b. ,H.HwdBS ILL
I
1
JOAN P, JASS
T H E CRAYFISHES
.-
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& SHRIMP:
OF WISCONSIN
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,
MILWAUKEE PUBLIC MUSEUM
*
*
THE CRAYFISHES 6SHRIMP
OF WISCONSIN
(Cambaridae, Palaemonidae)
H, H. Hobbs 111
Department of Biology
Wit t enberg University
Joan P. Jass
Invertebrate Zoology Section
Milwaukee Public Museum
Milwaukee Public Museum
@
1988 Milwaukee Public Museum
ISBN 0-89326-152-1
Library of Congress Cataloging-in-Publication Data
Hobbs, H. H.
The crayfishes and shrimp of Wisconsin (Cambaridae,
Palaemonidae)
(Special publications in biology and geology; no. 5)
Bibliography: p.
Includes index.
1. Decapoda (Crustacea)-Wisconsin. 2. CrustaceaWisconsin. I. lass, Joan P., 1944. 11. Title.
III. Series.
QL444.M33H6 1988 595.3’841
87-31415
ISBN 0-89326-152-1
TABLE OF CONTENTS
................................................
INTRODUCTION .....................................................
NOTES ON PRESENTATION ............................................
HISTORY OF WISCONSIN CRAYFISH INVESTIGATIONS .....................
METHODS ...........................................................
Collecting Techniques .................................................
Cambaridae .......................................................
Palaemonidae ......................................................
Water Samples .....................................................
Preservation Procedures ................................................
Identification Procedures ...............................................
Cambaridae .......................................................
Palaemonidae ......................................................
PHYSIOGRAPHY OF WISCONSIN ........................................
GEOLOGY OF WISCONSIN .............................................
Bedrock Geology .....................................................
GlacialHistory ......................................................
SURFACE DRAINAGE IN WISCONSIN ....................................
LIST OF WISCONSIN DECAPODS ........................................
KEY TO DECAPODS OF WISCONSIN .....................................
ACKNOWLEDGMENTS
.................................................
Cambaridae .........................................................
Cambarus (Lacmicambarus) diogenes Girard .................................
SPECIESTREATMENT
v
1
2
2
4
4
4
9
9
9
9
9
12
13
16
16
17
19
19
22
29
29
29
...................................
Orconectes (Oremicambarus) immunis (Hagen) ................................
Orconectes (Crockerinus) propinquus (Girard) .................................
Orconectes (Procericambarus) rusticus (Girard) ................................
Orconectes (Oremicambarus) virilis (Hagen) ..................................
Procambarus (Girardiella) gracilis (Bundy) ...................................
Procambarus (Ortmannicus) acutus acutus (Girard) .............................
Palaemonidae .........................................................
Palaemonetes kadiakensis Rathburn ........................................
GLOSSARY ..........................................................
LITERATURE CITED ..................................................
Fallicambarus (Creuserius) fodiens (Cottle)
APPENDIX I: WISCONSIN DECAPODS AND THEIR COUNTY AND
DRAINAGE OCCURRENCES
................................
APPENDIX 11: DISTRIBUTION MAPS .....................................
INDEX ...............................................................
39
43
52
66
79
97
105
114
114
123
127
141
143
173
ACKNOWLEDGMENTS
Many persons have assisted us in virtually all
parts of this study. Special thanks goes to Stanley
Rewolinski, University of Wisconsin Milwaukee, whose tireless efforts in the field helped
us fill many locality gaps. He also was most helpful
in plotting locality records supplied by the
Wisconsin Department of Natural Resources.
The cooperation of the Wisconsin Department
of Natural Resources was continuous, and we
extend our appreciation to Donald Fago, David
Siegler, Ruth Hine, and George Boronow for
their assistance in supplying maps, identifying
localities, and for their interest and patience.
George Becker of Stevens Point also was most
helpful in sharing maps and advice that were
invaluable. For supplying locality data for
Minnesota species, we thank Judy Helgen of St.
Olaf College. For lending specimens and
volunteering locality data, we are indebted t o
Marian Havlik and David Heath of La Crosse.
Karl Korschgan and John Caruthers are thanked
for their assistance and advice in field work in
the environs of La Crosse. We extend our
gratitude to Edward M. Stern of the University
of Wisconsin
Stevens Point for loaning
specimens, for supplying us with working space,
and for permitting us to examine the collection
of decapods at Stevens Point. For supplying
specimens and for informative discussions,
appreciation is extended to Steve Yeo of the
Center for Great Lakes Studies, Milwaukee.
James G. Lorman of Edgerton College is thanked
for supplying specimens and locality data
particularly important in determining the range
of Orconectes rusticus in Wisconsin. Appreciation
-
-
is extended to Jean Manley of the Milwaukee
County Printing Office, for reproducing state
maps for plotting of locality records. We are
indebted to James J. Flannery of the University
of Wisconsin Milwaukee Geography Department for supplying assistance and data concerning drainage basins. Thanks to Janice Mahlberg
of the Milwaukee Area Technical College for
photographing the majority of crayfishes shown
in this publication. We extend our appreciation
to Howard Mead, James A. Hilliard, and their
associates a t t h e University Cartographic
Laboratory, University of Wisconsin Madison,
for the plotting of and final listing of locality
data (available from the Milwaukee Public
Museum).
Many individuals in the Milwaukee Public
Museum were most helpful throughout the
study. Appreciation is extended to volunteers
Harold Bauers, Barbara Klausmeier, and Mary
Lou Ryan who assisted in proofreading, plotting
locality data, and maintaining a systematic
organization of specimens in the Museum. For
encouragement, interest, and assistance in the
field, we extend special appreciation to Leon
Zukrow, also a volunteer and member of the
Friends of the Museum. For field assistance and
continued encouragement we are grateful to
Susan Borkin and Robert Murray. Drew
Hildebrandt is thanked for plotting locality data.
We are indebted to Patricia Laughlin, Museum
librarian, for obtaining references and maps for
us. Appreciation is extended to Kurt Hallin of
the Geology Section for reading the sections
treating the physiography of Wisconsin. The
-
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V
advice of Teresa Noeske was invaluable in
statistical analyses and in entering data in the
Museum’s computer system. Me1 Scherbarth,
photographer, generously advised us on photographic matters and additionally produced
photographs of a number of our crayfish
specimens. Special thanks is given to Museum
volunteer Harry Pease, who supplied continual
encouragement. Had it not been for Max A.
Nickerson, Curator of Vertebrate Zoology, who
suggested t o the Wisconsin Department of
Natural Resources that they save all the
crayfishes and shrimps collected during the
Wisconsin Fish Distribution Study, we would not
have been able to be as complete in assessing
t h e geographic distribution, ecology, life
histories, etc. of Wisconsin decapods.
For their assistance with proofing we are
grateful to C. Beth Sunderland, Donna D’Angelo, and John F. Wing of Wittenberg University.
To Georgia B. and Susan K. Hobbs, we extend
our thanks for proofreading synonymies, locality
records, and the bibliographic citations. Horton
H. Hobbs, Jr. of the Smithsonian Institution,
assisted us in many phases of this study,
Collections and working space in the Museum
of Natural History were made available as was
his reprint collection and locality data. We are
deeply grateful for continual discussions and for
his criticisms of preliminary drafts of the
manuscript.
Funds for various aspects of this study were
received from The Institution for Museum
Services, Washington, D.C., from Friends of the
Museum (we wish to thank Kenneth Starr,
Director of the Milwaukee Public Museum and
Allen Young, Curator of Zoology, Milwaukee
Public Museum, for allocating these funds), and
from a grant from the Faculty Research Fund
Board, Wittenberg University. Finally, we would
like t o thank Mary Garity, Publications Supervisor, Milwaukee Public Museum, for her
interest, patience, and assistance demonstrated
in editing this manuscript.
The Crayfishes & Shrimp
of Wisconsin
LAKE
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O
W
A
\ I
92'
SU PERlOR
91.
L
L
90'
I
N
8 9.
0
I
S
8 8.
Figure 1. Localities in Wisconsin from which crayfishes have been collected; solid circles-specimens
open circles-from literature; circles represent one or more localities.
viii
/
87.
examined by us;
INTRODUCTION
The first official list of Wisconsin decapods
was published by Bundy in 1876. Although
numerous publications appeared subsequent to
that, it was not until 1932 when Creaser
presented a comprehensive survey of t h e
distribution of Wisconsin crayfishes and shrimp;
no such studies have been conducted since that
date. During the elapsed 56 years, many events
within the state have altered decapod distributional patterns. Most relate to man’s activities
which have caused local extinctions, range
extensions, and the establishment of an exotic
species, Orconectes (Procericurnburus) rusticus
(Girard) in portions of the state. Creaser’s
account of the decapods was based on holdings
derived primarily from the extensive collections
of the Wisconsin Geological and Natural History
Survey from all counties within the state
excepting several from along the Lake Michigan
shore. T h e present distribution differs
considerably and reflects the extensive impact
that man has had on the aquatic environment
during the ensuing 56 years.
Crayfishes and, to a much lesser degree,
shrimp, are important members of Wisconsin’s
aquatic communities. They are used as bait for
certain game fishes, are valuable components of
aquatic ecosystems in that they serve as major ‘
processors of detritus and can greatly affect the
biomass of primary and secondary producers,
have value as game fish forage, and are consumed
by man on a small scale in various parts of the
state. In addition several species of burrowing
crayfishes cause problems in agricultural areas
by damaging both irrigation and earthen dam
structures. When these aspects are coupled with
the need to conserve threatened, endangered
(see Bouchard 1976b), or endemic species, the
need for a compendium on the ranges, abundances, life histories, and ecology of Wisconsin
crayfishes and shrimp is obvious.
The distribution of many North American
cambarid crayfishes reflects both alterations in
drainage patterns and the climatic changes
associated with Pleistocene glaciation (Ortmann
1902, 1905, Rhoades 1962a, Fitzpatrick 1967,
and others). In Wisconsin, glaciation and
associated events altered climate and drainage
patterns, and through the establishment of
postglacial drainages allowed for (re)invasion of
aquatic systems by crayfishes and other organisms. The extent to which the Driftless Area
might have provided refuge for crayfishes is
unclear. Although speciation has not been
detected in Wisconsin since the retreat of the
glaciers, suspected hybrids have been observed.
Such apparent stability is in sharp contrast to
the high species richness of crayfishes found in
the southeastern United States, the proposed
center of origin of American cambarids where
crayfishes have maintained a much longer period
of continuous habitation than in Wisconsin.
With an increase in mobility man has become
a much more effective dispersal agent for a
number of organisms, including crayfishes. As
has been demonstrated in various areas of the
world, man has introduced crayfishes which have
subsequently become not only firmly established
1
but often have replaced native species throughout drainage systems (Schwartz et al. 1963, Aiken
1965, Hobbs and Walton 1966, Crocker and Barr
1968, Abrahamsson 1973, Huner 1977, Berrill
1978). Such changes have occurred within this
group of crustaceans in Wisconsin and are
treated in greater detail below.
In the accounts that follow, we have attempted to compile the known information concerning decapods within the political boundaries of
Wisconsin (see Fig. 78 for a map of county
locations). We are most fortunate to have had
collections from many areas of the state that
were procured between 1974-1981by the Bureau
of Research, Wisconsin Department of Natural
Resources. While collecting fishes in various
watersheds (Fish Distribution Study), the
Department also retained crayfishes and shrimps
that were gathered with the fishes and housed
them in the Milwaukee Public Museum. Our
concentrated field efforts from July to November
1982 filled many locality “gaps” for various
species. In addition, collections maintained in
the National Museum of Natural History
(Smithsonian Institution), The Ohio State
University Museum of Zoology, and at the
University of Wisconsin Stevens Point were
examined. Most of the total of 13,656 specimens
examined during this study were obtained from
the localities noted in Fig. 1.
+
NOTES ON PRESENTATION
Literature pertaining to crayfishes and shrimp
in Wisconsin is summarized within and followed
by a discussion of the geology, physiography, and
relatively recent geologic history of the state.
The two major drainages (St. Lawrence and
Mississippi) are subdivided and pertinent data
are presented. An overview of crayfish and
shrimp taxonomy is given and a key to the
Wisconsin species is provided. A section
discussing habitats sampled, methods of collection, preservation, and identification is included,
and a key to the decapods of the state is
presented. In the treatment of each species,
synonymies, with the exception of the initial
citation, are restricted to references pertaining only
to Wisconsin, and precede diagnoses (based
2
primarily on Form I males). The discussion of
each species is extended to include variations,
measurements, and color observations. The typelocality is listed and size data are presented in
tabular form. Information on the ecology of each
species, including observations from throughout
its range, is given. Life history data are presented
and summarized in tables. The geographic
distribution is stated and figured, and a spot map
of the location from which specimens were
collected is included. These represent collections
primarily from the University of Wisconsin
Stevens Point (now housed in the Milwaukee
Public Museum), the Wisconsin Department of
Natural Resources, and our field work during
the summer and fall of 1982.
A list of the Wisconsin decapods and the
counties and drainage basins in which they occur
appears in Appendix I. This appendix is preceded
by a glossary and a bibliography.
Numerous articles have adequately treated the
natural history, general and specific ecology, and
general biology” of these organisms (Turner
1926, Hobbs 1942a, Williams and Leonard 1952,
Crocker 1957, Waterman 1960, 1961, Crocker
and Barr 1968, Capelli and Magnuson 1983, and
Hazlett 1983), thus we shall not duplicate those
efforts herein. The reader is directed to these
works as well as to the species treatments below
for these kinds of information.
-
((
Subgenera used herein from:
Fitzpatrick, J. F., Jr. 1987. The subgenera of the crawfish
genus Orconectes (Decapoda: Cambaridae). Proc. Biol.
SOC.Wash. 100(1):44-74.
HISTORY OF WISCONSIN
DECAPOD INVESTIGATIONS
Literature citations are included in the
treatment of each species, yet they do not furnish
an integrated historical summary of the contributions to the knowledge of the decapods of
Wisconsin. In the following account we have
not included every isolated reference that has
been made, yet we have attempted to cite most
original studies as well as monographic and
summary articles.
The first mention of a decapod in Wisconsin
was in 1870 when Hagen published a monograph
on North American crayfishes. He indicated that
“. , . fresh-water crabs are found near Milwaukee
. ..”
(p. 101) and that C. virilis [=Orconectes
(Gremicambarus) virilis] was found in the Sugar
River in southcentral Wisconsin (pp. 64,97). He
also listed C. propinquus, [ =O. (Crockerinus)
propinquus] C. rusticus, and C. virilis as having
been collected by L. Agassiz from Lake Superior
but the precise locality within the lake is
unknown.
In 1874 Smith (p. 638) indicated that C.
propinquus occurred in Wisconsin and that the
species was present in a lake at Madison. Two
years later the first official list of Wisconsin
crayfishes appeared (Bundy 1876). In this
publication Bundy described Cambarus stygius
[=P. (Ortmannicus) a. acutus] and Cambarus
wisconsinensis [=O.virilis] from Racine County,
and Cambarus debilis [=O. virilis] from the
Baraboo River, Ironton, in Sauk County; this
latter species was also reported from the
Wisconsin River. In addition he indicated that
C. virilis [=O.virilis] was found in the Rock River,
and, although he described C. gracilis [=P.
(Girardiella) gracilis] from Illinois, he reported
that it occurred on the prairies in the environs
of Racine. Bundy (1882) published an updated
tally of the state’s crustaceans, presenting
additional descriptive information and geographical data for the 11 species he listed. In addition
to those species previously mentioned, Bundy
reported: 1) C. acutus [=P. (0.)
a. acutus] from
Racine and Sauk counties; 2) C. placidus
[misidentified, as this crayfish (0.
placidus) is
restricted to the
. Cumberland, Duck, and
Tennessee drainage systems in southern Kentucky, Tennessee, and northern Alabama”
(Hobbs 1974b:38) - possibly the single specimen
from the Fox River was confused by Bundy with
Orconectes rusticus?]; 3) C. rusticus [=O.rusticus]
from Lake Superior (reported earlier by Hagen,
1870, to have been caught by Louis Agassiz in
Lake Superior); 4) Cambarus barconii [=C.
(Cambarus) b. bartonii] from Lake Superior about
which he states, “I do not think this species
has been found in the interior of the state” (p.
183); and 5) C. obesus [=C.(Lacmicambarus) d.
diogenes], which is described by him (p.183) as
being one of the largest and most abundant species.
“. .
Bundy made a similar listing the following year
adding that C. rusticus has also been collected
from Beloit, Rock County. This may be the first
bona fide record of 0. rusticus in Wisconsin.
All previous references to this species in the state
are based on Hagen’s (1870) account of the Lake
Superior specimen.
Rathbun (1884) discussed crayfishes as part
of the “fisheries business” and indicated that
Wisconsin supplied the city of New York with
excellent tasting crayfishes. Hay (1896:5OO)
stated that in the lake regions of Wisconsin and
Minnesota C. virilis is used extensively as food.
This was one of the first accounts of the
economic importance of crayfishes in Wisconsin
and it also was the last t o treat Wisconsin
crayfishes prior to the turn of the century.
Much of the literature that appeared after
1900 pertaining t o Wisconsin decapods is
descriptive and treats primarily the geographic
distribution of species. Of importance, in 1932
Creaser published “The Decapod Crustaceans
of Wisconsin,” the only definitive work on this
group in the state. He discussed information on
the distribution and ecology of crayfishes and
shrimps the doubtful occurrence of C. rusticus
in Wisconsin, and presented synonymies of
crayfishes.
Cahn published an in depth ecological study
of the Wingra Springs region southwest of
Madison in 1915, listing Cambarus diogenes and
C. propinquus, indicating that in the Spartina
(marsh-grass) “society” of the area, “The most
abundant crustacean is Cambarus argillicola
[=Fallicambarus (Creaserius) fodiens] , though in
the drier places of the society Cambarus diogenes
is to be found” (p. 136). Finding this reference
was particularly exciting to us as it represents
the only record of this species in the state. We
have been unable to locate his specimens;
however, we feel that since he mentioned
collecting C. diogenes, it would have been
unlikely that he would have confused the two
species. It is difficult to conceive that other
Wisconsin crayfishes could be mistakenly
identified as C. argillicola; however, Turner
(1926:187) noted that Cambarus argillicola “. .
is tentatively regarded as a local variation of C.
.
3
diogenes.” Unfortunately, field efforts in October
1982 and July 1984 failed to produce any
specimens (as one might expect, the area of study
has been altered greatly in the 73 years since
his work was published), yet we consider this
to be a valid report of the occurrence of this
species in the state.
Ellis (1919) noted a number of species of
branchiobdellid worms associated with crayfishes, and reported infestations of C. virilis and
C. diogenes in Rhinelander, Oneida County.
Other investigators to note the occurrence of
parasites or commensals on crayfishes in
Wisconsin are: Marshall 1903, Tressler 1947,
Hart and Hart 1974 (entocytherid ostracods);
Creaser 1932 (discodrilid oligochaetes); and
Hoffman 1963 (branchiobdellid worms).
Much research has been conducted on several
Wisconsin crayfishes in the past decade. Stein
(1975a,b, 1976) reported on many aspects of the
biology of 0. propinquus, as did Capelli and
Magnuson (1976). Lorman (1975) discussed the
biology (primarily feeding activity) of 0. rusticus
in Wisconsin, He (1980) presented results of a
detailed population study (1976-1978) of 0.
rusticus from a Vilas County lake. Lorman and
Magnuson (1978) demonstrated the role of
crayfishes in aquatic ecosystems and noted that
0. rusticus significantly reduced the abundance
of macrophytes in many northern Wisconsin
lakes in which this crayfish has been introduced
in the last “3-15 years” (p. 8).Lodge (1984) found
that herbivory and predation by natural densities
of 0.rusticus contribute to the reduction of the
abundance and diversity of benthic flora and
fauna. Concern over the introduction of 0.
rusticus into the state initiated several studies.
Capelli ( 1980)discussed how hybridization and
mating interference could affect species displacement among these three species. The same year
Capelli and Capelli presented morphological
evidence for hybridization between 0. propinquus and 0.rusticus in Wisconsin and suggested
ten morphological characters to describe and
separate 0.propinquus, 0.virilis, and 0.rusticus.
Also, in 1981, the Wisconsin Department of
Natural Resources (Anonymous 1981) published
regulations concerning the capture of crayfishes
4
in Wisconsin waters.
Capelli (1982a) presented a discussion on the
displacement of the native crayfish (0.
uirilis)
by 0. propinquus and 0. rusticus in northern
Wisconsin. He (p.742) suggested that 0.
propinquus was introduced prior to 0. rusticus,
that 0. propinquus is capable of displacing 0.
virilis, and that 0. rusticus is able to displace
both species. In December of 1982 the Department of Natural Resources (Anonymous
1982c:224) prohibited the introduction of
crayfishes into the state, the ultimate goal being
curtailment of the spread of 0. rusticus within
Wisconsin. Hayes (1985)and Lodge et al. (1985)
presented the most recent review of the impact
that 0.rusticus is having on northern Wisconsin
lakes.
After 118 years of research on Wisconsin
decapods, the number of species reported stands
at nine (yet, the presence of F. fodiens has not
been substantiated). Data gathered on these
species are summarized below. Of particular
importance is the relatively recent invasion
(introduction?) of 0. rusticus into Wisconsin
lakes and streams and its impact on aquatic
communities. Much future effort will be required
to understand the mechanisms involved in so
successful a series of established populations.
Continued distributional spreading should
precipitate state funding for crayfish investigations. We recommend that research should be
directed towards control of the spread of this
exotic species in Wisconsin waters.
METHODS:
Collecting Techniques
Cambaridae:
Crayfishes are found in a diversity of habitats
and a variety of techniques are employed in
attempting to capture them. In shallow streams
(less than 1.5m) that have little vegetation (Fig.
2), a 1/4-inch mesh seine (2m or longer) can
be handily used. The seine (Fig. 3) should be
anchored approximately 2-3m downstream of
the area to be sampled; enter the water and
position the net from downstream. One end of
the seine should be placed against the shore or
bank; take care to anchor the bottom of the
net and position the opposite pole slightly
upstream. Areas in the vicinity of bridges,
particularly those immediately downstream,
should be kicked with caution as much debris
(glass, wire, pieces of metal, etc.) may be
encountered. Various versions of the “Crawdaddy Stomp” may be used in kicking the stream
bank and in agitating the stream bottom.
Crayfishes dislodged from debris piles, (Fig. 4)
shallow streambank burrows, and from the
hyporheic zone will be carried by the current
or may “back-stroke” into the net.
Figure 3. Root River in Milwaukee County; inhabited by
Orconectes (C.)
popinquus, 0. (P.) rusticus, 0. (G.) virilis,
and Procambarus (0.)
a. acutus.
Figure 2. Hibbard’s Creek in Door County; inhabited by
Orconectes (C.) propinquus.
Figure 4. Hibbard’s Creek in Door County; inhabited by
Orconectes (C.)
propinquus.
5
bank; take care to anchor the bottom of the
net and position the opposite pole slightly
upstream. Areas in the vicinity of bridges,
particularly those immediately downstream,
should be kicked with caution as much debris
(glass, wire, pieces of metal, etc.) may be
encountered. Various versions of the “Crawdaddy Stomp” may be used in kicking the stream
bank and in agitating the stream bottom.
Crayfishes dislodged from debris piles, (Fig. 4)
shallow streambank burrows, and from the
hypotheic zone will be carried by the current
or may “back-stroke” into the net.
Figure 3. Root River in Milwaukee County; inhabited by
Orconectes (C.)
Propinquw, 0. (P.) rwticus, 0. (G.)
virilis,
and Procambarus (0.)
a. acutus.
Figure 2. Hihhard’s Creek in Door County; inhabited by
Orconectes (C.)propinquus.
Figure
4. Hihhard’s Creek in Door County; inhabited by
Orconectes (C.)propinquw.
5
Figure 5. Chippewa River in Sawyer County; inhabited by
Orconectes (G.)
uinlis.
Figure 6. Littoral zone of Lake Michigan in Door County;
Orconrctes ( C ) propinquus and 0 (G.) ouilis collected in
this locality.
6
Pooled areas of streams (Fig. 5), shallow ponds,
and the littoral zone of lakes (Fig. 6) may be
sampled effectively by pulling the seine across
them, taking care to insure that the anchored
margin of the seine remains in contact with the
substrate.
Use of various wire traps is recommended;
inverted cones with openings enlarged slightly,
4 to 5cm diameter, seem to give the best results
in vegetation-choked streams (Fig. 7), ponds, or
lakes or when sampling deep waters of lakes. A
residence time of one to two nights has proven
to be most productive for traps baited.
A sturdy dipnet (Fig. 8) is very useful for
sampling ditches (Fig. 9),overhanging vegetation,
mud-bottomed pools, and areas choked with
vegetation. A dipnet also can be used to collect
specimens from the rocky substrates of lakes (Fig.
10) and streams by placing the net adjacent to
a rock, lifting it gently, and by either forcing
the animal into the net with the hand or by
netting it as it swims from its place of hiding.
Crayfishes in most environmental situations
are more active at night than during the day.
They can be collected in shallow water with the
aid of a headlamp and a dipnet; the light reflected
by their eyes is ruby-colored, thus making them
easy to detect.
In the Fish Distribution Study conducted by
the Wisconsin Department of Natural Resources, small-meshed seines and four different types
of electrofishing equipment were utilized (see
Norotny and Priegel 1971, 1974, and Fago 1982,
1986).
Probably all Wisconsin crayfishes burrow at
least occasionally for one or more reasons. Some
species are rarely found outside of burrows,
others leave them for short periods (hours or
for parts of seasons), whereas others spend only
a small part of the year in these subsurface
habitats (see discussion of burrowers in various
species treatments below). Data gathered by one
of us (HHH) on C. (L.) d. diogenes from a wetland
meadow in southern Indiana indicate (as one
might assume) that the diameter of the burrow
cavity gives a good indication of the size of the
crayfish inhabiting it (Fig. 11); chimney pellet
size IS also a useful index (Fig. 12) (substrate
Figure 7. Yellow River
(G.)
wirilis.
in Burnett County; inhabited by 0.
Figure 8. Raccoon Creek in Rock County; inhabited by
Orconectes (C.) propinpus and 0. ((3.1wirilis.
Figure 9. Ditch in Milwaukee County; burrows of
Procanburus (G.)gracilis.
Figure 10. Venus Lake in Oneida County; inhabited by
Orconectes (P.) rusticus.
7
texture also may influence species). One then
may glance at a burrow opening on the surface
and assume that the smaller openings (burrows)
contain juveniles and thus concentrate on
burrows of larger diameter in order to sample
the adult population. After selecting a promising
burrow, the chimney(s), if present, should be
removed carefully and any loose soil brushed
from the opening. If water is in the tube, the
mouth of the burrow should be enlarged to a
level several cm below the water surface, forming
a submerged platform. The water should be
agitated vigorously and then left undisturbed for
several minutes. Individuals of many species will
surface after a minute or two (e.g., C. diogenes,
not P. gracilis - commonly a great portion of
the burrow must be dissected in order to capture
the latter species); antennae will be observed
waving at the air-water interface. Be certain that
the crayfish is at least partially out of the burrow
proper and on the platform. At this time an
open hand should be thrust swiftly into the
+
[
r=0.72 r=0.86
5
4.0
3.5
m
“F
LL
0
3.0
25
d
-
”
10
20
30
40
50
P O S T - O R B I T A L C A R A P A C E L E N G T H (mrn)
Figure 11. Plot showing relationship of post-orbital
carapace length and burrow diameter of Cambarus (L.)
diogenes from Monroe County, Indiana; females (open
circles), males (filled circles).
8
opening, pinning the animal to the side of the
burrow and/or onto
the platform. With
careful
manipulation, the individual can be seized with
the fingers and extracted. If the crayfish retreats
into the burrow before it can be pinned, usually
it returns again to the surface within a few
minutes. If, however, one touches the crayfish
but is unable to capture it, normally it will retreat
into the burrow and will not resurface for a
considerable period of time. In this case a
dissection of the vertical tunnel may be
necessary.
Crayfishes, but few Wisconsin species,
construct complex ramifying burrows and
apparently an inverse relationship may exist
between burrow complexity and the depth of
the water table; that is, the greater the depth
of the water, the less complex the burrow (see
Hobbs 1 9 8 ~ 3 3 )Occasionally
.
crayfishes will not
be found in the vertical tunnel but in one of
the side branches.
More often than not, when a chimney is
initially removed no water is observed in the
burrow, indicating a low water table, and either
water should be poured into the burrow to raise
the level, or soil removed until the water level
is reached. Since it is difficult to maintain the
level of the water by the pouring technique, a
hole may be dug parallel to the burrow to prevent
soil particles from tumbling into and clogging
the crayfish’s excavation; this is a particular
problem when digging in sandy soil. The burrow
may then be dissected, allowing the dirt to fall
away from the tunnel. This may require digging
to a depth of 2m or more below the surface!
Once water is reached the procedure described
above is employed to capture the resident. The
use of gloves is of no help, and, if used, the
crayfish is often crushed before one realizes that
it has been cornered.
O n warm, rainy, humid nights, collections can
be made more easily. The burrowing crayfishes
come to the mouths of their burrows and often
leave them t o forage for terrestrial and
subaquatic macrophytes, invertebrates, etc.
With the aid of a headlight occasionally they
can be collected in large numbers.
Palaemonidae:
Shrimps are found in sluggish lotic or lentic
environments and are most easily collected with
the aid of a fine-meshed dipnet or a 1/4-inch
mesh seine. The use of fine-meshed wire traps
with inverted cones baited with meat (chicken
and fish scraps, liver, etc.) is particularly useful
when the area is heavily choked with vegetation
or if the water is deep ( >1.5m).
Water Samples:
All water analyses were made in the field. A
Yellow Springs Instrument Company (YSI)
Oxygen Temperature Meter (Model 54), a YSI
Salinity Conductivity Temperature Model 33
Meter, and a Markson pH Meter (Model 85)
were used for obtaining physicochemical data.
Probes were inserted directly into the lake,
stream, or burrow water and resulting data
recorded. These are presented in the “Ecology”
section of each species treatment.
-
-
-
Preservation Procedures
Figure 12. a) Burrow chimney of Cambarus (L.)diogenes
on French Island, La Crosse County; b) burrow chimney
of C. (L.)diogenes along shore of Phantom Lake Flowage
in Burnett County.
Individual crayfishes and shrimps should be
killed in 5 6 % neutral formalin and remain in
the solution from 1 2 hours to a week, depending
on the size. Specimens should then be washed
in running tap water for several hours and
transferred to 70% ethanol (ethyl alcohol). If
epizooites are to be saved the formalin in which
the decapods were killed should be filtered and
the exoskeleton should be rinsed and the rinse
water poured through a fine sieve; the symbionts
(see Sprague and Couch 1971 and Hart and Hart
1974) should be preserved in 70 75% ethanol
and carefully labeled.
Crayfishes should be stored preferably in
clamp-top glass jars with gaskets. The organism
should be placed in the jar anterior end down
and with ample room for preservative. A label,
using insoluble ink, should always be placed inside
the jar.
Identification Procedures
-
Cambaridae:
Many adult crayfishes, including the Wisconsin species, can be identified simply by gross
examination and/or with the aid of a hand lens.
Many smaller individuals cannot be observed
9
adequately without the aid of forceps, needles,
and a stereoscopic microscope. Waywell and
Corey (1970, 1972) suggest that pteridines
(pigments) in conjunction with other taxonomic
criteria (e.g., morphology), can be used as species
characteristics in crayfish systematic studies.
Of the several morphological characters used
in the identification of crayfishes in Wisconsin
and other regions, the first pleopod of the male
(Fig. 14) and the annulus ventralis (seminal
receptacle) of the female are the most reliable
(e.g., Fig. 23a,d,f). Although intraspecific
variations in these features occur, they are
minimal and the structures are apparently little
affected by adaptation to the environment
(Hobbs 1942a: 25).
The Cambarinae exhibit a cyclic dimorphism
associated with the reproductive cycle (Hobbs
197210). I n the more northern species this occurs
generally as an annual cycle. At the end of their
first season, the breeding (“Form I” or “first
form”) males of the population molt and are
transformed essentially into a juvenile morphology (“Form 11” or ‘(second form”). This form
is retained until the beginning of the next
breeding season when the second semiannual
molt returns them to the adult form (Form I).
Indeed Form I males can molt into Form 11, both
in the laboratory and in natural systems, yet this
“cycle)’ is not always predictable. Taylor (1985)
points out that for Procambarus (Pennides)
spiculifer (LeConte), Form I-Form I1 alternation
does not occur. His data suggest that Form I
males either remain Form I or die but that most
Form I1 males arise from juveniles and subsequently molt to Form I. Since individuals increase
in size with each molt, it is to be expected that
quasi-juveniles (Form 11) following their first or
second breeding seasons are larger than an adult
(Form I) male in its first. Thus, Hobbs (1972b)
points out that relative lengths alone cannot
be used in distinguishing between first and second
form males (e.g., that a n individual having a
carapace length of 20mm does not necessitate
it being a juvenile). The same author (1981:9)
indicates that the first pleopod of juvenile males
has a characteristic oblique suture in the
proximal half of the shaft but that Form I1 males
occasionally exhibit this “juvenile suture.”
<
Figure 13. Generalized diagram of palaemonid shrimp (ai = appendix interna, a m = appendix masculina, as = antennal
spine, b = basis or basiopodite, bg = branchiostegal groove, bs = branchiostegal spine, c = coxa or coxopodite, cp =
carpus or carpopodite, d = dactyl or dactylopodite, end = endopod or endopodite, ex = exopod or exopodite, i = ischium
or ischiopodite, m = merus or meropodite, p = propodus or propodite, and sc = scaphocerite or antennal scale).
10
men of rostrum
Hepatic spines
Cervical spine
Outer ramus
Figure 14. Generalized diagram of cambarid crayfish: a, ventral view; b, dorsal view (after Hobbs et al. 1977).
Although the dichotomous key (see below) is
appricable to both adult males and females, often
Form I males are required for positive species
identification. First form males may be distinguished from juvenile and second form males
by the presence of one or more corneous yellow
to brown (sclerotized) terminal elements on the
distal ends of the first pleopods. First pleopods
of juvenile and Form I1 males have much less
well defined terminal elements that are never
sclerotized. Secondary sexual characters of Form
I males include well developed hooks on the
ischiopodites of one or more of the third and
fourth pereiopods (Fig. 15) (in Wisconsin
crayfishes) and often bosses from the bases of
the coxopodites of the fourth and/or fifth
pereiopods (Figs. 30h, 69h).
In Wisconsin both members of the genus
Procambarus possess subterminal setae on or near
the distal end of the first pleopod of Form I
males which partly or completely obscure the
terminal elements. These are particularly
characteristic of P. u. ucutus and it is recommended that the setae be carefully removed in
order t o view these elements clearly. To
accomplish this it is customary that the crayfish’s
left pleopod be removed and placed in a dish
of 70% ethanol under a stereoscopic microscope.
The pleopod should then be held at its base with
fine forceps and the setae removed with a small
needle. Take care not to injure or break one
of the terminal elements. Both lateral and mesial
views can then be compared with illustrations
included in the text. Identifications of crayfishes
11
of other genera may be facilitated also by removal
of the left pleopod for microscopic examination.
References and illustrations of the major
characters used in identification are referred to
throughout the text (see also Figs. 14, 63). For
descriptive purposes, t h e first pleopod is
considered to hang pendant from the abdomen.
The attached end is considered proximal; the
opposite end, distal; the side toward the anterior,
cephalic; that toward the telson, caudal; the
surface facing the “middle” of the body and
corresponding paired pleopod, mesial; and that
facing the side of the body, lateral. When the
appendage is in normal position against the
sternum of the cephalothoracic region, the side
in contact with the thorax is the cephalic surface
and the side viewed in ventral aspect (from
below) is the caudal surface.
Hobbs (1981:ll) presented a brief explanation
of the descriptive terminology applied to the
annulus ventralis. He stated that the annulus
ventralis of more primitive species (members of
the genus Procumbarus) is surrounded by a
flexible cuticle, rendering it “freely movable.’’
The most advanced species (members of the
genera Orconectes and Cumburus) have the
cuticle joining the annulus to the sternum
Figure 15. Basal podomeres of peteiopods of male crayfishes
(after Hobbs I11 et al. 1976).
w
12
immediately anterior to it sclerotized so that little
if any motion is possible between them. In some
members of Orconectes, no motion between the
fused elements is possible, and the annulus is
described as being “inflexibly fused with the
sternum.” In those species belonging to the genus
Cambarns, the annulus itself is not uniformly
sclerotized. Even though the annulus is fused
with the sternum, a slight hingelike action occurs
between the anterior and the more heavily
calcified posterior part. Annuli such as these are
“capable of a slight hingelike motion.”
Palaemonidae:
Reliable identification of freshwater shrimps
can be made only if appendages are removed
and examined with the aid of a microscope. A
single species of shrimp is known from Wisconsin;
nevertheless, we advise removing a second
pleopod of adult males and mounting it on a
glass slide (water or glycerin) for examination.
Adult males are recognized by possessing a welldeveloped appendix masculina on the second
pleopod (Fig. 13). It is critical that the distal
portion of the appendix masculina be viewed
and that the apical and subapical setae be
counted; note the location of setae on the
appendix and their relative positions on the
endopod (Fig. 73j ,k). For additional discussion
of pleopod modification and sexual dimorphism
in the genus Palaemonetes, see Fleming (1969).
In order to observe structures clearly, all
appropriate appendages of shrimp should be
cleared. The easiest method is to place the entire
organism in a lactic acid-chlorozol Black E stain
solution (3.5 drops of 1% ethanol 95% stain
solution per 20ml of lactic acid) or a 1%solution
of fast green and heated to 150” C for 15 minutes;
larger specimens may require longer periods. This
should remove the soft parts and result in the
shrimp being translucent purple or green. It
should then be washed and transferred to
glycerine. Separate appendages can be dissected
away from the animal and a semi-permanent slide
can be made by mounting the appendage in
glycerine and “ringing” the cover slip with a
sealing compound (Hobbs 111 et al. 1976).
PHYSIOGRAPHY OF WISCONSIN
~
~
:
,
i
I
I
Wisconsin lies within two physiographic
provinces: the Superior Upland, a southerly
extension of the Canadian Shield, and the
Central Lowland (Fig. 16). Because of Wisconsin’s complex geologic history, provinces are
commonly divided into four or five physical
provinces (Fig. 17). In this study we have
employed: 1) the Northern Highland, the largest,
a part of the Superior Upland Physiographic
Province; 2) the Western Upland, 3) the Central
Plain, and 4) the Eastern Ridges and Lowlands
parts of The Central Lowland Physiographic
Province (see Martin 1932, Fenneman 1938,
Black 1964, Hunt 1967, and Paull and Paull
1977).
The Northern Highland Province is underlain
by generally massive igneous and metamorphic
Precambrian rock covered with a veneer of
glacial deposits. Some of the lowland areas along
Lake Superior are underlain by sandstone that
may be Lower to Middle Cambrian. The highest
points in this region are generally resistant
bedrock hills on the upland surface; the highest
point in the state, Tim’s Hill in Price County,
is capped with glacial deposits and is 596 m above
mean sea level. The Northern Highland is the
headwaters region for all the major river systems,
which drain into Lake Superior or Lake Michigan
(the St. Lawrence drainage) or the Mississippi
River. From a center near Land O’Lakes in Vilas
County, rivers flow to the southwest, south,
southeast, and north. Two areas of this physical
province are densely covered with glacial lakes:
the northwestern lake district extends northeasterly from Polk County to Bayfield County
in northwestern Wisconsin at the headwaters
of the St. Croix and Chippewa rivers; Highland
Lake District is centered in Vilas and Oneida
counties in the extreme northern part of the
state at the headwaters of the Flambeau branch
of the Chippewa and the Wisconsin, Wolf, and
Menominee rivers. High lake density in this
province results in “. . . 40% of the area being
either water or swamp” (Paull and Paull 1977:80).
More than 800 of Wisconsin’s 14,000+lakes are
located in Oneida County and literally hundreds
of bogs and marshes provide evidence of primary
succession occurring in other former lake basins.
These lakes are generally small, closely spaced,
irregular in outline, and connected by streams.
Most are glacial; however, the origins of the
basins are diverse. Few, if any, are in glacially
excavated rock basins as the bedrock is deeply
buried beneath glacial drift. Some are kettles,
some are in shallow depressions in ground
moraine, some are held by recessional moraines,
and many are in depressions in the outwash
gravel plains (Martin 1932). The surface deposits
are generally infertile, sandy, pitted glacial
outwash, or boulder and clay morainic deposits.
In summary, poor soils and inadequate drainage
are common characteristics of this province.
The remaining three provinces to the south
are underlain by nearly horizontally-bedded
Paleozoic sedimentary rocks (sandstone, dolomite, limestone, shale). The Western Upland, a
narrow band that trends northwest-southeast,
is located in the western and southwestern part
of the state. Much of this province coincides
with the aforementioned Driftless Area; therefore little Pleistocene glaciation masks the
bedrock, and this results in a region of high
topographic contrasts. In addition to the upland
itself, the most outstanding topographic features
are the deep “trenches” carved by the Wisconsin
and Mississippi rivers and their many tributaries
(e.g., the gorge of the Mississippi is incised more
than 150 m below the peaks of the upland ridges).
Solution features, such as caves and sinkholes,
are present in this region, yet are rare in other
provinces. Because of t h e well-developed
drainage, generally porous bedrock, and the
absence of young glacial deposits, very few
natural lakes, marshes, and swamps exist here.
A number of man-made lakes are found, some
having been formed by the damming of bedrock
river valleys.
In contrast, the Central Plain is identified by
moderate topography, developed largely on
Cambrian sandstone rock. It is the smallest of
the provinces and is somewhat crescent shaped,
being situated among the other three provinces
(see Fig. 17). The topography is flat or rolling
with little relief. In the unglaciated region, this
low relief resulted from nearly uniform erosion
13
I
O
W
A
Figure 16. Map of Wisconsin, illustrating physiographic provinces (Superior Upland and Central Lowland), and Driftless
Area (after Paul1 and Paul1 1977).
14
I
O
W
A
I
j
Figure 17. Map of Wisconsin, illustrating physical provinces of state (after Frye et al. 1965 a n d Paul1 and Paul1 1977).
15
of the soft, almost horizontally-bedded sandstone
and from deposition (filling irregularities) in
Glacial Lake Wisconsin. The glaciated region
exhibits a rolling cover of ground moraine, sandy
outwash, and clay-rich lake deposits. The most
obvious landforms of the Central Plain are the
steep-sided sandstone mesas and buttes (“cas.
tellated mounds”) located in the Driftless Area,
remnants of preglacial erosion.
Within the Central Plain the Wisconsin River
is the largest of a series of streams coursing across
the sandstone lowland and is characterized by
a gentle grade and few tributaries. T o the west,
the Black and Chippewa rivers flow from the
Northern Highland to the Mississippi River.
These streams carve through the Cambrian
sandstone and expose Precambrian rocks within
the province. Even though the Fox-Wolf river
system and the Oconto River pass across the
eastern part of the Central Plain Province, much
of this area is poorly drained, thus giving rise
to another lake district.
The Eastern Ridges and Lowlands Province
encompasses a narrow strip of land, underlain
by Paleozoic rocks, located on the eastern border
of Wisconsin along a north-south axis. It is a
region of uplands and lowlands: the uplands
consist of resistant Lower Ordovician Prairie du
Chien and Silurian dolomites while the Upper
Cambrian sandstones, the Middle and Upper
Ordovician sandstones, dolomites, and shales and
Devonian formations permeate the lowlands.
The lowest part of the state is located in the
province along Lake Michigan at Milwaukee
(176m). This province, with its varied landscape,
demonstrates well the results of Pleistocene
continental glaciation (see Martin 1932 and Paull
and Paull 1977 for additional discussion).
The drainages in the Eastern Ridges and
Lowlands Province are postglacial and are
controlled mainly by late Wisconsinan glacial
erosion and deposition; their significance as
dispersal routes for decapods and other organisms
is quite apparent (see below for further discussion
of drainage systems in Wisconsin).
16
GEOLOGY OF WISCONSIN
Bedrock Geology:
Wisconsin is generally a rolling plain with low
relief, having maximum elevation differences of
only approximately 425 meters. It is underlain
entirely by Precambrian igneous (e.g., granites)
and metamorphic (e.g., slate, schist, gneiss) rocks
that surface only in the northern part of the
state. These Precambrian rocks have undergone
complex processing, being originally flat-lying,
subsequently upthrust, and folded, and experienced periods of subsidence; erosion sheared off
the folds, resulting in anticlines, synclines, or
faulted portions of either. These older rocks are
covered to the south, east, and west by flatlying, younger, Paleozoic sedimentary rocks that
have been warped or arched into a broad
anticlinal fold; the axis of this fold trends
generally north-south. Because the axis pitches
southward, the sedimentary rock layers in the
central part of the state also slope (dip)
southward. Those near the Mississippi River and
near Lake Michigan tilt at a low angle to the
southwest and to the southeast, respectively.
These Paleozoic bedrock layers consist of
Cambrian sandstones, Ordovician dolomites,
sandstones, and shales, Silurian dolomites, and
Devonian dolomites and shales. The Devonian
rocks are the youngest in Wisconsin and appear
only in the southeastern part of the state. West
of the Devonian rocks, the surface becomes
Silurian dolomite which extends westward on
a gently rising slope and disappears at the Niagara
escarpment which faces west, thus forming the
lowland and cuesta pattern of the Eastern Ridges
and Lowlands Province. West of this escarpment
and slightly lower t h e surface rocks are
Ordovician dolomites; the Ordovician shales and
sandstones are restricted mostly to the bases of
escarpments or in deeply incised valleys; farther
west the Upper Cambrian sandstones are
exposed.
From this brief account, it becomes evident
that the major part of southern and eastern
Wisconsin is covered with limestones, central
Wisconsin by sandstones, and the northern part
of the state by Precambrian igneous rocks. The
acid character of the Precambrian granitic rocks
and the Cambrian sandstones has considerable
influence on the chemical composition of the
groundwater. This is demonstrated in the
podzolic soils and in the poorly buffered,
predominately soft waters of the central and
northern lakes. In distinct contrast, the waters
of lakes and streams in those areas underlain
by carbonate rocks are hard and generally well
buffered (Table I); the reader is referred to Poff
(1970), Gorham et al. (1983), and Lillie and
Mason (1983) for further data on the chemical
composition of Wisconsin lake waters.
The Paleozoic rocks previously mentioned
accumulated as sediments beneath a shallow
Paleozoic sea. Following their deposition,
Wisconsin’s geological history remained relatively stable (minor uplift, subsidence, and
erosion) for a period of more than 200 million
years (during the Mesozoic and most of the
Cenozoic eras).
Glacial History:
During the Pleistocene Epoch of the Quarternary Period, which began approximately two
million years B.P., the hiatus of significant
geological activity ended and Wisconsin, as well
as many other parts of the world, was invaded
at least four times by major continental ice
sheets: the Nebraskan, Kansan, Illinoian, and
Wisconsinan glaciers. These masses of ice
accumulated in Canada and spread southward,
covering much of the northern regions of the
United States. During this period (up to 10,00012,000 years ago) the area can best be characterized as having various periods of growth,
advance, stagnation, and retreat of the ice sheets.
These dynamic processes greatly affected the
topography of the regions they covered by
downwarping, scouring, and depositing deep
layers of glacial drift. Much of the overlying
Paleozoic rocks and considerable Precambrian
bedrock were eroded by glacial advances and
retreats. Glacial drift and the glacial ice at various
times dammed previous drainage routes and
created lakes (no longer extant) such as Glacial
Lake Wisconsin (dammed Wisconsin River) in
the central part of the state (parts of Adams,
Juneau, Monroe, Sauk, and Wood counties) and
Glacial Lake Oshkosh (dammed Fox-Wolf River
system) in Winnebago County. The Great Lakes
themselves have complex and distinct histories
but share their origins which resulted from the
effects of glaciers during the Pleistocene Epoch.
The Driftless Area (first described by Irving,
1878), a region of 33,670 km2 (13,000 mi2) in
the southwestern part of the state (Fig. 16) and
a small corner of northeastern Iowa, a thin strip
in southeastern Minnesota, and part of northwest Illinois, is considered to have “escaped”
much of the effects of glaciation during the
Pleistocene.
This region is surrounded by glaciated areas,
yet it was not an “island” perched high above
the continental ice. It occupies parts of three
of the four physical provinces: Western Upland,
Central Plain, and Northern Highland. Its
present topography resulted chiefly from
weathering: wind work, subsurface water, and
stream erosion. In other areas of the state, glacial
erosion and deposition, wave work, postglacial
stream erosion, and other processes have altered
greatly the earlier topography which was
influenced by weathering and preglacial stream
work. Here, the general topography, limited
glacial deposits, and presence of residual soil over
much of the Driftless Area gives credence to
the theory that little glacial activity took place
and that the area demonstrates a longer
weathering interval than its environs. The entire
area was once covered by Niagara (Silurian)
dolomites but erosional processes have removed
all but a few isolated outliers (e.g., Blue Mounds).
Some erratic fragments of Precambrian and
Paleozoic rock are known from the area,
suggesting a limited Wisconsinan ice cover;
however, currently there is a lack of total
agreement as to the Pleistocene history of this
region. Additional information concerning the
geology of Wisconsin is presented in Martin 1916,
1932, 1982, Fenneman 1938, Leighton 1957,
Wright 1957, Curtis 1959, Black 1964, Hunt
1967, and Paull and Paull 1977.
17
nr
Figure 18. Major drainage systems in Wisconsin.
18
I
G
SURFACE DRAINAGE IN WISCONSIN
LIST OF WISCONSIN DECAPODS
The dynamics of the events of the Pleistocene
are certainly complex and obviously glacial
activities obliterated any evidence of the Tertiary
distribution of crayfishes and shrimps in
Wisconsin. As the “last” continental ice sheet
retreated, surface drainage patterns began to
become established and gradually both aquatic
and terrestrial ecosystems developed. The
complex aquatic community assemblages observed today are thus geologically young and
represent immigration of biota into relatively
recently established drainage systems. Regression
of the Pleistocene glaciers allowed organisms to
move into the ecological vacuum created by the
retreating ice sheet. To understand present
distributions of crayfishes and shrimps it is
necessary to examine the drainage patterns (the
dispersal routes for these decapods) and
physicochemical characteristics of lotic and
lentic environments in the state.
Wisconsin is divided into two major surfacewater drainage basins: the St. Lawrence and
Mississippi, both resulting from post-glacial
events. Waters flow north and east into the
Atlantic Ocean by way of Lake Superior or Lake
Michigan (the St. Lawrence watershed); a large
number of streams flow south into the Gulf of
Mexico via the Mississippi River (Fig. 18). The
drainage area of t h e Mississippi basin is
considerably larger than that of the St. Lawrence,
100,707 km2 (38,883 mi2) and 43,882 km2
(16,943 mi2), respectively. Table 1 presents
chemical data for various portions of each
watershed; refer to Lillie and Mason (1983) for
additional limnological characteristics of Wisconsin lakes.
Hobbs (1974a) presented a composite summary of the families and genera of crayfishes
of the world and recognized two families
(Astacidae and Cambaridae) that constitute the
holarctic forms and a single family, Parastacidae,
whose members are confined to the Southern
Hemisphere. All the known (and expected)
species of Wisconsin crayfishes belong to the
family Cambaridae, Hobbs 194213, and the single
shrimp is assigned to the family Palaemonidae,
Rafinesque 1815.
A. CRAYFISHES
1. Cambarus
Cambarus (Lacunicambarus)
diogenes Girard
2. Fallicambarus
Fallicambarus (Creaserim)
fodiens (Cottle)
3. Orconectes
Orconectes (Gremicambarus)
immunis (Hagen)
Orconectes (Crockerinus) propinquus (Girard)
Orconectes (Procericambarus)
rusticus (Girard)
Orconectes (Gremicambarus)
virilis (Hagen)
4. Procambarus Procambarus (Girardiella)
gracilis (Bundy)
Procambarus (Ortmanni
cus) acutus acutus (Girard)
-
’
-
-
B. SHRIMP
1. Palaemonetes kadiakensis (Rathbun)
19
TABLE 1: Physicochemical Data for Selected Localities in the Major Drainage Systems of
Wisconsin during 1981 (data from Anonymous 1982b).
I. MISSISSIPPI RIVER DRAINAGE
Temp. (“C)
(mg/l)
0 2 Sat. (%)
PH
Specific
Conductance
(pmhos/cm)
Hardness
(mg/l)
Alkalinity
0 2
MISSISSIPPI RIVER
PROPER
(Range)
0.5 - 28.0
3.4 14.3
7.4 - 8.5
310 - 625
CHIPPEWA RIVER WISCONSIN RIVER
(Range)
0.0 27.0
7.3 12.7
81 - 113
6.4 7.9
81 * 319
-
26 - 57
(Range)
0 25.0
7.1 12.9
75 105
7.3 8.5
165 310
-
-
54 - 90
35 77
76 120
TREMPEALEAUBLACK
(Range)
0 26.5
9.1 11.6
81 - 117
6.0 7.7
80 170
PECATONICASUGAR
(Range)
0 23.0
ROCK-FOX
ST. CROlX PROPER
(Range)
0 21.0
9.2 11.9
62 87
7.3 7.9
130 - 300
(mdl)
Temp. (“C)
0 2 (mg/l)
0 2 Sat. (%)
PH
Sp. Cond.
(pmhodcm)
Hardness
(mg/l)
Alkalinity
(mg/l)
30 - 63
24 - 50
-
(Range)
1.0 25.5
310 670
-
580 1100
CLAM RIVER
(Range)
4.5 22.0
8.1 JJ.4
92 96
6.9 7.3
144 - 178
-
APPLE RIVER
(Range)
1.5 19.5
8.1 - 12.2
91 103
7.3 - 7.7
230 270
-
110 - 120
-
-
ST. CROIX BASIN
Temp. (“C)
(mgll)
0 2 Sat. (%)
PH
Sp. Cond.
(pmhos/cm)
Hardness
(mdU
Alkalinity
(mgN
0 2
Temp. (“C)
0 2 (Mg/l)
0 2 Sat. (96)
PH
Sp. Cond.
(pnhos/cm)
Hardness
(mg/l)
Alkalinity
(mg/l)
20
-
62 92
NAMEKAGON RIVER
(Range)
2.0 - 16.5
8.2 12.9
87 - 101
6.8 7.8
120 - 152
-
YELLOW RIVER
(Range)
6.0 23.0
8.3 12.8
100 108
7.1 7.9
145 - 170
-
54 69
46 - 70
68 - 73
110 - 130
11. ST.LAWRENCE RIVER DRAINAGE
Temp. ("C)
(mg/l)
0 2 Sat. (%)
PH
Sp. Cond.
(pmhos/cm)
0 2
Hardness
(mg/l)
Alkalinity
GREEN BAY
(Range)
0 - 20.0
6.9 13.2
75 95
6.9 8.2
< 50 240
-
LAKE MICHIGAN
(Range)
0 21.5
6.4 15.0
75 136
6.7 8.6
500 990
25 - 110
240 - 450
20 - 110
180 - 420
BAD RIVER
(Range)
0 - 28.0
6.6 - 13.0
79 102
6.2 - 7.6
46 - 261
NEMADJI RIVER
(Range)
0 .26.0
7.2 - 12.4
74 102
6.9 7.2
155 340
-
(mg/l)
LAKE SUPERIOR BASIN
Temp. ("C)
0 2 (mg/l)
0 2 Sat. (%)
PH
Sp. Cond.
(pmhos/cm)
Hardness
(mdl)
Alkalinity
-
31 - 92
22 - 89
70 - 150
62 - 140
(md)
21
Key to the Decapods of Wisconsin
1
Abdomen and rostrum compressed laterally, latter with multiple dorsal
and ventral teeth (Fig, 13); third pair of pereiopods never chelate
(Fig. 13): PALAEMONIDAE
.Palaemonetes kadiakensis Rathbun, 1902 (Fig. 73)
Abdomen and rostrum compressed dorsoventrally, latter lacking dorsal
and ventral teeth (excluding distal tip of acumen) (Fig. 23); third
pair of pereiopods bearing chelae (Fig, 14) , , CAMBARIDAE , , 2
Male (both Form I and Form 11).
.3
Female
10
Areola broad or narrow but always with room for at least two
punctations in narrowest part (Figs. 32,36) . . . . . . . . . . . . 4
Areola obliterated along part of length or so reduced in width as
to accomodate only one punctation in narrowest part (Figs. 23,63) .8
First pleopod terminating distally in two elements (Fig.
32)
.Orconectes.. . 5
First pleopod terminating distally in more than two elements (Figs.
69a,e)
Procambarus (Ortmannicus)
acutus acutus (Girard, 1852) (Fig. 69)
Terminal elements of first pleopod with apices directed caudally at
angle of 90 degrees to principal axis of appendage (Fig, 32); dactyl
of first chela with distinct excision on opposable margin (Fig.
32m)
Orconectes (@micambarms) immunis
(Hagen, 1870) (Fig. 32)
Terminal elements of first pleopod straight or bent caudally but apices
directed at angle distinctly less than 90 degrees t o principal axis of
appendage (Fig. 46); dactyl of first chela lacking excision on opposable
.6
margin (Fig. 46m)
Cephalic surface of first pleopod with shoulder at base of central
projection (Fig. 46a); distal incisor region of mandible blade-like (Fig.
46k); lateral margins of rostrum distinctly concave (Fig.
.............
1'
..................................
..
..
.........................
............................................
3'
4(3)
4'
..
....................................
.........................
..................
5'
....................................
463)
6'
....................
Orcmctes (Procericumburus) rusticus
(Girard, 1852) (Fig. 46)
Cephalic surface of first pleopod lacking shoulder at base of central
projection (Fig. 36a); distal incisor region of mandible irregularly
dentate-crenate (Fig. 36h); lateral margins of rostrum subparallel or
convergent (Fig. 36j)
.7
Central projection of first pleopod straight, comprising less than
113 total length of appendage (not necessarily true in Form 11) (Fig.
36a,g); rostrum usually with median carina (Fig. 36j); third maxilliped
Orconectes (Crockerinus) propinquus
sparsely setiferous
(Girard, 1852) (Fig. 36)
Central projection of first pleopod curved (weakly so in second form
male) comprising more than 113 total length of appendage (not
necessarily true in Form 11) (Fig. 52a); rostrum without median carina
(Fig. 52k); third maxilliped densely setiferous
Orconectes
(@micambarus) oririlis (Hagen, 1870) (Fig. 52)
..................................
..........
7'
..........
22
Opposable margin of dactyl of chela with deep excision in proximal
1/2of opposable margin (Fig. 30n) . . . . . .Fallicambarus (Creasetius)
fodiens (Cottle, 1863) (Fig. 30)
8‘
Opposable margin of dactyl of chela without deep excision in proximal
1/2 of opposable margin (Fig. 23m) ............................
.9
9(8’) Areola obliterated (Fig. 23j); first pleopod terminating in two elements
bent at approximately 90 degrees to principal axis of shaft of appendage
(Fig. 23a,f). .........................
Camburus (Lucunicambarus)
diogenes Girard, 1852 (Fig. 23)
9’
Areola narrow but accomodating one punctation in narrowest part
(Fig. 63k); first pleopod terminating in more than two elements (Fig.
63d) ........................... Procambarus (Sirardiella) gracilis
(Bundy, 1876) (Fig. 63)
lO(2’) Areola broad to narrow but always with room for at least two
punctations in narrowest part (Fig. 36j) . . . . . . . . . . . . . . 13
10’
Areola obliterated along part of length or so reduced in width as
to accomodate only one punctation in narrowest part (Fig. 30j) ...I I
1I( 10’) Opposable margin of dactyl of chela with distinct excision in proximal
half of opposable margin (Fig. 30n); rostrum spatulate, nearly as wide
as long (Fig. 30j); cervical groove discontinuous laterally (Fig.
30c) ..........................
Fullicumburus (Creaserius) fodiens
(Cottle, 1863) (Fig. 30)
Opposable margin of dactyl of chela lacking deep excision (Fig. 23);
11’
rostrum spatulate but clearly longer than wide (Fig. 63k); cervical
groove continuous laterally (Fig. 63c)
.12
12(11’) Annulus ventralis freely movable, cephalic portion bearing conical
tubercles (Fig. 631); postannular sclerite considerably less than twice
as broad as long (Fig. 631); areola narrow (Fig. 63k)
Procamburus
(Cjirardielkx) gracilis (Bundy,1876) (Fig. 63)
Annulus ventralis not freely movable; cephalic portion without
12’
tubercles (Fig. 23d); postannular sclerite almost twice as broad as
long (Fig. 23d); areola obliterated (Fig. 23j) ........... .Cumburus
(Lucunicamburus) diogenes Girard, 1852 (Fig. 23)
13(10) Length of palm subequal to length of dactyl (Fig. 69j); usually distinct
tubercle on opposable margin of distal third of immovable finger
(propodus) (Fig. 69j); annulus ventralis freely movable ...Procumbarus
(Ortmnnicus) acutus acutus (Girard,1852) (Fig. 69)
Length of palm 1/3 to 1/2 that of dactyl (Fig. 32m); tubercle lacking
13’
on opposable margin of distal third of propodus (Fig. 36m); annulus
.14
ventralis not freely movable.
14(13’) Dactyl of first chela excised near base on opposable margin (Fig.
32m); tuft of setae at junction of dactyl and propodus; annulus ventralis
asymmetrical with deep fossa extending beneath high dextral wall
(Fig, 32h)
Orconectes (@micamburus) immunis
(Hagen, 1870) (Fig. 32)
Dactyl of first chela lacking excision on opposable margin and without
14’
setae at base (Fig. 36m); annulus ventralis symmetrical with fossa
centrally located (Fig. 36n) ............................
.I5
8(3’)
.....................
....
...........................
...............
23
15(14’) Annulus ventralis not strongly sculptured and with two low cephalic
tubercles never fused at midline (Fig. 36n); rostrum usually with
median carina (Fig. 36j) . , . Orconectes (Crockerinus) propinquus(Girard, 1852) (Fig. 36)
15‘
Annulus ventralis strongly sculptured and with two cephalic tubercles
fused (or nearly so) at midline (Fig. 46d); rostrum without median
carina (Fig. 46j).
.16
16(15’) Distal incisor region of mandible blade-like (Fig. 46k); fossa of annulus
ventralis moderately small and extending directly beneath anterior
fused cephalic lobes (Fig. 46d)
Orconectes
(Procericumbuw) rusticus (Girard,1852) (Fig. 46)
Incisor region of mandible irregularly dentate-crenate (Fig. 520;
16’
annulus ventralis with very large and deep centrally situated fossa
(Fig. 521)
.Orconectes (Q-emicumburus) virilis (Hagen, 1870)
(Fig. 52)
....................................
.....................
......
In addition to using a key (gross morphology)
to separate individuals of different species, one
can also employ various measuremendratios.
Total gonopod length/postorbital carapace
length ratios as a function of postorbital carapace
length are plotted in Figs. 19 and 20. Ratios
of individuals of representative populations of
0. propinquus, 0. rusticus, 0. virilis, and C.
diogenes are shown in Fig. 19 and even though
there is some overlap between species, this ratio
can be used reliably to separate most individuals
of these species. Ratios of individuals of 0.
immunis, P. gracilis, and P. a. acutus are shown
in Fig. 20. Ratios of 0. immunis overlap those
of 0. propinquus and thus are not plotted on
the same graph; however, the distinct color
pattern of 0. immunis and the conside’rably
different morphologies of the gonopods and
annuli of both species make recognition of each
rather simple.
Overlap of ratios of P. gracilis
and P. a. acutus is pronounced (Fig. 20), yet these
ratios can be used in concert with examination
24
of gonopods and annuli, Of interest, populations
of 0. immunis, 0. propinquus, 0. rusticus, and
0. virilis show a significant (p 0.05) negative
correlation between total gonopod length/
postorbital carapace length ratio and postorbital
carapace length; that is, t h e gonopod is
proportionately shorter in larger individuals. This
trend was also noted by Capelli and Capelli
(1980:125) for 0. propinquus, 0. rusticus, and
0. virilis in Wisconsin populations.
Areola width/carapace length ratios can be
used to separate individuals of 0. propinquus,
0. rusticus, and 0. virilis (Fig. 21). Minimal
overlap occurs among the species and this should
pose no problems in separating these crayfishes.
This ratio for 0. immunis overlaps the values
for 0.rusticus and 0.virilis and thus examination
of other morphological characters and color
patterns is required to distinguish the species.
The same patterns are demonstrated when areola
width/areola length ratios are examined (Fig. 22).
<
.65
A
A
A
A
A
..
....
.. ..
%
AA
.55
O
.
A
A
0
0
p:
:A
A
AA
A
00
0
0 0
0 0
-45
f
Q
O
U
0
O
A
A t
OA
A
0
,.
00
.Q.n
a.
0
n
U
0
0
\
J
Q
I-
.35
0
.25
0
0
0
0
0
0 .
0
0
0
0
0
0
I
I
I
I
I
15
25
35
45
55
Postorbital C a r a p a c e Length (mm)
Figure 19. Total gonopod length/postorbital carapace
length ratios as a function of postorbital carapace length
for Cambarus (L.) diogenes (filled circles), 0.((2.1 propinquus
(filled squares), 0. (P.) rusticus (open circles), and 0. (G.)
virilis (filled triangles).
25
.55
0
0
0
0
.45
-I
0
0
0
n
0
\
08
A
0
0
4
(3
I-
.35
A
A
A
A
00
0 O A 0
'Ao
A
0 0
0 0
0
0
0
A
0
V
0
'
1
1
1
25
35
45
P o s t o r b i t a l C a r a p a c e L e n g t h (mm)
Figure 20. Total gonopod length/postorbital carapace
length ratios as a function of postorbital carapace length
a. acutus (open
for 0. (G.) immunis (filled circles), P. (0.)
circles), and P. (G.)grucilis (filled triangles).
26
0
025
.050
-075
.loo
Areola Width/Carapace Length
Figure 21. Frequency plot of areola width/carapace length for O c m c t e s (C.) popinquus (solid), 0. (P.) rwticus (stippled),
and 0.(G.) virilis (horizontal stripe).
N
41
I,JI
.120
20
15
a
U
E
LL
10
5
0
.05
.10
.15
.20
.25
.30
.35
-40
Areola WidthIAreola Length
Figure 22. Frequency plot of areola widthlareola length for Orconectes ((2,)propinquw (oblique stripe), 0. (P.) rwticus
virilis (horizontal stripe).
(stippled), and 0.(G.)
28
SPECIES TREATMENT
CAMBARIDAE
Cambarus (Lacunicambarus) diogenes Girard
(Figures 11,12,19,23 - 29,5759)
Cumburus diogenes Girard 1852:99.
Cumburus obesus
Bundy 1882:178,180,183;
1883:403; Creaser 1932:335.
Cumburus diogenes Faxon 1884:144, 1885b:71,
75; Underwood 1886:368; Harris 1903a59,
85, 150, 155; Ortmann 1906407, 459, 462;
Cahn 1915136,174; Andrews 1915:200,207,
210, 211; Ellis 1919:261; Turner 1926:187;
Newcombe 1929b:280; Creaser 193113269;
1932:321, 324, 325, 330, 331, 335, 336;
Creaser and Ortenburger 1933:41; Walters
1939:22; Goodnight 1 9 4 0 4 0 ; Threinen
1958b:2; Marlow 1960229; Crocker and Barr
1968:128 (Fig. 84); Hinkelman 1970:7, 29,37,
39, 40, 41, 44, 45, 47, 49; Capelli 197539,
44, 48, 53, 71, 199, 200, 205, 207; Sheffy
1978:222; Capelli 1982a:74 1; Threinen
19821333; Capelli and Magnuson 1983548,
-
549, 551, 556, 558, 559, 563, 564.
-
Cumburus Diogenes Faxon 188513313.
Cumburus (Burtonius) diogenes
Ortmann
1906:407, 459; Graenicher 1913:118, 120,
121, 122, 123.
Cumbarus diogenes sspp. - Penn 1950:646.
Cumburus ( C U ~ ~ U Tdiogenes
US)
Ortmann
-
-
1931:152.
Cumburus diogenes diogenes Hobbs 1942a:166;
Williams and Leonard 1952:1010; Marlow
1960:230, 232, 234; Boronow 1980:l.
Cumburus (Lacunicumburus) diogenes - Hobbs
197210 (1976a):146, 156.
DIAGNOSIS (refer to Table 2): Carapace (Fig.
23cj) laterally compressed, vaulted dorsally and
without cervical, branchiostegal, or hepatic
spines, more represented by prominent tubercles;
post orbital ridge short , terminating anteriorly
without spine. Rostrum lacking marginal spines
or tubercles and without median carina; upper
surface concave with somewhat thickened
margins; acumen very short, triangular, and tip
slightly upturned. Cervical groove moderately
deep and sinuous. Suborbital angle absent. Areola
(Fig. 23j) obliterated or linear, constituting 36.6
to 43.5% (mean 40.1%) of total carapace length
(42.8 to 49.3%, mean 46.2%, of postorbital
carapace length) with no room for punctations
in narrowest part (see Fig. l i in Hobbs 197613).
Antenna1 scale (Fig. 23g) nearly 3 times as long
as wide, broadest slightly distal to midlength;
distal portion tapering, acute. Cephalic lobe of
epistome (Fig. 23i) generally rounded, slightly
longer than broad with margins convex. Distal
incisor region of mandible (Fig. 23h) irregularly
dentate-crenate. Abdomen narrower and
subequal in length to carapace. Cephalic section
of telson with 2 spines in each caudolateral
corner; proximal podomere of uropod with
caudal spine on mesial lobe, lateral lobe rounded;
mesial ramus of uropod with prominent submedian dorsal keel terminating distally in strong
distomedian spine; distolateral spine of ramus
distinct. Chela (Fig. 23m) large, approximately
2.2 times as long as broad, swollen, and punctate;
mesial surface of palm with 2 to 3 rows of
tubercles, mesialmost consisting of 5 to 7, usually
5; both fingers tuberculate, dactyl not deeply
mesially excised at base. Carpus and dactyl of
cheliped deeply furrowed dorsally.
Ischium of only third pleopod (Fig. 23k) with
simple, slightly recurved hook extending
proximally over basioischial articulation. First
pleopod of male (Fig. 23a,b,e,f) with convexity
near midlength of cephalic surface, terminating
in short, only slightly tapering central projection
and distinctly tapering mesial process, both
directed caudally at angle of about 90 degrees
to axis of appendage; mesial process extending
distally t o beyond central projection; caudal knob
absent.
Annulus ventralis of female (Fig. 23d)
subquadrangular and deeply embedded in
sternum; cephalic region low with shallow
submedian trough, its fossa situated at midlength
extending beneath inflated dextral wall; caudal
region high, with dextral and sinistral regions
converging at midline.
TYPE-LOCALITY:
Environs of Washington, D.
c.,
USA.
29
TABLE 2. Range of measurements (in mm) of various diagnostic structures for Wisconsin
C. diogenes (N = number of specimens measured).
N
Minimum
Maximum
Mean
Standard
Deviation
f
MALES ( 8 I)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
Pleopod
Length
MALES ( 8 11)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
Pleopod
Length
-
-
FEMALES
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
-
30
11
11
11
38.2
32.9
15.9
58.9
51.9
25.9
49.3
43.3
21.1
6.9
6.3
3.4
11
0
14.7
25.6
20.1
3.4
11
11
7.4
12.9
16.3
23.0
11.9
18.3
2.8
3.7
11
1.3
13.5
10.7
3.4
25
25
25
23.1
19.8
9.0
57.5
49.2
24.1
32.9
28.4
13.7
7.9
6.9
3.4
25
0
9.2
24.1
13.2
3.5
24
24
4.3
6.8
13.5
21.1
6.5
10.0
2.3
3.3
7
6.6
13.5
9.1
2.4
50
50
49
22.7
19.2
8.9
49.5
42.8
20.9
34.5
29.9
14.9
6.7
6.0
3.3
50
0
8.7
20.4
13.8
2.9
41
41
3.1
5.3
10.5
16.1
6.5
9.9
1.9
2.8
e
I
Figure 23. Cambarus (L.) diogenes (b,e, second form male from Mann Creek, Vilas County; d, from female from burrow
on Willow Glen Road, 1.6 mi (2.5 km) southwest of US. Hwy. 94, Jefferson County; h, first form male Lake Wingra,
Dane County; 1, female from Warner Creek, Grant County; all others first form male from Richland Creek, Green
County): a,b, mesial view of first pleopod; c, lateral view of carapace; d, annulus ventralis; e,f, lateral view of first pleopod;
g, antenna1 scale; h, incisor margin of right mandible; i, epistome; j , dorsal view of carapace; k, proximal podomeres
of third, fourth, and fifth pereiopods; l,m, dorsal view of distal podomeres of cheliped, cp, central projection, mp, mesial
projection.
VARIATION. Since this is one of the most widely
distributed species of North American crayfishes,
its morphological variation is considerable. In
fact, this is a “species complex” needing
considerable taxonomic revision (see Hobbs
1974b).
Among variations noted are composition of
the rows of tubercles on the mesial margin of
the palm of the chela (Fig. 23m) in which the
number of tubercles per row varies from 5-7.
Much variation is apparent in the gonopod of
Form I males (see Fig. 24a,b,c,d); the length and
shape of the mesial process is particularly
variable. The annulus ventralis (Fig. 23d) is
variable and some fossa extend beneath an
inflated sinistral, rather than dextral, wall.
Occasionally abnormalities occur in various
structures; one is depicted in Fig. 231.
Without a larger number of specimens from
a single locality it is not known whether
differences noted are individual ones or
characteristic of local populations or even larger
geographical areas.
COLORNOTES (refer to Figs. 25, 26). Ground
color of carapace olive-brown to rich reddishbrown fading ventrally on hepatic and branchi-
ostegal regions to tan or pale green. Rostra1
margins and postorbital ridges orange or scarlet.
Abdomen similarly olive-brown t o reddishbrown with caudal margin of terga red. Tan to
grayish middorsal stripe originating at anterior
region of fused margins of areola (usually faint
coloration) extending posteriorly across terga of
abdomen (where more distinct), and terminating
on sixth abdominal segment, generally not
extending onto telson; stripe considerably more
pronounced in immature individuals commonly
displaying paired brown to dark olive stripes,
narrowing posteriorly, flanking median stripe.
Antennular and antenna1 peduncles dark brown
or dark olive with dark lateral margin. Cheliped
with distal third of merus, dorsal surface of
carpus, and lateral half of palmar area of propodus
dark olive to dark brown; distal part of all three
podomeres, mesial part of palmar area of
propodus, as well as dactyl suffused with orange
or maroonish-red; this color may replace olive
or brown on dorsal parts of both fingers from
near midlength almost to end of fixed finger,
and ventrally on dactyl; tips of fingers bright
orange to red with orange to yellow, occasionally
brown, corneous extremities; distinct lateral
Figure 24. Lateral view of first pleopod of first form male of Cambarus (L.) diogenes (a, Monroe, Green County; b, Starkweather
Creek, Dane County; c, Little Waumandee Creek, Buffalo County; d, Waumandee Creek, Buffalo County).
32
Figure 25. Photograph of female Cambarm (L.)
diogenes from Wisconsin River, Grant County.
margin of propodus orange to yellow, ventral
surface tan t o yellow; all ridges, spines, and knobs
of merus, carpus, and propodus scarlet to orange.
Pereiopods 2-5 with merus through dactyl olive
t o brown dorsally fading to pale tan (occasionally
pink or light bluish-green) ventrally, and distal
margins of ischium, merus, carpus, and propodus
orange t o scarlet. Telson and uropods brown t o
olive; margins of latter and all spines scarlet.
Kent ( 1 9 0 ~ 9 3 4reported
)
an unusual (questionable?) observation concerning color in this
crayfish. “They burrow during winter and come
out in the spring with more or less of the color
of the soil. The colors are gradually turned to
red in the open sunlight.”
ECOLOGY. Habitats. Cambarus diogenes is one
of the three most widely ranging species of North
American crayfishes. Such a large distribution
reflects a wide ecological tolerance and thus the
species exhibits innovations (e.g., enlargement
of the gill chambers) that allow it t o exist in
a broad spectrum of ecological conditions (see
Hobbs 1977).
The earliest observations of this crayfish in
Wisconsin were those of Bundy (1882:183).
Concerning this species (his C. obesus) he stated
that it is one of the largest and most abundant
crayfishes, preferring stagnant water, frequenting
ponds and meadow ditches and often wandering
far from surface water, burrowing in wet fields.
Graenicher (1913) reported a number of
localities (streams, ditches, ponds) and presented
some life history data and Andrews (1915:210)
mentioned a population in the Lake Butte des
Morts Bog region, Winnebago County. In 1932,
Creaser reported C. diogenes t o occur “throughout the entire state” of Wisconsin and remarked
on the high density of burrows along stream
banks. Threinen (195810) reported this species
from burrows near streams or marshes and
indicated that individuals move to open water
in late summer or fall to breed. Hinkelman
(1970:37) noted its occurrence in Turtle Creek
and from burrows (up to six mounds per m2)
in marshy and spring-fed areas on the east side
of Comus Lake in Walworth County. No other
ecological information has been published for this
species in Wisconsin.
Field observations. Based on our field observations and on the results of studies of C. diogenes
33
Figure 26. Photograph of juvenile female Cambarus (L.)diogenes, showing prominent dorsal stripe characteristic of immature
C. diogenes (specimen from Menominee Creek, Grant County).
in other states, we can say that this is a most
efficient burrowing crayfish. It has been found
in varied habitats: burrows in flood plains, waterlogged and dry fields, marshes, wet meadows,
riverbanks (Fig. 57), ditches, and in open water
of sloughs, streams (Fig. 59), rivers, lakes, ponds,
and in stagnant sewage.
Burrows. Typically individuals occur singly or
in colonies in burrows alongside some water body.
These burrows are most efficiently constructed
in fine-grain soils (Grow 1982), each extending
into the soil for approximately 1 1.5m (15cm
to 5m depth range) and often topped with a
“chimney” or a mud cap (Fig. 12). Several factors
restrict or necessitate the depth of burrows: 1)
type of soil (i.e., fine grain particles such as clay
are better supportive materials than coarse grain
particles such as sand, which tend to collapse
readily), 2) depth to “ground water,” 3) depth
to which soil freezes during cold months. Bundy
(1882:183) dissected a burrow to a depth of
approximately four meters without locating its
occupant or reaching the bottom of the tunnel.
Of importance, Grow and Merchant (1980)
showed no significant differences in the depth
-
34
of burrows seasonally.
Commonly, two (sometimes three) entrances
converge approximately 1/3m from the surface
and t h e single, generally vertical tunnel
terminates in a flask-shaped enlargement. Each
burrow is usually occupied by a single individual
that generally leaves to forage at night. Ground
water may be found very close to the mouth
of the burrow or some distance below the surface.
In several locations along stream banks, we
observed that burrows had one opening below
the stream surface; this penetrated for 10-25cm
into the bank and intersected the main burrow,
presumably supplying the occupant with an
additional escape route, This is similar to the
observation made by Hobbs (1942a) for a small
stream environment in Early County, Georgia.
Hobbs and Hart (1959:189) noted that
burrows may branch in several directions and
have several chimneys when they occur in
seepage areas. In southern Door County in Wisconsin, a roadside seepage area was sampled by
us on 14 July 1982. Very shallow, ramifying,
complex burrows were found, and dissected; one
burrow ramified through an area of approximately
12m2. All specimens either seen or captured were
located in a vertical tunnel which occurred
usually at one end of the branched burrow
system.
Hobbs (1981:29,31) reviewed the arguments
of Tarr (1884) and Abbott (1884) and others
(e.g., Shufeldt 1896, 1897, Engle 1926, Ortmann
1906) concerning the function of the chimney
portion of the burrow of C. d. diogenes. He
presented a strong case (enlightened by the work
of Vogel, 1978) supporting Abbott’s view that
the chimneys of crayfish burrows are a functional
part of the burrow system and that the burrow
is, at least in part, designed. The importance
of the structure is ensuring air flow through the
tunnels, particularly since the oxygen concentration of water in the burrow system is often
less than 2mg/l (Hobbs 1981:31). Hobbs and Hall
(1974:196) reported that crayfishes are somewhat unique in that they have an ability to live
out of the water for extended periods of time.
As long as the gills are covered by a film of
moisture most crayfishes in aerial environments
demonstrate no apparent ill effects. If oxygen
concentrations in the water are very low the
crayfish moves to the air-water interface, where
the gills on one side of the body may be exposed
to the air. This is a particularly important strategy
utilized by burrowing species. By climbing above
the water in the tunnel the individual can expose
its gills to air with virtually 100% relative
humidity, without the potential threat of
predators.
Grow and Merchant (1980) reported oxygen
values in burrow water on Theodore Roosevelt
Island in the Potomac River to range from 0.1
to 6.1 mg/l. A 1981 summer study of a population
of C. d. diogenes from a meadow in Monroe
County, Indiana by one of us (HHH, unpublished) indicated oxygen concentrations of
burrow water ranging from 0.1 to 1.4 mg/l (mean
0.61mg/l) with a mean saturation value of 7.5%.
Crayfishes were commonly observed throughout
the day positioned at the mouths of their
burrows, well above the water. Waters in the
burrows of Wisconsin C. diogenes were found
to have oxygen values of 1.1 to 3.6 mg/l (see
also Jaspers 1969).
Crayfish burrows serve as refugia, hibernacula,
etc. for other organisms during dry periods and
during winter. It is quite probable that many
burrows play an important role in the reestablishment of populations of invertebrates
and vertebrates particularly in temporary ponds
or intermittent streams. Trautman (1981:50S)
suggested that in Ohio the ictalurid, Noturus
trautmani Taylor, “. . . normally inhabits unseinable situations such as crayfish burrows or
other holes in the bank. . . .” Creaser (1931a:244)
listed a copepod, an ostracod, and an amphipod
(over 6000 individuals from a quart of water
from a burrow) from a C. diogenes burrow in
a dried slough in Missouri. While dissecting a
crayfish burrow in Racine County 19 October
1982, we found a leopard frog (Rana pipiens
Schreber); C. diogenes was collected from other
burrows in the stream bank, thus it is probable
that this crayfish constructed the tunnel
occupied by the frog.
Physicochemical parameters. During our field
studies we observed C. diogenes burrowing
predominantly in sandy and clay soils or in slowly
to moderately flowing streams with silt and
“muck” substrates. Approximately 60% of the
burrows examined had chimneys. In the water
within burrows specific conductance ranged from
28 to 730 pmhos/cm (mean = 395), pH varied
from 5.2 to 9.0 (mean = 7.4), and oxygen concentrations ranged from 1.1 (27.3”C)to 8.8 mg/
1 (17.1”C) (mean = 3.6 mg/l).
LIFE HISTORY. Life history data for C. diogenes
are summarized in Table 3.
Form I males. Since copulation usually occurs
within the burrow environmefit during late fall
and winter, many Form I males are observed
in the population in the spring prior to the molt
to Form I1 (see Table 3 the apparent low
incidence of Form I males during winter is
undoubtedly a reflection of the lack of field work
during the cold months).
Egg production and hatching time. During early
spring, females within the burrow, each occupying a separate one, lay their eggs. Prior to egg
laying, cement glands are prominent on the
ventral portion of the abdomen, telson, and
uropods (see Fig. 27). A single female collected
-
35
11 April 1911 from Milwaukee County carried
approximately 80 eggs, each about 1.9mm in
diameter. In contrast, Hobbs (1981226) found
an ovigerous female carrying 40 eggs (diameter
about 3mm) in September in Georgia, During
the period of March to May females with eggs
or young leave their burrows for open water;
once young are released in lakes, streams, or
ponds, females return to the burrows.
Young of the year. Young-of-year (YOY) are
observed in both the open water and in the
burrow water from March until June. Engle
(1926) noted that in Nebraska during late August
YOY were found only in open water, not in
burrows. In most instances, the occurrence of
YOY in either open water or burrows is a function
of the distance between the burrow and an open
water habitat. In West Virginia, Newcombe
(192913) noted that by midsummer, juveniles
were beginning to construct their own burrows.
Ortmann (1906) indicated that in Pennsylvania
by the end of July young are approximately 20mm
in length and are living in individual burrows
away from the mother. Phillips (1980) collected
a female with attached eggs on 22 April in Iowa.
In the laboratory they hatched by 16 May, the
young remaining with the female for approximately three weeks. By 8 June all had left the
female; on 11 June she molted. Ortmann
(1906:481) reported that by June, young attain
a length of 30 t o 40mm.
Fom II males. Males spend the summer as
Form 11, molt to the sexually active Form I in
the fall, and copulate, generally in burrows,
TABLE 3. Life history data for C. diogenes (* from references only, ** from literature and
from our collections, or USNM or OSU collections; those without asterisk(s) from USNM
collections only). The following abbreviations are used in one or more of the life history
tables provided for all decapod species discussed in this book: $I = Form 1 male; $11 = Form
I1 male; 9 e = ovigerous female; ~y = female with young attached to abdomen; yoy = youngof-year; copul = copulating pair observed; open water = individuals captured away from burrow
in open water; 9sp = female with sperm plug.
Month
$1
$11
9e
9y
'Wis.
'Wis.
yoy
copul
open
water
Jan.
Feb.
March
April
'Wis.
May
Wis.
Wis.
June
Wis.
Wis.
Wis.
July
Wis.
Wis.
Wis.
Aug.
Wis.
"Wis.
Sept.
**Wis.
'Wis.
Wis.
Oct.
"Wis.
Wis.
Wis.
Nov.
Dec.
36
Wis.
Wis.
'Wis.
'Wis.
Wis.
Wis.
during late fall and winter.
Summary. Copulation occurs during fall and
winter. Females lay eggs in t h e spring, release
their young in open surface water, and return
t o the burrow. Young-of-year remain in open
water for part of t h e summer but begin burrowing
as early as midsummer. Adult males usually spend
t h e summer as Form I1 and molt to Form I in
the fall.
DISTRIBUTION:
c.diogenes (see Fig. 28) is found
in every major drainage system in t h e state
(Appendix I) and is certainly tolerant of a variety
of environmental conditions (see Fig. 29 for total
geographical distribution).
CRAYFISH ASSOCIATES. Associated with C.
diogenes in one or more localities in Wisconsin
were Orconectes propinquus, 0.rusticus, 0.virilis,
Procambarus gracilis, P. a. acutus.
Figure 27. Photograph of ventral surface of female Cambarus (L.)diogenes dug from
burrow in roadside ditch in Green County (note cement glands).
37
Figure 28. Distribution of Cambarus
circles-from literature.
38
iL.1 diogenes in Wisconsin; solid circles-specimens
examined in this study; open
Figure 29. Geographic distribution of Cambarus
(L.)diogenes.
Fallicambarw (Crcmerius) fodiens (Cottle)
(Figures 30, 31)
Astacus fodiens Cottle, 1863217.
Cambarus argillicola - Cahn 1915:136,174.
Cambarus fodiens Hobbs 1948:229;Crocker and
Barr 1968:133 (Fig. 86); Radaj 1978:l.
-
DIAGNOSIS: Carapace (Fig: 30c,j) laterally
compressed, vaulted dorsally and lacking cervical,
branchiostegal, and hepatic spines or prominent
tubercles; postorbital ridge short, terminating
anteriorly without spine. Rostrum directed
somewhat ventrally, short, broad, and spatulate,
lacking marginal spines or tubercles and without
median carina; upper surface of anterior portion
concave with somewhat thickened margins;
acumen very short, triangular, and tip slightly
upturned. Cervical groove shallow, sinuous, and
discontinuous 1a.terally.Suborbital angle absent.
Areola (Fig. 30j) obliterated or linear, constituting approximately 40% of total carapace
length (47% of postorbital carapace length) with
no room for punctations in narrowest part.
Antenna1 scale (Fig. 30 1) approximately 2.3 times
as long as wide, broadest distal to midlength;
distal portion tapering to relatively blunt spine.
Cephalic lobe of epistome (Fig. 30g) triangular,
slightly broader than long with convex margins.
Distal incisor region of mandible (Fig. 30i)
dentate-crenate. Abdomen narrow and subequal
in length to carapace. Cephalic section of telson
with two prominent spines in each caudolateral
corner; proximal podomere of uropod with
caudal spine on mesial lobe, lateral lobe rounded;
39
Figure 30. Fallicumbarus (C.)diem (b,e, second form male Jasper County, Illinois; d, adult female Clark County, Illinois;
k,m, first form male Jasper County, Illinois; all others first form male from Washentaw County, Michigan): a,b,k, mesial
view of first pleopod; c, lateral view of carapace; d, annulus ventralis; e,f,m, lateral view of first pleopod; g, epistome;
h, proximal podomeres of third, fourth, and fifth pereiopods; i, incisor margin of right mandible; j, dorsal view of carapace;
I, antenna1 scale; n, dorsal view of distal podomeres of cheliped.
mesial ramus of uropod with prominent submedian dorsal keel terminating distally in distomedian spine; distolateral spine of ramus distinct.
Chela (Fig. 30n) heavy, approximately 2 times
as long as broad, palm swollen, and punctate;
mesial surface of palm with two rows of tubercles,
mesialmost consisting of 7-8, usually 7; less
distinct, more dorsal row of small tubercles
(usually 5); setal tuft situated between fingers,
tuft particularly dense on ventral surface; both
fingers tuberculate; dactyl with distinct excision
on opposable margin, fixed finger with one
prominent tubercle opposite excision of dactyl.
Carpus of cheliped furrowed dorsally with one
or more prominent mesial spines. Ischium (Fig.
30h) of only third pereiopod with simple, slightly
recurved hook extending proximally over
basioischial articulation; coxa of fourth pereiopod with boss not conspicuously large. First
pleopods of male (Fig. 30a,b,e,f,k,m)symmetrical,
contiguous basally with convexity near midlength, and terminating in two elements bent
caudally at least 90 degrees to shaft of appen.,
dages; arched central projection of first form
individuals corneous, bladelike, flattened
laterally, and with distinct subapical notch;
mesial process noncorneous, often appearing
twisted and usually with eminence on ventral
border distal to base; some specimens with one
or more small protuberances on mesial process
(Fig. 30k,m); mesial process extending distally
beyond central projection.
Annulus ventralis (Fig. 30d) not overlapping
postannular sclerite, subovate to subquadrangular and deeply embedded in sternum; fossa large
with deepest portion situated dextral or sinistral
to midline and swollen laterally and posteriorly
on U-shaped ridge, latter cut medially by narrow
sinuous sinus.
Because we have no material the sections
treating variation, color notes, and measurements are omitted.
TYPE-LOCALITY:Upper
Canada (Ontario?).
this species has not been
confirmed as occurring in Wisconsin, it is highly
likely that with additional field work in the
southern tier of counties it will be found. The
ECOLOGY. Although
Wingra Springs locality reported by Cahn (1915)
has been discussed above; ecological data were
supplied by him indicating that F. fodiens was
burrowing in moist soils supporting the marshgrass, “Spartinu.” As early as 1863, Cottle (p.
2 18) discussed the burrowing habits of this
species in Canada. Faxon (1884) reported this
species in Michigan to burrow to slightly less
than two meters. At the terminus of the burrow
he often found a pocket of loose gravel and clay
where, immediately above the water-line, it
formed a shelf on which the animal rested.
Crocker and Barr (1968) described the typical
burrow of F. fodiens as having one to three short
entrance tunnels that converge on a narrow
vertical tunnel. This tunnel usually extended
down to the water table. Williams et al.
(1974:367,369) discussed the burrow construction of adults and immature crayfishes alongside
an intermittent stream in Ontario. They
reported alterations to the general pattern occur
as young-of-year burrow and when females
become ovigerous. Maple (1968) and Reinert
(1978) demonstrated that the Massasauga
Rattlesnake, Sistrurus c. catenatus, in northeastern Ohio hibernates in the burrows of F. fodiens;
Maple presented a good description of the burrow
structure of this crayfish.
In addition to being found in burrows,
individuals are often seen in roadside ditches,
woodland or meadow ponds, swamps, marshes,
or backwaters, and very rarely from lotic
environments (see references listed below). In
Illinois, this species lives on wooded floodplains
or where wooded floodplains formerly occurred;
it lives in deep burrows and is in surface waters
only during floods (Page 1985:424). Maude and
Williams (1983) found this species t o have a
strong preference for mud substrates and to have
a mean slip speed of 26.7 cm/sec; that is,
individuals are able t o hold their positions on
a streambed with a current speed of no more
than 26.7 cmisec. They also indicated that the
body (cephalothorax) was laterally compressed,
an adaptation to burrowing that allows the
organism to escape into the burrow from any
angle. They also suggested that “streamlining”
was an adaptation for burrowing. Bovbjerg (1952)
41
looked at isolation among crayfishes with
overlapping ranges and showed that ecological
isolation (intolerance) played an important role
in enforcing segregation of species. F. fodiens (his
C. fodiens) was unable to tolerate rapid currents
and was thus excluded from streams by 0.
propinquus. 0.propinquus was not able to tolerate
temporary aquatic habitats and low oxygen
conditions that are typical of shallow ponds
occupied by F. fodiens.
Kiley and Dineen (1982:212) noted that in
Indiana man’s activities have reduced the
number of suitable habitats for this crayfish.
Obviously habitat alteration and destruction are
widespread problems affecting not only this
species.
Figure 3 1. Geographic distribution of Faliicamburus
42
(C.)
fodms
LIFE HISTORY. Although the life history of this
species is not fully known throughout its range,
Crocker and Barr (1968) presented a general
summary for the species in Ontario (based in
part on Creaser 1931b). They indicated that
adults move to open water in the spring after
snow melt. During March and into April the
females are ovigerous and young hatch in April.
Young become free-living during the latter half
of May when the adults return to their burrows.
The young-of-year begin burrowing by midsummer. Individuals overwinter in burrows and do
not emerge again until after the spring thaw.
Crocker and Barr (1968) suggest that there may
be a fall mating period but Williams et al. (1974)
did not find this to be the case in Waterloo
County, Ontario. Cummins (1921) discussed
fnigrations of this species in Michigan during
March and April. Migrations to ponds occurred
at irregular intervals and were coincident with
high humidities; most of the individuals involved
were adults, with some being ovigerous females
as well as some carrying young. Page (1985) noted
that size-frequency distribution data from
southern Illinois populations suggest that F.
fodiens reaches sexual maturity and dies in its
second year.
DISTRIBUTION: The distribution of F. fodiens is
shown in Fig. 3 I. See Cahn (19 15) for additional
information concerning the occurrence (?)of this
species in Wisconsin.
Orconectes (Cjremicambarus)immunis (Hagen)
(Figures 32 -35)
Orconectes immunis Hagen,1870:71.
Cambarus immunis Faxon 1884:146; 1885b399;
-
1885c:359; Underwood 1886:370; Ortmann
1902:280; Harris 1903a:102, 155; Faxon
1914:379; Creaser 1931133263; 1932:323, 324,
326, 328, 329, 335, 336; Creaser and
Ortenburger 1933:38; Turner 1935:875;
Walters 1939:23; Hinkleman 1970:7.
Cambarus (Faxonius) immunis
Graenicher
-
1913:118,120,121,122,123.
-
Orconectes immunis sspp. Penn 1950:646,648.
Orconnectes immunis Threinen 19581332.
Orconectes immunis
Wiens and Armitage
196150; Crocker and Barr 1968:107; Hinkleman 1970:49; Radaj 1978:Z; Capelli
1975:39,40,206,207; Crocker 1979:235,247;
Capelli 1982a:74 1; Threinen 1982a:78,79;
Capelli and Magnuson 1983548, 551, 558,
559, 563, 564+
-
-
DIAGNOSIS (refer to Table 4): Carapace (Fig.
32c,j) subcylindrical, somewhat compressed
laterally; rostrum longer than broad, excavate
dorsally with marginal rostra1 spines indistinct
or absent; postorbital ridge with blunt spine.
Cervical groove moderately deep and discontinuous laterally; distinct but short cervical and
branchiostegal spines present. Areola (Fig. 32j)
narrowest in anterior half of length with 1 or
2 punctations in narrowest point; areola
constituting 8.2 to 37.9% (mean 33.9%) of total
carapace length (10.5 to 49.2%, mean 44.4%,
of postorbital carapace length) and 2.7 to 20.5
(mean 11.0) times as long as broad. Antenna1
scale (Fig. 32i) generally 2.2 times as long as broad,
widest proximal to midlength, apex with small
terminal spine. Cephalic lobe of epistome (Fig.
3211) broader than long, rounded with uneven
convex margins. Distal incisor region of mandible
(Fig. 32g) irregularly dentate-crenate. Length of
abdomen nearly equal to that of postorbital
carapace length; cephalic section of telson with
two spines in each caudolateral corner, distal
podomere of lateral lobe of uropod rounded;
mesial ramus of uropod with distomedian spine
never reaching distal margin; small spine situated
on distolateral corner. Chela (Fig. 32m)
punctate, slender (approximately 2.5 times as
long as broad), tuberculate along mesial margin
of movable finger and palm; one prominent
tubercle on opposable margins of fingers,
prominent tubercle on mesial side of dactyl either
proximal or distal to that on mesial surface of
fixed finger; dactyl with distinct but shallow
excision on opposable margin; mesial surface of
palm with one distinct (mesialmost) row of
tubercles (consisting of 5-8) and less distinct row
of 3 t o 6 small tubercles; setal t u f t (not
illustrated) situated between fingers, tuft
particularly dense on ventral surface of immovable finger. Carpus of cheliped deeply grooved
dorsally with one prominent spine and several
small spines mesially, 1 or 2 large median ventral
spines. Ischium of only third pereiopod (Fig. 32d)
with large hook extending proximally over
basioischial articulation. First pleopods of first
form male (Fig. 32a,f) reaching coxae of third
pereiopods when abdomen flexed; terminal
elements subparallel and with apices directed
caudally at angle of approximately 90 degrees
t o principal axis of shaft with noncorneous mesial
process slightly longer and more strongly bent
caudally than central projection (Fig. 32a,b,e,f).
Annulus ventralis (Fig. 32h) subovate,
distinctly broader than long and with caudal
margin of nontuberculate sternum overhanging
its cephaloventral region; annulus ventralis with
deep sinuous depression across midlength,
43
deepest portion situated dextral to midlength;
postannular sclerite approximately 213 as broad
as annulus.
VAR~ATION. T h e most obvious morphological
variations occur with reference to the gonopods
of Form I males. In “typical” individuals, the
terminal elements (central projection and mesial
process) of the first pleopod are bent caudally
at a n angle of approximately 90 degrees to the
Principal axis of the appendage with both
elements subparallel to one another (Fig. 32a,f)*
Table 4. Range of measurements (in mm) of various diagnostic structures for Wisconsin 0.
immunis.
N
Minimum
Maximum
Mean
Standard
Deviation
Jr
MALES ($1)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
Pleopod
Total length
Central projection
Mesial process
Condyl length
-
23
25
25
23.0
17.2
9.7
38.5
40.1
19.6
31.4
24.1
14.4
4.6
25
25
3.0
0.6
18.5
1.7
11.0
1.0
2.8
0.2
18
18
4.6
5.9
14.3
21.3
8.2
10.3
2.2
3.4
23
23
24
23
7.9
1.2
1.0
7.6
12.4
2.5
2.6
11.3
10.7
2.0
1.6
9.8
1.3
0.3
0.4
1.2
31
31
31
17.4
12.3
7.7
48.0
39.1
20.7
29.2
22.2
12.8
6.8
5.6
2.9
31
31
5.8
0.6
18.2
1.8
10.1
1.o
2.6
0.3
15
15
3.4
3.6
9.9
13.4
5.4
6.6
1.8
2.5
10
7.3
21.4
10.4
4.0
62
64
64
19.3
14.1
8.5
41.8
32.2
19.5
29.2
22.3
12.9
5.2
4.1
2.4
64
64
6.3
0.4
14.6
1.8
9.8
1.o
1.8
0.3
37
37
2.1
2.3
8.3
12.2
4.7
6.2
1.4
1.9
4.7
2.3
MALES (811)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela .palm
Length
Width
Pleopod
Length
FEMALES
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela .palm
Length
Width
44
P
k
n
Figure 32. Orconectes (G.) immunis (b,e, second form male Otter Creek, Rock County; h , female also from Otter Creek;
I, first form male Birch lsle Lake, Burnett County; all others first form male from Allen Creek in Jefferson County):
a,b, mesial view of first pleopod; c, lateral view of carapace; d, proximal podomeres of third, fourth, and fifth pereiopods;
e,f,l, lateral view of first plcopod; g, incisor margin of right mandible; h, annulus ventralis; i, antenna1 scale; 1, dorsal
view of carapace; k, dorsal view of anterior portion of carapace; m, dorsal view of distal podomeres of cheliped; n, epistome;
cp, central prqection; mp, mesial projection; cdl, condyl.
The mesial process is more sharply bent in some
specimens (Fig. 32 1) and usually extends slightly
beyond the apex of the central projection; in
some, both elements are equal in length.
The acumen of the rostrum is commonly
distinct (Fig. 32j); however, in some specimens
it is greatly reduced (Fig. 32k). In addition, the
number of tubercles on the mesial margin of
the palm of the chela varies from 5-8 in the
mesialmost row. The setal tuft, situated between
the fingers, is variable in its extent but generally
is denser on the mesoventral surface of the
immovable finger.
COLORNOTES(refer to Fig. 33). Carapace brown
to olivaceous dorsally fading ventrally to lighter
shades. Carapace and chelipeds possessing or
lacking dark brown irregular splotches over entire
surface; rostra1 margins and postorbital ridges tan
to light olive. Abdomen similarly brown to olive
but lighter; caudal portion of terga margined in
dark brown or dark olive. Tan middorsal stripe
originating between postorbital ridges (here
stripe faint), extending posteriorly along areola
and along terga of abdomen (where more
distinct), terminating o n sixth abdominal
segment, generally not extending onto telson;
stripe not of consistent width; hourglass shaped
over areola and with expanded blocks of tan
centrally located on each tergum; each pleuron
with olive green to brown stripe at base. Telson
and uropods tan with small, irregular splotches
of dark brown. Antennular and antenna1
peduncles dark brown or dark olive with darker
lateral margins. Cheliped with dorsal surface of
carpus, merus, and propodus tan to light olive
bearing irregular brown to dark olive splotches;
some specimens with reddish-purple propodus
(see Threinen 1982a:78); distal part of fingers
with bright orange t o red tips. Pereiopods 2-5
light tan to cream with irregular dorsal olive
splotches. Sternal area of cephalothorax cream
or pale tan,
Kent (1901) indicated that individuals take
o n t h e color of t h e “environment” and
attributed red color of those found in shallows
to light exposure. He noted that young also were
predominently red but that within two months
of hatching they changed to match the dominant
color of the substrate.
TYPE-LOCALITY:Lawn
Ridge, Marshall County,
Illinois, USA.
ECOLOGY. Habitats. 0. immunis is found in a
wide variety of habitats and over its range is
generally found in either standing or slowly
moving, clear (or turbulent) water (permanent
or temporary) with a soft substrate usually
Figure 33. Photograph of Form I male Orconectes (G.)
immunis from unnamed creek in Ozaukee County,
46
supporting vegetation (e.g., Elodea, Potumogeton,
Marsilea - Tack, 1941). In addition this crayfish
may move to deeper water as the habitat begins
to dry (Phillips 1980) or burrow to the water
table. As early as 1876 Forbes (1876:4) reported
that this species in Illinois frequented muddy
ponds of the prairies. Creaser (1932:327-328)
indicated that 0. immunis is found in rivers and
streams where it avoids strong currents, in small
lakes, especially those with muddy bottoms, and
sometimes in temporary ponds. Maude and
Williams (1983) also demonstrated that it could
not tolerate high velocity streams (mean slip
speed of 26 cm/sec) and that it showed a strong
preference for mud substrates. However, Hobbs
and Marchand (1943:27) stated that in the
Reelfoot Lake area of Tennessee 0.immunis was
found high up the slopes in rapid lotic habitats
but occurred only in small numbers in sluggish
sections of valley streams.
Clark and Rhoades (1979) observed this
crayfish in wet plains and swamp forests of Ohio.
It dwells in streams, lakes, or ponds with an
abundance of vegetation. Osburn and Williamson (1898) indicated that 0. immunis was even
found in Lake Erie, 22 miles from shore. Barr
(1969:93), in discussing this species in eastern
Canada, stated that it is found in “. . shallow
ponds . . with high temperature, low oxygen
concentrations, high density of suspended mud
. . . .” Caldwell and Bovbjerg (1969) reported 0.
immunis in northwest Iowa from temporary and
permanent ponds, large shallow sloughs, small
muck-bottomed lakes, and slowly moving softbottomed portions of rivers and smaller streams.
In addition, Phillips (1979) found 0. immunis
in roadside ditches, overflow pools, marsh areas,
natural and artificial lakes, and from the
headwaters of small, warm water streams. He
(1979:51) indicated that 0. immunis demonstrated a distinct preference for a mud substratum since 91% of the lots containing this species
were collected over mud. Of these, 74% were
collected from sites with dense aquatic vegetation. Nevins and Townes (1935) noted that 0.
immunis is tolerant of low oxygen concentrations;
according to laboratory experiments conducted
by Park et al. (1940) and Bovbjerg (1970a), the
.
.
species can tolerate low oxygen concentrations
(<3.1 mg/l) for considerable time. Wiens and
Armitage ( 1 9 6 ~ 2 0 3 )demonstrated a direct
relationship between oxygen consumption and
increased water temperature and showed that
there is a decrease in consumption w i t h
decreasing saturation. They also noted an inverse
relationship between metabolic rate and body
weight. Helff (1928) proposed that tissue cells
of 0. immunis (his C. immunis) could store
oxygen, thereby creating a reserve oxygen supply.
When a certain low point in the reserve is
reached, the crayfish replaces it by increased
respiration (an adaptative strategy to living in
waters with low oxygen content that may be
shown for other species in similar habitats).
Aiken (1965:241) reported the presence of this
species in Newfound Lake in New Hampshire,
a clear (compensation point at 17m), well
oxygenated (8mg/l oxygen at 15m), relatively
unproductive lake where 0. immunis was found
beneath flat stones in less than one meter of
water.
In addition to these habitats, this species
frequents temporary bodies of water. Harris
(1903b:603) stated that individuals burrow when
ponds begin to dry and upon the approach of
winter. He (1901) indicated that the burrows
are variable, with a crescent-shaped pile of dirt
covering the opening or chimneys lOcm high
and 13cm in diameter. He noted the tunnels
are up to 38cm deep by 4cm in diameter with
a 9 lOcm wide chamber at the bottom. Some
were reported to be over 1.2m in depth, generally
vertical, and often with a dead-end side branch.
In a locality in Ontario Huntsman (1917:131)
noted the burrows are generally (‘tortuous,’)
having a single or multiple entrances. The
openings are sometimes entirely on the bank
above water, at other times some above and some
below, and occasionally all below. Maude and
Williams (1983) indicated the cylindrical-shaped
carapace was an adaptation to a fossorial
existence, this shape allowing an individual to
escape down its burrow from any angle. Kenk
(1949), studying populations in southern
Michigan, noted that 0. immunis does not
burrow when it occurs in permanent waters and
-
47
Steele (1902~26)observed during rainy weather
that individuals are often found moving about
in meadows and fields and along roads. Crocker
and Barr (1968) reported that at night some may
travel from pond to pond; during the winter
individuals burrow below the frost line (Phillips
1980 and Caldwell and Bovbjerg 1969). Rhoades
(1944a) noted that where gravel bottomed
streams occur it becomes a burrower in muddy
banks. This species will burrow singly (Ortmann
1906) or in pairs (Engle 1926).
Physicochemical garumeters. Concerning
response to environmental conditions, Crawshaw (1974) showed that during the day 0.
immunis was relatively inactive, selecting a mean
water temperature of approximately 18°C; during
the night individuals were active and selected
a mean water temperature of approximately
22°C. At night they wandered about through
a relatively wide range of temperatures, avoiding
only the extremes. This type of orientation was
termed klino-kinesis by Fraenkel and Gunn,
1961. During the day the crayfish selected cooler
water in which to become inactive (also noted
by Crocker and Barr, 1968), which in the natural
environment would necessitate their migration
into deeper water. Crawshaw (1977) also
demonstrated that individuals acclimated to 7°C
readily entered water much warmer than 15°C
soon after placement in a temperature gradient.
Discussing the effects of power plants on several
species of crayfishes (0.
immunis, 0.gropinquus,
0. virilis, and others) Crawshaw et al. (1980)
noted that the most obvious by-product of power
plant operation is the heat put into the
environment. This can have far-reaching effects
by altering growth and reproductive timing, by
changing resources upon which crayfishes
depend, and by altering the density of crayfish
predators. They indicated increased predator
density , . is potentially the greatest source of
imbalance for the crayfish as well as the
remainder of the community if refugia for young
crayfish are limited." In response to other
environmental
perturbations,
Creaser
immunis as being tolerant
(1934b~161)noted 0.
of sulfite pollution in Raquette River in New
York. Bovbjerg (1970a) discussed its interaction
".
48
with 0. virilis in Minnesota and Iowa and
concluded that it is excluded from streams by
0.uirilis. Caldwell and Bovbjerg (1969)also noted
this exclusion by the latter in Iowa and indicated
that 0. virilis eliminates 0. immunis from rivers
and streams and restricts it to temporary or
permanent ponds.
LIFE HISTORY.Egg production and hatching time.
The first observations concerning the life history
of 0. immunis in Wisconsin appeared in 1913
when Graenicher reported females with eggs in
the spring of 1910 from a pond in Wauwatosa
(Milwaukee County) on 20 March. Creaser
(1932:328) noted females with eggs from
November to February and Threinen (1982a)
reported Form I males from Polk County during
September. Although the life history is not
completely understood for 0. immunis in
Wisconsin, it is well known in various parts of
its geographical range.
The most detailed studies on the life history
of this species were conducted by Tack (1941)
in New York, and by Caldwell and Bovbjerg
(1969) in Iowa. Tack reported copulation from
mid-June t o early October, mostly among
yearling individuals and Caldwell and Bovbjerg
noted this occurring from late June to the
following April, except during the winter
months. Fasten (1914), working with Nebraska
populations, demonstrated a chromosomal
number of 208 and indicated that during June
and July the testes undergo greatest proliferation
and increase to maximum size; during this period
all stages of spermatogenesis are observed. Tack
found that eggs are laid in late October or early
November, carried through the winter, and
hatch in mid- to late May. Berrill (1978:167)
also noted that in Ontario, some females extrude
their eggs in late autumn and carry them over
winter. Females are in berry in Illinois from
January through April (see Fig. 5 Page 1985)
and carry young from April through June (Rietz
1912, Brown 1955). Threinen (1958b) reported
that in Wisconsin eggs were laid in the fall and
that the young hatch in the burrow; Forney
(1956, 1957) reported the number of eggs varies
from 60 300. Phillips (1980) collected a female
in berry as early as 22 April in Iowa. This pattern
-
-
has been noted in other parts of its range (Engle
1926, Nebraska - April: Creaser 1931b, Michi-
-
-
gan Nov., Feb.; Creaser 1932, Wisconsin
November to February; Creaser and Ortenburger
1933, North Dakota
June; Williams and
Leonard 1952, Kansas October; Forney 1956,
1957, New York mid-October; Aiken 1965,
New Hampshire early June; and Crocker and
Barr 1968, Ontario May). Hatching time varies
considerably depending on when the eggs are
laid and on water temperature. Tack (1941)
noted that after hatching the young remain
attached through the first two stadia, after which
they leave the female but stay in close proximity
during the third stage. Caldwell and Bovbjerg
(1969) reported that the young remain attached
to the mother for a total of seven to 19 days,
and Forney (1957) listed the time as ranging from
one to two weeks; females molt shortly after
the young hatch and leave.
Young of the year. The young molt frequently,
eight or more times in the first year (Forney
1957), and growth is highly variable depending
in part on the abundance of food and whether
or not they are forced to burrow due to summer
drying. Wiens and Armitage (1961) showed that
as individuals increase in size (body weight) there
is a corresponding decrease in metabolic activity.
This species is a variable but productive crayfish,
ranging from 51.5 to 1345 kg/ha of biomass
produced per year in ponds (see Lydell 1938,
Tack 1941, and Goellner 1943).
0.immunis may grow rapidly enough to reach
the minimal size necessary for sexual maturity
by their first autumn, this being 22mm as
reported by Tack (1941) and 23mm by Crocker
(1957). Berrill (1978:172) noted the range of
carapace lengths of Form I males in Ontario to
be 16-35mm, with a mean of 23mm. Females
with eggs or young range in size from 16-33mm,
with a mean carapace length of 23mm. In ponds
that become dry in midsummer, only a small
number of young mature by the end of their
first summer, whereas in permanent ponds most
individuals become mature during their first
growing season. Forney (1957) reported that
some females lay eggs five months after hatching
but that most do not reach sexual maturity until
-
-
-
-
-
the second summer. Caldwell (1969) noted that
flexibility in the eggdlaying season and the time
required for sexual maturation allow 0. immunis
to meet the common yet unpredictable occurrence of pond habitat dessication.
Tack (1941)reported that by September, when
the majority stop growing, the range in length
of the carapace of young-of-the-year varies from
13 t o 29mm. Virtually all individuals stop molting
from mid-November until late March or April.
The life span of both sexes apparently depends
upon the time of maturation. Those individuals
that mature during the first summer generally
die following the spring molt of the second year.
Those individuals remaining sexually immature
during the first summer usually live two full years.
Molt cycle. In addition to the above comments
concerning molting, the following summarizes
what is known of the molt cycle for this crayfish
based primarily on data from New York (Tack
1941). Form I males begin molting to Form I1
as early as mid-April and continue through midMay. Another molt which returns Form I1 males
to Form I generally begins in mid-June and
continues through mid-July. Spring molting is
a period of high mortality for adult males and
probably accounts for the large reduction of adult
males from the population during this season.
After the young leave the females in spring a
similar period of high mortality occurs among
the females. Males also are lost from the
population in August and September following
breeding (Tack 1941).
Scudamore (1948) noted that the delay in the
spring molt of berried females is regulated by
the action of the molt-inhibiting hormone of
the sinus gland. Prins et al. (1972) demonstrated
that there was no significant photoperiod effect
and thus no interaction between photophase and
temperature in affecting the molting process of
this species. Molley and Prins (1973) showed a
significant direct relationship between temperature and molting. Talton (1977) and Talton
and Prins (1978), using individuals from
Michigan, conducted laboratory experiments
that showed that 0. immunis does not use an
“Hourglass model” for photoperiodic time
measurements. They pointed out that it is not
49
known whether or not the stage of sexual
development affects t h e measurement of
photoperiodic time since no significant interactions between sex and photoperiod were
detected. They demonstrated significantly
greater numbers of molts at higher light
intensities and observed an increased molting
frequency with increased water temperatures.
Life history data are summarized for 0.
immunis in Table 5.
DISTRIBUTION:
In Wisconsin the distribution of
0. immunis is disjunct, with populations found
in the southern tier of counties, the northwestern part of the state (Vilas County), and central
Wisconsin (Wood County) (see Fig. 34).
Immigration into Wisconsin appears to have
been from two directions: one from the south
and east (led to the establishment of populations
in the southern counties) and those populations
in the northwest were established as the species
moved across the Mississippi and up the St. Croix
River. Its presence in the extreme southeastern
portion of Bayfield County (see Fig. 34) suggests
that this species is probably located throughout
the Chippewa River drainage. It is undoubtedly
widely distributed in the Namekegon drainage
as well. 0.immunis is present in all the Mississippi
River basins except the Trempealeau-Black and
is known only from the Lake Michigan drainage
in the St. Lawrence system (see Appendix I).
Additional field work, particularly sampling
ponds, should show this species occupying many
more drainages throughout the state. The known
total geographical distribution of this species is
shown in Fig. 35.
CRAYFISHASSOCIATES.In Wisconsin 0. immunis
was associated in one or more localities with C.
diogenes, 0.propinquus, 0.rusticus, 0.uirilis, and
P. a. acutus.
TABLE 5. Life history data for 0. immunis (* from references only; ** from literature and
our collections or USNM collections; those without asterisk(s) from USNM collections only).
$I
$11
Oe
Jan.
"Wis.
'Wis.
'Wis.
Feb.
'Wis.
'Wis.
'Wis.
March
'Wis.
'Wis.
'Wis.
'Wis.
April
"Wis.
"Wis.
'Wis.
'Wis.
May
"Wis.
"Wis.
"Wis.
June
'Wis.
"Wis.
July
"Wis.
"Wis.
'Wis.
Wis.
Aug.
"Wis.
"Wis.
'Wis.
Wis.
Sept.
'Wis.
'Wis.
'Wis.
Oct.
'Wis.
'Wis.
'Wis.
Nov.
'Wis.
'Wis.
'Wis.
Dec.
'Wis.
'Wis.
Month
50
~y
yoy
copul
"Wis.
molt
'Wis.
'Wis.
Wis.
'Wis.
burrow
L
920
910
L
9'
0
I
N
89'
Figure 34. Distribution of Orconectes (G.) immunis in Wisconsin; solid circles-specimens
circles-from literature.
I
Baa
87'
examined in this study; open
51
Figure 35. Geographic distribution of Orconectes (G.)immunis
Orconectes (Crockerinus) propinquus (Girard)
(Figures 36 - 45)
Cambarus propinquus Girard, 1852:88.
Cambarus propinquus - Hagen 1870:69; Smith
1874:638; Bundy 1882:177, 178, 179, 181;
1883:402, 403; Faxon 1884:147; 1885b:7, 13,
14, 91; Underwood 1886:371; Harris
1903a:59, 120, 150, 155; Ortmann 1905:115;
1906:363; Cahn 1915:174; Muttkowski
1918:393, 424, 474, 477, 480; Pearse
1918:252; Pearse and Achtenberg 1920:312;
Baker 1924:119, 138, 144; Pearse 1924:255,
256; Turner 1924:263-277; 1926:154, 158,
183; Creaser 1931b:266; 1932:325, 329, 335,
336; Turner 1935:865-870,872,873,875,876;
Van Deventer 1937:11, 12, 33; Walters
52
193926, 110, 168; Scudamore 1948230-232,
236; Hinkelman 1970:7; Hart and Hart
1974:85.
Cambarus (Faxonius) propinquus - Ort mann
1906:363, 433, Fig. 3; Graenicher 1913:118,
119, 120; Ortmann 1931:66.
Orconectes propinquus sspp. - Penn 1950:646,
648, 649.
Orconectes propinquus propinquus - Penn
1950:644; Crocker 1957:50,75; Crocker and
Barr 1968:73.
Orconectes propinquus - Bovbjerg 1952:42;
Fitzpatrick 1967:146, 166, 167; Crocker and
Barr 1968:72 (fig. 70); Hobbs 1968:K13;
McKnight 1969; Hinkelman 1970:7, 29, 31,
32, 33, 35, 36, 44, 45, 47; Avault 1971:6;
Bell 1971:16; Fielder 1972:142; Hobbs
1972b:83; Avault 1973:244; Hart and Hart
1974:24, 85, 120, 138; Hobbs 1974b:39;
Magnuson et al. 197567, 68, 69, 70, 71, 72;
Stein 1975a:20 [by implication]; Capelli
1975:5, 9, 39, 40, 44, 46, 48, 50, 53-58, 60,
61, 63-72, 79-83, 85-130, 134-158, 163-196,
199-201, 206, 207; Hobbs 1976a:83; Capelli
and Magnuson 1976:415,416;Stein 1976:221227; Stein and Magnuson 1976751-760; Stein
and Murphy 1976:2450, 2451, 2453, 2454,
2455, 2456; Stein 1977:1237-1253; Stein et
al. 1977:495-502; Lorman and Magnuson
1978:9; Capelli 1978:59; Payne 1978:6; Radaj
1978:2; Sheffy 1978:222; Malley and Reynolds
!979:313 [by implication], 316; Stein
1979:345, 346 [by implication]; Crocker
1979:243; Capelli 1980:83-86; Capelli and
Capelli 1980:121 132; Phillips 1980:87; Zaret
1980:130, 131; Horns and Magnuson
1981:299-302; Capelli 1982a: 741-745;
Tierney and Dunham 1982:547; Welles
1982b:99; Mather et al. 1982552; Maude and
Williams 1983:76; Capelli and Magnuson
1983:548, 549, 551-554, 557-561, 563, 564;
Serns and Hoff 1984:1@, 19; Lodge et al.
198533; Berrill 1985:347,348; Magnuson and
Beckel 198510; Butler and Stein 1985:169;
Lodge et al. 1986; Jezerinac 1986:178; Lodge
and Lorman 1987:591,592.
Orconnectes Propinquus Threinen 195813:1,2;
Threinen 19821333.
Orconectes Propinquis - Jaeger 1977:2571-B;
Dodson and Cooper 1983:347,350.
Crayfish Stamm 1977:1@4,105,108,109.
-
.
-
DIAGNOSIS (refer to Table 6): Carapace (Fig.
36c,j) subovate, depressed dorsally; cervical
groove moderately deep and discontinuous
laterally; carapace with distinct cervical spine,
branchiostegal spine absent; rostrum narrow, its
sides subparallel with marginal spines, excavate
dorsally but usually with prominent median
carina, postorbital ridge with sharp to blunt
spine. Areola (Fig. 36j) narrowest in anterior half
of length with 4 to 5 punctations in narrowest
part; areola constituting 31.0 t o 36.1% (mean
33.6) of total carapace length (40.1 t o
45.8%, mean 43.0, of postorbital carapace
length) and 2.8 to 5.6 (mean 4.3) times as long
36;) generally 2.3
times as long as broad, widest about midlength
tapering anteriorly t o small terminal spine.
Cephalic lobe of epistome (Fig. 361) slightly
broader than long, subtriangular but rather
rounded anteriorly. Distal incisor region of
mandible (Fig. 36h) irregularly dentate-crenate.
Length of abdomen subequal to postorbital
carapace length; cephalic section of telson with
large (lateral) and small (mesial) spine in each
caudolateral corner; distal podomere of lateral
lobe of uropod usually rounded; mesial ramus
of uropod with distomedian spine terminating
immediately proximal to distal margin of ramus;
small spine situated distolaterally. Chela (Fig.
36m) moderately punctate, slender (approximately 2.3 times as long as broad), two distinct
rows of tubercles along mesial margin of movable
finger and palm, rows on latter consisting of 7
to 10 tubercles; opposed margins of fingers with
5 to 7 prominent tubercles. Carpus of cheliped
moderately cleft dorsally with one prominent and
2 or more smaller spines or tubercles mesially;
3 to 5 median ventral spines. Ischium of only
third pereiopod (Fig. 36k) with hook, latter
extending proximally over basioischial articulation. First pleopod of first form male (Fig.
36a,d,e,g) reaching coxa of second pereiopods
when flexed; terminal elements of first pleopod
of males parallel, short, usually subequal in length
(Fig. 36a,b,d-g), mesial process only slightly
shorter than central projection, moderately
slender, tapering distally to acute tip.
Annulus ventralis (Fig. 36n) distinctly less
than twice as broad as long; not partly obscured
by caudal margin of nontuberculate sternum
immediately anterior to it; ventral face bearing
deep, sinuous, transverse depression across
midlength, fossa situated dextral to midlength;
anterior fourth of postannular sclerite sometimes
overlain by annulus ventralis and about half as
broad as annulus.
as broad. Antenna1 scale (Fig.
TYPE-LOCALITY: Girard
(1852) originally listed
three localities; Ortmann (1905) designated
Oswego, Oswego County, New York, USA and
this selection was accepted by Crocker (1957:35)
and Fitzpatrick (1967:146).
53
a
I
Figure 36. Orconecres (C.) propinquus (a,g, from first form male from Little Lake o n Washington Island, Door County;
d, from first form male from East Twin River, Kewaunee County; 0,from first form male from Branch River, Manitowoc
County; n , from adult female from Allen Creek, Rock County; b,f, from second form male from Allen Creek; all others
from first form male also from Allen Creek): a,b, mesial view of first pleopod; c, lateral view of carapace; d-g, lateral
view of first pleopod; h,o, incisor margin of right mandible; i, antenna1 scale; j , dorsal view of carapace; k, proximal
podomeres of third, fourth, and fifth pereiopods; I, epistome; m, dorsal view of distal podomeres of cheliped; n , annulus
ventralis.
Table 6. Range of measurements (in mm) of various diagnostic structures of Wisconsin
0.
propinquus.
N
Minimum
Maximum
Mean
Standard
Deviation
*
MALES ($1)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
Pleopod
Total length
Central Projection
Mesial Process
Condyl length
17.2
13.0
6.1
34.7
26.8
13.5
26.2
20.5
10.3
4.1
3.4
1.3
42
5.4
1.3
11.9
3.3
8.8
2.2
1.6
0.4
41
41
3.9
4.5
11.3
14.5
7.3
9.7
1.7
2.4
42
42
42
42
7.0
1.8
1.6
6.0
12.1
2.9
3.0
10.0
9.8
2.3
2.3
8.0
1.3
0.3
0.3
1.1
38
38
38
18.3
13.9
6.6
34.9
29.7
14.0
28.0
21.9
10.6
3.8
3.3
1.8
38
38
6.1
1.6
12.6
2.9
9.5
2.2
1.5
0.3
36
36
4.0
4.6
10.7
14.1
7.1
9.3
1.7
2.3
39
40
40
22.1
17.4
8.1
40.5
31.4
15.2
27.6
21.5
11.0
3.4
40
40
7.3
1.5
13.8
3.2
9.2
2.3
1.3
0.5
37
37
3.7
5.7
9.2
13.3
5.7
8.1
1.1
1.4
42
42
42
42
MALES ($11)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
FEMALES
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela .palm
Length
Width
'
VARIATION. Although Orconectes profiinquus is
not an extremely variable species, individual
variation was noted among and between
populations within the state. The relative size
and elevation of the rostra1 carina is probably
more variable than any character examined.
Variations in morphological features measured
are indicated in Table 6. Some differences were
2.7
1.6
noted in color (not color pattern) but considerable variation was found in the gonopods of
Form I males. Usually, the terminal elements
are subequal in length, the mesial process being
only slightly shorter than the central projection
(Fig. 36a,g). Some specimens however exhibited
a much shorter and blunter mesial process (Fig.
36e) or possessed a slightly longer one (Fig. 36d).
55
Comparison of Figs. 36h and 360 demonstrates
some of the variation that was noted among
mandible structures.
Using gel electrophoresis, Nemeth and Tracey
(1979) conducted a study of allozyme variability
among six species of Cumburus and Orconectes
from Ontario. For 0.propinquus they estimated
a mean heterozygosity of 4.3 to 8.4% (mean 6.0
1.7), with a mean genetic similarity (or
identity) of 0.946
0.04; for 0. virilis the
estimated mean heterozygosity was 2.9% (+
11.4); and the estimated mean heterozygosity for
0. immunis was 2.1 to 8.1% (mean 4.2 f 3.4)
with a mean genetic similarity of 0.997 f 0.003.
They concluded that the estimated mean
heterozygosity was low for crayfishes in comparison with averages calculated for other invertebrates; this may indicate low mutation rates
or low intra-cistronic (unit of function in a DNA
system) recombination rates.
Gynandromorphs are not uncommon in
crayfishes. Faxon (1884, 1885b), Benham (1891),
and Andrews (1909) reported such isolated
I1
abnormalities” occurring in various crayfishes
over the world. Hay (1905) quoted Faxon
(188513) and discussed gynandromorphs from
Washington, D. C., Ohio (0.propinquus), and
Tennessee, and Penn (1957) reported on this
phenomenon in Cambarellus (Dirigicumbarus)
shufeldtii (Faxon) and Procamburus (Pennides)
dupratqi Penn. Turner (1924, 1935) presented
the most detailed account of crayfishes having
abnormal sexual characters: 1) cases in which
individuals of one sex have one or more
additional characters of the opposite sex (e.g.,
females with first pleopod of male); 2) cases in
which individuals have supernumerary secondary
sex characters or in which characters are lacking
(e.g., females lacking an oviducal pore on one
side); 3) “miscellaneous types” (e.g., females
utilizing male genital pores on fifth pereiopods
as oviducal openings). See Turner (1935) for a
listing and discussion of Wisconsin gynandromorphs, many of which are representative of
*
*
0.propinquus.
t o Fig. 37): Ground color
of carapace tan, gray-green, or olive-brown,
fading ventrally on hepatic and branchiostegal
regions t o slightly lighter tan or gray.
COLOR NOTES (refer
56
Posterodorsal portion of cephalic section
(between and over mandibular adductor area)
dark brown, or more commonly, black.
Posterodorsal portion of thoracic section dark
brown or black, dark color extending laterally
over posterior part of branchiostegal region
joining longitudinal dark band extending along
entire ventral branchiostegal region; often most
of areola dark, darker in posterior region.
Abdomen tan to olive-brown with dark brown
to black mid-dorsal stripe, most pronounced
anteriorly on terga, fading laterally, much lighter
on last abdominal segment, and never extending
onto telson; each pleuron tan or with dark brown
or black splotch. Telson and uropods tan to olivebrown, slightly darker than ground color of
carapace, uropods with pale tan margins and
spines. Cheliped tan t o olive-brown dorsally,
cream ventrally; distal part of merus and carpus
slightly darker brown or olive-brown; distinct
lateral margin of propodus black to tan; tubercles
of chela black to dark brown; tips of fingers bright
orange to red. Pereiopods 2 to 5 with merus
through dactyl tan to olive-brown dorsally fading
to pale tan (sometimes pale pink or light bluishgreen) ventrally. Sternal area of cephalothorax
and most of ventral surface cream or pale tan.
Loeb (1967) reported a blue specimen of 0.
p. propinquus from the Susquehanna River in
New York. Dunham et al, (1979) described a
“new color morph” of 0. propinquus from
Ontario: white with a broad black middorsal
abdominal stripe. They noted that the acumen
length and the areola width are absolutely greater
in the white morphs than the more typical brown
morphs but proposed no explanation for the
occurrence of the white form. Jordan and
Dunham (1981) reported “chestnut” and
“brown” color morphs in immatures from Lake
Simcoe in Ontario, Canada. During our field
work in July 1982, we collected a light bluewhite colored first form male of 0. propinquus
from Clark Lake, Door County (Fig. 38); also
very light gray or pale blue colored individuals
were obtained in other localities in Door County,
including Lake Michigan (Fig. 6). The Clark Lake
specimen was the only extremely light individual
noted and may, indeed, represent another “white
morph” as described by Dunham et al. (op. cit.).
Figure 37. Photograph of “normal” color morph of Orconectes
Ozaukee County.
(C.)
propinquus
(Form I male) from unnamed creek in
Figure 38. Photograph of Form I male Orconectes (C.) prtrpinquus from Clark Lake, Door County; this may be same
“white” color morph reported by Dunham et al. (1979) from Ontario, Canada.
57
Of note, in all localities in which we collected
pale forms, the substrate was very light; in many
cases, stones were covered with a pale gray marl
encrustation. With no detailed study, we hesitate
to claim this color trend to be an ecophenotypic
adaptation, but note this as a possible explanation for the presence of light colored morphs
present on light colored substrates. No light
colored crayfishes were observed on dark
substrates. Bowman (1942), studying P. (Scupulicumbarus) clurkii, demonstrated that chromatophores change on the basis of time spent on
a black or a white background.
Color anomalies, although not particularly
common, have been noted in the literature for
various species of crayfishes: Ortmann 1906,
Kent 1901, Lereboullet 1851, Harris 1903b,
Newcombe 1929a, b, Evermann and Clark 1920,
Bowman 1942, Penn 1951, Volpe and Penn 1951
(first published study on crayfish genetics), Hand
1954,Waldo 1957, Anonymous 1965,Loeb 1967,
Crocker and Barr 1968, Dowel1 and Winier 1969,
Smiley and Miller 1971, and Momot and Gall
1971 (good summary of literature concerning
blue color mutants). The first breeding experiments involving color mutants were conducted
by Black (1975) with P. a. acutus, and additional
investigations were made by Black and Huner
(1976a,b), Black and Latiolais (1977), and Black
(1977, 1979, 1980). Most of these studies treated
P. clarkii and disclosed that various blue color
variations were due t o genetic mutations
transmitted as simple recessive alleles t o normal
body color, some being known only in females,
and homozygosity for one color (French blue)
allele is apparently lethal in males. Additional
observations of color morphs have been reported
by Clark (1979)) Dunham et al. (1979), Hazlett
et al. (1979a), and Fitzpatrick (1987).
ECOLOGY. Habitats.
Although this species was
reported to occur in Wisconsin as early as 1870
(Hagen), virtually n o ecological information was
presented until 1913. Graenicher (1913)
reported 0. propinquus from lakes and permanent streams, being quite common in the eastern
part of the state, especially in bodies of water
draining into Lake Michigan. Muttkowski
(1918:393) stated that 0.propinquus was
58
restricted to the “stony and gravelly portions”
of the shore of Lake Mendota. Pearse (1910)
reported it from rivers, small brooks, and lake
shores in Michigan, and Engle (1926:97) found
it to inhabit quiet waters in small Nebraska
creeks. He observed it sometimes to make a
shallow excavation but more often during the
day hid in old cans or beneath rocks in the stream
bed. Evermann and Clark (1920) reported that
in Indiana localities, 0. propinquus was found
among rocks and often in Cham beds, and Baker
(1924) stated that it was observed in boulder
habitats in Lake Winnebago. Creaser (1932)
propinquus from Wisconsin lakes
reported 0.
with stony bottoms, from clear streams, and from
dense mats of aquatic vegetation. Hinkelman
(1970) discussed two Wisconsin lakes supporting
populations of 0. Propinquus that were weedy,
silty, and stagnant. Others who noted its
preferred habitat as being rocky portions of lakes
and/or streams are Threinen 195813, Hinkelman
1970, Magnuson et al. 1975 (all for Wisconsin);
Page 1974 for Illinois; Momot 1975 for Michigan;
Pickett and Sloan 1979 for New York; Dean 1969
for the Great Lakes; and Crocker and Barr 1968
and Judd 1968 for Ontario.
Judd (1968) reported that in some Ontario
streams this crayfish was most common on mud
and gravel substrates without vegetation. In
contrast, Terman (1974) noted that it was rare
in mud-bottomed ponds or lakes, that it does
not burrow and cannot cope with fluctuating
water table levels and seasonal drying of habitats;
thus, 0. propinquus was believed by him to be
restricted t o comparatively non-fluctuating
stream or lake environments. Berrill ( 1 9 7 8 ~ 6 9 )
summarized its habitat diversities in southern
Ontario and he and Chenoweth (1982) noted
that juveniles burrow and seal the entrance
whereas large adults are unlikely t o construct
burrows. Nevins and Townes (1935) pointed out
that 0. propinquus was found in polluted areas
of some New York streams and that it was
tolerant of low oxygen concentrations and silt;
Momot (1975)found that well oxygenated waters
were required by this species as did Magnuson
et al. (1975). Berrill (1978) noted that 0.
propinquus was second only to 0. rusticus in its
range o f exploited habitats in Ontario where it
inhabits slow creeks, swift streams, rapids,
shallow eutrophic lakes, and deeper mesotrophic
lakes. Jezerinac (1982) found 0. propinquus in
pools in headwater streams of the Chagrin River
in Ohio where the smallest individuals were
restricted to shallow water and the large crayfish
were in deep pools. Using individuals collected
from southern Ontario, Maude and Williams
(1983) concluded in a laboratory study that this
is a fast-water species, having a mean slip speed
of 34.7 cm/sec but that it showed no strong
preference for mud or gravel substrates. Crocker
and Barr (1968) noted that larger populations
occur in ponds than in rivers in Ontario, and
Page (1985) reported 0. propinquus is the most
common crayfish in clear, rocky riffles of Illinois
streams.
Physicochemical parameters, Park et al. (1940),
Bovbjerg (1952), and Spoor (1955) showed that
heat toleration by crayfishes cannot be compared
concisely but it is apparent that adult 0.
propinquus (and 0.rusticus) have a lower
tolerance to elevated temperatures than do many
other species (e.g., C. diogenes, F. fodiens, and
0. virilis). Bovbjerg (1952) found that those
individuals of 0. propinquus and F. fodiens
acclimated at 18-28°C had a 12-hr median heat
tolerance of more than 35°C. Bott et al. (1973)
noted that 0.propinquus burrows into the stream
bed in the fall when water temperature reaches
10°C and becomes active again when it rises to
10-12°C in mid-April. Crawshaw e t al.
(1980:251) pointed out that thermal pollution
could initiate early development and lengthen
the yearly active period. Gillespie et al. (1977)
noted that when individuals are exposed to tap
water with a cadmium chloride concentration
of 10 ppb, they accumulated a mean cadmium
‘concentration of 18.4 ppb after 190.5 hr. These
researchers observed that where temperatures
in a stream or lake increased due to thermal
pollution, a hastening of the concentrating
process might be expected.
Egg production and hatching time. The timing
of oviposition appears to be related to temperature. Van Deventer (1937) reported egg laying
in early April in central Illinois; however, it is
delayed until early May in populations in
northern Wisconsin lakes where water temper-
atures are lower during most of the year (Capelli
and Magnuson, 1976); further discussion of
ambient temperature and photoperiod on egg
laying in this crayfish follows.
Behavior. An aggressive species (Lunt, 1962),
0. propinquus appears to be active throughout
the die1 cycle (Van Deventer, 1937) with peaks
of activity at night (Capelli, 1975); Dean
(1969:15) reported it t o be most active during
the day. Collins et al. (1982) showed that in
Ontario lakes where bass (predator) densities
were relatively high, 0. propinquus spent more
time in shelters and less time in locomotion than
those in predator-free lakes. Working with
populations of 0.propinquus from Vilas County,
Wisconsin, Stein (1977) showed that within any
size class of adults, life stages demonstrated
differential susceptibility to predators. Recently
molted individuals were the most easily preyed
upon by the smallmouth bass, Micropterus
dolomieui, whereas females with eggs were least
susceptible, Other life stages ordered from low
to high susceptibility to predation were Form
I males, Form I1 males, and females. Thus, life
stages with the greatest reproductive potential
(ovigerous females and Form I males) are least
susceptible t o smallmouth bass predation.
Juveniles, females, and recently molted crayfish
appear to modify their microdistribution and
behavior resulting in reduced predation risk.
Stein and Magnuson (1976) demonstrated the
behavioral response of individuals of 0.
propinquus to predation by M. dolomieui in
laboratory studies. They showed: 1)the predation
rate on sand, gravel, and pebble substrates was
inversely related t o particle size (interstitial
spaces provide refugia from predators); 2) the
response of crayfish seemed to be correlated with
liability to predation (the small, more vulnerable
individuals respond more quickly than larger, less
vulnerable crayfish); 3) the frequencies of various
patterns of behavior of all class sizes were
influenced by the presence of the fish predator
(active behavior patterns such as walking,
climbing, and feeding were suppressed while
behavior that probably reduces vulnerability,
such as chelae display and burrowing, increased);
and 4) juvenile and small adults spent less time
feeding on the detrital food supply when
59
predators were present (fish suppress grazing by
juveniles and small adults). Ameyaw-Akumfi
(1979) demonstrated that during aggressive
encounters with conspecific crayfish, repeated
waving of antennae and movement of mouthparts and the ambulatory appendages by one of
t h e individuals appeared t o be signals of
appeasement. Similar observations were made on
0. widis, P. clarkii, and C. robustus (see also
Rubenstein and Hazlett, 1974). Tierney et al.
(1984) determined that the site of pheromone
reception is the “outer” (lateral) antennule and
‘ I . . . that in 0.propinquus these structures alone
detect the behaviorally active substances or
mediate searching and attraction behavior” (p.
558).
Diet. Crocker and Barr (1968) regarded 0.
propinquus as a generalized feeder, Dean (1969)
reported it to be a vegetarian of lakes and
streams, and Capelli (1980) referred to it as an
opportunistic scavenger. Capelli indicated that
in Trout Lake in Vilas County, Wis. this crayfish
ate diatoms primarily and associated algae, midge
larvae, mayfly nymphs, and other crayfish.
Cannibalism peaked in late June and was related
to increased numbers of newly hatched young
in shallow water. He (1980:85) stated that
seasonal variation in diet is not only related to
the seasonal availability of food items, but
apparently is the result of selection in favor of
some items when they are available.
Field observations. During our field studies we
observed 0. propinquus in clear, to stained, to
turbid streams and lakes that ranged from
moderately swift to virtually no measurable flow
(Figs. 3, 39-43). Individuals were on varied substrates (silt, sand, gravel, pebble, cobble) but were
most commonly found on large particle substrates.
The pH of the waters sampled varied from 5.0
to 7.7 (mean = 6.3), specific conductance ranged
from 76 to 1160 pmhosicm (mean = 370)) and
oxygen concentrations varied from 8.1 (19.oOC)
to 14.1 mg/l (23.3”C) (mean = 10.7 mg/l). In
addition, the relatively abundant aquatidsemiaquatic plants observed were: Carex sp., Equisetum sp., Lemna sp., Mimulus sp., Nasturtium sp.,
Potumogeton sp., Ranunculus acris, Sugittaria sp.,
Typha latifoliu, and Vallisneriu sp.
Summary. 0.propinquus is an aggressive species
that feeds on a wide variety of food items and
is known from an extensive range of environmental situations.
Figure 39. Jackson Harbor, Lake Michigan in Door County;
Orconectes (‘2.1 popinquus and 0. (G.)
virilis collected in
this locality.
60
Figure 40. Namekagon River in Washburn County;
inhabited by Orconectes (C.) propinquus and 0. (G.) virilis.
Figure
42. Kangaroo Lake in Door County; inhabited by
Orconectes (C.) propinquus.
Figure 41. Butternut Creek in Price County; inhabited by
Orconectes (C.) propinquus and 0. (G.)vinlis.
Figure 43. Long Lake in Vilas County; inhabited by
Orconectes (C.) propinquus.
61
LIFE HISTORY. Mating, egg production, and
hatching time (see also above). The life history
of 0.propinquus is fairly well known throughout
its geographical range (see below). The earliest
and one of the best documented studies was by
Van Deventer (1937) in which he described the
life history of this species in a central Illinois
stream. He noted that copulation occurs in late
fall and early spring and remarked (p.33) that
in northern regions, such as Michigan and Wisconsin, copulation probably begins in July and
August, continues until November, but does not
occur during the spring. This has also been shown
for New York populations (Crocker 195750).
Creaser (1933) reported copulation occurring in
southern Michigan only in October and
November whereas Hinkelman (1970) reported
that copulation occurs from August to late
October in Wisconsin. Capelli (1975) found that
in northern Wisconsin mating extends from late
summer, through fall, winter and spring, and
ceases only when females go into seclusion for
egg laying and brooding. Berrill (1978) noted
that females remain hidden and sedentary when
carrying eggs and young. As mating continued
throughout the season, Capelli noted the
percentage of adult females that had mated
successfully gradually increased (determined by
presence of sperm plug) to 50% by December
and reached a maximum of nearly 80% of adult
females by May. Crocker (1957) indicated that
by late fall females in New York populations are
rarely found without a sperm plug. In Indiana
Evermann and Clark (1920) reported copulation
to occur as early as 27 April and also during
late November and early January.
Van Deventer indicated that eggs were laid
in late March or early April and were carried
for 4-6 weeks (dependent on temperature) and
that young hatched in May or June and remained
attached to the mother for 1-2 weeks. Scudamore
(1948) reported eggs hatched in mid-May in
Wisconsin and the young remained attached for
“several days.” Van Deventer noted that the
number of eggs produced ranged from 40 to 250
but he did not state sizes of the females carrying
them. Capelli (1975: 107) indicated the number
of eggs per female ranged from 28 (18mm cl)
62
to 263 (38mm cl), with a mean of approximately
60 eggs per individual. We examined a female
(2 1.3mm cl) collected from the Mississippi River
in Crawford County, 18 April 1981, that carried
60 eggs (egg diameter 2.0mm). In Illinois after
the second molt, the young become freeswimming. Stein and Magnuson (1976) indicated that individuals in Wisconsin are around
4mm cl at this time and the mother molts
immediately following detachment of the young.
Delay of the spring molt by ovigerous females
or those carrying young has been shown to be
regulated by the action of the molt-inhibiting
hormone of the sinus glands (Scudamore 1948).
He also reported that in Wisconsin populations,
these females molt approximately 3 weeks after
the males or non-reproducing females molt, the
latter two beginning around 1 May, with many
molting by 13 May and the remainder before
3 June. Stevenson and Cohen (1965) discussed
various characteristics for recognizing pre-,
inter-, and postmolt stages.
Young of the year. In Illinois (Van Deventer
1937) the young undergo 6 10 molts during
the first growing season and attain a carapace
length of 12-27mm by late September or early
October. Sexual maturity is reached when the
individuals grow to a 20mm cl (exceptions of
15.6mm, 18mm, etc.) and thus many become
sexually mature by their first fall after hatching
(little growth occurs during the winter see also
Creaser 1934a). Berrill (1978) also noted that
some individuals in southern Ontario populations reach maturity at the end of their first
summer. The smallest size of sexually mature
males and females was 16mm cl while the mean
size of both sexes was 23mm cl. Capelli (1975)
reported that the shorter growing season and
colder temperatures during the growing season
in northern Wisconsin prolonged the time
needed to reach maturity; thus, he found no
young-of-year reaching maturity by the end of
the first growing season in Trout Lake. Mating
may occur during the first fall after hatching
in Illinois; these individuals produce a brood and
usually die as yearlings.
Molt cycle. Individuals that were not sexually
active in the fall undergo approximately four
-
-
additional molts and attain a maximum length
of 35-40mm cl during the succeeding summer.
Males are Form I1 during the spring and in
summer (their second), molt to the breeding
Form I (true also of Wisconsin populations
Scudamore, 1948). Females produce a brood and
most die as two year olds. A few individuals,
mostly females, survive to a third year and
produce a brood of young during their third
spring; however, most produce only a single brood
of young in their life span. Fielder (1972)reported
a similar reproductive pattern for 0.propinquus
in Ashtabula County, Ohio.
Stein (197513) and Stein et al. (1977) noted
external morphological changes in primary and
secondary sexual characteristics of both males
and females as they mature and become sexually
active. In addition to the well known changes
in the first pleopods of the male, they noted
that the chelipeds of the first form males become
larger (see below). Mature females have two
pronounced tubercles, on the annulus ventralis
on either side of the anterior groove, whereas
juveniles have annuli that are flat (no relief),
have no tubercles, and no anterior groove. The
external openings of the oviducts appear white
and are covered by a flexible, clear or yellow
convex membrane. The abdomens of adult
females are wider than in juveniles.
Stein and Murphy (1976:2450) reported that
individual proximate composition (e.g., percent
water, percent inorganic material, and chitin)
varied with size, sex, and life stage.
“Understanding how these proximates vary with
life stage is necessary for adequately describing
transfer of energy and biomass between
populations.”
Capelli and Magnuson (1976:418) reported
that temperature is likely more important than
photoperiod in inducing egg-laying. Aiken
(196913) reported that 0. virilis requires a
minimum 4.5 month winter period (darkness and
4°C) for proper ovarian development, but that
the spring stimulus for egg-laying is temperature
and not photoperiod. Van Deventer (1937) also
indicated that temperature is important for the
process to occur. Capelli and Magnuson reported
that a temperature of 7°C was sufficient to
.
induce large numbers of females to produce eggs
in Trout Lake and noted that a few individuals
laid eggs at only 6°C.
Stein (1975a) called attention t o sexual
dimorphism in chelae size of 0. propinquus,
whereby male chelae are larger and heavier than
those of females. He indicated that chelae are
used in defense against predators, for capture
and manipulation of prey, in inter- and
intraspecific interactions, and for reproductive
activities (male male interactions for females
as well as in sexual bouts with females). Stein
(1976) also found that chelipeds constitute 2550% of the dry weight of individuals and that
those of Form I are proportionately twice the
weight of chelipeds of females. In sexual
encounters, males use their chelae to grasp and
hold the female during copulation (Capelli and
Mclntire, 1980, observed the mean duration of
copulation to be 53 min.). Males with large chelae
can interact more successfully with larger (more
fecund) females, thus these males may contribute more genes to future populations than males
with smaller chelae; selection should, then, favor
larger chelae in males (see Fig. 6 in Stein, 1976).
Based on laboratory and field experiments, he
(Stein 1975a) concluded that chelae are most
important for reproductive activity and hypothesized that large chelae are only advantageous
for reproduction and not necessarily required for
other activities.
Hybridization. Tierney and Dunham (1982,
s 0.
1984) demonstrated that 0. ~ r o ~ i n q u uand
virilis from Ontario are able t o distinguish
members of their own species from those of the
other in the laboratory (see section treating 0.
rusticus for discussion). Even though recognition
by crayfishes of conspecific members in mixed
populations has been demonstrated in the
laboratory, interspecific copulation is far from
unusual in natural situations; yet, hybridization
between any crayfish species has not been
conclusively demonstrated. Crocker (1957)
presented morphological evidence for hybridization between Cambarus Tobustus (Rafinesque)
and C. bartonii (Fabricius), between 0. propinquus and 0. limosus (Rafinesque), and between
0. propinquus and 0. obscurus (Hagen) in New
63
York. Dowel1 and Winier (1969) noted that male
0. virilis copulated with 0. immunis in the
laboratory in Iowa. Also Crocker and Barr (1968)
presented data suggesting hybridization between
sympatric populations of 0. propinquus and 0.
obscurus and 0. propinquus and 0. rusticus in
Ontario. Fielder (1972) noted that a male of
0.propinquus and a female of 0. sanbornii from
Ohio successfully mated under laboratory
conditions but did not determine if viable
offspring were produced. Smith (1979) suggested
that hybridization occurs between C. bartonii and
C. robustus in New York. Capelli and Capelli
(1980) conducted studies on 0. propinquus, 0.
rusticus, and 0.virilis from seven lakes in Vilas
County, Wisconsin. Turtle Lake supported
sympatric populations of all three species and
they noted apparent hybrids between 0.rusticus
and 0. propinquus but not among the other
species combinations. Smith (1981) commented
on hybridization among cambarids and presented
evidence that hybridization may occur between
~
0. rusticus and 0. limosus in Massachusetts.
Jezerinac (1982), working in the Chagrin River
in Ohio, observed individuals that he considered
t o be possible hybrids between 0.propinquus and
0.rusticus.
Life history data are summarized for 0.
propinquus in Table 7.
DISTRIBUTION: Even though 0. propinquus is
known from every major watershed in the state
(Appendix I), it is certainly not evenly distributed
(Fig. 44). Since Creaser's report (1932) the range
of this species has been extended considerably
over the northern tier of counties, yet its absence
is particularly evident in the lower St. Croix,
lower Chippewa, and the central Wisconsin
drainages. Only a few occurrences are known
in the Trempealeau-Black watershed. Total
geographical distribution is shown in Fig. 45.
CRAYFISHASSOCIATES. In Wisconsin, 0. propinquus was found in company with C. diogenes,
0.rusticus, and 0. virilis.
~~~~~~
TABLE 7. Life history data for 0. propinquus (* from references only; ** from literature and
from our collections or USNM collections; those without asterisk(s) from USNM or OSU
collections only).
Jan.
Feb.
Mar.
**Wis.
**Wis.
.
"Wis
"Wis.
"Wis.
May
"Wis.
"Wis.
"Wis.
'Wis.
Jun.
'Wis.
"Wis.
"Wis.
"Wis.
JuL
"Wis.
"Wis.
'Wis.
Aug.
"Wis.
"Wis.
'Wis.
Sep.
Wis.
Wis.
'Wis.
Oct.
Wis.
"Wis.
Nov.
"Wis.
'Wis.
Apr
Dec.
64
s9
Figure 45. Geographic distribution of Orconectes (C.) propinquus.
Orconectes (Procericambaw) rusticus (Girard)
(Figures 46 - 51)
-
Cambarus rusticus Girard 1852:88.
Orconectes rusticus
*Lorman 1975:i,ii, 1-56;
Magnuson et al. 197567-72; Capelli 197544,
48, 50, 53, 58-61, 161, 188, 198-201, 206,
207; Stein and Magnuson 1976:759;Bouchard
1978:16; Huner 1978:2-4; Lorman and
Magnuson 1978:8; Capelli 1978:59; Momot
et al. 1978:18; Payne 1978:7;Sheffy 1978:222;
Phillips and Reiss 1979:18; Stein 1979:345;
Capelli and Capelli 1980:121- 132; Lorman
198O:l-227; Gallepp and Lorman 1980:18;
Smith 1980:1,2,3; Boronow 1981:1, 2, 8, 9;
Kienitz 1981:9; Boronow 1982:1,3; Capelli
1982a:741-745; Tierney and Dunham
66
-
1982547; Threinen 1982a:78; Maude and
Williams 1983:76; McBride 1983:42; Stamm
1983:23; Capelli and Magnuson 1983548,
549,551-554,557,559-561,563,564; Dodson
and Cooper 1983:347, 350; van Goethem
1984; Momot 1984a:51; Lodge 1984:61;
Capelli and Hamilton 1984:251-259; Hayes
1985:2; Magnuson and Beckel 1985:lO; Lodge
et al. 198533; Butler and Stein 1985168, 169;
Page 1985412, 413; Berrill 1985347, 348;
Lodge et al. 1986, Jezerinac 1986:179; Lodge
1986:4; Lodge and Lorman 1987591-596.
Orconnectes rusticus Threinen 1982b:3.
oronectes rusticu - Lang 1977:lO.
orconectes rusticus McBride 1983:42,44,45.
rusticus - McBride 1983:46; Stamm 1983:23-25.
Rusty crayfish - Anonymous 1982a; McBride
-
1983:42-46; Hacker 1983:31; Pivar 198359;
van Goethem 1984; Magnuson and Beckel
19859; Lodge et al. 198532-37; Butler 1986:4.
Exotic species Leys 1980:8.
-
*
All previous records for 0. rusticus in
Wisconsin were published errors in determinations or repetitions of the original error.
DIAGNOSIS (refer to Table 8): Carapace (Fig.
46c,j) subovate, depressed dorsally; rostrum
longer than broad, deeply excavate dorsally with
distinctly concave margins; marginal spines acute
to reduced; acumen triangular tapering to acute
apical, slightly upturned spine; postorbital ridge
short, terminating anteriorly in short, acute to
blunt spine. Cervical groove deep, somewhat
sinuous, and discontinuous laterally. Distinct but
generally short cervical and branchiostegal spines
present; without hepatic spine. Areola (Fig. 46j)
narrowest near midlength with three or more
punctations in narrowest part; areola constituting 31.7 to 44.4% (mean 34.9%) of total carapace
length (39.9 to 64.8%, mean 44.2%, of postorbital carapace length) and 3.8 to 11.0 (mean 6.9)
times as long as broad. Antenna1 scale (Fig. 468)
2.3 times as long as broad, widest at about
midlength, distal portion tapering, acute.
Cephalic lobe of epistome (Fig. 46h) slightly
broader than long, rounded, with unevenly
concave margins. Mandible (Fig. 46k) with
straight, sharp, corneous edge on distal incisor
margin. Bouchard (1977a) observed that masticatory surfaces of mandibles of Holarctic
crayfishes are useful characters for assessing
relationships because similarities in structure of
the mandibles are indicative of phylogenetic
homogeneity among certain genera and subgenera. Conclusions based on mandible form agree
with previous assessments based on secondary
sexual characters. Abdomen shorter than
postorbital carapace length; cephalic section of
telson with one large (lateral) and two small
(mesial) spines in each caudolateral corner, distal
podomere of lateral lobe of uropod rounded;
mesial ramus of uropod with distomedian spine
reaching distal margin; one small (lateral) and
one large (mesial) spine situated distolaterally.
Chela (Fig. 46m) moderately punctate, slender
(approximately 2.3 times as long as broad), only
moderately tuberculate along mesial margin of
movable finger and palm; opposed margins of
fingers lacking prominent tubercles; mesial
surface of palm with 7 (range 6 to 8) distinct
(mesialmost) tubercles. Carpus of cheliped
moderately cleft dorsally with one prominent
spine on mesial surface; 2 to 3 median ventral
spines. Ischium of third pereiopod only (Fig. 46
1) with proximal hook extending slightly over
basioischial articulation. First pleopods of first
form male (Fig. 46a,f) symmetrical and reaching
coxae of second pereiopods with abdomen flexed;
terminal elements of first pleopod of male (Fig.
46a,b,e,f) subparallel, slender, tapering distally to
acute tips; corneous, spiculiform central
projection longer than mesial process; cephalic
surface with distinct shoulder at base of central
project ion.
Annulus ventralis (Fig. 46d) distinctly broader
than long and with caudal margin of nontuberculate sternum anterior to it overhanging its
lateral margins; ventral face bearing deep
transverse depression across midlength; postannular sclerite movable, approximately two-thirds
as broad as annulus (annulus of juvenile female
shown in Fig. 46i).
VARIATION: Being a widely distributed species
probably introduced into Wisconsin from a
number of different localities, it is not surprising
to find inter- and intrapopulation variations.
Differences in color are treated under “Color
Notes.” T h e most obvious morphological
variations occur in the absence (typical) or
presence (atypical) of a rostral carina and in the
relative length of the mesial process and central
projection of Form I male pleopods. Generally,
rostral carinae occur only in 0. propinquus in
Wisconsin. However, occasional individuals (Fig.
47) have well to poorly developed carinae (see
also Capelli and Capelli, 1980).Crocker and Barr
(1968:42) mentioned atypical characters demonstrated b y 0. rusticus where it occurs
sympatrically with 0. propinquus in Ontario: 1)
presence of rostral carina, [If indeed hybridization does occur between 0. rusticus and 0.
propinquus in the state, it appears this is one
feature that will be characteristic of the hybrids.
67
TABLE 8. Range of measurements (in mm) of various diagnostic structures of Wisconsin
Orconectes rusticus.
N
Minimum
Maximum
Mean
Standard
Deviation
-t
MALES ($1)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
Pleopod
Total length
Central Projection
Mesial Process
Condyl length
-
30
31
31
24.2
18.8
8.6
49.0
38.8
18.9
32.0
25.6
12.2
6.5
5.4
31
31
7.9
0.9
17.6
2.7
11.2
1.6
2.4
0.5
31
31
4.9
7.5
16.9
25.0
8.5
12.9
2.9
4.2
30
31
31
30
10.2
3.7
3.2
9.2
18.7
7.8
6.1
17.0
13.5
12.0
2.4
1.0
0.7
2.1
25
27
19.4
10.5
27
7 .O
46.5
38.4
17.0
31.4
24.6
11.6
7.2
6.7
2.9
26
27
6.6
0.6
17.1
3.0
10.9
1.6
2.8
0.6
24
24
2.6
4.2
12.9
15.9
6.8
9.7
2.8
3.7
61
62
62
18.5
14.5
7.0
44.5
35.4
18.5
29.7
23.8
11.6
5.4
4.5
2.3
62
62
6.3
0.8
15.1
2.8
10.4
1.7
2.0
0.5
57
57
1.8
5.9
11.3
15.5
6.0
9.2
1.8
2.4
5.0
4.2
2.7
MALES ($11)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
-
FEMALES
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
-
68
f
Figure 46. Orconectes (P.) rusticus (d, adult female Silver Lake, Forest County; i, juvenile female South Turtle Lake,
Vilas County; b,e, second form male Lake Metonga, Forest County; all others first form male from Lake Metonga, Forest
County): a,b, mesial view of first pleopod; c, lateral view of carapace; d,i, annulus ventralis; e,f, lateral view of first
pleopod; g, antenna1 scale; h , eplstome; j , dorsal view of carapace; k, incisor margin of right mandible; 1, proximal podomeres
of third, fourth, and fifth pereiopods; m, dorsal view of distal podomeres of cheliped.
69
Figure 47. Photograph of atypical Form I1 male Orconectes (P.) rusticus from Otter Lake, Langlade County (note rostra1
carina).
:;I
Figure 48. First pleopod of first form males of Orconectes (P.) rusticus (a-c, lateral view; ,d-h, mesial view): 3,' Twin Bear
Lake, Bayfield County; b,e, South Turtle Lake, Vilas County; c,d, Long Lake, Chippewa County; f , Upber Sugarbush
Lake, Vilas County; g,h, Manitowish River, Vilas County.
70
We do not, however, speculate that all “0.
rusticus” individuals possessing a carina are
indeed hybrids; much study is required concerning hybridization of crayfishes.] 2) reduction of
the right-angle shoulder on the anterior margin
of the gonupods of Form I males, 3) reduction
of the median ventral carpal spine, and 4)
lowering and separation of the anterior tubercles
of the annulus ventralis. Variation in the first
pleopod of Form I males is shown in Fig. 48.
Generally, the 1:3 ratio of central projection
length to total gonopod length is an adequate
measurement; however, many values, both
higher and lower, were noted. In addition, the
relative length of the central projection and
mesial process were variable (Fig. 48a-h), with
some gonopods more closely resembling those
of 0. propinquus or 0. uirilis. Also, typical 0.
rusticus specimens have a pronounced “shoulder” on the gonopod (Fig. 46a,f), yet many
specimens either lack this or have a distinctly
sloping shoulder (Fig. 48h); see also Capelli and
Capelli, 1980. We also noticed that in some
localities the carapace appears slightly more
compressed dorsoventrally.
The mesial margin of the palm has two distinct
rows of tubercles but the number of tubercles
on each row is highly variable: mesialmost row
generally from 7-10 but occasionally with as few
as 5; this is particularly variable for regenerated
chelae.
Bouchard (1976a:581) stated that 0. rusticus
bears a straight, sharp corneous edge on the
incisor portion of the mandible. He also
presented a detailed discussion of the mandible
in Holarctic crayfishes, noting again the bladelike incisor of 0. rusticus (1977a). The majority
of specimens we examined possessed the typical
form of the incisor (Fig. 46k); however, some
specimens had the blade-like edge interrupted
by crenulations.
Lorman (pers. comm.) also noted behavioral
differences among populations, indicating that
in Chippewa County groups of individuals often
are found clinging to each other; this is certainly
unusual for the agonistic 0. rusticus. Based on
morphological and behavioral differences
between populations in Chippewa County and
and those in other areas, he (1980) suggested
two separate origins for this species in Wisconsin.
See Capelli and Capelli (1980) for additional
discussion of variations among 0. msticus, 0.
propinquus, and 0.uirilis.
COLOR NOTES (refer to Fig. 49): Ground color
of carapace reddish-brown to reddish-tan,
“rusty,” fading ventrally o n hepatic and
branchiostegal regions to tan or cream; carapace
sometimes speckled with small tan or grayish
splotches of color; posterior region of cephalic
section between mandibular adductor muscles
with black splotch, sometimes U-shaped; dorsal
surface of thoracic section usually dark reddishbrown or gray, posterior part sometimes with
black saddle; posterolateral portion of carapace
(branchiostegal region) usually with distinct spot
of red or rust-brown, shape, size, and location
of spot quite variable (Lorman, pers. comm.,
reported specimens from Chippewa County with
crescent shaped spot); parts of antenna1 and
mandibular as well as orbital and hepatic regions
of cephalothorax often with dark red to reddishbrown stripe or band. Abdomen dark reddishbrown; pleura slightly lighter in color and
occasionally with small red dot on each. Telson
and uropods lighter shade of reddish-brown to
tan. Cheliped grayish-green, pink, or light
reddish-brown dorsally, often mottled with
varying shades of brown; dorsomesial portion of
merus dark gray or black; anterolateral portion
of merus sometimes with prominent longitudinal
“rusty” band; ventral surface pink, tan, or
sometimes cream; distinct subterminal black
band on fingers of chelae; apex of fingers usually
bright red, sometimes orange or scarlet.
Pereiopods 2-5 grayish-green, tan, or very light
reddish-brown, tips red to orange. See Capelli
and Capelli (1980) for further notes on color
variation.
TYPE-LOCALITY:
Ohio River at Cincinnati,
Hamilton County, Ohio, USA.
ECOLOGY.
Hubitats. Much conjecture has been
associated with the presence or absence of 0.
rustus in Wisconsin waters prior to recent
introductions. Creaser (1932:335) discounted
previous reports as being based upon “aberrant
71
Figure 49. Photograph of Form I male Orconectes (P.) rusticus Plover Creek, Marathon County (note regenerated right
chela and large left chela, latter typical of breeding males).
forms of C. propinquus,” and no validated reports
of the species in Wisconsin occurred prior to
1975 (Lorman 1975, Capelli 1975). As noted
below, this crayfish may have been introduced
into the state 30-35 years ago, presumably by
fishermen.
Orconectes rusticus inhabits a wide variety of
aquatic situations, including both lotic and lentic
systems. It has been found in both hard and
soft waters, in lakes as well as streams. Generally
it resides beneath or among rocks, logs, or other
cover on the substratum] which varies from clay,
silt, and sand to cobble and boulder, as well as
among vegetation. Creaser (1931b) noted its
occurrence in Michigan in small streams having
gravel and stone substrates. Rhoades (1944a)
collected 0.rusticus from swift, rocky bottomed
streams in Kentucky and reported a relationship
between the occurrence of limestone bedrock
and the distribution of this species; he also
recorded 0.rusticus in southern Ohio in streams
flowing through areas of limestone exposures
(1944b, 1962b). Others to report this species
72
from rocky-riffle areas in streams or from stony
lakes are: Crocker and Barr, 1968 (Ontario); Page
and Burr, 1973 (Missouri); Terman, 1974
(Michigan); Magnuson et al., 1975 (Wisconsin);
Bouchard, 1976a (Tennessee); and Pickett and
Sloan, 1979 (New York). Phillips and Reis (1979)
observed it in deep sand-bottomed pools in
headwater streams and Jezerinac ( 1982)indicated
that this was the dominant pool-dwelling crayfish
in the Chagrin River basin in Ohio. Maude and
Williams (1983) indicated this is a fast-water
species, having a mean slip speed of 40.2 cm/
sec; they suggested that this high slip speed may
play an important role in aiding it to expand
its range in areas where it has been introduced.
Turner (1926) and Eberly (1955) noted the
occurrence of 0. rusticus in ponds as well as
lakes, and Langlois (1935, 1937) collected it from
fish ponds in Ohio where it also burrowed in
the soil around the ponds. He (Langlois
1935:189-190) noted that in dense populations,
“. . the burrows may contain closely packed balls
of crayfish] one of which contained 67 individ-
.
uals.” Prins (1968) also noted that it burrowed
in Kentucky and Berrill and Chenoweth (1982)
indicated that in the laboratory juveniles burrow
and seal the opening but that adults are unlikely
to demonstrate fossorial behavior; also McMahon and Wilkes (1983), Phillips (1979) in Iowa,
and Momot (1975) in Michigan noted no
burrowing activity but Phillips (op. cit.) reported
that individuals made excavations under flat
stones in streams. Phillips and Reis (1979) and
Claussen (1980)found 0.rusticus in warm water
streams with rock bottoms where it most
frequently was observed in shallow “hides” in
riffle sections of Minnesota streams, as did Prins
(1968) for juveniles in a Kentucky stream.
Lorman (1975, 1980) did not observe 0.rusticus
burrowing in or along the banks of northern
Wisconsin lakes nor did we encounter individuals
burrowing in any lotic or lentic habitats sampled.
Prins (1968), Phillips and Reis (1979), Clark and
Rhoades (1979), and Smith (1980) reported 0.
rusticus inhabiting areas of streams having
vegetation and several studies have shown that
it feeds on aquatic plants and that it is responsible
in some areas for the dramatic decline in
abundance of these plants (Magnuson et al.,
1975, Lorman 1980, McBride 1983).
Lorman (1975) noted that there was a
tendency (not statistically significant) for
individuals to shift from animal to plant food
as they grow larger. He indicated that adult males
display little die1 periodicity in activity, measured
by stomach fullness in the field, whereas adult
females and juveniles exhibit strong nocturnal
activity patterns; that is, they tend to forage
only at night. Stein (1979) suggested that this
foraging pattern, active primarily at night, could
be interpreted as a size-mediated response
directly related t o its vulnerability to fish
predators. Lorman (1975) also noted that adults
tend to feed more during the day in the summer
but restrict their activity to the night-time
during the colder months.
Physicochemical parameters. Claussen ( 1978)
demonstrated that 0.rusticus had a high thermal
tolerance and generally made a rapid acclimation
response to increased temperatures. Layne et al.
(1985), using 0. rusticus from Indian Creek in
Butler County, Ohio, investigated the critical
thermal maxima and minima (CTMax, CTMin).
They found that 0.rusticus CTMax and CTMin
were higher in summer than winter throughout
most of the low to high temperature time course.
Additionally, the seasonal adjustments of the
time courses of thermal acclimation enhance the
ability of 0. rusticus to compensate for changes
in its natural environment. A significant positive
correlation with increased temperatures and
oxygen consumption was shown (Eggleston and
Lustick 1981), yet there were no significant
differences in oxygen uptake between starved
and fed crayfish or between males and females.
Juberg (1982) indicated that bilateral sinus gland
cauterization (molt induction) resulted in an
increase in the oxygen consumption of males
(up as much as 107% as compared with male
specimens at intermolt stage) and he noted an
inverse correlation between oxygen consumption
and body weight of both males and females. He
proposed that injury alone may cause the crayfish
to increase its oxygen consumption as a reaction
to a stressed situation. Scudamore (1947),
however, showed that normal (non-induced)
crayfish have increased oxygen consumption
levels during ecdysis.
St. John (1982) noted that this crayfish is
relatively tolerant of an environmentally stressed
(channelization, industrialization, urbanization)
stream in southwestern Ohio. Winner et al.
(1980:47) showed that “. . chemical species of
copper other than the cupric ion, can be
absorbed and assimilated by the crayfish.”
Hubschman (1966) stated that there is no tissue
damage at high concentrations (up to 10.0 ppm)
of copper; however, he indicated that long exposure to low concentration led to the degeneration of cells of the antenna1 glands. He (1967)
also showed that when exposed to concentrations of copper greater than 1.0 mg/l, respiratory
enzymes are inhibited quite rapidly. He demonstrated the effects of copper contamination
on enzyme systems by utilizing one Krebs cycle
metabolite (succinate); succinate utilization is
inhibited with increasing copper concentration
( > 0.5 mg/l) long before endogenous oxygen
uptake is altered. At concentrations of 0.5 mg/
.
73
1 or less, neither succinate utilization nor
endogeneous respiration is measurably affected;
however, low concentrations do impose a chronic
effect; cell repair cannot keep pace with cell
activity (e.g., cells of the antenna1 gland).
McMahon and Morgan (1983) conducted a
laboratory study on individuals from southern
Ontario and found that in acute exposure (96hr),
crayfish are substantially more acid tolerant
(H2S04) than most fishes (LC50 for 0. rusticus
at pH 2.5). They also noted that physiological
responses to sublethal exposure (4 days at
ambient pH 3.8) resulted in the development
of severe hemolymph acidosis (see also Wood
and Rogano 1986).
Summary. 0. rusticus, in Wisconsin is an
exotic, aggressive, tolerant species that has been
extremely successful in the variety of habitats
into which it has been introduced. Not only is
it capable of living in soft-bottomed lakes and
pools (Fig. lo), but it is also quite successful in
rubble-bottomed lakes (Fig. 56a) and swift
streams (Fig. 3), with or without weed beds. We
collected it from waters varying from no flow
to moderately swift current velocities and in clear
to turbid waters. Specific conductance ranged
from 71 to 601 pmhodcm (mean = 193 pmhosl
cm) and oxygen concentrations varied from 7.0
(19.8"C) to 14.1 mg/l (23.3"C) (mean = 9.3 mg/
1). In addition, the following aquaticlsemiaquatic plants were common in the habitats
where 0. rusticus was found: Chara sp.,
Cladophora sp., Elodea sp., Lemna sp., Myriophyllum sp., Najas sp., Nuphar microphyllurn,
Pontederia cordata, Potamogeton amplifolius, P.
angustifolius, P. crispis, P. pusillus,' Spirea alba,
Spirogyra sp., Typha sp., Vallisneria americana,
and Zinnichellia palustris. Where 0. rusticus has
become established it has generally had considerable impact on the community it invaded.
Much is still lacking in our understanding of its
success over native crayfish species in areas where
it has been introduced. For additional ecological
data pertaining to this species in Wisconsin, see
Lorman (1975, 1980), Lodge et al. (1985), and
Lodge et al. (1986).
LIFE HISTORY. Mating, egg production, and
hatching time. Prior to the work by Lorman
74
(1980) the only detailed descriptions of the life
history of 0. rusticus were those of Langlois
(1935, 1937) which were based on populations
in Ohio and by Prins (1968) on a stream (Doe
Run) population in Kentucky. Langlois (1935)
reported the principal mating season to occur
during September and October (see also Berrill
and Arsenault 1984). Immediately following
copulation, females burrowed into the pond
banks a t t h e water line (burrows nearly
horizontal and no more than 2.5 feet back) where
they remained until the following spring. Most
of them waited until spring to oviposit (also see
Williamson 1899, in Ohio, and Williamson 1907,
in Indiana); however, some individuals deposited
eggs in late October (Langlois 1937 and Busch
1940),where they presumably carried their eggs
over winter (mean number of ovarian eggs being
276, greatest number, 574). Langlois (1935)
noted the number of eggs laid correlated
positively with the size of the female and that
eggs carried by females during the winter hatched
as early as 14 March. Most females waited until
April or May when approximately 20 days after
laying, the eggs hatched; following 3- 12 days,
t h e juveniles became free-living. Rhoades
(1944a) also reported young attached during
April and May. Most of the young-of-year
became sexually mature at the end of the first
summer and participated in the fall mating
period. Prins (1968) found that in Kentucky
mating occurred primarily in September and
October but also in February. Females with eggs
were observed in February t o June. He noted
that the number of ovarian eggs was directly
related to body size and that the number of
abdominal eggs was less than the number of
ovarian eggs: ovarian eggs ranged from 54 357
and the number of abdominal eggs varied from
42 231. Busch (1940) noted that mature eggs
were approximately 2mm in diameter. Prins
indicated it took 4-6 weeks for the eggs to hatch,
and Busch (op. cit.) found that only 20 days
were required at room temperature, both
considerably longer periods than that reported
by Langlois.
Young of the year. Prins observed that once
hatched, the young stayed with the mother
through 4-5 molts and left when they were 4-
-
-
5mm long, remaining in shallow areas along the
banks or in gravel beds. Those that hatched in
late May or early June attained 6-llmm during
the June to July period. A few large males became
Form I and very few females possessed large
oocytes. Most individuals spent the winter at
lengths of 10-17mm with no growth occurring
(Prins reported that at temperatures of 6-8°C
they become sluggish, at 4 degrees torpid, and
that molting occurred at 10-12°C or lower, but
stopped if above 12 degrees).
Sexually mature foms. By their first spring they
ranged from 10-19mm (in April), were 12-19mm
long in May, and 14-22mm by June, Generally
sexual maturity was attained at a carapace length
of 18mm; however, the smallest Form I male
had a carapace length of only 16mm and the
smallest mature female, 17mm cl. Most of these
matured by September and mated during
September and October, males molting to Form
I during July, August, and September, although
Form I and I1 males were present during all
months. Sadewasser and Prins (1973, 1979)
reported varying effects of temperature, photoperiod, and light intensity on the molting
frequencies of 0. rusticus. By September
individuals ranged from 19-27mm and underwent no additional growth during late fall and
winter. The following May many molted again
and ranged in length 20-30mm. By May, females
oviposited, some Form I males died or molted
to Form 11, and spent females died. Highest
mortality occurred from March t o May and
resulted from predation, “old age,” and from the
molar action of stones and rubble during periods
of high stream discharge. Most surviving
individuals ranged from 24-32mm by July, and
had attained a carapace length of at least 30mm
by September. They overwintered and by the
following spring reached a carapace length of
40mm (largest female .44mm). During the third
spring and summer nearly all individuals died,
maximum life span varied from 2.5-3 years.
In Iowa populations, eggs are laid in April and
juveniles were present in late May (Phillips 1980).
Berrill (1978) noted that in southern Ontario
the smallest sexually mature males and females
had a carapace length of 17mm and that the
mean size of sexually mature individuals was
considerably larger, 28mm. He and Arsenault
(1982) presented a discussion of the life history
of 0. rusticus in southern Ontario. They
indicated that when waters reached 4“C, males
and females became sexually active, primarily at
night, and that males were very aggressive and
fought with each other as well as separated
copulating pairs. Eleven to twelve days after
copulation eggs were extruded (late April).
Following this period of copulation, males spent
less time fighting and more time feeding, and
females wandered much less, devoting much of
their activity to feeding. The authors suggested
that there may be an interaction between
photoperiod and temperature in controlling the
onset of egg extrusion. Eggs hatch and young
remain attached until they reach stage 111, when
they disperse by mid-June. This is followed by
a molt by the mother. Shortly after copulation,
males molt to Form 11. They molt again to Form
I during the summer, thus mating can occur in
spring and late summer in southern Ontario.
Population dynamics. Houp and Kuehne (1980)
summarized the population dynamics of 0.
rusticus in a central Kentucky stream. Young are
recruited into the population in late May, the
natality rate being 4O/adult female/year. They
have an expected life span of 2.5 years and reach
a maximum density of 2.6 individualdsquare
meter.
Butler and Stein (1985) investigated mechanisms governing replacement of native 0.
sanbornii by 0. rusticus in Ohio. They found
that the two species had similar life histories,
habitat preferences, and feeding habits in
allopatric and sympatric stream areas, yet 0.
rusticus replaced 0. sanbornii, They concluded
that 0. rusticus probably maintains greater
population growth because: more gravid 0.
rusticus females occurred in sympatry, 0.rusticus
produced more young than 0.sanbornii, and 0.
rusticus young-of-year grew faster. Reproduction
interference coupled with differences in aggressive dominance and young-of-year susceptibility
to predation were suggested as probable major
regulating mechanisms in the replacement of 0.
sanbornii by 0. rusticus in Ohio streams.
75
Recent field studies. A detailed study of the
life history of 0.rusticus in Wisconsin conducted
by Lorman (1980) is summarized below. Data
presented were primarily from a population in
Upper Sugarbush Lake (2.5 ha) Vilas County
where overall sex ratios were approximately
5050and total population estimates ranged from
672,000 in June 1978,to 774,000 in August 1977,
and 1,275,000 in July 1978. The mating season
begins in late summer, when males molt to Form
I, and extends through fall into winter. Berrill
and Arsenault (1984) studied a natural stream
population in southern Ontario and noted this
species has a promiscuous mating system
characterized by intense intermale aggression.
They also suggested that female choice may occur
in the breeding process. Capelli and McIntire
(1980) indicated that 0. rusticus exhibits the
shortest mean mating duration (40 min.) when
compared with those of 0. propinquus and 0.
wirilis. Sperm are stored in the seminal receptacle
of the female following copulation until the time
of egg-laying in spring. Females overwinter in
deep water but move into the littoral zone and
extrude their eggs very close to the time of ice
thaw. The earliest record of ovigerous females
is 19 April but most females lay eggs during late
April and early May. Not all females successfully
reproduce (of 1000 mature females examined
during 1976-1978, 82.1% bore eggs or young).
Females generally carry 50-200 eggs (linear
relationship between female cl and number of
eggs; also, inverse relationship between number
of eggs for female cl and individual egg weight).
Mean hatching dates were 7 June in 1976, 22
May in 1977 (warm spring), and 5 June in 1978.
Newly hatched young remain with the female
and undergo three or four molts prior to
becoming independent. Growth is highly
variable and may be controlled by densitydependent factors (availability of food resources).
Males generally undergo 8 molts before reaching
maturity and females have a similar growth-molt
pattern but fewer molts. The molting frequency
for both sexes decreases with the drop in
temperature in September. Total seasonal
growth ranges from 5.18 to 12.64mm; within
an age group the males are usually larger than
76
the females (particularly apparent in older age
groups). The growth increment at molting
generally decreases with increase in size and age
of the individual; growth of chelae at maturity
molts, particularly males, is apparently made at
the expense of growth in carapace length.
Survival is very low for newly hatched young,
only approximately 4% remaining two months
following hatching. Approximately 3% of the
estimated number of hatched young survive to
become mature at the end of the second growing
season and only about 20% of these survive and
reproduce a second time. Lorman also noted an
inverse relationship with survival of young-ofyear with increasing densities.
Some young-of-year may become sexually
mature at the end of their first summer, but
most are not mature until the following spring.
Maximum life expectancy is 3-4 years.
Lorman noted densities are much higher in
areas where rocks are abundant, and the largest
individuals observed are abundant in areas
covered with logs. Densities are highest in littoral
zones (up to 400,0001ha or 2-4 times as high
as densities of 0.wirilis) in three Michigan lakes
(Momot and Gowing 1977a). Lorman computed
the crayfish biomass for the entire lake at 10.4
kg/ha, with the littoral zone supporting 27.0 kg/
ha; biomass was higher where there was a hard
rocky substrate (39.6 kg/ha). 0. rusticus has a
mean caloric mass of 3.4 kcal at maturity.
Life history data of Wisconsin 0. rusticus are
summarized in Table 9.
DISTRIBUTION:
The distribution of 0. rusticus
in Wisconsin is disjunct (Fig. 50) and encompasses all major drainage basins except the
Trempeleau-Black (see Appendix I). T h e
occurrence of this species in the state is
“artificial” and closely approximates the location
of lake districts and/or areas of dense human
populations. The known geographical distribution is shown in Fig. 5 I.
CRAYFISH ASSOCIATES.In Wisconsin 0. rusticus
has been collected together with C. diogenes, 0.
immunis, 0.propinquus, and 0.uirilis.
TABLE 9. Life history data for 0. rusticus (* from references only; ** from literature and
from our collections or USNM collections; those without asterisk(s) from USNM collections
only).
Jan.
Feb.
Mar.
Apr.
"Wis.
'Wis.
June
Wis.
"Wis.
July
Wis.
Wis.
Aug.
Wis.
Wis.
Sept.
Wis.
Wis.
"Wis.
'Wis.
**Wis.
'Wis.
'Wis.
Oct.
Nov.
Dec.
77
I
O
W
A
Figure 50. Distribution of Orconectes (P.) rusticus in Wisconsin; closed circles-specimens
circles-from literature.
78
examined in this study; open
Figure 5 1. Geographic distribution of Orconectes (P.) rusticus.
Orconectes (Cjremicumburus) virilis (Hagen)
(Figures 52 - 62)
Cambarus virilis Hagen 1870:63.
Cambarus virilis - Hagen 1870:64, 65, 97, 101;
Bundy 1876:4; Forbes 1876:4; Bundy
1882:177, 178, 181; 1883:402, 403; Faxon
1884:147, 148; 1885b:7, 96, 97, 98; Underwood 1886:374; Forbes 1888:474; Hay
1896500; Harris 1903a:59, 134, 150, 155;
Fasten 1914587, 601, 621, 623-627; Muttkowski 1918:393; Ellis 1919:257; Pearse and
Achtenberg 1920:312; Pearse 1921:41;
1924:255, 256; Turner 1924:264; 1926:154,
156, 157, 158; Creaser 1931b3263; 1932:322,
323,324,326,327,330,331,335,336;Turner
1935465, 867, 868, 870, 871, 872, 873, 874,
876, 877, 881; Walters 1939:28, 110, 168;
Goodnight 1940:34,35; Roberts 1944:376 [by
implication]; Stephens 1952:70-83; Hinkelman 1970:7.
Cambarus viriles Bundy 1882:180.
Cambarus debilis - Bundy 1876:24,25 [type, MCZ
3449 male I1 (type-locality, Baraboo River,
Ironton, Sauk County, Wisconsin)]; Forbes
1876:25; Bundy 1882:178, 181; 1883403;
Faxon 1884:147, 148; 1885b:6,7; Underwood
1886:374; Creaser 1932:335.
Cambarus rusticus - Bundy 1882:178, 181;
1883:402;Faxon 1884:148; 1885b:7,108, 110,
113; Underwood 1886:372; Harris 1903a: 60,
122, 150, 155; Faxon 1914:418; Creaser
1932:335; Walters 1939:27.
-
79
Cambarus (Faxonius) rus ticus - Graenicher
1913:118.
Cambarus (Faxonius) rusticus rusticus - Ortmann
1931:83.
Cambarus wisconsinensis - Bundy 1876:4 [type,
MCZ 3448 male I1 (type-locality, Racine,
Racine County, Wisconsin)]; Forbes 1876:4;
Bundy 1882:178, 179, 181, 182; Underwood
1886:372; Faxon 1914:418; Creaser 1932:335;
Hobbs 197410342. (This was a name Faxon
-
believed to be a synonym of 0.rusticus; Hobbs,
197413, synonymized it with 0. uirilis.)
Cambarus Wisconsinensis Bundy 1883:402,403;
Faxon 1884:148; 1885b:6, 7.
Cambarus cousii??* Bundy 1883:402.
Cambarus (Faxonius) virilis
Graenicher
-
-
1913:118, 119, 120, 121.
Faxonius uirillis Penn 1943:7.
Orconectes virilis - Penn 1950:646, 648; Crocker
195759, 72; McWhinnie and O’Connor
1967:132; Momot 1967b:69; Thompson
1967:47,49,50,53 [by implication]; Crocker
and Barr 1968:28, 46, 47, 92, 95 (fig. 76);
Barr 1969:93; Caldwell and Bovbjerg
1969:463, 469; Heckenlively 1970:180, 181;
Hinkelman 1970:7, 8, 29, 33, 35, 36, 44, 45,
47; Avault 1971:6; McWhinnie e t al.
1972:357-372;Momot and Gowing 1972:482,
483; Weagle and Ozburn 1972:366; Avault
1973:244; Hart and Hart 1974:120, 138;
Magnuson et al. 197567, 70, 71, 72; Capelli
1975:2, 12, 39, 40, 44, 46, 48, 50,54.58, 60,
61, 63, 64, 71, 72, 79-83, 95, 106, 111, 112,
151, 199, 200, 206, 207; Momot and Gowing
1976:39; Capelli and Magnuson 1976:416;
Bouchard 1978:16; Capelli 197859; Huner
1978:2, 3,4; Momot et al. 1978:15, 18;Sheffy
1978:2 19-225; Unger 1978:3; Crocker
1979:235,244; Boronow 1980:l; Capelli and
Capelli 1980: 121-132; Claussen 1980:377,
379-383; Phillips 1980:88, 89; Horns and
Magnuson 1981:299-302; Boronow 1981:1, 2,
8, 9; 1982:1, 3; Capelli 1982a:741-745;
Jezerinac 1982:187; Maude and Williams
1983:76; Capelli and Magnuson 1983548,
549, 551-554, 557-561, 563, 564; France
1983:99; Momot 1984a:40; Lodge et al.
1985:33; Page 1985:422; Magnuson and
-
80
Beckel 1985:lO; Lodge et al. 1986; Jezerinac
1986:179; Lodge and Lorman 1987591, 592.
Orconnectes virilis
Threinen 1958a: 1 13;
-
-
1958b:1,2; 1982b:3.
-
Orconectes Stamm 1977:40,41.
Crayfish Stamm 1977:42,43, 104, 105.
-
DIAGNOSIS (refer to Table 10): Carapace (Fig.
52c,k) subovate; rostrum longer than broad,
excavate dorsally with distinct marginal spines;
postorbital ridge with prominent spine. Cervical
groove excavate and discontinuous laterally;
distinct but short branchiostegal spine or
tubercle; prominent cervical spine present.
Areola (Fig. 52k) narrowest at midlength or in
anterior half of length, with no more than two
punctations in narrowest part; areola constituting 23.4 to 39.2% (mean 36.1%) of total carapace
length (28.7 to 49.3%, mean 45.7% of postorbital
carapace length) and 5.7 to 23.8 (mean 11.8)
times as long as broad. Antenna1 scale (Fig. 52j)
approximately 2.6 times as long as broad, widest
at about midlength, tapering to acute spine
distally. Cephalic lobe of epistome (Fig. 52n)
slightly broader than long, subquadrate with
uneven convex margins. Distal incisor region of
mandible (Fig. 52f) irregularly dentate-crenate.
Postorbital carapace length subequal to length
of abdomen; cephalic section of telson with one
large (lateral) and one or two slightly smaller
(mesial) spines in each caudolateral corner, distal
podomere of lateral lobe of uropod rounded; two
spines, large lateral and smaller more mesial spine
in each caudolateral corner; mesial ramus of
uropod with distomedian spine never reaching
distal margin; small spine situated distolaterally.
Chela (Fig. 520) moderately punctate, slender
(approximately 2.4 times as long as broad),
tuberculate; mesial surface of palm with two
distinct rows of tubercles, each with 5 to 7
tubercles. Carpus of cheliped moderately cleft
dorsally with one prominent and 2 or more
smaller spines mesially; 3 to 5 prominent ventral
spines, 7 to 9 small median ventral spines.
TABLE 10. Range of measurements (in mm) of various diagnostic structures of Wisconsin
Orconectes virilis.
N
Minimum
Maximum
Mean
Standard
Deviation
zk
48
48
48
27.8
21.3
10.6
59.3
47.4
24.2
40.4
32.0
16.3
7.1
5.9
3.0
48
48
9.6
0.7
22.3
1.9
14.9
1.2
2.9
0.3
39
39
5.9
8.9
15.1
23.4
10.3
14.8
2.2
3.4
47
48
48
48
13.1
5.2
3.3
12.0
26.8
8.9
7.1
22.0
18.9
7.3
5.4
16.9
2.8
1.o
0.9
2.4
24
24
24
23.7
18.9
9.0
52.0
41.9
21.1
36.1
28.4
14.2
5.9
3.0
24
24
9.0
0.9
19.2
I .6
13.0
1.2
2.8
0.2
19
19
4.9
12.5
17.9
7.6
10.9
2.5
3.4
48
48
48
21.5
15.9
7.2
60.3
49.3
25.7
38.0
30.0
15.1
7.6
6.2
3.2
48
48
7.2
0.4
23.1
1.9
13.5
1.2
3.1
0.3
34
34
3.7
4.3
14.6
21.4
7.7
11.4
2.4
3.7
MALES ($1)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
Pleopod
Total length
Central Projection
Mesial Process
Condyl length
-
MALES ($11)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
-
Chela palm
Length
Width
7.2
7.2
FEMALES
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
-
Chela palm
Length
Width
81
C
n
k
Ischium of only third pereiopod (Fig. 52m) with
hook, latter extending proximally slightly over
basioischial articulation. First pleopods of first
form male (Fig. 52a, e, g, h, i) reaching coxae
of first or second pereiopods with abdomen
flexed; terminal elements of first pleopod of male
(Fig. 52a, b, d, e, g, h, i) subparallel, elongate,
slender, tapering distally to acute tips, both
directed caudally at considerably less than 90
degrees to main axis of shaft; central projection
longer than mesial process.
Annulus ventralis (Fig. 521) 1.5 times as broad
as long, with caudal margin of nontuberculate
sternum anterior to annulus not overhanging
cephaloventral region of annulus; very deep
transverse depression extending across midlength
of annulus; anterior fourth of movable postannular sclerite overlain by annulus ventralis,
approximately 2.6 times as broad as long, and
approximately two-thirds as broad as annulus.
TYPE-LOCALITY:
“Lake Superior,” USA; restricted by Faxon (1914:420).
VARIATION. As one might speculate with such
a widely distributed species (see below), considerable morphological variation is noted for 0.
virilis in Wisconsin. Measurements of various
diagnostic characters are presented in Table 10
and ranges of measurements are useful in
recognizing variations of structures (size related
differences obviously are included). Considerable
variation is apparent in the first pleopod of first
form males. Typically, the terminal elements
(central projection and mesial process) are long
and subparallel with the central projection
slightly widened proximally and extending well
beyond the mesial process (Fig. 52a, e, i). Some
specimens have a very short, broad, blunt mesial
process (Fig. 52s) or a very short and narrow
mesial process (Fig. 52h). The “shoulder” of the
gonopod varies from distinct (nearly rightangular) (Fig. 52h) to gently sloping (Fig. 52i),
or rarely absent (Fig. 52a, g). The amount of
setation on the third maxilliped varies, but
virtually all individuals exhibit a dense cover.
The width, length, and shape, as well as the
development of spines on the rostrum vary
somewhat but generally most individuals
resemble the one depicted in Fig. 52k.
The number of tubercles on the inner surface
of the palm is variable. Generally there are two
rows, the mesialmost better defined and ranging
in number from 5 -7. The subparallel second
row is less distinct and consists of 4-9 tubercles.
The areola is relatively narrow with 2-3
punctations, sometimes only one, in its narrowest
part .
A brief discussion of gynandromorphs is presented in the section devoted to 0.propinquus.
Turner (1935) listed a number of examples of
gynandromorphs of 0.
virilis from Wisconsin (pp.
865,867,868,870 874,876, 877, 881). Hazlett
et al. (1974:304) noted a gynandromorph in a
small stream in Livingston County, Michigan.
We observed only one such specimen (Fig. 53)
which was collected from the Wolf River in
Langlade County, 25 August 1982. Except that
the left first pleopod was modified into that of
a relatively typical first form male, the individual
was a mature female with cement glands.
-
COLOR NOTES (refer to Figs. 54, 55). Ground
color of carapace brown or reddish-brown to
olive-brown fading ventrally on hepatic and
branchiostegal regions to tan or dark brown;
carapace sometimes mottled with light and dark
reddish-brown splotches, particularly pronounced over mandibular adductor muscle areas;
posterior portion of cephalic section with paired
dark brown splotches situated posterolaterally in
areola, extending onto first abdominal segment.
Abdomen similarly reddish-brown to dark tan,
mottled with distinct dark brown splotches
located slightly lateral to midline on each side
of terga but usually absent on last abdominal
segment; each pleuron sometimes with small,
dark marking anteriorly; telson and uropod tan
with margins outlined in light to dark reddishbrown or scarlet. Cheliped steel-gray to bluegray on all dorsal surfaces, often mottled with
small black to blue splotches; all spines and
tubercles and lateral margin of propodus yellow
to cream; distal part of fingers often orange
dorsally; ventral surface of all segments cream
to pale yellow; some individuals with sky-blue
on ventral surface of anterior portion of merus
and entire ventral surface of carpus and chela.
83
Figure 53. Photograph of ventral surface of female Orconectes
County (note modified left first pleopod).
(G.)
virilis gynandromorph
from Wolf River in Langlade
Figure 54. Photograph of “normal” color morph of Form I1 male of Orconectes (13.1virilis from Lake Michigan, Milwaukee
County.
84
Pereiopods 2-5 olive-green to bluish-gray dorsally
fading to cream ventrally.
Hazlett et al. (1979a) noted a prominent color
spot on the ventralsurface of the sixth abdominal
segment of many individuals inhabiting a stream
in Livingston County, Michigan.
Blue color morphs have been noted in the
literature (see discussion under 0. propinquus
above and Fitzpatrick 1987) and several have
been observed for this species in Wisconsin. A
male was found on 5 July 1971 by David Zalewski
in Booth Lake in Walworth County; and an
immature female was collected by Don Rose on
23 August 1971 from a silty creek on the outskirts
of Fond du Lac, Fond du Lac County. These
specimens were sky-blue dorsally and pale
whitish-blue ventrally. An albinistic specimen
was reported by Anderson (1975) to have been
taken from the Kishwaukee River in Dekalb,
Illinois. He indicated that it was a “chalky” color,
lacking pigments except for pigmented eyes. An
albino specimen was collected from a depth of
“300 feet” in Torch Lake in Antrim County,
Michigan (Hobbs, pers. comm.). We observed
no albino specimens but a light colored Form
I male was collected from Lake Michigan (Fig.
55).
ECOLOGY.Hubitat. One of the first to mention
the habitat of 0. virilis in Wisconsin was Bundy
(1883:402) who noted that it was one of the
most abundant species and frequented lotic
Figure 55. Photograph of light colored Form 1 male Orconectes (G.) wirilis from Lake Michigan, Milwaukee County.
85
habitats. Graenicher (1913:120-121) indicated
this to be the most common and widespread
Wisconsin crayfish and should be expected to
occur in any suitable habitat. Muttkowski
(1918:393) hoyever, claimed that in Lake
Mendota 0. wirilis appeared to be restricted to
the deep water. Turner (1926) reported that 0.
wirilis populations in Wisconsin were found in
streams, rivers, and lakes and occasionally in
smaller pools, ponds, and streams. In 1932
Creaser stated that 0. virilis was the most
common crayfish in the state and indicated that
typically it was found under stones in lakes,
streams, and rivers but that it also lived in muddy
creeks and in aquatic vegetation. Capelli (1975)
and Magnuson et al. (1975) echoed these
observations. Maude and Williams (1983) noted
its preference for soft mud substrate and stated
that it could not maintain its position in fast
currents. They pointed out that it had a mean
slip speed of 28.4 cm/sec; that is, it is able to
hold its position on a stream bed with a current
speed of no more than 28.4 cm/sec. Engle (1926)
noted that 0.virilis was not an effective burrower
in eastern Colorado and Nebraska as did Momot
(1975) in Michigan; however, Hazlett et al.
(1974:304-305) observed them t o burrow
extensively both in loose soil and in packed clay
along the banks of a Michigan pond and stream.
They rarely observed two individuals occupying
t h e same burrow but occasionally noted
copulating pairs together in a burrow. Phillips
(1980) found 0. wirilis to be an occasional
burrower in Iowa and described a burrow along
the shore of Split Rock Lake, Chickasaw County,
to be 30 cm deep with a 5 cm tall chimney
and slanted at an angle of 45 degrees, terminating
in two chambers. Berrill and Chenoweth (1982)
also reported 0. wirilis to burrow in Ontario.
During our studies in Wisconsin, we never noted
truly excavated burrows, but did observe that
in many localities, individuals made shallow
excavations beneath stones in stream beds or
on lake bottoms.
Physicochemical parameters. Overall, 0. wirilis
is considered to be a fairly tolerant species to
environmental perturbations. Its successful range
expansion naturally and in those areas where
86
it has been introduced can probably be related
to its high tolerability (see below). Schwartz et
al. (1963) noted that in the Patapsco River,
Maryland, this crayfish tolerated the following
variations: pH 5-9.5, temperature 5-31°C,
dissolved oxygen
2 12 mg/l, and coliform
bacteria counts 23,000 2,400,000+. Bovbjerg
(1970a) conducted experiments t o demonstrate
the oxygen requirements of both 0. wirilis and
0.immunis and noted that when individuals were
limited to oxygen concentrations of less than
1 mgll, 75% of 0. wirilis died within 5 hours,
whereas 20 hours elapsed before 75% of 0.
immunis died. Momot (1975) also noted that 0.
virilis is intolerant of low oxygen levels. It ,can
be concluded that unless individuals of 0.virilis
are able to expose their gills to the air-water
interface or move onto land, it is likely that
they will be unable to tolerate overnight lowering
of oxygen concentrations that occurs frequently
in shallow lentic habitats with dense vegetation
or with a thick covering of organic sediments.
Using a closed system, Jones and Momot (1975)
measured the oxygen consumption of 0. virilis
at various temperatures (9, 14, 19, 24°C). They
found that the correlation coefficient for mg
oxygen/hour/animal versus total body weight
was highly significant at all four temperatures
< -
-
(0.80 - 0.93).
Claussen (1980:38l), using adult specimens
from Lake Butte des Morts in Winnebago
County, Wisconsin, demonstrated that 0.virilis
(and 0.rusticus) is quite resistant to high water
temperatures. Their data suggest that 0. virilis
shows a heat resistance equal to that of the more
southern species. Peck (1985) suggested that the
ability of 0. virilis to select favorable temperatures and segregate according to social rank may
be a strong cue for habitat selection. Additional
data related to varied temperatures are presented
below in the discussion of its life history.
VerMeer (1972) worked with populations of
0. wirilis in Manitoba, Canada and showed it
to be a good indicator of mercury levels in the
environment. He indicated that at low levels
of contamination the abdominal muscles appear
to furnish more reliable data for indicating
mercury levels than does the whole animal. In
1978, Sheffy sampled populations of 0. virilis
along the Wisconsin River and reported the
crayfish to be a good indicator of mercury
pollution because individuals accumulate organic
mercury by scavenging on fishes, individuals are
more restricted in their movements than are
most vertebrates, and because they are very
abundant in the Wisconsin River.
France (1983) and Davies (1984) reported on
the effects of experimental lake acidification in
Lake 223 in northwestern Ontario, a simulation
of the effects of acid precipitation. They found
the combination of depressed pH and the low
calcium content of lake water inhibited calcium
uptake. Severe recruitment failure has resulted;
in addition, an elevated body burden of heavy
metals (Mn and Hg) and increased susceptibility
to the microsporidian parasite, Thelohania sp.,
have also contributed to a decrease in the
population size of 0.virilis.
France (1985)indicated that adult and youngof-year crayfish from Lake 240 (pH 6.6-6.8)
displayed a strong avoidance to potentially lethal
water oLpH 4.5 and below whereas crayfish from
Lake 223 (experimentally acidified) had reduced
avoidance, being significant at only pH 4.0. He
concluded that the prolonged survival of this
species in water receiving acid runoff (spring melt
of acid snow) seems doubtful. Tierney and Atema
(1986) showed that acid exposure may interfere
specifically and quantitatively with chemoreceptive processes.
Behavior. Bovbjerg and Stephen (1975430)
summarized the general ecology of 0. virilis by
stating that it is, “Nocturnal, agonistic, and
intolerant of neighbors, it spaces during the day
as an occupant of a crevice in the rubble of
the substratum; it wanders extensively and
forages during the night (Roberts, 1944;
Bovbjerg, 1953, 1970a, 1970b; Momot, 196713;
Caldwell and Bovbjerg, 1969; Heckenlively,
1970).” In Massachusetts, Camougis and Hichar
(1959)demonstrated that 0.virilis does not have
a small, confined home range, but moves freely
about the bottom, showing no evidence of
territorialism. Hazlett et al. (1974:305) presented
data suggesting that individuals infrequently
move but when they do, it is for some distance.
They demonstrated a statistically significant
positive correlation between length of carapace
and distances moved by females, but no such
correlation was shown for males; the home range
varies considerably, ranging from 0-30m along
a stream. During the day, individuals remain in
burrows or under rocks but after 2100hr they
become active, an observation also made by
others (e.g., Roberts 1944, Dean 1969). Hazlett
et al. (1975) conducted laboratory experiments
to measure the relationships among starvation,
energy reserves, aggression, and locomotion in
0.virilis; those individuals that were starved one
week were significantly more active than
organisms that had fed; starved crayfish increase
aggressive activity and locomotion, and their
protein and carbohydrate levels become lowered.
Hazlett et al. (197913) showed that 0. virilis
undergoes seasonal upstream migrations, presumably to prevent extensive downstream displacement. Minckley (1964), Momot (1966), Elliott
(1971), Hobbs 111 (1978), and Hobbs 111 and
Butler (1981) have also demonstrated that
certain aquatic invertebrates exhibit extensive
upstream movements, although not necessarily
as a compensatory movement for downstream
displacement following spates. See life history
section for further discussion.
Pearse (1909), in a laboratory study, found
that 0.virilis females were passive and that males
lacked any sex discriminatory ability and thus
copulated with any crayfish, male or female.
Bovbjerg (1956), however, provided data to
indicate sex recognition. He demonstrated that
movements of individuals are density related and
that males dominate females. Camougis and
Hichar (1959) and Abrahamsson (1966) noted
that mature males are highly aggressive and
showed that they repel smaller males and females.
Lunt (1967) determined that chela length is
proportional to carapace length and is positively
correlated with dominance; males are larger than
females and thus dominate them by aggressive
behavior. Fast and Momot (1973), working in
northern Michigan lakes, suggested that intersex aggression is temDerature related. They, and
Momot and Gowing 1972), postulated a density
87
related emigration of females from the littoral
zone to the deeper waters of lakes, the shore
region having the higher temperatures as well
as better conditions for food and shelter, which
are seized by males. Although Fast and Momot
( 1973:99) did not conduct laboratory experiments, they suggested that aggressive behavior
of adult males is sufficient to force females out
of the preferred warmer, shallow-water areas.
Jones and Momot (1981) discussed the distribution of 0. virilis in an oligotrophic lake in
Otsego County, Michigan in which allochthonous import is a significant source of benthic
food (energy). West Lost Lake has a steep basin
slope and thus the littoral leaf fall distribution
is extended to the deeper portions, where,
because of the social dominance of males, adult
females are forced to reside, but an adequate
allochthonous food supply supports the brooding
stock.
Heckenlively (1970) studied the intensity of
aggression in 0. virilis from Wisconsin. He
concluded the position of antennae of the
winning crayfish seems to be important in
crayfish aggression, as an indicator of the
aggressive state of the individual and as a threat
display to the opponent. Rubenstein and Hazlett
(1974) discussed the possible role of repeated
antenna1 wave as an indication of the acceptance
of defeat during fighting in 0. virilis. AmeyawAkumfi (1979) also noted these movements as
well as movements of mouthparts and ambulatory appendages and noted that these movements were appeasement displays by the “loser”
in the aggressive encounter. Fast and Momot
(1973) found that if populations became
crowded, fighting could increase and continue
until maximum spacing is attained. Bovbjerg and
Stephen (197 1) also showed that “ordinarily”
spacing may regulate the density of individuals;
however, they suggested a passive aggregation
(stacking) may have survival value during
population concentration at times of low water.
Ellis and Wellins (1973) noted that under
laboratory conditions, 0. virilis quickly established and maintained territories for several
weeks. They also showed territory size and the
number of aggressive encounters were related
88
to the size of individuals and they suggested that
conspecific recognition was acquired rapidly and
that this recognition was mediated, in part, by
pheromonal cues. Hazlett (1985) tested individuals for responses to water containing conspecific
individuals of several sex-status categories.
Females showed little difference in response to
waters from different categories and males did
not react to self water but did demonstrate
aggressive postures when nonself male and female
water was introduced.
Fast and Momot (1973) suggested that
cannibalism will limit population density similar
to any controls imposed by reduction in fecundity
or increased predation by other species. Momot
(1965a,b; 1967a) discussed the importance of
brook trout (Salvelinus fontinalis) predation on
crayfish populations. He (1965b:40) noted the
effect of trout predation and cannibalism on 0.
oirilis in West Lost Lake and made a strong case
that predation does not appear to be as important
in regulating population size as does behavior
at the time of molting; this is coupled with certain
physiological and mechanical problems associated with molting.
In 1982 Tierney and Dunham conducted
laboratory studies on 0.virilis and 0.propinquus
showing that individuals react to chemicals from
both species and do indeed distinguish conspecific members from the other species. Interspecific interaction between 0. virilis and other
species has been studied sparsely. Bovbjerg (1953)
was one of the first investigators to demonstrate
the aggressive nature of this species. He (1970a)
provided strong (albeit indirect) field evidence
that the competitively dominant species, 0.
virilis, through agonistic interactions, excludes
0. immunis from streams, the latter a species
more physiologically tolerant of a variety of
environmental conditions.
Magnuson et al. (1975) noted that 0. virilis
is the native species in northern Wisconsin and
when it coexists with either 0. propinquus or
0.rusticus, it is found in relatively low numbers,
suggesting competitive exclusion. Jezerinac
(1982) noted that when 0. virilis and 0.
propinquus occur sympatrically in the Chagrin
River basin in Ohio, 0. virilis occurs only in
the pooled sections of streams. Capeili and
Capelli ( 1980) studied hybridization in crayfishes
and presented evidence for hybridization
between 0.
propinquus and 0.rusticus but found
n o indication that 0. virilis “hybridizes” with
either. There is no evidence that 0. plirilis
hybridizes with any other species, although
interspecific (0.virilis and 0. popinquus) copulation was observed by us when crayfishes were
placed in collecting buckets after removing them
from bodies of water; Dowel1 and Winier (1969)
also noted 0. virilis copulating with 0. immunis
in t h e laboratory. For fu rth er discussion
concerning hybridization in crayfishes see the
Life History treatment of 0.propinquus.
In some of the population studies, various
forms of capture and tagging have been used.
Capelli (1975) and Davies et al. (1977) found
that trap catches were strongly biased toward
large males. Davies et al. (op. cit.) also noted
that population estimates based on these catches
were tenfold lower than those actually observed.
Hobbs 111 (1978) presented a review of various
techniques utilized by researchers t o mark,
temporarily and permanently, individuals for
population studies. Goellner (1943), Hazlett et
al. (1974), and Hazlett et al. (197913) used a
pleural clip technique for marking crayfishes.
Duke (1979618) concluded “. , . that if radiotracking is to be introduced into the investigation
of home ranges in crayfish, the effects of the
weight of transmitters o n the mobility of the
species during foraging and shelter - seeking
activities must be considered.”
Trophic relationships. A review of the trophic
relationships of 0.wirilis is in preparation (Hobbs
and Jass). Generally, this species is considered
to be a scavenger, feeding on virtually any organic
material. Boronow (1980, 1981, 1982) noted 0.
virilis in Turtle Creek and Eagle Lake and found
an apparent inverse relationship between the
abundance of crayfish and aquatic vegetation
in the lake. Other studies demonstrating the
effect of crayfishes grazing on the abundance
of aquatic macrophytes are reviewed by Magnuson et al. (1975).In addition, Dye and Jones
(1975) conducted field and laboratory studies and
showed the impact that Aeshna sp., as a predator,
has upon various sized young-of-year 0. uirilis:
they “. . . were highly effective predators on 46mm crayfish but less effective o n 7-12m
crayfish” (p. 532).
Jones and Momot (1983) calculated th e
lifetime energy expenditure per individual
crayfish to be 54.8 kcal in two pothole lakes
in Michigan. They also estimated the net growth
efficiency (K2) of individuals was 23.5%.
Summary. 0. viritis inhabits a wide variety of
habitats. During our field study we collected
representatives of this species from habitats
ranging from clear swift currents t o very turbid
waters with n o measurable flow (see Figs. 3, 58, 40, 41, 56-60). Animals were found on
Figure 56. a) Lake Minoqua in Oneida County inhabited
by Orconectes (P.) rusticus and 0.
(C.) virilis; h) tributary
of Tiffany Creek in St. Croix County; inhabited by 0.
(G.)
virilis.
89
substrates consisting of silt, sand, gravel, cobble,
and concrete blocks. Waters ranged in pH values
from 5.5 to 7.7 (mean = 6.9), specific conductance varied from 69 to 730 pmhos/cm (mean
= 282 pmhos/cm). The following plants were
commonly found in the aquatic situations in
which 0. virilis was observed: Carex sp.,
Ceratophyllum sp., C h a r a sp., Elodea sp.,
Equisetum sp., Najas sp., Nuphur microphyllum,
Pontederia cordata, Potamogeton crispus, P.
pusillus, Ranunculus acris, Sagittaria sp., Spirea
sp., Spirogyra sp., Tygha sp., Vallisneria americana,
and Zannichellia pulustris.
Figure 57. Rlchland Creek in Green County; inhabited
by Cambarus
(L.)diogenes, Orconectes (C.)propinquus
and
0.
(G.)wirilis.
Figure 58. Chippewa River in Pepin County; inhabited by
Orconectes (G.)virilis.
LIFE HISTORY. Egg production and hatching time.
Numerous studies have treated the life history
of 0. virilis in various parts of its geographical
range. One of the early observations was made
by Harris (1900) when he noted that 0. virilis
laid eggs in the spring in Kansas. Steele (1902)
reported that in Missouri approximately nine
days after hatching the first molt of juveniles
occurs and within 14-15 days after hatching the
yolk has been used and young leave the female.
In Michigan, Pearse (1910) noted Form I males
as early as 14 April and females with eggs on
30 April. Fasten (1914) was the first investigator
to study spermatogenesis in the species. Using
crayfish from the environs of Madison, he
determined the chromosomal number to be 200,
that during June and July the most rapid rate
of proliferation occurred in which all stages of
spermatogenesis were observed and testes
obtained their maximum size, and that there are
two periods of copulation in Wisconsin: April
and May and September and October.
Spermatogenesis also was discussed by McCroan
(1940). Creaser (1931b) reported that females
lay eggs in Michigan before the last of April and
indicated that in Wisconsin (1932) females lay
their eggs in early spring, usually before the last
of April.
Following is a summary of the life history of
0.oirilis in Wisconsin with variations that have
been noted in other states. Threinen (1958a,
b) and Momot (1964) reported a maximum life
span of three years. Adults generally mate in
late summer (August October) of their second
year. Capelli and McIntire (1980) reported a
-
90
mean mating duration of 71 minutes, the longest
time compared to that noted in the pairing of
0.propinquus and of 0.rusticus. Following copulation most males die but some individuals survive and may be reproductively active for one
more season. Threinen (1958a,b), Crocker and
Barr (1968), and Fasten (1914) also reported a
spring mating period; this has been observed by
Caldwell and Bovbjerg (1969) in Iowa. However,
Weagle and Ozburn (1972) and Aiken (1969a)
reported no copulation occurring in the spring
in Ontario. Weagle and Ozburn (1972) noted
that in the McIntyre River in Ontario copulation
was restricted to those adults with a carapace
length of at least 24mm. Berrill (1978) found
that in southern Ontario the smallest sexually
mature male had a 26mm cl, the smallest sexually
mature female, 27mm cl, while the mean size
of sexually mature males was 34.5mm and that
of females, 33.5mm cl. Eggs are not laid until
the spring (hibernation period beginning in late
October) with peak activity occurring in midJuly; Phillips (1980) reported females with eggs
in Iowa as early as 15 April. Weagle and Ozburn
(1972) indicated the peak period to be between
May and June in Ontario when water temperatures warmed from 10 15°C. That all of the
ovarian eggs of crayfishes do not become
attached to the pleopods of the female is well
documented (Penn 1943, Goellner 1943, Smart
1962, Mason 1963, Momot 1967b, Prins 1968,
Payne 1971). Momot (1964)reported the average
number of ovarian eggs in 0. pririlis to be
approximately 162 with a maximum number of
276; he also indicated that the number of eggs
affixed to the pleopods ranged from 1 to 220.
The number of eggs produced is a measure of
t h e potential reproductive capacity, which
increases linearly with the size of the female.
He reported 83 eggs laid by one-year old females,
107 by the two-year old females, and (1967b)
an ovigerous female carrying 94 eggs. Weagle and
Ozburn (1972) reported a mean of 214 eggs.
Momot and Gowing (1977a) stated ovarian egg
counts to range from 92 to 156 and pleopodal
eggs to range from 66 to 145 in three Michigan
lakes. We examined a small female (19.3mm cl)
carrying 46 eggs (egg diameter 1.75mm) collected
from the Black River in La Crosse County 24
May 1978.
Once the eggs are laid, they are carried for
several weeks. Stephens (1952) suggested that
the female reproductive cycle is under endocrine
control: that four hormones control ovarian
growth, maturation, and egg-laying and that
these hormones are regulated in turn principally
Figure 59. Little Waumandee Creek in Buffalo County;
inhabited by Camburus (L.) diogenes and Orconectes (G.>
Figure 60. South Fork Flambeau River in Price County;
inhabited by Orconectes (G.)
uirilis.
-
uirilis.
91
by photoperiod. Egg laying in spring is induced
by exposure to long daily light periods. Aiken
(1969b)) however, demonstrated that temperature plays a more important role than Stephens
hypothesized. He indicated that both photope.
riod and temperature control ovarian growth and
maturation, and, that in spring increased water
temperature rather than a long day photoperiod
induces egg laying. He (1968b) also showed that
female 0. virilis in Alberta do not extrude eggs
at temperatures below 10" C. While brooding,
the female continues to feed. Hazlett et al.
(1979b)noted that females carrying eggs or young
were not very active in a Michigan stream.
Caldwell (1969) noted that in an Iowa stream
the rigid reproductive pattern of later egg laying
and prolonged immaturity minimizes the hazard
of the predictable spring flooding period. Hazlett
et al. (1974) noted that ovigerous females that
burrow tend to remain in the burrow and as
the eggs mature, the females move to the deepest
portion. Hazlett et al. (1979b) observed that
following the release of young, the mothers tend
to move downstream, whereas males or females
without eggs or young show no tendency to move
either up or downstream. Caldwell and Bovbjerg
(1969) and Phillips (1980) reported females with
young as early as mid-May in Iowa with young
leaving the female by early June, slightly ahead
of Wisconsin crayfish. Little (1975401) conducted laboratory studies using 0.virilis and two
other species and demonstrated that brooding
females release chemical stimuli into the water
that attract young to them, thus facilitating the
protective furictions of the brooding interaction.
He suggested that these cues are species-specific
but pointed out that they are not brood specific;
that is, young are attracted to females with young
other than their own mother, Weagle and
Ozburn (1972) indicated that the young undergo
at least five molts during the first year but that
there may be as many as seven; they reported
growth increments ranging from 2.0 to 2.7mm
cl for all age groups. Stephens (1955) noted a
positive correlation of molting with day length
(see Aiken 1968a for a discussion of ecdysis in
0. v i d i 3 ) . The relationship of the molting cycle
to tissue metabolism has been demonstrated by
92
McWhinnie and Kirchenberg (1962) and Travis
(1960), McWhinnie (1962))and McWhinnie and
Corkill (1964) further discussed the moltintermolt cycle in this species. Of note, certain
types of pollutants such as cadmium probably
act to depress molting in 0. virilis (Leonhard
1978:3). With the onset of low temperatures,
molting ceases. Generally, sexual maturation is
not reached in the first summer and, as
mentioned above, copulation occurs late in their
second summer. In Ontario, however, Weagle
and Ozburn (1972) noted that 50% of males
reached maturity by the first year as did 65%
of females. Momot (1984b, 1986) reported that
the low temperature regime at high latitudes
inhibited rapid juvenile growth, inducing an early
maturity molt.
Life spun. Following the release of young, most
adult females die; a few survive and mate again
the third fall. Momot (1964) determined that
females usually produce one brood in a lifetime
(two seasons of growth required for maturation),
with approximately 15% surviving to produce
a second brood. He (196713) found that mortality,
highest in young-of-year, peaks during the middle
of the second year for females, and is highest
for males during the fall of the third year
following mating.
A number of investigators have shown a
seasonal shift in the depth distribution of 0.
virilis in lakes. The studies conducted by Momot
(1967b) and Momot and Gowing (1972) in
northern Michigan lakes indicate that male and
female 0. virilis are concentrated in shallow
water during May and June, Following the
hatching and release of young, females molt and
migrate to deeper water by late June; adult males,
however, remain in shallow water all summer.
In Alberta, both sexes move to deeper water
in late summer, females preceding males (Aiken
1968b). At the same time, in both areas, young
of both sexes also migrate to deeper waters. Both
Aiken (196813) and Fast and Momot (1973)
postulated that migration to greater depths was
related to sexual maturation. Aiken (1968b) and
Ryck (1970) found in the laboratory that juvenile
maturation occurred only when crayfish were
kept in total darkness at temperatures below
10°C for a minimum of four months. Severe
winter conditions would eliminate a population
if they failed to migrate to a depth greater than
that at which the water freezes to the bottom.
Thus, rather than burrow into the substrate to
escape the threat of freezing, they accomplish
the same by moving to deeper water. Youngof-year appear to have higher mortality rates
during the winter than adults (Aiken 1968b).
This species has been noted from varying depths
in lake basins: 31.7m in Lake Michigan (Creaser
1934a), 20m in Lake Winnipesaukee in New
Hampshire (Aiken 1965), 9.2m in Shirley Lake
in Ontario (Crocker and Barr 1968), and 91.5m
in Torch Lake, Michigan (Hobbs, pers. comm.).
At spring thaw both males and females return
to the shallow zones of the lake.
Molt cycle. Several authors have noted a slower
growth rate in female 0. virilis than in males
(Momot 196713, Aiken 196813, Caldwell and
Bovbjerg 1969) and presented theories to explain
this observation. Females undergo fewer molts;
this is associated with egg production and
brooding, superior male aggression, and, since
females are found in deeper waters with lower
temperatures, their rate of growth is less than
that of males. Hazlett and Rittscholf (1985)
suggest that the difference in yearly size
increment between males and females in a
southeastern Michigan population is due to the
frequency of molting and not to the increment
per molt. Like 0. propinquus, 0. virilis demonstrates sexual dimorphism in the chela (Weagle
and Ozburn, 1970). See discussion under 0.
propinquus that treats importance of chelae size
in intraspecific interactions.
Orconectes virilis from Lake 223 (see above)
were subjected to varying pH levels in the
laboratory. Malley (1980) found that uptake of
Ca++ by postmolt crayfish, measured by the use
of 45Ca as a tracer, was inhibited by pH below
5.75 and ceased completely below pH 4.0 when
these levels of acidity were applied acutely.
Schindler et al. (1985) reported on the longterm ecosystem effects of the %year acidification
of Lake 223, which began in 1976. They reported
that by 1979 the exoskeleton of 0. virilis
hardened more slowly after molting and animals
remained softer. By 1980, in addition to the
recalcification problem, the population was
infested by a microsporozoan parasite
(Thelohania) and no young-of-year were observed. In 1981, a noticeable drop in the crayfish
population size was observed and no crayfish were
present by fall 1983. France (19841suggested that
a gradual acidification of a lake to an average
annual pH below 5.5 could result in eventual
population extinction as a result of mortality
of the young.
Production. Threinen ( 1958a) indicated that
0.virilis is the most common and valuable species
in Wisconsin, producing up to 1000 lbs./acre.
Camougis and Hichar (1959) also noted its high
level of production, estimating 8300 individuals/
acre (approximately 226kg/ha). Momot
(1967b:78) hypothesized that the high production rate (207.2kg/ha)results from the crayfishes’
ability I ‘ . . . to utilize marl-producing algae and
associated organisms of the aufwuchs as a food
source . . .” Other estimates of production in
its range are: from “One of the best fishing sites
in Wisconsin. ..,” 37.3kgIha (Momot and
Gowing 1976:39); from three lakes in Michigan,
60-142kg/ha (Momot and Gowing 1977a); from
two lakes in Ontario Dock Lake, 52.5kg/ha
(mean biomass of 28.7kg/ha) and Shallow Lake,
43.5kg/ha (mean biomass of 29.6kg/ha) (Momot
1978). Momot (op. cit.) noted lower mean
biomass and annual production in populations
in Ontario lakes than in similar Michigan lakes
despite faster growth rates, higher fecundity, and
an earlier age at maturity. He indicated that
in those Ontario lakes the instantaneous growth
rate declined with age, contributing to the low
annual production and mean biomass. Higher
annual mortality rates affect the higher growth
rate in the Ontario lakes. Momot and Gowing
(1977a) determined that fluctuations in mortality rather than in growth rates produced most
of the year-to-year differences in biomass
accumulation; that is, density-dependent control
of mortality and fecundity precipitated changes
in yearly biomass. Momot and Gowing
(1977b:2057) conducted a management study on
two lakes and found that any strategy based on
maximum sustained yield must be compromised
.
-
93
to reduce the risk of catastrophic loss. Further
discussion of the interrelationship of biomass,
growth rate, and annual production of 0. virilis
is presented in Momot and Jones (1977); see also
Momot and Gowing (1983).
Life history data are presented in Table 11
for 0.virilis.
DISTRIBUTION: This is the only crayfish that
has been found in every county in the state (Fig.
61); hence it is found in every major drainage
system in both the St. Lawrence and Mississippi
river basins. Orconectes virilis is the northernmost
distributed North American crayfish and its
geographical distribution in North America is
shown in Fig. 62. This species also has been
introduced into Europe.
Schwartz et al. (1963) noted that 0. virilis
may have been introduced into the Patapsco
River in Maryland as early as 1885. Aiken (1968~)
noted that it may have been brought into
Alberta, and that introductions in other areas
have been implicated in its range extension:
California (Daniels 1980), Colorado (Unger
1978), New Mexico (Bouchard 1977c), and
Tennessee (Bouchard 1976a,b); see also Hobbs
et al. in press). Bouchard (1976b:13) suggested
that introductions of 0. virilis into Tennessee
were made by fishermen using non-native
crayfish as bait. He (1977133418) also noted that
0.virilis is a very successful and aggressive species
which has become well established in other
localities outside its natural range. Schwartz et
al. (1963) cited several areas in Maryland where
0.virilis has greatly reduced the diversity of the
crayfish fauna and where it has restricted the
distribution of certain native crayfishes that were
once commonly widespread. Undoubtedly “bait-
TABLE 11. Life history data for 0. virilis (* from references only; ** from literature and
from our collections or USNM or OSU collections; those without an asterisk(s) from USNM
collections only).
$1
Month
Jan.
“Wis.
Feb.
“Wis.
$11
P
9y
copul
~~
Mar.
“Wis.
Apr.
“Wis.
Wis.
‘Wis.
May
“Wis.
Wis.
‘Wis.
Jun.
Wis.
Wis.
“Wis.
Jul.
“Wis.
“Wis.
“Wis.
Aug.
“Wis.
“Wis.
‘Wis.
Sep.
‘*Wis.
Wis.
‘Wis.
Oct.
“Wis.
Wis.
‘Wis.
~~
94
Nov.
‘Wis.
Dec.
“Wis.
Wis.
‘Wis.
‘Wis.
‘Wis.
L
92'
91.
L
90-
I
N
89"
Figure 61. Distribution of Orconectes (G.) wirilis in Wisconsin; closed circles-specimens
circles-from literature.
I
88'
examined in this study; open
95
bucket” transferals have occurred in Wisconsin,
but because this is the “native” species, being
found throughout the state, it is highly unlikely
that its transfer from one place to another would
significantly affect the distribution of any other
species in Wisconsin except 0. immunis, In fact,
recent studies (e.g., Capelli 1982a,b) have shown
that in populations of crayfishes in northern
Wisconsin, large shifts in favor of 0.propinquus
at the expense of 0. virilis and in favor of 0.
rusticus at the expense of both species, have
occurred in the last 30-50 years. Both Huner
(1978) and Bouchard (1978) also mentioned the
displacement of 0. virilis by 0. rusticus in parts
of Wisconsin.
CRAYFISHASSOCIATES. In Wisconsin, 0. virilis
has been found together with C. diogenes, 0.
immunis, 0. propinquus, 0.rusticus, P. a. acutus,
and P. gracilis.
Figure 62. Geographic distribution of Orconectes (G.)
virilis in North America.
96
Procumburus (Sirurdielka)grucilis (Bundy)
(Figures 63
- 68)
irregularly dentate. Abdomen narrow and
distinctly shorter than carapace. Cephalic
-
Cambarus gracilis Bundy 18765; Forbes 18765;
Bundy 1882:178, 179; 1883:403; Faxon
1884:141; 1885b:6, 7, 58; Underwood
1886:369; Ortmann 1902:279; Harris 1903a:
58, 98, 99, 155; Ortmann 1905104; Graenicher 1913:118, 119; Faxon 1914:412;
Ortmann 19183347; Creaser 1932:321, 323,
325, 326, 331, 332, 335, 336; Creaser and
Ortenburger 1933:44; Turner 1935:876;
Walters 1939:23; Ortmann 1945847; Hinkelman 1970:7, 49.
Cambarus gracillis Bundy 1882:182, 183.
Cambarus (Cambarus) gracilis Turner 19352381.
Procambarus gracilis
Williams and Leonard
1952:981, 983; Pennak 1953:462; Williams
1954:828; Hobbs 1959:887; 1968:K9; Reimer
196967; Bell 1971:17; Hobbs 1972b:47; Page
1974:97; Capelli 1975:39,40, 205, 207; Hayes
1977:444; Hobbs 1976a:47; Bouchard
1978:14; Pennak 1978:482; Phillips 1980:90;
Capelli 1982a:741; Page 1985:371.
Procambarus (Girardiella) gracilis
Hobbs
1974b:47; Hayes 1976:39; Fitzpatrick
1983:206; Hobbs 1984:16, 17 [by implication];
Hobbs and Rewolinski 1985:26-33.
+
-
-
DIAGNOSIS (refer to Table 12): Carapace (Fig.
63c,k) laterally compressed, vaulted dorsally and
lacking cervical, branchiostegal, and hepatic
spines or prominent tubercles; postorbital ridges
without spines or tubercles. Rostrum, deeply
excavate dorsally and lacking median carina, with
lateral margins subparallel, converging anteriorly
in small acumen. Cervical groove moderately
deep, somewhat sinuous and continuous laterally. Areola (Fig. 63k) obliterated or almost so
near midlength, constituting 37.4 to 41.3%
(mean 39.3%) of total carapace length (44.4 to
48.6%, mean 46.1%, of postorbital carapace
length) with room for no more than one
punctation in narrowest part. Antenna1 scale
(Fig. 63m) approximately 2.5 times as long as
wide, broadest near midlength; distal portion
tapering, acute. Cephalic lobe of epistome (Fig.
63h) generally rounded, slightly broader than
long, rounded anteriorly with margins convex.
Distal incisor region of mandible (Fig. 63i)
section of telson with one conspicuous and one
very small (mesial) spine in each caudolateral
corner; distal podomere of lateral lobe of uropod
rounded; distolateral spine of proximal podomere
distinct; mesial ramus of uropod with distomedian spine never reaching distal margin. Chela
(Fig. 63n) slender, little more than twice as long
as broad, somewhat inflated and tuberculate
dorsally; mesial surface of palm with one
(occasionally two) very distinct row of tubercles,
mesialmost consisting of 6-9; little more than
proximal half of mesial margin of dactyl also
tuberculate, mesialmost forming row of 4-6
tubercles, opposable margin of dactyl with
distinct, shallow excavation at base; fixed finger
with tubercle on mesial margin of basal 1/3.
Carpus of cheliped deeply furrowed dorsally with
very prominent spine distomesially. Ischium of
only third pereiopod (Fig. 633’)with simple hook,
that extending proximally over basioischial
articulation. First pleopods of first form male
(Fig. 63a, d, e, g) symmetrical; lacking preapical
setae; distinct angular shoulder at base of
terminal elements; mesial process, longest of
terminal elements, slender, acute, and directed
caudodistally; cephalic process short, acute,
directed distally; caudal element consisting of one
quadrangular corneous plate (“caudal knob”)
and small spiniform process (“caudal process”)
at its mesial base; central projection bladelike,
curved caudodistally and slightly overreaching
caudal knob; first pleopod of second form male
illustrated in Fig. 63b, f.
Annulus ventralis (Fig. 631) somewhat subquadrangular and completely exposed, cephalic
region with deep median longitudinal trough
bordered by tuberculate elevations; sinus curving
caudodextrally, making hairpin turn before
curving transversely and crossing to sinistral
region, then turning caudodextrally, on to broad
relatively flat caudal region.
TYPE-LOCALITY: Normal, McLean County,
Illinois, USA; restricted by Hobbs (197413347).
97
VARIATION. On the basis of the few specimens
examined, gross morphology of P. gracilis does
not appear to be particularly variable. Relative
lengths of the terminal elements of Form I male
gonopods are somewhat variable but generally
are consistent with those shown in Fig. 63d,e.
Chelae, particularly those of Form I males,
demonstrate some variation with respect to
number of tubercles on the mesial surface of
the palm (usually 6-7, occasionally as many as
9). Variation in excavation of the rostrum is
apparently not regionally restricted, and
although the degree of excavation of the annulus
ventralis is variable, so few specimens are
available that one cannot be certain of its being
consistent in any one locality.
TABLE 12. Range of measurements (in mm) of various diagnostic structures of Wisconsin
Procambarus gracilis.
N
Minimum
Maximum
Mean
Standard
Deviation
f
MALES ($1)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
*
Pleopod
Length
-
6
6
6
29.4
25.3
12.4
39.0
33.7
16.5
34.3
29.5
14.5
3.3
2.9
1.3
6
4
11.6
0.3
15.7
0.4
13.7
0.4
1.5
0.0
6
6
8.4
10.2
11.6
14.2
10.0
12.5
1.1
1.4
4
8.1
11.2
10.3
1.5
13
13
13
18.2
15.3
7.7
36.8
32.1
15.1
24.6
20.9
10.2
4.7
4.3
1.9
13
11
6.8
0.3
14.7
0.7
9.6
0.4
2.0
0.1
12
12
3.0
3.7
11.4
13.8
5.7
7.1
2.2
2.6
5
5.9
11.1
7.7
2.1
7
7
7
31.3
26.7
12.9
42.1
36.0
16.8
35.2
30.2
14.8
4.0
3.3
1.5
7
5
12.0
0.3
16.9
0.4
14.0
0.4
1.7
0.0
6
6
7.4
10.0
12.1
8.1
10.2
1.o
9.4
MALES ($11)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
P1eopod
Length
-
FEMALES
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
~
98
1.o
v
n
Figure 63. Procamburus (Girardiella) gracilis (the Oakwood site, Milwaukee County; I, adult female; b,f, second form male;
all others first form male): a,b,e, mesial view of first pleopod; c, lateral view of carapace; d,f,g, lateral view of first pleopod;
h , epistome; i, incisor margin of right mandible; j , proximal podomeres of third, fourth, and fifth pereiopods; k, dorsal
view of carapace; I, annulus ventralis; m, antenna1 scale; n, dorsal view of distal podomeres of cheliped.
99
Figure 64. Photograph of Form I male Procumbarm (G.) gracilis dug from burrow in Kenosha County.
COLORNOTES (refer t o Fig. 64). Ground color
of carapace scarlet, reddish-brown, or olivebrown fading ventrally to tan or cream. Downing
(1924) mentioned a green ground-color; Harris
(1903b) noted that in Kansas females are olive,
green and males are “almost a salmon red”;
mottled pattern of similar colors overlying
mandibular adductor region. Rostra1 margins and
postorbital ridges pink, orange, or tan. Abdomen
similarly reddish-brown to olive-brown, with
some specimens demonstrating a mottled pattern
of deeper browns; some individuals have a
middorsal, tannish stripe extending posteriorly
along terga of abdomen onto telson; cream
ventrally. Antennular and antenna1 peduncles
brown to tan. Cheliped with dorsal surface of
carpus, distal third of merus, palmar area of
propodus, and both movable and fixed fingers
olive-brown to tannish-brown; distal part of all
three podomeres green t o grayish-blue; all spines
and knobs of merus, carpus, and propodus cream;
tubercles on propodus and on movable and fixed
fingers black to dark olive; ventral surface
orangish-pink to cream. Pereiopods 2-5 with
merus through dactyl olive to brown dorsally
100
fading to pale brown or light orange ventrally,
and distal margins of ischium, merus, carpus, and
propodus brown. Telson and uropods scarlet,
reddish-brown, or olive- brown, all spines of
latter scarlet.
ECOLOGYAND BEHAVIOR. P. gracilis, a primary
burrower, has been noted by several authors (e.g.,
Bundy 1882, Steele 1902, Downing 1924, Creaser
1932) to inhabit prairie regions throughout its
range, this distribution being similar to that of
the eastern tall grass prairies. It is in the
northeastern extremity of this area, referred to
as the Prairie Peninsula (Transeau 1935), and
in its forested ecotone, that P. gracilis is found
in Wisconsin. Presumably, this species entered
Wisconsin from Illinois by dispersing northward
along contiguous drainages (e.g., Sugar, Rock,
Fox, and Des Plaines) (Hobbs and Rewolinski
1985).
This crayfish has been found in ponds, roadside
ditches, small creeks, marshes, swamps, small
artificial lakes, and it frequents burrows in wet
pastures and flat fields in prairies, and the banks
of sluggish streams and ponds (Figs. 9,65).
Additional ecological observations are reported
b y Bundy 1882, Hargitt 1890, H a r r i s
1900,1902,1903b; Graenicher 1913, Creaser
1932, Creaser and Ortenburger 1933, Williams
and Leonhard 1952, Bliss 1968, Reimer 1969,
Page 1974,1985, Pennak 1978, and Phillips 1979,
1980. See Hobbs and Rewolinski (1985) for a
discussion of a population from the southsouthwestern portion of Milwaukee County,
Wisconsin with particular emphasis on chimney
(Fig. 66) and burrow construction. Data for these
burrow waters are presented in Table 13. Note
that the values for oxygen are considerably
higher than those obtained from burrows in
other parts of this and other states (e.g., Jaspers
1969, Jaspers and Avault 1969, Grow and
Merchant 1980, and Hobbs 111, unpublished
data).
Creaser (1932332) stated that in Missouri he
excavated a burrow in
which the
water
level
was two meters below the surface. Creaser and
Ortenburger (1933) noted that burrows in
Oklahoma may be greater than two meters in
depth and that most were characterized by an
enlarged pocket at the bottom. Phillips (1980)
stated that burrows in Iowa were vertical and
usually greater than l m in depth, being 2-3cm
in diameter and terminating in a flask-shaped
enlargement 1Ocm or less in diameter and
partially filled with mud and water. Wisconsin
burrows are similar in form and as mentioned
above, many are bifurcate; in several instances
an individual was captured not in the main
tunnel but in the side branch above the water
level.
TABLE 13. Physicochemical data from water in burrows of P. grucilis at t h e Milwaukee Park
System, Milwaukee County, Wisconsin, 2 July 1982 (n=5) after Hobbs and Rewolinski 1985.
~
Min
Max
Mean
Depth to
water
(cm)
Temp
(" C)
35
52
12.8
17.0
46
14.6
Specific
Conductance
pmhosicm
PH
61.9
698
785
59.1
730
7.35
7.40
7.36
0 2
0 2
(mgil)
%Sat.
8.1
9.0
8.6
Figure 65. Field near Oakwood, Milwaukee County; habitat
of Rocambarus (G.)
gracilis.
55.7
Figure 66. Chimney of burrow of Procamburus (G.)gracilis,
Milwaukee Countv.
101
Williams and Leonard (1952) stated that
individuals are active at night in Kansas. In
Oklahoma, “During rainy periods and also at
night it leaves the burrow for repair or deepening
of its home or for feeding” (Creaser and
Ortenburger 1933:43). Hayes (1975) also stated
that P. gracilis in eastern Oklahoma constructs
simple burrows, with or without chimneys, and
that individuals leave the burrow only during
rainy periods or on warm, humid nights. He
observed that the greatest social activity occurs
from late April to late June or early July. Shortly
after sunset individuals leave their burrows, when
aggressive and sexual encounters are common.
He reported eight behavioral components for the
activity of P. gracilis during social interactions:
1) alert
2) approach
3) threat
4) combat
5) submission
6) avoidance
7) escape
8) courtship
Courtship results in the occupation of prime
burrow sites by dominant breeding pairs. See
Hayes (1977:447) for further discussion of the
behavior of this primary burrower in Oklahoma.
LIFE HISTORY. Hobbs and Rewolinski
(1985)
observed young-of-the-year in pools in April and
during July in burrows alongside pools and
streams. A female with eggs was collected on
17 July, and Form I males were observed from
July until October. O n 2 July, two Form I males
and a mature female were collected from the
same burrow.
Steele (1902) noted that this crayfish was
seldom seen except during February and March
in Missouri. She stated that females moved with
their newly hatched young from their burrows
along the banks of creeks and ponds out into
the water. The adults returned to their burrows
by the end of March but the young were often
found in open water during April and May. She
also noted that males were rarely found in open
water, as did Faxon (1885a) and Harris (1903a).
Both Creaser (1932), in Missouri, and Williams
(1954:827) in Missouri and Oklahoma noted
females with young as late as October. Likewise,
Creaser and Ortenburger observed females with
young as late as October in Oklahoma and said
102
“.
(1933:43-44)that this crayfish . . probably also
leaves the burrow to breed but this phase of
its life is not known.”
Hargitt (1890) stated that breeding males and
females were found during the spring, and Harris
(1900) and Williams and Leonard (1952)
observed females with attached young during
early spring in Kansas; breeding males were seen
only during June. Juveniles were observed
(Phillips 1980) in May, June, and late September
in Iowa. Refer to Table 14 for a summary of
life history data.
This primary burrower obviously spends most
of its life in the burrow habitat, the females
moving to open water for a relatively short period
in the spring, summer, or fall where their newly
hatched young are released. It is not known
where copulation takes place but it is probable
that it occurs in the burrow. Lending credence
to this supposition are the capture of Form I
males and a female in the same burrow (see
above), Harris’ (1900) observation that embryos
develop while females are in burrows, Steele’s
(see quote above) note that females enter creeks
and ponds with their newly hatched young, and
Hayes (1975) behavioral observations. Page
(1985) indicated that length-frequency data
suggest that males may live longer than two years
in northeastern Illinois.
DISTRIBUTION: WISCONSIN.
Procambarus (G.)
gracilis is the rarest Wisconsin crayfish and
appears to be restricted to the southeastern
counties. It is thus known only from the RockFox and Lake Michigan drainage basins (see
Appendix I) and has been collected from the
following counties in the state (Fig. 67): Kenosha,
Milwaukee, Ozaukee, Racine, Walworth, and
Waukesha (total geographical distribution shown
in Fig. 68).
CRAYFISHASSOCIATES. In Wisconsin C. diogenes,
0.uirilis, and P. (0.)
a. acutus have been collected
together with P. (G.)gracilis in a few localities.
TABLE 14. Life history data for P. (G.) gracilis (from literature and from our collections
or USNM collections)
Month
YOY
open
water
Jan.
Feb.
March
Wis.
April
Wis.
June
Wis.
JU ~ Y
Wis.
Wis.
Aug.
Wis.
Wis.
Sept.
Wis.
Oct.
Wis.
Nov.
Wis.
Wis.
Wis.
Wis.
Wis.
Wis.
Wis.
Wis.
Wis.
Dec.
103
A '
42.
910
L
L
30'
Figure 67. Distribution o f Procumburus (G.)gracilis in Wisconsin.
104
I
N
89"
0
I
S
88'
Figure 68. Geographic distribution of Rocambarus (G.)gracilis (modified from Hobbs and Rewolinski, 1985).
Procumbarus (Ortmannicus) acutus ucutus
(Girard)
-
(Figures 69 72)
Cambarus acutus Girard 1852:91.
Cambarus stygius Bundy 1876:3; 1882:177,179,
180; 1883:402; Harris 1903a:130, 155; Creaser
1932:335; Hobbs 1981:372.
Cambarus Stygius Faxon 1884:140; 1885b:7.
Cambarus U C U ~ U S - Bundy 1882:177, 179, 180;
1883402; Faxon 1884:135, 136; 1885b:7.
Cambarus blandingii - Faxon 1884:135, 136.
Cambarus blandingii acutus - Underwood
1886:368; Harris 1903a:50, 80, 150, 155;
Creaser 1931b3271; 1932:325, 326, 332, 333,
335, 336; Creaser and Ortenburger 1933:42;
Gander 1927:222; Walters 1939: 21; Hinkelman 1970:7.
Cambarus Blandingii acuta - Faxon 1885b:20.
Cambarus (Cambarus) blandingi acutus - Grae-
nicher 1913:118.
Cambarus blandingi acutus
Graenicher
1913:119, 120, 121, 122.
Cambarus (Ortmannicus) blandingi acutus
Ortmann 1931:63.
-
-
Procambarus blandingii acutus Hobbs 1942a:95;
Williams 1954:832; Threinen 1958b:2;
Crocker and Barr 1968: 135; HinkeIman
1970:29, 41, 43, 44, 45, 47, 48; Threinen
198210:3.
105
- Capelli 1975:39, 48, 50,
53, 58, 199, 200, 205, 207, 208; Capelli and
Magnuson 1983:548,551,558, 559,563, 564.
Procambarus acutus
-
Procambarus acutus U C U ~ U S Capelli 1982a:741.
Procambarus (Ortmannicus) acutus acutus Hobbs
1972a:9 [combination used herein]; Hobbs
1984:14 [by implication].
DIAGNOSIS(see Table
-
15):Carapace (Fig. 69c,g)
with one pair of prominent cervical and
branchiostegal spines or tubercles; suborbital
angle generally acute; cervical groove shallow and
discontinuous laterally. Rostrum shallowly
excavate dorsally, lacking median carina, long,
broad at base, tapered, usually with marginal
spine, or distinct angles at base of sharp acumen;
postorbital ridge prominent, with cephalic spine
or tubercle. Areola (Fig. 69g) 4.8 to 18.6 (mean
10.0) times as long as broad, constituting 20.9
to 38.0% (mean 34.6%) of total length of
carapace (26.8 to 47.4%) mean 44.5%, of
postorbital carapace length) with room for 3
4 punctations in narrowest part. Antenna1 scale
(Fig. 69i) approximately 2.2 times as long as wide,
broadest at midlength. Cephalic lobe of epistome
(Fig. 691) triangular, distinctly broader than long,
pointed anteriorly. Third maxilliped densely
setiferous, lateral half of ventral surface of
ischium often with long plumose setae on
proximal half. Distal incisor region of mandible
(Fig. 69k) irregularly dentate-crenate. Abdomen
subequal in length to postorbital carapace length.
Cephalic section of telson with two spines in
each caudolateral corner; distal podomere of
lateral ramus of uropod rounded with indistinct
median ridge never reaching distal margin;
distolateral spine of proximal podomere equal
in length to other distal spines; mesial ramus
of uropod with distomedian spine never reaching
distal margin. Chela of male I (Fig. 69j) long,
narrow, densely tuberculate, approximately 3.2
times as long as broad; mesial surface of chela
with mesialmost row of tubercles 7 to 9; fingers
slender; carpus shallowly furrowed dorsally; basis
of cheliped without mesial spine. Male with
simple hooks on ischia of third and fourth
pereiopods extending proximally over basioischial articulation (Fig. 69h); prominent
caudomesial boss on coxa of fourth pereiopod.
-
106
First pleopods of first form males (Fig. 69a,f)
asymmetrical, reaching coxopodites of third
pereiopods with abdomen flexed and terminating
in five distinct elements; mesial process long and
slender, subcylindrical or somewhat flattened
and directed distolaterally; cephalic process
bladelike, acute, directed caudodistally, and
hooding base of central projection; central
projection most conspicuous of terminal
elements, strongly corneous, acute, bladelike,
compressed laterally and directed caudodistally;
caudal process also sclerotized, flattened and
tapering distally; caudal knob situated at lateral
base of cephalic process and covered with
subapical setae; setae largely obscuring cephalic
process and much of central projection; first
pleopod of Form I1 males illustrated in Fig. 69b,e.
Annulus ventralis (Fig. 69d) subovate with
greatest length in transverse axis, longitudinal
S-shaped medial furrow flanked by prominent
dextral elevation near midlength; tuberculate
caudal margin of sternum immediately cephalic
to annulus, overhanging cephaloventral region
of latter; postannular sclerite approximately twothirds as broad as annulus.
Hobbs ( 1981:376) discussed
morphological variations of this species in
Georgia; many of those he mentioned are present
in Wisconsin populations: position and convergence in marginal spines of the rostrum, length
of acumen, width of areola, antenna1 scale length,
and annulus ventralis.
VARIATIONS:
COLOR NOTES (refer to Fig. 70). Carapace tan,
orange, or scarlet dorsally, fading laterally to
slightly lighter shades. Dark brown or black flecks
and paler brown splotches present over entire
carapace; small tubercles covering lateral portion
of carapace cream; margins of rostrum cream
dorsally, brown laterally and ventrally. Abdomen
with broad, longitudinal, dorsomedian, brown
stripe narrowing posteriorly, dark anteriorly and
generally fading posteriorly; each tergum,
however, darker at anterior end with caudal
margin pale tan, cream, or pink; paired tan to
pinkish longitudinal stripes flanking median dark
one; pleura dark brown, red, or dark orange,
presenting overall appearance of scalloped, dark
stripe. Telson about same color as carapace with
brown to black flecks and irregular dark tan
splotches. Antennular and antenna1 peduncles
pinkish tan with dark brown markings. Dorsal
surface of ischium of third maxilliped pinkish
cream distally and splotched with tan. Dorsal
surface of cheliped from distal part of basis to
pale tips of fingers pinkish tan, orange, or scarlet,
with irregular light brown to almost black
markings. Most tubercles on palm of chela and
carpus very dark. Remaining pereiopods pinkish
tan with irregular darker bands and splotches;
sternal area of cephalothorax cream to pinkish
cream.
In 1969 Dowel1 and Winier reported that
occasional blue specimens of this species were
found. Smiley and Miller (1971:221) noted that
in a population of P. a. acutus in Fairbanks, La.
blue color morphs occurred
. . only about 1
in 50,000 . .
Black (1975) conducted breeding
".
. ."
TABLE 15. Range of measurements (in mm) of various diagnostic structures of Wisconsin
Procambarus a. acutus.
N
Minimum
Maximum
Mean
Standard
Deviation
4-
16
16
16
36.5
28.4
7.9
54.9
44.2
25.1
43.3
34.3
18.7
5.2
4.5
3.8
16
16
12.3
1.o
20.8
2.2
15.6
1.5
2.2
0.3
15
15
9.7
8.1
21.1
16.1
13.8
11.6
3.0
2.3
15
4.2
14.9
10.9
2.3
37
38
38
26.2
20.4
9.5
43.5
38.7
19.2
34.9
27.1
14.8
4.4
4.3
2.2
38
38
9.3
0.5
15.2
2.0
11.9
1.2
1.6
0.2
20
20
4.2
3.2
10.7
9.6
6.9
6.2
1.8
1.8
24
31.9
23.3
13.9
52.7
43.3
23.5
41.7
32.3
18.5
4.1
24
10.0
1.0
20.0
2.4
14.7
1.5
2. I
0.3
7
7
7.3
7.9
10.0
8.5
9.4
1.o
1.1
MALES ($1)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
Pleopod
Length
-
MALES ($11)
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela .palm
Length
Width
FEMALES
Carapace
Total length
Postorbital length
Height
Areola
Length
Width
Chela palm
Length
Width
24
24
24
4.4
2.3
~
11.1
107
Figure 69. Procambarus (Ortmannicus) acutus acutus (b,e, second form male Oak Creek, Milwaukee County; d, adult female
Root River, Racine County; all others first form male Root River, Racine County): a,e, mesial view of first p l e o p d ;
c, lateral view of carapace; d, annulus ventralis; b,f, lateral view of first pleopod; g, dorsal view of carapace; h, proximal
podomeres of third, fourth, and fifth pereiopods; i, antenna1 scale; j, dorsal view of distal podomeres of cheliped; k,
incisor margin of right mandible; I, epistome.
108
experiments with P. a. acutus and concluded that
t h e passages leading from them uniting 15 t o
the blue color morph (variant) is due to a
35cm beneath the surface. The tunnel extends
mutation of the genes responsible for pigment
development and behaves as a simple recessive
t o its normal allele. No blue variants of this
species have been reported in Wisconsin.
vertically down for another 30 t o 5Ocm and
terminates in a slightly expanded chamber which
is usually filled with water and loosely packed
mud. Males have been collected that were
situated higher in the burrow t h a n females and
vice versa; n o general trend exists. Commonly,
both are tightly encased in mud at or near t h e
terminus of the burrow. It is assumed that some
members of a population copulate within the
burrow habitat.
This species is quite “plastic” in its habitat
requirements, demonstrating a large range of
tolerance for factors as pH, pollution, temperature, vegetation, turbidity, and bottom composition (Francois 1959:112). In Illinois Brown
(1959) found this species in waters with t h e
following characteristics:
TYPE-LOCALITY: Tributary t o Mobile River,
Kemper County, Mississippi,
USA.
In Wisconsin, P. a. acutus
has been taken from ponds, creeks, rivers, lakes,
ditches, sloughs, temporary pools, and burrows.
W e collected it from waters varying from no
t o slow flow, that were usually moderately turbid
t o very turbid, with t h e substrate consisting of
mostly silt, “muck,” and sometimes gravel. Some
of the plants observed where this species ’was
collected are: various sedges, Nuphar sp., Elodea
sp., Typha sp., and Potamogeton sp. Individuals
burrow in response to dessication of the water
source but limited data suggest that during t h e
fall, breeding males and females pair and enter
a burrow for the winter (=tertiary burrower).
T h e burrow generally has two entrances, both
usually capped with mud (no chimney or pellets),
ECOLOGY. Habitats.
PH
Dissol. oxygen
Dissol. COZ
Total alkal.
6.3 - 8.2
1 12mg/l
0 - 50mgil
50 - 463mg/l
-
mean
mean
mean
mean
7.2
5.0 mg/l
10mg/l
190mg/l
Figure 70. Photograph of Form I male of Procambarus (0.)
a. acutus dug from burrow, Kenosha County.
109
I
Abbott (1873:80) stated that individuals in
New Jersey are found on dense vegetation usually
near the surface of running streams. Hobbs and
Hall (1974:199)noted that very large populations
occur in roadside ditches heavily laden with clay
particles, with the secchi disk disappearing at
5cm below the surface. Meyer (1965) studying
a population from a pond in Arkansas found
individuals heavily infested with eggs of the
corixid, Ramphocorixa acuminata. Hinkelman
(1970) indicated “P, blandingii acutus” occurs
in weedy, near-stagnant water, muddy creeks,
clear, gravel bottom streams, and burrows in
stream banks in Wisconsin. Penn and Hobbs
(1958:466), based on extensive collections in
Texas, stated that this crayfish I ‘ . . . occurs more
frequently in shallow water, i.e. less than 15
inches deep (54%), that is turbid (52%),
permanent (77%), flowing (54%), and exposed
to full sunlight (71%) ... with mud bottoms
(62%), and without aquatic vegetation (57%).”
Hobbs and Marchand (1943) found this species
in woodland swamps, bogs, swiftly flowing
streams, creeks, rivers, ditches, and ponds of
western Tennessee. Meredith and Schwartz
(1960) even found it in burrows in Maryland
salt marshes. Phillips (1980) found P. a. acutus
in Iowa bottomlands associated with t h e
Mississippi River and suggested that its distribution may have climatic rather than habitat
limitations.
Brown (1981), using horizontal starch gel
electrophoretic techniques, found that P. a.
acutus was polymorphic at four of the 19 loci
(21.1%) tested. Compared with five other
crayfish species in South Carolina, this species
was the most heterozygous, correlating well with
its having one of the most extensive geographic
and habitat ranges. This study further substantiated previous reports of low genetic variability
in decapod crustaceans as compared with other
invertebrate and vertebrate organisms (Powell
1975, Nevo 1978).
Physicochemical parameters. This is a ubiquitous species with a wide tolerance for numerous
environmental parameters (eurytopic). For
example, Huner (1984) found that under
laboratory conditions P. a. acutus, a “warm110
water” species, is more tolerant of dissolved
oxygen stress than a “coolwated’ species, Astacus
astacus. Because of its tolerant (adaptable)
characteristics, P. a. acutus has been highly
successful, as can be noted by its widespread
geographic distribution (Fig. 72).
LIFEHISTORY. For a species as widely distributed
as P. u. acutus, it is somewhat astonishing that
so little is known concerning its life history; life
history data are summarized in Table 16.
Breeding males are found throughout the year
(from April to November in Wisconsin probably
limited by lack of collections during the cold
months). Females with eggs attached have been
observed from March until December and
females carrying young have been reported from
May until October in various parts of North
America. Albaugh (1973) noted that sperm may
remain viable for several months as females
produced viable offspring following confinement
within a laboratory aquarium for seven months.
It is highly probable that following copulation,
some females overwinter in burrows and in the
spring lay and thus fertilize their eggs. Turner
(1926) was the first to suggest that there is no
restricted “breeding season” for this species. He
based his remarks on Ohio populations, but this
has been noted in other parts of its range (Creaser
1932 Wisconsin, Phillips 1980 Iowa). In
contrast, Dickson and Giesy (1982) noted that
the major breeding season is in the fall in South
Carolina (based on the relative abundance of
Form I males). They determined the seasonal
variability of phosphoadenylate concentrations
and adenylate energy charge in the “dorsal tail
muscle.” They found that seasonal peaks of tail
muscle ATP concentration coincided with the
population breeding period, possibly reflecting
greater energy production associated with
increased muscular activity involved in mating
and agonistic encounters. Penn (1956) noted
that copulation probably began early in the year
and peaked during June and July in Louisiana.
Laboratory studies (Thorp and Ammerman,
1978) showed that individuals were significantly
more agonistic (meral and pincer spread - see
Hayes, 1977 for a discussion of behavior of this
-
-
-
species, particularly with reference to response
to a predator) when receiving chemical information from a stressed conspecific member than
when inflow water lacked this chemical cue.
They suggested the “ecological importance” of
pheromones is species-specific in crayfishes and
one can only speculate on the value of this
recognition strategy in the reproductive processes of this and other crayfishes.
Although no distinct molting period(s) is
known for this species, Thorp and Wineriter
(1981) reported the molting frequency for
juveniles is directly related to temperature.
Romaire et al. (1977) not only reported that
growth is allometric ‘‘. with weight increasing
faster than the cube of the length,” but they
also provided regression equations for Form I,
Form I1 males, and females.
..
DISTRIBUTION: WISCONSIN: The distribution of
this crayfish in Wisconsin is discontinuous (Fig.
71), it being known from t h e Chippewa,
Trempeleau-Black, Wisconsin, PecatonicaSugar, Rock-Fox, Menominee-Oconto-Peshtigo,
Fox-Wolf, and Lake Michigan drainages (see
Appendix I). Creaser (1932) noted it from several
localities in the Wisconsin River, and it is
probable that P. (0.)
a. acutus still resides in
that drainage. Threinen (195813) stated that this
adaptable “marsh crab” is found up t h e
Mississippi River as far north as Lacrosse, but
Phillips (1979, 1980) showed it slightly farther
to the north in Pierce County. Additional
collecting efforts, particularly in ditches and
temporary pools, will probably extend its range
farther to the north and west.
UNITEDSTATES:
P. (0.)a.
acutus may be the
most adaptable of all North American crayfishes
(Pickett and Sloan 1979), and its geographical
distribution is known from the “Coastal plain
TABLE 16. Life history data for P. (0.)
a. acutm in Wisconsin (** from literature and from
our collections or USNM collections; those without asterisk(s) from USNM collections only).
Jan.
Feb.
March
~
~~
April
“Wis.
May
“Wis.
“Wis.
June
“Wis.
Wis.
July
“Wis.
“Wis.
“Wis.
Wis.
Sept.
“Wis.
Wis.
Oct.
“Wis.
Wis.
Nov.
“Wis.
Aug.
~
“Wis.
~
Dec.
111
/
I
O
W
LAKE
SUPERIOR
A
Figure 71. Distribution of Procambarus (0.)a. ucutus, Wisconsin; closed circles-specimens
circles-from literature.
112
examined in this study; open
and piedmont from Maine to Georgia, from
Florida panhandle to Texas, and from Minnesota
a.] cuevachicae
to Ohio; intergrades with [P. (0.)
in southwestern Texas and northern Mexico”
(Hobbs 197413353). Its geographical distribution
is shown in Fig. 72. The introduction of this
species into San Diego County, California was
first noted by Gander (1927) and Bouchard
(1977b)further remarked on its occurrence there
but failed to locate any populations. Crocker
(1979) reported P. (0.)
a. acutus to have been
introduced into Maine and stated that its
presence in Massachusetts and Rhode Island
represent natural expansions, except where it
was introduced (Smith 1982) into west-central
Massachusetts. Hobbs (1981) indicated that it
was introduced into the Georgia Piedmont
Province in Meriwether County.
CRAYFISHASSOCIATES.
C. diogenes, 0. virilis, P.
gracilis.
Figure 72. Geographic distribution of Procambarus (0.)
acutus acutus.
113
PALAEMONIDAE
Palaemonetes kadiakensis Rathbun
(Figures 73 77)
Palaemonetes kadiakensis - Rathbun 1902:93,
Palaemonetes exilipes
Creaser 1931b:273;
1932:321, 332, 333, 334, 336; Creaser and
Ortenburger 1933:45.
Palaemonetes (Palaemonetes) kadiakensis Holthuis 1952:216, 217.
Palaemonetes kadiakensis - Phillips 1980:91.
-
DIAGNOSIS (see Table 17): Carapace (Fig. 73a)
vaulted, bearing distinct antennal and branchiostegal spines, latter lying ventral to branchiostegal groove and reaching to, or slightly
anterior to, margin of carapace, Rostrum (Fig.
73a-h) compressed, straight, reaching to distal
tip of scaphocerite (= antennal scale); upper
margin somewhat convex, bearing 5 t o 8,
generally 7, evenly spaced teeth; proximal-most
tooth always situated posterior to orbit of
carapace; ventral surface of rostrum usually
bearing 2, occasionally 3 (sometimes 0), teeth
situated in distal third of rostrum. Firhpereiopod
\
(Fig. 73a) distinctly shorter than seconkwith
fingers of chela equal in length to that of palm;
finger of second pereiopod three-fourths length
of palm, opposable margins lacking teeth;
propodus of fifth pereiopod with transverse row
of setae on posterodistal surface. Telson (Fig.
13a) with two pairs of dorsal spines; position of
anterior pair variable but usually located
posterior to midlength and posterior pair situated
near posterior margin of telson. Appendix
interna absent only on first pleopod; appendix
masculina (second pleopod Fig. 73i-k; Table
17) distinctly overreaching appendix interna;
appendix masculina with one to three subapical
seta proximal to three apical setae.
-
TYPE-LOCALITY:Kodiak Island, Alaska (dubius
locality see Holthuis 1952217).
ECOLOGY. Palaemonetes kadiakensis is one of
three members of the genus found in epigean
fresh waters of the United States (Strenth 1976);
three additional species assigned to this genus
are known from hypogean waters in Florida (one
species) and Texas (two species). As is the case
in Iowa (see Phillips 1980),P. kadiakensis is found
primarily in the Mississippi River in Wisconsin.
In addition it has been taken from the St. Croix
River and from the Wolf River, the latter
representing a disjunct population (possibly
established subsequent to introduction). It is
typically found among aquatic vegetation in
sluggish regions of lotic systems, the above Wolf
River locality being a lagoon used as a boat harbor
supporting
Lemna minor L., Ceratoghyllum
\.
demersum L., and Elodea canadensis Michx. (Fig.
77). Q.caser (1931b, 1932) indicated that this
species prefers slowly moving rivers and streams,
or lakes and ponds with dense vegetation and
attains maximum numbers in backwaters of
TABLE 17. Range of measurements (in mm) and numerical variation of certain diagnostic
structures of Wisconsin Palaemonetes kadiakensis.
N
Minimum
Maximum
Mean
11
11
11
4.43
6
1
6.34
5.12
No. dorsal rostral teeth
No. ventral rostral teeth
FEMALES
Postorbital carapace length
No. dorsal rostral teeth
No. ventral rostral teeth
No. eggslfemale
13
13
13
3
4.63
5
MALES
Postorbital carapace length
114
8
7
3
2
8.24
6.35
7
2
79
0
8
2
49
119
Figure 73. Palaemonetes kadiakensis (a,e,f,j, from males a t Ferry Lake, Mississippi River, Grant County; g,h, from female
Glen Lake, Grant County; b,i,k, from males and c,d, from females Guth’s Harbor, Wolf River lagoon, Waupaca County):
a, lateral view of shrimp; b-h, lateral view of rostrum; i, second pleopod; j,k, distal portion of appendix masculina.
115
rivers where fish populations are small. Creaser
and Ortenburger (1933)reported that the shrimp
was found in the Poteau River in LeFlore
County, Oklahoma, where there was a moderate
current and a pH of 7.5. Meehean (1936a)
considered this shrimp to be “fairly abundant”
in quiet water in the environs of Natchitoches,
Louisiana. He observed them among the
emergent vegetation, on the branches of
submerged trees, or clinging under the duckweed
at the surface of ponds. In high water he noted
shrimp in inundated terrestrial vegetation along
the shore, especially among high grass and bushes;
they were also collected among the floating
vegetation in lakes. Pelton (1954) noted that
in Ohio this crustacean was particularly
abundant in pools where leaves and sticks provide
cover (see also Gebhart 1936). Page (1974,1985)
described the habitat of the species in Illinois
as sluggish water, and Phillips (1980) reported
it to be abundant above navigation dams,
indicating that particularly large numbers of
specimens were found associated with Potumogeton crispus L, He suggested that the absence
of large, sluggish rivers with dense stands of
aquatic vegetation is probably the factor limiting
the distribution of this species in Iowa. Bouchard
and Robison (1980) provided data for Arkansas
populations and Nelson and Hooper ( 1 9 8 2 ~ 8 3 )
conducted thermal preference experiments on
P. kudiakensis from Monroe Co., Michigan.
LIFE HISTORY. Only scattered data are available
for this widely distributed shrimp (Table 18).
Shelford (1913:129) noted t h e springtime
occurrence of females with eggs in Illinois and
Creaser (1931133274) indicated that in Michigan,
females carry eggs in April, July, and August.
Creaser (1932:334) stated that females with
attached eggs are found in the spring and fall
in Wisconsin, suggesting two breeding seasons
per year. During this investigation we observed
ovigerous females in June. The most complete
accounts for the species are presented by
~~~
~
~
~
TABLE 18. Life history data for Palaemonem kadiakensis (from literature except for the June
entry).
Month
6
copul
9e
Jan.
Feb.
March
April
Wis.
May
Wis.
June
Wis.
July
Wis.
Aug.
Wis.
Sept.
Oct .
Nov.
Dec.
116
Wis.
9y
YOY
Meehean (1936a,b), White (1949), by Broad and
Born (1963),and by Nielsen and Reynolds (1977).
In Louisiana, Meehean (1936a,b) reported the
life span to be one year. The largest individuals
( >40mm total length), observed in late March,
were ovigerous females. He reported that the
number of eggs varies from about 40 to 160,
the average number between 50 and 80 and is
dependent upon the size of the individual.
Within a few days of the late March date the
large-size-class individuals disappeared from the
population. The average size continued t o
decrease until ovigerous females as small as 23mm
total length were observed in July; no adults with
eggs were subsequently seen. Thus, by August
no adults remained and individuals of the
youngest cohort ranged in total length from 1225mm.
White (1949) studied P. kadiakensis in
Louisiana, making “regular collections” from
lakes in the environs of Baton Rouge in order
to study gametogenesis. Ovigerous females were
collected from early February to late October
(ova production ceased from November to
January). White assumed external fertilization
and noted that females are capable of producing
two (possibly three) broods “in a season.” The
eggs are carried 12-16 days, after which hatching
occurs. The female usually molts within 24 hours
of hatching.
Broad and Born (1963523) discussed the life
history of this shrimp in a northern Ohio
impoundment. Individuals spawn from late May
until August, with an obvious pulse in mid-June.
Eggs are laid and hatch shortly thereafter, with
the major annual population recruitment of
juveniles occurring in July. Young-of-the-year
undergo rapid growth through October and by
the latter part of that monthgrowth ceases. They
remain active during the winter but growth does
not resume until April. By June (following
copulation) males become relatively rare in the
population, and by August, year-old individuals
of both sexes are scarce; however, some survive
to spawn the second year. Although data are
few for Wisconsin populations, it is presumed
that the life history for those is somewhat similar
to that of Ohio shrimps, with the major
difference being that in Wisconsin ovigerous
females appear and remain in populations slightly
later than those in Ohio. Much remains to be
learned of the life history of this shrimp (see
Table 18 for a summary of life history data).
Nielsen and Reynolds (1977) studied populations of P. kadiakensis from three ponds in
Boone County, Missouri. Individuals began
breeding in mid-May when waters reached 1520°C and the process terminated in August. They
suggested that cessation of breeding resulted from
post-reproductive mortality of breeders rather
than from direct environmental stimuli, and they
noted that shrimp generations were essentially
non-overlapping. They reported a mean fecundity of 40 eggs (range 20 76) for females (2536mm in total length). A number of females
collected from the Guth’s Harbor locality in
Wisconsin in June 1984 were ovigerous and
carried a varied number of oval eggs: 49 eggs
on female with POCL of 5.82mm eggs 1.12
x 0.85mm; 68 eggs on female with POCL of
5:81mm eggs 1.12 x 0.93mm; 119 eggs on female
with POCL of 8.24mm eggs 1.21 x 0.84mm.
Nielsen and Reynolds (1977) proposed that
females are capable of producing multiple broods
in the same year (ovigerous females were
collected bearing maturing ovarian eggs). Growth
in length was not constant but exhibited three
distinct phases. Shrimp grew rapidly from
hatching to about 50% of ultimate length in
three months (mid-June through mid-September). Twenty percent of ultimate length was
added in six months, and rapid growth resumed
in April. By June shrimp attained maximum
length. In all populations, the average length and
weight of females consistently exceded that of
males. They also noted that P. kadiakensis
demonstrates a latitudinal gradient toward longer
growing seasons and larger ultimate sizes in
southern habitats. Nielsen and Reynolds (19756)
reported summer densities to range from 72 to
340 individuaMm2.
Molt cycle. Also, very little is known about
the molting cycle of this crustacean. Meehean
(1936a) noted that adult females molted within
three days (generally within 24-hrs) after the
young are released. The newly hatched, first stage
-
-
-
-
117
larvae molt 12 to 36 hrs after hatching, and
subsequent development is discussed by Broad
and Hubschman (1963). They report, as do
Hubschman and Broad (1974), that the total
length of P. kadiakensis at hatching is 4.4mm.
They indicate that there are 5-8 larval molts
prior to metamorphosis which occurs when a
total length of 7.5mm is attained, Virtually no
other data are available concerning molting
habits.
Stoffel and Hubschman (1974),in a laboratory
study of an Ohio population, demonstrated that
the loss of four pereiopods induced molting.
Previously it had been demonstrated that several
species of Reptantia (including “C. propinquus”)
respond to limb loss by molting; that is, the
molting process is accelerated by loss of limbs
(see Zeleny 1905; Bliss 1956, 1959; and Skinner
and Graham 1970). Stoffel and Hubschman were
the first to show the same reaction for Natantia.
They suggested that pereiopod loss stimulates the
neurosecretory cells of the X-organs via nervous
impulses to stop releasing the molt-inhibiting
hormone.
Of note, a discussion of the trematode infestation of P. kadiakensis in Louisiana is presented
in Font and Corkum (1975, 1976).
Further information for the genus Palaemonetes can be obtained from Beck and Cowell
(1976) and Kushlan and Kushlan (1980) who
discuss life history data on the closely related
P. paludosus (Gibbes) in Florida.
COLORNOTES. Individuals are translucent with
a slight pinkish, yellowish, or tannish hue due
Figure
118
to numerous chromatophores present over entire
body (Fig. 74). Eyes dark brown to tan. Eggs
attached to pleopods pale green to lime green
in color.
DISTRIBUTION: WISCONSIN (Fig. 75) - Palaemonetes kadiakensis is known from the St. Croix,
Trempealeau-Black, Wisconsin, and PecatonicaSugar watersheds in the Mississippi drainage basin
and only from the Fox-Wolf system in the St.
Lawrence drainage (see Appendix I). It has been
collected in only the following counties in the
state: Buffalo, Crawford, Grant, Pierce, St.
Croix, Vernon, Waupaca.
UNITED STATE. Fresh waters of central U.S.A.
west of the Alleghenies from the Great Lakes
south to the Gulf Coast (Alabama,Louisiana,
Texas); also in N.E. Mexico and S. Ontario,
Canada (Holthuis 1952, Bouchard and Robison
1980). Bell (1971:lO) reports this species also to
be present in Vermont. Strenth (1976) pointed
out that the members of the genus Palaernonetes
are not territorial and proposed that this factor
may possibly explain the current geographical
distribution of the genus. He suggested that the
distribution is the result of competitive exclusion
of a once widespread genus by more recent,
successful genera (e.g., Macrobrachiurn); widespread introductions also have probably occurred, although this has been documented only
in central Missouri (Nielsen and Reynolds 1977).
The known geographical distribution for this
shrimp is shown in Fig. 76.
74. Palaemonetes kadiakensis (female with eggs) from Guth’s Harbor, Waupaca County.
Figure 75. Distribution of Palaemonetes kadiakensis in Wisconsin; closed circles-specimens
circles-from literature.
examined in this study; open
119
Figure 76. Geographic distribution of Palaemonetes kadiakensis.
Figure 77. Guth’s Harbor, Wolf River lagoon, Waupaca County. Habitat of
PalaemoneteA kadiakensis.
120
LAKE
SUPERIOR
SUPERIOR
Sayfield
I
O
W
C
!
A
meen
Rock
'
WnLVorih
I
,
~
Kenosha
1
Figure 78. Map of Wisconsin illustrating location of counties.
121
GLOSSARY
(Utilize figures for clarification of
morphological characters)
ABDOMEN. Region of body posterior t o
cephalothorax, consisting of six body segments
and telson (see Fig. 13).
ACUMEN. Pointed apical part (tip) of rostrum,
frequently delineated basally by marginal
spines (see Fig. 14).
ALLOCHTHONOUS MATERIALS. Various
forms of organic matter produced within
catchment basin and brought into stream,
pond, or lake.
ANNULUS VENTRALIS. Crayfish seminal
receptacle, consisting of median sclerite
between fourth and fifth pereiopods; the
spermatophore receptacle on sternum of
female (see Fig. 23d).
ANTENNA. Whiplike, generally long sensory
organ arising from anterior region of cephalothorax (see Figs. 13,14).
ANTENNAL SCALE. Bladelike exopod of
antenna situated at base of antenna (see Figs.
13,14).
ANTENNAL SPINE. Spine on, or adjacent to,
anterior margin of carapace and ventral to
orbit (see Fig. 13).
ANTENNULAR PEDUNCLE. Proximal segments of antennule, from which flagella arise.
ANTENNULE. One of pair of appendages of
first cephalic somite; “first antenna” (Figs.
13,14).
ANTERIOR. Front end of organism; “head end.”
APPENDIX INTERNA. Slender, rodlike structure on mesial border of endopod of second
through fifth pleopods of shrimps.
APPENDIX MASCULINA. Lobe or rodlike
structure, bearing setae, situated between
appendix interna and mesial margin of
endopod of second pleopod of male shrimps.
The presence or absence of this structure
provides easiest means of distinguishing males
from females; a n important taxonomic
character for palaemonids (see Fig. 73j,k).
APICAL. Apex; tip.
AREOLA. Dorsomedial area (usually hourglass-
shaped) of thoracic region of carapace of
crayfishes, bounded laterally by paired arched
(branchiocardiac) grooves delineating
dorsomedial limits of gill chamber (see Fig.
14).
AUTOCHTHONOUS MATERIALS. Various
forms of organic matter produced within water
body where found (lake, stream, pond).
BASIS. Second segment (from proximal end) of
segmented appendage.
BENTHOS. “Bottom” of a body of water (e.g.,
lakes, streams, and ponds), providing habitat
for various aquatic organisms on or within.
BOSS. An expanded portion (rounded protuberance) on mesial surface of coxa of fourth
pereiopod of male crayfishes (see Fig. 14).
BRANCHIOSTEGAL SPINE. Short spine
situated on or near anterior margin of
carapace, ventral to antenna1 spine in shrimps;
in crayfishes, located immediately ventral to
anterior extremity of cervical groove (see Fig.
13).
BURROWER: Crayfishes (usually all) that spend
some or nearly all of their life history in
confines of excavated burrow.
PRIMARY. Crayfishes that spend almost
entire lives in subterranean galleries.
SECONDARY. Crayfishes that spend much
of their lives in burrows but frequently move
into open water during rainy seasons.
TERTIARY. Crayfishes that live in open water
and retreat to burrows in response to several
factors: a) to remain below frost line during
winter, b) females enter as period of ovulation
approaches and remain in burrows to lay and
brood eggs, c) to find protective cover and
to avoid dessication as water bodies disappear.
CAMBRIAN. One of earliest geologic Periods
(began nearly 600 million years ago and lasted
approximately 70 million years), rocks in
which fossils of higher forms of life are fairly
abundant.
CARAPACE. “Shield” (exoskeleton) overlying
123
cephalothoracic somites of body (see Figs.
13,14).
CARAPACE LENGTH. In shrimps, distance
from posterior margin of orbit t o midcaudodorsal margin of carapace; in crayfishes,
distance from tip of rostrum to midcaudodorsal margin of carapace (see Fig. 14).
CARINA. Median middorsal ridge on rostrum;
oriented parallel to lateral margins of rostrum
(see Figs. 37,47).
CARPUS. Fifth segment from proximal end of
segmented appendage.
CAUDAL. Posterior (rear, tail) end of organism.
CENOZOIC. Geologic Era including Tertiary
and Quarternary Periods (65 to 2.5 million
years ago).
CENTRAL PROJECTION. Terminal projection
on crayfish male gonopod formed by fusion
of two processes; poorly developed in second
form male; located centrally on apex of
Procmnbarus gonopod, distally on that of
Cambarus, Fallicambarus, and Orconectes
(Figs. 23,32,63).
CEPHALIC. Pertaining to head; of head.
CEPHALOTHORAX. Portion of body bearing
eyes and all appendages through fifth pereiopod (fused head and thorax).
CERVICAL GROOVE. Major arclike suture
(groove) on carapace of crayfish, dividing it
into anterior (cephalic) and posterior (thoracic) regions (see Fig, 14).
CERVICAL SPINE. Spine on lateral surface of
carapace immediately posterior to cervical
groove of crayfishes (see Fig. 14).
CHELA. Forcepslike structure (“claw” or
“pincer”), consisting of two opposed distal
podomeres of first, second, and third
pereiopods of crayfishes, first and second
pereiopods of shrimps; dactyl (moveable
finger) and propodus (see Figs. 13,14).
CHELIPED. Pereiopod bearing chela (see Figs.
13,14), in crayfish literature applied almost
exclusively to first pereiopod.
COLLECTOR. Functional group category based
on feeding mechanisms into which immature
and adult aquatic insects are generally placed;
crayfishes herein are also assigned to this group
to indicate their functional role in aquatic
124
community trophic relations; that is, they
consume decomposing fine particulate organic
matter (FPOM) and associated microbes.
CONDYL. Knob; rounded process at base of
central projection of male first pleopod
(gonopod); “condyl length” as presented in
Tables 4,6,8,10is measured distance between
base of condyl and distal tip of central
projection (Fig. 32).
COPULATION. Joining of male and female for
transference of sperm; sexual union.
CORNEOUS. Structures that are horny (sclerified), particularly as related t o reproductive
appendages (modified first pleopod) of first
form males.
COXA. Proximal (first) segment of segmented
appendage.
DACTYL. Distalmost segment of usually 7segmented appendage; smaller, mesially
situated, and moveable part of chela (moveable finger) (see Fig. 14).
DEVONIAN. Geologic Period beginning 395
million years ago and lasting approximately
50 million years; “The Age of Fishes.”
DETRITUS. Non-living particulate organic
matter inhabited by decomposer organisms;
also, including dissolved carbon-containing
substances excreted by plants and animals as
well as soluble organic materials released from
decaying plant and animal tissues.
DISTAL. Away from body mid-line; toward apex
(tips) of appendages.
DORSAL. Top or back of shrimp or crayfish.
DRIFTLESS AREA. Unique feature in that area
(33,670 km2) of southwestern Wisconsin,
northwestern Illinois, northeastern Iowa, and
thin strip in southeastern Minnesota cloaked
by loess; although not formerly an island in
ice, it was not covered by glaciers during
Pleistocene and thus was affected by periglacia1 phenomena.
ENDITE. Mesial lobelike extension of podomere
or of axial part of unsegmented appendage.
ENDOPOD. Mesial ramus of biramus appendage,
originating on basal segment (basis) (see Fig.
13).
EPIGEAN. Referring to above ground habitats
as opposed to hypogean.
EPISTOME. Transverse plate (usually somewhat
triangular in shape) situated ventrally and
anterior to gnathal appendages of crayfishes
(see Fig. 23i).
EURYPHAGOUS. Employing wide range of
living and decaying plant and animal material
as food; “scavenger” strategy.
EXOPOD. Lateral ramus of biramus appendage,
originating on second segment (basis) from
base (see Fig. 13).
EXOSKELETON. Outer covering of arthropods;
“shell.”
EXTENSOR SURFACE. Unopposed surface
(face) of flexible podomere of an appendage
(in opposition to opposed, flexor surface).
FINGER. One of two rami of chela; movable
finger (dactyl) and immovable finger of
opposable part of propodus.
FIRST FORM MALE. Sexually functional male
crayfish (Cambaridae); at least one terminal
element of first pleopod usually corneous
(81).
FLAGELLUM. Ant ennal endopodit e; thin,
tapering, multiarticulate, elongate extension
of antenna and antennule (distal to peduncle).
FLEXOR SURFACE. Opposable surface (face)
of flexible podomere of appendage (in
opposition to non-opposed extensor surface).
GNEISS. Metamorphic rock characterized by
banded structure (due to separation of light
and dark minerals in crude layers).
GONOPOD. Pleopod in male modified for
reproductive purposes.
GYNANDROMORPH. Abnormal individual
exhibiting secondary characters of both sexes;
particularly well known in insects; occurs also
in Crustacea, birds, mammals.
HEPATIC AREA. Paired cephalolateral region
to either side of carapace between orbit and
cervical groove (in crayfishes) (see Fig. 14).
HEPATIC SPINE. Spine situated near anterior
margin of hepatic area of carapace of certain
shrimps, or on hepatic area of cambarids (see
Fig. 14).
HYPOGEAN. Subterranean; referring to beneath surface of earth; burrow or cave
environment.
HYPORHEIC ZONE. Interstices formed in
substrate of streams (between coarse sand,
gravel, and cobble).
INCISOR PROCESS. Cutting surface (lobe) of
mandible (as opposed to molar or grinding
lobe) (see Fig. 23h).
ISCHIOPODITE. Third segment from base of
a segmented appendage (see Fig. 13).
ISCHIUM. See ischiopodite.
KARST. Terrain (or topography) characterized
by numerous sinkholes, swallowholes, springs,
and caves formed by extensive dissolution of
carbonate rocks.
LATERAL. Referring to or toward side.
LENTIC. Denotes standing water environments
(e.g., lakes, ponds, bogs).
LIMNETIC. Upper circulating, open-water zone
of lakes beyond direct influence of shore or
bottom.
LITTORAL ZONE. Shore region of lake;
peripheral shallows of lake.
LOTIC. Refers to running water environments
(e.g., springs, rivers).
MANDIBLE. Most anterodorsally situated of
gnathal appendages; one, of a pair, of heavily
calcified jaws; the “teeth.”
MAXILLA. One of a pair of two sets of gnathal
appendages situated immediately posteroventral to mandibles.
MAXILLIPED. One of pair of three sets of
gnathal appendages lying immediately posterior to second pair of maxillae (see Fig. 14).
MERUS. Fourth segment from proximal end of
segmented appendage.
MESIAL. Referring to or toward middle.
MESIAL PROCESS. Terminal process located
mesially on modified first pleopod (gonopod)
of male cambarid crayfishes (Figs. 23, 32, 63).
MORPHOLOGY. Study of form and structure.
MOVABLE FINGER. (=dactyl); distalmost
podomere of segmented appendage.
ORBIT. One of pair of excavations on anterior
margin of carapace bordering eye (or
remnant).
PALEOZOIC. Second of five major subdivisions
of geological time scale; geological Era
beginning approximately 600 million years ago
and lasting until approximately 225 million
years ago.
125
PALM. Expanded portion of propodus of chela
ROSTRUM. Dorsomedian anterior projection of
situated proximal to “fingers” (Fig. 14).
PALP. Relatively slender 2- or 3-segmented
appendage of mandible.
PEREIOPOD (PEREOPOD). One of five pairs
of appendages (legs) supporting cephalothorax;
walking and chelate legs.
PLEISTOCENE. Earliest Epoch of Quarternary
Period beginning approximately 2.5 million
years ago and lasting until 10,000 years ago.
PLEOPOD. One of five pairs of appendages on
first five abdominal segments (“swimmerets,”
or modified into male gonopod ).
PLEURON. Lateral overhanging plate of
abdominal segments.
PODOMERE. Single segment of any appendage.
POSTANNULAR PLATE. Medially situated
sclerite (plate) immediately posterior t o
annulus ventralis.
POSTERIOR. Toward caudal or “hind” end.
cephalothorax, in crayfishes partly covering
eyestalks and bases of antennae and
antennules (Figs. 13, 14).
SCAPHOCERITE. Platelike exopod of antenna
of shrimps (see “Antenna1 scale”) (Fig. 13).
SCHIST. Metamorphic, crystalline rock formed
by intense heat, foliated, and cleavable into
many thin plates.
SCRAPER. Functional group category (see
“Collector”); herbivores that graze on algae
attached to stony or organic surfaces (periphyton feeders),
SECOND FORM MALE. One of two morphological forms of male cambarids; sexually
nonfunctional male ($11) lacking corneous
terminal element on first pleopod (gonopod),
SHREDDER. Functional group category (see
“Collector”); herbivores and detritivores that
masticate coarse organic particulates
POSTORBITAL CARAPACE LENGTH.
(CPOM).
SILURIAN. Geologic Period beginning 430
million years ago and ending 395 million years
Distance from orbit t o mid-caudodorsal
(posterior) margin of carapace.
PRECAMBRIAN. First (oldest) Period of
geological time; preceeding Paleozoic Era
(Archaeozoic and Proterozoic Eras) and
extending up to approximately 600 million
years ago (origin of earth to beginning of
Cambrian Period represents approximately
88% of geologic time).
PROPODUS. Penultimate segment (sixth from
base) of segmented appendage.
PROXIMAL. Toward the body; that portion of
appendage nearest body.
PTERYGOSTOMIAN REGION. Paired anteroventral area of carapace ventral to antenna1
region.
PTERYGOSTOMIAN SPINE. Spine (in
shrimps) located on anteroventral margin of
carapace and ventral to branchiostegal spine.
PUNCTATION. Small depression (pit).
QUARTERNARY PERIOD. Most recent
(youngest) Period (2 million years ago to
present); subdivided into Pleistocene and
Recent Epochs.
-
126
ago,
SPELEAN. Referring to subterranean habitats
(caves).
STADIA. A stage in the life history of an
organism; interval between successive molts.
STERNUM. Midventral platelike surface of
body, situated between coxae of appendages.
SUPRAORBITAL SPINE. Spine on carapace of
shrimps, situated posterodorsal to orbit.
TELSON. Terminal (posterior) portion of
abdomen; caudomedian element of “tail fan”
(see Figs. 13,14).
TROGLOBITE. Obligate inhabitant of subterranean habitats (caves).
TUBERCLE. Low protuberance on exoskeleton.
UROPOD. Paired biramous appendage on sixth
abdominal segment, lateral parts of “tail fan”
(see Figs. 13,14).
VENTRAL. On or toward underside.
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140
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APPENDIX I
WISCONSIN DECAPODS AND
THEIR COUNTY AND
DRAINAGE OCCURRENCES
5. Orconectes (Procericambarus) rusticus (Gi-
List of Species:
I. Cambarus (Lacunicambarus) diogenes GirardAshland, Barron, Bayfield, Brown, Buffalo,
Burnett, Chippewa, Clark, Columbia,
Crawford, Dane, Dodge, Door, Douglas,
Dunn, Eau Claire, Forest, Grant, Green,
Green Lake, Iowa, Jackson, Jefferson,
Kenosha, La Crosse, Lafayette, Manitowoc,
Marathon, Milwaukee, Monroe, Oneida,
Outagamie, Ozaukee, Pepin, Pierce, Polk,
Portage, Price, Racine, Rock, St. Croix,
Sauk, Sawyer, Sheboygan, Trempealeau,
Vernon, Vilas, Walworth, Washburn,
Washington, Waukesha, Winnebago, and
Wood
2. Fallicambarus (Creaserim) fodiens (Cott1e)Dane (?)
3. Orconectes (Gremicamburus) immunis
(Hagen)-Bayfield, Burnett, Dane, Dodge,
Door, Fond du Lac, Grant, Green, Jefferson,
Kenosha, Milwaukee, Ozaukee, Pierce, Polk,
Racine, Rock, Sheboygan, Vilas, Walworth,
Washburn, Waukesha, and Wood
4. Orconectes (Crockerinus) propinquus (Girard)-Ashland, Barron, Bayfield, Brown,
Burnett, Calumet, Columbia, Crawford,
Dane, Dodge, Door, Florence, Fond du Lac,
Forest, Grant, Green, Green Lake, Iowa,
Iron, Jackson, Jefferson, Juneau, Kenosha,
Kewaunee, Lafayette, Langlade, Manitowoc,
Marinette, Menominee, Milwaukee,
Monroe, Oconto, Oneida, Outagamie,
Ozaukee, Polk, Portage, Price, Racine,
Richland, Rock, Rusk, Sauk, Sawyer,
Shawano, Sheboygan, Taylor, Vernon, Vilas,
Walworth, Washburn, Washington, Waukesha, Waupaca, Waushara, and Winnebago
6.
7,
8.
9.
rard)-Adams, Barron, Bayfield, Burnett,
Chippewa, Columbia, Douglas, Florence,
Forest, Green Lake, Iowa, Iron, Jefferson,
Lafayette, Langlade, Lincoln, Marathon,
Marinette, Milwaukee, Oconto, Oneida,
Ozaukee, Polk, Portage, Racine, Rock, Rusk,
St. Croix, Sauk, Sawyer, Shawano, Sheboygan, Vilas, Walworth, Washburn, Mashington, Waukesha, Waupaca, and Winnebago
Orconectes (Gremicambarus) virilis (Hagen)All counties of the state
Procamburus (Girardiella) gracilis (Bundy)Kenosha, Milwaukee, Ozaukee, Racine,
Walworth, and Waukesha
Procamburus (Ortmannicus) acutus acutus
(Girard)-Buffalo, Crawford, Dane, Dodge,
Grant, Jefferson, Kenosha, La Crosse,
Langlade, Marinette, Milwaukee, Outagamie, Pierce, Racine, Richland, Rock, Sauk,
Vernon, Vilas, and Walworth
Palaemonetes kadia kensis (Rathbun)-Buffalo,
Crawford, Grant, Pierce, St. Croix, Vernon,
and Waupaca
List of Counties (abbreviations as follows: Cd
Cambarus (L.) diogenes Girard; Ff = Fallicambarus (C.) fodiens (Cottle); Oi = Orconectes (G.)
immunis (Hagen); O p = Orconectes (C.) propinquus (Girard); Or = Orconectes (P.) rusticus
(Girard); Ov = Orconectes (G.) virilis (Hagen);
Pg = Procamburus (G.) gracilis (Bundy); Pa =
Procambarus (0.)acutus acutus (Girard); Pk =
Palaemonetes kadiakensis (Rathbun)):
=
1. Adams - Or, Ov
2. Ashland Cd, Op, Ov
3. Barron Cd, Op, Or, Ov
4. Bayfield - Cd, Oi, Op, Or, Ov
5. Brown Cd, Op, Or
6. Buffalo Cd, Ov, Pa, Pk
-
-
-
141
7. Burnett - Cd, Oi, Op, Or, Ov
8. Calumet Op, Ov
9. Chippewa - Cd, Or, Ov
10. Clark Cd, Ov
11. Columbia Cd, Op, Or, Ov
12. Crawford Cd, Op, Ov, Pa, Pk
13. Dane Cd, Ff(?), Oi, Op, Ov, Pa
14. Dodge Cd, Oi, Op, Ov, Pa
15. Door - Cd, Oi, Op, Ov
16. Douglas Cd, Or, Ov
17. Dunn Cd, Ov
18. Eau Claire Cd, Ov
19. Florence * Op, Or, Ov
20. Fond du Lac Oi, Op, Ov
2 1. Forest - Cd, Op, Or, Ov
22. Grant Cd, Oi, Op, Ov, Pa, Pk
23. Green Cd, Oi, Op, Ov
24. Green Lake Cd, Op, Or, Ov
25. Iowa Cd, Op, Or, Ov
26. Iron Op, Or, Ov
27. Jackson Cd, Op, Ov
28. Jefferson Cd, Oi, Op, Or, Ov, Pa
29. Juneau Op, Ov
30. Kenosba Cd, Oi, Op, C)v, Pg, Pa
3 1. Kewaunee Op, Ov
32. La Crosse Cd, Ov, Pa
33. Lafayette Cd, Op, Or, Ov
34. Langlade Op, Or, Ov, Pa
35. Lincoln Or, Ov
36. Manitowoc Cd, Op, Ov
37. Marathon Cd, Or, Ov
38. Marinette Op, Or, Ov, Pa
39. Marquette Ov
40. Menominee - Op, Ov
41. Milwaukee * Cd, Oi, Op, Or, Ov, Pg, Pa
42. Monroe Cd, Op, Ov
43. Oconto Op, Or, Ov
44. Oneida - Cd, Op, Or, Ov
45. Outagamie Cd, Op, Ov, Pa
46. Ozaukee Cd, Oi, Op, Or, Ov, Pg
47. Pepin Cd, Ov
48. Pierce Cd, Oi, Ov, Pa, Pk
49. Polk Cd, Oi, Op, Or, Ov
50. Portage Cd, Op, Or, Ov
a
-
-
-
-
-
-
-
-
-
.
-
-
-
-
-
-
-
-
142
-
-
-
I
-
5 1 . Price Cd, Op, Ov
52. Racine Cd, Oi, Op, Or, Ov, Pg, Pa
53. Richland Op, Ov, Pa
54. Rock Cd, Oi, Op, Or, Ov, Pa
55. Rusk Op, Or, Ov
56. St. Croix Cd, Or, Ov, Pk
57. Sauk Cd, Op, Or, Ov, Pa
58. Sawyer Cd, Op, Or, Ov
59, Shawano Op, Or, Ov
60. Sheboygan Cd, Oi, Op, Or, Ov
61. Taylor Op, Ov
62. Trempealeau Cd, Ov
63. Vernon Cd, Op, Ov, Pa, Pk
64. Vilas Cd, Oi, Op, Or, Ov, Pa
65. Walworth Cd, Oi, Op, Or, Ov, Pg, Pa
66. Washburn Cd, 'Oi, Op, Or, Ov
67. Washington Cd, Op, Or, Ov
68. Waukesha Cd, Oi, Op, Or, Ov, Pg
69. Waupaca Op, Or, Ov, Pk
70. Waushara Op, Ov
71. Winnebago Cd, Op, Or, Ov
72. Wood Cd, Oi, Ov
-
-
-
+
-
r
-
-
I
-
-
List of Drainage Basins:
Mississippi River Basin
I. Chippewa Cd, Oi, Op, Or, Ov, Pa
2. Pecatonica4ugar Cd, Oi, Op, Or, Ov, Pa,
Pk
3. Rock-Fox Cd, Ff(?), Oi, Op, Or, Ov, Pg,
+
Pa
4. St Croix - Cd, Oi, Op, Or, Ov, Pk
5. Trempealeau-Black - Cd, Op, Ov, Pa, Pk
6. Wisconsin Cd, Oi, Op, Or, Ov, Pa, Pk
-
St. Lawrence River Basin
1. Fox-Wolf Cd, Op, Or, Ov, Pa, Pk
2. Lake Michigan Cd, Oi, Op, Or, Ov, Pg,
Pa
3. Lake Superior Cd, Op, Or, Ov
4. Menominee-Oconto-Peshtigo Cd, Op, Or,
Ov, Pa
-
-
-
-
APPENDIX 11
Maps computer-generatedby the University of Wisconsin Cartographic Laboratory, 1984.
143
144
StT
Cambarus diogenes
146
147
148
149
150
15 1
152
153
154
155
156
157
158
159
160
161
162
163
164
Orconectes virilis
-
165
166
167
168
169
-----------------
Procambarus acutus
?IC*IE I 1.0e0.000
0
170
10
20
30 XILOMETERS
171
INDEX
anis, Ranunculus, 60,90
acuminata, Ramphocorixa, 110
acuta, Cambarus Blandingii, 105
acutus, Cambarus, 3,105’
acutus, Cambarus blandingi, 105
acutus,Cambarus blandingii, 105
acutus, Camburus (Cumbarus) blandingi, 105
acutus, Cambarus (Ortmannicus) blandingi, 105
acutus, Procambarus, 106
acutus, Procambarus acutus, 11, 24, 37, 50, 58, 96, 107 (Table
IS), 106, 107, 109, 110
LIcutus, Procambarus
blandingii, 105, 110
a t u s , Procambarus (Ortmannicus) acutus, 3, 5, 22, 23, 26 (Fig.
20), 102, 105, 106, 107, 108 (Fig. 69). 109 (Fig. 70), 110, 111
(Table 16), 112 (Fig. 71), 113 (Fig. 72), Appendix I(141, 142)
Aeshna species, 89
alba, Spirea, 74
americana, Vallisneria, 74, 90
amplifolius, Potamogeton, 74
angustifolius, Potamogeton, 74
argillicola, Cambarus, 3, 39
Astacidae, 21
astacus,Astacw, 110
Astacus, 110
astacUS,110
fodiens, 39
bartoni, Cumbarus, 3,63,64
bartonii, Cambarus (Cambarus) burtonii, 3
blandingii, Cambarus, 105
Branchiobdellid worm, 4
Burrows, 1, 6,8, 33, 34, 35, 36
Cambarus (Lacunicambarus) diogenes, 6, 8, 33, 34, 35, 36
Fullicambarus (Creaserius) fodiens, 41, 42
Orconectes (Oremicambarus)immunis, 47, 48
Orconectes (Oremicamburus) uirilis, 86
Procambarus (Girardiella) gracilis, 8, 100, 101 (Figs. 65, 66), 102
Procambarus (Ortmannicus) acutus acutus, 109
Cumbarellus (Dirigicambarus)shufeldtii, 56
Cambaridae, 4,9, 21, 22, 29
Cumbarus, 12,21,56
acutus, 3, 105
argillicola, 3, 39
bartonii, 3,63, 64
(Burtonius) diogenes, 29
blandingi acutus, 105
blandingii, 105
Blundingii acutu, 105
blandingii cutus us, 105
(Cambarus) bartonii bartonii, 3
(Cambarus) blandingi acutus, 105
(Cambum) diogenes, 29
(Cambarus)gracilis, 97
cowii ? ? , 80
debilis, 3, 79
Diogenes, 29
diogenes, 3, 4, 8, 24, 29, 30 (Table 2), 33, 35, 36 (Table 3),
37, 50, 59,64, 76,96, 102, 113
dwgenes diogenes, 29, 35
diogenes sspp., 29
(Faxonius) immunis, 43
(Faxonius) propinquus, 52
(Faxonius) rusticus, 80
(Faxonius) rusticus rusticus, 80
(Faxmius) uirilis, 80
fodiens, 39
grucilis, 3, 97
gracillis, 97
immunis, 43,47
(Lacunicambarus) diogenes, 8, 9, 21, 22, 23, 25 (Fig. 19), 29,
30, 31 (Fig. 23), 32 (Fig. 24), 33 (Fig. 25), 34 (Fig. 26),
35, 36, 37 (Fig. 271, 38 (Fig. 28), 39 (Fig. 29), 90 (Fig.
57), 91 (Fig. 59), Appendix I(191, 142)
Color Notes, 32, 33 (Fig. 25), 34 (Fig. 26)
Crayfish Associates, 37
Diagnosis, 29
Distribution, 37, 38 (Fig. 28), 39 (Fig. 29)
Ecology, 33,34,35
Life History, 35, 36 (Table 3), 37
Type-locality,29
Variation, 32 (Fig. 24)
(Lacunicumbarus) diogenes diogenes, 3,6
obesus, 3,29, 33
(Ortmannicus) blandingi m t u s , 105
placidus 3
propinquus, 3,52,72, 118
robustus, 60,63,64
wticus, 3,66, 79
stygius, 105
Stygius, 3, 105
uiriks, 79
uirilis, 3, 79
wisconsinensis, 3,80
Wisconsinensis, 80
canadensis, Anacharis, 1 I4
cutenutus, Sistrurus cutenutus, 41
Curex species, 60,90
173
Central Lowland Province, 13, 14 (Fig. 16)
Central Plain Province, 13, 15 (Fig. 17), 16, 17
Ceratophyllum species, 90
demersum, 1 14
Chara species, 74,90
Cladopohora species, 74
clarkii, Procambarus (Scapulicambarus), 58
Collecting techniques, 4, 9
Cambaridae, 4, 5 (Fig. 3),6, 7 (Figs. 8, 9)
Palaemonidae, 9
cordata, Pontederia, 74, 90
cousii ? ? , Cambarus, 80
crispis, Potamogeton, 74,90, 116
cuewachica, Procambarus (Ortmannicus) acutus, 113
gracilis, Cambarus (Cambarus), 97
gracilis, Procambarus, 8, 24, 37, 96, 97, 98 (Table 12), 101 (Table
141, 113
gracilis, Procambarus (Girardiella), 3, 7, 21, 23, 24, 26 (Fig. 20),
97, 98, 99 (Fig. 63), 100 (Fig. 64), 101 (Figs. 65, 66), 102,
103 (Table 14), 104 (Fig. 67), 105 (Fig. 68), Appendix I(141,
142)
gracillis, Cambarus, 97
Green Bay, 19 (Table 1)
Green Bay Drainage Basin, 19 (Table 1)
Gynandromorphs, 56
Orconectes (Crockerinus) propinquus, 56
Orconectes (Gremicambarus) uirilis, 83, 84 (Fig. 53)
Historical Account of Wisconsin Decapods, 2, 3 , 4
Hybridization, 1,4, 63,64,67, 71,89
debilis, Cambarus, 3, 79
Decapoda, 30,46
demersum, Ceratophyllum, 114
diogenes, Cambarus, 3, 4, 24, 30 (Table 21, 33, 35, 36 (Table 3),
37, 50, 59,64, 76,96, 102, 113
diogenes, Cambarus diogenes, 29, 35
diogenes, Cambarus (Bartonius), 29
diogenes, Cambarus (Cambarus), 29
diogenes, Cambarus (Lacunicambarus), 8, 9, 21, 22, 23, 25 (Fig.
19), 29, 30, 31 (Fig. 23), 32 (Fig. 241, 33 (Fig. 25), 34 (Fig.
26), 35, 36, 37 (Fig. 27), 38 (Fig. 28), 39 (Fig. 29), 90 (Fig.
57), 91 (Fig. 59), Appendix I(141, 142)
diogenes, Cambarus (Lacunicambarus) diogenes, 3, 6
Diogenes, Cambarus, 29
diogenes sspp., Cambarus, 29
Driftless Area, 1, 13, 14 (Fig. 16), 16, 17
dupratei, Procambarus (Pennides), 56
Identification Procedure,
Cambaridae, 9, 10, 11 (Fig. 14), 12 (Fig. 15)
Palaemonidae, 9 (Fig. 13), 12
Introductions, 1, 2
Crayfishes, 1, 2 , 4
Shrimp, 119
immunis, Cambarus, 43,47
immunis, Cambarus (Faxonius), 43
immunis, Orconectes, 24, 43, 46, 48, 49, 50, 6, 64, 76, 86, 88,
89,96
immunis, Orconectes (Grmicambarus), 21, 22, 23, 26 (Fig. 20), 43,
44 (Table 4), 45 (Fig. 32), 46 (Fig. 33), 47, 48, 49, 50 (Table
5), 51 (Fig. 34), 52 (Fig. 3 3 , Appendix I(141, 142)
immunis, Orconectes, 43
immunis sspp., Orconectes, 43
Eastern Ridges and Lowlands Province, 13, 15 (Fig. 17), 16
Elodea species, 47, 74,90, 109
Elodea canadensis, 114
Equisetum species, 60,90
exilipes, Palaemonetes, I 14
kadiakensis, Palaemonetes, 21, 22, 114 (Table 17), 115 (Fig. 73),
116 (Table 18), 117, 118 (Fig. 77), 119 (Fig. 74), 120 (Figs.
75, 76), Appendix I(141, 142)
kadiakensis, Palaemonetes (Palaemonem), 114
Key to Wisconsin Decapods, 22, 23,24
Fallicambarus, 2 1
(Creaseriw) fodiens, 3, 21, 22, 23, 39, 40 (Fig. 30), 41, 42 (Fig.
31), 43, Appendix l(141, 142)
Diagnosis, 39, 41
Distribution, 42 (Fig. 31), 43
Ecology, 4 1, 42
Life History, 42, 43
Type-locality, 41
fodiens, 4, 41, 42, 43, 59
Faxonius uirillis, 80
fodiens, Cambarus, 39
fodiens, Fallicambarus, 4, 41, 42, 43, 59
fodiens, Fullicambarw (Creaserius), 3, 21, 22, 23, 39, 40 (Fig. 30),
41, 42 (Fig. 31), 43, Appendix I(141, 142)
fontinalis, Salwelinw, 88
Lake Michigan, 13, 16, 19 (Table l), 21,93
Lake Michigan Drainage Basin, 13, 20 (Fig. 18), 50, 102, 111,
142 (Appendix I)
Lake Superior, 13, 21
Lake Superior Drainage Basin, 20 (Fig. 18), 142 (Appendix I)
latifolia, Typha, 60
h n a species, 60, 74
Lemna minor, 114
limosus, Orconectes, 63, 64
Life History,
Cambarus (Lacunicambarus) diogenes, 35, 36 (Table 3), 37
Fallicambarus (Creaserius) fodiens, 42,43
Orconectes (Oremicambarus) immunis, 48,49, 50 (Table 5)
Orconectes (Crockerinus) propinquus, 59, 62, 63, 64 (Table 7)
Orconectes (Procericambarus) rusticus, 74, 75
Orconectes (Oremicambarus) wirilis, 90
Procambarus (Girardiella) gracilis, 102, 103 (Table 14)
Procambarus (Ortmannicus) acutus acutus, 110
Palaemonetes kadiakensis, 116 (Table 18)
Geological Setting of Wisconsin, 16, 17
Glacial History of Wisconsin, 1, 16, 17, 21
gracilis, Camburus, 3, 97
174
Macrobrachiurn, 119
Marsifea, 47
Micropterus dolomieui, 59
Mimulus species, 60
Mississippi Drainage Basin, 13, 21, 118, 142 (Appendix I)
Myriophyllum species, 74
Nujus species, 74, 90
Nasturtium species, 60
Northern Highland Province, 13, 15 (Fig. 171, 17
Noturw truutmani, 35
Nuphur species, 109
Nuphar microphyllum, 74,90
obesus, Cambarus, 3,29, 33
obscurus, Orconectes, 63, 64
Oligochaetes, discodrilid, 4
Orconectes, 12, 21, 22, 56,80
(Crockerinus) propinquus, 3, 5, 6, 7, 21, 22, 24, 25 (Fig. 191,
27 (Fig. 21), 28 (Fig. 22), 52, 53, 54 (Fig. 36), 55 (Table
6), 56, 57 (Figs. 37, 381, 58, 59, 60 (Fig. 391, 61 (Figs.
40-43), 62, 63, 64 (Table 7), 65 (Fig. 44), 66 (Fig. 451,
90 (Fig. 57), Appendix I(141, 142)
Behavior, 59, 60
Color Notes, 56, 57 (Figs. 37, 38), 58
Crayfish Associates, 64
Diagnosis, 53
Distribution, 64, 65 (Fig. 441, 66 (Fig. 45)
Ecology, 58, 59,60
Life History, 62, 63, 64 (Table 7)
Type-locality, 53
Variation, 55, 56
(Gremicumbarus) immunis, 21,22, 23,26 (Fig. 20), 43,44 (Table
4), 45 (Fig. 32), 46 (Fig. 331, 47, 48, 49, 50 (Table 51,
51 (Fig. 34), 52 (Fig. 35), Appendix I(141, 142)
Color Notes, 46 (Fig. 33)
Crayfish Associates, 50
Diagnosis, 43
Distribution, 50, 51 (Fig. 341, 52 (Fig. 35)
Ecology, 46,47, 48
Life History, 48, 49, 50 (Table 5)
Type-locality, 46
Variation, 44, 46
(Gremicumbarus) uirilis, 3, 5, 6, 7, 21, 22, 24, 25 (Fig. 19), 27
(Fig. 21), 28 (Fig. 22), 60 (Fig. 39), 61 (Figs. 40, 41), 79,
80, 81 (Table lo), 82 (Fig. 52), 83, 84 (Figs. 53, 54), 85
(Fig. 55), 86, 87, 88, 89 (Fig. 56), 90 (Figs. 57, 581, 91
(Figs. 59, 60), 92, 93, 94 (Table l l ) , 95 (Fig. 61), 96 (Fig.
62), Appendix I ( l 4 1 , 142)
Behavior, 87,88,89
Color Notes, 83, 84 (Fig. 54), 85 (Fig. 55)
Crayfish Associates, 96
Diagnosis, 80, 83
Distribution, 94, 95 (Fig. 61), 96 (Fig. 62)
Ecology, 85, 86, 87,89, 90
Life History, 90, 91,92,93,94 (Table 11)
Type-locality, 83
Variation, 83, 85
immunis, 24, 43, 46, 48,49, 50, 56,64,16,% 8% 89, 96
immunis sspp., 43
limosus, 63, 64
obscurus, 63,64
placidus, 3
(Procericambarus) rusticus, 1, 5, 7, 21, 24, 25 (Fig. 19), 27 (Fig.
21), 28 (Fig. 22), 66, 67, 68 (Table 8), 69 (Fig. 46), 70
(Figs. 47, 48), 71, 72 (Fig. 49), 73, 74, 75, 76. 77 (Table
9), 78 (Fig. 50), 79 (Fig. Sl), 89 (Fig. 56), Appendix I(141,
142)
Color Notes, 71
Crayfish Associates, 76
Diagnosis, 67
Distribution, 76, 78 (Fig. 50), 79 (Fig. 51)
Ecology, 71, 72, 73, 74
Life History, 74, 75, 76, 77 (Table 9)
Type-locality, 7 1
Variation, 67, 71
propinquis, 53
propinquus, 4, 24, 37, 42, 48, 50, 52, 53, 55, 56, 58, 59, 60,
62,63, 64, 67, 71, 76, 83, 85, 88,89,91, 93, 96
propinquus propinquus, 52, 56
propinquus sspp., 52
rusticus, 3, 4, 24, 37, 50, 58, 59, 63, 64, 66, 67, 71, 72, 73,
74, 75, 76, 77 (Table 9), 86,88, 89, 91,96
sanbmii, 64, 75
uirilis, 3, 4, 24, 37, 48, 50, 59, 60, 63, 64, 71, 80, 83, 85, 86,
87, 88, 89,90,91, 92,93,94,96, 102, 113
Orconnectes immunis, 43
Orconnectes propinquus, 53
Orconnectes rusticus, 66
Orconnectes uirilis, 80, 81 (Table 10)
orconectes rusticus, 66
orconectes rusticu, 66
Ostracods, entocytherids, 4
Palaemonetes, 12, 118
exilipes, 1 14
kadiukensis, 21, 22, 114 (Table 17), 115 (Fig. 73), 116 (Table
18), 117, 118 (Fig. 74), 119 (Fig. 75), 120 (Figs. 76, 771,
Appendix I(141, 142)
Color Notes, 118
Diagnosis, 114
Distribution, 118, 119 (Fig. 75), 120 (Fig. 76)
Ecology, 114, 115
Life History, 116 (Table 18), 117, 118
Type-locality, 114
Variation, 115 (Fig. 73)
(Pulacmonetes) kadiakensis, 114
~ U ~ U ~ O S U1S18
,
Palaemonidae, 9, 12, 21, 22, 114
paludosus, Palaemonetes, 118
pulustris, Zannichelliu, 74, 90
Parastacidae, 2 1
pH (acidity), 87, 93
175
Physicochemical Data, 18 (Table 11, 19.(Table 11, 35, 47, 48, 59,
73,86,90, 101 (Table 13), 109, 110
Physiography of Wisconsin, 13, 14 (Fig. 16), 16
pipiens, Rana, 35
placidus, Cambarus, 3
Pontederia cordata, 74,90
Potamogeton species, 47,60,109
amplifolius, 74
angustifolius, 74
crispis, 74,90, 116
pusillus, 74,90
Preservation Techniques, 9
Procambarus, 11,12,21
acutus, 106
acutus acutus, 11, 24, 37, 50, 58,96, 107 (Table 151, 106, 107,
109, 110
bfandingii acutus, 105, 110
clarkii, 58,60
(Girardiella) gracilis, 3, 7, 21, 23, 24, 26 (Fig. 20), 97, 98, 99,
(Fig. 63), 100 (Fig. 64), 101 (Figs. 65, 66), 102, 103 (Table
14), 104 (Fig. 67), 105 (Fig. 68),Appendix l(141, 142)
Behavior, 100, 101
Color Notes, 100
Diagnosis, 97
Distribution, 102
Ecology, 100, 101,102
Life History, 102, 103 (Table 14), 104 (Fig. 67)
Type-locality, 97
Variation, 98
gracilis, 8, 24, 37, 96, 97, 98 (Table 12), 101 (Table 141, 113
(Ortmannicus) acutus acutus, 3, 5, 21, 22, 23, 26 (Fig. 201, 102,
105, 106, 107,108 (Fig. 69), 109 (Fig. 70), 110,111 (Table
16), 112 (Fig. 71), 113 (Fig. 72),AppendixI(141,142)
Color Notes, 106, 107, 109
Crayfish Associates, 113
Diagnosis, 106
Distribution, 111, 112 (Fig, 71), 113 (Fig. 72)
Ecology, 109
Life History, 110, I 1 1 (Table 16)
Type-locality, 109
Variation, 106
(Ortmannicus) acutus cuevachicae, 113
(Pennides) dupratzi, 56
(Pennides) spiculifer, 10
(Scapulicambarus) clarkii, 58
propinquis, Orconectes, 53
propinquus, Cambarus, 3,52,72, 118
propinquus, Cambarus (Faxonius), 52
propinquus, Orconectes, 4, 24, 37, 42, 48, 50, 52, 53, 55, 56, 58,
59,60,62,63, 64, 67, 7 1, 76, 83,85, 88,89, 91 96
propinquus, Orconectes (Crockerinus), 3, 5, 6, 7, 21, 22, 24, 25
(Fig. 19), 27 (Fig. 21), 28 (Fig. 22), 52, 53, 54 (Fig. 361, 55
(Table 6), 56, 57 (Figs. 37, 38), 58, 59, 60 (Fig. 391, 61 (Figs.
40,43), 62, 63, 64 (Table 7), 65 (Fig. 441, 66 (Fig. 45), 90
(Fig. 57), Appendix I(141, 142)
propinquus, Orctrnectes propinquus, 52, 56
176
propinquus, Orconnectes, 4, 24, 37, 42, 48, 50, 52, 53, 55, 56,
58, 59, 60, 62, 63, 64, 67, 71, 76, 83, 85, 88, 89, 91, 93,
96
propinquus sspp., Orconectes, 52
pusillus, Potamogeton, 74,90
Ramphocorixa acuminata, 110
Ram pipienr, 35
Ranunculus amis, 60,90
River,
Apple, 18 (Table 1)
Bad, 19 (Table 1)
Black, 16, 18 (Table l), 20 (Fig..18), 50, 76, 111, 118, 142
(Appendix I)
Chippewa, 13, 16, 18 (Table l), 20 (Fig. 18), 50, 111, 142
(Appendix I)
Clam, 18 (Table 1)
Fox-Wolf, 13, 16, 17, 18 (Table l), 20 (Fig. l6), 111, 118, 142
(Appendix I)
Menominee, 13, 20 (Fig. 181, 111, 142 (Appendix I)
Mississippi, 13, 16, 18 (Table l), 21, 50, 114
Namekagon, 18 (Table l), 50
Nemadji, 19 (Table 1)
Oconto, 16,20 (Fig. 18), 111,142 (Appendix I)
Pecatonica, 18 (Table I), 20 (Fig. 18), 111, 118, 142 (Appendix
1)
Peshtigo, 20 (Fig. 18),111,142 (Appendix I)
Rock-Fox, 18 (Table l), 20 (Fig. 18), 100, 102, 111, 142
(Appendix I)
5\'. Croix, 13, 18 (Table l), 20 (Fig. 18), 50, 114, 118, 142
(Appendix I)
Sugar, 18 (Table l), 20 (Fig. 18), 100, 111, 118, 142 (Appendix
1)
Trempealeau, 18 (Table l), 20 (Fig. 18), 50, 76, 111, 118, 142
(Appendix 1)
Wisconsin, 13, 16, 17, 18 (Table l), 20 (Fig. 18), 33 (Fig. 25)
Wolf, 13, 16, 17, 20 (Fig. 18),83, 114
Yellow, 18 (Table 1)
robustus, Cambarus, 60,63,64
rusticu, oronectes, 66
rwticus, 66
rusticus, Cambarus, 3,66,79
rusticus, Cambarus (Faxonius), 80
rwticus, Cambarus (Faxonius) rusticus, 80
rusticus, Orconectes, 3, 4, 24, 37, 50, 58, 59, 63, 64, 66, 67, 71,
72, 73, 74,75,76,77 (Table 9),86,88,89,91,96
rusticus, Orconectes (Procericamburus), 1, 5, 7, 21, 24, 25 (Fig. 19),
27 (Fig. 21), 28 (Fig. 22), 66, 67, 68 (Table 81, 69 (Fig. 46),
70 (Figs. 47, 48), 71, 72 (Fig. 49), 73, 74, 75, 76, 77 (Table
9), 78 (Fig. 50), 79 (Fig. 51), 89 (Fig. 56), Appendix 1 (141,
142)
rusticus, Orconnectes, 66
Sugittmiu species, 60,90
Suluelinus fontinulis, 88
sunbornii, Orconectes, 64, 75
shufekitii, Cambarellus (Dirigicambarus), 56
Sistmrus catenatus catenatus, 41
Spartinu, 3,41
spiculifer, Procambarus (Pennides), 10
Spirea species, 90
alba, 74
Spirogyra species, 74
St. Lawrence Drainage Basin, 13, 21, 50, 118, 142
(Appendix I)
stygius, Cambarus, 105
Stygius, Cambarus, 3,105
Superior Upland Province, 13, 14 (Fig. 16)
viriles, Cambarus, 79
wirilis, Cambarus, 3, 79
virilis, Cambarus (Faxmius), 80
virillis, Faxonius, 80
wirilis, Orconectes, 3, 4, 24, 37, 48, 50, 59, 60, 63, 64, 71, 80,
83, 85,86,87,88,89,90,91,92,93,94,96, 102, 113
wirilis, Orconectes (Gremicambanrs), 3, 5, 6, 7, 21, 22, 24, 25 (Fig.
19), 27 (Fig. 21), 28 (Fig. 22), 60 (Fig. 391, 61 (Figs. 40, 411,
79, 80, 81 (Table lo), 82 (Fig. 52), 83, 84 (Figs. 53, 54), 85
(Fig. 55), 86, 87, 88, 89 (Fig. 56), 90 (Figs. 57, 58), 91 (Figs.
59,60), 92,93,94 (Table 1l), 95 (Fig. 61), 96 (Fig. 621, Appendix
Thelohania species, 93
Typha species, 74, 90, 109
latifolia, 60
Western Upland Province, 13, 15 (Fig. 17), 17
wisconsinensis, Cambarw, 3,80
Wiscminemis, Cambarus, 80
Vallisnen'a species, 60
americana, 74,90
Zannichellia palustris, 74, 90
l(141, 142)
177