface of
the anterior segments, are segmental, vary in number and arrangement.
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Eyespots (Ocelli): Photoreceptors of the cephalic region, lateral region
of trie urosome, or caudal sucker of certain leeches.
Ganglia (sing. Ganglion): Concentration of nerve cell bodies In the
ventral nerve cord. Excluding the supra- and subesophageal mass (six
ganglia) and the caudal mass (seven ganglia), there are 21 ganglia In the
ventral nerve cord, labeled in Roman numerals VII-XXVII. The neural
annulus of segment (somite) 10 would, therefore, be expressed as Xa2 of
the triannulate somite.
Gonopores: External openings of the reproductive tracts, located on or in
the furrow of the annulus on the midventral surface of somite XI and XII
(about one third from tip of cephalic region). The male gonopore is
anterior to, larger than, and more conspicuous than the female gonopore.
The male and female gonopores generally are separated by 1-7 or more
annuli but in a few species open into a common gonopore.
Glosslphom'ld (pi. s): A leech in the family Glossiphoniidae.
Hirudim'd (pi. s): A leech in the family Hirudinidae.
Hiradtm'dae: The widely accepted familial name based etymologically on
the stem hlrudlnls, the Latin genitive singular of hirudo. It should
always be used in preference to Hirudidae (Art. 29a of international Code),
Hirudofauna: All the leeches peculiar to a country, area, or period. A
treatise on leeches.
Incomplete (abbreviated) somites: Occur at both ends of the body of
leeches and may have any number of annul 1 less than the complete somites
into which they grade.
Integument: A general term for the covering, or outer layers of an
animal. The purpose of the integument is for defense and sensing
environmental conditions.
Internal ridges: Fleshy anatomical structures of the pharynx in
Hirudinidae. Sometimes called pharynx folds or pods.
Jaws: Three large, oval bladelike jaws shaped like a half circular saw
occurs just within the mouth cavity of some species of GnathobdelHda.
Each bears along the edge a large number of small teeth (denticles). The
three jaws are arranged in a triangle, one median dorsal and the other two
ventrolateral. They have a covering of cuticle, and along the free edges
of the disc it 1s thickened to form rows of numerous, minute teeth
(monostichodont or distichodont).
Lateral ocel11: Eyespots on the urosome of Cystobranchus meyerl.
Leech: Any segmented worm with terminal suckers used for attachment and
locomotion; the various species may be parasites (blooding sucking),
predators, or scavengers; most are aquatic in North America.
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Hetamere: A body segment or somite.
Metameric: Referring to metamerism or segmentation.
Metameric dots, spots, patches, or prominences: Yellowish or whitish
areas usually metamerically (segmentally) arranged; some represent
sensillae or segmental receptors.
Neural annul us (Sensory annul us): The annul us aligned with the ganglion
in the central nervous system. In the 3* or 5-annulate somites, it is the
middle annulus.
Nephridia {Sing. Nephridiuml: Excretory organs, usually that of
macroinvertebrates.
Nuchal plate: See scute.
Ocelli (sing. Qcellus): Eyespots found in cephalic, caudal ends, or
lateral margins of body of certain leeches. They may be found singly in
any body part, including the caudal sucker and lateral margins, and
aggregations of them in a common network of chromatophores usually form
eyes of piscicolids. Eyes of leeches are confined to the cephalic "head"
region and are segmental in some species. Such eyes are simple when
single, and compounded, such as Placobdella, when one or two small ones
are attached to the main one.
Oculiform spot: See punctiform ocelli,
Oral sucker: Anterior sucker.
Papillae: Small to large protrusible sensory organs scattered or in a
metameric series on the dorsal surface of the leech and thought to be
tactile in function. Large papillae are sometimes called "tubercules."
Pedicel (Peduncle): A stem or stalk of annuli supporting the caudal
sucker in some leeches.
Peduncle: See pedicel.
Pharynx: A muscular tube, anterior part of alimentary canal, following
the buccal cavity.
Pharynx folds: Internal muscular ridges or fleshy pods of the pharynx.
See internal ridges.
Pigment: A cutaneous structure of brown, black, red, green, blue coloring
in the integument of leeches. They are found in chromatophores and
arranged between muscle strands and other organs to give a characteristic
color pattern.
Piscicolid (pi. s): A leech in the family Piscicolidae.
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Proboscis: A tubular structure with a crown-like anterior end found in
leeches of the families Glossiphoniidae and Piscicolidae. It is a tube
lying within the proboscis cavity, which is connected to the yentrally
positioned mouth by a short narrow canal. The proboscis is highly
muscular, has a triangular lumen, and is lined internally and externally
with cuticle. This organ is an anatomical characteristic of the
rhynchobdellids.
Posterior sucker: Caudal sucker.
Preatrial loops: Vas deferens or ejaculatory ducts conveying spermotozoa
of certain erpobdellids.
Pulsatile vesicles: Eleven small hemispherical vesicular structures
filled with coelomic fluid and found along the lateral margins of
Cystobranchus and Piscicola species. They pulsate rhythmically and
function as respiratory organs.
Punctiform ocelli: Eyespots or dot-like structures on the caudal sucker
or lateral margins of some piscicolids.
Rays: Distinct pigmented areas on the caudal sucker of certain
piscicolids.
Rhynchobdellid: A leech in the Order Rhynchobdellida.
Scute (Nuchal plate): A chitinous scale-like structure found on the
dorsum of the neck region" of Helobdella stagnalis; function unknown.
Segment: A somite, metamere, or a series of anatomical divisions of the
body.
Sensillae (Segmental receptors): Whitish or yellowish, rounded, oval,
metameric dots, spots, patches, prominences (sometime raised), which may
resemble minute, inconspicuous papillae on neural annuli containing
several types of sensory cells that are receptive to light and water
movement.
Setae: See chaetae.
Somite (Segment, Metamere): A true body segment or metamere made up of
superficial transverse annuli or rings. All leeches have 34 somites, each
of which corresponds to a nerve ganglion.
Stomach: See crop,
Taxon (pi. Taxa): Any taxonomic group, for example, a race, subspecies,
species, genus, family, order, and so forth.
Teeth: See denticles.
Testisacs: Coelomic sacs containing the testes.
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Trachelosome: Narrow neck region of some piscicolids.
Tubercles: Large papillae.
USNM: An abbreviation, usually associated with museum specimen catalogue
numbers, for the United States National Museum (National Museum of Natural
History).
Prosome: The thicker or wider body region of some piscicolids.
Velum: A flap of skin which separates the buccal cavity from the cavity
of the oral opening (mouth) in Hirudinidae.
White prominences: The raised, irregular-shaped areas on the dorsum of
certain leeches and functioning as segmental receptors.
Whitish or yellowish dots, patches, spots (sometimes raised): Sensillae,
segmental receptors usually metamerically arranged.
100
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SECTION 8
SPECIES DISTRIBUTION
The distribution and abundance of aquatic leeches, like other groups
of organisms, are affected by biological, chemical, and physical
characteristics of the environment. The important ecological factors
include food organisms, substrate requirements, habitat type (lentic and
lotic), water depth, pH, temperature, dissolved oxygen, dissolved salts,
turbidity, salinity, and organic and chemical pollutants. Information and
discussions on the effects of these environmental factors and other topics
at the species level can be found in Mann (1962), Herrmann (1970), Klemm
(1972a, 1975, 1976) (also cf. Selected Bibliography), and in the review
paper on pollution ecology of leeches by Sawyer (1974). However,
relatively little has been written specifically on the natural history,
ecology, pollution tolerance, and distribution for many of the North
American species of leeches.
Zoogeographic information of the hirudofauna is rather sketchy because
our current knowledge on the systematics and ecology of leeches is
deficient. However, existing information on the zoogeography of leeches
has been discussed for North America (Sawyer, 1972), for the world (Soos,
1970), for the physiogeographic provinces of North and South Carolina
(Shelley, 1975; Sawyer and Shelley, 1976), for Colorado (Herrmann, 1968;
1970), and for South America and MesoAmerica (Ringuelet, 1980).
Nevertheless, the zoogeography of leeches still presents numerous and
enormous gaps in North America and on a world scale, and a warning should
be given against making generalizations that are too broad.
The distribution records for the leech species in Maps 1-32 (pages
105-136) are based on the published literature (Herrmann, 1970; Klemm,
1972a,b, 1977; Sawyer, 1972; Davies, 1973; Sawyer and Shelley, 1976; and
Klemm, et al. 1979), and the records of the authors listed with an
asterisk in the Selected Bibliography. In addition, the distribution of
each species is based on my unpublished personal records, on the
identification or verification of specimens sent to me by the many
individuals mentioned in the Acknowledgment, and the distribution data
with identification or verification of voucher specimens from the
reputable museums that are also listed in the Acknowledgment. Many of the
distribution records on the maps are new for the species, but they will
not be indicated there specifically. Information on specific locality
records for some species can be found in the papers given in the Selected
Bibliography.
In an earlier study (Klemm, 1972b), I erroneously reported
Glossiphom'a complanata from Florida and Mississippi, Alboglossiphom'a
101
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heteroclita from Mississippi, Batracobdella picta, Hacrobdella sestertia,
and PlacoTxfella ornata from Louisiana, and trpobdella triannuiata from
California and North Dakota. Furthermore, the species, Placobdel'la
multilineata, that I reported from Michigan (Klemm, 1972a,b)9 is changed
to P. ornata.
The symbols on the distribution maps only indicate that the species
has been reported from the state in the United States and province or
territories in Canada. Information on abundance, host preference, and
ecological restrictions are also given where appropriate for each species
in the legion of each map.
The distribution patterns for all North American leech species are
still not completely known, due primarily to inadequate collecting, to the
small number of sites and different habitats sampled, and to the
infrequencies in reporting the record at the species level. However, this
manual has extended the ranges for many species.
The Great Lakes region of North America is probably the only area that
has been extensively surveyed for leeches. There are several published
records of leeches from the southern states, especially North and South
Carolina (Sawyer and Shelley, 1976), but the southern states are in need
of more extensive sampling for leeches. Very few records are available
for the western United States, and the distribution data for leeches from
this area are badly needed. In Canada, several areas have been surveyed
for leeches (Davies, 1973), but not intensively, and more information is
still needed on leech distribution in Canada.
At first glance, the distribution maps (1-32) will indicate that
several species are widely distributed. Some are predominatly northern or
southern, and others are geographically restricted. There is sufficient
evidence to indicate a northern distribution for the following species:
Glossiphgnia complanata, Helobdella papillata, Marvinmeyeria lucida,
Placobdella ornata, Theronjyzon biarinulatum, T. rude, T. tessulatum,
Pisci'cola geometra, P. mi'lnerT, Haemppis grandis, H. "plumbea, Macrobdella
decora, [Tina dubfa, IT. parva, Mooreobdella fervTda, and Nephelopsis
obscura.
Macrobdella ditetra, Oligobdella biannulata, Philobdella gracilis,
PlacobdeTIa multilineata, and P. trans!ucens appear to have a southern
distribution pattern. IToth Placobdella multilineata and Phi'lobdella
qraci'lis, however, extend northward through the Mississippi Valley.
Erpobdella punctata coastal is, Mooreobdel1 a melanostpma, and H. tetragon
have an Atlantic coastal distribution pattern, but the habitat range for
these recently described species (Sawyer & Shelley, 1976) is not known.
The northern or southern limits of the distributional ranges for some
species is not known, and some closely related species are known to have
overlapping ranges.
The discontinuous or restricted distribution of some species, which
have been reported only from a single collecting site or state, may be a
function of low abundance or insufficient sampling. Leech species which
102
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have only been reported from a single area, may have a restricted
distribution due to host preference or some unknown specific chemical,
physical, or other biological factors. With proper collecting and greater
sample frequency, the leech fauna for some areas will undoubtedly be found
to be much richer and varied than now apparent.
Leeches may be dispersed actively or passively. Some species in the
genus Theromyzpn are distributed by their food source (waterfowl), which
they infest.However it is not known why Theromyzpn spp. are not found in
the southern states since many waterfowl migrate there in winter. Fish
leeches of the families Glossiphoniidae and Piscicolidae are dispersed
passively by migration of the fish host from one body of water or drainage
system to another. Other species of glossiphonids migrate passively by
the movements of reptiles and amphibians. There have also been several
reports that species of the families Hirudinidae and Erpobdellidae make
seasonal upstream migrations. The maps are incomplete. As leeches are
collected and identified to species, they can be added to the maps.
At one time Hirudo medicinal is (Linnaeus, 1758), the European and
western Asian medicinal leech, was imported into the United States by the
thousands for medicinal purposes specifically, phlebotomy and research.
Davies (1973) indicated that hL medicinal is was available in certain drug
stores in Calgary, Alberta, CanacIiT He reported collecting specimens from
a slough north of Calgary, apparently abandoned there, presumably after
being used medicinally. He never indicated, however, the establishment
there of a wild population. Sawyer (1972) and I have never found or
confirmed a wild population record of H_. medicinal is for North America,
and we do not consider it as being established here. I have included this
species because it is still possible to purchase it from a few drug stores
in distinct, ethnic areas of North America.
The record for Batracpbdella paludosa (Carena, 1824), another European
leech, is based on a single specimen from Nova Scotia (Pawlowski, 1948).
The specimen is no longer available for verification. This species,
however, has never been confirmed here, and Sawyer (1972) and I do not
considered it as being established in North America. I have included _B.
paludosa in this manual only because of Pawlowski's (questionable)
identification. If specimens of this species are collected, they must be
confirmed and reported by someone familiar with the species before it can
be considered a valid North American species.
Erppbdella triannulata Moore (1908), a poorly described species, was
originally taken from Lake Amatitlan, Guatemala, Central America (Moore,
1908). Moore (1936) also reported it from the Yucatan and stated that
this species was also found distributed extensively throughout Central
America, Mexico and the Pacific United States. Soos (1966a, 1968) and
Klemrn (1972b) reported it from southern California on the basis of J.P.
Moore's old record. However, for reasons unknown, Moore (1959) and Mann
(1962) excluded this species, E. triannulata, from their keys to North
American leeches. Ringuelet (T976) stated its distribution only as
Guatemala and Mexico. I have examined the poorly preserved type material
(ANSP 2389) at the Academy of Natural Sciences of Philadelphia but was not
103
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able to assess or describe its anatomy further. Furthermove, I have not
been able to obtain fresh material from southen California or elsewhere.
Until £. triannulata is fully described, and additional specimens of this
species are confirmed by a specialist, it is best to consider Moore's
record for southern California as erroneous.
Ringuelet (1976) reported Erpobdella octoculata (O.F. Muller, 1774)
from Mexico and the U.S.A. from an old record.The record for the United
States is erroneous. Davies (1971) included this species in his key of
Canadian leeches on the basis of a questionable identification made by
J.P. Moore of Erpobdella atormaria (Cavena, 1820) as reported in Rawson
(1953) for the Northwest Territories. I questioned the validity of these
records and considered them erroneous (Klemm, 1972b). Later, Davies
(1973) concurred that the Canadian record for E. octoculata was doubtful.
Soos (1966a) viewed E. atomaria as a variety oT E. octoculata and recorded
its distribution in Europe and Japan, and stated that E. octoculata was
restricted only to the Palearctic Region, and there is no valid record of
this species occurring outside of that region (Soos, 1966a, 1968),
Ringuelet (1976, 1980) reported from a single old record the species,
Haementeria officinal is de Fillippi (1849), from New Orleans which may be
a specimen in the Paris museum. The only known distribution of this
species is Paraguay, Venezuela, and Mexico. Sawyer (personal
communication) suspects that the specimen might be Placobdella papillifera
which it resembles. Until the specimen in the Paris museum is confirmed
by examination, I consider this record erroneous.
104
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70
GO
MAP 1.
Acanthobdella peledina
Comments:
A peledina is widely distributed in the USSR and Europe, but this
species is found infrequently. It has been recorded as indigenous to
Norway and Sweden. Hauck, et al. (1979) reported it from Alaska, where it
was found parasitizing the least Cisco (Coregonus sardinella). Holmquist
(1974) reported finding a juvenile specimen of an unidentified species of
Acanthobdella in a benthic sample also from northern Alaska (cf. Notes on
Identification). A. peledina has been collected on fish hosts of the
genera Salmo, Salvelinus, Coregonus, Stenodus, Thymallus, and Lota (Koli,
1961; Anderson, 196Z; Dahm, 1962; Andersson, 1965; Nurnrinen, 1965, 1966;
Borgstrom and Halvorsen, 1972; and Solem, 1975).
105
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ISO 150 140 130 120 110 100 90 8C 7C 60 50
20
10
60
MAP 2.
Actinobdella annectens
Acti nobdel1 a inequiannulata
Comments:
A* annectens 1s described from a single specimen collected on a snapping
turtle (Chelydra serpentina) from Lake Erie, Ontario, Canada. It has not
been reported since its original description (Moore, 1966).
£. inequiannulata is widely distributed but not frequently encountered.
Daniels and Freeman (1976), however, found it abundantly infesting the
white sucker (Catostomus comrnersoni) at Lake Louisa, Algonquin Provincial
Park, Ontario, tanada.
106
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60
160
150
140 130 120 1JO 100 90 BO 70 60 50
30
10
60
MAP 3. • Alboglossiphonia heterocllta
Comments:
A' heterocllta is found rarely. It is usually collected in benthic grab
samples, especially in the Great Lakes region. This scarcity of reports
may merely reflect the difficulty in finding this small species. I have
examined specimens collected from a marsh area of the Piscataway Creek in
southern Maryland. This species also occurs in Eurasia, (cf. Notes on
Systematics).
107
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160 15P 140 130 120 110 100 SO 80 70 60 50 40
30
20
10
10
70
60
MAP 4. * BatracobdelTa cryptobranchii
Batracobdella michiganensis
• Batracobdella phalera
Comments:
JL- cryptobranchii is known only from the Ozark hellbender
fCryptobranchus alleganiensis bishopi) and reported from Ozark County in
southeastern Missouri (Johnson and Klemm, 1977).
B. michiganensis has been reported by Sawyer (1972) from St. Joseph
County, Michigan. I have collected several individuals (USNM 49959) from
Emmet County, Michigan.
EL phalera is widely distributed but not frequently encountered. I have
found this species free-living and parasitizing the bluegill (Lepomis
macrochirus).
108
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140 130 120 110 100 90 80
10
10
80
70
60
MAP 5.
Batracobdella paludosa
Batracobdella
Comments:
£.• paludosa is common in some localities of Eurasia. Pawlowski (1948),
based on a single specimen which is no longer available for examination,
reported it from Pottle Lake, North Sydney, Nova Scotia in Canada (nee
Newfoundland reported in Sawyer, 1972; Elliott and Mann, 1979). UntTT
additional individuals are collected, it is believed to be a doubtful
species in North America, (cf. Discussion in Distribution Section).
£• PJcta is common in some localities and widespread in northern North
America, and it is usually found inhabiting ponds which contain its
favorite hosts, amphibians. Sawyer and Shelley (1976) reported that the
range of this northern species also extends south in the Appalachian
Mountains. I have examined specimens found parasitizing the tiger
salamander (Ambystoma tigrinum) in ponds in Utah.
109
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70
60
MAP 6.
Boreobdella verrucata
Glossiphom'a complanata
Comments:
£.• verrucata Is reported for the first time from North America (Alaska;
Bering and Kodiak Islands, Alaska; Vancouver Island, British Columbia).
Some reports of £. complanata from Alaska and northern areas of Canada may
actually be J5. verrucata (cf. Notes on Identification).
&. complanata Is common and widely distributed in the northern half of
the United States and Canada. This species also occurs in most parts of
Eurasia.
110
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MAP 7. •Helobdella elongata
Helobdella fusca
Comments:
il* e]oflfl.ata is widely distributed and is found infrequently throughout
North America. The small size and benthic habit of the species can make
it difficult to collect without the aid of grab sampling devices.
Ringuelet (1976) reported it from Cuba.
H. fusca is common in the Great Lakes region and widely distributed. It
is~less common in the southern United States. Some published records of
H- triserial Is in Canada may actually be H. fusca (cf. Notes on
"Systematic*). I have identified specimens found inside the operculate
snail (Goniobasis virginica) and inside the shell of Ferrissia sp. This
species also parasitizes a variety of pulmonate snails (Sarah, 1971;
Klemm, 1975, 1976).
Ill
-------
160
HO 130 120 110 100 90 80 70 60
10
MAP 8.
Helobdella pap ill a ta
Harvlnmeyeri'a luclda
Comments:
H_. papillate is uncommon and has only been collected in the Great Lakes
regi on.
M. luclda is widely distributed in Cananda and common in some
localities. The species is also common in certain localities of
southeastern Michigan (Sawyer, 1968, 1972; Klemm, 1972a,b; 1975; 1977).
Welch (1975) and Brook and Welch (1977) reported it from Nebraska, but I
have examined some of the specimens and was unable to confirm them. Until
additional individuals are collected and positively identified, the record
for Nebraska is erroneous.
112
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60
160
150 140 130 120 110 108 90 80 70 SO 50
40
30
MAP 9. • Helobdena stagnalis
Helobdella transversa
Comments:
_H. stagnalis is reported from every continent except Australia. It is
found abundantly throughout the northern half of the United States,
including Alaska and Canada, but is less common in the southern states.
H. transversa has been reported only from southern Michigan (Sawyer,
1972}":
113
-------
180 150 140 130 120 110 ISO SO 80 TO 60 50 AD
30
20
10
80
70
60
MAP 10. • Helobdella triserialls
Comments:
Jl* triserialls 1s widely distributed and Is frequently collected In some
localities.TRTs species Is also found abundantly In Central and South
America, (cf. Notes on Systematlcs).
114
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20
ID
6Q
MAP 11. • Oligobdella blannulata
Placobdella hollensis
Comments:
£. biannulata is rarely encountered; Sawyer and Shelley (1976) reported
that It may be host specific for desmognathine salamanders and that it is
restricted to small streams in the montane areas of the Carolinas.
£.• hpTlensis is uncommon and occurs in certain woodland ponds of the
Great Lakes region and is reported from a few other localities in North
America. One record of a single specimen, collected on water hyacinth
(Eichhornia crassipes) and identified by J. P. Moore, exists for the
Withlacoochee River, Florida (USNM 44005). I have since examined
specimens resembling this species from the Little Withlacoohee River,
Sumter County (USNM 62828) and Taylor Creek, Okeechobee County, Florida
and from Screven amd Effingham Counties, Ogeechee River, Georgia,
115
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ISO
ISO
140 130 120 110 100 90 80 70 60 SO
30
60
MAP 12.
Placobdella montifera
Placobdella multillneata
Comments:
P. montifera is widely distributed throughout the United States and
Canada, but otherwise uncommon.
£.• multilineata is common in the southern half of the United States and
uncommon elsewhere. Beck (1954) collected one specimen from Currant
Creek, Wasatcn County, Utah. The specimen is not available for
verification, but the validity of this record needs to be substantiated by
further collections.
116
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140 130 120 110 1QO 90 8G 70 60
60
MAP 13. • Placobdella nuchal is
Placobdella ornata
Comments:
£.• nuchal is has been reported from several localities of the coastal
plains in the Carolinas (Sawyer and Shelley, 1976). I have identified
specimens of £. nuchal is. collected from the chin of a channel catfish
(Ictalurus punctatus), from Kentucky Lake near Paris Landing, Tennessee.
White (1977) (nee P. parasitica Say, 1824) found an individual attached to
the eye of a carp TCypnnus carpio) from the Ohio River, Kentucky. These
fish are the first known hosts of the species. I have examined several
specimens collected free-living from the Susquehanna River, Three Mile
Island, Pennsylvania, and the Ogeechee and Canoochee Rivers in Georgia.
P. ornata is common and widespread throughout the northern half of the
UnTted States and Canada; there are few reports of this species from
southern localities. It has been reported from Mexico (Caballero, 1940b;
Ringuelet, 1976).
117
-------
60
MAP. 14. •Placobdella papillifera
Placobdella pediculata
Comments:
f.- pap111 Ifera is widely distributed, common in some areas and uncommon
elsewhere.Sawyer (1972) reported 1t from the musk turtle (Stenothaerus
odoratus).
P. pediculata 1s uncommon and reported mainly from the Midwest. Records
indicate that this species has a high degree of host specificity for the
freshwater drum (Aplodinatus grunniens) and surely will be found throughout
the distribution of its host. Data show that the isthmus and the inside of
the opercula (Fig. lla) are the usual sites of attachment. Adult specimens
that I have examined were also found parasitizing £. grunm'ens from the
Mississippi River in Illinois, Kentucky Lake in southwest Kentucky and
northwest Tennessee, and Lake St. Clair in Michigan. This leech has never
been collected in the free-living state.
118
-------
16D 150 HO 130 120 110 100 90 80 70 SO 50 40
3D
20
70
SO
GO
MAP 15. "Placobdella parasitlca yPlacobdella translucens
Comments:
£. parasitica is common and widely distributed throughout the United
States and Canada.
P. translucens is uncommon and known only from the Gulf States and coastal
areas of the southern United States. I have examined specimens from North
Carolina, Georgia, Alabama, and Louisiana.
119
-------
140 130 120 110 100 90 80 70 60
10
80
70
60
MAP 16. • Theromyzon biannulatum • Theromyzon rude *Theromyzon tessulatum
Comments (cf. Notes on Systernatics.):
T. biannulatum is uncommon and known only from the central and eastern
UnTted States.
T. rude is uncommon and apparently distributed in the central and western
UriTted States and Canada, including Alaska, and is infrequent elsewhere. I
have examined specimens from British Columbia.
T. tessulatum 1s uncommon and characteristic of Eurasia but has been
reported infrequently from North and South America. I have listed its
distribution in North America as questionable (Klemm, 1977), but I have
since examined specimens from Quebec and British Columbia In Canada. Davies
(1973) has also reported it from British Columbia and Saskatchewan, and
Pawlowski (1948) from Nova Scotia. Herrmann (1970) collected it from three
sites near Boulder, Colorado. The South America records are Blanchard
(1892) from Chile, Oka (1932) from Peru, Moore (1911) and Ringuelet (1944a)
from Argentina.
120
-------
HO 130 120 110 100 90 80 70 60
10
10
SO
MAP 17. • Cystobranchus mammillatus
Cystobranchus meyerl
Comments:
£. mammillatus is uncommon but widely distributed throughout north and
central Europe. In North America this species has only been reported from
the Northwest Territories of Canada (Meyer and Roberts, 1977).
c- rceyeri *s uncommon and reported only from eastern New York (Hayunga
an? Grey, 1976). I have also examined specimens collected from the fins
of the white sucker (Catostoma commersoni) from southeastern Lake Ontario.
121
-------
160 ISO 140 130 120 118 101 90 80 TO 60 50 40
30
2D
10
80
70
10
60
MAP 18. •Cystobranchus verrilli * Cystobranchus vlrgim'cus
Comments:
C. verrilli is uncommon with several reports from the Great Lakes region
aricT southern Canada; the species is less common elsewhere.
C. virginicus is uncommon and has been reported from Virginia (Hoffman,
1974) and West Virginia (Putz, 1972). (cf. Notes on Systematics).
122
-------
MAP 19. •Myzobdella lugubris
Piscicola geometra
Comments:
M. lugubris is common and widely distributed throughout North America in
both fresh and brackish waters. Also, Caballero (1940a) described a
closely related species, ^. patzcuarensis, from Lake Patzcuaro in Mexico.
£. geometra is common but not found abundantly in Eurasia. It is
considered by some authors to be holarctic in distribution. The
occurrence of this species in North America is open to question (cf. Notes
on Systematical. Sawyer et al. (1975) reported that this species also
occurs in brackish waters of the upper Baltic Sea and in many types of
freshwater habitats throughout Eurasia.
123
-------
160
150 U?___!30 120 110 100 30 8C 70 60 __50
30
20
70
60
MAP 20. • Piscicola punctata
Piscicola salmositica
Comments:
£.• punctata is widely distributed in North America, chiefly reported
from the northern United States and Canada, but infrequently encountered
elsewhere.
£• salmositica is locally abundant in western United States, including
Alaska, as well as British Columbia in Canada, and uncommon elsewhere.
124
-------
60
160
150 140 130 120 110 100 90 80 70 BO 50 40
3D
20
10
80
70
60
MAP 21.
Piscicola milneri
Piscicolaria reducta
Comments:
£.* mi'\"er? ^s uncommon and reported from the Great Lakes region, the
eastern United States, including Alaska, and throughout Canada, (cf.
Notes on Systematics).
P_. reducta is uncommon in some localities, elsewhere common, especially
in the Great Lakes region. I examined a specimen collected on the arrow
darter, Etheostoma sgg'jtta, from the Poor Fork of the Cumberland River in
Kentucky'Pearse (1936) reported a juvenile Piscicolaria sp. from Carteret
County, North Carolina, but Sawyer and Shelley (I97bj considered it a
dubious record.
125
-------
ISO 150 140 130 120 HO 100 90 80 70 GO 50 40
30
20
70
50
60
MAP 22. • Heflnopis grandls » Haemopls kingi
Comments:
H. grand1s is common 1n the Great Lakes region, eastern United States
aricT Canada; there are few records of this species elsewhere.
H. kingi is uncommon and known only from northwest Iowa (Mathers, 1954}
an? southwest Colorado (Herrmann, 1970). I have examined specimens,
resembling this species, from Seven Springs, Arizona and the Guadalupe
Mountains, New Mexico.
126
-------
160
ISO Hfl 130 120 110 100 90 80 70 60 50
30
20
70
EO
MAP 23. • Haemopls 1aterpmaculata
Haemopis marmorata
Comments:
H. lateromaculata is uncommon and recorded only from northwest Iowa
(Mathers, 1963) and Minnesota (Mathers, 1963; Fish and Vande Vusse, 1976}
11' mannorata is common in some localities and widely distributed
throughout North America. Richardson (1971) described a new species,
Percymoorensis cabal 1eroi, from Mexico that closely resembles the North
American H. marmorata.
127
-------
140 130 120 110 100 S8 3D 70 60
MAP 24. • Haemopis piumbea * Haemopis septagon • Haemopis terrestris
Conments:
H. piumbea is uncommon and known from Minnesota, Wisconsin, Michigan and
OhTo in the United States; Ontario and Quebec in Canada. I have examined
specimens from Michigan, but old records of this species occurring in
Wisconsin, Ohio, and Ontario need to be substantiated by new records.
jj. septagon has been reported from the coastal plain and eastern
piedmont areas of the Carolines and Virginia (Sawyer and Shelley, 1976).
I have identified specimens also collected in North Carolina. This
species may be difficult to find because of its terrestrial existence.
il- terrestris is uncommon and known from the southern Great Lakes
region, Mississippi Basin to Colorado, and southward to the Gulf States.
This species has been reported from both aquatic and terrestrial habitats.
128
-------
160
ISO
140 no 120 no 100 90 so TO eo
10
10
70
GO
25.
Macrobdella decora
Macrobdella dlplotertia
Comments:
M. decora is common In ponds in the northern half of the United States
aricT Canada. This species is less common in the southern states; Sawyer
and Shelley (1976) regarded it as being restricted to the piedmont and
montane areas of the Carolines and Georgia. This species is also known to
occur in northern Mexico (Caballero, 1952).
M. diplptertia has been reported only from western Missouri (Meyer,
1975). Klemm, et al. (1979) identified several specimens of this species
from Chautauqua County in southeastern Kansas, thus extending its range.
129
-------
MAP 26. »Macrobdella ditetra
Macrobdella sestertia
Conments:
M.- ditetra 1s common and widely distributed In the Gulf States and
coastal plain areas of the Carolines; it appears less common elsewhere.
M. sestertia is uncommon and known only from eastern Massachusetts
(WFitman, 1886; Moore, 1959; Klemm, 1972b; Sawyer, 1972; Smith, 1977).
130
-------
MAP 27. • Philobdella floridana
Philobdella gracili's
Comments:
P. floridana has been reported from Lake Okeechobee in Florida (Verrill,
1972)1I have examined specimens that resemble this species from
Jacksonville and West Palm Beach, Florida, (cf. Notes on Systematics).
£.* gracilis is common and widely distributed throughout the southern
United States and is less common elsewhere-. Individuals of this species
have been reported from Otis Lake, Barry County, Michigan (Kopenski, 1969;
Klemm, 1972a,b, 1977), and I have examined specimens collected in southern
Illinois, (cf. Notes on Systematics).
131
-------
140 13D 120 110 ICO 90 80
10
70
60
MAP 28. "Dina anoculata
Dina dubia
* Dina parva
Comments:
£, anoculata is not a well known species (cf. Notes on Systematics).
Moore (1898) collected specimens (USNH 4844) from a mountainous area of
San Diego County, California. He also identified specimens (USNM 36938
and USNM 36874) from Oregon at elevations of 2,830 and 3S287 feet.
Scudder and Mann (1968) collected a single specimen from a lake in the
Southern Interior Plateau region of British Columbia, but I consider this
a doubtful record.
J), dubia is widely distributed throughout the northern United States,
including Alaska and Canada, but it is collected infrequently.
£. parva is widespread throughout the northern United States and Canada,
but it is not frequently encountered.
132
-------
UP 15B HO !30 1ZO 113 1BO SO 80 70 GO 50 40
30
60
MAP 29. "Erpobdella punctata coastal is • Erpobdella punctate punctata
Comments:
JL» £• coastal is is known only from the coastal areas of the Carolines
(Sawyer and Shelley; 1976).
E_. p. punctata is common throughout most of the United States and
Canada, especially the Great Lakes region. This species is also reported
from Mexico (Caballero, 1941; Sawyer, 1972).
133
-------
160
150
140 130 120 110 100 SO 80 70 6D
50
40
30
20
TO
BO
MAP 30.
Mooreobdella bucera
• Mooreobdel 1 a fervida
Mooreobdella melanostoma
Comments:
M_. bucera is uncommon and reported only from one locality in
southeastern Michigan (Kenk, 1949; Klemm, 1972a,b, 1977; Sawyer, 1972).
M. fervida is common in some localities but widely distributed in the
northern half of the United States and over much of Canada.
M. melanostoma has been reported from the Carolinas (Sawyer and Shelley,
1976)*;; I have examined specimens from Massachusetts, North Carolina and
Louisiana in the United States and Quebec in Canada.
134
-------
160
ISO
UO 130 120 110 100 90 80 70 60 50
\"
40
30
20
10
70
60
MAP 31.
Mooreobdella microstoma
Mooreobdella tetragon
Comments:
M. microstoma is common and widespread throughout the southern United
States; it is less common in the northern United States and Canada. The
species, Hooreobdella ochotherenai (Caballero, 1932) which closely
resembles M. microstoma, lives in Mexico (Sawyer & Shelley, 1976).
!!• tetragon is known only from the Atlantic Coast areas and Gulf States
of the United States. I have examined specimens from Massachusetts, New
Jersey, North Carolina, Georgia, Florida, and Alabama.
135
-------
60
ISO 150 140 13D 120 110 100 90 80 70 6D 50 40
30
20
10
BO
70
60
MAP 32. * Nephelopsi's obscura
Comments:
M^ obscura 1s widely distributed In the northern half of the United
States, Alaska, and Canada, and It Is common in some localities.
136
-------
SECTION 9
PARTIAL SYNONYMY
The section which follows contains the majority of important and
widely used synonyms found in the literature. An authority on which the
synonymic placement is based is listed beside each species name. The
primary sources for the synonyms are adopted from Harding (1910), Autrum
(1936), Klemm (1972b, 1977), Sawyer (1972), Sawyer, et al. (1975), Soos
(1965, 1966as 1966b, 1969as 1969b).
FAMILY GLOSSIPHONIIDAE
Actinobdella annectens Moore, 1906
Actinobdella inequiannulata Moore, 1901
Actinobdella triannulata Moore, 1924
Actinobdella triannulata Daniels and Freeman, 1976; Klemm, 1977;
Ringuelet, TM5
Alboglosslphonia heteroclita (Linnaeus, 1761)
Hirudo heteroclita Linnaeus, 1761
Hfrudo hvalina O.F. Muller, 1774
Hirudo ~swampina Bosc, 1802
Hirudo paplllgsa Braun, 1805
Hi rudo tr1ocu1 ata Carena, 1820
Hirudo "arcuata Fabricius, 1826
Clepsine~hyalina Moquin-Tandon, 1827
Clepsine carenae Moquin-Tandon, 1827
Glossobdella hyalina de Blainville, 1827
Glossobdella triocufata de Blainville, 1827
GlossobdeTTa carenae de Blainville, 1828
Clepsine carenae de Filippi, 1839
Clepsina Tiyalina Brightwell, 1842
Glossiphonla heteroclita Moquin-Tandon, 1846
Glossoppra hyalina Johnston, 1846
GlosslpholTia carenae Moquin-Tandon, 1846
Glossiphonla arcuata Moquin-Tandon, 1846
Clepsine swamplna Diesing, 1850
Clepsine papillosa Grube, 1851
Glossiphonla hyaTTna Thompson, 1856
Clepsine paTlida var. a Verrill, 1874
Clepsine heteroclita WTfttman, 1878
137
-------
Clepslne strlata Apathy, 1888
Clepsine palom'ca Lindenfeld and Pietrusuzynskl, 1890
Glossiphorna tnoculata Blanchard, 1893
GlosspslpHonla heterpcllta Blanchard, 1894
Glossiphonia heterocllta Castle, 1900
Glosspslphpm'a heterocllta Harding, 1910
Glosslphoni'a heterpcllta Ryerson, 1915
Glossiphonia pappllosa Pawlowskl, 1936
Glossi'phom'a nyallna Pawlowsk 1, 1936
GlosslphonTa striata Pawlowski, 1936
Glossi'phom'a swampi'na Sawyer, T973
GlosslphonTa swampina Sawyer & Shelley, 1976; Ringuelet, 1980
Glossiphonia LAIboglosslphonla] heterocllta Luken, 1976
Batracobdella cryptobranchli Oohnson and Klemm, 1977
Batracobdella michlganensls Sawyer, 1972
Batracobdella paludosa (Carena, 1824)
Hlrudo paludosa Carena, 1824
Clepsine paludosa Moquln-Tandon, 1827
Glossobdella paludosa de Blainville, 1828
Clepsfne succfnea de Ff7fppf, 1837
Clepsine paludosa de Filippl, 1839
Glosslphom'a paludosa Moquin-Tandon, 1846
Glossiphom'a succinea Moquln-Tandon, 1846
Glossiphonia paludosa Blanchard, 1894
Glossiphonia smargadina Oka, 1910
^emlcTepsIs "geel Oka, 1926
Clepsimdes paludosa Augener, 1892
HemlclepsTs" smargaoTna Oka, 1928
Batrachopflella paludosa Johansson, 1929
Batracobdella paludosa Autrum, 1936
Batracobdella phalera (Graf, 1899)
Clepslne phalera Graf, 1899
Placobdella phalera Moore, 1906
Placobdella phaleria Mullln, 1926
HaementerTaT LPIacoD^eTla] phalera Autrum, 1936
Batracobdella phalera Moore, 1959
Batracobdella plcta (Verrlll, 1872)
Clepslne picta Verrlll, 1872
Ila pic
Placobdella picta Moore, 1906
PlacobdelTa picta Ryerson, 1915
Glosslphonla picta Moore, 1923
Haementarla LPIacobdella] picta Autrum, 1936
Batrachobdella picta Richardson, 1949; Moore, 1952; Beck, 1954
Batrachobdella picta Meyer and Moore, 1954
138
-------
Bactacobdella picta Mann, 1962
Boreobdella verrucata (Fr. Muller, 1844)
Clepsine verrucata Fr. Muller, 1844
Glossiphonia verrucata Johnston, 1865
Glosslphpm'a moTh'ssima Moore, 1898
?GlQss1phon1a gram'fers Johnston, 1865
?G1ossiphgnia verrucata Johansson, 1909
Boreobdella verrucata Pawlowski, 1936
Batracobdella verrucata Autrum, 1936
Glosslphonia complanata molllssima Moore & Meyer, 1951; Klemm,
1972D
Borebodella verrucata Soos, 1969
Glossiphonla complanata (Linnaeus, 1758}
Hirudo sexoculata Bergmann, 1757
Hirudo complanata Linnaeus, 1758
Hirudo laterlbus attenuatis Hill, 1759
Hirudo crenata Kirby, 1794
Hirudo crinata Pennant, 1816
Glossiphonia tuberculata Johnson, 1816
Glossiphonia complanata Johnson, 1816
Glossopora tuberculata Johnson, 1817
Erpobdella complanata de Blainville, 1818
Gossopora~"complanata Fleming, 1822
Clepsi'ne complanata Savigny, 1822
Sangui'suga complanata Bruguiere, 1824
Glossobdella complanata de Blainville, 1828
Erpobdella crenata Tempieton, 1837
Glosslppra" tuberculata Thompson, 1841
Glosslpholh'a sexoculata Moquin-Tandon, 1846
Gossiphom'a cimiciformls Baird, 1869
Clepsine "pallida var. b Verrill, 1872
Clepsine elegans VerriTl, 1872
Clepsine patellifprmis Nicholson, 1873
Clepsine elegans Verrlll, 1874
Clepsine pam'da Verrill, 1875
Clepsine sex-puncto-lineata Sager, 1878
Clepsine sabartensis OrleyT 1886
Clepsine sexoculata Apathy, 1888
Glossiphonia palllda Vaillant, 1892
Glossiphom'a cpmplanata Blanchard, 1894
GlossiphonTa elegans Castle, 1900
Glossiphom'a concolor Johansson, 1909
Glossosiphonia cpmplanata Harding, 1910
Glosspsiphpm'a elegans Pinto, 1923
Glossiphonia complanata Moore, 1901
Glosspsiphom'a mollissina Pinto, 1923
Glossiphonia paludosa Ussing, 1929
Glossophiona complanata Mason, etal., 1970
139
-------
Glosslphom'a [Glosslphonia] complanata Lukin, 1976
Helobdella elongata (Castle, 1900)
Clepsine nepheloidea Graf, 1899 (nomen nudum)
Glosslphom'a elongata Castle, 19(50
Glosslphonia nepheloidea Moore, 1906
Glpssqsiphom'a elongata Pinto, 1923
Helobdella nepheloidea Moore, 1924
Helobdelta" elongata Autrum, 1936
Glosslphom'a nepheToldae Miller, 1937
Helobdella fusca (Castle, 1900)
Glosslphom'a fusca Castle, 1900
Glosslphonia fusca fusca Moore, 1906
Glosslphom'a Utelobdellaj fusca Moore, 1922 (? in part)
HelobdellaTusca Moore, 1$59
Helobdella" tri serial is Soos, 1969; Davies, 1971 (in part)
Helobdella papillate (Castle, 1900)
Clepsine papillifera var. b Verrill, 1824
Helobdella fusca var. papiTlata Moore, 1906
Helobdella fusca Moore, 1918 (in part)
Helobdella paplllata Moore, 1952
Helobdella stagnalis (Linnaeus, 1758)
Hirudo biocluata Bergmann, 1757
Hirudo "stagnalis Linnaeus, 1758
Hirudo Moculata 0. F. Muller, 1774
Hirudo pulllgera Daudin, 1800
Hirudo "ciirculans Sowerby, 1806
Helluo THirudo] bioculatus Oken, 1815
Glosslphom'a per'ata Johnson, 1816
Glossoppra punctata Johnson, 1817
Erpobdella Dioculata de Blainville, 1818
Clepsine bipculata Savigny, 1822
Glossopora bloculata Fleming, 1822
Clepsine sowerbyi Moguln-Tandon, 1827
Hirudo [Glosspbdella] pulligera de Blainville, 1827
Glossobdella bloculata de Blainville, 1828
Erpobdella stagnalis Tempieton, 1836
Clepslna stagnalis de Filippi, 1837
Glosslphom'a circulans Moguin-Tandon, 1846
Glosopora circularis Johnston, 1846
Glossiphonia bloculata Moguin-Tandon, 1846
Glosslpora bloculata Thompson, 1856
Clepsine Til Ippl Polom'o, 1863
Clepsine modesta Verrill, 1872
Clepsine submodesta Nicholson, 1873
140
-------
Clepsine viridissima Picaglia, 1877
Clepsine bioculata Levi'nsen, 1883
Glossiphonia modesta Vaillant, 1890
Glgsspsiphonia stagnalis Blanchard, 1894
Helobdelja stagnalis Blanchard, 1896
Helobdella" bioculata Bayer, 1898
Glossiphom'a stagnaTis Moore, 1898
Glossiphonia sraagnalis Castle, 1900
Glossiphom'a LHelobdella] stagnalls Moore, 1922
TiTpssqppra p'unctata Johnston, 1925
Helobdeltir stagnalls Andre, 1930
BakedebdeTla gibbpsa" Sciacchitana, 1939
Erpobdella stagnalls Oliver, 1958
Helobdel1"a stagnaTTs" Moore, 1952
Helobdella trans versa Sav/yer, 1972
Helobdella triserialis (E. Blanchard, 1849}
Glossiphom'a trjseri'ali's Blanchard, 1849
Clepsine trlsen'alls Grube, 1859
Clepsine llneolata Grube, 1871
Clepslne papilllfera var. b Verrill, 1872
Clepsine papilllfera var. Tineata Verrill, 1874
Glpsslphpm'a lineoTata Vaillant, 1890
Helobdella trl sen'alls Blanchard, 1896
GlosslphoTa lineata Moore, 1898
Glossiphonla fusca"Cast!e, 1900
Glosslphonia fusca 11neata Moore, 1906
Anoculobdella tntuberculata Weber, 1915
Placobdella triserialis Apathy, 1917
Glossiphom'a fusca Moore, 1918 (in part)
Glosslphom'a LHelobdella] fusca Moore, 1922 (in part)
Glossosiphona fusca Pinto, 1923
Helobdelja fusca Moore, 1924
He!obdei1 a punctata-1ineata Moore, 1939
HelobdelTa" m'^ricans Ringuelet, 1943
Helobdella triserial is lineata Ringuelet, 1943; 1980
Helobdella striata RTnguelet, 1943
Helobdella unilineata Ringuelet, 1943
Helobdella' lineata 5oos, 1969; Davies, 1971 (in part)
Helobdella fusca Soos, 1969 (in part)
HelobdelTa trialbolineata Klemm, 1974
Helobdella 11neata Klemm, 1977
He]obdel1 a punctata-1i neata Moore, 1939; 1959
Helobdella punctatalineata Klemrn, 1972b; 1976;1977
HelobdelllT punctatolineata' Sawyer and Kinard, 1980
Marvinmeyeria lucida (Moore, 1954)
Oculobdella lucida Moore, 1954
Harvinmeyeria lucfda Soos, 1969
141
-------
Oligobdella bi annul ata (Moore, 1900)
Microbdella biannulata Moore, 1900
OligobdelTa' biannulata Moore, 1918
Placobdella hollensis (Whitman, 1892)
Clepslne hollensis Whitman, 1892
Placobdella hollensis Moore, 1906
HaementerTa LParabdeTla] hollensis Autrum, 1936
ParabdellaThpllensis Meyer and Moore, 1954
Placobdella hollensTs Moore, 1952
Placobdella montifera (Moore, 1906)
?Glossiphom'a trisulcata Balrd, 1869
Clepslne papillifera var. carinata Verrill , 1874
Hemiclepsis can nata Moore, 1901
PlacobdelTa montifera Moore, 1906
Haementaria LPlacobdella] montifera Autrum, 1936
PI acobdetTa montifera Moore, 1952
PlacobdelTa" parasitica Am1n, 1977 (Specimen examined) (nee P.
parasitica (Say,
Placobdella multilineata Moore , 1 953
Placobdella multilineata Sawyer, 1972; Sawyer and Shelley, 1976
(southern variety)
Placobdella nuchal is Sawyer and Shelley, 1976
Placobdella parasitica White, 1977 (Specimen examined) (nee P.
parasitica (Say,
Placobdella ornata (Verrill), 1872
Clepsine ornata Verrill, 1872
Clepslne ornata var. rugosa Verrill , 1874
Clejjsine ornata var. stellata Verrill, 1874
Glossiphonla parasitica var. rugosa Castle, 1900
P1acobdelTF"ruoosa Moore, 190i
Haementarla LPiacobdella] rugosa Autrum, 1936
PI acobdetTa' ornata Moore, 1952 (northern variety)
?Placpbde]Ta multilineata Beck, 1954
Placobdella multilineata Klemm, 1972a
Placobdella papillifera (Verrill, 1872)
Clepsine papi Hi fera Verrill, 1872
Placobdel I ja papilTT7era Moore, 1952
PI acobdelTa" jaapi 1 1 i fera Meyer and Moore, 1954
Placobdella papillifera Sawyer and Shelley, 1976
142
-------
?Haementeria officinal is Ringuelet, 1976
Placobdella parasitica (Say, 1824)
Hirudo parasltica Say, 1824
Clepsine parasitica Dleslng, 1850
Clepsine ornata var. rugosa Verrlll, 1872
Clepsine marmarata Sager, 1878
Clepsine chelydrae Whitman, 1889
Clepsine plana Whftman, 1891
Glosslphoma parasitica Moore, 1898
Glossiphpnia parasitica var. plana Castle, 1900
Placobdella parasitica Moore, 1901
GlossoslpHonia parasitica Pinto, 1923
Haementaria [PlacqbdenaJ parasitica Autrum, 1936
Placobdella parasitica Moore, 1952
Placobdella pedi'culata Hemingway, 1908
Haementeria [Placobdella] pedlculata Autrum, 1936
PlacobdelT? pediculata Moore, 1952
Placobdella translucens Sawyer and Shelley, 1976
Theromyzon bi'annulatum Klemm, 1977
Hemiclepsis occidentalis Moore, 1912
Protoclepsls occidental is Moore, 1922
Theromyzon occidentale Autrum, 1936
Theromyzon occidentalis Sooter, 1937
Theromyzon meyeri Moore, 1959, Sawyer, 1972
Theromyzon maculosum Klemm, 1972a,b; Davies, 1971, 1973
Theromyzon" blannuTatum Klemm, 1977
Theromyzon rude (Baird, 1869)
Glosslphom'a rudls Barid, 1869
Clepsine occidentali's Verrill, 1874
Glossiphonia occldentalis Vaillant, 1890
Theromyzon occidentalis Bere, 1929
TheromyzorT rude Hoore and Meyer, 1951
Theromyzon occidentale Meyer and Moore, 1954
Theromyzon tessulatum (0. F. Muller, 1774)
Hirudo tessulata 0. F. Muller, 1774
Hirudo tessulata Gmelln, 1788
Hirudo tesselata Bosc, 1802
Hirudo "tessulatum Braun, 1805
Erppbdella tessulata Fleming, 1822
NephelIs tessel1ata Savigny, 1822
Ichthyobdella tesselata de Blainville, 1828
143
-------
Erpobdella vulgarIs var. tessellata de Blainville, 1828
Clepslne Tanguinea F11ippl, 1837
Clepslne tessulata Fr. Muller, 1844
Erpobdella tessulata Thompson, 1844
Glosslpholna tesseTata Thompson, 1846
Glossiphoma eacheana Thompson, 1846
Glosslphonla tessellata Moguln-Tandon, 1846
Gpssi'pnom'a sangulnea Moquln-Tandon, 1846
Clepslne tessulatum Dieslng, 1850
Hlrudo yltrlna Dalyell, 1853
Glosslphorn'a eacheana Thompson, 1856
Haemocnarls eacheana Thompson, 1856
Gloss1pho"nTa yitrina'Johnston, 1865
Theromyzon pallens Phtlippl, 1867
HemiclepsTs tesseflata Vejdovsky, 1884
Clepstne "tesselata Weltner, 1887
Gosslphom'a tessuTatum Blanchard, 11892
HemiciepsTs tesselata Scharff, 1898
Protoclepsis tesselata Livanow, 1902
Protoclepsls tessellata Harding, 1910
Theromyzon tessulatum Pawlowskl. 1936
TheromyzolT tessulata~Autrum, 1936
Protoclepsis granata Endri'gkeit, 1940
FAMILY PISCICOLIDAE
Cystobranchus mammniatus (Malm, 1863)
Cystobranchus [Platybdella] mamm'llatus Malm, 1863
Cystobranchus meyerl Hayunga and Grey, 1976
Cystobranchus verrilli' Meyer, 1940
Cystobranchus virgim'cus Hoffman, 1964
Myzobdella lugubris Leidy, 1851
Ichthyobdella punctata Verrill, 1871 (In part)
?Icnthypbdella fundull Vern'11, 1872
Plsclcola 7un?ul1 Pratt, 1935
Illlnobdella alba Meyer, 1940
Illlnobdella elongata Meyer, 1940
Illtnobdella n'chardsom' Meyer, 1940
minobdena moorel Meyer, 1940
Mysobdella lubrlgls Pearse, 1948
Myzobdella fundim "Moore, 1952
MyzobdelI"? moorei Meyer and Moore, 1954
Ichthyobdella rapax Wass, 1972
Cystobranchus virginlcus Paperna and Zwerner, 1974 (Specimen
examlned)(nee C. vtrglm'cus, Hoffman, 1964); Sawyer, Lawler,
Overstreet, 1975
144
-------
Myzobdella lugubris Sawyer, Lawler, and Overstreet, 1975
Piscicola geometra (Linnaeus, 1758)
Hirudo alba perexigua piscibus adhearens Aldrovandus, 1602
Hirudo ore caudaque ampla Frisch, 1729
Hirudo teres extremitatibus dilatis Linn., 1746
Hirudo piscium Roesel, 1747
Hirudo geometra Linn, 1758
Hirudo "galearia Braun, 1805
Piscicofa piscium Blainville, 1818
Hacinocha'ris piscium Savigny, 1820
Piscicola geometra Moquin-Tandon, 1826
Ichtjiyobdella geometra Blainville, 1827
Ichthlobdella piscium Egidy, 1844
Piscicola percae Johnston, 1846
Piscicola piscium Tauber, 1879
Piscicola" lavereti Tauber, 1879
Piscicola" perspicax Olsson, 1893
Piscicola" lippa Olsson, 1893
Piscicola" volgensis Zykoff, 1903
Piscicola" geometra~"Harding, 1910
Piscicola milneri (Verrill, 1874)
Ichthyobdella milneri Verrill, 1874
Piscicola milneri Ryerson, 1915
Piscicola" milneri Meyer, 1940
Piscicola punctata (Verrill, 1871)
Ichthyobdella punctata Verrill, 1871 (in part)
Piscicola punctata Moore, 1912
Piscicola salmositica Meyer, 1945
Piscicola salmositica Meyer, 1946
Piscicola" salmonsitica Moore, 1959
Piscicolaria reducta Meyer, 1940
FAMILY HIRUDINIDAE
Haemopis grandis (Verrill, 1874)
Semiscplex grandis Verrill, 1874 (in part)
Haemopis grandis Moore, 1912
Mollibdella grandis Richardson, 1969; Davies, 1971; Klemm,
197Za,b; Ringuelet, 1980
Haemopis kingi Mathers, 1954
145
-------
Percymoorensis kingi Richardson, 1969; Klemm, 1972b
Haemopis lateromaculata Mathers, 1963
Haemopis latero-maculata Mathers, 1963
Percymoorensis lateromaculata Richardson, 1969; Davies, 1971;
Klemm, 1972b
Haemopis marmorata (Say, 1824)
Hirudo marmoratis Say, 1824
Democedes maculatus Kinberg, 1867
Aulastomum lacustris Leidy, 1869
Aulastomum lacustfe~Verril1, 1872
Hexobdella depressa Verrill, 1872
Aulostoma lacustris Forbes, 1893
Haemopis~sanquisuga Blanchard, 1896
Haemopis marmoratis Moore, 1912
Haemopis marmorafuT Hanklnson, 1908
Haemopis marmorata Moore, 1923
Haemopis marmoratis Hull in, 1926
Percymoorensis marmoratis Richardson, 1969
Haemopis marmorate Gates "and Moore, 1970
Haemopis marmorata Moore, 1952
Percymoorensis marmorata Davies, 1971; Ringuelet, 1980
Percymoorensis marmoratis Klemm, 1972a,b
Haemopis plumbea Moore, 1912
Haemopis plumbeus Moore> 1912
?Haemopis plumbeus Mullin, 1926
Haemopis plumbeous Miller, 1937
Haemopis plumbea"Moore, 1959
Bdellarogatis plumbeus Richardson, 1969; Davies, 1971
Bdellarogatis plumbea Klemm, 1972a,b; Ringuelet, 1980
Haemopis septagon Sawyer and Shelley, 1976
Haemopis terrestris (Forbes, 1890)
?Hirudo lateralis Say, 1824
?Au1acostomum oenops Grube, 1871
Semiscolex terrestris Forbes, 1890
Haemopis Tateralis Moore, 1898
Haemopis lateral!? terrestris Moore, 1918
Haemopis lateralis" Miller, 1937
Percymoorensis lateralis Richardson, 1969; Davies, 1971; Klemm,
1972a,b
Haemopis terrestris Sawyer, 1972
Percymoorensis terrestris (Forbes, 1890) Ringuelet, 1980
Hirudo medicinal is (Linnaeus, 1758)
146
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Hirudo major et van'a Gesner, 1558
Hirudo varla Aldrovandus, 1602
Hirudo minor variegata Mural to, 1685
Hirudo medicinal Is Ray, 1710
Hirudo medicinal fs" Linnaeus, 1758
Hirudo venesector Braun, 1805
Hirudo venaestector Carena, 1820
Hirudo "verbana Cafena, 1820
Hirudo "provincialis Carena, 1820
Sanguisuga medicinal is Savigny, 1822
Sanguisuga officlnalis Savlgny, 1822
Hirudo officinal is Derhein, 1825
Sanguisuga offlcinalls Moquln-Tandon, 1826
Sanguisuga obscura Moquin-Tandon, 1826
Sanguisuga verbana Moquin-Tandon, 1826
Sanguisuga" meridional 1s Risso, 1826
Sanguisuga carena Risso, 1826
n"
carena Risso,
TatrobdelTa LHirudo] medicinal is
de Blainville, 1827
1833
Sanguisuga chlorgaster Brandt, 1833
Sanguisuga provincialis Brandt and Ratzeburg,
Macrobdella decora (Say, 1824)
Hirudo decora Say, 1824
Hirudo ornata Ebad, 1857
Macrobdella decora Verrill, 1872
Macrobdella diplotertia Meyer, 1975
Macrobdella ditetra Moore, 1953
Macrobdella sestertia Whitman, 1886
Macrobdella testertia Moore, 1953
Philobdella floridana Verrill, 1874
Macrobdella [Philobdella] floridana Verrill, 1874
PhilobdellT floridana Moore, 1901
Philobdella gracills Moore, 1901
Philobdella floridana Moore, 1898 {in part)
PhilobdelTa grac1lTs~Moore, 1901
PMlobdella graclTTs Moore, 1952
PhilobdeTTa" graciTTs' Viosca, 1962
Phllobdella gracllfs Richardson, 1972
FAMILY ERPOBDELLIDAE
_D_ina anoculata Moore, 1898
147
-------
Dina dubia Moore and Meyer, 1951
Dina parya Moore, 1912
Erpobdella punctata coastal is Sawyer and Shelley, 1976
Erpobdella punctata punctata (Leidy, 1870)
Nephelis punctata Leidy, 1870
Nephelis lateral is Verrlll, 1871 (in part)
NepheMs quadrestriata Verrill, 1872 (in part)
Nephelis marmorata Verri 11,1872
?NepheTts yenm'formis Nicholson, 1873
?Nephelis 4-striataTorbes, 1893
Nepnelfs alterais Brfstol, 1898 (in part)
Herpobcfella punctata Moore, 1898
ErpobdelTa punctata Moore, 1901
Erpobdetta" punctata annulata Moore, 1922
?Dfna fery'ida Miller, 1929
Herpbbdel1a punctata Meyer, 1937
Dina lateral is Moore. 1952
Erpobdella lateral is Moore, 1952; 1959
Erpobdella puctata Mann, 1961
ErpobdellT annulafa Mason and Gates, 1970
Erpobdella punctata annulata Klemm, 1972a,b; Sawyer, 1972;
RlngueYet, 1980
Erpobdella punctata punctata Sawyer and Shelley, 1976
Mooreobdella bucera (Moore, 1949)
Dina bucera Moore, 1949
Mooreobdella bucera Moore, 1959
Dina LMoofeobdellaj bucera Klenm, 1972a,b
Mooreobdella fervida (Verrill, 1781)
Nep_h_elis feryida Verri 11. 1871
NephelIis yermiformis Nicholson, 1873 (in part)
TmTa fervida Moore, 1901
TTTna TervTga Mathers. 1945
Hooreobde'lla fervida Pawlowski, 1955
Dina [Mooreobdella] fervida Klemn, 1972a,b
Mooreobdena melanostoma Sawyer and Shelley, 1976
Mooreobdella microstoma (Moore, 1901)
Dina micrpstoma Moore, 1901
Erpobdella [Mooreobdella] microstoma Pawlowski, 1955
Mooreobde'na microstoma Moore, 1959
Mooreobdella microstoma Soos, 1966
Dina LHooreobdellaJ microstoma Klemm, 1972a,b
148
-------
Mooreobdella tetragon Sawyer and Shelley, 1976
Nephelopsis obscura Vern"11, 1872
?Nephells obscura var. macu]ata Forbes, 1892
?Nephells maculuta Forbes, 1893.
149
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-------
SELECTED BIBLIOGRAPHY
This bibliography lists all the original and important taxononric
references for scientific names of taxa now accepted as valid for the
leeches of North America. Other references listed are cited throughout
the sections of this manual. References proceeded by an asterisk (*) are
additional distribution records or are not included in Klemm (1972b).
*Aliff, J.V., D. Smith, and H. Lucas. 1976. Some metazoan parasites from
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1977. Some metazoan parasites from fishes of middle
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*Amin, O.M. 1977. Helminth parasites of some southwestern Lake Michigan
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* . 1981. Leeches (Hirudinea) from Wisconsin, and a description
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Anderson, C. 1962. Noen iakttagelser over biologien til borsteiglen,
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quences their interlake transfer through a proposed water diversion
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151
-------
Baird, W. 1869. Descriptions of some new suctorial annelides in the
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*Basch, R.E., C.H. Pecor, R.C. Waybrant, and D.E. Denaga. 1980.
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* . 1975b. Notes on some Louisiana freshwater leeches
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155
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., and M.F. Vidrine. 1976. New freshwater mussel host records
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* ., and M.F. Vidrine. 1977. New freshwater clam host records
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. 1972. Annotated bibliography to the freshwater leeches
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156
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Antlcostl Island, Quebec. Can. Field-Nat. 93(3): 310-313.
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173
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INDEX TO SCIENTIFIC NAMES
Acanthobdella, 11, 23, 31, 105
1ivanowi,~?3
peledina. 11, 23, 31, 105
Acanthobdel H dae, 11, 31, 95
Actlnobdella, 11, 25, 51, 96
annectens, 11, 51, 52, 105, 137
Inequl annulata, 11, 51, 52, 105,
T37
Alboglossiphom'a (Glosslphom'a),
11, it), 59, T37
heteroclita, 11, 15, 16, 23,
27, 36, 59, 60, 102, 107, 137
annectens, Actinobdella, 11, 51,
5?7105, 137
annulata punctata, Erpobdella, 1 9 ,
anoculata, DTna, 1 3, ZO, 23, 84 ,
5S7 877T32, 147
atomaria, Erpobdella, 104
Batracobdel la, 11
cryptobra"nc"hii, 11, 25, 51, 53,
108, 138
michlganensis, 11, 50, 108, 138
paludosa, 11, 23, 36, 59, 103,
TW7138
phalera, 11, 49, 50, 108, 138
pjcta, 11, 53, 56, 101, 109, 138
Bdellarogatis (Haemopls), 18
plumbea, T?
bi annulata, Ollgobdella, 12, 33, 55,
55775 , 102, 115, 1 42
biannulatum, Theromyzon , 12, 16, 17,
6TT52, 10Z, 127, 143
Boreobdella, 11, 59
verrucata. 11, 23, 36, 60, 61,
mn39
bucera, Hooreobdella, 14, 19, 83, 84
55, 87, 88, 134, 148
Calliobdella, 18
T8"
caballeroi, Percymoorensis, 127
(Haemopis), 18
coastal is punctata, Erpobdella, 83,
5*7~85, 86, 102, 133, 148
complanata, Glosslphom'a, 11, 15,
23, 24, 36, 60, 61, 101, 102,
110, 139
cryptobranchll, Batractobdella, 11,
25, 51, 53, 108, 138
Cystobranchus, 2, 12, 25, 65, 99
mamnrillatuT, 12, 67, 68, 121, 144
meyeri, 12,' 66, 67, 97, 121, 144
verrnii, 12, 39, 67, 68, 122, 144
vlrglnlcus, 12, 18, 66, 67, 122,
m
vi'vidus, 18
decora, Macrobdella, 13, 28, 70,
71, 73, 102, 129, 147
Dina, 13, 19, 21, 28
anoculata, 13, 20, 23, 84, 85,
87, 132, 147
dubia, 13, 83, 90, 91, 93, 101,
T32, 147
lateral Is, 19
parva, 13, 83, 90, 91, 93, 102,
"^TJ2, 147
diplotertia, Macrobdella, 13, 28,
70, 7l 73, 129, 147
ditetra, Macrobdella, 13, 28, 70,
7T, 72, 102, 130, 147
dubia, Dina, 13, 83, 90, 91, 93,
107TT32, 147
elongata, Helobdella, 12, 33, 45,
T57 111, 140
Erpobdella, 14, 19, 28
atomaria, 104
lateralIs, 19
octoculafa, 104
punctata,~14, 19
annulata, 19
coastal is, 14, 83, 84, 85, 86,
102, 133, 148
punctata, 14, 19, 83, 84, 85,
86, 87, 133, 148
triannulata, 102, 103, 104
Erpobdellidae, 6, 13, 19, 29, 40,
44, 83, 96, 144
fervida, Mgoreobdella, 14, 19, 83,
S?, 88, 89, 90 102, 134, 148
174
-------
floridana, Philobdella, 13, 18S 19,
74, 75, 131, 147
fusca, Helobdella, 12, 16, 45, 46
111, 140
geometra, Piscicola, 13, 17, 69,
TT5?, 123, 145
Glossiphom'a (Glossiphonia), 136
Glossiphom'a, 11, 15, 59
complanata, 11, 15, 23, 24, 36,
60, 61, 101, 102, 110, 139
heteroclita (Alboglosslphonia),
11, 15
swampina, 15, 16
Glossiphoniidae, 11, 15, 33, 34, 35,
36, 97, 137
gracills, Philobdella, 13, 18, 19,
707 74, 75, 102, 131, 147
grandis, Haemopis, 13, 18, 26, 27,
40, 79, 81, 82, 102, 126, 145
(Mollibdella), 18
Haementeria,
pfficinaTis, 104
Haemppis, 6, 13, 18, 21, 26, 74, 96
caballeroi fPercymgorensis), 127
grandis, 13, 18, 26, 27, 40, 79,
81, 82, 102, 126, 145
kingi, 13, 18, 77, 78, 126, 145
Kingi
Tateri
ateromaculata, 13, 18, 77, 80,
127, 146
marmorata, 13, 18, 27, 70, 77, 79,
80, 127, 146
plumbea, 13, 18, 26, 40, 79, 81,
5?7~102, 128, 146
septagon, 6, 13, 18, 70, 74, 76
128, 146
terrestrls, 6, 13, 18, 70, 77, 78
128, 146
Helobdella, 12, 24, 43,
elongata, 12, 33, 45, 46, 111, 140
Tusca, 12, 16, 45, 46, 111, 140
papillata, 12, 24, 45, 47, 102,
112, 140
punctatolineata, 16
stagna1is,"T?7"43, 44, 99, 113,
™
transversa, 12, 45, 46, 113, 141
trlserTalTs, 12, 16, 24, 32, 45,
47, 48, 114, 141
heteroclita, Alboglossiphonia, 11,
15, 16, 23, 27, 36, 59, 60,
102, 107, 137
(Glossiphom'a), 15
Hirudinidae (Hirudidae), 6, 13, 18,
40, 41, 70, 97, 145
Hirudo, 13, 96, 97
medlclnalis, 13, 74, 103, 146
hollensls, Placobdella, 12, 36,
2F?7~43, 44, 95, 115, 142
inegui annulata, Acti nobdel1 a, 11
51, 52, 10S7T37
kingi, Haemopis, 13, 18, 77, 78,
126, 145
(Percymoorensis), 18
lateralI's, Dina, 19
lateral is", Erpobdella, 19
1ateromaculata, Haemopis, 13, 18,
77, 80, 127, 146
(Percymoorensis), 18
1ucida, Marvinmeyeria, 12, 27, 43,
£4, 102, 112, 141
livanowi, Acanthobdella, 23
lugubris, Myzobde11arT2, 25, 38,
39, 64, 65, 123, 144
Macrobdella, 13, 18, 70, 96
decora, 1'3, 28, 70, 71, 73, 102,
T?S, 147
diplotertia, 13, 28, 70, 71, 73
129, 147
ditetra, 13, 28, 70, 71, 72, 102
T307 147
sestertia, 13, 28, 70, 71, 72,
102, 130, 147
maculpsum, Theromyzon, 16
mammillatus, Cystobranchus, 12, 67,
68, 121,~TP
marmorata, Haemopis, 13, 18, 27, 70,
777~79y 80, 127, 146
(Percymoorensis), 18
Marvinmeyeria, 1?
1ucida, 1?7 27, 43, 44, 102, 112,
TTT
medicinal is, Hirudo, 13, 74, 103,
U5
melanostoma, Mooreobdella, 14, 19,
83, 86, 102, 134, 148
meyeri, Cystobranchus, 12, 66, 67,
57, 121, 144
michiganensis, Batracobdella, 11,
50, 108, 1"38
microstoma, Mooreobdella, 14, 19,
8TT34, 88, 89, 135, 148
milneri, piscicola, 13, 17, 69,
TTT2, 125, 145
Mollibdella (Haemopis), 18
175
-------
grandIs, 18
montlfera, Placobdella, 12, 25, 33,
*?7 96, 116, 142
Mooreobdella, 14, 19, 21, 28
bucera, Tfr, 19, 83, 84, 85, 87,
837 134, 148
fervida, 14, 19, 83, 84, 88, 89,
5U7102, 134, 148
melanostoma, 14, 19, 83, 84, 86,
102, 134, 148
mlcrostoma, 14, 19, 83, 84, 88,
89, 135, 148
tetragon, 14, 19, 83, 88, 89,
TOT135, 149
multilIneata, Placobdella, 12, 24,
57, 58, 102, 116, 142
Myzobdella, 12, 25, 63
lugubrls, 12, 25, 38, 39, 64,
65, 123, 144
patzcuarensls, 123
Nephelopsis, 14
obscura,"l4, 28, 83, 90, 91,
9?H02, 136, 149
nuchal is, Placobdella, 12, 25, 33,
*?7 96, 117, 142
obscura, Nephelopsls, 14, 28, 83,
9ff, 91, 92, 102, 136, 149
occidental Is, Theromyzon, 16
octoculata, Erpobdella, 104
offlci'alf?, HaementerTa, 104
011gobdena,~T2
biannul afa, 12, 33, 55, 56, 95,
102, 115, 142
ornata, Placobdella, 12, 24, 56,
57, 102, 117, 142
paludosa, Batracobdella, 11, 23,
357 59, 103, 109, 138
papillata, Helobdella, 12, 24, 45,
2F77~"IOZ, 112, 140
papillifera, Placobdella, 12, 24,
57, 58, 104, 118, 142
parasltlca, Placobdella, 12, 54,
parva, Dlna, 13, 83, 90, 91, 93,
10I7T32, 147
patzcuarensls, Myzobdel1 a, 123
pedlculata, Placobdella, 12, 25,
3571)1, 53, 118, 143
peledina, Acanthodella, 11, 23, 31,
Percymoorensis (Haemopls), 18
caDalleroi, 127
klngi, 18
marmorata , 18
lateromaculata, 18
terrestrls, 18
phalera, Batracobdella, 11, 49, 50,
TU8,~735
Phllobdella, 13, 18, 19, 28, 71, 96,
florldana, 13, 18, 19, 74, 75, 131,
T77
gracilis, 13, 18, 19, 70, 74, 75,
102, 131, 147
pi eta, Batracobdella, 11, 55, 56,
101, 109, 138
Plsclcola, 13, 25, 63, 65, 99
geometra, 13, 17, 69, 102, 123, 145
mllnerl, 13, 17, 69, 102, 125, 145
punctata, 13, 67, 68, 124, 145
salmosltlca, 13, 68, 69, 96, 124,
TC3
zebra, 17
Plsclcolaria, 13, 25, 63
reducta.TS, 25, 33, 38, 39, 64,
T257 145
PisclcoHdae, 6, 12, 17, 33, 37, 38,
66, 98, 144
Placobdella, 12, 24, 98
hollensls, 12, 36, 42, 43, 44, 95,
115, 142
montlfera, 12, 25, 33, 42, 96,
116, 142
mul til Ineata, 12, 24, 57, 58, 102,
116, 142
nuchal Is, 12, 25, 33, 42, 96,
TT77142
ornata, 12, 24, 56, 57, 102, 117,
~
papllllfera. 12, 24, 57, 58, 104,
118, 142
parasltlca, 12, 54, 55, 119, 143
pediculata, 12, 25, 39, 51, 53,
118, 143
translucens, 12, 50, 102, 119, 143
plumbea, Haemopis, 13, 18, 26, 40,
75, 81, 82, 102, 128, 146
(Bdellarogatis), 18
proplnquus, Theromyzon, 17
punctata punctata, Erpobdell a, 14,
T?7 83, 84, 85, 86, 87 133, 148
annul ata, 19
coastal Is, 14, 83, 84, 85, 86,
102, 133, 148
punctata, Plsclcola, 13, 67, 68,
176
-------
124, 145
punc-tatolineata, Helobdella, 16
reducta, Piscicolaria, 13, 25,
33", 38, 39, 64, 125, 145
rude, Theromyzon, 12, 16, 17, 62,
102, 120, 143
salmosltica, Pisclcola, 13, 68,
69, 96, 124, 145
septagon, Haemopi's, 6, 13, 18, 70,
74, 76, 128, 146
sestertia, Macrobdella, 13, 28, 70,
7T7"72, 102, 130, 147
stagnalis, Helobdella, 12, 43, 44,
99, 113, 140
swampina, Glpsslphom'a, 15, 16
terrestris, Haemopis, 6, 13, 18, 70,
77778, 128, 146
(Percymoorensis), 18
tessulatum, Theromyzon, 12, 16, 17,
S?7T3, 10Z, 120, 143
tetragon, Mooreobdella, 14, 19, 83,
5S7 89, 10Z, 135, 149
Theromyzon, 12, 16, 17, 27, 36, 59,
TGT~
biammulatum, 12, 16, 17, 61, 62
102, 120, 143
maculosum, 16
occldenta'lis, 16
proplnquus, 17
rude, 12, 16, 17, 62, 102, 120,
T43
tessulatum, 12, 16, 17, 62, 63,
102, 1ZO, 143
translucens, Placobdella, 12, 50,
102, H97T53
transversa, Helobdella, 12, 45, 46,
trl annul ata, Erpobdella, 102, 103,
HE
triserlalls, Helobdella, 12, 16, 24,
32, 45, 47, 48, 111, 114, 141
verrilli, Cystobranchus, 12, 39,
577 68, 1ZZ, 144
verrucata, Boreobdella 11, 23, 36,
^7"61, 110, 134
vlrginicus, Cystobranchus, 12, 18,
66, 67, 1ZZ, 144
vivlda, Calllobdella, 18
vlyldus, CystobrancFus, 18
zebra, Piscico1a,T7
177
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