United States Environmental Monitoring and Support EPA 600 4-80-OjJ/
Environmental Protection Laboratory June 1980
Agency Cincinnati OH 45268
Research and Development
A Guide to the Naididae
(Annelida: Clitellata:
Oligochaeta) of North
America
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RESEARCH REPORTING SERIES
Research reports of the Office of Research and Development, U.S. Environmental
Protection Agency, have been grouped into nine series. These nine broad cate-
gories were established to facilitate further development and application of en-
vironmental technology. Elimination of traditional grouping was consciously
planned to foster technology transfer and a maximum interface in related fields.
The nine series are:
1. Environmental Health Effects Research
2. Environmental Protection Technology
3. Ecological Research
4. Environmental Monitoring
5. Socioeconomic Environmental Studies
6. Scientific and Technical Assessment Reports (STAR)
7 Interagency Energy-Environment Research and Development
8. "Special" Reports
9. Miscellaneous Reports
This report has been assigned to the ENVIRONMENTAL MONITORING series.
This series describes research conducted to develop new or improved methods
and instrumentation for the identification and quantification of environmental
pollutants at the lowest conceivably significant concentrations. It also includes
studies to determine the ambient concentrations of pollutants in the environment
and/or the variance of pollutants as a function of time or meteorological factors.
This document is available to the public through the National Technical Informa-
tion Service, Springfield, Virginia 22161.
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EPA-600/4-80-031
June 1980
A GUIDE TO THE NAIDIDAE (ANNELIDA: CLITELLATA: OLIGOCHAETA)
OF NORTH AMERICA
by
Jarl K. Hiltunen*
Great Lakes Fishery Laboratory
U.S. Fish and Wildlife Service
Ann Arbor, Michigan 48105
and
Donald J. Klemm*
Aquatic Biology Section
Environmental Monitoring and Support Laboratory
Cininnati, Ohio 45268
*co-authors
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 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,
periphyton, macrophyton, macroinvertebrates, and fish.
* Automate the measurement of physical, chemical, and biological
quality of water.
* Conduct an Agency-wide quality assurance program to assure
standardization and quality control of systems for monitoring
water and wastewater.
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. Naidid worms are important components of the
benthic fauna and are frequently abundant in a variety of freshwater
habitats. The varied response of naidid 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
Naididae have been published, this report 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 naidid oligochaetes.
Dwight 6. Ballinger
Director
Environmental Monitoring and Support
Laboratory - Cincinnati
iii
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ABSTRACT
In North America the aquatic annelid worms (Clitellata: Oligochaeta),
belonging in the family Naididae, are composed of 21 genera and 62
species. All taxa can be identified by external morphological features.
This guide presents the following: an introduction to the general
biology of the Naididae, collecting and processing methods, a species
list, an 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 naidid
taxonomy and ecology. /
iv
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CONTENTS
Foreword "Hi
Abstract iv
List of Figures vi
Acknowledgments ix
Section 1. Introduction 1
Section 2. Methods 5
Section 3. Species List 7
Section 4. Key to the Naididae of North America (North of Mexico,
Excluding Alaska) 10
Section 5. Glossary 30
Section 6. Annotations 32
Selected Bibliography 40
Index to Scientific Names 46
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LIST OF FIGURES
Number Pages
1 Generalized naidid, lateral view, anterior segments 2
2 Generalized forms of chaetae or bristles 3
3 Chaetogaster diaphanus, lateral view; anterior end "II"
and "III" indicate approximate position of segments
II and III 10
4 £. setosus, chaeta 11
5 £. limnaei, chaeta 11
6 Prostomium with a proboscis 12
7 Prostomium without a proboscis 12
8 Pristina aequiseta, enlarged ventral chaeta of segments
IV and/or V 13
9 P_. aequiseta, unenlarged ventral chaeta of segments other
than segments IV and/or V 13
10 P_. foreli, acicular chaeta 14
11 P. breviseta, acicular chaeta 14
12 Ophidonais serpentina, dorsal chaetae 14
13 A sigmoid-shaped chaeta, apex bifurcate 14
14 Amphichaeta sp., lateral view, anterior segments with
chaetae 15
15 A^ americana, chaeta 15
16 /L leydigi, chaeta 15
17 Dero sp., branchial apparatus with digitiform lobes 16
18 Dero sp., branchial apparatus with digitiform lobes and
palps 16
vi
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LIST OF FIGURES - continued
Number Pa9es
19 JD. trifida, acicular chaeta cleft, apex with an intermediate
~ tooth 17
20 JD. obtusa, slightly expanded digitiform lobes (gills) of
the branchial apparatus 17
21 JD. pectinata, acicular chaeta with few intermediate
teeth 18
22 2* flabel!iger, acicular chaeta 18
23 2- vaga, acicular chaeta with impression of fused
intermediate teeth 18
24 Sty!aria lacustris, showing proboscis and invagination of
the prostomium 19
25 Acicular chaeta with teeth equally long (Nais magnaseta) . . 20
26 Acicular chaeta with equal teeth (Dero abranchiata) 20
27 Allonais paraguayensis, acicular chaetae with unequal
primary teeth . . 20
28 2- abranchiata, ventral chaetae in segments II, VI,
and X, respectively 21
29 Specaria josinae, acicular chaeta 21
30 Slightly bent acicular chaeta (£. fraseri) 21
31 Nais communis, acicular chaetae with divergent teeth .... 22
32 _N. elinguis, acicular chaeta with teeth not divergent .... 22
33 N. variabilis, ventral chaeta of segment VI 23
34 N^. variabilis, acicular chaeta with short,
often obscure, essentially parallel teeth 23
35 _N. bretscheri, acicular chaeta 24
36 _N. bretscheri, ventral chaeta of segment VI, posteriad ... 24
37 N. bretscheri, giant ventral chaeta of segment VI,
posteriad 24
vii
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LIST. OF FIGURES - continued
Number Pages
38 N^. pardalis, acicular chaeta 24
39 N^. pardalis, ventral chaeta of segment VI, posteriad .... 24
40 N^. behningi, ventral chaeta of segments II-V 25
41 Unmodified ventral chaeta of segments II-V 25
42 Acicular chaeta with acutely attenuated apex 25
43 Acicular chaeta with somewhat short ("obtuse") apex .... 25
44 Uncinais uncinata, chaeta of segment II 26
45 Piguetiella michiganensis, chaeta 26
46 Bratislavia unidentata, ventral chaetae of
segments II and VI 27
47 B_. unidentata, acicular chaeta 27
48 Stephensoniana trivandrana, chaeta 27
49 £. longisoma, ventral chaeta 27
50 Acicular chaeta with divergent teeth 28
51 Acicular chaeta with divergent teeth 28
52 Acicular chaeta with parallel teeth (Pristina idrensis) . . 28
53 £. sima, acicular chaeta 29
54 £. idrensis, penial chaeta 29
55 P. longidentata, acicular chaeta 29
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ACKNOWLEDGMENTS
The authors thank Walter J. Harman and Kurt S. Stimpson 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 the Smithsonian Institution, U.S.
National Museum of Natural History, Department of Invertebrate Zoology,
Division of Worms, Washington, D.C.; The Academy of Natural Sciences of
Philadelphia, Department of Invertebrate Zoology, Philadelphia,
Pennsylvania; and the Naturhistoriska Riksmuseet, Sektionen for
Evertebratzoologi, Stockholm, Sweden, for permitting us to examine their
collections of Naididae. We are also indebted to the following
individuals for supplying distribution records: Michael T. Barbour,
James R. Brice, Michael S. Loden, Anthony F. Maciorowski, Bruce E.
Markert, Robert Schmal, Douglas R. Spencer, Kurt S. Stimpson, and David
R. Wefring.
ix
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SECTION 1
INTRODUCTION
The composition and abundance of benthic animals are commonly used to
demonstrate the effects of pollution on the biological integrity of
surface waters and changes in the biotic community resulting from the
destructive activities of man. The segmented worms, or Oligochaeta, are
one of the important components of the fauna collected during biological
investigations of surface waters. This group of invertebrates is
composed of fourteen families (Brinkhurst and Jamieson, 1971), including
the Naididae. The naidids are, in general, relatively small, commonly
1 mm to 10 mm long, and can easily be overlooked or ignored in sample
analysis, lost during sample reduction (even under standard sieving
procedures), or misidentified by investigators not familiar with their
morphology. Frequently the authors of environmental studies have
recorded the group only by subclass (Oligochaeta), family, genus, or
merely as "worms". The improper or inadequate treatment of the Naididae
is attributable, at least in part, to the lack of a practical key to
species. To perceive water quality requirements and pollution tolerances
of aquatic organisms, the animals must be identified to the species level
(Resh and Unzicker, 1975 and Carricker, 1977). In the few studies where
naidids have been identified to species, a relationship between species
assemblages and water quality is apparent (Hiltunen, 1967; Learner et
al., 1978; Learner, 1979). For example, Wapsa mobilis is abundant in the
polluted Saginaw Bay (Lake Huron), Michigan, but uncommon elsewhere in
the Great Lakes. Published data, albeit meager, indicate that some
species have a discontinuous or restricted distribution; e.g. a number of
species have been reported only from the Gulf States. In some instances
the apparent discontinuities may, however, be a function of insufficient
sampling.
Thus, the present guide to the ordinarily free-living naidid species
found in the United States and Canada was composed to assist aquatic
biologists in Federal, state, and private water monitoring agencies in
identifying specimens to the species level and to increase our knowledge
of the relationship between species composition and water quality. The
guide includes a species list, an illustrated key, a glossary, and an
annotation for each species. The drawings of the generalized naidid
(Fig. 1) and of the chaetae or bristles (Fig. 2) are especially important
in the family and species identification. The morphological terms
employed in the differentiation of species are defined in the Glossary.
A brief discussion of methods for processing specimens is also
presented. Dimensions and illustrations are based on preserved
material. A Selected Bibliography section was assembled from
1
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Prostomium
Eye
ro
Proboscis
Dorsal fascicle of chaetae
n
1 <
ffl
Ventral fascicles of chaetae
Anteriad Posteriad -
FIG. 1. Generalized naidtd, lateral view, anterior segments.
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Serrulate Smooth Simple-pointed Bifurcate
Capilliform chaetae
Acicular chaetae
FIG. 2. Generalized forms of chaetae or bristles (cf. Glossary).
-------
literature pertaining directly to the systematics and ecology of the
Naididae of North America. Distribution data are those of the authors as
well as others who have graciously made their records available to the
authors.
Inasmuch as new zoogeographical records of species are published
frequently, the present key may not include the latest 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.
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SECTION 2
METHODS
Naididae may be collected by any sampling method suitable to the need
of the collector, but inasmuch as many species are small and delicate,
care should be exercised in collecting and handling samples containing
naidids. Ideally the sample material should not be sieved. However,
abstention from washing the samples is often impractical. In most
instances it is desirable to wash mud from the sample, and this is best
done by putting small amounts of a sample in the sieve and agitating it
gently with the mesh submerged in water. The resultant residue con-
taining the oligochaete specimens is transferred into a widemouthed
container and fixed with 10% formalin. For studies where quantitative
retention of naidid specimens is desirable, a U.S. Standard No. 60 sieve
(60 meshes per inch, 0.250 mm openings) is recommended.
In the laboratory a dissecting microscope is required to discern
individuals in the sample. Forceps or medicine droppers can be employed
to remove specimens from the sieved residue. The oligochaetes can be
stored in a vial containing 10% formalin or, if desired, the material can
be mounted directly on microscope slides using a non-resinous mounting
medium containing a clearing agent (e.g. Hydramount*). In lieu of non-
resinous media the worms may be placed temporarily in Amman's lactophenol
(100 g phenol, 100 ml lactic acid, 200 ml glycerine, 100 ml water), a
medium which also clears tissues and eliminates the risk of specimen
dessication if a permanent mount cannot be prepared immediately following
extraction from the sample. The clearing process usually takes a few
hours to a few days depending on the size of the specimen. Gentle
application of heat will speed the clearing process. If the specimens
were preserved in 70% alcohol, they should be placed 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.
They can be held indefinitely in Amman's lactophenol or 10% formalin for
later processing and mounting.
1Bio/Medical Specialities (Box 1687, Santa Monica, CA 90406)
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Optimal resolution and longevity of mounted materials are achieved
only in resinous media (e.g. Canada Balsam, Harleco's Xylene
Coverbond2, etc.). These mounting media require dehydration of the
specimens through the alcohol series and clearing before using the
mountant, but they produce the best permanent mounts. The method can be
found in any standard biological techniques book (e.g. Knudsen, 1966;
Meyer and Olsen, 1971). Non-resinous media are recommended for rapid
processing of large numbers of specimens. For extremely important
reference specimens, a permanent resinous mounting medium is best.
A 12 or 18 mm diameter, No. 0 or 1, round cover glass is appropriate
because it will adequately accommodate the entire size range of naidids,
and the shape allows for maneuvering the specimen to rest in the most
desirable position by gentle rotation of the cover glass. When preparing
a temporary or permanent mount, an attempt should be made to place the
specimen on its side, thereby, revealing both dorsal and ventral
fascicles of chaetae. A variation from this is allowed with specimens of
Dero which must be viewed from the dorsal aspect, revealing the
arrangement of the branchial apparatus. The identification of species
requires a compound light microscope with oil immersion (1000X).
Caution must be taken in identifying naidids to species. Because
they undergo asexual reproduction by architomy (budding), daughter zooids
may not have the proper order of, or bear a normal complement of, full
sized chaetae. Observing unseparated zooids will provide the
investigator a clearer contrast between the features of a fully developed
parent and its daughter zooids.
Naidid material no longer needed in a study should be deposited in an
appropriate museum. In North America, the specimens with proper collec-
tion data can be sent to the Worm Division, Department of Invertebrate
Zoology, U.S. National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560.
Scientific Products (1430 Waukegan Road, McGaw Park, IL 60085)
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SECTION 3
SPECIES LIST *
Phylum Annelida
Class Clitellata
Subclass Oligochaeta
Oder Haplotaxida
Family Naididae
Genus Allonais Sperber, 1948
Allonais paraguayensis (Michaelsen, 1905)
Allonais pectinata (Stephenson, 1910)
Genus Amphichaeta Tauber, 1879
Amphichaeta americana Chen , 1944
Amphichaeta leydigi Tauber, 1879
Genus Arcteonais Piguet, 1928
Arcteonais lomondi (Martin, 1907)
Genus Bratislava Kosel. 1976
Bratislavia bilongata (Chen, 1944)
Bratislavia unidentata (Harman, 1973)
Genus Chaetogaster von Baer, 1827
Chaetogaster cristallinus Vejdovsky, 1883
Chaetogaster diaphanus (Gruithuisen, 1828)
Chaetogaster diastrophus (Gruithuisen, 1828)
Chaetogaster limnaei von Baer, 1827
Chaetogaster setosus Svetlov, 1925
Genus Dero Oken,
Dero abranchiata Harman, 1977
Dero digitata (Miiller, 1773)
Dero f label liger (Stephenson, 1931)
Dero furcata (Miiller, 1773)
Dero nivea Aiyer, 1930
Dero obtusa d'Udekem, 1855
Dero pectinata Aiyer, 1930
Dero trifida Loden, 1979
Dero vaga (Leidy, 1880)
*The year of publication for some old species descriptions varies in the
literature, and. we have chosen the dates given in Sperber (1948).
Hie maintain Aulophorus as a subgenus under Dero. Consequently, all the
species names conform to the feminine gender of Dero.
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Genus Haemonais Bretscher, 1900
Haemonais waldvoge/i Bretscher, 1900
Genus Homochaeta Bretscher, 1896
Homochaeta naidina Bretscher, 1896
Genus Nais Miiller, 1773
Nais alpina Sperber, 1948
Nais barbata Miiller, 1773
Nais behningi (Michaelsen, 1923)
Nais bretscheri (Michaelsen. 1899)
Nais communis Piguet, 1906
Nais e/inguis Miiller, 1773
Nais magnaseta Harman, 1973
Nais pardalis Piguet. 1906
Nais pseudobtusa Piguet, 1906
Nais simp/ex Piguet, 1906
Nais variabilis Piguet, 1906
Genus Ophidonais Gervais. 1838
Ophidonais serpentina (Muller, 1773)
Genus Parana!s Czerniavsky, 1880
Paranais litoralis (Muller, 1784)
Genus Piguetiella Sperber, 1939
Piguetiella michiganensis Hiltunen, 1967
Genus Pristina Ehrenberg, 1828
Pristina acuminata Liang, 1958
Pristina aequiseta Bourne, 1891
Pristina breviseta Bourne. 1891
Pristina foreli (Piguet. 1906)
Pristina idrensis Sperber, 1948
Pristina longidentata Harman, 1965
Pristina longiseta leidyi Smith, 1896
Pristina longiseta longiseta Ehrenberg, 1828
Pristina longisoma Harman, 1977
Pristina osborni Walton, 1906
Pristina plumaseta Turner, 1935
Pristina sima Marcus, 1944
Pristina synclites Stephenson, 1925
Genus Ripistes Duj, 1842
Ripistes parasita (Schmidt, 1847)
Genus Slavina Vejdovsky, 1883
Slavina appendiculata (d'Udekem, 1855)
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Genus Specaria Sperber, 1939
Specaria fraseri Brinkhurst, 1978
Specaria josinae (Vejdovsky, 1883)
Genus Stephensoniana Cernosvitov, 1938
Stephensoniana tandyi Harman, 1975
Stephensoniana trivandrana (Aiyer, 1926)
Genus Sty/aria Lamarck, 1816
Sty/aria fossularis Leidy, 1852
Sty/aria lacustris (Linnaeus, 1767)
Genus Uncinais Levinsen, 1884
Uncinais uncinata (0rsted, 1842)
Genus Vejdovskyella Michael sen, 1903
Vejdovskyella comata (Vejdovsky, 1883)
Vejdovskyella intermedia (Bretscher, 1896)
Genus Wapsa Marcus, 1965
Wapsa grand is Harman, 1977
Wapsa mobilis Liang, 1958
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SECTION 4
KEY TO THE NAIDIDAE OF NORTH AMERICA (NORTH OF MEXICO, EXCLUDING ALASKA)
1 With dorsal chaetae (in few or all segments) 7
Without dorsal chaetae 2
2(1) Ventral chaetae present in all segments; mouth ventral,
clearly surpassed by the conspicuous prostomium (Fig. 1)
Ophidonais serpentina
Ventral chaetae present in all segments except segment III;
mouth terminal, or subterminal; without prostomium (Fig. 3)
or, at most, prostomium slightly developed
Chaetogaster 3
FIG. 3. Chaetogaster diaphanus,
lateral view; anterior end
"II" and "III" indicate
approximate position of
segments II and III.
n m
%i;silll
?%&&
" '" - :*x$f>n'ifo\$
10
-------
3(2) Apex of chaetae simple or, if bifurcate (teeth visible under
ca. 1000X), then the proximal tooth is extremely slender,
appearing rudimentary and adherent to the slender distal
tooth (Fig. 4) Chaetogaster setosus
Apex of chaetae with a pair of distinct teeth (evident
under 400X or less magnification) 4
4(3) Teeth of chaetae distinctly reflexed (uncinate) (Fig. 5) ...
Chaetogaster limnaei
Teeth of chaetae not especially reflexed, similar to the teeth
on ventral chaetae in Fig. 1) 5
5(4) Length of chaetae in segment II 120 ym or greater; mouth
terminal, prostomium not developed 6
Length of chaetae in segment II less than 120 ym; mouth
subterminal, prostomium conspicuous
Chaetogaster diastrpphus
6(5) Length of chaetae in segment II less than 200 ym
Chaetogaster cristallinus
Length of chaetae in segment II greater than 200 ym
Chaetogaster diaphanus
FIG. 4. Chaetogaster setosus.
chaeta.
FIG. 5. Chaetogaster limnaei,
chaeta.
11
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7(1) Some or all dorsal fascicles with capilliform chaetae .... 8
All dorsal fascicles devoid of capilliform chaetae 16
8(7) Segment II with dorsal chaetae 9
Segment II without dorsal chaetae 24
9(8) Prostomium produced into a proboscis (Fig. 6) 10
Prostomium not produced into a proboscis (Fig. 7) 57
proboscis
FIG. 6. Prostomium with a
proboscis.
FIG. 7. Prostomium without a
proboscis.
12
-------
10(9) Apex of acicular chaetae appear simple under 1000X
Pristi'na longiseta longiseta
Apex of acicular chaetae appear bifurcate under 1000X ....
11
11(10) Ventral chaetae of segments IV and/or V conspicuously
larger, (Fig. 8) than those in other segments (Fig. 9) . .
Pristina aequiseta
Ventral chaetae of segments IV or V not conspicuously
larger than those in other segments 12
FIG. 8. Pristina aequiseta, enlarged
ventral chaeta of segments
IV and/or V.
FIG. 9. Pristina aequiseta, unenlarged
ventral chaeta of segments
other than segments IV and/or
V.
12(11) Capilliform chaetae in segment III conspicuously longer
(at least 1.3X) than those in other segments
Pristina longiseta leidyi
Capilliform chaetae in segment III not longer (may even be
shorter) than those in other segments 13
13(12) Teeth of acicular chaetae unequal in length 14
Teeth of acicular chaetae equal in length 15
13
-------
14(13) Capilliform chaetae serrulated (approximately
as in Fig. 2) ' . . . Pristina plumaseta
Capilliform chaetae not serrulated Pristina synclites
15(13) Teeth of acicular chaetae tiny, their length not more than
the thickness of the chaeta (Fig. 10). . . . Pristina foreli
Teeth of acicular chaetae long, their length about
twice the thickness of the chaeta (Fig. 11)
Pristina breviseta
16(7) Segment II with dorsal chaetae Homochaeta naidina
Segment II without dorsal chaetae 17
17(16) Dorsal chaetae straight, spiciform, apex simple or
minutely cleft (Fig. 12) Ophidonais serpentina
Dorsal chaetae at least slightly sigmoid, the apex distinctly
bifurcate (e.g. Fig. 13) 18
10
11
12
13
FIG. 10. Pristina foreli. acicular chaeta.
FIG. 11. P. brevisetaTacicular chaeta. (Modified from Bourne, 1891)
FIG. 12. Ophidonais serpentina, dorsal chaetae.
FIG. 13. A sigmold-shaped chaeta, apex bifurcate.
14
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18(17)
21(20)
With dorsal chaetae in segments III-IV; ventral chaetae
in these segments directed distinctly posteriad
(Fig. 14) Amphichaeta
Without dorsal chaetae in segments III-IV; ventral
chaetae in these segments nearly perpendicular to
the body (Fig. 1)
19
20
19(18) Six chaetae per fascicle in III, three per fascicle
in IV, distal tooth about twice as long as the
proximal tooth (Fig. 15) Amphichaeta americana
Five chaetae per fascicle in III, two per fascicle
in IV, distal tooth only slightly longer than proximal
tooth (Fig. 16) Amphichaeta leydigi
20(18) Segment V with dorsal chaetae 21
Segment V without dorsal chaetae 56
Length of distal tooth of anterior (ventral) chaetae about
twice that of the proximal tooth (similar to Fig. 15) . .
. Wapsa
22
Length of the distal tooth of (ventral) chaetae anteriad to
segment V, at most only slightly longer than the proximal
tooth (similar to Fig. 13) 23
15
16
FIG. 14. Amphichaeta sp., lateral view, anterior segments with chaetae,
FIG. 15. A_. americana, chaeta.
FIG. 16. y\. leydigi. chaeta.
'
-------
22(21) Length of chaetae in segment II 128-148 ym, about 7 chaetae
per fascicle Wapsa grandis
Length of chaetae in segment II 94-106 urn, 4 chaetae
per fascicle Wapsa mobilis
23(21) Length of ventral chaetae in segments II-IV about 60 ym; no
eyes Paranais litoralis
Length of ventral chaetae in segments II-V 100 pm or more;
older individuals with eyes Uncinais uncinata
24(8) Dorsal chaetae not present anteriad segment X
Haemonais waldvogeli
Dorsal chaetae present anteriad segment X 25
25(24) Digitiform lobes (Fig. 17) or both lobes and elongate
palps (Fig. 18) present on anal segment; without eyes . .
Dero 26
No digitiform lobes or palps on anal segment; with or
without eyes 33
digitiform lobes
FIG. 17. Dero sp., branchial
apparatus with digiti-
form lobes.
FIG. 18. Dero sp., branchial
apparatus with digiti-
form lobes and palps.
digitiform lobes
ie
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26(25) Only digitiform lobes present on anal segment 27
Both digitiform lobes and elongate palps present on
anal segment 30
27(26) Four or more pairs of gills; acicular teeth cleft, forming
two or more teeth of unequal length 28
Three pairs of gills; teeth of acicular chaetae essentially
equal in length 29
28(27) Apex of acicular chaetae cleft, forming two teeth,
distal tooth being longer than proximal tooth
Dero digitata
Apex of acicular chaetae cleft, forming two teeth
with a slender intermediate tooth (Fig. 19)
Derp trifida
29(27) In preserved material, end of anal segment exceeded
by the digitiform lobes (Fig. 20) Dero obtusa
In preserved material, end of anal segment clearly
surpasses the tips of the digitiform lobes (Fig. 17)
Dero nivea
19
digitiform lobes
FIG. 19. Dero trifida. acicular chaeta cleft, apex with an intermediate
tooth.
FIG. 20. £. obtusa, slightly expanded digitiform lobes (gills) of the
"branchial apparatus. (Modified from Sperber, 1948).
17
-------
30(26) Apex of acicular chaetae with one or more intermediate
teeth between the lateral pair, or the entire
tip expanded into a blade 31
Apex of acicular chaetae merely bifurcate
Dero furcata
31(30) Intermediate teeth at the apex of acicular chaetae
distinct (Fig. 21) Dero pectinata
Intermediate teeth at the apex of acicular chaetae
appearing fused to form a blade (Figs. 22, 23) 32
32(31) Blade-like apex of the acicular chaetae asymmetrical,
impression of fused teeth not clearly evident (Fig. 22)
Dero flabelliger
Blade-like apex of the acicular chaetae essentially
symmetrical, impression of the fused teeth evident
(Fig. 23) Dero vaga
21
22
23
FIG. 21. Dero pectlnata, aelcular chaeta with few intermediate teeth.
FIG. 22. £. flabelliger, acicular chaeta.
FIG. 23. J). vaga, acicular chaeta with impression of fused Intermediate teeth,
18
-------
33(25) Prostomium produced into a proboscis (Fig. 6 or 24) 34
Prostomium not produced into a proboscis (Fig. 7) 37
proboscis
FIG. 24. Stylaria lacustris,
showing proboscis and
invagination of the
pro s torn i urn.
invagination
34(33) More than three capilliform chaetae per fascicle 35
Not more than three capiTliforms per fascicle 36
35(34) Length of capilliform chaetae in segments VI-VIII
more than three times the length of those in segment IX . .
Ripistes parasita
Length of capilliform chaetae in segments VI-VII
less than three times the length of those in segment
IX Arcteonais lomondi
36(34) Proboscis originates in an invagination of the
prostomium (Fig. 24) Stylaria lacustris
Prostomium not invaginated, proboscis appears
as a narrow elongation of the prostomium
(Fig. 6) Stylaria fossularis
37(33) Segment III with dorsal chaetae
Bratislavia bilongata
Segment III without dorsal chaetae 38
38(37) Capilliform chaetae in segment VI conspicuously (2 times),
longer than those in other segments
Slavina appendiculata
Capilliform chaetae in segment VI not conspicuously
longer than those in other segments 39
19
-------
39(38) Apex of acicular chaetae bifurcate or pectinate 40
Apex of acicular chaetae simple 50
40(39) Acicular chaetae pectinate 41
Acicular chaetae merely bifurcate 44
41(40) All teeth of the acicular chaetae equally long or
nearly so (Fig. 25 or 26) 42
One tooth of the acicular chaetae distinctly longer than
the other teeth (Fig. 27) Allonais paraguayensis
25
26
27
FIG. 25. Acicular chaeta with teeth equally long (Nais magnaseta).
FIG. 26. Acicular chaeta with equal teeth (Dero abranchiate).
FIG. 27. Allonais paraguayensis, acicular chaeta with unequal primary teeth.
20
-------
42(41)
43(42)
44(40)
Shape of ventral chaetae in Segment II distinctly dissimilar
to those in Segment VI and posteriad (Fig. 28); lateral
teeth on the acicular chaetae not different from the
intermediate teeth (Fig. 26) Dero abranchiata
Shape of ventral chaetae in segment II essentially
like those in segment VI and posteriad; lateral teeth
on the acicular chaetae distinctly thicker than the
intemediate teeth (Fig. 25)
43
The distance from the nodulus to the apex in the acicular
chaetae is approximately 20 ym; the length of the acicular
teeth is about 4 ym, the lateral two teeth slightly divergent
(Fig. 25) Nais magnaseta
The distance from the nodulus to the apex in the acicular
chaetae is approximately 13 ym; the length of the acicular
teeth is about 3 ym, the lateral two teeth diverge slightly
more than those shown in Fig. 25 Allonais pectinata
Acicular chaetae gently sigmoid (Fig. 29); without
eyes Specaria josinae
Acicular chaetae straight or slightly bent (Fig. 30);
commonly with eyes
45
28
29
30
FIG. 28. Dero abranchiata. ventral chaetae in segments II, VI, and X,
res pectively.
FIG. 29. Specarla josinae, acicular chaeta.
FIG. 30. Slightly bent acicular chaeta (£. fraserl).
21
-------
45(44)
46(45)
Ventral chaetae in segment II thicker and longer than
those in segments III and IV ...;.... Specaria fraseri
Ventral chaetae in segment II identical to those in segments
III and IV 46
Size and shape of ventral chaetae in segment VI
are essentially like those in segments II-V 47
Size and shape of ventral chaetae in segment VI abruptly
dissimilar to, and shorter than those in segments II-V
48
47(46)
Teeth of acicular chaetae, short, appearing divergent
(Fig. 31); distal tooth of ventral chaetae in segment VI
and posteriad equal to or slightly larger than
proximal tooth Nais communis
Teeth of acicular chaetae not divergent, appearing parallel
(Fig. 32); distal tooth of ventral chaetae 1.5-2 times
longer than proximal tooth in segment VI and posteriad . . .
Nais elinguis
31
32
FIG. 31. Nais communis, acicular chaetae with divergent teeth.
FIG. 32. N. elinguis. acicular chaeta with teeth not divergent.
22
-------
48(46) Length of distal tooth of all ventral chaetae in segment VI
and posteriad equal to, or slightly shorter than,
that of the proximal (Fig. 33); acicular teeth short,
often obscure, appearing essentially parallel (Fig. 34) . .
Nais variabilis
Length of distal tooth of (at least some) ventral
chaetae in segment VI, and posteriad, 1.5 to several
times longer than that of the proximal tooth;
acicular teeth appearing either parallel or slightly
divergent 49
33
FIG. 33. Nais variabilis. ventral chaeta of segment VI.
FIG. 34. N_. variabilis. acicular chaeta with short, often obscure,
essentially parallel teeth.
23
-------
49(48) Teeth of acicular chaetae distinct, appearing to diverge
slightly (Fig. 35); in segment VI and posteriad, one or
more ventral chaetae per fascicle have relatively
thick distal tooth which is commonly directed strongly
posteriad, the proximal tooth is relatively small to
rudimentary (Figs. 36, 37) Nais bretscheri
Teeth of acicular chaetae obscure and appearing parallel
(Fig. 38); in segment VI and posteriad, one or more
ventral chaetae per fascicle, with distal tooth
longer than proximal, but not strongly bent
posteriad (Fig. 39) Nais pardalis
35
36
38
39
FIG. 35. Nais bretscheri, acicular chaeta.
FIG. 36. N_. bretscheri, ventral chaeta of segment VI, posteriad.
FIG. 37. N_. bretscherT, giant ventral chaeta of segment VI, posteriad.
FIG. 38. N^. parda1isr~aci'cular chaeta.
FIG. 39. _N. pardali's, ventral chaeta of segment VI, posteriad.
24
-------
50(39) More than three capilliform chaetae per anterior (i.e. segments
II-VI) fascicle 51
Not more than three capilliform chaetae per anterior (i.e.
segments II-VI) fascicle 52
51(50) In segment VI and posteriad, ventral fascicles composed
of three or more chaetae; eyes present; all ventral
chaetae alike Vejdovskyella comata
In segment VI and posteriad, ventral fascicles reduced
to a single chaeta; without eyes; often with enlarged
ventral chaetae in anterior segments (i.e. VI-VII)
Vejdoyskye11 a i ntermed i a
52(50) Distal and proximal teeth of ventral chaetae in segments II-V
slender, extremely reduced in thickness (Fig. 40) ....
Nai s behningi
Distal and proximal teeth of ventral chaetae in segments II-V
not disproportionately reduced (Fig. 41) 53
53(52) Apical part of acicular chaetae acutely attenuated (Fig. 42),
mean length of acicular chaetae, ca. 90 v\n 54
Apical part of acicular chaetae somewhat short ("obtuse")
(Fig. 43), mean length of acicular chaetae, ca. 60 ym
55
40
41
42
43
FIG. 40. Nais behningi, ventral chaeta of segments II-V.
FIG. 41. Unmodified ventral chaeta of segments II-V.
FIG. 42. Acicular chaeta with acutely attenuated apex.
FIG. 43. Acicular chaeta with somewhat short ("obtuse") apex.
25
-------
54(53) Ventral chaetae in segment VI short and stout,
their teeth equally long Na1s barbata
Ventral chaetae in segment VI thin, distal tooth
1.5 times as long as proximal tooth .... Nais pseudobtusa
55(53) Distal tooth of ventral chaetae about equal to
the proximal tooth Nais simplex
Length of distal tooth of ventral chaetae, ca. 1.5 times
the length of the proximal tooth Nais alpina
56(20) Distal tooth of chaetae in segment II distinctly
longer than proximal tooth (Fig. 44); older
individuals with eyes Uncinais uncinata
Distal tooth of chaetae in segment II equal to or
shorter than the proximal tooth; never with eyes (Fig. 45)
Piguetiella michiganensis
57(9) Apex of acicular chaetae simple 58
Apex of acicular chaetae bifurcate 60
FIG. 44. Uncinais uncinata, chaeta of segment II,
FIG. 45. Plquetiella michiganensis, chaeta.
26
-------
58(57) Anterior ventral chaetae dissimilar to those in segment VI,
(Fig. 46); acicular chaetae nearly sigmoid-spiciform
(Fig. 47) Bratislavia unidentata
All ventral chaetae essentially alike among the
segments; acicular chaetae slender 59
59(58) Distal tooth of ventral chaetae twice as long as
the proximal tooth (Fig. 48) ... Stephensoniana trivandrana
Distal tooth of ventral chaetae less than twice as
long as the proximal tooth (Fig. 49) . . . Pristina longisoma
46
47
48
FIG. 46. Bratislavia unidentata, ventral chaetae of segments II and VI,
FIG. 47. B. unidentata, acicular chaeta.
FIG. 48. Stephensoniana trivandrana, chaeta.
FIG. 49. Pristina longisoma, ventral chaeta.
27
-------
60(57) Teeth of acicular chaetae appearing divergent (Figs. 50, 51) 61
Teeth of acicular chaetae appearing parallel (Fig. 52) ... 62
61(60) Teeth of acicular chaetae distinctly unequal in size
(Fig. 51) Stephensoniana tandyi
Teeth of acicular chaetae essentially equal in size
(exclusive of intermediate teeth, when present) (Figs. 50,
53) 64
62(60) Capilliform chaetae serrulated 63
Capilliform chaetae not serrulated; penial chaetae
present in sexually mature individuals (Fig. 54); (cf.
Annotations) Pristina idrensis
50
51
52
FIG. 50. Acicular chaeta with divergent teeth.
FIG. 51. Acicular chaeta with divergent teeth.
FIG. 52. Acicular chaeta with parallel teeth (Pristina Idrensis),
28
-------
63(62) Dorsal fascicles are composed of one capilliform chaeta
and one acicular chaeta (Fig. 55) ... Pristina longidentata
Dorsal fascicles are composed of more than one
capilliform chaetae and one acicular chaeta
Pristina acuminata
64(61) Apex of the acicular chaetae with one or more small
intermediate teeth (Fig. 53)
Pristina sima
Apex of the acicular chaetae without intermediate teeth
(Fig. 50) Pristina osborni
53
55
FIG. 53. Pristina sima, acicular chaeta.
FIG. 54. £. idrensis. penial chaeta.
FIG. 55. P_. longidentata, acicular chaeta,
29
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SECTION 5
GLOSSARY*
Acicular chaeta: A dorsal needle-like chaeta usually with the apex
simple or cleft, forming two or more teeth, but sometimes greatly
modified into various other forms. Its length approximates the
thickness of the animal and ordinarily does not exceed half the length
of a capilliform chaeta.
Anal segment (Periproct); The posterior terminal segment in whole
(unbroken) individuals.
Anteriad: Direction toward the anterior of an organism.
Architomy: Reproduction by fission (budding) with subsequent regeneration
of tissues and organs.
Bifid; A condition where the apex of a chaeta is cleft, forming two
equal teeth.
Bifurcate: A condition where the apex of a chaeta is cleft, forming two
unequal teeth.
Branchial fossa: A pit or hollow of the anal segment in which the palps
and digitiform lobes or gills of Dero originate.
Capilliform chaeta: A dorsal hair-like chaeta; longer and usually more
flexuous than an acicular chaeta (cf. acicular chaeta).
Chaeta (pi. chaetae): A bristle, which in various forms, aids primarily
in locomotion or functions in connection with the reproductive
organs. (In literature, "seta" is frequently used instead of
"chaeta," but here the former term is reserved for its application in
arthropod morphology).
Clitellum: 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.
Crotchet: A somatic chaeta, commonly bifurcate, with conspicuous
teeth.
*Additional morphological terms applied in oligochaetology can be found
in Reynolds (1977).
30
-------
Digitiform lobes: See Gills.
Paired masses of pigmented photoreceptor cells in Segment I.
Fascicle: A cluster or "bundle" of chaetae.
Genital chaetae: Spermathecal and penial chaetae.
Genital segments: Body segments (ordinarily, V-VII) which bear the
reproductive organs.
Gills: The collection of protuberances or lobes that lie in the
branchial fossa of Dero. When distended in live material the lobes
may be digitiform.
Modulus: A knob or enlarged region on crotchets and acicular chaetae.
Palps: A pair of elongate projections in the anal segment of some species
of Dero. Their length clearly exceeds that of the gills, even in
preserved specimens.
Pectinate chaeta: A dorsal chaeta with apex cleft into few to several
spine-like teeth, thereby forming a comb.
Penial chaetae: Chaetae associated with the penes, ordinarily their shape
Ts~lmlike that of the somatic chaetae.
Posteriad: Direction toward the posterior of an organism.
Proboscis: A nonretractile, narrow elongation of the prostomium.
Prostomium: The antero-dorsal 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).
Simple chaeta: A chaeta with an uncleft apex.
Somatic chaeta: A chaeta which functions in connection with the somatic
segments only.
Spermathecal chaetae: Chaetae associated with the spermathecae, their
shape is unlike that of the somatic chaetae.
Spiciform: Spike-shaped as exemplified by acicular chaetae of Bratislavia
um'dentata and Ophidonais serpentina.
Trifid: A condition where the apex of a chaeta is cleft, forming three
teeth.
31
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SECTION 6
ANNOTATIONS
1. Allonais paraguayensls
Distribution: Lincoln Parish, Louisiana, and southward; adventive in
aquaria.
2. Allonais pectinata
Distribution: St. Clair River, Ontario, Canada; Illinois River near
Marseilles, Illinois; Mississippi River near Cordova, Illinois;
Mahoning and Ohio Rivers, Ohio; Monongahela River, Pennsylvania;
Hudson River near Albany, New York, and coastal Georgia.
3. Amphichaeta americana
Distribution: Union Lake, near Millville, New Jersey (type
locality); Siskiwit Lake, Isle Royale, Lake Superior; St. Marys
River, Chippewa County, Michigan; South Carolina.
4. Amphichaeta leydigi
Distribution: Widespread.
5. Arcteonais lomondi
Distribution: Widespread. Swims with tortuous or wriggling
movements.
6. Bratislava bilongata
Distribution: Hardingville, New Jersey and Indian River County
Florida.
7. Bratislavia ym'dentata
Distribution: Lake Erie; Illinois River to Oklahoma, Alabama,
Florida, Louisiana, Mississippi, North Carolina, and Texas. This
species was first included under Pristina because acicular chaetae
were present in segment II (in fact, acicular chaetae are found also
in segment I in most individuals). Following the discovery of
sexually mature specimens, Harman and Loden (1978) transferred the
species to the present genus.
32
-------
8. Chaetogaster cristallinus
Distribution: Yukon and Northwest Territories, the Great Lakes and
eastward. The validity of distributional records for the species is
uncertain because of its occasional confusion with £. diaphanus.
9. Chaetogaster diaphanus
Distribution: Widespread. The species' predatory and even
cannibalistic behavior is unusual among oligochaetes.
10. Chaetogaster diastrophus
Distribution: Widespread. This species is difficult, if not
impossible, to distinguish from C. langi; consequently, the latter is
maintained in synonomy under £. 7iastrophus following the rules of
nomenclatorial priority. The first author has examined some of
Christina Sperber's specimens to which she applied either name. The
distinction between the two taxa in her material was not clear.
Recent collections of Great Lakes oligochaetes made by the first
author have yielded few specimens that resemble £. diastrophus except
that they bear not only the normal compliment of ventral chaetae, but
also opposing dorsal chaetae; the later features being incongruous
with the definition of the genus.
11. Chaetogaster limnaei
Distribution: Widespread, usually associated with mollusks.
Specimens of C,. limnaei, taken off Lymnaea stagnalis (Gastropoda) at
Charlevoix, Michigan, August 6, 1894, may be the earliest collection
of Naididae from the St. Lawrence Great Lakes. The specimens are
housed at the Academy of Natural Sciences of Philadelphia,
Pennsylvania. In literature (e.g. Gruffydd 1965a, b) some subspecies
have been recognized but these are not treated here.
12. Chaetogaster setosus
Distribution: St. Marys River, Chippewa County, Michigan; Saginaw
Bay, Lake Huron; Lake Erie; Lake Ontario; Hudson River near Albany,
New York.
13. Dero abranchiata
Distribution: Louisiana and Texas.
14. Dero digitata
Distribution: Widespread.
33
-------
15. Dero flabelliger
Distribution: Orange County, Florida; Hudson River, New York; Lake
Wylie, North Carolina.
16. Dero furcata
Distribution: Ontario and Wisconsin to Texas, Alabama, Maryland,
Virginia and New York.
17. Dero nivea
Distribution: Widespread.
18. Dero obtusa
Distribution: Massachusetts to Washington, including James River
Virginia; Orange County Texas; Oconto River, Wisconsin. Distinction
of this species from £ nivea is often difficult.
19. Dero pectinata
Distribution: Orange County, Texas and other southern states. This
species should not be confused with ]) pectinata which enters the
genus through the subordination of Aulophorus.ID. (A,.) pectinata has
not been reported from North America.
20. Dero trifida
Distribution: Louisiana; Cape Fear River, Chatham County, North
Carolina.
21. Dero vaga
Distribution: Washtenaw and Livingston Counties, Michigan; Hancock
County, Mississippi; Young County, Texas; Massachusetts.
22. Haemonais waldvogeli
Distribution: Michigan and Wisconsin to Louisiana, Texas,
Mississippi, Maryland, Virginia, and New York.
23. Homochaeta naidina
Distribution: The Richmond, Virginia record is apparently the only
report of the species in North America. Voucher specimens of Falls
(1974) are reported lost.
24. Nais alpina
Distribution: Lake Ontario (Judd and Bocsor, 1975); Chippewa County
and Presque Isle County, Michigan.
34
-------
25. Nais barbata
Distribution: The Great Lakes and Fox River, Wisconsin, to the
Mississippi River, Illinois River, and Jackson Parish, Louisiana;
Penobscot River, Maine.
26. Nais behningi
Distribution: Widespread, frequently in lotic habitats. The
morphological features of this species and N_. pseudobtusa appear to
overlap.
27. Nais bretscheri
Distribution: Widespread.
28. Nais communis
Distribution: Widespread.
29. Nais elinguis
Distribution: Widespread.
30. Nais magnaseta
Distribution: Bee County, Texas. The species resembles Allonais
pectinata and may better be placed in Allonais.
31. Nais pardalis
Distribution: Widespread. The species closely resembles _N.
yariabilis and is often difficult to distinguish from that species.
Morphology of its chaetae can vary with respect to season and with
habitat.
32. Nai s pseudobtusa
Distribution: Richmond, Virginia; Lincoln Parish, Louisiana; St.
Marys River, Chippewa County, Michigan; British Columbia and
northwest Canada.
33. Nais simplex
Distribution: Widespread east of the Mississippi River.
34. Nais variabilis
Distribution: Widespread. One of the most commonly occurring
naidids in North American freshwater environments.
35
-------
35. Ophidonais serpentina
Distribution: Widespread. The compliment of dorsal chaetae on an
individual, as well as between individuals, may vary from none to
several chaetae.
36. Paranais litoralis
Distribution: Lake Michigan (?); Elizabeth River, Virginia; Slack
Reach (Cooper River), Berkeley County, South Carolina, and Flag
Creek, South Carolina; Piscataway Creek (Potomac River tributary),
Maryland; Nova Scotia, New Brunswick, Ontario, British Columbia,
Yukon and Northwest Territories. Re-examination of the voucher
specimen on which P_. literal is was reported from Saginaw Bay, Lake
Huron (Brinkhurst, 1967) revealed that the individual is Piguetiella
michiganensis.
37. Piguetiella michiganensis1
Distribution: Great Lakes and Mississippi River to Susquehanna and
Chemung Rivers, New York.
38. Pristina acuminata
Distribution: Lake Erie and Ohio River. The record of this species
in North America is attributable to Spencer (1978b). When Spencer's
voucher specimens were examined, few of their features (e.g. lack of
serrulations on the capilliforms) did not fully agree with the
description of P. acuminata. A comparison of specimens collected
from the type locality (China) and North America may be necessary to
resolve the morphological differences.
39. Pristina aequiseta
Distribution: Michigan to Texas and California, east to Alabama and
New York.
40. Pristina breviseta
Distribution: Widespread.
41. Pristina foreli
Distribution: Yukon and Northwest Territories; Great Lakes and
Mississippi River to Nebraska; Ohio River; Chemung, Susquehanna, and
Hudson Rivers in New York; Penobscot. River, Maine.
^After the manuscript went to press, we discovered that benthic samples,
collected from Lake Michigan by the Great Lakes Research Division of the
University of Michigan, contained some individuals that had chaetae in
segment V and posteriad.
36
-------
42. Pn'stina idrensis
Distribution: Yukon and Northwest Territories; St. Marys River and
Lake Huron, Michigan; Buckhorn Creek, Chatham County, North Carolina;
Cedar River, King County, Washington; Mississippi, Ohio, Muskingum,
Monogahela, Susquehanna Rivers, east to the Hudson River and Lake
Umbagog, New Hampshire. On 29 September 1968, the first author
collected, from Lake Huron, two sexually mature or elite!late
specimens, one of which bears a pair of penial chaetae (Fig. 52).
Sperber (1948) contended, however, that P_. idrensis lacks genital
chaetae.
43. Pristina longidentata
Distribution: Garfield County, Oklahoma; Louisiana.
44. Pristina longiseta leidyi
Distribution: Widespread.
45. Pristina longiseta longiseta
Distribution: Several streams and lakes in Michigan, and likely
elsewhere in the Great Lakes drainage. Historically, the binominal,
P. longiseta, has been cleaved into subspecies or even new species
Fased upon the presence of bifid or simple acicular chaetae. Inas-
much as that acicular feature is the only significant character
employed to subdivide P_. longiseta, we are maintaining the form with
bifid acicular chaetae as the subspecies P. J_. leidyi.
46. Pristina longisoma
Distribution: Arkansas and Louisiana to Florida.
47. Pristina osborni
Distribution: Great Lakes and Wisconsin to Louisiana, east to
Virginia, Pennsylvania and Maine. P_. minuta has been reported from
Bee County, Texas (Harman, 1973), but we excluded it from the key
because of its uncertain validity. Sperber (1948) considers £.
minuta and £. osborni as probable synonyms.
48. Pj-istina plumaseta
Distribution: Ontario, Georgia, Virginia, Mississippi and the
Potomac River, Virginia.
37
-------
49. Pristina sima
Distribution: Mississippi, Susquehanna, Chemung, and Hudson Rivers.
If the intermediate teeth on the aciculars are not visible, the
individual is indistinguishable from P_. osborni.
50. Pristina synclites
Distribution: Mississippi and Illinois Rivers; Bayou Deview,
Arkansas; Chemung River at Corning New York; Susquehanna River at
Binghamton, New York; Kanawha River, West Virginia.
51. Ripistes parasita
Distribution: Known in Europe but reports of the species in North
America are unconfirmed.
52. Slavina appendiculata
Distribution: Widespread.
53. Specaria fraseri
Distribution: Fraser River, British Columbia.
54. Specaria josinae
Distribution: Widespread.
55. Stephensoniana tandyi
Distribution: Louisiana.
56. Stephensoniana trivandrana
Distribution: Kanawha River, West Virginia; Illinois River at
Marseilles, Illinois; Ohio River; Pennsylvania; Piscataway Creek
(Potomac river tributary), Maryland; Colorado River, Texas.
57. Sty1 aria fossularis
Distribution: Great Lakes and Wisconsin to Louisiana, Alabama, South
Carolina, West Virginia, and Maine. Specimens of S^. fossularis,
collected at Erie, Pennsylvania in 1896 (under the supervision of
Jacob Reighard) are held in The Academy of Natural Sciences of
Philadelphia, Philadelphia, PA.
38
-------
58. Sty1 aria lacustris
Distribution: Widespread. Specimens of S_. lacustris, collected from
Lake St. Clair and western Lake Erie in 1899, are deposited in The
Academy of Natural Sciences of Philadelphia, Philadelphia, PA.
59. Uncinajs uncinata
Distribution: Widespread. Superficially similar to Picjuetiella
michiganensis with which it is often sympatric in the Great Lakes.
The relatively long distal teeth in anterior chaetae help distinguish
it from P. michiganensis. Vernal populations in the Great Lakes (and
ManitobaJ compose individuals with chaetae in segment V and posteriad.
60. Vejdovskyella comata
Distribution: Wisconsin and Menominee Rivers, Wisconsin; Bush River,
Hartford County, Maryland; Penobscot River, Maine; Nova Scotia,
Canada. Eyes present.
61. Vejdovskyella intermedia
Distribution: Widespread in Great Lakes drainage; Hudson River, New
York; Piscataway Creek (Potomac River tributary), Maryland; Muskingum
River; Illinois River. Never with eyes.
62. Wapsa grandis
Distribution: Terrebone Parish, Louisiana.
63. Wapsa mobilis
Distribution: Widespread, often in estuaries and rivers, component
species of brackish or freshwater. Species of Wapsa are morpho-
logically similar to those in Paranais; consequently, VJ. mobilis is
sometimes included under Paranais as P. frici.
39
-------
SELECTED BIBLIOGRAPHY
Adams, D.E., and R.D. Kregear. 1969. Sedimentary and fauna! environments
of eastern Lake Superior. Int. Assoc. Great Lakes Res., Proc. 12th
Conf. Great Lakes Res. 20 pp.
Barbour, Michael T. 1977. Chaetogaster limnaei limnaei (Oligochaeta:
Naididae) inhabiting the mantle cavity of the pill clam Sphaerium.
Trans. Am. Microsc. Soc. 96(1): 141-142.
Barton, O.R., and H.B.N. Hynes. 1978. Wave-zone macrobenthos of the
exposed Canadian shores of the St. Lawrence Great Lakes. J. Great
Lakes Res. 4(1): 27-45.
Brinkhurst, Ralph 0. 1964. Studies on the North American aquatic
Oligochaeta I: Naididae and Opistocystidae. Proc. Acad. Nat. Sci.
Phi la. 116(5): 195-230.
1967. The distribution of aquatic oligochaetes in
Saginaw Bay, Lake Huron. Limnol. Oceanogr. 12(1):137-143.
. 1975. Oligochaeta. In: F.K. Parrish, ed.
Keys to the Water Quality Indicative Organisms of the Southeastern
United States, U.S. Environ. Prot. Agency, Environ. Monit. & Support
Lab., Cincinnati, OH. pp. 69-85
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45
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INDEX TO SCIENTIFIC NAMES
abranchiata (Dero), 7,21,33
acuminataTPristina), 8,29,36
aequiseta (Pristina'), 8,13,36
AllonaisT 7,35
paraguayensis, 7,20,32
pectinata, 7721,32,35
alpina (Nais), 8,26,34,
americana (Amphichaeta), 7,15,32
Amphichaeta, 7,15
americana, 7,15,32
leydigi, 7,15,32
Annelida, 7
appendiculata (Slavina), 9,19,38
Arcteonais, 7
lomondi, 7,19,32
Aulophorus, 7,34
barbata (Nais), 8,26,35
behningi jNaTs), 8,25,35
bilongata (Bratislava), 7,19,32
Bratislava, 7
bilongata, 7,19,32
unidentata, 7,27,31,32
bretscheri (Nais), 8,24,35
breviseta""(Pristina), 8,14,36
Chaetogaster, 7,10
cristallinus, 7,11,33
diaphanus, 7,11,33
diastrophus, 7,11,33
lanqi, 33
langi,
limna<
imnaei, 7,11,33
setosus, 7,11,33
Clitellata, 7
comata (VejdoyskyelTa), 9,25,39
communis (Nais), 8,22,35
cristall'inus (Chaetogaster), 7,11,33
Dero, 6,7716".30,31
abranchiata, 7,21,33
digitata. 7,17,33
flabel!iger, 7,18,33
furcata, 5718,34
nTvea7~8,17,34
obtusa, 8,17,34
pectinata, 8,18,34
trifida, 8,17,34
vaga, 8,18,34
diaphanus (Chaetogaster), 7,11,33
diastrophus (Chaetogaster), 7,11,33
digitata (Dero), 7,17,33
elinguis (NaiT), 8,22,35
flabelliger (Dero), 7,18,33
foreli (PrTstTnaT, 8,14,36
fossularis (Stylaria). 9,19,38
fraseri (Specaria), 9,22,38
frici (Paranais), 39
furcataTpero), 8,18,34
grandis (Vlapsa), 9,16,39
Haemonais, 8
waldvogeli, 8,16,34
Haplotaxida, 7
Homochaeta, 8
naidina, 8,14,34
idrensis (Pristina), 8,28,37
intermedia (Vejdoyskyella), 9,25,39
josinae (Specaria), 9,21738
lacustris (Stylaria), 9,19,39
langi (Chaetogaster), 33
1 imnaei (Chaetogaster), 7,11,33
litoralis (Paranais), 8,16,36
longidentata (Pristina), 8,29,37
longiseta leidyi (Pristina), 8,13,37
longiseta loncpseta (Pristina), 8,13,37
longisoma (Pristinir), 8,27,37
leydigi TArcphichaeta), 7,15,32
lomondi (Arcteonais), 7,19,32
Lymnaea, T3
stagnalis, 33
magnaseta (Nais), 8,21,35
michiganensis (Piguetiella), 8,26,36,39
minuta (Pristina), 37
mobilis TWapsa), 1,9,16,39
Naididae,"/
naidina (Homochaeta), 8,14,34
NaTsT
alpina, 8,26,34
barbata, 8,26,35
penning i, 8,25,35
46
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bretscheri , 8,24,35
communis, 8,22,35
elinguis, 8,22,35
magnaseta, 8,21,35
pardalis, 8,24,35
pseudobtusa, 8,26,35
simp Tex, 8,26 , 35
variab ilis, 8,23,35
nivea (bero), 8,17,34
obtusa TDero), 8,17,34
Oligochaeta, 7
Ophidonais, 8
serpentina, 8,10,14,31,36
osborni (PrTstina), 8,29,37,38
pardalis (Nals). 8,24,35
paraguayensis (Allonais), 7,20,32
Paranais, 8,39
frici, 39
litpralis, 8,16,36
parasita (Ripistes), 9,19,38
pectinata (Allonais). 7,21,32
ectinata (Dero), 8,18,34
~
michiganensis, 8,26,36,39
plumaseta (Pristina), 8,14,37
Pristina. 87?5
ac urn in at a. 8,29,36
aequiseta, 8,13,36
breviseta, 8,14,36
foreli, 8,14,36
idrensis. 8,28,37
longidentata, 8,29,37
longiseta leidyi, 8,13,37
longiseta Tongiseta, 8,13,37
longisoma, 8,27,37
minuta, 37
osborni, 8,29,37,38
plumaseta, 8,14,37
sima, g,29,38
synclites, 8,14,38
unidentata, 32
pseudobtusa (Nais), 8,26,35
Ripistes, 9,19
parasita, 9,19,38
serpentina (Ophidonais). 8,10,14,31,36
setosus (Chaetogaster), 7,11,33
sima (Pristina), 8,29,38
simplex (Nais), 8,26,35
Slavina, 9
appendiculata, 9,19,38
Specaria, 9
fraseri, 9,22,38
josinae, 9,21,38
stagnails (Lymnaea), 33
Stepnensoniana, 9
tandyi, 9,28,38
trivandrana, 9,27,38
Stylaria, 9
fossularis, 9,19,38
1acustris7 9,19,39
synclites (Pristina), 8,14,38
tandyi (Stephensoniana), 9,28,38
trifida (Dero), 8,17734
triyandrana (Stephensoniana), 9,27,38
Uncinais, 9
uncinata, 9,15,26,39
uncinata (Uncinais), 9,15,26,39
unidentata (Bratislavia), 7,27,31,32
vaga (Dero), 8,18,34
yarTabTTTs (Nais), 8,23,35
Vejdovskyella, 9
comata, 9,25,39
intermedia, 9,25,39
waldvogeli (Haemonais), 8,16,34
Wapsa, 15,39
grandis. 9,16,39
mobilis, 1,9,16,39
47
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TECHNICAL REPORT DATA
(Please read Instructions on the reverse before completing)
REPORT NO.
EPA-600/4-80-031
3. RECIPIENT'S ACCESSION-NO.
TITLE AND SUBTITLE
A GUIDE TO THE NAIDIDAE (ANNELIDA:CLITELLATA:
OLIGOCHAETA) OF NORTH AMERICA
5. REPORT DATE
June 1980 issuing date
6. PERFORMING ORGANIZATION CODE
. AUTHOR(S)
8. PERFORMING ORGANIZATION REPORT NO.
Jarl K. Hiltunen and Donald J. Klemm, Co-authors
PERFORMING ORGANIZATION NAME AND ADDRESS
Great Lakes Fishery Laboratory, U.S. Fish Wildlife
Service, Ann Arbor, Michigan 48105 and Aquatic Biology
Section, Environmental Monitoring & Support Laboratory
U.S. Environmental Protection Aaencv. Cincinnati. Ohio
10. PROGRAM ELEMENT NO.
PE 1BD612
11. CONTRACT/GRANT NO.
12. SPONSORING AGENCY NAME AND ADDRESS
Environmental Monitoring and Support Laboratory
Office of Research and Development
U.S. Environmental Protection Agency
Cincinnati, Ohio 45268
13. TYPE OF REPORT AND PERIOD COVERED
14. SPONSORING AGENCY CODE
EPA/600/06
15. SUPPLEMENTARY NOTES
Supplement to "Biological field and labortory methods for measuring
the quality of surface waters and effluents." EPA-670/4-73-001.
16. ABSTRACT
In North America the aquatic annelid worms (ClitellatarOligochaeta), belonging in
the family Naididae, are composed of 21 genera and 62 species. All taxa can be
identified by external morphological features. This guide presents the following:
an introduction to the general biology of the Naididae, collecting and processing
methods, a species list, an 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 naidid taxonomy and
ecology.
17.
KEY WORDS AND DOCUMENT ANALYSIS
DESCRIPTORS
b.lDENTIFIERS/OPEN ENDED TERMS
COSATI Field/Group
Aquatic Biology
*Freshwater Biology
Indicator Species
*Benthos
Taxonomy
Segmented worms
Distribution
Collection
Preservation
Aquatic Fauna
*Annelida
Water Pollution
Macroinvertebrates
*Naididae
*Clitellata
*01igochaeta
North America
Species List
*I1lustrated Key
identification Manua
6C
6F
13. DISTRIBUTION STATEMENT
RELEASE TO PUBLIC
19. SECURITY CLASS (ThisReport)
CURITY CLASS (Thi
UNCLASSIFIED
21. NO. OF PAGES
58
20. SECURITY.
is page)
22. PRICE
EPA Form 2220-1 (9-73)
48
US GOVERNMENT PRINTING OFFICE 1980-657-146/5693
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