United States
Environmental Protection
Agency
Environmental Monitoring and
Support Laboratory
Cincinnati OH 45268
EPA-600/3-82-033
April 1982
Research and Development
x>EPA
A Guide to the
Fresh water Tubificidae
(Annelida: Clitellata:
Oligochaeta) of
North America
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EPA-600/3-82-033
April 1982
A GUIDE TO THE FRESHWATER TUBIFICIDAE (ANNELIDA: CLITELLATA: OLIGOCHAETA)
OF NORTH AMERICA
by
Kurt S. Stimpson
Ecological Analysts, Inc.
Suite 306, 1535 Lake Cook Road
Northbrook, Illinois 60062
Donald J. Klemrn
Aquatic Biology Section
Environmental Monitoring and Support Laboratory
U.S. Environmental Protection Agency
Cincinnati, Ohio 45268
and
Jarl K. Hiltunen
3160 Pitts view Drive
Ann Arbor, Michigan 48104
ENVIRONMENTAL MONITORING AND SUPPORT LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U.S. ENVIRONMENTAL PROTECTION AGENCY
CINCINNATI, OHIO 45268
U.S. Environmental Protection Agency
Region 5, Library (PL-12J)
77 West Jackson Boulevark 12th Floor
Chicago, IL 60604-3590
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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.
11
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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 conducts research to develop, evaluate, 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,
peri phy ton, 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 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. Tubificid worms are important components of the
benthic fauna and are frequently abundant in a variety of freshwater
habitats. The varied response of tubificid species to different kinds of
pollution and toxic substances makes them very useful as water quality
indicator organisms. While several regional keys to North American
Tubificidae have been published, this manual is the first to contain an
illustrated key and the distribution for all taxa. The publication was
developed to assist aquatic biologists in evaluating data collected in
studies of the effects of toxic substances and other pollutants on the
communities of benthic macroinvertebrates.
Robert L. Booth
Acting Director
Environmental Monitoring and Support
Laboratory - Cincinnati
iii
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ABSTRACT
In North America, the freshwater annelid worms (Clitellata:
Oligochaeta), belonging in the family Tubificidae, are composed of 18
genera, 54 species, one subspecies, and several variant forms. All taxa
can be identified by external and internal morphological features. This
guide presents an introduction to the general biology of the Tubificidae,
collecting and processing methods, a species list, an artificial
illustrated key, a glossary, an annotated systematic list, and a selected
bibliography which includes the references cited in the text and other
publications which provide additional information on tubificid taxonomy
and ecology.
iv
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CONTENTS
Foreword Ill
Abstract iv
Figures v1
Acknowledgments x
1. Introduction 1
2. Methods 3
Collection, Preservation, and Preparation
of Tub1f1c1dae Whole Mounts 3
Depository for Tublflcld Worms 4
3. Species List 5
4. Key to the Freshwater Tub1f1c1dae of North America
(North of Mexico) 9
Introduction 9
Key to the Tub1f1c1dae 12
5. Glossary 41
6. Annotations 44
Selected Bibliography 53
Index to Scientific Names 60
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FIGURES
Number Pages
1 Generalized tubificid, illustrating external and internal
anatomical terminology, lateral view 10
?. Generalized tubificid, illustrating types of somatic and
genital chaetae and anatomical terminology 11
3 Branchiura sowerbyi, dorsal and ventral gill filaments on
posterior segments 13
4 Granular-opaque, densely papillate dermis of Spirosperma
nikolskyi 13
5 Quistadrilus multisetosus: (a,b) slightly opaque dermis with
scattered papillae 13
6 Clear dermis, without papillae 14
7 Ouistadrilus multisetosus, dorsal pectinate chaeta 14
8 £. multisetosus, posterior ventral chaeta 14
9 j). multisetosus: (a) anterior ventral chaeta; (b) posterior
ventral chaeta 14
10 Spirosperma ferox, dorsal pectinate chaeta 15
11 S^ beetoni: (a) anterior ventral chaeta; (b) posterior
ventral chaeta 15
12 _S. nikolskyi, posterior ventral chaeta 16
13 _S. carolinensis, posterior ventral chaeta 16
14 Ilyodrilus mastix, anterior end illustrating enlarged
mouth 16
15 Aulodrilus pigueti, posterior dorsal crotchet chaeta:
(a) facial aspect; (b) lateral aspect 17
16 Potamothrix vejdovskyi: (a) anterior dorsal crotchet and
capilliform chaetae; (b) spermathecal chaeta 18
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Number Pages
17 Rhyacodn'lus brevidentatus, dorsal crotchet chaeta 18
18 Aulodrilus plurlseta: (a) ventral crotchet chaeta;
(b) dorsal pectinate chaeta 19
19 Rhyacodrilus coccineus: (a) dorsal crotchet chaeta;
(b) penlal chaetae 19
20 Psammoryetides barbatus, pectinate anterior dorsal crotchet
chaeta 20
21 Rhyacodrilus montana: (a) anterior dorsal pectinate chaeta;
(b) anterior ventral crotchet chaeta 20
22 Tubifex harmani, posterior dorsal crotchet chaeta 21
23 Rhyacodrilus punctatus, anterior ventral chaeta 21
24 Tubifex nerthus, anterior ventral chaeta 22
25 T_. tubifex, penis bulb, atrium, and vas deferens 23
26 Haber cf. speciosus: (a) penis sheath; (b) penial chaeta;
(c) spermathecal chaeta 24
27 Potamothrix bedoti, specialized chaeta 24
28 Psammoryctides minutus, penial chaeta 25
29 Rhyacodrilus soda!is, dorsal pectinate chaeta 25
30 Psammoryctides convolutus, spermathecal chaeta 26
31 £. californianus, spermathecal chaeta 26
32 Potamothrix hammoniensis, spermathecal chaeta 26
33 £. bavaricus, spermathecal chaeta 26
34 Varichaeta pacifica: (a) penis sheath inverted;
(b) penis sheath everted 27
35 Tubifex superiorensis, penis sheath 28
36 Ilyodrilus templetoni; (a) penis sheath with distal end
absent; (b) penis sheath with distal end present 28
37 Tubifex kessleri americanus, penis sheath 28
38 Ilyodrilus frantzi form capillatus, penis sheath 28
vii
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Number Pages
39. Telmatodrilus vejdovskyi; (a) anterior ventral chaeta;
(b) posterior ventral chaeta 29
40 Aulodrilus amerlcanus, posterior dorsal crotchet chaeta .... 29
41 Aulodrilus limnobius, posterior dorsal crotchet chaeta:
(a) facial aspect; (b) lateral aspect 30
42 Monopylephorus he! obi us, crotchet chaeta 30
43 Bothrioneurum vejdovskyanum, SEM of prostomium showing
sensory cavity 31
44 B. vejdovskyanum; (a) ventral crochet chaeta of segment II;
Tb) ventral crochet chaeta of segment XXX; (c) penial chaeta. . 31
45 Isochaetides curvisetosus, posterior dorsal chaeta 32
46 Limnodrilus udekemianus: (a,b,c) ventral chaetae in segment
II, illustrating variation in shape 32
47 Isochaetides freyi; (a) penis sheath; (b) spermathecal
chaeta 33
48 Rhizodrilus lacteus; (a) specialized chaeta in segment IX;
{b) specialized chaetae in segment XI 33
49 Potamothrix moldaviensis: (a) spermathecal chaeta;
partially everted penis; (c) fully everted penis 34
50 Phallodrilus hallae, penial chaeta 35
51 Rhyacodrilus falciformis, penial chaeta: (a) lateral
aspect; (b) facial aspect 35
52 Limnodrilus rubripenis, penis sheath 37
53 L_. angustipenis, penis sheath 37
54 L_. hoffmeisteri (spiralis form), penis sheath 37
55 _L. psammophilus, penis sheath 37
56 U udekemianus, penis sheath 37
57 ^. profundicola, penis sheath 37
58 L. silvani, penis sheath. 39
viii
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Number Pages
59 ±. hoffmeisteri (variant form), illustrating two pern's
sheath types:(a) scalloped margin; (b) scalloped margin
with irregular distal projection 39
60 L_. hof fmei steri, penis sheath 39
61 L_. claparedianus, penis sheath 39
62 L_. maumeensis, penis sheath 39
63 L- cervix (variant form), penis sheath 39
6* i.' cervix, penis sheath 39
ix
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ACKNOWLEDGMENTS
The authors thank Ralph 0. Brfnkhurst, Thomas V. Clevenger, and
Michael S. Loden for their reviews of the technical contents of the
manuscript. We also thank Cornelius I. Weber for reading the manuscript
and making valuable suggestions. We are grateful to Kristian Fauchald of
the Smithsonian Institution, U.S. National Museum of Natural History,
Department of Invertebrate Zoology, Division of Worms, Washington, D.C.
for permitting us to examine the collections of Tubificidae. We are also
indebted to the following individuals for supplying specimens and
distribution records: Ralph 0. Brinkhurst, Dale F. Kriewall, Michael S.
Loden, Anthony F. Maciorowski, Bruce E. Markert, Bruno Vincent, and Mark
J. Wetzel. We extend special thanks to Douglas R. Spencer for his
contributions in early stages of preparation of the guide. We would also
like to thank Peter M. Chapman for the use of Figure 43.
The secretarial assistance of Cordelia Nowell and Diane Schirmann is
gratefully acknowledged.
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SECTION 1
INTRODUCTION
The relationship between benthic macroinvertebrate community
structure and the physical and chemical characteristics of aquatic
habitats has been examined in great detail in the past several decades.
The body of knowledge relating the qualitative and quantitative responses
of benthic communities to changes in habitat has grown to the point where
an examination of benthic community structure has become a valuable tool
for regulatory agencies, water resource managers, and aquatic ecologists
in assessing and monitoring water quality and detecting pollution
sources. The Oligochaeta, or segmented worms, are an important component
of the benthic community in nearly every freshwater biotope. The group
comprises 24 families, among which the Tubificidae are the most widely
distributed and abundant oligochaetes in freshwater systems. The utility
of the Tubificidae for monitoring and detecting changes in water quality
and physical habitats has been illustrated by several authors (Brinkhurst,
1966, 1970, 1974a,b; Lang and Lang-Dobler, 1980; Stimpson, et al.,
1975). Compared to other macroinvertebrate groups, the Tubificidae are
particularly well-suited for use in biological assessments of water
quality, water pollution, or other changes in aquatic ecosystems
resulting from natural causes or man's activities. Members of the family
live longer than many other invertebrate forms; are generally sedentary
infaunal forms and, as opposed to, for example, most aquatic insects, are
restricted to the aquatic environment throughout their life cycle. Most
importantly, the habitat and water quality requirements as well as the
pollution tolerance of many species have been documented in the
literature (Chapman, Parrel 1, and Brinkhurst, 1982a,b; Hiltunen, 1967,
1969a, 1969b; Howmiller and Beeton, 1970).
Unfortunately, the Tubificidae have often been overlooked or ignored
in sample processing or misidentified by investigators not familiar with
their taxonomy. In many instances, investigators have recorded the group
only as subclass (Oligochaeta), family, or merely as "worms." The
inadequate treatment of the family represents a loss of valuable
ecological information that may be attributed, at least in part, to the
misconception that the group is taxonomically very difficult, and to the
lack of a practical key to the species.
Thus, the guide was composed to assist the USEPA and other biologists
in Federal, state, and private water monitoring organizations in
identifying specimens of tubificid worms to species. The guide will aid
in the monitoring and detection of changes in water quality and the
effects of toxic substances and other pollutants on macroinvertebrate
1
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community structure. It will also facilitate an expansion of our
knowledge of the ecological requirements of some of the lesser known
tubificid species. The guide includes a discussion of methods for the
collection, preparation, and examination of specimens, a species list, an
artificial illustrated key, a glossary, an annotation for each species,
and a selected bibliography. Morphological features utilized in the
differentiation of species are defined in the glossary. The reader is
referred to Brinkhurst and Jamieson (1971) and Reynolds (1977) for
additional morphological terms. The selected bibliography was assembled
from literature pertaining directly to the systematics and ecology of the
Tubificidae of North America.
Inasmuch as new zoogeographical records are published frequently,
this guide may not include the latest North American discoveries. In
instances where a species is not keyed in the present work, the reader is
directed to consult Brinkhurst and Jamieson (1971), Aquatic Oligochaeta
of the World. Complete descriptions that may be helpful in confirming
the identity of most species in this guide, may also be found therein.
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SECTION 2
METHODS
COLLECTION, PRESERVATION, AND PREPARATION OF TUBIFICIDAE WHOLE MOUNTS
Tuhificidae may be collected using any of a variety of methods,
ranging from qualitative hand-picking and dip-netting to quantitative
dredge, grab or core sampling. Using most conventional collection
methods, it is necessary to reduce the volume of sediment in order to
discern and sort the organisms. Although Agency procedures for
macroinvertebrate studies (Weber, 1973) recommend screening the sample on
a U.S. Standard No. 30 mesh sieve (28 meshes per inch, 0.595 mm
openings), it should be noted that many small individuals may be lost
during the sieving process and that use of a finer sieve (for example,
No. 60 mesh) or no sieving at all may be required to insure collection of
all individuals. Great care should be taken in sieving samples
containing worms; too vigorous agitation may break specimens or damage
body parts required for species identification. Tubificidae collected by
selective hand-picking in the field may be fixed and preserved by placing
them directly in 5% formalin. Sieved samples may, depending on the
volume of sediment retained, require a slightly greater concentration of
formalin (10%) to assure adequate preservation. Use of alcohol as a
fixative should be avoided because worms initially preserved in alcohol
without first being fixed in formalin tend to deteriorate.
The initial sorting of specimens from sediment residue should be done
at 5-10X using a dissection microscope or magnification lens. Even when
sieving has been accomplished carefully, some individuals will fragment.
To avoid possible overcount in quantitative studies, only head-end
sections and whole worms should be enumerated. Forceps are commonly
employed to remove specimens from the sieve residue. The sorted
oligochaetes may be stored indefinitely in sealed vials containing 5-10%
formalin. Additional instructions for sorting macroinvertebrate samples
can be found in the USEPA Biological Methods Manual (Weber, 1973).
To identify oligochaetes to species, they must be cleared and mounted
on glass slides for examination under a compound microscope capable of
magnification up to 1000X. An 18 mm diameter, No. 0 or 1 round cover
glass is appropriate because it will adequately accommodate nearly the
size range of tubificids and the shape allows for maneuvering the
specimen into the most desired position by gentle rotation of the cover
glass. When preparing a temporary or permanent slide mount, an attempt
should be made to place the specimen on its side, thereby, revealing both
dorsal and ventral fascicles of chaetae. It may be necessary to apply
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gentle pressure on the cover glass to flatten the specimen to render the
important anatomical features more readily visible. Depending on the
need of the investigator, a variety of clearing agents, media, and
mounting procedures may be employed. The simplest method is to mount
specimens directly in a permanent nonresinous medium (for example,
Hydramountl, or CMC^) that contains a clearing agent. This method
allows rapid processing of specimens, but further handling of specimens
is rendered difficult because of the permanence of the mount.
Alternatively, specimens can be cleared in a solution of Amman's
lactophenol (Brinkhurst, 1968), prepared by combining lOOg phenol, lOOmL
lactic acid, 200mL glycerine, and lOOmL water. Clearing of specimens in
Amman's lactophenol may be accomplished either in a stoppered vial or by
mounting them temporarily on glass slides and covering with a round
coverglass. The clearing process usually takes a few hours to a few days
depending on the size and preservation of the specimens. Gentle
application of heat will speed the clearing process.
If the specimens are preserved in 70% alcohol, they should be placed
in 30% alcohol and then in water for a short time to leach out the
alcohol. The alcohol retards the clearing process of Amman's
lactophenol. However, do not leave specimens in the water too long (not
more than 2 hours) because the worms will begin to deteriorate.
Tubificids can be held indefinitely in Amman's lactophenol or 10%
formalin for later processing and mounting. After clearing, the
specimens can be examined directly or removed and mounted in a permanent
medium. Optimal resolution and longevity of mounted material are
achieved only in resinous media (for example, Canada balsam, Harleco's
Coverbond^ for xylene, and so forth). These mounting media require
dehydration of the specimens through the alcohol series before using the
mountant, but they produce the best permanent mounts. This method can be
found in standard biological techniques sourcebooks. Nonresinous media
are recommended for rapid processing of large numbers of specimens, but
for a few or for important reference materials, resinous mounting media
are best. Longevity in the quality of a mounted specimen can be
increased by sealing the margin of the cover glass with clear fingernail
lacquer.
DEPOSITORY FOR TUBIFICID WORMS
Tubificid material no longer needed in a study should be deposited in
an appropriate museum. In North America, the 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. 20560.
iBio/Medical Specialities (P.O. Box 1687, Santa Monica, CA 90406)
2Master's Chemical Co. (P.O. Box 2382, Des Plaines, IL 60018)
Scientific Products (1430 Waukegan Road, McGaw Park, IL 60085)
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SECTION 3
SPECIES LIST
Phylum Annelida
Class CHtellata
Subclass Oligochaeta
Order Haplotaxida
Suborder Tubificina
Family Tubificidae
Genus Aulodrilus Bretscher, 1899
Aulodrilus americanus Brinkhurst and Cook, 1966
Aulodrilus' limnobius Bretscher, 1899
Aulodrilus pigueti Kowalewski. 1914
Aulodrilus' pluriseta (Piguet, 1907)
Genus Bothrioneurum Stole, 1886
Bothrioneurum vejdovskyanum Stole, 1886
Genus Branchiura Beddard, 1892
Branchiura sowerbyi Beddard, 1892
Genus Haber Holmquist, 1978
Haber cf. speciosus (Hrabe, 1931)
Genus Ilyodrilus Eisen, 1879
Ilyodrilus frantzi Brinkhurst, 1965
Ilyodrilus' frantzi form capillatus Brinkhurst and Cook, 1966
Ilyodrilus rnastix Brinkhurst, 1978
Ilyodrilus tempietoni (Southern, 1909)
Genus Isochaetides Hrabe, 1966
Isochaetides curvisetosus (Brinkhurst and Cook, 1966)
Isochaetides freyi (BrTnkhurst, 1965)
Genus Limnodrilus Claparede, 1862
Litnnodrilus angustipenis Brinkhi
Limnodrilu? cervix Brinkhurst, 1963
Limnodrilus angustipenis Brinkhurst and Cook, 1966
Brii
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Limnodrilus cervix (variant form)
LimnodriTu? claparedianus Ratzel, 1868
Limnodrilus hoffmeisteri Claparede, 1862
Limnodrlfu? hoffmeisteri (spirails form)
LimnodriTu? hoffmeisteri (variant form)
Limnodrifus" maumeensis Brinkhurst and Cook, 1966
Limnodrilu? profundicpla (Verrill, 1871)
Limnodritui? psammophllus Loden, 1977
Limnodrilus rubripenis Loden, 1977
Limnodrilus silvani Eisen, 1879
Limnodrilii? udekemianus Claparede, 1862
Genus Monopylephorus Levinsen, 1884
Monopylephorus he!obi us Loden, 1980
Genus Phallodrilus Pierantoni, 1902
Phallodrilus hallae Cook and Hiltunen, 1975
Genus Potamothrix Vejdovsky and Mrazek, 1902
Potamothrix bavaricus (Oschmann, 1913)
Potamothri'x bedoti fPiguet. 1913)
Potamothrix hamrnoniensis (Michaelsen, 1901)
Potamothrix' moldaviensis Vejdovsky and Mrazek, 1902
Potamothrix vejdovskyi (Hrabe, 1941)
Genus Psammoryctides Hrabe, 1964
Psammoryctides barbatus (Grube, 1861)
Psammoryctides califprnianus Brinkhurst, 1965
Psammoryctides convolutus Loden, 1978
Psammoryctides minutus Brinkhurst, 1965
Genus Quistadrilus Brinkhurst, 1981
Quistadrilus multisetosus (Smith 1900)
Genus Rhizodrilus Smith, 1900
Rhizodrilus lacteus (Smith, 1900)
Genus Rhyacodrilus Bretscher, 1901
Rhyacodrilus brevidentatus Brinkhurst, 1965
Rhyacodrilus coccineus (Ve'jdovsky, 1875)
RhyacodrilTi? falciformis Bretscher, 1901
RhyacodriTuT montana (Brinkhurst, 1965)
Rhyacodrilus punctatus Hrabe', 1931
RhyacodriTirs" soda!is (Eisen, 1879)
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Genus Splrosperma Brinkhurst, 1981
Splrosperma beetoni (Brinkhurst, 1965)
Splrosperina" carol1nens1s (Brinkhurst, 1965)
Splrosperma ferox (Elsen, 1879)
Spirosperma~ nlkolskyl (Lastockln and Sokolskaya, 1953)
Genus Telmatodrllus E1sen, 1879
Telmatodrilus vejdovskyl Elsen, 1879
Genus Tublfex Lamarck, 1816
Tublfex harmanl Loden, 1979
Tublfex Ignotus (Stole, 1886)
Tublfex kessleri amerlcanus Brinkhurst and Cook, 1966
Tublfex nerthus (Michael sen, 1908)
Tublfex superlorensls (Brinkhurst and Cook, 1966)
Tubifex tubifex (Muller. 1774)
Genus Varlchaeta Brinkhurst, 1981
Varichaeta nevadana (Brinkhurst, 1965)
Varlchaeta paclflca Brinkhurst. 1981
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SECTION 4
KEY TO FRESHWATER TUBIFICIDAE (ANNELIDA: CLITELLATA: OLIGOCHAETA)
OF NORTH AMERICA (NORTH OF MEXICO)
INTRODUCTION
Phylogenetic relationships within the Tubificidae are poorly known.
Generic limits within the family are based primarily on the configuration
of internal reproductive organs observable only in dissected or
differentially stained and sectioned material. A natural, phylogenetic
key to the Tubificidae, therefore, would require extensive preparation of
individual specimens and would be wholly unworkable for the aquatic
biologist interested only in determining the identity of specimens within
a given collection. The key provided herein was constructed, therefore,
on artificial bases using only external and internal features readily
observable in simple whole-mounted material using a conventional light
microscope. The key allows the user to proceed directly to the species
level of classification without keying out the genera. For discussions of
phylogenetic relationships and taxonomic limits of genera, the reader is
instructed to consult recent taxonomic literature; for example, Baker and
Brinkhurst, 1981; Brinkhurst, 1979b, 1979c, 1981; Brinkhurst and Jamieson,
1971; Holmquist, 1978, 1979; and Loden, 1977, 1978, 1979) cited in the
Selected Bibilography.
The principal features used to identify tubificids in this guide are
the somatic chaetae, specialized genital chaetae, and structures of the
male reproductive system. Chaetae occur in paired fascicles or bundles
located ventro-laterally, "ventral chaetae," and dorso-laterally, "dorsal
chaetae" (Figure 1). A variety of chaetal types occur. Generalized
chaetae and their designated terminology are illustrated in Figure 2.
Segments are identified by Roman numerals. An important point to note is
that segment I, which includes the peristomium, and mouth, and to which
the prostomium is connected, is devoid of chaetae. Therefore, to
determine the numerical position of each segment, one must count chaetal
fascicles, either dorsal or ventral, posteriad beginning in segment II.
This is particularly important in locating the genital segments X and XI
that bear the spermathecae and male pores, respectively.
A number of tubificid species must be sexually mature to be
identified to species. At two points within the key, the necessity for
mature specimens is noted. In each case, if specimens in question are
immature, unidentifiable immature reporting categories are provided. Two
points need to be emphasized in this regard: first, the majority of
individuals within any assemblage may be immature and, therefore, reported
-------
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11
-------
only as unidentifiable; second, the composition of the immature components
can generally be assumed to reflect the species composition of the mature,
identifiable individuals, present in the same collection.
KEY TO THE TUBIFICIDAE
1 Capilliform chaetae (Fig. 2) present in anterior (segments II-V)
dorsal fascicles 2
Capilliform chaetae not present in anterior dorsal fascicles. . . 37
2(1) Each segment, posterior to approximately segment XXX, bears a
dorsal and ventral gill filament (Fig. 3)
Branchiura sowerbyi
Posterior segments bear no gill filaments 3
Fig. 3. Branchiura sowerbyi, dorsal and ventral gill filaments on
posterior segments.
12
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3(2) Dennis opaque or nearly opaque, appearing to be densely papillate
and encrusted with sediment materials (Fig. 4) £r dermis coarse
perhaps slightly opaque and beset with some scattered papillae
(Fig. 5a,b). Prostomium often retractile and capilliform
chaetae frequently stout and sabre-like 4
Dermis clear (Fig. 6), or at most, slightly opaque but without
encrusted sediment materials or papillae. Prostomium rarely
retractile, capilliform chaetae rarely sabre-like 9
5a
5b
Fig. 4. Granular-opaque, densely papillate dermis of Spirosperma
nikolskyi.
Fig. 5. Quistadrilus multisetosus: (a,b) slightly opaque dermis with
scattered papillae.
Fig. 6. Clear dermis, without papillae.
13
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4(3)
Dermis coarse but not opaque and without appearance of encrusted
sediment materials; variously beset with scattered short,
finger-like papillae. Papillae often most readily visible
posteriad to approximately segment XI. Anterior dorsal
fascicles composed of stout sabre-like capilliform and
distinctly pectinate chaetae (Fig. 7)
Dermis opaque or nearly opaque, appearing encrusted with sediment
materials, densely papillate. Dorsal crotchet chaetae often
difficult to differentiate
5(4)
Anterior and posterior ventral chaetae similar in shape, distal
tooth longer than proximal tooth (Fig. 8). [See Annotations.] .
Qulstadrilus multisetosus (in part)
Anterior and posterior ventral chaetae differ in shape (Fig. 9a,b).
Posterior chaetae thick, with distal tooth shorter and much
thinner than proximal tooth. [See Annotations.]
Quistadrilus multisetosus (in part)
8
9a
9b
Fig. 7. Quistadrilus multisetosus, dorsal pectinate chaeta.
Fig. 8. Q. multisetosus, posterior ventral chaeta.
Fig. 9. Ij. multisetosus: (a) anterior ventral chaeta; (b) posterior
ventral chaeta.
14
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6(4) Dorsal fascicles composed of capilliform and distinctly pectinate
chaetae (Fig. 10) Spirosperma ferox
Dorsal fascicles composed of capilliform chaetae and other shorter,
much thinner capilliform-like chaetae, which may have pectinate
tips, but are difficult to differentiate 7
7(6) All anterior ventral chaetae bifurcate, 2 per fascicle, teeth
small (Fig. lla). Posterior ventral chaetae 1 per fascicle,
distal tooth reduced or rudimentary (Fig. lib). Lake Tahoe. . .
Spirosperma beetoni
Anterior ventral fascicles contain at least some simple-pointed
chaetae 8
lla
lib
Fig. 10. Spirosperma ferox, dorsal pectinate chaeta.
Fig. 11. J5. beetoni: (a) anterior ventral chaeta; (b) posterior ventral
chaeta.
15
-------
B(7) Ventral fascicles of segments II, III and often segment IV include
1 to 3 simple-pointed crotchet chaetae and 1 to 3 bifurcate
chaetae. Posterior ventral chaetae with teeth subequal or
distal tooth longer than proximal tooth (Fig. 12)
. Spirosperma nikolskyi
Ventral fascicles of segments II-VIII or IX composed of 1
simple and 1 bifurcate crotchet chaeta. Posterior ventral
chaetae 1 per fascicle, strongly recurved with thin distal tooth
(Fig. 13) Spirosperma carolinensis
9(3) Segment I with large mouth and enlarged eversible pharynx
(Fig. 14). Dorsal and ventral chaetae of segment II usually
single with short, thin distal tooth; chaetae of segment II and
often segment IV lacking. Posteriad, dorsal fascicles composed
of capilliform and bifurcate crotchet chaetae with distal tooth
thinner but about as long as proximal tooth
Ilyodrilus mastix
Segment I with normal mouth, without an enlarged eversible
pharynx, anterior dorsal and ventral chaetae other than above.
10
12
13
Fig. 12. Spirosperma nikolskyi, posterior ventral chaetae.
Fig. 13. S. carolinensis, posterior ventral chaeta.
Fig. 14. Tlyodrilus mastix, anterior end illustrating enlarged mouth.
16
-------
10(9) Anterior dorsal fascicles composed of capilliform and bifurcate
crotchet chaetae (Figs. 15, 16a, 17, 18, 19a) without
intermediate teeth 11
Anterior dorsal fascicles composed of capi111 form and pectinate
chaetae (Figs. 20, 21a, 29). [Intermediate teeth of pectinate
chaetae may require approximately 1000X magnification to be
observed.] 16
11(10) Dorsal crotchet chaetae from approximately segment VII posteriad
modified to a spatulate shape when viewed In facial aspect
(Fig. 15a) and appearing compressed when viewed from lateral
aspect (Fig. 15b) Aulodrilus pigueti
Dorsal crotchet chaetae not modified into a spatulate form as
above 12
12(11) Anterior ventral crotchet chaetae with greatly reduced distal
teeth, essentially simple-pointed. Each posterior segment
bearing dorsal and ventral gill filaments (Fig. 3)
Branchiura sowerbyi
Anterior ventral crotchet chaetae distinctly bifurcate (although
distal tooth nay be reduced), ^r if ventral fascicles contain
simple crotchet chaetae, then no gill filaments on posterior
segments 13
a b
Fig. 15. Aulodrilus pigueti, posterior dorsal crotchet chaeta: (a)
facial aspect; (b) lateral aspect.
17
-------
13(12) Teeth of anterior dorsal crotchet chaetae equal in length,
somewhat ovate and not strongly divergent (Fig. 16a).
Anterior capilliform and crotchet chaetae appear bent to
nearly sigmoid in shape or form (Fig. 16a). Mature specimens
with spermathecal chaetae in ventral fascicles of segment X
(Fig. 16b) Potamothrix vejdovskyi
Teeth of anterior dorsal crotchet chaetae unequal in length (Figs.
17, 18a), or if teeth are similar in length, then strongly
divergent (Fig. 19a) 14
14(13) Dorsal fascicles with 1-2 capilliform chaetae, 2-3 bifurcate
crotchet chaetae with distal tooth shorter and thinner than
proximal tooth (Fig. 17). Ventral fascicles with up to 4
chaetae, either simple-pointed or with distal tooth shorter
and thinner than proximal tooth. With coelomocytes. Lake
Tahoe Rhyacodrilus brevidentatus
Dorsal fascicles not as above, ventral crotchet chaetae never
simple-pointed 15
16 a
16 b
17
Fig. 16. Potamothrix vejdovskyi: (a) anterior dorsal crotchet, and
capilliform chaetae; (b) spermathecal chaeta.
Fig. 17. Rhyacodrilus brevidentatus, dorsal crotchet chaeta.
18
-------
15(14) Distal tooth of ventral crotchet chaetae in segment VIII
distinctly shorter and/or thinner than proximal tooth (Fig.
18a). Dorsal crotchet chaetae also with distal tooth shorter
and thinner than proximal tooth, occasionally dorsal crotchet
chaetae may be minutely pectinate (Fig. 18b). Capilliform and
crotchet chaetae somewhat arcuate or sigmoid
Aulodrilus pluriseta
Distal tooth of ventral crotchet chaetae in segment VIII neither
shorter nor obviously thinner than proximal tooth. Distal
tooth of dorsals equal to or slightly longer than proximal
tooth (Fig. 19a). Mature specimens with several simple-
pointed penial chaetae (Fig. 19b). With coelomocytes
Rhyacodrilus coccineus
18 a
18 b
19 a
19 b
Fig. 18. Aulodrilus pluriseta: (a) ventral crotchet chaeta; (b)
dorsal pectinate chaeta.
Fig. 19. Rhyacpdrilus coccineus: (a) dorsal crotchet chaeta; (b)
penial chaetae.
19
-------
16(10) Anterior dorsal fascicles with broadly expanded pectinate crotchet
chaetae (Fig. 20) Psammoryctides barbatus
Anterior dorsal fascicles with unexpended pectinate crotchet
chaetae (Figs. 21 a, 29) 17
•
17(16) Teeth of anterior dorsal pectinate chaetae long, distal tooth
often much longer than proximal tooth (Fig. 21a). Capilliform
chaetae in segment II frequently much longer than poster!ad.
Anterior ventral crotchet chaetae with distal tooth 1 1/2
times longer than proximal tooth (Fig. 21b). With
coelomocytes Rhyacodrilus montana
Distal tooth of anterior dorsal pectinate chaetae not distinctly
longer than proximal tooth. Capilliform chaetae,in segment II
not distinctly longer than poster!ad and ventral chaetae may
or may not be pectinate 18
20
21 a
21 b
Fig. 20. Psammoryctldes barbatus, pectinate anterior dorsal crotchet
chaeta.
F1g. 21. Rhyacodrilus montana; (a) anterior dorsal pectinate chaeta;
(b) anterior ventral crotchet chaeta.
20
-------
18(17) Posterior dorsal and ventral fascicles containing strongly
sigmoid, arcuate crotchet chaetae (Fig. 22)
Tubifex hartnani
Posterior dorsal and ventral fascicles containing no strongly
sigmoid or arcuate crotchet chaetae 19
19(18) Anterior and posterior ventral chaetae pectinate (Fig. 23). With
coelomocytes. Lake Superior
Rhyacodrilus punctatus
Anterior and posterior ventral chaetae not pectinate.
20
22
23
Fig. 22. Tubifex harmani, posterior dorsal crotchet chaeta.
Fig- 23. Rhyacodrilus punctatus. anterior ventral chaeta.
21
-------
20(1P) Capilliform chaetae very long and thin, poster!ad from segment X
capilliform chaetae 2-3 times longer than diameter of
respective segment. Long thin worms Tubifex ignotus
Capilliform chaetae not especially long and thin and progressively
diminish in length posteriad to segment X 21
21(20) Anterior ventral chaetae with distal tooth approximately twice as
long as reduced proximal tooth (Fig. 24)
Tubifex nerthus
Teeth of anterior ventral chaetae not as above. [Henceforth
specimens must have attained sexual maturity in order to be
identifiable to species, otherwise specimens can be
categorized only as "Unidentifiable immature Tubificidae with
capilliform chaetae"] 22
Fig. 24. Tubifex nerthus, anterior ventral chaeta.
22
-------
22(21) Mature individuals bear a muscular penial bulb attached to a
crescent-shaped atrium which constricts into very long
annulated vas deferens (Fig. 25). Penis sheath a granular
membrane, very thin-walled and often indistinct. No modified
genital chaetae present Tubifex tubifex
Penial apparatus, atrium and vas deferens not as above.
Individuals bear a translucent penis sheath and/or modified
genital chaetae in at least one segment between segments VII
and XI 23
Penis Bulb
Vas Deferens
Penis Sheath
Fig. 25. Tubifex tubifex, penis bulb, atrium, and vas deferens.
23(22) Mature individuals bear a short, broad penis sheath in segment
XI (Fig. 26a). Penial and spermathecal chaetae long, thin,
parallel-sided (Fig. 26b,c), originating in glandular sacs. .
Haber cf. speciosus
Mature individuals bear either a penis sheath or spermathecal or
penial chaetae but no combination of a penTs sheath and both
spermathecal and penial chaetae 24
23
-------
26 a
26 b
26 c
Fig. 26. Haber cf. speclosus; (a) penis sheath; (b) penial chaeta; (c)
spermathecal chaeta.
Fig. 71. Potamothrix bedoti, specialized chaeta.
24(23) With or without a cuticular penis sheath. Specialized genital
chaetae present in one or more segments ranging between
segments VII and XI 25
With a translucent cuticular penis sheath in segment XI. No
genital chaetae in segment X. Ventral chaetae in segment XI
similar in size and shape to somatic chaetae, or at most, only
slightly thickened 32
25(24) Seal pel-shaped specialized chaetae (Fig. 27) present in any or- all
segments ranging between segments VII and XI
Potamothrix bedoti
Genital chaetae present only in segment X or XI and not seal pel -
shaped 26
26(25) With penial chaetae in segment XI, without spermathecal
chaetae in segment X 27
Without penial chaetae in segment XI, with spermathecal chaetae
in segment X 29
24
-------
27(26) Penial chaetae enlarged with bifurcate tip (Fig. 28). Penis sheath
tub-shaped, indistinct. Pectinate chaetae range from bearing
parallel lateral teeth to slightly palmate and sigmoid with
distal tooth thinner and slightly longer than proximal tooth.
Posterior dorsal fascicles consist only of bifurcate crotchet
chaetae, no capilliform chaetae. Without coelomocytes. . . .
Psammoryctides minutus
Penial chaetae blunt-tipped and simple-pointed (Fig. 19b); no
penis sheath, chaetae not as above. With coelomocytes. .
28
28(27)
Teeth of dorsal crotchet chaetae long, thin, and parallel,
intermediate teeth fine (Fig 29). Frequently, distal tooth of
dorsal chaetae in segments VIII and IX twice the length of
proximal tooth Rhyacodrilus soda!i s
Teeth of dorsal crotchet chaetae not long and parallel and distal
tooth of dorsal chaetae in segments VIII and IX not twice the
length of proximal tooth Rhyacodrilus coccineus
29(26)
Spermathecal chaetae in segment X parallel sided with teeth
fused forming a groove (Figs. 30, 31)
Spermathecal chaetae not parallel sided, but are elongate
spade-shaped, broadest near the middle and tapering to a
pointed tip (Figs. 32, 33)
30
31
28
29
Fig. 28. Psamnioryctides minutus, penial chaeta.
Fig. 29. Rhyacodrilus soda!is, dorsal pectinate chaeta.
25
-------
30(29) Anterior ventral chaetae with teeth subequal or distal tooth
slightly longer than proximal tooth. Posteriad, ventral
chaetae thicker, often with distal tooth more than two times
longer than proximal tooth. Penis sheath indistinct, if
visible, tub-shaped with reflexed head
Psammoryctides convolutus
Anterior ventral chaetae with distal tooth longer and thinner than
proximal tooth, posteriad, ventral chaetae slightly shorter,
but also with distal tooth longer and thinner than proximal
tooth. Penis sheath indistinct, tub-shaped but without
reflexed head Psammoryctides caliform'anus
31(29) Spermathecal chaetae in segment X often two per fascicle, much
longer than wide (Fig. 32). [See annotations.]
Potamothrix hammoniensis
Spermathecal chaetae in segment X usually single, short and
broad (Fig. 33) Potamothrix bavaricus
30
31
32
33
Fig. 30. Psammoryctides convolutus, spermathecal chaeta.
Fig. 31. P. californlanus, spermathecal chaeta.
Fig. 32. Tbtaniothrlx hammoniensls, spermathecal chaeta.
Fig. 33. P. bavarfcus, spermathecal chaeta.
26
-------
32(24) Penis sheath well developed, cylindrical, conical or conical
with an elongate tip. (Figs. 34-37) 33
Penis sheath indistinct or short and tub-shaped (Fig. 38). . . . 36
33(32) Penis sheath only slightly longer than wide (Fig. 34a,b). Ventral
chaetae of segment XI similar in shape to those in adjacent
segments but slightly thicker Varichaeta pacifica
Penis sheath other than above. Ventral chaetae of segment XI
similar in shape, size and thickness to those in adjacent
segments 34
34(33) Penis sheath a cylindrical tube with a reflexed basal portion
(Fig. 35) Tubifex superiorensis
Penis sheath other than above 35
35
Fig. 34. Varichaeta pacifica: (a) penis sheath inverted; (b) penis
sheath everted.
Fig. 35. Tubifex superiorensis. penis sheath.
27
-------
35(34) Penis sheath conical, uniformly tapered from base to apex
(Fig. 36a,b) Ilyodrilus templetoni
Penis sheath with a conical base abruptly tapered to an elongate
apex (Fig. 37) Tubifex kessleri americanus
36(32) Penis sheath tub-shaped (Fig. 38), spermatozeugmata "frying pan"
shaped Ilyodrilus frantzi form capillatus
Penis sheath indistinct, delicate. Lake Tahoe.
Varichaeta nevadana
36 a
36 b
37
38
Fig. 36. Ilyodrilus templetoni: (a) penis sheath with distal end absent;
(b) penis sheath with distal end present.
Fig. 37. Tubifex kessleri americanus, penis sheath.
Fig. 38. Ilyodril'us frantzi form ca"p"illatus, penis sheath.
28
-------
37(1) All chaetae anterior to approximately segment VII simple to
minutely bifurcate with much reduced distal tooth (Fig.
39a,b) 38
Anterior chaetae never simple, teeth distinct 39
38(37) Anterior chaetae simple. Posteriad from approximately segment
VII, dorsal chaetae laterally expanded with several minute
teeth (Fig. 40) Aulodrilus americanus
Anterior chaetae appear simple (Fig. 39a) to minutely bifurcate
with minute distal tooth. Posteriad, dorsal chaetae bifurcate
with reduced distal tooth (Fig. 39b)
Telmatodrilus vejdovskyi
39 3
39 b
Fi9- 39.
Fig. 40.
Telmatodrilus vejdovskyi; (a) anterior ventral chaeta;
(b) posterior ventral chaeta.
Aulodrilus americanus, posterior dorsal crotchet chaeta.
29
-------
39(37) Posteriad from approximately segment VII dorsal crotchet chaetae
spatulate when viewed from facial aspect (Fig. 41a); viewed
laterally, each chaeta appears compressed with tiny teeth
(Fig. 41 b) Aulodrilus limnobius
Posteriad from approximately segment VII dorsal chaeta not
laterally expanded, teeth not reduced 40
40(39) All chaetae similar; bifurcate crotchet chaetae with distal tooth
thinner and shorter than proximal tooth (Fig. 42), anteriorly
3-4 per fascicle, 2 per fascicle posteriad. With
coelomocytes Monopylephorus he! obi us
All crotchet chaetae not similar, or of different form and number
than above 41
b
Fig. 41. Aulodrllus limnobius, posterior dorsal crotchet chaeta:
facial aspect; (b) lateral aspect.
Fig. 42. Monopylephorus he!obi us, crotchet chaeta.
(a)
30
-------
41(40) Prostomium with a ciliated sensory cavity with an irregular opening
on the dorsal surface of the prostomium (Fig. 43). Viewed
laterally, the cavity appears as a depression and specialized
cells lining the pit are apparent. Chaetae of anterior
segments II to approximately IX generally 3-4 per fascicle
(Fig. 44a) and often thinner than those posteriad. Posterior
chaetae usually number tv/o per fascicle, thicker than anterior
chaetae (Fig. 44b). Mature individuals bear hook-shaped
penial chaetae in segment XI (Fig. 44c). With coelomocytes. .
Bothrioneurum vejdovskyanum
Prostomium without a sensory cavity, chaetae other than above.
42
Fig. 43. Bothrioneurum vejdovskyanum, SEM of prostomium showing sensory
cavity (from Chapman, 1979).
Fig. 44. B. vejdovskyanum; (a) ventral crotchet chaeta of segment II;
Tb) ventral crotchet chaeta of segment XXX; (c) penial chaeta.
31
-------
42(^1) Dorsal crotchet chaetae posteriad from approximately segment XX
sharply arcuate and larger than opposing ventral chaetae (Fig.
45). Mature individuals bear short, thin, parallel-sided
spermathecal chaetae in segment X
Isochaetides curvisetosus
Dorsal crotchet chaetae posteriad from approximately segment XX
neither sharply arcuate nor larger than opposing ventral
chaetae 43
43(42) Distal tooth of ventral chaetae in segments II through IV longer
than proximal tooth and deflected at an angle of approximately
90° to axis of chaetae (Fig. 46a,b,c)
Limnodrilus udekemianus
Distal tooth of ventral chaetae in segments II through IV not
deflected 90° to axis of the chaetae. [Henceforth specimens
must be sexually mature to be identifiable to species,
otherwise specimens must be classified as "Unidentifiable
immature Tubificidae without capilliform chaetae"] 44
46a
46 b
415 c
Fig. 45. Isochaetides curvisetosus, posterior dorsal chaeta.
Fig. 46. LimnpdriluT'udekemianus; (a,b,c) ventral chaetae in segment
II, illustrating variation in shape.
32
-------
44(43) With pern's sheath in segment XI (Fig. 47a), and spermathecal
chaetae long thin, parallel-sided (Fig. 47b) in segment X. . .
Isochaetides freyi
With spermathecal chaetae in segment X and/or penial chaetae in
segment XI or penis sheath but no combination of genital
chaetae and a penis sheath 45
45(44) Genital chaetae in segments IX and XI; spatulate and single
in segment IX (rarely segment X) (Fig. 48a), 4-5 knobbed
chaetae in segment XI (Fig. 48b). With coelomocytes. . . .
Rhizodrilus lacteus
Spermathecal chaetae in segment X £r penial chaetae
in segment XI but both spermathecal and pem'al chaetae not
present. Penis sheaths present or absent 46
47 a
47 b
48 a
48 b
Fig. 47. Isochaetides freyi; (a) penis sheath; (b) spermathecal chaeta.
Fig. 48. Rhizodrllus lacteus; (a) specialized chaeta in segment IX; (b)
specialized chaetae in segment XI.
33
-------
46(45) With penis sheaths 49
Without penis sheaths 47
47(46) With penial chaetae in segment XI, no spermathecal chaetae in
segment X 48
Spermathecal chaetae present in segment X (Fig. 49a), and eversible
penes present in segment XI (Figs 49b,c)
Potamothrix moldaviensis
b
Fig. 49. Potamothrix moldaviensis: (a) spermathecal chaeta; (b)
partially everted penis; (c) fully everted penis.
34
-------
48(47) Pem'al chaetae in segment XI simple-pointed, distally hooked
(Fig. 50). Lake Superior Phallodrilus hallae
Penial chaetae in segment XI sickle-shaped, twice as long and much
thicker than somatic chaetae (Fig. 51a,b). With coelomocytes.
Rhyacodri1 us falci f omri s
49(46) Penis sheath tub-shaped (Fig. 38). Spermatozeugmata "frying pan"
shaped, distal tooth of anterior chaetae longer than proximal
tooth Ilyodrilus frantzi
Penis sheath elongate and cylindrical 50
50
51 a
51 b
Fig. 50. Phallodrilus hallae, penial chaeta.
Fig. 51. RhyacodriTu? falciformis, penial chaeta:
(b) facial aspect.
(a) lateral aspect;
35
-------
50(49) Margin of head of penis sheath set at essentially right angles to
axis of shaft (Figs. 52-57) 51
Margin of head of penis sheath slanted, hooded or otherwise, not
at right angles to axis of shaft (Figs. 58-64) 56
51(50) Penis sheath reddish-brown; with two tapered lateral
projections (Fig. 52) Limnodrilus rubripenis
Penis sheath clear; distal end without two tapered lateral
projections 52
52(51) Distal 3/4 of penis sheath cylindrical, tube-like, basal 1/4
conical, shaft long in fully developed sheaths (Fig. 53). . .
Limnodrilus angustipenis
Penis sheath not distinctly differentiated into cylindrical
distal and conical basal portions, shaft short or long in
fully developed sheaths 53
53(52) Fully developed sheaths long (generally greater than 300 urn), shaft
uniformly cylindrical with head a round plate set at right
angles to shaft. Margin of head may be slightly up-turned on
one side (Fig. 54)
Limnodrilus hoffmeisteri (spiralis form)
Fully developed sheaths short, head of sheath other than above. .
54
54(53) Shaft broadly expanded basally, head a round plate (Fig. 55)
Limnodrilus psamnophilus
Shaft not broadly expanded nasally, head of sheath other than
above 55
55(54) Anterior ventral chaetae of segment II to approximately segment
IV with distal tooth longer than proximal tooth and bent at a
90° angle to axis of chaetae (Figs. 46a,b,c). Penis sheath
with a round reflexed head (Fig. 56)
Limnodrilus udekemianus
Anterior ventral chaetae with distal tooth subequal or only
slightly longer than proximal tooth and teeth set at 45° or
less to axis of shaft. Shaft of sheath thin, head small (Fig.
57) Limnodrilus profundicola
56(50) Mature shaft short, head of sheath spade-shaped (Fig. 58).
Limnodrilus silvani
Mature shaft long, (generally greater than 300 urn) and
head of sheath not spade-shaped 57
36
-------
52
53
55
56
57
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
52.
53.
54.
55.
56.
57.
Limnodrilus rubripenis, penis sheath.
L. angustTpenis, penis sheath.
r. hotfmeisteri (spiralis form).
U. psammophilus, penis sheath.
L. udekemianus, penis sheath.
L_. profundicoTa, penis sheath.
penis sheath.
37
-------
57(56) Head of penis sheath with margin scalloped (Fig. 59a), or
scalloped with an irregular distal projection (Fig., 59b) . . .
Limnodrilus hoffmeisteri (variant form)
Head of penis sheath not scalloped 58
58(57) Shaft wall not excessively thickened (Figs. 60, 61) 59
Shaft noticeably thick-walled (Figs. 62-64) perhaps appearing
as two layers 60
59(58) Head of penis sheath with an overhanging hood and a broad, flat
proximal lip, distally the diameter of the shaft flares into
the head; mature tube length 300-600 urn (Fig. 60)
Limnodrilus hoffmeisteri
Head of penis sheath without an overhanging hood. Shaft
slender, long (600-700 urn) sigmoid with head equilaterally
triangular and hence bilaterally symmetrical (Fig. 61). ...
Limnodrilus claparedianus
60(58) Head of penis sheath broad, asymmetrically triangular and distal
end of shaft canted (Fig. 62) Limnodrilus maumeensis
Head of penis sheath longer than wide, shaft not canted
distally 61
61(60) Head of penis sheath small, pear-shaped, bilaterally
symmetrical, apical portion of head often reflexed over
orifice, lower proximal portion small (Fig. 63)
Limnodri1us cervix (variant form)
Head of penis sheath asymmetrical, with an offset proximal
digitate lobe (Fig. 64) Limnodrilus cervix
38
-------
58
59 b
61
63
Fig. 58. Limnodrilus silvani, penis sheath.
Fig. 59. L. hoffmeTsteri (variant form), illustrating two penis sheath
types!(a) scalloped margin; (b) scalloped margin with
irregular distal projection.
Fig. 60. L. hoffmeisteri, penis sheath.
Fig. 61. r. claparedianus, penis sheath.
Fig. 62. r. maumeensis, penis sheath.
Fig. 63. r. cervix (variant form), penis sheath.
Fig. 64. U. cervix, penis sheath.
39
-------
SECTION 5
GLOSSARY*
Anteriad: Direction toward the anterior of an organism.
Arcuate; Curved or shaped like a bow.
Bifurcate: A condition of a chaeta where its apex is cleft, forming two
unequal teeth.
Capi11iform Chaeta: A dorsal, uncleft, hair-like, somatic chaeta; longer
and usually more flexuous than a crotchet chaeta (sometimes called a hair
seta).
Chaeta (pi. Chaetae); A bristle, which in various forms, aids primarily
in locomotion (somatic chaetae) or can function in connection with
reproduction (genital chaetae). (In the literature, "seta" is frequently
used instead of "chaeta," but here the former term is reserved for its
application in arthropod morphology).
C1itel1 urn; A differentiation of the epidermis in the genital region into
a somewhat verrucose "sleeve" which will transform into a cocoon that
serves as a repository for the eggs following their fertilization.
Coelomocyte: Free cells within the body cavity (coelom). Granular in
appearance, these spherical to ovoid cells may completely fill the body
cavity in certain species. A characteristic feature in four of the North
American tubificid genera: Bothrioneurum, Monopylephorus, Rhizodrilus,
and Rhyacodrilus.
Cosmopoliton: World-wide in distribution.
Crotchet chaeta (pi. Chaetae); A somatic chaeta sometimes simple, but
usually bifurcate, with two primary lateral teeth, with or without
intermediate teeth between the lateral teeth. Generally refers to simple
or bifurcate chaetae without intermediate teeth. See also Pectinate
Chaeta.
*Additional morphological terms applied in oligochaetology can be found
in Brinkhurst and Jamieson (1971) and Reynolds (1977)
41
-------
Fascicle: A "cluster" or "bundle" of chaetae. Generally four fascicles
per segment, two located dorso-laterally and two located ventro-laterally.
Genital Chaeta: A chaeta which functions in sexual reproduction;
spermathecal and penial chaeta.
Genital Segments; Body segments (ordinarily segments X and XI) which
bear the reproductive organs.
Gill Filaments; Paired finger-like extensions of the body wall that
function in respiration. Located on the posterior segments of Branchiura
sowerbyi, two per segment, one positioned dorsally and one ventrally.
Modulus: A knob or enlarged region on a crotchet chaeta generally
located at or near the midlength of the chaeta where it leaves the body
wal 1.
Papilla (pi. Papillae): Projections of the dermis or body wall.
Frequently heavily cuticularized and obscured by accumulated foreign
particles, thereby giving the worm a very opaque appearance.
Pectinate Chaeta: A crotchet chaeta generally equipped with two primary
lateral teeth and few to several finer intermediate teeth. Nearly always
dorsal and in fascicles with capilliform chaetae. Intermediate teeth may
require 1000X magnification to be resolved.
Penial Chaeta; A chaeta associated with the penes occurring in the
ventral fascicles of Segment XI, ordinarily their shape and size is
unlike that of the somatic chaetae.
Penis Sheath; A translucent cuticular covering of the penis in segment
XI, ranging from a thin, often obscure ring to an elaborate, thick tube.
Posteriad: Direction toward the posterior of an organism.
Prostomium: The anterodorsal part of the cephalic segment (Segment I).
Segments: A series of anatomical divisions of the body (somites or
compartments), each usually separated from its neighbor by a septum
(partition).
Seta (pi. Setae); See Chaeta.
Simple Chaeta; A chaeta with an uncleft apex.
Somatic Chaeta: A chaeta which functions in connection with locomotion;
the somatic segments only, as opposed to specialized genital chaetae that
are associated with the genital segments.
42
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Spermathecal Chaeta; Chaeta associated with the spermatheca, generally
in the ventral fascicles of Segment X. A characteristic feature of
certain genera and species, their shape is usually unlike that of a
somatic chaeta.
Spermatozeugma (pi. Spermatozeugmata); Organized bundles of sperm found
in the atrium prior to copulation and in the spermatheca following
copulation. Of various, often diagnostic shapes and sizes, but generally
with an inner lumen surrounded with sperm heads oriented inward and sperm
tails radiating outward.
43
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SECTION 6
ANNOTATIONS
1. Aulodrilus americanus
Largest species of the genus. Reproduces primarily by budding and
fragmentation. Moderately tolerant to organic enrichment and common
in productive habitats. Dwells in tubes that often stay with the
individual after sieving. Nearctic. Known from St. Lawrence Great
Lakes, the eastern United States, Cayuga Lake, New York, British
Columbia, Manitoba, New Brunswick, Northwest Territories, Ontario,
Quebec, and Saskatchewan.
2. Aulodrilus limnobius
Exhibits a preferance for silty substrates in mesotrophic habitats.
Moderately tolerant organism that reproduces primarily by budding
and fragmentation. Often collected within a tube of agglutinated
organic materials. The species is often represented in the drift of
rivers. Usually sympatric with £. pigueti. Cosmopolitan.
Widespread in the United States. Reported from Alabama, Louisiana,
North Carolina, South Carolina, Tennessee, Wisconsin, and Canada.
3. Aulodrilus pigueti
Usually sympatric with A. limnobius and with a similar distribution,
life history and habitat preferences. Cosmopolitan.
4. Aulodrilus pluriseta
Locally abundant in enriched habitats in silty substrates. Forms a
mucoid tube similar to other members of the genus. Reproduces
sexually and asexually by budding. Cosmopolitan. In North America,
reported from St. Lawrence Great Lakes, Illinois, Indiana, Cayuga
Lake, New York, North Carolina, Pennsylvania, Tennessee, Hudson
River, Ohio River, Wisconsin River drainage, and British Columbia,
Quebec in Canada.
5. Bothrioneurum vejdovskyanum
A widespread species. Generally most abundant in large rivers in
coarse sand substrates. Rarely abundant in lentic habitats. Has
likely been frequently overlooked in many North American collections
due to confusion regarding the characteristic sensory prostomial pit
44
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and infrequent occurrence of sexually mature individuals within a
population. Cosmopolitan. Known from St. Lawrence Great Lakes,
Indiana, Illinois River, Cayuga Lake, New York, Ohio River,
Wisconsin River, Lake Washington, British Columbia and Quebec.
Note; M.S. Loden (Personal Communication, 1981) reported^.
americanum from Georgia and Louisiana. In the immature state, it is
indistinguishable from EL vejdovskyanum and mature specimens lack
penial chaetae.
6. Branchiura sowerbyi
Large worms that are rarely abundant. Known to thrive in heated
effluents in Europe. Great Lakes records are from Lake St. Clair to
Lake Erie. Frequently fragments. Cosmopolitan. Widespread
throughout North America, generally in rivers.
7. Haher cf. speciosus
Only two recorded specimens in North America; Susquehanna River, New
York, and Hudson River, New York.
8. Ilyodrilus frantzi and Ilyodrilus frantzi form capillatus
Ilyodrilus frantzi was originally described by Brinkhurst (1965)
from Suisan Bay, California. The species was redescribed to include
l_. frantzi form capillatus by the same author in 1978, based upon
specimens collected in the Fraser River, British Columbia. No other
known distributional records.
9. Ilyodrilus mastix
Recently described by Brinkhurst (1978) from the Fraser River,
British Columbia, the only known North America record.
10. Ilyodrilus tempi etoni
Common in both lake and river habitats, attaining greatest densities
in enriched habitats. One of the most common tubiflclds in North
America. Often sympatric with Limnodrilus hoffmeisteri in highly
enriched waters. Easily confused with Tubifex tubifex. Holarctic.
Widely distributed throughout the United States and is probably
widespread in Canada.
11. Isochaetides curvisetosus
Transferred from Peloscolex by Brinkhust (1981). An occasional
species apparently not occurring in eutrophic habitats. Nearctic.
Known from St. Lawrence Great Lakes, Alabama, Indiana, North
Carolina, and Pennsylvania.
45
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12. Isochaetides freyi
Transferred from Peloscolex by Brinkhurst (1981). An occasional
species generally occurring only in mesotrophic habitats. Rarely
very abundant. Nearctic. Known from St. Lawrence Great Lakes,
Mississippi River, Alabama, Indiana, Louisiana, Cayuga Lake, New
York, North Carolina, Pennsylvania, Tennessee, Chippewa River and
Red Cedar River in Wisconsin, Manitoba and Fraser River, British
Columbia, Canada.
13. Limnodrilus angustipenis
An uncommon taxon usually collected in oligotrophic lakes.
Nearctic. Known from St. Lawrence Great Lakes, Alabama, Louisiana,
North Carolina, Tennessee, New Brunswick, Ontario, and Manitoba.
14. Limnodrilus cervix and Limnodrilus cervix (variant form)
The distinction between the typical and variant forms was first
noted by Hiltunen (1969). The taxonomic status of the variant form
is a subject that has, to date, not been adequately resolved, and a
state of confusion exists as to the correct rank of the variant
form. In this treatment, the authors defer from making a taxonomic
decision as to the status of variant forms, whether a subspecies,
ecomorph, hybrid or other entity, but have maintained the variant
form herein based upon apparent distributional discontinuities
between the typical and variant forms in relation to habitat and
water quality. The typical form is common, although rarely
abundant, through large areas of the United States. It is often
common in productive, enriched habitats, although it occurs
infrequently in grossly degraded situations. In contrast, the
variant form has been collected primarily in areas sustaining high
degrees of enrichment or in grossly polluted lakes, rivers and
harbors in the United States. It is often one of the predominant
tubificids, attaining large population densities in such
situations. In such cases, the variant form has been collected both
sympatric with, and in the absence of typical form. Hoi arctic.
Widespread throughout the United States, including the St. Lawrence
Great Lakes, Mississippi River, Illinois River, Ohio River, Hudson
River, and known from Ontario, Canada.
15. Limnodrilus claparedianus
A common species in both lakes and rivers. Generally most abundant
in enriched habitats but also known from clean water areas.
Cosmopolitan. Recorded from throughout the United States and Canada.
16. Limnodrilus hoffmeisteri
A ubiquitous species that is probably the most abundant and
widespread species in North America. Known from habitats ranging
from pristine to grossly polluted. An extremely tolerant species
46
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that is frequently the most abundant organism in areas sustaining
high degrees of organic enrichment. Several forms exist. See U
hoffmeisteri (variant form) and L_. hoffmeisteri (spiralis form).
Cosmopolitan. Widely reported in the United States and Canada.
17. Limnodrilus hoffmeisteri (spiralis form)
This form was first noted by Hiltunen (1967) in the Great Lakes and
applies to individuals possessing a penis sheath with a flat,
rounded head set perpendicular to the long axis of the sheath. The
name spiral is is applied to individuals of this type due to their
similarity to Eisen's (1885) Camptodrilus [aLimnodrilus] spiral is.
The correct rank of this taxon has not been satisfactorily
resolved. The authors have maintained this form as a distinct taxon
because of apparent differences in ecological requirements between
the typical and spi rali s form. The spi rali s form has been reported
from a variety of habitats but is generally most abundant in grossly
polluted habitats often attaining large population densities in the
absence of typical L_. hof fmei steri. Widespread? Known from St.
Lawrence Great Lakes, Mississippi River, Ohio River.
18. Limnodrilus hoffmeisteri (variant form)
Herein, this name is applied to any of several forms that possess a
scalloped margin on the head of the penis sheath. The status of
this taxon has not been satisfactorily resolved. The authors defer
from making a decision regarding taxonomic rank of these forms, but
concede that the morphology of these types is likely ecologically
related. Several authors make no distinction between the variant
form and typical specimens. In this treatment the variant form is
maintained as a distinct taxon for two reasons: First, the authors
have noted a discontinuous distributional pattern between variant
and typical forms, and secondly, the distinction is maintained in
hopes that ongoing and future investigations will elucidate the
ecological factor or factors affecting the occurrence of variant
types. Discontinuous distribution. Cosmopolitan? Known from
Hudson River, Mississippi River, Missouri River and St. Lawrence
Great Lakes.
19. Limnodrilus maumeensis
This species has been reported only from habitats sustaining high
degrees of organic enrichment. Frequently very abundant in polluted
harbors and river mouths. Nearctic. Known from St. Lawrence Great
Lakes, Illinois River, Mississippi River, and Louisiana.
20. Limnodrilus profundicola
Generally restricted to cold oligotrophic habitats. Rarely a
dominant species in any tubificid assemblage. Cosmopolitan. Known
from St. Lawrence Great Lakes, Nebraska, South Dakota, Wisconsin,
47
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Alberta, British Columbia, Manitoba, New Brunswick, Northwest
Territories, Ontario, and Saskatchewan.
21. Lirnnodrilus psammophilus
Described by Loden (1977) from material collected in Florida,
Louisiana, and Tennessee.
22. Limnodrilus rubripenis
Described by Loden (1977) from Louisiana. Subsequently reported by
Wetzel (1980) from a stream in southern Illinois.
23. Limnodrilus silvani
Distributional and ecological information sparse. A western species
known only from California (Brinkhurst, 1965) and one unpublished
record from Lake Ontario at the mouth of the Niagara River.
Hoi arctic.
24. Limnodrilus udekemianus
A long worm, rarely very abundant, usually littoral. Found in
organically polluted waters as well as oligotrophic habitats.
Cosmopolitan. Widespread in United States and Canada.
25. Monopylephorus helobius
A coastal species described by Loden (1980) from marine, estuarine
and freshwater habitats in the southern United States. Not likely
to be collected inland from the coast.
26. Phallodrilus hallae
Oligotrophic waters of Lake Huron and Lake Superior.
27. Potamothrix bavaricus
An uncommon species generally collected in littoral habitats.
Hoi arctic. Known from several localities in North America, ranging
from Lake Michigan (Green Bay), Illinois, Indiana to Utah,
California, and Oklahoma.
28. Potamothrix bedoti
This species, until Spencer's (1978) work, was considered a variant
form of £. bavaricus. It appears to prefer a silty, muck, substrate
in profundal areas of lakes. Holarctic. In North America known
only from St. Lawrence Great Lakes and Cayuga Lake, New York.
48
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29. Potamothrix hammonlensis
A common European species reported only twice in North America, from
Green Bay, Lake Michigan and one questionable record from Ontario in
Canada. The authors have not been able to locate specimens
collected in North America. Its occurrence, although possible,
awaits confirmation. The figure of P. hammoniensis (Fig. 32)
included herein was drawn based upon~~European material supplied by
R.O. Brinkhurst.
30. Potamothrix moldaviensis
A common and often abundant species in mesotrophic waters of the St.
Lawrence Great Lakes and its drainage. Often sympatric with £.
vejdovskyi. Hoi arctic.
31. Potamothrix vejdovskyi
A common and often abundant species in mesotrophic waters of the St.
Lawrence Great Lakes and its drainage. Often sympatric with P_.
moldaviensis. Hoi arctic.
32. Psammoryctides barbatus
Known only from Europe until reported by Vincent (1979) in the
freshwater St. Lawrence River upper estuary. Hoi arctic.
33. Psammoryctides cali form'anus
Uncommon. Originally described in a collection from California
(Brinkhurst, 1965). Has subsequently been reported from the St.
Lawrence Great Lakes (St. Marys River), Black River, Michigan, and
Cayuga Lake, New York. Nearctic.
34. Psammoryctides convolutus
Recently described by Loden (1978) in collections from swamp
habitats with fine organic muck substrates. Nearctic. Gulf Coast
swamps from Florida to eastern Mexico.
35. Psammoryctides minutus
A western species known only from Lake Tahoe in the United States and
Sturgeon Lake in Alberta, Canada.
36. Quistadrilus multisetosus
Common North American species that occurs in a wide range of
habitats but is found in greatest densities in organically enriched
habitats. Two subspecies, Quistadrilus multisetosus multisetosus
and £. m. longidentus originally described by Brinkhurst and Cook
(1966), have been maintained by many investigators but have not been
49
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included here because of the questionable validity of their
subspecific rank (Loden and Dugas, 1978). Specimens with anterior
and posterior ventral chaetae of similar shape are referable to Q.
m. longidentus; those with posterior ventral chaetae bearing reduced
distal teeth are referable to (). m. multisetosus. Nearctic. Known
from the St. Lawrence Great Lakes, Mississippi River, Ohio River,
Wisconsin and throughout the eastern United States.
37. Rhizodrilus lacteus
This nearctic species is rare in North America, known only from
Illinois and South Carolina.
38. Rhyacodrilus brevidentatus
Endemic to Lake Tahoe; not likely to be found elsewhere.
39. Rhyacodrilus coccineus
An occasional species, never very abundant. Ecological requirements
poorly known. Cosmopolitan. In North America known from the St.
Lawrence Great Lakes and its drainage and the Wisconsin River.
40. Rhyacodrilus falciformis
Small worms, ecology poorly known. Until recently, reported only
from Europe. Only one United States record; from Hudson River, New
York. In Canada, four specimens reported from Airport Creek,
Victoria, British Columbia.
41. Rhyacodrilus montana
An uncommon species generally found only in cold oligotrophic and
profundal habitats. Nearctic. In North America, known from the St.
Lawrence Great Lakes and its tributaries, British Columbia, Manitoba,
Northwest Territories, and Saskatchewan.
42. Rhyacodrilus punctatus
Recently reported in Lake Superior by Cook (1975). Prior to this
record, the species was known only from Europe.
43. Rhyacodrilus soda!is
An uncommon species with extremely variable chaetal morphology that
suggests taxonomic uncertainty. This species needs considerable
attention in a generic revision to verify its correct taxonomic rank
or position. Environmental requirements poorly documented.
Holarctic. Known in the United States from Alabama, Lake Ontario,
Louisiana, Tennessee, and in Canada from British Columbia, Manitoba,
Northwest Territories, and Saskatchewan.
50
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44. Spirosperma beetoni
Endemic to Lake Tahoe.
45. Spirosperma carolInensis
Known only from the southeastern United States in North Carolina.
46. Spirosperma ferox
A common, often abundant species in moderately enriched habitats,
and uncommon in oligotrophic or grossly polluted areas. Individuals
with a non-granular epidermis occur occasionally. Holarctic. In
North America patchily distributed in the Great Lakes. Louisiana,
and eastern North America.
47. Spirosperma nikolskyi
Based upon a recent revision by Brinkhurst (1981), this taxon
includes two formerly well known North American species, Peloscolex
oregonensis and _P. variegatus (sensu Brinkhurst, 1979c). An
uncommon species, generally most abundant in cold, oligotrophic
profundal habitats. A dominant organism, abundant in oligotrophic
profundal areas of the St. Lawrence Great Lakes. Holarctic. Known
from Alabama, Alaska, Great Lakes and the type locality of P_.
variegatus (sensu Brinkhurst, 1979c) Sckuylkill River, Pennsylvania,
Georiga, Louisiana, Mississippi, North Carolina, Oregon, Tennessee,
and Washington in North America.
48. Telmatodrilus vejdovskyi
A western species known only from California and British Columbia.
49. Tubifex harmani
Recently described by Loden (1979) from material collected in
Louisiana. Reproduces primarily by asexual fragmentation; sexually
mature individuals rare. Ecological preferences undocumented.
Nearctic. Known only from Florida, Indiana, Lousiana, Wisconsin,
and British Columbia.
50. Tubifex ignotus
An uncommon species that in North America is found primarily in
harbors and in lakes near river mouths. Holarctic. In North
America, known from Alabama, Lake Michigan, Lake Ontario, and the
St. Marys River.
51. Tubi fex kessleri americanus
An uncommon subspecies found in cold, oligotrophic, profundal areas.
Known from Lake Michigan, Lake Superior, and Wisconsin in the United
51
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States and from Alberta, New Brunswick, Northwest Terrestories,
Ontario, Saskatchewan, and the Yukon 1n Canada.
52. Tubifex nerthus
Recently combined with J. newfei by Brinkhurst (1978). A tolerant,
coastal species known in both brackish and freshwater. Holarctic.
Known from Newfoundland and New Brunswick.
53. Tubifex superiorensis
Transferred from Peloscolex to Tubifex by Brinkhurst (1981). A
small worm, occasional in cool, clean waters. Nearctic. Known
primarily from the Great Lakes and recently from the Mississippi
River, Wisconsin River and Red Cedar River in Wisconsin.
54. Tubifex tubifex
Indifferent to water quality, locally abundant in habitats ranging
from grossly polluted and organically enriched, to pristine alpine
and suhalpine lakes and streams. Often abundant in habitats
supporting few other species in both clean and perturbed waters.
Frequently confused with Ilyodri1us tempietoni. Cosmopolitan.
Widespread throughout northern North America, apparently rare in the
southern United States.
55. Varichaeta nevadana
Recently transferred from Isochaeta by Brinkhurst (1981). Endemic
to Lake Tahoe.
56. Varichaeta pacifica
Described by Brinkhurst (1981) from Washington and Alaska.
Apparently a cold stenotherm.
52
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59
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INDEX TO SCIENTIFIC NAMES
americanum (Bothrioneurum), 45
americanus (Aulodrilus), 5 29,44
amerjcanu?, (TubifexT~kessleri, 7,28,
51
angustipenis (Limnodrilus), 5,36,37,
Aulodrilus, 5
americanus, 5,29,44
Iimnobius7 5,30,44
pigueti, 5,17,44
pluriseta, 5,19,44
barbatus (Psammorycti des). 6,20,49
bavaricus (Potamothrix), 6,25,48
bedoti (Potamothrix), 6,24,48
beetoni (Spi rosperma), 7,15,51
Bothrioneurum, 5
americamim, 45
vejdovskyanum, 5,31,44,45
Branchiura, 5
spwerbyi, 5,12,17,42,45
breyidentatus (Rhyacodrilus), 6,18,
50"
cali form anus (Psammoryctides), 6,
Camptodrilus (Limnodrilus), 47
(form spiral is), 47
capi11atus (ITyodri1 us frantzi
form), 5,28,45
carolinensis (Spirosperma), 7,16,51
cervix (Limnpdrilus), 5,38,39,46
("variant form), 6,38,39,46
c1aparedianus (Limnodrilus), 6,38,
coccineus (Rhyacodrilus), 6,19,25,50
conyulutus (Psammorycti des), 6,25,
curvisetosus (Isochaetides), 5,32,45
falciformis (Rhyacodrilus), 6,35,
50
ferox (Spirosperma), 7,15,51
form capillatus, 5,28,45
(Ilyodrilus frantzi), 5,35,45
frantzi (Ilyodrilus), 5,35,45
form capi11atus , 5,28,45
freyi (Isochaetides), 5,33,46
hallae TPhallpdrilus). 6,35,48
hammpni ens i s (Potamothri x), 2,25,
Haber, 5
cf. speciosus, 5,23,24,45
harmam'ttubi f ex), 7,21,51
he!obi us (Mpnppylephorus), 6,30,48
hoffmeisteri (LimnodriTus), 6,38,39,
(spiralis form), 6,36,37,47
(van' ant form), 6,38,39,47
ignotus (Tubifex), 7,21,51
Ilypdrilus, 5
frantzi', 5,35,45
form capi11atus, 5,28,45
mastix, 5,16,45
tempietoni, 5,28,45,52
Isochaeta, 52
Isochaetides (Peloscolex), 5,45
curvisetosus, 5,32,45
freyi. 5,32,46
kessleri americanus, 7,28,51
TTubTfexTTT
lacteus (R"hizpdri1us), 6,33,50
limnobius (Aulodrilus), 5,30,44
LimnodriTus, 5
angustipenis, 5,36,37,46
ceryix, 57M, 39,46
Tvariant form), 6,38,39,46
claparedianus, 6,38,39,46
hoffmeisteTTT 6,38,39,45,46
(spirialis form), 6,36,37,47
(variant form), 6,38,39,47
maumeensis, 6,38,39,47
profundicp'la. 6,36,37,47
psammophilus, 6,36,37,48
rubripenis, 6,36,37,48
silvani, 6,36,39,48
udekemianus, 6,32,36,37,48
60
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longidentus, 49,50
(Quistadrilus multisetosus), 6,13,
mastix (Ilypdrilus). 5,16,45
maumeensis (Limnodrilus). 6,38,39,47
mi nutus (Psammoryctides), 6,25,49
moldaviensis (Potamothrix), 6,34,49
Monopylephorus, 6,
he!obi us. 6,30.48
montana (RFyacodrilus), 6,20,50
multisetosus (QuistadVilus), 6,13,14,
49
longidentus, 49,50
multisetosus, 49,50
nerthus (TubTFex), 7,22,52
nevadana (Varichaeta), 7,28,52
(Isochaeta), 52
newfei (TuSFex), 52
nikolskyi (Spirosperma), 7,13,16,51
oregonensis (Peloscolex), 51
pacifica fVarichaeta), 7.27.52
(Isochaeta), 52
Peloscolex, 45,46,52
curvisetosus. 45
freyi, 46
oregonensis, 51
superiorensis, 52
yariegatus, 51
Phallodmus. 6
hallae, 6,35,48
pigueti (Aulodrilus), 5,17,44
pluriseta (Aulodrilus). 5,19,44
PotamothfixTl>
bavaricus, 6,25,48
bedoti 7^.24,48
hammoniensis, 6,25,49
moldaviensis, 6,34,49
vejcfovskyi, 6,18.49
profundicpla (Limnodrilus), 6,36,37,47
psammophilus (Limnodrilus). 6,36,37,48
Psammprycti des, 6
barbatus, "g, 20,49
caliform anus, 6,25,49
convolutus, 6,28,49
minutus,T, 25,49
unctatus (Rhyacodrilus), 6,21,50
uistadrilus, 6
multisetosus 6,13,14.49
longidentus, 49,50
multi setgsus, 49,50
Rhizodrilus, 6
lacteus. 6,33,50
Rhyacodrilus. 6,
brevidentatus, 6,18,50
coccineus, 6,19,25,50
falciformls, 6,35,50
montana, "6720,50
punctatus, 6,21,50
sodalisT'6,25.50
rubripenis (Limnodrilus), 6,36,37,48
silvani (Limnodrilus). 6.36.39,48
soda!is (RhyacodrilTTs). 6,25,50
sowerbyi TSranchiura)'. 5,12,17,42,45
speciosus cf. (Haber), 5,23,24,45
(spiralis form), 6,36,37,47
(Limnodrilus hoffmeister), 6,46
47
spiral is (Camptodrilus). 47
Spirospenria, /
beetpnTT 7,15,51
carolinensis. 7,16,51
ferox, 7,15,51
nTkoTskyi, 7,13,16,51
(Limnodrilus), 6
superiorensis (Tubifex), 7,27,47
(Peloscol¥x), 52
Telematodrilus, 7
vejdovskyiT 7,29,51
tempietoni (Ilyodrilus). 5,28,45,52
Tubifex, 7.5T^
liamiani, 7,21,51
ignotus. 7,22,51
kessleri.7.28
americanus, 7,28,51
nerthus, 7,22,52
newfei, 52
superiorensis, 7,27,52
tubifex, 7,23,45,52
tubifex (Tubifex). 7,23,45,52
udekemianus (Limnodri1 us). 6,32,
36,37,48
(variant form), 6,38,39,47
(Limnodrilus hoffmeister), 6,45,46
(variant form), 6,38,46
(Limnodri 11 us cervjx), 6,46
Varichaeta (Isochaeta), 7,52
nevadana. 7,28,52
pacifica, 7,27,52
van egatus (Pel oscol ex), 51
vejdovskyanum (Bothrionerum), 5,31,44
?T"^
vejdovskyi (Potamothrix), 6,18,49
vejdovskyi (Telematodrilus), 7,29,
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