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DISTRIBUTION OF LYCOSIDAE AMONG VARIOUS HABITATS
IN MACKINAC COUNTY, MICHIGAN,

INCLUDING RECORDS OF DIURNAL AND NOCTURNAL ACTIVITY

By

Fred Edward Doepke, Jr.

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE

Department of Zoology

1981

ABSTRACT
DISTRIBUTION OF LYCOSIDAE AMONG VARIOUS HABITATS

IN MACKINAC COUNTY, MICHIGAN,
INCLUDING RECORDS OF DIURNAL AND NOCTURNAL ACTIVITY

By

Fred Edward Doepke, Jr.

Portions of a square mile in the Upper Penninsula of
Michigan were sampled over a period of three months in an effort
to determine the distribution of species of Lycosidae among various
habitats and to observe their diurnal-nocturnal activities.

Randomly placed pitfall traps provided a weekly means of
sampling populations in seven habitats within the section. Regularly
placed pitfall traps provided recordings of diurnal activity in two
habitats.

Individuals of Pardosa moesta Banks were captured in the

 

greatest number. Tarentula aculeata (Clerck) and Trochosa pratensis

 

 

(Emerton) were the only species captured in all seven habitats. The
evidence of activity during the day and inactivity at night, as in-
dicated in the diurnal-nocturnal study, is attributed to the tactile

cueing of these spiders in motion in darkness.

Dedicated to

Gale R. Gleason, Jr.

Geolzcosa missoutiensis, with young,
the entrance to her burrow

iii

guarding

 

ACKNOWLEDGEMENTS

I wish to extend my most sincere thanks to Dr. T. Wayne
Porter for his wisdom and guidance in the preparation of this
manuscript, and to Dr. Louis Wilson and Dr. Richard Snider for
their helpful suggestions concerning the experimental design of
this project and serving as members of my guidance committee.

I am also deeply indebted to my father, Fred, for contributing
countless hours to helping me with all aspects of my field work
during the spring and summer of 1978, and to my mother, Anabel, for
tolerating the littered situation this three month study created in
her otherwise immaculate home.

Others I would like to thank for their help in checking and
picking up the pitfall traps include Marty Brawley, Gretchen and
Kurt Doepke, Gary Ehn, Austin and Adam Hancock, Todd Hines, Nikki
Tremblay and Gary Salter.

Rhonda Hines was responsible for the excellent pictorial summary
of the field work, and to her I extend my heartfelt thanks. Jim
O'Neil took and presented to me the introductory picture of the
wolf spider, and to him I am also deeply grateful.

Lastly, I would like to thank my wife, Jolene, for the many

unselfish hours she devoted to helping prepare this entire manuscript.

iv

LIST OF TABLES . .

LIST OF FIGURES .

INTRODUCTION . . .

STUDY AREA . . . .

HABITAT STUDY . .

Methods . .

Results and

Discussion

TABLE

OF CONTENTS

DIURNAL-NOCTURNAL ACTIVITY STUDY . . . .

Methods . .

Results and Discussion

SUMMARY AND RECOMMENDATIONS

REFERENCES CITED .

Page
vi

vii

l4
14
15
29
29
29
32

34

LIST OF TABLES

Table Page

I: SPECIES OF LYCOSIDAE REPRESENTED BY LESS THAN
1% OF TOTAL CAPTURE AND HABITATS IN WHICH FOUND. . . . 19

II: COMBINED RECORD FOR SIX LEAST COMMON LYCOSIDAE
SPECIES OF DIURNAL-NOCTURNAL STUDY 0 o o o o O o o o o 19

vi

Figure

10:

ll:

12:

13:

LIST OF FIGURES

Study Area in Newton Township, Michigan, used for
studying habitats and activities of the Lycosidae

Study Area #1 used for Lycosidae habitat study;
major ground cover was grass and goldenrod . . .

Study Area #2 used for Lycosidae habitat study;
major ground cover was sugar maples . . . . . . .

Study Area #3 used for Lycosidae habitat study;
area was completely submerged two months prior to
time of photograph . . . . . . . . . . . . . . .

Study Area #4 used for Lycosidae habitat study;
permanent pond in background is not visible in
photograph O O C O O O O O I I C O O I I O O O 0

Study Area #5 used for Lycosidae habitat study;
major ground cover was rock and grass . . . . . .

Study Area #6 used for Lycosidae habitat study;
major ground cover was cedar . . . . . . . . . .

Study Area #7 used for Lycosidae habitat study;
major ground cover was sand . . . . . . . . . . .

Site of diurnal-nocturnal Lycosidae activity study

in field; plot measures six meters by six meters

Site of diurnal-nocturnal activity study in hard-
woods; plot measures six meters by six meters . .

Initial placement of pitfall trap in Lycosidae
habitat study showing antifreeze and flag . . . .

Pitfall trap in Habitat 7 of the Lycosidae habitat
study; rainwater dilution of antifreeze and labelling

technique can both be seen . . . . . . . . . . .
Species and percentages of Lycosid spiders by

frequency of occurrence, captured in all areas
throughout StUdy O O O O O O O O O O O O O C O 0

vii

Page

11

ll

12

12

13

13

l6

l7

LIST OF FIGURES (Continued)

Figure Page

14: Lycosidae species composition of (a) grass field,
(b) hardwoods, (c) vernally flooded field and (d)
pond edge habitats of the Lycosidae habitat study.
The total number of individuals captured from June
through August is given for each habitat . . . . . . . 21

15: Lycosidae species composition of (a) rock outcrop-
ping, (b) softwoods and (c) sand pit habitats. The
total number of individuals captured from June through
August is given for each habitat . . . . . . . . . . . 22

16: Distribution of (a) Pardosa distincta, (b) Pardosa
milvina, (c) Pirata minutus and (d) Trochosa
pratensis among habitats of the Lycosidae habitat
study . . . . . . . . . . . . . . . . . . . . . . . . . 23

 

 

17: Distribution of (a) Pardosa moesta, (b) Pardosa
modica, (c) Schizocosa crassipalpis and (d) Tarentula
aculeata among habitats of the Lycosidae habitat
study . . . . . . . . . . . . . . . . . . . . . . . . . 24

 

 

18: Frequency of (a) Pardosa distincta, (b) Schizocosa
crassipalpis and (c) Tarentula aculeata capture in
24-hour periods - six collections June 8 - August
31 for grass plus hardwood plots . . . . . . . . . . . 3O

 

 

 

 

viii

INTRODUCTION

Wolf spiders constitute a family of hunting spiders in the
Order Araneae of the Class Arachnida. Unlike most other families,
these spiders do not construct a web for the capture of food, but
instead stalk their prey on the ground. As a retreat, some con-
struct tubular tunnels in the soil, and others utilize natural
depressions under rocks and debris. Others never use a retreat,
but are found running through grass, among dead leaves on the forest
floor, or over sandy and stony areas (Kaston, 1978).

Sometimes considered as pests in human dwellings, wolf spiders
are important in helping control other invertebrate pests, partic-
ularly insects; Dondale (1962) has speculated on their use in pest
control. Enemies of the Lycosidae include other spiders and some
species of insects and birds, but parasites and availability of food
also help control spider populations (Levi, Levi and Zim, 1968).

An understanding of the role of the environment in the natural
selection of these creatures can be enhanced by knowledge of their
optimal habitats. Collectors generally record the habitats in
which they find specific spiders. However, to this author's knowledge,
no one has yet attempted to experimentally determine whether or not
specific spiders are restricted to specific habitats.

Furthermore, taxonomic studies on the Lycosidae Egggu
strictu have not been performed in Michigan prior to this undertaking.
Indeed, it appears to have been the rule that individuals of the

1

2
Lycosidae have been reported either incidentally or in the context
of larger studies involving all the Araneae. Also, studies of
the Lycosidae in Michigan reflect heavy bias toward collecting in
the Lower Penninsula.

Chickering (1932; 1933; 1935) and Chickering and Bacorn (1933)
published early taxonomic studies on six genera and 32 species of
Lycosidae, all from the Lower Penninsula. Drew (1957) found four
genera and eight species of Lycosidae in a woodlot near Lansing, and
Hindle (1972) listed five genera and eight species from the area
of Bog Lake, Michigan. One early study (Chickering, 1934) involved
a subset of two genera and four species in an Upper Penninsula
collection near Marquette, Grand Marais, and Ishpeming. Vail (1974)
reported five genera and 16 species in his statewide survey of
spiders associated with dwellings. While Drew (1965) tabulated
six genera and 23 species in his Beaver Island survey, and Bixler
(1967) reported six genera and 14 species from Isle Royale, no
Lycosidae have yet been reported from Mackinac County.

The objectives of this study were:

1. to determine the species and frequency of species of
Lycosidae in the area chosen in Mackinac County, and thus
add to the knowledge of Lycosidae fauna in the Upper
Penninsula of Michigan;

2. to examine the distribution of Lycosidae among diverse

habitats.

3
3. to study some habits and diel activity the Lycosidae

demonstrate.

STUDY AREA

All species of Lycosidae reported in this study were obtained
from a one square mile area in Newton Township (T43N, R11W: 529),
Mackinac County, Michigan. The study area (Fig. 1) is immediately
west of Gould City, and bounded on the north by U.S.—2, east by
Gould City Road, and south by Corinne Road. The western edge of
this section has no visible natural or artificial boundary. The
grade for the 800 Line Railroad cuts from the southwest corner of
the section diagonally to north of the middle of the eastern
boundary. The only inhabited houses in the section are those that
border the western side of Gould City Road.

A man-made sandpit, centered in the southeast quarter of the
section, is accessible from Corinne Road via a seldom used two—track
road. A permanent pond approximately 75 meters in diameter is
located in the northeast quarter of the southeast quarter of the
section. East of this pond, and bordering on Gould City Road, a
temporary pond approximately 120 meters in diameter appears in the
spring of the year.

The area is approximately three-quarters grassland. The north-
west quarter of the section is occupied by a stand of hardwoods, approx-

imately 90% of which are sugar maple (Acer saccharum Marshall). A

 

stand of conifers, primarily cedars (Thuja occidentalis Linnaeus),

 

occupies the southwest quarter of the southeast quarter of the section.

Scattered throughout the section are several stands of scotch pine

4

  
 

 

4Q

  

Gould City Road

 

 

 

Corinne Road

 

 

I Permanent pond Distribution study areas

'2 Softwoods T m Hardwoods
N /

" " " Two-track trail

\
(DJ > Activity study areas

I Dwellings

FIGURE 1: Study Area in Newton Township, Michigan, used for studying
habitats and activities of the Lycosidae.

6

(Pinus sylvestris Linnaeus) that were planted in the late 1950's.

 

In this study, nine plots in seven different habitats were
set up and numbered. Plots one through seven corresponded to the
habitat study; plots eight and nine corresponded to the diurnal-
nocturnal study. In a format similar to that of Boyd (1960) and
Duffey (1966), the seven habitats containing these plots were
selected on the basis of their superficial differences. These
habitats represented areas of 1) grass, 2) hardwoods, 3) forb-
grass vernal flooding, 4) pond shore, 5) rock terrain, 6) soft—
woods, and 7) sand. A description of these habitats:

Habitat 1;

This area was located in the middle of a grassy field in the
center of the section's northeast quarter. A mat approximately three
centimeters thick was formed by short grass (Gramineae) mixed with
previous years residue. Clumps of goldenrod (Solidago spp.) were
scattered throughout. A thin layer of mosses (Musci) covered the
ground beneath the mat (Fig. 2).

Habitat_2:

This area was located near the center of the deciduous forest
that covered the greater portion of the northwest quarter of the
section. Sugar maples approximately ten meters tall were the major
flora. Other deciduous trees scattered throughout included a few

paper birch (Betula papyrifera Marshall), beech (Fagus grandifolia

 

 

Ehrhart), ironwood (Ostrya virginiana Miller), and yellow birch (Betula

 

lutea Michaux). Some aspen (Populus spp.) were also present. The

 

 

FIGURE 2 (ABOVE): Study Area #1 used for Lycosidae habitat study;
major ground cover was grass and goldenrod.

FIGURE 3 (BELOW): Study Area #2 used for Lycosidae habitat study;
major ground cover was sugar maples.

8
most abundant saplings and seedlings were maple and beech. The soil
was extensively covered with leaf litter, small twigs, and sticks
(Fig. 3).
Habitat 3:

This area lies in the northeast corner of the southeast corner
of the section and represents a low, grassy field that floods each
spring when the permanent pond adjacent to it overflows. At the
beginning of the study, after the flood water had dissipated,
approximately 90% of the mud floor was covered with moss. As
summer progressed, grasses sprouted, and near the end of the study
this habitat resembled Habitat 1. During summer this area supported
goldenrod, milkweed (Asclepias spp.), Clover (Trifolium spp.), and
miscellaneous forbs (Fig. 4).

Habitat.£:

Located in the southeast quarter of the section, this area was
the eastern border of the permanent pond, and supported a variety of
vegetative types. Among these were aspens of various sizes, apple

trees, a few spruce (Picea_glauca Moench) saplings, some willow (Salix

 

.gpp.), cedar saplings and seedlings, and various forbs. Goldenrod
and clover were both fairly abundant. The floor was typically
grasses and forbs growing through the litter-covered moss (Fig. 5).
Habitat_§:

Located in the southeast quarter of the southeast quarter of
the section, this area is characterized by its exposed rock floor.

It is on a ledge overlooking Gould City Road. Sugar maples in

 

FIGURE 5 (BELOW):

   
   

FIGURE 4 (ABOVE):

1 it
Study Area #3 used for Lycosidae habitat study;

area was completely submerged two months prior
to time of photograph.

  

 

gr, 1
' ‘H'C’D {‘76.

A

Study Area #4 used for Lycosidae habitat study;
permanent pond in background is not visible in
this photograph.

10
this area were approximately 12 meters tall. Grass in this area
was sparsely distributed, as were the seedlings and forbs (Fig. 6).
Habitat-Q:

Located on the western side of the southeast quarter of the

 

southeast quarter of the section, this area consisted of cedars
approximately nine meters tall interspersed with a few spruce,

cedar saplings and several paper birch trees. The area had a floor

 

covering of cedar needles approximately two centimeters deep. Much
of the floor was shaded, and a few scattered grasses were evident
(Fig. 7).

Habitat Z;

This area consisted of a region in which sand had been removed
from the side of a hill in the southwest quarter of the southeast
quarter of the section. The excavated area measured approximately
25 meters in diameter. Although mature aspen surrounded the area,
the pit was relatively free of vegetation. There were scattered
goldenrod, grasses and forbs, and numerous aspen seedlings; however
the area has the appearance of a sandy hillside on the edge of a
field (Fig. 8).

Areas eight and nine, which contained the plots used in the

 

diurnal-nocturnal activity study, were located in Habitats 1 and

2 respectively (Figs. 9, 10). Therefore, each of these two habitats
contained two sets of plots; one for the habitat study, and one

for the diurnal-nocturnal activity study. Each of the five other

habitats contained only a plot for the habitat study.

11

 

FIGURE 6 (ABOVE): Study Area #5 used for Lycosidae habitat study;
major ground cover was rock and grass.

FIGURE 7 (BELOW): Study Area #6 used for Lycosidae habitat study;
major ground cover was cedar.

 

 

 

FIGURE 8 (ABOVE): Study Area #7 used for Lycosidae habitat study;
major ground cover was sand.

FIGURE 9 (BELOW): Site of diurnal-nocturnal Lycosidae activity study
in field; plot measures six meters by six meters.

FIGURE

FIGURE

 

10 (ABOVE):

11 (BELOW):

Site of diurnal—nocturnal activity study in
hardwoods; plot measures six meters by six meters.

Initial placement of pitfall trap in Lycosidae
habitat study showing antifreeze and flag.

HABITAT STUDY

The habitat study was designed to satisfy the objective of
determining the species of Lycosidae in each of the seven selected
habitats within the study area. Satisfying this objective allows

a comparison of the species of Lycosidae of each habitat.

Methods

Within each of the seven selected habitats, ten pitfall traps
were randomly placed on a north-south oriented grid in a ten by
ten meter area which had been selected for its characteristic
habitat.

This investigator used pitfall traps similar to those used by
Fichter (1941), Mitchell (1963) and Best (1977). Each trap con-
sisted of a 600 ml white plastic food container having a diameter
of 11.4 cm across the open top, a diameter of 9.0 cm across the
bottom, and having a depth of 7.2 cm. Containers were obtained from
Michigan State University Food Stores and came equipped with clear
plastic snap-on lids.

Traps consisting of containers were set in the ground with
their lips flush to the substrate. No protective covers were used.
Each trap was accompanied by a red marker flag to help locate traps
which would later be concealed by undergrowth (Fig. 11).

Each of the traps was filled with approximately 200 ml of

®

Chemcraft antifreeze. This ethylene glycol based commercial
solution has a low evaporation rate, kills and preserves spiders, and

14

 

 

 

15
is less expensive than laboratory grade ethylene glycol recommended
for pitfall traps (Best, 1977). In addition, the blue hue of the
solution provides a visual measure of rainwater dilution (Fig. 12).

Each of the seventy traps was picked up and replaced each
week during the study. Each lid was labeled with a habitat, trap,
and collection date number. In the laboratory, the contents of
each trap were sorted. Spiders in each trap were transferred to
six-dram vials containing approximately 75% ethyl alcohol and three
drops of glycerin. All spiders were preserved for later identifi-
cation with this technique.

All wolf spiders in this study and in the diurnal—nocturnal
activity study were identified to genus and species using Kaston
(1948). In some instances Kaston (1978), Levi and Field (1954),
and Levi, Levi and Zim (1968) were used in making determinations.
Spiders of the genus Pirata were identified using keys provided by

Wallace and Exline (1977).

Results and Discussion

 

A total of 2218 wolf spiders were identified to species (Fig.
13). Although the eight genera and 22 species of Lycosidae reported
in this study is less than the nine genera and 50 species this author
has determined to have previously been reported from Michigan by
Chickering (1932; 1933; 1934; 1935), Chickering and Bacorn (1933),
Drew (1957; 1965), Bixler (1967), Hindle (1972) and Vail (1974), these

findings add to the Checklist of Lycosidae from the mainland

l6

 

FIGURE 12: Pitfall trap in Habitat 7 of the Lycosidae habitat
study; rainwater dilution of antifreeze and labelling
technique can both be seen.

Species

Pardosa moesta
Tarentula aculeata
Schizocosa crassipalpis

 

17

 

 

19.572
15.692

 

Pardosa distincta

15.242

 

 

Trochosa pratensis

 

 

Pardosa milvina
Pirata minutus
Pardosa modica

Pardosa xerampelina
Lycosa frondicola
Schizocosa saltatrix
Pardosa lapidicina

Pirata piratica

Lycosa modesta
Pirata insularis
Arctosa rubicunda
Pirata sedentarius
Pardosa floridana

 

 

 

 

 

 

 

Pardosa saxatilis
Pirata montanus

 

 

Pardosa mackenziana
Geolycosa missouriensis

FIGURE 13:

F

V V V V V V v ‘ '

 

._ 1.712
-— 1.442
- 1.082
- 0.902

0.862
0.592
0.452
0.322
0.232
0.182
0.182
0.182
0.182
0.142
0.142
0.092
0.052

6.992

33.812

N - 2218

 

0

I
200

l l
400 600

Frequency

r
800

Species and percentages of Lycosid spiders by frequency

of occurrence, captured in all areas throughout study.

18
Upper Penninsula of Michigan established by contributions from
Chickering (1934) and Vail (1974).
The species most prevalent in the area studied was Pardosa
moesta. Individuals of this species were captured in numbers almost

twice that of the next most common species, Tarentula aculeata.

 

These two species, along with Schizocosa crassipalpis, Pardosa

 

distincta and Trochosa pratensis, accounted for over 902 of all in-

 

dividuals captured in this study. Species with numbers of individuals
totaling less than 12 of all individuals captured, and habitats
in which found, are listed in Table I.

A summation of species:

Arctosa rubicunda (Keyserling)

Geolycosa missouriensis (Banks)

 

 

Lycosa frondicola Emerton

 

modesta (Keyserling)
Pardosa distincta (Blackwall)
floridana Banks

lapidicina Emerton

 

mackenziana (Keyserling)

 

milvina (Hentz)
modica (Blackwall)
moesta Banks
saxatilis (Hentz)

xerampelina (Keyserling)

 

Pirata insularis Emerton

19

TABLE I: SPECIES OF LYCOSIDAE REPRESENTED BY LESS THAN 12 OF TOTAL
CAPTURE AND HABITATS IN WHICH FOUND

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Total Area
Species Captured 1 2 3 4 5 6
Arctosa rubicunda 4 2 2
Geolycosa missouriensis 1 1
Lycosa frondicola 19 1 1 10 4
Lycosa modesta 5 2 2 1
Pardosa floridana 4 3 1
Pardosa lapidicina 10 l 3 3 2
Pardosa saxatilis 3 1 1 1
Pardosa xerampelina 20 2 12 5
Pardosa mackenziana 2 2
Pirata insularis 4 2 2
Pirata montanus 3 3
Pirata piratica 7 3 4
Pirata sedentarius 4 4
Schizocosa saltatrix 13 3 6 1

 

 

TABLE II: COMBINED RECORD FOR SIX LEAST COMMON LYCOSIDAE SPECIES OF
DIURNAL-NOCTURNAL STUDY

 

 

 

 

 

 

 

Time (hrs)
c> c> c> c: c: c: c: c:
c> c: c> c: c> c> c> c:
co ~o ox ca aw a: '4 ~¢
O O O v-I H H N N
I I I I I I I I
Total 0 o o o o o o o
Captured 38888882
Species Woods + Field N o o o .-I .—I .—I N
Lycosa frondicola 2 1 l
Pardosa moesta 4 3 1
Pardosa xerampelina 3 2 1
Pirata minutus 2 1 1
Schizocosa saltatrix 1 1
Trochosa pratensis 4 1 2

 

 

20
minutus Emerton
montanus Emerton

piratica (Olivier)

sedentarius Montgomery

 

Schizocosa crassipalpis (Emerton)

 

saltatrix (Hentz)
Tarentula aculeata (Clerck)
Trochosa pratensis (Emerton)
Kaston (1948) indicated the habitats from which his specimens
were obtained. In general, species collected in this study were

found in the habitats he described for the same species. For

 

example, Kaston (1948) and Bixler (1967) both list Pardosa distincta
as an open field species, and the results presented here (Figs.
14a, 16a) agree with their studies. It is notable that Barnes (1953)
did not report this species in his study of non-forest spiders in
North Carolina. According to Kaston (1978), the range of this species
is New England and adjacent Canada south to North Carolina and west
to Arizona and Montana. Therefore, the results of Barnes (1953)
reflect either the range of this species or the distribution of
collectors.

In his study of spiders in an overgrazed pasture in eastern

Ontario, Turnbull (1966) cites Pardosa saxatilis, Pardosa milvina

 

 

and Pirata minutus as three of his most commonly found species. This

 

study is in concurrence; Pardosa saxatilis (Table I), Pardosa milvina

 

 

(Figs. 14c, 16b) and Pirata minutus (Figs. 14c, 16c) were all

 

21

 

 

 

 

 

 

 

 

 

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The total number of in-

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Lycosidae species composition of (a) grass field, (b) hard-
each Habitat.

FIGURE 14:

 

 

22

Habitat 5 d Habitat 6
Rock Outcropping (N - 67) ” Softwoods (N e 80)
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Lycosidae species composition of (a) rock outcropping,
(b) softwoods and (c) sand pit habitats. The total
number of individuals captured from June through August
is given for each Habitat.

Frequency

200 4‘

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23

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(a)

T_"’

 

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Frequency

 

Habitats

Pirata minutus

(e)

 

 

 

FIGURE 16:

Habitats

Frequency

frequency

 

 

 

 

301
Email. We
(b)
20'
10"
J 4 3 6 7
Habitat.
Trochosa pratensis
(d)
‘0 4

20‘

 

 

 

 

 

 

Habitats

Distribution of (a) Pardosa distincta, (b) Pardosa
milvina, (c) Pirata minutus and (d) Trochosa pratensis
among habitats of the Lycosidae habitat study.

Frequency

24

 

 

 

 

 

 

m d
Pardosa moesta
(a)
200 '1
3 b S 6 7
Habitats
200 d
Schizocosa crassipalpis
(e)

1.. .

 

 

 

 

Habitat.

Frequency

Frequency

10 ‘

 

Pardosa modica

(b)

 

 

300 4

 

“Abltlt.

Tarentula aculeata

(d)

 

 

LililL

Habitats

FIGURE 17: Distribution of (a) Pardosa moesta, (b) Pardosa modica,
(c) Schizocosa crassipalpis and (d) Tarentula aculeata
among habitats of the Lycosidae habitat study.

25

found in relatively open areas. Individuals of Pardosa saxatilis

 

were captured only in the spring of the year in areas of pond edge
and vernally flooded field, a time during which the vernally flooded
field was void of grasses and the pond edge had little ground cover.

Pardosa milvina was in the vernally flooded field only in the spring

 

of the year, just after the water had dissipated and the area was

void of grass. Kaston (1948) reports Pirata minutus as a marsh

 

lowland inhabitant. This study found Pirata minutus to be ex-

 

clusively an inhabitant of habitats three and four, the vernally
flooded lowland and pond shore habitats (Fig. 16c).

Schizocosa crassipalpis was found predominantly in the grass

 

field (Fig. 17c). It became increasingly abundant in the vernally
flooded area as the summer progressed and this area dried and became

more like area one. Schizocosa crassipalpis is clearly a meadow

 

species, being absent in both hardwoods and softwoods.
As indicated in Table I, several species of Lycosidae were
found in only a single habitat. These species included Geolycosa

missouriensis, Pardosa mackenziana, Pirata montanus and Pirata

 

sedentarius. Since they were represented by only a few individuals,

 

it is difficult to make a valid assignment of specific habitats to

these species. Another individual of Geolycosa missouriensis, for

 

example, was observed in another habitat but did not fall into a
pitfall trap. Therefore, it was not included in Table I. As might
be expected, no species represented by less than 12 of the total

capture was found in all seven areas studied.

26

Species commonly trapped (greater than 12 of the total
number of captured individuals) in this study generally tended
to be absent from at least one habitat. For example, Pardosa
distincta (Fig. 16a) is rare in the hardwoods and softwoods.
Because a number of biotic and physical factors may determine the
habitat of a particular species (Lowrie, 1948), it is difficult to
properly assess one variable as the cause of a particular species
being present or absent in a given habitat. It is possible that

Pardosa distincta has a light or temperature requirement that

 

eliminates this species from the woods. Predation may reduce the
number of individuals of this species in these two areas, or perhaps
the darkness in the woods causes these spiders to rely heavily on
tactile cues, thus avoiding pitfall traps more readily. Curtis
(1980) has compared the efficiency of different types of pitfall
traps in spider community studies, and it is possible the type of
pitfall trap used in this study is inefficient in capturing this
species of Lycosidae.

Pardosa modica was found almost exclusively in area three

 

(Fig. 17b), the vernally flooded field, but a few were collected

in areas one and four, the grass field and pond shore areas.

Kaston (1948) describes these habitats in his reference to collecting
this species, and therefore these data agree.

Trochosa pratensis (Fig. 16d) and Tarentula aculeata (Fig. 17d)

 

 

are the only species found in all seven of the habitats examined.

While the reason for this is not entirely clear, the large number

27

of individuals of both of these species also reflects their adaptive
success. Apparently, they can not only survive in the variety of
habitats examined, they also do well reproductively. Kaston (1978)
describes the ranges of both of these species as extending across
the northern United States and southern Canada, with Trochosa
pratensis found as far south as North Carolina. It appears these
spiders are in the middle of their geographical range in the Upper
Penninsula of Michigan, and thus might be expected to be surviving there.

Other species of Lycosidae represented by large numbers of
individuals in this study were not ubiquitous. However, the areas
in which these other species appear to be absent were generally
low yield areas for all species of Lycosidae. Therefore, there is
a possibility that biases associated with the pitfall traps are
responsible for their absence in the pitfall traps in those areas.

Interestingly enough, Bixler (1967) found Tarentula aculeata

 

to be a forest species. The data from this study indicate Tarentula
aculeata is clearly a field species (Fig. 17d). It is possible that

on Isle Royale Tarentula aculeata could not adopt its normal habitat

 

and therefore adapted to the forest. Recalling the biotic factors

mentioned by Lowrie (1948), it is possible that Tarentula aculeata

 

survived on Isle Royale because a woodland competitor had failed to
colonize the island. Equally plausable is the possibility that a

predator of Tarentula aculeata had not colonized Isle Royale. The

 

absence of interspecific competitive or predatory factors may have

insured the survival of Tarentula aculeata on Isle Royale.

 

28
The most intriguing spider associated with the habitat study

was Geolycosa missouriensis. The photograph at the beginning of

 

this manuscript shows a female of this species at the entrance to

a burrow in the sand of habitat seven. This spider, with her young,
was the only animal observed at the entrance to that burrow, and

she was never observed away from the burrow. She retreated into the
burrow when approached, and quickly cleared the entrance of debris,
such as straw, twigs, and pebbles, placed deliberately to observe
her reaction.

Two other geolycosids were observed in the rocky outcropping
area, where they had somehow managed to construct a burrow in the
soil between the rocks. While it was originally thought that a
Geolycosa population had entered the area via a belt of sandy soil
extending nine miles from the beach of Lake Michigan, the burrows
in the rocky substratum indicated Geolycosa did not need a continuum
of sand to extend its range. Furthermore, the fact that no geo-
lycosids were captured in pitfall traps suggests they remain close

to their burrows, at least during the season of this study.

DIURNAL-NOCTURNAL ACTIVITY STUDY

Past research has demonstrated the feasibility of inverte-
brate activity studies based on pitfall analyses of different habitats
(Williams, 1959; Pedigo, 1970). However, Mums and Mama (1949) suggest
caution in interpreting diurnal pitfall data because of their findings
that some species of spiders exhibit vertical daily migration as well

as, or other than, horizontal daily migrations.

Methods

Areas eight and nine, which represented areas in the grass
field habitat and deciduous forest habitat, were examined for
diurnal-nocturnal movement of Lycosidae. For a 24 hour period
beginning every two weeks - six times from June through August - 36
pitfall traps were placed evenly in a six by six meter area in each
of the two habitats (Figs. 9, 10). All 72 traps were emptied and
recharged every three hours, so that eight collections were made
during each 24 hour period. Spiders of each collection time were
separated into vials corresponding to their respective areas of

capture.

Results and Discussion

 

Assuming pitfall traps can indicate periods of activity in the

Lycosidae, the results of this study indicate Pardosa distincta and

 

Schizocosa crassipalpis are most active in the mid—afternoon (Fig. 18).

 

Tarentula aculeata (Fig. 18c) appears to be most active in the mid-

 

afternoon until just before dust. Pitfall data indicate these three

29

Frequency

Frequency

30

 

 

 

 

 

 

+ Pardosa distincta Schizocosa crassipalpis
23
(a) (b)

a. ..I

15 J

10: EI- 10'

51

a g § § § § § §§

§§§§§§§

2400
0300

Tina (Hrs)
Tins (Hrs)

Tarentula aculeata

(e)

101

 

 

8 8 8 8 8 8
S 3 g 2 g 2 a :3

0300

Tina (Hrs)

FIGURE 18: Frequency of (a) Pardosa distincta, (b) Schizocosa

crassipalpis and (c) Tarentula aculeata capture in
24-hour periods - six collections June 8 - August 31
for grass plus hardwood plots.

 

31
species are all inactive at night. Even in the instances of those
Lycosidae that were not numerous in the diurnal-nocturnal study
(Table II), the available data indicate activity during the day and
inactivity at night.

Authors have suggested hunting wolf spiders at night with a
flashlight (Levi, Levi and Zim, 1968; Kaston, 1978). After dark,
they are supposedly out of hiding and actively seeking prey. In
addition, the reflective quality of their eight eyes simplifies
the task of locating them in the dark.

While these spiders may reach peak activity during the day
as indicated in this study, a bias associated with pitfall traps in
diurnal—nocturnal study may be responsible for these results.
Haynes (1981) suggests that these spiders operate under different
stimuli at various times of the daily cycle. During the day, they
probably use visual cues for orientation. At night, they may feel
their way over the substrate, advancing cautiously and thus detecting
pitfall traps more readily. For this reason, care should be exer-
cised in interpreting invertebrate activity based on diurnal—

nocturnal pitfall results.

SUMMARY AND RECOMMENDATIONS

(1) A square mile in the Northern Penninsula of Michigan was studied
in an effort to determine the distribution of species of Lycosidae
among various habitats and to observe the diurnal activities these
species exhibit.

(2) This study took place during the period from May 25 to August 31,
1978.

(3) Randomly placed pitfall traps provided a weekly means of sampling
populations in seven habitats within the section.

(4) Regularly placed pitfall traps provided recordings of diurnal
activity in two habitats.

(5) A total of 2218 individuals of the Lycosidae were identified to
species. This number represented 8 genera and 22 species.

(6) Pardosa moesta Banks was found to be the species most frequently

 

captured, and accounted for 33.82 of the total capture.

(7) Individuals of Tarentula aculeata (Clerck), Schizocosa crassipalpis

 

(Emerton), Pardosa distincta (Blackwall) and Trochosa pratensis

 

(Emerton) comprised 57.52 of the total capture.

(8) Tarentula aculeata (Clerck) and Trochosa pratensis (Emerton) were

 

the only species represented by pitfall captures in all seven habitats
sampled.

(9) Activity during the day and inactivity during the night was
evidenced by those species of Lycosidae captured in diurnal-nocturnal
study, but this is attributed to tactile cueing of these spiders in
motion in darkness.

32

33
(10) A device for automatic timed emptying of pitfall traps

(Williams, 1958) is recommended for activity studies in the future.

REFERENCES CITED

REFERENCES CITED

Barnes, Robert D. 1953. The Ecological Distribution of Spiders
in Non-Forest Maritime Communities at Beaufort, North Carolina.
Ecol. MOno. 23: 315-338.

Best, Richard L. 1977. The Pitfall Trap. Carolina Tips 15(5):
17-18.

Bixler, David E. 1967. Spiders gf Isle Royale, Michigan. M.S.
thesis. Michigan State University. 56 pp.

 

Boyd, J. Morton. 1960. Studies of the Differences Between the
Fauna of Grazed and Ungrazed Grasslands in Tiree, Argyll.
Proc. Zool. Soc. Land. 135: 33-54.

Chickering, Arthur M. 1932. Notes and Studies on Arachnida. II.
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Chickering, Arthur M. 1933. Notes and Studies on Arachnida. IV.
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Chickering, Arthur M. 1934. Araneae from the Northern Penninsula
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Chickering, Arthur M. 1935. Further Additions to the List of Araneae
from Michigan. Pap. Mich. Acad. Sci. 17: 583-587.

Chickering, Arthur M. and Gertrude Bacorn. 1933. Notes and Studies
on Arachnida. V. Additions to the List of Araneae from Mich-
igan. Pap. Mich. Acad. Sci. 17: 521-528.

Curtis, David J. 1980. Pitfalls in Spider Community Studies (Arach-
nida, Araneae). J. Arach. 8: 271-280.

Dondale, C. D. 1962. Spider Research in Canada. Proc. Ent. Soc.
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Drew, Leslie C. 1957. The Spider Fauna 9: Baker Woodlot and
Vicinity, Michigan State University, Including Life Histories
g§_Selected Species. M.S. thesis. Michigan State University.
46 pp.

 

 

 

Drew, Leslie C. 1965. The Spider Fauna pf Beaver Island, Its
Composition, Ecological Distribution and Origin. Ph.D. disser-
tation. Michigan State University. 89 pp.

 

 

 

34

35

Duffey, Eric. 1966. Spider Ecology and Habitat Structure. Senck.
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Fichter, Edson. 1941. Apparatus for the Comparison of Soil Surface
Populations. Ecol. 22(3): 338-339.

Haynes, D. L. 1981. Dept. of Entomology, Michigan State University,
East Lansing, Mich. Personal communication.

 

Hindle, Joseph A. 1972. Some_§piders from Bear Lake Bog. M.S.
thesis. Michigan State University. 12 pp. '

Kaston, B. J. 1948. The_§piders 2f_Connecticut. Hartford. 874 pp.

 

 

Kaston, B. J. 1978. How £2 Know the Spiders. W. C. Brown, Co.,
Dubuque. 272 pp.

 

Levi, Herbert W. and Howard M. Field. 1954. The Spiders of Wisconsin.
Am. Mid. Nat. 51(2): 440-467.

Levi, Herbert W. and Lorna R. Levi and Herbert S. Zim. 1968. ‘A
Guide £2_Spiders and Their Kin. Golden Press, New York. 160 pp.

 

Lowrie, D. C. 1948. The Ecological Succession of Spiders of the
Chicago Area Dunes. Ecol. 29: 334-351.

Mitchell, B. 1963. Ecology of Two Carabid Beetles, Bembidion
lampros (Herbst) and Trechus guadristratus (Shrank); II.
Studies on Populations of Adults in the Field With Special
Reference to the Technique of Pitfall Trapping. J. Anim.
Ecol. 32: 377-392.

 

Muma, Martin H. and Catherine E. 1949. Studies on a Population of
Prairie Spiders. Ecol. 30: 485-503.

Pedigo, Larry P. 1970. Activity and Local Distribution of Surface
Active Collembola (Insects): 1. Woodland Populations. Amer.
Mid. Nat. 83(1): 107—118.

Pedigo, Larry F. 1970. Activity and Local Distribution of Surface
Active Collembola: II. Pond Shore Populations. Ann. Ent. Soc.
Am. 63(3): 753-760.

Turnbull, A. L. 1966. A Population of Spiders and Their Potential
Prey in an Overgrazed Pasture in Eastern Ontario. Can. J. Zool.
44: 557-583.

36

Vail, Harry D. 1974. Spiders Associated With Dwellings 13
Michigan With Special Emphasis gn_the Predatory Behavior
2£_Chiricanthium inclusum Hentz (Araneae: Clubionidae).
Ph.D. dissertaEion. Midhigan State University. 141 pp.

 

 

 

 

 

 

Wallace, H. K. and Harriet Exline. 1977. Spiders of the Genus
Pirata in North America, Central America, and the West Indies
(Araneae: Lycosidae). J. Arach. 5(1): 1-112.

Williams, G. 1958. Mechanical Time Sorting of Pitfall Captures.
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Williams, C. 1959. The Seasonal and Diurnal Activity of the Fauna
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28: l-13.

         

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