THE SPIDER FAUNA OF BAKER WOODLOT AND
VICINlTY, MICE-(5%“ $TATE UNIVERfilTY; iNCLUDmG
LIFE HIS'EDEEES OF SELECTED SPECIES

Thesis {or ‘3er Degree 0? M. 5.
MICHHGAN SMTE UNIVERSE”
Leslie C. Drew
1957

 

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THE SPIDER FAUNA OF BAKER WOODLOT AND VICINITY,
MICHIGAN STATE UNIVERSITY, INCLUDING LIFE

HISTORIES OF SELECTED SPECIES

By

Leslie C. Drew

AN ABSTRACT OF

A thesis submitted to the College of Science and Arts
Michigan State University of Agriculture and
Applied Science in partial fulfillment of
the requirements for the degree of

MASTER OF SCIENCE
Department of Zoology

East Lansing, Michigan
1957

Approve a M, Li-

THESIS ABSTRACT

The purpose of this study was to survey the spider fauna
of a designated area and to reveal the life histories of three
species of spiders.

The survey was restricted to a 250 acre plot located on
the property of Michigan State University. Collecting was
conducted over a period of 19 months. The spiders were col-
lected by visual searching, sifting leaf litter, shaking
bushes, sweeping with an insect net, and by placing pitfall
traps in selected areas. The survey included data concern-
ing habitat preference and distribution. The spiders were
determined to the species level by use of available keys and
publications. The determinations were confirmed by competent
aranealogists or by comparison with confirmed specimens.

The life history studies were conducted under labora-
tory conditions in which the external environmental factors
were not rigidly controlled. Throughout the course of these
studies each spider was centained in a glass vial. Food,
vestigial—winged Qgpsophila melanogaster Meigen, and water
were added regularly. Data were recorded concerning the num-
ber of moults, the duration of stadia, the characteristics of
the egg cases and eggs, and measurements of the body lengths
of the spiders at various instars. The effectiveness of the

methods used in rearing spiders under laboratory conditions

was tested by comparing results obtained for two species of
spiders whose life histories were known with the results of
other investigations of the same two species.

The total number of families of spiders represented in
the area was 20. The results of the survey were presented
in tabular form. Included in the table of species repre-
sented were data concerning the habitat, distribution, and
number of mature specimens collected.

The developmental data concerning Philodromns pernix
Blackwall. Dictyna sublata (Hentz) and Theridion frondeum

Hentz were also presented in tabular form.

THE SPIDER FAUNA OF BAKER WOODLOT AND VICINITY,
MICHIGAN STATE UNIVERSITY, INCLUDING LIFE

HISTORIES OF SELECTED SPECIES

By
Leslie C. Drew

A THESIS
Submitted to the College of Science and Arts
Michigan State University of Agriculture and
Applied Science in partial fulfillment of
-the requirements for the degree of

MASTER OF SCIENCE

Department of Zoology

East Lansing, Michigan
1957

./ - ..
dices/a?
.5 /as30

ACKNOWLEDGMENTS

The author wishes to express his sincere appreciation
to the following members of the Michigan State University
faculty: Dr. K. A. Stiles, Head of the Zoology Department,
for the use of the laboratory space and facilities; Dr. J. L.
Cantlon for assistance in describing the vegetative ecology
of the collecting area; and to Drs. R. L. Fischer and M. M.
Hensley for constructive criticism of the manuscript.

The author is indebted to the following investigators
for their confirmation of determinations and helpful sugges-
tions throughout the course of this study: Dr. R. D. Barnes,
Gettysburg College; Dr. H. L. Levi, Harvard University; and
Dr. M. H. Mums, University of Florida.

To Dr. A. M. Chickering, Albion College, for access to
his extensive collection of Michigan spiders and for his
interest, I extend my thanks and appreciation.

To Dr. T. W. Porter, for his guidance, patience and
evaluation throughout the course of this investigation and
to the late Professor J. W. Stack who provided the opportu-

nity for graduate study, go my heartfelt thanks.

TABLE OF CONTENTS

ACKNOWLEDGMENTS

LIST OF FIGURES

LIST OF TABLES
INTRODUCTION................................... ...... ...... 1
THE COLLECTING AREA........................................ A

THE SPIDER FAUNA OF BAKER WOODLOT AND THE HORTICULTURAL
EXPERIMENTAL PLOTS.......................................lO
Methods..................................................lO

Habitats investigated..................................lO
Types of collecting....................................ll
Procedure in the field.................................ll
Determinations...........................................l2
Procedure..............................................12
Taxonomic characters and available keys................12
Confirmation of determinations.........................13
Tabular presentation of determinations.................13
Discussion...............................................20

LIFE HISTORY STUDIES.......................................26
Purpose and Objectives...................................26
MethOdSooooooooo000000000000000.000.000.000...o.noto0000026

Treatment of egg cases.................................26
Isolation of spiderlings...............................27
Materials and their use................................27
Care of the spiderlings................................28
Control groups.........................................30
ResultSOOOOOOOOOOOOOOOO000.00.00.00...00.0.0000000000000031
Philodromus pernix Blackwell...........................31
Dictyna sublata (Hentz)................................32
Theridion frondeum.Hentz...............................3h
Discu881on0000000.00.00.00.00...0.0...0.0.0.0...0.0.0.0003;

SUMMY.....C....O....O..O00.0.00...00.......C.‘O..........39
' SELECTED REFERENCESLLO

LIST OF FIGURES

Site of the collecting area........................... h
The collecting area................................... S
The divisions of Baker Woodlot........................ 7

Plug of the "feeder bottle"...........................29

II.

III.

IV.

V.

VI.

LIST OF TABLES

A list of families represented in the collecting

area..............................................lh
A list of the species collected.....................lS
Productivity of the habitats........................23
Developmental data of Philodromus pernix............33

Developmental data of Dictyna sublata...............3h

 

Developmental data of Theridion frondeum............36

INTRODUCTION

That spiders are of scientific significance is verified
by the number of publications concerning these animals. -The
publications can be divided into four main categories. First,
are the works which provide a basic foundation in Araneology.
Among these are the works of Hentz (1875), Emerton (1878),
Petrunkevitch (1911, 1923, 1933. 1939, 19h2), Bristowe (1939,
19hl), Cmmstock (19h0), Keston (19h8), and Gertsch (19h9).
The second group, the taxonomic investigations, contains
numerous separate papers on infrafamilial categories. Ex-
amples are Bishop (192A), Exline (1936), Gertsch (1939), and
Levi (1953-1957). Ecological studies on spiders comprise
the third category and include such workers as Lowrie (19h2,
19h8) and the Barnes' (1953, 195k. 19553). Last to be men-
tioned are the life history studies. These studies or rear-
ing experiments are largely restricted to the isolation of
spiderlings and observations on their development with little
reference to environmental factors. Studies by Ewing (1919),
Gabritschevsky (1927) and Baerg (1928) are of this type.
However, that such factors as food, light, temperature, and
humidity affect the number of moults and the length of the
stadia has been confirmed by Browning (19h1, 19h2), Jones
(l9hl), and Deevey (19h9).

The spider fauna of Michigan first received attention
in 189k at Agriculture College (East Lansing) by C. F. Baker.
His list included kl species of 33 genera representing ten
families. During the past 25 years, A. M. Chickering of
Albion College has investigated the diverse fauna of the
state. He has compiled faunal lists for various geographi-
cal areas, listed families represented throughout the state,
and prepared keys to the species in five families. Noland
(1953) investigated the spider fauna of the Proud Lake Rec-
reation Area, Oakland County, Michigan, with reference to
ecological distribution.

The objectives of this study were to collect and deter-
mine the species of spiders of a designated area on the
property of Michigan State University, East Lansing, Ingham
County, Michigan, and to reveal the life histories of three
species present in the area.

Included in the survey are data concerning habitat pref-
erence and distribution. .Contained in the life history in-
vestigations are descriptions of egg cases and eggs, the
number of moults from egg to sexual maturity, and the dura—
tion of the stadia.

Both aspects of the study were begun in June of 1955.
Collecting for the survey was terminated in January of 1957.
- During this time, representatives of 20 families were ob-

ftained. The rearing experiments, which continued until April

of 1957, provided the data necessary to record the life his-

tories of Theridion frondeum Hentz, Dictyna sublata (Hentz),

 

 

and Philodromus pernix Blackwall.

J:-

THE COLLECTING AREA

The collecting area (Fig. l), which was located about

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Figure l. Site of the collecting area with

reference to the center of the Michigan
State University (Beaumont Tower).

 

 

one and one half miles southeast of the center of the Michigan

State University Campus, contained approximately 250 acres.

It was bounded on two sides by well-traveled roads and on the
remaining two sides by railroad tracks. The larger portion,
or 200 acres (Fig. 2), was known as the Horticultural Experi-

'mental Plots. The remaining 50 acres was the Baker Woodlot.

 

 

    
      
 

 

   

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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Figure 2. The collecting area

The experimental plots were not entirely under cultiva-
tion. A portion was covered by an open woods of oak and
sumac. Nine permanent outbuildings were located on the area;
and during the growing season, several temporary structures
were erected in the fields. The area also included a tree
nursery, an apple orchard, a vineyard, and extensive lawns

which surrounded the main buildings. The majority of the

plot area was, however, given over to cultivated fields and
meadows. Dissecting the entire area were roadways bordered
by occasional hickory and oak trees. Collecting was also
conducted in the bordering railroad beds and embankments.
Baker Woodlot received considerable attention because
of the diversity of habitats present there. This woodlot
did not fit easily into a definite ecological description
because of the heterogeneity of the vegetation. For this
study, the area was divided into three sections on the basis
of the drainage property of the predominating soil types,
namely: well-drained, wet, and very wet (Fig. 3). The
following paragraphs are devoted to an analysis of the vege-
tative strata which occurred on these soils.
The well-drained portion consisted of soils of the Hills-
dale and Hillsdale-Miami complexes and was the largest of the
three areas. The upper story was composed of red oak, ggggggs

rubra Linnaeus; sugar maple, Acer saccharum Marshall; beech,

 

Fagps grandifolia F. Ehrhart; basswood, Tilia amerigana Linnaeus;

 

 

and black cherry, Egunus serotina F. Ehrhart; with occasional
tulip poplar, Liriodendron tulipifera Linnaeus; white ash,
Eraxinus americana Linnaeus; white oak, Quercgg alba Linnaeus;

American elm, Ulmus americana_Linnaeus; and slippery elm,

 

H. rubra Muhlenberg. The understory had, along with tree

reproduction, witch hazel, Hamamelis Virginians Linnaeus;

 

arrowwood, Viburnum recognitum Fernald; elderberry, Sambucus

Rubens Michaux; Virginia creeper, Parthenocissus guinguefolig

 

Linnaeus; poison ivy, Rhus radicans Linnaeus; and some run-

ning strawberry, Euonymus obovatus Nuttall. The herbaceous

 

 

E
Farm Lane

 

 

 

 

(/4

...Dry

 

 

 

 

 

\

\
\
\ x

...Wet

 

..Very Wet

 

 

 

Figure 3. The extent and location of
divisions of Baker Woodlot.
This is the spring arrangement.
As the summer approached and
progressed, the dry area in-
creased in extent.

layer of this soil type was composed of bedstraw, Galium

_§giflorum.Michaux; true Solomon's seal, Polygonatum pubescens

 

Willdenow; along with spring flowers, e.g., adder's tongue,

Erythronium albidum Nuttall; Viola spp.; dutchmen's breaches,

 

 

Eccentra cucullaria Bernhardi; Trilium gr_andiflorum Michaux;
which gave way, primarily to Aster spp. and goldenrod, _S_o_li-
dagg spun, as the season progressed. This succession also
occurred on the following soil types.

The wet portion of the woodlot had the Wauseon soil
which was very poorly drained. The trees of this section
were basswood, sugar and red maple, Acer rubrum Linnaeus;
beech, American elm, white ash, bitternut, Carya cordiformis
K. Koch; American hornbeam, Carpinus caroliniana Walter; and
hop hornbeam, Ostrya vigginiana K. Koch; with occasional
swwmp white oak, guercus bicolor Willdenow; slippery elm and
rock elm, Ulmus thomasi Sargent. The shrubby understory had
tree reproduction of white ash, red maple, black cherry,
alternate leaf dogwood, Cornus alternifolia Linnaeus, and

elderberry. Species of Viola, Gramineae and Claytonia to-

 

gether with bedstraw, sweet cicely, Osmorhiza longistylis
Torrey, and black snakeroot, Sanicula trifoliata Bicknell,
were the predominating herbs.

The division classed as very wet (Fig. 3) was charac-
terized by Carlisle muck which in this woodlot supported the
vegetation in and around one vernal and two permanent ponds.

The surrounding trees were cottonwood, Populus deltoides

 

Marshall; black ash, Fraxinus nigra Marshall; black gum,
Nyssa sylvatica Marshall; red maple, basswood, white ash,

American elm, swamp white oak and burr oak, Quercus macro-

 

carpe Michaux. The shrubby layer around or in the ponds was

button bush, Cephalanthus occidentalis Linnaeus; American
cranberry, Vaccinium macrocagpon w. Aiton; and black currant,
Eipgg americanum P. Miller. The herbs in this division were
sedges, Carex spp., and floating aquatics such as duckweed,
EEEEE ERE-

The border of the woodlot differed from the interior in
that sassafras, Sassafras albidum Nees von Esenbeck; black
locust, Robinia pseudo-acacia Linnaeus; prickly ash, Xantho-
.xylum amegiganum; grapes, Vitis aestivalis Michaux; bitter-
sweet, Celastrus scandens Linnaeus; and abundant tree repro-

duction were present.

 

10

THE SPIDER FAUNA OF BAKER WOODLOT AND THE

HORTICULTURAL EXPERIMENTAL PLOTS

The purpose of this portion of the study was to reveal
the spider fauna of the area by extensive collecting and
determination of specimens to the species level. In con-
junction with the collecting, data concerning habitat pref—
erence and distribution of the species was recorded.

Sexually mature individuals were used in determining the
species of spiders. The basis of species determination, in
most cases, is the arrangement of the structures composing

the external genitalia.

Methods

Habitats investigated. After consulting the literature
on the habits of spiders and before beginning collecting,
this investigator analyzed the study area and selected certain
habitats in which the collecting would be done.

The habitats selected in the experimental plot area were:
in the soil; on the soil; permanent and temporary outbuild-
ings; flower heads; herbaceous vegetation of the nursery,
orchard, open woods and vineyard; vegetation of the culti-
vated fields and meadows; tree bark of the open woods, or-
chard, nursery and occasional trees along the roads; and

leaves of shrubs and trees.

11

The habitats selected in the woodlot were: in the soil;
in leaf litter; on stumps and fallen logs; on herbaceous
vegetation of the woods, vernal pond, and in and surrounding
the permanent ponds; on leaves of shrubs and trees; on and
‘beneath bark of shrubs and trees; and on flower heads.

Types of collecting. Visual searching, which enables
one to observe the exact habitat occupied by the spider,
served as the primary method of collecting. In addition to
visual searching and in order to secure a more complete sam-
ple of the spider fauna, shrubs were shaken over a cloth,
areas of uniform vegetation were swept with an insect net,
pitfall traps were placed in selected areas of the woodlot,
and leaf litter which had to be thawed in the winter was
sifted for spiders. Specimens of Agelenidae were frequently
collected by blocking the escape tunnel of their funnel-
shaped web with a small hand hatchet. Also, certain other
spiders were collected as they dropped on a line of silk
after their web had been lightly tapped.

Procedure in the field. Initially, the spiders were
collected in a dry shell vial as recommended by Kaston
(l9h8). By collecting in this manner, one could ascertain
whether the spider was mature or not without causing injury.
If the specimen was mature, it was transferred to a two—inch,
screw-top vial containing a solution of 95% alcohol. Addi—

tional specimens taken in the same habitat were placed in

 

 

12

this vial. By this method, one screw-top vial contained all

the spiders collected in a specific habitat on one date.

Determinations

Procedure. The process of determination began with the
vials containing the collection from a specific habitat for
one date. As the specimens were separated to families and
lower taxonomic categories, the ecological data contained in
the above-mentioned vials accompanied each specimen. This
resulted in working with an increased number of vials; but
each specimen when finally determined to species level had
with it the collection data as originally recorded.

Taxonomic characters and available keys. The taxonomic
characters used at the family level are: the arrangement of
eyes as in Salticidae, Lycosidae, Pisauridae, Scytodidae and
Oxyopidae; leg armature in Theridiidae and the cribilates;
spinneret arrangement of Agelenidae, and Hahniidae; general
body shape as in Thomisidae and Tetragnathidae; size; and
various combinations including the above and other charac-
ters such as tarsal claws and spiracle placement. The key
used at the family level of determination was that contained
in the monograph by Kaston (l9h8) which separates 29 families
and is easily followed. The works by Comatock (l9u0) and
Chickering (1952) also contain keys to the family level.

On the generic level, the taxonomic characters used in
identification may be classed as finer divisions of those

used for family determinations. Again, Kaston (19h8) was

 

13

used as were the papers by the following authors: Bishop
(192A); Chickering (1939, l9h0, l9hh); and Archer (1951,
1951a).

Keys to the species level were found in several papers,
e.g., Seeley (1928), Exline (1936), Chamberlin and Ivie
(19h0), and others. Kaston (l9u8) was particularly useful
here as it contained numerous illustrations which aided in
determination. Also, at this level, one has recourse to the
revisionary papers by Gertsch (l93hyl951), Levi (1953-1957),
and others.

Confirmation of determinations. After using the above

 

works to identify the specimens, the author was fortunate in
being able to compare some of his determinations with those
specimens contained in Dr. Chickering's extensive collection
of Michigan spiders. Several identifications were confirmed
in this way. Still other determinations were confirmed by
the following araeneologists: Dr. R. D. Barnes, Gettysburg
College; Dr. H. w. Levi, Harvard; and Dr. M. H. Mums, Univer-
sity of Florida.

Tabulargpresentation of determinations. The results of
the determinations are included in Tables I and II. The first
contains the families recorded and their breakdown into the
number of genera and species represented. Table II is a
complete listing of the species found together with the speci-
fic area where collected and the habitat occupied at the time

of collection.

 

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TABLE I
A LIST OF THE FAMILIES REPRESENTED IN THE HORTICULTURE
EAPERIMENTAL PLOTS AND BAKE WOODLOT WITH THE NUMBER

OF GENERA AND SFECIES PER FAMILY

 

 

Families Genera Species

 

 

Scytodidae l 1
Pholcidae 1 l
Theridiidae h h
Linyphiidae l l
Micryphantidae 2 2
Epeiridae 7 l2
Tetragnathidae 2 3
Agelenidae h 11
Hahniidae l l
Pisauridae 2 3
Lycosidae u 8
Oxyopidae l 1
Gnaphosidae l l
Clubionidae 2 2
Anyphaenidae 2 2
Thomisidae 6 10
Salticidae 7 8
Dictynidae l h
Uloboridae 2

Amaurobiidae 1 1
Total 20 S2 78

 

15

TABLE II
A LIST OF THE SPIDERS FOUND IN THE HORTICULTURE
EXPERIMENTAL PLOTS AND BAKER WOODLOT SHOWING

HABITATS OCCUPIED AT TIME OF COLLECTION

 

Abbreviations of the productive habitats of the two sec-
tions of the collecting area:

 

 

Experimental Plots Baker Woodlot
VF - vegetation of fields LL - leaf litter
VOW - vegetation of open woods VW - vegetation of the woods
FH - flower heads VVP - vegetation of vernal pond
OB - outbuildings SFL - stumps and fallen logs

SB - shrub bark
LS - leaves of shrubs
TB - tree bark

The figures listed with the abbreviations of the habitats
refer to the number of mature specimens collected in that par-
ticular habitat.

 

 

 

 

 

 

 

 

Baker Hort.
Species Woodlot Exp. Plots

Scytodidae I I. I _ -

l. Scytodeg thoracica (Latreille) OB u
Pholcidae

2. Pholcus phalangiqides (Fuesslin) CB 1
Theridiidae

3. Enoplggnatha tecta (Keyserling) SFL 2 CB 1

u. Speatoda borealis (Hentz) VW h OB 2

S. Theridion frondeum Hentz VW 6

6. Achaearaneg tepidariorum

 

(c. L. Koch) 0B 2n

TABLE II continued

16

 

Species

Baker

Hort.

Woodlot Exp. Plots

 

Linyphiidae

7. Frontinella communis (Hentz)

 

Micryphantidae

8. ggraticelus fissiceps

 

(O. P. Cambridge)

9. Erigpne atra Blackwell

 

Epeiridae
10. Argigpe aurantia Lucas
11. Mastqphora bisaccata (Emerton)

12. Cyclosa conica (Pallas)

 

l3. Mangers maculata (Keyserling)

 

1h. Neoscona minima F. O. P. Cambridge

 

15. N. benjamina (Walckenaer)

l6. Aranea solitaria (Emerton)

 

17. Epeira dumetorum (Villers)
18. E. raji (Scopoli)

l9. trifolium Hentz

 

 

E.
20. E. displicata Hentz
21. E.

pegnia Walckenaer
Tetragnathidae
22. Leucauge venusta (Walckenaer)
23. Tetragnatha laboriosa Hentz

2h. 1. versicolor Walckenaer

LL

LL

VW

VW

VVP

VW

VW

r4 nu n: r4

VOW 3
VF 1
VF 1
QB 2
VF 1
VF 30
VF 2

17

TABLE II continued

 

Baker Hort.

Species Woodlot Exp. Plots

Agelenidae
25. Coras juvenilig (Keyserling)
26. g. medicinalis (Hentz)

27. g, lamellosus (Keyserling)

 

28. Wadotes calcaratus (Keyserling)

 

29. E. new species
30. fl. hybridus (Emerton)
31. Circurina robusta Simon
32. Agelenopsis pennsylvanica
(C. L. Koch)
33. A. utahana (Chamberlin and Ivie)
3h. A. emertoni Chamberlin and Ivie

35. A, potteri (Blackwell)

Hahniidae

36. Neoantistea radula (Emerton)

 

Pisauridae

37. Pisaurina mggg (Walckenaer)
38. 2. mira var subinflata (Hentz)
39. Dolomedes tenebrosus Hentz
Lycosidae

no. Pirata arenicola Emerton

ul. g, maculatu§_Emerton

 

M2. Schizocosa crassipes (Walckenaer)

TB
TB
SFL
VW
VW
SFL

SFL

VW

VW

VW
VVP

LL

VW

VVP

LL
LL
LL

F4 r4 a: r4 a) #4 ta

Hrwww

13

VF

TABLE II continued

18

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Baker Hort.
Species Woodlot Exp. Plots

h3. ngosa punctulata Hentz VVP l

uh. L. helluo Walckenaer VW 2

MS. Pardosa moesta Banks VVP 3

E6. g. milvina (Hentz) LL 1

A7. E. saxatilis (Hentz) VF 3
Oxyopidae

M8. Oxyopes salticus Hentz VF 2
Gnaphosidae

E9. Herpyllus vasifer (Walckenaer) VW 1
Clubionidae

50. Clubiona obesa Hentz LS l

51. Trachelas tranquillus (Hentz) L8 1
Anyphaenidae

52. Alggg gracilis (Hentz) L8 12

53. Anyphaena celer (Hentz) LS 3
Thomisidae

5h. Misumena calycina (Linnaeus) FH h
55. Misumenops asperatus (Hentz) VW 3 EH h
56. Coriarachne versicolor Keyserling LL 1

57. Mysticus triguttatus Keyserling vw l

58. g. funestus Keyserling vw 1

59. Philodromus pernix Blackwell TB 2

60. E, placidus Banks vw l

19

TABLE II continued

 

 

 

 

 

Baker Hort.
Species Woodlot Exp. Plots
61. g, imbecillus Keyserling VF 6
62. Tibellus oblongus (Walckenaer) VVP 5 VF l
63. 1; maritimus (Mange) VW 2
Salticidae
6h. Habrocestum pplex (Hentz) LL 1
65. Habronattus viridipes (Hentz) VF l

 

66. Metaphidippus protervus (Walckenaer) VW 5
67. M, exiguus (Banks) VW 1

68. Paraphidippus marginatus

 

 

 

 

 

(Walckenaer) TB 1
69. Phidippus audax (Hentz) VVP 5 VF u
70. Icius hartii Emerton TB 1
71. Maevia vittata (Hentz) LL 1
Dictynidae
'72. Dictyga sublata (Hentz) LS h
73. Q, volucripes Keyserling VW 3
7h. 2, muraria Emerton VW 1
75. Q, hentzi Kaston VW 2
Uloboridae
76. Uloborus americanus Walckenaer vw 3
77. Hyptiotes cavatus (Hentz) SB 9
Amaurobiidae
78. Amaurobius bennetti (Blackwall) VW 3

 

Discussion

Twenty families were obtained in the collecting area
described above. Comparing this total with the 2h families
found in Michigan (Chickering, 1952), one can readily see
the diversity of fauna in this limited area. This comparison
needs further explanation as two systems of family classifi—
cation were used. The author followed Kaston (l9h8) while
Chickering (1952) used the work of Petrunkevitch (1939) in
part. The latter pair of workers considered Argiopidae as
one family; whereas Kaston classified it as three separate
families, Epeiridae, Theridiosomatidae and Tetragnathidae.
If one employed the classification of Kaston, the total for
the state would number 26 families. or this total, the
family Dysderidae has yet to be recorded for the state bring-
ing the actual number of families for Michigan to 25.

Using the family classification of Kaston (l9h8), the
five families not represented in this survey are Mimetidae,
Atypidae, Oeoobiidae, Oonopidae, and Theridiosomatidae.

Of these listed above, only Mimetidae would occur with
any regularity as two genera and four or five species are
represented in the state (Chickering, 1952). A possible
explanation for the absence of this family could be directly
related to the lack of food. Bristowe (19h1), reported that
species of Theridiidae, specifically Theridigg spp., and

Linyphiidae were the principal source of food for mimetids.

21

Tables I and II indicate that only four genera of the theri-
diids and one genus of the linyphiids were collected in this
area.

Each of the four remaining families have been recorded
three or fewer times in the state. Atypidae, the only Mygal-
omorphae known to occur in Michigan, is represented by a sin-
gle species Atypus milberti (Walckenaer). Lowrie (l9h8)
recorded a specimen from Berrien County and Chickering (1952)
recorded the same species from Jackson County. The distribu-

tion of Atypus milberti as given by Gertsch (1936) included

 

"Northeastern United States from Massachusetts to Wisconsin
and south to Ohio, Pennsylvania, and North Carolina." It
would appear that the occurrence of this species in Michigan
should be more common than has been recorded. Oeoobiidae,
Oonopidae and Theridiosomatidae, all small spiders not ex-
ceeding 3 mm., have one genus recorded for each in the state.
In addition to their size, the secretive habits of the Oeoo-
biidae and Oonopidae may have resulted in my inability to
detect them. According to Kaston, the preferred ecological
niche of the Theridiosomatidae is ". . . damp, dark situ-
ations such as bushes and rocks on the banks of creeks and
in wet moss on the faces of cliffs." Such habitats were
absent in the collecting area of this study.

In Table II, which gives the genera and species of the
families, Gnaphosidae and Clubionidae are sparsely repre-

sented. Both are quite abundant in the state (Chickering,

22

1939, 1952). These families, together with Anyphaenidae, are
part of the group known as "running spiders" (Gertsch, 19h9).
The habitats preferred are on the soil and on leaves of vege-
tation. The collecting methods used in this study were ef-
fective in obtaining representatives of Lycosidae and Pisau-
ridae which occur in the same general habitats. Two species
of Anyphaenidae represented in the state by only three or
four species (Chickering, 1939) were also collected. The
Gnaphosidae and Clubionidae are essentially night hunters
(Gertsch, l9h9) and hide in retreats in debris and vegeta—
tion during the day. If these families were strongly repre-
sented in the area, they would have been more abundant as
were the specimens of Lycosidae and Pisauridae.

Spiders were found in only 11 of the 17 habitats ini-
tially selected. These productive habitats with the percent-
age of the total number of spiders collected in each are
listed in Table III. This table is based on the 270 mature
Specimens collected in this survey.

The burrowing Lycosidae, Geolycosa, was not found in
the area. The author has collected specimens of Q, wrightii

(Emerton), one of two species of Geolycosa occurring in

 

Michigan (Wallace, l9u2), in Clinton and Kent Counties.

A factor influencing the spider population of the herbs-
ceous vegetation of the vineyard, nursery, and orchard was
the frequent use of insecticides. A few immature specimens

of Tetragnathidae, Salticidae, and Thomisidae were collected

TABLE III
PRODUCTIVE HABITATS OF THE EXPERIMENTAL

PLOT AREA AND BAKER WOODLOT

 

 

Experimental Plot Area Baker Woodlot
Percent Percent
of totall Habitat of total2 Habitat

 

56.6 Vegetation of fields h7.5 Vegetation of woods

33.0 Outbuildings 13.8 Vegetation of vernal
pond
7.6 Flower heads 10.9 Leaf litter
2.8 Vegetation of open 10.9 Leaves of shrubs
woods

7.1 Stumps and fallen logs
h.9 Bark of shrubs

h.9 Bark of trees

 

L-lThe total number of specimens collected in the
Experimental Plot Area was 105.

2The total number of specimens collected in
Baker Woodlot was 165.

here. The spraying either reduced the insect population
which served as food or was effective in killing spiders.

From the totals of each section of the collecting area
(Table III), it can be seen that Baker Woodlot supported over
25% more spiders in its 50 acres than did the 200 acre experi-
nmntal plot. This was due in part to the multiple vegetative
strata of the woodlot which increased the area of the substra-

tum and with it the number of ecological niches occupied by

21;

species of spiders. In comparison, three of the four produc-
tive habitats of the experimental plots were limited to the
single herbaceous vegetative stratum. This correlation be—
tween vegetative stratification and increased spider fauna
has been recorded by Lowrie (19h8), R. D. Barnes (1953),

B. M. and R. D. Barnes (l95h), and R. D. and B. M. Barnes
(1955) in their respective ecological studies.

Seasonal variation in the species complex of the mature
spider fauna of the woodlot was observed. The collecting
period for this survey extended from June,l955, through
January, 1957. When the collection was examined at the end
of the 1955 season, it was noticed that the web-weaving
species predominated. During the following spring and first
weeks of summer, the non-web weaving spiders including the
Lycosidae, Pisauridae, and others, were the dominant types.
There were very few mature web-weaving species present at
this time. Again, during the latter portion of the summer
and throughout the fall, it was the web-weaving varieties
that predominated. The underlying basis for such a phenome-
non is the fact that most of the web-weaving spiders mature
during one season, mate, lay their eggs and die; and the
majority of non-web weavers do not mature until very late
in the fall or in the spring of the following year, then
mate, lay their eggs and die. The web weavers overwinter
as eggs and the non-web weavers as mature spiders or imma-

tures in the penultimate or earlier instars.

25

Included in the collection of the woodlot is a mature
female Agelenidae belonging to the genus Wadotes. At pre-
sent, this specimen appears to be a new species. Dr. M. H.
Muma who confirmed the determinations of species for this
family, recognized the specimen as being closely related to
Hgdotes calcaratus (Keyserling). It is hoped that mature
male specimens of this species will be found during the pre-
sent growing season (1957). These would enable the author

to describe the species.

26

LIFE HISTORY STUDIES

Purpose and Objectives
The life history facet of Araneology has not received

as much attention from investigators as the taxonomic phase.
No doubt, the primary reason for this is the amount of time
involved in the rearing of spiders. According to Deevey
(19h9), life history studies for 21 species of spiders have
been completed. The present study contains the life histo-
ries of three species which have not been previously recorded.

AThese are Philodrompg pernix, Dictyna sublata, and Theridion

 

frondeun .

The purpose of this part of the investigation was to
study the developmental sequence of the spider from egg to
maturity. The primary objectives were to observe and record
the number of moults from.the egg to the sexually mature
adult and the duration of the stadia. Secondary objectives
were to record the characteristics of the egg cases, their
contents, and the measurement of body length of the spiders
at various instars.

Methods

Treatment of egg cases. When collected, the egg case
and its substrate were placed in a suitable bottle contain-
ing a strip of tissue paper. The bottle was loosely capped.

Each bottle and its contents were assigned a letter for

27

recording purposes. The bottle was then placed in a position
to receive indirect natural light. Every two days, a few
drops of water were placed on the tissue paper in an attempt
to keep the humidity somewhat constant.

Isolation of spiderling_. The spiderlings were isolated
and put into separate vials not later than two days after
they had emerged from the egg sac. Separation made observa-
tion of the individual spider possible and it also eliminated
cannibalism.

Each spider was given a number which in conjunction with
the letter of its egg case served as its label throughout the
study. This label was printed in permanent ink on the cork
of the vial in which the spider was housed. A three-by-five
file card was kept for each specimen.

After the separation of the entire brood was completed,
measurements and descriptive notes were taken. The first
live food was then introduced into each vial.

Materials and their use. Two sizes of glass, shell
vials were used in the study. The small vial measured
50 mm. x 15 mm. (1% dram capacity), and a larger vial
70 mm. x 31 mm. (b dram capacity). Initially, the smaller
vial was used and as the spiderling grew, a larger vial was
substituted. In the early stages, the smaller vial limited
the area in which the flies, used as food, could move making

them easier prey for the young spiderling.

28

Appropriate sized corks used as stoppers were not forced
into the vial. Rather, they were loosely placed insuring an
air supply for the spiderlings.

Each vial contained a strip of two-ply tissue paper which
was narrower than the diameter and shorter than the length of
the vial. The purpose of the tissue was to absorb any excess
moisture from the bottom of the vial, thus preventing mold.

Care of the spiderlings. The factors of temperature,

 

humidity and light were neither rigidly controlled nor were
they allowed to fluctuate from one extreme to the other.
The temperature did not exceed 85°E.nor did it fall below
68°F. The relative humidity of the vials was different than
the outside atmosphere due to the fact that water was intro-
duced every three or four days for drinking purposes. The
vials were so placed as to avoid direct sunlight. Artificial
light was not shielded from the specimens as no effort was
made to control the period of time light reached the vials.
Vestigial-winged Drosophila melanogaster Meigen were
used as the food source in these studies. Other investi-
gators (Crane, l9u8 and Deevey, 19h9) have used this species
and secured favorable results. In the present studies, the
flies were not anesthetized before being placed in the vials;
nor was the amount consumed rigidly controlled. Every three
days, several live flies were placed in the vials. The num-
ber of flies depended upon the size and condition of the

8Pider. If the spider was in the process of moulting or

29

appeared swollen in readiness to do so, no flies were intro-
duced. For in the former case, they may have caused the
spider to injure itself and in the latter case, the flies
would have been wasted.

The method used to introduce live flies into the vials
was devised by the author during the course of this study.
In methods described in the literature (Brown l9h6, Crane
19h8, Deevey 19h9), either the flies were allowed to crawl
into the vial containing the spider or the flies were anes-
thetized. In the present studies, the flies were trans-
ferred from.the culture bottle to an empty half-pint milk
bottle. This bottle was stopped with a plug as shown in

Figure h. The corks were removed from the vial and spout of

   

Cork

Spout of heavy paper

Figure h. Plug of the "feeder bottle"

the "feeder bottle," the spout was fitted into the mouth of

the vial, the bottle was tapped lightly until a sufficient

3O

anber of flies were in the vial and then both the vial and
spout were again corked.

Water was supplied by placing a drop on the bottom of
the cork that had been removed for feeding. When the cork
was replaced, the drop was pendant inside the vial and acces-
sible to the spider by means of the tissue paper strip used
as a walking surface.

The specimens were fed and watered frequently so that
any change in their condition was easily observed. The cast
skins or exuviae were of major importance and were the basis
of the treatment given a spider. That is, the size of the
vial was changed or more food was added. Each time an exu-
vium.was removed, an entry for that date was made on the
file card of this particular specimen. The data recorded
included the measurement of body length taken as accurately
as possible with a millimeter scale while the spider was
next to the inside surface of the vial. Two days after the
spider had shed, it was again examined and notes on its
coloration were taken. The lapse of time allowed the colors
to deepen. At maturity, the spider was preserved in 95%
alcohol, labeled and placed in the collection.

Control grogp_. In an attempt to check the effective-
ness of the methods used in this study, Achaearanae tepi-
dariorum (C. L. Koch) and Latrodectus mactans (Fabricus)
whose life histories had previously been investigated by

others were reared as control groups. Because external

31

environmental factors such as humidity, temperature and food
can cause variation in the number of moults (Browning 19hl,
Jones l9hl, Deevey 19h9), and because such factors were not
rigidly controlled in the present studies, control groups

were necessary for comparison of results.

Results

Philodromus pgrnix Blackwall. A total of four egg sacs

 

of this species were collected. Of these, two were left
undisturbed and two were dissected. All four sacs of white
silk, collected in October, 1955, were located in forked
twigs of trees. The outer coverings were oval, taut, and
measured 13 mm. x 10 mm., 13 mm. x 9 mm., 11 mm. x 9 mm.,
and 10 mm. x 8 mm. respectively. The two sacs which were
examined had a layer of loose white silk beneath the outer
cover and contained 12 and 17 eggs which measured less than
1 mm. each. These were yellowish—white and non-agglutinate.
One of the two sacs which were not dissected was col-
lected on October 21, 1955, and 12 spiderlings emerged from
it on November h, 1955. The other sac was collected on
October 23, 1955, and 15 spiderlings emerged on November 1h,
1955. Cast skins found in the remains of the two egg sacs
were evidence that when the spiders emerged they were in
their second instar. According to Gertsch (l9h9), all

spiders undergo their first moult in the egg sac.

32

The changes that occurred in this species from the
second instar spiderling to the mature adult were primarily
in size. The emergent spiderlings measured approximately
0.8 nmu and the adults averaged h.53 mm. in body length.
The coloration of the legs darkened from a faint yellow to
amber and the brown spots on this ground color remained more
or less concentrated at the distal ends of the metatarsal
and tarsal segments. The coloration on the dorsal surface
of the emergent spiderlings was predominately a dull gray
with numerous brown speckles. The coloration in the adult
was essentially the same except for a chevron pattern on the
posterior abdomen. The ventral surface was gray at all
stages of growth but the demarcation between the dorsal and
ventral abdominal coloration became sharply defined as the
spider moulted. The ratio of leg lengths of the emergent
spiderlings was 2-l-h-3. This was also the ratio of the
adult leg lengths.

Of the 27 spiderlings, three reached sexual maturity,
two females and one male. Table IV contains the develop-
mental data on Philodrpmus pernix Blackwall.

Dictyna sublata (Hentz). A female of this species was

 

found in the rolled edge of a dead shrub leaf early in July,
1956. She was put in a vial and placed in the laboratory
where she produced the two egg sacs used in this study.

Both of the egg sacs were white and lens-shaped. The

female placed the first egg sac directly on the tissue paper

TABLE IV

THE DURATION OF STADIA AND NUMBER OF

MOULTS FOR PHILODROMUS PERNIX

 

33

 

 

 

 

 

 

 

 

Sex and Duration of stadia in days No. of Measurement

label 3rd hfh 5th 6th 7th Bt moults of adult1
Male 3 13 23 23 20 20 S7 7 u.3 mm.
Female S l 28 19 23 2O 28 7 h.8 mm.
Female P l 27 33 2h hh 22 29 8 h.5 mm.

 

 

 

 

 

 

 

 

 

 

1Measurement was taken at last ecdysis.

in the vial and the next partially overlapping the first egg
sac. When separated from one another, the egg sacs measured
u.8 mm. x h mm. and h-3 mm. x 3 mm. Each was about 1.5 mm.
high.

The examined sac contained 20 agglutinated, grayishswhite

eggs which measured about 0.7 mm. in diameter. The other egg

sac produced 11 spiderlings. When the remains of the sac were
examined, seven apparently infertile eggs were found. This
brought the total of eggs in this sac to 18.

At emergence the spiders measured approximately 1.3 mm.
and were a uniform dull yellow. The adult spiders averaged
3.52 mm. in body length. In coloration the legs and ventral
surfaces of the body segments of the adult were uniformally

gray. The cephalothorax was yellow to brown in front, darker

brown on the sides and thoracic portion.

3h

The dorsal surface

of the abdomen had a mottled appearance of brown on gray.

Of the 11 spiderlings, six reached maturity.

The three

males matured on the average of 208 days after the second

moult whereas the females averaged 186 days.

Table V con-

tains the developmental data of the study on Dictyna sublata

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

(Hentz).
TABLE V ‘
THE DURATION OF STADIA AND NUMBER OF
MOULTS FOR DICTYNA SUBLATA

Sex and Duration of stadia in days No. of Measurement
*label 3rd hth 5th 6th 7th moults of adult]-
Male H 1 15 11 19 98 h2 7 3.3 mm.
Male H 3 ll 15 31 151 6 3.5 mm.
Male H h 10 11 19 92 92 7 u.0 mm.
Female H 2 i 11 15 27 1&3 6 .3.0 mm.
Female H 7 ' 17 55 lb 106 6 3.3 mm.

.—— as -—

Female H 9 2h 58 73 17 6 h.0 mm.

 

 

 

 

 

 

 

1Measurement taken at last ecdysis.

Theridion fgpndeum Hentz. A mature female of this spe-

 

cies and her single egg sac were taken from the lower surface

of a shrub leaf. The female produced three more sacs later

35

in the laboratory. These were white, circular, and measured
3.9 mm., 3.8 mm., and 3.5 mm. in diameter. The eggs measured
approximately 0.6 mm. in diameter, were gray white, non-ag-
glutinated, and totaled 5E, E3, and E1 in the three sacs.

Two weeks after it had been collected, the egg case
that was taken with the female produced 65 spiderlings. The
emergent spiderlings measured approximately 0.7 mm. in body
length and in profile the cephalothorax was of the same pro-
portion as the abdomen. The coloration was a nearly uniform
gray white with the anterior portion of the cephalothorax
being somewhat darker. After the second moult, the abdomen
became globular in outline with the spinnerets in a mid-ven-
tral position, and the color of this body region became pale
orange-tan which was darker than the cephalothorax. The
adults averaged 3.20 mm. in body length and were white with
nearly black markings on the abdomen. The cephalothorax had
a mid-dorsal dark stripe and dark spots on tarsal and metatar-
sal segments. Males and females each took an average of E3
days to reach maturity after the second moult.

Of the original number of 65 spiderlings, ten reached

sexual maturity. Table VI contains the data for these.

Discussion
The studies of Browning (l9El), Jones (19E1) and Deevey
(19E9) have demonstrated the fact that certain external en-

vironmental factors can influence the number of instars or

TABLE VI
THE DURATION OF STADIA AND NUMBER OF

MOULTS FOR THERIDION FRONDEUM

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

SIX and Duration of stadia in days No. of Measuremegt
__ abel 3rd ‘Eth 5th 6th 7th moults of adult
Male L 2 11 I 8 8 1E 6 3.0 mm.
Male L E l7 l7 8 5 3.5 mm.
Male L 8 3E r 3 10 5 3.0 mm.
'Male L 10 21 8 l3 5 3.0 mm.
Female L 1 2 9 8 8 1E 7 3.5 mm.
Female L 5 8 7 35 5 3.0 mm.
Female L 7 11 8 28 5 E 3.0 mm.
Female L 11 1E 7 21 5 3.5 mm.
Female L l3 l7 8 l8 5 I 3.0 mm.
Female L 1E 1E 8 13 5 3.0 mm.
I

 

 

 

 

 

 

 

 

1Measurement taken at last ecdysis.

stadia and moults. Jones (19E1) found that temperature
affects the rate of growth and the duration of the stadia;
also that humidity influences the rate of growth as well as
the rate of mortality with a greater effect on mortality.
Deevey (19E9) investigated the effect of different rates of

feeding on Latrodectus mactans. Her results showed a

37

relationship between the rate of feeding and the number of

moults and length of instars. Browning (19E1) studied the
effect of the factors mentioned above and included light in
an attempt to explain the variation in numbers of moults and
length of instars. It is evident then that the life history
of a spider is influenced by the external environment. In
conjunction with this are the inherited factors (Browning,
19El and Deevey, 19E9) which enable some spiders to with-
stand a greater degree of variation in external environ—
ment.

If the number of moults and stadia can fluctuate under
laboratory conditions, it is probable that spiders do not
have a constant number but rather a range in number of
moults and stadia.

The results obtained by this author for the two control
species closely paralleled and often duplicated the results
of other investigators. In the investigations of Latrodectus
mactans, Thorp and Woodson (19E5) gave the number of moults
as from three to six for males and six to nine for females,
and Deevey (19E9) reported three to eight for males and six
to nine for females. The results obtained on this species
in the present study were five to six for males and eight
moults for females. For Achaeranea tepidariorwm, the second
control species, Kaston (19E8) recorded six or seven moults
for males and seven for females. Ewing (1919) found that

males moulted five times and females four, six, seven, and

38

eight times. The results of the present study were that
males matured in five to six moults and females in seven to
eight moults. Using these favorable comparisons, it was
apparent that the rearing methods used in this study were
effective in obtaining reasonable data on life histories of
spiders.

Throughout the course of the rearing investigations,
the author observed spiders drinking from the drop of water
that had been placed on the cork of the vial. This behavior

was not limited to any particular species of spider.

39

SUMMARY

1. Twenty of the 25 families of spiders known to occur
in Michigan were collected in the Horticultural Experimental
Plots and Baker Woodlot located on the property of Michigan
State University, East Lansing, Ingham County, Michigan.

2. Seventy-eight species belonging to 52 genera of the
20 families were represented in the collection.

3. Stratification of the spider fauna was observed in
the 50 acres comprising the Baker Woodlot. This area sup-
ported over 25% more spiders than the 200 acre experimental
plot area.

E. Seasonal variation of the woodlot spider fauna was
observed over the 18 month collecting period.

5. A mature female Agelenidae belonging to the genus
Wadotes is new. This specimen is closely related to Wadotes

calcaratus (Keyserling).

 

6. The females of Philodromus pgrnix Blackwell moult
seven to eight times. The males moult seven times before
becoming sexually mature.

7. In Dictyna sublata (Hentz), maturity is reached by

 

males after six to seven moults and in females after six
moults.

8. The males of Theridion frondeum Hentz moult five
to six times and the females five to seven times before

becoming sexually mature.

to

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1951. Studies in the orbweaving sgiders (Argiopidae) 1.
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1951a. Studies in the orbweaving spiders (Argiopidae) 2.
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Baerg, W. J.
1928. The life cycle and mating habits of the male
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Baker, C. F.
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Barnes, R. D., and B. M. Barnes
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El

Bristowe, W. S.
1939-19El. The comity of spiders. Ray Society of Lon-
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19E2. The inte ent and moult cycle of Te enaria atrica
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Chamberlin, R. V., and W. Ivie
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Chickering, A. M.
1932. Araneae from the Douglas Lake region of Michigan.
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3’4-9‘3550

1933. Araneae from.the Douglas Lake region of Michigan.
II. Ibid. 1932 XVII: 515-520.

193E. Araneae from the northern peninsula of Michigan.
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1935. Further additions to the list of Araneae from
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1939. An haenidae and Clubionidae of Michigan. Ibid.
193 XXIV: E9-8E, 90 figs.

l9E0. The Thomisidae (crab spiders) of Michigan. Ibid.
1939 xxv: 189-237. 87 figs.

19EE. The Salticidae (jumping spiders) of Michigan.
Ibid. 19E3 XXIX: 139—222, 16E figs.

1952. A revision of families of the spiders in Michigan.
Ibid. 1950 XXXVI: 119-139, 21 figs.

1957. The genus Tetragnatha (Araneide, Argiopidae) in
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Chickering, A. M., and G. Bacon

1933.

Comstock,
19Eo.

crane, JO

19E8.

Crompton,
1950.

Additions to the list of Araneae from.Michigan.
Papers Michigan Acad. Sci. for 1932 XVII: 521-528.

J. H.

The spider book. Revised and edited by W. J.
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J.
The spider. Collins: London. 25E pp., 2E figs.

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1925.

1928.

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Revision of the spider genera Erigone, E eri one
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98 pp., 151 figs.

 

Deeve , G. B.

19 9.

The developmental history of Latrodectus mactans
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Emerton, J. H.

1878. The structure and habits of spiders. S. E.
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Ewing, H. E.
1919. The life and behavior of the house spider. Proc.
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Exline , H.
1936. Nearctic spiders of the genus Cicurina Mange.

Amer. Mus. Novitates No. 850. 25 pp., 2E figs.

Fabre, J. H.

1925.

The life of the spider. Translation by Alexander
Teixeira Da Mattos, Dodd, Mead and Co.: New York,
New York. EOE pp.

E3

Gabritsehevsky, E.
1927. Experiments on color changes and regeneration in
the crab spider Misumena vatia. Jour. Expt. 2001.
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Gertsch, W. J.
193E. Notes on American Lycosidae. Amer. Mus. Novitates
No. 693. 25 pp.

193Ea. Some American spiders of the family Hahniidae.
Ibid. No. 712. 32 pp., E2 figs.

1936. The nearetic Aytpidae. Ibid. N00 8950 19 PP.,
29 figs.

1939. A revision of the typical crab spiders (Misume-
ninae) of America north of Mexico. Bul. Amer.
Mus. Nat. Hist. LXXVI: 277-EE2.

19El. New American spiders of the family Clubionidae. l.
Amer. Mus. Novitates No. 11E7. 20 pp., 53 figs.

19Ela. New American spiders of the family Clubionidae. 2.
Ibid. No. 11E8. 18 pp., 5E figs. ‘

19E2. New American spiders of the family Clubionidae. 3.
Ibid. No. 1195. 18 pp., 37 figs.

19E6. Five new spiders of the genus Neoantistea. Jour.
New Yerk Ent. Soc. LIV: 31-37. 8figs.

19E6a. Notes on American spiders of the family Dictyni-
dae. Amer. Mus. Novitates No. 1319. 21 pp., 30
figs.

19E9. American spiders. D. Van Nostrand Co.: New York,
New York. v-xii+285 pp., 32 pls., 32 illus.

1951. New American linyphiid spiders. Amer. Mus. Novi-
tates No. 151E. 11 pp., 16 figs.

Gertsch, W. J., and A. F. Archer
19E2. Descriptions of new American Theridiidae. Amer.
Mus. Novitates No. 1171. 16 pp., 3E figs.

Gertsch, W. J., and W. Ivie
1955. The spider genus Neon in North America. Amer.
Mus. Novitates No. l7E3. 17 pp., 2E figs.

Gertsch, W. J., and H. K. Wallace
1935. Further notes on American Lycosidae. Amer. Mus.
Novitates No. 79h. 22 pp., E5 figs.

1937. New American Lycosidae with notes on other spe-
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Hentz, No Ho
1875. The spiders of the United States. Edited by J. H.
Emerton, Occ. Papers Boston Soc. Nat. Hist. II.
171 PP.

Jonas, S. E.
19E1. Influence of temperature and humidity on the life
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Ann. Ent. Soc. Amer. 3 : 7-571.

Kaston, B. J.
19E5. New Micryphantidae and Dictynidae with notes on
other spiders. Amer. Mus. Novitates No. 1292.

1A pp.. nu rise-

19E8. Spiders of Connecticut. Bul. Connecticut Gaol.
Nat. Hist. Surv. 70. 87E pp., 21EE figs.

Levi, H. W.
1953. Spiders of the genus Dipoena from.America north of
Mexico. (Araneae, Theridiidae) Amer. Mus. Novi-
tates No. 16E7. 39 pp., 121 figs.

195E. Spiders of the new genus Theridiotis (Araneae:
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195Ea. The spider genera Episinus and Spintharus from
North America, Central America and West Indies
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LXII: 65-90, 55 figs.

195Eb. Spiders of the genus Euryo sis from North and
Central America (Araneae: heridiidae). Amer.
Mus. Novitates No. 1666, E8 pp., 138 figs.

195Ec. The spider genus Theridula in North and Central
America and the West Indies (Araneae: Theridiidae).
Trans. Amer. Microscopical Soc. LXXIII: 333-3E3,
25 figs.

1955. The spider genera Coresia and Achaeranea in Ameri-
ca north of Mexico raneae: Theridiidae). Amer.
Mus. Novitates No. 1718. 33 pp., 8E figs.

 

 

AS

l955e. The spider genera Oronuta and Stemmops in North
America, Central America and the West Indies
(Araneae: Theridiidae). Ann. Ent. Soc. Amer. E8:

333-3142, 36 figs 0

1956. The spider .enus Mysmena in the Americas (Araneae:
Theridiidae§. Amer. Mus. Novitates No. 1801.

13 pp., E8 figs.

1957. The spider genera Enoplognatha, Theridion, and
Peidisca in America north of Mexico (Araneae:
Theridiidae). Bul. Amer. Mus. Nat. Hist. 112.

123 pp., L121 figs.

Levi, H. W., and H. M. Field
l95E. The spiders of Wisconsin. Amer. Midl. Net. 52:

14110-1167, 113 £183.

Lowrie, D. C.
19E2. The ecology of the spiders of the xeric dunelands
in the Chicago area. Bul. Chicago Acad. Sci. VI:
161-189.

19E8. The ecological succession of spiders of the Chicago
Dune area. Ecology 22: 33E-351.

Muma, M. H.
19E6. North American Agelenidae of the genus Cores Simon.
Amer. Mus. Novitates No. 1329. 20 pp., 52 figs.

19E7. North American Agelenidae of the genus Wadotes
Chamberlin. Ibid. No. 133E. 12 pp., EE figs.

Noland, G.
1953. The systematics and ecology of the spiders of
Proud Lake, Michigan. Unpublished manuscript.

Petrunkevitch, A.

1911. A synonymic index-catalogue of spiders of North,
Central and Sonth America with all adjacent is-
lands, Greenland, Bermude, West Indies, Terra del
Fuego, Galapagos, etc. Bul. Amer. Mus. Nat. Hist.
XXIX: iii-v+809 pp.

1923. On families of spiders. Ann. New York Acad. Sci.
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1933. An inquiry into the natural classification of
spiders based on a study of their internal anato-
my. Trans. Connecticut Acad. Sci. XXXI: 299-389.

 

 

 

E6

1939. Catalogue of American Spiders. Ibid. XXXIII:
135-33 0

19E2. A study of amber spiders. Ibid. XXXIV: 119-E6E.

Seeley, R. M. -
1928. Revision of the spider enus Tetragnatha. New
York State Mus. Bul. 27 : 99-1E7, 58 figs.

Thorp, R. W., and W. D. Woodson
19E5. Black widow America's most poisonous spider.
Univ. North Carolina Press, Chapel Hill. v-xi+
222 pp., illus.

Wallace, H. K.
19h2. A revision of the burrowing spiders of the genus
Geolycosa (Araneae, Lycosidae). Amer. Midl. Net.
27. 62 pp., 116 figs.

 

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