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AMERICAN SPIDERS

AND

TEER SPINNINGWORK.

——

A RMATURAL HISTORY

ORBWEAVING SPIDERS OF THE UNITED STATES

WITH SPECIAL REGARD TO THEIR INDUSTRY AND HABITS.

———= (j=

HENRY OC. McCOOK, D. D.,

Vick-PRESIDENT OF THE ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA;
VICE-PRESIDENT OF THE AMERICAN ENTOMOLOGICAL SOCIETY 5
AuTHoR oF “Time AGRICULTURAL ANTS OF TEXAS,”

“Tine Honky AND OccIDENT ANTS,”

ETC, ETC.

VOL. IT.

PUBLISHED BY THE AUTHOR,
ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA.

A. D. 1890.

A G12
/ CORNELL
| UNIVERSITY)

a 2
\. LIBRARY 7

AUTHORS: E.DETTOm,

This Edition is limited to Two Hunprep anp Firry copies, of

which this set is

Subscription No..... haa

Autior’s SIGNATURE,

THE PRESS OF
ALLEN, LANE & SCOTT,
PHILADELPHIA,

See eee eS “a = ’

—-— Sr
a . my 4

PREFACE.

Wir the completion of the second volume of “American Spiders and
their Spinningwork,” I feel that I have substantially ended the task which
many years ago I proposed to myself. That task, as it lay in my pur-
pose, was the description and illustration, in as large detail as possible, of
the spinning industry and general habits of true spiders.

Subsequently, as announced in the first volume of this work, my plan
was so far modified as to make the spinningwork and habits of Orbweay-
ers the principal theme, and to group around the same the industries of
other spiders in such relations and proportions as seemed practicable, In
the present volume I have adhered to this modified plan, but less closely
than in the preceding one, having made large use of the natural history
of other tribes than the Orbitelaric.

It is probable that this volume will be more interesting than Volume I.
both to the scientific and general public. It takes up the life history of
spiders, and follows them literally from birth to death, from the cradle to
the grave; more than that, it goes beyond the sphere of existing faunal
life into the geologic periods, and touches upon the history and destiny of
ancestral araneads who lived in the strange surroundings of prehistoric
continents, the sites of which are embosomed in the rocks, or, like the
amber forests, are now beneath the ocean. The courtship and mating of
these solitary creatures; their maternal skill, devotion, and self sacrifice ;
their cocoon life and babyhood; their youth and old age; their means of
communion with the world around them; their voyages through the air
and dens in the ground; their allies and enemies; their fashion of death
and its strange disguises—these and other facts I haye tried to bring be-
fore the reader in the following pages.

Moreover, my studies have necessarily brought me face to face with
many of the interesting problems, theories, and speculations of modern

science. I have had no pet theory to approve or oppose, and haye not
(3)

4 ‘ PREFACE,

sought to marshal the facts in hand for or against this or that philoso-
phy of life and its origin, Indeed, my aim has been to write a natural
history, and not a philosophy thereof. Yet I have here and there alluded
to matters with which current thinking has much to do. This fact may
also tend to make this volume more generally interesting than the preced-
ing or sueceeding one.

I have not found the difficulties of my task lessened, but rather in-
creased in treating these features of the history. Spiders are solitary and
secretive at the best, and these characteristics have reached their highest
expression in those acts—cocooning, for example—with which a large part
of Volume II. is concerned. It has thus been unusually difficult to secure
a continuous authentic record of habits. Then, again, these studies have
necessarily been only the recreations of a busy professional life, whose en-
gagements have rapidly multiplied, and been more onerous and exacting
in the last six years than ever before. These off labors have, therefore,
continually receded or been suspended before the pressing and more se-
rious obligations of duty. Nevertheless, I am glad to have done so much,
and have great satisfaction in the hope that others, stimulated by my
labors, may pass on through the vestibule where I must stop, and explore
the vast temple of aranead lore that lies beyond.

I have spoken of my task as substantially completed. I do not forget
that the Third Volume yet remains to be finished, and that it is the most
costly and, in some respects, the most difficult of all. But much of the
work thereon is already done, and I feel justified in finishing it in a more
leisurely way. That volume, with the exception of two chapters, will be
devoted to species work, and will present, as far as it seems to me neces-
sary for identification, descriptions of the Orbweaving fauna of the United
States. These will be illustrated by a number of lithographic plates,
drawn in the best style of art and colored by hand from Nature. Plate
IV. of the five colored plates in the present volume will best illustrate
the character of those which are to follow. To the above I will add some
species of other tribes whose habits have had especial notice in this work.

I have now said all that I expect to make public of my observations

of spider manners, with the exception of one chapter on General Habits,
which I have reserved for the opening pages of Volume III., and, per-
haps, a second chapter, which may be necessary for the explanation and
enlargement of matters to which attention may be called by those: who
have followed me in the preceding studies.

——— | ——— Toe

site wit ao" on ml, il im a ie, ls aad 7 ————

PREFACE,

In these opening chapters of Volume III. I shall consider the toilet
habits, manner of drinking, methods of burrowing, moulting and its con-
Sequences, prognostication of the weather, some of the superstitions associ-
ated with spiders, spider silk and its commercial value, and some other
points in the natural history of spiders not embraced in the preceding
volumes.

I again make my thankful acknowledgments of the assistance cordially
given me by various friends and fellow laborers. Dr. George Marx, of
Washington, has been especially helpful by generously placing at my dis-
posal his entire collection of spider cocoons, and also by notes upon the
habits of some of the species whose life history I have described. ‘To Prof.
Samuel H. Scudder I am indebted for various references and hints in pre-
paring the chapter on Fossil Spiders, and for the use of his own publi-
cations. Mrs, Mary Treat and Mrs. Rosa Smith Eigenmann have both
helped me with valuable material sent by the one from the Atlantic coast,
by the other from the Pacific.

H. C. McC.

Tue Manse,
Puivapevputa, July 3d, 1890.

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Urdaneri i? Bera vate i>

TAbin OF CONTENTS OF VOLUME II.

PART I.—COURTSHIP AND MATING OF SPIDERS.

CHAPTER I.
WOOING AND MATING OF ORBWEAVERS.
PAGES
The Mystery of Mating—The Male searching for his Mate—Males relatively Fewer—
Males before Mating—Argiope cophinaria—Stages of Courtship—Aranead Lovers—A
Lover’s Peril—Relative Sizes of Sexes—An unequally matched Couple—Nephila
and Argiope—Sexes that live together—The Water Spider—Quarrels of Males—Fe-
male Combativeness—Methods of Pairing among Orbweavers—A Love Bower. . 15-40

CHAPTER II.
COURTSHIP AND PAIRING OF THE TRIBES.

Love Dances of Saltigrades—Pairing of Linyphia marginata—The Period of Union—In-
terruptions—Agalena neevia pairing—Love beneath the Waters—Caressing—Pairing
of Laterigrades—Lycosids—Love Dances of the Saltigrades—Love Dances of Birds—
Displays are to attract Females—A Saltigrade Harem—Color Development. . . . 41-60

CHAPTER ITI.
COMPARATIVE VIEWS OF VARIOUS MATING HABITS.

Value of general Habits—Value of spinning Habit—Maternity inspires Insect Archi-
tecture—Spider Industry influenced by Maternity—By sexual Feeling in Males—
Disproportion of Size in Sexes—Sexes of equal Sizes—Numerical Proportion of
Sexes—Relative Activity of Sexes—Spermatozoa—Agamic Reproduction . . .. . 61-74

PART II—MATERNAL INDUSTRY AND INSTINCTS.

CHAPTER IV.
MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS.

Cocooning Sites—Argiope’s Cocoons—Leafy Canopies—Contents of Cocoons—The Egg
Mass—Argiope cophinaria—Epeira Cocoons—Cocooning Tents—Cocoons of Zilla—
Cocoon of Nephila—Gasteracantha—Spiders with several Cocoons—Tetragnatha
extensa—Cyrtarachne’s Cocoon—The Cocoon String of Labyrinthea—Cyclosa bi-
furca—Basilica Spider’s Cocoon—Plumefoot Spider’s Cocoon—Uloborus—Double Co-
COOMINnbineAIIONE: sa 8 6 a wy sce leo ee pest aces Pnta a's ofc he hg ROEM IR Che mtn 437 WG)

8 TABLE OF CONTENTS.

CHAPTER V.

GENERAL COCOONING HABITS OF SPIDERS.
PAGES
Cocoons of Theridium—Argyrodes trigonum—Cocoons of Ero—Theridium frondeum or

Theridiosoma radiosum ?—Cocoons in Nests—Carrying Cocoons in Jaws—Pholeus—
Upholstered Cocoon of Agalena—Medicinal Spider—The Water Spider’s Cocoon-—
The Parson Spider—Brooding Cocoons—Mud plastered Cocoons—Cocooning Nest of
an English Drassid—Cocoons in Tubes—Segestria canities and her Cocoons—Dic-
tyna philoteichous—Cocoons of the Territelarie—Trapdoor Spiders—Cocoon of the
Tarantula—Lycosa carrying her Cocoon—The Leaf thatched Cocoon Nest of Dolo-
medes—Pucetia aurora—Nesting Cocoons of Saltigrades—Cocoons of Laterigrades—
The Huntsman Spider and her Ege Cradle—Cave Spiders—Origin of Cave Fauna—
Effects of iCawe ‘iife: a2 008 ao Se ee oat ea » 7 110-158

CHAPTER VI.
COMPARATIVE COCOONING INDUSTRY.

How Argiope weaves her Cocoon—Use of the Legs in Spinning—Equalizing the Output
of Thread—Epeira’s Method—Weaving a Cocoon—Theridium—Agalena nievia—
Beating down the ,Thread—General Spinning Method—Composition of Cocoons—
How Cocoons are disposed of—Protection of Cocoons—Cocoon Forms—Variety and
Complexity—Number of Cocoons ............55. 55 24 was . 159-177

CHAPTER VII.
MATERNAL INSTINCTS: MOTHERHOOD,

Cocoon Sites—Feeding Limits—Secreting Cocoons—Night Cocooning—Ovipositing—Cali-
fornia Trapdoor Spider’s Eggs—Shape of Cocoon—Maternity and Cocoon Structure—
Complexity and Maternal Care—Cocoon Vigils—Multifold Cocooning—Number of
Eggs—Fertility and Exposure—The Mother Turret Spider—The Watch of Dolo-
medes—British Spiders—Special Cases of Mother Care—Feeding the Young—Per-
sonal Care of Young—The Spiderlings—Strength of Maternal Feeling—Mistakes of
Mothers—Unintelligent Instinct—Intuitive Skill—Marks of pebviy: Bere Mud
Cradle Maker—Man’s Method and the Spider’s ............... . 178-205

PART III.—EARLY LIFE AND DISTRIBUTION OF SPECIES.

CHAPTER VIII.
COCOON LIFE AND BABYHOOD.

Adult and Young—Period of Hatching—First Moult—Cocoon Canni! Jism—Escape from
the Cocoon—Delivery by Birds—By Mother Aid—First Days e Outdoor Life—Gre-
garious Habit—Movement Upward—A Tented Colony—Dispersions—The Children
of the Spider Web—Mortality among Spiderlings—Assembly of Spiderlings—Bridge
and Tent Making—A Cantonment and Tower—Argiope aurelia and her Young—
Spider Communities—Spider Colonies—Darwin’s View Examined—Accidental As-
semblage—Squatter Sovereignty—A Cellar Colony—A Camp of Juveniles—Young
Water Spiders—The Spiderlings Pick-a-back—The Turret Spider’s Young—A young
Tower Builder—Follow the Leader—The Young of Atypus—Nurture in the Nest—
Young Tarantulas—Young Trapdoor Builders—Nest Development—Marvels of In-
stinct—Dew covered Webs—Character Habits Inmate ........... . . 206-2655

—~

>
F

TABLE OF CONTENTS. 9

CHAPTER IX,

AERONAUTIC OR BALLOONING HABIT,
PAGES

Flying Spiders—Velocity of Flight—Attitude of the Aeronautic Spider—Frolicsome Spi-
derlings—In the. Air—Controlling the Descent—The Height of Ascents—Floating
Gossamer—Aeronautic Orbweavers—Flossy Balloons—Modes of Ballooning—<Aerial
Navigation—The Huntsman Spider—Around the World by the Trade Winds—Spi-
ders at Sea—Distribution of Species—Gossamer Showers—Their Origin—Dr. Jona-
than Edwards—His boyish Studies of Spider Life—Professor Silliman’s Tribute . 256-282

PART IV.—SENSES OF SPIDERS AND THEIR RELATIONS
TO HASIT.

CHAPTER X.
THE SENSES OF SPIDERS, AND THEIR ORGANS.

Spider’s Eyes—Ocellus—Structure of Eyes—Orbs made in the Dark—Cocooning in the
Dark—Sighting Prey—Night Habits—Color of Eyes—Night Eyes and Day Eyes—
Atrophy of Eyes—Cave Fauna—Sensitive to Light—Limited Vision—Good Sight in
Saltigrades—Lubbock’s Experiments—Eye Turrets—Eye Tubercles—Sense of Smell—
The Peckhams’ Experiments—Olfactory Organs—Sense of Hearing—Organs of Hear-
ing—Effects of Sound—A Disgusted Spider—Communication by Touch—Sensitive-
ness to Music—Attracted by a Lute—The Violin—Beethoven and the Spider—A
Natural Explanation —Auditory Hairs—Wagner’s Studies—Are Spiders Mute ?—A
Male Love Call—Stridulating Crustaceans—Scorpions—Westring’s Discoveries—Strid-
ulating Theridioids—How Sounds are Made—Mygale stridulans—Uses of Stridula-
tion—Mute Mygale—How Tarantula Strikes... ..........-25245 283-322

CHAPTER XI.
COLOR AND THE COLOR SENSE.

Facts of Spider Colors—Beautiful Spiders—Attoid Jewels—Metallic Hues—Colors of the
Shamrock Spider—Color Development in Young—Color and Sex—Moulting Influ-
ences—Colors of Age—Effects of Muscular Action—Influence of Sex—Color Con-
sciousness—Climatie Influences—Influence of Enyvironment—Mimetic Harmonies—
Color of Cave Spiders—Bleached by Sunlight—Color Utility—Industrial Compensa-
tions—Warning Colors—Unconscious of Danger—Color Consciousness—Color Sense
of Spiders—Spiders prefer Red—Mimiery and Colors—Cocoon Colors—Prevailing
Spider Colors—Color of Silk—Metallic Hues—Color Seales... 2... . . . . 828-351

PART V.—HOSTILE AGENTS: THEIR INFLUENCE.

CHAPTER XII.
MIMICRY IN SPIDERS.

Industrial Mimiery—Cutting Ants—Mimetic Trapdoors—Self Protection—Trapdoor
Architecture—Moggridge and his Trapdoor Spiders—Tree Trapdoors—Form Mimi-
ery of Animals—Ant like Spiders—Value of slight Variation—Darwin’s Theory—
Sight of Birds—Birds eating Ants—English Game Birds at Linton Park—The

10 TABLE OF CONTENTS.

PAGES
Great Ant Thrush—Raiding the Driver Ants—Are Wasps Mimicked ?—Ants eat
Spiders—Form Mimicry of Environment—Tetragnatha extensa—Mimicking Knots
and Buds—Color Mimiery—Ambush in Flowers—Misumena vatia—Mimicking Wild
Flowers—English Mimies—Mimicking Bark and the Ground—Natural Selection and
Mimicry—Metallie Colors—Cocoon Mimicry—Cyclosa caudata and her Cocoon—
Young Mimics—A Savyage’s Decoration—Protective Resemblance. .. . . . . . 852-377

CHAPTER XIII.
ENEMIES AND THEIR INFLUENCE ON HABIT.

Perils of Spiders—Season Changes—Animal Destroyers—Rats eat Spiders—Sheep also—
Cannibalism—Goethe on Robber Wasps—The Mud Dauber’s Nest—The Blue Wasp
hawking for Spiders—A Waspling Larva at work—Wasp Poison—The Cicada
Wasp—Pipes of Pan—The Tarantula Killer—Special Selection—Characteristics of
Captives—Social Wasps—Nest Parasitism—The Pirate Spider—A Spider Feud—
Spider Duels—Body Parasites—Parasites on HBggs—Saltigrade Guests—Parasitized
Cocoons—Mold, Flies, and Birds—Foolishly Hostile Man—Arachne as a Forest
Keeper—Arachne a Philanthropist—Influence of Enemies on Industry—Moulting
Tents—Climate Coyers—Self Protective Industry—The Tiger Spider and her
Bower—Elis 4-notata—The Burial of Lycosa—Lycosa tarentula—Trapdoor Spi-
ders—Strange Towers and their Builders—Secret Chambers in the Earth—A new
Use of the Abdomen—Shaping the Cocoon ................. . 378-418

CHAPTER XIV.
DEATH AND ITS DISGUISES, HIBERNATION AND DEATH FEIGNING.

The Decline of Argiope—Fashions of Death—First Stages of Mortality—Sexta’s Death
Record—Death after Cocooning—Limit of Life—Lubbock’s old Ant Queen—The
oldest Spider—Tarantula’s Age—Winter Habits—Winter Dens—Winter Tents—
Hunting on the Snow—Hibernation—Sudden Resuscitation—Death Feigning—Not
Fear Paralysis, but ’Possuming—The Peckhams’ Studies—Wonderful Shamming—

A Spider Stoic—Darwin’s View—Origin of Death Feigning—Voluntary Hypnotism
among Men—Purpose of the Habit—Fabre’s Studies... ......... . 419-445

PART VI.—FOSSIL SPIDERS.
CHAPTER XY.
ANCESTRAL SPIDERS AND THEIR HABITS.

Sites of American Fossils—Scudder’s Studies—Lake Florissant—Cause of Entombment—
Manner of Kntombment—Volcanic Showers—Oeningen Spiders—Fossil Nephila—
Climatic Conditions—European Fossil Spiders—Fossil and Existing Fauna—The
Oldest known Spider—Koatypus woodwardii—Fossil Tunnelweavers—Geological
Position of American Fossil Spiders—Fossil Spinningwork—Fossil Cocoons—The-
ridiosoma Cocoon—Unmodified Industry—The Amber Tree—Sources of Amber—
Amber Land—Amber Bay—Trees running Amber—Deep Sea Storehouses—Break-
ing up the Storehouses—Climate of Amber Land—Insect Food of Amber Spiders—
Spiders of the Amber—Archea paradoxa—Embalming Amber Insects—A Romance
Of Arner Toads co 8 ca i) inl ve) ay wt, ers cab cc Pie Senne ce eee ae SL ea AG)

— 2 ee

be Piare I.—Cotors or Eperka prero.tum.
- aha Facing page 48. See Chapter XI., page 326.

: ; Prare Il.—Minicry or Exyironment—Trarpoor Sprpers.
Sade’ Facing page 128. See Chapter XII, pages 353, 354.

Puiare I1t.—Minicry or ENvinonéenr.
Facing page 192. See Chapter XIL., page 366, sq.

" ve Piare TV.—Corors or Sprpers anp tien Cocoons.
Pees Facing page 288. See Chapter XL, page 323.

Age ‘ Pruatre V.—Some Hymenorrerous Enemies of Spipers.
p Facing page 368. See Chapter XIIL., page 383, sq.

a,
area © WK Py

ew ar

CONTENTS OF VOLUME III.

PART I—GENERAL HABITS AND SUPPLEMENTARY NOTES.

Toilet Habits—Toilet Implements—Toilet Methods—Hair-dressing the Feet—The Tarantula’s
Toilet—Compared with Ants—House Cleaning—Working from a Swinging Platform—
Peril of Untidiness—Purseweb Spider’s Cleanliness—Drinking Habits—Tarantula at the
Bowl—Lugging Drops of Mist—Drinking the Dew—Swaying the Body—Pholeus as a
Dervish—Night Habits—Prowling—Sitting in the Hub—Water Habits—Rafting Dolo-
medes—Burrowing Methods—The Tiger Spider—Turret Spider—Tarantula’s Pick and
Wheelbarrow—Tigrina’s Courtship—Mating of Dictyna philoteichous— Moulting Habits
in various Tribes—Wagner’s Notes—Renewal of Lost Limbs—The Process Described—
Weather Prognostication—Stories and Traditions—Records of Several Years—Arachne as a
Weather “Indicator”—Superstitions about Spiders—Good Luck— Money-spinners—Spi-
der Silk—Its Use in the Arts—Its Economical Value.

PART IIT—DESCRIPTION OF ORBWEAVING SPECIES.

PART III.—COLORED LITHOGRAPHIC PLATES AND EXPLANATIONS.

PART L—COURTSHIP AND MATING OF SPIDERS.

CHAPTER. I.
WOOING AND MATING OF ORBWEAVERS.

qT,

THERE is nothing in the life history of spiders that seems to me more
mysterious and wonderful than the faculty by which the male finds the
female to fulfill his office in Nature and fertilize the eggs. Over all diffi-

culties and distance, through the midst often of a multitude of

ae ae individuals of various families and genera, and with apparently
ery 0 ea Hate :
Mating unfailing accuracy, the males of the several species find their

appropriate mates.

It is impossible to determine definitely how wide is the circuit over
which is scattered any single brood of spiderlings after its exode from the
cocoon. Circumstances may confine all the individuals to a comparatively
narrow space. More commonly, perhaps, through the aeronautic habit, by
the agency of passing winds, they are dispersed throughout a wide region.
Under ordinary circumstances, at least, the space is practically impassible
by spiders whose habits are as sedentary as those of Orbweavers. Yet such
is the power of the marital sense, and so strong and true the guidance of
sexual feeling, that, over all barriers of environment the male reaches his
proper consort. As far as I know, he never makes a mistake by falling
upon the web of an.alien species. At all events, if such error occurs, he
knows enough to promptly turn away. :

The partner whom the Orbweaver gallant seeks is commonly seated in
a well isolated nest, or at the hub of her snare, separated by a distance of

several inches from him as he travels over the leaves, twigs, and

na ’ other material upon which the foundations of the orb are hung,
Pee i (See Fig. 1.) The errant lover’s difficulty in finding a mate must

His Mate. Certainly be increased by this fact, for in his cautious approaches
he is not able to draw very near, but must determine through a
distance relatively great the question of identity: “Is this a partner of my
species or not?” He touches the outer foundation line of the orb, and
determines the question from that position. If he is satisfied, he settles
near or upon the web, and awaits the issue of his courtship.
And now, how has he determined, simply from contact with the snare
spun by his chosen spouse, that this is the individual whom he seeks?
(15)

16 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

What trace has the female left of her identity? By what subtle influence
does she attract her wooer to settle in her vicinity? By what strange
responsive power does he know the signs, and discern that his mate and
the mating hour are nigh? ‘There is no fact in the life of spiders that
has struck me with greater force as an unsolved mystery of Nature than
this. I have no suggestions to offer in answer of the queries raised, but
proceed to give such facts about the pairing of spiders as have passed
under my observation, and been gathered from the records of others.

To arachnologists such studies are of special value. In the systematic
grouping of spiders, among the characters to which later students give
greatest. force are the distinctive organs of the male and female. The
characters of the palp on the one, and the epigynum on the other, dom-
inate the decisions by which species are determined. It is certainly reason-
able to infer that if the external forms of these organs are of such con-
trolling value in determining species, the use of the organs, or, in other
words, the manner of pairing, might be expected to show characteristic
differences. In point of fact we so find it; and the reader will be able to
determine how closely the one may correspond with the other. I venture
to add the suggestion that habits which stand at the very gates of life
must have especial value in the natural history of such creatures as we
are studying, and no artificial delicacy should turn aside the student.

It seems probable that fewer male spiders than females are hatched
from the eggs; or, that fewer reach the adult state. At least, one finds

not only in collections, but in field observations, that females
Males = commonly greatly outnumber males. It would follow that one

ee male spider probably serves as gallant for several females, a
Fewer, ‘species of polygamy which reminds us of the barnyard chanti-

cleer. This fact, as has been said,! would indicate that the peril
which an aranead husband is commonly supposed to undergo during
courtship has been considerably exaggerated by writers. According to
De Geer, in his observation upon Linyphia montana, a single male suffices
for many females, to whom he pays his respects consecutively in the same
hour.2. Mr. Campbell saw one male in union with three females of Tege-
naria guyonii during twenty days in August.® Professor Peckham records
similar facts among the Saltigrades. Thorell speaks of the male as “the
rarer sex,’ and Darwin was informed by Blackwall that males are more
numerous than females with a few species, but that the reverse appears to
be the case out of several species in six genera. On the other hand, Mr.
Campbell captured ten spiderlings of Tegenaria and found that seven of
them showed the swollen palps of the immature male.®

' Emile Blanchard, quoted from Reyue des Deux Mondes in “Popular Science,” Octo-
ber, 1888. 2 Vide Walck., Apteres, Vol. II., page 411, suppl.

® Linn. Soe. Jour. Zool., Vol. XVII, “ Pairing of Tegenaria guyonii,” page 167.

4“On European Spiders,” page 205. °“ Pairing of Teg. guyonii,’ page 168.

— a

_—

WOOING AND MATING.

Fic. 1, Snare and nest of the Shamrock spider. The orb, nest, and surroundings

show the field of courtship among Epeiroids.

18 AMERICAN SPIDERS AND THEIR SPINNINGWORK,

It is perhaps not strange that there should be such wide differences of
opinion, since the conclusions are based chiefly upon the indications of
collections. Now, in Nature, the males show themselves in great-

sani est numbers at the pairing period. They appear to mature a
Mating. little earlier than the females, and their solicitations have begun

even before there is reasonable hope for favorable response.
Thus, at this particular time they may be found by a collector more
readily than at any other, and would show in larger numbers in his col-
lection. As most males disap-
pear shortly after maturing, and
are probably not long lived, while
the female survives until after
cocooning, collections made after
the mating time would be lack-
ing in males.
I have seen four males of the
Banded and three of the Bas-

es ket Argiope respective-
ne. N
Wee
Ey

AY /S
AS) (\
Za ANY,

Several
Gallants.

Dene
R\ ly hanging at the same
tt IN
IK UX

time upon the margin
4 IRAP 2
WARS /
WN 7

of one female’s snare. I have
A

observed two and three males of
the Labyrinth spider waiting in
the outer courts of the habitation
of the female of that species, and
the same number of the Insular
spider ranged near the leafy bow-
er of my lady Insularis. I have
seen two males of Agalena nevia
approaching at one time the door
of their lady’s silken chamber,
| * although it must be said that one
of them promptly ran away when
he found that his rival had come
nearer than he. It is not unlikely that many females deposit their eggs
without previous fertilizing; at all events, I have frequently found cocoons
containing infertile eggs. But in the long run, in view of such facts as
the above, it is scarcely to be questioned that Nature, who always knows
how to hold an eyen balance in the product of her living creatures, pro-
vides a master for every mate.

‘Fic. 2. Males of Argiope cophinaria courting the female.

Tl.

The males of Argiope begin to mature about the middle of July, and
they anticipate somewhat the maturity of the female. They may be found

WOOING AND MATING. 19

at this period occasionally occupying separate webs, but more frequently
domesticated upon the orb of the female, upon which several will be found
congregated. For example, in a clump of grasses I found the

The __ web of an apparently mature female, to whom three males were
Bs, ee paying attention. wo of the males were established upon the

ope. outer margins of the female’s snare, upon small rudimentary

webs. The third had built a separate snare immediately behind
the female. There he hung in the usual position at the hub, which was
covered with light straggling lines, a kind of imitation of the ordinary
shield. Above and below were two
faint, irregularly formed ribbons, mere
suggestions of the beautiful ribbon
spun by the female. This snare had
about twenty-one radii and twelve or
thirteen spirals beaded apparently in
the ordinary way. he web was about
four inches in length and about two
inches wide.

On the same day several males
were found on separate webs. These

webs are ordinarily quite ru-
ee ek dimentary. In one the upper

ale Ar- : “ae

giope. part consisted principally of

a mass of straggling lines
somewhat resembling a shield of the
female when it is first spun. The
lower part had ten radii concentrated
upon the hub and all of them crossed
by beaded interradials. The occupant
hung to the upper part of his snare
and stretched his legs over the lower
part. The snare in width was little
greater than the spider’s length meas-
ured from the tip of the hind legs to the feet of the fore legs. In other
words, he spanned his entire web.

Another and similar male snare was found spun into the protective
wings of a mature female snare. A figure of this rudimentary web is
given, (Fig. 3.) On the whole, my observations justify the conclusion that
after the male spider matures, the character of lis web is rudimentary, after
the manner above described and sketched. Previous to that period he ap-
pears to form the characteristic web of the species, quite like young females.
In the immature state, the male Argiope differs from the mature individ-
ual; it then resembles more closely the female in shape and the markings
upon its back. Indeed, at first glance, it would be taken for a young female.

Fig. 3. Male Argiope cophinaria upon his snare.

20 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

In other species the conditions of male spinningwork are different. For

example, I found,’ in a grove of young oak trees, a number of males of

Epeira insularis established in nests, and with perfect orbs spread

Snare and}eneath them. The nests were well sewed, and like those of

ia a the females, which were numerous in the vicinity. The orbs

Insularis, Were also perfect, and of the typical sort. These males were

mature; some had their nests built close to those of females,

upon whom they were evidently in attendance. In several cases two and

three males were seen in the same neighborhood, occupying nests or hang-
ing about the margins of the same female’s snare.

One male of Argiope cophinaria was found on the same day (August
28th), which had spun a tolerably perfect snare twelve inches behind the
orb of a mature female. ‘This snare had twenty-eight radii and nine
spirals, and the flanks were protected by wings or fenders of the typical
sort heretofore described.2 Thus, there appears to be a striking difference
in the character of the web made by the male of this species and that
woven by the male Insularis. In Cophinaria the orb is certainly not per-
fect after the type of the species, but in Insularis it appears in every re-
spect to conform to the type, as does also the leafy nest or tent. It may
be added, as perhaps throwing some light upon such a difference, that the
male Insularis is a larger and more formidable animal than the male
Cophinaria, and relatively much more equal in size and strength to his
mate.

The first stages-of courtship have already been indicated. Having found
the snare of his partner, the male Orbweaver stations himself upon the
: outer border and awaits results. It is not difficult for him to
First communicate his presence. Indeed, he must take his place deftly
Stages of P : ‘ 4 ; F
oink. and keep it quietly upon the snare, or he will quickly bring
ship. down upon him the voracious lady of the house, A touch of
his claw upon a radius would telegraph to the female the fact
of his presence; and I believe, from what I have seen of the operations
of the male in this preliminary courtship, as well as from the recorded
observations of others,? that he does thus intimate his presence, and that
the first stages of the engagement are consummated by these telegraphic
communications back and forth between male and female over the delicate
filaments of the silken snare.

If matters be favorable, the male draws nearer, usually by short ap-
proaches, renewing the signals at the halting places. Sometimes this pre-
liminary stay is brief; sometimes it is greatly prolonged. I have known
it to be continued during several days, in which the male would patiently

1 August 28th, Niantic, Connecticut. 2 Vol. I., chapter vi., page 105, Fig. 96.
2 See the statements of Waleckenaer, Menge, and Emerton, further on.

WOOING AND MATING. 21

wait—sometimes, but not always, changing his position—until his advances
were favorably received, or were so decidedly repulsed that he was com-
pelled to retire. With Labyrinth spiders I have generally seen the male
stationed upon the maze, or that part of the snare which consists of
crossed lines. Here he would make for himself, as he hung back down-
ward, a little dome of spinningwork, which spread above him
like a miniature umbrella. (Fig. 4.) The male of Argiope
cophinaria feels the web with his feet for some time! before
the final approach. The male of Linyphia marginata, as he cautiously
approaches, pulls upon the threads connecting his own with his lady’s
bower.2. The male of Epei-
ra diademata commences his
courtship by touching with
one leg a thread of his lady’s
web.?

Professor Peckham’s ob-
servation upon the courtship

. of Argiope coph-
Argiope’s inaria is to the
ae a same effect. When

advancing towards
the female, the male seems to
pause and pull at the strands
of the web, as though to no-
tify her of his approach.
When he comes toward her
from the front she imparts
a slight motion to the web
with her legs, which seems
to serve aa & warning, ae hé "4, Tce note Tayrhin mites
either moves away or drops male is the upper figure, in the nest.
out of the web. When he
comes from behind, she pays no attention to him until he begins to creep
up on her body, when she slowly raises one of her long legs and brushes
him off.

The same author watched the successive and unsuccessful approaches
of three males who were paying their court to a female Argiope argyraspis,
The warning vibration of the web as the males approached was noted in
this species also, and Professor Peckham believed that the female recog-
nized from the character of the vibration the advent of a male, distin-

guishing the movement of the lines from that made by a struggling
a ES ES a ae ee eee os
* Emerton, “Habits and Structure,” page 87.
* See my description of the pairing, hereafter.
* Termeyer: Proceedings Essex Institute, Vol. I, page 71,

Love
Signals.

22 AMERICAN SPIDERS AND THEIR SPINNINGWORK.,.

insect entangled in the meshes of the snare.'| Of this there can be no
doubt, the female appears to be always conscious of the presence of a
male of her species, as distinguished from all other intruders.

The period of approach or courtship is generally terminated by a sud-
den rush, which brings the partners into union. The advances, as far as
I know, are made by the male; rarely by the female—directly,
sey A at least. They are not always received with favor; and it is
‘ undoubtedly true that the male is sometimes sorely put to it to
make his escape from the premises of an unresponsive female, and occa-
sionally prosecutes his amours at the cost of life. Menge, in the course
of his experimental observations, lost many males, after feeding them un-
til mature, by introducing them into a cylinder containing females.? ‘Ter-
meyer records, with a “surprise and indignation” which seems refreshing
to modern observers, that a male Diadem spider, after the act of union
with the female, was attacked by his spouse, and, happening to be in such
close quarters that he could not escape, was deliberately enveloped in her
threads and deyoured.*

I have watched this point with great interest in the experimental colo-
nies upon my vines. Many males of Argiope cophinaria have been found
trussed up and suspended on the snares of females upon whom I had seen
them in attendance but a little while before. Two males were thus de-
stroyed by the same female in one day. In some cases the males would
be tolerated for several days, even though they hung quite near, and then,
without any apparent reason, would be suddenly found killed and hung
up in silken bonds close by my lady’s bed at the hub of her orb. In
these cases there can be no doubt that the female knew the character
of her visitor during all his stay. Any other creature thus intruding would
at once have been attacked. The amatory feeling was evidently strong
enough to tolerate her lover’s presence for several days, but not sufficiently
warm to encourage the further advances which he made, and which cost
him his life.

One female was attended for a number of days by a male who kept
near and just above her, often feeling her gently with his fore legs. I
supposed the female to be mature, but could not decide without capturing
her. However, I one day found her moulting, apparently the last moult
preceding complete maturity. A few hours after the moult I found my
patient gallant trussed up and hanging close by his lady love, who had
not deigned to eat him. (Fig. 5.) In spider world, at least, it would
sometimes seem an ill advised action to “haste to the wedding.” I have,

1 Sexual Selection in Spiders, page 55. 2 Preussische Spinnen.
* Proceedings Essex Institute.

WOOING AND

MATING,

Fic. 5. Female Argiope with a fresh moult and slaughtered mate
langing to her web.

24 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

but less frequently, noted similar treatment of the male Insularis by his
mate. He is better equipped for taking care of himself than the male
Cophinaria, but, nevertheless, sometimes pays the penalty of his rashness
and importunity.
Notwithstanding the above facts, I have reason to know that matters
are sometimes reversed, and the femal’ is the victim of the cannibal ap-
petite of the male. Among my own specimens, for example, I
Males have had a male of the Furrow spider, which was enclosed in a
Destroy . : : : :
Females, Jar along with two females, satisfy his hunger by devouring one
of his partners. Baron Walckenaer saw a male of Epeira incli-
nata take advantage of a female of his species, which was not able to stir
without difficulty, being full of eggs, to attack, garrote, and eat her.1
Mr. Campbell observed the male of Tegenaria guyonii destroying the
female. Of one pair which he placed together, the male at once began to
pay his addresses. Shortly afterward he rapidly applied one of his palps
to the female, in the manner elsewhere described, and, apparently, with her
consent. Five hours afterward he charged his partner, tore away two legs
below the trochanter, and began to suck one, using the mandibles to hold
the limb, just as a human being would a stick of asparagus. The female
died an hour afterward. This female lacked one moult of being mature ;
but her killing cannot be explained by her supposed sexual incapability,
for Mr. Campbell says he saw two males similarly dismember their spouses
an hour after union. .Hunger could not have been the cause of this feroc-
ity, for they were well fed. In fact, males in confinement take their food
much better than females, which may be due to their being accustomed to
feed, during their sexual excursions, in places which are strange to them.
Only twice did Mr, Campbell see a female of Tegenaria drive the male
away. In both cases this occurred immediately after union. On the other
hand, as illustrating the difference which individual disposition or circum-
stances may produce, the same observer kept together an adult pair of
this species from the 22d of August to the 28th of October, more than two
months, and they lived in perfect unity. The male never ceased paying
unrequited attentions, except to feed.?
Excepting one spider, Argyroneta aquatica, whose male is larger than
his mate, all those found in Great Britain have the female either equal
; in size to, or else larger than, the male. (See Figs. 9, 10.) The
Relative difference, however, between the sexes in these northern regions
Sizeof . : pels :
Sexes, 18 not carried to the extreme limits which are frequently reached
in the tropics. For example, Nephila chrysogaster Walck., an
almost universally distributed tropical Epeiroid, measures two inches in
length of body, while that of the male scarcely exceeds one-tenth of an
inch, and is less than one thirteen-hundredth part of her weight. In other

! Aptéres, I., page 148. * Pairing of Tegenaria guyonii, page 168.

Fic. 6,

WOOING AND MATING.

Female and male of Nephila nigra. Natural size.

(After Vinson.)

26 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

words, the female is twenty times as long and thirteen hundred times as

heavy as her partner.' Dr. Vinson? strikingly represents this disparity ~

of size in the species Nephila nigra (Vinson), which is here presented,
(Fig. 6), with both sexes natural size. A full grown female of our Basket
Argiope bears about the relative proportion to the size of her male, of a
horse to a large dog. The largest female Argiope measures in body length
one inch, in spread of legs three inches. Her abdomen is thick in pro-
portion. A male has a body length of one-fourth inch, the spread of legs
being one inch and a quarter. Fig. 14 will show the relative body lengths
and sizes of the sexes of Argiope cophinaria.

This disproportion, however, in the size of the sexes is not universal. In
some species, as will be found by a reference to the plates in Volume III., the
difference is slight, and, indeed, is sometimes on the side of the male, even
among Orbweavers, as in the case of Epeira strix. Moreover, the males
have relatively longer and apparently more powerful legs than the female.
The increased length must be serviceable in the preliminary courtship,
when the males stand off and solicit or test the feeling of
their mates by touches of the fore feet. These features are
also beneficial in clasping their mates during amatory em-
brace, and must add to their muscular vigor both in conflict
and retreat. This difference in the legs, I have no doubt,
fully compensates for difference in body size in the case of
many species. Especially is this true in the case of the Wan-
derers, with the exception, perhaps, of some of the Thom-
isoids. Moreover, the legs of some Orbweavers are armed
ub with formidable weapons in rows of strong spines arranged
Fic. 7. Tibial spines. along the inner surface of the tibia. (Fig. 7, Tibial spines

(After Emerton.) of Kpeira domiciliorum, a, and Epeira trivittata, b.)
There is also a difference in size among the individuals of any one
species. I have found females (Epeira vertebrata) quite mature, making
: cocoons, who were scarcely more than half as large as others of
Size Va- the same species, and to some extent a like difference prevails
riation in . . .
Slee in the sizes of the other sex. It may readily occur, therefore,
Species, that a large male and a small female will come together, and
thus, in point of strength, be placed more nearly upon an equal-
ity, or even give the preponderance to the male. In such a case his oppor-
tunity for feeding upon his partner is quite as good as hers. For these
reasons I am disposed to think that the perils of courtship dé¢pend, first,
upon the relative size of the individuals, and, second, upon the chances
of arousing the voracious appetite of one or the other partner by unusual
movements. In other and ordinary cases, Nature provides a sentiment

1 Cambridge, “Spiders of Dorset,” introduction, page xxvii.
2 Spiders of Réunion, Maurice, and Madagascar, Pl. VI.

coalite
—

WOOING AND MATING. oT

strong enough to protect the origins of life. It may be added here that
the only two species of Orbweavers which are said never to repel the ad-
vances of the male are those belonging to the genera Tetragnatha and
Pachagnatha—genera, by the way, which are marked by the most formidable
developments of the mandibles, particularly in the male, the chief organs
of attack and defense among spiders. This would seem to be an example
of the theory that the best way to insure peace is to be thoroughly pre-
pared for and formidable in war.

While the above facts represent the relations between the sexes of spiders
as they generally exist, there are some exceptions which present our araneads
under a more domestic character. Among the Retitelaria and
many families of the Theridioids, the males dwell a long time

with the females on the same web. I have fre-
More Do- quently observed the male of Theridium tepedari- =
mestic . ;
Habits. OU, the most ferocious and formidable of our

common species, stationed with comparative secur-
ity upon the cobweb of the female. The interesting little
black Lineweaver, known as Steatoda borealis, I have very
often found underneath stones, or in webs of crossed lines in
other situations, where the two sexes dwelt together in ap-
parent harmony. I believe that this is an habitual domestic

i Ae . . m : : fic. 8. Male (up-
characteristic of this species. The pretty Lineweaver, Liny- “jer figure) and
phia costata,' also belongs to the few American species known prone oe
to me of which the male and female occupy a common home. :
I have nearly always seen them together upon the same slight snare of
lines stretched between the edges of a leaf, or in like situations.

Dolomedes mirabilis of Europe is said to share with the female the care
of their posterity. He will take up the cocoon which the female drops,

place it under his breast, and defend it until the little ones are
Sexes —_ hatched.2 The males of Epeira apoclisa of Europe are said to
Living ; : : aes
Mopether: dwell with the females in the same nest, without inflicting or re-

ceiving injury. According to De Geer the male and female of
Epeira fusca dwell harmoniously near each other, at least in the spring-
time. ‘The male is a little apart from the female, and sometimes ventures
to promenade the common web without receiving any injury from his
partner.® Walckenaer confirms this observation.4 The little English spider,
Ergatis benigna, has earned its pleasant specific name “benigna” by the
fact that she lives peaceably with her husband in the same nest. This is

‘The Linyphia phrygiana of Europe, according to Emerton. 2 Walckenaer.
‘ L’Hist. Nat. des Insectes, Tom. 7, page 236, + Apt., Vol. IT., page 85.

28 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

constructed by the joint labors of the wedded couple, and is a loosely
framed den, with quite open meshes, spun upon the blossoming top or be-
tween the stalks of grasses.1
Of the beautiful European Orbweaver, Epeira quadrata, Menge states
that towards the end of July he observed five nests in which the two sexes
lived together peacefully. ‘These nests are similar to those made
Tempo- by our Insular and Shamrock spiders, which are dens of folded
2 plete leaves, whose interiors are tapestried with silk. 'The female Quad-
‘rata occupied the upper part of the nest, having her fore legs
doubled up so that the knees projected above the head. The male occu-
pied the opposite part of the tent, and kept his legs folded as conveniently
as was possible under the circumstances without elevating the body. This,
however, was not a permanent arrangement, but only a preliminary stage
of courtship, and doubtless terminated when the act of pairing occurred.?

Fig. 9. Fie. 10.

Fic. 9, female, and Fic, 10, male Water spider, Argyroneta aquatica. (After Blackwall.)
ry

I have seen the male and female of our Epeira insularis and trifolium occu-
pying the same tent, apparently under similar circumstances, and regarded
the situation as exceptional. Certainly these species habitually live separate.

The Abbé de Lignac, having placed a large number of Argyronetas in
a bottle, found that they devoured each other. The male, says he, which
was perhaps the only one, had been sacrificed to the jealousies
of the females, who after him were mutually destroyed.* Baron
Walckenaer records a fact which appears to be contrary to this.
He put a number of Water spiders in a glass vase along with some gold
fishes. Within the vase he placed a bunch of coral, and observed a female
make her bell shaped nest and attach it to a branch of the coral, and a

Domes-
ticity.

1 Staveley, British Spiders, page 120.

* Menge, Prussian Spiders. Under Epeira quadrata.

*’ Memoire pour servir 4 commencer l’Histoire des Araignées Aquatiques, page 52. Paris,
1748. By Joseph Albert de Large de Lignac. I quote here and elsewhere from the original
edition in the library Acad. Nat. Sci. of Philadelphia.

WOOING AND MATING. 29

large male construct his domicile at the side of this female. He was wit-
ness of their caresses and their amours, but having been forced to be ab-
sent he could find upon his return only the male and a few young spiders.
All the females, to the number of seven or eight, had disappeared. He
was not able to recover them, and supposed that they had been devoured
by the male, who was in good condition and very lively. However, he
never could find any of the débris of the legs and mandibles, and a spider
is not able to devour these hard parts.!

Clerck kept together one male and ten females of these spiders for many
successive days, during which they were not provided with food, without
having observed the least disagreement.2 De Geer placed many males and
females in the same vase, and they never attempted to injure each other,
He observed that when they encountered one another in the water they
mutually felt each other with their legs, embracing with some vigor, and
whether male to male or female to female, they opened wide their formi-
dable mandibles with such force that for the moment the observer expected
to see them give the death stroke. But they did nothing. After having
felt one another for a long time, they separated and swam each to his own
cell. De Geer placed water insects into the vessel. The same spiders, who
had been so tolerant of one another, instantly seized and devoured these
creatures, their natural prey. It seemed to Baron De Geer that the Argy-
ronetas were less cruel than terrestrial spiders.?

Cambridge states that the two sexes of Agalena labyrinthea may be
found in great amity together in their tubular retreat; so also the sexes of
Meta segmentata, Linyphia marginata, and other species inhabit
the same web when adult. This statement is made without any
qualification, but I am inclined to think, judging from what I know of the
American congeners of these species, that the inhabiting of the web by the
two sexes is not in any proper -sense a dwelling ‘together, but is confined to
the period of pairing, when the males seek the web in courtship and remain
sometimes hanging about the snare for several days.

Mr. Enoch® found on July 7th a male and female of Atypus piceus dwell-
ing together in the same tube, which was a large one. He had no doubt
that they had been thus living together since October of the pre-
ceding year. If this be so, Atypus presents one of the most
striking examples of conjugal domesticity and fidelity thus far
observed among aranead tribes. The tubes of the males were ‘generally

Agalena.

Atypus
piceus.

' Walckenaer, Apteres, Vol. II., page 390.

? Svenska Spindlar, ete., page 148. Aranei Svecici, Descriptionibus et Figuris, ete. Caroli
Clerck, Reg. Soc. Scient. Upsal Memb. Stockholmis, MDCCLVII. In Swedish and Latin. I
quote here and elsewhere from the Latin version.

*Memoire pour servir l’Histoire des Insectes, par M. le Baron Carles De Geer. Tome
Septidéme, Ouyrage Posthume, page 308. A Stockholm, MDCCLX XVIII.

* “Spiders of Dorset,” page xxxiii.

* Life History of Atypus piceus, Trans. Ento. Soc. London, 1885, page 402.

30 AMERICAN SPIDERS AND THETR SPINNINGWORK.

found near those of the females, and Mr. Enoch believes that they find
their way into the female’s quarters the same night that they emerge from
their own nests.

Darwin learned from Mr. Blackwall that he had not seen the males of
any spider fighting together for the possession of the female. He further
expresses the opinion that, judging from analogy, it is not prob-
able that such a habit exists, for the males are generally much
smaller than the females, sometimes to an extraordinary degree.
Had the males been in the habit of fighting together, Mr. Darwin argues,
they would, it is probable, have acquired greater size and strength.’ But
later observers have been more fortunate than Blackwall, and their obser-
vations reverse the judgment of Darwin. We now know that, as with many
other animals, the mating period of spiders is marked by frequent and
vigorous conflicts among the males, who are thus not only exposed to peril
from the voracity of their mates, but also from the jealousy of their rivals.
It might indeed seem, in view of the fact that a number of males may be
found at one time quietly hanging about the lady’s bower, that they are
not a very combative generation. Doubtless, the males of Sedentary species
do have, occasionally, to secure their marital rights by battle. But such
combats are probably far less frequent than among the Wanderers. In
point of fact, the conditions are such that it is not so easy for them to come
in personal contact with one another. When several Orbweayers attend
one female they prefer different parts of the web, and evefi when I have
seen them grouped tolerably near one another they showed no disposition
to quarrel.

With the Wandering tribes the conditions are different and such as to
compel personal contact with rivals, and thus it may be that a more com-
bative habit has grown up. This is well illustrated by the obser-

Fights of
Males.

ene vations of Professor and Mrs. Peckham upon the Saltigrades.
Quarrels, These naturalists have given a number of interesting and valu-

able notes upon the combats between males. The various atti-
tudes were both photographed and figured from Nature, and as a result
we have not only attractive descriptions but characteristic illustrations.
Some of these I have copied. The males of our little Zebra spider,
: Epiblemum scenicum, which may be seen in the early spring
Epible- skipping about on walls, fences, and outhouses, were found fight-
mum . ; : :
aca ing on a brick wall. They held up the first pair of legs and
oun. moved rapidly in front of each other, now advancing and now
retreating in a half circle, distant from each other about four
and a half inches. There was little real earnestness in the affair, and it

' Descent of Man, Vol. II., chapter ix., page 329, Amer, Hd.

WOOING AND MATING. 31

reminded one of the bluster of two boys each threatening and daring the
other, and neither willing to be the aggressor. In a few minutes, however,
they both wandered away.!
Several males of a species of Icius when placed.within boxes proved to
be very quarrelsome, and had frequent fights, but were never found to be
: injured. Indeed, after having watched hundreds of similar bat-
eee tles between the males of this and other species, Professor Peck-
Duels, am has reached the conclusion that they are sham affairs, gotten
up for the purpose of displaying before the females, who com-
monly stand by, interested spectators. This harmless nature of the conflicts
of spider duelists is in accordance with my own observations, and also in
accord with the few statements that have been made by other observers.
The males of Dendryphantes capitatus are very quarrelsome—sparring
whenever they meet, chasing each other about, and sometimes
clinching. The Peckhams put eight or ten males into a box,
and they fought; and, although it seemed cruel sport, it was
soon apparent that they were very prudent little fellows, and were fully
conscious that—

Dendry-
phantes.

“He who fights and runs away
Will live to fight another day.”

In fact, after two weeks of hard fighting, the observers were unable to
discover one wounded warrior. When approaching for combat the males
hold the first legs up in a vertical direc-
tion. Sometimes they drop the body upon
one side, as they jump about each other.
These movements are very quick, and they
are always ready for a passage at arms.?

Two males of Zygoballus bettini, while
executing a dance before a female, engaged
in a quarrel. They ran savage-
ly upon each other and fought Fic. 11. Position of two male Saltigrades,
twenty-two minutes, during one Zygoballus bettini, when fighting. (After
round, remaining clinched for six minutes, P°**’™
When fighting, the abdomen is held nearly at a right angle with the
cephalothorax. (Fig. 11.) The combatants appeared tired at the close of
the battle, but after a short rest were perfectly well and fought a number
of times subsequently. *

Several males and* females of Phileus militaris were placed together
in a box. Among the males was a large fellow, who proved to be a
universal bully. In the course of time another male, almost his size, was

Zygo-
ballus,

‘ Observations on Sexual Selection in Spiders of the Family Attidse. Oceas. Papers Nat.
Hist. Soc. Wis., Vol. T., 1889, page 89.
* Peckham, id., page 45. * Idem, page 48.

32 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

introduced, and he also adopted the role of a bully. After driving his
smaller companions about for a time, he was engaged in devouring a gnat,
when the original bully emerged from some leaves, got sight of
the neweomer, and at once approached, bristling with pride and
ire. His first legs were raised high as if to strike, his palps
vibrated with excitement, his abdomen dragged first on one side and then
on the other. Number Two was evidently of good courage, for he held
his ground and, not relinquishing the gnat, raised his legs and clinched
with his antagonist. The battle raged for five minutes, and resulted in
Number One robbing his antagonist of his dinner and chasing him igno-
miniously away. For several days following, life in the mating box was
robbed of its monotony by perpetual battles among the males. The
females, in eluding them, jumped and suspended themselves from threads.
On one occasion, the big bully who had now lost his mate, invaded the
home of the lesser bully while the owner was out seeking food. The first
time this happened the rightful proprietor, upon his return, ejected the
invader without ceremony. ‘The second time the two had a prolonged
struggle, clinching, and falling thus hooked
together a distance of about twelve inches,
the height of the box. (Fig. 12.) Some time
afterward the two males wandered about,
fighting whenever they met.!

These notes give a brief picture of the

general character of the observa-
ga _ tions made by Professor Peckham.
Py vohnios “i ae They indicate, first, that the males
after maturity, and during the

mating season especially, are addicted to frequent quarrels.

Second, their mode of combat consists in thrashing each other with their
fore legs, clinching with the mandibles, tugging and dragging each other
about, and generally tumbling and scratching one another with their claws.

Third, these conflicts, although they present the appearance of extreme
ferocity and deadly purpose, rarely, if ever, result fatally.

Fourth, the females are usually disinterested witnesses of the duels be-
tween their attendants, although, in point of fact, the victorious rival re-
ceives whatever favors she may have to bestow.

Fifth, the combativeness, or, at least, the actual combats of male spiders
are much more frequent between the Wanderers, who, by reason of their
errant mode of life, necessarily come into close contact with one another
during their rival courtships. ‘The habits of the Sedentary tribes, which
keep them stationary at fixed points of the snare, tend to hold the males
separate from one another, and thus conduce to peace.

Philzus
militaris.

1 Tdem, pages 52, 53.

> je cata tit ty es ints

WOOING AND MATING. 33

Combativeness among spiders is not limited to males; the females also
fight, and with great ferocity, not only with one another, but with the op-
posite sex. Professor and Mrs. Peckham haye contributed to our

coer knowledge of this trait as displayed by females among the At-
tidee, to which brilliant family their studies have been chiefly

iveness. ; : :
; directed. They found that the females are, with few exceptions,

larger, stronger, and much more pugnacious than the males. They placed
two females of Phidippus morsitans together in a glass jar. No sooner
did they observe each other than both prepared for battle. Eyeing one
another with a firm glance, they slowly approached, and in a moment were
locked in deadly combat. Within a few seconds the cephalothorax of one
was pierced by the fang of the other, and with a convulsive tremor it re-
laxed its hold and fell dead. In all, four females were placed together,
and in each instance the fight was short, but to the death. Subsequently,
the observers admitted a well developed male, which, though smaller, was
compactly built and apparently strong enough to bring the virago to terms;
but, to their surprise, he seemed alarmed and retreated, trying to avoid
her; she, however, followed him up, and finally killed him. They ob-
served the same habits in Phidippus rufus.
In Dendryphantes elegans the female is nearly a third larger than the
male. A number of this species, males and females, were kept together in
a large mating box, and their behavior demonstrated the greater
poe quarrelsomeness of the females; they would frequently go out of
Females, their way to chase one another, and they were much more cir-
cumspect in approaching each other than were the males. In
Icius mitratus neither sex was especially pugnacious, but the male was as
little so as the female. In Synageles picata the females never came near
each other without some display of hostility, though they did not actually
fight. In several species of Xysticus, as ferox and gulosus, the females
are savage and ready to attack anything that comes in their way, while
the males are smaller and more peaceable.

VII.

From these more general facts we may now pass to the detailed descrip-
tions of the act of conjugation in such species as have been studied. I have
never been fortunate enough to observe the actual pairing of Orbweavers,
my only opportunities of study having been with Linyphia marginata and
Agalena nevia. I am therefore dependent upon the observations of others
for the pairing habits of the Orbweavers.

Termeyer, nearly a century ago, thus correctly noted some points in the

* Observations on Sexual Selection in Spiders of the Family Attidse, by George W. and
Elizabeth G. Peckham, pages 10, 11.

34 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

courtship of Epeira diademata: The male approaches little by little with
much caution, doubtful of the reception which he is to meet in the web
of the female, who occupies the centre, intent only on her prey. He
= commences by touching with one leg a thread of her web. The
a enae female approaches him. He flies, allowing himself to hang by
demata, ® dropline. Soon he reascends, being assured in some way that
he will not be ill received. Then he approaches his mate, and
with one of the palps touches her abdomen quickly many times.*
The pairing of the Diadem spider is also described by Menge.” ‘The male
accomplishes his approach to the female by means of a strong thread fast-
ened immediately above her, which thread is, in fact, the strongest
Menge’s that he spins. This becomes his love bridge, over which he passes
Descrip- E : ; : : :
Hoa with trembling and uncertainty as to his reception, his feet ex-
pressing both invitation and fear. If his reception is friendly,
he passes under the body of the female, with his fore feet folded to allow
her expanded feet to encircle his, while their faces and partly their breasts
touch. He now quickly touches the vulva of the female with his palps,
and instantly drops to the ground by a thread ; however, he soon returns,
and the deposition of the semen is continued until finished. During Sep-
tember, one year, Menge observed a male thus approach a female about
twenty times. Finally he caught the ovipositor or hook of the vulva, de-
taining the same and turning so that the two abdomens and their adjoin-
ing parts touched, and the posterior parts of the same were no longer sep-
arated, but pressed closely together. The connection continued for over a
minute, when the male dropped backwards to the ground, and remained
there for some time as if dead.
Menge also observed the pairing of Epeira marmorea on a warm August
evening. The female left her web and advanced towards the male. The
movements of the latter were very careful, and when sufficiently

Epeira —_Glose he touched the vulva of the female in passing, and instantly
mar- : “ A : :
morea, Withdrew. As the female remained quiet and did not attack him,

the act was renewed the second and third time. The third time

the female retired to her dwelling, and the male dropped down by a thread.
The pairing of Tetragnatha extensa has been described by several writers.
Lister, the pioneer of English arachnology, says that May 25th at sunset
he saw the pairing of many spiders of this species. The two
sexes were suspended by means of a thread placed upon their
webs. The male was below, having his body stretched upon a
straight line. The body of the female, on the contrary, was
doubled, and her abdomen touched the fore part of the abdomen of the
male. He continually thrust a little horn, remarkable by its tubercle, upon

Tetragna-
tha ex-
tensa.

1 Translation of Prof. Wilder. Proceedings-Essex Institute, Vol. V., pages 71-3.
2 See Prussian Spiders, under Epeira diademata.

WOOING AND MATING. 35

the superior part of the abdomen of the female. The feet and mandibles
of the one were interlocked with those of the other.1

Walckenaer has given a complete and graphic description of the loves
of Tetragnatha. His observation was made on the 26th of May, when the
weather was serene and moderately warm. A male was stationed under a
quite large orbweb spun in an inclined position. The female was below,
suspended by the hindermost feet. Her body was bent double, her abdo-
men in a sense horizontal, so that her cephalothorax was bent
back upon the male in a vertical position. Her fore feet were
entangled in the fore feet of the male, but gently and without
stiffness. Her mandibles were opened, as were also those of the male, and
the extremities were supported one upon the other, and presented the form
of a trapeze, like the four open blades of two pairs of scissors if joined
at their points. The male had his body stretched upon the same line in
a horizontal position, but reversed; that is to say, the sternum of the
cephalothorax and the venter or lower part of the abdomen were turned
towards the sky, and the dorsum or back towards the ground. It resulted

‘ from this position that, from beneath, the male, although much
rite smaller than the female, appeared to surpass her in length by

‘half of his abdomen. Further it resulted that the vulva of the
female fell exactly beneath the palps of the male. He was suspended from
his snare by the fore feet, which were entangled in those of the female.
His two hindermost feet were posed upon the abdomen of the female, and
served to press her lightly against himself, while he applied the palps to
the vulva, The valve of the palpal bulb during the act of pairing was
swollen, brilliant, and the color of yellow amber.

This pairing lasted more than a quarter of an hour, and although the
observer came very near in order to see more distinctly, the mates did not
separate. Once he touched a part of the web and caused it to vibrate.
The partners recoiled, but still remained coupled. The female then made
some efforts to disengage herself, but the male prevented her. The ob-
server’s attention was diverted at this point, for the space of two minutes,
to make another observation. When he turned his eyes again upon the
pair, only the female remained ; she was at the centre of her snare in the
accustomed position, that is to say, with the body and feet stretched out.
The male had disappeared, and was searched for in vain; but during the
search Walckenaer observed another female engaged in spinning her snare
While another male waited upon an adjoining branch.

It will be seen from the above account, which describes the entire pro-
cess, that, making allowance for less’ skill in observation, the English ob-
server had correctly seen what the French naturalist so correctly reports.
Lister, however, represents the male as stretched below the female, while
Walckenaer reverses the attitude.

Position
of Female

‘ Lister, Historie Animalium Anglie (Araneorum Anglise), 1678, page 31.

io = " ss —— os a ee a ee

36 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Emerton, in his chapter on the growth of spiders,' presents some inter-
esting facts upon pairing. Two of these, the pairing of Linyphia and Ag-
alena, I am able to confirm, from my own observations, as sub-
stantially accurate. According to this author, the male of Argi-
ope, which is very small, stands on the upper edge of the web,
while the female occupies her usual position in the centre. After feeling
the web with his feet for some time, he runs down the centre so lightly
as not to disturb the female, and climbs over her body for some minutes
in an apparently aimless way. She takes no notice of him at first, but at
length, especially if he approach the under side of her abdomen, she turns
and snaps at him with her jaws. He is usually nimble enough to dodge
and drop out of the web. Not discouraged, however, he climbs up to the
top and begins over again. In these encounters the males are
often injured. They frequently lose their legs, and one fellow
was seen with only four left out of ‘eight, but still maintaining
his embrace. At length the male succeeds in getting under his
mate and inserts his palpi under her, into the epigynum. (Fig.
13.) In the meanwhile the female hangs in the web, while the
male holds by his legs to the under part of her abdomen. (See
Fig. 14.2)

Mr. Emerton thus describes the courtship of Epeira sclope-
taria. The approaches of the male were always seen in the

evening, just before dark, when the females usually
Courtship make or repair their webs, and the males are wander-
PE a ing about visiting. The male would climb carefully
Poae over the edge of the web, and, finding that the female

took no notice of him, would run suddenly towards
rie. 18, Mare the middle of the snare, on the side opposite to that occupied
Argiopecoph- hy the female. His head was towards her head, and, if she
Se ge permitted him to approach near enough, he would slap one of
male. (After his palps upon her epigynum. By this time the female would
Emerton.) i sos

strike viciously at her lover, and he would drop suddenly out
of the web by the usual dragline.*

September 24th, on the banks of the river Arduson, Baron Walckenaer
found a reed, the leaves of which were spun together into a nest by Epeira
apoclisa. He saw the male of this species mounting towards the nest.
Wishing to make observations at his leisure, he broke the reed a little be-
low the place where the male stood, and carried the whole to his chamber.
The male did not attempt to run away or quit the neighborhood of the
nest. Walckenaer placed the reed in a large box. The following morning
he saw a long thread stretched and the female issuing from her nest upon

Pairing of
Argiope.

1 Structure and Habits of Spiders, page 87. 2 Drawn by Mr. Emerton for this work.
* Letter to the author, October 17th, 1888.

WOOING AND

MATING,
NIRV
UI: ~ i \

AS
RUDY

>
————

——

Nf u

—————— ns
—— SS

—BZ

——

————

(|
\
mn

Hd)
(iil

fj WHEN

nd figures

in part just under the abdomen

ria embracing the female. Snare a

Argiope cophina
ize. The male is seen

life size

Fic, 14,

38 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the border of the box and in the act of pairing. But the male, forthwith
frightened, quitted his hold and ran rapidly away. The female, on the
ae contrary, remained motionless. The observer captured the two
=m A of and placed them in a covered glass bottle with the nest which
Spot the female had temporarily abandoned to meet her partner. He
also introduced into the bottle living flies. During three days the
mates ceased not to caress each other.. The female did not return to her
nest, but kept below in a reversed position. The male approached her
from the side, with head elevated, stretched out his feet and spread them
gently and slowly upon the back of the abdomen of the female, sometimes
touching the fore feet with his own by a slight and very quick movement.
Then the female leaned to one side, in such a way as to expose her yenter,
against which the male stretched his palps, and the union took place by
means of the palps. It was between 5 and 6 A. M. when this act occurred,
and it was repeated many times. During the remainder of the day the
couple remained separate. The female rested in the same place almost
wholly motionless, without doing anything. The male, more lively, more
wandering, more active, constructed a little web, placed himself in the mid-
dle and caught some flies. Again, he sometimes promenaded the glass,
stretching threads; but always after this momentary absence he returned
to a position in front of his spouse, appearing to contemplate her, laid his
feet against her own, his head vis-d-vis with her head, in a reversed po-
sition.

Finally the female constructed a tube of silk, within which she retired.
The male penetrated this; the tube was large enough to contain the two;
and they remained therein for ten days in perfect harmony, with-
out attempting to issue from their love bower. During all this
time the female continued to rest wholly tranquil, and did not
attempt to escape. It was not so, however, with the male, who frequently
wandered abroad into the bottle. The two did not attempt to make orbic-
ular webs, but stretched irregular lines, upon which they caught flies, a fact
which shows that spiders are able to adapt themselves and their industry
to various circumstances. On the twelfth day, October 4th, the female Apo-
clisa became invisible, and it was observed that she had returned to her
original nest, first built at the top of the reed. The tube was abandoned by
her and remained vacant. The male did not attempt to rejoin his com-
panion, but wandered about the glass, occupied in seeking an opportunity
to escape therefrom. At this point Baron Walckenaer was compelled to
interrupt his observation.

I make the following summary of the various stages as above described :
First, the female comes out to meet the male; second, she hangs below a
few stretched lines, with her back downward; third, the male touches her

A Love
Bower.

* Walckenaer, Aptires, Vol. IL., pages 63, 64.

el i i es | a ae” teed Oe a a a ee) =

WOOING AND MATING. 39

back and feet; fourth, the male palpal bulbs are applied many times to the
epigynum ; fifth, the embrace ceases, the female remains stationary in the
same place, the male wanders about, makes a straggling web and
catches flies; sixth, the male comes before the female, touches her
feet, and remains vis-d-vis; seventh, the female makes a tube and
enters it; eighth, the male penetrates the tube; ninth, the pair remain
for ten days domiciled within this bower, the female staying persistently
within, the male making frequent excursions; tenth, on the twelfth day
the female leaves the tubular bower and returns to her leafy nest; eley-
enth, the male makes no attempt to follow her, but wanders around the
bottle, seeking to escape. At this point observation ended, but there was
probably nothing further to observe.

Zilla callophylla is lacking in ferocity, and lives on good terms’ with

Summary
of Stages.

Fic. 15. Fic, 16. Fig. 17.

Argyroepeira hortorum. Fig, 15, male; Fig. 16, female; Fig. 17, the male palps,
much enlarged. (After Emerton.)

her mate. September 30th a male and female of this species were taken
by Walckenaer, and placed in a glass bottle. The male, after
the ordinary preliminary caresses, such as touching with the ex-
tended feet, stretched some threads in the manner of those which
served the female to descend from the top of the bottle to the bottom, where
he was. Then, by the movement of her feet, she excited her spouse to
approach her. Every part of the male’s body trembled in a sensible man-
ner. He advanced towards his mate, not without appearance of fear, since
she received him with open mandibles. Three times he essayed, always
while advancing, to introduce the genital organ of one of his palps into
the vulva of the female, and he succeeded at the fourth attempt with the
digital bulb of his left palp. Then was manifest in the male, as in the
female, a convulsive trembling of all the limbs and of all parts of the
body, which evidently announced that the union was accomplished. Four

Zilla cal-
lophylla.

40 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

other movements of the same nature, separated only by very short. inter-
vals, followed the first. After these five acts of union the male retired to
a short distance.

About half a minute afterwards he approached as on the first occasion,
introduced the bulb of his right palp into the epigynum of the female;
then, after the fifth movement of convulsive trembling, he again retired.
He continued in this manner during the space of twenty minutes. In
these twenty minutes he made thirteen embraces, or thirteen introductions
of the digital bulb of one of his palps into the vulva of his partner. After
these acts the male retired and went away. The female rested for about
a half hour in the same position, as if she awaited the return of the male,
who did not come back. Then she decided to remount to the top of the
bottle. !

1 Walckenaer, Apt., Vol. II., pages 71, 72.

OR APE LT.

COURTSHIP AND PAIRING OF THE TRIBES: LOVE DANCES
OF SALTIGRADES.

Havine thus considered the methods of pairing which prevail among
Orbweavers, we may note some of the modes which obtain among repre-
sentatives of other aranead tribes,

On the afternoon of June 14th? I witnessed the pairing of a male and

female of Linyphia marginata. The spiders were first observed at a

quarter before four o’clock. They were hanging inverted in the

Line- dome shaped nest of the species, in line with each other and

weavers: about three-quarters of an inch apart. Each hung within a

ae smaller dome which was formed by the outspread feet drawing

nan down the inner surface of the snare. The nest was hung from

the under surface of a plank that jutted over from a pile of

lumber, and was about two and a half feet from the ground; so that,
seated before the nest, my face was on a level with the spiders.

The male cautiously extended one foot towards the female, and pulled
upon the intervening threads. I turned a moment to adjust the block
on which I sat, and, on looking again, the two were in embrace. The

_ female was suspended as before, although turned at right angles to her
first position. The male’s head was laid against the sternum of the
female, his abdomen inclined a little upward, the fore legs interlocked
with, or, rather, interlaid upon those of the female. Both spiders hung
by threads, in the normal way. (Fig. 18.2) This was nine minutes be-
fore 4 P. M.

After a moment’s embrace the pair separated; the female made a cir-
cuit of the lower part of the dome, moving in an excited, jerking manner,
then returned to the summit. The male approached, the female stretching
out her fore legs somewhat as he laid his fore legs within them, which po-
sition was maintained, as was the relative position of the two, during the
entire period of union. The female, during the act, remained perfectly
motionless, except an occasional twitching of the apex of the abdomen.

The two terminal bulbs upon the male palps were laid upon the epigy-
num of the female, and pressed downward. From one of these issued the

1 The same species was again seen pairing in August, in Connecticut.
* Drawn by Mr. J. H. Emerton for this work.
(41)

42 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

palpal sac, a bean shaped organ, of a bright amber color, and translucent,
which shone brilliantly in the sun that fell full upon it from the west. It
remained thus projected for a brief space, held between the finger like tufted
horn of the palpal bulb, and was then gradually contracted and withdrawn
within the black corneous bulb, which was meanwhile pressed eagerly

against the vulva. A small elbow or projection upon the upper
Useof part of the bulb. seemed to press within the spermatheca. The
i Ss two bulbs were laid simultaneously upon the epigynum, but the

ulb. : i ;

inflated palpal sac appeared in but one bulb at a time, alter-

nately. There was a prolonged squeezing motion of the bulbs, as though

Yai vane

PIII

= SS

thi aes ea

————
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PN I Mi :
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Heian av? Hi i. AUTH TG i \ a et Ml ih fia)
Ie i Dek Mi. . ch. he 4 a Wd A UNG Wty) ni i
oe nee Dak we Biii beth ue Bi })
i)

===>
———

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—

=—
SSS

———

s AMIN NN ayy; /
HON oe a ie fe ae
iv . i ( \ mn Maa ini CaN? Ny Hn au AN Ihe WW
ANH I ail HAN (ri il NZ

Ki I We ail HN

Fic. 18. Pairing of Linyphia marginata. The figures much enlarged. The little dome caused by
the pulling down of the feet is represented, and a part of the snare proper.

pressing into the spermatheca, and at times a corresponding motion in
the abdomen of the female, especially at the apex. With this exception
the female remained motionless during the whole period. After applica-
a tion as above the palpal bulb was slowly, for the most part,
Biting the 4 sometimes rapid] d by th le, bent d, and
Bulbs. pidly raised by the male, ben upward, and ap-
parently clasped upon the falces or lower margin of his face,
which parts of course were upward. Three or four movements back and
forth in this clinched position followed, when the series of motions above
described “was repeated.

PAIRING OF SPIDERS. 43

In the meanwhile the second bulb remained upon the other tube until
the first bulb began to descend, when it in turn was elevated and the
same motion made, As the bulb descended, its sac began to inflate and
issue. ‘This process was quite regularly repeated. Sometimes, however,
both bulbs were clinched upon the falces at the same time; sometimes the
moyements of the bulb were more rapid than at others. The bulbs had
the appearance of having been moistened by some secretion, presenting
the peculiar gloss which a colorless liquid gives to a black surface, but I
could see no secretion otherwise, although I was able at any time to use
my pocket lens with the exercise of a little care.1

At twenty minutes before six o’clock I was compelled to leave, at which
time the pair had been in embrace one hour and forty-nine minutes. At
six o’clock twenty-eight minutes I returned, and found the pair in
precisely the same positions. I remained five minutes, and then
left an intelligent young man at the post, with full instructions as
to points of observation. He reported that at thirteen and a half minutes
past seven, afternoon, the pair parted suddenly. The male ran down to the
lower margin of the dome, pursued by the female, who stopped suddenly
just above, and turned back to the central point in the summit. Shortly
after receiving this report I visited the web, and found the female sus-
pended motionless in this position, and the male at the point to which he
had fled, feeding upon a small fly. The next morning at seven o’clock the
female was in the same position, and the male had disappeared. I attempted
to capture the female, but she ran among the boards and escaped. The pair
had thus been in union two hours and fifty-five and a half minutes.

During this period they were separated a number of times. Nineteen
of these interruptions were noted; one was caused by a small fly striking

the snare, at which the male darted in a fierce manner, but
Interrup- failed to seize, as the fly broke loose before he reached it. Others
See nd ere caused by the observer touching the foundation threads or
tions other parts of the web. Toward the close of my observations I

accidentally broke the suspending lines nearest me, and caused
one side of the dome to fall in. This made only a momentary interrup-
tion. Many of these separations were, however, apparently without any
extraneous cause.

Twice the male ran to one side of the domed snare, made a web at-
tachment to a bit of leaf hanging therein, drew out a thread about two
and a half inches long, which he overlaid a couple of times, and then
made the following motion: First, the body was placed erect, that is, back
Epyans; and was moved back and forth along the line, rubbing the points
or “nippers” of the palps at the same time; then the spider swung over

Period
of Union.

1T did not at the time suspect that the palpal bulb oitaht have been applied to the ab-
dominal organ of the male, and did not look for this act. But subsequently I have imagined
that such might have been the case.

ae _————- == ve -_ % — iy ee ee ee

44 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

until the body made an angle of about forty-five degrees with the line,
and while holding on thus the palps were rubbed back and forth alter-
nately along the line as before. The process was repeated during another
of the intermissions, as described above. It was conjectured that the pur-
pose of this movement might have been, the distribution of the seminal
fluid into the palpal bulbs. It has been supposed that this is taken up by
the saes, by the inflation and contraction of whose membraneous coats it is
forced into the spermathecse of the female.
Mr. Emerton! observed the pairing of the male and female of Steatoda
borealis in April, and again in May. The female was in a scant web under
a fence cap. The pair stood head to head, as far apart as pos-
PA sible. The left palpus was kept in an hour and a quarter after
the couple were first seen. The male contracted his body sud-
denly, and swelled up the base of the palpal organ once every two or three
seconds. Two days afterwards Emerton saw the right palpus used by the
same pair for an hour. The adult males and females of this species occur
at all seasons, differing in this respect from many others.

TUL

Among the Tubeweavers I have observed the pairing of our common
Speckled Agalena. The male cautiously approaches over the broad sheeted

Fig. 19. Agalena nevia pairing. Fic. 20. Agalena nevia applying Fig. 21. Agalena applying the
Front view. (After Emerton.) the right palp in pairing. left palp in pairing.

web which forms a sort of front yard or plaza before the tube in which
the female waits. He is usually larger than the female, and is, therefore,
better able to compel a respectful reception. In the act of union
eae he takes his partner in his mandibles, turns her upon one side,
Agalena, 2 Which position she lies perfectly motionless, and with her

legs somewhat doubled together, as in the attitude of feigning
death. (Fig. 19) The male rests upon the side of the female, in a posi-
tion nearly at right angles with her prostrate body, and, while holding
her still with his fore feet, applies the palps alternately to the vulva.

(See Figs. 20, 21.)

' New England Theridide, Trans. Acad. Conn., 1882, page 19.

PAIRING OF SPIDERS. 45

According to Walckenaer, the union of the male and female of the

European Agalena labyrinthea takes place in the tube which serves as the

dwelling place for the female. In France this act occurs about

ae the middle of July. The female turns herself upon her side, al-

rinthea, ™ost upon the broad of her back. The male places himself

upon her in such a position as to hide from the observer his

head and cephalothorax.!_ It will thus be seen that the method entirely

corresponds with that of our own Agalena neevia, which this familiar Eu-
ropean spider so closely resembles.

The male of Clubiona constructs a web for union with the female, and
prepares, as one may say, the marriage couch, to which he admits the
female when the propitious moment has come.’

The interesting habits of Argyroneta aquatica, the well known Water
spider of Europe, were first fully made known by De Lignac, a priest of

the Oratoire, Paris, A. D. 1748. He not only observed the man-
so ner of making the nest beneath the water (Fig. 22), which has
Spider, been frequently confirmed since, but also the act of pairing.

When the male wishes to pair, says De Lignac, he constructs
near the nest of the female, and by the same means, a nest resembling
that of his spouse; but the nest is somewhat smaller. When the male has
completed the construction of his domicile, he makes a long canal, which
joins his cell to that of his spouse. He then cuts through the wall of the
latter, and introduces his body into the strange apartment. This vehicle
of communication being made, he strengthens it on the roof and _ sides.
He plasters this, as he does the rest of his nest, with silk, white and im-

permeable, and thus extends this corridor until it may be as
ieee large as the two apartments. Sometimes one sees, but only oc-
Corridor, ©asionally, as many as three lodges, which communicate with each

other. As these cells have been thus easily united, they also
sometimes separate, as, for example, when they are too lightly united, or
by the movements of the spiders when they engage in combats, for it ap-
pears that during the time of amour they are somewhat irascible. Often-
times one sees a strange spider making an effort to enter into one of these
lodges; but the inmate, who keeps its feet outside, guards, as a watchful
sentinel, the safety of its domicile, and drives the intruder from the door.

Baron Walckenaer confirmed these observations of De Lignac and added

some interesting details. On the 27th of July he placed together

Walek- in glass vessels males and females of Argyroneta, On the fol-
enaer’s : . ;
Obwsrvn: lowing day he saw in one of the silken bells woven by them a

tions. male caressing the female with his feet, and carrying his palps
to her abdomen. The two spiders were then upon the same line,

1 Aptéres, Vol. II., page 22. 2 Walck., I. Apt., page 143.
° L’Histoire des Araignées Aquatiques, page 43. De Geer in Holland as early as 1736 had
observed the curious industry of the Water spiders.

46 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

and stood face to face. The male carried his head under the body of his
mate in a reversed position. He stepped aside, and the female with her
feet tickled the apex of his abdomen.

The next day at 6 A. M. he saw a little web constructed by one of his
Argyronetas. He gradually filled the bottle with fresh water, whereupon
the couple began to work with extraordinary activity, and in less than an
hour’s time had formed a cell which looked like a bubble of air and had
the form of a subterranean vault. The male and female kept together.
As soon as the cell was finished, and on the same morning, the female
made a web at the surface of
the plant which had been in-
troduced into the
bell glass. Much
to the baron’s sur-
prise, she immediately de-
posited her eggs and envel-
oped them in a silken co-
coon. ‘The cocoon was placed
near the surface of the water,
and upon the very walls of
the vessel. The eggs, which
were of a beautiful orange
yellow, could be seen through
the fine, white tissue of the
cocoon.

July 29th, at six o’clock
morning, Walckenaer saw the
female near her eggs; then
she ascended to the surface
and dived. The male joined
himself to his companion.
The two spiders gently rub-
Fic. 22. The subaqueous nest of the Water spider, Argyroneta bed the extremities of their

aquatica, within which the cocoon is woven. anterior feet one against the
other, having the air of ca-
ressing. Soon this movement of the feet became more brusque and ap-
peared menacing. The male, struck by the feet of the female, suddenly
leaped aside, but the pair presently sought each other anew.
They interlaced their legs, the one within another, and gradually
approached nearer and nearer, head against head, The man-
dibles were opened; they flung themselves one upon’ another ; afterwards
recoiled, separated instantly, and sprang aside as if they had suddenly been
seized with fear. Thereupon the female returned to her position near her
eggs,

| ==>] The

= Fiy

Cocoon.

Caress-
ing.

PAIRING OF SPIDERS. 47

The next day Walckenaer renewed the water in the vessels, and saw the
couple approach one another, lightly touch their feet, swim without stretch-
ing out any thread and without touching the insects which had
been placed in the water for them, but which were all dead. At
five o’clock in the evening again the observer saw the male and
female upon the cocoon, drawn near together, the feet interlaced and mo-
tionless. On opening the bottle they separated. He was then astonished
to observe that the web that had surrounded the cocoon had disappeared.
Had it been employed to strengthen the cocoon?

The cocoon was a silken flask, attached to a plant by a short pedicle.
It was in part immersed within the water. It was rounded, flat-
tened, about three lines in diameter, was formed of a fine thread
of a very compact tissue, thin as an onion peel, and difficult to
tear. It contained forty eggs, not agglutinated, globular, of a pale yel-
low color.

On the first of April Walckenaer again observed in the jar where the
spiders were confined a little bubble of air and a web larger than the
former had been. After five days’ absence, April 6th he observed that
the spiders had detached the cocoon, in order to sink it to the bottom of
the bottle. The water was changed in the vessel and immediately they
swam about with delight, refreshed themselves, reunited near the cocoon,
and caressed each other with their feet. On the 7th of April he decanted
the water of the jar into a cistern. The Argyronetas, troubled by the
sudden movement of the flood, swam with great rapidity, and the female
having recovered her cocoon in the midst of the water, seized it, embraced
it with her feet and sought to buoy it up.

One of the most interesting and satisfactory accounts of the act of pair-
ing among 'Tubeweavers is given by Mr. Campbell from observations on

* . egenaria guyonii.1 The male was placed in a bottle contain-
steal ing a female which had been mature for a fortnight. He was
left within the vessel in which he had been lodged, but the

cover was removed therefrom. Notwithstanding the glass wall which sep-
arated him from the female, he soon became conscious of her presence,
and issuing from his own quarters approached her. The following morn-
ing he was standing with the first pair of legs over the female, and
his maxille resting on her abdomen, while she was crouching motion-
less, with her head in an opposite direction. Both were in

aah ae the same position the next morning, August 7th, 7 A. M. At
Male. 10 A. M. the male became restless, and wandered about the
bottle with spinnerets extended, returning every now and then to

place his palps upon the female. After each action he jerked his abdomen

Hmbrac-
ing.

The
Cocoon.

1On the Pairing of Tegenaria guyonii Guer., with a Description of certain Organs in the
Abdominal Sexual Region of the Male. By F. Maule Campbell, F. L. 8. Linn. Soe. Jour.
Zool., Vol. XVI., page 163.

48 AMERICAN SPIDERS AND THETR SPINNINGWORK.

upwards and downwards, a movement which often may be observed in
males, and which gives an idea of an expression of impatience.

The next observation, after a few minutes’ absence, showed the male
about two inches behind the female, standing as it were on tiptoe. His
palps were placed alternately and nervously to his maxille. On their re-
moval the whole body was raised still higher, and the abdomen
brought nearly to a right angle with the cephalothorax, with con-
siderable muscular effort in the basal portion, and with violent
tremulations. The movements, which were repeated four times, had the
effect of throwing the spider slightly forward, while the palps were
shaken in that peculiar manner which denotes great muscular tension in
some other part than that in visible motion. The palps were now gener-
ally alternately placed under the sternum and moved backward and forward,
upward and downward, with a scooping motion. In five minutes these move-
ments of the abdomen and palps were repeated ten times in regular succes-
sion, only varied by an occasional transfer of the digital organs to the
mouth,

Mr. Campbell observed thirteen couples pairing in confinement from the
middle of July to the end of August; and the following account may be
taken as typical of the species, with the exception that the union
does not necessarily occur so quickly after the female has gained
maturity. On the 13th of August he placed together a male and
female. On the 17th the latter cast her last skin. Up to that time, 6 A. M.,
they had taken no notice of one another. At 9.45 P. M. the two were so
close together that the femora of the first pair of legs of each were
almost in contact. After a few convulsive twitches of the legs the male
pressed forward, moving his palps up and down, when, as they touched the
palps of the female, the pair played with these organs like two friendly
bees with their antenne. After a few minutes the female raised herself,
leaning a little on her left side, and the male crept forward until his head
was under the sternum of his mate, while his first pair of legs were rest-
ing upon hers. He then advanced his right palp, leaning a little to the
left, and using the left palp as part of his support.

The male now rapidly raised his palps up and down for four or more
seconds, and with such energy as to compel the female to assume a verti-
cal position, He then retired, and again approached her, repeat-
ing the movements a greater or less number of times, occasion-
ally pausing before he withdrew his palps with a slight twist
inwards. At times he would leave the female for five minutes, and strut
with straightened legs around the vase, wagging his abdomen. Now and
then he would remain perfectly still with the palp withdrawn, or play with
the palps of the female, who seemed in a comatose state. He would then re-
new the union with undiminished vigor, appearing on each occasion less
desirous of changing his position.

Use o
Palps.

Ap-
proaches,

Use of
Palps.

ee ee

PLATE |

COLORS OF EPEIRA TRIFOLIUM.
1—9, VARIATIONS IN COLOR OF FEMALES, 10-11, THE MALE.

PAIRING OF SPIDERS. 49

The observer left them at 12.30 A. M. and returned at 7 A. M. The
male was still using his right palp. He saw no application of the left
palp, but had no doubt that it was employed during the night,
as in other cases. He had never observed the pairing interrupted
for a fresh collection of semen, although there is no reason to
think that this may not occur, The duration of the pairing is long, but
he was inclined to think it is more dependent on the difficulty in inserting
the embolos than on sexual endurance.

Impreg-
nation.

III.

The pairing of Xysticus trivittata Keyserling has been briefly described
by Mr. Emerton, and figured.1_ The spiders were seen on the 5th of June
among the short grass in an open pasture
in New England. The female held: herself

head downward on a blade of °
Pairing Of srass, with the abdomen turned
Tati away only enough for the male
grades. pia :

to reach under it with his palps.
There did not appear to have been any web
on the grass, though there may have been
a few threads for the female to hold by.

Among Lycosids we have the descrip-
tion given by one of the earliest natural-
ists, Clerck, the Swedish observer.2 -He
saw the pairing of Lycosa sac-
cata about the middle of June,
upon a rock exposed to the sun.
The two sexes approached by jumps, which
became fewer and slower as they drew near.
The male ended these preliminary stages
of courtship by suddenly leaping upon the
female. He then passed one of his palps un-
der her abdomen, and, holding and inclin-
ing her body with the other, inserted first
one and then the other palp. When the NY
pairing was ended, the two sexes separated Fie. 23. The pairing of the Laterigrade

species, Xysticus trivittata. (From Nature.)
and promptly ran away from one another.®

Emerton * says of the same family that the male leaps upon the back
of the female, and is carried about by her. He reaches down at the side
of her abdomen and inserts his palps into the epigynum underneath. The

Pairing of
Lycosids.

‘ Psyche, Vol. V., 1889, page 169.
* Clerck, Aran, Svee., pages 91, 92, pl. 4, Tab. 5, Figs. 1, 2, male.
* Walck., Apt., I, page 328, * Habits and Structure, page 95.

Te ee ee a ee re - 4

50 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

accuracy of the early observation made by Clerck is thus abundantly con-
firmed, The attitude of Lycosa is represented in Fig. 24, which has been
drawn from Nature for this work by Mr. Emerton.
Among the Attide, De Geer has described the pairing of Epiblemum
scenicum (Attus scenicus), which occurred upon a wall. The male mounted
uke upon the body of the female, passing over her head towards her
overs of .bdomen, under which he advanced one of his palps. He gently
* raised the abdomen by upward pressure of his legs, and then ap-
plied the extremity of his palp to the vulva. An instant afterward the two
spiders separated and removed a little distance from one another. The male
did not wait long before again approaching, and he repeated many times the
action above described. The female did not offer the slightest opposition,
but, on the contrary, seemed to greatly enjoy the act."

IV.

The mating of the Attoids, as told in the delightful pages of Mr. and
Mrs. Peckham’s Observations on Sexual Selection in Spiders, presents one
of the most important chapters in
the life history of araneads. It
is a strange and interesting story,
a romance of natural history as
fascinating as any love story of
modern fiction. These accom-
plished arachnologists, who have
Fic. 24. Male of Lycosa saccata embracing the female. carried on all their studies to-

From Nature. (Drawn by Emerton.) : 5
gether, have given special atten-
tion to the Saltigrades, and they were led into the study of the courtship
of these interesting creatures by a desire to solve some of the current
problems in natural and sexual selection. Independent of this, the facts
recorded are extremely valuable.

The first group of observations uncovered the habit of the males to
exhibit themselves before the females in a series of varied movements,

which may be generally characterized as dancing. ‘The purpose
Love of this appears to be, beyond doubt, to attract the attention of
Dances of :
the Males. the female, and render her complaisant to the addresses of her

lover. The courtship of Saitis pulex was thus conducted: The
male, when placed in a box with a mature female, at once observed her,
although she was twelve inches away. At the distance of four inches he
stood still; and then began the most remarkable performances that an
amorous male could offer to an admiring female. She eyed him eagerly,
changing her position from time to time, so that he might always be in
view. He, in the meantime, extended the fore legs upon one side of the

1 De. Geer, L’Hist. des Insectes, page 90.

ee ee ee ee ee
: =)

LOVE DANCES OF SALTIGRADES. 51

body in such wise as to elevate that side and correspondingly to depress the
other. The legs and palpus of the lower side were folded under, and upon
these the spider sidled along, moving in a semicircle for about two inches.
He then instantly reversed the position of the legs, and circled in the
opposite direction, gradually approaching nearer and nearer to the female
in the course of these oscillations.

The female dashed toward him, while he, raising his first pair of legs,
extended them upward and forward, as if to hold her off, but withal slowly
retreated. Again he began his oscillating movements until one
hundred and eleven circles had been counted. The female in
the meanwhile gazed toward him, apparently in a softer mood,
evidently admiring the grace of his antics. When he had approached
almost within reach of her, he whirled madly around and around her,
she joining and whirling with him in a giddy maze. He then fell back,
and resumed his semicircular motions, with his body tilted over. She, all
excitement, lowered her head and raised her body, so that it was almost
vertical. ‘lhe two then drew nearer. ‘The female moved slowly under
the male, he crawling over her head, and the mating was accomplished. !

A male of Synagales picata executes his love dance with all his feet
on the ground. He raises himself on the tips of the six hindermost legs,
but slightly inclines his head downward by bending his front

Saitis
pulex.

Love legs, their convex surface being always turned forward. His ab-
Dance of cole F ate :
Benes domen is lifted vertically, so that it is at a right angle to the

gales. plane of the cephalothorax. In this position he sways from side
to side. After a moment he lowers the abdomen, runs a few
steps nearer the female, and then tips his body and begins to sway again.
Now he turns in one direction, now in another, pausing every few moments
to rock from side to side, and to bend his brilliant legs so that she may
look full at them. He could not have chosen a better position than the
one he took to make a display, and the observers were impressed by the
fact that the attitude taken by the males served perfectly to show off their
fine points to the female.”
Marptusa familiaris is an Attus of sombre gray and black colors, that
may be frequently found on trees, fences, and like positions in the neigh-
borhood of Philadelphia. It is apparently a widely distributed

Mita species. When the two sexes were placed together, the female
Ae “*P" saw the male as he entered at the opposite side of the box, thir-

familiaris. teen inches away. Eyeing him attentively, she slowly changed
her position to keep him in sight, and kept her palps moving
rapidly, a characteristic action of the species. As the male neared her,

‘ Observations on Sexual Selection in Spiders’ of the Family Attide. By George W. and
Elizabeth G, Peckham. Occasional Papers of the Natural History Society of Wisconsin,
Vol. I, 1889.

* Idem, page 43.

a ae a. as

52 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

he stretched the first and second pairs of legs sidewise, but after a moment
backed away. These manceuvres were repeated many times, the attitude
assumed during them being as represented by Fig. 25. Occasionally he
would bend the tip of the abdomen
down, lifting the body up on the
last joints of the two hindermost legs.
The female always paid the greatest
attention to his movements, lying on
= the ground with all the legs flattened
Fic. 25. Positions in courtship of Marptusa familiaris, and the palpi slightly raised, the

Male on the right hand. (After Peckham.) only movement visible being the vi-
bration of the palps. There is a certain slowness and dignity about the
wooing of this species, almost ludicrous. !

The males of Dendryphantes capitatus approach the female rapidly,
until within two to five inches, when they stop and extend the legs di-
rectly forward close to the ground, the legs being slightly curved, with the

_ tips turned up. This position serves admirably to
Showing expose the whole of the bronze and white face
ha es (Fig. 26) to the attentive female, who watches him

closely from a little distance. As the wooer grows

more excited, he lies down on one side, with his legs still %
extended. These antics are repeated for a very long time, fie, 26. Bronze
even for hours, before the female accepts his addresses. tee g
yphan

The male of Habrocestum splendens is a magnificent fel- tes. Male. (At
low, having an abdomen of glowing pink, and bronze ceph- ‘* Peexhem?
alothorax tinted with reddish brown. He began his addresses by advanc-

ing a few inches towards the female and then backing off again.
Habro- ‘This movement was repeated many times. After awhile he set-
cestum 3 é me
oleridate tled down under a little web in a corner. The female, troubled

by this indifferent treatment, advanced toward him, whereupon
he came out and she fell back. This play was kept up for some time,
and at length the male began his courting in
earnest. When within a few inches of her, he
commenced a rapid dance from side to side,
raising the whole body high on the tips of the
legs, the first pair being directed forward, and
the palps clasped together, with the abdomen
Fic. 27. Male Saltigrade, Habroces. turned to one side and lifted up. (Fig. 27.)
tum splendens, when approaching After a short dance he stood motionless,
female. (After Peckham.) Aap 5 ;.

striking an attitude as shown in the figure, re-

maining quiet for half a minute. Then he turned his back on the female,
moving irregularly about with his legs forward and his palps vibrating.

1 Tdem, page 44.

LOVE DANCES OF SALTIGRADES. 6

Again he danced sidewise before her, strutting and showing off like a
peacock, whirling around and around. Professor Peckham at first supposed
that this turning around was accidental, but it hap-
pened so regularly at a certain stage of
oe courtship, that he concluded that it was an
rule important part of his display, serving the
3 better to show his brilliant abdomen.

In approaching the female the males of Phileus
militaris were very eager and fairly quiy- =
ered with excitement. The first two legs

‘aiged he head d ed d Fic, 28. Position of male
were raised over the head and curved toward — phijeus militaris when ap-
each other, so that the tips nearly met, and the palps Proaching the female. (Af-

q 2 ter Peckham.)
were moved up and down. (Fig. 28.)

Astia vittata is peculiar in the fact that it has two well marked male
forms, which shade into each other, but maintain at least one characteristic
distinction, namely, three tufts of hair which mark
the black form, niger. Mrs. Peckham was kind
enough to send me a box in which were packed

: a number of specimens of the female
Dervish Bec . :

and both varieties of the male, in order
Dance of : :
Rata. that I might witness these remarkable
courtship dances. This was prior to the
receipt of the work from which I have been quot-
ing, and I had but a hint of what I might expect,
and how best to proceed. Moreover, my specimens
unfortunately, arrived in a bad condition. All
were dead except one female and two males, and
Fic. 29, Male Astia vittataindanc- the latter were much dilapidated, one of them par-
a eae before female. (From ticularly being apparently in a dying condition. I
ature, . . .
succeeded, however, in resuscitating both males by
doses of water and good nursing. One of them
in a short time seemed quite well.

I placed the three together in a box, and had
the privilege of observing, in some degree, what
the Peckhams have so fully described. The most
lively male at once began animated movements,
which were evidently induced by the presence of © —
the female, who, however, ran away and kept cir- = ~ a
cling around the box, running over the walls and ig. 90. The male of Astia vittate
climbing upon the glass cover without showing im the act of vaulting during a
¢ qs ore love dance. (From Nature.)
any disposition to respond to the advances made.

The male threw himself into what may be described as a rampant position

Phileeus
militaris.

1 Idem, page 49, 2 Tdem, page 51,

54 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

(Fig. 29), that is, the body was thrown into a position at about an angle of
forty-five degrees, the abdomen almost in the line with the cephalothorax,
but a little bent under at the apex, which nearly or
quite touched the ground. The two hind pairs of
legs were thrown outward from the body, the fourth
or hindermost pair being well curved, the third or
next pair somewhat bent, but more extend-
oo ed. The second pair (next to the foremost)
ih fete My confirmed Was quite extended in a line without much
noticia = curvature, except towards the last two joints.
ee) hee froak legs were extended in a line somewhat
Fic. 31. Love dance of Astia
vittata. Male with front curved, and well thrown upward above the head, and
es poise. (After Peck- the palps, which are black, were stretched out in a
corresponding position, and continually rubbed one
upon the other in an excited manner. In this attitude the male moved
backward and forward with a lively sal-
tigrade movement, whirling around a little
as he leaped upward (Fig. 30) and brush-
ing the tips of his palps together in the
meanwhile. This was about all I was
permitted to see, but it at least confirms
in part what the Peckhams have so care-
fully recorded.
The first male form, which corresponds
in appearance to the female, when he ap-
proaches his mate, raises his first
mutt of legs so that they point either
tie, forward or upward, keeping his
palps stiffly outstretched, while
the tip of his abdomen is bent to the Nes
ground, This position he commonly takes fic. 32. Position of male Astia vittata when
when three or four inches away. While ‘%PProsching the female. Much enlarged.
: (After Peckham.)
he retains this attitude he keeps curving
and waving his legs in a very curious manner. Frequently he raises only
, one of the legs of the first pair, running all the
time from side to side. As he draws nearer to
_ the female, he ee his at to the ground and,

pairs so that the Hips Menbhe in front (Fig. 32),
ee we eee ae mitratus ‘0€ proximal joints being turned almost at an

dancing before female. (After angle to the body. Now he glides in a semicir-
Peckham.)

times receding, until at last she accepts his addresses.
The Niger form is much the more lively of the two, and whenever the

cle before the female, sometimes advancing, some-'

. 5 ee.

LOVE DANCES OF SALTIGRADES. 55

two varieties were seen to compete for the female, the black male was suc-
cessful. He is bolder in his manners, and was never seen to assume the
prone position as did the red form when close to the female. He always
held one or both of the first legs high in the air (Fig. 31), waving them
wildly to and fro; or, when the female became excited, he stood perfectly
motionless before her, sometimes for a whole minute, ateming to fascinate
her by the power of his glance.

The male of Icius mitratus is quite different from the female, especially
in his slender tapering body and long first legs. The female is remark-
able for her indifference, and takes less interest in the male’s display of
his personal charms than any spider observed. In courting and fighting,
the position of the male is the same; the body is somewhat raised; the
first legs are held at a right angle to the cephalothorax ; the abdomen is
twisted to one side, and, as he dances before his lady love, is changed now
to the right, now to the left.’ (Fig. 33.)

It is interesting to find that these amorous displays on the part of
males have recently been observed in other invertebrates. Mr. 'T. H. Mor-

gan thus describes the performance of a male crab (Platyonychus
Love ocellatus) in paying his courtship to the lady crab. The specimens
rani were confined together in an aquarium. While sketching some
Grals: hermit crabs which had previously been placed in the same tank,

the observer was attracted by the movements of the male Platy-
onychus. Without apparent cause he was seen to rise upon the third and
fourth pairs of legs; his large chelee were thrown above his head, with the
claws open and their points touching in the middle line; his fifth pair of
feet were held horizontally behind, and his body perpendicular to the floor
of the aquarium, or at right angles to the normal position.

The posture was ludicrous, and when he began slowly to gyrate, hjs
movements and attitude were the cause of much merriment upon the part
of the spectators. At times he balanced on two legs of one side, again on
two legs of opposite sides. Now he advanced slowly and majestically, and
now he wheeled in circles in the sand on the floor of the aquarium, and
now for a few moments he stood as if transfixed in this unnatural posi-
tion. An electric light hung above and to one side of the water, which
suggested the possibility that it might be the exciting cause. It was turned
out, and still the dancing went on. At last, from sheer exhaustion, Mon-
sieur Crab sank down to the sand in his usual attitude.

But now the female, who had all this time remained tucked away in
the sand, came forth and began to move about the aquarium; soon she
came near to the male crab, who instantly rose to his feet and began to
dance. Again and again the performance was repeated, and each time the
approach of the female was the signal for the male to rear upon his hind
feet, and reel about the aquarium as if intoxicated.

1 Tdem, page 50,

— OO ae ee me “. a

56 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

At times, when the female approached him as he danced, he was seen
to make attempts to enclose her in his great chelate arms, not with any
violence, for the claws never snapped or closed violently. She was coy,
however, and refused to be won by his advances; for the dance may haye
been nothing new to the lady crab, nor half as interesting as it was to the
two spectators outside the water. Later the male also buried himself in
the sand, and the performance came to an end.1

The love dances of Saltigrade spiders also suggest a similar habit record-
ed of certain birds. Familiars of our American woods and fields will recall

the well known partridge dances. Among the Chatterers the beau-
Love tiful bird known as the Cock of the Rock (Rupicola rocia) is
a a of pamous for its saltigrade performances at the mating time. In-

deed, the action of our domestic pigeons and barnyard fowls,
although not so decided as these, yet suggest a like tendency.?

Mr. Wallace has given an account of similar actions by the beautiful
Birds of Paradise in the Aru Islands. They moult about January or Feb-
ruary; and in May, when in full plumage, the males assemble in the morn-
ing to exhibit thernselves in a most singular manner. ‘These are what are
called their “ sacaleli,” or dancing parties, and they occur in certain trees in
the forest, which are not fruit trees, but have an immense head of spread-
ing branches and large but scattered leaves, giving a clear space for the
birds to play and exhibit their plumes. On one of these trees a dozen
or twenty full plumaged male birds assemble together, raise up their wings,
stretch out their necks, and elevate their exquisite plumes, keeping them
in a continual vibration. Between whiles they fly across from branch to
branch in great excitement, so that the whole of the tree is filled with
waving plumes in every variety of attitude and motion.

The bird itself is nearly as large as a crow, and is of a rich coffee
brown color. The head and neck are a pure straw yellow above, and rich

metallic green beneath. The long, plumy tufts of golden orange
Displays feathers spring from the sides beneath each wing, and when the
cae bird is in repose are partly concealed by them. At the time of
Paradise, lts excitement, however, the wings are raised vertically over the

back, the head is bent down and stretched out, and the long plumes
are raised up and expanded until they form two magnificent golden fans,
striped with deep red at the base, and fading off into the pale brown tint
of the finely divided and softly waving points. The whole bird is then over-
shadowed by them, the crouching body, yellow head, and emerald green
throat forming but the foundation and setting to the golden glory which
waves above. When seen in this attitude the Bird of Paradise really

'T. H. Morgan, Popular Science Monthly, February, 1889, “The Dance of the Lady Crab.”
* For further material on the display of their charms by the males of birds see Darwin’s
Descent of Man, Vol. II., chap. xiii, Am, Ed.

LOVE DANCES OF SALTIGRADES. 57

deserves its name, and must be ranked as one of the most beautiful and
wonderful of living things.!

This habit enables the natives to obtain specimens with comparative
ease. As soon as they find that the birds have fixed upon a tree on which
to assemble, they ambush themselves in the neighborhood. <A boy waits at
the foot of the tree, and when the birds come at sunrise, and a sufficient
number have assembled and have begun to dance, the hunter shoots the
bird with a blunt arrow with sufficient force to stun it. It is then secured
and killed by the boy without its plumage being injured by a drop of
blood. The rest take no notice of the loss, but continue their amatory
dance, and fall one after another until some of them take the alarm.

Thus in these widely separated orders of animal life the excitement of
the mating hour influences the males in substantially the same manner.

That is, the sexual agitation finds vent in saltigrade movements,
Displays }efore and around the female, of various forms and degrees of

ree intensity. These movements appear to be directed towards the
Females, female with a view to attract her attention, excite her affection,

and win her favors. As far as I can judge, there is no reason
why this apparent purpose should not be regarded as the real one, and that
these devices, common to spiders, crabs, birds, and doubtless other animals,
are really prompted by the wish to secure marital favors from the female,
and that they do have a sensible influence. upon her.

Another interesting habit described by the Peckhams is the overspinning
of the female by the male with a little tent or love bower, within which the
two remain together, sometimes for several days. Three pairs
of the Zebra spider (Epiblemum scenicum) were placed together
in a box, and after two hours they had all come to an agree-
ment and mated, the male in each case getting his partner in the corner
of the box and spinning a cover over and around her. Sometimes, while
the male was working, the female would wander off several inches, but
when the bower was nearly completed he would seek her and half lead
and half drive her home, when he would follow her into the nest. Here
the mating would be accomplished after some slight preliminaries. The
female seemed to have some difficulty in choosing from among the males,
but after a decision had been reached and a mate accepted, there appeared
to be complete agreement, and the male commenced to build his house.

The habit of secluding and protecting the female has developed an
even more striking trait in at least one species. The males of Phileus
militaris were observed to select immature females, overspin them with a
’ little sheeted tent, then spin a second sheet above this as a cover for

A Love
Bower.

The Malay Archipelago, by Alfred Russel Wallace, pages 466, 467.

58 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

themselves, and remain quiet for a week in the little nest thus formed.
During this time every spider that approached was driven away. The
males went out occasionally for food, but were not seen to carry
Tending jn any for their mates. At the end of a week one of the males
Seas was observed to be pairing with his female, which had moulted
and was now mature. Successive observations showed that this
marital seclusion of young females’ was not an accidental result of artificial
conditions, but is a fixed habit of the males. It must be acknowledged
that it displays a remarkable degree of foresight and thoughtfulness—the
immediate product, no doubt, of the emotional conditions of courtship.
In all these various movements the position of the female of most
species was simply one of watching. She followed the movements of her
dancing partner, evidently with keen interest; sometimes took
The Fe- herself out of the way, but ordinarily was quite attentive until
male Qui- P fe ts : :
escent, ‘he entire rejection of the suit or the acceptance of the suitor.
Two species formed striking exceptions to this rule, as far as
the attitude is concerned. In one, the female lay close to the ground
with her first legs directed forward and upward, while her second legs
were held on the ground and stretched forward in front of her face. In
another species, Marptusa familiaris, a similar attitude was assumed by
the female, who lay on the ground with all the legs flattened out and
the palps slightly raised, the only movement visible being the vibration
of the palps. (See Fig. 25.)
The attitudes of the males were far more varied. A reference to the
details of the notes as given will show that at least seven characteristic
attitudes are assumed, namely :—
Sum- First, the legs of one side are bent over, doubled under,

mary: : 7 Selo ine obs ts :
eee and so kept while the male engages in his semicircular dance.

of Males, (Saitis pulex.)
Second, the body is well elevated, the abdomen lifted verti-

cally, all the legs upraised and stretched out, and the entire eight legs
touch the ground during the dance. (Synagales picata.)

Third, the male, like the two females referred to aboye, lies flat on his
venter, keeping the tips of the fore legs touch-
ing (Icius); or the male lies flat, wriggling his
abdomen and frequently turning from side to side,
his legs held up over his head, slightly diverging,
and often twisted, waved, or turned about. (Zygo-

ballus bettini.) (See Fig. 34.)
stat hd panes en Bets Fourth, the two front pairs of legs are stretched
ballus bettini approaching fe- out in a straight line from the cephalothorax,
male, (After Peckham.) Xi ete ‘
while the remaining legs are raised and curved
and used for moving the body forward in its whirling dance. (Marptusa
familiaris.)

LOVE DANCES OF SALTIGRADES. 59

Fifth, the first legs are extended directly forward, close to the ground,
the legs being slightly curved, with the tips turned up (Dendryphantes
capitans), or again he lies down on one side with the legs well extended.

Sixth, the fore legs are elevated high above the head and curved towards
each other, while the body is sustained upon the remaining feet during
the saltigrade movement (Philezus militaris), or again the fore legs are
extended and the abdomen turned up. (Habrocestum splendens.)

Seventh, the spider maintains a rampant attitude, something like the
position last mentioned, with the fore feet raised high and curved forward,
instead of toward each other. (Astia vittata.)

These are the most characteristic positions, and they are maintained
during the courtship dance with more or less persistence, according to the

various species. The position after the consummation of the
Position wooing is much the same in all species. In mating, the male

on. usually crawls over the female, or the female crawls under the
Mating. ™ale, and the palps are applied to the vulva while in this atti-

tude. An exception was observed in two species, where the male
jumped upon his partner from a distance of one or two inches, the ap-
proach being per saltem, instead of by the gradual crawling movement
above indicated.

For the most part the female appeared to be complaisant or, at the
furthest, indifferent. She maintained herself in a position to
watch the antics of her lover and to be influenced by them.
Sometimes she ran away and avoided the advances of her suitor,
but showed no disposition to attack or annoy him.

At least one exception, however, to this general complaisance was ob-
served in the case of Phidippus rufus, who is a ferocious creature, haying
a great advantage in size over her partner. It happened to one
assiduous male that in an unguarded moment he was pounced
upon and eaten up by the lady whom he was wooing. Another
species of Phidippus showed the same ferocity. This is our
large black Phidippus morsitans, a creature not in good repute in certain
parts of the country, it being regarded as one of our poisonous species.!
The single female which the Peckhams caught during the summer was a
savage monster. The two males provided for her had offered her only the
merest civilities when she leaped upon them and killed them.

The male of this species has the first pair of legs much longer than the
corresponding legs of the female, and also it is thickly adorned

Female
Attitude.

A Fero-
cious
Female.

Me with white hairs, some of which are long and others short and
Signals, Scale like. It was while one of the males was waving these

handsome legs over his head that he was seized by his mate and
devoured, This love signal was evidently not sufficiently attractive to win

‘Vol. I, page 276. A letter just received from Prof. Peckham denies this accusation.

60 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the consent of his unesthetic partner, and, no doubt, had he been less
concerned to produce a fine effect upon his lady love, he could have used his
legs to better purpose in running away. Nevertheless, the manner in which
the snowy legs, as well as the white hairs upon the palps, were displayed
before the female’s eyes, indicated that he had natural confidence that the
lady was to be won in this way, if won at all.

The Peckhams appear to have no doubt that the purpose of this re-
markable display on the part of the dancing males is to win the favor of
their chosen partners. It seems to them beyond question that, during all
these complicated movements, the portions of the body which are most
highly decorated are exhibited to the watching female, and are displayed
in such wise as to give the most heightened effect to the coloring. It is
noted that the portions of the body among Saltigrade spiders which are
commonly most highly ornate, are those which are placed at the front of
the body, as, for example, the mandibles, which are often of bright metallic
colors, green or blue. The arrangement of the hairs upon the face, and

also the coloring of the fore legs, appear very commonly to be more attract-

ive than in any other portions of the body.
Indeed, the Peckhams associate the development of this coloring upon
the fore parts of the body with the dancing habits which they so attract-
ively describe. They believe it to be the result of sexual
Color selection. In other words, the males which have the brightest
Develop- ;
aes colors upon the fore parts of the body are precisely the ones
which have survived; since, being more pleasing to the females,
they were the partners most frequently chosen, and thus the individuals
so marked more frequently transmitted their peculiarities to the offspring.
The males will pair as soon as they have the opportunity, and, as the
mating season lasts for two or three weeks, the most brilliant males may
easily be selected again and again.

lee oi

CORAPTER. LET,
COMPARATIVE VIEWS OF VARIOUS MATING HABITS.

From the mating habits of the various tribes of spiders, as described in
the preceding chapters, a number of generalizations may be drawn with
more or less confidence. These I have thought well to place in a separate
chapter, together with several facts connected with reproduction, but not
heretofore alluded to.

We may begin by noting the influence of the general habits and char-
acteristics of the various species upon the manner and conditions of mat-
ing. Several conclusions and inferences appear.

First, a marked difference is observed between the methods
of the Sedentary and the Wandering groups; and this difference
is characterized by the radical difference in their manner of living and
capturing prey. The Sedentary spiders carry their persistent habit of
dwelling upon the snare into the act of pairing, and the snare is with
them constantly the scene of lovemaking. Their courtship and mating
proceed while they are hanging to the lines of their snares in the natural
attitudes of ordinary life.

On the other hand, with the Wanderers the courtship is in the open,
and the male directly places himself upon the body of the female. In
this group, also, the power of ordinary habit is seen directing the act of
mating, although, of course, in an exactly opposite mode. In other words,
species that do not live by webs dispense with webs in mating.

With the Tubeweavers, again, we see the same influence of general
habit. This tribe is properly classed with the Sedentary spiders, for they

dwell persistently within their webs, by which they capture their
Value of : cate :
General Prey in large measure. Yet they do not maintain upon their
Habits, Webs, for purposes of feeding, the inverted position that charac-

terizes Orbweavers and Lineweavers. In other words, instead of
hanging to their snares head downward, they rest upon their snares in a
position entirely similar to that of the ordinary attitude of individuals of
the Wandering group, and rush out upon the prey entangled within or
near their webs, which they seize and devour, generally without swathing
them, as also do the Wanderers. We might, therefore, reasoning from the

(61)

Generali-
zations.

ee
? _ I7— F Tag

62 AMERICAN SPIDERS AND THEIR SPINNINGWORK,.

influence of general habit, naturally suppose that the method of union
would be a compromise between the two already described. So we find it.
Among the Tubeweavers the mating occurs within the tube, into which
the male penetrates. The bodies come into close contact, and the female
remains in a state of absolute quietude.

Second, the differences in mating habit among Sedentary spiders are
characterized and evidently modified by the characteristic differences in
their snares,

1. With Orbweavers making a vertical web the male approaches from
the under side of the female’s body, applying the palps in that position,

clinging meantime to the female. Sometimes, though probably
ee rarely, the male hangs on the opposite side of the round snare,
Sunes * and from. this position applies the palps through the open

meshes, or through the free zone, clinging the meantime to the
web, as in the case of Epeira sclopetaria.

2. Orbweavers making a horizontal snare, and probably all making
composite snares, as Epeira labyrinthea, etc., mate upon the snare, hang-
ing to the crossed lines, back downward; the male above, and face up-
ward; the female beneath, with face upward. In other words, in the case
of both these groups of the one tribe the mating occurs while the spiders
are in the positions most natural to them during their hours of capturing
prey—the one maintaining the vertical position, and the other the hori-
zontal.

Third, the Lineweavers assume precisely the same attitude during court-
ship that is observed by Orbweavers which make horizontal orbs, and,
probably, by those which have a retitelarian annex, like the Labyrinth
spider. Here the common habit in the two separated groups has operated
to produce a common habit in the act of mating, for the Orbweayers with
horizontal snares habitually hang back downward upon their webs, just as
do the Lineweavers.

Fourth, in the proportion that spiders come directly in contact with
each other during mating without the aid of a snare, does the disparity
in size between the two sexes seem to disappear. Among Tubeweavers the
male is generally equal, and sometimes superior, in size to the female.
The same rule applies to the various genera of the Wandering tribes. The
fact of direct contact would seem at once to suggest the necessity of equal-
ity in size between the sexes, or a greater degree of complacency on the
part of the female. One is not able to speak concerning the latter factor,
but certainly the former seems to be reasonably well established.

It is perhaps worth noticing that the greatest disparity in size between
the sexes is seen among Orbweavers, and the larger the species, as repre-
sented by the female, the smaller does the male become. In the smaller
species of Orbweavers the difference between the sexes is not so great, and,
indeed, is often scarcely noticeable.

“a

= ——<——

COMPARATIVE, VIEWS OF VARIOUS MATING HABITS, 63

Fifth, amorous solicitations proceed from the male, and as a rule the
female is apparently indifferent to, or a passive recipient of, his advances.
There are, however, in this respect, differences among the various species,
some females being more complaisant than others.

Sixth, the male is frequently less vigorous in physical organization than
the female, is generally shorter lived, and is provided, in a less degree,
with those habits which secure prolonged activity and greater security. In-
the matter of spinningwork his acquirements are incomplete or rudimentary
with many species; with some, however, this exception does not exist, and
the webs spun are as perfect as those of the female. Generally speaking,
there is a tendency among males of the Sedentary tribes to defective spin-
ning industry, while among females the habit is invariably complete after
their kind. Among the Wanderers, of course, the chief spinning industry
is cocoon making and is thus confined to the female.

Seventh, in the case of some species, particularly among the Saltigrades,
male spiders have the habit of attracting the notice of the females by
certain saltigrade or dancing movements, which appear to be conducted
with a view of displaying to better advantage certain attractive colors or
markings. This habit is noticeably limited to spiders belonging to the
group of Wanderers. In the nature of things it could scarcely exist in
the case of Sedentary tribes, since the opportunity to display the person is
excluded by reason of the habit which limits their life to snare and nest.

HKighth, in the period of courtship it frequently occurs that several
males attend upon one female at the same time. This rivalry is often
without any special demonstration of hostility between the attending gal-
lants, but sometimes results in quarrels which, for the most part, appear
to be without serious harm to either combatant. The quarrelsomeness of
rival males seems to be limited to or greatest in the Wandering tribes, a
fact which again probably depends on characteristic habits. Such con-
flicts are possible with Wanderers, as rival males must come in contact with
each other upon the open field in which their loves are prosecuted. But
as the amorous movements of Sedentary species must be limited to the
snare or nest of the female, opportunities for personal contact are much
more circumscribed,

Ninth, as a rule, the general solitary habit of spiders is manifest also
in mating habits. With most species there is no such permanent relation
of mates as that which one often sees in vertebrate animals. But to this
there are some striking exceptions. Certain species seem to have acquired
a degree of domesticity, so that the two sexes are quite invariably found
together at all seasons after maturity. In a few cases, it has been asserted
that the male possesses so strong a domestic character that he will share
with the female the care of the egg cocoons, thus approaching the habit
of certain birds, fishes, ete., who unite with their partners in providing for
and protecting the offspring.

64. AMERICAN SPIDERS AND THEIR SPINNINGWORK.,.

The maternal and sexual instincts exercise a decided influence upon
the industrial activity and art of spiders.

The influence of the maternal feeling is decidedly manifest in the spin-
ningwork of the female spider. I have often observed that with insects,

as ants, wasps, and bees, the habits by which their wonderful
Maternity architecture is created are prompted by and revolve about the
saree care of the young. It is most manifestly so with solitary insects
erine such as the Carpenter bee and the Mud-dauber wasp, but it is
tecture, none the less true with such social insects as ants, social wasps,

etc. Among these creatures the workers or neuters, as they are
popularly called, are undeveloped females, and possess all the instincts of the
female of their species. Upon them devolves the work of the colony. They
are the nurses of the formicary, as well as its architects, scavengers, soldiers,
and purveyors. The whole care of the eggs, larvae, and pups rests with them,
and with the greatest enthusiasm and self devotion they exercise that care,
venturing their lives freely on all occasions for the welfare of their wards.

With female spiders a like maternal devotion exists. Their cocooning
industry is the most intricate and ingenious of their spinningwork, and

this is directly the product of the maternal instinct. ow varied,
Industry complicated, and ingenious this spinningwork is will be shown
area ae in the series of illustrations given in the following chapters.
Maternity Numerous as they are, they but imperfectly represent the indus-

try of the aranead mother; and I am confident that, when this
field shall have been fully explored, my studies will be found to reflect but
imperfectly the actual facts as they exist in the aranead world.

It is highly probable that not only the cocooning, but also the nesting,
industry is under the influence of the same maternal sentiment. Certainly
many of the admirably constructed nests described and figured in Volume I.,
Chapters XVII. and XVIII., are used as home shelters for the cocoon and the
young; as, for example, the nest of our Theridium zelotypum, the won-
derful domicile of the English Theridium riparium, and the various sub-
terranean nests of the Lycosids. This would seem to be true also of the
remarkable nesting industry of the Tunnelweavers.

Abbé Sauvages expresses the belief that the Trapdoor spider’s nest is
primarily designed for the preservation of the young, rather than for the
preservation of the individuals themselves. Certainly these ingenious struct-
ures do serve as a nursery for the spiderlings, as will be shown in Chapter V.
The cocoons are suspended within them under the vigilant watch care of
the mother, and therein the younglings are hatched and dwell for a con-
siderable period of time. Their habit appears to be to leave the maternal
nest only when they are abundantly furnished with strength to enter upon
housekeeping for themselves. Then they migrate, and, establishing them-

COMPARATIVE VIEWS OF VARIOUS MATING HABITS. 65

selves in the neighborhood, pierce the earth with tiny tubes, which in their
silk lining and hinged trapdoor are tiny miniatures of the maternal dom-
icile. It certainly is in the line of that influence upon architecture and
spinningwork generally, which is associated with, and probably incitive of,
the maternal industry, that this remarkable talent for house building should
have been developed by the Trapdoor spider. But I am disposed to think
that the protection of the spider itself, from certain enemies which are
not as yet well known, has much to do with the structure.

Coming now to the male spider, it may be observed, in certain species
at least, that the sexual feeling serves, to some extent, a like purpose

with the maternal instinct in exciting the animal to a higher
Sexual order of industrial art. As a rule, the spinning abilities of male
bt i spiders, as far as they relate to the capture of prey, have been

shown in Volume I. to be less decided than with females. The
rule is not absolute for all species, as in some cases the snare spun by the
male is precisely like that woven by the female. But in certain other
genera, as, for example, Argiope and probably Nephila, the snares of the
male are rudimentary, and do not compare in perfection with those of
the female.

Yet, in the hour of courtship, and under the influence of amatory ex-
citement, the male of the Water spider, Argyroneta aquatica, will be in-
cited to weave a silken cell close by that of his lady love, and resort to
the unusual device of uniting this with the domicile of his spouse by a
silken vault, which is so admirably arranged as to permit communication
without inviting destruction in the midst of an element ill calculated to
preserve intact a flimsy material like spider’s silk.

So, again, it will be found that among the Saltigrades the male of Phi-
leus militaris is prompted by sexual excitement to the remarkable habit of
preparing a special silken bower for his chosen mate, to which he
leads her, and in which he confines her until the nuptial hour.
What is yet more remarkable, the males of this species have ac-
quired the habit of selecting immature females, and secluding
them under a silken tent until maturity prepares them for nuptial rites.

It will thus be seen that, under the powerful influence of sexual feeling,
the male responds to a higher type of industrial art, and that to some
extent this feeling operates upon his organization in the same manner
that the maternal instinct influences the habit of the female.

Influenc-
ing In-
dustry.

III.

It has been shown that in some species of Orbweavers the females will
seize and deyour the male even immediately after the exercise of his natural
office, which indeed he has to undertake with great self control and care
to be able to accomplish it at all. From this propensity of the female,

_>:h ee

—_—

66 AMERICAN SPIDERS AND THEIR SPINNINGWORK,.

Rey. O. P. Cambridge! accounts for the great lessening in size of some
male spiders, as Nephila, in comparison with that of the female,

Dispro- hy a kind of sexual selection. It is obvious, he reasons, that

chagrin the smaller the male the better his chance of escape, and thus

in Sexes. Selection would operate until males became so small as only just
to be able to fulfill the office of impregnating the female,

It is perhaps difficult to reason upon this subject without a much larger
array of facts than we at present possess, but there are some points which
may be remarked upon with advantage. And it is to be noted that, in the
case of Orbweavers, the extremely diminutive size of the male obtains in
those species whose females have acquired the largest development. For
example, our indigenous Argiopes are among the largest of the tribe, and
their partners are very small, not exceeding one-fourth the female’s size.
The same is true of Nephila wilderi of our Southern seaboard, and in the

case of the large Nephilas of tropical countries, as, for example,
Disad- the Nephila nigra of Vinson (see Fig. 6, page 27), the differ-
vantages once ig even more remarkable. Now, it is certainly true that,
of Large 5 i , ; :
Size. were the male of a size corresponding with the female, his weight

upon the orbicular snare of the genus would appear at first sight
to be a disadvantage in several respects. First, it might break down and
injure the snare, and thus place a serious obstruction in the way of nat-
ural union.

Again, the advent of such a bulky creature upon the snare would at
once advise the female of invasion by a most formidable stranger, and
the natural instincts of the occupant of the web would be to regard that

stranger as hostile, so that her natural ferocity would be awakened, and,

the chances for combat and loss of life, or the prevention of sexual union,
would be a pretty certain result. From this view of the subject, Mr. Cam-
bridge’s suggestion, that the diminution of size would be a great advantage
to the male of these large species in accomplishing his amatory purpose
and protecting his life, has somewhat to support it.

But, on the other hand, it may be said that the immense snares of
Nephila and Argiope are no more fragile in proportion than those of
smaller sized Orbweavers, in whose case the sexual disparity’ does
not exist. Again, it might well be reasoned that natural selection
might have operated quite as favorably by maintaining the pro-
portionate size of the male or even preserving the largest examples of that
sex, inasmuch as increased strength would make him more formidable and
thus better fitted to accomplish his purpose. In other words, there is no
reason why Nature should not have preserved or bestowed the adyantage
of superior strength, as well as the advantage of insignificance in size and
therewith, perhaps, corresponding caution and ingenuity in approach. As

Per
Contra.

1 Zoologist, 1868, page 216, and Proceedings Zool. Soc. Lond., 1871, page 621.

COMPARATIVE VIEWS OF VARIOUS MATING HABITS. 67

to the latter point, however, I must say, as a result of many: observations,
that I have not been at all impressed with the ingenuity of the male sex of
Argiope in approaching his lady’s premises. On the other hand, I have
rather acquired the impression that he shows a remarkable degree of stu-
pidity or, at least, stolidity.

Moreover, Mr. Cambridge’s argument implies the fact that at one time
the sexes were of equal size, and that natural selection operated in the way
of producing a diminution of size in the male, to his advantage.
But this hypothesis, in its first particular, is not admissible, by
the very nature of the reasoning, which implies the necessity for
a reduction in size in order to preserve the male, and thus facilitate the
preservation of the species. It seems difficult to convince one that Nature,
having at the outset provided a comparative equality between the species,
or wrought the sexes up to such an equality, should have felt compelled
to reverse her decision and her processes, and reduce the size of one of the
sexes to such ridiculously small proportions. In other words, if Mr. Cam-
bridge’s theory starts out, as it seems to do, with a comparative equality of
the sexes, there appears to be no reason why that equality, having once
obtained, should not have continued; for the fact that it had once ob-
tained forbids the hypothesis that any necessity existed, or would be likely
to arise, for reducing the original equality of size.

Still further, it is a very common thing to find Orbweaving species
whose sexes are of nearly equal size and vigor. Such, for example, are

Epeira strix, Epeira sclopetaria, and Epeira labyrinthea, which
Sexes of i + ae . .
Equal 2% among our most common indigenous species. The same is
Size: true of many Lineweavers, as, for example, Theridium tepidari-

orum, Steatoda borealis, Linyphia marginata, and Linyphia com-
munis. The reasons for difference in size between the two sexes would
seem to be equally potent in the case of the above species, and all others
of like habit. Yet we see that natural selection has not operated along
the line of action supposed in the case of Nephila and Argiope. Cer-
tainly these exceptions are too numerous not to be regarded as throwing
much discredit upon the theory or, at least, laying upon its supporters the
burden of further proof.

It is pertinent to ask, do those Orbweaving species which habitually oc-
cupy nests or tents, in which the pairing occurs, at least occasionally, show
a disproportion of size on the part of the sexes? One would
reason that in such cases a substantial equality of size, or even
the superior strength of the male, would work to his advantage,
and so to the propagation of the species. That is to say, it has been
found that among the Tubeweavers and Wanderers, and all other species
where courtship and mating are conducted by direct contact, and not by
the act of suspension within the snare, as is habitual with Lineweavers
and Orbweayers, the male is of equal, or even superior, size. In the case

Natural
Selection.

Nesting
Species.

0
a. ‘

68 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of the nesting species of Orbweavers does the same rule obtain, thus fol-
lowing the line of similar mating habit?

With Epeira trifolium the males observed by me are very much smaller
than the females. The disproportion is nearly as great as between the
male and female of Argiope. In the case of Epeira insularis the male is
smaller than the female, but the disproportion is not great. ‘The male of
Epeira domiciliorum is not greatly inferior to the female. The male of
Epeira cinerea is smaller in abdomen, but in the size of the cephalothorax

and mouth parts is about equal, and the legs, if anything, are

geen q more powerful.. So far as our American species throw light
Disparity Upon the question, it would seem that the Orbweavers who per-

sistently dwell within tents, show no very marked disparity of
size between the sexes. This is the rule, with occasional exceptions. Among
British spiders, as described by Blackwall, the case is as follows: In Epeira
quadrata the male is about one-half the size of the female; the female of
Epeira apoclisa is about one-third longer than the male.’ The proportions
are nearly the same as those prevailing between the sexes of American
nesting species; but there is apparently a greater tendency in the former
than the latter toward diminished size of the males. On the whole, it can
hardly be inferred that the facts among Orbweavers indicate that equality
of size results from contact of male with female without the mediation of
a snare.
It has already been intimated that, ordinarily, where the sexes habitu-
ally come into direct contact, by reason of their natural habit, the dispar-
ity in size between them is less noticeable. Kyen among species
Sexual of the Sedentary tribes, such as the Tubeweavers, where union

Size in ; : :
Tube does not occur during suspension upon the web, this rule ob- oe
weavers, tains. For example, the male of Agalena nevia is quite as

large and powerful as the female. In the case of the remark-
able Water spider of Europe, Argyroneta aquatica, the male is even larger
than the female. There is, perhaps, some relation between this fact and
the necessity which seems to be compelled by the element in which the
life of the species is spent, and which appears to preclude anything like
cautious approach. The male Argyroneta must necessarily adopt for his
motto the Virgilian sentiment “Fit via vi” in his approach to his lady’s
domicile. It thus falls out that the superiority of size proves to be a con-
siderable advantage to him.

Among Tubeweavers, generally, there exists no very marked disparity
of size. With the immense creatures popularly known as Tarantula, the
Mygalidee, the male is certainly somewhat smaller than the female, but the
difference is not sufficient, as far as my observation extends, to make any
great disproportion in vigor. In fact, in this and in all other cases, it

1 Spi. Gt. Brit. and Ir, plate xxii., Figs. 236, 237.

_ 4 oe le.

COMPARATIVE VIEWS OF VARIOUS MATING HABITS. 69

must be remembered that the abdomen of the male is always, and neces-
sarily, by reason of difference in organization, much smaller than that of the
female, which gives him, to the current observer, the appearance
of being less in general size. But the organs of offense and
defense are not situated in the abdomen. The legs and mouth
parts of the male, including the cephalothorax, are often equal or superior
in size to those of the female, even when he seems at a casual glance to
be much smaller by reason of difference in size of the abdomen.

Among the Wandering tribes the same rule holds good. The Attoids
show a substantial equality in size between the two sexes, there being an
average difference of about one-sixth in favor of the female.
But this difference in size does not necessarily imply a difference
in physical vigor. It must also be remembered that at the mat-
ing period the sexual excitement of the male imparts an abnormal degree
of strength, which helps greatly to overcome any disparity in physical or-
ganism which may exist. The Lycosids follow the same rule of substan-
tial equality in size, with a slight difference in favor of the female.

In the case of the Laterigrades the rule holds good, as an examination
of the superb plates of Blackwall’s “British Spiders” (Plates IV and V),
will show. There are some remarkable exceptions, for which I am not
able to account. With a few Thomisoids there is said to exist the same
immense disparity in size that has already been noted between the sexes
of the Orbweaving Nephila and Argiope.?

Male
Anatomy

Wan-
derers.

LY.

The numerical proportion of the two sexes is a matter of great interest,
not only in itself, but in its bearing upon certain theories, especially relat-
_. ing to development. The facts are so insufficient, and at many
Numeri- points so conflicting, that it is a difficult matter to arrive at any-
cal Pro- Fs ‘ :
nota thing like a settled conclusion.
of Sexes. M. Eugene Simon remarks that in the genera where the in-
equality between the sexes is slightest the number of males ap-
pears to be equal to that of the females, since at the period of amour Orb-
weavers having this characteristic regularly come together in pairs. But
in the genera where there is a great disproportion of size the number of
males is much more considerable, since it is not unusual to see four or five
individuals of the sex courting one female. These little males mature first,
but the duration of their life appears very short, for after the period of re-
production they completely disappear. ‘They do not construct a proper
web, but keep in the neighborhood of those woven by the female, watching
for the propitious moment for union.?

1 Cambridge, Art. Arachnida, Brit. Eneye.
2 Les Arachnides de France, I., page 20.

o— 2 eo) ee ae a

70 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Blackwall and Mr. C. Spense Bate reported to Darwin that the males
of spiders are very active and more erratic in their habits than those of
females.1_ This appears to be a general opinion among araneolo-
Are Males pists, upon what ground as to the matter of activity I am not
pein able to perceive. One, of course, is compelled to ask what is
Sex ? meant by activity and inactivity as applied to spiders. Cer-
tainly the words must be regarded as relative terms. here is
a sense in which the females of Sedentary spiders are not as active as the
females of the Wandering groups. They may not, indeed, be able to
make way over the ground and among herbage with the same facility
that marks the Saltigrades, Laterigrades, and Citigrades; but the activity
in spinningwork of the average female Orbweayer is simply
Female enormous. One who has watched the method by which the
ee great round webs of our common indigenous species are spun,
tivity, Will certainly agree that the operator is one of the most active
of creatures in that department of work, at least. The rapidity
with which the threads are woven, the unceasing play of the hind legs
in pulling out the thread, and the striding of the other limbs around
the circle, together with the active exercise of the remaining organs, are
evidences of immense vigor and activity. The fact that such a large
and intricate web as Epeira spins can be wrought out in the course of
half an hour or forty minutes, is proof enough of this activity. These
snares will be reproduced several times a day if necessary, and the repro-
duction continues day after day throughout the lifetime of the aranead.
So also the same vitality of the female Orbweaver appears in the con-
struction of nests, which is not an inconsiderable work, involving no
slight exercise of strength, as well as of ingenuity, as any one will see
by turning to the chapter upon Nesting Habits, of this work, Chapter
XVII., Volume I.
Again, this activity appears in the capture of prey. If any one will
take his stand before an average orbweb of almost any common species,
_, say Epeira strix or Epeira sclopetaria, or Argiope cophinaria,
Activity at a season when flies and other insects abound, and in a site
ee where they are plenty, he will be surprised at the intense ac-
tivity displayed in the capture of insects. One after another
these victims are seized, swathed, dragged to the hub or den to be de-
voured, and that with a display of vigor in capturing, in swathing, in
cutting out the captive, and repairing the web, which must strike the
most casual observer. The feast will be left a number of times to seize
and truss up in like manner other victims who happen to strike the snare,
and on each successive capture the same tremendous rush and energy of
action will be noticed.

1 Descent of Man, chapter ix., Vol. IL., page 329.

COMPARATIVE VIEWS OF VARIOUS MATING HABITS. 71

I scarcely know a limit to the voracity of these orbweaving spiders
when full opportunity is given them to feed upon their natural prey ;
and I can certainly appeal to any one who has observed the
actions referred to, whether the whole demeanor of the aranead
is not such as to impress him with the sense of a vast store of
vitality, and an almost exhaustless activity. Taking, then, the spinning-
work and the ordinary action in capturing prey by means of nets as the
standard, it cannot be affirmed with truthfulness that female Orbweavers
are inactive, or that they suffer in respect of this element from compari-
son either with the Wandering tribes or with the males of their own
species.

I might go further and say that when a female Orbweaver is placed
upon the leaves of a plant, or even upon the ground among the grasses,
she will display an amount of activity in getting from leaf to leaf, and
limb to limb, and from point to point, which is surprising in a creature
whose habits are so generally sedentary. I have often been amazed at the
rapidity and facility with which the largest Orbweavers, as Argiope cophi-
naria and argyraspis, could make the circuit of a bush, or travel over a
plane surface.

As to the males of Orbweavers generally, it is certainly not in accord-
ance with my observations that they are more active than the females. On
the contrary, I am disposed to think them rather lethargic and

Female
Activity.

mae ., sluggish fellows. I am aware that it has been said, in corrobora-
uggish- |. j Page a
neuEr tion of the theory that the female is more inactive than the

male, that she will hang to the hub of her orb, or remain motion-
less within her tent for hours and perhaps even days. It is true; but
that action is quite as characteristic of the male as of the female. I have
seen the males of Cophinaria, in attendance upon a female, hanging upon
the outer courts of their lady love’s snare, apparently entirely inactive, for
as much as two or three or four days in succession. They are very patient
in their waiting, and make few movements during the courting period.
So also it may be said that those Orbweaver males which spin webs
that are as perfect after their kind as those of the female, show precisely
the same degree of patience in managing their snares and watching for
the advent of insects, as is shown by the female.
If we turn now to the Wandering groups, and make comparison be-
tween the males and females of the species of these tribes, I am certain
that it will be found that the females are as active as, or even
Wander- more active than, the males. During certain seasons of the year,
a=. as, for example, when they are carrying their cocoons, they do
roups . j ;
Gane indeed prepare for themselves a little cave or silken cell where-
pared, in they live until their young are hatched. But during that
period, even, the Lycosids may be found running around upon
the rocks and over the fields, dragging their egg bag after them. When

ne i A

ia AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the young are hatched, it is not uncommon to observe the mother wan-
dering over the fields with all her offspring piled upon her abdomen and
the lower part of the cephalothorax—a strange, and, to most beholders, a
horrible sight, since the ordinary observer is not apt to associate the un-
couth vision with the beautiful maternal devotion which the spider thus
shows, and which has its analogue in the human mother bearing her
child in her arms, or carrying it upon her bosom. Moreover, the exca-
vating and fitting up these subterranean homes is a strong proof of a de-
cidedly industrious character, and the act requires the exercise of great
vigor, which, of course, is exclusively by the female.

As a matter of fact, therefore, I am compelled to think that among all
Wandering groups the difference between the activity of male and female
is certainly not in favor of the former. Whatever conclusions, therefore,
are drawn from the belief that the male is possessed of greater activity
and vital force than the female, must, in my judgment, be regarded as
erroneous. That he is more erratic, in certain species, is true.

There appears to be little doubt that previous to the act of pairing, the
fertilizing fluid is extruded from the sexual organs of the male upon par-
ticles of spinningwork, thence is transferred to the dig-
ital joint of the male palps (Figs. 85 and 36), whence

it is conveyed to the epigynum of the female. ,
Convey- he alternate inflation and contraction of the
niga hehi ah palpal bulb is probably the means by which

the fertilizing fluid is forced into its proper
receptacle. Various naturalists have been able to estab-
lish this fact. Menge has observed, in the cases of Lin-
yphia, Agalena, and Lycosa, this fluid collected from
the sheet like spinningwork.!

Mr. Campbell, in the case of Tegenaria guyonii above
cited, was able to confirm this account. It was only dur-
ing the last moments of the process that reflected light
permitted him to see a triangular silken sheet attached
to the spider behind the abdominal sexual organ by its

ee , apex, and by its external angles to the mesh across
aaa a ee the bottle in which the aranead was confined. The
vittata, Fic.36. Epeira sheet extended from under the abdomen to the anterior
domiciliorum, largely

magnified. (After Em- Part of the sternum, and lay above the palps. The male
erton.) now left the sheet and approached the female; but she
appeared heedless of his addresses. The observer seized him, and in his
attempts to evade capture he injured the silken sheet. An examination

7 Preussische Spinnen.

COMPARATIVE VIEWS OF VARIOUS MATING HABITS. fs:

showed that the sides consisted of many shreds (Fig. 37, A), while the
intervening space was covered with an irregular mesh, which was doubtless
originally more systematically arranged. Here and there was a mass of
semen containing a fine, granulated substance (Fig. 37, B) of great refract-
ive power. The whole was in a very liquid state, and spermatozoa were
arranged singly above the threads.

Immediately after the spider was secured, one of its palps was removed.
The vesiculum seminis was charged with spermatozoa, even to the embolos,
where they were plainly seen at intervals. He could not, however, discover
any on the external parts of the palpus. Menge is entitled to the credit
of discovering the relation between the male palpus and the male abdom-
inal sexual organ.!

Ausserer confirmed the observations of Menge in studies of Linyphia
triangularis and Dictyna benigna.? Bertkau, following the same line of
investigation with Linyphia montana and another
species, corroborated these statements. *

In the act of copulation Blackwall observed, what
I have recorded of Linyphia marginata, that the
palps were frequently conveyed to the mouth. He
saw a male Lycosa lugubrius apply his palps eighty

times to the vulva of the female without

Applica- the possibility of bringing it into contact
tions of : = : :
Palps with the inferior surface of its abdomen,

except by a very conspicuous change of *
position. As an equal number of similar acts was
performed by the left palp, we have the extraor- yy. 97. spermatozoa of Teg:
dinary fact of the palps being employed one hun- cette) X 570. (After Camp-
dred and sixty times during this greatly protracted ;

process, unaccompanied by any contact whatever with the part where the
seminal ducts are considered to terminate.*

VI.

Whether or not spiders may be propagated by agamic reproduction is
an open question. I am at least satisfied that females will produce cocoons
without previous pairing with the male. It is reasonably certain,

male also, that in some species a single act of fecundation suffices for
mo all the eggs laid for several years in succession, where life is
duction. 88 y

continued that long. Audebert brought up and kept for some
years many individuals of the domestic spider, probably Tegenaria domestica.

* “Ueber die Lebensweise der Arachniden,” Neuste Schriften der Naturf. Gesellsch, Danzig,
Vol. IV., 1848.

ae Beobachtungen uber die Lebensweise, Fortpflanzung und Entwicklung der Spinnen,”
Zeistchr. Ferdinendeums, 1867.

*“Ueber den Generationsapparat der Arachniden,” Arch. Nat. Gesch., 1875, page 254.

* Zoolog. Researches, page 3165.

74 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Some females which had been isolated, produced in succession several gen-
erations, each in its order being equally fruitful.

Mr. F. Maule Campbell records an example of probable parthenogenesis
in the common English house spider. An immature female of Tegenaria
guyonii was taken and boxed in May. It moulted twice, and survived
during the winter confined within a large glass bottle. Herein she quite
domesticated herself, and was fed throughout the winter. - In the early part
of April she spun the ordinary cocoon of her species, within which she
deposited eggs, and about a month thereafter she died. On the 7th of
June, Mr. Campbell, observing some movement in the cocoon, separated the
sheets enclosing the eggs and found that two spiders had been hatched.
Twelve eggs still retained vitality, while the rest were hard and shriveled.
The fertility of this spider, after a confinement of eleyen months, during
which time she twice moulted, can only be explained by one of the follow-
ing alternate causes: First, that she was impregnated previous to the cast-
ing of the two exuvie in an early, and therefore immature, stage; second,
that parthenogenesis occurs in true spiders.”

Mr. Campbell is inclined to believe that the case recorded by him is
one of agamic reproduction, inasmuch as he could find no lumen in the
exuvie through which impregnation could have taken place. This is jus-
tified by the opinion of Bertkau, which expresses the general belief of ara-
neologists: this much is certain, that spiders immediately upon or shortly
after the final moult become sexually mature.®

1 Cuvier, Animal Kingdom, Lond. Ed., Vol. XIII., page 468, supplement.
2 Jour. Linn. Soe. Zool., Vol. XVI., page 538.
3 “Ueber den Generationsapparat der Araneiden,” page 253.

PART II—MATERNAL INDUSTRY AND INSTINCTS.
CHAPTER IV. ?
MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS.

Tue maternal industry of spiders is concerned chiefly in the preparation
of the silken sac within which the eggs are deposited. It includes also the
various methods by which this sac, when woven, is disposed of in order
to secure a greater protection for its contents from exigencies of climate
and weather, and assaults of enemies. I shall treat this part of my subject
after the methods previously adopted, and describe in detail the cocooning
habits of Orbweayers, and then present brief studies of the cocoonery of
typical species of other tribes, with a view to comparison as to various
points, such as the form, number, modes of preservation, and construction.

Among, Orbweavers, the largest cocoon known, to me is that of Basket
Argiope. It is usually a pyriform or globular flask or sac of stiff, parch-
.__,. ment like, yellowish silk, suspended in various sites by a series
eae of short lines passing from all parts thereof to surrounding ob-

jects. These lines, at the points of attachment to the cocoon,
diverge into minute conical or pyramidal deltas, similar to those formed to
anchor the usual dragline when the spider walks.

The objects upon which the cocoons are hung depend, of course, upon
the local habitat of the individual. For the most part, Argiope spins her
web in low positions; on the tall grasses growing in the angles of a rail
or “worm” fence; on the miscellaneous shrubbery that will be seen along
the edge of a New England stone fence; in the low bushes of various sorts
found in fields, lanes, the skirts of woods, and out of the way places—one
will be sure to meet these pear shaped objects in October or early November.

A collection that lies before me as I write will be sufficiently typical
of the positions in which Argiope spins her cocoons. Here is a cluster of
tall grasses, upon which two cocoons are hung. One, with a brown
external case, is suspended within a series of closely intersecting
yellowish threads, which are lashed to the stalks of the grass
eight inches from the roots. Just within the little concavity formed by the

stems as they have been pulled together in a circular position, the little
(75) :

Cocoon-
ing Sites.

76 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

flask, with its precious contents, is swung. At the top of this clump a
second cocoon is placed. It is of a yellowish white color, and, in order to
give it a proper site, the tops of the spears of grass have been pulled down
and twisted together, so that the capsules, or graceful clusters of seed vessels,
hang around the cocoon on every side, giving it a beautiful setting. These
cocoons are eleyen inches apart, and were probably spun by two spiders.
Another example is hung
in the very midst of a tall
field chrysanthemum. The
cocoon is much larger than
those just described, and is
of a rounder shape. ‘Two
branches of the plant have
been drawn towards each oth-
er, and these again
Among towards the cen-
Fhocrcmna, tral stalk. With-
in the space thus
circumscribed the egg sac is
suspended in the midst of
a maze of lines attached at
one end to the cocoon, and
at the other to various parts
of leaves and stems of the
plant. It is about eighteen
inches from the ground, and
forms a pretty object amidst
the balled white blossoms of
chrysanthemum. (Fig. 39.)
A third and fourth spec-
imens are hung in similar
positions within the out-
branching limbs of a wild
flower unknown to me,
which is thick set with little
white blossoms. Still an-

Fic. 38, Cocoon of Argiope cophinaria, hung in the tops of grasses. other is hung within a little
canopy formed by the leaves

of a blackberry vine, that have the beautiful hues with which, in our
climate, the autumn is wont to paint the foliage. Still another is suspended
beneath a similar canopy, formed of leaves on a young maple

Leafy bush. Another has a similar site within the clustered leaves of
Canopies. PUsh nother has a similar site within the clustered leaves o
a fragrant honeysuckle vine; and yet one more has been sus-

pended upon the leaf stalks and under the leaves of our well known

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS.

Fig. 39. Cocoons of Argiope cophinaria, swung among field grasses and wild flowers.

Virginia creeper. A _ pretty
environment, indeed, this last
one, but of less stability than
beauty; for, as the autumn ad-
vanced, and the leaves of the
ampelopsis dropped to the
ground, the egg case, so care-
fully wrought by the mother
while expending upon it the
last energies of her life, fell
to the ground, and probably
would have soon mingled with
mother earth had it not been
rescued by the collector’s hand.
These cases will sufficiently il-
lustrate the natural sites chosen
by this spider upon which to
suspend her cocoon.

The hanging of the silken
flask is not without an evi-
dence of nice care and discrim-
ination in the adjustment of its
supports. The guy lines are
commonly so placed upon the
different parts of the cocoon,

es a a a
' :

78 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

and so stretched and fastened to adjacent objects, that the mother leaves
her precious casket so well poised and finely hung that even the strongest
wind fails to disturb its balance when a good position has been selected.
In this position it will commonly remain until the brood is hatched; but,

as we have pare seen, sometimes the mother’s care is misplaced.
Stability T+ sometimes happens that the cocoon is simply anchored to
eis leaves, and, when the autumn brings the usual fall of foliage, it
is carried down to the ground. There, buried among rubbish, covered with
snows and rains, the chances for development of the young are seemingly
not very good. Yet even thus it is possible that, in sites comparatively
undisturbed by tramping feet of men and animals, the eggs may remain
healthful throughout winter, and yield their
broodling Argiopes when spring suns dissolve
the snow and the spring wind has scattered
the leaves.

It is not an unusual thing for Cophinaria
to hang her cocoon in the angle of walls in
a house or outbuilding. (Iig. 40.)
I have met a number of such cases
in the outlying parts of Philadel-
phia, as, for example, Germantown and West
Philadelphia. There still remain in those
sections a number of gardens and spacious
yards, within which this large and beautiful
creature has maintained her position against
all encroachments of civilization since the
landing of the Swedish pioneers. Their snares
are woven upon the vines which cluster about
Fic. 40. Cophinaria’s cocoon suspendea arbors, outbuildings, and verandahs; and it is

in the angle of a wall in the midstof 4 common thing for the mother, when the
crossed lines. =
cocooning time has come, to slip underneath
a roof or cornice, and there suspend her egg sac.

In this case she protects it by a slender encasement of retitelarian lines
spun entirely around it. A cocoon thus disposed is represented at Fig. 40,
as it was found in the early summer in the basement of a hotel at Atlantic
City. The enclosing lines were from seven to eight inches high, and of
about equal width. The lines were much soiled by dust, the accumulation
of winter and spring, but the cocoon proved to contain many healthy
spiders, although in the lower part it was infested with parasitic ichneu-
mon flies.

Another case of suspension within doors offered an interesting exception
to the usual mode. This cocoon was hung in the angle of the walls of a
room in Sedgley House, at Fairmount Park, Philadelphia, the headquarters
of Captain Chasteau, of the Park Guard, who said that it was made about

Indoor
Sites.

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS, 79

October 1st. When first observed, it was a round ball, which was gradually
wrought into a pear shaped object. This, when I saw it, was hung from

the under side of a
Cocoon sheeted curtain (Fig.
ae *41), that curved over

and extended like a
bridge from the shield shaped
hub of the snare to the adjacent
wall. The curtain terminated in
a pocket, from the bottom of
which the cocoon was suspended.
The cocoon was thus just behind
the orb which was spun across
the angle about seven feet from
the floor. The characteristic zig-
zag ribbon of the web extended
well downward, and a number of

lines stretched from side to side Aly
across the angle, nearly to the ye

“

floor, forming a convenient gang-
way for the spider.

Immediately after finishing
her work the mother spider be-
gan to languish. She would not
take flies as aforetime when of-
fered to her. Once she tried to
escape from the room into the
Park, but was brought back, and Fra. 41. Cocoon of Argiope suspended from a curtain

behind her snare in Sedgley House.
placed upon her lower gangway
lines, which she mounted, with great apparent difficulty, to the central
shield, behind which she stationed herself. She was found dead upon the
floor one morning, haying lived only a few days after the
completion of her cocoon.

The cocoons of Cophinaria vary in length from five-
eighths of an inch to one inch and five-eighths. Three meas-
urements between these limits are one and a half, one and
a fourth, and one and one-eighth inches. The bowl is gen-
erally about one inch wide, and the flask one-eighth inch
Fic. 42, Around co. Wide at the tip of the neck. The bowls are for the most

coon of Argiope part decidedly pyriform in shape, but sometimes are spher-
ba" ical instead of oval. As the spiderlings grow a little within
the sac after hatching, the bowl somewhat expands, or rather fulls out, but
the original shape remains substantially unchanged.

The structure of the cocoon is as follows: First, the outer case or shell

80 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

(Figs. 43, 44, 0.c) is usually a thin, stiff, parchment like substance, that
feels dry, and crackles under the touch, as though glazed. It is substan-

Fie, 43. Section view
of cocoon of Argiope
cophinaria. fe, flossy
envelope inside the
outer case, oc; p.d,
the brown padding;
e.u, the cup or dish
against which the
eggs (e) are deposited;
c.a, cap covering the
egg cup; ¢c.s, suspen-
sion cord.

is filled with a

tially water tight. I have found several cocoons of a softer
material, and thicker, much like a delicate yellow felt.
The glazing above mentioned is not the result of ageing
or weathering simply, but is produced by the action of
the spider herself, perhaps by the overspreading of the
viscid secretion which forms the beads on the spirals of
a snare.

When this outer case is cut away there is first pre-
sented a flossy envelope (f.e) of soft yellowish silk, which
quite surrounds the contents of the bowl. Next is a dark
brown pyriform or spherical pad of spinningwork (p.d),
which swathes the eggs completely, interposing a thick,
warm, silken blanket between them and the external case.
On the upper part of this pad is a plate or cup (c.u), of
like color and closer texture, with the concavity down-
ward. I have at least once found this to be a whitish
disk of stiff silk. The neck or stalk (nk) of the cocoon
compact silken cone (cs), of a yellowish or brown color,

which is united at the base to the egg plate (c.u), and at the top terminates
in a strong twisted cord (c.s), which sometimes extends upwards and forms
the central support to the cocoon. Next to the brown pad is often a thin
flossy envelope, which surrounds the egg sac. :
The latter is a rather closely spun pouch of
variable tenacity, and whitish or pinkish
white color, that encloses the thousand or
more eggs which lie in a globular mass with-
in the heart of the cocoon. The inner egg
sac (e) is attached above to the plate or cup
(c.u), which, after the spiderlings hatch, is
pushed upward by them not unlike a trap-
door, permitting them to creep out into the
surrounding padding, leaving their white
shells within the sac.

The plate serves to support the eggs, which
are probably oviposited upward against it.
One female, confined within a box, got so far
in the construction of her cocoon as to spin
the plate, but went no farther, leaving, how-
ever, this evidence of the point at which her ovipositing would have begun.

The genus Argiope is widely distributed throughout the globe, and the
cocooning habit of the species has elsewhere the same characteristics as in
America. Argiope fasciata of Southern Europe and Northern Africa makes

Fic. 44, Cocoon of Cophinaria dissected
to show the parts. Letters as in Fig. 43.

“ a

z< yiee. cys, nH ——— es ee ee ee ers
Winey, < Ly ’ 7 . aa <

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 81

a cocoon much like that of our Cophinaria. Fig. 46 shows the external
case, and -Fig. 45 gives a section view of the central egg sac, supported in
the midst of a bunch of loose flossy silk.1
IT have found numbers of Cophinaria’s cocoons on vacant city lots in
Philadelphia, strung to the stems of tall weeds on either side of a well
traveled footpath. The mothers had safely passed through the
betel perils of assaulting boys and voracious birds, and left these
i tokens of their maternal care in this conspicuous spot. As far
as examined the cocoons contained broods of healthy spiders, One excep-
tion, however, permitted me to see the position and structure of the egg
mass. It is a hemispherical mass five-sixteenths of an inch high and
wide. he eggs are bright yellow, contained within a delicate white or
pink hued membranous silken sac, through which they can be seen in
outline. -
It is interesting to observe that there is some variety among the mother
Argiopes in the manner of preparing a cocoon. I have one before me
which is composed,
Variation first, of a soft silken
in Struct- :
ear exterior case; then,
of three easily sep-
arated layers of delicate yellow
silken tissue, extremely soft
and beautiful. Next to these
layers is the loose yellow flossy
mass hitherto described, and
then the brown padding which
surrounds the egg sac proper.
This brown padding is not as

Fa, 45, Fie, 46,
abundant as I commonly find Fig. 46. Cocoon case of Argiope fasciata. Fie. 45. Section of

it f or th e reason p erh aps th at same, to show the central egg sac. (After Cuvier.)
) ) ’

the yellow silken envelope is so much more pronounced. Another cocoon
before me has in it nothing but the brown padding, scarcely a trace of
yellow floss, and no layers such as above described. I account for the
distinet layers by supposing that they were woven between well marked
intervals of resting.
The Banded Argiope is not as common a spider, at least in the imme-
diate vicinity of Philadelphia, as her congener Cophinaria. Her life ap-
‘ pears to be prolonged a little further into the autumn, for I find
pele her upon the bushes when the Basket Argiope has entirely dis-
raspis, 2Ppeared. Her cocoon is therefore made, as a rule, somewhat
later; but it is suspended in a similar manner and in similar
sites. I do not find it often, and, as compared with the cocoon of Cophinaria,

1 Cuvier, Regne Animal, Arachnides, pl. ii.

82 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

it is rare. It seems to be less fond of human society, or else less able
to stand the exigencies of civilization than Cophinaria. In outlying
sections, where Nature has been less disturbed
by men, it may probably be found more readily.
It is suspended by means of silken guys to the
leaves and stalks of grass or low growing plants,
which are bent over and also lashed together
above the swinging egg nest in the manner rep-
resented at Fig. 47. Again, it may be found as
at Fig. 48, swung in the midst of a retitelarian
maze woven amidst the branches and leaves of a
bush, or, as at Fig. 49, seated and suspended in
the crotches of a wild meadow flower.

The shape of her cocoon differs from Coph-
inaria’s in being hemispheroidal instead of pyri-
form; in other words, it resembles the lower half
of a spheroid. Across the wide top is stretched a
circular piece of silk, like the head of an Indian
drum. (Fig. 50.) The outer case is of stiff yellow
silk, as is also the head or top; this part, in a
cocoon now before me, is somewhat darker in color

Fic. 47. Banded Argiope’s cocoon
beneath a canopy of leaves and than the rest of the case. A marginal flap sur-

grass tops. A a .
rounds the head, and has various points to which

guy lines were attached in site. (See Fig. 50.) The height and width of the
cocoon are about the same—one-half inch. When the outer case is cut aside,
as at Fig, 51, the interior is seen to con-
sist, first, of a yellow flossy envelope,
which is packed between the inner wall;
and, second, an egg pad, which is not
composed of purple silk as in Cophinaria,
but of yellow silk plush loosely woven,
and is three-eighths of an inch long.
Within this are the eggs. Immediately
above is the egg cover of white silk
plush, which is commonly flat, not con-
cave as with Cophinaria. It is about
one-eighth inch thick, and is attached
firmly by silken threads to the inside
of the top of the case. Against this
cover, no doubt, the eggs are oviposited Fi. 48. Cocoon of Argiope argyraspis, suspended
amidst supporting cross lines upon a bush.

upwards, and are then covered by the

mother spider. The portion of the egg cover is shown at Fig. 51, where
one edge adheres to the remaining part of the top of the case, and also
at Fig. 52, where the object is viewed from the side.

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 83

Among various other examples of California spinningwork received
from Mrs. Higenmann and Mr. R. L. Oreutt, of San Diego, were several
cocoons of rare beauty. They were lenticular or hemispheroidal
Argiope masses, of a yellowish, yellowish green, and green color. (See
ee figures, Plate IV.) They were pulled out into angles at the flat
f side, as though they had been suspended by threads at the an-
gular points. ‘They varied somewhat in
size, from three-fourths of an inch to an
inch long, one-half inch wide, and three-
eighths high. It was long a matter of
wonder and discussion with me what spe-
cies formed these beautiful egg nests. Mr.
Oreutt finally attributed them to Argiope
argenteola, without giving a reason for his
opinion. The question was at last settled
by a living female specimen of that spider
sent me by Mrs. Eigenmann, which, hap-
pily, reached me alive, but very feeble. I
placed her under a trying box, fed her
with water and flies, and she revived.
The following morning a cocoon was hung
within the box, whose shape and color
solved the mystery, and proved that Mr.
Oreutt was correct in attributing the co-
coon to Argenteola.

This cocoon was a keystone shaped
patch of white sheeted silk, upon which
was raised a greenish button that enclosed
the egg mass. (Fig. 53.) The white color
of the sheet can hardly be characteristic,
for in specimens before me this part is
green.

The whole was suspended between lines
that were attached above to the lower foun-
dation lines of the orb, and to the sides and
bottom of the box beneath. Evidently the Fe. 49. Cocoon of Argiope argyraspis,

: » A ‘ hung upon the stalks and leaves of a
spider, in spinning her cocoon, had first — \iia fower.
stretched the sheet, and against or within
this had placed her eggs, which she then proceeded to overspin in the usual
manner, though, of course, it is not impossible that in this and like cases
the cocoon may be framed upon a flat surface and then raised and sus-
pended in the above described position. In general appearance this cocoon
resembles that of Epeira rather than the typical Argiope cocoons as rep-
resented by our two familiar species, Cophinaria and Argyraspis. But in

EE a ee eee

84. AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the manner of suspension, as well as the character of the egg case, Argen-
teola resembles her congeners. !

A cocoon, when dissected, shows two principal parts—the basal sheet
above referred to, and the cup or case which is set upon it. Both these
parts consist of closely
woven silk, like that
which forms the out-
er case of Cophinaria
and Argyraspis, the
latter of which it most
resembles, This cup
is of a yellow or yel-

lowish

Fria. 52.
Fig. 51. Interior, showing
Fa. 62. Side view of the egg cover.

Fie, 50. Fig. 51.
Fic. 50. Cocoon of Argiope argyraspis. X 2.
padding, eggs, and the egg cover.

green color,
and the deep green
tints appear most de-

cidedly in slight flossy tufts, which here and there overspread it. ‘The
inner surface of the basal sheet is overspread with
white silk. Within the case is a ball of
white flossy curled silk, which forms the
inner upholstery of the nest. It thus ap-
pears, that while the cocoonery of this remarkable
spider resembles that of Epeira in its external shape
and the nature of the interior furnishing, yet in
the texture of the case and manner of suspension
it is like the cocoonery of its congeners. In the
example produced in my trying box the basal
sheet is hung vertically. If it were suspended hor-
izontally, with the egg case downward (Fig. 54), it
would closely resemble an Argyraspis’ cocoon.

Mrs. Higenmann tells me that Argenteola makes
more than one cocoon. A specimen which had spun
a web in her sitting room placed a cocoon upon it somewhat in the posi-
tion observed by myself, as above described; but
shortly after (the time is not specified) a second
cocoon was formed upon the web about two inches
below the first one. A few days previous to this
iis thee td Raabe aaa ae cocooning the spider neglected to eat, and paid

genteola; side view. Aboutnat- no attention to the flies placed upon her web.
ry in The discoverer had concluded that the creature’s
mission was ended and death would soon ensue, but was surprised to find

Internal
Structure

Fia. 58. Cocoon of Argiope ar-
genteola; front view. Natural
size.

! Koch has described species from South America which closely resemble A. argenteola,
and perhaps may be the same, See Arachniden, Tafeln 5-8, Fig. 860, Arg. argentatus, and
Fig. 361, Arg. fenestrinus.

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 85

its lethargy only the condition naturally preceding cocooning. ‘he sec-
ond cocoon was a little larger and more flocculent than the first.

After this maternal duty the mother
disposed of the flies that were entangled
in her web, without any hesitation. This
was not the end of the matter, however,
for on the 14th of December, just three
weeks after the second cocoon had been
spun, a third was made, which was like-
wise attached to the web. On the after-
noon of January 6th, three weeks after
this last maternal act, the spider lost her
grip upon the meshes of her web and
fell dead to the floor, having been in the
possession of the observer three months.

The genus Epeira, which includes our
best known and most numerous species
of Orbweavyers, has little va-
riety among its most typical
species in the form of its co-
coons, the manner of protection, and
nature of sites selected for them. The
general form is that of a ball, hemi-
sphere, or semiovoid mass of thick, silken
floss, that enswathes a white silken bag,
within which a number of eggs, usually
yellow, are massed. ‘This is fastened in
any convenient and eligible position, at-
tached directly to the surface or hung
amid supporting threads. I have strip-
ped from a decaying trunk a bit of bark
eighteen inches long, on which one could
count forty or fifty of these cocoons in-
termingled with those of Agalena nevia
and other Tubeweayvers, and of Lateri-
grades, as well as the white silken tubes
of Saltigrades. (Fig. 55.) Often the
dried bodies of the mothers, who had
died shortly after their last maternal
care and work, were found clinging to

Epeira
Cocoons.

Fie. 55. Cocoons of Epeira underneath the
bark of an old tree.

the nurseries of their young. When deposited in such sites the eggs
rarely have any other protection in the way of spinningwork than the

EE ————————

86 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

—
ZN
A
Ss

|
AM

UY, f panel) Mi
at “me EM Mi ;

Fic. 59.

Fic. 56. Globular cocoon of Insular spider, spun in a paper box.
woven against a leaf within a glass tumbler. (Natural size.)
orum, within a curled leaf.

Fic. 59. Epeira cocoon enclosed within a curled leaf.
Epeira cocoons overspun with a common tent.

BiG. 56.

Fia. 57. Cocoon of Hpeira insularis,
Fic. 58. Cocoon of Epeira domicili-

Tia, 60.

—- - 3 _

MATERNAL INDUSTRY : COCOONS OF ORBWEAVERS. 87

flossy cocoon case, the shelter of the bark being, no doubt, sufficient barrier
against assault of enemies and stress of weather. A favorite site of this
gort is the trunk of an old hickory tree, whose flaky outer bark, curled
up at the free ends, offers an acces-
sible retreat.

A cocoon of Insularis, in my col-
lection, spun within a small paper
box, is a globular ball of
yellow silken plush three-
fourths of an inch in di-
ameter and of a light yellow color.
(See Plate IV., Vol. IL) It is hung
against the side of the box (Fig. 56)
in the midst of a maze of short right
lines an inch and a half wide and
high. These lines. are knotted to-
gether at innumerable points, which are marked by little white dots. ‘This
meshed envelope extends nearly to the cocoon, and certainly appears to
be a sufficient barricade against hymenopterous invaders, although it was
not able to save the eggs from those universal and well nigh irresistible
pests of collections, the Dermestide. I have another cocoon of this species
similarly disposed within an inverted glass tumbler, under which" the
mother had been confined. She attached herself to the bottom of the
glass (the top when inverted), and, as is the custom of her kind, hung
there back downward until the period of cocooning. (Fig. 57.) Not long
after she died, and her dried
up form is partly shown in
the drawing. The spots
upon the glass represent the
points of attachment for the
supporting lines of the co-
coon, and are little pats of
adhering silk.

Sometimes cocoons are
found laid against a leaf
which has been drawn
around it, as at Figs. 58
and 59, reminding one of
Fig. 62. Cocoon of Epeira domiciliorum, woven upon a pine tree. the manner in which cer-

tain lepidopterous larvee
protect themselves before they pass into the pupa state. When this sort
of protection is secured for the eggs, viz, a leafy covering around the
egg pad, no further envelope is added, precisely as when the eggs are
laid upon the under side of bark and stones.

Cocoon of
Insularis.

Fic. 61. Epeira cocoon in angle of a wall, protected
by a tent or palisade of lines.

88 , AMERICAN SPIDERS AND THEIR SPINNINGWORK.

In other cases, however, as in the angles of walls, porches, outhouses,
etc., the silken egg pad is itself enclosed in a tent of spinningwork more
or less open. (Fig. 60.) In some cases this tent is little more than a
series of lines drawn across the angle at a little distance from the cocoon,
as at Fig. 61. Strix, Sclo-
petaria, and Domiciliorum
are all in the habit of weav-
ing around their cocoons
such a tent.

A Domicile spider, which
I found in the act of com-
pleting her cocoon, was con-
tent with a scantier coyer-
ing than this. Her ege sac
was an oval mass of yellow-
ish brown silk one and one-
fourth inch long by three-
Fic. 68. Epeira cocoon protected by a tent of close spinningwork, fourths of an inch wide. It

was fastened upon a twig of
a pine tree. At one end short lines were thickly strung across from the
needle like leaves, making a sort of “fly” or awning. This
was repeated at the other end, thus about half covering the
cocoon. The mother spider hung to a few threads above (Tig.
62) her egg nest, with shrunken abdomen, and so much exhausted as to
be little inclined to move. This cocoon was made September 24th.

For the most part the outer tent is of closer texture than those above
described, being in fact an enclosing curtain of silken cloth, through which

the outline of the cocoon within may be traced. (Fig. 63.)
pres Great numbers of these tent enclosed

cocoons may be seen at the boat houses
near the Inlet of Atlantic City and Cape May.
They are made during the last days of May
and to the middle or last of June, and again
in the fall.1. The cocoons measure seven-eighths
of an inch long by six-eighths of an inch wide,

and less. The enclosing tent measures Fic. 64. Egg mass of Epeira, show-
Foreordi- two and two and a half inches long by Mee uae oa
nation in ‘ : : :
Nature, ue and three-eighths inch wide. Fre-

quently the tents are overlaid one upon another, or spun close
to each other, as at Fig. 58. I have found three large cocoons thus

Cocoons
in Tents.

? Of two specimens of Epeira sclopetaria kept by me, one cocooned May 22d; the other
May 26th; a third about the middle of June. An Epeira domiciliorum cocooned Septem-
ber 16th.

ss samme tm iil et ial

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 89

overlaid, and the outer tent, four inches long, covered the others so com-
pletely that one might have supposed the whole to be the work of one
spider. Undoubtedly, these works are precau-
tions against both enemies and the weather,
which, although without experience of the ef-
fects of either upon her offspring, the mother
takes as though she really foresaw the danger. jag, 65, gg mass of Epeira, showing
If an egg nest of this class be opened the under sheet and outer covering
‘ : removed.

there will be found, in order, first, the outer

tent, separate from the covering of the cocoon; second,.a thin white
silken sheet, which is the outer envelope of the cocoon proper ;
third, the thick egg pad of curled silk, usually yellow; fourth,
the eggs, a conical or hemispherical or spherical mass of small
yellow globules. (Fig. 64.) When the spider oviposits against a flat sur-
face, the eggs are generally laid upon a coating or sheet of silk spread
upon the surface, and the padding is then woven over it in the manner
of Argiope cophinaria. If the cocoon is suspended within a maze of lines,
the eggs are laid in the midst of the curled nest or egg pad, which is after-
wards completed. .

The cocoon of Epeira cinerea shows a variation from the common type
of her congeners, The egg pad is a large flattened hemisphere, an inch
in diameter, and one-fourth to three-eighths of
an inch thick. This is spun against some flat
surface, the boards of a shed, as I have seen it,
upon a light cushion of curled yellow silk. Over
and around this, on all sides, is woven the egg
pad, which is flattened down quite compactly,
and the whole mass lashed at the edges to the surface. The entire
cocoon has a diameter of one and five-eighths inch or more, and is a
quarter or three-eighths of an inch thick at the centre. (Fig. 66.)

Interior
Structure

lic. 66. Cocoon of Epeira cinerea,

III.

Epeira triaranea makes a cocoon of the common type, but smaller. Of
two now before me, spun in bottles, one measures one-fifth of an inch,
and the other about half that. They are both round or ovoid
flossy masses, protected by a maze of intersecting lines spun
around them. This maze is often thickened into a tent, in which
condition I have observed numbers spun in the angles of the joists of a
cellar at Atlantic City, in the early spring (May 22d), full of young spider-
lings just ready to emerge. These cocoons measured one-half inch long,
which is somewhat above the normal length.

One female was observed (New Lisbon, Ohio), whose cocoon was wrapped
up within a rolled leaf. This was swung to a cord, attached at one end

Epeira
triaranea.

———

——S ae S

EE ee a
~*

90 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

to the silken, bell shaped tent within which the spider nested, and at the
other end to the fence top against which the tent was placed. (Fig. 67.)
In this way the mother had her future progeny literally “cradled,” and in
good position also to be freely “rocked.” What freak had caused her to
make this divergence we can only conjecture; probably the cocoon had
first been spun upon the leaf, which, becoming loose, and threatening to
fall, was secured in the manner described.

A familiar resort of Triaranea in New England is the stone wall, char-
acteristic of that section. Underneath the irregular slabs or boulders of

granite which are heaped, one upon the other, to form the diyi-
Stone sion fences between meadows, ete., I have found large numbers
Fence : - oe ‘ ; oe :
Colony. of this species. The orb, which is usually about six inches in
diameter, is woven within the interspaces of the rocks, and the
spider has her resting place against the rough surface, or within the little in-
dentations of the stone which
forms the top of the cavity.
77 Against this surface the moth-
a nies
Amin] ev Triaranea weaves her bowl
i shaped tent, and against the
same surface, an inch or two
away, she spins her cocoon.
This is about a quarter or
three-eighths of an inch in
diameter; is a hemispherical
disk of flossy white silk, which
is overspun by a stiff, taut,
close, but transparent tent of
white silk about three-fourths
of an inch long. This may
be considered the typical co-
. coon of the species.
The number of eggs in
x three cocoons counted was,
Fic. 67. Leaf enclosed cocoon of Epeira triaranea, swung respectively, forty-five, forty-
to her silken nest and above her snare. :
two, and thirty-two. They
were of a gray color. Little spiders had just developed in one, and these
had yellowish abdomens, round, and very slightly oval, with the legs white.
The egg skin had just been cast, and the little fellows were stretching them-
selves and straggling about in a feeble manner.

One female was resting within a circular depression underneath a rock,
and had spun a few silken lines, forming the foundations of a little circular
tent, the framework of which extended downward toward her snare. Within
this was an old empty cocoon, against which the spider rested. Near by
was a fresh cocoon, nearly one-fourth inch in diameter, overspun by a

moe =e =a

4 R rs

~—S
gp

; SUOTTTENT
|

lon

SS ee ee ee ee ES eee ae phd ~~ =

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 91

tough silken tent, and this appeared to belong to the spider, who, moreover,
looked as though she might soon make another cocoon. The question was
started, but was not solved, does Triaranea weave more than one cocoon?
The cocoon was a little flossy
ball, flattened, of course, on
the side attached to the rock.
I captured one of the fe-
males, which cocooned in a
box, thus showing that the
cocoons above described were
those of this species.

A cocoon of Epeira thad-
deus was sent to me from
Vineland, by Mrs. Mary
Treat. It had been spun upon some potted ferns within her lodgings. It
is a subglobose sac, of a delicate pearl gray color, one-fourth
inch (six millimetres) in diameter. It is attached at the top to
a strip of silk ribbon, or rather it widens out at the top into
two triangular points, by which it is fastened upon a cord stretched between
two sprigs of fern. The egg ball thus swings free. (Fig. 68.)

I have secured cocoons of this species, by confinement within the trying
box, which differ from the above. They are globular or subglobular masses
of flossy yellow silk, about three-eighths of an inch in diameter. I believe
that, ordinarily, Thaddeus will be found to weave a cocoon of this sort upon
a leaf or other surface, probably enclosing it within a curled leaf, or over-
spinning it in the manner of Epeira triaranea.

I have not been fortunate enough to identify the cocoons of our common
Zillas; but a species which I observed in Florida made a cocoon shown
at Fig. 69, top of the cut. It was a flossy
ball, about three-eighths of an inch thick,
and was woven within the silken tent
which formed the spider’s domicile. It
was placed in the top of the tent, and
against the twigs, which formed a sort
of framework for it. After the cocoon
had been made the spider shifted her
domicile to a lower point, and gradually
spun a new dome shaped tent just be-
neath her cocoon, within which she con-

Fic. 69. Cocoon (top of figure) and tent tinued to dwell.
of a Florida Zilla.

Fic. 68. Cocoon of Epeira thaddeus, swung upon a line.

Epeira
thaddeus.

Le il ieee
|g iit i
Za" ay

Ky, Uh yFZ
ny RY f

The cocoon of Nephila wilderi, accord-
ing to Professor Burt Wilder,' is a large flossy hemisphere of silk, which is
usually spun upwards against a leaf or similar surface. The spinningwork

1Trans. Am, Assoc. Advance. Sci., 1873, page 263.

AMERICAN SPIDERS

Fig, 71.

AND THEIR SPINNINGWORK.

Fic. 70. Cocoon of Nephila wilderi, woven against a leaf. (After Wilder.)
Fic. 71. Cocoons of West Indies Nephilas spun on plants. (After Wood.)

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 93

is of a yellow color, and so slight as to show the loose mass of eggs within,
(Fig. 70.) It appears to resemble quite exactly the cocoon of its congeners
in Africa and the West India Islands.
For example, the cocoon of Nephila ni-
gra, according to Dr. Vinson,’ is of a
beautiful yellow color, and is attached to the bark
of trees, or spun against the surface of some re-
cess. Nephila maurata spins a large cocoon, of a
beautiful orange yellow color. This is not attached
to her snare, but is woven against any adjacent
recess, or in some shaded place near to her, al-
though sometimes she goes quite a distance from
her web to find a cocooning site. The orange
colored egg sac is enclosed in a flossy envelope
of a paler color.”

If we may credit the statement, or rather the
illustration of Mr. Wood, the Nephilas of the
West Indies, which are there known as the Tufted
spider, spin a cocoon similar to that described, but
suspended to the stalks of various plants, instead
of being hung beneath leaves or woven against
hard surfaces. The figure presented by Mr.
Wood, and which is here reproduced, is said by ye, 72, cocoon of a California
the author to be made from specimens in the asteracantha, woven upon
British Museum, although I do not remember to Nea ie ae
haye seen these when examining the collections of spinningwork at Ken-
sington several years ago.

Nephila
Cocoons.

ENS.

T have several cocoons of our American Gasteracantha, two of which
were sent from Southern California by Mrs. Eigenmann. A third was
woven by a living female sent from the same section; and a
fourth was received from Dr. George Marx, of Washington. The
latter is attached to the bark of a twig, upon which it is spun.
It is a flossy button or wad of a bright yellow color. The outer strands
of the spinningwork have a glossy appearance. It is about three-fourths
inch long and one-half inch wide. (Fig. 72, and Plate IV., Vol. II.) The
California examples are smaller but similar.

These cocoons are, in structure, like those of their African congeners
-as described by M. Vinson.4 This author describes a cocoon of Gastera-

cantha bourbonica as an ovoid, round and flattened, w oolly wad of a a yellow

: Avaneides des Madoegage ar, ete., page 191. 2 Idem, page 186.
3 “Homes Without Hands,” page 584.
+ Araneides Réunion, Maurice, et Madagascar, page 238.

Gastera-
cantha.

—_— =.” ©

94. AMERICAN SPIDERS AND THEIR SPINNINGWORK.

and green color. The case which enyelops it is twenty millimetres long,
and the central egg mass measures four-fifths of an inch (ten millimetres)
in width. The centre, which contains
the eggs, is white, but grows brown from
the moment of enclosure.

The cocoon of Meta menardi, as I
have found it, is a somewhat oblong
roll of brownish silk, not very com-
pact in texture, but sufficiently open to
allow one to see the eggs enclosed with-
in. It is deposited near the snare of
Fig. 78. Cocoon of Meta menardi. (About one- the female, and simply attached to some

third larger than natural size. .

surface by a rather sparing system of
supporting lines. According to Blackwall, the species (Epeira fusca) as
observed by him in North Wales makes a cocoon somewhat different
from this. In autumn the female fabricates a large oviform
cocoon of white silk, of so delicate a texture that the eggs,
connected together by silken lines in a globular mass a quarter
of an inch in diameter, may be seen distinctly within it. Its transverse
axis measures about eleven-tenths of an inch, and its conjugate axis
eight-tenths. It is attached by numerous lines, generally forming a short
pedicle on one extremity to the walls or roofs of the places it inhabits.
(See Fig. 74.) The eggs, which are yellow and spherical, are between four
and five hundred in number.? The general characteristics of the cocoon
as thus described by Blackwall correspond with those of
the American species, except in the habit of suspending
the cocoon by a short pedicle. However, a wider obser-
vation of the American species might show even closer re-
semblance in cocooning habit. One or two of my specimens
have a little tuft at one pole, as though a slight stalk or
attachment had been there made.

The cocoon of Tetragnatha extensa is a pretty object.
I have never seen the mother weaving it, nor have I ob-
tained it by confining the female within my trying boxes. A
But I have found it in the fields, where one may identify Ya

it by its resemblance to that spun by European ra, 74, Cocoon
Tetrag- individuals of the species; and, moreover, I have °% Enslish Meta
natha’s menardi or Epei-
Cocoon. batched the young, and thus demonstrated the true ra fusca, (After
cocoon. It is an ovoid object, about quarter of Blckwall)
an inch long and three-sixteenths of an inch wide and thick, and is com-
monly woven against a leaf, or twig, or bit of bark, or other convenient

Meta
menardi.

* Mr, Isaac Banks has also found it thus placed in Central New York.
* Blackwall, “Spiders of Great Britain,” page 350; and pl. 26, Fig. 252, g.

ee ae ee ee -_—*. - ™» i

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 95

object. (Figs. 75, 76.) I have found what I suppose to be this cocoon,
suspended by four diverging lines within an open space, as, for example,
in the post hole of fences, as shown at Fig. 77. The cocoon varies some-
what in color, being usually of a cream white tinted with green. The silk
looks almost like wool. The exterior is covered with little points or minute
projecting rolls, in this respect somewhat approximating the cocoon of
Uloborus. Within this exterior case are found the eggs, which are over-
spun by a slight flossy covering.
The English species forms its cocoon in June. It is described as round-
ish, less than one-fourth inch in diameter, fine and slightly woven; and
is either whitish with greenish tufts, or greenish with whitish
The Eu- tufts upon its surface. The cocoon is fixed to some object near
pee a the web, and contains pale yellow eggs.! This corresponds sub-
stantially with the account of Walckenaer, who describes the
threads of the interior as of a bluish green color, but the exterior as a
little browner in hue, and presenting inequalities as of little globules pro-
duced by the eggs.? Lister also describes the cocoon, which he frequently
found attached to the joints of twigs and to the leaves of plants. Thus
it was nearly or quite the first example of spider cocooning to attract the
notice of naturalists.

Most Orbweavers habitually make but one cocoon. There are some ex-
ceptions, however, among them two species very common in the United
States, viz., the Labyrinth spider and the Tailed spider, which

rig distribute their eggs in several cocoons, as does also Epeira bi-
Revatet furca of Florida. A rarer species having the same habit is the

Cocoons. Basilica spider; Uloborus plumipes and Cyrtarachne complete the
list of Orbweayers known to me to habitually construct a string
or cluster of egg sacs, These species represent groups having well defined
differences in structure and decided differences in the characteristics of
their snares.
The genus Cyrtarachne is remarkable by the peculiar form of the body,
and is distributed quite extensively throughout the United States. There
are probably two species, the Bisaccata of Emerton and Corni-

pel gera of Hentz. The cocoon made by the two species is similar
cee in general form, but there appears to be a marked difference in

the mode of attachment. Moreover, Cornigera apparently spins
but one cocoon, while Bisaccata, as its name implies, spins at least two;
and I have had cocoon strings sent me from California by Mrs. Eigen-
mann containing three. Thus Emerton’s specific name appears to be a
misnomer, I have a number of specimens; one collected by Dr. Marx

' Staveley, “British Spiders,” page 268, * Aptéres, Vol. IT., page 207.

96 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

hy,

x
WYSE

Nb i WM
ji ‘ % teh A ay Ali ik Men
Lt agli Ny "4 Go

NUM: ON diy,
( LN), Y Jur
CTX. \|

KOSS

Fic. 77,

Cocoons of Tetragnatha extensa: Fic. 75, spun upon a leaf, x 3; Fia, 76, woven against a twig, x 3;
Fig. 77, suspended within a fence post hole, the last about natural size.

MATERNAL INDUSTRY : COCOONS OF ORBWEAVERS. 97

at Washington, D. C., a single cocoon; another containing two cocoons,
gent to Dr. Marx from Fort Yukon, Alaska. Still others were forwarded
to me from various parts of the country.
The range of the species is, therefore, evi-
dently from the southern extremity of Cali-
fornia to the Alaskan peninsula on the west, and in
the east along the New England coast, and as far south
at least as Washington.

Several of my specimens are fastened to the twigs
upon which they were woven, and give a correct idea
of the ordinary manner of attachment. The cocoons
are about three-eighths of an inch in length, with a
foot stalk of varying length, which gradually ends in
a fine thread stretched upward along the twig. One
example, containing two cocoons, is lashed against a
twig by an overlying cord of yellowish silk five inches
long. The cocoons are composed of dark brown or
bluish silk, with overspread tufts or patches of: white.
They are separated by a space of nearly half an inch,
and the foot stalk of the lower cocoon is united to the
bottom of the upper one by a thick, stiff, blackish
cord.

The lower portion of the ball of the egg sac has a ria. 78. Cocoons of Cyrta-
scalloped fringe with blunt points or processes, which, — Tapimestspmded Against
as far as my specimens show, have nothing
to do with the manner of suspension. Nevertheless, they may
serve some useful purpose in anchoring the egg sac to the twig.
This description will fairly represent the form and mode of suspension
of all my specimens.

Emerton found his specimens at New Haven, Connecticut, on a beech
tree. They were dark brown, as dark as the bark of the tree, and as
hard. Around the middle of each was a circle of irregular
points. One of his cocoons was attached by a string to the
bark, and the other was attached in the same way to the
first cocoon. The spider held on to one of the cocoons,
which, therefore, had probably been recently spun. We may

safely conjecture the date of this observation, October 22d,
ena. to be the cocooning period of this species. The following
tarachne bi- Spring another similar pair of cocoons was found on a low
in OF oak tree in the same vicinity, still firmly attached to the
bark. From these the young came out in June.

In my specimens there is much difference as to the regularity of the
little exterior processes or points alluded to. In some specimens they are
quite regularly formed, and make a very pretty ornament upon the

Distribu-
tion.

Scalloped
Fringe.

98 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

cocoon. In others they are quite irregular, not only in their shape, but
in the mode of arrangement, being little more than irregular nodules upon
the surface. One of the
specimens from California
consists of three cocoons, the
first of which has the points
arranged with considerable
regularity, while the others
are less in size and are al-
most without rugosities. All
have little openings towards

Fic. 80. Cocoon string of a California Cyrtarachne, seen from two the top, through which, no
sts ge ek doubt, the spiderlings made

their escape. (See Fig. 80, which shows the cocoons natural size.)
Cyrtarachne cornigera is quite as remarkable in the character of its
cocoon as in its own structure. This cocoon is a flask shaped object, re-
sembling that of Argiope riparia, but with a neck relatively

Cyrtar- much longer. ‘Two examples before me differ greatly in size,
achne

one being more than one-
Cocoons. 5 ?

third larger than the
other.! In the former the stalk
or neck is of uniform thickness ;
in the latter it is twice as thick
at the mouth as at the bowl. (Fig.
81.) The cocoon is lashed at the
base of the bowl to a twig by a
number of silken threads, which
are attached to one side, carried
quite around the twig, and simi-
larly fastened to the opposite side.
The entire lower half of the bowl
is thus covered by the attached
wrappings, which are drawn so
tightly that the flask sits quite
firmly upon the twig. At the op-
posite end the cocoon is stayed by
lines that pass from the tip of the

stalk to the snare of the spider or Fig. 81. Cocoon of Cyrtarachne cornigera, lashed to
a twig. x 2.

other support. The attachments of
these guys are shown in Fig. 81, which is drawn twice natural size.
In the Camden cocoon (Fig. 82, natural size), the lashings are of a
1 No. 1, collected by Mr, Isaac Martindale, Camden, N. J.; length, 19 mm.; bowl, 10 mm. long,
9mm. wide; stalk, 9 mm. long, 8 mm. wide. No. 2, collected by Dr. George Marx, Washington,
D. C.; length, 12 mm.; bowl, 6 mm. long, 5 mm. wide; stalk, 6 mm. long, 14 to 8 mm. wide,

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 99

yellow, glossy silk, and so abundant as to make quite a ribbon. Here the
threads are carried around both sides of a projecting twig, as though the
spider mother had purposely availed herself of this mechanical advantage,
and are additionally strengthened by being crossed or twisted as they pass
around the branch to which the cocoon
is attached. The outer envelope is in
color a very dark yellowish brown, and
is of extraordinary stiffness. When cut
open the bowl is found to contain a
ball of white silken floss, within which
the eggs are deposited. This ball is anes see
fastened to a very tough twisted cord, IG. 82. Cocoon of Cyrtarachne cornigera (natural
that passes up through the neck (Fig. ene en eee ere
83), and which is the line by which the

egg ball was suspended before the outer flask was spun around it. The
texture of the external shell has every appearance, under the lens, of
having been hardened by means of a viscid secretion applied to it by the
spider; the toughness is evidently not the result of simple weaving.

Another example of Cornigera’s cocoon is drawn at Fig. 84. The
manner in which the bowl of the vase shaped object is seated upon the
twig and lashed by a ribbon is there well shown. The top of the stalk
is stayed by various lines wrapped about a neighboring twig.

Epeira labyrinthea belongs to the small group of Orbweavers that spin
compound snares; that is, snares in which the orb is associated with a

well developed retitelarian snare.'| The labyrinth of crossed lines
Laby- —_ igs placed behind and above the orb, and within this the spider
rinth :
older has her dwelling, commonly beneath a dry leaf; here also she
suspends her string of cocoons, placing them near her tent, and
usually above it and to one side, as represented in Fig. 85.

It consists of several, usually
five, lenticular or semiglobular
vessels, of a yellowish, tough
texture, about one-fourth inch
‘long and one-sixth wide. These
may be properly described as
woven dishes with covers. Each
cocoon consists of two disks
joined together at the edges
tightly enough to cause them to
adhere until the parts are grad-
ually loosened before the strain of the growing spiderlings, and finally
Open up and permit the inmates to escape.

Iie, 84. Cocoon of Cyrtarachne cornigera, with ribbon
lashing. and stay lines.

‘See Vol. I, page 131, and Fig. 115.

—_ ~

—-

100 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Ugniii\
yyy
Go

TINNYGN IRINA NO AD
These disks, on
examination, pre-
sent very uniformly
the appearance shown
at Fig. 88, a, b. The
lower part of the cup,
a, is an oval dish twice
as long at the top as at the

the form of a portable bath
tub much in vogue. The
upper disk, the cover or cap,
b, is in shape a miniature
soft slouch hat with
a rounded crown and
turned up rim. ‘The
rim of the cap fits
upon a minute cor-
responding lip of the cup.
When the eggs are first laid
the cocoon has a somewhat
flattened appearance, which

ON
LRUUESSS,

Fic. 85. The Labyrinth spider’s cocoon string, suspended within the maze above
her leaf roofed tent.

bottom, reminding one of

Le eT eee t—™

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS, 101

in many cases (not all) becomes much rounded as the spiders grow. If
the cap be lifted up or pulled off, as may readily be done when the
young are nearly ready to emerge, a
ball of yellow silk will be found inside,
amidst which the eggs are originally
deposited, and in whose fibres the spi-
derlings burrow. The cocoons are in
number about five, more or less, and
each one contains about twelve to twen-
ty eggs, so that the aggregate number Fic. 88. The

, . dish, a, and
of eggs is about equal to that found gover, b, of
in the single cocoons of some other ®Labyrinth

P spider’s co-
species. ean

For the most part the cocoons over-
lay one another, the top of each projecting one-
third to one-half its length over its neighbor, as
shown at Fig. 89, i, front view; ii, back view.
They are held together chiefly by a band of loose
threads (0, ii) which are stretched along the back

Fig. 86. Fic. 87. parts of the cups, although at the points where the
Cocoon strings of Labyrinth cocoons overlap they are also lightly attached. The

ay, Sa age eee band upon which the cocoons are thus strung is
above. Fic. 87. The tiled fastened to a strong, thick, branching white cord,
position of the cocoons. P 5
which is anchored above and below
to the network of cross lines. This cord is usually longest
above, deltated and often suspended upon a similar trans-
verse cord. (See Figs. 85, 86.) When the cocoons are
opened in October, the spiderlings are found fully de-
veloped, lively, and ready to escape. They resemble the
adult form in markings.

The cocoons are sometimes separated from each other,
as at Fig. 86, but again are all overlaid, Fig. 87, being
lashed together by the band of threads upon which they
are strung. Occasionally, the spider will spin her tent
beneath the lowest cocoon of the series, instead of the
usual leaf or other débris, and will be found backed up
against the same, holding to the trapline of her snare, fio. 89. Two overlaid
(Fig. 90.) The full page cut (Fig. 85) shows Labyrin- — (yen°™Snowing the
thea’s cocoons strung in natural site, above and behind cord i, and ii, o, x,
the leaf-roofed tent. sae ieee

The mother begins to spin her cocoons in August,
adding one every week, or thereabouts, until the tale is complete. The
suspensory cords that support the cocoon string are strong, thick, and of a
pure white color, I have found numbers of the empty cocoon shells in

— OE

102 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the early spring, hanging intact upon the bushes where they had been
placed, although, of course, the snare had entirely disappeared.

The Tailed spider, Cyclosa caudata, differs from Labyrinthea in the
mode of hanging her string of egg sacs. This is suspended within the
limits of her orb, above the central space, along the line of the
perpendicular. As the cocoons increase in number, the adjacent
radii and the connecting spirals are cut out, leaving a clear seg-
ment resembling that in the snare of Zilla, in the middle of which the
cocoon string hangs. (Fig. 92.) The number of cocoons
appears to vary much; I have usually found from three
to five; Hentz never observed more than five. They

are generally in shape a double cone, although often
round or roundish, and are from three-sixteenths
to quarter of an inch (five to seven millimetres)
long and one-eighth inch (three millimetres)
wide. A cocoon is not composed of two dis-
tinct parts, like one of Labyrinthea’s, but
is spun in a single piece of soft yellow-
ish floss, externally close enough to be
\ weatherproof, but which ravels out
into woolly threads when picked
with a needle.

Within, the sac is filled abun-
dantly with delicate, flossy, yellow
silk, in which the eggs are de-
posited. These vary in number;
for example, three now before me,
opened in succession, contain, re-
spectively, twenty-two, two, and
ten; certainly a remarkable differ-
ence. On one occasion a female
enclosed within a paper box began
to make a cocoon, but proceeded
no further than to weave a tiny
saucer, similar to that spun by Ar-
giope riparia. This would, there-

fore, appear to be the commencement of her cocoon, and it may be that
against such a disk Caudata habitually deposits her eggs before enclosing
them. However, I have not found this within her cocoons, as is the case
with Argiope’s, and conclude that the disk is made the basis of the external
sac, into which it is woven as the spider proceeds. The cocoons are often
well separated upon the string, but also are found touching and even over-

Cyclosa
caudata.

fh
ff

Fic. 90. Labyrinthea’s snare, viewed from behind, with
two cocoons in site above the tubular nest.

1 “Spiders United States,” page 127.

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS, 108

lapping one another like tiles. Sometimes nodules of flossy silk, or of silk
mixed with the débris of captured and devoured insects, are irregularly
interposed between the cocoons. This is, indeed, a fixed
and most interesting habit of the species, which will be
described in a succeeding chapter.

During a temporary stay in Florida, April, 1886, I
found nested upon the porch of Dr. Wittfeld’s place,
Fairyland, Merrit’s Island, on the Indian River a little
way below Rockledge, a new spider, which I named Cyr-
tophora bifurca. Its snare resembles that of Cyclosa
caudata. It also resembles that spider in the manner of
hanging its cocoon string in the vertical axis of its orb
just above the hub. ‘The character of the cocoon, how-
ever, differs entirely from that of Caudata. It is, in
shape, a somewhat irregular octagon, and is of a dark
green color. I have found as many as fourteen cocoons Be ete ea ee ciing
in one string, overlapping one another in the manner of manner of suspend-
cocoons of the Labyrinth spider, and which may also be = "8, = ae =
seen at times with the cocoons of Caudata, although for
the most part, the latter are arranged at intervals
along the string. (See Figs. 96, 97.)

The cocoon strings collected varied in the
number of cocoons attached thereto, probably ac-
cording to the period of advancement in the proc-
ess of ovipositing on the part of the mother.
Of the specimens collected one string contained
fourteen, another twelve, and another ten cocoons.
They are bound together, along one side, by con-
tinuous series of thick white threads, which ex-
tend from the top to the bottom of the string.
Each cocoon consists of two parts, which have
evidently been fastened together by a selvage.
These parts present the appearance of two dishes
placed together edge to edge. They are woven
of a soft, but rather tough, texture. A very slight
tuft of flossy white silk is found inside, and with-
FG. 93. Bra. 95. ma.ss, in this the eggs are deposited. In one cocoon of
Fic. 93. Cocoon string of Caudata, g string of thirteen, twenty-five minute dead spi-

with silk nodules interposed. (Nat- ° :

ural size.) Fic. 94(upper). Cocoons. Gers were counted, which had passed their first
fe he ooo Ee moult. In another cocoon, taken from a string
Epeira bifurca, showing shape and of five only, there were twenty-six. The num-
Bais gee dare ber varies a good deal, however. ‘The cocooning
period appears to extend into May; at least I have received from Miss
Anna Wittfeld, as late as the middle of June, a string, in which were

104

AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Fic. 97. Snare and cocoon string of Epeira bifurca, to show the
manner of suspension.* .

+ The shape of the cocoons is not well represented in the cut.

(See Fig. 96.)

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 105

some cocoons empty, one with spiderlings passed the first moult several
days, and another with young who had just broken the egg. There was
no trace of the bifurcated abdomen upon these younglings. The spider
is of a uniform light green color, about the shade of its cocoon.

Another Orbweaver that makes several. cocoons is Epeira basilica. I
am indebted to Dr. George Marx, of Washington, for the specimens from

which the following studies and drawings have been made, as
Cocoon of wel] as for the information concerning Basilica’s habit of caring
Basilica 4: her eggs. The number of cocoons is five, thus corresponding
Spider. ge ve, esponding

with that of Labyrinthea, and generally with Caudata. They
are round, covered on the outside with gray spinningwork, and united by
a cordage so stiff that the series stands out like a stick. They are attached
to a triangular patch of yellowish white silk, which is an expansion of a
long, glossy, strong linen like cord, composed of many
threads, by which the string of egg balls is suspended.
(Fig. 98.)

According to Dr. Marx, whose observations were made
at Washington, the string is hung just above the centre
of Basilica’s peculiar domed snare, and wholly or in part
within the dome, as represented at Fig. 99. The mother
has position beneath her egg bags, back downward, as
is the habit of Orbweavers making horizontal snares.!

When the cocoon is dissected, it is found to consist,
first, of an exterior sac of gray material; within this is
next enclosed a round black case (Fig. 100), four or five
millimetres in diameter, having a thin shell of remark-
able hardness, in this respect resembling the cocoon of Fis. 98. Cocoon string
Cornigera. When illuminated and examined under the cite cade
microscope this egg ball is seen to be composed of yellow
silken fibre of exceeding fineness, and so closely woven that, looked at
when within its bag, it is quite black. The paper like stiffness of the ball
could hardly be caused by even such fine spinning, and I believe that the
fibres are smeared with a viscid secretion, which gives them their peculiar
stiffness. When this black case is cut open it is seen to contain flossy silk
(Fig. 101), which forms the customary wrapping of the eggs and nest of
the young spiders.

The cocoon of Uloborus is about one-fourth inch long, and one-eighth
thick. It is drawn out at either pole into a point, and the surface is
covered with small pointed or blunted processes. (Fig. 102.) It
is made of a pure white silk, quite stiff of texture. Several of
these cocoons (I have never found more than three) will be found united
together so closely that they appear to be but one object, and not strung

Cocoon.

‘See for further details Vol. I:, Chapter [X., especially page 170, Fig, 159,

106 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

y i Wy
Supa, on NU
NYA

Me annie AC =

WW)
.;

\
4

WZ,
Ui / Yj

PN niin too Nee
i ON
eer

==
=
YSN
Za
S—

HN\\)
Wie

Fig. 99. Dome shaped snare and suspended cocoon string of the Basilica spider.

a et

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 107

loosely, by attaching threads, as is the case of some other spiders that

make several cocoons. However, in this respect, the habit may differ. As

a rule these cocoons are stretched like those of

Cyclosa caudata, along the axis of the mother’s

horizontal orb, and are thus im-

mediately under the maternal

care. (Fig. 103.) In this posi-
tion I have seen them in New

Jersey, and thus Mrs. Treat has F Soci eae

observed them, and so also Mr. to show the surface

Fra. 100. Fi, 101. Emerton has described them, ?%*

Oe man tb peat hel alae (Fig. 104.) Our American species appears in this
egg ball; Fra. 101, the ball open yespect to have the same habit as the European
to show the inside structure. : =e

species, Uloborus walckenaérius.

This modé of disposing of the cocoon, however, cannot be universal,
for I possess a specimen, received from Dr. George Marx, which is stretched
along a little twig, to which its orb was attached, at a point slightly above
the cocoon string. (Fig. 105.)

Hentz describes the cocoon of Uloborus mammeatus as tapering at both
ends, in color whitish, with veins of brownish black, and with many small
tubercles. He collected it in Alabama in dry places.!

VI.
The division here indicated between species habitually making a single
cocoon and species habitually spinning several is, on the whole, a natural
one; but there are certain facts to be noted which throw a measure of

Fic. 108. Cocoon string of Uloborus in position upon the snare.

uncertainty around any such generalization. For example, it has long been
supposed that Argiope cophinaria spins but one cocoon; and, judging from

1 “Spiders of the United States,’ page 129, plate xix., Fig. 126.

— eS ee oe
an i ’ Life "= iw
. - Fs

108 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

its size and the number of eggs that are found therein, one would seem
to be sufficient to guarantee the continuance of the species. I have no
‘ doubt that, as a general rule, Coph-
inaria makes but one cocoon, but
that there are exceptions is very
certain.

Several years ago a clerical
friend brought me two cocoons of
this species, which had been spun
on his premises by the same spi-
der. Mrs. Mary 'Treat has discoy-
ered what appears to her to be a va-
riety of Argiope cophinaria, which
makes four cocoons, and which she
accordingly named Argiope multi-
concha.! She sent me a string of
yi these cocoons, of which there were
four, of the general shape and about the usual size, strung within a few
inches of each other, They had been spun against the wall of a kitchen
in a house in Western Missouri. The spider mother was also sent, but the
specimen was much dried up, and in such a condition that it could not
be very satisfactorily studied. It seemed to differ in no particular from
Argiope cophinaria. If it be indeed the same species, what are the pecul-
iar circumstances that have caused
such a remarkable variation in
habit? Is it true that Cophinaria
does, more frequently than has
been supposed, indulge in the lux-
ury of an additional egg case?
Two cocoons of this lot were open-
ed and found to contain young spi-
ders that had hatched, but died
within the egg sac. The spider-
lings were not counted, but they
were very numerous.

During the summer of 1888 a

female Cophinaria was
a! discovered in the Farm-
ing Ar- ers’ Market of Philadel- Fic. 105. Uloborus snare and cocoon
giope. Phia upon the meat stall SANG On OE Oe
of one of the butchers.

She had probably been brought into the market from the country, hid-

Fic. 104. Cocoon string of
Uloborus; cocoons in the

snare. (After Emerton.)

1“American Naturalist,” December, 1887, page 1122.

MATERNAL INDUSTRY: COCOONS OF ORBWEAVERS. 109

den among vegetable leaves, as the huge tarantula and the large Lateri-
erade spider, Heterapoda yenatoria, are brought to our port from the
West Indies in bunches of bananas and other fruit. Or, she may have
floated in, as a young balloonist, from some city garden; for the species is
abundant in open grounds within the city limits. Instead of brushing
her down and killing her, after the usual manner of dealing with such
creatures, the farmer took a fancy to preserve her, and would allow no
one around his stall to inflict any injury upon her. She wove her char-
acteristic web against one of the iron rods for suspending meat, chickens,
game, ete., and there remained secure during the season.

Some time between the 10th and 20th of August she be-
gan to make a cocoon, which she enclosed within a little
tent of interlacing lines, after the manner of that repre-
sented at Fig. 40. About a week or ten days thereafter she
made a second cocoon, placing it in a position sixteen inches
above the other. Both of these co-
coons I saw precisely as they were
left by the spider. They were spun
within tents of crossed lines, five or
six inches long and four or five wide,
with a thickness of between two and
three inches. The lines constituting
the under edges of the tent were at-
tached to the post of the stall on
which the orb was spun. The upper
tent had its roof lines sustained and
drawn out from the post by the
foundation lines of the orb. (Mig.
106.) The lines composing the tents
were of a greenish yellow silk, sim-
ilar to that used in the construction of the cocoon cases.

I remoyed the cocoons and opened them. ‘The lower one was an
inch and a quarter long and seven-eighths of an inch wide; was com-
posed of a soft, yellow silken plush, and inside was constructed pre- |i
cisely like the ordinary egg sac of this species. It contained one hundred
and twenty eggs, all of them sterile. The only peculiarity was that the
stem which one usually finds at the top was missing. The second cocoon
was not quite so large, one inch long and five-eighths of an inch wide,
but was more perfect in shape, containing the usual stem. The eggs
within this cocoon were also sterile, and the number did not exceed fifty.
The number of eggs in both cases is small as compared with the usual
fecundity of the species.

We may probably account for the making of the second cocoon by
some abnormal condition of the ovaries, which prevented the ovipositing

aha ae
rare aE

——, Fe

ropes

ram *

aE:

aw ew
aw

Fie. 106. Double co-
coons of Argiope,
hung ina meat stall.

110 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of all the eggs at once. The first lot, when extruded, were protected in
the usual manner. Subsequently Nature compelled the mother to get rid
of the remaining eggs; and, moved by the same impulse which covered
the first lot, she was excited to oyerspin the second also.

This species will sometimes make a cocoon, or a part of one, in con-
finement, and I have observed that she will occasionally do the same in
natural site. I have the branch of a bush which shows the beginning of
a cocoon, being the little cup against which the eggs are spun, and also
what appears to be the inner egg bag. There is nothing more, and the
whole is stayed and shut in by the usual tent like spinningwork. Near by
is a perfect cocoon, secured in quite the same manner. If we suppose
that these two were made by the same spider, as is highly probable, we
may infer that the original cocooning purpose of the mother was diverted
in some manner, perhaps by alarm, which drove her from the spot. She
returned to enclose the work partially done, but, moved by the urgency of
motherhood, presently found a neighboring site, and finished her maternal
duties.

Epeira diademata habitually spins but one cocoon; but the Spanish
investigator, Termeyer,! in the early part of this century, discovered and
announced that she would spin as many as six cocoons when specially
nourished. ‘The fact strikes me as an extraordinary one, and I have never

felt quite free to fully admit it.

1 Walckenaer’s Aptéres, Vol. I., page 152.

L605 sl get gel ea
GENERAL COCOONING HABITS OF SPIDERS.

Havina considered in detail the structure of the cocoons of Orbweavers,
it is important for the sake of comparison that we should also consider
some of the typical cocoons of other tribes. It will not be practicable to enter
into details as fully as with the Orbweavers, nor to consider as many species
in any of the remaining tribes. But I will give a few examples, under each
tribe, of those species whose cocooning habits may be considered typical.!

Theridium tepidariorum is one of our best known Lineweavers. It
appears to be a native of America, and has been widely distributed by im-
migration throughout Europe. I judge that the course of immi-

based gration has been eastward, because in Europe the species is found
lum tepl- . . i
pad almost exclusively in hot houses, both in England and on the

continent, while in America it habitually lives in fields, forests,
ravines, among rocks, around outhouses, indeed everywhere that a cobweb
can be located. In short, in Europe the conditions of its life are artificial,
in America natural. It is a ferocious species and an expert trapper, prey-
ing upon some of the largest insects. It spins during the season from
three to five ovoid cocoons, often sharply pointed at one end, varying some-
what in size, but sometimes at least a third of an inch in the longest diameter.
These are woven within the retitelarian snare of the creature, and sus-
pended well towards the top.

Blackwall’s figure of the manner in which the cocoons are suspended
is erroneous, or the English spiders must differ in habit from the Ameri-
can. I have never seen any such sheeted, bell shaped tent as that which
this author represents as enclosing the cocoons.

The cocoon is rather simple in structure, consisting of an outer case of
yellowish brown material, well compacted, stiff, within which the
eggs are loosely placed without any or with but little interior pad-
ding. During the weaving process the cocoon is hung by a strong
thread, or series of threads, to the cross lines of the snare. ‘The spider

Cocoon
Weaving.

‘My systematic knowledge of the other tribes is far less than of Orbweavers, and I have
sometimes had difficulty in positively identifying the species whose habits I haye observed.
But I hope that I have not erred in many cases; certainly not in enough to materially affect
my statements and conclusions.

(111)

112 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

clings to her web by one long fore leg, while with other legs she revolves
her cocoon, using the hind legs, as is customary, to draw out the spinning
stuff. This issues in numerous diverging filaments, which bunch up in
minute loops as the abdomen
descends, and are beaten down
smooth by the spinnerets.

Our widely distributed Latro-
dectus mactans! quite resembles
Tepidariorum in cocooning habit;
but its ovoid cocgons are larger,
being a full half inch at the longer
axis, and somewhat more spheri-
Fie. 107. Cocoons of Theridium tepidariorum, hung in her eg] jn shape. She makes at least

snare. (About natural size.)
as many as four or five cocoons.

Theridium serpentinum Hentz? is one of our common Lineweaving spi-

ders, whose snares are found in dimly lighted cellars and in rooms aban-
} doned or rarely used. In the angle of a window or wall the
Therid- mother spreads her snare of intersecting lines, and establishes
aris herself at one end thereof, always well towards the top. In the
course of time she succeeds in thickening her dwelling place by

added threads, until it has formed a sort of shelter of lies much more
closely set than those of the rest of the snare. In the neighborhood of
this dwelling place and on
a line therewith, or just a
little above it and to one
side, she spins several co-
coons, in number four or
five usually, but sometimes
as many as eight, as shown
in the figure. (Fig. 108.)
They are little white, oblong
or flask shaped flossy balls,
about quarter of an inch in
diameter, in the centre of
which the eggs are depos-
ited. In the delicateness and
scantiness of the enveloping
tissue, this cocoon resembles

= . > Fic, 108. Cocoons of Theridium serpentinum in site at top of
Steatoda borealis and Phol etCacaaes (Snir tee)

cus phalangioides. The eggs
are distinctly seen through the silken envelope. When the spiders are
hatched they hang for a little while in clusters like minute swarms of

= ; =
1 Lathrodectus formidabilis Walck. See also Vol. I., page 274.
2T am not positive as to the identity of this species.

GENERAL COCOONING HABITS OF SPIDERS. 113

bees upon the adjoining lines, and soon thereafter distribute themselves, as
is the custom with Theridioids generally, to surrounding points, where
they construct webs like the mother’s.
Another Theridioid spider, whose specific name is unknown to me, spins
a similar snare in like localities, and deposits therein several eggs, almost
resembling those of Serpentinum, except that they are of a yellowish brown
color and more spherical in shape. They have a pretty appearance as they
hang amidst the crossed lines in the dusty and dusky sites which the
mother frequents.
Among Lineweavers making several cocoons is Argyrodes trigonum.
The species belongs to a genus quite famous for its habit of invading the
snares of other
Argy- __ species, particular-
rodes ba: ly those belonging
gonum. : ‘
to its own tribe of
Retitelariz, and those Orb-
weavers that make com-
pound snares and thus af-
ford a suitable dwelling place
in the labyrinth or maze of
crossed lines. I haye ob-
served this habit in Trigo-
num, but haye more fre-
quently found it in its own

WW Wy)
Vi ZA LL—__ YY We
Lita fone ae TA

Z3
the —— hy
SS Seed)

Wee a

Ald

alas / :

snare. It is an awkwardly LS

shaped creature, and its odd SS NGS! BZ v=
appearance is increased by — = - Ve
its habit of bunching its FiG. 109. Argyrodes trigonum in her snare, with three

‘ cocoons. (Natural size.
legs together, and hanging op

upon a few crossed lines in its snare, as represented at Fig. 109. In this
position it looks not unlike a trussed fowl in a green grocer’s stall.

Her cocoon is a pretty pyriform hanging basket, about one-fourth inch
in length and one-eighth in thickness, composed of stiff yellowish brown
silk. The upper part is a cone, rounded or tapering well to a point, at
which is attached a stiff white cord, by which it is fastened into its place
among the crossed lines of the snare. The lower part of the basket termi-
nates in a short projection from the middle. (Fig. 110.) The mother
makes several cocoons; I haye found as many as three (Fig. 109) hanging
within a snare, at one time, all of which were doubtless made by the little
mother. The cocoons are suspended by long, stout cords. When this
hanging basket cocoon is opened the eggs are seen loosely deposited in the
midst of a little puff of flossy silk. I sometimes find at the bottom of the
cocoon a little hole, through which evidently the young have escaped after
hatching.

114 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Ero thoracica, a spider common to Europe and America, weaves a
small flossy cocoon, containing about twelve eggs, which it suspends to
various objects, grass, twigs, etc, by a long thread. (See Fig.
eave Sao 111.) Emerton has found this spider in New England; it is
racica. 5 : .
common in winter under leaves; he has also seen cocoons like
those of the European Ero as above described, but has not identified them
with the American species, whose web he has not seen.
Ce rf) Something similar to this, but a little more complicated
, in structure, is the pretty orange brown cocoon of Theridium
frondeum, which is found suspended ordinarily to a stretched,
stiffened cord among rocks or leaves. It appears particularly
to love shady positions; at all events, I have found it most
frequently among rocks on banks of streams, in ravines, or
moist and secluded spots, as far west as the hills of Eastern
Ohio. It is about an eighth of an inch long, but varies some-
FiG.110. Cocoon What in length.
e eee On opening this pretty little cocoon of Theridium frondeum,
"it is found to be filled with a delicate white silken floss, in
the midst of which the eggs are deposited and the young will be found
after hatching. The number of eggs appears to differ a good deal. I
have counted as many as twenty-five in one cocoon, but many less than
this in others. The flossy padding is compacted well towards the top of
the cocoon, and passes out of a round opening therein in the shape of a
carded cord of straight lines of white silk, which gradually diminishes
until it is compacted into the stiff white cord by which the
whole is suspended. A curious arrangement is shown in the en-
larged figure of a dissected cocoon (Fig. 113), which is used by
the spider as a cap to the open top of her cocoon. In other words, the
cocoon, instead of being a continuous piece of spinningwork
gradually tapering into a point, as it appears at the first
careless glance, proves to be composed of two pieces. First
is the principal part or sac, which has already been referred
to as having a round opening at the summit. Fitted di-
rectly upon this, but easily separated from it by pulling, is
a conical cap, which surrounds the lower part of the sus-
pensory cord already described. This cap, by manipulation
under the microscope, can be unraveled so that it is seen ol
to have been formed by lapping the yellowish cocooning wi
thread, of which the main sac has been woven, around and ‘r'pro thoracten
around the base of the suspensory cord, after that has been slightly enlarged.
spun. (See Fig. 114.) The whole cocoon forms a very beau- ee ar
tiful and delicate bit of spinningwork, and shows considerable deftness
in weaving on the part of its little architect.
Somewhat similar to this is the cocoon of Ero variegata (Theridium

Cocoon
Structure

GENERAL COCOONING HABITS OF SPIDERS. 115

Fie, 112, Fic. 113.

Fic. 116,

‘ale pit ye)
Ae % errs oe
Fic. 114,

Fic. 112. Cocoon of Theridium frondeum, magnified. Fic. 113. The same, natural size, suspended
in natural site. Fig. 114. Cocoon of Argyrodes trigonum, much enlarged, to show the structure.
Fic. 115. The spiral thread on the cap and stalk. Fic. 116. Cocoons of Ero variegata, twice
natural size, (After Blackwall.)

116 AMERICAN SPIDERS AND THETR SPINNINGWORK,

variegatum), a little spider not uncommon in England, which would arrest
the attention of even an indifferent person. It is of an elegant pear shape,

fotmed of a strong yellow brown silk network, and attached by
ag a long elastic stem of the same material to stalks of dead grass,
variegata .. ° ; :

sticks, or other substances in shady places.1 It is often placed
on the under side of rocks, stones, etc. The envelope is double, an inner
sac being formed of soft pale brown silk, loosely woven and enclosed in a
coarse covering of dark reddish brown threads, which unite and form the
stalk. The diameter of the cocoon is about one-eighth inch, and the length
of the stalk is from one-tenth to one-half an inch. The cocoon contains
about six brown eggs. The mother is one-eighth inch long.”

Theridium pallens is a small English Lineweayer, about one-tenth of
an inch long, that makes a cocoon a little longer than herself, containing

about twenty pale yellow eggs. It is white, of a close, fine text-
Therid- yre, and somewhat pear shaped; with several conical prominences
SeTLew: disposed in a circle around its greatest cireumference. (Fig. 117.)

The sexes pair in May; the cocoon is formed in June, and is
found on shrubs and bushes, on heaths or near woods.®

A Lineweaver which I find in our fields, and which I take to be The-
ridium differens (Fig. 118) makes a globular cocoon, about one-eighth inch
in diameter, a little larger than herself, which she hangs within
her snare of crossed lines that may often be found spun in the
interspaces formed by bending down the top and edges of a leaf.
. The cocoon is rather flossy in its exterior. The cocoon of the
Fic. 117. Co. Same species, or one closely resembling it, I find within the con-
coonofThe- cavity of a leaf, formed by pulling the pointed ends inward, as
ridium pal- = 5 . : f
lens. x 4, at Fig. 119. The hollow is overspun with intersecting lines
(idee) Which form the spider’s snare and dwelling, and the lodging

place for her egg sac. The little mother is usually found near
her cocoon, which she often clasps with her legs, especially at any suspicion
of danger. She is apt to lug it about from point to point within the leafy
bivouac thus prepared.

A similar cocoon made by a Theridioid spider which I am unable to iden-
tify is represented at Fig. 120, The cocoon was a globular one, resembling in
appearance the last two described, but was hidden underneath a stone within
a little nest of characteristic spinningwork, but which on one side was protected
by a semicircular wall of clay, mingled with silk and attached to the under
surface of the stone. In this respect, the cocoon and cocoon nest resemble that
of Neriene rufipes and others of this genus as described by European writers.

Theridium lineatum Clerck is found among our American fauna, Emer-
ton* having taken it in Massachusetts. It is common in Kurope; its cocoon

1 Cambridge. 2 Staveley, “ British Spiders,” page 156.
8 Blackwall, “Spiders of Great Britain,” page 195. 4 New England Theridide, page 16.

, 2 ta

GENERAL COCOONING HABITS OF SPIDERS. 117

Vie. 119.

ic. 120,

Fie, 121. Fig. 122.
Vig. 118. Snare and cocoon of Theridium differens. Fie. 119. Cocoon of Theridium differens in a
leafy tent. Tic. 120. Theridioid cocoon under a stone. Fie. 121, Cocoon of Theridium linea-

tum in natural site on a leaf. (After Blackwall.) Fic, 122. Theridium varians and cocoons. (After
Blackwall.)

118 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

is formed in July and is round, one-fourth inch in diameter, and bluish or
greenish blue in color. It is loosely covered with silk and fastened to the
lower side of a leaf, the edges of which are bound together, so as to pro-
tect it.! (Fig. 121.)
Theridium varians pairs in June, and in July the female constructs
several globular cocoons of dull white silk, of a loose texture, the largest
of which measures about one-seventh of an inch in diameter.
Therid- They are attached to objects situated near the upper part of the
aes var snare, and contain, according to their size, from twenty to sixty
‘ spherical eggs, of a yellowish white color, not adherent among
themselves.2 (Fig. 122.) Withered leaves, dried moss, and particles of
indurated earth are generally disposed about the cocoons.* This habit,
which, as will be seen further on, prevails largely in other families, appears
to have but slight hold upon the cocooning instincts of the Lineweavers.
The little bronze colored spiders
belonging chiefly to the genus Eri-
gone, weaye their cocoons within
the balled mass of intersecting lines
which form their snare and abode.
I have seen numberless examples
of these webs, made manifest by the
morning dews along the Delaware,
shining over the entire external foli-
age of a large spruce tree from top-
most to lowest bough. Again, they
will be seen with other Theridioid
webs, glittering in the slanting sun-
light on myriads of bunched grass
tops, timothy heads, and weed tops.
Some species of Erigone make a lit-
Fic. 123. Cocoon of Erigone (?) suspended between tle balled cocoon similar to those
twigs of pine. eels .
of Theridium first described, and
similarly held within the snare. Another form of cocoon which I attrib-
ute to a spider of the same genus is a minute white button shaped or
double convex bag, from one-sixteenth to one-eighth inch in
diameter. It is suspended at the converging points of four lines
(Fig. 123), which are attached to the surrounding foliage, as in the ex-
ample shown of a cocoon hung between two twigs of pine, near a Theridioid
web, in which an Erigone was ensconced.

Hrigone.

' Staveley, Brit. Spiders, page 140; Blackwall, Spi. Gt. B. & L., pl. xiii., Fig. 111.
? Two small round cocoons are seen within the tent like structure in the cut, but in this
case, as with the figure of Theridium tepidariorum, as heretofore remarked, the artist has
erred by drawing in a sheeted tent instead of a structure of open lines. ~

* Blackwall, Spiders Gt. B. & I., page 189, pl. xiv., Fig. 120, d.

et. ee ae ee eS SS
2 a af ‘ — bs

GENERAL COCOONING HABITS OF SPIDERS. 119

Theridium zelotypum makes a flattened cocoon of soft silk, which she
establishes within her pretty nest, that has heretofore been described (Vol.
I., page 317) as a silken, bell shaped tent thatched with the leaves

Therid- of spruce, balsam, hemlock, or other plant on which it is built.
ee Within this the young are hatched, and here for a while after
‘their exode mother and young may be found dwelling together.

A like habit is possessed by the English nest making spider, Theridium
riparium, whose most remarkable nesting architecture is described Vol. L.,
page 318. The mother makes several yellowish white, round cocoons
about one-eighth inch in diameter.

Theridium sisyphum also shelters her reddish brown cocoons in a silken
tent which hangs in her snare, and is sometimes strengthened by the intro-
duction of dried leaves and other extraneous matter.?

Another English spider, Theridium nervosum, also* forms a silk lined
nesting tent, thatched with bits of dead leaves, flowers, or other particles,
including the débris of slaughtered insects. Within this tent the mother
spins a little round green cocoon, containing yellowish white eggs. The
cocoon is one-eighth inch long, the spider herself being one-sixth inch.
The mother is usually to be found in an inverted position, embracing her
treasure and covering it with her body.

It is probable that all the nest weaving species of Retitelariz place their
cocoons within their nests, in which habit they substantially agree with
their congeners, who suspend their cocoons upon the thickened cross lines
which form the resident part of their snares.

I have never been able to determine satisfactorily from observation the
cocoons of our common species of Linyphia, but the Linyphia montana of
Europe makes a flattened white cocoon, which it usually conceals
underneath a stone, remaining with it and guarding it with the

greatest care.4
Linyphia marginata, one of our most common American spiders, is also
a European species.© It pairs in May, and in June the female spins one
or two lenticular cocoons of white silk of a loose texture, which are at-
tached to withered leaves or other objects situated near the snare. The
larger of these cocoons measures half an inch in diameter and contains
’ about one hundred and forty spherical eggs of a palish yellow color, not

agglutinated. ®
The English Linyphia erypticolens is remarkable for the habit of car-
rying her cocoon fastened by threads to her spinnerets. It is globular and
of a diameter equal to the whole length of the mother, is formed in June

Linyphia.

1 Staveley, British Spiders, page 152. 2 Idem, page 148. 8 See Vol. I., page 317.
4 Staveley, “British Spiders,” page 165.

® Wqual to L. montana Sund., L. resupina Walck.

® Blackwall, Spiders Gt. B. & L., page 215.

SS ae

120 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

or July, is a pale brownish color, containing brown eggs. It resorts to dark

and damp places, as cellars and the under surfaces of stones. It is cer-
tainly remarkable to find a Line-
weaving species thus approximat-
ing the Citigrades, from which it

- so greatly differs in other respects,
in the manner of caring for the
egcoon.

But in this habit she is not
alone among her tribe. Theridium
carolinum forms in June a round
white cocoon one-tenth inch in di-
ameter, which she carries attached
by threads to her person! A
pretty little Theridioid, Steatoda
maculata (Theridium maculatum
Linn.), is also said to carry about
its egg cocoon suspended between
the legs, and only relinquishes it

Fic. 124. The mother Pholeus hanging in her snare, when force is used, regaining it
with cocoon held in her jaws. quickly if possible.

The cocoon of Pholeus phalangioides, which is perhaps the very simplest
in structure of all this tribe, and I may add of all the tribes, is simply a
Ph gauzy covering which encloses the eggs, the whole being gathered

olcus. * +e ae : ie

into a globular mass. This is held by the spider within her

jaws as she hangs in her ordinary position within her straggling web of
intersecting lines. In this portage of her egg case Pholeus approaches the
habit of the Citi-
grades and 'Tunnel-
weavers. (Fig. 124.)
~—— Seytodes thora-
cica Latr. (Seytodes
cameratus Hentz)
has been found by
Mr. Emerton, in
New England, as a
house spider, which
he supposes has
been imported from
Europe. European observers note that this spider carries her cocoon under
her breastplate, in which position it is not secured by silken threads, but
is held by the falées and palpi. In this habit it resembles Pholeus, with

lic, 125. English Pholeus phalangioides, with her cocoon. (After Blackwall.)

- - :
' Staveley, “ British Spiders,” page 141.

GENERAL COCOONING HABITS OF SPIDERS. 121

which it is closely allied structurally. It is found in houses, upon walls,
etc., In warm situations, It is described as slow and deliberate in its
motions, displaying somewhat of the action of a gnat in lifting
and poising its leg in the air when walking. The whole char-
acter of the aranead is mild and quiet. The poison fangs are
so feeble as to be of but little use in seizing its prey, which office is chiefly
performed by the maxilla. When taken, Scytodes offers no resistance and
attempts no flight, but, feigning death, resigns itself quietly to its fate.?
This tribe embraces the singular genus Walckenaéra, some of whose
species have the eyes placed upon little turret like elevations of the ceph-
alothorax. Their habits have not been carefully studied, and
their cocoons are little known. One European species, Walcke-
naéra acuminata, makes a cocoon flat on one side, rounded on the
other, about one-third inch in diameter, and composed of slightly woven
white silk. It is found in autumn on the under surface of stones and

Scytodes
thoracica.

W alck-
enaera.

Fic, 126. ] Fig. 127. Fie. 128.

Cocoon of Agalena nevia, spun upon bark.

I'iG, 126. Appearance of exterior, covered with brown sawdust. Figs. 127 and 128. Views after the outer
coverings have been removed.

other objects.2 Our American fauna has a number of closely related rep-
resentatives of this strange genus, which are relegated by Emerton to vari-
ous genera,® and it is probable that their cocoonery nearly resembles that
of the above species.

ET.

The most common Tubeweaver in the Eastern States is probably the
Speckled Agalena, Agalena neevia. Its funnel shaped nest, with its broad
sheeted top spread over the grass or hedges, or stretched in mis-

Tube- cellaneous sites, is one of the most familiar objects in our land-
weaving ee arnt
Agale- °°@Pe. Its cocoon is attached to some surface, as the leaf of a

nine. tree, a rock, or the under surface of a loose bit of old bark. In
this position Agalena spreads a circular patch a half inch or

* See his “New England Theridiide.”

eS ae

2, AMERICAN SPIDERS AND THEIR SPINNINGWORK.

again with a thin sheet, upon which the mother overlays a wadding of
sawdust or pulverized bark gnawed from the surrounding surface. In
the absence of such materials, the upholstery consists of any available
fibre furnished by the particular site. The whole is then overspun with
an exterior covering. It is thus one of the most elaborate of known
cocoons, and apparently is as well calculated to preserve the life concealed
within as any spinningwork that could be wrought by aranead spinning
organs.

When Agalena cannot conveniently obtain sawdust and like material for
the upholstery of her cocoon, she will overspin her eggs without such pro-

tection. For example, a female of this species was observed upon
Uphol- the window of a chicken house, with a pretty tubular snare hung
eae against the frame, and two cocoons woven upon the glass near
mitted. |_°
by. These were simply eggs of a pinkish hue, covered over with
silken spinningwork and no upholstery added. I have also found Aga-
lena’s cocoon woven upon the under side of a leaf, in which position it
contained no upholstery, and, indeed, quite resem-
bled the cocoon of an Epeiroid spider spun in
like situation, I suppose that in this case, as also
in the preceding one, the difficulty of gnawing off
the tough, green fibre of the leaf and branches, or
oe te tse Be (ooo one tie painted wooden frame of the window was an
the mode of uniting to the obstacle which prevented the mother from pursu-
opposite surface by a stalk. A A A .
ing her usual habit. Perhaps, indeed, it requires
the suggestion of near by and available material, like that of bark or
decayed wood, to induce this additional upholstering protection of the
cocoon.

On the other hand, a female of this species, which I kept within a
glass jar, having made her cocoon, proceeded to collect from the bottom of
the jar bits of débris of various sorts, which she placed upon it in the
usual position. There were only a few of these particles, not enough to
be of any value for the protection of the enclosed eggs, even if there had
been any exposure to danger under the circumstances. Of course, it could
hardly have been expected that this mother would understand that her
offspring, by reason of the situation within a glass jar, would be safe
from the enemies which usually assail the eggs of the species in natural
site.

Sometimes the cocoon of this species, when spun upon a loose piece of
bark, will have a thick stalk spun across to the opposite surface of the
tree to which it is united by a circular patch of thick silk. (Fig. 129.)
A like arrangement is found when the cocoon is woven up against the
lower side of a stone, the exterior or under part being then carried down-
ward by a stalk to the earth. This is not a common method, however, and
I can think of no good reason for such a variation.

GENERAL COCOONING HABITS OF SPIDERS. A.B}

Agalena labyrinthea of Europe resembles in its general habits the Aga-
lena nevia of America. According to Walckenaer the female makes her
single cocoon in the month of August, which she encloses within
Agalena 4 huge purse like web full of soil and vegetable detritus. When
ISB the web is removed, the cocoon is seen to be about the size of the
rinthea. : : y
end of one’s thumb, and woven of a fine silken tissue enveloped
by clods of earth. Next to these is another envelope of silk, and then,
finally, particles of soil so strongly adhering to the cocoon that they cannot
well be separated. When the cocoon is opened, it is found to be formed
of a thick, tough web. On the exterior it is beautifully white and perfectly
polished. It contains as high as one hundred and thirty-four eggs of a
greenish yellow color.?
The well known cellar spider, Tegenaria derhamii,? which is widely

Fie, 180. Fie. 181. ,

Fic. 180. Snare of Tegenaria derhamii in a cellar window, with three cocoons suspended thereto.
Fic. 181, One cocoon, natural size.

distributed over both hemispheres, conceals her eggs within a flattened
ball or hemisphere of soft silk, somewhere in the neighborhood
of her snare. Sometimes this is suspended by threads to the
snare itself (see Vol. I, page 2389, Fig. 221), or again is attached
directly to it, and the envelope interwoven with the fibre of the web, so
that it has much the appearance of a rounded button upon a coat. Fig. 130

Tege-
naria.

' Walckenaer, Aptéres, Vol. II., page 22.

* T have supposed that the Medicinal spider of Hentz, Tegenaria medicinalis, is identi-
cal with this species, and have so used the name in Vol. I. Mr. Emerton, however, in a
recent paper, declares Hentz’s Tegenaria medicinalis to be a Ccelotes, and separate from T.
derhamii. He classifies as Coelotes medicinalis the spider that I have heretofore considered
Hentz’s Tegenaria persica. See Trans. Conn. Acad., Vol. VIII., 1889-90, New Eng. Spiders
of the Families Drassidee, etc.

124 AMERICAN SPIDERS AND THEIR SPINNINGWORK.,

represents the snare and cocoon of one of these spiders. The mother
was hidden within a curtained screen or tower newly spun. On the beam
just above the snare hung two cocoons. They were attached above and on
the sides to the beam, and in front and on the sides to the flap of the
snare. ‘Their position was such that they were just above and in front of
the door of the den. One of them was coyered with black particles of
dust. They were about half an inch in diameter. Figs. 182 and 133 are
views of the manner in which the cocoons were suspended. One often
finds these cocoons woven into the texture of abandoned snares in cellars
and outhouses. Fig. 130 is a sketch of such a web hanging in a window
of my church cellar. The pouch like snare stretched upward to the window
roof, and at the bottom, on either side of the tube or tower, three button
shaped cocoons were inserted. They were still white when sketched in
midwinter, although the web was
‘ \wittessss much soiled with the cellar dust
ea \) and soot. I do not know that
all three cocoons were made by
one mother.

Coelotes medicinalis (Tegena-
ria persica Hentz) usually spins
her cocoons on or near her snare.
I have found in one snare two
globular cocoons covered with bits

Fic. 182. Fic. 188,
Fics. 182, 183. Suspended cocoons of Tegenaria derhamii. of clay. One contained round

(Natural size. mee
bie arr: whitish eggs; the other had liy-

ing spiderlings with white cephalothorax and greenish abdomen.

Agreca brunnea’ is an English species. The sexes pair in May,
and in the month of June the female constructs an elegant vase shaped
cocoon of white silk, of a fine compact structure, attached by a short foot
stalk to rushes, stems of grass, heath, or gorse. It measures about one-
fourth inch in diameter, and contains from forty to fifty yellowish spher-
ical eggs, enveloped in white silk, connected with the anterior surface of
the cocoon, contiguous to the foot stalk. Greatly to the disadvantage of
its appearance, the cocoon is smeared with moist soil, which when dried
serves to protect it from the weather, and, as an additional security, the
extremity is closed and directed downward.? In the illustration (Fig. 184)
the uppermost cocoon is shown as it is first spun, the two lower cocoons
as they appear when plastered. Another drawing (Fig. 135) of this beau-
tiful cocoon, which has attracted the attention of all English araneologists,
is taken from Rey. Pickard-Cambridge. With it is a similar cocoon of
an English congener, Agroeca proxima (Fig. 136), woven like the former
species upon a twig of heather.®

1 Agalena brunnea Blekw. “ Blackwall, Spid. Gt. B. & I; page 160, pl. xii., Fig. 102.
* Spiders of Dorset, Vol. I., pl. ii., Fig. 7.

— a

GENERAL COCOONING HABITS OF SPIDERS. 125

Coelotes saxatilis makes a cocoon half an inch in diameter, containing
yellowish white eggs. The external case is partly plastered with
Celotes earth. (Fig. 137.) Textrix lycosina has the same habit of pro-
we .. tecting her cocoon, which is usually woven to the under side of
extrix. ae : :
a stone near her tubular hiding place. It is white, flattened,
and about one-fourth inch in diameter. !

According to the Swedish naturalist Clerck? the eggs of the Water spider,
Argyroneta aquatica, are round, of a saffron yellow color, contained within

a globular silken cocoon, which occupies about one-fourth of the
ried subaqueous maternal cell. (Fig. 138.) The female remains con-
aquatica, Stantly near it, keeping her abdomen in the interior of her hab-

itation, and the fore part of her body in the water. The figures
of this cocoon (Figs. 189 and 140) are from Blackwall,® and represent a
hemispherical or disk like object resembling cocoons made by many terres-
trial Tubeweavers, especially the Clubionide. Argyroneta’s cocoon presents
the appearance of having been woven against a flat, solid surface, or per-
haps the silken walls of the cell. Other naturalists represent it as being
swung like a hammock across the cell, somewhat in the fashion of the
cocoons of various 'Tunnelweavers hereafter described.

Blackwall’s description of the cocoon, its site, and preservation is as fol-
lows: Argyroneta aquatica habitually passes the greater part of its life in
the water, not only pursuing its prey in that liquid, but constructing be-
neath its surface a drum shaped cell in which is placed its cocoon of white
silk of a compact texture and lenticular form, containing from eighty to
one hundred eggs of a yellow color, not agglutinated together. This is well
supported in a vertical position, the open part being directed downwards
by lines of silk connecting it with aquatic plants, and as it comprises a
considerable quantity of atmospheric air, the spider can at all times occupy
it without experiencing the least inconvenience. In swimming and diving
Argyroneta assumes an inverted position, and is more or less enveloped in
air confined by the cireumambient water among the hairs with which it
is clothed. The supply is always more abundant on the under than on
the upper part, in consequence of the greater length and density of the
hairs distributed over its surface.

Passing into the large and varied family of Drassids, we find a sub-
stantial uniformity in the general shape and structure of their cocoons.

These are usually lenticular or button shaped (plano convex) ob-

es jects woven against some solid surface in the vicinity of the
ras- : . . .
oie tubular nest or ordinary haunts of the species. The covering is

a close textured silk, as stiff as parchment. The circular piece
attached to the surface is of similar composition, and the eggs are

1 Blackwall, Spid. Gt. B. & I., pl. xii, Fig. 109. 2 Aran. Svecici., page 149.
’Sp. Gt. B. & IL, pl. vili., Figs. 87 g, h.

0 i i ee

_— — |

126

(After Cambridge.)
daubed on the surface.
(After Cuvier.)
Blackwall.)
show the flat bottom.

Fic. 134. Cocoons of Agalena brunnea, attached to moss. Slightly enlarged. (After Blackwall.)
135. Cocoon of Agroeca brunnea.

Fic. 189. Cocoon of Argyroneta aquatica, front view.
Fie, 141.

AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Fig, 136.

Fic. 140,

Fig, 142.

Fie. 141.

Fie,
Fie, 136. Cocoon of Agreeca proxima, attached to a sprig of heather.

Fie. 137. Cocoons of Ceelotes saxatilis, natural size, with particles of earth
(After Blackwall.) Fia. 138. Subaqueous cocooning nest of the Water spider.

Iie. 140. Side view. (After
Two Drassid cocoons woven against a board.

Fie. 143. Cocoon of Clubiona tranquilla (probably), woven upon bark,

Fig. 142. One detached, t

ee ee es

GENERAL COCOONING HABITS OF SPIDERS. 127

commonly deposited inside, without any or with only a little flossy pad-
ding. The exterior is frequently plastered more or less freely with mud
or the detritus of decayed wood.

Clubiona tranquilla makes a hemispherical or button shaped cocoon,
which is attached to various surfaces, as of rocks, bark, or boards. (Fig.
143.) One female confined within a jar for observation spun her co-
cgon upon a little twig placed for her
convenience within the vessel. As first
completed by the mother the external
covering was pure white silk. But, fol-
lowing her maternal instinct, she de- Via, 144. Cocoon of Clubiona tranquilla, woven on
scended to the earth upon the bottom Sey a eg ot
of the jar, collected pellets of mud between her mandibles, carried them. up
to her cocoon, and daubed the surface over in little ridges until the whole
was quite mottled with the plastered mud. (Fig. 144.)

Sometimes the Drassid’s cocoon is contained within the tubular domi-
cile of the mother, and this again will be overspread with a tent of deli-

‘cate texture, as in the case of the Parson spider, Prosthesima ecclesiastica
(Herpyllus ecclesiasticus Hentz). (Fig. 145.)

The Parson spider is a quite large species one half inch long, with a
black body, marked along the thorax and dorsum of the abdomen with

decided circular and oblong patches of white, to which peculiar

The markings it owes its specific name. Its habits are those of the
Parson Fi stage .
Spider Drassids generally, although it is not as sedentary as some others,

but wanders in search of prey. It is commonly found upon trees,
fences, etc., near some recess or opening into which it may retreat. Like
some of our common house Theridioids, it is fond of taking refuge under
the projecting parts of outhouses. In winter it is found wrapped in a
thick sheeted tube of silk under the bark of trees and like situations. It
is active in its movements, and prowls for its prey. It makes its cocoon
early in June. This is com-
posed of several layers of pure
white silk, between one of
which particles of dust are
Ss placed and quilted in with
sade ED 3 ea, y: spinningwork. I have found

ee aie ™" ~=6the chippings of the carpenter

bee among these particles.

An interesting and rather pretty little Tubeweaver, which appears to be
Micaria aureata, the Herpyllus aureatus of Hentz, conceals its cocoon with-
in a double tent. (Fig. 146.) The cocoon itself is a small, button shaped
object, containing a few brownish yellow eggs. The example illustrated in
the figure was spun within the angle of a wall, and covered over with a
tube such as the spider usually spins for a dwelling place. Openings were

° f+ = were oy Se

128 AMERICAN SPIDERS AND THETR SPINNINGWORK.

left at either end of this tube. Above the whole, and quite encompassing
it, was woven a large tent several times the size of the first tube, and
composed of spinningwork whose threads were quite closely placed, but of
so thin tissue that one could see through it without any difficulty. A
large opening appeared at one end of this external tent, but whether it
was left of purpose for a door, or, more probably, was the result of acci-
dent, I could not determine.

Among the Drassids which I have found in Colorado is a species of
Gnaphosa, which I took under a stone on the summit of the Snowy Range.
It was dwelling in a little tubular nest. This species, according
to Emerton,! is found all over New England, from the White
Mountains to New Haven. Professor Packard found a female
with a cocoon of eggs on Gray’s Peak, Colorado, over eleven thousand feet

He 1
PUMIAUIMCCTTELEAL dull a mn
P poo SG \\ ae a
ae ol
W \ I) \ \
Nc RA

Gna-
phosa.

SONNY)

=~ ne
iN eee WA a
AUN \

Ml:

ite
AY

Bi ZW)
Wa P if Py RY
NN Ps Ui Hp

Fic, 146. Cocoon of Micaria aureata within an interior and exterior tent.

high. It thus has a remarkably great geographical as well as vertical dis-
tribution. The spider lives under stones and leaves. The cocoon is white
and flat, with its diameter as great as, or greater than, the length of the
spider. Emerton says that the female stays near her cocoon, but makes
no nest. I would have expected her to make her cocoon within her cell.

Some of the Drassids, like the Agalenads, protect their cocoons by com-
pletely enclosing them in cases of mortar. Among these is a species sent
me for determination by Mr. F. M. Webster, assistant entomolo-
gist of the State of Illinois, through whose intelligent interest
the remarkable facts concerning this spider have thus been made
known. Mr. Webster has found these mud cocoons throughout
the whole range of Illinois, a State of great longitudinal extent. T'wo
balls om Routhern Illinois are larger than the others, and composed of

1 New Bog Dagettins Trans, Conn. Acad, Vol. VIIL, page 13.

Mud
Encased
Cocoons.

PLATE Il.

MIMICRY OF ENVIRONMENT. TRAPDOOR SPIDERS.

1, BURROW WITH DOOR OF DRY OLIVE LEAVES, CLOSED. 2, THE SAME, OPEN.
8, 4, 5, TRAPDOOR COVERED WITH MOSS,

GENERAL COCOONING HABITS OF SPIDERS. , 129

yellowish earth or clay; but balls from Central Illinois are made out of
the rich black soil common to the prairies. They vary in diameter from
one-half to one-fourth of an inch. (Figs. 147, 148.) From most of them
a slight silken cord protrudes (Figs. 147, 148, 153, 154), by which they are
often found attached to the under side of a board or stone. The cord is
sometimes thickened into a cup shaped patch at the point of attachment,
and is occasionally composed of several threads. When these mud balls
are softened in water one is able to open them, and in some cases the
mud peels off in little layers like the skin of an onion, indicating that the
method of structure is to plaster a thin coating of mud upon the entire
cocoon, and add successive layers, which likewise cover the whole surface
before another layer is begun. It is evident that no little mechanical skill
is involved in such even distribution of the mortar.

In the centre of the mud ball is found a cocoon of delicate structure
and pure white color (Figs. 151, 152), within which a few eggs are depos-
ited. This can be lifted out of its matrix, leaving the round

an concavity smooth and well defined, as shown at Figs, 149, 150.
Renee The stock of the cocoon is carried at one point entirely through

the mud ball, and issues at the surface in a thin cord whose
use has been alluded to above. This stalk or suspensory cord is, of course,
spun before the plastering begins, and is covered over gradually, an act
which must require delicate manipulation.

By keeping some of the cocoons in a moist condition, I was able to
hatch from one, May 80th, a brood of about thirty lively young Drassid
spiderlings. They apparently belong to the genus Micaria, and I therefore
named the species Micaria limicune,' although with much hesitation, as
it is difficult to determine species from young spiders.

These mud balls in external form closely resemble the spherical mud
egg nest of the wasp Eumenes, which I have often found attached to the
stalks of weeds, grasses, ete., in the neighborhood of Philadelphia. (Fig.
156.) It is certainly interesting to observe that this habit of concealing
the future progeny within a globular cradle of mud belongs to a spider
as well as to a wasp, and to note how maternal solicitude finds expression
in like forms among widely separated orders.

Limicune appears to be much subject to the attacks of hymenopterous
parasites. Mr. Webster found parasitic ichneumon flies in some of his

boxes, which had evidently crawled out of one of the mud balls.

Limiou- Some of the balls seen by him had openings in the side about
See one millimetre in diameter (Fig. 148), from which evidently

the ichneumon had escaped, since it contained the stiff white
silken case commonly spun by the larva of this insect. I secured from
one of my specimens, in the process of hatching the spiderlings, two of

* Proceedings Acad, Nat. Sci., Philadelphia, 1884, page 153,

yl ae Tye —

a ae

ee ee a a ee 4

1380 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

152

148

155

Mud plastered cocoons of Drassid spiders.

Tias. 147-152. Micaria limicune. x 2. I'19s. 153-155. Unknown species from Alexandria Bay. X 2.
Fic, 156. Mud nest of a wasp Eumenes. |

—" =

ae iy

GENERAL COCOONING HABITS OF SPIDERS, 131

these flies, which were determined by the eminent hymenopterist, Mr. Ezra
T. Cresson, to be Pezomachus meabilis Cresson.

I collected cocoons somewhat similar to those of Limicunze near Alex-
andria Bay, New York, on the St. Lawrence River. They were

wld 7 attached by very loose spinningwork to the under side of stones,
rate : : ;
ee but the external case, instead of being mud, was a mass of ag-

Ball. glomerated particles of, old wood, bark, leaves, blossoms, shells
and wings of insects, etc., which were held together by a deli-
cate weft of threads. (Figs. 153, 154, 157.)

Two of these balls contained whitish cocoons similar to those in the
mud balls of Limicune. (Fig. 155.) Another had within it the charac-
teristic cases of some hymenopterous insect, containing dried pupe. A
very thin veneering of soil immediately enclosed the silken egg pouch,
but otherwise no mud plaster was used. I did not succeed in hatching
spiders from the specimens, and could not therefore determine that these
cocoons were made by the same spider that constructs the mud balls of
Illinois, but I am inclined to think they were made by
the same or a closely related species. 1

This habit of protecting cocoons with an armor of
mud and agglutinated rubbish of divers kinds, is widely
spread, and is, no doubt, quite cosmopolitan. It is pos-

_ sessed by several of the European species. ‘Teg-
sonia ek enaria agrestis is found under rocks, in which
cms position the mother attaches her large cocoon, gccoon of Micaria,
about half an inch in diameter, formed of a triple or armored with chip-
. > : . ae pings, soil, etc. 2.

quadruple envelope. ‘The first are thin, white, containing
a layer of sand and the débris of insects agglutinated together, followed
by a third envelope of beautiful orange red, which contains a loose wad,
a little compacted where the eggs are. The mother makes several cocoons,
which she abandons and leayes isolated, or which she encloses under a
single web, fine and transparent. In France these cocoons are found in

July. and August, chiefly in woods.?

The cocoon of Tegenaria emaciata, as described by Walckenaer, is formed
of a round mass larger than a good sized pea. ‘This mass is composed of
soil agglutinated and mingled with the detritus of the bodies of small in-
sects, as beetles, ants, and others. In the midst of this mass of earth is
placed the cocoon, of a beautiful orange yellow color, but not perfectly
globular, having the shape of a little flask.

The particles of earth which enclose this are held together by filaments
of silk, but are not enveloped by white silk, as is the case with Tegenaria
agrestis. The immediate envelope of the cocoon is a pellicle so compact

Iie. 157. Globular

1 McCook: “A Spider that makes a Spherical Mud Daub Cocoon.” Proceed. Acad. Nat.
Sci., Philadelphia, 1884, page 151.
2 Walckenaer, Aptéres, Volume II., page 8.

132 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

that one can tear it. Walekenaer found, August 20th, twenty-six spider-
lings perfectly developed enclosed within a cocoon. Each was about a
millimetre long, of a milk white color, the eyes not very distinct. In
another cocoon, found at the same period, he counted twenty-three eggs.
He saw no web near the tube in the neighborhood of the cocoons. Ex-
amples of the same mode of treating cocoons by the European Agroeca
brunnea have already been given.

While walking through the fields near the home of Mr, F. M. Camp-
bell, at Hoddesdon, Hertz, England, I
noticed a number of pretty, spherical
nests which had been formed by mass-
ing together spikelets of a species of
grass. A ball about the size of a hick-
ory nut, that is to say, one inch in di-
ameter, was thus formed. At first sight
I took this to be the work of lepidop-
terous larve, but upon plucking some
nests the spinningwork which bound the
spikelets together appeared to be spider
silk rather than that of a moth larva.

One of the nests was therefore opened
and proved to contain a species of Dras-
sid which I took to be a Clubiona.
Unfortunately, the specimens which I
had preserved for further examination
Fic. 158. Cocooning nest of Were lost, and I can only give this

an English Drassid, woven general identification. The species, as I
upon tops of grass. (From :
Nature.) remembered it, seemed much like our
American Clubiona pallens, or the Eng-
lish Clubiona hollosericea, The drawings (Fig. 158) were made
from specimens which I brought home, and upon careful ex-
amination prove beyond doubt to be the home nest of a spi-
der, woven upon heads of a grass somewhat resembling maize,
probably Leersia oryzoides Swz., or Rice Cut-grass. When cut
open, a hollow sphere of white silk is disclosed, which is the dainty cell
in which the aranead lived. A veritable fairy palace! Among the British
Clubionide, as described by Blackwall and Staveley, I can only find one
species, Clubiona erratica, whose habits would suggest such a nest as this,
The cocoon of this species is white and nearly round. The mother places
it in a nest, around which she forms a guard by binding together the
branches of firs or other plants in the midst of which she is placed. She
remains in the nest with her young.”

‘ ! Aptéeres, Volume IT., page 14. * Staveley, British Spiders, page 110,

GENERAL COCOONING HABITS OF SPIDERS. 133

This species, however, as described by the English authors, does not
correspond with my recollection of the inhabitant of the pretty nest which
I have noticed. It is possible that my memory may be at fault,
Cocoon i ‘ : :
Nest. and that this cocooning tent was prepared by the female of Clu-
biona erratica. American Drassids, as we have seen, make simi-
lar spherical nests, but I know none that thus hangs them to foliage.
The substantial agreement in cocooning habit between the Drassids of
America and those of Europe may further be seen by comparing the fol-
lowing descriptions of English species. The female of Drassus ater con-
structs a large white cell of close texture, usually in a hole in the earth or
under a stone. Within this, in the month of May, she places a plano con-
vex cocoon, which is attached by its flat side to the stone or other sub-
stance on which the cell is formed. This cocoon is white or slightly yel-

Tia. 159. Vie. 160. Fic, 161.
De Geer's sketches of Clubiona cocoon nests.
Fie, 159. On birch leaves. Fic. 160. Cocoon of the same. Fic, 161. Nest on an apple leaf.

lowish at first, but afterwards becomes yellowish in color. The female re-
mains on guard by her eggs.

The cocoon of Drassus sylvestris is white, of a flattened shape, and a
little less than one-third inch in diameter. It is formed in July
and concealed in the silken cell in a hole in the earth under
stones. The mother is usually found with her cocoons.

Drassus lapidicolens conceals herself in a cell formed between, the sur-
face of the earth and the under side of a stone, near which she spins some
threads, forming an irregular square. In this cell, in the months of July
and August, she places her cocoon, covering it with dead leaves. This, at
first, is in the form of a flattened sphere, but becomes nearly round when
the young are about to escape. It is white and about one-half an inch in
diameter. The mother remains with her young some time after the eggs
are hatched. The cocoons formed by the beautiful little Drassus nitens
are about one-sixth inch in diameter, hemispherical, and white. The
mother inhabits a tube which proceeds from the upper side of the cocoon."

English
Drassids.

1 See descriptions of Blackwall and Staveley.

18 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

I present in this connection two of the earliest published figures rep-
resenting the spinningwork of spiders of this family, both of them prob-
ably belonging to the genus Clubiona. They were made by that
pioneer araneologist, Baron De Geer. Fig. 159! represents a leaf
nest with the spider within it, woven on the imner surfaces of
birch leaves. This constituted the mother’s dwelling and the egg nest of
her cocoon. The mother remained with most of her body concealed within
her nest, but her fore feet were held outside ready to seize whatever prey
might pass by. Fig. 160 is the cocoon separated from the enclosing nest.
Fig. 161 represents an apple leaf within the concave inside of which is seen
a white cell spun by the female of Araneus pallidus Clereck (“Araignee
tapissiere”’), apparently a species of Clubiona. It serves as a dwelling
for the mother and contains also her cocoon, within which the eggs
are deposited and the young hatched. The nest was sketched July 25th.
It was opened and the spiderlings found within
with their mother. The mother showed no fear,
but stayed by her little ones closely, even during
the process of tearing open the nest for examina-
tion.?

The Dysderads form one of the most interest-
ing families of the 'Tubeweavers, and are especially
distinguished by haying six instead of eight eyes,
six spinnerets, and four breathing holes. In their
general habits they are closely related to the Dras-
sids, living in tubes or cells of silk formed under
stones in cracks and crannies of walls, fence rails,
old trees, and similar places. Our most common
FiG, 162. Snare and nesting tube species in this geographical province is Dysdera

di det bicolor.® I have found it in great numbers occu-
. pying numerous interstices between the stones of an old barn in Delaware
County, and in the interspaces between door jambs and window frames of
the wall.

Tubes of all sizes, from those of baby spiderlings to grizzled adults’, had
their outlet upon the wall surface, at which points the tube widened out
into a rectangular margin or flap, by which it was attached
to the wall. The species is widely distributed over the adjoin-
ing fields, in fences, ete., and the accompanying figure was drawn
from a huge walnut tree that stood solitary in a meadow. The trunk was
cleft by a longitudinal fissure twelye feet or more in length and from an
inch to two inches wide. The bark was stripped off along the edges of
this fissure, and within the crevice ten or twelve tubes were spun, extending

De Geer’s
Figures.

Dysdera
bicolor.

* Mem. des Insect., Tom. VIL, plate xviii., Figs. 8-9. * Idem, page 268, pl. 15, Fig. 16.
* Ariadne bicolor Emerton, New England Drassidee, page 38.

GENERAL COCOONING HABITS OF SPIDERS. 135

inward for two inches and more. ‘he silk of the tube was fine, but the
flap of netted work by which it was attached to either side was of coarser
fibre. (See Fig. 162.) The tubes were spun all the way
up the fissure to the fork of the trunk.

The spiders watch near the orifice of their tubes with
the first three pairs of legs directed forward, an unusual
position, as spiders usually have only the first two pairs
thrust outward.

The cocoon, containing twenty or thirty eggs, is placed
within the inner part of the tube in July and August.
Emerton! saw one in this position July 10th, and an-
other under a stone with a cocoon containing thirty-four
eggs. The English Dysdera hombergii spins her egg sac SEM aeiitten
within her tube in June; it is an oval cell, within which
are from twenty to thirty pinkish eggs loosely bound together. The cell is
slightly woven, and is covered with particles of gravel or other extraneous
matter. It thus appears that the cocooning habits of the genus as rep-
resented in Europe are the same as those of our American species.

In material sent me from San Bernardino, California, by Mr. Wright,
were cocooning nests of a peculiar type made by a species of Segestria,
which appears to be new, and which I have named Segestria
canities. (Fig. 163.) The species was determined from young
spiders found enclosed in some of the cocoons. Subsequently, I
received from the same section, through Mrs. Eigenmann, two mature
females, which enabled me to confirm my previous determination, and
thus to identify the cocoons which are here described. The species is
shown at Fig. 163, and a view of the face at Fig. 164.?

The mother Segestria spins a series of flattened disks, which are over-
laid one upon another like tiles upon a roof, and are bound by silken

threads somewhat after the fashion of Epeira labyrin-

thea’s cocoons. his series of cocoons is sometimes three

= inches or more in length. The examples sent me were
covered (apparently intentionally) with leaves, from the

plant upon which the string had been suspended, resem-

bling the leayes of spruce or hemlock. Along the entire

Fro. 164, View ofeyes length of one side of the cocoon string the mother
ee of Segestria had spun a silken tube, within which she dwelt. The
manner in which the string is suspended-is represented

in Fig. 165. It hangs within a maze of intersecting cross lines like the

Segestria
canities.

' Notes, Hentz’s Spiders U.S., page 22.

2 The spider is about three-eighths inch long; the cephalothorax brown, the abdomen
brownish yellow covered thickly with white hairs, which also strongly mark the cephalo-
thorax, suggesting its specific name. The legs are yellow, with brown rings at the joints
and a similar ring in the middle of the tibia.

1386 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Fig, 167.

FIG. 168,

Tra. 166.

FG. 165. Cocoon string of Segestria canities, with domicile tube alongside. Via, 166. The same,
side view, and cocoons covered with leaves. Ita. 167. Cocoons of wall loving Dictyna within a
sewed leaf. (Natural size.) I'r@, 168. The same, snare and cocoons on inside of leaf.

iid ills

GENERAL COCOONING HABITS OF SPIDERS. 137

.
webs of Lineweavers; is attached above to a strong thread, and is stayed
and balanced by various guy les along the entire length. On opening
the several cocoons of one of these strings I found (in one of twelve co-
coons) the first seven contained only the first moults or shells of the escaped
spiders; the next three, young spiders in successive degrees of advanced
growth; and the last two, eggs alone.

The exterior case of the cocoons is a light straw color or creamy white.
It is made of two saucer shaped pieces well woven together at the edges,
and is about three-eighths of an inch in diameter. Fig. 166 gives a side
view of a cocoon string, showing the way in which the cocoons overlap one
another.

Dictyna usually makes several cocoons, small flattened globes of pure
white, about one-eighth inch diameter, which are placed within the snare,
usually grouped near one of circular doors on which the web
lines converge. (See Vol. I., page 349.) When she spins her
web along a brick wall or like surface, the cocoons are fastened to the wall,
arranged along the angle or clustered together loosely. When the spider
makes her snare within a leaf, as she frequently does, the cocoons are
placed upon the leaf, protected, of course, by the enclosing cross lines.
The mother is found near her cocoons, though apparently not exercising
any special vigil upon them. She simply lays her eggs in the position
most convenient to herself. The edges of the leaf are sometimes drawn
well together (Fig. 167) and sewed in the prevailing aranead style; but
more frequently the edges of the leaves are simply bent over by silken
lashings as in Fig. 168. This cut is drawn from a sketch made on the
grounds of the Smithsonian Institution, Washington.

Dictyna.

Of the typical cocoons of the Territelarise we may speak with some
positiveness; but the number of species whose cocoons are known is small.
However, it is highly probable that the variety of form and
method of suspension and care is not great, and we may per-
haps conclude that we possess a good knowledge of the general
cocooning habit of the tribe.

Mr. Enock determined the position in which the mother Atypus piceus
spins her cocoon. In a tube ten inches long and from a half to five-
eighths inch in diameter he found that about six and a half inches below
the surface the tunnel widened into a sort of pouch. On opening this he
saw the mother’s cocoon suspended in a beautiful hammock of silk one
inch long, the flat ends of which were about three-sixteenths of an inch
wide, and were attached to the top and bottom of the pouch.!

This description entirely corresponds with that previously recorded by

Terri-
telariz.

1 Life History of Atypus piceus, Trans. Ent. Soc. Lond., 1885, page 394.

—— Oe

——

138 AMERICAN SPIDERS AND THEIR SPINNINGWORK,.

M. Eugene Simon! and by Mr, Pickard-Cambridge.? Mr. Simon has made
a drawing of the cocoon as fourd by him in natural site, which
I reproduce from the paper just quoted. The earth is therein
shown dug away to disclose the burrow, and the projecting tube
is seen as laid along the surface. (See Fig. 169.) Instead of the ham-
mock which Enock describes, Mr. Simon says that a number of threads
are used to suspend the cocoon in the throat of the enlargement of the
burrow.

Mr. Enock found the male of Aty-
pus piceus in the tubular nest of the
female October 15th, and again Octo-
ber 20th, but the fertilization must.
have occurred earlier, for the same
writer, on August Ist and again on
September 1st, found the cocoons con-
taining eggs, and during the months
of September and October the young
were already found hatched. Accord-
ing to this observer, the number of
eggs in the cocoon of Atypus piceus
was usually over a hundred. On sey-
eral occasions he counted the number
of young living with a single female,
the sum always exceeding one hun-
dred, and sometimes as high as one
hundred and fifty-seven.? Blackwall,
however, states that the mother Aty-
pus deposits between thirty and forty
eggs,* but in view of the particular
and definite statements of Mr. Enock
Fre, 169. ‘The cocoon of Atypus piceus, suspend- we must conclude that this is a mis-

ed within her tunnel. (After Simon.)

take.

Abbot’s Atypus of Florida no doubt protects her egg sac in the same
manner as Atypus piceus, since, according to Abbot’s note, as re-
corded by Baron Walckenaer,® and which I have read in the
original manuscript, the young are found, like the offspring of
Lycosids, domiciled on the back of the mother after they are hatched.®

Atypus’
Cocoon.

Abbot’s
Atypus.

‘ Annals of the Entomological Society of France, fifth series, Tom. III., 1874, page 114
and pl. 4.

* Spiders of Dorset, page xxxiii., Introduction. 5 Op. cit., page 392.

* Spiders of Great Britain and Ireland, page 15.

* Hist. Nat. des Insectes, Aptéres, Vol. I., page 248.

® McCook, “Nesting Habits of the American Purseweb Spider,” Proceed. Acad, Nat. Sci.,
Phila., 1888, page 213.

GENERAL COCOONING HABITS OF SPIDERS, 1389

That accomplished French arachnologist, M. Eugene Simon, has recently
added largely to our knowledge of this interesting tribe. A visit to South
America enabled him to make personal studies of trapdoor nests,

eeu and these have happily found expression in admirably drawn
Spiders, Plates, some of whose figures I haye ventured to redraw for these

pages. Rhytidicolus structor is a common species in Venezuela,
particularly upon the slopes of compact and sandy ground. Its burrow is
the most complex that Simon observed. It is composed of three successive
spacious chambers, communicating one with another by straight openings,
which close by a hinged door.

The first chamber is largely dilated in the form of a pear, but quite
contracted at the two extremities. (See Fig. 176.) The second chamber is
more or less cylindrical, and termi-
nates in a cul de sac. The third
chamber communicates with the sec-
ond, not by its extremity, but upon
the side, which is dilated and oval,
like the first, and rounded at the bot-
tom. The walls of the entire burrow
are perfectly built, very smooth, and
draped with a white tissue, light,
transparent, and adhering. The three
doors are almost alike. They are
thick, cut like a stopple upon the edge,
and penetrate within the opening,
which is itself slightly widened and a
little prolonged beyond the surface.
They are semicircular, and cut in a
straight line on the side of the hinge.
Their superior faces are rough, like

at gate P 2 Fic. 170. Section in the earth, showing trapdoor
the adjoining soil, even with the in- _ nest of the female Rhytidicolus structor. (After

Simon.) Fic, 171. Outline of first chamber of

side doors ; sometimes at an ex ternal Rhytidicolus structor, to show location of cocoon.

opening the doors are a little swollen,
and very unequal, but always slightly concave on the internal doors. The
internal faces of the doors are convex, and have a silk drapery like that
of the walls. On the edge of the bevel are small holes for the attachment
of the claws when the trap is to be held down, and these are more distinct
on the entrance door. This swings naturally from within to the out-
side. The second door opens, on the contrary, from the outside inwardly
in such manner that in the first chamber the two doors show the inter-
nal faces equally smooth. ‘The arrangement of these doors is shown in
the figure.

The female deposits her eggs in the first chamber; they are not agglu-
tinated, and are enveloped in a cocoon of white, opaque tissue, much longer

“? ye i el siial ¢

140 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

than large, and are suspended obliquely, like a hammock, between the op-
posite walls, as shown in the outline sketch, Fig, 171.

Among the Venezuelan Avicularidee Simon discovered and describes an

interesting species, which he names Psalistops melanophygia. It is a com-

mon species in the neighborhood of Caracas, particularly in the

Burrow forest of Catuche. It digs a burrow in the ground six or seven

and ee inches in depth, garnished toward the top with a silken lining
Sheree slightly adherent. The burrow is quite straight in the upper

tops. part, from which proceeds a simple branch, straight and quite

long, cutting the main entrance at an acute angle, and mount-
ing near to the surface of the earth. (See Fig. 172.) Below the point at
which this side branch enters, the main
burrow is much enlarged and more or less
curved towards the bottom. From this
point also it is destitute of a silken lining.

The opening to the burrow is with-
out a trapdoor. It is slightly elevated
above the surface, where it is always gar-
nished by a collarette of dry leaves or
any other sort of débris retained within
the threads. The eggs, which were ob-
served on the 12th of January, are not
agglutinated. They are enveloped in a
simple cocoon of cottony tissue, white ahd
opaque; are placed near the bottom of
the burrow, and suspended from one of
the walls by a very short pedicle or
stalk. (See Fig. 172.)

A large female ‘Tarantula, probably
Eurypelma hentzii, or a closely related.
species, was sent to me from the West
Be Indies, and arrived at the Academy dur-
Fic. 172. The burrow of Psalistops melano- ing a prolonged absence. She died be-
eae on Aa Be oe suspended at fore my return, and was preserved in

spirits; but afforded me an opportunity,
which I had long desired, of determining the egg cocoon made by this family
of the Theraphosoide. While cleaning out the box in which
she had been sent I observed a piece of spinningwork within,
which proved to be an abandoned cocoon. When inflated it
showed a hollow spheroid composed of thick silken cloth, somewhat soiled
on the outside, but within clean and white. It measured two inches along
the epee axis and one and one-fourth inch along the shorter one. It

Tarantula
Cocoon.

* Simon, Arachnides de Venezuela, page 197, plate 3, Fig. 1.

a er t=. i i ge ee a hn ll i eee | ms
y ow er’ .?,

GENERAL COCOONING HABITS OF SPIDERS. 141

was empty of young, whose first moults, however, were within the cocoon,
as were also a few unhatched eggs, which are yellowish spheres three mil-
limeters in diameter. Three small openings in the case showed where the
spiderlings had escaped. Both cocoon and eggs are shown natural size in
the accompanying figure. (Fig. 178.

The interior of this cocoon was without any flossy lining or padding,
resembling thus the egg sac of the Lycoside generally. A curious flap
overlapped the cocoon at one side, whose use I could not conjecture, unless
it may have served to attach the object to the mother’s body, or suspend
it within her burrow; or perhaps it was simply a remnant of material
which had remained after the eggs were rolled up within the silken rug
upon which they are proba-
bly deposited after the man-
ner which I have shown to
exist in the genus Lycosa,!

The janitor who received
the box containing this spi-
der and placed it in my
room was at the time new
in his position, and did not
understand the importance of

observing all the
Mode of . ial
Cnriviin: particulars in the

habits of living
creatures sent to the Acad-
emy. He therefore failed to
make any notes, but told
me, when questioned, that
he believed the cocoon was
attached to the lower part
of the spider’s body when Fic. 173, Cocoon and eggs of the Tarantula (Mygale).
it arrived. No doubt this ss 7th
is a correct observation, and we may assume with some degree of certainty
that the large egg sac of the Theraposids is carried by the mother, lashed
to the spinnerets at the apex of the abdomen, precisely as in the case of
Lycosids, whose well known habit is familiar to every frequenter of our
fields.

This cocoon is exhibited in my collection of aranead architecture de-
posited in the Philadelphia Academy, and is the only one, as far as I
have been able to learn, exhibited in any similar institution. A second
specimen in my possession is similar to this, except that the silken sac is

' See Proceedings Academy Natural Sciences of Philadelphia, 1884, page 188, my note on
“How Lycosa fabricates her round Cocoon,”

142 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of much more delicate tissue, it probably having been made in confine-
ment. Termeyer speaks of cocoons of the Mygalide of South America
(“Aranea avicularia”) even greater than the above. They are three inches
long by one wide, and are placed in the fissures on trunks of trees. ‘They
contain thousands of eggs. This extraordinary size of the cocoon had made
the inhabitants, who do not observe carefully, imagine that this
spider would take the cocoon of “the bombice moth, del Guyayo
(Janus, Linn.),” and, having destroyed or eaten the chrysalis, would place her
own eggs therein, and then artificially close the hole by which she had pene-
trated it. One of these cocoons weighs as much as six cocoons of the silk
worm before they are washed, and as much as three or four after having
been washed.!

In San Domingo, according to Palissot de Beauvois, Mygale blondii is
found in the fields, where it prepares a hole in which it awaits its prey.
It does not confine itself to this manner of providing its food, but issues
forth eyening and morning, climbs up trees, and, penetrating into the nests
of small birds, sucks their eggs or the blood of their little ones. ‘The
female’s cocoon is the size of a pigeon egg.”

Walckenaer describes the cocoon of Mygale avicularia as composed of
three silken enyelopes, of which the middle one is thinner, and
does not contain a silken padding. ‘The female places her
cocoon near her tubular dwelling, and watches it assiduously.

M. Moreau de Joannés, as quoted by Baron Walckenaer, says that the
female of this species in Cayenne envelopes, in a cocoon of white silk, her
eggs, to the number of eighteen hundred or two thousand. He observes
that the red ants eat the little Mygalide when they issue from the cocoon.
M. Guérin had in his collection a cocoon of this Mygale which was covered
over with a multitude of very small parasitic Cynips. This cocoon was flat-
tened, rounded, and about three inches in diameter. It was opened in the
presence of Walckenaer, and the young spiders were found enclosed therein.*

Madame Merian, who first recorded a report that the Theraphosoide prey
upon small birds, must have observed the cocoon of these spiders, as it
seems to me. She indeed speaks of them as having their domicile in a
large round nest resembling the cocoon of a caterpillar; but the plate to
which she refers is a fairly accurate figure of a female tarantula with a
large oval cocoon attached to her abdomen, in the way usual to Lycosids.4
I haye the opinion that the egg cocoon of the spider was mistaken by
Mademoiselle Merian or her informants for a ‘“ domicile.”

Size.

Mygale
avicularia

x
0

Communications Essex Institute, Vol. V., 1866-67, page 61. “Researches and Experi-
ments upon Silk from Spiders and upon their Reproductions, by Raymond Maria de Ter-
meyer.” Translated from the Italian, and revised by Burt G. Wilder.

* Walckenaer, Aptéres, Vol. II., page 211. 5 Aptéres, I., 218.

* Desertation sur la Generation et les Transformations des Insects de Surinam. Marie
Sibillee Merian. A la Haye, MDCCOXXVI. Fig. 18 and explication.

ee Sl

GENERAL COCOONING HABITS OF SPIDERS. 148

At all events we may consider it fairly well assured that, in her cocoon-
ing habits, the female Tarantula throughout most, or perhaps all, species,
closely resembles the Lycoside, and the resemblance probably ex-
tends to all the Territelariz. In other words, the Theraphosid
cocoon is, first, round or ovoid; second, is carried about with the mother,
attached to her body, or kept under her care; and, third, the young for a
period longer or shorter remain with their mother. The affinity between
these two great groups of araneads is also marked in their nesting habits;
both burrow into the ground a cylindrical tunnel or shaft, within which
they domicile, sometimes lining it more or less completely with silk.

Summary

? IV.

Passing now into the group of Wandering spiders, we reach the co-
coonery of the Citigrades, and here find little variety in structure, with
scarcely an exception, The cocoons of this
tribe are round balls without any interior
furnishing, which are carried by the moth-
er within her jaws, as in the case of Dol-
omedes, or lashed to the spinnerets, as’
with the Lycosids and most other species. Al —
(Fig. 174.) The manner in which the co- SS
coon is made has been quite fully de- Fic. 174. Lycosa carrying her round cocoon
scribed by myself, 1 lashed to her spinnerets,

While walking in the suburbs of Philadelphia, I found under a stone
a female Lycosa (probably L. riparia Hentz), which I placed in a jar on

dry earth. For two days the spider remained on the surface
Lycosa’s pearly inactive. The earth was then moistened, whereupon
Mode of ; : sen Tae ;
tana: (May 2d) she immediately began digging, continuing until she
ing. had made a cavity about one inch in depth and height. The

top was then carefully overlaid with a tolerably closely woven
sheet of white spinningwork, so that the spider was entirely shut in. This
cover she fortunately made against the glass side of the jar, and the move-
ments of the inmate were thus exposed to view. Shortly after the cave
was covered the spider was seen working upon a circular cushion of
white silk, about three-fourths of an inch in diameter, which was spun
upwards in a nearly perpendicular position against the earthen wall of
the cave. The cushion looked so much like the work of Agalena nevia,
and the whole operations of the Lycosa were so like those of that spider
when cocooning, that I was momentarily possessed with the thought that
I had mistaken the identity altogether, and again examined her carefully,
only to be sure that she was indeed a Lycosid.

1“ How Lycosa fabricates her round cocoon.” Proceed. Acad. Nat. Sci., Phila., 1884,
page 138.

— i
at
ns

144 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

After an absence of a half hour I returned to find that in the interval
the spider had oviposited upon the central part of the cushion, and was
then engaged in covering the hemispherical egg mass with a silken
envelope, working like a mason spreading mortar with a trowel.

Unluckily, at this stage of work I had to leave for an imperative en-
gagement, and did not see my spider again for an hour and a half, when

: I was delighted to find a round silken ball dangling from the

co ae apex of her abdomen, held fast by short threads to the spin-
nerets. The cushion, however, had disappeared. It is not dif-

ficult to explain the intervening process. Within this circular cushion the
eggs are deposited, after which act the spider proceeds to pull the edges of
the cushion together until the whole is rolled ‘around the egg mass, after
the fashion of a schoolboy putting a leathern covering on a yarn ball. ‘This
done, the mother goes over the exterior of the ball, and spreads upon it an
outer layer of spinningwork, which is woven in the same manner as the

Fic. 175. The cocooning burrow of Lycosa saccata, made underneath a stone. The walls of mingled silk
and soil. This figure shows the nest as exposed when the stone was removed. Fic. 176. The stone
under which the burrow of Fig. 175 was made. The under part of the stone is shown turned upward.

original cushion. ‘Thereupon she attaches it to her spinnerets, where it is
varried until the young are hatched. I had often wondered how the round
egg ball of the Lycosid was put together, and the mechanical ingenuity and
simplicity of the method were now apparent. The period consumed in
the whole act of cocooning was less than four hours, and the act of ovi-
positing took less than half an hour. Shortly after the egg sac was fin-
ished the mother cut her way out of the silken cover woven over her
little cavern. She had evidently thus secluded herself for the purpose of
spinning her cocoon. This was in accord with a firmly fixed habit of the
Lycosids to exclude themselves, before making their cocoons, in a burrow
or cave which they form in the ground. This is often made under a stone
and is protected on the sides by a plastered wall of mud, and above against
the stone by a piece of spinningwork which thus forms an upholstered
roof to this pretty, cavernous home. An approach to the cave is:cut, which
often debouches among the grasses, clumps of clover, mosses, or wild flow-

GENERAL COCOONING HABITS OF SPIDERS.

Fie. 177. Leaf woven cocoon nest of Dolomedes sexpunctatus.

146 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

ers, that give a touch of natural beauty to the gateway. One of these
Lycosid cocooning caves is shown at Figs. 175 and 176. It was made be-
neath a stone, and when that was lifted up the spider, Lycosa saccata,
showed within as at Fig. 175. The roof of her den was broken off by
lifting and is shown in inverted position at Fig. 176. The use of this
special cocooning den is common with Lycosids; but some species, and
probably all at times, live within the home burrow while carrying their
cocoons. This is the habit of Lycosa arenicola, which may often be seen
on her turret with her egg ball at her spinnerets. (See Vol. I., page 314,
Fig. 289.)

There is no flossy wadding within the cocoon case of Lycosids, as is
common with Orbweaving spiders. Indeed, such a provision for the com-
fort and safety of the brood appears wholly unnecessary in the case of
younglings whose egg life is so brief, and of a mother who carries her
young about with her, and thus gives them the advantage of her personal
protection and care. The Orbweaving mother generally dies within a few
days after ovipositing. Personal protection of her offspring is therefore im-
possible, and the period of development is often greatly prolonged. Nature
has taught her to provide for them the necessary covering of a warm, flossy,
silken blanket beneath which they may outlive the changes of weather.

In the case of Dolomedes, the cocoon is carried by the mother until
shortly before the period of hatching, when it is generally deposited within
a pretty nest composed of leaves
drawn together and lashed at the
edges into the form of a tent.
(See Fig. 177; also Vol. L., Fig.
339.) Within this a mass of in-
tersecting lines is spun, upon
which the cocoon is hung. After
hatching the spiderlings occupy
the temporary home thus provid-
= ed for them, and hang in clusters
or individuals upon the intersecting lines.

Dolomedes differs from Lycosa in the mode of deporting her cocoon,
suspending it beneath the abdomen and sternum, so that it is surrounded
by the legs. (Fig. 178.) When walking, the mother Dolomede must -
straighten out her legs as much as possible, and carry her body high.
(Figs. 178, 179.) The. cocooning habits of the English Dolomedes mira-
bilis differ in no particular from those of our American species. She car-
ries her cocoon, which is large, globular, and of a dull yellowish color,
attached to her body during all her hunting expeditions, until the time
approaches for the hatching of the eggs. She then weaves a sheet of
close, fine silk upon grasses or the branches of bushes, forming a dome, of
which these supply the rafters.

Fig. 178. Dolomedes sexpunctatus carrying her cocoon.

eee. = =

GENERAL COCOONING HABITS OF SPIDERS. 147

Among the Citigrades, Pucetia aurora has a special interest, both from its
appearance and structure and from the peculiarity of its cocooning habit.
This spider was received in collections sent me by Mr. W. G.
Wright, of San Bernardino, California. Numerous specimens of
young and old were subsequently sent by Mrs. Eigenmann and
others from the same locality. The genus Pucetia belongs to the family
Oxyopoid of the Citigrade spiders, to which it is doubtless relegated in
spite of certain analogies with the Satigrades on the one hand and the
Laterigrades (Philodromine) on the other.1_ Mr. Wright describes the
specimens sent me as jumping spiders; and Hentz, who describes several
species under the generic name of Oxyopes, says that Oxyopes salticus leaps
with more force than Attus. This family is arboreal in habit; the spiders
are found on plants, with their legs extended, thus practicing Tetragnatha’s
form of mimicry, and thence spring-
ing upon their prey. The female’s
cocoon is usually conical, surrounded
with points, placed in a tent made
between leaves drawn together and
lashed, and is sometimes of a pale
greenish color. Oxyopes viridens
will make a cocoon suspended mid-
way by threads attached to these ex-
ternal prominences, and this she will
watch constantly from a neighboring
site. Hentz also thought that the
mother of« this species carries its Fia, 179. English Dolomedes mirabilis carrying

. her cocoon. (After Blackwall.)
young like a Lycosa.?

Pucetia aurora appears to be a new species.* The body length is four-
teen millimetres; the legs are long and tapering, except among the young,
The body is yellow and pale yellow; the cephalothorax striped longitudi-
nally with bright red streaks; the abdomen marked above with red streaks;
the sternum is red; the legs are yellow, with red rings at he joints.
These red streaks upon the yellow background suggested the specific name
of “aurora.”

The cocoon nests, according to Mr. Wright, are uniformly placed upon
bushes of Erigonum corymbosum. They are hung from three to four
feet above the ground, and, being upon the topmost twigs of the plant,
are easily seen from a distance. The cocoons, received by me in consider-
able number, are straw colored spheres five-eighths of an inch in diameter.
They are covered externally with various pointed rugosities, from which
numerous lines extend to the adjoining corymb of the plant upon which

Pucetia
aurora.

!'Thorell, On European Spiders, page 196. * Spiders of the United States, page 48.
, I I pag pag

* Proceed. Acad. Nat. Sci., 1883, page 276, “Notes on two new California Spiders.”

148 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

all the specimens sent are attached. (Fig. 180.) The retitelarian snare
which surrounds the whole doubtless serves as a temporary home for the
young spiders. The cocoon has no suture, and the spiderlings escape by
cutting the case, which is thick and closely woven. No flossy padding was
found inside of the case. (Fig. 181.) The
cocoon thus resembles that made by Cit-
igrades generally.

A fine, large species of Ctenus from
Central America, sent to me by Mr.
Samuel H. Scudder, carried its cocoon.
This was a large object, one inch and a
quarter long, constructed in the ordinary
manner of Lycosid cocoons, but differ-
ing somewhat in shape, being globular
instead of hemispherical. The mother
carried it for some time after she came
to me, and then fasteried it by threads,
in hammock fashion, to the side of the
box wherein she was confined. Shortly
thereafter a large brood of spiderlings
appeared, spread themselves over my lab-
oratory table, covering the books and
other objects thereon with a sheet of fine
spinningwork. They finally built upward
a huge bridge like structure, a sort of
aranead Hiffel Tower, which touched the
ceiling above the table, and at another
point diverged to the laboratory window.
Some further account of this brood, with
a figure of their bridge at a certain
stage, will be found in the subsequent
chapter on Cocoon Life and Babyhood.

Fic. 180. Duplex cocoons of Pucetia aurora, woven Among Saltigrades the cocoons close-
among the blossoms of Erigonum corymbosum. .

ly resemble those of many of the genus
Epeira. They are spun against some surface, as that of a rock or
tree, the eggs being overlaid by a thick blanket of white spinningwork.
Over this again is stretched a tent or cell of lighter structure, although
still of close and somewhat adhesive texture, but not so thick as to pre-
vent the cocoon from being seen through it. The eggs are heaped in a
hemispherical mass, and are of a pinkish or light rose color. In the case
of Phidippus morsitans the cocoon is from one-half to three-fourths of an
inch in diameter. ‘The spider dwells within her cell, and.of course close

Rs ee CCU CU CU — _ .

GENERAL COCOONING HABITS OF SPIDERS. 149

by her cocoon. The outer covering of the eggs is quite thick, very white,
and apparently a little viscid; at least, it is quite adhesive. The exterior
tent has something of the same qual-
ity. (Fig. 182.) I sometimes find the
cocoon of Saltigrades enclosed within
the nesting cell and spun up within a
rolled leaf, as shown at Fig. 183, a beau-
tiful example of aranead sewing. Fig.
184 shows the leaf opened up, disclos-
ing the tubular nest, and again the
mass of eggs much enlarged and dis-
played against the egg case thrown back,
the egg case, of course, being within
the cell.

One of the most elaborate cocoon nests woven by a Saltigrade spider
is that made by Phidippus opifex of California.! The examples both of
nests and spiders in my possession were sent me by Mr. W. G. Wright, of
San Bernardino, California. The cocoon nest is externally an egg shaped
mass of white spinningwork, sometimes three inches long by two and a
half inches wide, but often less, as in Fig. 185, which is drawn natural
size. The outer part consists of a mass of fine silken lines crossing in all
directions and lashed to twigs of sage bush, within which it is enclosed.
This maze surrounds a sack or cell of thickly woven sheeted silk, irregu-
larly oval in shape, two inches long by one inch in width, and also at-
tached to the surrounding twigs. At the bottom this cell or tent is pierced
by a circular opening which serves the spider as the door of her domicile.
Like others of her genus Opifex lives and hibernates within this silken
tent. Against an inner side of the
tent she spins a lenticular cocoon
(of double convex shape), consisting
of thick white silk, within which the
eggs are placed. When the cocoons
sent me reached Philadelphia many
young spiders had escaped and occu-
pied the package box. They were
about one-eighth’ inch long, resem-
bling the mother, but less heavily

Fic. 182, Cocoon tent and cocoon of Phidippus coated with gray. The spider her-

morsitans spun upon a rock. ers ~ :
self is a large example, five-eighths
of an inch in body length, stout, the legs of moderate thickness, the whole
animal covered closely with grayish white hairs, the skin beneath which is

Fic. 181. Cocoon of Pucetia aurora, opened
to show structure,

‘The spider and its habits were originally noticed by me in Proceedings Acad. Nat.
Sci., Philadelphia, 1883, page 276,

—— =F

150 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

‘ Fic, 186.

Fia. 185.
Typical Saltigrade Cocoons.
Fic. 188. Sewed leaf tent of a Saltigrade spider. Fic. 184. Leaf opened to show the silken cell
and the egg mass (enlarged) in the open cocoon. Fic. 185. Cocoon nest of Phidippus opifex.
(Natural size.) Fic. 186. Fac simile of a figure of an Attus cocoon nest, by Baron De Geer.

GENERAL COCOONING HABITS OF SPIDERS. 151

black. I named the species Attus opifex, but according to Professor Peck-
ham it belongs to the genus Phidippus.!

I present in this connection a fac simile drawing of perhaps the earliest
sketch of a Saltigrade cocooning nest. July 26th, 1745, Baron De Geer
found among the needle like leaves of a pine tree a large, oval cocoon
nest of white silk, which was woven around the branch and interlaced with
the leaves. (Fig. 186.) The spider was inside along with her little ones,
who were present in great number. In the middle of the cocoon nest, at
the side, was a door, at which the spider stayed on guard; but generally
she was within the tent with her little ones, preferring the back or middle
part thereof, near the supporting branch. De Geer found at the entrance
detritus of flies and other insects which had been devoured by the mother,
such as the legs, wings, etc.

The spiderlings accompanied the mother, and appeared to live on good
terms with her. They were about a line long, but dtherwise quite resem-
bled the mother, having black bodies and brown legs. They moved with
great vivacity and appeared to be nourished, in cormmon with their mother,
by the prey captured by the mother. The species appears to be Dendry-
phantes hastatus Clerck.?

¥I.

Among Laterigrades a very general habit is to spin a plano convex
cocoon of tough silk fibre, which is attached to some surface. Sometimes
a light shelter tent is spun over this, and the
spider will be found dwelling within. (See Vol. I,
page 347, Fig. 338.) Thomisus cristatus Clerck,
of Europe (Xysticus audax Koch), secludes her-
self in the leaves and stretches some isolated
threads around her, and there sometimes she sus-
pends herself. In this retreat the female lays her
eggs in a flat cocoon, one-fourth inch in diame-
ter, the tissue of which is swollen by the eggs,
and presents rounded eminences. The spider places a
herself upon the cocoon and does not abandon it “
when touched. The cocoon contains one hundred
eggs of yellowish white color.*

The eggs of Philodromus are usually enclosed within a cell which is
hung among the leaves or stretched between twigs. (Hig. 187.) The egg
sac is surrounded by a slight silken tent, wherein the mother dwells. An
example of Philodromus mollitor, in my collection, is woven in the angles

Fic. 187. Cocooning tents of
Philodromus mollitor.

1“North American Spiders of the Family Attidee, Phidippus opifex McCook.” Trans.
Wisconsin Acad. Sci. Vol. II., 1888, page 20.

2 De Geer, pages 286, 287. 8 Walckenaer, Aptéres, Vol. I., page 523.

4 This example was sent me by Dr. Geo. Marx as the cocoon of this species.

_— es? oe oe

152 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of forked twigs and are composed of very white stiff silk, the stiffness prob-
ably being caused by the tightness with which the lines were spun.
(Fig. 187.)

Misumena yatia is well known among the Laterigrade spiders by its
remarkable mimicry of the colors of flowers upon which it lurks for prey.
A fine example of its cocoon was brought to my notice by a lady who had
transported a specimen from the Wyoming Valley to her home in Phila-
delphia. Her attention had been arrested by the remarkable resemblance of
the creature to the bright golden yellow Coreopsis flower on which she discoy-
ered it. The spider was, placed in her bedroom chamber about the 28th of
August, and during the first week in September it wove the cocoon repre-
sented at Fig, 188, in a corner of the dressing bureau, just where a mirror
is fixed in the woodwork.
The cocoon consists of a
flossy mass, something
| after the fashion of that
Hy. of Epeira, which covers
= over the eggs. A tent of
close white spinningwork
encloses this, having at
the bottom a circular
opening one-eighth inch
in diameter, through
which the spider passed
to and fro. A thicker
band of silk, of the most
beautiful whiteness, look-
ing like spun glass, passes
across the centre of the
enclosing tent, joined at

FiG. 188. Cocoon and tent of the Laterigrade, Misumena vatia, one end to the mirror
woven upon a ladies’ dressing bureau. 2 :
and at the other to the
cabinet work, The tent and cocoon are partly woven upon the glass of
the mirror. The tent is about two inches long and one and a half inch wide,
and the cocoon, which is somewhat irregular in shape, is about three-fourths
of an inch in diameter. A few separate lines are stretched across the entire
spinningwork, and attached on either side of the angle or corner which
contains the cocoon. A few days after finishing this work of maternal
industry the mother died. According to Hentz! this species attaches its
cocoon to the under side of a leaf, and remains near it. :

There is evidently a good deal of variety among the cocoons of Lateri-

grades. Many of them consist of two stiff, paper like pieces, viz., first, a

WZ
( al

1 Spiders U.%., page 78, on Thomisus fartus.

———— rl <a

————— se |, lr tC avr

GENERAL COCOONING HABITS OF SPIDERS. 153

flat circular plate which is attached to the object, rock, bark, or wood on
which the cocoon is spun; and, second, a convex covering which fastens
above the eggs like a cap. The
inside is lined with pure white
silk, but the outside is often
of a grayish brown, and ap-
parently is purposely soiled in
order to subdue the color. Usu-
ally there is no flossy pad-
ding for the eggs. I am not
sufficiently acquainted with co-
coons of this tribe to enter
largely into a comparison with
those of others, but the forms *
known to me and above de-
scribed are probably typical, "1.45: are sd rage seaee coats ape caries
and substantially represent the
maternal industry of the Laterigrades of the United States, and probably
of the globe.
The well known tropical species, Heterapoda venatoria, or the Hunts-
man spider, is one of the largest of the Laterigrade species, and may
properly be classed with the spider fauna of the United States,

The as I have specimens from Florida. It abounds in the West
Hunts- 4s r . 7 & é

Indies. The cocoon is a large double convex or plano conyex
man Het- 3

erapoda, Object, resembling those of Thomisus and other species when
woven against various surfaces. It appears, however, to be car-

ried by the mother; at least, one female preserved by me in a box wove
a cocoon of this sort which she carried in the manner represented at Fig
189, which gives a view from
above, and Fig. 190 a view
from beneath. The button like
cocoon was put beneath the
body, which it almost entirely
covered; at one end it appeared
to be attached to the spinner-
ets, and at the other was held
tightly by the outspread palps.
The mother made an awkward
\ appearance as she straddled
about the box, holding her
Fic. 190. Cocoon of Huntsman spider, clasped by the mother. legs high up and outspread
Sey, ere Renee over her cumbersome cocoon.

If this be fairly representative of the prevailing habit of this widely
distributed species, we have among the Laterigrades also an example of

4s)

154 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

spiders that protect their cocoons by deportation, in this respect allying
the Laterigrades to the Citigrades, Territelaria, and the few known spe-
cies of Retitelaria, by whom cocoons are deported.

VII.

The cocooning habit continues, apparently without any marked change
from the normal methods, under the most decided changes of enyiron-
ment and eyen of life economy and constitution. This is shown
he well by some studies made of the cave fauna of the United States.
Prof. A. S. Packard, Jr.,1 accompanied by Profs. Shaler and
F. G. Sanborn, collected a number of spiders from Mammoth, Wyan-
dotte, Bradford, Carter, and Weyer’s Caves, obtaining about one hundred
specimens in nine species. Six of these, all belonging to the Theridioids,
were found in considerable numbers in the larger caves, where there is
little or no light, and the climate is little affected by outside changes.
These, judging from the printed descriptions, follow closely the character-
istic cocoonery of like species in the open air,

The well known cave species Anthrobia mammouthia was collected, to-

gether with cocoons, in the Mammoth Cave under a stone in dry, but not
the driest, places. On the bottom and at different points in the
Anthro- ¢aye sometimes two or three cocoons would be found under a
eee stone as large as a man’s head. The cocoons were orbicular,
* flattened, one-eighth inch in diameter, formed of fine silk, and
contained from two to five eggs. They occurred with eggs in which the
blastodemic cells were just formed, April 25th. The eggs were few in
number, and seemed large for so small a spider, being twenty-five thou-
sandths of an inch in diameter. Packard was unable to trace the devel-
opment of the young and ascertain if the embryos are provided with
rudimentary eyes. Two young Anthrobias were hatched out March 3d in
his room.

Nesticus pallidus? was found in Fountain Cave, Virginia, among sta-
lactites where there was no daylight. It spins a weak, irregular
web, consisting of a few threads, according to Packard.? Among
these threads several loose cocoons were found, one containing
thirty or forty young just hatched.

Several years ago I received from Prof. Joseph Leidy two spiders col-
lected by him in Luray Cavern, which appear to be Linyphia weyeri.
The specimens were not accompanied by any notes as to snares or location,
but I have recently received some information concerning Luray spiders

Nesticus
pallidus.

1“The Invertebrate Fauna of Kentucky and Adjoining States.” American Naturalist,
Vol. IX., page 274.

* Emerton, “Notes on Spiders from Caves, ete.,” Amer. Natr., Vol. [X., page 279.

3 Id., page 277.

GENERAL COCOONING HABITS OF SPIDERS. 155

from Mr, Clarence P. Franklin, of Philadelphia, who made an explora-
tion of that cavern in order to collect the fauna living therein. Among

the specimens collected and submitted to me are two which
Luray appear also to be Linyphia weyeri, and are probably identical
with the Linyphia found by Professor Packard in the caves of
Kentucky, Indiana, and Virginia, and which have been de-
scribed by Mr. Emerton.

These spiders were found in a chamber separate entirely from that
part of the cavern which is now illuminated by electric lights, and which
is entered by a narrow opening. The location is about half a
mile from the main entrance of the cave, and is in total dark-
ness. The araneads were found upon the banks of a pool, and
were seen crawling about in the light of the torches in the guide’s hand.
No webs were seen in this spot, though doubtless such must exist.

In another inner chamber, about one-fourth of a mile distant from
the entrance, Mr. Franklin found numbers of webs and at least one co-
coon. This spot is also in total darkness,

is beyond the influence of the
Cave electric lights, and in a com-
eee paratively dry part of the cay-

ern, among older formations of
stalactite and stalagmite. Upon one sta-
lagmite about ten feet high and two feet
to two and a half wide at the base, he
found at least fifty webs hanging in vari-
ous parts of the structure. This stalag- =Sis
mite was roughly fluted from the base to Fic. 191. Snare and cocoon of Linyphia

‘3 : weyeri (?) in Luray Cavern.

the apex, the fluting being from one and
a half to three inches in width. Within the interspaces thus formed
were stretched these webs, which appeared to be all of one character, as
represented at Fig. 191. This figure was not drawn upon the spot, but
is a memory sketch made from notes and recollection of the observation.
The web appears to be composed of simple lines stretched across the space
and placed between the walls of the fluting, and one above another, so
thickly that they touch and form an open hammock or sheet. This when
seen was depressed in the centre, and formed a shallow pocket or pouch.
In the midst of the intersecting threads, and at one side of the snare,
was hung a small cocoon. It is about one-eighth inch in diameter, is
composed of fine flossy silk, and contained a small number of eggs.

I conjecture that the original web consisted of retitelarian lines strung
across the flutings, and that these, by reason of use and age, had relaxed
and merged one with another, forming the rude sheet like snare described
by Mr. Franklin. This characteristic may often be seen in Theridioid
snares. Indeed, as I have heretofore shown (see Figs. 211 and 212, and

Their
Abode.

156 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Vol. L, page 223), it is a constant tendency of Lineweavers to thicken the
upper part of their snares until they sometimes have a quite sheet like
appearance.

Nothing could be more interesting than a study of the life of these
creatures thus doomed to perpetual darkness in the bowels of the earth.

: To a great extent their natural history must be a matter of
aa speculation. ‘That they make snares we know, and the character

; of those snares is without exception the most rudimentary spun
by spiders, namely, a series of intersecting lines drawn from point to
point. The fact that they make snares implies that there must be some
creatures to be ensnared upon which they feed, and these creatures must,
of course, be less in physical power than themselves. What insects form
the basis of their food? Among the insects occupying similar American
caverns are minute delicate mites, and Professor Packard conjectures that
these, together with young Podure, may form a portion of their food.

Moreover, these cave araneads have sufficient vitality to propagate their
own species. The old story of wooing and mating goes on in these regions
of unbroken darkness just as it does in the sunlight and among the flow-
ers and shrubs above. Mother love is not quenched by the endless gloom,
and the tiny creature of the cavern spins and cares for its tiny* cocoon
just as does its more favored sister of the sunlight. ‘The number of eggs
within the cocoons is very small in some species, according to the state-
ments of Professor Packard from two to five in Anthrobia, but in others
the excess of life is quite sufficient (there being thirty or forty in the co-
coons of Nesticus pallidus) to justify the inference that the chief supply
of food for these cave spiders is drawn from their own numbers. In other
words, they feed upon each other, and these Plutonic children of Arachne
must continue their generations largely by cannibalism.

The influence of this mode of existence upon the structure and senses
of these cave spiders will be considered elsewhere. It may be fitting, how-
ever, to add here an experience which throws some light upon the manner
in which the animals might have been originally conveyed to these sub-

ae terranean abodes. One summer while examining the limestone
Origin caverns in central Pennsylvania, located among the Allegheny
of Cave M ae ; : ‘
WMetuns. ountains, in what is known as the Scotch Valley, not far from

the city of Altoona, I stood in the channel of a small stream in
the neighborhood of Sinking Spring. Looking forward, towards the source
of the stream, I saw the waters flowing down towards me, but gradually
diminishing, with no apparent reason, until, near the spot where I stood,
the stream dwindled to a mere thread and disappeared. It produced a
curious sensation to stand thus in midchannel and see a flowing brooklet
lost to one’s sight before it reached the point where, under ordinary cir-
cumstances, it would haye swept around the feet of the observer. The
secret, however, was readily explained, for the whole section underneath

GENERAL COCOONING HABITS OF SPIDERS. 157

Ps — ———

the bed of the stream is hollow and connected with the cavernous forma-
tion which characterizes the entire region. ‘The water could be seen trick-
ling down through the pebbles, and, by putting the ear to the ground, it
could be heard dropping into the depths beneath.

I observed that, in the neighborhood of the place where the threaded
points of the stream thus sunk out of sight, numbers of little spiders

_ were running about, some of them probably seeking food, but
pig ea others appearing simply to be enjoying themselves like other
Brink, Young animals at play. These spiderlings seemed to be chiefly

‘Lycosids, though some of them were probably the young of
Agalena neevia, and, it may be, other species. They were running over
the bed of the brook and hiding under the pebbles at the very spot
where the water sunk out of sight; and it immediately occurred to me
that nothing was more likely than that numbers of these spiders might
be, and indeed had been, caught in the falling waters and carried down
into the cavern underneath.

They have sufficient vitality to endure, without much injury, such a
transition, but what would befall them when they reached their subter-
ranean prison? Would these lost spiderlings make themselves at home
and proceed to adapt themselves to their new environment? Many spiders,
we know, love gloomy abodes, dens in the earth and shaded spots. They
are nocturnal creatures, and go out at night to seek their prey. It would
not be so very difficult for creatures reared under these circumstances to
survive in total darkness, if only the means of livelihood could be found.

But what would be the influence of environment upon these unwilling
prisoners of the cave? How long would it be ere change in life compelled

a change in structure? To what degree would that structural
Bee change occur? What forms are those most likely to survive
ave Life. oe ;

such a transition? What species were the ancestors of our cave
fauna? As far as known, those now extant appear to belong chiefly to
the Theridioids, and their habits indicate the rudimentary form of spider
life which is possessed by that family. Have all other species accident-
ally introduced into caves perished, except these delicately organized Reti-
telarians. Or shall we think that some species have gradually been trans-
formed by the influence of their strange new life into those peculiar forms
which now inhabit our caves?

Judging from the well known habits of surface species, I would have
expected and predicted that the prevailing fauna of caverns would have

belonged to the Tubeweavers. Many species of this tribe live
Tube- in corners, crannies, dark rooms, under stones, rocks, fallen trees,
Peetu and like locations, from which light is largely excluded. Others
dox. - secure their prey during night hours, although in this respect
they are not peculiar from other tribes. The transfer from
such a habit to a life within a cavern to which no light ever penetrates,

i
= oe Te ee

. weg Wet 7" x
AMERICAN SPIDERS AND THEIR SPINNINGWORK. +

or which is but dimly illuminated, would seem to be less decided transition
than in the case of many other genera. Nevertheless, the facts at present
within possession of araneologists appear to indicate that the Tubeweavers
have secured no permanent representatives among our cave fauna, this
position being wholly occupied by members of the Retitelarize. We have
thus presented the curious fact that the species which by natural tendency
appear to be best adapted to survive cavern conditions have been thor-
oughly eliminated or excluded.

am oP etl ait Ate We 46

OMAPTER VI.

COMPARATIVE COCOONING INDUSTRIES.

In this chapter I propose to review the cocooning industry of spiders
as given in the last two chapters. In order to compare those methods of
spinning the cocoon which characterize the genera of the several Tribes,
I shall first describe in detail the manner in which Argiope cophinaria
constructs her cocoon. ;

No single point, in my study of aranead spinningwork, has been a
subject of more prolonged attention, and the cause of greater disappoint-
ment, than the mode practiced by the mother spider in overspinning and
protecting her eggs. I strongly desired to see and describe the entire
process on the part of at least one species, and, if possible, of several
species. To this end I have year after year sought the natural sites at
the cocooning time, and have kept watch, day and night, personally and
by proxy, over numbers of gravid females confined within various boxes,
jars, and other objects. I have tried to make the artificial conditions as
favorable and natural as possible. Notwithstanding all the patience and
ingenuity expended upon the observations, I am compelled to confess that
the secretiveness of the female spider has been a fair match for my curiosity.

It is true that I have made many observations of the process of cocoon
making at various points of the same, ranging all along from the first
stages to the last. Yet I have never had the opportunity to see the entire
process in any one species or genus, and, indeed, there are one or two
points of the process which I have never seen in any species of any
family. Nevertheless, I have observed nearly all the stages of construc-
tion in the case of Argiope cophinaria, and am therefore able to describe
methods of spinningwork which, if they have heretofore been observed,
have at least never been made known. The description of this industry
will form the first section of the present chapter. Thereafter will be in-
troduced some conclusions which a comparative view of cocooning indus-
try has suggested to my mind.

I observed one of my Argiopes, which I will distinguish as Prima for
convenience’ sake, shortly after she had left her snare upon an ampelopsis
vine, crawling along stems under leaves with that uneasy demeanor which

commonly indicates that the crisis of maternity is near. I placed her in
(159)

— ee ee eee

160 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

a trying box, and sat up with her tntil a late hour at night. During
a considerable portion of the evening she moved back and forth in the
box, spinning lines from one side to the oth-
er, and finally settled in one corner as though
to rest. Thereupon I retired.

Next morning at 5.35 o’clock I visited the
box, and found that the eggs had just been
laid, and enclosed within their first thin silk-
en pouch. A rectangular patch of yellowish
white silk was swung to the cross lines spun
nibh asin eau the night before. Immediately beneath this

Tinian ovipositing.. The spider is bee WaS & tuft of brownish silk, that enclosed a

neath the mass of newly laid eggswhich tiny silken dish three-eighths of an inch in
she has just begun to overspin. . % « :

diameter, which rested against the top piece,

with its concavity downwards. Against this dish Prima had oviposited

her eggs, forcing them upward evidently as she hung in position

Argiope )}eneath. At the time of my observation this was the position

cophi- of affairs. The eggs were in a hemispherical mass, and hung
naria. s . . .

downward, with no enclosure except the white silken pouch which

is the first covering the Orbweaver usually places upon the eggs. (Fig. 192.)

The mother remained for a few minutes beneath her eggs, and then

began spinning the brown covering. Her back was downwards, and her

feet curved upwards, holding to the supporting lines or to the
The edges of the top piece. Gradually moving herself around in a
tig horizontal plane, she spun the threads upward against the top

part of the egg mass, attaching threads to the overhanging edges
of the flossy tuft already described. This action and position are repre-
sented by Fig. 193. At six o’clock and eight
minutes she rested for a few moments, and
at that time her work presented the appear-
ance represented at Fig. 198.

Unfortunately, she was much hampered
by having lost two legs, which happened
to be the most important for her present
purpose, as one of them was the hind leg
used in spinning, and the other the first leg,
which is the guide, if I may so say, of the
spider’s motion, being continually used to .
feel the way as she progresses, and pilot ™ sae nies ete Ae eae
her into the proper course. Thus mutilated, — shown drawing out the silk, and the bunch
Prima probably was twice as long accom- % 1°°PS i# shown against the cocoon.
plishing her task as she otherwise would have been, since she only had
one leg with which to draw out and pack the silk as it issued from the
spinnerets. Nevertheless, she managed affairs very handsomely.

- on Se —— =

COMPARATIVE COCOONING INDUSTRIES. 161

During this and the earlier part of her weaying it seemed to me that
the silk escaped from the posterior pair of spinnerets alone. It came out
as white silk with a little yellowish cast in it, bearing a pretty

hoe es gloss. The spinnerets were widely flared, and the silk issued in

ind Leg. :

several filaments. The hind leg was thrown upward as the

spider moved and seized these filaments with the foot, apparently using
all the spines from the claws upward to the tarsal joint, and even part of
the metatarsus. The thread was carried away from the abdomen rather
slowly toward the cocoon, (Figs. 194 and 195.1) At the same moment,
also, the abdomen approached the cocoon without touching it. Between
the spinnerets and the spider’s foot the silken filaments were stretched
taut (Fig. 196), and after the first gentle motion of approximating the
cocoon the abdomen was swung in the opposité direction; that is, away

Fias. 194, 195, The action of Argiope in drawing out silk with the spinning legs. The
alternation of the legs appears by comparing the figures.

from the foot, so that the intervening stream of silken threads was drawn
out to a considerable distance, sometimes as far as three-quarters of an
inch. In the meantime that portion of the outspun threads between the
spider’s foot and the point at which the line was attached to the cocoon,
of course, relaxed and doubled up into a curled loop of several strings;
. and in this condition it was when the leg finally touched the
Laying ¢ocoon. (See Fig. 196.) With a quick movement the thread
eae was slipped off the leg and pushed into the mass of spinning-
work already accumulated. It at once adhered, though no viscid
material appeared to be intermingled therewith, and added its flossy loops
to the mass that had been spun before. The position of the leg and spin-
nerets during this action is represented at Fig. 197.

1 These two figures were made from a second spider with full complement of legs.

162 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

At occasional intervals the spinnerets were laid against the cocoon
and held there a brief space, while the spider pushed them into the
mass, attaching her thread precisely after the manner described as cus-
tomary when she is making a dragline anchorage. (See Vol. I, page 61.)
This, of course, held to its position and
prevented the raveling of the thread al-
ready accumulated. I was somewhat sur-
prised that more use was not made of the
spinnerets, as I had conceived the idea that
FiG. 196, The hind leg of Argiope stretched they were continually employed to beat

out and drawing the ray of threads. ; :

down and pack the cocooning material,
after the fashion of the long spinnerets of Agalena nevia and most of
the Tunnelweavers. On the contrary, the spinnerets rarely touched the
cocoon, at this stage of the work at least, and the entire process of packing
was accomplished by the pressure of the leg alone.

It seemed to me also that the palpi had something to do in packing
the flossy loops as they gathered upon the mass. At all events, they were
always held doubled under, as represented in some of the fig-
ures, and were moved continually in a way that gave me the
idea that the spider appeared to be kneading the silk with them, Per-
haps the reason of this was to prevent the palpal claw from fastening in
the threads, for care was also taken to bend the claws of the feet well
under, as though to guard them from that annoyance.

In this manner the spider proceeded, working her silken mass down-
ward, and gradually bringing it to a tolerably regular, oval shape. ‘This

_ was done entirely by so regulating the discharge of the silk and
Equaliz- the application of it to the cocoon that the surface was kept
siete even and regular. I cannot positively say upon what principle

this was accomplished, but I was made
aware of the fact that the aranead continually
changed her course as she moyed around the co-
coon, describing a complicated series of convolu-
tions. This was shown strikingly in the following
way: In order to make exact drawings of the va-
rious attitudes of the spider while spinning, I drew
a number of outlines of the cocoon at various stages
upon blank paper, and waited to insert the various
parts, as the legs, palps, spinnerets, etc., in proper }
sequence as the spider would from time to time re- F'6. 197. Manner of forming and
appear at the same point. My idea was that on boise pte
one round I would sketch one leg, on another the next leg, and so on,
supposing that Prima would appear substantially at or near the same
place a number of times during her numerous rounds, and thus I would
have many opportunities to catch her in the same attitude.

Packing.

_s "7.

COMPARATIVE COCOONING INDUSTRIES. 163

I was, however, made painfully conscious of the fact that she very
rarely presented the same attitude consecutively. As she made her rounds
} she would almost invariably appear at a different point each
Wind- time, now above, now below, now at the middle, and anon
te oe emerging from beneath and coming back upon her course. (Figs.
* 198, 199.) It thus became a matter of much greater difficulty
to secure good drawings than I had calculated upon, although I eventually
satisfied myself. But in the meantime it appeared that by this peculiar
mode of progress the spider equalized the distribution of spinning material
upon her cocoon, and prevented any part of it from growing dispropor-
tionately to other parts. The method was not unlike that of a person
winding a ball of silk or wool from a skein of thread.
One cannot but recognize in this action a manifest purpose, however
directed or originated, to build up her cocoon mass symmetrically, and
cover all parts thereof equally. While thus engaged in spinning, the feet

Fic. 198. Fie. 199.
Equalizing the output of thread.
FiG. 198. Position below the cocoon, Fic. 199. Position above on next round.

were extended upwards, grasping the shoulder of the flossy mass or the
edges of the supporting top piece. As the mass increased, the legs, of
course, were stretched out further, but at no time was there any difficulty
in enclosing the entire structure within the long legs of the animal.

Brief rests were taken at long intervals of time, but the periods were
very short, three or four minutes as a rule, rarely more. The spinning
continued without intermission, sometimes more slowly and again
more rapidly. At seven o’clock and eight minutes the cocoon
appeared to be completed, as far as its general shape and size
were concerned, but the spider continued working on it until ten o’clock
and fifteen minutes, when I was compelled to cease my observations.

During the last three hours the spinnerets were more frequently squeezed
against the cocoon, as though to pack the mass and fasten the threads
more closely. The filaments now, instead of being bent upon the surface

Brief
Rests.

es ne

164 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

in the form of flossy loops of curled thread, seemed to be laid down as
straight lines. As a consequence the surface after spinning did not show
the flossy appearance, for example, of a bit
of cotton wool, but rather the smooth and

wound sewing thread. Not that the co-
x aN coon thread was wrapped quite as closely
Fi. 200. Fre, 201. as the spool, but in a general way it pre-
Fics. 200, 201. Hpeira strix enclosing her eggs sented that appearance. This effect was
within silk floss. (After Emerton.)
promoted by the use made of the leg,
which was laid flat along the cocoon, and the last two joints pressed against
it, thus serving to compact the threads.

When I returned at twelve o’clock and ten minutes, work upon the
cocoon had ceased, and the spider was putting in the finishing lines of
the maze of interlacing threads within which the cocoon of this species is
ordinarily suspended. I was somewhat surprised, however, to find that no
change had occurred in the exterior appearance and character of the mass
since I had left it. I had supposed that some kind of a varnish would be
laid wpon the surface, having the idea that perhaps some modification or
degree of the material which composes the viscid beading of the snare
would be used to cover in the interspaces of the silk on the exterior, thus
making it partly weatherproof. But Prima’s cocoon showed only the
glossy white silken surface with a little tinge of yellow, and no trace of
anything but the original silk as it had issued from the spinnerets.

This was in sharp contrast with a cocoon in the trying box just above,
which had been made by another Argiope two days before, but whose
making I was not able to see. I had watched it late into the night, and
in the morning when I came to look at it the cocoon was entirely finished
and the spider engaged in weaving
around it its protection of netted lines,
But the surface of the cocoon had been
treated in the ordinary way, and pre-
sented the customary yellowish brown
tint, had the hard, dry, parchment like
feeling, and gave out the crackling
sound which is almost invariable in
cocoons of this species. I have little
doubt that it is treated in some pe-
culiar way, immediately after comple-
tion, in order to produce this effect, but Fie. 202, Female Theridium tepidariorum finishing

A a cocoon. Other cocoons hanging in the nest,
as yet the method is unknown to me.

The spider Prima probably began to make her cocoon shortly before
five o'clock of the morning, and must have continued weaving at least
until half past ten. She was therefore engaged five hours, at the least,

COMPARATIVE COCOONING INDUSTRIES. 165

in work. But as her mutilation prevented her from making the ordinary
speed in spinning, this cannot be taken as a fair test of the time which
an able bodied spider would require for the same work. In

See Fe point of fact, I think that half the time, or from two hours
Weaving. to two and a half, is the period commonly taken by the fe-

male Argiope cophinaria to construct her beautiful and intricate
cocoon. I had the opportunity, since making the above detailed observa-
tions, to see other Argiopes spinning their cocoons. The process is always
the same, except that those spiders which (unlike Prima) have both their
hind legs, use them alternately in drawing out and packing the spinning
stuff.

¥ Ty,

Without entering into details as fully as with Argiope, we may state
generally the methods of cocoon construction practiced by other species
and tribes. This will give sufficient material for comparison.
The same mode of weaving is used by Epeira and all other
Orbweavers whose habit in this regard is known to me. Mr.
Emerton! gives a brief description of the manner in which Epeira strix
weaves her cocoon. She touches her spinnerets as in Fig. 200, drawing
them away at a short distance, and at the same time pressing upwards
with the hind feet, as in Fig. 201. Then she moves the abdomen a little
sidewise and attaches the bands of threads so as to form a loop. She
keeps making these loops, turning at the same time, so as to form a
rounded bunch of them.

Of the Lineweavers, I have seen heridium tepidariorum engaged in
overspinning her eggs. Fig. 202 represents this spider in the act of put-
ting the finishing touches upon the outer case of one cocoon, while two
others are hanging within the intersecting lines of her snare, from one
of which a little brood of spiderlings has already escaped. The cocoon
was suspended by a stout thread to the thickened mass near the top of
the web, upon which the mother held with one long fore leg while she
clasped and revolved her cocoon with the other legs. The spinning ma-
terial was drawn out and laid on in loops as described at length of
Argiope.

The process of making a cocoon, as practiced by the Speckled Agalena,
was observed in the case of a female confined within a glass jar. A leaf

was laid against the inner side of the vessel as a suitable object

Epeira.
Theridium

pees! ,. upon which to place the cocoon should the mother be inclined
Gencae if use it. She presently availed herself of the opportunity, and

wove upon the leaf a cocoon of the ordinary sort. I did not
observe the whole process, but saw the finishing. The silken rug had

“Habits and Structure,” page 101.

See ee Tr. S:hCUemSt””C~”

—_——" oa ee ei i Ab

166 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

first been spun upon the leaf, within which the egg mass was oviposited.
Over that the external blanket was woven, and when my observation
began the mother was engaged upon this.

The method of proceeding did not differ from that of other species
and tribes in like work. The spider grasped the margins of the cocoons
with the claws of her fore feet, which she continually moved around
the margin as she spun. The third pair of legs, and occasionally one of
the fourth, were also used for grasping the cocoon and moving the spider’s
body. The remaining fourth leg, and sometimes both the hind legs, were
used for drawing out the spinning
threads. As the spider thus swung
around her cocoon, heavy filaments
of silk were extruded from all the
spinnerets, which were opened up
and somewhat elevated. The long
jointed third pair, particularly, was
constantly lifted up and dropped,
as though beating in the silken tis-
sue, after the manner described in
the case of Argiope cophinaria when
making her silken shield. (Vol. I.,
Chapter VI., page 100.)

We may confidently assert that
the Territelarizee form no exception,
for I have fully observed their mode
of spinning the material which cor-
responds with the silken cocoon.
The silken rug on which our great
Tarantula rests, the tube of the
Purseweb spider, etc., are almost
certainly woven precisely as is the

- ———— cocoon of those species; and, if so,
Fic, 208. Agalena nevia engaged in covering her eggs; the Tunnelweavers spin their co-
use of the long spinnerets. .
coons as do other tribes.

Turning to the Wanderers, we have in the case of the Lycosids an ex-
ample, to which I have heretofore referred (Vol. II, page 144), of the man-
ner in which Lycosa fabricates her round cocoon. She first
weaves a circular patch, which she afterwards forms into a hol-
low sphere surrounding her eggs. The mode of equalizing the spinning
thread is as follows: The feet clasped the circumference of the cushion,
and the body of the animal was slowly revolved. The abdomen, greatly
reduced in size by the extrusion of the eggs, was lifted up, thus drawing
out short loops of silk from the extended spinnerets, which, when the
abdomen was dropped again, contracted and left a flossy curl of silk at

=
N
N
>
N

Lycosa.

COMPARATIVE COCOONING INDUSTRIES. 167

the point of attachment. The abdomen was also swayed from one side to
another, the filaments from the spinnerets following the motion as the
spider turned; and thus an even thickness of silk was laid upon the eggs.
The same behavior marked the spinning of the silken rug or cushion in
the middle of which the eggs had been deposited, It will thus be seen
that the entire process of forming a cocoon, as wrought by Lycosa, resem-
bles in every particular the mode practiced by 'lubeweavers and substan-
tially by Orbweavers.

So also is it with the Saltigrades. I have observed Phidippus rufus
spinning its cocoon, and she proceeds after the same general method. A Salti-
grade mother is represeuted at Fig. 205, as sketched in the act
of cocoon making. The diverging linés of silken spinning stuff
are there seen proceeding from the spinnerets, while the abdomen
is lifted up at a considerable incline, and the feet clasp the borders of the
cocoon, As this Phidippus revolved she alternately dropped and elevated
the abdomen, while the silken loops thus formed curled down into the
mass already spun and were further beaten in by the spinnerets and legs.
It thus appears from personal observation of typi-
cal species in all the tribes, with the exception of
the Laterigrades, that the manner of outputting
the spinning stuff while weaving cocoons is prac-
tically the same. The only difference observable
is confined to the use of the spinnerets in beating Fic. 204. Phidippus rufus spin
down the outspun threads, these organs being ee eS
more freely used for this purpose among the Tubeweayers and Tunnel-
weayers, who possess long pairs of superior spinnerets, than among others.

Salti-
grades.

WUE.

Proceeding now to a comparative study of the cocooning industry of
spiders, we observe, first, in view of the preceding sections of
General : pees
eininning this chapter, that the general method of spinning the cocoon,
Method, 28 it has been observed in representative species of all but one
of the tribes, is substantially the same.

1. That method consists in drawing out thickened lines from the spin-
nerets while the body is slowly: revolved around the area to be occupied
by the cocoon; or, as in Theridium, the cocoon is revolved upon a sus-
pensory line. The loops thus drawn out are about the length of one-half
the distance between the surface points to which the cocoon is attached
and the point to which the spinnerets are raised by the elevation of the
abdomen. As the spinnerets drop after their elevation, the thread relaxes,
curls, and thus a soft loop of curled thread is left upon the growing
cocoon mass. In some cases this is beaten down by the feet and spinner-
ets, or spread over by them as a plasterer spreads mortar, until the cocoon

oil iad, a ee

168 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

case is quite hard. In others, it is left in the flossy condition in which
it is originally spun.

2. While the general method of spinning out the material, as above
described, is that which prevails among all Tribes, the composition of the

. cocoon, or general plan of architecture, may be properly sep-
veaee) arated into three distinct modes. In the first the eggs are made
Cocoons. the centre of operations, being first laid upon a circular patch,

covered by a mass of continuous floss, and thereafter usually
enclosed within a seamless case of thickened spinningwork. The protec-
tion to the eggs is thus a single and unbroken covering. ‘This method
prevails among Orbweayvers and Lineweayers. :

In the second method the cocoon covering is spun in two parts. There
is first woven a sheet to receive the eggs, and after the eggs have been
overspun and swathed a second and similar sheet is made as an outer
covering. This method is the prevalent one among Tubeweavyers, Salti-
grades, and Laterigrades.

Cocoons classified under the second mode may be subdivided into two
well defined groups, viz., first, those in which the covering consists of pure
silk ; and, second, those in which the silken covering is strengthened or
padded by bits of gnawed bark, sawdust, and various light chippage,
daubs or pellets of mud, and sometimes by an entire coating of clay.
This mode of providing an armor of extraneous material is most preva-
lent with Tubeweavers, although it occasionally appears among Orbweay-
ers. Sometimes the armor or upholstery is itself covered over with an
exterior silken case, as with Agalena nevia; but again it forms the outer
casing or plaster, as with Micaria limicunee and Clubiona tranquilla.

The third special mode is that which prevails, one may say almost
universally, among the Citigrades, and which is probably practiced by the
Tunnelweayers also. It consists in spinning a single sheet, within which
the eggs are deposited, which is subsequently pulled over the egg mass,
and pinched by the jaws into a globular covering, the selvage of which is
united with sufficient firmness to adhere until the spiders are ready to
leave the cocoon, when the seam yields sufficiently to allow the escape of
the inmates.

3. A third point of comparison is as to the disposition of the cocoon
by suspension or attachment. We may divide the cocoons of all tribes
broadly into two classes, (I.) hanging cocoons and (II.) fixed
cocoons. The hanging cocoons may be subdivided into those
(1) which are suspended within the snare and (2) those which are
suspended or attached outside of or near the snare. The latter class may
again be divided into (a) those which are suspended with external pro-
tection and (b) those which are suspended without external protection.

Of spiders that hang their cocoons within the snare, the Orbweavers
have a number of representatives, as especially Epeira labyrinthea, Cyclosa

Dispo-
sition.

———- ti

COMPARATIVE COCOONING INDUSTRIES. 169

caudata, Epeira bifurca, Uloborus plumipes, Epeira basilica. Among
Lineweavers may be found most of the genus Theridium, as T. tepida-
riorum and T. studiosum, Steatoda borealis, the various species
of Erigone and Argyrodes. The Tubeweavers have numerous
representatives, as it is a quite general habit for the species of
this tribe to deposit their cocoons within the tubular portion of their
snare, which forms also a nest. Among the Territelarie Atypus has the
same habit, so also have the South American species described by M. Simon,
and the immense creatures known as the Mygalidwe appear also to nurse
their cocoons within their burrows. This is the custom of our well known
‘Trapdoor spider, Cteniza californica. Among the snareless Wandering spi-
ders, Citigrades, Saltigrades, and Laterigrades, of course, there are no rep-
resentatives of this group.

Spiders that hang their cocoons outside their snares are largely repre-
sented among Orbweavers. Indeed, this may be said to be a general habit,
as most of such genera as Epeira, Argiope, Zilla, Acrosoma, ‘Tet-
ragnatha, Nephila, Meta, and Hyptiotes spin their cocoons sep-
arately from their snares. The habit prevails also among Line-
weavers, as is illustrated by the habit of Theridium frondeum in swinging
her pretty little orange colored cocoon to the under sides of leaves and
the surfaces of rocks. Among Tubeweavers, Segestria canities of California
suspends her string of clustered cocoons outside her nest, although she
does subsequently spin a tubular cell alongside the cocoon string, and
there dwells while completing the process of cocoon making and while
the young are being reared. ‘Tegenaria medicinalis also suspends her co-
coon most frequently to some object, as a log, or beam, or branch, out-
side of her snare, although sometimes she hangs it to the lower portion
of the sheeted pouch itself, or even interweaves it within the fibres of
the sheet. Among Citigrades, the Southern species, Pucetia aurora, and
all the known species of the genus Dolomedes swing their cocoons not,
indeed, outside their snares, since they are Wandering spiders, but in
special nests prepared for the purpose.

The spiders which attach their cocoons to fixed surfaces, instead of
swinging them among interlacing lines or suspending them with-
in their snares, are numerous and have representatives among
nearly all tribes. Such is the habit among Orbweavers, in most
of the genera, as Epeira, Zilla, Gasteracantha, Nephila, ete.

Among Lineweavers, species that dispose of their cocoons in this way
appear to be rare, although such a European species as Theridium denti-
culatum has this habit.1 Among Tubeweavers, the great mass of spiders
of all or nearly all the genera fasten their cocoons to various surfaces.
These plano convex objects may be seen in the autumn, for the most

Within
Snares.

Outside
Snares.

Fixed
Cocoons.

1 Staveley, “British Spiders,” page 147.

—

— = OO

170 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

part, attached to the under side of stones or spun within rugosities of the
bark of trees. They have often bright colors, and are found covered
with mud. Among Saltigrades and Laterigrades, all species appear to
have the habit of thus disposing of their cocoons.

IV.

4. A fourth point of comparison is the method of protecting cocoons.
When we come to consider the modes of protecting cocoons, we find much
variety, and the various methods well represented among all
Protec- ‘Tribes, modified by differences in habit characteristic of the
ag groups represented. The chief modes of protection are, first, by
Tinan, lines, within which the cocoons are spun and which form an
interlacing barrier of threads around them. This mode has many
representatives among Orbweavers and Lineweavers; is occasionally repre-
sented among Tubeweavers, as with Dictyna and Segestria; occasionally
among the Citigrades, as Dolomedes and Pucetia; but is unknown among
Saltigrades and Laterigrades.
The second mode of protection is by leaves, which are drawn over the
cocoons. ‘This is either done by attaching the cocoon to a single leaf and
then curling the edges thereof around it, or by forming a sort
aes of bower of several leaves united at the points and edges, and
spinning within this cavity a maze of interlacing lines, within
which the cocoon is hung. This method of protection is well represented
among Orbweavers. Among Lineweavyers it is seen in our Theridium dif-
ferens, and in certain European species of Theridium, as T. nervosum,
T. riparium, and T. lineatum, Among Tubeweavers it largely prevails, Aga-
lena, at least, practicing this method. Dolomedes represents the Citigrades,
making a beautiful bower, within which her cocoon is hung. The Salti-
grades frequently thus protect their cocoons; and among Laterigrades,
Thomisus, Sparassus, and Philodromus.
The third method of protection is by silken tents and tubes. This is
sparingly practiced among Orbweavers. Among Lineweavers, as far as I
know, it is only used when the silken tent is enclosed within an
aed and Cuter covering of leaves. Among Tubeweavers it has a wide use,
Tubes, Herpyllus, Ariadne, Drassus, Clubiona, and others of like spin-
ning habit practicing it probably without exception. I know of
no Citigrades that thus protect their cocoons, unless we consider the burrow
and home as a cocoon tent or den, and it certainly does serve that end.
But among the Saltigrades the habit is general, Phidippus, Attus, Syna-
gales, Synemosina, and all known genera protecting their cocoons with an
outer silken cell, within which the mother dwells. Among the Lateri-
grades, Thomisus and Sparassus have the same habit.
The fourth method of protection is by an armor of extraneous ma-

Se
: :

COMPARATIVE COCOONING INDUSTRIES. rel

terial, such as insect débris, plant chippage, sawdust, sand, and mud.
Among Orbweavers that practice this method of armoring their cocoons
are Epeira cinerea and Cyclosa caudata. I know no American
representative among Lineweavers, but there are no doubt such,
as some English species have the habit. Among Tubeweay-
ers the habit is most common. The cocoons of Agalena are frequently
found upholstered with sawdust and scrapings from bark, or bits of chip-
page plucked from surrounding vegetation. Clubonia frequently plasters
over with mud her beautiful white cocoon. Micaria limicunz completely
encloses her little egg sac within a thick ball of mud. Others of this
family make a spherical ball, composed of miscellaneous débris, within
the heart of which the cocoon is protected. The habit appears to have
secured no lodging among the Citigrades and Saltigrades, but is practiced
to a limited extent by some Laterigrades.

A fifth mode of protection is suspension within the snare. Some Orb-
weavers have this habit, as the Labyrinth spider, the Tailed spider, and

others. Many Lineweavers in the genera Theridium, Argyrodes,
5. Sus- ete,, thus protect their cocoons. Indeed, it is the well nigh uni-
pension versal method in this tribe. Among Tubeweavers the custom
in Snares. F fost j ‘

prevails, that is, if we consider the tubular dwelling cell as a
portion of the snare. The same remark applies to the Territelariz, as
Atypus, Cteniza, and the large Mygalide, who protect their cocoons within
their tubular dens, as do the Saltigrades also. The Citigrades and Lateri-
grades, of course, have no such habit, as they are not snare making tribes,
although they make a cell or den to contain and shelter their cocoons.

The sixth method of protecting the cocoons is by sentry, that is to say,
by watching on or near the cocoon—an action to which the term “ brood-

ing” has sometimes been applied. Of this method, the Orb-
atone _ Weavers have representatives among the genus Epeira, as, for

example, our American Epeira cinerea and several European
species, to which may probably be added Cyclosa and Uloborus and others
of like habit. Nearly all Lineweavers may be considered as_ practicing
this method. At all events, their cocoons are swung within their snares
and the mothers are frequently found embracing them and vigorously re-
sist any effort to deprive them of the treasure. Among Tubeweavers,
many genera keep faithful watch on their cocoons, as Agalena, Herpyllus,
and many of the Drassids. ‘The Tunnelweavers, who retain their cocoons
within their burrows, may be regarded as keeping sentry upon them, and
the habit is quite general among Saltigrades and Laterigrades.

The seventh mode of protection is by portage, that is, by carrying the
cocoons within the jaws or attached to the spinnerets. I know no rep-
resentative of this habit among Orbweavers. Among Lineweavers the
habit is universal with Pholeus, who carries her cocoon beneath her jaws.
At least one European species of Linyphia and one of Theridium have

4. By Ar-
morage.

i, ‘ -
ee ee eee ee ,

UTZ AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the habit of deporting their cocoons, like a Lycosid, tied to her spinner-
ets. No Tubeweavers are known to carry their cocoons. Among Citigrades
the habit is universal with Lycosids, and prevails with Dolo-
7. By medes until about the time when the eggs are ready to hatch,
satiate when she discards the cocoon and deposits it within a prepared
of Stones, Lest. Ctenus probably resembles Dolomedes in this peculiarity.
The Territelariz appear to carry their cocoons, although not to
the same extent as the Lycosids, owing to marked difference in mode of
living. None of the Saltigrades or Laterigrades are recorded as deporting
their egg sacs, except Heterapoda venatoria, who does so occasionally. The
eighth method of protection is sheltering cocoons beneath stones, bark,
ete. This is in use among all the tribes, with the probable exception of
the Tunnelweayers.
The ninth and last mode of protecting the cocoon is by simple suspen-
sory lines. This seems to be the simplest form of protection, and is
doubtless effective against such enemies as would be apt to find
9. By the eggs if placed upon a flat surface, but who would not venture

a se to assault them if compelled to creep along a fragile thread.
Tinese Among Orbweavers few species are known, but there are some

European representatives of this habit. Cyrtarachne may be
considered as thus providing for her cocoon. Among Lineweavers there
are a number of representatives, our most conspicuous American one being
Theridium frondeum. Of the Tubeweavers, Tegenaria medicinalis, at least
occasionally, thus disposes of her cocoons; and among European species
are Agalena brunnea and others that hang their flask like egg sac from
heather and other plants by means of a foot stalk. The other tribes, as
far as I know, do not swing their cocoons free in this manner.

The accompanying table will show at one view the comparative preva-
lence of these various modes of protection among the tribes, as far as
my observations and notes permit. It will be seen from this view that
all the modes of protection, with the exception of portage, prevail among
Orbweavers and Tubeweavers; that all the methods are represented among
Lineweavers, although some of them appear to be faintly developed and
sparsely represented therein.

5. A fifth basis of comparison is the form of cocoons. The greatest va-
riety obtains among Orbweavers. Round cocoons, hemispherical or plano
convex, pyriform, or stalked cocoons, with various modifications,
as illustrated in the accompanying group, are the forms that
commonly prevail among American Orbweayers. A comparison
with the cocoonery of European and exotic Orbweavers, as far as they are
known, shows that there is a substantial likeness between them and the
American spider fauna.

Cocoon
Forms.

173

COMPARATIVE COCOONING INDUSTRIES.

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*‘suadldg {0 SHdIayT, SQOLLVA
GHL dO VHUANGY) IVOIdAT, SNONV SNOOOOD) SNILOMLONT JO SAGOJT LNATVARYT SNIMOHS AIAV I,

Fig. 211. Fia. 212. Fig, 213. Fic. 214. Fig. 215.

‘ - LINEWEAVERS.

Fig. 216. Fig, 217. Fic, 218. FiG. 219. Fie. 220. Fic. 221. Fig. 222,

= 5 TUBEWEAVERS.

iG, 225. Fic. 226. Fia. 227.

TUNNELWEAVERS.

a Fic, 228. Fic. 230.

ba LATERIGRADES. SALTIGRADE. Crrigrapr.

Comparative map of aranead cocoon forms.
ta # <

COMPARATIVE COCOONING INDUSTRIES. 175

Next to Orbweavers, the Lineweavers exhibit the greatest variety of
form. A round or ovoid cocoon is the prevalent form, but the pyri-
form is well represented in this tribe. Among Tubeweavyers the almost
universal form of cocoon is the plano conyex or hemispherical. This re-
sults from the quite general habit of attaching the egg sac to the surface
of some object. In some cases, however, Tubeweavers suspend within their
nets a double conyex cocoon; and, again, hang to the foliage or other
surfaces a pyriform cocoon, as in the case of the European Agreeca brunnea.
Among Tunnelweavers there is apparently but one form, as is indicated
by the cocoonery of the few species known, This cocoon is a round ball
and is in every respect like, or at least closely resembles, that of Citigrades.

The Citigrades also have apparently one form, a globular silken case
within which the eggs are enclosed with little or no padding. In numer-
ous species of Lycosa, Dolomedes, Ctenus, etc., this form prevails. Among
Saltigrades, also, there is apparently but one form, a hemispherical or plano
convex cocoon, attached to some surface, the case being enclosed within a
soft, flossy, or thick netted covering of spinningwork. Among Laterigrades
there is greater diversity than among the last three mentioned Tribes. But,
for the most part, the cocoons consist of stiff hemispherical cases attached
to surfaces of rocks and trees; occasionally, however, as in the case of
Philodromus and some species of Thomisus, the cocoon is a double con-
vex covering hung between leaves or twigs.

It is thus observed that the greatest variety and complexity of cocoons,
as to form and structure, are to be found among the Sedentary tribes. The

very greatest is in the Orbweavers, where the variety of form is

iach Aah remarkable. Next in order are Lineweavers, although it is pos-
plexity. ‘ible that, if a wider study of this tribe were made, they might

be found to approach more nearly the Orbweavers in this re-
spect than we are justified at present in asserting. The Tubeweavers
follow in order. The Territelariee are classed ordinarily with Sedentary
spiders, and many of the species fully justify this classification, since, like
Atypus, they persistently dwell within their tubes. But they have also
many of the characteristics of the Wanderers, and therefore we find their
cocoons approaching those of Citigrades in simplicity of form.

In the comparative chart printed upon the opposite page I have tried to show at one
view the typical forms of cocoons known to be made by representative genera of the va-
rious tribes. The following is the explanation of the chart: Cocoontnc Forms or Orsweay-
prs: Figs. 205, 206, Epeira; 207, 208, Argiope ; 209, 210, Cyrtarachne ; 211, Epeira labyrinthea ;
212, Epeira bifurea; 218, Tetragnatha; 214, Uloborus; 215, Cyclosa caudata. LinzwEavers:
216, Argyrodes trigonum; 217, Theridium frondeum; 218, Steatoda and Theridium; 219,
Theridium; 220, 221, Theridium; 222, Pholeus. Tusrwravers: 223, 224, Agalena, Drassids ;
225, Segestria ; 226, Micaria limicune; 227, Tegenaria. TuNNELWEAVERS: 228, Mygalide, Eury-
pelma; 229, Atypus; 230, Nemesia. Larrricraprs: 231, Thomisus, XNysticus, and many
genera; 232, Heterapoda and others. Sauricraprs: 233, Attus, Phidippus, and all genera.
Crriarapes: 234, Lycosa, Dolomedes, and all known genera.

ii RO hg

176 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

The greatest general simplicity of structure appears among the cocoons

of the Territelarie, Citigrades, and Saltigrades, and the Laterigrades nearly

approach them in this combination of simplicity and uniformity.

stoi It may be said that the tribe which shows the greatest simplicity

plicity, 22d uniformity of cocoon structure is the Citigrades. The in-

ference may therefore be drawn, that the greatest general sim-

plicity of structure exists among the cocoons of those spiders which have

them most closely under their personal care. It is manifest that in the

case of Lycosa and other genera that attach their egg sacs to their spin-

nerets and carry them about until their young are hatched, there is less

necessity for complex cocoonery to protect the enclosed eggs than in the

case of Orbweaving spiders, like Epeira or Argiope, who hang their cocoons
in the shubbery and leave them to the watch care of Nature alone.

While this deduction is justified in the general view of the subject, it
must be allowed that there are some exceptions which cannot well be
explained. For example, the two cocoons which have absolutely
the simplest structure are made by members of the Retitelarie,
as Pholeus phalangioides and Steatoda borealis. The egg bags
of the latter species consist of a mere pinch of silk of such sparse weft
that the eggs are plainly seen through them. Pholeus, who carries her
cocoon underneath her jaws, while she hangs continually upon her snare,
holds her eggs together by little more than a netted bag of scant spin-
ningwork.

One who examines, even casually, these various forms will see that they
are determined substantially by the fact that the eggs, as they are extruded,
naturally form a spherical or hemispherical mass, according as
they hang free or are oviposited against some surface. Around
this mass the protecting spinning stuff is woven, and then the
external case. The addition of a foot stalk, more or less pronounced, ap-
pears to be determined by the act of suspending the cocoon during the
weaving thereof, and the subsequent covering in and thickening of the
suspensory cord so that the texture corresponds with the remainder of the
outer case. ,

The little conical or pointed processes which characterize several cocoons,
as those of Tetragnatha and Uloborus, probably originated in the same way,
namely, by the attachment of suspensory or broken threads to various points
of the external surface, the points of attachment being thickened into little
puffs or rolls or points of spinning stuff.

The introduction of extraneous material as an additional protection and
the encasing of the silken sack in mud, as with Micaria limicune, is a
habit to be accounted for altogether outside of the above; but the fact
that these mud protected cocoons preserve the general form of the spin-
ningwork which encloses the eggs, is undoubtedly determined by the same
causes that regulate the shapes of all other cocoons.

Excep-
tions.

Origin of
Forms.

eS lt(“‘“

i i i ik le al

COMPARATIVE COCOONING INDUSTRIES. iPr

Vi.

6. A sixth basis of comparison is the multiplex cocoonery of certain

species. The general habit among spiders is to make but a single cocoon

at a time, and most females probably limit their maternal duty

So to the production of one egg sac. But there are numerous ex-

coons. &Pptions, which have been noted. Among Orbweavers the Laby-

rinth spider, the Tailed spider, the Basilica spider, and some

others habitually produce several cocoons. These are not made contempo-

raneously, but are spun consecutively, with intervals of several days be-

tween each cocoon, so that the younglings will be hatched from the first
brood while the last is yet freshly laid.

It is to be noted, also, that even those spiders that ordinarily limit
themselves to one cocoon, as Argiope, under certain conditions, which are
not fully understood, produce two or more cocoons. Epeira, when specially
nourished, is said to produce several. The fecundity of the spider may
therefore be said to be subject to variation, and the disposition to multi-
ply cocoons is dependent, more or less, upon the fecundity.

Among the Retitelarize numerous species are found spinning several
cocoons, the most familiar example being Theridium tepidariorum and
Latrodectus. The Tubeweavers also have some remarkable representatives
of multiplex cocoonery, as, for example, certain species of Dictyna and
Segestria. The cocooning habits of the Territelaris are so little known
that one cannot speak positively, but it is probable that no Tunnelweaver
makes more than one cocoon. ;

Among the Wandering spiders the single cocoonery which characterizes
the Tunnelweavers is the rule. I know no Saltigrade and no Laterigrade
that produces more than one cocoon, although of the former Staveley says
that Epiblemum scenicum makes one or two, and of the latter that Philo-
dromus czespiticolis deposits two flattened cocoons in a large nest.1 Among
Citigrades I know no species except Pucetia aurora; this spider produces
at least two cocoons, that are concealed within a little nest of crossed lines,
very much after the fashion of that constructed by Dolomedes. No doubt,
however, a wider knowledge will compel us to include other species in
this group.

This summary of facts points to these conclusions: First, that the three
Tribes which are by especial eminence Sedentary possess the greatest
number of species that make more than one cocoon. Second, that the
three Tribes that are conspicuously Wanderers make but one cocoon, with
rare exceptions. Third, that the Tunnelweavers, whose habits sometimes
approach one group and sometimes another, but in the matter of cocoonery
resemble the Citigrades, as regards multiplex cocoonery are to be classed
with the Wanderers, apparently limiting themselves to a single egg sac.

and 86.

1 “Brit, Spiders,” pages 5

ee, Ee

. OHAPTER GL.

MATERNAL INSTINCTS : MOTHERHOOD.

In the chapters immediately preceding I have described the various
devices and forms of spinning industry prompted by maternal instinct
for preserving offspring. Apart from this—the mere industrial or archi-
tectural expression of motherhood—there are some facts in the natural
history of the maternal habit which may perhaps best be considered in a
separate chapter. Such, for example, are the motives which regulate the
choice of a cocoon site; the methods of ovipositing; the measure of ma-
ternal purpose as taken from the complexity, isolation, or vigil of the
cocoon; the causes regulating the number of cocoons and eggs; the motive
controlling the armoring and mud plastering of cocoons; brooding the
egg nest; the degree of and conditions limiting the maternal anxiety for
the eggs; and the intensity and intelligence of the maternal sentiment.
These are points of the greatest interest to all naturalists, and are well
worthy of a far more extended and philosophic treatment than I feel com-
petent to give. But it may be permitted me at least to open the way.

The sites which spiders choose for their cocoons are, of course, largely
determined by their habitat. The cocoons will always be found near by
the locality in which the mothers haye lived. Although some
of them do occasionally move from their native centres, the
migration is, as a rule, extremely limited; and Orbweavers, in-
deed all Sedentary spiders, may be considered as practically spending
their lives within the narrow compass of the spot where they chance to
pitch their first snare.

The favorite sites of Orbweavers are bushes, low trees, grass, weeds,
the angles of walls in the neighborhood of houses and outhouses, and like
situations which afford them facilities for hanging their snares. They are
frequently exposed to the full blaze of sunlight; some species appear to
love the most open exposures in woodlands; others, again, shun the sun-
light and are found in woods and forests, in obscure corners, hollow
trees, clumps of underbrush, and even, as in the case of Meta, in caves.
They hang their nets along the banks of streams, in glens and ravines,
on the seaside, on the lowest plains and prairies, and on the tops of the

highest mountains, as far up at least as the timber line extends. I have
(178)

Cocoon
Sites.

MATERNAL INSTINCTS: MOTHERHOOD. 179

taken them on the highest railing of the dome of St. Peter’s in Rome;
have seen their round webs swinging against the cliffs of Mosquito Mount-
ain Pass in Colorado, more than ten thousand feet high; have found them
upon the mountains of Scotland; and captured the British Epeira umbra-
tica from snares spun against the basaltic columns of Fingal’s Cave.

Their fixed positions are, of course, determined by their ability to
obtain food therefrom; and, as their food is insects, the limit of insect

life must also be the limit of spider life. For, although spiders
Food : :
Limits, 2° frequently at the mercy of the winds and are carried great
distances when they are young, during the aeronautic stage, they
cannot long sustain themselves and propagate their species if they chance
to fall upon positions where it is difficult or impossible to obtain generous
supplies of insect food.

In seeking a spot upon which to place their cocoons, most Orbweavers
go a little distance from their snares and construct the cocoon against the
outer surface of a bush or tree, rock or wall, or cover it up
within a leaf. As a rule, the disposition to find a secluded spot
is quite manifest, but there are many exceptions. Other species
deposit their cocoons within their webs, stringing them along one of the
radii of the orb, as in the case of Uloborus, Epeira caudata, and Epeira
bifurea; or suspend them within a maze of crossed lines which overhangs
the orb, as in the case of the Labyrinth spider. Others, again, as with
Argiope, will frequently swing their cocoons within a specially prepared
mass of crossed and netted lines, which are hung to branches or boughs,
leaves, or blades of grass.

What is said of Orbweavers as to cocooning site is substantially true
of the other Tribes, with, of course, such variations as are required by
essential differences of habit and structure. For example, those Seden-
tary spiders, as the Lineweavers, which suspend their snares in positions
quite like those of Orbweavers, also follow closely that Tribe in the gen-
eral principle of selection for cocoon sites. In other words, they ‘hang
their cocoons in some part of their snare, or somewhere near, hidden be-
neath a convenient cover, or in a neighboring retreat.

So also many Tubeweavers, and the Tunnelweayers even more persist-
ently, attach their cocoons to some part of their web, or weave one of their
characteristic tubes around the egg case when it is once spun. In these
cases the cocoon site is pretty sure to be identical with the dwelling place
and snare. —

Among Wanderers the home site has less influence upon the cocoon
site. As these animals pursue their prey over a more or less extended
range of territory, the site of the cocoon is dependent on the
place where the hour of maternity may overtake the females.
Wherever they happen to be, the Saltigrades and Laterigrades
will spin a tubular tent, enclose within it their cocoon, and there remain

Cocoon
Secreting.

W ander-
ers.

es a eee

——————=. Or

180 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

until the young are hatched. However, it must be said that, with Salti-
grades at least, there is a tendency before cocooning to prepare a perma-
nent dwelling tent, to which, when the proper time approaches, the mother
will resort to deposit her eggs. Lycosids also strongly incline to spin and
burrow a cocooning house after their kind. But inasmuch as they deport
their cocoons, they are apt to move about from site to site with their egg
bags dangling at their tails, stalking prey and bivouacking in any con-
venient refuge. :

Wt,

I infer that female spiders habitually prefer the night or early morn-
ing hours for cocooning. At least I have never been able to observe any
: species laying eggs, although I have frequently and quite per-
redikl sistently watched, both in artificial and natural sites, with a view
Cocoon- : : Prue Siete
ing. to such observation. I am satisfied that it is within the power
of the female to control the maternal function and compel Na-
ture to await her pleasure for a considerable length of time. IT cannot
otherwise well account for some experiences with my captives. Moreover,
I have spent many days during the last fourteen or fifteen years in wan-
dering among haunts of spiders, north, south, east, and west, in our own
country and Europe, but have never
once surprised a female in the act of
ovipositing. his leads me to the con-
clusion that spiders must commonly
choose the night or early morning as
the time for laying their eggs.

Others, however, have been more

: fortunate ; and, judging from their ac-

Fia. 235. Fia. 236. : . *

Fic. 285. Section views of abdomen, to show loca- counts, and reasoning, from the varl-

tion of eggs. Te. 236. Same, with eggs removed. og stages at which I have partially

(From alcoholic specimen.) ;

observed the process, by putting the

pieces of observation together, we obtain a tolerably accurate idea of the
mother spider’s mode of procedure.

Just before cocooning, the eggs will be found massed within the centre
of the abdomen, the ovaries being so greatly distended as to compress and
somewhat displace the surrounding and adjacent organs. (Figs. 2385 and
236.) They are in this state gelatinous bodies, but have a spherical shape
even in their soft condition. They are still jelly like objects when ex-
truded from the ovaries along the vulval hook or ovipositor, and do not
harden until shortly after they are laid.

When the mother is prepared to drop her eggs, and has satisfied herself
as to locality, the next step is to prepare either a little sheet, or dish shaped
dish, or a flossy tuft of spinningwork, against which the eggs are posited.
I believe that this is most frequently done upwards in the case of females

Le
Cae
ae 1a 53D
‘see ate
98, ae “9

SS ss S Se el,

MATERNAL INSTINCTS: MOTHERHOOD. 181

who swing their cocoons free, as Argiope and Theridium; that is to say,
the spider hangs with her back downward while ovipositing. But in many
cases of females that have cocooned for me in boxes, the eggs must have
been placed in the reverse position, since the
cocoon was attached to the bottom of the box.

Of course, the species that fasten their
cocoons to yarious surfaces, as do many

manifestly been placed upwards, as, for ex-
ample, those spun on the under surface of stones, fallen logs, ete. Others,
again, have been laid while the spider was in a vertical position, as when
cocooning upon loose bark of trees and similar vertical sites. The bodily
attitude appears to make little or no difference as to the facility with
which the female can deposit her eggs. Whether directing them upward
(with the dorsum towards the earth), or directing them downward (with
the dorsum towards the sky), or depositing them against a vertical surface,
with the head downward or upward, as the case may be, the mother is
able to empty the ovaries with equal comfort and ease.

Mr. Emerton has observed several species in the act of ovipositing, and
his brief notes upon the manner thereof are as follows:! Epeira strix first
spins a rounded bunch of loose threads, into the middle of which
she discharges her eggs, as shown in Fig. 287. The eggs, which
are little drops of jelly, are held up by the loose threads until
the spider has time to spin for them a covering of strong silk. It is to
be regretted that the description here is so indefinite, as the term ‘“ coy-
ering of strong silk” may imply either the flossy boll which is invari-
ably found to surround the egg mass of Epeira, or the smooth textured
silken bag which immediately encloses the eggs and against which the
flossy blanketing is laid. When a cocoon of Epeira strix and others of
similar habit is cut open, this silken encasement is invariably seen, and
it presents the appearance of having been
the original substance against which the
eggs were directly laid.

The same author has been fortunate
enough to observe the mode of positing
eggs with two other tribes. The female
Drassus (Fig. 238), spins a little web
Fra, ier Baan eee in seta of drop- across her nest and drops the eggs upon

: ; it. They are soft, and mixed with liquid,
and are discharged in one or two drops, like jelly. They quickly suck up
the liquid, and become dry on the surface, sometimes adhering slowly

Ovipos-
iting.

' Habits and Structure, page 101.

182 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

together. After the eggs are laid, the spider covers them with silk, draw-
ing the threads over from one side to the other, fastening them to the
edges of the web below. When the covering is complete, she bites off the
threads that hold the cocoon to the nest, and finishes off the edges with
her jaws.

Phidippus galathea (Attus mystaceus Hentz) spins, before laying her
eggs, a thick nest of white silk, usually on the under side of a stone. In

this she thickens a circular patch on the side next the stone, and
ote discharges her eggs upward against it. (Fig. 239.) They adhere,
Ane te and are subsequently covered with white silk, after the manner
common to Saltigrades. Mr. Emerton had a female of this species that
deposited her eggs in confinement; he records that, ‘instead of completing
the cocoon properly, she ate the eggs immediately after laying them,”! a
breach of maternal fidelity which I believe to be rare among araneads, even
when cocooning in the unnatural conditions of a forced imprisonment.

The eggs are deposited in a mass, cylindrical, conical, or hemispherical,
individuals of which are usually fastened together by a glutinous sub-

stance, but sometimes are deposited loose, so that they roll about
nsdn in the hand when the cocoon envelope is cut. We are indebted
ae to Menge for the following interesting observation: After all the
eggs are deposited the spider rests for a season, when she commences to
draw threads over the eggs, as if desirous of
covering them up; but it soon becomes clear
that something else is to follow. After a while
she returns to the cocoon and discharges a
cs ue rants re ta clear liquid over the eggs, which is absorbed
laying eggs within a silken cell. (After by them without in any way interfering with
meta the web. This causes the eggs to swell to
such an extent that they could no longer be contained within the animal.
Menge thinks that this fluid proceeds from the semen pockets, which at
this period are very much enlarged, and becomes mixed with the
male semen, so that in reality the fructification of the eggs is
completed by the female. he mother now appears very much
exhausted. She lays down for a while on the eggs, and, finally, com-
mences to spin them over, entirely covering them.?

Mr. Moggridge had the opportunity to observe the eggs laid by a
specimen of our Cteniza californica, which was sent to him from America
and kept for a while in captivity. The eggs were deposited in
several clusters, at various times, upon the under surface of a
gauze fastened upon the mouth of the box in which she was
imprisoned, The first of these groups was laid See the night, between

Fructifi-
cation.

Cteniza’s
Eggs.

1 Struatin, and Habits, pages 99, 100.
* Menge, “Preussische Spinnen.” The author adds “that it takes patience and perseyer-
ance to observe the spider during this entire process, and he had only succeeded twice.”

MATERNAL INSTINCTS: MOTHERHOOD.

Fig. 240. A, Cteniza californica; B, her trapdoor nest; C, group of eggs, natural size;
D, same, magnified; BH, a second group, magnified; F, the same, largely magni-
fied. (After Moggridge,)

183

184. AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the 12th and 13th of July, in a cluster shaped like a raspberry. The
eges were grayish white or pale brown, and varied in shape from globose
to oblong. All were very small, the largest one half a line in its greatest
length.

A fortnight later, July 27th, another cluster of eggs was laid, this time
between the hours of 5 and 8 P. M. When the lamp was brought in at
the latter hour, Mr. Mogeridge perceived what he took to be a drop of
water hanging from the gauze covering, above and rather in front of the
spider’s door, the position occupied by the clusters of eggs previously
described. On closer inspection this proved to be a drop of pellucid,
colorless liquid, in which some thirty eggs floated. One egg was laid on
the gauze at some distance from the main group, and several were also
attached to the inside of the tin box. At midnight he found that the
drop had coagulated and contracted, and by the following morning the
mass was quite dry and resembled the former group, only that it was
not quite so convex. Some of the eggs forming these clusters were much
larger than in the preceding one, and one measured as much as a line
in length by half a line in breadth.

Between the above date and the end of November, when the spider
died, eggs were laid on seven distinct occasions, namely, on July 81st,
August 11th, 15th, and 31st (when he found the eggs floating on a drop
of liquid, having been deposited on the gauze between two and _half-past
four in the afternoon), September 9th (twenty-three eggs laid on the
earth near the entrance to the nest), September 19th (about thirty eggs
on the gauze), November 4th (about thirty eggs on the gauze). Thus,
between July 13th and November 4th, this spider laid nine clusters of
eggs, all but one of which were placed on the same part of the gauze
cover, above and a little in front of the door, apd the total number of
eggs deposited cannot have been less than two hundred and fifty.1

Of course, it is difficult to account for the peculiarities of this female
in oviposition, for there is little doubt that this manner of laying eggs in
disconnected groups, at extended intervals of time, is quite foreign to the
usual habit of the species. During the long journey from her native home
she may have experienced a shock resulting in a morbid condition of the
ovaries. Undoubtedly, like her congeners, of whom Mr. Eugene Simon
gives an account (see Chapter V.), Cteniza californica lays her eggs in one
mass, and suspends them within her burrow. But the above facts at least
show the power of the female to control the function of ovipositing, and
indicate that there are certain irregularities in that function, more or less
under the control of the female, which may give a clue to the habitual
production by certain species of several cocoons, and the occasional multi-
plication of cocoons by other species.

* Moggridge, “Trapdoor Spiders,” Supplement, page 203 sq.

MATERNAL INSTINCTS: MOTHERHOOD. 185

When the eggs are laid the spider mother proceeds to spin the outer
envelope by which they are protected, and within which the progeny, when
hatched, may find a comfortable home until sufficiently matured
to begin life for themselves. This external structure differs,
among various species, in shape, size, interior arrangement, and
more or less in the character of construction. The details of these points
have appeared in the preceding chapters, and they form some of the most
interesting features in the life habits of araneads. The cocoon may be
described in general terms as consisting of a silken sheet or sac surround-
ing the eggs, a padding of greater or less compactness above that, and a
case of a more or less compact texture surrounding the whole.

The shape of the cocoon appears to have no special relation to the
maternal instinct, but is probably regulated by the habits of the particular
species and the character of the cocoon site chosen. It has already been
seen that the forms, although at first view they seem to be quite varied,
may, by analysis, be reduced to the round or hemispherical. In other
words, the eggs, as they drop from the spider’s ovaries, naturally assume
a more or less rounded form when the cocoon swings free; and when ex-
truded against a fixed surface as naturally form into a hemispherical mass.

This is simply the result of the law of equilibrium. As the maternal
care is directed solely to covering up and protecting the eggs, the shape
of the egg mass inevitably regulates the shape of the spinningwork woven
around it. It thus would seem that the maternal purpose is shown in the
fact of enclosing the eggs within the cocoon, and not in the external shape
which that cocoon assumes.

However, a measure, of maternal interest and intelligence is undoubt-
edly found in the architectural details of the cocoon. I have shown (Chap-

_ ters IV. and V.) that these have a tolerably wide range; that
aaah oes some cocoons.are extremely simple in their structure, and others
Structure (uite complex. To what degree are these differences regulated

by maternal affection and intelligence? This question cannot be
considered wholly from the standpoint of the cocoon structure itself, for
other elements enter into consideration, as the natural environment chosen
for a cocoon site, or the artificial environment prepared for it. That is to
say, a cocoon may be quite simple in its structure, having little spinning-
work to directly enclose the egg mass, but, as in the case of Dolomedes,
have a supplementary protection of a leafy tent, and an associated en-
closure of intersecting lines, which add materially to the protection of
the eggs. Of course, in thinking upon the degree of intelligence and
affection exhibited by such a mother, the external protection must be an
important factor,

What is the relation between the simplicity or complexity of a cocoon’s

Shape of
Cocoon.

er re ee ee

NN a a ee jh) gg

186 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

construction, and the amount of care which the mother gives it? ‘There
is much difference in the extent of elaboration of cocoons. The simplest

construction of which I have any knowledge is that of our com-
Complex- mon cellar spider, Pholeus phalangioides, which surrounds its
nt i ng a little cluster of agglutinated eggs with the barest filament of
Gare, silk through which the eggs are entirely visible. This rude co-

coon the mother holds underneath her jaws, and there carries it
until the spiderlings are ready to hatch out, when they take their place
upon the straggling lines at the top of the maternal snare. Steatoda bore-
alis spins a cocoon scarcely more elaborate than the above; she hangs it
within her snare of crossed lines and stays near it. The cocoons of Lycosa
and Dolomedes are also carried about by the mothers until they are hatched
or nearly ready to hatch. These cocoons are rather simple in structure,
consisting of a patch of spinningwork rolled up into a ball, without any
internal padding or protection whatever. The cocoons of many Tube-
weayers, the Drassids, for example, and the cocoons of Laterigrade spiders
are simple parchment like textures, spun against a surface, and are also
free from any internal padding or external protection.

These spiders are in the habit of watching their cocoons, remaining
near them until the little ones are hatched. Thus far it might be said
that there is some reason for the conclusion that lack of complexity in
the structure of a cocoon is supplemented by additional vigilance on the
part of the mother in watching the cocoon.

Let us see how it is among Orbweavers. The most complex cocoons

are found among these spiders. That of Argiope, for example,
ee exhibits remarkable regard for the protection of eggs and young,
rb- F 5 ; ; : ;
weavers, Py its tough external case, its thick lined padding of brown silk,
which nearly surrounds the egg mass, and the sac which con-
tains it. Argiope, as far as known, never watches her cocoon.

The same is true of most species of the genus Epeira, whose cocoons
are frequently enclosed within a tent of sheeted spinningwork or of closely
laid lines, and are themselves composed of several layers of spinningwork
of various textures. Most spiders of this genus give their cocoons no care
after they have made them. ‘There are, however, exceptions. Epeira cine-
rea, for example, not only encloses her eggs in a well furnished cocoon,
but adds to it scrapings from the bark of trees or the dry wood surface
upon which the cocoon may be fastened. Yet, according to Mrs. Mary
Treat, this spider is extremely watchful of its cocoon.

Cyclosa caudata provides the ordinary enswathment for her eggs, and
adds to that an exterior armor of the disjecta membra of in-
sects captured by her. Yet these cocoons are hung within her
snare, and during the cocooning season she is found constantly
clinging to the end of her cocoon string. However, that this contiguity is
an actual vigil is not proved.

Cocoon
Vigils.

MATERNAL INSTINCTS: MOTHERHOOD. 187

The Speckled Agalena makes a cocoon which equals in its complexity
the most carefully prepared of the Orbweavers. It not only surrounds its
eggs with several swathings of silken material, but adds a mattress of saw-
dust or bark chippings scraped from surrounding objects. Yet, according
to Mrs. Treat, a spider mother of this species kept watch over her cocoon
long after the frosts of winter had fallen, it being preserved in a suffi-
ciently protected spot.! Mr. Emerton attributes to this spider the habit
of remaining near her cocoon until she dies.” Nevertheless, my own nu-
merous observations compel me to believe that this species gives an example
of complexity associated with isolation of cocoon.

The interesting California spider, Segestria canities, spins a string of
ten or a dozen cocoons, which it suspends in the midst of a thick maze of
crossed lines, forming a strong protection, yet she keeps her home in a
silken tube spun along one side of the cocoon string.

Tegenaria agrestis of Europe makes a well protected and cushioned
cocoon for her young, yet she watches it carefully. The cocoons of all
known Saltigrades are all protected underneath a thick exterior tent and
by a stout case, but the mothers remain near, within the cell, although,
according to Professor Peckham,® underneath an extra covering. Such
examples as Segestria and the Saltigrades cannot positively be cited as
- eases of cocoon vigil, but at all events the mother’s domicile includes the
cocoon within its premises.

The above facts appear to indicate, first, that cocoons which are least
carefully protected by spinning industry have a supplementary defense in
the personal care of the mother; on the other hand, second, that
cocoons which are abandoned as soon as made, and are entirely
without maternal sentry, are protected by elaborate structures ;
but, third, in some cases the complex structure and the maternal vigil
exist together.

The In-
ference.

Orbweavers differ among themselves as to the number of cocoons spun
by females. Certain species, as the Tailed and Labyrinth spiders, habitu-
ally spin several cocoons; others, again, as most Epeiras, ordi-
Multifold narily spin but one. This habit must be subject to some va-
Cocoon- -_. |. : - :
wie riations, the reasons for which are not clear. Epeira apoclisa,
according to Lister, lays three and even four cocoons in the period
of a little more than two months. Termeyer makes the statement that
Epeira diademata, when well fed, will make six cocoons.
Several years ago a ministerial acquaintance, Rey. P. L. Jones, brought
me two cocoons of the Basket Argiope, both of which, he affirmed, had

1“My Garden Pets,” page 18. 2 New England Drassidie, page 200, (86).
’ Letter to the author.

188 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

been made by a single mother. It struck him as a strange circumstance,
and he reported the fact to me. Only recently Mrs. Mary Treat has pub-
lished a description of what she considers a variety of this spider, Argiope
multiconcha,! which habitually makes as many as four and sometimes five
cocoons. I have one of these strings, which was made in a kitchen where
a great cooking stove was in almost constant use to supply the demands
of a large family. It contains four cocoons, which were hung close to
each other, and precisely in the manner of those of Basket Argiope, which
they exactly resemble. The habitat of this spider, as far as now known,

is Missouri. The animal itself differs very little from Cophi- _
Argiope yaria. Unfortunately, the one specimen that I have seen was so
page much dried up that it could not be figured, nor could any dis-
tinctive features be readily traced; but it seems to differ in no
essential respect from Cophinaria. Thus, the interesting question emerges,
what are the conditions controlling this function in this spider? It can
hardly be quantity of food, as with Termeyer’s Diademata. If it be qual-
ity, upon what meat does this aranead feed, that she should so excel her
congeners in cocooning industry? <A tropical spider, Argiope aurelia, it
may here be stated, according to Mr. Pollock, makes ten cocoons.

The number of eggs within cocoons differs very much in different spe-
cies, and even among different individuals of the same species. Walckenaer
reports? that Epeira diademata has been found by him to con-
Number . : ;
of Higgs, tain from three to four hundred eggs, and again from six to
eight hundred, a wide range of difference certainly. I have
counted as many as eleven hundred and ten young spiders in the cocoon
of one Argiope cophinaria, and eleven hundred and fifty-two spiderlings
in another; and this is probably near the normal number. Some females
lay many less; as low, at least, as one hundred and fifty. The Peckhams
give the number of eggs laid by this species as varying from five hundred
to twenty-two hundred. According to Staveley, the cocoons of Argyroneta
aquatica range from forty to one hundred in number.

The reasons for this varying fertility are not certainly known. They
depend, no doubt, upon the female’s vital condition. A full measure of
health and abundant nutrition doubtless conduce to the normal number
of eggs, and this is probably diminished by physical weakness and lack of
nourishment. I believe also that the conditions surrounding the spider
influence the number of eggs, for females in artificial confinement seem to
deposit fewer than those in natural habitat.

Spiders that make a number of cocoons, as a rule deposit few eggs in
every one, so that the aggregate of eggs laid is about equal to the number

‘See “American Naturalist,” December, 1887, page 1122. * Aptéres, Vol. I., page 154.

MATERNAL INSTINCTS: MOTHERHOOD. 189

in the single cocoon of other species. The Labyrinth spider lays from
sixty to eighty eggs. The Tailed spider about the same. The Bifid spider
an aggregate of one hundred to one hundred and fifty in all her cocoons.
There is a marked difference in the number of eggs contained in the sev-
eral cocoons in one brood or cocoon string, as though the female issued an
‘unequal number of eggs at the various periods of ovipositing.

In tribes other than Orbweavers, the various species show the same
differences in the number of eggs contained within their cocoons; for ex-
ample, Tegenaria medicinalis has about sixty eggs; Agalena nevia, a hun-
dred or more; Dysdera bicolor, twenty to thirty. Walckenaer reports Aga-
lena labyrinthea to contain sixty eggs; Tegenaria domestica from sixty to
one hundred and eighty; Dolomedes mirabilis, one hundred to one hun-
dred and sixty; Lycosa narbonensis, five to six hundred; Lycosa agrestis,
one hundred and eighty. Dysdera hombergii (Staveley) lays from twenty
to thirty eggs.

It perhaps may be said, as a general rule, that the number of eggs
deposited by any species corresponds with the size thereof. Thus we have
seen that our large Argiope will lay as many as twenty-two hun-
dred eggs. Westring counted eight hundred eggs in a cocoon
of Epeira quadrata;! Epeira diademata will lay as high as eight

hundred eggs. Both of these are large species. The great tarantula, My-
gale blondii, deposits as many as three thousand eggs. On the contrary,
we find such a diminutive Saltigrade species as Synagales picata laying
but three eggs, while Phidippus morsitans, one of the largest of the Salti-
grades, lays one hundred and eighty eggs, thus being one of the most fer-
tile species.2, The Lineweaver Theridium variegatum has six eggs in her
cocoon. Oonops pulcher makes several cocoons, and deposits two eggs in
each one.’ ‘The cave spider Anthrobia mammouthia lays from two to five
eggs, while another cave species, Nesticus pallidus, deposits from thirty to
forty.4 It will thus be found, I think, that small and feebly organized
: species tend to deposit a smaller number of eggs, although there are some
marked exceptions to this.
The Peckhams give some interesting suggestions as to the relations
between fertility and exposure to peril. For example, the fact is pointed
out that while Argiope cophinaria is sufficiently well protected,
Fertility jer cocoons are exposed to serious loss through the assaults of
and HEx- . :
Gnas: ichneumon flies, and, perhaps, also through exposure to the
weather. Professor Wilder suggests that the immense fertility
of Nephila plumipes is counterbalanced by the destruction of its cocoons,
which are so placed, depending from leaves, that great numbers of them
are washed away and destroyed by rains.
The little Attid spider Synagales picata lays three eggs. Yet, beyond

Size of
Mother.

1 Aranew Svyecise, page 31. 2, The Peckhams. 8 Staveley. 4 Packard.

ae SE UP Se eee

190 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the fact that it is small and dark colored, this species has absolutely
nothing to protect it but the resemblance which it bears to an ant. Can
this alone give the species so great advantage that it is able to maintain
itself with as low a birth rate as three or four in a season? Considering
the direct relation between mortality and multiplication, it is plain that
no species could maintain itself at a low birth rate were not its mortality”
correspondingly low. It must then either practically have no enemies,
or its means of protection from enemies must be uncommonly efficacious. !
I state this theory without giving assent to it, and add the simple
remark that this species, or an Eastern species which greatly resembles
it in mimicry of ant forms, makes a cocoon of precisely the
Mimicry same character and protected in the same way as that of Phi-
ae “1:4,, dippus morsitans, one of the most fertile species among the At-
ertility. ~. 3
tide. Undoubtedly, cocoons of one species are exposed to the
same dangers as those of the other. Whatever advantage, therefore, Syn-
agales possesses in the way of protection is limited to the mature form,
and does not accrue to the eggs.

Wale

We come now to speak of the character of maternal solicitude as
shown by female spiders in the vigil of their cocoons. It is beyond
doubt that many species do guard their egg sacs with more or
ne less constancy during the period of hatching. The term “brood-
igs, =) : ; : osu
ing” has been applied to this action, but, of course, is inexact,
and only implies that the mother remains near or sometimes roosts upon
the cocoon, and, it is inferred, exercises some sort of protection against
the numerous enemies which assail the eggs.
Mrs. Treat? has observed that Epeira cinerea broods her cocoons for
a couple of weeks, and then drops dead from her maternal watch. Cer-
tainly she has good occasion for thus mounting guard, for of several
cocoons received from that lady, every one was infested by parasitic ich-
neumons, whose white pupa cases occupied portions of the egg padding.
I have never observed an orbweaying spider in what I could consider
an actual state of brooding her egg nest, although I have sometimes seen
female Orbweayers clinging to cocoons apparently lately made, This po-
sition seems to me to be rather due to indisposition to leave the vicinity
after the exhausting task of spinning and enswathing her eggs. But
various observers attribute the habit of brooding to some of the
Epeiroids of England. So, also, Menge, speaking of the ege nest of
Epeira diademata, says that the spider literally guards it with her life.

2 Communication to the author.

MATERNAL INSTINCTS: MOTHERHOOD. 191

fallen leaves, which are attached to the ground and covered with strong
threads. The mother lies down upon her cocoon, never leaving it to
take nourishment, thus starving herself to death in two or three

an weeks.! This author makes a like statement concerning Singa
eae albovitata. The phrase “starving herself to death” must, how-

ever, be taken in a figurative sense, for the fact is not one of
maternal sacrifice, but the inevitable debt of Nature after the maternal
functions are fulfilled.

Notwithstanding the above statements, I must say that I have met
with no evidence that any mother spider, during so called brooding, ever
actively exercised herself to protect her eggs against assault of
hymenopterous or other natural enemies. I once watched a fe-
male Herpyllus ecclesiasticus during more than two weeks’ vigil
of her cocoon. She never appeared to leave the vicinity of her button
like egg nest, which lay near a crevice into which she frequently retired,
and indeed was apt to retire at the sign of any disturbance made by my-
self. Sometimes she had her station upon the cocoon, embracing it with
her legs, but during all this period I failed to see the approach of any
natural enemies, and therefore was not able to record the fact that this
vigil resulted in any practical benefit to her embryo progeny.

With few exceptions, of which some have been noted, and which seem
to me to have doubtful features, Orbweavers hold no vigil over their

cocoons. As a rule the whole wealth of maternal care is ex-
oe pended upon the elaboration of the egg covering, after which,
Vigils, | in most species, cocoon and eggs are abandoned to the foster love

of Nature while the mother goes away about her business of
food gathering or falls upon death. It may be thought that those species
which make several cocoons present an evident exception, as in the cases
of Epeira basilica, caudata, bifurca, and labyrinthea, whose cocoons are hung
within their snares. But I have not been able to obtain any evidence
that the cocoons of these species receive special maternal care in the way
of personally protecting them from enemies, or aiding the young within
them to make their egress, or looking after the brood when escaped from
the cocoon.

Caudata may, indeed, often be seen hanging to the lowest cocoon of her
series, suspended along the vertical axis of her orb; but that is the natural
position for the spider under ordinary circumstances, and it may
have been assumed simply for convenience. I have never seen
the slightest indication of a desire on the part of the little mother to
mount from the lowest to the higher and highest ones of the string with
a view to protect them or oversee them.

In the case of Labyrinthea I have little doubt in saying that she

Parson
Spider.

Caudata.

1“Preussische Spinnen,” sub Epeira diademata.

192 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

exercises no care at all upon her cocoon string. This is ordinarily stretched,
as may be seen by consulting Chapter IV., page 100, at a position some-
what removed from her ordinary nesting site. It is true, her silken tent
is sometimes spun just beneath the lowest of the several cocoons which
she strings in a line one above another. But even in this case I have no
evidence that she exercises a personal vigilance upon them, or protects
them in any way.

The known species of Uloborus also suspend their several cocoons
along one of the radii of their horizontal orbs. Hentz, indeed, says that
Uloborus mammeata! watches her cocoon with incredible perse-
“verance, and shows great courage in attempting to defend it
against human disturbances. Fear seems to be wholly merged in maternal
solicitude, and as soon as the cocoon is torn from its place, the mother,
having remained firmly attached to it, proceeds to secure it with new
threads. I have observed similar behavior on the part of Theridioid
spiders, who hang their cocoons within the limits of their snare, and may
sometimes be seen embracing them or stationed near them.

Even this account may fail to assure one that the mother would have
~ repelled the attacks of natural enemies, and that she benefited her off-

spring by her stubborn keeping to her post. Nevertheless, it
Protect- seems to me a legitimate inference that the exercise of such
a maternal patience in vigil, and energy and courage in resisting
urpose. Coed

attempts to rob her of her treasure, indicate a purpose to pro-
tect the cocoon from natural enemies. Why does the mother so per-
sistently maintain her position close by her cocoon? Why is she ready
to undertake even such a hopeless venture as maintaining her egg sac
against the tremendous odds of a raiding naturalist? These questions
appear.to me to have but one answer. Motherhood is on guard beside
the cradle of her young, moved by a natural impulse to protect them
from the possibilities of natural peril. Even though no one has yet seen
or recorded the actual defense of the vigilant mother against such enemies
as ichneumon flies, egg loving spiders, etc., the circumstantial evidence
seems to be good that such defense must often occur.

It is known that outside of the Orbweavers some species of spiders
do persistently brood their egg nest, or rather keep: a close watch upon
them until they are hatched. I have already intimated that I
have seen this quite in detail with Hentz’s Herpyllus ecclesias-
ticus, and I have observed the same fact with Dolomedes tene-
brosus. Mrs, Treat notes the habit of a Thomisoid mother, probably
Misumena vatia, to remain near her cocoon, watching it with untiring
patience until the young are hatched. Hentz also speaks of this species?
as watching its cocoon until the eggs are hatched.

Uloborus

Other
Tribes.

1 Spiders of United States, page 129. * Thomisus fartus, “Spiders U, 8.,” page 78.

PLATE Ill,

MIMICRY OF ENVIRONMENT.

1, 2, MISUMENA VATIA, 3, A LATERIGRADE ON BARK. 4, EPEIRA STRIX. 5, EPEIRA PARVULA ON
LICHEN, AFTER PECKHAMS. 6, TETRAGNATHA EXTENSA AND THE ORCHARD SPIDER. 7. PURSE
WEB SPIDER’S TUBE.

MATERNAL INSTINCTS: MOTHERHOOD. 193

Theridium studiosum, when its web is destroyed, does not abandon
the cocoon, which is orbicular and whitish and is placed in the central
part of the web. The mother then grasps it with her mandibles and de-
fends her progeny while life endures. Her maternal solicitude is not
limited to her cocoon, but she also takes care of her young, making a
tent for their shelter and remaining near them until they can protect
themselves. !

Toward the last of July the female Turret spider appears at the top
of her tower with a cocoon of eggs, about as large as a hazel nut, at-
tached to the spinnerets. She exercises the greatest care over

Pe her cocoon, On cool days she keeps it out of sight down in her
Spider tube, which is about eight inches in depth, including the tower.

But when Mrs. Treat set the jar in the centre the mother spider
soon came up and put the cocoon in the sunshine. When the weather was
cool enough for fire in the room, if the jar were placed near the fire the
spider placed her eggs on the side next the stove. If the jar were then
turned around, the mother presently moved the cocoon around to the warm
side, letting it hang outside of the walls of her tower. On the 6th of
October the young spiders were hatched, and at once perched upon the
mother’s back, and even on her head and legs. She carried her cocoon
two months before the eggs hatched.

The Lycosid Oxyopes viridans makes a conical cocoon haying small
eminences, to which are attached the threads that hold it suspended firmly
in the air. After it is finished the mother watches it constantly,
never leaving its unprotected family.? Professor Hentz, speak-
ing of the general maternal instincts of the Lycosids, says that the mother
defends her progeny to the last, and her feet can be torn from her one
by one before she can be compelled to abandon her treasure. Thus can
maternal tenderness be exhibited in beings which are relentless to their
own species, and even to the sex which gives life to its progeny.? I must
say that my own experience gives no such examples of persistent devotion
under attempts to separate Lycosids from their cocoons. If the female of
Lycosa lenta be caught or wounded, the little ones escape rapidly in all
directions, but the mother is faithful to her duties and defends her progeny
while life endures.# 7

A female of Dolomedes albineus was captured by a child, who trans-
fixed her cephalothorax with a pin. The creature was placed in a glass
jar, and the wound, instead of proving mortal, healed rapidly.
After remaining inactive about three days, the spider made an
orbicular cocoon of light brown color, in which her eggs were
placed. She held it constantly grasped in her mandibles, and seemed in-
tent on watching it to the last; but the effort of cocooning once made,

Lycosids.

Dolo-
medes.

' Hentz, id., page 106. 2 Id., page 48. *Td., page 25. *Td., page 28.

ee. a

194 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

her strength failed. The wound opened again, and, the fluids running
freely, she gradually lost her muscular power. But faithful to her duties,
the last thing which she held was the ball containing her future family.
Can maternal tenderness be more strikingly exhibited ?!

Dr. T. W. Harris, whose work on “Injurious Insects” is well known,
found in Massachusetts a female Dolomedes lanciolatus on a large, irreg-
ular, loose horizontal web, at one extremity of which was situated her egg
bag with her young, which the parent appeared to be watching.? Micro-
mata marmorata remains constantly by its round white cocoon, which it
embraces closely with its long legs, while it hangs suspended by one thread
in the middle of its snow white tent.

Many British spiders have the same habit of caring for their cocoons.*
The female of Philodromus ceespiticolis conceals herself with usually two
flattened white cocoons in the large nest, which she forms upon
the end branch of some shrub, drawing the leaves into a con-
venient position with silken threads, which form a close tissue
of a somewhat gray color. The cocoons are frequently of unequal size,
the largest being about one-fourth inch in diameter. If the cocoon be
touched the mother will not take flight, but will defend it with all her
power.*

Drassus ater makes a plano convex cocoon, which is attached by its
flat side to a stone or other substance, on which the cell is formed. This
cocoon is white or slightly yellowish at first, and afterwards becomes red-
dish in color. The female remains on guard by her eggs.° The female
of Drassus lapidicolens conceals herself in a cell formed between the sur-
face of the earth and the under side of a stone, near which she spins some
threads, forming an irregular snare. In this cell, in the month of July or
August, she places her cocoon, covering it with dead leaves. This cocoon
is at first in the form of a flattened sphere, but becomes nearly round
when the young are about to escape. It is white, and about half an inch
in diameter. The mother remains with her young for some time after the
eggs are hatched.

Clubiona holosericea makes a white flattish cocoon one-fourth inch in
diameter in June, and places it in a long tube shaped cell, formed on the
under side of a leaf, or in some crevice, as of the bark of a
tree. ‘The female remains in this cell except when she leaves it
to pounce upon an insect passing near its opening, and which she carries
into the cell. The cell is divided into two chambers, in which, in the
month of June, male and female may be found each occupying one. The
spider is timid until she becomes a mother, when she will face any danger
rather than abandon her cocoon. Before that time, ¢f driven from her

British
Examples

Clubiona.

' Hentz, page 39. *Id., page 41. * Staveley, British Spiders, page 168.
4Id., page 85. 5 Td, page 91. 6 Id., page 97.

= i ll ee ie APA ee aS

_ MATERNAL INSTINCTS: MOTHERHOOD. 195

cell, she falls to the earth without drawing a line with which to suspend
herself, feigns death for some time, and then, making a rapid flight, sets
to work to build a new house in a fresh place.!

VIF.

Mrs. Mary Treat, in a little work designed for popular use entirely,
gives’ several extraordinary examples of maternal care on the part of
spiders. Ome of these was a female of Dolomedes scriptus, which
Special - first attracted attention by the fact that she was carrying a bag
reais of eggs about the size of a small cherry, with which she planted
Gare herself on top of a leaf nest of a Shamrock spider (Epeira tri-
folium). One morning Dolomede was missed from her accus-
tomed place, but upon searching some adjoining ferns the characteristic
cocoon tent of the species was discovered. It was three or four inches
in length and from two to three in breadth, composed of ferns bent over
and fastened together. ‘Through one of the openings between the leaves
the cocoon was seen suspended from the ceiling, precisely as I have myself
observed it, and as is represented in the sketch Fig..177, Chapter V.

In two or three days thereafter the young Dolomedes were hatched
and swarming all over the outside of the cocoon. When the leafy domi-
cile which enclosed them was touched the little ones ran down the lines
in the direction of Mrs, Treat’s finger, as if they expected something, and
reminded the observer of young birds, which always open their mouths
to be fed whenever they are approached by a human being, not having
yet learned to recognize their parents.

This behavior led Mrs. Treat to suspect that the youngling spiders were
fed by their mother, and she accordingly kept watch upon the colony.

One evening, not long afterwards, the mother Dolomede was seen
speak’ with a large fly in her mandibles taking long strides in the
ing the : : iS Bie :
Young. direction of her. domicile. She was soon inside, and the little

ones thronged around her and sucked the juices of the fly while
she held it. The fly had previously been crushed in the mother’s jaws as
though to make the food available for her nestlings. How long this
process continued is. not stated. It is a great pity that the details were not
given, and the lack of these details leaves in my mind the question, was
_ this really a case of feeding the young? Or did the Dolomede simply
return to her nest to prey upon the food which she had gathered for
herself, and permit, without interference, her brood to share in the repast?
I have seen a mature Argiope (see Vol. I., page 256) feeding upon a blue
bottle fly, while a number of small Diptera were sharing in the feast,
having crowded up to the very jaws of the spider to sip the juices of the
carcass, Of course, no purpose to feed the little flies could be inferred on

' Staveley, page 100.

ee ee ee a

ss ee re

~~ a i i

196 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the part of the big spider. Might not a deliberate intention to feed her
young be excluded from the act of this mother Dolomede on precisely
the same ground ?

Quite as extraordinary as the above is the behavior of a little Jumping
spider, Attus nubillus, related by the same observer.'| This spider de-
posited her cocoon, after the manner of her genus, within a couple of
curled leaves of prickly Smilax rotundifolia. Mrs. Treat opened the
nest and found that the spiders were apparently just hatched, and were
of a pale green color. The mother was not then in sight, but knowing
that Attus remains with and cares for her young until they leave the
nest, the observer waited and was rewarded by witnessing the little mother’s
return. For a time she seemed to look with dismay upon her pretty home
torn asunder, and her spiderlings scattered around, but soon proceeded to
gather the younglings together and tuck them back under the

ae 1 silken canopy. One spiderling, which had wandered farther than
the rest to the verge of the leaf, was picked up bodily, as a cat
Young. g ) if } y

would carry its kitten, and put back into the flossy interior of
the cocoon. Then the mother set about repairing her damaged cocoon ;
and after the rent was mended the young were not visible. She also
tried to bring the enclosing leaves together again, but presently abandoned
that effort.

She remained on the outside of the nest, and no threatened danger
would induce her to leave. She sprang towards the observer’s hand, and
fiercely grasped the point of a pencil thrust near her. Several times
daily the nest was visited, and the mother was found persistently pres-
ent until the third day, when she was missed. A second time the cocoon
was opened, and the spiderlings found to have made the first moult, and
were crawling about slowly. When the mother came back and perceived
her young disturbed again, she varied her behavior so far as to look
around for the cause of the disaster—spying around leaves, and over and
under them. Finding nothing, she soon became quiet, put her brood
_, within the cocoon once more, and again repaired the damage.
cae This completed, she went to work to bring the leaves together.

The tips now stood two inches apart, while at the base or stem
end the space was half an inch. The leaves were thick and leathery, and
the petioles stiff and firm. She fastened a thread of silk to one leaf and
then to the other, and went back and forth strengthening and shortening
the lines, and slowly bringing the leaves together. The next morning they
were found quite joined, and the interior entirely hidden.

A third time, during the mother’s absence, the leaves were separated
without disturbing the young within their cocoon. When the mother re-
turned she did not attempt to reconnect the leaves. In a day or two

1 “My Garden Pets,” pages 64-68.

MATERNAL INSTINCTS: MOTHERHOOD, 197

thereafter a door was seen in one side of the nest, out of which the spi-
derlings soon made their exit. They were quite lively; several were on a
leaf; they seemed to be playing, springing at each other, then back into
the nest and out again. When the leaf was touched every one instantly
disappeared.

At the next visit Mrs. Treat moved her pencil over the nest a little
harshly. Instantly the spiderlings all fled from the abode, springing in
every direction; but, before leaving, every one must have fixed
a thread to the leaf, for all soon returned, slowly ascending,
taking in. their tiny cables, until they reached the leaf, when
they cautiously approached the nest. “I was somewhat puzzled at their
return, after such a flight,” says the writer, “until I saw the mother with
a fly and the little ones all around her sucking its juices. ‘This, then, was
the reason of their remaining together—they were fed by her.”

This account is so fully detailed, and contains the evidence of such re-
peated and careful observation, that we have no room to doubt the fact
that the mother Nubillus maintains a remarkable degree of oversight and
care in behalf of her young during the period of their early life and
growth. The manner in which the spiderlings were covered again and
again within the silken egg sac, especially the extraordinary fact of one
being carried in the mouth and placed along with its comrades, suggests
to us a degree of maternal solicitude on the part of this aranead which
falls little short of that exhibited by the mothers of vertebrate animals.
If we add to this the fact, of which Mrs. Treat appears to have no doubt,
that the mother actually brought food to the nest and bestowed it upon
her offspring, we shall be compelled to place the mother aranead even yet
nearer to her vertebrate sisters in the quality of her maternal affection
and care. It is: greatly to be regretted, however, that in this case; as well
as that of Dolomedes scriptus, only one example of so called feeding could
be observed and recorded. :

These detailed observations confirm the statements of Blackwall, that
“the young of some species live together for a considerable time, and in
many instances are supplied with sustenance by the mother;”1 and again,
that the young of Theridium riparium “remain with the mother for a
long period after quitting the cocoons, and are provided by her with food,
which consists chiefly of ants.”? I have observed the young of spiders,
particularly of a small species of Dictyna,* feeding upon parts of a dead
fly which the mother was eating. At least the young seemed to be feed-
ing, and I have the belief that they were doing so, although I could not
positively declare it, since they may have simply been resting upon the
limbs and other parts of the dead insect, after the manner of young

Spider-
lings.

1 Spiders Gt. B. & LI, Intro., page 7.
2 Td., page 9, and Researches in Zoology, page 356.
® Dictyna philoteichus, Vol. I., page 354.

198 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

spiders to get themselves upon all objects and in all manner of positions
within their neighborhood. Certainly these observations must open up
before our minds more clearly than ever the existence of a high degree of
maternal instinct in spiders, and the possibility of discovering yet higher.

VIII.

Among experiments on the mental powers of spiders, made by Professor
and Mrs. Peckham, were some on the strength of the maternal feeling. They
proceeded by removing their cocoons from the mothers and then

Strength noting with what degree of eagerness they sought to regain them;

ste ree and also by determining for how long a time they would remem-
ings. ber the cocoons after they were separated from them. The Ly-

cosids were selected as the principal objects of study, because
these spiders keep the egg sac attached to the spinnerets until the young
are hatched, and thereafter carry the spiderlings on their backs for a cer-
tain length of time, until they are able to shift for themselves, It was
thought that the lengthening of the period of infancy during which the
female cares for her young, might, in the case of araneads, as in that of
more highly organized animals, have produced a greater development of
maternal instinct than in other species of spiders where the eggs receive
little or no attention from the spider after she has deposited them. The
following will show the character of some of the experiments :—

The cocoon of a female Pirata piraticus was removed from her. Dur-
ing this act the mother seized the egg sac with her falces several times and
tried to escape. After the removal she seemed much affected
and searched about in all directions to find her lost treasure.
In an hour and a half the cocoon was restored to the mother, who im-
mediately took it between her falces and passed it back to its proper place
beneath the abdomen. It was again removed and returned in three hours.
The mother did not seem as ready to receive it as in the first instance,
but after a little hesitation took it up and carried it off. From three
spiders of the same species cocoons were removed, and retained for the
space of thirteen, fourteen and a half, and sixteen hours respectively. All
remembered them and took charge of them when they were returned.

The same cocoons were again removed and retained for twenty-
Absence four hours, when they were restored. Two of the mothers re-
Weakens 6 od to re 5 het parivall eae : fe :
Interest, ‘sed to resume their maternal duties, seeming not to recognize

their cocoons. The third, however, after her cocoon had been
placed in front of her seven times, slowly resumed charge of it, but
with none of the eagerness before displayed.

Experiments with a female Lycosa of an unknown species resulted as
follows: After being separated from her cocoon for a whole day the mother

Lycosids.

* Mental Powers of Spiders, page 397.

MATERNAL INSTINCTS: MOTHERHOOD. 199

recollected it, and promptly took it up. A second individual, after an ab-
sence of forty-three hours, had apparently forgotten all about her cocoon,
since, although she touched it five times with her legs, and it was four
times placed directly under her, not until the fifth time did its presence
recall her to a sense of duty. She then very slowly and languidly took
it up and passed it to the usual place. From another individual the
cocoon was kept forty-eight hours, but the little spider could
not remember so long, and, although the observers worked long
and patiently to make her recollect, she would have nothing
more to do with it. Notwithstanding many efforts, no female among the
Lycosids was found constant in her affection to her cocoon after as long
a period as forty-eight hours.

Several species of the Attidee and Thomisidee did not remember their
cocoons for twenty-four hours, On the other hand, a female of Clubiona
pallens remembered her eggs for the space of forty-eight hours, and when
they were returned to her spun a web over them in the corner of the
box in which they were placed. Of all the spiders experimented upon
by the Peckhams, the little Lineweaver Theridium globosum had
the best memory for her cocoon. This was returned to her after
Mother. fifty-one hours’ separation. She at once went to the eggs, touched

them with her legs, then left them to improve her web, every
now and then running back to see if they were safe, and presently set-
tled down near them.

Here, again, our ideas of what might be expected in the ordinary course
of Nature are thrown into confusion. The Lycosids, who carry their co-
coons about their persons until their young are hatched, and then person-
ally conduct their broodlings until they are strong enough to take their
chances in life apart from maternal care, appear to have a weak memory,
and a comparatively feeble maternal affection for their offspring. So also
the Attids and Thomisids, who remain near their cocoons, brooding or
guarding them for the space of fifteen or twenty days, were found so de-
fective, either in memory or maternal feeling, that they lose interest in

Two Days
Limit.

A Faith-

their cocoons if separated from them for the space of twenty-four hours.

No doubt these experiments need supplementing; and when the patient
observers who have given us these results shall have wrought
longer upon the same field, we may come to different conclu-
sions; but at present it would seem that the development of
maternal instincts appears to be quite independent of those causes which,
according to the theory of evolution at least, we might have expected to
affect them most vigorously.

Develop-
ment.

IX.

The Peckhams found, as others had discovered, that it is not a difficult
matter to deceive spiders as to their proper cocoons. A ball of cotton they

— ee a

200 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

refused, but a little pith ball led them entirely astray. The following will
indicate the nature of some of their experiments. A pith ball three times

as large as the cocoon of Pardosa pallida was refused by the
Mistakes mother. When reduced in size she took it between her falces
ele and attached it to her abdomen. The bit of pith appeared to

give as much satisfaction as the egg sac. When the cocoons
were nearly of a size, one mother would take that of another, although of
a different genus, just as quickly as she would her own.

As a further test of general intelligence, the outer case was taken from
a cocoon of Pallida and slipped over a lead shot of the same size, but three
or four times as heavy. Much of the silk envelope was broken
away in thus covering the shot, but when offered to the spider
she at once seized it and after a good deal of trouble fastened
it to her abdomen. The load was so heavy that the mother had great
difficulty in walking up the side of a board. While transferring this spec-
imen to another box, the shot from its weight fell from the abdomen, and
the mother spent over thirty minutes, working with all her might, fas-
tening it on again. Once more it fell off, and this time she carried it
about between the falees and the third pair of legs. A second specimen
of this species was tried with the plain shot, but would have nothing to
do with it. The web covered shot was then removed from the first speci-
men, and the plain shot offered to her instead, but this was stubbornly
refused, whereupon the web covered shot was returned and was taken back
with every evidence of tender emotion.

Another test was made by offering a cocoon and a pith ball together.
The two objects were placed side by side. The mother, approaching one

side, first touched the pith ball, and at once seized it with her
Touch —falces. But as she moved away one of her fore legs touched the
ae cocoon. She stopped, remained quiet a moment or two, then

dropped the pith ball, took wp the cocoon, and moved away with
it. The next day the two objects were again placed in front of her. This
time she also happened to meet the pith ball first, and, as before, took it
up at once. She ran off with it, and it was some time before the experi-
menters managed to place the real cocoon just in front of her. As soon
as her legs touched this she stood still, and then after a few minutes
dropped the pith ball and took up her eggs.

From these observations it is evident that the spiders experimented
upon, when allowed to choose, can distinguish their own cocoons and have
a preference for them. But in the absence of their cocoons they content
themselves either with a pith ball or, more strangely still, with a web
covered shot. The presence of the web upon the shot makes a marked
difference in the disposition of the spider towards it. The contact of the
pure metal causes the rejection of the object, while contact with the web
covering thereof produces that complacency which leads the mother to

Lugging
a Shot.

MATERNAL INSTINCTS: MOTHERHOOD. 201

adopt the shot in place of the cocoon, The fact that the spider will carry
about so comparatively heavy an object as a lead shot instead of its cocoon
certainly argues a poorly developed muscular sense.' Sir John Lubbock
appears to have made some experiments in the line marked out by the
Peckhams, as a result of which he concludes that examples of Lycosa
saccata did not appear to recognize their own bags of eggs, but were equal-
ly happy if they were interchanged.?

A gravid female of Argiope cophinaria sent to me enclosed in a paper
box was found dead, having left an unfinished cocoon. She was cling-

ing to one end of a thick patch of white spinningwork that
Covering quite enclosed a small tuft of fern leaves, at the other end of
Empty é ty ; areal
Hipe weed, which was a roll of purple swathing, corresponding with a

purple pad of a complete cocoon. I expected to find the eggs
within this roll, but was surprised upon opening it to see a yellowish
ball of silk, and nothing more. Where the eggs should have been was
only a round silken wad. Nevertheless, the expiring energy of the spider
had been spent in spinning a protecting cover around this mock egg
mass.

An anonymous observer® records somewhat similar cases. He found
one Cophinaria cocoon in which there were no eggs, and another con-
taining but three. The eggs were on some boards beneath the cocoon
site, having fallen from the first receptacle before they were covered. In
each case the mother went on with her work and carefully finished the
egeless flask. If the loss was discovered the knowledge made no differ-
ence in the exercise of her maternal functions, which, apparently, were
controlled by an instinct or feeling quite independent of knowledge.

A like example of mental abstraction (if one may be allowed such a
phrase) was reported to me by Mrs. Mary Treat as having occurred at Vine-

land, New Jersey, with a large Florida Dolomede spider, probably

ee Dolomedes tenebrosus Hentz. This mother, after the habit of
igen 7. . ny ' . y . A 46 ? “{ 4 ]
Tustinet, her genus, carried her cocoon under her jaw, but sometimes

shifted it to a position beneath the abdomen. Yet there were
no eges in the cocoon—a fact which thus came about: When first caught
the Dolomede was confined in a tin can, which so surprised or frightened
her out of her maternal propriety that she deposited her eggs in the
can without attempting to protect them with a cocoon. She was removed
to a natural environment upon the ground, whereupon she spun a web
and gathered up sundry materials, which she managed to make into the
form of a cocoon, which, with this species, is a round sae about the size
of a boy’s playing marble. This she hugged to her body and lugged
about with as zealous care as though it were filled with eggs.

' Mental Powers of Spiders, pages 417-419.
* On the Senses, Instinets, and Intelligence of Animals, Sir John Lubbock, page 179.
5“ Katydid,” Chicago Tribune, September 11th, 1881.

— a

202 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Evidently in this case, as with the Argiopes, the mother acted under
the impulse of a mentalism that was without reasoning; or, if we may
suppose that she reflected upon the loss of her eggs, such reflection had
not sufficient influence upon her will to resist the instinctive impulse—
or shall we say the simple functional impulse ?—to brood upon something.
These cases remind one of the well known persistence of setting hens in
brooding over an empty nest.

The English observer, Mr. F. M. Campbell, records a like example.
While watching a female Tegenaria guyonii lay her eggs, it occurred to
Tegenaria “i to see what she would do if these were removed. Accord-

ingly the eggs were deftly lifted away, but somewhat to the
disturbance of the mother. After a few seconds she began to overspin the
spot where she had just placed her eggs, and completed her cocoon.’ Mr.
Campbell’s suggestion, that the force of habit urged forward the aranead
to act as though the eggs were in the proper site, must be qualified by
the fact that “habit” cannot have much influence in a maternal act which
is repeated so few times as with this species. There is more plausibility
in the physiological aspect of the act which he suggests, viz., that the
maturity of the eggs may have been correlated with greater activity in the
collection of fluid by the spinning glands; and, as in the case of the mam-
mee of a vertebrate, the discharge of their contents may have been neces-
sary for the comfort of the creature.

It is impossible not to note the many evidences of exact mechanical
skill and design shown by spiders in the act of cocooning. We speak of
this, no doubt properly, as instinctive. Certainly it is not the
result of experience, for even in the case of those spiders that
make several cocoons, the first one is finished with the same
accuracy, and indeed after the same methods, as the last. It cannot result
from instruction, for in the great majority of cases the young are never
associated with their mother; and in those cases, as among Saltigrades and
Lycosids, where the spiderlings are under maternal watch for a little season
after hatching, the idea of instruction in the art of cocoon making is not
to be thought of. Neither can we suppose this remarkable mechanical ac-
curacy to be the result of observation, though no doubt it might happen
that immature females observe the methods of cocooning practiced by
mature specimens of the various families. In short,- the only admissible
conception is that the act, including all the methodical details, is intuitive,
and springs into being in full operation at the moment that it is needed,
and that without any previous preparation or knowledge of any sort on
the part of the aranead mother.

Intuitive
Skill.

1 On Instinct, Trans. Hertfordshire Natural Hist. Society, Vol. IIL. page 3, Dec., 1884.

MATERNAL INSTINCTS: MOTHERHOOD. 2038

Nevertheless, it must be admitted that the entire actions of the spider
are such as one would expect from an individual that had been thoroughly
instructed, had acquired skill from experience, and was pursuing
ae of 4 matured plan with all the evidences of forethought and skill.
thought. To illustrate this fact, one may refer to the method practiced by
Argiope in making her cocoon. Nothing could be more indic-
ative of forethought than the manner of placing the eggs at the outset
upon the little saucer shaped disk prepared for their reception, and which
are retained in their place by means of the silken sac swiftly woven
over them. ‘The preparation of the thick padded purple mass which next
envelops this strikes our attention as indicating wise skill. The manner
in which this is woven so as to produce the loose substance that envelops
the eggs, and at the same time is compacted into a solid and shapely mass,
is certainly what the artisan might denominate a ‘mechanical job.”
Then, again, contrast the mode of weaving this object with that prac-
ticed upon the external case of the cocoon. As has been shown in full
detail, the method of spinning is here quite different, and corresponds.
closely with the character of the fibre to be spun. In other words, as it is
the intention of the mother to make the outer case a closely woven tissue,
instead of a loose mass like the purple pad above alluded to, she proceeds
to tighten the threads, bracing them upon each other and compacting
them by all the methods familiar to her spinning art.
Nor can we fail to mention here the manner in which the service is
equalized so that every part is of nearly equal thickness and evenly dis-

tributed, so as to form the shapely pyriform cocoon familiar to most '

wanderers in our fields. As has been explained, this is done by carrying
the silken filaments as they are outspun from point to point while the
spider circumambulates the surface. That such a practice requires a di-
recting purpose, and that such a method is the outcome of an express
design, seems the most manifest conclusion. The spider spins her cocoon
case very much after the manner in which a lady winds up a ball of silken
thread or of embroidering wool. One would think it as unreasonable to
say that the manner in which the ball in the lady’s hand is shaped into
its globular form is without any directing purpose on her part, as that
the mother Argiope, in forming her cocoon ball, wraps the silken material
of which it is composed without any guiding purpose to keep it even.
Take another example, to illustrate this point, the remarkable mud ball
of Micaria limicune. The cocoon sac has been shown to be a white oval
case within which the eggs are spun, and which is connected to
The Mud some surface by a little silken cord or pedicle attached to the
Cradle saat : ; ‘
Maker, ‘P: ‘This sac made, the spider proceeds to cover it with mud,
and so works that, when she has finished, the enclosing shell of
mortar is, with scarcely an exception in a large number of specimens ex-
amined, a well rounded globe. Now, how has the spider proceeded to

ee a
, ’

204 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

accomplish this result? If we were to suppose a human plasterer given
the task of covering a sack hung to the ceiling of a roof with a coating of
mortar two or three times the bulk of the sack, and to have the same, at
the conclusion of the work, in the shape of a globe, how would he proceed?
Manifestly by placing successive layers around the sack, suffering one to
harden before the other would be laid on, rounding each up with his
trowel as he proceeded, building the mortar around the cord by which
the sack was suspended, and all the while so guiding his implement that
the object would gradually assume the globular shape required.
Precisely such is the method pursued by our little Limicune. A dis-
section of the cocoon shows that the spider placed her mud upon the silk-
pee en sac in numerous successive layers; that she permitted one
Limi- layer to harden before the other was laid on; and that, as she
cune’s .
Arata: gradually proceeded, she built her mortar around the suspensory
cord or pedicle, and shaped the whole with her mandibles and
feet until it assumed the form of the smooth, round object represented in
Figs. 147 and 148 on page 130.
One cannot venture to think that the process by which the human
plasterer arrives at his method of work is identical with that pursued by
this spider plasterer. In the one case it is the result of educa-
Man’s tion and experience, and of the application by reasoning of

Method : oat ;
and the Previous training to the problem in hand. In the case of the
Spider’s, Spider no such education or experience, and probably no such

process of reasoning, can be predicated. What mental processes
has she gone through, if indeed she has passed through any? Can we
ascribe to her, under the circumstances, the credit of reasoning upon -her
work and ‘adapting her methods thereto? ‘lo do this would seem to me
to place her thinking abilities and natural mechanical capabilities above
those of man. ‘That there have been design and forethought somewhere
behind all the processes of the spider mother one cannot doubt; yet with
equal certainty we must refuse to attribute them solely to the spider her-
self. Forethought and mechanical skill abide in Nature, whose formative
forces have wrought out the structure of the spider and guided all its
functions. But forethought and skill are the attributes of mind, of per-
sonality ; and how shall we denominate this Personal Thinker? How can
we deny His Presence?

Perhaps a third illustration may be added. Cyclosa caudata has the
curious habit of attaching to the exterior of her cocoons carcasses of in-
sects from which she has sucked the juices, instead of casting
them from the snare, the usual aranead mode of disposing of
such material. Given the habit of suspending the cocoon with-
in the disc of the orbicular snare, and also the habit of protecting the
same by an armor of extraneous material, it is, perhaps, inevitable that
the mother should be compelled to resort to some such method. It is

Cyclosa
caudata.

MATERNAL INSTINCTS: MOTIHERIIOOD. 205

obviously impracticable to descend to the ground and secure mud, vege-
table mould, and chippage, as is the custom with those species whose
cocoons are fixed upon various surfaces, and whose makers can conveni-
ently resort to terra firma, In the case of our little Caudata, whose net
swings in the open air, the chippage of slaughtered insects is after all
the most convenient material at hand. Necessity here, as among human
creatures, appears to have been the mother of invention, aided much by
opportunity. The hard, dry shells are cut up into pieces, which are stuck
to every part of the egg sac until the whole is covered, often very closely.
Thus, in a single cocoon one will be able to detect the wings, head, elytra,
abdomens, and other parts of various orders of insects, many of them
having bright colors.

In these various methods of exercising this general habit one can
find no motive which meets the facts of the case as well as that of ma-
ternal solicitude. Mother love has found expression in the armoring of
the silken vessel within which the eggs are enclosed, thus protecting
them from the enemies which are to beset them. The motive is none
the less potent, and none the less to be recognized, because of the fact
that the mother herself could have had no knowledge of the character
of those enemies to which her progeny would be exposed, and acted in
obedience to an impulse within which we can trace no factor of personal
reasoning.

a ae

"
‘j

PART [I.—EARLY LIFE AND: DISTRIBUTION
OF SPECIES. |

GAAPTER VITI.

COCOON LIFE AND BABYHOOD.
Te

THE tyro in arachnology experiences his first and greatest difficulty in
the attempt to separate between the mature and immature spiders collected
by-him, There are resemblances between the young of various
species, particularly of the same genera; and the differences be-
tween the young and the adult of any one species are, in certain
cases, so great as to produce confusion. In point of fact, except for pur-
poses of special study in life economy, young spiders are not worth collect-
ing or retaining in a collection. The valuable specimens are only those
which are mature.

Now, it must be remembered that spiders do not undergo a metamor-
phosis—a fact which is continually forgotten because of their classification
with insects by the earlier writers, and the frequent treatment of them
under Entomology even at the present day. Certain orders of insects, as
the Lepidoptera, undergo a complete metamorphosis. The butterfly ar-

: rives at maturity through the well marked stages of the cater-
Spiders pillar and chrysalis. Other orders, as the Orthoptera—locusts
bk hee cand grasshoppers, for example—haye an incomplete metamor-
phosis. Phosis. But a spider is a perfect animal from its birth, and

only requires the general growth and strengthening of its mem-
bers, together with the development of the sexual organs, to complete its
maturity.

This maturity is reached after several successive moultings of the skin.
An important outward structural change takes place at the final moult,
at which time male spiders get their complete armature of spines, bristles,
and hairs, according to their species. Moreover, the last or digital joints
of the palps, which, to quote the language of Cambridge,' have been up
to that time tumid and homogeneous, break up into the digital joint, so

Adult and
Young.

1 “Spiders of Dorset,” Introduction, page 26.
(206)

—

ieee

COCOON LIFE AND BABYHOOD. 207

called, and the curious and more or less complete congeries of lobes, bulbs,
and spines known as the palpal organs. The full dimensions of the legs
. . are also sometimes attained at the same period. The female
mip spider at her last moult merely develops the genital aperture
turity. With its external processes. Up to this time the aperture is in-
visible, though, like the palpal organs of the male, it has been
gradually developing beneath the cuticle.

Of spider life within the cocoon our knowledge must necessarily be
limited. The period of hatching differs according to the species, the time
of the year, and the nature of the season. The eggs in many
be aie autumn cocoons do not hatch until spring, say from the middle
ife. 5 ;
of April to the middle of May. I haye gathered many cocoons
that have wintered out of doors, of Agalena nevia, of various Laterigrades,
and several species of Orbweavers, which contained unhatched eggs from
which young spiders were subsequently bred. After hatching, the little
creatures remain massed within the cocoon along with the white shell of
the egg or the first moult. At times they spin delicate threads, which add
to the flossy nest within which they domicile, so that after a cocoon has
been opened for examination, the fracture will be closed up by such spin-
ningwork,

The spring or summer cocoons are hatched at periods varying from
fifteen to thirty days. According to Professor Wilder, the eggs of Nephila
plumipes laid in September were hatched in about thirty days.!
A cocoon of Epeira cornigera, taken in April and having the
eggs then unhatched, I found to contain fully hatched young on
May 15th. A female Epeira sclopetaria cocooned in a trying box May
26th, and on June 13th, eighteen days thereafter, the young brood issued
from the cocoon,

I have opened cocoons of Argiope cophinaria in the early winter, and
found the young within crawling about in a sluggish way among the
silken fibres of the interior enswathment, or massed inside the central,
common pouch along with the white skins of their first moult. On the
contrary, I have found cocoons in which, as late as April 20th, the young
had just cast off the egg shell, and were beginning their first, feeble
movements in struggling with the silken lines of their enswathment. I
have little doubt that the young of Argiope are generally hatched from
the egg within a month or six weeks after the cocoon has been made.
They, therefore, remain within the cocoon during the winter and until
the season is sufficiently advanced to make their egress safe.

Period of
Hatching.

' Proceedings American Academy of Arts and Sciences, Vol. VIL., 1866, page 56.
page

a aa

208 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

But in the case of females who, for whatever reason, haye been be-
lated in positing their eggs, the frosts of early autumn probably have
the effect of retarding the process of development; and when
ares a the later autumn frosts and the winter cold follow, the eggs of
altel such cocoons remain unhatched until the first warm days of
coming spring quicken their vitality. This is probably true of other spe-
cies than Argiope. I have never made any experiments upon the effect
of frost to retard or prevent the hatching of spider eggs, but am inclined
to think that cold has this effect upon them, as it is known to have upon
the development of insects.

On May 22d, one exceedingly cold season, I found the young of Epeira
sclopetaria, at Atlantic City, all escaped from their cocoons, great num-
bers of which were fixed upon the cornices of various buildings around
the Inlet wharf. At the same time many cocoons of Epeira triaranea
had the young still within them. I have had young Insular spiders
colonized upon my vines make their exode May 19th.

M. Vinson says that July Ist Gasteracantha bour-
bonica, a Madagascar species, enclosed in a flagon, had
fixed her cocoon against the side. On the 25th the
little spiders were hatched. They were perfect as to
their forms, but were still imprisoned within the co-
Fic. 241. Young Agalenn COON. ‘They presented a blackish appearance. They
stripping off the first jgsued from the cocoon and scattered on the 11th of

moult. « :

August, a period of seventeen days after hatching. So
that under an African winter the hatching of eggs and escape of the
young occupies a period of forty days.

On the disengagement of young spiders from the egg every part is en-
closed in a membranous envelope; they are embarrassed in their move-
ments; are unable to spin or seize prey, and seem indisposed
to action, For the unrestrained exercise of these functions it
is requisite that they should extricate themselves from the cover-
ing which impedes them. This operation, or, as it may be termed, their
first moult, occurs after a period whose duration is regulated principally
by the temperature and moisture of the atmosphere. The first moult in-
variably takes place in the cocoon, or general enyelope of the eggs, and
the young spiders do not quit the common nest until the weather is
mild and genial.!

Once, while peeping inside a cocoon of Agalena nevia, I was fortu-
nate enough to observe a spiderling in the last stages of this first moult.
While it held on to the flossy nest with the two front and third pairs
of legs, the hind pair was drawn up and forward, and the feet grasped
the upper margin of the sack like shell, which, when first seen, was

First
Moult.

' Blackwall, “Spiders of Great Britain and Ireland,” Intro., page 6.

a se ree

COCOON LIFE AND BABYHOOD. 209

about half way removed from the abdomen, The feet pushed downward,
and at the same time the abdomen appeared to be pulled upward, until
the white pouch was gradually worked off. (Fig. 241.) The motion was
not unlike that of a child stripping off its night dress by pushing it down

the body and stepping out from the drapery.
Life within the cocoon is not wholly destitute of “moving accidents”
and “hairbreadth scapes,” if we may believe Professor Wilder, who ar-
gues that the young of Argiope cophinaria eat one another while

Cocoon yet within cocoon limits, His reason is that a comparison of
roe’ the contents of cocoons opened early in the season with those

opened later showed that the spiderlings were fewer in number
but larger in size.'| He infers the same thing from the fact that after
egress the young do prey upon one another, but without sufficient ground,
as the one fact by no means implies the other. My own observation has
been, of all species, that the young live together peacefully while within
the cocoon. However, I have chiefly studied the cocoonery of our more
northern latitudes.

In southern latitudes, where the hatching probably occurs earlier with
some species, and the period of confinement after hatching is thus much
prolonged, or the appetite of the young quickened by the climate, hunger
may assert its supremacy. Yet, even in the case of some southern spiders,
as examples of Zilla from southern California, reared during winter in my
study under conditions of temperature not very different from their native
latitude, there never appeared a trace of cannibalism until after the young
araneads had woven their first independent snares. In the case of most,
probably of all, species in our more northern climate, during the greater part
of the four months intervening between hatching and egress, the young
are probably more or less torpid by reason of the cold, and thus with
natural appetite still in abeyance. Even in our Southern States the influ-
ence of season is seen by a general suspension of activity in the insect
and aranead world; and, independent of climatic influence, Nature doubt-
less gives a semidormant tone to the spider young.

Whatever may be the truth as to Argiope and Nephila, I am certain
that many species do not have this cannibal habit within the cocoon, nor,
indeed, for some time after egress therefrom. Mr. Pollock’s observations 2
of Argiope aurelia, of Madeira, quite correspond with this statement, for
the broods were always friendly within the cocoon, and indeed for a fort-
night after leaving it.

It.

The spiderlings themselves procure exit from the cocoon in most spe-
cies. This is frequently accomplished by cutting a small opening through

‘ Proceed. Am. Asso., VITI., 1873, page 260. ? Ann. and Mag. Nat. Tist., 1865, page 460.

ihe i Natit 2

—=eo— =: rt” =.

eee

210 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the outer envelopes. (Figs. 242, 243.) I have noted these openings in co-
coons of Agalena nevia which were under observation for that purpose ;
the period at which the openings were cut was identified, and the
Egress ittle inmates seen peeping out at the round doors, of which there
ae were, in some cases, a number opened, from which also they
ocoon. : Bike :
escaped when the cocoon was agitated. Similar openings have
frequently been observed in the cocoons of Argiope cophinaria, Epeira
cornigera, Argyrodes trigonum, and in numerous examples of Epeiroid,
Tubitelarian, and Laterigrade cocoons. In these, however, as a rule, there
was only one opening, but sometimes two.
Professor Wilder has recorded some facts upon this point.’ Cocoons
of the Basket Argiope kept by him in South Carolina were never seen to
be pierced by the inmates. Of four hundred and six cocoons
Argiope obtained on James Island in the spring of 1865, only one hun-
pa dred and thirty-four were entire, presenting no opening what-
Cocoons. ever. Of the others one hundred and ninety were pierced when
found, but no spiders came out of these before May 10th. The
openings in them were similar to that made in a New York cocoon June
14th, by the inmates themselves. This hole was near the pedicle or stem
of the cocoon, and from it
the young escaped. Of the
remaining eighty-two co-
coons fifty-nine were torn
in one or more places, and
loose silk proceeded through

Fia. 242. ; Fie. 243. Ki
Drassid cocoons, to show the openings out of which the young the rents. Professor Wilder

d. . .
meveieaane once saw a little bird, about

Fic. 242. Front view. Fic. 243. Side view. A
the size of a sparrow, fly

at a cocoon hanging in a tree, make one or two quick pulls and then re-
treat. He is therefore inclined to think that all the above rents were so
caused; and, as these attacks would usually open the cocoon without in-
juring the inmates, he drew the inference that this might be a provision
of Nature, somewhat like the fertilization of flowers by insects, by which
the invasion of the cocoon should really permit the continuance of the
species.

There may be some ground for this inference, but it is certain that in
ordinary cases no such external provision is required. Birds are much
disposed to use the silken material of spider cocoonery for their
nest building operations. Mr. Thomas Meehan, the botanist, has
seen the pewit engaged in collecting spider’s spinningwork on his
grounds at Germantown. Hummingbirds are known to make large draughts
upon spider webs for nest building material. I have in my collection

Delivery
by Birds.

1 Proceed, Am, Assoc, 1873, page 260.

“— eee

COCOON LIFE AND BABYHOOD, 211

several nests built by a Vireo, the white eyed Vireo probably (Vireo novo-
boracensis), which are largely composed of the thick sheetings taken ap-
parently from the cocoons of various Orbweavers and the Speckled Agalena,
which may all have been abandoned cocoons. However, it is extremely
probable that some of them were filled with young spiders when seized.
Such seizure would not necessarily prove fatal to the young, as I have
demonstrated by experiment, substituting my fingers for the bill of a bird.

At the first pull, or as soon as a fracture had been made, a number of
the wee fellows would run from the cocoon hurry-skurry and take refuge
under surrounding objects. When a pinch or two more had widened the
fracture so as to allow the brood to escape freely, and the hand was swung
upward through the air as nearly as might be after the manner of the
supposed robber bird, a long trail of young spiders floated behind, all
hanging on as for dear life to the filaments that streamed backward like
a kite tail, and which were the united threads of the whole evicted ten-
antry forced into the utmost activity of their spinning organs.

Nearest to the fingers the filaments were thickly placed, and here the
young balloonists were massed. Further on they were less in number, and
so to the end of
this curious pennant,
where one or two
clung to the taper-
ing point of gossa-
mer. Of course, dur-
ing the rapid motion
some of the spider-
lings were detached
from the mass and
floated away upon
single or manifold
strands. It is thus easy to see that a bird carrying a torn cocoon under
similar circumstances might distribute a large portion of a brood along
the course of her flight without destroying many. Even for those re-
maining within the cocoon or clinging to the shreds thereof, there would
be good chance to escape scot free after the work of weaving the silken
material into the nest should begin. The action of birds in opening
cocoons is an accident of which spiderlings doubtless avail themselves, but
it probably goes for little or nothing in the natural delivery of the brood;
and the peculiar spinning habit of spiders tends to protect them from the
violence of such attacks when made.

Mrs. Mary Treat has informed me that the young of Argiope cophi-
naria have been observed by her escaping through the pedicle or stem
of the outer cocoon case. A reference to the figures of the cocoons of
this species in Chapter V. will show how this is done. The pedicle of

Fig. 244. Young Agalenas escaping from a plundered cocoon.

O18 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the cocoon is a short hollow tube. Just below this tube on the inside
is hung a funnel shaped silken cap, which is attached above to a strong
silken cord composed of numerous fibres, which cord passes up-
Hgress of ward through the hollow stem, sometimes forming an outward
SretoPe’s attachment to some external object. It would not be a diffi-
oung. f :
cult task for the young Argiopes to work their way between
the inner wall of the cocoon case and this cap above described, and so
along the cord and out into the air through the pedicle. If Mrs. ‘Treat’s
observation should be established as a common habit, it would, of course,
account for the fact that Professor Wilder found so many of these co-
coons without any external opening. Simply, the spiders had crawled
out through the pedicle; but I believe this is not common,
In the case of many cocoons spun by Epeira, and, indeed, by Orb-
weavers generally, there always is a selvage uniting the upper to the
lower portions of the outer case. As the spiders grow and the
The period for egress approaches, this selvyage appears to open, a
er result which is perhaps due in large part to the influence of
weather and time in loosening the tension of the threads which
close the edges of the parts. Through this open selvage the spiders are
enabled to escape with comparative ease. Eyen were there no relaxing
of tension in the uniting threads, it would be easier for the spiderlings
to cut their way out from this part of the cocoon than through the un-
broken parts. A reference to several of the cocoons described in Chap-
ter V. will show this.
It remains to be determined whether the mother in some species may
not be an active agent in delivering or aiding the deliverance of the
brood. Emerton once noticed a small Theridium gnawing at its

oe soft cocoon, and found that one side had in this way been made
eee much thinner than the remaining parts. He placed the spider
Aad, with her cocoon in a bottle, where he could watch her. She

soon recommenced the biting, and kept it up during the re-
mainder of the day. The following night the young came out. Of course
such a habit could only appear among those species that brood or watch
over their cocoons until the young are hatched, or among those who, like
various Theridioids and such Orbweavers as the Labyrinth and Tailed
spiders, make several cocoons and string them within their snares. As
most cocoons are abandoned by the mother immediately after spinning,
the enclosed young must escape without maternal aid.?

Menge observes that the warm rays of the spring sun awaken the germ
of the eggs, and by the time Mother Nature has provided a plentiful supply
of flies and mosquitoes, the young hatch. It is a peculiarity of spiders
that they do not leave the egg nest at once, but remain until legs, palps,

1 “Structure and Habits of Spiders,” page’ 104.

Pt ee , = m il * i -/

COCOON LIFE AND BABYHOOD. 213

skin, and all parts are perfected. By this time the body is covered with
hair and they possess claws and bristles; they crawl about and begin to
spin, but remain in the neighborhood of the cocoon. They have as yet
no need for food, as sufficient yelk is deposited in their bodies for present
wants.

After six or eight days the second moulting takes place, and they now
begin to feel hungry, and, when nothing else offers, attack each other, the
strong devouring the weak. Menge also noticed that when kept impris-
oned they will even eat the old skin; but when at liberty neither of these
extreme measures take place, as a general thing, inasmuch as plenty of
food is found around the place of their birth. At this time each aranead
supports itself as Nature ordained, and, its appetite becoming ravenous, it
rapidly increases in size and development. For this reason Menge never
succeeded in carrying young ones, hatched in a glass, over this period and
he doubts whether it can be done at all, even taking foreign varieties (such
as American) for the purpose. He tried Le Bon’s experiment, feeding them
from quills filled with blood of young pigeons, but without success. A few
of them may suck the blood, but most of them pay no attention whatever
to this unnaturally served food. Most grown spiders present the same dif-
ficulty, preferring to starve to death rather than accept food which they
do not fancy; even the very insects on which they live when free are re-
fused if not caught by themselves.

Menge often tried to bring to maturity a yet undescribed spider (Me-
lanophora), which he found rarely, and always full grown; but in this he

_..., failed. Although the glass was filled with flies, mosquitoes, po-
Artificial dura, etc., the spider left them untouched, and finally both in-
Rearing : j 5
Difficult. Sects and spider died. The same result attended efforts with

Saltigrades. Lineweavers and Tubeweavers were much easier to
feed, as they attack everything that falls into their web when not too large
or too much against their taste. The easiest to keep in captivity are the
Lycosids, which become tame and will take flies offered to them in the
hand.!

IV.

After the rigors: of winter have been successfully endured, the warm
days of spring first hasten the process of hatching, and then tempt the
.. Spiderlings from their cocoon. I have repeatedly observed, dur-

ae me ing a series of years, the issuing of broods and their behavior
immediately thereafter. The observations have been under fa-

vorable conditions within doors, and also out of doors from cocoons trans-
ferred from their original site and affixed to branches of shrubbery, and a
few in original site. The young of various species representing Orbweavers,

1 Menge, “Preussische Spinnen.”
ge,

214 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Tubeweavers, and Laterigrades, especially, have been studied. ‘The results
from experimental hatching are but little different from those which
everywhere transpire in Nature, and, taken together with numerous facts
noted afield, enable us to accurately sketch the life of the infant spider
just after deliverance from the cocoon.
One example, followed consecutively, will illustrate the habits of Orb-
weavers. Early in May a cocoon of Epeira insularis was taken from a
tree on the banks of the Schuylkill. It had been placed by
First the mother spider on the under side of a branch, where it

ee was best protected from the weather, and consisted externally
Tike of a ball of thick, yellow, curled floss, about one-half an inch in

diameter. (Fig. 245, C.) This was attached to the limb by a
thin coating of white tissue, from which short, strong cords entered the
ball. Within the ball were about one hundred young spiders, just fully
hatched. The cocoon was placed in a paper box, and the spiderlings re-
mained shut up in it until May 13th. Meanwhile they had made their
first moult. This cocoon was now opened and put within a large covered
paper box, which, by a dent in the side, had free communication with
the outside. Next morning I found that
the spiderlings had issued from the box
and woven a mass of delicate webbing
over the surrounding objects upon the
table. The lines were most closely spun
near the points of exit, where they re-
sembled a delicate tissue web. They were
Fig. 245. Cocoon (C) of Insular spider,on carried along the table on one side to

IR One Re Sitar a distance of five feet, on another of
two feet, and the lines decreased in number as the distance increased.
Where threads were dense the spiderlings were massed (O, Fig. 246) in
large numbers, and as the lines thinned out the numbers decreased, until
at each of the two points where the spinningwork ceased were one or two
pioneers engaged in pushing the lines further from the centre.

In point of fact, this last sentence expresses the general instinct which
controls the young on their first issue from the cocoon—they spin away,
and away from the home cradle, restlessly further and further, until they
are arrested by satisfactory surroundings and further flight is hindered,
or until they pause to undergo another moult. ‘This is undoubtedly the

impulse bestowed by Nature for the dispersion of the brood,

path a with a view to the distribution and preservation of the species,
aoocie primarily, perhaps, to the preservation of the young from their

own cannibal propensities. In order to test this matter and de-
cide the mode of procedure, I fixed attention upon one of the outposts.
Three feet from the main assembly (O, Fig. 246) a single straggler had
carried or followed a line.

me ssw —

—————— a a S| ee ee ee —

COCOON LIFE AND BABYHOOD, O16

A toy column from a box of a child’s building blocks was placed
eight inches from the point reached by the spiderling, in order to arrest
the thread which I believed she would emit. Directing a magnifying
glass upon her, I presently saw her assume the attitude common to her

Fic. 246. Assembly of spiderlings when first escaped from cocoon. O, the maze of
crossed lines found outside of box; V, the furthest limit of same.

order when about to take aeronautic flight. The eight legs were spread
in a circle, the abdomen elevated, and from the spinnerets issued a deli-
cate gossamer line, which was carried to and fro in the slight currents
prevailing even in a closed room. Quite soon the line entangled upon
the top of the column. Just as the spider was about to adventure upon
her tiny bridge, a sister broodling reached her, at the touch of whose
foot she instantly dropped downward along the side of the table and
hung, back underneath, by the emitted line. (Fig. 247, 1.) Meanwhile
the new comer unhesitatingly mounted the bridge line and crossed over the
column. (Fig. 247, 2.) The journey was made “hand over hand,” to
use a not inappropriate figure, and with the back downward, the invari-
able posture of all spiders on like occasions. The original pioneer now
reascended, and straightway followed her predecessor.
At this stage I was summoned from the room by a vis-
itor, and when I returned, in half an hour, a colony of
fifty-three spiders had been drained from the mass meet-
ing at O, Fig. 246, four feet distant, and were spread
over a series of open lines woven into a triangular net-
work fence (Fig. 248, F), into which the original line
had now expanded.

This illustrates another marked tendency of the earliest
movements, viz., the bulk of the colony follow the pio-
neers, and group themselves near together; in other words,
they are at this stage gregarious. This action was re-

™ peated asnumber of times during the next three days.

sivacticing the arop 1 found that I could always transfer the group to any
ee « chosen spot by placing thereon some elevated object.
For example, I put a second column at y (Fig. 248),

eight inches from the first column (x), and then pushed a toy dancing
puppet (z) across the table eighteen inches distant from x, In order to

216 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

test the effect of a current of air, I slightly raised an adjoining window,
admitting a light play of wind across the fence on the column x. In
three minutes two lines
were fastened upon the cap
of the puppet, and two spi-
ders had begun to cross
from the points marked 2,
3. These lines were so del-
icate that I had not seen
them until the motion of
the spiders along invisible
bridges directed particular
attention to the spots.

Fic. 248. Migration instinct. F, fence of netted lines; 1, 2, 3, Within an hour all the
points of first departure; B, bridge lines for transit; n, final colony but two had crossed

ee ae over the fence (I) to the
puppet, and were swarmed around the head, face, and chest of the figure,
and upon a mass of lines (n) that stretched to a wire (w). A triangular

bridge of lines (B) had now been formed, whose apex was
Grega- the head of the puppet (z), and which broadened out, touching
rious : : §
Habit. the columns (y and x) and connecting with the first perpendic-

ular bridge (IF) by the three principal points (1, 2, 3) from which
the migration had proceeded.

In the course of three days, by arranging various elevated objects over
the table, and breaking off the threads that floated beyond the prescribed
limits, I had induced the brood to cover a space having a linear boundary
of about twelve feet. The greater portion of the area thus bounded be-
‘ame at last sheeted by a web composed of the innumerable lines emitted
by the little spinners, so that the whole presented a quite good miniature
of the canvas tents of a traveling circus company.

For long periods the little creatures would hang quite still, separated
from each other by distances varying from three-fourths of an inch to
one, two, and three inches. In these rest-
ing moments they hung inverted between
two lines which they grasped re-
spectively by the four feet on
either side; the abdomen was
elevated somewhat, a short thread issued
from the spinnerets, and was attached to
an upper line, thus helping to support the ™% ™*
body. (Fig. 249, 1.) Occasionally the two
hind legs grasped a cross line hung upon or above the parallels, and the
thread from the spinnerets was also attached to the cross line. (Fig.
249, 2.)

Position
in Rest.

Position of spiderlings when at
rest upon assembly lines.

COCOON LIFE AND BABYHOOD. 217

A number of cocoons forwarded to me in the early spring, by Mrs.
Eigenmann, from San Diego, California, gave me an opportunity to note
the tendency of young Orbweavers in outdoor site. ‘The co-
Eseap- goons were fixed upon bushes .within the forks of branches, at
teal the time when the young were just ready to escape. They evi-
’ dently felt the fresh air of the open, as contrasted with the boxes
in which they had been confined, and at once
pushed their way from the flossy interior to the
outside of the cocoon. Then one adventurous
spirit scrambled to a branch and began to as-
cend a stem. Another and another followed,
each trailing a dragline along the surface, until
at last several threads’ were merged into one,
which the little creatures laid hold of as suc-
ceeding numbers emerged from the cocoon.
Thus a long line of them appeared climbing
up the thread, which at places swung free
from the stem, and at others hugged it closely.
(Fig. 250.) They reminded me of a watch of
sailors following each other up the shrouds of
a ship.
Here and there, at various points, individ-
uals would strike out an independent line of
progress, and would be sure to be

elas followed by some of their comrades.
ane One might be seen dangling from a

Ascend, leaf by a slender filament; another
with elevated abdomen sending out
the first lines of a tentative balloon; a third
already embarked on an aeronautic venture,
swinging free and swaying in the breeze. (Fig.
250.) The general tendency was to ascend ;
scarcely a spider went below the point at which
the egg sac was fixed. Here and there little
groups would form and hang back downward
for a while by a few crossed threads; these
again would break up, and at last, well toward a sda amen ducting Wines,
the summit of the bush, the colony, with the ex- from. To show tendency to ascend
, . = and migrate,
ception of a few independent characters, massed
themselves under a leafy shelter, and so remained pendent like a ball—legs,
palps, heads, and abdomens mingled in a confused mass. (Figs. 251, 252.)
This I suppose to be a good example of the general habit at this period.
The “balling” or “snugging” of the brood is quite sure to suggest to the
observer the appearance of a swarm of bees just escaped from the home hive.

{
n
J ‘
OO —_— ae

a a

218 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

How long does the brood remain thus massed? ‘This depends greatly
upon circumstances, particularly the velocity of the wind and
temperature of the air, A brisk wind and fair day tended to
scatter my experimental spiderlings very rapidly; indeed, during the after-
noon and night. This will best be illustrated by the following case.

Balling.

An interesting example of the habit of young spiderlings immediately
after escape from the cocoon, was seen May 23d, 1887, in a ravine upon
the ground of Ogontz, a young ladies’ school in the vicinity of
A Tented Philadelphia. When observed, the little creatures were snugged
Selon together in a ball underneath a large leaf of Indian turnip or
Jack-in-the-pulpit (Arisema triphyllum). Two smaller individuals of the
same plant stood on either flank. The tall central plant served as a sort
of tent pole, and from the margins of the broad top leaves a delicate
silken tissue spread downward to the edges of the shorter Jacks mentioned.
There was thus formed a symmetrical pavilion, within which the spider-
lings were contained, and which presented all the appearance of haying
been constructed intentionally. I am confident, however, that the deli-
cate canvas wall of this tiny tent was simply formed by the immensely
multiplied threadlets which the colony continually dragged after them as
they moved back and forth, up and down, in the preparatory stages of
settling themselves.

When first observed, the whole colony was massed in a ball as large
as a walnut underneath one of the top leaves. The spiders were of a

, yellowish brown color, and gave a pretty appearance as seen
eee through the silvery white of the silken wall against the green

f background of their tent roof. When I tapped lightly upon
the top of the leaf beneath which they were snugged, the ball instantly
broke up, and a hundred or more of the little fellows dropped swiftly
downward. Every one dragged after it a silken attachment, which filled
the inside of the pavilion with perpendicular lines. Most of the number
returned in a little while to their position. Some remained hanging at
various distances; a few who had fallen quite to the bottom of the tent,
which was limited by the top leaves of the two flanking Jack-in-the-
pulpits, ran out from under the edge of the tent and extended their ex-
cursion for a little distance beyond.

When I left the brood, Miss Skinner, the teacher of natural history in
Ogontz, kindly consented to keep it under observation, and I am indebted
to her for the following history prolonged through a period of ten days:
The colony was first observed on the morning of May 28d. The next
day was rainy and windy. On the 25th it was found that great rifts had
been made in the overhanging web or pavilion wall on the leeward side;

COCOON LIFE AND BABYHOOD.

—————— a

219

“yi

\)
Reig
ie
nN

> |

Fic. 251. The tent and assembly of young spiders beneath a leaf of
Jack-in-the-pulpit, on the grounds of Ogontz Seminary.

Ee oes a
: e ’
’

220 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

while on the windward side it was quite swept away. ‘To quote the moral-
izing sentiment of the journalist, “their frail house was more ragged than
good resolutions after a week’s wear!” ‘The spiderlings remained snugged
underneath their leaf as when first seen.

On the afternoon of May 27th the little fellows had “outgrown their
clothes, and hung them on the line, while they looked very smart in their

... new clothes, over which no one had toiled. Their change of gar-
Shedding vents had led to no change of habits,” for they were snugged to-
First ;

Coats. gether in a ball as when first observed. In other words, the

spiderlings had undergone a moult, and their white casts of skins
clung to the lines upon which the moult had been effected. This is usual
among young spiders. Mrs. Treat has even obseryed the shed skins of
baby Turret spiders! clinging to lines stretched across the top of the
mother’s abdomen, upon which the younglings had unfrocked themselves.

May 29th, 9 A.M. The colonists were still closely snugged. They had
grown some, and had thrown out a few cables to support their tent, which

‘ was then quite rickety. At five o’clock in the evening they were

eet in the same condition. May 30th, 5 P. M. A few individuals

were found spinning webs on an adjoining tree, but the majority

were “ wandering in the wilderness of life, and could not be found.” 'T'wenty-

one still clung to the old home. * * * May 81st, at 2 P.M., only five

spiderlings could be found. “These wandered about in a forlorn way like
pilgrims preparing to seek a shrine beyond the known country.”

June Ist, at 3 P. M., not one of the colony was to be found. The frag-
ments of the web and “the old clothes” were all that were left. About a
rod beyond the site of this colony Miss Skinner found a new ball of spider-
lings, apparently quite recently made; I quote the conclusion of her journal,
which relates to this second colony: “June 2d. Something has happened
to them, I know not what! Not a trace is to be found. So perish great
nations!”

Two of the young ladies of the seminary made sketches of the colony
two or three days after the first observation. At that time the enclosing
pavilion had been Blown away, nothing remaining but ‘a few straggling
lines. I have restored the pavilion from my own sketch, presenting it thus
as when first seen. (Fig. 251.) There is nothing to show how many of
the two Ogontz colonies may have survived. It is not unlikely that a few
scattered into the surrounding foliage and might have been found quietly
ensconced beneath leaves or any other sheltered position, but the proba-
bility is that most, if not all, of them perished. Such is certainly the
fate of multitudes of young Orbweavyers.?

* I reserve for the chapter on General Habits (under Moulting) the history of a brood
of Epeiras hatched upon a honeysuckle arbor in my manse yard, whose fortunes I followed
with particular interest. .

ade sities ree ey

COCOON LIFE AND BABYHOOD. 221

One of the young ladies in the natural history class of the school wrote
and published in the ‘Ogontz Mosaic” a versified account of the above
colony, which I yenture to add, as a pleasant description of and happy
comment upon the incident. It may at least serve to brighten for a mo-
ment the dullness of these pages eof details, and show that one may
find a gleam of poetic fancy even in the babyhood of despised Arachne’s
ehildren.

THE CHILDREN OF THE SPIDER WEB.

Unper a Jack-in-the-pulpit’s care,
Where the shadows are deep, and the sunlight rare
Tenderly kisses the maiden hair,
A loving mother made her nest,
And neyer did rest
Till flossy blankets and silken sheet
Enclosed her eggs in a safe retreat.
The brood was safe, but the mother dead,
For loye’s last act spent life’s last thread,
And the fair cocoon was left to swing
Till winter’s snow dissolved in spring.
The air was warm and the sunshine soft;
To and fro the breezes tossed
The tiny hammock of shining threads,
Of shimmering, silvery spider webs.
Far from the sounds of war and strife
Were the spider babies wooed to life.
On one bright day they all awoke,
Their prison doors they burst and broke ;
And, peeping through the barriers white,
Discovered a wonderful world of light.
With glad surprise they looked around,
Then a daring one, with a single bound,
Went dancing down on a tiny thread,
Making his own little spider web.
Graceful and airy,
A real fairy,
He entered this new found land of glory.

The days went by, and the babies grew. —
Were their pleasures many, their sorrows few?
Or within the silken canopy

Was there acted out a tragedy?
* * * * * *

Shall we e’er know the source
Of that wonderful force
By which the good little mother wove
Her babies’ cradle with threads of love ?
Why the eggs are laid by the little wife?
Tlow the sunlight laughs them into life ?
Where the shadows are deep, and the sunshine rare
Tenderly kisses the maiden hair,
Beneath the Jack-in-the-pulpit rest
The mysteries of the spider’s nest.

CC ————

222 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

About the middle of May, the same spring, I watched the egress of a
whole colony of the young of Epeira insularis from a cocoon which I had
hung upon an ampelopsis vine outside my study window. They
Tendency moved with great celerity and soon were widely scattered over
bar the vine. All mounted upwards, not a single one descending be-
* low the site of the cocoon; which habit, as I have observed, is
quite common to all species. A few days thereafter their tiny filaments
could be traced stretched from leaf to leaf over a large surface of the vine,
as high as ten and a half feet from the ground. But not a single web
was afterwards formed during the whole summer and autumn, and, as far
as I know, every individual perished. Those who are familiar with like
facts will readily perceive the necessity for the immense fecundity of fe-
male spiders in the production of eggs. Only under favorable circum-
stances can considerable numbers of any single colony reach maturity. My
observations on colonies of Epeira labyrinthea and Epeira triara-
Mortality nea show that twenty, thirty, or fifty may survive for a short
Saal period, and construct in the same vicinity their little orbicular
lings. snares. But these, too, soon perish under the combined assaults
of their natural enemies and unfavorable weather. It is probable,
indeed I believe that it is quite certain, when cocoons are located in spe-
cially favored spots, and the young inhabitants issue forth under specially
favored circumstances, that the majority of them pass beyond the period
of babyhood and attain middle growth, and reach in goodly proportion
mature life; but these examples must be comparatively rare.

VI.

My observations of the habits of spiderlings immediately after egress
are confirmed by such brief notes as other observers have made in natural
site. Emerton says (speaking apparently from observation) that
Other Ob- . may 5
servers, @ brood of young Epeiras may often be seen living in a common
web, and looking like a ball of wool in the top of a bush; while
below them, connected by threads to their roost, are the skins left at their
second moult, and further down, also connected by threads, the cocoon.!
I have often seen the young of Theridium tepidariorum, and of the long
legged cellar spider, Pholeus phalangioides, hanging in these cottony clus-
ters at the top of the maternal snare, the mother herself suspended beneath.
The Orbweavers thus appear to agree in this habit with these Lineweavers.
Wilder also has a brief reference in the same direction to the young of
Nephila plumipes, which, he says, even after leaving the cocoon, are more
or less gregarious, always keeping in companies, and preserving good order
while moving.”

1 Structure and Habits, page 110.
* Proceed. Amer. Acad. Arts and Sciences, VII., 1865, page 56.

COCOON LIFE AND BABYHOOD. 223

Fic. 252. Assembly of young spiders, just after escape from cocoon,
balled beneath a rose bush leaf. ;

224 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

The young of Epeira diademata, as observed in Europe,! have a like
habit. In the spring, when the spiders are newly hatched, almost as
soon as they’ leave the eggs they spin a small irregular mass of
English almost invisible lines, in the middle of which they cluster to-
ga dean gether, forming themselves into a ball about the size of a cherry
stone.? This hangs apparently in midair, and an obseryer ap-
proaching it to discover its nature touches some one of the slender lines
by which it is suspended, or some twig near enough to communicate mo-
tion to them. In an instant a hundred living atoms begin to disperse,
the solid little ball seeming for a moment to be turning into smoke, so
minute are the animals, so rapid are their motions, and so invisible the
means of their dispersion. After a few seconds, if the disturbance be not
repeated, the little creatures begin to subside again into a cluster, which is
not at once restored to its former small size, since a thousand legs, how-
ever minute, require a little time for the necessary curling, packing, and
‘settling by which this animate sphere of snugging spiderlings is formed.

_A series of careful observations, made and communicated to me by

Mrs. Treat, confirm the above records and furnish some interesting details.

Females of Epeira harrison *® were brought from New Hamp-
The | shire to Vineland in October, and there made their cocoons in
Side the same month. ‘These the mothers fastened to the ceiling after

the fashion of the Domicile spider, and as long as life lasted
manifested an unvarying love and care for the future offspring. As soon
as a cocoon was completed the mother addressed herself to protecting it
from insect foes and frost. For this purpose she scraped weather beaten
boards with her mandibles, and made little pellets of the gray chippings,
with which she covered the cocoon, which thus resembled somewhat a nat-
ural inequality in the wood.

The younglings did not leave the cocoon until the following spring.
When they first came out they moved about six inches distant and
formed a compact mass like a miniature swarm of bees, in which con-
dition they remained a day or two, Finally, the mass broke up and
formed four groups, in which they remained another day. Then they
separated, and the united spinning labors of the entire brood made a
thick web five or six inches in length and breadth. Herein they left
their first baby clothes strung thickly along the innumerable lines. 'There-
after they began to disperse, scattering everywhere around the house, each
spinning a perfect little orb not much larger than a silver dollar. At
this stage the observer began to look upon her spiderling emigrants with
dismay. Several hundred must have emerged from each cocoon; and,

1 Staveley, British Spiders, page 239.
, i ’ ] f=)
* There must be a mistake here as to size; the clusters of Diademata would surely be
much larger. * Epeira cinerea Emerton.

COCOON LIFE AND BABYHOOD. 225

besides, a number of half grown specimens brought from New England
with the colony, would be mothers in the fall. Thus, with the actual
and prospective issue, an aranead inyasion seemed imminent, carrying
therewith the prospect that house, vineyard, and grounds would be en-
swathed and shrouded in cobwebs.
Mrs. Eigenmann has informed me of like behavior on the part of the
young of Epeira gemma, at San Diego, California. A number of females
had been placed, about the 1st of November, in tin cans, where
California they deposited their large tawny brown cocoons, The cans with
fe their enclosed. cocoons were placed aside, and when opened Feb-
ruary 5th following, an interval of three months, they contained
numbers of little yellow spiders, marked with a black spot posteriorly on
the abdomen. One can was put out of doors and opened. In a few hours
a silken ladder of delicate lines had been made from the tip upward
eighteen inches to the buds and flowers of Encelia californica growing
in the garden, At the top the ladder was attached to a bud which was
bent downward, and between it and the stem of the plant some filmy
spider weaving served as a scaffold. Upon this the spiderlings had as-
sembled in three separate bunches, somewhat triangular in outline, which
suggested to the observer tiny bunches of very prolific grapes. Mrs. Eigen-
mann reinclosed the spiders within the tin, in order to ship them to me,
but in the act many escaped. The rest arrived safely, and immediately
upon the opening of the can issued forth and began to spin their delicate
filaments.

VII.

The brood fraternity of spiderlings, in connection with their rapidly
developed tendency to spin themselves away from the home centre, leads
: to the accidental formation of objects that curiously resemble
Bridge ridges, canopies, and tents. When they begin to move they
and Tent drag after them fine filaments of silk. A hundred spiderling
Making, 8g after them fine filaments of silk. undred spiderlings,
more or less, passing from point to point and back and forth
by single bridge lines, and keeping close together, will not be long in
laying out a series of lines and ribbons that suggest miniature roadway
trestles and cables of a wire bridge.

One of the most curious miniatures of this sort which I have known
was once made in my library. A package of cocoons of Zilla x-notata,
sent to me from California by Mrs. Eigenmann, was laid upon a long
table. One morning, upon entering the room, I found that the spiders
had hatched and issued from the openings in the lid of the package, a
large cylindrical fruit can. From the summit of this can, as from a
bridge pier, the spiderlings had strung their lines to books and paper
boxes laid upon the table, and thus formed a series of piers and abut-
ments. They had already woven a sheeted way several inches wide, that

eee ee | ~~

226 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

stretched above the middle of the table for five feet. ‘Thence it spread
upward, in diverging threads, to the window curtain, on which many of
the wee adventurers hung. (Fig. 253.) I kept the bridge for several
days, during which time the roadway received many additional strings,
and some of the baby bridge builders spun delicate little cobwebs along
the edges and among the trusses of their bridge, and, separating them-
selves from their fellows, set up housekeeping for themselves.

Another example shows that precisely the same habit exists among

Baby RoD
BribseIuildg
Zi

Fig. 258. Bridge of spinningwork laid by a brood of Epeiroid spiderlings.

spiders widely separated in structure. A large specimen of Ctenus was
sent to me by Prof. 8. M. Scudder, who had received it from a
Young = friend. The animal had come from Central America, and had
y liis brought her cocoon with her. This was a large conical object
* nearly an inch in diameter, constructed like the ordinary Lyco-
sid cocoon. The mother with her egg bag was placed in a box, and after
a few days, tired of lugging her cradle, hung it to the side of the box in
a hammock of loosely meshed lines. It was not long before an immense
host of little Ctenids, several hundreds in number, issued from the cocoon,
crawled out of an opening in the cover of the box, and distributed them-
selves over a large study table in my room at the Academy of Natural
Sciences.

—

SS sr eS rrmU rmCmUmUr LA ~ ———

COCOON LIFE AND BABYHOOD. 227

On opening the door one morning I was surprised to find every object
upon the table—books, manuscript, pamphlets, bottles, inkstand—including
the box in which the mother spider was contained, literally covy-

ee, ered with a mass of sheeted spinningwork, which lay over the
saa tops of the objects on the table like a thin silken cloth. It

Tower. Showed the inequalities of those objects, thus presenting a good

miniature model of the immense cantonment of a modern tray-
eling circus company, This remarkable structure concentrated upon the
tallest object on the table, a large box standing at one corner. ‘lo this

Fig, 254, Bridge lines, canopies, and turret spun by a brood of young Citigrade spiders (Ctenus).

point, evidently, the migrating brood had drifted, and here a strange sight
was presented. Favored by the breeze, one adventurous spider had ap-
parently found its output line borne upward until it caught upon the ceil-
ing. Up it mounted, and in a little while was followed by others, each
spiderling dragging after it a similar thread, until at last a tower like
structure was formed, the base of which is represented in the drawing,
Fig. 254, reaching entirely to the ceiling of the room, a distance of eight
or ten feet. At several places along this were lines which issued towards

228 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the window and other parts of *the room, marking points where little ad-
venturers, following their inherent tendency, had departed from their
“ Biffel Tower” of spinning threads, and dispersed into other parts of the
building.

VIII.

A valuable account of life within the cocoon of a mother Argiope is
given by Frederick Pollock, Esq. The cocoon, which resembles substan-
tially that of Argiope argyraspis, contained from six hundred to
Argiope one thousand bright yellow eggs glued together in the shape of
ARE a bean. The egg shells burst at the end of the fourth week.
Tike, The spiders at that time were helpless and nearly transparent.
At the end of the fifth week they cast off their first skin and
became quite lively and active. Their color at this time was a bright
yellow, with darkish legs. Their bodies were about the size of an ordi-
nary pin head. Three or four dark spots gradually developed down each
side of the abdomen. At about the end of the seventh week the spider-
lings emerged through a small hole probably gnawed by them.

After departure from their cocoon their habits, as reported by Mr. Pol-
lock, agree with those of young Epeiroids as heretofore described. ‘They
club harmoniously together, hanging closely packed in a ball, upheld by
numerous lines attached to adjacent objects. This community life con-
tinues for ten days or a fortnight, the spiders occasionally separating them-
selves from their snugged or balled estate, but always reverting to it. Dur-
ing this time they eat nothing.

At the close of a fortnight this friendly condition ceases. ‘The indi-
viduals of the brood scatter abroad, and each individual makes a round
web about the size of a penny. Mr. Pollock conjectures that on
account of the extreme weakness of these webs few insects are
held by them, and that in consequence hundreds of spiderlings
at this precarious period of their existence perish from starvation or other
causes. He thinks that not more than one or two out of the entire brood

survive. In this estimate of mortality he is doubtless correct as
Mortality fay as certain seasons are concerned. A heavy storm will destroy

First
Webs.

Amon “ne C
hee & a whole brood. The presence of some skillful enemy will work
Young. %® similar destruction, but under favorable circumstances quite a

number of the brood will survive. The contingencies, however,
are uncertain, and the life of baby spiders during the first few weeks of
their existence hangs by an even weaker thread than that which they spin.
Their little webs are strong enough to hold microscopic insects, the only
kind that spiderlings could prey upon at their time of life.

1 Qn the History and Habits of Epeira aurelia, Annals and Magazine of Natural His-
tory, page 459.

COCOON LIFE AND BABYHOOD. 229

When young Aurelia begins to construct snares it also begins to feed,
to grow, and become darker. Mr. Pollock thinks that in a month or two
from that time, according to the food it gets, the spiderling
changes its skin. The females have nine changes after leaving
the cocoon. From the first to the eighth moult these changes
take place pretty regularly, under favorable circumstances, at periods in-
creasing gradually from about fifteen to twenty-five days. For about two
days preceding each change the spider seems to eat nothing, and remains
motionless.

The operation of getting out of the old skin isa strange looking per-
formance, and is thus effected: The spider is fastened firmly, by a thread
from the spinnerets, close to the under side of the web; the
ie I legs are all gathered together, and appear to be fixed to a spot

oulting. : ; :
close by; the body hangs downwards, the skin begins to split
at the sides, and the spider, by a succession of powerful efforts, lasting
about an hour, gradually draws its legs out of the old skin. When fairly
freed, its former attitude is reversed, for it hangs with the end of its
abdomen uppermost and its legs dangling loosely down; they are now
quite soft, flexible, and semitransparent, the abdomen slender, and the
spider feeble and exhausted. It can scarcely crawl or exert itself in any
way. It remains stationary for about an hour, then turns its legs up,
and climbs by its attaching line to the web, where it remains motionless
for some forty-eight hours, after which it resumes its usual habits.

Should it at any time whilst young lose a limb or part of one, nothing

appears to occur towards its reproduction until at least one subsequent
change of skin has taken place; the new leg is not much more
Lost than half the length of the corresponding perfect part, and is
Pea: of a somewhat lighter color. These stunted limbs Mr, Pollock
thought of little use to the spider; and he could not notice that
there was any reproduction of limbs lost after the seventh change of skin.

The moults take place regularly from the first (after leaving the co-
coon) till the eighth. Then the spider is adult, and begins making

cocoons, the first in a month’s time, and others at periods within

Moulting
Periods.

Begin- from about fifteen to twenty-five days apart. About a week
ning to : a F ee
Cocoon, “ter the fifth cocoon has been made the spider changes its

skin for the last time, rests from its egg laying for about thirty
days, makes five more cocoons at intervals of from fifteen to twenty-five
days, and dies a week or so after making its last one.

The spots on the sides of the abdomens of young Aurelias gradually
disappear, and give place to handsome markings of regular transverse
bands across the abdomen of silver and orange alternating with black, a
silver thorax, and transverse stripes of brown and black on the legs.'

1 Ann, and Mag. Nat. Hist., 1865, pages 460, 461.

i
i _

230 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

TESS
Naturalists have at various times recorded descriptions of “ gregarious
spiders,” which have attracted especial interest by their singularity. Dar-
win mentions a “gregarious Epeira” found in great numbers
Grega- near St. Fe Bajada, the capital of one of the provinces of La

rious f : ;
ipidpre Plata. The spiders were large, of a black color, with ruby marks

: on their backs, and were all of one size, so that they ‘could not
Darwin. ,

have been a few old individuals with their families.”! The ver-
tical webs were separated from each other by a space of about two feet,
but were all attached to certain common lines of great length, that extend-
ed to all parts of the community. In this manner the tops of some large
bushes were encompassed by the united nets. These gregarious habits in
so typical a genus as Epeira seemed to the distinguished author to “ pre-
sent a singular case among insects which are so bloodthirsty and solitary
that even the sexes attack each other.” In point of fact Mr. Darwin
had only come across a brood of Epeiroids, who, for some reason of en-
vironment, as protection from the wind, freedom from enemies, or abun-
dance of food, or from sluggishness of nature, had kept within a com-
paratively limited space after egress from the cocoon. It is therefore
not allowable to speak of this colony as a “community,” in the ordinary
sense of the word as applied to such social insects as ants, termites, bees,
and wasps.
Don Felix de Azara had the same misconception, if indeed it be one.
Although the family of spiders, he says, is for the most part regarded as
of solitary habit, there is one in Paraguay which lives in a com-
Spider munity to the number of more than a hundred individuals.

mmu- * . ‘
aire Each spider builds a nest larger than a hat, and suspends it aloft
Azara. %t the canopy of a high tree or the ridge piece of a roof, in such

a manner as to be a little sheltered from above. From this a
great number of threads issue in all directions, into every available part.
The lines, in fact, are fifty or sixty feet long, white and thick. They are
traversed by other threads of great fineness, upon which are entangled
winged ants and other insects, which serve as food for the community of
spiders, each individual of which eats what itself had trapped. These spi-
ders all die in autumn, but leave in their nest eggs which are hatched out
the ensuing spring.” In both the above cases the facts are undoubtedly
recorded correctly; but the inference from them can scarcely be justified.

Darwin, who briefly refers to the account of Azara, appears to be quite
right in thinking the Spaniard’s “community” to be of the same species
as his own, although Walckenaer gives in a note the opinion that the

' Voyage of Beagle, Vol. IIL, Zoology.

* Voyages dans L’Amérique Meridionale. Par Don Felix de Azara. Tome Premier, page
212, 1806. Walckenaer’s French edition.

4 COCOON LIFE AND BABYHOOD. 231

y aH aaa! Si
. ie a (Sp BS
. Paes aunt HS Za Be He HH
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Waal Maar ae :
VSS, aly SS: \ SY
P< S| as:
! 4
A MAIR
i] <A —s
hie «| wr HE |X
: [| Pegi} itt ) ‘ ne na
XUN \S
WEE ON LI KWESY
SSESF
| K S \ | Woseze
LAs a i i (ae aa FE
Wl P ed
, O\ SS (|
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it Th]
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=
——

rudimentary nest is shown in the angles, and the orientation of the froe

Via. 255. A colony of young Epeira triaranea upon a lattice screen. The

radius illustrated.
|
|

Zou AMERICAN SPIDERS AND THEIR SPINNINGWORK.

spinningwork indicated a Lineweayer—Theridium, perhaps. Darwin, how-
ever, saw no “central nest” in which the eggs were laid; and here I
think he misreads Azara, who appears to me to mean that every
orbweb has a cocoon or string of cocoons attached to it, pre-
cisely as is the case with Cyclosa caudata, or hung in the retitelarian
labyrinth aboye the orb, as is the case with the Labyrinth spider. That
Darwin saw no cocoons is not strange, for his observation was made in
spring (“May-June”), and as the colony was evidently a spring brood,
doubtless immature, the pairing had not begun, and the eggs would not
have been deposited until autumn, which in fact was the time when Azara
saw them. The two accounts do not, therefore, contradict, but confirm
each other. All the details of these two narratives—the number of the
brood, the uniformity of the size, the distance by which the individual
webs were separated, the straggling uniting threads, which were probably
simply incidental to the Orbweavers’ habitual behavior, and not an essen-
tial part of the snare—seem to me to justify the conclusion that these
were not “communities,” but simply accidental assemblages of individuals,
each one of which still maintained its solitary habit. Nevertheless, one
should express this opinion with some reservation in view of the possibili-
ties of Nature.

The opinion here expressed is largely based on studies of broods both in_
artificial sites and afield. I have often found small groups of the Laby-
rinth spider, which have been spoken of as “colonies,” occupy-
ing one bush, and presenting an appearance in kind the same as,
but greatly less in degree, than the broods described by Azara
and Darwin. I have seen snares of young Triaraneas hung along the
strips of a latticed chicken house, in great numbers and close contiguity,
more than forty of such webs appearing within a space of fifteen feet.
Another similar colony appeared in the latticed screen of a cottage kitchen
at Asbury Park, a section of which is given at Fig. 255. The rudimentary
nests appear in the angles; and the tendency of the species, at the begin-
ning of life, to preserve the characteristic open sector and free radius at
the top of the orb, is well shown, as also the disposition to vary the loca-
tion of the nest to right or left, according to convenience or whim.

An old stone barn in the vicinity of Philadelphia has at times pre-
sented to me an appearance most interesting and beautiful, by reason of the

immense number of orbwebs spread over one of the gables.

Darwin.

Spider
Colonies.

sacl Placing the face close to the wall so as to get the right reflec-
Barn: tion of light, I saw the whole surface of the building, from foun-

dation to roof, covered with orbs as closely set as space would
well allow. Along the cornice of the roof they were especially massed
in manner not at all unlike the “community” of Azara. As the morning
light played upon the beaded spirals and white strands, or flashed in rain-
bow colors from gathered dewdrops, the whole showed a natural decoration

COCOON LIFE AND BABYHOOD.

233

Fic. 256. A colony of Orbweaving spiders, formed on lines spun between boat houses
extending into an inlet of the sea.

234 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

quite beyond the power of human art. These orbs were not all of one
dimension, although multitudes did agree in size, but they were nearly
all of two species, Epeira strix and Epeira triaranea, principally the former.

They were undoubtedly composed of several broods of these ara-
Acci- neads, of an equal age, who had, under favorable circumstances,

pane been distributed in the same vicinity.! I may here again refer
blage. (see Vol. I., page 64) to the numerous colonies of Epeira sclo-

petaria which domicile within a limited area upon the surfaces
of the boat houses at Atlantic City and Cape May, as another example
of accidental rather than gregarious assemblage. These colonies spin their
orbs between the outer walls,
above the Inlet waters and
hang the snares to foundation
lines ten and fifteen feet long.
(Fig. 256.)

My notes show several ob-
servations of this kind: At
the summit of a tall branch-
ing weed had been woven a
large orbweb, which, probably
after it had been abandoned,
was occupied by a group of
young Epeiroids, Furrow spi-
ders. These little settlers, with
a fine acquisitiveness that sug-
gested the once famous Amer-
ican theory of “squatter sov-
ereignty,” had seized upon the
araneal commons, and every
one appropriating to itself a
corner or segment of the ter-
ritory, had woven a small orb-
web, These snares were pitched
between the radii, which in

Ii. 257. Young Orbweayers nested on an adult snare. places were cut away, and

which made excellent founda-
tion lines. (Fig. 257.) This certainly seemed a canny operation, and might
have been held to savor of economy did not one know the prodigality of
spiders in the matter of their spinningwork.

This use of large abandoned webs I have elsewhere seen afield and also
around houses, once in a hotel outbuilding, once in a broken window of a

‘IT have observed the same phenomenon at the Fish House of the historic club in the
“State of Schuylkill,” on the banks of the Schuylkill River in Philadelphia.

COCOON LIFE AND BABYHOOD. 235

tannery. (Fig. 258.) I had never raised the thought of a “community ”
to Account for these groupings, for I knew that the species represented
therein had the solitary habit characteristic of Orbweavers.

Another example fell under my observation, which more closely resem-
bled those cited by Darwin and Azara. I once found on the slopes of
Brush Mountain, Pennsylvania, just above the banks of the Juniata River,
a large colony of the young of Uloborus plumipes. Their pretty horizontal
webs were spread over the tops of a clump of low laurel bushes covering
an area ten or twelve feet in diameter. It needed only increased size and
more vigorous spinningwork to establish a close correspondence between
the appearance of this brood’s en-
‘ampment and the ‘‘ community ”
of La Plata.

A case somewhat similar to this
is recorded by Vinson as observed
in the African island of Réunion.!
In the great net of the Epeiroids,
stretched between trees of Panda-
nus, one might count the inmates
living in colony (en famille), and
in real harmony. ‘There were
found spiders of all ages and sizes;
there were Nephila nigra and N.
inaurata, messmates so hearty; and
there came the Linyphize to estab-
lish themselves upon these huge
snares in order to glean the petty
prey. It is Vinson’s opinion that
these little aranead parasites sought
the protection of the large Orb- Fic. 258. “Squatter sovereignty.’ A colony of young
weavers by suspending themselves pt pas Pa Me eee WIR
thereto in innumerable quantities,
in order to avoid the birds and other enemies. Probably the “ seeking”
consists in the simple and natural fact that the young were bred in the
neighborhood of the webs, and continued where they were hatched, avail-
ing themselves of the spare spaces in the webs of their gigantic kindred,
precisely as the little Furrow spiders of our figures. (Figs. 257, 258.) The
Linyphias, however, apparently presented a case of real nest parasitism.

Thus far our observations upon the habits of young spiders have been
chiefly confined to the broods of Orbweayers. We turn now to consider

' Araneides des Isles de la Réunion, ete., pages xix., xxi.

236 AMERICAN SPIDERS AND THEIR SPINNINGWORK,

the habits of the younglings of other tribes. We shall find that, in propor-
tion as the general habits of the species approach one another, there is a
likeness in the behavior of the young, Between Lineweavers
Young = and Orbweayers there is little difference. Their cocoons are com-
Bea monly suspended within the intersecting lines that constitute the
’ regular snare. The little ones issue from the cocoon and arrange
themselves in fluffy masses, following the tendency, which has already been
noted, to climb as far towards the top as they can. Here they remain for
a little while undisturbed by the mother and, as far as I know, unre-
garded by her. Soon they spin themselves away to various convenient
sites in the neighborhood, and establish housekeeping for themselves. Thus,
in the case of those spiders which weave several cocoons, one brood after
another will appear and disappear.

Pholeus phalangioides, the “daddy longlegs” or cellar spider of our
province, carries her bundle of eggs in her jaws until the little ones are
ready to hatch, when she abandons them and they
take their place, in accordance with the custom of
other Lineweavers, at the top of the home snare.

It will thus be seen that the young Lineweavers
reared within the limits of the maternal snare have
precisely the same habit as Orbweavers, like Epeira
labyrinthea, that deposit their cocoons near their
orbs within a supplemental snare of retitelarian lines.

The young of Agalena nevia remain within the
cocoon until they are lively little creatures covered
AOE bearer Mantes with black hairs, apparently well able to skirmish

Saltigrades, Bpiblemum scen- for themselves. ‘They then issue forth, and may be
a a bark. (After found in great multitudes upon a dewy morning

hanging beneath little sheeted webs spun upon the
grass, leaves, upon the roadside, and even within the furrows of newly
plowed fields. They are pretty little snares when thus covered
with the beaded drops of morning dew, forming beautiful ob-
jects for study under a common pocket lens.

Tegenaria medicinalis presents little difference from Agalena in the gen-
eral habit of the young. They leave the egg nest, rapidly disperse, and
spread themselves into the neighborhood and immediately construct their
characteristic webs. :

The tendency of young spiders of the Wandering tribes to form colo-
nies is not very decided, as, of course, the manner in which the young-
lings are reared within the mother’s nest until they are able to set up
housekeeping for themselves precludes such special habits as we find in
the assemblages of Orbweavers and Lineweavers. But when the young
Saltigrades have abandoned the maternal cell, groups of them may be
seen underneath a bit of bark occupying their own tiny cells, which lie

Agalena.

COCOON LIFE AND BABYHOOD. 237

close to each other, forming thus a miniature colony. One of these settle-
ments I have redrawn from Mr. Otto Herman's description of the Hun-
garian spider fauna.!

Bl.

The disposition of some young spiders to settle in colonies in the
neighborhood of their maternal origin may well be seen in the case of
the Medicinal spider. For example, in my church cellar several
cies windows have been left undisturbed, by my directions, in order
that the various species inhabiting them might have free op-
portunity to multiply and build in a natural way. On one window, which
is represented in the accompanying cut (Fig. 260), an interesting spectacle
is presented to the observer. The opening for the window is a deep one,
the wall being four feet in thickness. The glass opens into an area exca-
vated from the embankment outside, and through which light falls, dimly
illuminating the window space. The’ whole place is occupied by spiders
of several species.

In the forefront may be seen the web of intersecting lines spun by
Theridium serpentinum. The mother has disappeared, but her eight co-
coons of flossy white silk still (in midwinter) hang in the midst of the
maze of crossed lines, almost as spotless as when spun, appearing to have
little capacity to gather the dust and muck of the cellar. Just beyond,
and almost filling the capacious opening, the long cables of Theridium
tepidariorum are stretched. Here the mother had her home, and she
has left a dozen of her pear shaped, yellowish brown egg bags within the
meshes of the snare. Beneath this a species of Linyphia has stretched
her sheet like web, and as late as Christmas (1889) was found hanging
beneath it, apparently patiently waiting to pick up such chance prey as
the late season might bring her. Small snares of young individuals of
the two species of Theridium above mentioned are woven at various points
in the intervals. In a few the proprietors may be seen hanging back down-
ward; from others the spinners have disappeared into various crevices and
rugosities of the rough plastered window.

Further on we reach the glass window frames close against the area.
In either corner, and occupying the angle for a considerable distance on
either side, are stretched the triangular shaped webs of Tegenaria
medicinalis. Some of them are quite large. All are covered
with cellar dust and soot. Some of them look broken and aban-
doned. In others, if one follows the snare to the angle and runs his fin-
ger into the turret, he will find still living the sombre colored spider that
wove the web. These webs and towers are or were the snares and homes

Medicinal
Spider.

1 Wohnungscolonie yon Hpiblemum scenicum unter Rinden. Magyarorszig Pok-fadija
(Hungarian Spider Fauna), Vol. L, irta Herman Otto, pl. iii., Pig. 64.

238 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

COCOON LIFE AND BABYHOOD, 239

of the adult species, both male and female. Between these larger webs,
occupying the angles and spread along the window frames, one sees many
smaller webs. ‘They occupy the angles where the intersecting frames of
the sash cross one another. ‘hey are built just underneath the frames,
They are stretched from the top of the frame to the surface of the glass,
and some of them are woven upon the glass itself. They are small as
compared with the webs of the adults, and they are of a bright bluish or
lead colored silk, which has not been defiled by dust.

I counted on this window as many as one hundred and six of these
little tents, and in the neighborhood many spiderlings may be found. But

many more have disappeared. Whither have they gone? Alas,
A Camp there can be no doubt that many of them have fallen victims
se of that fratricidal strife which is sure to appear when the young
‘of any brood of spiderlings have once set up housekeeping for
themselves. Others, doubtless, have gone to satisfy the appetite of their
own mothers, who, when once their broodlings have left the maternal care,
make no distinction between their own and another mother’s offspring, but
eat all indiscriminately that fall within their maws, while on still others
alien species have preyed.

The window presents an interesting object as it is thus depicted, and
the carefully made photograph, which has assisted the artist’s study, accu-
rately presents to the reader what may be seen by the student who takes
his stand with the author and looks into this window. Elsewhere through-
out the cellar the same phenomenon is presented. On another window I
counted fifty-three of these youngling snares spread in like positions; but
the one here figured is the most interesting object, and presents the largest
exhibit I have seen of youthful spider industry intermingled on a natural
site with the webs of adults and of other species.

The Swedish naturalist Clerck saw many little Argyronetas swimming

in the month of July,’ which indicates that they are hatched
be! about that time, and appeared greatly to enjoy themselves in
Spiders. sporting through the element which forms the environment of

their home. The instinct of swimming is as fully developed in
these little ones at the very outset of life as in their parents.

According to De Lignac,? when the mothers of Argyroneta aquatica are
about to oviposit they construct a new silken bell or renew that which

: they have already made. ‘The eggs are enclosed therein, and
Swim- when hatched one may see issuing from the beautiful balloon,
ming Eix- f ‘ fale ; iia 5 i
cursions. Which is shining white, a prodigious quantity of little bubbles,

brilliant as quicksilver, which swim about in different. ways!
These are the young water spiders. One female, obseryed and reported by
this author made her cocoon on the 15th of April, and on the 3d of

' Aran. Svecici, pages 149, 150. 2 Op. cit., page 53.

ee Oe 7

as — ea eee ee ee

240 AMERICAN SPIDERS AND THEIR SPINNINGWORK.,

June following the little spiderlings issued forth. Their excursion was not

simply for observation. They mounted in search of air. Many made little

cells of their own upon a water plant

wwii which they found in the vase; never-

iat YW theless, they still continued to go into

and out of the maternal mansion. Some

of them threw themselves upon the

corpse of a dragon fly larva, each one

tugging at his own side in such a way

that they tore the body as ferociously as

two dogs engaged in dragging at a piece
of flesh.

Fig. 261. Lycosid mother, with her newly On the fifteenth day they changed
hatched brood upon her back. . s

their skin, and our observer saw a large

number of their moults floating upon the surface of the water. After

the young spiders had left the maternal cell it appeared transparent; but

two days after the advent of the family a part appeared to be renewed,

satiny, and opaque. When the balloon was deserted, the male, who had

constructed a beautiful cell upon the surface of the water, sometimes came

to visit the old apartment. hese spiders have a local attachment for the

neighborhood of their cells.

XII.

The Lycosid mother referred to (page 143) presented a good oppor-
tunity to observe the habits of her younglings. One month after her co-
coon had been made, June 4th, the
Spider- spider was found with the young
ae hatched and massed upon her body,
Hane from caput to abdomen. The empty
egg sac still clung to her spinnerets,
and the younglings were grouped upon the
upper part of the same. The abdomens of the
little spiders were of a light yellow color, the
legs of a greenish brown or slate color, and the
brood were tightly packed upon and around
each other, the lower layers apparently holding
on to the mother’s body and the upper upon
those beneath it. Twenty-four hours thereafter
the cocoon was dropped, and the spiderlings
clung to the mother alone. An examination of Fic. 262. The site of brood of Dolo-
the cocoon showed that the young had escaped {164s between a pont landing and
from the thin seam or joint formed by the
union of the egg cover and the circular cushion when the whole was
pulled up at the circumference into globular shape.

Pe ae eS ee ee eee ——— vo = -“ 7),

COCOON LIFE AND BABYHOOD. 241

On June 11th, one week after the hatching of the young Lycosids, one
hundred had abandoned the maternal perch and were dispersed over the
inner surface of the jar and upon a series of lines stretched from side to
side. About half as many more remained upon the mother’s back, but
by the 18th, two days thereafter, all had dismounted. In the meantime
they had inereased in size at least half, apparently without food.!

One summer, at the steamboat landing of Lake Saratoga, New York,
between the platform and the logs driven as piles to protect it, I observed

a large nest of interlacing lines within which hung a round co-

Young oon from half to three-fourths of an inch in diameter, Imme-
Dee diately beneath the cocoon many young spiders were massed in
seaduel ately bene: any y g@ spiders > mass

colony, hanging inverted, in the usual posture, from the crossed
lines of the maze. These were
the little fellows who had been
hatched within the swinging
ege bag, and who had doubt-
less issued therefrom within
the last week or ten days. At
least, they were so well grown
that they might have been of
that age.

; il
the habit of Dolomedes, I i oe
the habit of Dolomedes, I in- ‘ini
ferred that this may have been
the cocoon nest of one of the
large Dolomede spiders that
frequent the borders of our i
American lakes and other wa- ae i i ea a HSH j
ters. I captured some of the iN Nt “Al MTT
young spiders, with some diffi- Fis. 263. View of Dolomede cocoon in site, and part of the
culty however, for they were brood hanging to the supporting lines.

old and active enough to scamper away upon the least agitation of the
snare. An examination showed that they were young Dolomedes, proba-
bly Dolomedes tenebrosus, a spider that attains great size under favor-
able circumstances. No doubt, the mother had carried her cocoon along
the shore, hiding among rocks or underneath the platform of the boat
landing, until Nature prompted her to the last action characteristic of her

' Proceedings Acad. Nat. Sci., Phila., 1884, page 138, “ How Lycosa fabricates her round
cocoon.”

242 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

species. ‘Thereupon she swung it within a tented maze as described and
figured, and probably set herself to watch in some convenient position.
She, however, had disappeared when the naturalist came upon the scene,
and may have perished or returned to her life occupation of capturing
insects upon the blue waters of Lake Saratoga. Fig. 262 shows the posi-
tion of the snare and cocoon in site, and Fig. 263 shows the snare enlarged,
with the cocoon about natural size, and the young clustered beneath.

When one approaches the cocooning nest of Pucetia aurora he usually
sees the mother hovering over her offspring, or starting a new sac of eggs.
She makes two and sometimes three cocoons on one twig. Some-
times the young ones will still be within an old cocoon while
the mother is enclosing a new bundle of eggs immediately ad-
joining the old one. The young were not seen upon the mother’s body
by Mr. Wright, who forwarded the specimens to me from California. The
mother stays close by her cocoon nest. If the spiderlings be hatched out,
she will perhaps drop down a foot or more. If the first effort to capture
her be not successful, she will not drop to the ground unless forced to do
so. If engaged in carrying her cocoons, it requires force to separate her
from them. (See Chapter V., page 147, and Fig. 180.)

The young ones take alarm sooner than their mother. ‘They drop down
a few inches from their perch on the maze of intersecting lines surround-
ing the cocoon, or, at times, as far as two feet, each one suspended to a
tiny thread, forming thus a pretty swinging fringe to the overhanging nest.
In a few moments, if no further alarm be created, the younglings reas-
cend by their traplines to their perch, but, if they be frightened again,
will drop entirely to the ground and run into concealment. In such cases
the little ones were not observed to jump, as is the habit with the mother,

The young of Lycosids generally escape from their cocoons through
the seam which extends around the central part. It is thinner at this
point and splits nearly around the whole circumference, so that
the young come out in a body. But with the Turret spider,
Lycosa arenicola, the young cut a smooth, round hole in the
cocoon just large enough for them to come out one by one. The first
three weeks the little objects are piled all over the head and thorax of the
mother, often completely blinding her. They seem ambitious to reach
the highest point, and jostle and crowd one another in various efforts to
be at the head of the heap. The mother patiently endures this for a
time, but when her younglings become too thick over her eyes, she takes
her long fore legs, which she uses as feelers or hands, reaches up and
scrapes off an armful and holds them straight in front of her. Soon she
gently releases them, slowly opening her arms, and they quietly take their
places around the edge of the tower, where they usually remain until the
mother goes below, when they all follow. Upon her reappearance they are
again stationed upon her back.

Pucetia
aurora.

Turret
Spider.

i | 7
a a

COCOON LIFE AND BABYHOOD. 243

The young do not all leave the mother at the same time, but go out in
detachments, when about three weeks old. When three or four weeks old
the mother manifests a disposition to send them adrift. She
is no longer quiet and patient, but frequently picks up one of
her babies and throws it across the jar, yet seems to be careful
not to injure it. She behaves much in the same way that the higher
animals do in weaning their young.

When the spiderlings leave the mother’s back they run up a tree or
some neighboring plant and are lost to sight. Some linger with the mother
until the cold weather begins. ‘The mother clears the ragged webs and
moults from her body and looks plump and bright. She sits on the top
of her tower with the remaining little ones stationed around the edge.
They now seldom rest upon her, and when she goes within her burrow
they all follow. Upon her reappearance a few spiderlings, it was observed,
had availed themselves of the opportunity of being carried up upon her
back, but they did not remain there.

One of this brood was observed making a small burrow in the jar in
which it was confined. The tube was less than one-fourth inch in diam-

eter, and the spiderling was two days in excavating an inch below
ae the surface. On the top of this burrow it built up a tiny tower

uilding. : A 5 .

fully half an inch high, which was made wholly of earth inter-
mingled with web. In digging, the diminutive architect brought the little
pellets in its mandibles, and those which it did not wish to use in the
tower it let fall by the side. It did not shoot the earth to a distance as
the adult Turret spider does, but stood on the top of its tower, opened its
mandibles, and let the pellet drop. At the same time it threw apart its
legs as if that would help it to dispose of the earth, a movement which
Mrs. Treat speaks of as being very baby like.

The actions of this little Turret builder showed emphatically that she had
shut herself apart from the rest of the family and would not be annoyed by
them, Frequently one of her brothers or sisters, meandering about, came
to its little tower, and not often would one pass by without going up and
looking in. ‘This always seemed to exasperate the small householder. She
dropped her work, sprang from the top of her tower, and sometimes chased
the fleeing brother half across the jar, then turned and went back to her
work. No such disposition was manifested, as far as Mrs. 'Treat observed,
as long as the younglings remained with the mother. During that period
they crowd together, walk over each other, and never have any quarrels.

At the end of the sixth week after birth most of the brood, if per-
mitted, would abandon the maternal nest and build tubes and towers for

_ themselves. ‘Two had erected their towers within two inches of
ea one another, and they sat on the tops of their turrets, often
facing each other and watching the tiny scavenger beetles that

bred in the jar and lived on the refuse insects thrown out by the mother,

_ Leaving
Home.

244 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

These beetles were food for the young housekeepers, but Mrs. Treat
thought that they were not to their taste as much as flies, although they
dared not as yet take a living house fly, and if one came near them they
quickly dodged within their burrows. If a fly were killed and laid on
the tower, however, they would try to take it within, but it being impos-
sible to do this with the wings and legs adhering, they made many in-
genious but futile attempts to get the large carcass inside the burrow. If
the wings and legs were removed from the insect, and laid upon the tower,
the carcass was soon carried below and after a few hours was brought up
to dry and thrown out.!

In November the Tiger spiders all hermetically close their doors and
keep them shut until the following April, when they again come forth,
the females each with a cocoon of eggs attached to the spinner-
ets. The eggs hatch in May, and the young spiders crawl upon
the mother’s back, literally covering her body. After a few days
they leave her, and all at once come rushing out of the burrow. For two
or three months these young spiders flit about here and there over bushes
and on the lower branches of trees, seemingly ambitious to get to higher
places.

Toward the end of July their roving lives cease, and they settle down
and dig little burrows in the earth, which they do not conceal the first
season. The wasps do not molest the young ones. The following spring,
when a year old, they are little more than half grown, and during the
summer they grow rapidly and moult several times, each time changing
their appearance. By August they seem to be nearly full grown, when
their enemy, the wasp, makes havoe among them. By thus tracing the
life history of this spider we find it to be two years old before the first
brood of young are hatched, and, if no accident befalls it, it probably lives
several years.?

Lycosa
tigrina.

XIII.

Mr. Frederick Enock * determined the manner in which the young of
Atypus piceus issue from the parental nest, and their subsequent behavior.
October 15th he dug up five tubes, each containing a male arid
female. The males were removed, and the tubes containing the
impregnated females were reset in a bank at the bottom of a
garden, and were kept under daily notice during the seasons following.
March 28th of the next year the aerial extensions of the tubes, which dur-
ing the winter had laid nearly flat upon the bank, showed signs of being
repaired by the inmates. On the next day in the apex of each of the
five tubes there was observed a small round hole one-sixteenth of an inch

Young
Atypine.

* Mrs. Mary Treat, “Home Studies in Nature,” Harper’s Magazine, May, 1880.
* Idem, page 712. * Trans. Ento. Soc. Lond., 1885, page 395.

2 dete ts

COCOON LIFE AND BABYHOOD. 245

in diameter. The succeeding day, March 30th, was warm, and at ten
o’clock morning a young Atypus was observed to emerge from one of
these holes. It was shortly followed by others, until ten had left the home
of their birth never more to return, A few younglings also issued from
two others of the tubes.

The first young Atypus that emerged walked a short distance to the
foot of a grass stem, up which it crawled, leaving its silken thread as

it went along. When it had climbed about an inch high an-

“ Fol- other young one came out, took hold of the first line, adding its

ret He thread to it; and so on, each successive youngling following the

‘leader, which, by the time the tenth one emerged, had mounted

up several inches. As the leader climbed from stem to stem it bridged

over the intervening space with the never ending silken cord, along which

each successive spiderling followed, strengthening. it as they passed, until
it became quite visible, glistening in the sun.

The spiderlings above alluded to kept mounting up higher and higher,
and ascended to the top of a number of pea vine sticks which had been
planted in the neighborhood, and were about three feet and a half above
the ground. The first adventurer, having arrived at the top of one of
these sticks, walked around and around it. The others soon joined the
first, and none seemed inclined to descend by the way the.party mounted.
The rising wind gently swayed the sticks about, until some of the spiders
were blown off into midair, but still keeping hold upon their endless
silken cord until they became attached to other sticks. These they mount-
ed as they did the first; but were again and finally carried off by the
breeze at five o’clock afternoon, and landed upon the ground, where they
hid themselves among the grass and rubbish, no doubt taking lodgings
therein for the night, during which there was a sharp frost.

The next morning all the small outlets of the tubes were carefully
spun up, and, judging by the character of the web, Mr. Enock thought
that the mothers had closed the openings to prevent the remaining mem-
bers of the family leaving the parental nest until more favorable weather
would permit them to do so with safety.

The above behavior appears to represent accurately the ordinary habit
of the spiderling Atypus immediately after exode. That is to say, it first

; seeks a position at the summit of any neighboring plants or
First elevated objects, from which it is carried away by the wind
ae upon an aeronautic expedition of greater or less extent. Fall-
Habits. img upon the earth, it conceals itself for a little while, and then

proceeds to dig in the sand or soil a tube which is extremely
minute, corresponding in size to the spider digging it. As the spider-
ling grows it enlarges its tube, or, removing from the one in which its
child life was passed, prepares another nest better adapted to its mature
condition.

—_—

- — ee
ni ee

246 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

In these particulars the young of Atypus differ little, perhaps, I may
say, not at all, from the habits of Lycosids, after they have left their
mother’s back and started housekeeping for themselves. Indeed, the re-
semblance has a wider range among the tribes, inasmuch as Orbweavers,
Laterigrades, and Saltigrades show the same disposition to seek elevated
objects immediately after exode, and thence procure dispersion by means
of the wind.

The mother Atypus may occasionally carry its young upon its back
during residence within the parental nest, but has not been seen doing
this outside of its cave. This fact is not strange, since it rarely leaves
its tube at all, but spends its entire life within its silken domicile, which
is for it alike home, snare, nursery, and grave. According to Mr. Enock,
maturity is not reached until the Atypus is at least four years old.

The young of Atypus piceus were seen by Mr. Enock, September 25th,
in the same nest with the female, looking very white and moving feebly,
as evidently just hatched. He found the young nested with the mother
at various dates through September, October, November, and again in
March and April of the year following. It is thus established that after
the young leave the cocoon in August and September, they remain with
their mother during the entire autumn and winter, and during the early
spring until the weather is mild enough to justify their leaving the ma-
ternal home and establishing nests of their own.

What they feed upon during this period is not known. Much of the
time, no doubt, they are in a torpid condition, requiring no food. There

is not the slightest evidence that they prey upon one another.
Nurture [i is possible that the mother may provide food for them, and,
aoe indeed, this is highly probable. If so, these troglodyte spiders

furnish a beautiful example of domesticity; and the maternal
care shown by creatures so unprepossessing in personal appearance and
occupants of such gloomy homes, is not excelled by that of any of the
known lower animals. I might, perhaps, truthfully add that the more
highly organized vertebrates scarcely exhibit a greater amount of maternal
tenderness and care.

The immense cocoon of Mygale, sometimes as large as a hen’s egg, is
stocked with as many as two thousand eggs. In Cayenne the little My-

galide, when issuing from the cocoon, are attacked and de-
Young voured by red ants, and are too feeble to offer effectual resist-
Taran- ;
ree ial ance. Walckenaer describes the contents of a cocoon of Mygale

avicularia from Cayenne, which was infested by a multitude of
parasitic Cynips. Numbers of young spiders were found therein. They
were about two lines long, of uniform yellowish white color, except at the
eye space, which was brown. The long spinnerets showed at the apex
of the abdomen. The mandibles were prominent and curved, the eyes
very apparent. All the characteristics of the genus were well developed.

COCOON LIFE AND BABYHOOD. 247

The inner intermediate eyes were large and of a reddish brown, The
first pair of legs were longer than the fourth pair.!

It is a suggestive fact in the natural history of these immense repre-
sentatives of a race so destructive to insect life to find them the victims
of such puny creatures as parasitic Ichneumon flies and Cynips, and to
see their young devoured in multitudes as a delicate morsel by little
red ants. It is thus that Dame Nature knows how to keep an equilib-
rium in the thronging life of the insect world, and, moreover, to bring
it about by what seems an apt and admirable stroke of justice well in
accordance with ‘the eternal fitness of things.”

XIV.

Mr. Moggridge was fortunate enough to see the female of Nemesia me-
redionalis constructing a trapdoor in captivity, after having been placed in
a flower pot full of earth, in which a cylindrical hole had been
made in order to forward the spider’s operations. She quickly
disappeared into this hole, and during the night following made
a thin web over the aperture, into which she wove any materials that came
to hand. At this stage the trapdoor resembled a rudely constructed hori-
zontal orbweb, attached by two or three threads to the earth at the mouth
of the hole. In this web were caught bits of earth, moss, leaves, etc.,
which the spider had thrown into it from above. On the second night
the door was nearly the normal texture and thickness, but in no case would
it open completely. Mr. Moggridge believed that when a door is fastened,
the few threads which serve as supports and connect it with the earth on
either side, are severed.

Young Trapdoor spiders, both of the cork and wafer kind, when taken
from the nest of the mother, will make their own perfect little dwelling
in captivity, and Moggridge observed them construct tube and door within
fifteen hours. This may be favorably compared with the work of the adult
Cteniza moggridgii, which the same observer saw make a perfect tube and
furnish it with a movable door in a single night when confined under
gauze or moist earth.”

The same author has enabled us to decide that the young Nemesia
proceeds in precisely the same manner as the adult when it builds a nest.
While engaged at night in sketching, he detected something mov-
ing at the mouth of a tiny hole just large enough to admit a
quill pen, in a mass of earth near where he sat. The lamplght
fell full upon it, and he soon saw that the moving object was a very small
spider, which was at work in the mouth of its tube. The opening of the
tube was completely uncovered, and it soon became apparent that the little
aranead was intent upon remedying this deacianoy, After a few threads

Making a
Trapdoor.

Young
Builders.

1 Walckenaer, Apttres, Vol. I., pages 218, 219. Grandoor Gaiden, Supplement, page 245.

bo
=
e oe)

AMERICAN SPIDERS AND THEIR SPINNINGWORK,

had been spun from side to side of the tube, he watched the spider
making one or two hasty sorties, apparently spinning all the while; and
finally saw her gather up an armful, as it were, of earth and lay this on
the web.

After this the occupant of the tube was concealed, but the observer
could see from the movement of the particles of earth that they were be-
ing consolidated, and that the weaving of the under surface of the door
was being completed. Next morning he could lift up the door, which
had the form of a small cup of silk, in which the earth lay. It was then
soft and pliant, but in ten days’ time it had hardened and become a very
fair specimen of a minute door of the “cork” type.1

He had watched the proceedings of young spiders when taken from
the mother’s nest in the following species: Nemesia manderstjernze, Nemesia
eleanora, Nemesia congener, and Nemesia moggridgii, the first three con-
structing wafer doors and the last a thick beveled or cork door nest. All
of these very young spiders will excavate their own tubes and bring out
pellets of earth, which closely resemble those carried out from their galler-
ies by ants.

The young brood while still in the mother’s nest will often comprise
individuals of different sizes, and, though a majority are no larger than
one-fourth ‘of an inch long, some may occasionally be found that are fully
twice as large. The little nests which they make in captivity vary accord-
ingly in size. A large number made in captivity varied in size from two
lines (one-sixth inch) to three lines (one-fourth inch) in width.

These little spiders need to be kept constantly supplied with flies,
which should be killed and placed near their nests. They are often so
greedy that they will try to drag a house fly into their tubes, for which
it is much too large, and when the door is pushed open the fly remains
sticking in the entrance of the nest, with its legs up in the air. One may
often feed these by approaching carefully without causing any vibra-
tion, pushing the fly, placed on the end of a pencil, within reach of the
spider. ?

Mr. Moggridge entertains the opinion that, as a rule, the mature trap-
door nest with its hinged lid is the result of many successive enlarge-

ments, beginning with the diminutive tube of the baby spider,

Nest De- ‘ A a ates: 5
salen: which is no bigger than a crow quill. This infantile home is
ony not abandoned, but is enlarged from time to time according to

the growth of the inhabitant, and becomes the abode of the

full grown spider.
Of course, this must require a series of months, and possibly of years,
for its accomplishment, and it is not unlikely, judging from what we know
of the prolonged life of some of the Territelarie of other families (for

' Moggridge, Trapdoor Spiders, page 119. * Trapdoor Spiders, Supplement, 245.

COCOON LIFE AND BABYHOOD. 249

example, that of Atypus piceus, as shown by Mr. IF. Enock, and that of Eury-
pelma hentzii, as I have demonstrated by several species), that the Trap-
door spiders may live for several years at least. Mr. Moggridge was
inclined to think, judging from the character of the nest and its sur-
roundings, that some which he saw had been occupied more than a year.
Evidence of enlargement of the door is not rare to meet with, though, as
a rule, the new piece is woven on to the old with such neatness as more
or less to obscure these. Examples were found in which the old and
smaller door of Nemesia meredionalis was partially attached to the large
new door which had been constructed below it.

This view is borne out by the fact that a cork trapdoor may be readily
separated into a number of layers of silk, with more or less of earth be-

’ tween every one. These layers decrease in size from without
sie and inwards, and together form a sort of saucer in which the small
Doors, Central mass of earth

lies. (See Fig. 264.)!
By moistening a series of the
cork trapdoors of Nemesia ce-
mentaria, Moggridge was able to
detach, in one of medium size,
from six to fourteen circular
patches of silk, of which the
outermost, or that which forms
the lower surface of the door,
was the largest, and the inner- Fic, 264. Successive layers in formation of a trapdoor.

(After Moggridge.)

most the smallest, thus being in-
termediate in size as in position. The last and smallest appears to be the
first door the spider ever made, and the consecutive layers mark successive
stages in the enlargement of the nest. Baron Walckenaer found more than
thirty alternate layers of silk and earth in the nest of Cteniza fodiens.?

Moggridge was confirmed in his opinion that these layers mark a suc-
cessive enlargement of the nest, by the additional fact that in very small
doors they are few or single, and a proportion is observable between the
size of the door and the number of layers of which it is composed.*

In order to test whether the doors were enlarged or not, Moggridge
measured the surface doors of seven double door nests, and one minute
cork door, on April 30th. On the 8th of October following he measured
all these nests once more and found that they all were enlarged, the aver-
age rate of increase being one and seven-tenths lines in the five and one-
half months which had elapsed. ‘The highest increase of the eight was
from five lines across to seven and one-half lines across. In none of the

1 After Moggridge, pl. xiv., and page 193. 2 Apt., Vol. I., page 228.
* Trapdoor Spiders, page 125, and table from twenty-eight specimens examined, page 150.

A, ee ee

250 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

series had the increase been less than one line in width, which was equal
to an increase of one-fourth the original width of the door.

We can scarcely venture from such limited premises to draw any precise
conclusions. But if we suppose that during the entire course the nests
increased on an average by about four lines in diameter, and assume that
the rate of growth continues the same, the nest of the infant spider, whose
surface door measures scarcely a line across, would still require four years
to attain the dimensions of some of the largest double doors, whose surface
doors measure ten lines across.1

In the nests of several females of Cteniza ariana Walck., on the island
of Niros, in the Grecian Archipelago, Mr. Erber found eggs at the bottom

of the tube attached by separate threads, and not placed in
Grecian ¢ocoons. The young spiders when hatched were turned out from

se ie the asylum of their mother’s nest, and these creatures were
Species. found, scarcely two lines long, already established in nests three

inches deep and furnished with perfect trapdoors, specimens of
which were collected.”

Costa states that the young of Nemesia meredionalis, observed by him in
the neighborhood of Naples, remain in the bottom of the maternal tube.
The mother herself stands
at the door, holding the
lid raised by means of the
four anterior feet and the
palpi, the curved extremi-
ties of which she inserts
between the rim of the
tube and the door. Some-
times the limbs do not ap-
pear, but the spider leaves

FiG. 265. The trapdoor and burrow of a young Nemesia only a chink for observa-

meredionalis. Natural size. (After Moggridge.) tion. He also observed

the fact that the young spiders make perfect little tubes entirely inde-
pendent of the maternal nest.*

XV.

Most persons who consider the above facts will cordially join with Mr.
Moggridge in thinking that these very small trapdoor nests, built as they
are by minute spiders probably not very long hatched from the

Marvels ooos, must rank among the most marvelous structures of the
kind with which we are acquainted. That so young and weak a
creature should be sable to excavate a tube in the earth many

1 Moggridge, Trapdoor Spiders, page 127.
? Verhand. der k. k. Zoologish-botanischer Verein in Wien, Vol, X VIII. (1868), page 905.
® Costa, Fauna del Regno di Napoli, Aracnidi (1861), page 14, tab. i., Figs. 1-4.

— <_< = — os |

COCOON LIFE AND BABYHOOD. , 5

times its own length, and know how to make a perfect ‘miniature of the

nests of its parents, seems to be a fa
Nature. (See Fig. 265.)
When we remember how difficult

ct which has scarcely a parallel in

a thing it is for even a trained

draughtsman to reduce by eye a complicated drawing or model to a greatly

diminished scale, we must own that
the performance of this feat by a baby
spider is so surprising as almost to
exceed belief. And yet even the most
complicated form of trapdoor nest,
namely, that of the branched double
door type, is perfectly reproduced in
miniature by these tiny architects,
with the upper door, the lower door,
the main tube, and the branched
body accurately placed.!

Mr. A. R. Wallace shows that there
is some reason to doubt whether birds,
which are so frequently said to build
by instinct, would construct the nest
proper to their kind if they were sep-
arated from the mother at the earliest
age and reared apart from her or oth-
ers of her kind. He states that birds
brought up from the egg in cages do
not build the proper specific nest; nor
do they even sing their parent’s song
without being taught.2, Whatever may
be the case with birds or other highly
organized animals, there is not the
slightest reason to doubt that, with
spiders, all forms of nests are built
in the most perfect condition by the
young as soon as they are able to do
any work at all after being hatched
from the eggs. There is no fact which
I have more frequently observed and
demonstrated than that all the inter-

, il Hg

‘ Os
We

ly,

v]
m4
—

Fic. 266. The spinningwork commons of a brood of
young Agalenas, made in confinement.

esting habits of spiders, including those which would appear to require the
greatest reasoning powers, or the exercise of faculties that in highly or-

ganized animals would imply the po

ssession of experience and cunning

' Trapdoor Spiders, page 127. See Fig. b, plate ix., page 98.
* Contributions to the Theory of Natural Selection.

252 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

skill, are exercised in their utmost plenitude by baby spiders fresh from
their cocoons. A few additional illustrations may be here grouped to-
gether, although many examples are scattered throughout these pages.

A brood of Agalena neevia hatched within a fruit jar, showed in a rather
curious way the tendency of young spiders to imitate the parental snare.

A leaf or two and several dry twigs had been placed within the
Bou ee bottle, and these formed points of support for the delicate, sheeted

spinningwork which the young Agalenas were not long in spin-
ning. Soon a hollow cylinder of silk was woven inside the jar, quite near
the glass. Now, the habit of this spider in natural site is to pierce her
sheeted snare with a circular opening, to which is attached a funnel like
tube leading downwards into the grass. The limitations of our imprisoned
spiderlings would not permit them to form such a structure; but, yielding
to the tendency of inherent instinct, they penetrated the sheeted cylinder
with circular holes, which, curiously, were placed in little groups at various
points. (See Fig. 266.) Through these openings the spiderlings came and
went, and, although they were continually adding to the texture of the
sheeted common by the draglines which they carried after
them, I never observed that the circular holes were closed.

When these little Agalenas make their exode in natural
site, and have the opportunity to pursue unobstructed their
natural tendency, they spin a little miniature of the maternal
snare, except that, as a rule, the funnel like tube is not quite
as distinctly marked, and does not form so prominent a part
Fic. 267. Ayoung of the web. At the period when the Agalena broodlings are
Agalenanevia. issuing from their cocoons they may be seen dispersed over
all manner of surrounding surfaces, upon which they have spun their
peculiar snares. They hang them between blades of grass, stretch them
across the surfaces of leaves, weave them within the angles of houses and
walls, in all kinds of crannies and corners, upon rocks, and boards, and
logs, and bits of dry wood; and I have often observed them by scores
and hundreds spun during an evening over the broken clods of a recently
spaded garden patch, or along the furrows of a plowed field. ‘These tiny
sheeted nests, when seen of an autumn morning covered with the beaded
drops of dew and glistening in the early sunlight, present a remarkably
beautiful appearance. A sketch of one of these dew covered nests is given
at Fig. 268.

M. Lucas observed on the part of certain young ‘Trapdoor spiders,
Cteniza moggridgii, a behavior somewhat resembling that of these young
Agalenas, but displaying even more decidedly the specific industrial char-
acteristics. Mr. Moggridge sent some of the Ctenizas by mail to M. Lucas,
at the Jardin des Plantes in Paris, enclosed in little, wide mouthed, cylin-
drical glass bottles. The young Trapdoors, in transitu or shortly there-
after, lined the bottles with silk and then proceeded to close them at the

COCOON LIFE AND BABYHOOD. 253

Fra, 268. Dew covered web of a young Speckled Agalena.

a = -

254 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

mouth with a door fitting accurately into a beveled lip. In the manufac-
ture of these doors fragments of moss, the only material at the spiderling’s
disposal, were used in place of earth.!

The behavior of two of the brood of Epeira sclopetaria referred to (Vol.
I., page 150), was notable as showing in its plenitude the presence of the

strongest instincts immediately after egress. A small insect, while
Sone hovering around the lamp, was snared in the straggling lines. A
Epeira. 3 : F ;

spiderling near by instantly ran to it, threw out from its wee
spinnerets jets of filaments, and completely enswathed the creature precisely
in the manner of an adult.

Another of the brood began in a few minutes after its coming to make
an orbweb. The foundations were attached to the end of one of the lines
hanging to the lamp globe by dropping a thread to the table, a distance of
eighteen inches; then a triangular frame was formed by uniting a point
of this thread to the opposite end of the upper line; within this frame a
perfect orb was spun. (See Fig. 141, page 151, Vol. I.) I
observed the whole process, laying in the radii, spinning the
notched zone, the foundation spirals, the beaded spirals; all was
complete, and an exact likeness of a perfect adult web. Neither
of these young spiders could have been more than half an
hour out of the natal tent; nor had they any previous ex-

perience, having been excluded from all spinningwork what-

peta soever; nor had they taken food of any sort. There was no

ing in a foot cannibalism within cocoon or tent before the egress of the
basket. * +s :

brood, as not a single dead individual remained; every egg

had hatched a perfect spider, and all the brood were gone, except three

living ones, who remained within the tent until the next day. Nothing

could more fully demonstrate the facts that the perfect exercise

Charac- of the function of spinning, and the full possession of the char-

ae acteristic habit of capturing prey, are innate with the spider-
Innate. ling, and dependent upon and influenced by nothing external

whatsoever. These facts, indeed, I have often demonstrated in
the various families and species by experiments quite as conclusive as the
above.

A curious deviation from the harmony which prevailed throughout this
Epeira brood was shown by the spider which made the aboye mentioned
web and another who chanced to straggle upon it. The intruder passed
along a radius toward the hub where the Orbweaver hung awaiting prey.
The latter immediately turned and seized the radius with her feet, her
little frame meanwhile showing in every part the vigor and expectancy of
her kind when a victim strikes the web.

A series of pulls and counter pulls ensued; then the two araneads ap-

1M. H. Lucas, Bull. des Seances de la Soc. Entom. de France, No. 27, page 107, 1874.

COCOON LIFE AND BABYHOOD. 255

proached. ‘There was a sharp contact, a momentary whirl of confused legs,
a retreat by the maker of the orb, who dropped from her snare quite to
the table, where she lay in the characteristic mimicry of death.
This behavior—conscious feigning or unconscious paralysis, as the
case may be—is shown by the youngest spiders when they are
touched upon their webs, or handled when off them. Like the
aeronautic habit, swinging by dropthread and foot basket (Fig. 268), snare
weaving, and enswathing the prey, it also springs into being as a_per-
fectly developed instinct.

The intruder upon the snare followed the owner a little way towards
the confines of her abandoned domain, then returned to the hub, and de-
liberately settled herself in the natural attitude, as much at home as though
she had herself spun the orb. The little exile meanwhile recovered from
her paralysis and climbed over to the standard of the lamp, where I left
her. The actions of these two spiders showed the most determined hostility,
and I have no doubt that, had either gained the mastery, the other would
have been fed upon. On the contrary, those of the brood hanging upon
the commons swung cheek by jowl without the slightest demonstration of
a cannibal propensity. I believe that the ordinary brood fraternity is
broken with the spinning of the first snare, at whose construction the
natural solitary and ferocious character of the creature, and all its wonder-
ful instincts, heretofore dominant, are vivified and spring into active exer-
cise. Possibly the little chappies are as much surprised as their human
observer to find themselves possessed of such strange powers.

Feigning
Death
Innate.

"ss

OR APT HE ae
THE AERONAUTIC OR BALLOONING HABIT.

Many accounts have been published, more or less valuable, of what
are popularly known as “flying spiders.” As the natural habits of familiar
a animals have come to be better understood, this popular phrase
Ph hr aed has yielded to the more accurate one, “ballooning spiders.” How-

ever called, the habit referred to has been and remains interest-
ing and attractive to the ordinary scientific observer. The fact that an
animal which has none of the natural provisions for progress through
the air granted to winged creatures, should, nevertheless, be able to over-
come gravity, mount into the atmosphere, and accomplish aerial jour-
neys, sometimes of immense distances, is certainly well suited to capti-
vate the imagination, awaken curiosity, and stimulate research. This
interest is quickened by the fact that the mode by which the spider
aeronaut reaches these results bears a marked likeness to the artificial
means by which man has himself solved the problem of aerial naviga-
tion. The thought that the invention of Mongolfier’s mind possesses this
striking analogue in the natural history of an inferior creature, strikes
into a profounder depth than curious wonderment, and touches the prob-
lem of a Supreme Mind over Nature.

I have studied the aeronautic habit of spiders from representatives of
the Orbweavers, Tubeweavers, Citigrades, Laterigrades, and Saltigrades,
and have not been able to note any difference in the mode of flight as
practiced by all. It is probable that the young of most spiders, and many
of the small species of all the great groups, are more or less addicted to
such mode of motion. Certainly the habit is very strongly fixed in Orb-
weavers. Epeiroid spiderlings just out of the cocoon lift themselves into
the air and sail away, precisely in the manner hereafter described. In-
deed, the infant aranead, when separated from its fellows and exposed
to a strong puff of air, seems instinctively to throw out its spinnerets and
send forth jets of silken filament, just as a human baby sets in motion
its feet and hands.

As the jets almost instantly acquire sufficient buoyancy to counter-
balance the spider’s weight, the creature becomes an aeronaut, nolens
yolens, and one can see how readily the deliberate habit of ballooning

(256)

THE AERONAUTIC OR BALLOONING HABIT. 257

could have been formed and fixed by heredity. The largest Orbweaver
that I ever saw taking flight was a partly grown Domicile spider about
the size of a marrowfat pea, say one-fourth inch long. After having
floated over a field and above a hedge row, it crossed a road and anchored
upon the top of a young tree. It never attained a height of over twenty
feet, but moved quite as fast as I could run.

Young and small spiders fly rapidly, their motion depending, of
course, upon the state of. the breeze, although they do not appear to

undertake their aerial voyage when the wind is strong. How-

aire ever, even when the air seems quite still to the observer, the

little aeronauts find a sufficient current in the height to which

they immediately ascend to bear them along with a good degree of speed.

Indeed, I have been surprised at the velocity of their progress in the
midst of what might be called a dead calm.

Spider ballooning is not limited to a special period of the year, but
may be practiced at any time. In point of fact, however, the seasons
when it most prevails are the spring or early summer, and the
autumn after the young have been hatched. The fall of the
year is more especially the season for “ flying spiders,” and Oc-
tober the month most favored. But in early November also the
balloonists are abroad, particularly during the Indian summer, or when
a series of cool days is succeeded by a warm day.

Seasons
and Con-
ditions.

II.

The following studies! were made during October, in fields adjacent
to Philadelphia and in the adjoining Delaware County. The days were
warm and bright, with a soft wind from the west, or a gentle breeze
blowing, but not steadily from any quarter. Stooping low and glancing
along the meadow, the eye caught the sheen of myriads of fine silken
filaments glistening in the sunlight. The tops of grass spires and the
bushy heads of tall weeds were netted together by innumerable threads,
and ‘from many points of the same filaments were streaming out at
various lengths into the air. Numerous small spiders, chiefly Orbweavers,
especially the young of Tetragnatha extensa, were rising from these plants
and sailing over the field.

The finest exhibition of the aeronautic flight was seen along a post
and rail fence which divided the meadow, and the description of this may

be considered as covering the like behavior among all balloon-
Eleva- —_ ists scattered over the fields. The tops of the fence posts were
pies * the favorite ascension points, and upon these clusters of young
Lycosids were gathered, sometimes eight or ten in a group. The
purpose in choosing these elevated spots is quite apparent, the currents of

" Proceedings Academy Natural Sciences, Philadelphia, 1877, page 308, sq.

ee ee ‘ ™

ee ee er ae

258 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

air being stronger there than close to the surface of the earth, and conse-
quently affording much better facility for flight. The presence of a delib-
erate and wise volition seems evident from the fact that the Lycosids are
eround spiders, and not found habitually in such positions as the above.
They had certainly mounted to the top of the fence with the settled pur-
pose of taking advantage of the stronger breeze and better “ send off”
which the superior height afforded.

At least, it was easily determined that such an advantage did ensue
from elevation. I selected some of the
lower stalks of grass from which silken
streamers were fluttering quite lazily.
Close up to the stalk or blade I saw the
spider placed back downward clasping
the thread with its claws. Sometimes a
thickened conical or flattened piece of
silk marked this end of the line. When
these grass stalks were broken off and
lifted into the air the streamers fluttered
out briskly and were soon snapped off,
carrying the young araneads away with
them. ‘These experiments showed that
the act of ascension is aided by eleva-
tion, both in these cases and in those
where the spider mounts directly from
the perch,

The young Lycosids had generally
chosen the yery tops of fence posts as
points of ascent, and fortunately this
site suited the observer’s convenience as
Aq much as the spider’s, and I could there-
FiG. 270. Attitude of aeronautie spider just fore notice with comparative ease the

ss aa methods of the miniature balloonists.

The spider’s first action was to turn its face in the direction from which
the wind was blowing. Then the abdomen was elevated to an angle of

about forty-five degrees, and at the same time the eight legs

sway were stiffened, thus pushing the body upward. In order to per-
Flight mit this movement the claws were brought in somewhat, but not

beneath the body, so that when the legs were stiffened the body
stood high above the surface. From the spinnerets at the apex of the
abdomen a single thread or ray of threads was exuded, and rapidly drawn
out by the breeze until, by reason of its delicacy, it was lost to sight.
Four, five, even six or more feet of the lines would at times be in view.
Gradually the legs were inclined in the direction of the breeze, and the
joints straightened out. The foremost pair of legs sank almost to the

THE ABRRONAUTIC OR BALLOONING HABIT. 259

level of the post; and these especially, but indeed all the legs and the
entire attitude of the creature, presented the appearance of an animal
resisting with utmost force and tension of muscles the effort of some su-
perior power to snatch it away.

Suddenly and simultaneously the eight claws were unloosened, and the
spider mounted with a sharp bound into the air, and went careering away
across the meadow, at a rate more or less rapid according to the
velocity of the wind. The utmost care was used to determine
whether in this upward bound the volition of the spider had any further
agency than the simple unclasping of the feet from the post. Owing to
the extreme difficulty of such an observation, I cannot speak with absolute
confidence, but was able to satisfy my own mind that the aeronauts always
vaulted upward and clear of the post at the moment of releasing their
hold. I can hardly be mistaken in the belief that this was so in many
cases, at least.

A similar action was frequently observed during the preliminary and
tentative movements in which the spiderlings indulged prior to the final

flight. Something was noticed among them not unlike the frol-

icsome pranks of kittens or lambs. One would rush up to an-
Spider- other, who thereupon would immediately change position, either
lings. by running or quickly vaulting to another part of the post. At

times a leap would be made quite away from the post, but the
buoyancy of the thread which had been exuded being insufficient to over-
come the weight of the animal, instead of rising into the air, the creature
returned to thé post or struck upon the adjoining rail. In these and sim-
ilar movements I was able to detect distinctly the vaulting action of the
spider, and the eye, being thus familiarized with the movement, was less
liable to be deceived in the more difficult observation of the quick spring
at the time of the aerial flight.

The posts and parts of railings adjoining were covered with threads
adhering to the wood and streaming out into the air. These were the

result in part of the feints at flight just referred to, but were
A der partly owing to another cause. The spiders, previous to flight

or vaulting, attached themselves to the post in the manner com-
mon to most of their order. The apex of the abdomen was thrust down
upon the surface, and the liquid silk at the same time exuded from the
spinnerets was thus caused ‘to adhere thereto. As the creature moved away
the thread was run out into line, and gave the spider a firm attachment.
It is a question whether this anchorage is always made previous to flight,
and whether the thread is cut immediately before the ascent. The obser-
vations made all pointed to an affirmative answer, but the matter was not
positively settled.

The attempt was made to follow some of the aeronauts beyond the
point of ascent. The difficulty in getting the minute objects in position

Vaulting.

Frolic-

-_ a

260 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

relative to the sun favorable for such observation, the motion of the
air which carried them upward, as well as the rapidity of flight, frustrated
many attempts. A position was finally taken beside one of the side posts
of the sliding “bars,” which being opened gave a point of observation with
the back to the sun, the eye upon the object, and a fair opportunity to
follow it without the delay of leaping over a high fence, which before had
been between the observer and the course of the aeronaut sailing before the
wind, Fortune favored patience, and at last a spider took flight in a line
which was a little higher than the face.

Following the aranead at a moderate run, with the eye held closely
upon it, I observed that the position of the body was soon reversed; that
is, the head was turned in the direction toward which the wind was blow-

ing, instead of the point from which it
| blew, as before the ascent. Thus the long
thread which streamed out above the aero-
naut inclined forward, and at the top was
in advance of its head. I also observed
that the legs were spread out, and that
they had been united at the feet by deli-
cate filaments of silk. The action by
which the spinningwork was accomplished
was not noticed, owing to the smallness of
the creature, the’ rapidity of its move-
ments, and the difficulty of such an excep-
tional mode of observation.’ But the fact
was noted. The reason naturally suggested
Fie. 271. Fic. 272. for it is the increased buoyancy resulting
Fro. 271, Attitude of ballooning spider just from the increased surface thus offered to
andine botreute base the Tepistemes) Of) the ait, provided, “ct
course, any reason be required beyond the
animal’s need of some sort of foothold while afloat. Mr. Emerton,! in the
course of some accurate observations of ballooning spiders, says that the
most of them while afloat hung by their spinnerets only, and drew their
legs close against their bodies, a posture which I have also sometimes ob-
served,

The spider whose behavior I am now describing was followed for a dis-
tance of eighty feet, when it gradually settled downward upon the meadow.
Before, or rather during, this ascent a small, white, flossy ball of silk was
seen accumulating at the mouth, which, with the peculiar motion of the
fore feet, palps, and mandibles, at once suggested the drawing in of a
thread. This behavior is not infrequent with spiders under other cireum-
stances; indeed, it may nearly always be observed when webs are being

—

' “lying Spiders,’ American Naturalist, 1872, pages 168-9.

THE ABRONAUTIC OR BALLOONING HABIT. 261

cleared away, and during ascent upon a dropped dragline after a spider has
thrown herself from her snare. But it became especially interesting at that
moment, for at once it suggested an act of volition on the part

age of the Lycosid, by which, in a measure at least, it might control
canoer its descent. Evidently the shortening of the overhanging thread

operated like the furling of sails upon a vessel, and decreasing the
motion of the spider increased the influence of gravity upon the body,
which thus sank toward the ground. At the same time, the diminution of
the surface of the thread above, and the increase of bulk at the mouth
(trifling as it might be), tended to increase the buoyancy of the whole, and
allowed the creature to fall. The same effect was thus produced by the
spider aeronaut, and by a strikingly analogous mode, as that which the
human aeronaut accomplishes when he con- \]
tracts the surface of his balloon by causing
the inflating gas to escape.

The manner in which the lines of spi-

ders are carried out from the spinnerets by

a current of air appears to be
How Fila- thus: As a preparatory measure,
ments are : ; -
Fmittea, 'e spinnerets are brought into

close contact, and the liquid silk
‘is emitted from the spinning tubes; the
spinnerets are then separated by a lateral
motion, which breaks up the silk into fine
filaments; on these filaments the air current
impinges, drawing them out to a length
which is regulated by the will of the ani- Fic. 278. Fia. 274.
mal; and, on the spinnerets being again ian Se ee ea hee ee
brought together, the filaments coalesce and looning spider gathering in its threads
form a compound line.1 According to Mr, *™ ‘eseen®
Emerton,? the line seems to come from the middle pair of spinnerets
only, but the posterior pair were in constant motion, folding together
over the middle ones and then spreading apart as if to help throw out
the threads,

rT.

It will here be in place, and will add to the understanding of the
reader, to insert a few field notes giving in detail the above and some
further facts as to the posture and action of spiders before and during
flight.

'“Blackwall on the Structure, Functions, and Heonomy of the Araneidea,”’ Ann. and
Mag. of Nat. Hist., Vol. XV., page 241, 1845.
* “lying Spiders,” American Naturalist, 1872, page 168.

262 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

There is no difference between the aeronautic habit of these araneads
and that of spiders in other parts of the United States. Moreover, obser-
vations of naturalists on ballooning spiders in various quarters of the globe
show that the same methods everywhere prevail. It will be further ob-
served that the notes relate chiefly to Lycosids, which appear to be univer-
sally addicted to the ballooning habit. This is probably true of all Citi-
grades. It is worthy of special notice that these ground spiders, when
seeking aeronautic flight, take pains to seek some elevated spot as a point
of departure. This is not limited to the Lycosids, for Mr. Enock speaks
of young Atypine in Eng-
land securing an easy and un-
obstructed flight in the same
way. The instinctive impulse
which urges spiderlings to
leave their resorts on the
ground and seek spots essen-
tial for favorable ascent, cer-
tainly has the appearance of
reasoning intelligence. At all
events, the younglings, by
whatever process they reach
the conclusion, do the best
thing possible to aid their
ballooning enterprise.

Example No. 1. A young
Lycosid, apparently Lycosa
scutulata Hentz, was posed
on the side of a fence post

Nee DNS aN 3 ; i ;
Ja ee; WAGS opposite the wind, face down-
Ny, E ca

wards, abdomen elevated, the
body raised by the legs. I
Fic. 275. Ballooning Lycosids ascending from a fence post, followed it after flight for two

and floating before the wind. Fe i
hundred feet; it rose as high

as thirty feet before it was lost to sight. Its flight was across a wide
meadow, and promised to be a long one. Several threads were streaming
out and up behind and before the spider.

No. 2. A Saltigrade, probably the young of Astia vittata, was posed
on the side of a fence board opposite the wind. Its legs were elevated,
thus raising up the body; the abdomen was turned well nigh straight
upward; a long thread floated out and up from the spinnerets. The
spider walked several inches upward along the rail, keeping its body in
the same stilted position, the thread meanwhile flying. Then it was off,
rather slowly, and about on a line with my face. It showed, in motion,
one small thread in front and one (or more) behind. It moved straight

THE AERONAUTIC OR BALLOONING HABIT. 263

forward for about fifty feet, and then rose suddenly upward, as though
it had passed into an ascending current of air.

No. 3. Lycosa; observed at 2 P. M. Pose and actions as No. 1. After
flight I distinctly saw one thread before and (apparently) two behind ;
the head was toward the wind. After sailing fifteen feet it rose up and
out of sight, a long stretch of meadow before it. Once, before it mounted,
it lifted up one hind foot, as though laying hold upon the stay thread.

No. 4. Lycosa; this example was followed for a distance of forty or
fifty feet; in front of it there appeared to be but one thread, a ray of
several fine diverging threads floated behind from the spinnerets. Its
back was toward the ground. Its abdomen seemed, but could not be
certainly determined, to be riding in front, 1. e., toward the direction of
the wind. The body of the spider was thus at the apex of the angle
formed by the fore and hind filaments, the free points of which were
quite far apart. The balloon struck a tree, and part of it went on, the
spider apparently staying on the tree.

No. 5. Lycosa; this specimen floated with the abdomen toward the
point of departure. Several threads ascended from it, one thread in front;
the feet were gathered together; but, apparently, the back was upward.
It crossed the highway, and a carriage just then passing interfered with
the observation.

No. 6. The head rode in front, the back was certainly toward the ground.
A fourfold streamer of threads was thrown out before mounting. At first
the spider moved off slowly, but soon climbed up the fore thread, the
“bow,” so to speak; further on it climbed up the rays of threads a dis-
tance of several inches. The balloon, when lost sight of, had at least three
separate filaments. It was followed one hundred feet before it rose out
of sight.

No. 7. Lycosa; riding back downward; it sailed sidewise part of the
time; afterward the head seemed to be directed toward the course of the
wind,

Before vaulting into the air many of the spiderlings turned their ele-
vated abdomens first to one point then to another; repeating the action

many times, as though testing the direction of the wind. The
The Etoe- hols process of aeronautic flight, as it has been described, may
ess Sum- - : : : : :
marized, be briefly given as follows: First, the spider seeks a high posi-

tion, such as the top of a bush, grass stalk, or fence post, as the
point of ascent. Second, the abdomen is elevated to as nearly a right angle
with the cephalothorax as may be. Third, a ray of threads is issued from
the spinnerets, the face being, meanwhile turned to various points; the
legs are stretched upward, thus raising the body; fourth, they gradually
incline in the direction of the breeze, the joints straighten out, the legs
sink forward and down until the first pair are almost on a level with the
surface, the whole attitude of the animal being that of one resisting some

264 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

force exerted from above. Fifth, suddenly and simultaneously the eight
claws are unloosed, and the spider mounts with a sharp bound, apparently,
and (sixth) floats off with the back downward, usually, but sometimes with
this position reversed. Seventh, at first the abdomen seems to be in ad-
vance, but generally the body is turned so that the head rides in front.
Eighth, the ray of threads is apparently grasped with the feet and floats
out in front, upon which (ninth) sometimes the spider will climb upward,
as though to adjust the centre of gravity. Meanwhile (tenth) a thread or
cluster of threads issue from the spinnerets and float out behind, leaving
the spider to ride in the angle of the two diverging rays, or, as it some-
times happens, of three, which are widely separated at the upper free
ends. Eleventh, the feet seem to be united by delicate filaments, which
would serve to increase the buoyancy of the balloon. ‘Twelfth, the spider
is now carried forward by the wind, riding for long distances in an open
space, and often borne high upward upon ascending currents. Thirteenth,
the anchorage of this miniature balloon appears at times to be within the
spider’s own volition, by the fact that it can draw in with its claws the
forward ray and gather it in a white roll within the mandibles. But
most frequently the balloonist is stopped by striking against some ele-
vated object, or by the subsidence of the breeze. A bright warm day in
October is commonly chosen for the ascent, and judging from the pres-
ence of a number of dry moults, apparently of the same species of spider
observed in flight, the animals had recently cast their skins.

IV.

The greatest height to which I have seen spiders ascend is about one
hundred and fifty feet; but, undoubtedly, they often rise much higher.
Dr. Lincecum observed the gossamer balloons of certain Texas

ioe ee species floating at an altitude of one to two thousand feet.!
inoenes Blackwall found ascending currents of air acting with such
force upon the gossamer streamers as to raise them in the atmos-

phere to a perpendicular height of at least several hundred feet.2 Dr.
Martin Lister, the earliest observer of the habit (A. D. 1670), says: “As
to the height they are able to mount, it is much beyond that of trees or
even the highest steeples in England. his last October the sky here
upon a day was very calm and serene, and I took notice that the air was
very full of webs. I forthwith mounted to the top of the highest steeple 3
in the Minster [York], and could thence discern them yet exceeding high
above me; some that fell and were entangled upon pinnacles, I took and

* “The Gossamer Spider,” American Naturalist, 1874, page 592.

* Trans. Linn. Society, Vol. XV., page 453.

* The central and two western towers are 201 feet high. Cathedrals and Abbeys of
Great Britain, Dr. Wheatley.

eek imerl a —

ee i a i i oe ad ek ee

THE AERONAUTIC OR BALLOONING HABIT. 265

found them to be lupi [Lycosids], which seldom or never enter houses,
and cannot be supposed to have taken their flight from the steeples.” I
once found a number of half grown Epeiras upon their round webs on
the topmost railing of the dome of St. Peter’s at Rome (Italy), whither
they or their maternal ancestor had doubtless been carried by the wind
from the surface of the earth.

October 25th, 1883, was a bright day following a series of cold, wet
days caused by a severe northeast storm. At noon, while crossing the
Chestnut Street Bridge, Philadelphia, I saw a great number of
aeronautic threads floating in the air, streaming from the tips of
the bridge balustrade and lodged upon the piers. One of the
threads, a long filament, was sailing slowly toward the river as a Pennsyl-
vania Railroad train dashed along the river track beneath the bridge. It
was low enough to strike the cars as they rolled by, and so was carried
on southward with its tiny voyager—another illus-
tration of how artificial habits of man tend to
the geographical distribution of life. The filaments
were long, pure white, curled or wrinkled, about
one millimetre wide or less, occasionally expanded
into thicker wads, and frequently carried attached
to them minute insects which had doubtless en-
tangled in the fibres as the threads floated in the
air, (Fig. 280.) On one thread I found three, 5. a6 Fochuclites wea
on another two small flies. The young balloonist — out aeronautic threads while
is thus provided with food upon his landing, if "™#'™s "Po" * weP-
he choose to avail himself of these chance supplies. The insects are sim-
ply entangled, as the fibre is without viscidity.

The field observations recorded above have been confirmed by numer-
ous studies made with spiderlings reared in the house, especially the young
of Epeira sclopetaria, Epeira domiciliorum, Epeira insularis, and
Agalena neevia. As the results obtained were not different from
those already given, they require but brief mention. When let
loose into the air from the finger tip, the spiderlings floated out by a sin-
gle thread, which was always and instantly first attached to the finger.
At first the head was outward, the abdomen being turned toward the hand,
from the apex of which the long superior spinnerets of the tubeweavers
diverged. Presently the little creature turned and cast out a thread be-
hind, when, if permitted, it would usually clamber up the original thread
to the finger. When this was broken off, the spider, seated midway of
the two filaments, floated off and outward, and was lost to sight. Again,
by an eddy of the air, the thread would be thrown backward and upward
and catch against the wall, upon which the little voyager would anchor.

Floating
Gossamer

Young
Spiders.

' Correspondence of John Ray, page 77. Lister to Ray, January 20th, 1670.

266 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

At other times, much to my surprise, after the thread had been quite lost
to view, the spider was supposed to be far away upon its flight, it would
descend as from the clouds, and send out its silken grapnels against the
cheek or nose. The will of the little spider seemed to have no control
over these movements, which apparently were always wholly at the mercy
of the wind. However, the manner of accomplishing aerial flight by
means of the buoyancy of a single thread, or rather of two threads united
at or near the middle, was quite in accord with the methods above de-

scribed.
V.

While the young balloonists were adventuring their flight in the fields
in the manner heretofore described, several species of small Orbweavers were
making or waiting for their
ascension in a manner so dif-
ferent that it requires espe-
cial notice. ‘These were sta-
tioned upon the small grass-
es and weeds, from which
innumerable cords of spider
silk were streaming, and up-
on which similar threads
were twisted and meshed by
the eddies of the wind and
the passing of the spider-

y) oats Sih : I i ad a Dh
t | A ir if il
aN ad

Fic. 277. Aeronautic Orbweavers preparing to ascend from
floating threads.

Bee ; j lings from point to point.
ve ( (fi ff hai The attitude of most of
“AT ane oT hie "cH these was one of expecta-
VAT RR I iW i iy fay aii #
Han) AN VN i aes Gi wie My yi tion. Only two were ob-
Hal AN \ i) a wh | i) Wl served in actual flight, and
|

one of these I assisted. ‘The
nearness to the ground and
the shelter of surrounding
herbage doubtless retarded
the process. However, this
greater deliberateness is quite
in harmony with the more

phlegmatic Orbweavers, just as the energy of the Lycosids in mounting the
fence and their haste to be off are characteristic of that group.

Varia-
tions :
Orb-
weavers.
end of the thread.

The little Orbweavers were hanging upon the lower part of the
floating strings near the point of attachment to the grass.
backs were downward and their heads outward, or toward the free
(Fig. 276.) The first, second, and fourth pairs

Their

of legs were stretched along the thread, and the third and shortest pair

THE ABRONAUTIC OR BALLOONING HABIT. 267

were held off, curved, the feet apparently united to the main thread by
taut filaments. This position, as far as could be determined, was main-

tained after flight. In some cases a series of two or three puffs or pellets
of floss were gathered around the thread between its free end and the
spiderling. ‘They were generally cone shaped, the apex being turned toward
the animal. In form they were not unlike the pellets which one used to
see gathering upon the roll of wool as it passed from the fingers of our
maternal ancestors into the whirling “flyers” of an old fashioned spin-

ning wheel. (Fig. 277.) Perhaps they may have been wrought

by a similar process, the twisting of the loose threads through
‘the action of the wind and the counteraction of the spider. The
continuation of such twisting must presently break the thread, and thus
set the occupant afloat. The greater force of the wind secured by gently
breaking a stalk and lifting it into the air soon snapped off a thread, car-
rying the little aranead away with it.

T am inclined to think that this mode of ballooning prevails, particu-
larly among Orbweavers ; that is to say, the spider, having spun out a long
thread, sometimes thickened at the attached end, lays hold upon it and
waits for the wind to pull it loose, when it is borne away and aloft. It
is even probable that the spider may cut the thread, and thus procure
her own release. This would place the moment of ascent within her own
volition, and the fact (should it be established) would add greatly to the
interest with which one must regard this variation in the aeronautic habit
of these interesting araneads.

Dr. Gideon Lincecum has put upon record a case in point.' He de-
scribes the balloon of a Texas Orbweaver, which he calls the “ Gossamer
Spider,” as follows: A lock of white gossamer five or six inches long and
two inches wide in the middle, tapering toward the ends, is attached to

a stalk, bush, or other elevated object by a thread two or three
cae inches long. At the free end or “bow,” two lines thirty or forty
Balloon, feet long are spun out, and one twenty or thirty feet long is

spun from the attached end or stern of the aerial craft. All
being ready for ascent, the voyager cuts the cable which holds the balloon,
and floats briskly upward and forward on an inclined plane, or bounds aloft
with a sharp spring that eludes one’s efforts to stop it. Lincecum’s descrip-
tion of the hammock shaped balloon and its float lines answers very well to
the above described acronautic spinningwork of Orbweavers (Fig. 277), and
I am disposed to accept as quite trustworthy the statement that the attached
end was actually severed by the spider, who thus controlled, in some
measure, the period*of her ascent.

Blackwall had already observed that occasionally spiders may be found
on gossamer webs after an ascending current of rarefied air has separated

Flossy
Balloons

1American Naturalist, 1874, page 595.

268 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

them from the objects to which they’ were attached, and has raised them
into the atmosphere. He, however, added the opinion that, “as they never
make use of them intentionally in the performance of their aero-
a ed nautic expeditions, it must always be regarded as a fortuitous
looning. ©ircumstance.”! This opinion, I think, must be abandoned, and
the conclusion reached that there are two modes of ballooning
practiced by spiders, viz.: First, ascent by means of the buoyancy of lines
issuing directly from the spinnerets, the aranead vaulting upward from
its perch; and, second, the ascent upon lines, sometimes thickened by flossy.
tufts or strands, which are first spun out and attached to fixed objects, and
afterward released by the force of the wind or cut loose by the spider.

VI.

While arranging a collection of spiders in the Academy of Natural
Sciences of Philadelphia, I discovered a number of specimens of a large
Laterigrade, the Huntsman spider, Heterapoda venatorius, from

Aerial various localities, as represented upon the accompanying tables

nea and chart. (Fig. 278.) Starting with the specimens in my private
tenn collection, the line of distribution was traced from Santa Cruz,

Virgin Isles, to Cuba, to Florida, across Central America, Yucatan,
and Mexico; across the Pacific Ocean by way of Sandwich Islands, Japan,
and Loo-Choo Islands; and thence across the continents of Asia and Africa
to Liberia. The line thus indicated extends from the extreme eastern limit
of North America to the extreme western coast of Africa, thus girdling the
globe, with the exception of 54° of longitude. This excepted area ex-
presses substantially the width of the Atlantic Ocean.

It occurred to me, when this fact became apparent, that this line of
distribution is within the belt of the North Trade Winds; and, further,
that there might be some connection between the two facts and the fact
that Laterigrade spiders, to which group this animal belongs, are among
those which are most addicted, in the earlier stages of growth, to balloon
migration. Thereupon I referred to the general course and limits of the
North Trades, which are roughly indicated in the chart (Fig. 278) by the
two upper lines of arrows, marked (at the ends) A A and BB. In the At-
lantic Ocean the North Trade Winds prevail between latitude 9° N. and
30° N.; in the Pacific between 9° N. and 26° N. We now may turn to
the chart, in which the following geographical points (shown by black
spots and figures) are represented by our spider. The specimens which
have been examined in the Academy, and my own collections, whose habi-
tats are personally known, are marked by an asterisk (*).

The species is credited to the other localities named on the authori-
ties given therewith.

' Blackwall, Spiders of Gt. Br. and Ir, Introduction, page 12.

THE AERONAUTIC OR BALLOONING HABIT, 269

A comparison of this table with the chart will at once show that the
dotted lines in the latter, which indicate the geographical belt over which
Venatoria is distributed correspond, with remarkable general exactitude,
with the belt over which the North Trades blow. It is not, therefore, an
improbable conjecture that this distribution has been accomplished by
means of those winds and the spider’s habit of aerial flight. It is, of
course, supposable that commerce, following largely the same belt, may
have originated or aided this distribution. But certain facts in the history
of the spider seem to forbid this hypothesis.

Some of the facts are: First, the early discovery of the species as al-
ready widely distributed ; second, its presence at so many different insular

points nearly or altogether contemporaneously with first visits
Not Ar- }y commercial nations; third, the existence of the species or its
ater .close allies among the fauna of the tropical interiors of conti-
bution. ents far distant from coast lines; fourth, the variations, chiefly

in color, which have been observed, and which would seem to
require for their development a longer period than that which has tran-
spired since the commencement of commercial communication with the
localities in which the variations have been wrought. While one may
not conclude with absolute certainty from these facts, they warrant the
theory that the Huntsman spider has become cosmopolitan by the action
of Nature, independent of the aid of man.

TaspLp or Distripution North or THE EQuarTor.

Locaniry. LATITUDE, LONGITUDE (GrR.). AUTHORITY.
;
GSE boggy hl sl eke eas ea ty bs SON 6° N. 163° W. *
DOPED We Sietl Cs sn an cle @ogaurans Wie eet a a! 7°- 8° N. 134° E. | L. Koch.
Sy Mlgo-ChooWeands= e608 mega ee 25°-29° N. 128° EF. *
WO MDHT chien psciainki is SR Eb AR sye! wo ketice | 80°-40° N. 130°-140° FE. | *
Di NICOMARAIBIAIARS peak ctahus fay x tactics Yas | |. 6°=10° IN, 96°- 97° FE. | Bock.
6. Tranquebar, India ........... L2P ON, 80° E. Fabricius.
TMA WSR WPAUEIOH aretha keen ate wie ok 5°- 9° N. 10° W. *
8. Senegal, Africa. 6. we ee [ale ay |. 16? W. Walckenaer.
9. Martinique, North America. ..... .| 15° N. 61° W. | *
TO atthe entie eed Aman ls Wye ice & 18° N. | 65° W. | *
Be HIMALGR sts ein suite tet os cre yo 8 se TS2SN. lee Me | Walckenaer.
UC ie lett ss Niten ta ane var te aha | 20°-28° N. | 74°85° W. | *
(EROU OMG Gers, byte eiec tina hase vec scones tn | 30° N, |) 2SS- We *
hel UCC) Oe ae) ere ctl Sane icy oy ccd | 20° N. | 82°-91° W. | *
DB; eMew1co) eee ope x) eo eae ees 20° N. O72 W. *
Afr e GalItOMN Tae era hae eeu ah ah aD aay ? 109°-117° W. | L. Koch.
*

17. Oahu, Sandwich Islands ........ 20° N. 55°-160° W.

I was so impressed by the above chain of facts, and so confident of
the inference therefrom, that I ventured to predict that corre-
sponding results would follow a comparison of specimens collect-
ed from all quarters; that is to say, they would be found to lie
within the belt of the North or South Trade Winds. The only specimens at

A Pre-
diction.

270 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

hand were those cited above, and from Zululand and Surinam. But I
was able to pursue the matter by reference to locations given by a number
of naturalists. I was aided in this by references kindly sent me by Mr.
William Holden. Some of the localities thus obtained have been named
above, and others were found to correspond with the points represented
by the specimens examined. So far my conjecture was verified.

The two lower arrow lines in the chart, C C and D D, give a general
view of the course and limits of the South Trades, which prevail in the
Atlantic Ocean between latitude 4° N. and 22° S., ond the Pacific between
latitude 4° N. and 235° S.! It is, of course, understood that these limits
are not stationary, but follow the sun, moving northward from January
to June, and southward from July to December; an oscillation which is
also indicated in the zone of distribution. They are, however, substantially
as above given, and may be compared with the following table, which
shows the southern geographical distribution of this species, according to
the authorities cited therein :—

Tape or Distripurion SoutH or THE EQuaTtor.

LocaLitry. LATITUDE. LONGITUDE (GR.). AUTHORITY.
1. Viti Levu, Fejee Islands ........!| 16°S. | 180° W. L. Koch,
9, Mew Caledouin . .. - . is g oe ome | 20°-22° s, 163°-162 BE, |
5. Sidney, Australia... 2% 5 oe eo aoe 150° BE. Bock,
4. Australia: 205 sy) 4) kde pele len ee ee Le ae 105°-115 EB. L. Koch.
PSUR DAONG ponte ise peels io ern att Peel 104° Ki. Walck.
6. Zanzibar, Africa... PAM eee 6° S. 40° EK. Gerstaecker,*
7. Southeast Equatorial Africa. |... . . | 10°-20°S, (?) 30°-50° KE. =| Blackwall.
8. Mauritisss 2. . 5 si es 8) ay ee eco ere 56° E. Walckenaer.
0; Madagnecni = bz \.) cig teen aeeine mrat 8°-26° S. 43°-50° Ee. | Vinson.
10; Zululand: 935 << we avs ane eaten BM oR RO aE 28° E. *
Mo Pernanip0o ss” «0 atm, cee We tae eae 7° 8. 37° W.
LDN IBEAC ees he ayer a Rat decdang wee he mre 37°-70° W. Simon, Walck.
13: Rio VaOeItO vices poe eee thane eke al ey 50° W. Walck
14. Surinam. .... tras ee Sebel fea ah cs ; GFN, 55° W. bal
16. Valparaiso, Chili. “seo. 0. -fs ae tty on BOD Seey | 202 We. L. Koch,
16. Tahiti, Huaheine, Society Islands... .| 18° S. 150° W. a
17. Rarotonga, Cook’s Tale Rrcomk fe! -ail| een 162° W. u;
18. Upolu, Navigator Island ..... . . .| 184°-143°S, 168°-173° W. | se
19. Tongatabu, Friendly Islands ..... .| 20°S8, 172°-176° W. | ue

This table shows a distribution corresponding with the limits of the
South Trades, with, in three cases, viz., Sidney (3), Surinam (14), and Val-
paraiso (15), a slight oscillation in ane with a fact above stated. Thus

‘The arrow line which indicates the course of the Trades is intended to give Sank the
general direction. In point of fact, however, that course, in the case of the Southern Trades,
is more nearly conterminous with the line: of distribution than here shown. The arrow
line should not run directly westward from Valparaiso, Chili (No. 15), but from a point 10°
above it, passing just south of Friendly Isles (No. 19).

* Gerstaecker speaks of species as distributed over a large part of Africa, Asia, and South
America. See Von der Decken’s Travels in East Africa, ITIL, ii., page 482.

Cure

271

THE AERONAUTIC OR BALLOONING HABIT,

ae ee WY y ae, ) ral
D meee eee Yj <_—_—_— —<—— 4+ eee ae eee <=. —«-f y Ye |
G YY

“HW
WY

—<— << —<—— >

-"IGe

ay
»

Yj UY. oo Ue o ||
oe LH Wy =< ae YX
4 ree

“S

Fic. 278. Chart to show the cireumnavigation of the globe by the Huntsman spider, in the course of the Trade Winds.

——- a a

272 AMERICAN SPIDERS AND THETR SPINNINGWORK,

was entirely fulfilled the expectation with which I entered upon the prep-
aration of these comparative tables.!

It may not be without interest, and may, perhaps, have some bearing
upon the above theory of distribution, to remark that the genus (or a
closely allied genus) to which Heterapoda venatoria belongs is probably
one of the oldest known forms of the spider fauna, 'Thorell? places the
now existing genus Heterapoda (Ocypete, Koch; Oxypete, Menge) among
those which are represented in the amber spiders. Amber probably be-
longs to the tertiary (oligocene) period, and in it numerous spiders are
found, generally well preserved. How far any supposed contiguity or
closer approach of continents now separated might have facilitated or oc-
casioned the world round distribution of our Huntsman spider, is a point
upon which geologists may more properly express an opinion.

The question, what variation of species, if any, occurs in the course
of this distribution, is of great interest. The specimens examined by me

show no variations which may not come within the range of
Variation those natural differences which obtain in many species. Most of
ete ini, tHe specimens had been so long in alcohol as to obliterate any
bution. ‘ifferences in color and markings which might have existed.

The normal color is a uniform tawny yellow, varied upon the
cephalothorax by a circular patch of blackish or blackish brown color
covering nearly two-thirds of the space; and, further, by a white or whit-
ish marginal band quite or nearly girdling the same. In some of the
specimens this circular patch seems to haye been more or less of a brown-
ish color. Gerstaecker* speaks of this species as distributed over a large
part of Africa, Asia, and South America. Specimens were examined by
him from Dafeta, Mombas, and Zanzibar. In these there was some varia-
tion in the coloration of the maxillary palpi: on the one hand, from a
light rust color to brownish red and pitch brown; on the other hand, to
a more or less sharp division or limitation of the light yellow color of
the anterior and posterior borders of the cephalothorax. There was also
a browning of the region about the eyes. But the araneologist will not
regard such differences as having any special value as specific characters.

1 When these studies were originally announced in the Philadelphia Academy, I had
no specimens from the South Pacific Islands within the same general belt; nor from the
chain of small islands between the Sandwich Islands and Asia, viz., Philadelphia, Drake, and
Massachusetts Islands, Anson and Magellan Archipelagoes; nor the Cape Verde and St.
Helena Islands, off the west coast of Africa. Nevertheless, I expressed the belief that these
had all been stations in the line of migration, the latter across the Atlantic Ocean as the
Antilles have been; the former across the Pacific, as the Sandwich Islands, Loo-Choo Island,
and Japan haye been, and as Mauritius and Madagascar Islands have been across the In-
dian Ocean. Moreover, I ventured the prediction that a more diligent search would proye that
this cosmopolitan species exists, and probably had already been collected at some of the
above points.

* European Spiders, page 231, Noy. Acta. Reg. Soe. Sci., Upsal., 1870.

* Von der Decken’s Travels in Bast Africa, III, ii., page 482. Vv

THE APRONAUTIC OR BALLOONING HABIT. 273

VII.

There seems nothing improbable in the theory of aerial circumnaviga-
tion suggested to explain the series of facts above presented. There are
not, indeed, many recorded observations of the distances to which
spiders are carried out to sea in their aeronautic flights. But,
before a strong steady wind, or in cases of storm, it is possible
that the greatest distances which appear in the tables could be overcome.
An observation of Mr. Darwin is the only recorded one to which I can
refer! At the distance of sixty miles from land, while the “ Beagle” was
sailing before a steady, light breeze, the rigging was covered with vast
numbers of small spiders with their webs, The little spider, when first
coming in contact with the rigging, was always seated upon a single
thread. While watching some that were suspended by this filament, the
slightest breath of air was found to bear them out of sight. I have ob-
served similar single threaded “balloons” sailing at considerable height

Spiders
at Sea.

Fic. 279. The Huntsman spider; a male. ©, the female’s cocoon,

above the surface of the earth, and know no reason why, with a favorable
breeze, they might not have been carried hundreds of miles. That they
were carried at least sixty miles, as Mr. Darwin’s testimony shows, and
that before a light breeze, gives great probability to such a conjecture. It
is to be noted, moreover, that the spiders arrested by the “Beagle’s” rig-
ging were evidently moving on when so stopped, and some of them, when
arrested, soon resumed their flight across the main,

I am able to add a valuable observation in the same line as that of
Dr. Darwin’s. The late Capt. George H. Dodge, of the American Line
steamer ‘‘ Pennsylvania,” informed me, during a voyage across the Atlantic
in the winter of ’81-2, that he had found the masts and rigging of his

' Voyage of the Beagle, Vol. III., page 187.

— ie |

oe Pe 2 Se ee ee ey eee ee

274 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

vessel covered in the same way with innumerable webs of spiders, while
sailing during the month of March along the eastern coast of South
America. His ship was more than two hundred miles from land and
about four hundred miles south of the equator. The wind at the time, ac-
cording to his recollection, was blowing from the westward ; that is, from
the continent. Captain Dodge, at my request, communicated the facts
in writing, the incident haying been impressed upon his memory by the
strangeness of seeing such creatures so far out at sea. “The spiders
seemed like little elongated balls, with a sort of umbrella canopy above
them. They settled upon the sails and rigging, and, finally, disappeared
as they came.’”’!
The purpose of such a remarkable habit as these facts exhibit is,
ie doubtless, to secure the distribution of species throughout wide
means? regions. The buoyant filaments of spider gossamer serve the
1 tiny arachnid the same good office that is rendered the dandelion
and thistle seed by the starry rays of down surrounding them.

Vill.

The ballooning habit of spiders gives a complete explanation of a nat-
ural phenomenon which has attracted the attention of men from an early
period, and which has been variously
alluded to in prose and poetical writ-
ings, viz., Showers of Gossamer.

One who walks the open fields in
the latter part of September or in the
soft bright days of October, which
is the most delightful period of our
Fic, 280. A flocculent thread of gossamer, with American year, will notice great quan-

small flies entangled. aha ; 5 ‘ as, ;

tities of spider silk trailing and float-
ing from the stalks of weeds and grasses, and indeed from all elevated
objects. In the early morning, when the dew deposited upon these fila-
ments betrays their presence, one will be surprised at the vast
amount visible. Further on in the day he will observe quantities
of this threaded spinningwork sailing through the air. (Fig. 280.)
A great excess of these floating tufts and filaments constitutes what is com-
monly known as a gossamer shower. Doubtless Pliny alluded to such a
phenomenon in the statement which he makes? that “in the year that
L. Paulus and C. Marcellus were consuls it rained wool about the castle
Carissa, near to which, a year after, T. Annius Milo was slain.”

Gossamer
Showers.

1 Captain Dodge adds, very significantly: “You know that it is not unusual for birds to
be blown out to sea. How much easier for a spider, provided he had the means to keep
himself suspended in the air!”

* Natural History, I1., 54. Holland’s translation, page 27.

—"

THE AERONAUTIC OR BALLOONING HABIT. 275

In later days, among our English ancestors, an explanation of this
phenomenon even stranger than Pliny’s prevailed and found expression
through some of the English bards. For example, Spenser writes :-—

“More subtle web Arachne cannot spin;
Nor the fine nets, which oft we woyen see,
Of scorched dew, do not in th’ ayre more lightly flee.”

Still later Thomson in his “Seasons” utters the same idea :—

“How still the breeze! save what the filmy threads
Of dew evaporate brushes from the plain.’?

We have, however, passed beyond the period when so simple a natural
phenomenon could be accounted for on such an impossible theory as that
of autumnal dews scorched by the sun.

I have never been so fortunate as to observe anything that could be
called a “shower” of gossamer, although I have seen quantities of the
material afloat in the air or fluttering from the foliage. I will therefore
quote from others a description of the phenomenon. Mr. Kirby describes
the gossamer observed by him early in the morning as spread over stub-
bles and fallows, sometimes so thickly as to make them appear as if coy-
ered with a gauzy carpet, or rather overflown by a sea of gauze, presenting,
when studded with dewdrops, a most enchanting spectacle.*

Rey. Gilbert White, whose “Natural History of Selborne” has been so
long and deservedly popular, describes such an incident as occurring in
England on September 21st, 1741. At daybreak he found the stubble
and clover grounds matted all over with a thick coat of cobwebs, in the
meshes of which a heavy dew hung so plentifully that the whole face of
the country seemed covered with two or three fishing set-nets drawn one
over another. The dogs were so blinded by this deposit that they could
not hunt, but lay down and scraped the encumbrances from their faces
with their fore feet. ‘As the morning advanced,” writes the author, “the
sun became bright and warm, and the day turned out one of those most
lovely ones which no season but autumn produces, cloudless, calm, serene,
and worthy of the south of France itself. About nine, an appearance very

.., unusual began to demand our attention—a shower of cobwebs
ee falling from very elevated regions, and continuing without any
Shower, imterruption, till the close of the day. These webs were not

single filmy threads, floating in the air in all directions, but per-
fect flakes or rags; some near an inch broad, and five or six long, which
fell with a degree of velocity that showed they were considerably heayier
than the atmosphere.

1 Faerie Queene, B. 2, XII., s. 77. ? Seasons: Summer, I., 1209.
* Kirby and Spence, Introduction to Entomology, Vol. IT., 841, Letter XXIII.

. eel ee

276 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

“On every side, as the observer turned his eyes, he might behold a
continual succession of fresh flakes falling into his sight, and twinkling
like stars, as they turned their sides towards the sun.”

This shower extended over at least eight miles of territory, for Mr.
White received an account from a trustworthy gentleman living that dis-

tance from his house, corroborating his own observation. This
Extent gentleman met the gossamer shower while he was riding abroad,
ech and, concluding that he could escape it by mounting a hill
‘above his fields, which was three hundred feet in height, rode
to that point. But, to his astonishment, when reaching this lofty spot,
he found webs apparently still stretched as far above him as before, still
descending into sight in a constant succession and twinkling in the sun
as they fell. Neither before nor after, says Mr, White, was any such a
fall observed; but on this day the flakes hung in the trees and hedges
so thick that a diligent person sent out might have gathered baskets
full.!

Another account, quite as noteworthy as the above, was reported in
the ‘London Times” on October 9th, 1826, which I quote from Mr.
Frank Cowan’s interesting and valuable “Curious Facts.”? “On Sunday,
October Ist, 1826, a phenomenon of rare occurrence in the neighborhood

of Liverpool was observed in that vicinage, and for many miles
Another istant, especially at Wigan. The fields and roads were covered
English : . : ; :
Shovwvei: with a light filmy substance, which, by many persons, was mis-

taken for cotton; although they might haye been convinced of
their error, as staple cotton does not exceed a few inches in length, while
the filaments seen in such incredible quantities extended as many yards.
In walking in the fields the shoes were completely covered with it, and
its floating fibres came in contact with one’s face in all directions. Every
tree, lamp post, or other projecting body had arrested a portion of it.
It profusely descended at Wigan like a sheet, and in such quantities as
to affect the appearance of the atmosphere. On examination it was found
to contain small flies, some of which were so diminutive as to require a
magnifying glass to render them perceptible. ‘The substance so abun-
dant in quantity was supposed by the writer who described the phenom-
enon to be the gossamer of the garden or field spider, often met in fine
weather in the country, and of which, according to Buffon, it would take
663,552 spiders to produce a single pound,”

An English writer? describes what he calls a “Visitation of Spiders,”
which occurred at Newcastle-on-Tyne. ‘Three miles of iron railing in the
writer’s neighborhood was covered with the little creatures. They were
equally numerous about one mile north of Newcastle, and, in fact, covered

1 Natural History of Selborne, Letter LXV.
* Curious Facts in the History of Insects, including Spiders and Scorpions.
5 “Science Gossip,” December Ist, 1865, page 282.

= Ras) oh py od i

THE AERONAUTIC OR BALLOONING HABIT. 277

the entire town. A gentleman from Hexham, a town twenty miles from
Newcastle, reported that they were abundant there also. The spiders were
unknown up to that time, Mr. Blackwall not having described them in
his elaborate work on the “Spiders of Great Britain and Ireland,” only
having noticed them in the “Annals of Natural History” in 1863, previous
to which time they had not been observed in England. No one had ob-
served this spider in the neighborhood of Newcastle up to the time of
their appearance, and they disappeared as suddenly as they came. Ac-
cording to Mr. Blackwall, the spider is an aeronautic species, Neriene
dentipalpis.
One of the most temperate descriptions of a gossamer shower I quote
from Mr. Blackwall. A little before noon on an October day which was
remarkably calm and sunny, with the thermometer in the shade

Black- ranging from fifty-five to sixty-four degrees, Mr. Blackwall ob-
bidet aed served that the fields and hedges in the neighborhood of Man-

chester, England, were covered over with a profusion of fine, glossy
lines, intersecting one another at every angle and forming a confused kind
of network. So extremely numerous were these slender filaments that ‘in
walking across a small pasture his feet and ankles were thickly coated
with them. It was evident, however, notwithstanding their great abun-
dance, that they must have been produced in a very short space of time, for
early in the morning they had not attracted his notice.

A circumstance so extraordinary could not fail to excite the curiosity
of so keen an observer. But what more particularly arrested his attention
was the ascent of an amazing quantity of webs of irregular and compli-
cated structure, resembling raveled silk of the finest quality and clearest
white. They were of various shapes and dimensions, some of the longest
measuring upwards of five feet in length and several inches in breadth in
the widest part, while others were almost as broad as long, presenting an
area of a few square inches only. Mr. Blackwall quickly perceived that
these gossamer threads were not formed in the air, as was gen-
erally supposed at that time (1826) even among naturalists, but
at the earth’s surface. The lines of which they were composed
being brought into contact by the mechanical action of gentle airs, adhered
together, until by continual additions they were accumulated into flakes or
masses of considerable magnitude. On these masses of spinningwork the
ascending current, occasioned by the rarefaction of the air contiguous to
the heated ground, acted with so much force as to separate them from
the objects to which they were attached, raising them in the atmosphere
to a perpendicular height of at least several hundred feet.

About midday Mr. Blackwall collected a number of these webs as they
arose, and again in the afternoon, when the upturned current had ceased
to support them and they were falling. Scarcely one in twenty contained
a spider, though on minute inspection he found small winged insects,

Forma-
tion.

a _—

278 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

chiefly aphides, entangled in most of them. This flight of gossamer ap-
pears to have been quite general throughout Great Britain, as it was no-
ticed in England, Wales, and even in Ireland.!

Mr. Blackwall is undoubtedly correct in the suggestion which he makes
as to the origin of gossamer showers. My own observations, at least, are

¥ precisely in the direction of his conclusion. As has already been
eed i said, the aerial excursions of spiders in the United States usually

‘ occur in the soft, balmy days of early autumn, during the months
of September and October, although they occur in a less degree during the
first warm days of June. The reasons for this are manifest. In the first
place the conditions of the atmosphere are favorable. ‘The balmy weather
invites the spiders to issue from their hiding places and attempt aerial
flight. The wind is not high enough to disturb their excursions, and yet
the temperature is sufficiently high to cause ascending currents of air.
Were the weather cold or rainy spiders would not venture forth. Were
the wind high its violence would greatly interfere with their excursions.
Were the air perfectly still it would be impossible for them to mount
above the earth. But the conditions being favorable, as they generally are
in the haleyon days of our American autumn, immense numbers of spi-
ders, but particularly the young, may be found upon all manner of elevat-
ed objects—blades of grass, weeds, bushes, fences, and what not—essaying
an aeronautic flight.

In many, and I would venture to say in the great majority of cases,
before a successful ascent is accomplished many unsuccessful attempts are
made. A spider will assume the proper position and spin out
a long thread. Jor various reasons, which we are not able to
explain, it fails to mount aloft. The thread floats in the air
until it is whipped off by the breeze. One, two, or a dozen attempts of
this sort produce as many floating filaments. These while waving to and
fro in the eddying air are sometimes tangled together before they are loos-
ened. Others, again, are united in the air after release. If now we think
of the unnumbered myriads of young spiders who are abroad at this sea-
son, all moved by the common imptise to fly away from their present
site, and all making the unsuccessful efforts described, we can imagine
the enormous quantity of loose filaments of gossamer threads which would
thus be set afloat within a short period of time.

These, no doubt, ascend to a certain height, at which they become
more or less united into a loose, flocculent mass, and from which, in the
cool of the evening, or on the cessation of the air currents, they slowly
descend, and add to the quantity already fluttering from all points of
the herbage on the surface.

Trial
Threads.

“Researches in Zoology,” by John Blackwall, F.L.8., second edition, London, 1873,
page 258, sq.

———————— ee hl ee

.
THE ABRONAUTIC OR BALLOONING HABIT. 279

This is a natural, and undoubtedly is the true, explanation of gossamer
showers. ‘The theories which have attributed them to electrical phenomena,
or to the shooting out of threads from the spinnerets by the physical
power of spiders before their ascent, must be dismissed as having no foun-
dation in fact. They are really no more worthy of credit than the popu-
lar superstition that these fleeing cobwebs are—

“Caused by the autumnal sun,
That boils the dew that on the earth doth lie.”

The French naturalist Mr. Virey made certain observations and ex-
periments which led him to conclude that spiders ‘swim in the air” by
approximating their limbs and striking the air as birds or insects do their
wings. Moving the feet with incredible agility, they are able by means
of the vibration to propel themselves through the atmosphere.' In this
bold but fanciful conjecture, as Blackwall properly terms it, Mr. Virey was
anticipated by Dr, Lister. ‘Certainly this is a rope dancer,” he writes,
“and itself effects its ascent and sailing. For, by means of its legs, closely
applied to each other, it balances itself, as it were, and promotes and di-
rects its course no otherwise than as if Nature had furnished it with
wings or oars,” ?

Notwithstanding the importance which such names give to the suppo-
sition, it is thoroughly unworthy of belief. The only movement which I
have ever perceived on the part of spiders is a momentary adjustment of
their bodies, so as to swing them between the two floating rays of threads
that constitute their balloon; and, also, to spin the little foot basket or
support for their feet, which I have heretofore described. Otherwise they
appear to remain perfectly quiet until they reach the ground and escape
from their aeronautic threads.

It is hardly worth while to more than mention the theory of Murray
that the ballooning ascents of spiders are caused by electricity.? The the-
ory was much mooted at one time, and had some worthy names
to endorse it. It is, of course, not impossible that a material
composed of silk, as is the spinningwork of spiders, may be in-
fluenced more or less, and in one way or another, by electricity. But as the
result of careful, long continued, and wide observation and study I have
no hesitation in saying that electricity has nothing (or next to nothing)
to do with the ballooning of spiders, and that the ascending and moving
currents of air are entirely responsible for aeronautic phenomena.

There appears to be a special tendency on the part of certain species
to undertake aeronautic flight, and certain species appear to be destitute of

Elec-
tricity.

1 Bulletin des Sciences Naturelles, October, 1829, page 183. * De Araneis, page 85.
* John Murray on the Aerial Spider, London Magazine of Natural History, November,
1828, pages 320, 324.

280 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the power, or at least the wish, for such excursions. Among the latter
Blackwall! ascertained that Tegenaria civilis and Ciniflo atrox are to be
reckoned; among the former, the most skillful balloonists ob-
Balloon- served by him were Thomisus cristatus and Lycosa saccata. The
prehl largest individuals of the first named species seen to take aerial
; journeys measured one-sixth inch between the extreme points of
the head and abdomen, one-tenth inch across the broadest part of the ab-
domen, and weighed about a quarter of a grain. The largest individ-
uals of Lycosa saccata seen floating in the air were of similar weight and
dimensions.

Most readers of general and theological literature possess some knowl-
edge of the position held by Dr. Jonathan Edwards as a philosopher. His
work on “The Will” still ranks as one of the greatest books
Dr. Jona- written by an American; but the fact that Jonathan Edwards is
than Ed- : . :
wards, Cntitled to a place among the pioneers of natural history has
heretofore been limited to a small number of persons specially
interested in science. ‘To that little band it gives particular pleasure to
note the recognition of that fact which the last few months have brought.
In the first volume of this work I have already alluded to the observations
of Master Jonathan Edwards upon spiders, and have credited him with an-
ticipating by at least one hundred and sixty years some of the most inter-
esting observations which I have made and published under the supposition
that they were original with myself.?

It is proper at this point to call attention to some facts in the aero-
nautic habits of spiders which this lad made known. Dr. Sereno E.
Dwight, the editor of the “Life and Works of Jonathan Edwards,” appears
to have been the first to publish a letter written by him, when a boy of
twelve or thirteen years old, to an English correspondent of his father’s,
in which letter he describes what he has seen of the habits of “ flying
spiders.” The scientific world was made acquainted with the matter as
early as 1832 by the editor of “Silliman’s Journal,”® who published in full
the above named letter as printed by Dr. Dwight.

The January number of the “Andover Review” takes up this subject
anew, and in a valuable paper’ Professor Smyth covers the whole ground
of Edwards’ studies, and permits us to look into the operations
of the young mind while pursuing his remarkable observations
and experiments. An unpublished manuscript is therein edited,
which appears to have been the original record of the boy’s studies, from

Boy Nat-
uralist.

1 Researches in Zoology, page 275. 2 Volume I., page 69.

5’ American Journal of Sciences and Arts, Vol. XXII., 1832, pages 112, 113.

*“The Flying Spider: Observations by Jonathan Edwards when a Boy,” Andoyer Re-
view, 1890, Prof. Egbert C. Smyth.

a a Ol ee

THE AERONAUTIC OR BALLOONING HABIT, 281

which record the letter of the English correspondent was probably con-
structed.

Young Edwards appears to have made a rude division of various tribes
of spiders, which, as far as it goes, is accurate, at least sufficiently so for
all popular purposes. In a general way this lad as early as A. D. 1716
had hit upon the foundation principle of classification of the distinguished
naturalist Latreille, who, just a century later, divided spiders into seven
groups, based upon those very habits which young Edwards notes, al-
though, of course, with more careful characterization.'

Edwards had found that on a dewy morning towards the end of Au-
gust or beginning of September one has the best opportunity to study
field spider webs. He had further discovered that spider webs which are
ordinarily unobserved may readily be brought into view by putting one’s
self into such position that the rays of the sun shall fall
upon them against some opaque body.

Once more, the boy naturalist had discovered that the aero-
nautic habit of spiders is closely associated with those bridge
lines which are continually observed in summer stretched from
tree to tree across roads, between fences, and in like position.

Again, he appears to have discovered that the spider, while
engaged in casting out these bridge lines, often swings itself

_.* in a little basket of lines held between the bunched
Swinging feet, I have particularly alluded to this in Volume
Foot .

Basket. I., page 69, when speaking of the use of what I have

called the swinging foot basket, a
habit of which I had supposed that I was the
original discoverer. The drawing in Volume L.,
Fig. 65, was taken from what I supposed to be
an accurate fac simile in “Silliman’s Journal;”
but, in point of fact, Edwards’ drawings, as
given by Professor Smyth, are far more accu-
rate than those, particularly in the outline of
the spider’s body and legs, and I therefore re-

FiG. 281. Edwards’ Ballooning Spiders. 1,
dropping from twig; 2, swinging from
produce them here, after the drawings in the line: 3, sending out threads, bc; 4, a,

% abandoned thread ; ¢ b, spider in fli is
“Andover Review.” iia Sieaaee

Again, Edwards defined correctly the manner in which the spider’s
thread is formed. He could make no studies of the interior structure
of the animal. It was reserved for the age of the microscope
_ to do this, but this boy of thirteen years old reasoned that the
spinning stuff must be contained in liquid form within certain
appropriate organs in the abdomen, from which it is expressed, escaping
from the spinnerets as a liquid, and immediately hardening by contact

A Pre-
diction.

1 See Cuvier’s “Le Regne Animal,” edition 1817, Paris.

282 AMERICAN SPIDERS AND THEIR SPINNINGWORK.,

with the air. I quote his language: ‘Seeing that the web while it is
in the Spider, is a certain cloudy liquor with which that Great bottle
tail of theirs is filld which immediately upon its being Exposed to the
Air turns to A Dry substance, and Exceedingly Rarifies and extends it
self”. . . . “Now if it be a liquor it is hard to Conceive how they
should let out a fine Even thread without Expelling a little Drop at the
End of it but none such Can be Discerned, but there is no need of this.”

Young Edwards also perceived that the spider had no direction of its
frail aerial vessel after it had once embarked, but was compelled to go
at the will of the wind, and to disembark and settle wherever its balloon
might find an entanglement. He correctly discerned and explained the
theory of equilibrium by which the spider navigates the air. This is
his explanation: “If there be not web more than enough Just to Coun-
terbalance the gravity of the Spider the spider together with the web
will hang in equilibrio neither ascending nor Descending otherwise than
as the air moves but if there is so much web that its Greater Rarity Shall
more than Equal the Greater Density they will ascend till the Air is so
thin that the Spider and web together are Just of an equal weight with
so much air.” This statement substantially expresses the opinion of all
students at the present day.!

This review of the studies in natural history of the boy Edwards will
suffice to justify the language used nearly sixty years ago by Prof. Ben-

jamin Silliman, one of the most eminent of America’s men of
Prof. science: “The observations recorded by him present a very curi-
Silliman’s ; ‘ : ; ; ;
Tribute, US and interesting proof of philosophic attention in a boy of
twelve years, and evince that the rudiments of his great mind

were even at that immature age more than beginning to be developed.”
Even with the more perfect light of the present there will be found few
to question the further words of the same distinguished authority, that
“had he devoted himself to physical science, he might have added another
Newton to the extraordinary age in which he commenced his career; for
his star was just rising as Newton’s was going down,’”?

"See a paper by the author on “Jonathan Edwards as a Naturalist,” in Presbyterian and
Reformed Review, July, 1890.
* American Journal of Sciences and Arts, 1832, page 110.

PART IV.—THE SENSES OF SPIDERS AND THEIR
RELATIONS TO HABIT.

On APYTE RX.
THE SENSES OF SPIDERS, AND THEIR ORGANS.
iL

A SPIDER’s eye is a globose object or capsule, lined internally with pig-
ment and having the outer surface transparent and conyex, constituting
the cornea or corneal lens. Behind the lens is the optic cham-
ber, filled with a semiliquid and lined on the hinder part with
the retina, which receives and reverses any image, as in the
-human eye,

The eye of a spider corresponds with the ocellus of insects, and speak-
ing generally, the ocellus may be regarded as consisting, first, of a lens,
forming part of ,the general body covering; second, of a layer of trans-
parent cells; third, of a retina or second layer of deeper lying cells, each
of which bears a rod in front, while their inner ends pass into the fila-
ments of the optic nerve; fourth, of the pigment. From the convexity of
the lens it would have a short focus, and the comparatively small number
of rods would give but an imperfect image, except of very near objects.
Though these eyes agree so far with ours, there is an essential difference
between them. It will at once be seen that the pigment is differently
placed, being in front of the rods, while in the vertebrate eye it is behind
them. Again, the position of the rods themselves is reversed in the two
cases, !

In details, the structure of fully formed ocelli presents many differences;
and it is remarkable that in some species this is the case even with the
eyes of the same individual, as in those of the well known En-
glish Orbweaver, Epeira diademata. (Fig. 282). The figure,
which is taken from the admirable work of Grenacher,? represents a section
through the front (A) and hinder (B) dorsal eyes. In this case the eye B
would receive more light and the image therefore would be brighter, but on
the other hand the image would be pictured in greater detail by the eye, A.

Spiders’
Hyes.

Ocellus.

1 Sir John Lubbock, “On the Senses, Instincts, and Intelligence of Animals,’ page 147.

* H. Grenacher, “Untersuchungen tiber das Sehorgan der Arthropoden,” Goéttingen, 1879.
I have made the cut from Lubbock’s book. )
(283

eS he Oe .

284 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Fig. 288 is a drawing of a section through an anterior median eye of
Agalena nevia eight days after hatching multiplied about three hundred
and fifty times. The retinal portion has not reached its full development, but
on the whole in this stage the essential features of the eye are established. 1

It will be seen that in this species, according to Locy, the eye consists,
first, of the lens, which a few days after hatching assumes the form of the
cuticular lens of the adult; second, the vitreous body, which is the
magnified portion of the hypodermis, with which it has never
ceased to be continuous; third, of the hypodermis; and, finally, of
the retina, consisting of the first or inverted layer of optic invagination,
and the second or non inyerted layer of optic invagination.

Speaking roughly, the eyes of spiders and ocelli of insects may be said
to see as our eyes do; that is to say, the lens throws on the retina an
image, which is per-
ceived by the fine
terminations of the

optic nerve.

Or : x From the nature
TIAN of the external in-

ee : ' tegument, the eyes of
spiders would seem
to be fixed in one di-
rection. Yet micro-
scopic examination
of them when alive
Fic. 282, Long section through the front (A) and hinder (B) dorsal eyes of appeared to satisty
Epeira diademata. (After Grenacher.) A, anterior eye; B, posterior eye; Mr. Campbell? that
Hp, hypoderm ; Ct, cuticle; ct, boundary membrane; K, nuclei of the cells spiders not only have

of the retina; M, muscular fibres; M, M', cross sections of ditto; St, rods;
Pg, P*, pigment cells; L, lens; Gk, vitreous body; Kt, crystalline cones; 4) adjusting power

Rt, retina; Nop, optic nerve.

over the lenses, but
that they also can move the eye itself within the cavity covered by the
transparent cuticle. This appeared to the author to be the only way to
account for the frequent changes of color, as well as of the form and posi-
tion of the color, which take place in spiders’ eyes, and which resemble
that of a moving liquid globule. This opinion seems to me unfounded.

Il.

How far Orbweavers may be guided by sight in making their webs,
and how far by touch, is an open question. The organs of sight do not
seem to be as highly organized in these and other Sedentary spiders as
in the Wanderers.

Structure
of Eyes.

}

Uf ww

* William A. Locy, Observations on the Development of Agalena nevia, Bulletin Mus.
Compar, Zool. Hary. Coll., Vol. XII., No. 3, plate x., Fig. 69,
? Observations on Spiders, page 42.

— t= *r =

THE SENSES AND THEIR ORGANS. 285

Professor Wilder, when studying the habits of Nephila plumipes,! came
to the conclusion that spiders of that species cannot see anything at all,
whether near or remote. I do not accept the conclusion, but
give some facts on which it was based. The spiders paid no at-
tention to an object put close to them, nor to the quiet movements
of any one about them. An individual would often rush by an insect en-
tangled in her net, if it chanced to cease its struggles before she had accu-
rately determined its position; she would then slowly return to the centre
of the web, and wait until another vibration indicated the whereabouts
of the insect. A fly offered upon the point of a needle would not be no-
ticed until it began to buzz, when it would be seized at once. Nephila,
however, always prefers the light, and constructs her large orbs where the
sun can reach them. The young mani-
fest the same instinct, and in confine-
ment seek the sunny side of a glass
vessel.

In order to test the ability of the
Furrow spider to work upon her web

without the aid of daylight,

Quality
of Vision.

Orbs —_ I secluded one within a large
‘Made in Gel] with a sliding glass door
the Dark. sree Beg

She soon spun an _ orbweb
across the cell as close to the glass door
and as far toward the light as could well fy. 293. sagittal section through an anterior
be. Fortunately not a line of the snare ™odian eye, eight days after hatching; the

- retinal portion has not yet reached its full
was attached ‘to the glass itself, so that development. x about 350. Ins, lens; vit,

Ivéould! draw it back amd: forth at wi eee ee cea tener aP enkle tare.
I next cut away the lower foundation ination; rtn”’, second (non inverted) layer.
line, broke up the entire lower part of © “%* "°°
the web, and the box was then completely darkened. Next day, when
the covers and screens were removed, the web was found thoroughly
mended, every part being so neatly and accurately repaired that it was
scarcely possible to determine which was the patchwork and which the
original snare. Even if we admit that some particle of light may have
entered the cell, the sense of touch in this case must have been the chief
reliance,
In other experiments the spider’s sight, in so far as that sense can be
apprehended by human experience, could have had no part in
Cocoon- directing her work. I have repeatedly confined female Orbweay-
ihe mae ers, Epeira strix, sclopetaria, insularis, domicilioram, and triara-
nea, Acrosoma rugosa, and others, in paper boxes absolutely im-
pervious to light, and opening them the next day have found eggs deposited

1 Proceed. Boston Soc, Nat. Hist., Vol. X., page 208.

a 0

286 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

upon the inside, sealed, covered, and swathed within the characteristic co-
coon, and all done as accurately as though the creature had wrought under
the most satisfactory conditions. Readers who have followed the methods
of cocoon weaying as detailed in the preceding chapters will be able to
appreciate the acuteness and accuracy of that sense of touch, and what-
ever other faculty may be associated therewith, by which such an in-
dustry was wrought, and share the wonder that it could have been
done in total darkness. Like facts are true of other tribes than Orbweay-
ers, particularly Tubeweavers and Saltigrades, with which I have experi-
mented.

The fact that spiders are able to spin their cocoons not only in the
dark, but without apparent organs of vision, is demonstrated by the case
of the well known caye spider, Anthrobia mammouthia. Professor Pack-
ard found this species in Mammoth Cave, and was fortunate enough to
collect a cocoon belonging to it. This, of course, shows that the delicate
work required in constructing this object can be wrought by touch alone.
So also the cavern Linyphians, although having eyes, dwell in total dark-
ness and weave snares and cocoons.

I have often found the peculiar tubular nest of the Saltigrades spun
within boxes in which they had been captured, These spiders have, per-

i See haps, the most perfect organs of vision of any of the order. Yet
eae they are able to do quite as good weaving in the dark as in the
: light. No one who has watched them stalking prey during the
day could well fail to conclude that they are guided by a tolerably accu-
rate sense of sight. Their rapid and marked change of manner when
prey is “sighted,” the mode of approach, like the action of a cat creep-
ing upon a bird, the peculiar behavior displayed when the final spring
is made, are not to be accounted for on any theory other than a keen
sense of sight. ‘So also with Citigrades. I have seen young Dolomedes
sexpunctatus leap from the side of a box and catch a fly “on the wing,”
and return to its perch by the rebound of its dragline. Such an act not
only shows ability to see, but also some faculty to estimate distance, unless
we suppose it to have been a chance shot.

In Laterigrades similar action may be observed. Like Saltigrades, they
are arboreal in their habits, and crouch for prey and steal upon it. One
of our largest indigenous Laterigrades, the Huntsman spider (Heterapoda
venatoria), received from Florida and kept in captivity, permitted a large
fly placed in her cell to run between the legs, fly into the face, alight on
the back, without any attempt to capture it. In the course of time, how-
ever, the fly lit on the side of the box a short distance in advance of the
Huntsman. She perceived it, crouched, slowly moved her limbs, stealthily
and by almost imperceptible advances approached, then swiftly shot forth
her claws and secured her victim. The behavior was quite similar to that
shown under kindred conditions by animals with well organized sight.

THE SENSES AND THEIR ORGANS. 287

Rey. Mr. Pickard-Cambridge! records that he has more than once seen
an English Orbweaver, Meta segmentata, drop from her web upon an insect
which it had espied on the ground a little way below it, and ascend again
with its prize by means of the line drawn from its spinnerets in the de-
scent. This is certainly a remarkable degree of keensightedness for an
Orbweaver, one, by the way, that habitually affects a shadowed habitation.

Many species of Orbweavers are found upon their snares during day-
time in normal posture for capturing prey. They seem to have little
cessation of activity in the hours of light, nor does there appear
to be special increase of activity during evening. Other species,
as the Furrow spider, quite habitually exclude themselves from
the orb during the day and hide in the vicinity until the approach of
evening, when they come forth and take position at the hub of the orb.
With most species this is the time when such work will be done. As the
light begins to diminish over the landscape an increased activity may be
observed throughout a large part of spider world, and everywhere indi-
viduals may be seen flying tentative filaments, restlessly pioneering the
neighboring shrubbery, running foundation lines, weaving webs, swathing,
trussing, eating insects. Something of the same sort may be seen in the
early morning, when snares broken by the night’s work are renewed or
replaced.

Night
Habits.

Ta:

I am not prepared, as yet, to say whether a comparison of the species
which show morning activity with those which display activity at night
would justify a separation into nocturnal and diurnal spiders. Certainly
the line would not be a rigid one; but there is some ground to suppose
that there are occasional tendencies to this side or that, more or less de-
cided. There is probably a difference in this respect among the several
eyes of any one spider; some are undoubtedly organized to re-
ceive more light than others, while some receive the image pict-
ured in greater detail. (Fig. 282.) We may therefore attribute
different powers of vision to the different eyes. The eyes of spiders vary
both in shape and color in the same individual. Some are pearly white;
others yellowish, amber, dark gray, or black. Some Saltigrades have eyes
that shine like precious stones, reflecting various brilliant hues, as the
emerald, the amethyst, the opal—which may be due, in part at least, to
reflection from the brilliantly colored mandibles. These hues, according
to M. Simon, indicate different powers of vision and qualities of service.
Quoting favorably a remark of Dr. Vinson, he would divide a spider’s
eyes into “diurnal” and “nocturnal.” The diurnal eyes are brilliant,

Color of
Hyes.

' Spiders of Dorset, Vol. II., page 241.

_ ne a

288 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

transparent, and seem to have a pupil and iris; the nocturnal are, on the
contrary, dull and opaque.!

Lebert expresses the same opinion,? adding further that those which are
most convex and brightly colored, serve to see during daylight, and the
others, which are flatter and colorless, serve during the dusk. It occurred
to me to test this theory by making such division on the basis of habit,
as above referred to, and observe whether the spiders which incline to
nocturnal habits have the nocturnal or white eyes, and vice versa. Such
a comparison, extended to a few species, shows the following results :—

Among our indigenous Orbweayers, those whose night habits are most
pronounced, or whose ordinary habitat is dark or shaded places, proved to

have light colored eyes. For example, I rarely find Epeira strix
Night upon her web in daytime; she comes out for prey in the early
Aa ae evening or twilight, and remains quite persistently throughout
Byes. the day in her nest of curled leaf or in some conyenient den.

The eyes of this species have the entire rear row and the side
eyes of the front row a light gray color. The middle front eyes are of the
same color, but a darker shade. Meta menardii, which persistently inhab-
its shaded places, and which I have found in caves in central Pennsylva-
nia, has all its eyes a light pearl or gray color. The specimens of this
species examined were found in Sinking Spring Cave quite distant from
the mouth. The Ray spider I have always found in ravines or well shaded
spots under the cover of ledges or foliage. Its eyes are all a light pearl,
the middle front eyes having a little darker shade.

Turning to other tribes I found, for example, that the Medicinal spider
(Tegenaria medicinalis) has all its eyes yellow, with the exception of the
small midfront pair, which are dark with a marginal ring of light color.
Its dwelling place is habitually cellars and dark corners. I have often
found it living quite in the shade.

Clubiona palens, which lives within a silken tube and is frequently
found underneath stones, has pearly white eyes, the middle front being a
slightly darker shade. The eyes of Agalena neevia are all a uniform bril-
liant yellow (amber), rather darker than Tegenaria medicinalis. This spi-
der inhabits a tube which expands into a sheeted web. She keeps habit-
ually on guard within the mouth of her tube, but sallies forth at all hours
of the day after the prey which drops or alights upon her web. She evi-
dently possesses good day sight. So far, therefore, one might say that the
conjecture that the white eyes are used for seeing in the dark, are nocturnal
eyes, in fact, is corroborated.

But a further examination introduces facts which are in serious conflict
with the theory. For example, Argiope cophinaria lives persistently in the

1 Simon, Histoire Naturelle des Araignees, page 35.

2 Die Spinnen der Schweiz, page 6, quoted by Lubbock, “On Ants, Bees, and Wasps,”
Linn. Soc. Jour., Vol. XX., 125.

COLORS OF SPIDERS AND THEIR COCOONS.

1, ARGIOPE COPHINARIA; 2, EPEIRA INSULARIS; 3, EPEIRA DOMICILIORUM; 4, 5,6, COCOONS AND FEMALE
OF ARGIOPE ARGENTEOLA; 14, MALE OF SAME; 15, MALE ABDOMEN; 7, 8, COCOON AND FEMALE OF
GASTERACANTHA; 9, NATURAL SIZE OF FEMALE; 10, COCOON STRING OF CYCLosa BIFURCA; 11, FEMALE,
SIDE VIEW, ENLARGED; 12, DORSAL VIEW, ENLARGED; 13, NATURAL SIZE.

Auth- Del. in part Edw. Sheppard, Lith.& Del.

vpn ad als
: , bee it >
T fe WE

sc ites ith

. 4
ran) ot

THE SENSES AND THEIR ORGANS. 289

light. I have always found it hanging on the central shield of its web in
broad daylight and at all hours of the day. Its eyes are a light yellow
color. The same is true of Argiope argyraspis. Acrosoma rugosa I have
always found upon its web in daytime. This is a wood loving species, but
commonly spins its web in open places. Its eyes are light gray, the mid-
dle front pair having a little darker shade. Epeira labyrinthea is also a
diurnal spider, selecting, as a rule, a position upon branches stripped of
foliage or dead limbs. Its rear eyes are light colored, pearl gray or a del-
icate amber, but those of the front row are black. Epeira insularis habit-
ually occupies its nest of sewed leaves during the daytime and often at
night also, but it takes prey quite freely during all hours of the day. Its
eyes are all light colored. Two gravid specimens of this species which I
examined had eyes decidedly lighter than other specimens, and the ques-
tion occurred to me whether it might not be that the color of the eyes is
affected during the period of gestation. I had not sufficient specimens,
however, to follow this inquiry, which, perhaps, is not worthy of further
attention. .

The eyes of Linyphia weyerii, which I have examined from several
specimens received from Luray Cave, are of light color, the two central
eyes being white. The latter is a marked variation from the general con-
dition of this pair of eyes, which appear to be darker, as a rule, in all
‘terrestial species, and to be obliterated in some cavern Poaunes

I submitted a few European species to a similar examination.’ 'Tetrag-
natha extensa (Russia) has the side eyes a yellowish brown, the side rear
eyes of lighter hue than the side front. The midrear eyes are
dark yellow, and the midfront eyes are darkest of all. Epeira
sclopetaria (Ireland) has the side eyes light colored, the rear eyes
being lightest. ‘The midrear eyes are a dark yellow, and the midfront
darkest of all. Epeira sclopetaria of Russia is colored in the same way.
Epeira quadrata (Russia) has the side rear eyes light colored, the side front
eyes a little darker hue, and the front eyes tolerably dark. Epeira dia-
demata (Russia) has the rear eyes brownish yellow, the front eyes a darker
hue of yellow, and the front side eyes a darker yellow, and the midfront
eyes darkest of all, almost black.

None of the above species can be classified as nocturnal in their habits,
although all of them, of course, are able to capture prey at night. 'Tet-
ragnatha extensa and Epeira sclopetaria are continually seen upon their
webs in broad daylight. Epeira quadrata and Epeira diademata belong to
nesting species, having habits similar to our Epeira insularis and trifolium.
‘They live in dome shaped tents, roofed and walled by clustered leaves or
by a single rolled leaf, Their faces are towards the opening, looking upon

‘The Russian species were received from Mr. Waldemar Wagner, of Moscow, and the

European
Species.

British species were collected partly by myself and partly by Mr. Thomas Workman, of

Belfast.

i ia pie ae

290 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

their webs, which are usually spun in well illumined places. They may
be regarded as diurnal in their habits quite as much as nocturnal.

Of Epeira cornuta I examined specimens from Moscow (Russia) and
Treland. The side eyes are an amber yellow of a rather dark hue. ‘The
middle eyes are still darker; the midfront ones the darkest of all. The
side rear eyes have the lightest hue. This species resembles our Epeira
strix in its habits, and is much inclined to live in dark places, and for
the most part retires to its cell or den, or some secluded retreat, during the
day, showing its greatest activity at night. It is not exclusively a nocturnal
species, but approaches nearly that habit.

Of Epeira umbratica I examined two species captured by me upon the
outer basaltic columns of Fingal’s Cave and one from England. The side

eyes are amber yellow, of a darkish hue. The midrear eyes
Epeira have a little darker color, and the midfront darker still. In
raracas the English specimens the colors were similar, but a little darker,

the midrear eyes being quite dark and the midfront almost
black. This species, as is well known, is nearest a nocturnal species of all
the Orbweavers of Europe. It quite frequently seeks shaded places, although
this is not its exclusive habit. The webs of the Fingal’s Cave spiders were
exposed to the light, although the individuals were hidden within a little
recess of the rock. I have seen numbers of the webs of these species on
the grounds of Tatton Hall, near Manchester, the estate of Lord Edgerton,
swung between the railings of a rustic bridge, shaded only by foliage.

These two spiders present the strongest testimony in contradiction of
the theory that the white eyes are most useful to those species that are
nocturnal in habit. Judging by their habits, their eyes should have been
the lightest of any Orbweavers of Europe, but the contrary appears to be
the case. I am not able to solve such contradictory facts. Quite at the
opposite extreme, and in line with the general tendency, are the eyes of
cavern fauna. The eyes of Linyphia weyerii, which I have examined from
several specimens received from Luray Cave, are all light colored, the two
central eyes being white.

The above facts appear to point to the conclusion that eyes of a light
color are better suited for seeing in the dark, but that dark colored eyes
are not necessarily especially valuable to the species having diurnal habits.
In short, there does not appear to be a corresponding difference between
the nocturnal and diurnal habits of spiders, and the supposed nocturnal
and diurnal eyes, sufficiently marked to justify a division on that basis.

An examination of the above facts also shows that there is a quite

persistent tendency on the part of the side eyes to be lighter in
Most color than the middle eyes; and, of the side eyes, the rear ones
Persistent : : P
Hives, are generally the brightest. It also appears that the middle

group of eyes tend to be darker colored, and, of these, the front
pair are darkest of all.

—_—_—

THE SENSES AND THEIR ORGANS. 291

On the theory that the dark colored eyes are of the greatest advan-
tage in the light, and the light colored eyes most valuable in the dark,
one would expect that in the case of cave species the eyes first to disap-
pear would be the middle ones, and those longest to persist the side ones;
the rear eyes longest of all. I was anxious to test this theory, but unfor-
tunately had but a scant amount of material to do it. However, the few
facts at hand are valuable for comparison, and are quite in harmony with
the above inference.

Pavesi has observed! that while the species of Nesticus possess nor-
mally eight eyes, in a cave dwelling species, Nesticus speluncarum, there
are only four, the four middle eyes being atrophied. This suggests that
the four central eyes serve especially in daylight.

The above observation of Pavesi corresponds substantially with Emer-
ton’s studies of the spider fauna of some of the large caverns of America.?
Out of six species of Lineweayers described, five
show some unusual condition of the eyes. Three
species haye the front middle pair very small ;
“one has all the eyes small and colorless, with the
front middle pair wanting in the males and some
females; and one species is entirely without eyes.
The complete obliteration of all the front middle

“ pair in some specimens, and the partial atrophy
of the same eyes in others, would seem to indi-
cate that the organs so situated are of most ben- jy. ogs, Face of Linyphia inserta,
efit in full sunlight, or, at all events, that sun- _ with two eyes wanting. Fiq. 285.

if 1 A . t h a ‘ . } Byes of another individual, same
1g. nt 1s more necessary to their preservation than species, all present. Fic. 286.
the others. Face, of Anthrobia mammouthi,

with eyes atrophied.

Fia, 284. FiG, 286.

Several figures are here presented, made from
Emerton’s drawings,* which will illustrate the progressive atrophy of the
eyes in the case of some of the spider fauna of the caverns of Kentucky
and Virginia. Fig. 286 shows the face of a female Anthrobia mam-
mouthia, from which the eyes have been entirely obliterated. Fig. 285 is
a drawing of the eyes of a female Linyphia inserta from Fountain Cave.
Here the middle eyes of the front row are extremely small, but, neverthe-
less, are quite manifest. In contrast with this is Fig. 284, which repre-
sents the head and mandibles of a male of the same species (Linyphia
inserta), from which the middle front eyes have entirely disappeared. The
absence of this or any pair of eyes, so far as my knowledge extends, is in
no case a sexual characteristic, so that the disappearance of these eyes, if
we suppose the figures to have been drawn correctly, can only be attributed

‘Sopra una nuova specie de Ragni appartenente alle collezioni dei Museo Civico di
Genoya, Ann, Mus. Civ., 1873, page 344.

* Notes on Spiders from Caves, Am. Naturalist, Vol. IX., page 278.

8 Op. cit., plate i., Figs. 5, 18, 21.

—— - _ r a

292 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

to the gradual progress of the atrophy, or to one of those natural freaks
which occasionally occur with spiders as well as other living things.

Occasional irregularities in the number of eyes are not wholly due to
causes which produce the atrophy of those organs. For example, Black-
wall! records that an adult female Epeira inclinata captured in August
was entirely destitute of the left intermediate eye of the posterior row,
and the right intermediate eye of the same row was not the usual size.
In another adult female taken in the autumn of the same year the right
intermediate eye of the posterior row had not one-eighth of the usual size,
being merely rudimentary. This spider abounds in many parts of Great
Britain and Ireland, and seems to prefer districts which are well wooded,
but otherwise has no habits which would account for such irregularities.
It is simply an abnormal state of the eyes, resulting from some morbid
condition.

Concerning Linyphia inserta, drawings of whose eyes are shown at Figs.
284 and 285, Emerton says that the eyes are small and colorless and sep-
arated from each other. The front middle pair are very small, hardly
larger than the circles around the base of the hair by which they are sur-
rounded, and only distinguished from them by wanting the dark ring
which surrounds the hair circles. In five females from Fountain Caye all
the eyes are present, (Fig. 285.) In one female one eye of the front mid-
dle pair is wanting. In three males from the same cave both front middle
eyes are wanting, as in Fig. 284. In one male only one of the front mid-
dle pair is wanting. In four females and one male from Bat Cave, Carter
County, Kentucky, the front middle eyes are wanting.? This irregularity
in the number of the eyes indicates with little doubt the fact that the in-
fluence of environment has been strongly felt in producing a greater or less
atrophy of these organs of sight.

IV.

That spiders have accurate perception of the direction and intensity of
light, one may easily determine by experiments with the young. A great
number of such experiments I have made, but will content myself with
an illustration or two which fully typify the universal tendency. A brood
of young Zillas heretofore described (Volume I., page 148) habitually

.,,.. Placed themselves upon the illuminated side of their common
Sensitive Frcs fis _ :
to. Tait. eb, This position during the day looked toward a bay win-

dow a few feet from the table on which the colony was settled,
and at night was on the opposite side and toward the lamp on my desk.
In the morning, if the day were bright so that. the sunlight streamed in
at the bay window, the colony invariably migrated to that side. If the
day’ were dull, inasmuch as a side window shed some e light over the table,

‘ Spiders Gt. Bt. & ik, .» page 3 BG 2 ire cit., page 280.

THE SENSES AND THEIR ORGANS. 293

the movement was not so decided. At night the direction was reversed,
and the migration set toward the library lights, and the colony settled as
near them as possible.

This behavior was so often and uniformly observed that the conclu-
sion was quite satisfactory, but I nevertheless made a series of experi-
ments which fully confirmed it. For example, one evening I found the
brood massed at two points (a and b, Fig. 287) on either side of a toy
wooden column, hanging in a dim light which fell from a lamp that
previously had been turned down. An oil lamp giving a bright light
was now lit, and so placed (beyond the point c) that one cluster (at a)
was in shadow, the other (at b) faintly illuminated. In twenty minutes
twelve individuals had passed over from the partly shaded spot (b) to
the illuminated point (c), and about half the group in the shadow (at a)

5 ne eerie meen sees me:
# ee rE re

a. b c

Fic. 287. Experiment to show the perception of light by young spiders. The group in shade were all
transferred to the shelf by lighting the lamp thereon.

had crossed to the better light at b. The lamp was then removed to
the opposite side of the table, reversing the conditions of light, casting ¢
in the shadow, b partly in the shade, and throwing full light upon a.
Instantly a movement began among the spiders now in the dark at ¢,
who turned and ran rapidly along the lines communicating with the col-
umn. In less than two minutes only four of the twelve remained; one of
these soon followed, and shortly the other three. In the meanwhile the
shaded group at b was being broken up by an active transfer to the illu-
minated section at a.

Another experiment gave the following results: I captured a female
Kpeira sclopetaria at Atlantic City and placed her within a box to cocoon.
May 26th, a cocoon was formed in the angle of the box, over which was
placed a triangular piece of sheeted spinningwork attached at the three

294. AMERICAN SPIDERS AND THEIR SPINNINGWORK.

margins to the sides and bottom of the box. (Fig. 288.) This formed the
cocoon tent, whose dimensions were two inches on the floor and along the
sides. June 13th, a small cluster of yellowish white spiderlings appeared
at the bottom of the cocoon, showing that the young had already hatched
and found their way outside within the intervening eighteen days. They
had then the appearance of having been hatched a day or two.

During the ensuing week they gradually darkened in color and were.
joined by their fellow broodlings, who gathered in a semicircle around the
upper edge of the cocoon on the box. Here they remained six days upon
the top of a case of drawers near an open window. While reading on
the evening of June 19th by the light of an argand burner, I glanced up-
ward and observed that the lamp was covered with web lines that fringed

; ipods ‘the bottom of the por-

i Ae Wa We We iy ge celain shade and met-

Wey My al stand. Upon these
is Le L

i 3 lines forty or fifty spi-
Zp Ny. derlings hung, in the
full blaze of light.
They had _ evidently
just issued from the
cocoon tent, and had
been carried by the
wind along a bookcase
and across the desk to
the lamp, a total dis-
tance of fourteen feet.
A bridge line four feet
long was strung from
the bookcase to the
lamp, along which the brood had clambered, attracted undoubtedly by
the light. There was no reason why they should have sought that particu-
lar spot, and many reasons why they should have gone elsewhere, but the
light dominated their action. (See Volume I., Fig. 141.)

A portion of these I removed to a table, where, during the night, they
set up a cobweb commons of the kind heretofore described, and remained
grouped thereon until next morning. Then they and nearly all their fel-
lows were dispersed by the breeze when the windows were opened. It thus
appeared that exposure to and the force of the wind determined the fact
of a-quick and wide distribution of spiderlings immediately after egress.
In the case of the other broods that were protected from the effects of
strong winds, the young remained within a limited space for two or three
weeks. Most of them gradually disappeared by aeronautic flight, mount-
ing in that way to the ceiling and walls; some of them spun small orbs
in the vicinity, and some remained upon the common web to the end.

Ai
ify

Fic. 288. Cocoon tent of Epeira sclopetaria.

THE SENSES AND THEIR ORGANS. 295

¥.

The vision of spiders is evidently limited in extent, although it remains
to be determined what that extent may be. Prof. Auguste Forel, so
widely known for his distinguished studies of ants, records in a
paper on the “Senses of Insects,” that if a cocoon be removed
from a ground spider (Lycosa) to the distance of two or three
inches, she will hunt about for it and have great difficulty in finding it.
He states, moreover, that jumping spiders (Saltigrades) cannot perceive
their prey at a distance greater than two or three inches.!

Professor Peckham, as the result of his experiments, concludes that
spiders, at least the Lycosids, even when they see their cocoons, are not
able to recognize them except through the medium of the sense of touch.?

But, on the other hand, the Peckhams’ observations make it plain that
Saltigrades can see objects at a distance of at least ten inches, ‘They fre-
quently saw them stalk their prey at a distance of five inches. They re-
peatedly held Astia vittata on one finger and allowed it to jump to a
finger of the other hand, gradually increasing the distance up to eight
inches. As the distance increased, the spider paused a longer time before
springing, gathering its legs together to make a good “ready.”

I have repeatedly verified this experiment with an adult female of
Phidippus morsitans. Holding the spider upon a box in which she had
been taken, I approached a finger within an*inch of her face, until her
attention was evidently attracted. Presently she leaped the space, alight-
ing upon the finger. I then restored her to her position upon the box,
and by manipulation again tempted her to escape by vaulting twice the
distance. By gradually increasing the space, she finally jumped a distance
of from three and a half to four inches. Her whole action showed that
she had seen the object before her, had discerned the fact of an interven-
ing space, had carefully measured the distance, and then yaulted, success-
fully reaching the object. As usual on such occasions, she always kept
herself secure by a dragline attachment to the box from which she jumped.

Twice the Peckhams saw a male Astia vittata chasing a female upon
a table covered with jars, bags, and boxes. The female would leap rapidly

from one object to another, or would dart over the edge of a
Good — ook or box so as to be out of sight. In this position she would
Sight in : . : : mates ;
Salti- remain quiet for a few moments, and then, creeping to the edge,
grades, Would peer over to see if the male were still pursuing her. If

he happened to be hidden, she would seem to go to him even
when ten or twelve inches away, and would quickly draw back. But in
case he was hidden behind some object, she would hurry off, seeming to

Limited
Vision.

1 Sensationes des Insectes, I. Recueil Zoologique Suisse, Tome TV., No. 1, pages 18, 19.
2 Mental Powers of Spiders, pages 401, 402. -

oe a a i

296 AMERICAN SPIDERS AND THEIR SPINNINGWORK,

think she had a good chance to escape. The male in the meantime fre-
quently lost sight of the female. He would then mount to the top of the
box or jar upon which he found himself, and, raising his head, would
take a comprehensive view of surrounding objects. Here he would remain
until he caught sight of the female, which he often did at a distance of
at least ten inches, when he would at once leap rapidly after her.?

These observations certainly show a well developed power of vision,
sufficient at least for all purposes of the active life led by these wandering
Saltigrades. They confirm the opinion elsewhere expressed in this work
(Volume I., page 19), that individuals of this tribe possess more highly
developed vision than those of any others. One might almost infer this
from the appearance of their eyes, the seeming expression which rests
upon their faces, and the general intelligence that marks their demeanor
and movements.

Sir John Lubbock appears to have tested the experiment made by the
Peckhams upon sight of spiders. Lycosa saccata, a familiar European

species, was selected for his observations. A female from whom

Lub- an egg sac was removed was placed upon a table, about which
bock’s the- ram fora adie aaa Tastes Peak dient WI i
Experi. ‘%2¢ tan for a while as though looking for her eggs. 1en she

ments. became still, the cocoon was placed about two inches in front

of her. She evidently did not see it. It was gradually pushed
towards her, but she took no notice until it nearly touched her, when she
cagerly seized it. :

The cocoon was again removed, put in the middle of the ‘table, from
which all other objects had been taken. The spider wandered about, some-
times passed close to the egg sac, but took no notice of it. She spent
an hour and fifty minutes in this aimless wandering before she found the
cocoon, and then apparently by accident. A third time it was removed,
placed upon the table as before, and an hour was spent in wandering before
it was discovered. The experiment was tried with other individuals, and
with the same results. Sir John’s conclusion is that “it certainly appeared
as if they could not see more than half an inch before them; in fact,
scarcely further than the tips of their feet.’

It is impossible, however, to admit the explanation which the learned
author has made of this inability promptly to recognize the cocoon. “It
must be remembered,” he says, “that the sac is spun from the spinnerets,
and that the Lycosid perhaps had never seen the bag of eggs.” On the
contrary, the manner in which the Lycosid prepares her round cocoon,
as it has been quite fully shown and described by myself, compels
the conclusion that the mother Lycosa has perceived her cocoon both by
the touch of sense and sight from the beginning of its fabrication to the

‘ Mental Powers of Spiders, page 402.
* Senses, Instincts, and Intelligence of Animals, 1888, page 179.
® See Chapter V., page 144.

THE SENSES AND THEIR ORGANS. 297

time when, having deposited the eggs within the little circular patch,
she rolls the cover around them into a ball, and then attaches the ball
to her spinnerets. This is undoubtedly the universal method with spiders
that carry about their egg sacs. The cocoon covering is first spun upon
some surface, the eggs enclosed, the ball prepared, and the last act is attach-
ing it to the spinnerets.

Of course, during this process, a spider whose eyesight is so good that
it can perceive its prey at a distance of several inches, must of necessity
have seen its cocoon. It would be impossible for us to reach any other
conclusion. The confusion in maternal recognition and selection of her
offspring cannot, therefore, be accounted for by defective sight.

Among the Theridioid spiders is a large group of species which Black-
wall has placed under the genus Walckenaéra, which present some re-
markable peculiarities in the location of the eyes. These are
distributed on the anterior part of the cephalothorax, which
sometimes in females and usually in males is remarkably: ele-
vated. The drawings here presented (Figs. 289-293) are taken from Black-
wall’s descriptions of
Walckenaéra acumina-
ta,! and represent one
of the most remarkable
of these turret like de-
velopments of the ca-
put for the reception of
eyes.2 The length of
the female (Fig. 292)
is about one-seventh
of an inch. The sexes
are similar in color, but
the male (Fig. 289) is
smaller than the fe-
male, and the anterior
prominence of its ceph-
alothorax is much more
elongated and slender,

measuring about one- ‘

: l £ : | . Fic, 289. Male. Fic. 290. Eye turret of male, greatly enlarged; front
twentiet 10F an men mn view. FiG, 291. The same, side view. Fic, 292. Female. Fic, 293.
length. This eye tur- Same, side view of cephalothorax, with outline showing natural size.

ret is elevated vertical. ‘“™" Bickel)

ly and dilated near the middle and at the apex (see Figs. 290, 291), the
latter dilatation being separated by a transverse groove into a superior and
inferior segment, both of which are rough, with short, strong hairs. On

Hye
Turrets.

Bde SS
a Try ty SSS

Walckenaéra acuminata. 291

' Spiders Gt. Br. & In, pl. xx., Fig. 208.
* See also Mr. W, Simon’s Arachnides de France, Vol. V., part IIL, page 820.

298 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

these enlargements the eyes are seated. Both sexes of this curiously con-
structed spider were taken in England in October under stones and on‘rails,?

It is to be observed that the greatest prominence of the eye turret of
Acuminata belongs to the male, and this appears to be the rule with simi-
larly constructed species. What can be the cause or the use of

toe of this? One would think that such a remarkable development
Turrets, must be intended to serve the spider some special advantage in

making its way around its natural site. But until we know
something more in detail of the habits of the species we can only venture
a suggestion. The probability is that all these small Theridioid spiders,
like many of the minute species dwelling in the United States, spend their
life upon or near the surface of the earth, where they weave their loose
webs of lines around the bases and among the roots of grasses. It may
be taken for granted that the male, in his amorous search after the female
during the period of courtship, would find such an eye turret advantageous
in detecting the home web and person of his mate amidst such an entang-
ling environment. At least, no other advantage can be suggested for this
strange exaggeration of the tubercles on the eye space.

A difficulty which at once arises, is the fact that so many other spe-
cies haying like habits and location show no peculiarity of a like kind,
and appear to have no need therefor. Great numbers of species have
eyes placed upon tubercles or slightly elevated parts of the caput, usually
of a rectangular or quadrangular shape. In some Epeiroids these tubercles
are quite pronounced, and many of the genera are sharply distinguished

thereby. In many more genera, and indeed it may be said in

aang almost all, the eye or ocellus proper is quite commonly raised
and Rows'pon or within a little cup of black chitinous formation, which

thus slightly elevates it above the surface of the face. As far
as I know, there are not many eyes that appear to be set immediately into
the eye space without this enclosing cup.

Among Saltigrades and Citigrades one pair of eyes is commonly placed
at considerable distance behind the others, giving in this way an addi-
tional advantage to the species by the seeming ability to observe, to some
extent, objects lying behind it. The breaking up of the eyes into rows,
generally two, but sometimes three, may serve the same useful purpose.
This whole subject is an interesting one, but the facts in my possession
are so few that I can do little more than open it for the consideration of
future students.

Vi.

In considering the sense of smell in spiders two questions require at-
tention. In the first place do spiders possess this sense at all, and, if so,

1 Blackwall, idem., page 290.

Ee ne

> .
a

— eS

THE SENSES AND THEIR ORGANS. 299

to what extent? And, in the second place, where are the olfactory organs
located? The conclusion which I had reached, as the result of experi-
ments and observations’ of my own, is that spiders have little
The sense of smell, although they are in some way affected by certain
Sense of : ; ae
Smell, dors. I have long entertained the opinion that the sense of
smell in spiders, like that of hearing, abides entirely in the deli-
cate hairs which constitute the covering and armature of the creature. I
have thought that in some way the nervous system receives through these
organs or appendages impressions that may be considered analogous to
hearing and smelling in the higher animals; but, further, that both these
senses are in an extremely rudimentary condition.

These conclusions are substantially confirmed by the experiments re-
corded by Professor and Mrs. Peckham, which were carefully performed
and continued through a number of examples large enough to
justify a conclusion, Their method was to place a rod dipped
in various essential oils, cologne, and several kinds of perfumes,
close to the various parts of the spiders, and note the effect.
The results were carefully tested in all cases by first presenting a clean rod.
Among the essential oils used were oil of peppermint, of lavender, of cedar,
of cloves, and of wintergreen. The first experiments made were upon some
tame Attide that had taken up their abode with them. These were fear-
less little creatures, ready to jump upon the finger, catch the gnats that
were offered them, or drink from a spoon. They showed the same facility

in smelling that they exhibited in seeing, and were quick to re-
Effects of :
Oden spond to any test of their sense of smell. The most common

effect. produced by an odor was that it caused the spiders to raise
their fore legs and palps, which sometimes they also moved up and down.
In one case great excitement was caused by the approximation of pepper-
mint. In several cases the spiders, after indicating that they noticed the
scent, moved away from it.

With Orbweavers the effect of the various perfumes was to cause an
upward jerk of the abdomen and a movement of the legs. Sometimes the
tips of the legs were rubbed between the palps and the falces. The result
of two hundred and twenty experiments may be summed up as follows:
Three species, Epeira hortorum, Dolomedes tenebrosus, Herpyllus ecclesias-
ticus, did not respond to the test. In all other cases it was evident that
the scent was perceived by the spiders, although it may be noted that
among spiders of the same species great differences of degree exist in their
sensitiveness to odors, ‘The spiders exhibited their sensitiveness to the
various perfumes by movements of the legs, palps, and abdomen; by shak-
ing their webs; by running away; by seizing the rod which had been
dipped into the scent, enswathing it as they would insects; in the case of
the Attide, by approaching the testing rod with the first leg and palps
held erect, but whether in the way of attacking it, or, as it sometimes

The Peck-
hams’ Eix-
periments

300 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

seemed, because the smell was pleasant to them, the observers could not
determine. !
As to the olfactory organs, the experiments would indicate that they are
distributed more or less over the entire surface of the body, especially at
the tips of the feet and at the apex of the abdomen, but that
Olfactory they probably are more highly developed in the fore part of the
alana body and in the organs immediately surrounding the face. In
order to test the value of the palps as olfactory organs, those
parts were dissected from two females of Argiope cophinaria. The result-
ing tests indicated that the araneads had suffered no apparent loss of sen-
sitiveness. In one case the application of the oil of lavender at the front
of the body caused the spider instantly to contract her legs and rub the
tips thereof, one at a time, upon the falces. The other spiders responded
to heliotrope and Chinese bouquet by quickly jerking the abdomen and
rubbing the tips of the legs over the falces,

VII.

A number of experiments, prolonged through several years, have been
made with a view to determine the extent to which spiders hear, and the
location of the auditory organs. I have found myself continually
thwarted, or at. least confused, by doubts lest the various re-
sponses made were caused by independent movements of the air,
which, operating on the delicate body armature, of course produced sensa-
tion and excitement. I made many experiments upon the tarantula “ Lei-
dy,” which I had in my keeping for more than five years, and whose life
I have elsewhere recorded.2. These experiments were made with tuning
forks, with several kinds of musical instruments, and by sounds of all de-
grees of sharpness and dullness made by the human voice and various
sonorous objects.

Once I had nearly concluded that the great creature was immensely
excited by my flute. Certain tones, when the instrument was brought close
to the vessel in which the tarantula was confined, caused her at once to
rear upon her hind legs in that rampant attitude which this creature as-
sumes when about to strike its prey. During one experiment, however,
something occurred which induced me to drop my flute and make a light
puff of air with my mouth over the edge of the glass cage, so that the
wind thus produced would be reflected against the animal. At once she
assumed the rampant position precisely as before. Repeating this, I found
that it was simply the motion of the air over the mouth hole of the flute,
which was carried into the cage, that had agitated the tarantula, In other

Sense of
Hearing.

1 Mental Powers of Spiders.
2 Proceedings Acad. Nat. Sci., Phila., 1887, page 869, sq. “Prolonged Life in Invyer-
tebrates: Notes on the Age and Habits of the American Tarantula.”

——— ee Pl rt =
acre 4 "*" ’ ‘ .

THE SENSES AND THEIR ORGANS. 301

words, when I blew upon her, the action of the wind excited her, and
caused her to rear upon her hind legs as though some enemy were ap-
proaching or some victim coming within reach.

This will- illustrate some of the difficulties in the way toward a just
verdict. Of course, all sound is produced by vibrations of the air; but it
seems possible that the movements produced by tuning forks and other
instruments, sounded, as they must be, in so close proximity to the spider,
may and probably do mechanically agitate the hairs upon the body, and
thus effect the sense of touch alone, producing an excitement which I have
often observed and sometimes attributed to hearing. The difficulty has
been to separate between these two sensations and decide whether my ex-
periments had not simply excited the spider by touch. My conclusion, as
the result of independent observations, is that if spiders have
any sense of hearing proper, that sense is distributed, like the
sense of smell, over the entire body; and, further, that it can
scarcely be distinguished from the sense of touch as it is known to us.
No doubt, however, the aranead has some of the advantages within its
limited sphere that auditory organs proper give to higher animals.

On this point the Peckhams also made a number of experiments, with
some interesting results, as follows: Certain spiders indicate that they hear
a vibrating tuning fork, by characteristic movements of the legs; others by
signs of alarm, dropping from the web and keeping out of sight for a
longer or shorter time. One spider, at least, Cyclosa caudata, when sub-
jected to frequent approximations of a tuning fork, seemed to become grad-
ually accustomed to the sensation, and, instead of dropping from her web
as at first, remained immovable and apparently undisturbed. One of the
most interesting points developed is that orbmaking spiders appear to be
most sensitive to the vibrations of the tuning fork. All these responded
promptly, being evidently alarmed by the sound.

On the contrary, spiders that make no webs gave not the slightest heed
to the sound. Among those species that proved unresponsive were two
Tubeweavers (Herpyllus), several Lycosids, and the familiar Dol-
omedes tenebrosus. Professor Peckham suggests that this differ-
ence may be partially explained by difference in the feeding
habits of the two groups, an explanation which leaves much to be ex-
plained. May we venture to suppose that, in the case of the Orbweavers,
the particular effect produced by the vibrations of the tuning fork upon
the spider hanging on her web, or upon the delicate filaments of the web
itself, is very much the same as that produced by the rapidly vibrating
wings of an insect when humming around the snare or when struggling
within it? Certainly Orbweavers are dependent upon some such agitation,
especially of the web, for the intelligence that their snares have succeeded
in trapping a victim. On the other hand, we know that Lycosids, for ex-
ample, which stalk their prey in the open field, instead of ensnaring them

Organs of
Hearing.

Effects of
Sounds.

302 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

upon silken nets, are chiefly dependent upon sight for knowledge of their
victim’s presence and power to secure it.

A few experiments were made to determine where the organ of hearing
is located, which, as far as they go, seem to confirm my own opinion as
expressed above, that the auditory apparatus is but little specialized, and
is distributed over a considerable portion of the epidermis. The removal
of the palps appeared to make no difference in the power of individuals to
respond to the vibrations of the fork. So also the removal of the first
pair of legs seemed to leave the auditory organs intact, at least made no
impairment of power to respond to sound.

I will now describe two of my own experiments, as illustrative of the
method pursued and the grounds for reaching my conclusion, An adult

; female Domicile spider, hanging in the centre of her orb, was
ee tested by an “A” tuning fork. The fork in rapid vibration was
periments moved all around her, gradually approaching until the instru-

ment was within a few millimetres of her person. All parts of
the body were thus tested and no signs of excitement appeared. ‘The fork
was then touched to the top of the web, when Domicile immediately
showed signs of excitement, acting precisely as if an insect were entangled
at that point. She turned herself in the direction of the fork, grasped
the radii leading outward to the point of contact, pulled upon them in the
usual way as though testing the strength or entanglement of the supposed
insect, and then gradually approached the point of agitation, I withdrew
the fork, and, as the spider came up to the margin of the web where the
instrument had been, she turned around, made several motions as though
examining the strands, spun out a few lines, and went back to her hub,
dragging a thread after her.

The fork was then placed at the bottom of the web, then at the sides,
and so successively to various points on the circumference of the orb.
The same action substantially resulted, the spider always going to the spot
where the fork was vibrating against the lines of her web. Finally, I suf-
fered the fork to remain as the spider approached. She touched it with

her fore feet; at once showed tokens of surprise, indeed of some

pitrins stronger emotion; she seemed to be expressing the feeling, “I
Spider. have been fooled;” turned her back upon the fork, shot out a

thread from her spinnerets, and scampered away to the hub,
where she curled herself up, drawing her legs toward her face until the
knees projected above the head, exhibiting what I cannot express by
another phrase than tokens of disgust.

I then laid the vibrating fork upon the outer lines, but Domicile would
not respond. I revolved the fork around the hub, close to her, as at the
beginning of the experiment. This time, instead of remaining motion-
less, she waved her fore feet back and forward, as though she had observed
the vibration and were feeling the situation. After a few moments’

THE SENSES AND THEIR ORGANS. 303

interval I again tried the fork to the sides of the web. This time Domicile
was again deceived, and turned towards the point of agitation as in pre-
vious cases. She had evidently forgotten her former experience in the brief
intervening space.

The conclusion which I draw from such an experiment is that the
spider was affected by the vibratory motions of the fork, communicated

through the taut elastic line to the hairs and spines of the feet,
pope The sense of touch was the only means of communicating the
by Touch, *gitation, and no other indications of the spider having heard

the sounds of the fork were here shown than appear in the cap-
ture of an insect under well known and ordinary circumstances. Certainly
the theory has never been advanced, and could not be maintained, that the
spider hears the motion of an entangled insects wings and runs to secure
the victim at such a signal. In such case it is manifest that the spider
feels the action of the struggling captive as it is communicated over the
vibrating radii to her feet, which grasp them at the centre of her snare.
If this be so, it seems to me equally manifest that the same sense was
brought to bear in determining the position of the tuning fork in the
above and like experiments.

I may venture to give the record of another experiment with the tun-
ing fork, which was also wrought upon a Domicile spider, The vibrating
fork was placed near her as she sat upon the hub, and moved around her
four times, the spider showing no symptoms of perceiving the vibrations.
The fork was approached within a few millimetres of the hind legs, where-
upon she showed excitement. This was again repeated a number of times,
the spider showing no signs. I quote the notes: “The vibrating fork is
next touched to a radius on one side of the web. The spider turns and
runs out towards the point of contact. After one minute’s interval the
orb was touched on the opposite side. Domicile leayes the hub, runs out
a little ways towards the fork, hangs upon the radial line, waves one fore
foot around through an open space torn by the rain just below the hub,
then returns to the hub.

“One minute interval. The fork is applied to a radius at the top of
the web. Domicile makes same demonstrations as before. One minute
interval. The fork is applied within a half inch of the spider’s face as
she hangs upon the hub. She stretches out one fore leg as on guard.
One minute interval. The experiment is repeated. The fore legs are both
thrown out quickly, violently, as though to grasp something. After one
minute the experiment is repeated three times. The first two are unno-
ticed. At the third application Domicile shoots out her fore legs. After
the same interval the fork is tried at one side and at the top of the web
several times. No response. It is then placed upon the web at the other
side. The spider runs towards the fork. Various trials are made at the
same and other points, and all fail to elicit a response.”

304. AMERICAN SPIDERS AND THEIR SPINNINGWORK.

The result of this experiment, as in the case above detailed, and indeed

in all other cases tried, is the conclusion that the symptoms of hearing, as

they have ordinarily been described, seem to appear in the spider

Conclu- only when the vibration of the sound instrument is communi-

sine cated along the line of her web to the hairs which form the
ments, ®mature of her body.

I ean reach no other conclusion from the experiments so ad-
mirably portrayed by Mr. Peckham.’ His experiments upon Orbweavers
enabled hinxy to get results which, to his mind, indicated the sense of hear-
ing. That the vibrations produced a state of excitement is true, but the
question is, what was the mediate cause of that excitement? Did it result
from hearing a sound, or was it simply caused by feeling a vibration
similar to that made by an insect captured or hovering near? Mr. Peck-
ham’s experiments indeed seem to me clearly to indicate the conclusion
which I myself have reached.

He used the tuning fork upon a half dozen species taken from different
groups of spiders, making ten or twelve trials on each one. No individual
gave the least intimation of hearing anything. These unresponsive species
belong to the genus Herpyllus among the Tubeweavers, and the genera
Pardosa, Piratica, Lycosa, and Dolomedes among the Citigrades. None of
these spiders, so far as known, ever capture their prey by means of webs—
a fact which struck Professor Peckham. It seemed to him “ remarkable
that while all the Epeiroids responded promptly, being evidently alarmed
by the sound of the tuning fork, the spiders that make no web, on the
contrary, gave not the slightest heed to the sound. This may perhaps be
partially explained by the difference in the feeding habits of the two
groups.”

Does this explain anything? ‘The difference here indicated certainly lies
in this, that the excitement of the Epeiroids was produced by the agita-
tion of the hairs upon their feet, and that agitation was awakened by vibra-
tions of the fork along the lines of the web. That the web is affected by
these vibrations I thoroughly satisfied myself by experiment. For exam-
ple, a vibrating fork, when approximated to the broad, zigzag ribbon upon
the orb of Argiope, would cause it to sway back and forth as though
agitated by the motion of the air, which, beating upon it, alternately re-
pulsed and attracted it.

It would indeed be a remarkable fact were it to be established that;
those spiders which, like the Lycosids, are dependent upon keenness of
the senses for their success in capturing prey, should prove to be destitute
of the valuable sense of hearing; while the webmaking spiders, who are
so little dependent upon the sense of hearing, and are enabled to accom-
plish the most important functions of life by the sense of touch alone,

1 Mental Powers of Spiders, pages 396, 397.

belt akc t a pm

—T- Te

THE SENSES AND THEIR ORGANS. 305

should be found to possess hearing in a degree of acuteness. It is not
often that one finds a contradiction like this in natural history, viz., that
those animals that most need a certain organism or sense have none, while
those which are in least need are highly sensitive.

The experiments of Mr. C. V. Boys! would really lead to the same
conclusion. He notes that after a spider has been dropped from its web
by bringing a tuning fork near it, if the fork is made to touch
any part of the web the spider is aware of the fact, and climbs
the thread and reaches the fork with marvelous rapidity. Mr.
Peckham observes and records a similar fact in the case of Epeira
strix and Epeira labyrinthea.? How shall we account for these actions? It
appears to me clear that when the fork was placed near the animal its vi-
brations agitated the hairs upon the body and the spinningwork immediately
under and around, just as a large insect hovering near in the same posi-
tion would have done. ‘The spider, therefore, did in the case of the tun-
ing fork what it would almost certainly have done in the case of the in-
sect—it dropped from its hub as a measure of defense.

In fact, a spider seated upon its hub is ordinarily at a decided disad-
vantage when an insect enemy, such as a wasp, approaches near it. Its
best defense, therefore, is to get out of the way. But it is quite a differ-
ent thing when its enemy appears at any viscid part of the web and by
the agitation thereof gives indications that it is captured. This is a signal
which the spider understands to mean, in almost every case, that its vic-
tim is ensnared and it can approach it with comparative safety. For this
reason the spider that would run from an insect or a supposed insect that
seemed to threaten it, would run towards the same when it appeared to be
captured and harmless. In these experiments, therefore, I see simply dif-
ferent manifestations of the same sensation of touch under different excit-
ing causes,

Mr. Boys’
Exper-
iments.

VIII.

In connection with these observations upon the auditory powers of spi-
ders, one must at least glance at the numerous stories about, and prevalent
beliefs in, the sensitiveness of spiders to music. There is such a

Sensi- discrepancy between belief in this commendable trait and the
tiveness 7 : :
to Music, 2°neral contempt and disfavor with which araneads are regarded,

that one might incline to think there is good grounds for the
tradition, since it would hardly have arisen under the circumstances with-
out some basis of truth. Certain it is, the opinion is quite ancient and is
widely distributed. Nor are there lacking incidents of seeming historic
verity to be cited in confirmation thereof. It may be of value, it will at
least be interesting, to quote a few of these.

1 “Nature,” XXIII, pages 149, 150. 2 Op. cit., page 411.

306 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

The anonymous author of an ancient history of music records the fol-
lowing examples among others which tend to illustrate the effect of harmo-
nious sounds upon the lower creation.! An officer of a Navarre (Irench)
regiment was committed to prison for having spoken too freely of M. de
Louvois. In order to brighten his prison life he sent for his lute. He was
astonished, after four days, to see that when he played, the spiders would
descend from their webs in his cell and form in a circle around him in
order to listen.

On the first occasion he was so greatly surprised that he remained per-
fectly motionless, when, having ceased to play, all the spiders retired qui-

etly into their dens. This strange assemblage caused the officer
Attract- { fall into a muse upon the accounts related by the ancients of
Se Orpheus, Arion, and Amphion. He assured the author of the

Histoire, who appears to have received his account at first hand,
that he remained six days without again playing, an abstinence which was
caused in part by his astonishment, and perhaps more especially by the
natural aversion he had for this kind of “ insects.” However, he began
anew to give a concert to these animals, who seemed to come every day
in greater numbers, as though they had invited others, so that in the
course of time he found a hundred gathered about him,

But this sort of society in such multitude proving in the end undesir-
able, Monsieur the Captain got a cat from his jailer. ‘This animal he would
shut up in a cage when he wished the presence of his aranead admirers ;
on the contrary, when he would dismiss them, he let the cat loose. The
particular actions of pussy are not described, but the narrator alluded to
them as “making a kind of comedy that alleviated his imprisonment.”

The author of this history from which I quote long doubted the truth
of the above story, but declares that he was confirmed therein by subse-
quently hearing a gentleman of position, merit, and probity, who
played very skillfully upon several instruments, relating an inci-
dent of the same tenor. This person said that he once went into
his chamber to refresh himself after a walk, and took up a violin to amuse
himself with music until supper time, a light being placed upon the table
before him. He had not played a quarter of an hour before he saw sev-
eral spiders descend from the ceiling, who came.and arranged themselves
round about the table to hear him play. He was greatly surprised at such
a demonstration, but did not interrupt the music, having the curiosity to
see the end of so singular an occurrence. The spiders remained on the
table very attentively until some one entered to call the musician to sup-
per, when he ceased to play. Thereupon, as he informed the author, the
spidens remounted to their webs, and, very much to his credit be it said,

The
Violin.

1“TVlistoire de la Miusigte et de es Effets,” edition Paris, 1715. I am nduitad to the
musical library of Mr. 11. C. Wilt, the organist of my chureh, for references to this book and
the work of Sir John Hawkins.

THE SENSES AND THEIR ORGANS. 307

he added that he would suffer no injury to be done them. Subsequently
he declared it was a diversion with which he often amused himself and
gratified his curiosity.!

A somewhat similar incident is associated with the distinguished musical
composer Ludwig van Beethoven. According to Schindler, the story, if not
originated, was generally spread by a biographical paper on Bee-
thoven by Dr. Christian Miiller, of Bremen. The tradition runs
that as often as the little Ludwig played his violin in his
little room a spider, enamored of the strains, let itself down
and sat upon the instrument. When his mother discovered her son’s
strange companion she killed the spider, whereupon the little fellow broke
his violin, Upon this fairy tale Schindler comments: “The great Ludwig
could not recall such a fact, as much as this fable amused him. On the
contrary, he said that everything, even flies and spiders, would have fled
before his terrible scratching.”? Of course, in view of such statements,
not the slightest credence can be given to “this pretty fairy tale of a
poet’s invention,” and it shows how little credit is often due to these pop-
ular fancies that associate themselves with distinguished characters.

The well known anecdote of Pelisson, as described by Abbe Olivet, is
another example in point. This gentleman was confined in the Bastile

, ,, during the reign of Louis XIV., and amused himself by feed-
ee ing a spider, which, from the description, must have been one

pider. 4 - é ;
of a Tubemaking species. The hour of feeding was timed to
the rude music played by a Basque, who was the companion of his cell.
The spider in time learned to distinguish the sound of the music, and to
associate it with the season for its special banquet. This story, with
various embellishments, has had a wide circulation and belief, though I
believe it is wholly discredited by modern historians.

Cowan quotes an account of a certain young ladies’ school at Kensing-
ton, England, in which an immense species of spider was said to be un-
comfortably common. When the young ladies were gathered for
their morning and evening worship, and engaged in singing their
accustomed hymn, these spiders made their appearance on the
floor, as the story goes, or suspended overhead from their webs in the
ceiling. ‘The obvious attraction, it was inferred, was the sweet singing by
the worshiping young ladies.

Walckenaer quoted Gretry as relating in his memoirs that at his coun-
try seat a spider would seat itself upon the table of his piano wheneyer it
was played, and would disappear therefrom when one ceased to touch the

Beethoven
and the
Spider.

Singing
and Piano

‘Sir John Hawkins’ History of Music, Vol. III., page 117, note.

* Biographie yon Ludwig van Beethoyen verfasst von Anton Schindler. Dritte, neu bear-
beitete und vermehrte Auflage. Erster Theil, page 8, Miimster, 1860. See also Thayer’s His-
tory: Ludwig van Beethoven, von Alexander Wheelock Thayer. Erster Band, page 112,
Berlin, 1866,

—=— ee OOO he

308 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

keys. He also states, as a fact which had recently come to his attention,
that a lady while playing a harp in the midst of her garden perceived a
spider fixed in the attitude of attention above her. Presently she trans-
ported her instrument to another part of the room, whereupon the spider
also changed its position, The lady’s curiosity being excited, several simi-
lar movements were made, which led her to conclude that the aranead was
affected by the sound of the instrument.!
Campbell had in confinement a Tegenaria domestica, which at a tune
from a music box would open her spinnerets, an act which is common to
: spiders when expecting food, and go to the centre of her web
oe where she used to be fed. It took some weeks thus to train
her, and the first sign of reconciliation to her imprisonment was
an angry movement of the falees when he touched her, whereas previously
she used to run away.? He also speaks of startling spiders some distance
off by the banging of a door, and their agitation could not be explained
by supposing a current of air; nevertheless it easily might have been ex-
plained by the agitation of the door jambs communicated through the
wall to the web of the spider. Other examples might be cited, but the
above are sufficiently typical, and will answer all my purposes.
Now, the question naturally arises, what basis of fact is there for such
a consensus of belief? Shall we dismiss the matter by simply supposing
that all the individuals concerned deliberately drew upon their im-
A Natural yoination, or were deceived? There is no reason to doubt a certain
miiid part of the facts, at least. It is noticeable that in most of the
above accounts the hour at which the spiders appeared was the
evening, or just before evening. ‘This is a most natural occurrence. I
have sat upon an open porch or verandah, and as the afternoon waned
and sunset drew near, have seen spiders descend from the angles, cornices, ~
and crevices of the ceiling and roof, and spin their webs in the open
spaces above me. They seemed to hang suspended in the air, without any
special purpose, and, indeed, one who did not take pains to watch them
would naturally conclude that they were in a position of fixed attention.
It is customary for spiders, particularly Orbweavers and Lineweavers,
who are the ones to which such manners are to be attributed, to descend
; in this manner and present this attitude as the evening ap-
Evening proaches, which is the time for their ordinary activity. To be

me a sure, they do not remain thus long, but proceed to make their
bration. snares, yet, in so far, the stories which we have quoted may be

. considered as substantially true. The persons concerned may
have seen the spiders descend at the evening hour and hang in an ap-
parent attitude of attention.

1 Apteres, Vol. L., page 110. 2 Observations on Spiders, I’, Maule Campbell.
* Tbid., page 41.

ee a Se

‘

THE SENSES AND THEIR ORGANS. 309

The question may further be raised, did the music have any effect in
promoting this habitual behavior? I would not be willing to affirm it,
but it is not improbable. The vibrations of air caused by singing and
the sounds of flute or violin might affect spiders, as they rested upon their
webs in the angles and corners of the ceiling, to such a degree as to im-
press them with the idea that insects were near. No doubt there is some
similarity in the effects produced by the humming of insect’s wings and
the vibration of musical instruments. It is not irrational, therefore, to
conclude that certain spiders may have been influenced by musical sounds
to such a degree as to hasten their habitual action. But, for the most
part, as far as our stories are to be regarded as credible, I am inclined to
think that habit alone is sufficient to account for the alleged conduct of
the spiders.

That they were affected by the music to the degree believed and re-
ported is not credible; though it is perfectly natural that, under the cir-
cumstances, the observers should have so believed. Imagination could
have gone a far way to supply the details and picture the spiders as
gathering around the table or head of the performer in rapt attention to
the “concord of sweet sounds.” As for the rest, one knows how stories
grow—how a spider or two can be multiplied into a dozen, and how a
dozen can readily grow into a hundred, and a natural action be involved
in mystery or exaggerated into marvel. But, however we dispose of these
‘ widely disseminated traditions, one thing is certain, I have never been
able, after many experiments and observations, with all sorts of music,
good and bad, and with divers instruments, to see the slightest evidence
that spiders are in the least sensitive to music.

IX.

Spiders are well provided with the means of feeling the slightest move-
ments of their webs or other objects. On their legs and palps are long,
slender, silken hairs, which differ from others in that they are attached
to a small disk on the integument. !

It is not my purpose to present anatomical details of the organs through
which the sensations analogous to smelling and hearing are conveyed to
the nerves of spiders. But somewhat has been written upon the
matter, and a reference thereto will be of interest. Dahl has
opened the way in a contribution upon the auditory organs of
spiders,? and this has been freely commented upon by Mr. Waldemar
Wagner, of Moscow.®

Auditory
Hairs.

? Campbell, Observations on Spiders, Trans. Hertfordshire Nat. His. Soc., Vol. I., 1880, page 40.

2 Das Gehor- und Geruchorgan des Spinnen, Zool. Anz., 1883.

5 Des Poils Nommés Auditifs chez les Araignées. Bull. de la Societe Imper. des Natural-
istes de Moscou, 1888.

— —_———

310 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Several illustrations of the types of hairs known as auditory are here
reproduced from the latter writer. Their character is well enough ex-
plained in the legends of the cuts, but a brief description may be added.
The two parts of the hairs are distinguished as the root and the stalk or
shaft. By the root is understood that. portion which enters the cuticle,
and is inserted into an appropriate pit; by the stalk the free part of the
hair which extends above the cuticle. The hairs of spiders, both by their
structure and their root, appear to be divided into two principal types,
perfectly distinct. One sort is distinguished by a root which is much
larger than the portion of the stalk im-
mediately above it. In other words, the
stalk narrows at its foot to swell out
again into a much enlarged root, thick-
ened into the form of a button and in-
serted into a sac like cavity of the skin.
(See Fig. ae) This is what Wagner
denominates a Tactile hair, proper. The
roots of the other sorts of hair are ordi-
narily much smaller, as compared with
their stalks, than the type above named.
(See Figs. 295 and 297, r, r, compared
with Fig. 294, r, r.) The hair pits or
follicles enclosing the roots are also more
simply constructed.

Tactile hairs (poils tactiles) are en-

Fic. 294. Transverse section of a Tactile hair in
the foot of a spider. (After Wagner.) ch, 1,

2, 3, 4, 5, layers of chitine; mt, the matrix of
the hair; t, tube formed by the inferior layer
of the cuticle (ch. 5), and filled with plasm,
pls; pl, fold formed by the tube (t) at the
level of the first layer of the cuticle; t.r.s.,
inferior part of the basal thickening of the
fold; t.r.a., its superior part; r, central part
of the radix of the hair; ec, papilla; 0, orifice
of the root by which the plasm passes from
the cavity of the tube into the cavity of the
hair, x; N, the nerve; p, the stalk of the
hair; b, the annular thickening of the supe-
rior layer of the cuticle surrounding the root
of the thread.

dowed with extreme sensibility, as is
manifest from the fact that the lightest
filament of silk can at once be detected
by them and communicated to the ani-
mal. The other types are simpler in
their structure and, perhaps, their func-
tion. Dahl does not make any distinc-
tion between the hairs of the different
types, and names them all auditory; but
Wagner distinguishes the hairs into three

principal types, the Tactile hair, including one of finer structure (poil tactil
fin), the Beaded hair (poil a chapelet), and the Clubshaped hair (poil

cucurbitiforme).

The principles that led Dahl to attribute to his auditory hairs this
function rests alone upon the fact that the waves of sound set them into
motion, which movement is borne along the extremity of the nerves ound

provokes the sensation of sound.

He appears to attribute the same function to all the types of hairs

distinguished by Wagner.

In this opinion the latter author cannot agree,

wry ‘ a

‘THE SENSES AND THEIR ORGANS. 311

but thinks that the functions of these three types are not identical, since
being found upon the same individual, one cannot well admit that three
different organs are constructed for the same physiological role. Might
they not, however, serve for different degrees of the same function ?

Wagner does not doubt that spiders have a delicate sense of hearing,
but the objective ground on which he rests it is unreliable. He states
it as “a fact known to all biologists,” that in order to entice a spider
from its nest or den it is only necessary to cause a fly to buzz near it,
while an unskillful imitation of the
buzzing sound fails to deceive the ara-
nead. On the contrary, even the pres-
ence of flies in the web often fails to
tempt the spider forth; and I know
that unskillful imitations of insects have
often drawn them forth; but such imi-
tations I have never conffned to sounds.
They are only or chiefly successful when
the movement is communicated to the
threads of the web itself.

Wagner admits that if the movement
of hairs of any type under the influence
of sound could be proved, that would
suffice to assign to that type the role of
an auxiliary auditory organ, at least.

But, in fact, ib is far from being proved! "Satin of « Pete tal, Me mrs
that sound sets the so called auditory marked by the same letters. z, an eminence
hairs into movement. At least his OW ee ne aap ero oe or ae ea anc ans

ing of the fold by which the free borders are

experiments failed to show this. By the bent against the stalk of the hair, and form a
‘ d fe ] ee l bl little external pouch which is situated in front
al or an ¢ ectric lantern he Was able to of the large external pouch; bo.e., cavity of the
throw distinctly upon a screen figures of large external pouch; r, root of the hair with
‘ . its thickening; p.t.r., section of the walls of the

a row of auditory hairs, enlarged to the external pouch by which the inferior part of
size of from three to six inches He had the basal thickening of the fold (t.r.s.) is set
" J : below; b, wall of the large external pouch, cor-

prepared a fine section of that portion of responding to the annular elevation of the first

the foot which is provided with auditory ¥°F of chitine in @ Tactile hair, b, Fig, 204
hairs, and this had been so placed as to allow free movement. Sounds of
various, sorts and tones were then produced without having any effect upon
the shadows on the screen.

Suspecting that his lack of success might be due to the dryness of his
preparation, he replaced it with a foot freshly cut from a living spider.
The figures of the hairs were thrown clearly upon the screen, and, again,
every effort to cause them to move by the aid of sound waves was unsuc-
cessful. The size of the hairs upon the screen was so considerable that
the least vibration would be perceived, and, therefore, he could hardly ad-
mit that the failure was the result of defect in his experiment. From the

pls,

312

AMERICAN SPIDERS AND THEIR

SPINNINGWORK.

above facts he concluded, first, that the function of the three types of hair
above described cannot be recognized as identical; and, second, that no one
of these types can be regarded as an auditory

Fic. 296. Section of a fine Tactile hair.
p, the stalk of a hair; r, the superior
thickening of the root in the cavity of
the large external pouch ; bo.e., cavity
of the large external pouch; ry, basal
thickening of the root at the point of
its junction with the basal fold; z, ele-
vation upon the superior part of the
basal thickening of the fold by which
the free borders are bent against the
stalk of the hair and form a little ex-
ternal pouch; p.t.r., section of the
walls of the internal pouch, by which
the inferior part of the basal thicken-
ing of the fold (t.r.s.) is inserted be-
neath; b, wall of the large external
pouch, corresponding to the annular
elevation of the first layer of chitine
in the Tactile hair. The other parts
corresponding to those of the Tactile

organ.

But, if their function is not identical, their
fundamental likeness in anatomical structure
gives one a right to suppose that they do have

a function more or less analogous.
does that function consist ?

Wherein
Mr. Wagner con-

siders that the function of the Tactile hairs
ought to be more perfect than that of the

other types, because
of their more per-
fect structure, and
that, therefore, they
should be capable
of receiving more
delicate excitations
than the ordinary
Tactile hairs. One
use, he thinks, may
have been indicated
by Dahl, who re-
marks that the
slightest breath of
air is able to move
the auditory hairs, a

hair are marked by the same letters. Ho. 2by¥-—Sectlon sof thaigntiotn

fact which may be
readily observed ; at least, that spiders are always
extremely sensitive to the slightest puff of wind
made by the human mouth. There may, therefore,
be assigned to the auditory hairs the function of
transmitting the mechanical movements of the air,

Wagner again raises the conjecture that the
Beaded and Clubshaped hairs may be used to in-
dicate the state of the weather; a conjecture which
he bases upon what he supposes to be an ac-
cepted fact, namely, that spiders are so sensitive
to weather changes as to be able to anticipate
them, and, indeed, to prognosticate them by their
behavior. hat this is a widespread belief I else-
where indicate (in the chapter on General Habits) ;

Clubshaped hair. m, the club-
shaped blade of the hair; st, lon-
gitudinal striations of the blade;
z, elevation upon the superior
part of the basal thickening of
the fold by which the free borders
are bent against the stalk of the
hair and form a little external
pouch; r, root of the hair, with
its thickening; p.t.r., section of
walls of the internal pouch by
which the inferior part of the
basal thickening of the fold (t.r.s.)
is inserted beneath; b, walls of
the large external pouch, corre-
sponding to the annular eleva-
tion of the first layer of chitine
in the Tactile hair. The other
parts corresponding to those of
the Tactile hair are marked by
the same letters.

but that it is without foundation, I think, I also show; so that Mr. Wagner's
suggestion must fall to the ground before the presence of facts of habit,

Oe ee ——— << TLC

THE SENSES AND THEIR ORGANS. 313

My own opinion is that all these various types of hairs may be re-
garded, generally speaking, as Tactile hairs, and that they serve to com-
municate to the spider the sensations which are included by more highly
organized animals in the distinct senses of touch and hearing, and, I
might add, of smell.

It seems to me that there can be nothing contrary to this view in the
fact of differences in the forms of hairs, if we suppose that the several
types may indicate some differentiation in the character of touch sensa-
tions communicated by them, so that a spider may be able to distinguish
between the agitations of the air caused by ordinary movements of the
wind and the impressions of waves of sound, and those sensations which
result from touch proper, as the undulatory motion of surfaces on which
a spider rests, or the agitation of the web upon which it hangs and the
trapping thread to which it holds as it lurks within its den. In other
words, there are differences in the sensations produced by the organs of
touch, but these have not been so far differentiated as to justify us in
distinguishing any of them as organs of hearing.

Mr. Wagner calls attention to facts which may lead up, after wider
study, to important conclusions. He says that the Orbweavers (Epeiride)
and Lineweavers (Therididee), for example, only possess these hairs upon
the tibia and metatarsus; while the Wanderers have them not only more
numerously on the tibia and metatarsus, but also upon the tarsus. We
perhaps may not accept Mr. Wagner’s opinion that the Wanderers are ex-
posed to far greater dangers than the Sedentaries, but certainly there is a
difference in the form in which the dangers approach them, as well as in
the character of the dangers. The greater number of Tactile hairs on the
legs of Wanderers may perhaps be associated with the fact that they do
not rest upon a web, but come in contact with the ground and the vari-
ous surfaces on which they lurk for prey. Their feet also are used, at
least in some cases, for digging holes in the earth and for other uses which
are not habitual to Orbweavers and Lineweavers. Moreover, Sedentary spi-
ders, hanging on their webs by their feet, need a concentration of sense
organs in the neighborhood of the claws or tips of the tarsus; and it
seems to me that the Sedentaries are well provided in this respect, and
are thus able to detect the slightest motion that runs along the lines of
their snares when agitated by insects or by raiding enemies. However,
we must confess that here we are largely in the region of conjecture, but
the manner of life among Wanderers, one would suppose, naturally re-
quires a better physical organization, inasmuch as they are not provided
with the habit which constructs trapping instruments for the accession of
prey and the defense of their persons. In other words, it may be that the
presence of additional sensation hairs upon the Wanderers is a compensa-
tion for the lack of industrial facilities.

Mr. Wagner has also found some interesting facts concerning the

—=_ = Te

314 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

development of these Tactile hairs: Immediately after hatching from the
eggs, Attus terebratus has none of these organs upon its tarsus or metatar-
sus, and only one upon the tibia. Lycosa saccata when first hatched has
not a single Tactile hair. After the second moult, however, both these
species acquire one hair upon the tarsus, two upon the metatarsus, and
two upon the tibia. Lycosa saccata when adult has four hairs upon the
tarsus, nine of such hairs on the metatarsus, and seven on the tibia. This
would seem to indicate that with the development of the spider, and thus
with the approach of need for sensation organs, Nature causes those organs
to appear. The young spider has no need of food, as it subsists upon the
nourishment provided by the mother in the egg. It is not until after its
first moult or two that Nature requipes it to set up housekeeping for itself,
and capture its own prey. This fg oe of the Sedentaries. The Wander-
ers, at least some of them, live with the mother until the first moult has
been made.

Are spiders mute? The question is one of much interest, whether con-
sidered from the standpoint of the relation between the sexes, or
Are the number and nature of the senses. The amount of informa-
Spiders : : ; ] oye
Mute? tion possessed upon this subject is scarcely sufficient to warrant
a decided opinion, but such as I have will be presented.

At the outset, it may be suggested that, reasoning from analogy, we
would expect to find in spiders some mode of stridulation. The subking-
dom of Arthropoda, to which they belong, has at its head the Insecta,
among which are many genera whose species are characterized by their
power to stridulate. In illustration of this, any frequenter of our fields
and forests will recall the rolling drumming of the harvest fly or cicada,
which may be heard in yast and confusing notes when the seventeen year
locust, as it is popularly called (Cicada septendecim), makes one of its
periodical appearances, and covers the trees with hosts of insects. The
cheerful creaking of “the cricket on the hearth,’ which has passed into
our proverbs and poetry, is an example of stridulation. The shrilling of
the grasshopper, locust, and field cricket are other well known examples.
Professor Wood-Mason has discovered stridulating organs in the Phasmidee.!
These were seen in a species of Pterinoxylus, the stridulating organs being
fixed partly on the wings and partly on the tegmina, like the Orthopterous
Edipoda described by Seudder.?

In these cases it has commonly been regarded, and is probably true,
that the stridulating instruments are exclusively possessed by the males,
and that the sound is in some way intended as a call to his mate. This

1 Proceedings London Entomological Society, 1877, page xxix.
2 American Naturalist, Vol. II, page 113.

EE Te ee

| a

THE SENSES AND, THEIR ORGANS. 315

fact has long been known, as is evident from the old witticism attributed
to the incorrigible Rhodian sensualist Xenarchus, who alludes in
o Male * 5 the following terms to the cause of that great happiness which
Love Call. : é ,
was popularly attributed to these insects, and which seemed to
the common folk to make them apt images of the gods :—

“lappy the Cicadas’ lives,
Since they all have voiceless wives! ” !

The fact here noted is probably true of insects generally, as in most
cases females cannot utter sounds, and stridulating organs are limited to
males. Yet there are some exceptions which add perplexity as
well as interest to the problem. For example, the stridulating
organs possessed by the Phasmide above alluded to are, accord-
ing to Professor Wood-Mason, found in the females, thus furnishing a case
in which functional stridulating organs are present with that sex. Another
example of power to stridulate on the part of female insects is that of
Cicada montana?

Passing to the other extreme of the Arthropods, we find examples of
stridulating among Crustaceans and Scorpions. Mr. Darwin, alluding to
stridulation among spiders as recorded by Professor Westring, makes the
remark that this is the first case known to him, in the ascending scale of
the animal kingdom, of sounds emitted for this purpose.* But we are
now able to embrace other Arthropods among the music making animals
belonging to the lower orders. ;

Mr. J. Sackville Kent discovered sound producing properties in a Crus-
tacean, a species of Spheroma. He was not able to ascertain the exact
method in which the sound was produced, nor whether the ani-

Hixcep-
tions.

rad mal has organs specially adapted for the purpose. On numerous
moe ee occasions, however, he heard the sound made by this little creat-

ceans. Ure, a Crustacean scarcely one-fourth of an inch long. The ani-

mal was confined within a glass jar, of which it was the only
occupant, and the noise made was a little sharp tapping sound, produced
three or four times consecutively, with intervals of about one second’s
duration.

The observer could almost exactly imitate it by striking the side of
the jar with the pointed end of a pipette. The character and intensity
of the sound produced, associated with the small size of the animal, in-
duced him to believe that it was caused by the sudden flexion and ex-
tension of the creature’s body.*

1 See Cowan’s Curious History of Insects, page 250.

2 See Trans. Lond. Soc., 1877, pagé xvi.

8 Descent of Man, Vol. II., American Edition, Chapter IX., page 330.

4 Nature, November Ist, 1877, page 11. See also Proe. Lond. Ent. Soe., 1877, page xxvii.

— ee ee eee eT

316 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

On this communication Professor Wood-Mason remarks that the sound
producing organs in Crustacea are paire organs, as in Scorpions, Mygale,
and Phasmide; that is to say, organs working independently of
Stridulat- each other on each side of the body. They are differently seated
ing AP- oy situated in various genera, but in all cases appear to consist
paratus. : 5
of what may be called a scraper and a rasp, and the sound is
produced by rubbing together these two organs, which constitute the strid-
ulating apparatus.

Professor Mason has also announced the discovery of stridulating organs
in Scorpions. This appeared from the study of the anatomy, but the matter

was placed beyond doubt by observations made at Bombay. Two
Scorpions Jarge living scorpions, procured from Hindustani conjurors, were
hiatag fixed face to face on a light metal table and goaded into fury.

At once they commenced to beat the air with their palps, and
simultaneously to emit sounds which were distinctly audible, not only to the
observer, but also to bystanders. They were heard above the flutter made
by the animals in their efforts to get free, and resembled the noise pro-
duced by continuously scraping with one’s fingers bits of silk fabric or a
stiff tooth brush. The stridulating apparatus in this species is developed
on each side of the body; the scraper is situated upon the flat outer face
of the basal joint of the palp fingers, the rasp on the equally flat and pro-
duced inner face of the corresponding joint of the first pair of legs.!

It is thus found that from one extreme of the Arthropods, the Insecta,
where stridulation is frequent, through the Scorpions, and to the opposite
extreme, the Crustaceans, the habit of producing sounds, for whatever pur-
pose, is to be found. We -therefore have a strong basis in analogy for the
belief that similar organs might be found among the spiders, animals that
rank between these extremes.

XI.

In point of fact, such organs have been found. The Swedish nat-
uralist Westring was the first to discover them, and his observations are
accessible to the general reader in his valuable work upon Swedish
Spiders.? He appends this observation to his description of “Theridion
serratipes.” The abdomen of the male, around the cord by which it is
united to the thorax, is armed with a denticulated coat, whose use West-

ring had often puzzled over. At length he fortunately discoy-
West- . ered that this valve is an instrument for stridulation. At the
mine's base of the thorax the aranead is armed with t
covery. P#se of the thorax the aranead is armed with transverse, most
delicately wrinkled striations, which are applied by the animal
for the producing of sound, as among insects. This sound Westring heard
when the spider was squeezed slightly ; then, either freely or when touched

' Proc. Ent. Soc, Lond., 1877, xviii. 2 Aranew Syecicee, page 175.

coal LS ee em

THE SENSES AND THEIR ORGANS. 317

with the fingers near the apex of the abdomen, he moved his abdomen up
and down, and its base or the serrated valve near the base of the thorax
was rubbed upon. The female ‘of the species does not possess these or-
gans.!
Mr. F. Maule Campbell? has taken up these observations of Westring,
and in a valuable and interesting paper added much to our information.
He made special studies, both of the male and female of Stea-
Stridula- toda guttata and Steatoda bipunctata. In the fore extremity of
er een the abdomen he found, in the male of Guttata, that the socket
is a complete ring with some strong chitinous spurs on the in-
side of its external edge (Fig. 299), which is also roughly serrated. ‘That
of the female is divided into two parts, the inferior being smallest, while
the superior, as in the male, is the deepest. In the female (Fig. 298)
there are no spurs. The inner edge, however, is undulated, and in points
becomes angulated, while a little below are stiff hairs on small protuber-
ances. The chitinous thoracic extension of the male is marked on its

Fic. 298. Z Fig. 299.

The stridulating organs of Steatoda guttata, male and female. (After Campbell.)

Fic. 298. Female; view from above, of chitinous ring or socket attached to abdo-
men, covering the union with thorax. Fic, 299. The same part of the male.
Fia. 800. View from above of chitinous extension of thorax of female.

superior surface with many fine, parallel, transverse grooves, which are
absent in the female. (Fig. 300.) In the same position on both sexes are
several ridges which are less numerous in the male. Thus, it appears
that while it is likely that individuals may vary in details, the female of
Steatoda guttata has organs adapted to stridulation, as well as the male.
Mr. Campbell also examined both sexes of Steatoda bipunctata, and
found that the socket of the male is much shallower than those of the
male and female of Guttata. The inside of the external edge is rough,
and the sides are lined with a row of bristles seated on prominences. The
only opposing surface is a spinate ridge on the base of the thorax, which

1 Since this original discovery Westring had heard many males of Theridium and Stea-
toda stridulating. Among these he mentions Theridium castaneum Clerck, Theridium (Stea-
toda) bipunctatum (page 185), Theridium hammatum, Theridium albumaculatum (page 186),
Theridium (Steatodos) guttatum (page 188).

2On Supposed Stridulating Organs of Steatoda guttata and Linyphia tenebricola, Linn,

. Soc. Jour, Zool., Vol. XV., 1880, page 152.

318 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

has no chitinous extension covering the abdominal union. In the female
of this species, unlike Guttata, there is no trace of these organs.
Of course, any sounds which might be produced by the organs thus
described, must be occasioned by the flexion and extension of the two
principal parts of the body; that is to say, by the drawing back
Boe and forward, within the socket, of the cartilaginous pedicle which
connects the thorax with the abdomen. But Mr. Campbell has
‘also discovered an apparatus which he ventures to call stridu-
lating, seated on the faleces and palps in both sexes of Linyphia tene-
brosus.' These are of a different structure from those
heretofore described. On the outer side of the basal
joint of each falx are about twenty parallel transverse
chitinous bands, placed so that their inferior edges are
free. (Fig. 301.) The effect, when viewed from the
front, is that each falx has a distinctly serrated outer
edge, which becomes more and more decided towards

Fic, 301. FG, 302. the base
Fic. 301. Right falx of Lin- s < 5 Pome!
ppliet tenebrtania amiales The opposing surface is that of the humeral joint

viewed obliquely fromthe of each palpus (Fig. 302), which is marked with a

right side. Fic. 302. Hu- :

meral joint of left palp, More or less regular series of curved grooves deep

showing spines on tre enough to give the appearance of serration on its side

top the horny plate. (Af under a two-third objective. On the under side of this

es Bee joint, close to its base, is a curved enlargement, and on
the top a prominent, horny, somewhat triangular, knob like plate, with a
rounded apex. This differs in form, size, elevation, and position from the
chitinous prominences usually seen in connection with spines, of which
there is one near each side, but of which, in some individuals, it is inde-
pendent.

The above described organs persist in all adult members of this species;
but those on the palps of females are not so highly developed, the chief
difference being the size of the enlargement at the base of the third joint.
When confined in a glass tube, Mr. Campbell observed that these spiders
often move their palps backward and forward, with a slight rotary motion,
in such a manner that the horny plate crosses the bands on the falces.
But he had been unable, even with the aid of the microphone, to detect
sounds in connection with these movements.

Mr. Campbell adds the suggestion that the absence of specialized stridu-
lating organs in most Aranez does not imply that they are mute. It is a
common practice with many to rub the falces against the maxille; and
were the serrated edge of these latter found in another part of the body,
similarly opposed to a hard, toothed, chitinous surface, it is most likely

1 Linyphia terricola Blkw., or Linyphia tennuis Blkw. 2 Op. cit., page 155.

THE SENSES AND THEIR ORGANS. 319

Another account of organs of stridulation in spiders is that observed
by Mr. 8. E. Peal in the great stridulating Mygale of Assam, and brought
to notice by Prof. James Wood-Mason.! Mr. Peal’s account is
eee that the noise is made by the tarantula when in a state of great
Take’ excitement, particularly at the presence of some enemy. When
thus roused, the spider usually rested on the four posterior legs,
raising the other four and shaking them in the air, with the thorax
thrown up almost at right angles to the abdomen, and the palps in rapid
motion. The noise made is both peculiar and loud. It resembles that
made by pouring out small shot upon a plate from a height of a few
inches, or perhaps by drawing the back of a knife along the edge of a
strong comb. ‘The stridulation was very distinct, and had a ring about
it which the observer had never noticed in the stridulation of orthopterous
insects, wherein it more closely resembles a whistling sound,

Professor Wood-Mason, who reported Mr. Peal’s statement to the London
Entomological Society,? remarks that the sound apparatus in Mygale stridu-
lans has been found to consist, first, of a comb composed of a number of
highly elastic and indurated, globe shaped, chitinous rods, arranged close
together on the inner face of the basal joint of the palp; and, second, of
a scraper formed by an irregular row of sharp erect spines on the outer
surface of the penultimate joint of the palps. He further states that it
is equally developed in both sexes, the first specimen met with by Mr.
Peal having been a gigantic female.

In the spiders alluded to by Westring, the stridulating apparatus con-
sists, as we have seen, of a serrated ridge at the base of the abdomen,
against which the hard hinder part of the thorax is rubbed, and of this
structure not a trace could be detected in the females.

Professor Mason agrees with Mr. Darwin and Professor Westring in
feeling almost sure that the stridulation made by these spiders serves as
a call to the female. It is manifest, however, that if the sound serve this
purpose in the Mygale, it must serve as a mutual call, the apparatus being

present in both sexes. Professor Wood-Mason further ventures
eis so the suggestion that the sounds are emitted by the spider in self
lation, ‘efense; that is, to render itself terrible in the eyes of its ene-

mies; or, it may be from fear. He thinks that they may. also be
serviceable to the spider in terrifying its prey; and, further, that during its
nocturnal rambles in quest of food, it may warn the creatures that it
preys upon of its dangerous and deadly nature, as, for example, is the
case with the rattles upon the tail of our American rattlesnake.

Of course, the presence of stridulating organs, if they be regarded as
sound producing organs, naturally infers the presence of auditory organs.

1 Proc. Asiatic Soe. Beng., 1876, and Ann. Mag. Nat. Hist., 1876.
2 Transactions, 1877, page 282.

i a i i i Bi

320 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Via, 3083 (upper figure). A Tarantula rampant, just before striking.
Fic, 304 (lower figure). Tarantula in act of striking.

THE SENSES AND THEIR ORGANS. 321

If we suppose that the species of spiders in which the male is provided
with stridulating apparatus, possesses it for utility in courtship, and uses
it for love calls, as is generally thought is the case with orthopterous in-
sects, then we must also suppose that auditory organs exist, at least in the
female spider. In other words, spiders are not deaf; they can hear. It
is true that distinct auditory organs have never been found, at least have
never been recognized as such, and, if spiders hear, they must hear by
means of sense organs widely different from any possessed by animals that
have the power of hearing.

If, further, we suppose that those species wherein stridulating organs are
possessed in common by male and female, use them to make mutual calls,
like the notes of birds or like the sounds uttered by higher vertebrate an-
imals, then we are also to infer the power of hearing in the male as well
as in the female.

-This much, at least, appears reasonably certain, that the theory that
the organs above described are proper organs of stridulation, whose pur-
pose is to produce sounds that will be heard by the opposite sex, is de-
pendent upon the demonstration of the fact that spiders possess organs of
hearing. If we are able to affirm the presence of auditory organs in spi-
ders, we may then conclude that the way at least is open for the theory
that stridulating organs are common for mutual communication by sound
between the sexes. Until this be established, the theory-rests upon a very
uncertain foundation.

Concerning the observations upon the stridulating Mygale described by
Mr. Peal, and announced by Professor Wood-Mason, I would remark that
I have kept for many years in succession living species of both
males and females of the large Mygalide. One of these (Eury-
pelma hentzii) I had in my possession for a period of nearly six
years, and one living at this date has been with me about five years. I
have often seen them assume the attitude described by Mr. Peal. When
I have tested their appetite for small vertebrates by putting mice into their
artificial home, or have given them large insects, as locusts, or when I
have teased them with a pencil, or annoyed them in any way, it is their
invariable habit to throw themselyes into the rampant position which
Mr. Peal has described and illustrated. This position I have frequently
sketched from various points of view, and from some of these sketches
Figs. 303 and 304 have been engraved. But in all these cases I have
never heard any other sound than that which I regarded as the clatter-
ing of the fangs as they were struck together in the movements of the
mandibles under the powerful influence of hunger or fear. No sound that
T could at all regard from any other standpoint has it ever been my op-
portunity to detect. Such negative evidence, of course, amounts to little—
amounts to nothing, indeed—in the face of positive testimony. I only
state it as serving to qualify any conclusions which we may be disposed

My Mute
Myegalidez

322 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

to make as to the actual cause and intention of the sounds which some-
times are heard to issue from spiders in a condition of excitement.

The method of the tarantula in. attacking its prey is similar to that
of Lycosids and other spiders. It throws itself upon the four hind legs,

draws back its cephalothorax to a greater or less degree, accord-
How Ta- ing to the nearness of its adversary, raises the two front pairs
Penne of legs and the palps, and, holding them well together, throws

trikes. :

them backward, opens wide the tremendous fangs and the man-
dibles, which are held straight out from the face, and then at the proper
moment launches itself forward (Fig. 304), striking its adversary with its
fore claws and fangs. The stroke will be repeated a number of times
with great rapidity, and after each stroke the tarantula falls back into the
rampant position above described. (Fig. 303.)

I am hardly able to give serious credence to Professor Mason’s theory
that the stridulating organs, like the rattles upon a rattlesnake’s tail, are
intended to give warning to victims. Even if we were to suppose that
the large insects and other creatures fed upon by these spiders are able
to detect such sounds and recognize their meaning, I cannot think, in the
face of ‘my long continued observations of living species in confinement,
and the few observations made in the state of Nature, that they do utter
sounds sufficiently distinct to cause anything like terror on the part of
intended victims. I never saw an insect fed to my tarantulas that showed
the least sign of fear or even consciousness of the presence of an enemy.

OP APT HR: x1.

COLOR AND THE COLOR SENSE.

Tue popular impression that spiders are extremely ugly is deeply
seated. Even specialists in other branches of natural history are apt to ex-
press surprise when one speaks of high ornamentation among araneads.
Butterflies are commonly thought to have special claims to beauty, and
without disputing these one may truthfully say that as fair and brilliant
colors may be found among the Aranew as among the Lepidoptera. I
suppose the popular impression to the contrary is largely due to the fact
that the spiders which frequent our cellars and outhouses, and straggle
occasionally inside our homes, belong to the genera whose colors are
rather inconspicuous. Possibly, contact with human beings has tended to
demoralize these species, and thus disrobe them of colors which once may
have made them attractive!

One does not need to go to the tropics for examples of richly colored

spiders. Our indigenous Orbweavers furnish species whose coloring may

well challenge the admiration of lovers of the beautiful, This

ecciteag will be abundantly illustrated by the plates prepared for Volume

Colors. III. of this work, but several -examples are presented in this

volume, as those on Plates I. and IV. Our two indigenous spe-

cies of Argiope have bright colors, Cophinaria being at once distinguished

by her size and prominent, black, yellow, and brown markings, and Argy-

raspis adding to these a metallic white which in earlier stages of her life
has a noticeable lustre.

Epeira insularis is well known among familiars of our fields by her
attractive yellow and orange colors; and the varied and beautiful robing
of the Shamrock spider is well illustrated by the specimens presented in
Plate I.

Yet these are far excelled in beauty and brilliancy by the Orchard
spider, and the remarkable aranead, Argiope argenteola (Plate IV., Fig. 6),
which is found in the southwestern portions of the United States, The
genus Acrosoma also presents several species whose attractive coloring makes
them worthy of notice in this connection, and Gasteracantha (Plate IV.,
Fig. 8) is often well decorated.

(323)

—=— Ys VT e)|0Ul

324 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Many Theridioids also bear beautiful and delicate colors, the varied
hues and shades of green, yellow, and brown being particularly noticeable
in this group. Some of the genera, as the parasitic species of
Argyrodes, are covered with burnished silver. Of these two
tribes of Sedentary spiders it may be affirmed that they contain,
in all portions of the world, and particularly in tropical countries, exam-
ples of as delicate and brilliant coloring as may be found elsewhere in
Nature.

But the coloring of Orbweavers and Lineweayers is probably even ex-
ceeded by that of the Saltigrades, which is as rich as that of humming-
birds or beetles, according to Professor and Mrs. Peckham. The most bril-
liant family of this tribe, the Attide, especially, contains examples of
brilliant ornamentation. Of some of these araneads Wallace says that they
are noticeable for their immense numbers, variety, and beauty. They fre-
quent foliage and flowers, run about actively in pursuit of small insects,
and many of them are so exquisitely colored as to resemble jewels rather
than spiders.'” Elsewhere he speaks of the abundance and variety of the
little jumping spiders which abound on flowers and foliage, and are often
perfect gems of beauty.? Most travelers in South America who haye care-
fully observed aranead life, agree with Bates that the number of
spiders ornamented with showy colors is remarkable.* Professor
Peckham makes the strong assertion, which my own experience
confirms, that a large collection of spiders from the tropics is almost
certain to contain as great a proportion of beautifully colored specimens
as would be found among an equal number of birds from the same re-
gion.

Some of the Laterigrades also are richly colored. We have several
species in the neighborhood of Philadelphia that would attract the admi-
ration of any observer. The yellow and brown markings, varied with red
and purple, which characterize the familiar Misumena vatia (Plate IIL,
Fig. 1), may often be observed in the midst of wild flowers of our fields.
A small species, apparently of Philodromus, which I am not able to iden-
tify, is remarkable for its pleasant grass green hues, with markings of
bright red and brown upon the legs and palps. The most brilliant color-
ing appears to be confined to these four tribes, namely, Orbweavers and
Lineweavers among the Sedentaries, and Saltigrades and Laterigrades among
the Wanderers. Tubeweavers and Tunnelweavers among the Sedentaries,
and Citigrades among the Wanderers, are, for the most part, distinguished
by dull and inconspicuous coloring, though it is highly probable that a
wider knowledge of the species of these three tribes will uncover many
decorated species.

Beautiful
Spiders.

Attoid
Jewels.

' Tropical Nature, page 97. * Malay Archipelago, page 437.
® Naturalist on the Amazons, Vol. I., page 106. * Sexual Selection in Spiders, page 10.

= ee
j
ad ‘TS =

COLOR AND THE COLOR SENSE. oul

Among Orbweavers and Lineweavers there appears to be a preponder-
ance of yellow hues, and the metallic species of these tribes are generally
marked by a metallic white or silver. The Saltigrades have a
tendency to somewhat darker colors, the reds and browns being
more generally prevalent in this tribe; and where metallic colors
occur they are usually metallic green, or occasionally blue. Yellows, greens,
and dark browns prevail in the Thomisoids. Uniform browns, grays, blacks,
and lead colored or neutral tints are most common among Tubeweavers,
Tunnelweavers, and Citigrades.

It will thus be seen that spiders present a sufficient number and variety
of facts in coloration to occupy the attention of naturalists. It is to be
regretted that these facts have not been so systematized and presented in
connection with the habits, industry, and structure of the species as to
enable one to consider them with accuracy and satisfaction in their bear-
ings upon many problems that now occupy the thought of scientific ob-
servers. Nevertheless, something may be attempted; and even the imperfect
contributions of this chapter may, in the future and in other hands, be
found helpful.

How shall we account for this variety of coloration? And what under-
lying causes have influenced the special colors of particular species? In
point of fact, color appears to belong to the natural constitu-
tion of the spider, being imparted to it at its birth, and pre-
served through life by the power of heredity. It seems to be
an accident or incident of physiological changes which have not been ac-
counted for; and as such it can hardly be considered to have special
regard to utility in one direction or another. To quote the language
of Mr. Wallace, ‘Color per se may be considered normal and needs no
accounting for. Amid the constant variations of animals and plants it
is ever tending to vary, and to appear when it is absent,” 4

No doubt it is modified by food, habit, environment, variations of heat,
cold, moisture, light, and darkness; but the strong hereditary tendency
by which it is controlled is dominant, even amidst the abnormal influ-
ences: which sometimes more or less modify it. Nevertheless, it may be
worth while to attempt to present some of the facts in habit, environ-
ment, and structure which seem to be most closely related to the colors
and color changes of spiders.

Some of the most remarkable and perplexing facts in aranead colora-

tion are seen in Epeira trifolium, and these have been repre-
Color Va-sented in Plate I., Volume II., wherein several variously hued
rieties of . . ; ; z Saye se
Trifolium, 8Pecimens of this species are given, colored from the individ-

uals themselves, as they were collected from one field in Niantic,
Connecticut. The locality is described at length in Volume L., page 292,

Metallic
Hues.

Color is
Normal.

1“ Essay on Colowrs of Animals.”

326 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

to which the reader is referred. The specimens were all near neighbors,
exposed to the same influences of habitat, food, sunlight, ete., and most
of them were taken from bushes of the same plant.

They were all domiciled in nests of clustered leaves or of single leaves
rolled and sewed together. Let us examine some of these specimens, all
of which are females, as we pluck them from their homes, and note their
colors. Beneath this nest is a spider whose feet are black and whose
legs are white, ringed with black at the feet and around the joints. The
body too is white, with only here and there faint black lines bringing
out more distinctly the trifolium markings. In the next bush is another,
differing from the first only in the fact that the annuli of the legs are
brown instead of black. Here is another (Plate I., No. 1), pale yellow
on the abdomen, deepening into orange towards the spinnerets under-
neath the body. The trifoil markings on the abdomen are very faint,
indeed, scarcely distinguishable. The legs are a pale, transparent yellow,
with red brown rings at the joints. Another specimen (No, 2) resembles
No. 1, except that the front of the abdomen is orange below and greenish
yellow at the top, the face being light brown. This spider is drawn in
the position which it usually assumes when sitting in its nest, or when
it rests upon a branch, with the knees bunched up against the abdomen.
Still another specimen (No. 3) is dark yellow brown on the dorsum of
the abdomen, growing into a deep chocolate at the sides and wunder-
neath; the trifoil patterns and spots on the abdomen are chalk white.
The legs are orange with brown. The next specimen (No. 4) is drawn
as viewed from underneath, the abdomen and sides showing there dark
orange, with crimson stripes through the centre and yellow hues along
the sides. The legs have deep orange rings on transparent pale yellow.

Still another (No. 5) is colored yellow, the top of the abdomen deep-
ening into yellow brown along the sides and beneath, and has the out-
lines of the trifolium spots distinctly marked and of a pale yellow. Two
short, greenish, longitudinal bars mark the tip of the abdomen, The legs
are pale yellow with brown rings.

In the next specimen (No. 6) the trifoil spots are yellow on a green-
ish yellow abdomen, the latter deepening to orange on the sides and be-

neath. The legs have dull brown rings. Yet another specimen
Straw- is of a bright strawberry tint, the abdominal patterns being a
nueas bright yellow, the legs yellow with red brown rings. It is a beau-
tiful object, certainly, as it lies bunched up in the palm of one’s
hand, and no one looking upon it could deny that spiders are sometimes
attractively clothed. Still another specimen (No. 9) has the dorsum of
the abdomen orange, which deepens to crimson red below and at
The . é wae
Male. the.sides, and has light yellow trifoils and spots. The legs are
white, with dark brown rings at the joints. A male, Figs. 10 and
11, which we find in the nest of one of the females, is colored yellow, the

oe) a “ei add a UF

COLOR AND THE COLOR SENSE. 327

legs and cephalothorax having brownish rings and bands, and the abdo-
men being a lighter yellow with brownish spots.

Thus the colors run, with varieties of tints and hues that confound
the observer. Most of these spiders appear to be of one age and at the
same period of gestation. Those that are least advanced, perhaps, may be
said to have the white colors. The next degree of maturity in mother-
hood shows the yellow tints. ‘The next the deepening brown, and so as
the creature ages the colors seem to deepen and brighten. When the last
stage of maturity has been reached, and the spider mother has spun her
beautiful silken cocoon, depositing therein her eggs neatly and securely
blanketed against assaulting enemies and winter frosts, these colors will
gradually merge into the dull, dark hues of the sere and yellow leaf of
which her nest is built, and so her life will fade away.

The physiological causes of this change in the colors of Trifolium
present an interesting study. Other species known to me are subject to
changes. In some the change is quite marked. In some there is a great
variety of coloring, and particularly of dorsal patterns, as in the case of °
Epeira patagiata and Epeira parvula, but the Shamrock spider exceeds all
species which I have ever observed in the puzzling variety and contrasts,
as well as beauty, of the colors it assumes in the closing weeks of its life.

The color of young spiders is almost without exception light yellow or
green, whitish or livid, tints that blend well with the prevailing greens of
foliage and young twigs, and the grays of bark on trees, of rocks
Color De- and soil. This is probably due largely to the fact that the tis-
pee sues are at that time translucent. The effect may also be caused
Young. Py the absence of acquired food in the alimentary tract and lack
of distribution throughout the system of other than the prena-

tal nutriment.

As young spiders advance in age the color deepens, which is caused, no
doubt, by gradual hardening of the tissues, thus making them more opaque.
Up to this period no food has been taken, hence the absence of food alone
is not sufficient to account for the lighter colors of the first stages after
exode. Yellows and browns in various tints occur at this period, and
in some cases—though not generally, I believe—color patterns which are
characteristic of the various species in adult life begin to appear with
more or less distinctness, or at least suggestively. It is not until Seden-
tary spiderlings have established themselves upon their own webs, and, so
to speak, have set wp housekeeping for themselves, that the characteristic
colors and markings of the species begin to appear with positive degrees of
distinctness.

The Attide, like birds, moult frequently, and at each moult the mark-
ings may change, so that some of the older writers have formed several

~~ a

328 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

species for the different moults of one. These difficulties are increased by
the fact that the adult males and females of a species usually differ con-
siderably in appearance. ! '

This is not entirely in accord with the statement of Mr. Cambridge that
the pattern of a spider—that is, the design formed by its colors and mark-
ings—differs in general but little in immaturity and maturity, excepting
that it is usually more distinct in the young and in the female exam-
ples. The first of these statements appears to me to be too sweeping,
unless the period of youth referred to be placed well on toward maturity.
In some species there are striking differences between the colors and mark-
ings of the very young spider and those which it attains after one or
two changes of skin. Epeira diademata and Zilla x-notata, two of the
commonest English spiders, are conspicuous examples of this.”

According to Peckham, the young spiders often differ from adults, and
in many species when the sexes differ when adult, the male being brighter,
they are alike until they reach maturity, when the male, along
with his sexual development, acquires his brilliant color. Again,
soon after hatching, young spiders, probably at the third or
fourth moult, begin to show color more decidedly, and the colors are dis-
tributed in the patterns characteristic of the species, and as the spiders
continue to advance in age and make their successive moults, other and
more marked changes may be noted. The truth appears to be that there
are differences among species in the degrees of resemblance between imma-
ture and adult forms, but that generally the likeness strengthens from the
time of hatching onward to maturity.

I give a few observations upon the appearance of spiderlings during
and shortly after their cocoon life. These, however, can hardly be fully
appreciated by those who do not know the adult species, without consult-
ing the plates in Volume III. But the following species may be com-
pared with figures or descriptions in this volume.

Just after its escape from the egg shell the young of Argiope coph-
inaria is about two millimetres long. The cephalothorax is a grayish

white color, translucent, upon the fore part of which the eyes,

Color
and Sex.

Sauer which are a brownish color, stand out vividly, seeming to form
Se hee aa large part of the face. The legs are white, translucent, as
Young. ; ‘ “ ‘

are also the palps; as the spider sits upon a surface both legs
and palps are doubled under the body. In this position the palps seem
to be a shorter pair of legs, so that as thus viewed the animal really
seems to have ten legs. The abdomen is a yellowish color, except that
in the places where the peculiar yellowish irregular marks of the dorsum
are seen upon the adult, may be seen irregular markings of pure white.

* North American Spiders of the Family Attidee, page 5. * Spiders of Dorset, xxvi.
* Peckhams, “Sexual Selection,” pages 14, 15.

COLOR AND THE COLOR SENSE. 329

Fig. 305 is drawn from a young Cophinaria just out of the shell, and
Figs. 306 and 307 from the same a few days older.

When the young Cophinarias have advanced in age a few days the fo-
lium upon the dorsum of the abdomen assumes a distinct shield shaped
outline resembling that which is common upon the adult forms
of Epeira insularis, sclopetaria, etc., the color of the same being
a darkish green, and the scalloped margins being surrounded by a white
band which extends quite around the fore part of the abdomen. The
sides also have a greenish band, the same color prevailing around the
spinnerets. The usual aspect of the abdomen is thus green in the centre
and lower part of the dorsum, and white along the fore part of the abdo-
men and the sides. The hairs are quite prominent both on the abdomen
and legs. The eyes have a darker hue, and little processes on either side
of the base of the abdomen distinctly appear. The abdomen is now in
general shape a miniature of the adult form. ‘The legs are covered with
greenish bands closely placed. When separated from the mass of its fel-
low broodlings, a single spider will throw out
a thread from which it will hang down, sus-
pending itself by its dragline and weaving a
little foot basket, precisely in the manner of
the more matured spiders. Three longitudinal
bands appear upon the cephalothorax, one in

Argiope.

i R ;, pr Fic. 305. Fic. 306. Fig. 307.
the median line and one on each side. The jaye, 395,

youngling looks plump, as though well nour- fter leaving the shell. Fics. 306,

Young Argiope cophinaria

r 807. Appearance after first moult.
ished.

The young of Epeira strix shortly after its advent from the cocoon
(April 14th) often presents a uniform glossy black appearance. After an-
other moult this appearance is somewhat changed, the legs have
black annuli around the joints, and the interspaces are of a yel-
lowish brown hue well covered with black spines. The folium
upon the abdomen is along its margins jet black, with a median cross like
figure of a dark yellowish brown. Bands of the same color surround the
scalloped margin along the sides. The cephalothorax has the same gen-
eral hue of glossy black.

At the time of hatching, the young Gasteracanthas of Africa, accord-
ing to Dr. Vinson, are round and black, without the pointed spines pecul-
iar to the adult, and with a triangular white spot upon the abdomen.
These peculiarities are also characteristic of our California species. Among
those sent to me by Mrs. Eigenmann were a number of young in various
stages of growth. They are all quite black, and the spines are either lack-
ing or just beginning to push out slight angles upon the otherwise rounded
abdomen. (Compare with Plate [V., Volume II., Fig. 8.)

The black color of these young Gasteracanthas is a singular variation
from the ordinary color of spiderlings, which is quite light, the colors

Epeira
strix.

= ae

—— a

330 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

being nearly always white, or a faint livid, or a delicate hue of pink or
yellow. I do not know whether the absence of spines characterizes the
young of those species that show these peculiarities in adult life.
Abdom- Tt may be that the development of these thorn like processes is
ae in some way connected with the development and growth of
’ the young spider, and is only completed at maturity. It would

be interesting to know the physiological causes of this vital phenomenon.

Some of those species which have soft conical tubercles upon the fore
part of the abdomen show these very early in the young. I have observed
them distinctly formed upon at least two of the Angulata group of our
American Epeira, gemma and bicentennaria. In the case of Argiope coph-
inaria the spiderling immediately after escape from the egg (the first moult)
appears to be without the processes or bifurcations which mark the base
of the abdomen of that species, but after the next moult these show plainly.
(Compare Fig. 305 with Figs. 306 and 307.)

When the young of Tegenaria medicinalis first break from the shell,
the legs and palps are white and semitransparent. The eyes stand out
brown and distinct upon the face. The cephalothorax in the
fore part has a slight bluish or lead colored tint, with a touch
of yellow at the posterior part near the abdomen. The mandibles are the
color of the cephalothorax, but with the fangs prominent, feeble looking,
whitish, instead of the dark, horny appearance of the adult. The abdo-
men is a uniform yellowish hue, at the apex of which the spinnerets ap-
pear lead colored, the long, jointed pair quite prominent. The spines are
quite manifest on the legs, and hairs are seen on the abdomen. ‘The fo-
lium or dorsal figure can be traced, together with the transverse bars, on
either side of the median line. In a day or two the color of the legs
deepens until they have a leaden hue, upon which the black spines stand
out more prominently. The abdomen is a little brighter yellow, and the
cephalothorax corresponds in color with the legs. In two days more the
yellowish tint has faded from the abdomen, the whole spider has a black-
ish appearance, caused by the dark hairs upon the lead colored body; the
transverse markings stand out more prominently upon the abdomen.

The young of Epeira cucurbitina (English) when extracted from the
egg have the cephalothorax and legs of a pale yellowish white color, that
of the abdomen being reddish brown. But after their first change of in-
tegument they acquire an olive or brownish green tint, the upper part of
the abdomen being metallic with whitish spots on each side, with a longi-
tudinal stripe of the same hue parallel with it. On the upper side there
is a series of minute black spots.' | These examples will be ample, when
compared with adult forms, to enable the student to note the color changes
that occur during the growth of spiders.

Tegenaria

1 Spiders of Great Britain and Ireland, page 3438.

COLOR AND THE COLOR SENSE. 381

TI.

As spiders further advance in age and make their successive moults,
various color changes may be noted. Immediately after moulting the
color is always lighter, which is probably due to the fact that
ae the harder skin just cast off prevented the passage of light
ancen: through the tissues. The new skin is thinner and more trans-
lucent. Moulting produces changes in color patterns of a de-

cided kind, at least in certain species.

Phidippus rufus when mature is a dark red spider, the male consider-
ably brighter than his consort. When about one-seventh grown and after
the third or fourth moult, the young are dark brown with light yellow
legs. Some moults later they are reddish, with narrow, oblique, whitish
bars on the sides of the abdomen, and two dark bands on the dorsum, on
each of which is a row of white dots. The appearance of the spider
changes but little during the next four moults, but after the last, the
tenth, both male and female become mature, and acquire the adult color.
The appearance of the female after the fifth moult is similar to that of
many other females in the genus.!

The female of Phidippus johnsonii has the abdomen red and_ black
with a white base and some white dots, while the male abdomen is bright
vermilion red, with sometimes a white band at the base. The young of
both sexes resemble the mother, until the last moult, when the males as-
sume their bright livery.?

In old age the color changes are often quite decided. In some, as Epeira
trifolium and Epeira thaddeus, the changes give added brilliancy to the

color at certain parts of the body. Some of the color changes
ue a of Trifolium are remarkably beautiful, and the same is true of

Thaddeus. But advanced age, as a rule, brings darker colors.
Orange and brown then have a ruddier hue; yellows darken into orange
and brown. Sometimes the yellow patterns are entirely lost, and the spi-
der becomes dark, almost black. There is a grizzled appearance about the
animal in this stage which reminds one of vertebrate animals at the cor-
responding period. These last named changes are manifest in the female
spider after the final deposit of eggs.

In gravid females changes of color are sometimes noticeable. Some of
the bright colors upon Trifolium and Thaddeus are doubtless due to this
condition. However, other and perhaps most species during ges-
tation have a lighter color, which may be the result of mechan-
ical changes in structure. The skin becomes distended and more transpa-
rent, the pigment is thereby distributed, and thus centres of color are
broken up and the coloring matter diffused. Not only the skin, but other

Gestation

! Peckhams, Sexual Selection, page 25. 2 Idem, page 17.

— ll

> — pe TE ee

Os
v

332 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

parts of the abdomen are distended during gestation, and this distension
produces changes in the color of the animal, it may be by modifying in
some way the various secretions from the liver and other organs, and in
some cases, perhaps, widening the intervals between color centres and color
hairs, and breaking up groups of the same.

The little pits or dark spots upon the dorsum of the abdomen, which
mark the attachment of the muscles within, seem to me to be centres for

the aggregation of coloring material. At least the dorsal pat-
Effects of terns appear to be grouped in some regular way around these
rer muscular attachments. Thus the action of the muscles on the

skin and chitinous shell or walls seems to compel certain aggre-
gations along the lines of use that form these colors and patterns. It.
might be important in this connection to consider what is the ordinary
effect of muscular action upon the distribution of pigment and colored
hairs in vertebrate animals.

The color rings or annuli around the joints of the legs of spiders may
be influenced by action of the muscles. The tendency of these darker and
more vivid colors is towards the ends of the joints, as though by the at-
tachments and prevailing outward action of the muscles the pigment were
forced mechanically or otherwise attracted toward these points. The foot
or terminal joint is usually dark, and often black. In the. cephalothorax
may be noted the same tendency of color to group itself somewhat sym-
metrically around the points of muscular attachment, particularly the cen-
tral depression.

IV.

Color and markings are undoubtedly influenced by sex. Peckham, after
summarizing the Attide of France from the studies of M. Simon, finds
that in thirty-nine species the male is plainly unlike the female,
Influence }eing in twenty-six instances much more conspicuous, while in
ee se °” fifty-five the sexes are similar, or, if they differ, the male is no
more conspicuous than the female. These facts make it clear
that the sexes commonly differ, the male being brighter than the female.
Peckham considers it not too much to say that in the Attidee at least
two-fifths of all the species have the male more conspicuous than the
female. !

Menge, in referring to the greater brilliancy of the male of Micromata
ornata, says that it only assumes its bright color as a bridal adornment,
and in this connection makes a statement that in the Thomiside and Sal-
tigrade the males are generally more beautifully colored than the females.?

Phileus militaris, a common American Attoid, is another illustration.
In the male the cephalothorax and abdomen are bright bronze brown, the

1 Sexual Selection, page 20. * Menge, Preussische Spinnen, II., page 396.

COLOR AND THE COLOR SENSE. 330

former with a wide, pure white band on each upper side and a white spot
on the centre of the head, the latter with a wide white band around the
base and sides. The female has the brown all covered over with white and
gray hairs, which form a more or less distinct pattern of lines and spots.!

In Habrocestum splendens, while the young males are not exactly like
the adult female, they resemble her much more closely than they do the
adult male. This is one of our most beautiful male Saltigrades. The
highly iridescent scales which cover the entire body make it impossible
to give in a painting a correct idea of its brilliancy, since the color changes
in eyery light. The male only gets his gorgeous livery at the last moult,
just as he becomes mature, though in some species the nuptial moult is
acquired one moult before maturity.”

This prevalent condition of the relative brilliancy of coloring between
the sexes of the Attidee is entirely reversed among Orbweavers. In this
tribe there is a strong tendency to inconspicuous colors in males, and fre-
quently in the degree that the females are conspicuous for size and color-
ing, the males are diminutive and dull.

We have already seen (see Chapter II., page 60), from our examination
of the interesting studies of the Peckhams upon the courtship of Salti-

grade spiders, that there is a close relation between mating

oat habits and the brilliant colors prevalent among males. In other
on- F ae x
schae words, the favors desired from the female are solicited with such

Wasa? a display of the ornamented parts of the male body, as to jus-

tify the conclusion that the ornamentation is pleasing to the fe-
male, and is presented in the way of soliciting her favors. Of course, if
we accept this fact, we also admit that there must be, on the part of both
sexes, a consciousness of the presence of color, and the fact that the female
at least is so sensitive to the differences in color ornamentation as to be
moved towards this wooer or that according to the splendor of his physi-
cal finery.

That climate and favorable environment sometimes exert modifying in-
fluences upon the general facies and, to some extent, the industry of spi-
ders, is illustrated by Epeira labyrinthea. I have specimens of
this species from almost every part of the United States where
collections of araneads have been made. It is distributed from
the far Northeast to the southwestern portion of California. I have also
received specimens in collections forwarded to me by Professor Peckham
from several South American States. These southern representatives of the
species are larger and decidedly more vigorous looking animals than the
northern specimens. he industry of the spider experiences no essential
change as far as I can learn. ‘The snare is, perhaps, larger with tropical
examples. Specimens of cocoons sent to me from southern California have

Climatic
Influence.

1 Peckham, Sexual Selection, page 17. 2 Id., page 18.

334 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

all the characteristics of our northern species, but are decidedly larger.
No marked influence appears to have been exerted upon the coloration of
the spider itself. But other collections indicate contrary conditions,

The species was collected pretty freely by the naturalists of the U. 8. 8.
“Albatross,” in its explorations along the shores of the southern Atlantic

and the Pacific, showing an immense distribution over the en-
Epeira tire Western Hemisphere. Dr. Marx remarks that specimens col-
ee lected at the Straits of Magellan are hardly recognizable, for its

color has greatly changed. The dorsal folium is nearly obliter-
ated. Only two lighter spots at each side remain, and the whole body is
covered with a long, dense pubescence. !

If Dr. Marx is correct in his surmise that Epeira cooksonii Butler,
which lives in great abundance on the Gallapagos Islands, is related to our
Epeira domiciliorum, and that the latter spider has undergone on the Pa-
cific coast so great a change in form and coloration as to be identical
with my Epeira vertebrata, this species will afford another example of the
influence of climate upon color. I am not satisfied, however, that the
last two named species are identical. On the other hand, certain species,
as notably Argiope cophinaria and argyraspis, have undergone a transcon-
tinental distribution, covering wide extremes of climate and conditions,
without experiencing any notable change in general appearance.

These examples will be sufficient as illustrations of the fact that the
influence of climate must vary according to species. The fact is that some
of the species probably are more elastic and impressionable in their nat-
ural constitution, and thus are more sensitive to radical changes in envi-
ronment, while others are able to resist such changes more vigorously, and
thus retain their characteristics through extreme changes.

Spiders that live upon plants, as a rule have colors that are har-
monious with the prevailing greens and yellows, and admixtures thereof,

of branches, leaves, and flowers. Spiders that nest in stables,
Influence houses, on fences, and like locations, ordinarily have dusky
pera colors, harmonious with the environment; as, for example, The-

ridium tepidariorum, Agalena nevia, Tegenaria medicinalis.
However, I do not find that any great difference in color is observable
in the above species when they nest in foliage, as is often the case, at
least with Agalena and Theridium. It might be said, perhaps, that there
is a slight tendency to darker hues and a more uniform color when the
spiders are found in the first named locations.

Ground spiders, as the Lycosids, generally have colors of neutral grays
that blend well either with the soil, with rocks, or with stalks of grass
and weeds, especially when the latter are somewhat dry. Lycosids found
in the neighborhood of streams do not seem to be especially influenced by

* Proc, U. 8. National Mus., Smithsonian Institn., Vol. XII, page 209. * Td., page 210.

? *
—_—S es cs — Oe

COLOR AND THE COLOR SENSE. 335

the natural color of water; but Dolomedes sexpunctatus, which is so con-
stantly found on the water, sometimes has a tint that at least well harmo-
nizes with that of the stream itself.
Saltigrades follow the rule of the Lycosids; their colors, being chiefly
black, gray, and brown, harmonize with the surfaces of rocks, trunks of
trees, etc., upon which they habitually seek their prey. Many
Mimetic of them are freely marked with yellow, and thus are also suffi-
ae ciently harmonized with the color of the leaves. The metallic
monies. .
green and blue on the fangs of some Saltigrades seem almost
like a leaf ambush to the body of the creature as it is observed stalking
its prey. This suggests the strategy most familiar from its association
with the lines of Shakespeare :—

“Macbeth shall never vanquished be, until
Great Birnam wood to the Dunsinane hill
Shall come against him.”

It is, perhaps, a not wholly untenable theory that some insects are
made less wary by the resemblance to surrounding foliage and the play of
iridescent hues from the mandibles of a stalking Phidippus morsitans,
for example, as it stealthily moves upon its prey. But independent of
the indifference of the ordinary insect to spider presence, the Peckhams
have taught us to find the chief service of these gorgeous frontlets in
courtship. But what can be their use in the female Morsitans? She is
such a ferocious virago that we might suspect in her an example of warn-
ing coloration as towards her own lovers.

According to Emerton,' in one species of Linyphia from Weyer’s Caye,
Virginia, the colors and markings of some specimens are as bright as on

spiders of the same family living in cellars and shady woods.
ae of ‘The other five species are pale in color. On the supposition
Spiders. that all these species drifted from the outside world into the

caverns, we must reason from such a fact either, that the species
retaining a normal color had been domesticated in the caverns at a much
more recent date than the others, or that it was possessed of greater power
to resist the changes consequent upon its changed environment.

The influence of cave life upon Anthrobia mammouthia appears to be
manifest in this lack of color. Two young Anthrobias were hatched May
3d for Professor Packard, who describes the whole body, including the
legs, as snow white, with the legs much shorter than in the adult state.
The ‘adult in life is white, tinged with a very faint flesh color, with the
abdomen reddish. In some specimens the abdomen has beneath several
large transverse dusky bands. Linyphia subterranea as observed living

“Notes on Spiders from Caves in Kentucky, Virginia, and Indiana,” American Nat-
uralist, Vol. [X., page 278.

0 SS

ESSN
: 7 “- a

336 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

in Wyandotte Cave is pale pinkish, horn brown on the thorax and legs,
while the abdomen is dull honey yellow.! \
Two specimens of Linyphia weyerii in my possession, collected by Dr.
Joseph Leidy in Luray Cavern, Virginia, are of a light honey yellow, the
abdomen of one individual being darkish brown. As this is a common
color for spiders of all conditions, after they have been in alcohol a little
while, I am not able to say what may have been the original color of
these, particularly as they are not adult.
In the case of Lycosa arenicola, whose habits were studied by me on
Coffin’s Beach, near Annisquam, Massachusetts, the undue presence of sun-
light and heat appears to have produced precisely the same
Pe eee results as the absence of sunlight in Mammoth Cave. The
light. eastern shore of the bay opposite Annisquam consists in part
of a stretch of sand hills known as Coffin’s Beach. The sand is
a bright white color, and is massed at places into elevations of consider-
able height. The fragrant Bay bush grows in clumps along the edges
and summits of these irregular sand hills, and this is intermingled with
patches of tough grass, among which are numerous burrows of the Turret
spider. These Lycosids are domiciled in the sand, and spread very gen-
erally over the dunes. The burrows are dug straight downward, penetrat-
ing the upper layer of loose sand, and striking the more compact and
moist strata below the surface. The spiders captured were of a light hue,
as compared with the same specimens found
in meadows, fields, and like environment in
the interior. Specimens almost identical with
these in color were found by Dr. Joseph
Leidy, and subsequently by myself, in the
sand at Beach Haven, New Jersey. This
pale coloring appears in all other littoral
specimens examined. The influence of envi-
ronment, as manifest in these spiders; was also
Fic. 808. Lycosa arenicola. (A dark seen in a. grasshopper or locust which is
specimen.) ; .
quite abundant on Coffin’s Beach, and is al-
most as white as the sand over which it was found hopping.?
It is certainly confusing to one who studies the influence of light upon
aranead coloration to find such contradictory facts as these, viz., that the
absence of light results in albinism in the spider fauna of cay-
Contra- erns, while the excessive sunlight which beats upon and is re-
dictory ; it ‘
Effects, {lected by the white sea sand produces the same condition. Evi-
dently something more than the direct influence of sunlight must

* McCook: The Turret Spider on Coffin’s Beach, Proc. Acad. Nat. Sei., Phila., 1888,
page 333.

COLOR AND THE COLOR SENSE. Bad

But confusing elements do not stop here, Any one who is familiar with
our ordinary cellar spider fauna must have observed among them a strong
tendency to black or dark colors. The most common cellar spider in the
neighborhood of Philadelphia is probably the Tubeweaver Tegenaria me-
dicinalis. It is a quite dark lead color, which is as near black as any of
our indigenous spiders. So also Agalena nevia, when it nests in dark
places, as it often does, seems to me to add to the somewhat sombre colors
which it bears in arboreal and sunny sites, several additional tints of
darkness.

It should be ‘noted that in the case of the Turret spider of Coffin’s
Beach a large portion of the year must be spent in a torpid condition
by such individuals as survive the winter, which is severe and long in
that vicinity, On the other hand, the white spiders of Mammoth Cave
live in a uniform and pleasantly warm temperature. The same, to some ex-
tent, is true of the Medicinal spider, which is kept reasonably warm when
living in our cellars and other unfrequented places in human habitations,
but, unlike the Mammoth Caye Anthrobias, are black. Thus similar con-
ditions of temperature, as well as of light, appear in these cases to issue
in opposite conditions of color.

I have already considered the theory of Peckham that the bright orna-
mentation of the male Saltigrade spider serves to attract the attention of
the female, and to secure him her favor in preference to duller

Color colored rivals. In the case of Orbweavers, where the conditions
Utility : : ;
Mating, ®° reversed and the female is the more highly ornamented, we

may suppose that the point of utility is also reversed, and the
brighter colors of the female serve to attract to her the attention of the
male. In the case of Orbweavers and Lineweavers, there would seem to be
some necessity for this, inasmuch as the females for the most part occupy
the centre of their webs, which are often of considerable size, and thus
they would require to be marked in such wise that the vagrant male on
his courtship excursions could discern his mate at the distance of at least
several inches. It seems not an overstrained supposition that both the in-
creased size and excessive coloration of the female would make her a
more conspicuous object and thus facilitate the union of the two sexes.
Again, in a large number of species which are influenced by what we
might call mimetic harmony, and to which allusion has already been
made, we can readily see how highly useful the general resem-
blance of environment would be in protecting spiders from vari-
ous enemies. All the Wanderers, and some of the Sedentary
spiders, appear to be subject to those influences which harmonize their color
with the surroundings of their daily life. This subject may be more fully
considered in the chapter on Mimicry, but it has been necessary to allude

Mimetic
Harmony

— = =

a_i Ae

338 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

to it in this connection. Here, perhaps, we have a view of the greatest
influence exercised by color and, as one may say, the absence of color, upon
the life of araneads.

When we come to consider the more highly colored species, particularly
in their relation to their habits, the question of utility is complicated by
many apparently contradictory facts. Such large and well marked species
as those of the genus Argiope, Acrosoma, Gasteracantha, and such brilliant
species as the Orchard spider, are found well nigh invariably hanging at
the centre of their webs in full view of all enemies and assailants. One
who is pursued by the theory of utility can scarcely fail to ask whether
these species are protected by their color from any enemies, and, if so, in
what manner are they protected? It is certainly natural to suppose that
they are more exposed thereby to raiding birds, digger and mud dauber
wasps.

Another question may be mooted, have these species, thus highly col-
ored, and thereby exposed to enemies, any industrial protection which

may be considered a compensation? I refer to a few exam-
Industrial ples bearing upon this inquiry. Argiope cophinaria and Ar-
ee. gyraspis sometimes have protective wings of retitelarian lines

thrown out on each side of their nets, which guard the dorsal
parts of their bodies, and a thick shield like sheeting, which protects the
under side. (See Volume I., Figs. 96 and 89.) These spiders are highly
colored and conspicuous by size. They dwell in shrubs, bushes, grasses,
low trees, and commonly are stationed in the centre of their round webs,
having no domicile or tent to which they retire. No doubt, the pro-
tective wings are serviceable in warding off attacks of raiding wasps, as
well as helping to secure insect food for the occupant. But I find that in
a large number of cases these fenders are omitted. The tendency to omit
them, if I am not mistaken, strongly increases as the spiders mature in
age. Perhaps in this case the size of the animal may be considered as
sufficient protection. At all events, I think that the protective wings are
rather characteristic of the immature state.

The Insular and Shamrock spiders, which are among the most brightly
colored of our fauna, do not hang habitually in the centre of their webs
during the day, but live in leafy tents, and their habitat is among bushes,
shrubs, and trees. The Insular spider inclines to groves much more strongly
than Trifolium. Epeira thaddeus has the same habit. Indeed, it may be
asserted generally that many of our most highly colored Epeiroids belong
to tentmaking species, a fact which is true not only of the American, but
of the European, spider fauna. The facts here seem to point to a special
development of industrial protection as a compensation for the exposure of
bright colors.

On the other hand, Epeira strix, which is not a bright colored spider
by any means, is one of the most secretive Orbweavers in its habits, dwelling

=~ : ee

el ee

COLOR AND THE COLOR SENSE. 339

in a domicile of rolled leaves, shrinking away into cavities and holes of
bark, in angles of walls, and like positions, occupying at times a well made
‘ leafy tent, after the manner of Insularis and Trifolium, and only
Seeming occupying its snare during the night. It is one of the rarest
eae things to find a Furrow spider, unless it be quite young, sus-
pended upon its orb, trapping flies, in daytime. Notwithstand-
ing its seeming protective color, and the additional protection of its secret-
ive habits, it is mercilessly pursued, in the vicinity of Philadelphia, by the
steel blue wasp, which stores numbers in its clay cells.

Epeira domiciliorum and cinerea are also spiders of rather inconspicuous
colors, the last named particularly so, and both of them screen themselves
in tents, though the Domicile spider at least not as habitually as some
others.

The Labyrinth spider and Epeira triaranea are among the most
strongly protected of our fauna by their industry, having, besides their

orbs and thick retitelarian
Domicile, snare, a dome shaped silk-

ei en tent as a domicile. In
rinth, atti RAE IR BEE
Orobara addition, Labyrinthea roofs

Spiders. her tent with a dry leaf, or

uses it as a shelter for her
body. These spiders are strongly
marked as to their patterns, and are
not inconspicuously colored, but do
not have the bright hues which characterize Argiope, Epeira insulatis, and
some others.

The Orchard spider is one of the most brilliantly colored of our indig-
enous species, although its hues harmonize well, particularly its green and
yellow, with its leafy surroundings. It rests beneath its horizontal orb,
where it is abundantly exposed to attack from above, but has straggling,
pyramidal retitelarian lines beneath it, which form some protection. It
dwells mostly in wooded places, or among shrubs and thick, leafy bushes.
Epeira gibberosa is also a brilliantly colored spider. Its industrial protec-
tion is more manifest, for it dwells beneath a sort of hammock or structure
of lines woven between the edges of a leaf. This hammock protects it
above, while the leaf affords good security from beneath (Volume I., page
154), and its green color completes the protection.

Our three indigenous species of Acrosoma, viz., Rugosa, Spinea, and
Mitrata, are all, particularly the first two, well marked and brightly colored
spiders. They are protected, Mitrata least conspicuously, by spin-
ous processes, if such can be called protections. They live in the
centre of their orbs as a rule, and their webs are most frequently
found stretched between the trunks of young trees, in openings of groves,
woods, and like spots. They seem to me more directly exposed to assault

Fig. 309, Leaf nest of Epeira.

Spinous
Species,

a |, ee

340 AMERICAN SPIDERS AND THEIR SPINNINGWORK,

than almost any other of our native Orbweavers, unless, indeed, their pre-
ferred site within the shaded walks of groves and woods may be consid-
ered a protection.

Gasteracantha, with its strongly developed spines, has much the same
habit as Acrosoma, but the spines appear to be wanting in the young of
this genus, the very age, one would think, at which they are most needed,
However, the young of Gasteracantha, at least with numerous specimens
sent from the Pacific coast, are almost black in color, a feature which must
certainly be regarded as protective, if bright colors best invite the observa-
tion of enemies.

On the whole, the conclusion seems to be justified that many spiders
which appear to be more exposed to enemies by reason of bright colors
or greater size, possess special variations in industry and habits that in
some degree are protective; but there are so many apparent exceptions to
this, which require more careful study, that no generalization can now be
warranted,

If we come finally to consider the bearing of what has been called
“warning coloration” upon spiders, there is little to be said. In the case
of those numerous species which belong to the Wanderers, the
colors cannot be considered as warning, but, as we have seen, are
rather protective. As to Sedentary species, I cannot think of any
animal that would avoid them as inedible on account of their color, or of
any influence that their color could have in warning insects of danger.
In point of fact, the colored spiders appear to be as delicate morsels to
those that feed upon them as any other sort, and they are just as fre-
quently, and perhaps I may say even more frequently, though by no
means exclusively, selected for such purposes.

That a warning should be conveyed to insects by the color of a spider
at the centre of its web seems to me wholly imaginary, since Sedentary
spiders do not capture food directly, but by means of their manufactured
trapping instruments. Indeed, I have little faith in the opinion that in-
sects are capable of experiencing anything like a warning, from color or

other causes, against the presence of spiders. I feel sure that
Insects this is the case with flies, the insects which perhaps more than

Warning
Colors.

Uncon- i ;
scious of @2Y other form the staple food of the various aranead tribes. I
Danger. have often been witness of the absolute indifference of various

species of flies to the presence of spiders. One remarkable ex-
ample is recorded (Vol. I, page 256), in which, during an entire season,
I observed numbers of a little black Diptera settling and feeding upon
the carcasses of large blue bottle and house flies which had been trapped
and trussed within the orbs of Argiope. In several cases these little creat-
ures were observed stationed within the open jaws of their gigantic enemy,
sipping juices which the spider was expressing from the fly on which she
was feeding.

COLOR AND THE COLOR SENSE. 341

A fly which had been put into a box with Epeira strix tempted her
in vain to make a breakfast upon it. Three times it flew into or against
the spider’s jaws and escaped. Twice it crept between the front pair of
legs; once it lit upon and crept up the hindermost legs; and all the time
was walking everywhere around her, utterly unconscious of the presence
of an enemy. The spider remained motionless, except when the fly flew
into her face, when she made an effort to seize it. At the time she was
seated upon the bottom of the box, separate from any snare or web, and
thus without ordinary means by which the presence and locality of in-
sects are determined. We may suppose that the spider was confused by
the unusual circumstance of separation from her web, and her vision
momentarily impaired; but the fly, at least, was in normal condition,
hunting food and otherwise acting in a natural way.

In numerous other cases when flies haye been placed within boxes
where spiders have been confined, I do not remember a single individual
that showed the slightest sense of fear, but on the contrary they would
run all around, and even settle upon the spider, apparently no more con-
scious of its presence or of any peril therefrom, than if it had been a
clod or chip. The same is true of grasshoppers, hundreds of which have
been fed to the large tarantulas that from time to time, during a number
of years, I have kept in confinement. Of such insects, at least, it would
be highly absurd to argue anything of service in the way of “ warning
coloration.” Bright or dull, large or small, they seem to be absolutely
without Consciousness of the presence, or fear of the power, of spiders.

In taking a summary view of the facts above recorded it may be said,
in general terms, that the influences which appear to modify the color of
spiders, in various degrees more or less known, are the follow-
ing: Moulting changes; the effects of advancing age and ap-
proaching dissolution; the disturbance of gestation; the distri-
bution of pigment and color hairs by muscular action; the effects of
food, environment, and general habit; sexual differences and the excite-
ments of courtship and mating; and, possibly, inimical influences, such
as natural enemies and weather changes.

General
Summary

VI.

How far are spiders conscious of the color elements in their surround-
ings? They are found among leaves, flowers, and blossoms of all the
: varied kinds and colors in the vegetable kingdom. Sedentary
Conscious: syiders hang their webs to the branches and leaves of trees,
ness of ; - :
Coles, and weave them amidst flowers, often selecting for them sites
which strike the observer as choice and notable for beauty.
One may find, for example, the pretty web of Linyphia communis hung
within an opening upon a morning-glory vine, the bright colored flowers

: Cd]
~“—aeen eee ee

342 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of which encircled the web like a charming frame to a picture.! Again,
one may see the round webs of Epeira spun among lilies,? and hung
within full blossoming sprays of honeysuckle. Indeed, at every point in
Nature where flowers appear, there also appear spiders erecting their domi-
ciles, weaving their snares, and spinning their cocoons.

In like manner many of the wandering tribes spend their lives in
arboreal situations, continually stalking their prey, and plying the varied
industries characteristic of their species among grasses, shrubs, blossom-
ing trees, vines, and beds of flowers. Are they utterly unconscious of the
color effects among which they continually move? Or if, on the other
hand, they have some sense of color, in what degree is it possessed ?
These are interesting questions, and to some extent they have been solved,
although much remains to be proved.

It cannot, of course, be known that the light waves of various lengths,
whose vibrations result in color, produce upon the spider’s organ of vision
effects similar to those known to man and many of the higher mammals.
But that some effect is produced seems clear, and that this is analogous
to the color sense in man, we may perhaps safely assume; for we can
only think and speak of the sensations of spiders in terms of our own
conscious states.

The late Prof. Paul Bert claimed that all animals see the rays of the
spectrum as we do; that beyond this they see nothing that is unseen by
us, and that, in the extent of the visible region, the differences
between the illuminating powers of the different color rays are
the same for them as for us.2 He rests these conclusions on
experiments made on a small fresh water crustacean belonging to the
genus Daphnia. Sir John Lubbock dissents from this generalization as
too sweeping and based upon an insufficient foundation,+ but, as the
result of numerous experiments with Daphnia pulex, concludes that while
it would be impossible to prove that these crustaceans actually perceive
colors, to suggest that the rays of various wave lengths produce on their
eyes a different impression other than that of color, is to propose an en-
tirely novel hypothesis. At any rate, he thinks that he has shown that
they do distinguish between waves of different lengths, and prefer those
which to our eyes appear green and yellow.°

On the other hand, M. Merejkowski denies to the crustaceans any
sense of color whatever. He thinks that they distinguish very well the
intensity of the ethereal vibrations, their amplitude, but not their number.
In the mode of their perception of light there is a great difference

On Crus-
taceans.

1Vol. I., page 344, Fig. 335. * Thid., Fig. 104.

’ “Archive de Phisiologie,” 1869, page 547. 4“Ants, Wasps, and Bees,” page 220.

° Sir John Lubbock: “On the Sense of Color among some of the Lower Animals,”
Linnzan Society’s Journal Zoology, Vol. XVII. (1883), page 214. See also “Senses and In-
stincts,” page 228.

COLOR AND THE COLOR SENSE. 343

between the lower crustaceans and men, as well as between those animals
and ants. While we see the different colors and their different intensi-
ties, the inferior crustaceans neither behold any color or the different
variations of intensity therein. We perceive colors as colors, they per-
ceive them only as light.!

Mr. Alfred R. Wallace does not admit that the fact that the lower ani-
mals distinguish what are to us diversities of color, proves that their sen-
sations of color bear any resemblance to ours. The insects’ capacity to
distinguish red from blue may be and probably is due to preceptions of
a totally distinct nature.?

We have much testimony that insects have a decided color sense.
Most important and decisive are, perhaps, the remarkable investigations

of Sir John Lubbock, whose experiments indicate that ants are
Color oa ,
Sense of *Msitive to the ordinary colors of the solar spectrum. It be-
Insects, Comes probable, moreover, that the ultra violet rays must make

themselves apparent to ants as a distinct and separate color, of
which we can form no idea, but as unlike the rest as red is from yellow
or green from violet. He adds, that as few of the colors in Nature are
pure, but almost all arise from the combination of rays of different wave
lengths, and as in such cases a visible resultant would be composed not
only of the rays which we see, but of these and the ultra violet, it would
appear that the colors of objects and the general aspect of Nature must
present to ants a very different appearance from what it does to us.*

Lubbock has also shown that bees have a decided preference between
colors, and that blue is distinctly their favorite, although yellow is much
liked.4 He also demonstrates that wasps are capable of distinguishing
color, although they do not seem to be so much guided by it as bees are.°
The fact having thus been established, that among two classes of the
Arthropods, namely, the Crustacea and the Insecta, there are found genera
which show a decided color sense, prepares us to expect the same fact in
the case of the Arachnida, and indeed of all other Arthropods.

The best sustained and most conclusive experiments upon spiders them-

selves, of which I have knowledge, were made by Professor
Spiders: and Mrs. Peckham in the neighborhood of Milwaukee, Wiscon-
Peck- sin. Their method of procedure was as follows: A cage was
hams’ é
Bixpert- constructed, formed of four differently colored compartments,
ments, ®ll made of glass and opening freely into one another. The

cage was placed on a table on,a covered porch, with the wall
of the house on one side, while the other sides were exposed to light. A

1“Tes Crustacés inférieurs distinguent-ils les couleurs?” (Do the inferior crustaceans
distinguish color?) Par M. Merejkowski. 2 Wallace, “Tropical Nature,” page 238.

8“ Ants, Bees, and Wasps,” page 220, 4 Thid., page 310. 5 Tbid., 316,

6 “Some Observations on the Mental Powers of Spiders,” Journal of Morphology, Vol. I.,
Deceniber, 1887.

i i

344 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

spider was then admitted to the cage and, after having become sufficiently
domesticated, was gently driven into a specially colored compartment, say
the blue. It was then left without interference to select such position
as it might prefer in any one of the four differently colored compartments.
When the spider had changed its position and remained therein a suffi-
cient time to indicate a preference for the color under which it rested, it
was again disturbed and moved to another color. If, for example, it set-
tled within the red compartment, it was transferred to the yellow, and so
on, a record being made of the various changes and preferences. his
process was continued during several days, in which several hundred ex-

l periments were made. As a result it was found that among all
SONHSIS: dia spiders experimented with 181 preferred the red, 32 the yel-
Prefer ;

Red. low, 11 blue, and 13 green. The preference of the spiders for

red was thus decidedly marked, resembling, although in a more
marked degree, the preference of ants for the same color, as demonstrated
by Sir John Lubbock’s experiments,! which appear to have suggested those
of Professor Peckham.

A test case was made which gave a striking result, quite in confirma-
tion of the experiments as above described. An individual of Lycosa
nigroventris, which had shown a strong preference for red; choosing that
compartment 383 times out of 41, was temporarily blinded by covering
its eyes with paraffine. When put within the cage it was found that
the spider remained quiet in whatever compartment it was placed until
it was driven out. If placed in the blue compartment, with its eyes
as close as possible. to the red, it showed no inclination to enter, al-
though this color had before proved so strongly attractive. Its prefer-
ences, or rather its locations, during the resulting experiments, are recorded
as follows :—

Preferences after blinding: Red 6, yellow 6, blue 6, green 5.

Preferences before blinding: Red 33, yellow 5, blue 0, green 3.

Such results leave scarcely any room for doubt that in some way the
spider had been influenced by a color sense, since, while it possessed
normal vision it expressed a most decided preference for the red color,
but when temporarily deprived of vision settled indifferently and about
equally in all the colors represented in the series, there being no stronger
preference for red than there had been in previous experiments for the
blue compartment, which it had entirely shunned. These results seem to
justify the conclusion that there exists a color sense in certain spiders.

It is to be remarked, however, that in all the cases recorded, and ap-
parently in all experimented upon, the individuals were chosen from the
Lycosids alone. These spiders undoubtedly have a keen sense of sight,
although I am inclined to think that in this respect they are inferior to

*“Xnts, Bees, and Wasps,” page 189.

COLOR AND THE COLOR SENSE. 345

some other groups. Their habit keeps them during much of their life
concealed within earth burrows, or little caves excavated and fitted up
by them under stones, logs, and like surroundings. They move over the
ground or water, where they stalk their prey, and are not as apt to be
found in arboreal situations among flowers, blossoms, and leaves, as other
tribes of the Wanderers, the Saltigrades and Laterigrades,

There is one well known species of Laterigrade spider, Misumena vatia,
whose habits have awakened in my mind the query whether it might not
be influenced by a decided color sense in the selection of certain
sites. Most araneologists have observed this species stationed
upon yellow flowers, as the golden rod and the brown daisy
which is popularly known in our section as “black eyed Susan.” Misu-
mena lurks upon this flower with its legs spread out within the very
centre, and so closely corresponding in golor to its floral site that one must
look closely ere he discovers it. The yellow centre of the common ox eyed
daisy is also a lurking place for this spider. I have found the same spe-
cies nestled within the petals of a half opened tea rose, and then its color
also corresponded with its environment, being white, with various delicate
shades of green and pink. (Plate III., Fig. 2.) In these cases we are
forced at least to face the question, was the spider moved in such selec-
tion by the color of the flower? If we say yes, then we are also con-
strained to the conclusion that, in some way, the aranead must have been
conscious of the fact that its peculiar color harmonized with the color of
the flower which it sought as a stalking point for the capture of its prey.

It is doubtless true for the most part that light is perceived by spiders,
and arthropods generally, by the eyes, and not chiefly by the skin. Sir
John Lubbock has shown, by a series of ingenious experiments, that ants
perceive the ultra violet rays with their eyes, and not, as suggested by
Graber, by the skin generally. These experiments haye been repeated and
the conclusions verified by an observer so careful and experienced as Dr.
Auguste Forel.! Nevertheless, it seems to me probable that there is some,
and it may be considerable, perception of light by the skin of spiders.

The abdomen of spiders is included within a soft integument which is
frequently covered heavily with hairs. May it not be that this soft skin
is far more sensitive than the hard chitinous enclosure of the abdomen of
insects? May it not, therefore, be that such a spider as Misumena vatia
is led to settle within those flowers which correspond in color to itself, by
that comfortable feeling which results from the harmony of an individual
with its environment, and which may be caused, for all we know, by the
fact that the yellow rays of the flower are perceived by and agreeable to
the sensitive skin of the spider? In thinking of the power of spiders to
distinguish the various hues, may we not be justified in calling into play

Misume-
na vatia.

1 Lubbock on the Senses of Animals, page 211.

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346 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

this sensitiveness of the entire skin, instead of limiting the perception to
the eyes alone?

There is indeed another theory which may be suggested, namely, that
the color surroundings of the spider, in some manner not now explicable,
so rapidly influence the organism of the creature that a change of color
is produced in harmony with its environment. Can we suppose, in this
case, that the spider possesses the power to influence at will the chromat-
ophores or pigment bodies, so that they may change her color with chang-
ing site?

There is another explanation of the above peculiar habit of Misumena.
Many insects are strongly attracted by yellow colors, and as insects are

the chief food of spiders, it is natural that the familiar resorts
Seeking of insects should be the places most affected by spiders. ‘That
pees insects have such attraction to colors has already been shown,

esorts.

and that they are drawn to yellow colored flowers has been fully
established by Miller in his remarkable volume on Alpine flowers.! ‘This
author gives a table recording the numerous visits of various insects to
flowers of different hues; and a study of the table shows that butterflies,
bees, flies, and gnats, and other insects manifest a strong preference for
yellowish white and for yellow flowers. With such a fact as this in view,
we may, perhaps, conclude that the habit of Thomisus and Misumena to
frequent flowers of the character above described, resolves itself into the
well known instinct of all animals to seek their food in those resorts
where the supply is most abundant and accessible. This explanation does
not, of course, exclude the fact that the spider, in seeking such favorable
site, may be guided by its sense of color, but it reduces it to a subordi-
nate rank.

VII.

Walckenaer ? advanced the idea that the form of the cocoon corresponds
with that of the abdomen of the mother. This is in some measure cor-
f rect, for the abdomens of spiders have most frequently an oval
Relation shape, and this is substantially the shape of the cocoon. But

a fig when one comes to compare the shapes of the abdomens of in-
coons, dividual spiders with the shapes of their cocoons, the excep-

tions are so numerous and decided that no such generalization
can be accepted.

The same author suggested that some correspondence exists between
the color of the cocoon and that of the mother’s abdomen. The facts,
however, at least as far as American spiders are concerned, will not sus-
tain this theory, except in a general way. For example, the colors of
the abdomen of Argiope cophinaria are yellow, black, white, and brown.

1 Miller, Alpen Blumen, page 487. * Aptéres, Vol. I., page 147.

COLOR AND THE COLOR SENSE. 347

The colors of her cocoon are yellow, white, and brown. Argiope argy-
raspis has yellow, black, and silvery white upon her abdomen, Her co-
coon is yellow and white. Argiope argenteola has an abdomen whose
colors are metallic white or silver, yellow, and black. Her cocoon is green
or yellow, or a combination of green and yellow on the outside with a
white tuft within. ‘The Insular spider has for its abdominal colors yellow,
orange, and brown. Her cocoon is a uniform yellow. The Bifid spider
has for its prevailing colors a light greenish hue intermingled with a livid
yellow and a little brown. Her cocoon is a dull green color.

The prevailing colors of Orbweavers’ cocoons may be said to be yellow
and white. Sometimes the yellow shades into green, sometimes into brown.

_. The dark or blackish cocoons, when examined carefully, are
Prevailing found to owe their shade to the compactness of the threads of
Spider p . ma
Colors. Which they are spun and the presence of gum. The above

colors, namely, yellow and white, are the prevailing ones among
Orbweavers themselves. The yellows sometimes shade into green, oftener
into brown, livid, and orange. The white frequently becomes metallic,
having a silver sheen. There is, therefore, some basis for suggesting a
correspondence between the color of a cocoon and that of the spider, or
abdomen of the spider, which spins it. The harmony is more apparent,
as far as my observation extends, among Orbweavers than other araneads;
but there are not enough facts in hand to justify a generalization.

Turning from the Orbweavers to other tribal groups, we find that the
exceptions are so many and striking that they appear at once to wholly
disparage the theory.* Most Citigrades and Tunnelweavers, and many Tube-
weavers, are dark colored, but their cocoons are quite uniformly white.
For example, the well known American tarantula, whose large cocoon is a
white ovoid ball three inches long, has a dark reddish brown and black
coat.

Tegenaria medicinalis is a quite dark, almost black spider, but she
spins a white cocoon. The same is true of Tegenaria persica, whose clus-
tered cocoons are white when originally spun, although the mother
covers them with dirt, and thus soils the appearance. The Speck-
led 'Tubeweaver is a dull creature, yet she spins a beautiful white
cocoon, although she also mars its whiteness by adding extraneous mate-
rial. With many of the Drassids the same rule obtains. Without multi-
plying examples, this may be said fairly to represent the color relations of
American araneads to their cocoonery.

An examination of the colors of European cocoons, as given by Walck-
enaer, Blackwall, or Simon, or by Staveley,! in her tabular arrangement
of cocoons and eggs, will show that the same fact obtains among the spi-
ders of Great Britain and Ireland. We learn from this table that a great

Cocoon
Colors.

1 British Spiders, pages 269-275.

e—

348 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

majority of British cocoons are white; green or greenish, yellow, and yellow
brown being the other colors represented. These colors are distributed
quite indifferently of the maternal colors.

The color of the silk extruded from the spinnerets of spiders of all
tribes in the construction of snares is, with few exceptions, white, some-
times having a steel blue tint, and often a lustre which gives it
the appearance of spun glass. There are some exceptions to this
rule, as, for example, the round web of Nephila is uniformly
woven with yellow silk; and perhaps a wider study of the spinningwork
of araneads will show that there are other exceptions, and perhaps many
of them. :

The differences of color in the spinning silk of araneads appear in the
construction of the cocoon. Many cocoons are composed of white silk, per-
haps it may be, said the majority of them, but others again show some
pretty varieties of color, and in some species several hues of silk will be
used in weaving one cocoon.!

Among Orbweavyers the colors used in cocooning are principally white ;
but one frequently finds yellow in various shades, green, and sometimes
brown. A few Lineweavers make colored cocoons, and among Tubeweavers
may be found a few species whose cocoons are various shades of red, some-
times quite bright. I often find these cocoons in the shape of little plates,
with the convexity upward, attached to bark and stones, and showing a
very dainty appearance, but have not been able to fully identify them with
the species making them.

An examination of the spinning glands of spiders under the micro-
scope will show, in some species at least, as Argiope cophinaria, several
colors represented in the liquid contents. The causes controlling the secre-
tion of these specially colored silks are, of course, physiological, but one
cannot presume to guess even what they may be.

Color of
Silk.

VIII.

The forms in which the coloring materials are arranged constitute the
various patterns or marks that characterize spider species. These patterns
are most varied and prominent upon the dorsum of the abdo-
men, although the venter and the sternum are also sometimes
highly colored. While there is much variety in the arrangement
of these patterns, there is, on the whole, a general tendency to form a
folium or leaf shaped outline, scalloped on the outer margin. This folium
outline appears to have some orderly relation to the little pits or points of
muscular attachment, and, in a general way, may be said to be regulated
by those sections of the abdomen which have commonly been held to

Color
Patterns.

1 See Chapter IV., page 80, and Plate IV., Figs. 3, 4, 5.

COLOR AND THE COLOR SENSE. 349

indicate a segmentation. In other words, some writers are disposed to con-
sider the abdomen as segmented, and, in a rough way, the symmetrical
divisions in the pattern folium may be said to outline the articulations of
the segments.

The cephalothorax is most frequently uniform in its color, commonly
with longitudinal stripes of different shade. It follows, in a general way,
the coloration of the legs, which it resembles in its chitinous
character, the entire fore part of the body having the hardness
which is characteristic of the enclosing walls of insects. The ab-
domen, on the contrary, is enclosed in a soft skin, a fact which
exposes it to injury, and causes its rapid decay after death, one of the
principal difficulties in the way of preserving specimens of spiders. On
the cephalothorax there are often several longitudinal bands, one on each
side, near and indeed quite surrounding the margin, and two drawn from
the suture of the caput backward. ‘There is also frequently a band of
color in the median line from the middle of the eye space backward. The
legs are usually colored like the cephalothorax, and are generally uniform
in hue, except that at the joints there are rings of color usually darker
than the rest of the leg. The feet are nearly always black or blackish.

In order to determine if possible the structural causes producing color
in spiders, I made studies from a number of dissections of various species
chosen with special regard to variety and brillianey of colors.
Among these are Argyroepeira hortorum, Argiope argyraspis, Ar-
giope cophinaria, Argiope argenteola, Epeira insularis, Gastera-
cantha cancer, and Phidippus morsitans.' I do not speak of the results
positively, for the studies are in a field where trained histologists alone
are competent to decide. But I venture to give some indications of what
appeared to me, in the hope that others may follow the path suggested,
and reach positive determinations.

The metallic white upon the cephalothorax of Argiope argyraspis is
produced chiefly by a vast number of white hairs. These are packed
closely one upon another and reflect white light, the combined
reflections forming the metallic appearance of the object. The
metallic hues of the abdomen of Argyraspis are produced in
part by closely thatched white hairs that reflect white light in the manner
of those upon the cephalothorax. The black transverse bands on the ab-
domen are produced by amorphous granules of black pigment just beneath
the skin, which thin out towards the margin of the band, becoming yellow
as they diminish. There appears also to be a diffused yellow stain in the
chitine, and, in addition, white pigment bodies which resemble the chro-
matophores that give the color in frogs and lizards, for example.

Cephalo-
thorax
and Legs.

Causes
of Color.

Metallic
Hues.

‘T had purposed to include in this yolume a plate of colored drawings representing, in
a general way, the indications of these studies, but sundry disappointments and delays have
compelled the postponement of this plate to the third volume.

ee eee ee

aia a ee

350 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

In Argiope cophinaria the metallic color of the cephalothorax is pro-
duced by hairs in the same way as with Argyraspis. ‘These hairs are
wavy; there appear to be two kinds, one flattened, with a wavy
outline, having somewhat the appearance of cotton fibre, which
may, however, be produced by irregular cavities or spaces within the
hairs. Others again, present a similar appearance, but are cylindrical.
On the abdomen of Cophinaria the colors are produced chiefly by pig-
ment granules beneath the epiderm, the chitinous layers of which are
arranged in beautiful undulating lines.

The pretty orange color upon the thigh of Epeira insularis is pro-
duced chiefly by vast numbers of pigment granules lying beneath the
epiderm, the secreted layers of which are arranged in diamond shaped
figures.

Argyroepeira hortorum, the most beautifully colored of our indigenous
spiders, makes a fine object for mounting in order to show colors. The

hairs have little or nothing to do in producing these varied
The hues, which are due to green and yellow pigment granules, and
ig to what appear to be chromatophores. These chromatophores are

white for the most part, though some of them are yellow tinted,
and they yield a strong white reflection, which, it seems to me, is a chief
agent in producing the brilliant silvery white of this aranead.,

In the case of Phidippus morsitans the color of the abdomen is due
to several causes. The black shades with dark green metallic reflection
on the sides are produced chiefly by dark green pigment granules under-
neath the skin, and in part by black hairs. The white spots on the sides
of the dorsum are composed of peculiar white lanceolate hairs laid one
upon another. They are marked by longitudinal striations on the surface,
which give it, under the lens, the appearance of a minute ear of Indian
corn. The little yellow lunettes of color on the dorsum near the apex
appear to be composed of somewhat similar hairs, of nearly the same
shape, but a little more elongated, yellow in color, and these, instead of
longitudinal grooves, have slight feather like projections or papille irregu-
larly distributed over the surface.!

The remarkable metallic green on the mandibles of this species is pro-
duced by a method quite different from any of those above named. ‘The
surface of the mandibles is broken up into a number of rugosities, ar-
ranged, though somewhat irregularly, in arcs of circles. These ridges
appear to act as prisms, refracting the light; and to this evidently is due
the brilliant metallic color which has attracted the attention of all obsery-
ers of the species. Under a microscope the minute lunettes and waves of
green light are readily distinguished; but the natural eye does not separate

Argiope.

' These hairs in the color patches on the abdomen of P. morsitans appear to be of the
type of Mr. Wagner’s clubshaped hairs. (Fig. 297.)

COLOR AND THE COLOR SENSE. 351

the several groups of refracted rays, and perceives them as an unbroken
band of metallic green color.
These cursory examinations appear to suggest that the structural causes
of color in spiders are probably the following: First, color stains diffused
throughout the tissues; second, pigment granules of various hues
Pam, distributed beneath the skin; third, pigment bodies or chromat-
Bao ophores; fourth, the reflection of light from the surfaces of
Causes. thickly overlaid or thatthed hairs; fifth, by hairs of various col-
ors and peculiar forms, in some degree analogous to the scales
of the Lepidoptera; sixth, certain colors, particularly the brilliant metallic
colors, are produced by refraction of light from broken or ridged surfaces
of the epiderm, that appear to act as prisms.
Little attention has been paid to the structural causes of color in spiders,
and scarcely more to the form of the color hairs, and the manner in which
they are grouped and overlaid in order to

form the various color spots and pattern
outlines produced exclusively or in part by
them. The subject might well repay the
careful study of the microscopist, and it

may often be found that these color hairs
will show many varying forms, correspond- yy. s10, Fic. 31. ee ee
Ing with genera or even species. Micaria longipes.

Mr. Emerton says } that the hairs or Fias. 310 and 818. White scales from spots

Z onabdomen. FG. 311. Scale from hind-
“scales” usually found on the Drassidee erhalfofabdomen. F1G.312. Scale from

and Agalenide are feathered.? Each scale, — *°m* of *Pdomen. (After Emerton.)

as far as he had noticed, is uniformly colored. Along the edges
of the red spot in Geotrecha crocata, for example, red and black
scales are mixed, but each scale is either all red or all black.
The scales of Micaria longipes* are either white or brown. The irides-
cence of the abdomen, which is very marked in certain lights, he had seen
on the individual scales. In general form these hairs resemble those which
I have seen on Phidippus:n morsitans.

Color
Scales.

1In a ister to the author. 2 fie New Eng. Dr pasties plate i iii., F ig. 3, e.
8 Td., plate iii, Fig. 1

PART V.-HOSTILE AGENTS: THEIR INFLUENCE.
CHAPT Hy Self.
MIMICRY IN SPIDERS.

Tre subject of mimicry among spiders, as with other animals, is most
interesting and yet most difficult to treat. I accept the word as one gen-
erally used among naturalists, to express certain resemblances, more or less
complete, between a spider and surrounding objects in Nature. I do not
include within the word the idea that the volition of the spider controls
these resemblances, except in a very limited degree, which will hereafter
be pointed out. The theories of the origin of mimicry, which have been
discussed by many naturalists, appear to me to rank little higher than
more or less ingenious suggestions unsupported by facts sufficient to justify
them as scientific inferences. But at present this condition of things
seems unavoidable, and by patient and careful accumulation of facts chaos
may at last yield to order and well defined law.

Among spiders the various kinds of mimicry may be divided into the
following: First, industrial mimicry of plants and other objects or envi-
ronment; second, form mimicry of animals; third, form mimicry of envi-
ronment; fourth, color mimicry; fifth, cocoon mimicry; and sixth, death
mimicry. The last of these will be considered in another connection.

The most remarkable examples of industrial mimicry of surrounding
objects are to be found among the Trapdoor spiders, as recorded in the
charming pages of Moggridge, some of whose figures I have
Industrial thought well to reproduce in Plate II. of this volume. These
Mimicry . . ; a
artnet: animals, which make burrows in the earth, whose openings are
ronment, Closed by doors swung upon a hinge of thickened silk, are in
the habit of covering the outside of their doors with dry leaves
or living moss, so that they resemble the surrounding site, in which they
are placed so closely that even Mr. Moggridge, when looking for them,
was often deceived.

Perhaps in no case is the concealment more complete than when dead
leaves are employed to cover the door, In some instances a single withered
olive leaf is placed in to cover the trap. In others several leayes are woven
together with bits of wood and roots, as seen at Plate II., Figs. 1 and 2,

(352)

MIMICRY IN SPIDERS. BDO

which represent the projecting entrance of a nest of Nemesia meredionalis
at Mentone. Fig. 1 shows the door closed and well disguised by re-
semblance to the dry olive leaves which cover the ground in the vicinity
of the nest. Fig. 2 shows the same tube with the door thrown open. The
effectiveness of this disguise is at once apparent. It may be questioned,
however, whether it is made with deliberate intent at mimicry. The spi-
der’s purpose in attaching leaves to the outer surface of its door is doubt-
less protective. But may it not be that, moved by ordinary self protective
instinct, it simply took the first available material, without regard to
mimetic resemblance ?

I may illustrate this by referring to my studies of the parasol or cut-
ting ant of Texas.’ My first experience of a formicary of these insects was
discouraging. I had encamped in its vicinity on the strength of
Kate information that it was a large and active hill; but at a morning

visit it seemed utterly abandoned, not a sign of life or activity
anywhere present. The mound was dotted over with forest chippage, bits
of twigs, dried leaves from an overhanging live oak, but no entrance into
the nest appeared. Returning to the place in the evening I found that
vigorous life had succeeded the semblance of death. Numerous openings
appeared all over the surface of the mound, out of which myriads of
insects were pouring, streaming away into the surrounding country, engaged
especially in harvesting leaves from the immense live oak tree that over-
hung. The change was readily explained by subsequent studies. I found
that the ants are in the habit of closing up their doors after their night’s
work, and when the upper part of the tubes which lead into the main
formicary have been filled to the distance of about an inch with various
chippage and sand, piles of dry leaves and twigs are erected above the open-
ing. These chips form a part of the permanent property of the ants, for
I observed that the same pieces were used day after day. The mimicry
in this case was complete. The surface of the mound was made to re-
semble a bit of natural soil covered over with piles of dried leaves and
twigs. It effectually deceived me, although I was on the lookout for the
insects. Yet I have never thought that the ants designed to produce such
a mimetic harmony. They simply gathered such dry leaves and other ma-
terials as were convenient, and the fact that these so closely resembled the
surface of the mound was accidental. May it not be that some of the
striking resemblances in the doors of Trapdoor spiders are produced in
the same way ?

Striking illustrations of this mimetic resemblance are shown in Figs.
3, 4, and 5 of Plate II. Fig. 3 represents a moss covered sod, pierced by
the tube of the nest of Nemesia coementaria, the door of which is entirely
concealed from view, and only discovered by one who happened to cut

" Proceedings Acad. Nat. Sci., Phila., 1879, page 33, sq.
si pag q

7 — ee

—_-™S~s—ss so — a a

354 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

through it in digging up a plant. The moss on the trap grew as vigor-
ously, and had “in every way the same appearance, as that rooted in the
surrounding earth, and so perfect was the deception that Mr. Mog-
Mimetic ridge found it impossible to detect the position of the closed
oa _ door, even when holding it in his hand, No doubt many nests
escape observation in this way, and the artifice is more surpris-
ing because there is strong reason to believe that this door garden is delib-
erately planted with moss by the spider, and not the effect of mere chance
growth. Figs. 4 and 5 represent a section of earth covered with a delicate
moss, which includes the trapdoor of Nemesia ccementaria. The door is
shown open at Fig. 4 and closed at Fig. 5, and the concealment of the door,
although not so striking as in Fig. 3, is nevertheless quite manifest.

In the case of Trapdoor spiders which make a thin or “wafer door,” as
Moggridge calls it, there is but a thin coating of earth on their upper sur-
face, since it is rare to find any of the larger mosses or lichens growing
upon them. But, as if to compensate for this deficiency, a variety of for-
eign materials is employed, which are scarcely ever found in the thick cork
doors, such as dead leaves, bits of roots, straw, of grasses, etc., and Mog-
eridge had seen freshly cut green leaves, apparently gathered for the pur-
pose, spun into a door which had recently been constructed.?

There is the widest possible difference between nest and nest in the de-
gree of perfection in concealment; and although, as a rule, the surface of
the upper door harmonizes well with the general appearance of
its surroundings, there are some individual nests in which it
readily catches the eye and even attracts attention. Mr. Mog-
gridge saw nests in mossy banks where the door, being made of nothing
but earth and silk, showed distinctly as brown patches against the green.
These doors even when surrounded by earth were often easily detected,
because when they dried up, as they quickly did, they became much paler
in color than the earth of the bank which retained its moisture.*

Thus it seems that the simple instinct to cover in the door, and so
protect the artificer from exposure to weather and enemies, was the domi-
nant motive. Material was chosen from the immediate vicinage suitable
and convenient for closure, and no purpose appears in the act to select
such material as would disguise the nest. In other words, a sense of se-
curity by means of sheltering barriers dominated the spider's mind, and
security by means of mimetic harmony or protective resemblance appears
to have had no place at all.

Some support for this opinion appears to me to be derived from Mr,
Moggridge’s statements as to the mode in which some of these Trapdoor
spiders work. He fastened back the doors of several tunnels, in order to
test the mode in which the inmate would deal with this difficulty. In one

Self Pro-
tection.

1 Trapdoor Spiders, page 97. 2 Ibid., page 103, * Tbid., page 103.

MIMICRY IN SPIDERS. ; 38D5

case he observed that a new covering had been cleverly extemporized out
of three fallen olive leaves, taken from the vicinity, which were loosely
spun together and attached by one or two threads to the margin of the
tube. This formed an admirable concealment, but did not move freely
as a door, the web being too imperfect. Two days later, however, it was
completed, and had become a perfect door, moving on a hinge just within
and below that of the former door, which still remained as it had been
fastened. The other nests remained in the same condition as before, only
that a little moss had been dragged into the mouth of one tube, which had
been partially closed with its own lip.

Moggridge further states that Trapdoor spiders, Nemesia meridionalis,
will make use of various objects strewed near their nests, in order to build

up a new door. This he tested by placing bits of scarlet braid
Nearby along with particles of moss and fragments of leaves, in a circle
ee around the opening of the tube, and about two inches away

from it. Nevertheless, it is apparent that these Trapdoor spiders
do exercise some discrimination in the choice of materials, for Mr. Mog-
gridge observed several instances in which, when the door of the cork
nest had been removed, if the door had been originally covered with moss,
it would again be used in its reconstruction, even though the mouth of
the tube were then surrounded by bare earth.

Thus, in reasoning upon the power of the spider to disguise the en-
trance to its habitation by mimicking surrounding natural objects, we are
brought in contact with this apparent defect of rational action. This point
Moggridge further illustrates by a case in which he had cut out a little
clod of mossy earth about two inches thick and three inches square on the
surface, containing the top of the tube and the moss covered cork door of
Nemesia ccementaria. He found, on revisiting the tube six days later, that
a new door had been made, and that the spider had mounted up to fetch
moss from the undisturbed bank above, planting it in the earth which
formed the crown of the door. Here the moss actually called the atten-
tion of an observer to the trap, which lay in the little plain of brown
earth made by the digging. He subsequently saw many examples of the
same sort, and purposely removed several cork doors from mossy banks,
in order to observe this point.?

If in the above cases the spiders appear to have been guided by the
simple wish to protect themselves with the first available material, in

other cases the mimicry seems to be due to the natural secret-
ae iveness of animals seeking prey. Mr. Moggridge detected the
iveness, /@™ains of insects, and especially ants, in the nests which he ex-
amined in situ. Frequently, however, one may open several in
succession without finding any of this débris, and at other times it will

* Trapdoor Spiders, page 121. * Tbid., page 120.

356 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

only be detected beneath the bottom of the tube, layers of silk having
been spun over successive layers of refuse. The horny crops of ants form
by far the largest proportion of these remains, and Moggridge was struck
by the number of instances in which, while digging out ants’ nests at
Mentone, he found Trapdoor burrows, especially those of Nemesia mander-
stjerne and Nemesia moggridgii in their midst, the tubes often traversing
the very heart of the ants’ colony, and coming into close contact with
their galleries and chambers.

In these instances the trapdoors had almost always escaped his notice,
and, indeed, they so closely resembled the surface of the ground that even
when he knew, from having accidentally cut across the tube below the
eround, that one of these doors must lie near a given spot, yet he could
only discover it by following the passage below upwards, ‘This perfect
concealment the discoverer thought of essential importance to the spider’s
success in life, for if they once alarmed the whole colony of ants and let
them know the exact whereabouts of their
lurking place, they would soon learn to
avoid it.?

The fact of mimetic resemblance in the
tubular snare of the Purseweb spider has
already been alluded to. I have seen hun-
dreds of these in various parts of Florida,
and have before me several score speci-
mens. These are covered on the outside
with particles of sand, and even more free-
ly with the brown wood mold which has
accumulated in large quantities around the
trunks of trees in Floridian forests where
the spider abounds. The resemblance of
the tube to the bark of the tree against
which it is planted is close, much closer,
i in most cases, than is represented in the
Pram: ith / Tila dete Deneibae MS Pe drawing Plate IL, Fig. 7. As the spider

seudidiops opifex. (After Simon.) A

is dependent for her supply of food upon
the number of insects that crawl upon her tube, we may suppose that
she derives considerable advantage from this resemblance, imas-
much as it allows her to creep upward to where her victim rests,
or encourages the victim to crawl towards the point where she
lies in wait to fling her web around it.

An example of nest architecture among Trapdoor spiders which may
be classed in the same category as the above, is a species which Mr. Ku-
gene Simon describes as Pseudidiops opifex. (See Fig. 314.) This aranead

Purseweb
Spider.

! Trapdoor Spiders, Supplement, page 237.

MIMICRY IN SPIDERS. 357

constructs a tube with a trapdoor opening, in every respect resembling
that of species which burrow in the ground, except that the tube is placed
upon the bark of various trees and ordinarily lodged within
Tree the furrows of the same. Mr. Simon! found specimens of these
are upon trees of Venezuela, South America, and I have seen a num-
: ber of examples in the collection of the British Museum at Ken-
sington, London. ‘These were of various ‘sizes, some of them with doors
no larger than a pin head, yet perfectly constructed and exact miniatures
of that made by the adult. The utility of this mimicry is apparent. The
insects which alight and walk on trees, must often go over and around
this trap, which in appearance and texture so closely resembles its site.
Thus opportunity and facility are afforded the spider, waiting at its partly
open door, to seize its prey.
I.

Among spiders, the form of other animals is sometimes mimicked. The
most striking example is that of the little group of araneads which, by
stricture of the abdomen and shape of the head, are made to
Form resemble the form of certain ants. Simonella americana Peck-
saree ham is an example of this form mimicry. (Fig. 315.) One
mals. specimen of this spider is recorded as haying been found run-
ning among leaf cutting ants.? Certainly there is a sufficient
resemblance between the two creatures to permit one to think that a spi-
der so formed might run upon the ground among a marching or working
column of these emmets without any great fear of detection, provided
the ants had no better means of discovering the presence of friend or foe
than their eyesight. As a matter of fact, however, their
principal means of observation in this respect appears to be a
the sensitive antenna. Having closely studied the habits of
the leaf cutting ants, Atta fervens,? in Texas, I cannot MS: An outline

A : - side view of Si-
readily think that any spider, or any other creat- monella america-

Ant, like | ure at all obnoxious to these ants, whatever might eer tape
Spiders. ham.*)

be its form, would have been permitted to remain

in the way of the immense swarms of insects that issue in the evening
from their formicary, and go forth upon their predatory expeditions into
the surrounding foliage.

Another of these ant formed spiders, which belongs to our indigenous
fauna, is Synemosyna formica Hentz. (Fig. 316.) The figure is drawn
from a specimen sent me by Professor Peckham, but the species is quite
widely distributed over the United States, having been originally de-
scribed by Hentz. Yet another spider which certainly bears a striking

1 Ann. Soc. Entom. de France, 1889, page 220, plate i, Fig. 3.
2 Spiders of the Subfamily Lyssomane, Trans. Wisconsin Acad. Sci., 1888, page 252.
’ Proceed. Acad. Nat. Sci., Phila., 1879, page 33.

* Proced. Nat. Hist. Soc. Wisconsin, 1885, plate i, Fig. 1.

SO — es a er

358 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

resemblance to hymenopterous insects, is drawn from a specimen sent me
by Professor Peckham, and was collected in South America. (Fig. 317.)
I do not know the particular genus to which this species be-
longs. The number of species having this characteristic form
appears to be quite limited in any given lecality, particularly
of temperate climates; but there are probably many more
than has been supposed, for Professor Peckham informs me
oder syne, that he has at least seventy-five species in his private col-
mosyna for- ection.
ik ak According to Mr. Cambridge, the most striking instance of
resemblance among British spiders is that of Micaria scintalans, found in
some abundance in the Isle of Portland. ‘Dis spider so nearly resembles a
large ant which abounds in th same locality, that it requires the
second look of even a practiced eye to vie sure whether it be really
spider or ant. The advantage, or ‘ protective effect,” afforded
to one thus resembling another is not always easy to be understood at
once, se eae often consist in the protection from certain Sangety to which

A Britis
Specimen

steccnlnn ‘the ants of Poranls “being: ‘of a hard ae horny nature,
may not be a favorite food for those enemies which would find an agree-
able morsel in the softer and more succulent spider. ‘The latter, therefore,
would deceive, and -so-escape such enemies, from its resemblance to the dis-
tasteful ants. In other cases (and, possibly, also in the one just mentionéd)
Cambridge thinks the resemblance may give the resembler a chance of ob-
taining its prey more easily. Thus, in the south of Africa there is a spider
resembling an ant even more closely than the Portland species. The habit
of this ant is to feed on honey dew along with mul-
titudes of insects of other orders. ‘These latter have
no dread nor suspicion of the ants, which, in fact,
have a common object in view, and do the other in-
sects no harm. But then, under cover of the close
resemblance to the ants, come the spiders, who, un-
suspected and unresisted, regale themselves at their
leisure upon the defenseless insect.

Having recorded the facts, it may be well to con-
sider some of the theories advanced to account for
them. In general, it may be said that this
mimicry is considered as protective. Its use
is, first, to enable the mimic to more readily
obtain necessary food among its natural prey; or, sec- Fie. 817. An ant like spider
ond, to protect it from natural enemies, particularly ee ee
(as in this case of form mimicry) from some enemy that especially threatens
its existence. Darwin applies to these analogical resemblances, or adapt-
ive resemblances, as he calls them, his theory of natural selection. He

Darwin’s
Theory.

MIMICRY IN SPIDERS. 359

cordially adopts the opinion expressed by Mr. Bates concerning the remark-
able mimicry between the genera of butterflies, Ithomia and Leptalis, as they
are found in Brazil.!_ Mr. Bates concludes that the Leptalis first varies,
and when that variety happens to resemble in some degree any common
butterfly inhabiting the same district, this variety, from its resemblance to
a flourishing and little persecuted kind, has a better chance of escaping
destruction from predaceous birds and insects, and is consequently oftener
preserved, the less perfect degrees of resemblance being from generation to
generation eliminated, and only the others left to propagate the kind.

In this connection. Mr. Darwin remarks: ‘Insects cannot escape by
flight from the larger animals, hence they are reduced, like most weak
creatures, to trickery and dissimulation.”? In what sense can it be true
that a resemblance in form, which must be the result of influences oper-
ating upon the very germ of life, acquired by and transmitted from an-
cestors, is traceable to the volition of the creature, and is an act of delib-
erate “dissimulation and trickery”? Whatever may have been the origin
of adaptive resemblances, certainly at the outset we may exclude any such
supposition as this. In the nature of things the cause of structural re-
semblance is beyond the individual control of the mocking or mimicking
species.

The most striking example of the mimicry of animal forms among
spiders, as has been said, is that of Ant spiders, of which Simonella ameri-
cana is an example. The theory which accounts for this on the
hypothesis of natural selection supposes that, through the nat-
ural tendency to vary, a spider in a brood acquired a slight re-
semblance to an ant. This slight resemblance protected the spiderling so
much as to give it an advantage over its fellows during the attacks of
birds that feed upon spiders, but do not feed upon ants. This protected
individual, having matured, transmitted its peculiarity to offspring, some
of whom, by the same tendency to variation, exaggerated the ant likeness;
and so, by infinitesimal increments, in the course of time Simonella amer-
icana and other species more or less closely resembling ants were produced.

Concerning this theory it may be remarked, first, that the real difficulty
seems to be in the supposition that such a slight variation as is supposed

could possibly be of any advantage to an individual spiderling

Value of in the midst of a large brood. The dangers to which these are
Sa a exposed are not chiefly from birds. They are very small, soft
bodied creatures, exposed to many perils. As soon as they set

up housekeeping, and even before it, they are preyed upon by their own
order, for large spiders unceremoniously eat. little ones, and small spiders
eat less ones. Among | Wanderers like Simonella and other Attide, the
—— : a === —

Ant like
Forms.

1 Bates, “ Naturalist on the Amazon.”
2 “Origin of Species,” Chapter XIII., page 386.

* =o a

360 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

young stay with the mother until they are somewhat grown, and then
have shelter beneath rocks or other secluded positions where birds are not
apt to find them, although, of course, some birds do mouse around and
pick up insects in the most secluded spots. I imagine, however, that the
dangers threatening young Attide are much greater from certain insects
and individuals of their own order, than from birds. I can, therefore,
hardly conceive what advantage it would be under such circumstances for
the spider to resemble an ant, even if we were to suppose that such a
minute resemblance as the hypothesis requires at the outset, would be of
advantage in any case. In point of fact, the theory is not workable, as it
seems to me. Any change of form to be effective must occur in the first
stages of life. But a minute resemblance could be of no advantage, as the
discriminating powers of enemies, whether insects, birds, or spiders, are
hardly so acute and delicate as to make an infinitesimal variation of
much importance in screening one individual spiderling in the midst of
a brood. In short, if we are to suppose that the birds are the real ene-
mies, or any other creature that is indisposed to feed upon ants, it seems
necessary, in order to justify anything like this theory of the origin of
mimicry, to suppose that the variation of the spiderling was, at the outset,
so great as to give it at least a reasonable likeness to the ant.

The theory takes for granted an accuracy of eyesight on the part of
birds that close observers will scarcely be willing to admit. How far can
birds distinguish color? This is a question which has scarcely
yet been fully solved. How far can insects distinguish color?
How far can birds and insects distinguish between such minute
variations in form as that which the above theory seems to require? In
accounting for the origin of cocoon mimicry we suppose that the eyesight
of birds and wasps is so defective as to form and color as to permit them
to be deceived by a difference as little marked as that which exists between
the cocoon of the Tailed spider, for example, and the spider herself. But,
in accounting for the mimicry of ant like forms, we are compelled to re-
verse this attitude, and suppose the eyesight of birds and raiding wasps
to be so accurate that it can distinguish between a slight variation on the
part of one spider of a brood towards an ant, and the normal form of
other spiders of the brood, and distinguish so accurately that it will
avoid the ant resembling spiderling and take others. Whatever theory
of the origin of mimicry we adopt, certainly must be free from inconsist-
encies such as this.

Moreover, the greatest destruction, as far as I am aware, wrought by
birds upon young spiders is accomplished under circumstances that pre-
clude any such an element as above. Immense numbers of spiderlings,
including, I believe, all species of Saltigrades, possess the acronautic habit,
and while they are flying through the air upon their tiny mimic balloons
they are devoured by swifts and swallows, who skim the air and gather

Sight of
Birds.

MIMICRY IN SPIDERS. 361

these flying spiders into their crops, where they have been found in quan-
tities. Certainly, a resemblance of an ant form could be of no avail here.

Again, the theory compels us to assert that ants are not subject to de-
struction by birds. Of course, unless this be true we can conceive of no

: advantage in the mimicry of an ant form as protecting the spi-
age der against birds that seek to devour it. Its safety lies in the
at Ants. ics P : ’
fact that it is covered from harm by its likeness to an insect
which birds avoid. On the contrary, I know that some birds certainly
do eat ants, and eat them greedily.

Mr. Carl Voelker, of Carlingdale, Delaware County, Pennsylvania, is a
taxidermist of large practical experience, and with a fondness for natural
history which prompts him to make and note observations upon the gen-
eral habits of birds and other animals. He has informed me that the
flicker, Picus auratus, at certain seasons of the year—in the spring, for
example—will station itself upon a dead stump, which in our American
woods is frequently infested with ants, and feed greedily upon them as
they pass to and fro. This he has observed many times, and_ believes
that at certain seasons this bird lives entirely upon ants. He has seen
two species devoured by them in the manner above described—a_ little
black ant and one about twice as large.

The pileated woodpecker, Picus pileatus, feeds quite habitually upon
the large black Pennsylvania carpenter ant, Camponotus pennsylyanicus.
He has frequently taken these insects, in various stages of decomposition,
out of the crops of these birds, at one time having counted nearly seven
hundred in the crop of a single woodpecker. The bird not only takes
the ants in the summer, but also in the winter, and Mr. Voelker has seen
them stationed upon trees, pecking at the gangways or gates into the for-
micaries until they had been hammered open, and then extracting the
ants while they were in a torpid state. As some of these homes in forest
trees are extensive bits of architecture and are inhabited by vast numbers
of insects, the amount of food thus obtained must be considerable. In the
forests of Pennsylvania I have seen formicaries of the carpenter ant six
feet in length and occupying the entire central part of a goodly sized
tree or branch.

The European woodpecker, Picus ater, according to the same gentle-
man, who has observed the same species in Germany, subsists

Ee a entirely upon ants, and the same fact is true of another Euro-
Retre pean species, Picus virens, popularly known as the Grass Wood-

pecker. Mr. Voelker has seen this bird on the hills of Germany

digging into the soil, and feeding eagerly, not only upon the larvee, but
upon the ants themselves.

It is generally known that the ordinary barnyard fowl will devour ants

without hesitation. Mr. Voelker states that once he was engaged in the

woodyard of his country residence in breaking up a log of decayed wood

——— <—" =

ee ss Oe el

362 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

which had served as a formicary for a large colony of ants. The insects
with their larvae and pupx were scattered over the ground, and the chick-
ens, getting scent of the game, flocked one after another from the barn-
ian in such numbers and with such greedy persistence that he had to

ease his labors out of regard to the safety of his fowl. The chickens de-
ie the larve and pupe and the ants.

The same gentleman says that he has found particles of ants in the
stomachs of grouse, although it is possible that they may have picked up
the ants while feeding upon the larvee, which latter may have been the
chief object of desire. The various song birds of America, as the mocking-
bird, catbird, thrushes, etc., eat the larve of ants, as is well known, but
Mr. Voelker had never discovered particles of the ants themselves in their
crops.

Mr. T. B. A. Cockerell! notes that Dr. Riley records that sparrows
(Passer domesticus) feed on certain Aculeta, Halictus, Typhia, Myzine, and
ants. Mr. Cockerell himself had found ants in the stomach of
Sialia arctica shot in Custer County, California. The stomach
of a woodpecker shot by Rey. A. Wright in the same locality
contained a number of ants, the majority apparently Formica fusca, with
a few of Formica integra.

I can certify by my personal observations, as well as by reports of
others, that some of the native birds of Fairmount Park (Philadelphia)
feed upon colonies of Formica integra, which are found in the neighbor-
hood of Rockland on the Schuylkill River and elsewhere. It is well known
that birds of all kinds are fond of “ant eggs,” by which popular name is
meant the pupe of those ants whose larve enclose themselves within a
cocoon, These ant pup are gathered in immense quantities from the
mounds of Formica exsecta, F’, fusca, F. rufa in various parts of Europe,
and are regularly sold in the markets as food for pet birds. These eggs,
if nothing else, would invite the attack of birds upon ant hills, and would
thus lead directly to devouring the ants themselves, who invariably rally
to defend their nurslings.

Certain game birds are extremely fond of ants. In the summer of 1887,
while visiting Mr. E. C. Cornwallis at Linton Park, Kent, England, I was

taken by my host to the gamekeeper’s lodge, on the grounds of
English which several hundred, perhaps a thousand or more, young par-
Game j - ; ; ‘
Birds. tridges were being raised for the purpose of stocking the shoot-

ing park. These little fellows had been hatched out under barn-
yard fowl, and were, when I saw them, turned loose upon a bit of sloping
ground that was literally honeycombed with the nests of a small species of
ant, apparently a Lasius. The whole slope had been torn up in order to
procure these nests as food for the young partridges. Mr, Cornwallis gave

Other
Birds.

1 Entomological News, Philadelphia, May, 1890, page 65,

MIMICRY IN SPIDERS. 363

a peculiar whistle, which was at once recognized by the flock of chickens,
who hurried together at the call, as tame birds always do when summoned
to feed. The gamekeeper tossed them several bits of sod containing ant
nests, which the birds attacked, tore in pieces, and in a moment had de-
voured all the insect contents. Simonella americana is not much larger
than these ants, and certainly there would be little protection from such
voracious creatures as these in the mimicry of ant forms. I have no doubt,
although I cannot speak positively, that our American quail are equally as
fond of ants as these English partridges; and as they are ground birds,
their habits of feeding would make them destructive enemies of all ant spe-
cies burrowing in the earth.
Again, it is well known that in the warm districts of South America,
and in other parts of the earth, there is a family of birds who are such
persistent destroyers of ants that they take their family name
Bae Wee from this habit, and are known as the Formicariide.! ‘These
ant thrushes, Pittas or Pittide, are also an Old World group,
being found in the Malay Islands. The Great ant thrush, which is also
called the Giant Pitta, is a native of Surinam, and is a bird about the size
of the English rook.?

I have heard, although I cannot now cite the authority, that in Africa, |

when the Driver ants go out upon their excursions, during which they will
prey upon all sorts of insects and small vertebrate animals, ant thrushes,
or some species of ant devouring bird, hover over the raiding column,
upon which they make their assaults, devouring immense numbers of the
drivers. Such are some of the facts which have fallen to my notice, or
under my eye; and while it is probably true that some birds avoid ants
as articles of food, I imagine that nearly all animal feeding birds will pick
them up whenever they have an opportunity. Such being the case, we can
hardly admit the force of an argument which is based upon the supposi-
tion that the form of an ant would protect a spider, or any other creature,
on account of the disrelish of birds for ants.
If we were inclined to accept the theory of natural selection, as above
outlined, as an origin for mimicry of ant forms, it would seem to me more
rational to suppose that the particular enemy against which the
Are mocking form is protected, is not the bird, but the wasp and
Wasps. = ; te
Mim- ichneumon fly. My chapter on the Enemies of Spiders shows
ickead? What depredation is wrought among araneads by various mem-
bers of the wasp family. As far as I know, wasps do not inter-
fere with each other, or with ants, who closely resemble them, being, in
fact, members of the same order of Hymenoptera, Anything that would

1 Alfred Wallace, “Geographical Distribution of Animals, Vol. IL, page 296; Wright,
“Animal Life,” page 271. 2 “Wood’s Natural History,” page 341.

*T think that this, or a similar fact, was told me by an African missionary to the Congo
region.

a ————= ss lh CU CU CU

—=

364 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

be likely to deceive these inveterate spider enemies would undoubtedly be
a protection to spiders of all tribes.

But then, with such a theory in mind, we are met at once by the fact
that those spiders which are most frequently found within the clay cells
of mud dauber wasps, and those which these insects most frequently col-
lect as food for their larvee, are the Sedentary groups such as Orbweavers
and Lineweavers. They do indeed take the Thomisoids, especially those
that lurk on flowers in pursuit of prey, and which, in turn, sometimes
rapture the wasps. The Saltigrades are also taken; but if I may judge
from my own observations, they are least numerously represented of all
the tribes except perhaps the Lycosids and the Tunnelweavers. This seem-
ing immunity is evidently not due to any likeness of Attide in general
features to wasps, but simply to their manner of life, which, in large meas-
ure, screens them from assault, and enables them to escape. Now, the
question must rise in considering such a theory, why does not natural
selection operate for the protection of those spiders which obviously need
protection the most ?

Is it not remarkable that during all the ages in which the forms of
Orbweavers have remained substantially unchanged, as well as the forms
of wasps, and during which period the habits of both creatures must have
been the same, Nature has refused to work in the direction of protecting
the exposed Orbweavers by providing them some analogical resemblance
such as that which we remark in the case of Simonella americana? It
seems to me illogical to expect a general law to account for the origin of
certain peculiarities in Nature, and yet to exclude this general law or force
from operation within the whole field of life with the exception of one
very small section. It seems further illogical to hold that this general
law would have failed to operate not only in the cases where it seems to
be most necessary, but in those wherein all the circumstances are best ar-
ranged for its most effective operation.

While spiders thus abundantly prey upon ants, sometimes the condi-
tions are reversed, for when the opportunity presents, the ants will feed
upon spiders. In certain cases this takes the shape of a system-
atic raiding of the whole section, as, for example, according to
Mr. Cambridge, the large red ant of the woods, Formica rufa,
destroys spiders so completely, that in localities thickly inhabited by those
insects, he had generally found it almost useless to search for spiders.1

Whether or not any ant like species are found among Sedentary tribes
I do not know. But it entirely passes my imagination to conceive what
possible advantage could accrue to an Orbweaver, for example, from resem-
blance to an ant. Orbweavers, and yet more frequently Lineweayers, prey
upon ants; but it is not necessary that there should be any resemblance to

Ants Hat
Spiders.

‘ Spiders of Dorset, Vol. I., page xxxi.

MIMICRY IN SPIDERS. 365

the emmet in order to accomplish the destruction of vast numbers of them,
as I can fully testify. As the Orbweavers and Lineweavers do not leave
their snares to capture prey and move among the ants after the fashion
of the prowling Saltigrades that do mimic ant forms, the fact of ant re-
semblance, should it exist among them, must have a quite different solu-
tion. One could suggest, in their case, a protective value in resemblance
to wasps, but none at all as against ants.

Ti:

Another style of mimicry among spiders is resemblance to forms of
objects among which they live, as with Tetragnatha extensa, This aranead
has a long and narrow body, of a cylindrical shape, not unlike
Local, or g small twig in appearance. Its colors are delicate green, yellow,
Mimicry . . .
of Envi- and gray, thus increasing its resemblance to the plant. But the
ronment. ™ost striking feature of the mimicry is the habit of drawing to-
gether the four hind legs until the joints closely approximate
each other, stretching them straight backward, and treating the two pairs
of fore legs in the same way, stretching them out forward. (Plate IIT.,
Fig. 6.) Thus the spider is extended along the stem of the plant in a
straight line, so that her body closely resembles the object upon which she
lies. ‘The habit prevails in all known species of the genus. In this case,
although we concede that Tetragnatha has had no control over her own
particular form and its resemblance to a small twig, we must allow that
her action is a matter of personal volition, and appears to be exercised
with deliberate purpose to conceal her presence. Yet, the behavior of the
spider frequently compels one to wonder how the supposition of mimicry
can be harmonized therewith.

For example, the Peckhams record that a male Tetragnatha grallator,!
when touched as he hung in the web, ran to a branch, whereon he stretched
himself. In this position he was almost indistinguishable, as his
color was exactly like that of the branch to which he clung.
The branch was gently shaken, but instead of keeping quiet he
ran a little way and then stretched out again. This he repeated, stupidly
betraying himself as often as the branch was touched.? One would think
that Nature, having taken pains to produce such a_ striking protective
mimicry, would not have failed to make it useful by imparting a corre-
sponding instinct that would adapt behavior to opportunities.

A Laterigrade spider, Tibellus, which has at least one quite common
representative in the United States, has a habit not very different from
Tetragnatha. Tibellus oblongus is‘ abundant in many marshy places in
the south of England. It has an elongated oval body, with longish legs,
and is of a uniform dull yellowish hue; it is an exceedingly active spider,

Tetrag-
natha.

''T. elongata Walck, 2 Mental Powers, page 411.

366 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

and, when running in autumn among the dull yellowish, decaying grass
and rushes, looks much larger than it really is. All of a sudden one loses
sight of it, and unless he is aware of its habits, is puzzled to know what
can have become of it; but there it is close by, stretched out at full length
along the similarly colored stem of grass or rush, with its first and second
pairs of legs put forward in a straight line, and its third and fourth pairs
stretched in the same way backwards, so as to be scarcely distinguishable
from the stem itself.?

In the case of Tetragnatha the mimicry of the twig on which she lies
appears at first sight very striking. I have no disposition to undervalue

the character or protective benefit of this mimicry, But it must
pone be remarked that, in point of fact, she simply assumes the posi-

abit. P : : ;

tion which she habitually takes when hanging on her web. Both
this genus and Uloborus stretch themselves out upon the central part
of their orb, or upon a string suspended from it, with their feet ap-
proximated in precisely the attitude above described as taken by them
when they stretch along the under part of a branch. We are therefore
compelled to inquire how far this attitude of Tetragnatha may be a de-
liberate attempt to shield herself, and how far it is the natural result of
habit prompting her, when attempting to screen herself, to drop into the
form natural because most common to her. The value of the form need
not be questioned, but in considering its origin we are required to con-
sider the habitual attitude of the spider upon her web as well as the atti-
tude of mimicry upon a twig during her occasional excursions. Some
other spiders have the habit of stretching themselves like Tetragnatha
upon foliage and twigs, as, for example, the Orchard spider, although not
to the same extent as Tetragnatha. (Plate III., Fig. 6, above.)

The ordinary, or at least the common, position of Epeiroids, when rest-
ing outside their nests or snares, is to draw up the legs so that the two
front pairs are humped up above the head and are nearly in a
plane with each other and the spider’s face. The hind legs are
drawn up against the abdomen. The spider thus forms a little
roundish bunch and as thus seen is not unlike one of the knots, warts,
or excrescences frequently seen upon plants. This habit universally pre-
vails among spiders, and one who follows their actions for a little while
will be certain to observe it. Thus they remain perfectly still, and at a
casual glance might be taken for a knot or other excrescence upon the
bark of a tree or plant, as in the case of Epeira strix, represented Plate
III, Fig. 4.

The mimicry of a wart, knot, or bud, or other natural irregularity,
would seem to be quite as useful as the special mimicry of Tetragnatha,
since it would be as likely to deceive the eye of a prowling enemy. In

Mimick-
ing Knots

’ Cambridge, Spiders of Dorset.

MIMICRY IN SPIDERS. 367

this bunching habit spiders, Orbweavers at least, follow the influence of
habit, for when at rest within their nests they almost invariably draw
themselves up in the manner described. However, the position is contrary
to that generally assumed by spiders making vertical orbs when hanging
in wait for prey upon the orb. Then the legs are stretched backward and
forward, not, indeed, in a straight line like Tetragnatha and Uloborus, but
somewhat in the position of a St. Andrew’s cross or the letter “x,” as
heretofore described.

IV.

Color mimicry in spiders is supposed, first, to facilitate taking prey, thus
promoting the life of the species, and, second, to protect the mimic from
assaults of enemies, who are deceived by the close resemblance
either to surrounding objects or to some animal obnoxious to
the assailant, An interesting example of color mimicry is the
common spider Misumena vatia. ‘This spider is generally yellow, mottled
upon the abdomen, and with darker rings on the legs. It will often
be found spread out upon the yellow heart of an ox-eyed daisy, or in like
position upon Coreopsis (Plate IIL, Fig. 1) or golden rod. Here it re-
mains and preys on insects frequenting the plant. It certainly closely re-

Color
Mimicry.

‘sembles the flower upon which it is ambushed, and the ordinary observer

might well fail to notice its presence. On one occasion I found this spe-
cies concealed underneath one of the outer petals of a half opened rose,
and, curiously, it had its prevailing yellow greatly modified by a pinkish
cast of color, more closely resembling the rose upon which it was lodged.
(Plate III., Fig. 2.)

Mrs. Treat gives an account of a Laterigrade that appears to be Misu-
mena vatia, whose lurking place was in the heart of roses, and was so
nearly the same shade of color as the flower as to make it dif-
ficult to see her. When the rose began to wither she took up
position on a fresh one. The spider was first observed in July,
and remained on the same bush about three weeks, and then moved to a
bright red tea rose, whose stamens were more conspicuous and numerous
than the other, and which was visited by a greater number of insects.
Misumena went to the centre of one of the flowers, but the stamens were
of a deeper yellow than her body, and the surrounding petals made her
easy to be seen. She seemed to know this as well as the looker on, and,
although more game visited these roses, she did not stay long. The ob-
server thought the spider conscious that her safety depended upon the re-
semblance, and therefore returned to her old home among the petals of
the light colored rose.

She made no web to entrap prey, but depended wholly upon strategy
and muscular strength. When waiting for prey she cuddled down in the
centre of the flower, and erected her long fore legs in such position that

Misume-
na vatia.

‘te ie an

——

sil canal

368 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

it was almost impossible to distinguish them from the imperfect scattering
stamens. If a wasp or humble bee alighted near her, she dropped her
stamen like legs and crouched down and concealed herself as much as
possible; but when these formidable insects departed, she resumed her ex-
pectant attitude.

Now a pretty butterfly comes flitting down, all unconscious of danger.

Misumena is perfectly motionless, but at the proper moment
oes shoots out her legs and grasps the insect in fatal embrace. The
Soares butterfly is often four or five times her own weight, yet she

manages to prevent her victim from mounting with her into the
air, probably by holding firmly with her hind legs to the flowers.!

The remains of night flying moths were often observed scattered near
this individual, which had evidently been captured during the night, but
her most frequent game was dipterous. The fact, however, that she does
prey so frequently upon night flying insects is a good indication that she
is able to acquire all the food needed without the aid of mimie colors.

I quote another popular account of the habits of a spider which I take
to be the same species. The description is from the pen of an intelligent
observer, but not a naturalist, who simply records, with great astonish-
ment, a first experience of a hitherto unknown fact, and therefore without
any predisposition to see a case of mimicry in a casual resemblance. The
account is taken from a description of a walk in the vicinity of Media (a
few miles from Philadelphia), and was published in a Friends’ educational
journal. ?

“In Bare Hill meadow was a garden of flowers such as no man ever
planted or ever shall. Asters, Golden rods, and Composite generally, were

Ete massed in such profusion that the meadow was like Joseph’s
coe aie coat, of many colors and bright ones. In some places the herb-
Flowers, #g¢ was higher than our heads, and passage through it was

difficult. Over all towered the spires of the purple Boneset or
Queen-of-the-meadow, so beautiful and graceful when seen thus, so coarse
and clumsy when examined closely. On a head of this species, and among
its purple flowers, we noticed a little purple spider with oval body and
peculiar markings. So closely was the color of the spider adapted to that
of his dwelling that we should have overlooked him entirely if he had
not moved,

“Shortly we found on the white panicle of the Boneset proper a spider
similar in size, shape, and markings, but pure white in color. We then
examined the Golden rods and found a third similar spider thereon of a
yellow color. A close examination of our flower garden revealed a number
of these variously colored insects, each simulating the color of its habitat.

* Mrs. Mary Treat, “My Garden Pets,” page 13.
*“The Student,” Philadelphia, Fourth Month, 1889, page 335. “The Banks of Crum,”
L. Chalkley Palmer.

PLATE

SOME HYMENOPTEROUS ENEMIES OF SPIDERS.

1, EURYPELMA HENTZII. 2, PEPSIS FORMOSA.

3, ELIS 4-NOTATA.

5, PEZOMACHUS DIMIDIATUS. 6, PEZOMACHUS GRACILIS.

8, TRYPOXYLON POLITUM.

4, PEZOMACHUS MEABILIS.
7, CHALYBION C/ERULEUM

' actin “4 ~

(GPa } ry ic
b, Sage wal, -- 7

MIMICRY IN SPIDERS. 369

It would be strange if all these were one species, differently colored accord-
ing to the place they dwell in, and having power to change from purple
or white to yellow at will; yet this seemed probable to us. Or was it
that a species had developed these different varieties, each adapted to live
on a certain plant? However this may be, the arrangement evidently
resulted in a twofold advantage, in that it enabled the spiders to escape
the peering eyes of birds, and at the same time to lie in wait unperceived
for the insects of various sorts that frequent such flowers in great num-
bers. In no case, it may be added, did we see a purple spider on a yellow
flower, or vice versa,”

Mr. Cambridge has observed and recorded like facts of Misumena vatia
in England! He says: “TI find this spider very commonly in the blooms

of the great mullein, Verbascum thapsus, to which the pale yel-
ae F lowish hues of the spider are well suited for its concealment in

nelish é : F 5

Species, the yellow blossoms. An allied spider, Thomisus onustus, found

on the heather blooms, and upon some other pink flowers, is
beautifully tinted with pink, chiefly in its younger and feebler stages.
The Rey. C. W. Penny (of Wellington College, Wokingham) tells me that
he has found examples of this spider on yellowish blossoms, and that
these examples are generally of a yellowish hue, quite devoid of the pink
color of those found on pink blooms. I am inclined to think that this
is not invariable, inasmuch as I have found here the more mature exam-
ples, which are generally devoid of pink coloring, also on the pink heather
blooms. The protective resemblance of color would not be so necessary,
in the above instance, for the protection of the more mature as for that
of the younger spiders, and therefore we might expect to find the former
on flowers of any color growing where the spiders are found; while I have
certainly only met with the younger pink colored spiders on the pink
heather blossoms.” It is important to note the above exception as to
adult forms,

Most other Laterigrade spiders known to me are of a dull gray or
brownish color. As they are frequently found resting upon the bark of
trees, over which they prowl seeking their natural prey, their resemblance
to the color of the bark is quite striking, and might serve to protect them
alike from the observation of enemies and of victims (Plate IIT., Fig. 3.)

Among the Citigrade or ground spiders, the same fact may be noted.

ere Their colors are generally neutral or dull grayish, mottled or
Mimick- striped with black. They thus blend easily with the colors of
i the ground and stalks of plants and grasses among which they
Ground. frequently move. Some of these spiders when found in littoral
sites take upon them the color of their surroundings. For ex-

ample, the Turret spider, which I have observed along the seashore from

1 Spiders of Dorset.

—

370 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Cape Ann southward, burrowing in her characteristic perpendicular holes,
has in such sites a color quite corresponding with that of the white sand
in which she dwells.1 The same species taken further in the interior is
found of a darker hue, thus resembling the soil in which it lives. This
would appear to be a decided example of the adaptation of color to envi-
ronment, or, as better stated, the influence of environment upon color.

Cambridge refers to like facts as marking English species. A Lateri-
grade, Xysticus sabulosus Hahn, so exactly resembles both in form and
color the little bits of gret, yellowish black, and red brown mottled stone,
found on the bare patches where turf has been pared off the heaths, that
until the spider moves it is almost impossible to detect it. Lycosa herbi-
grada, a gray spider marked with black and brown markings, is another
instance of exact adaptation to the gray, sandy heaths where it occurs;
while Philodromus fallax is equally well concealed by the perfect adapta-
tion to the coloring of the dull yellowish, sandy spots where alone it is met
with. The common and beautiful English Epeira cucurbitina, found on
rose and other bushes, in gardens and woods, is of a clear, bright green
color with a brightish red spot at the hinder extremity of the abdomen ;
this spider, when, as it often does, it sits tucked up between the green
shoot and the axil of the leaf, looks exactly like a young bud just ready
to burst.?

Mimicry and the survival of the fittest have been suggested to account
for this interesting habit. It is argued that those spiders, among the nu-
merous broodlings hatched out from the eggs, whose colors most
closely resemble those of the flowers alluded to above, are the
ones which survive, by reason both of the degree of protection
against enemies derived from their likeness to colors of the flower, and
their facility to capture prey because of the same resemblance, which would
naturally conceal their presence. In other words, those spiderlings which
by any chance happen to find lodging upon yellow flowers, or flowers most
closely colored like themselves, are the ones which survive the perils of
spider babyhood and grow to adult age.

Before one fully accepts this theory it will be well to consider certain
difficulties. ‘The most perilous age of spiders, as is well known, is that
which immediately follows exode from the cocoon. In a multi-
tude of cases in which these little ones entered life far removed
from any flowers corresponding with their normal color, how
are we to account for their preservation? Certainly they did live and
retain their natural colors in spite of the absence of golden rods, black-
eyed Susans, ox-eyed daisies, and flowers of like hue. Moreoyer, one is
compelled to establish the fact that the opening up of these flowers cor-

Natural
Selection.

Difficul-
ties.

‘See my notes on “The Turret Spider on Coffin’s Beach,” Proceed. Acad. Nat. Sci.,
Phila., 1888, page 333. ? Spiders of Dorset.

MIMICRY IN SPIDERS. OTL

responds with the entrance of the little fellows into life, and that the pe-
riod of flowering is contemporaneous with their growth. It seems neces-
sary, in order to sustain consistently the theory of survival, that a yellow
spiderling should have a yellow environment from the outset, and that
a white and pink spider should have a corresponding site from exode
to maturity. But, in point of fact, when we find the adult Misumena
upon a half opened rose, as in the cases above mentioned, we know that
the rose was opened up but yesterday, whereas the spider must have been
several weeks in maturing. This is true of all cases, or certainly of most
cases, in which we find adult spiders domiciled or ambushed upon flowers.
We are therefore compelled to the conclusion that the color did not
nourish the spider by providing for it a protective site, but that the
spider sought the flower and settled upon it, either accidentally or of
choice.

Epeira parvula is a spider remarkable for the variations it presents
in the dorsal markings of the abdomen. It is widely distributed through-
out the United States from ocean to ocean, and everywhere has

Hpeira the same characteristic. I have usually found it upon its orb

parvula.

waiting for prey, but like other Epeiras it undoubtedly rests —

upon adjacent objects. The Peckhams cite this species as an example of
protective resemblance in spiders.!. It is a common spider in Wisconsin,
and the Peckhams most frequently saw it on cedar bushes, where its color
harmonizes with the color of branch and fruit. During the day it usually
rests on the branch near its web. The back of the abdomen is a peculiar
bluish green, exactly like that of the lichens growing on tree barks. The
bluish color is broken up by waving black lines, which imitate the curl-
ing edges of the lichens. I reproduce the drawing given by the Peckhams
to illustrate this resemblance. (See Plate III., Fig. 5.) Undoubtedly, the
resemblance in this case is striking, but I take it to be simply an accident
of the situation. Parvula is found everywhere and upon all sorts of foli-
age, even where cedar bushes and lichens are not found. It is necessary
to remember this, although, of course, it does not gainsay the fact that
among Wisconsin cedar bushes it may have received some benefit from the
resemblance which the Peckhams note.

The suggestion has been raised that there may be some protective
value in the brilliant metallic colors which are possessed in a high de-
gree by some species of spiders. I have no observations to offer
on the subject, but quote a remark of Mr. A. G. Butler, of
the Kensington Museum. He says that metallic colors are not
a source of protection from birds, as birds know nothing of the nature of
metal, and whatever is brilliant and shining they make for at once, to see
whether it is good to eat.”

Metallic
Colors.

' Observations on Sexual Selection, page 83. * Jour, Royal Micros. Soe., 1889, page 633.

372 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Examples of cocoon mimicry are furnished by certain American Orb-
weavers, as, for example, Cyclosa caudata and Cyclosa bifureca. These
spiders make cocoons which in general shape and color closely
fine resemble the mother. The cocoons are hung in a connected se-
’ vies within the orb, a rather exceptional disposition. The mother
clings to the lower cocoon of the string, and might easily be confounded
with her cocoon. The conical shape of Caudata’s cocoon is paralleled by the
compressed apex of her abdomen, which has given her the name of the
Tailed spider. Her color also, a grayish white mottled with blackish mark-
ings, increases the resemblance between her and her egg sac, which is com-
posed of whitish silk covered over with the scalpage or débris of slaugh-
tered insects. Cyclosa bifurea is colored green, and her cocoon has a
greenish hue. (See Plate IV., Figs. 10, 11, 12.)

The suggestion has been made that placing cocoons of this particular
form within the limits of the spider’s snare, has a tendency to deceive
attacking insects, such as raiding mud dauber wasps or arachnophagous
birds. Professor Peckham alludes to the fact that Caudata, when a vibra-
ting tuning fork is placed near her, instead of remaining steadfast upon
her snare, drops from it in the way common to Orbweavers, and thus be-
trays her position and exposes her person.1. The implication is that, were
the resemblance really protective, the spider would have held steadfast
and not acted as she did.

On the contrary, it seems to me that this fact does not really break
down the force of the suggestion that such mimicry may be protective. For
we must conceive that a raiding bird or wasp, if deceived at all
by the appearance of the cocoons hanging in the snare, would
flutter from one cocoon to another until at last the spider would
be reached at the end of the string. The vibration of the wings of a bird
or insect would be the spider’s warning of the nearness of an enemy,
and her chance of safety would certainly be to drop from her web at
once. Of course, if the assailant should first strike the spider herself her
opportunity to escape would not be great; but supposing that there is about
an equal chance that the assailant would strike one of the cocoons, think-
ing it to be a spider, in that case the mother has a fair opportunity to
escape, and her chances are in proportion to the number of cocoons in the
string. Mr. Peckham’s experiment, therefore, instead of showing against
the suggestion that the cocoon mimicry is useful to Caudata, seems to me
to be entirely in harmony with it.

In this connection it is to be noted that the young of Caudata are in
the habit of hanging upon their orbs little puffs of silk closely resembling

Cyclosa
caudata.

1 “Mental Powers of Spiders,” page 395.

eee ar

374 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

the cocoons of an adult mother, and in the same position upon the orb.
These cocoons are also covered with the disjecta membra of minute insects

killed by the spiderlings. How shall we account for this strange
Young imitation of a maternal habit by the young? Are these puff
balls “dummies”? Is their purpose to deceive assaulting ene-
mies, and thus protect their maker? I know no examples of a similar
habit, except certain Gasteracanthas, that sometimes spread like objects at
various points upon their webs, and a single case of Acrosoma rugosa,
whose orb I found to be decorated in like manner.

Vig. 318 represents Caudata, much enlarged, clinging to a new made
cocoon, while two others hang above, covered with the disjecta membra of
slain victims. In nature the cocoons are often much more thickly covered
than here shown. A slight viscidity of the silken fibre of the sacs evi-.
dently assists this habit, although the scalpage is tied or lashed to the
surface by minute threads. By the time the maternal cares of the spider
are ended, if the season be one fruitful of insects, not only all the cocoons,
but the connecting parts of the supporting string, will be hung thickly
with this ghastly crop.

Dr. Martin Lister! observed the habit in Cyclosa conica, a European
species that corresponds closely with our Caudata, of thus stringing the
débris of its prey along the central vertical line of its snare. In
attributing the act to a sort of “pride of the chase” (venationis
gloriola), he gave a reason perhaps as near the truth as some other theo-
ries. It is at least sufficiently startling to find in the habits of an aranead
such a striking analogue of the customs of our savage human fellows who
decorate their persons, lodges, and villages with the scalps and skulls of
the unhappy victims of war and cannibal feasts. I am inclined to believe
that the habit is for the most part protective of the young, being intended
to guard the egg sac from the assaults of parasitic enemies. If so, it is a
convenient substitute for chopped straw, mud, gnawed wood, ete., with
which other araneads defend their eggs from enemies. But it has the dis-
advantage of depending wholly upon the somewhat uncertain chances of
the chase. ‘These chances, however, are the best, and indeed the only ones
at her disposal. The habit of suspending her cocoons within her viscid
orb well nigh estops her from descending to the ground or adjoining plants
to procure dirt or chippage, as species can readily do that attach cocoons
to various surfaces. I have seen only one case in which Caudata’s cocoons
appeared to be daubed with particles of mud. The general and special
habits are thus happily harmonized.

In addition to this the habit may also serve as a protection to the.
spider herself. At all events, as she hangs at the tip of one of these orna-
mented cocoons she is with some difficulty distinguishable from them,

Scalpage.

1 Hist. Animal Angl., page 33, 34, Tit. 4.

——— ll

MIMICRY IN SPIDERS. 375

since the colors of her body, as well as its shape, correspond well with
the colors of the egg sac. We might, therefore, regard this as a case of
protective mimicry. This scalpage is never in the shape of re-

picnic served stores of food, as Lister seems to think, and therefore
Leven ® cannot be cited, as by Kirby and Spence,! to show that Conica

blance. iS “more provident than its brethren.” Those distinguished en-
tomologists should have known that spiders do not feed upon
the hard shells of dead insects. :

This mode of disposing of the fragments of her feasts is not limited
to Caudata’s cocoons. Like her English congener Conica, as described by
Lister, she hangs those remnants upon her snare. I often see orbs through
the middle of which are stretched, above and below the hub, a perpendic-
ular ribbon of open fibre. Along this will be attached two or three little
conical balls above and below the hub (see Fig. 319), composed of the
members of dead insects cut into fine particles and lashed together by
threads. May we venture to suppose that this also is a
case of mimicry, that is to say, a purpose to set up “ dum-
mies” to distract the attacks of hymenopterous and other
enemies from her own person?

I have met a like behavior in Acrosoma rugosa. The
web was a small one, five inches in diameter, spun between

the branches of a fir tree. Around the margin

Acro- the remains of seven flies were threaded, much
ae in the fashion above described. Three of these
rugosa,

were above and three below the centre. They
appeared to be mere shells, not fresh insects trussed up for ,., 919 pats of
future use, as one often sees when flies are plenty. They — insect debris in
had not. simply become thus entangled when cast out from — S*ut™te’ssnere.
the web, for three of them were fastened above the centre, at which
the spider sits, and the four below were arranged along the are of a
circle in such order and position as to indicate design. I never met this
peculiarity in the snare of Rugosa except in this one case, and have not
observed anything like it in any other species. As a habit it exists in
Caudata alone.

The young of Caudata, as I have frequently noticed, have precisely the
same curious habit that marks the adults, to string along the central band
of their webs fluffy, loose bunches of silk, covered with little particles of
trapped insects, which increase in size as she grows. Mrs. Mary Treat has
observed the same habit.2 The transmission and early possession, in full
.force of such an exceptional and remarkable habit, is peculiarly note-
worthy, and, except on the theory of protective mimicry, it perhaps would
be difficult to suggest any useful purpose in the habit. A gentleman

1 Introd. Ento., I., 421. 2 “My Garden Pets,” page 42.
g

376 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

friend, who heard this statement, suggested that the spiderlings might be
“playing mother” and dandling their rag baby cocoons as our children
do their dolls!

Since the spiderlings thus have the habit, it may perhaps be consid-
ered as primarily for personal protection, and it is interesting to find it
transferred to the protection of the cocoon. Or, if we suppose that the
habit arose primarily to protect the cocoon, it is even more interesting to
think that it has been carried over by heredity to the young for their
own protection. It is impossible not to suspect that this habit may have
arisen from the prevalent custom of trussing up newly caught flies for food.

In the cases of Cyclosa caudata (Fig. 318) and Cyclosa bifurca (Plate
IV., Figs. 10 and 11) one must allow a striking resemblance between the
general appearance of the cocoons and the mothers
who make them. But when one comes to inquire
if the like resemblances prevail generally among
spiders, he finds that these two species, and a few
others, stand in a small group by themselves, As
shown in the preceding chapter on Color, there is
little resemblance between the great majority of spi-
der mothers and the cocoons which they make,
either in general shape or color.

However, it must be remembered that the value
of cocoon mimicry would naturally be limited to
those species which hang their cocoons in or upon
their snares, and to those which brood their cocoons
or watch upon or near them while the young are
being hatched. Of course, there would be no util-
ity in such a resemblance in species that make their
cocoons and forthwith abandon them or die. Ney-
ertheless, if we glance over the list of mothers that
Fic. $20, Collections of insect hang their cocoons in their snares, it becomes ap-

debts in the orb ofa young parent that even with them cocoon mimicry must
Cyclosa caudata. rae eer °

be limited. Argyrodes trigonum, as she hangs
bunched in her retitelarian web (Fig. 109, page 113), might be said to
have some resemblance to her basket shaped cocoon. The familiar The-
ridium tepidariorum is also a good example of resemblance between a
mother and her cocoons, both in general shape and color, (See Chapter V.,
page 112, Fig. 107.)

Uloborus plumipes is not unlike her cocoons as she hangs with
bunched and elongated legs beneath her orb. (See Fig. 104, page 109.)
In fact the Peckhams cite this species as an example of deceptive
resemblance.'| In Wisconsin these observers found Uloborus invariably

Vian J FE
SS ere; igany,
ASS
mS aii SB
<<
SS “SaaS LLL

PN
Ae

Observations on Sexual Selection, page 76, pl. iy.

——_——— SC oe . r ,

MIMICRY IN SPIDERS. 377

building in dead branches, where its wood brown or grayish color resem-
bles small pieces of bark or bits of rubbish entangled in deserted webs.
They also perceive a case of cocoon mimicry in her habit of so disposing
her grayish cocoons along the web as to look like a mass of rubbish. The
protective resemblance in this species, therefore, is twofold: that of the
spider to particles of dead wood entangled in its snare, and to the dry
branches among which the snare is spun; and again, that of the spider
to her cocoon. When Uloborus is found, however, as I often have found
it, in the midst of green laurel bushes or other verdant environment, the
fact of a protective resemblance disappears. If we concede the cause of
mimicry as urged by the Peckhams, we must go still further, it seems to
me, and suppose that the spider is endowed with a power, in one locality,
which forsakes her in another, and it may be a nearby one.

Theridium serpentinum! (Fig. 108, page 112), with her glossy brown
colors, can scarcely be considered as bearing a striking resemblance to the
snow white cocoons which she hangs within her snare; and Epeira laby-
rinthea (Fig. 85, page 100) can by no stress of imagination be reckoned
as bearing a resemblance to her cocoons.

1 This species Dr. Marx catalogues as a synonym of Teutana triangulosa Walck. “Cata-
logue of the Described Araneze of Temperate North America,” Proceed. U. S$. Nat. Mus., 1890,
No. 782, page 521.

CA AIP bt 2.

ENEMIES AND THEIR INFLUENCE ON HABIT.

We have considered the means by which maternal instinct secures the
life and growth of the spider young. It is necessary to study the natural
methods by which excessive reproduction is held in check, for Nature
presents to the observer a more or less continuous series of favoring and
adverse circumstances, a “ balance of power,” so to speak, by which on one
hand life is protected, and on the other is devoted to destruction. In
each case there is equal regard for the common good and the general har-
mony of Nature.

ure

The perils which beset the spider are many, serious, and diversified.
They besiege the very gate of being and cease not their relentless vigil
until the coveted life has been yielded. The “natural death ”
of the aranead is a violent one; comparatively few spiders, per-
haps, outside of those mothers who perish from inanition shortly
after the act of cocooning, have any other. It is this fact which compels
the great fecundity of the female, inasmuch as otherwise the species could
not be preserved.

It is possible for one to conceive how the protective habits which have
been heretofore described might have gradually resulted from the mother’s
struggle with her own enemies and those of her progeny. But it is far
otherwise when one asks, could this struggle have so reacted upon the
structure of the animal as to thus modify its organs of reproduction? If
no other obstacle presented, there would remain the seemingly insurmount-
able difficulty of accounting for the continuance of the species at all dur-
ing the long interval required for the supposed adjustment of the organs.
However that may be, we shall see that there is need for all the eggs laid
and young hatched, and all the protective instincts and industries by which
these ends are secured.

There is, of course, more or less irregularity in the operation of un-
friendly agents, which are themselves subject to laws of variation. In such
case there is a corresponding variation in the security of the species, and
so of their increase. The effects of a season unfavorable to spiders or

favorable to the growth of some enemy, or, on the contrary, advantageous
(378)

Diversi-
fied Perils

—— ee ——

ENEMIES AND THEIR INFLUENCE. 379

to the one and inimical to the other, may be seen in the number of ara-
neads in that or the succeeding year. This is also true of the abundance
or lack of a natural food supply. For example, the boat houses, fences, and °
outbuildings at Atlantic City fairly swarm with Epeiroids, especially Sclop-
etaria and Strix. This abundance is probably caused by the presence of
greenhead flies with which the district is infested and which, affording
an excess of food for the adult and partly grown spiders, relieves them
from the necessity of preying upon their own species, which thus increase
enormously as compared with sections a little distant.

But with these and such like exceptions, and notwithstanding all other
variations, the distribution of a given orbweaving species in a given sec-
tion will be found surprisingly uniform from year to year. The balance
of hostile and unfriendly influences is held well poised by Nature’s even
hand. The enemies of spiders
may be divided generally into
those which assail the animal
itself and those which affect
its eggs,

Among the enemies of spi-
ders, as of all other creatures,
may be placed the
changes of the sea-
sons. ‘The araneads’
power to endure cold is great,
but an unusually cold and
moist winter will destroy
many. Heavy rains prove fa-
tal, especially to the young,
and to females great with
eggs—beating down the foliage
in which they are ensconced, or sweeping the creatures themselves to the
ground. The extreme tension of the abdominal sac under the distended
ovaries makes fatal a shock that otherwise would work little harm.

It is well known that toads and lizards take kindly to a spider diet.
In southern Florida I once found a young lizard, while in the act of
shedding its skin, and with the white moult still adhering to it,
devouring a large Tetragnatha. (Fig. 321.) Many birds relish
spiders and pursue them at all seasons, plucking the Sedentary species out
of their very webs. In the autumn, when the broods of younglings are all
afloat upon their little aérial ships, swallows and swifts, birds that take
their prey upon the wing, have been seen skimming the tiny balloonists
into their bills as they coursed the air. A specimen shot for examina-
tion showed the accuracy of the observation by the presence of spiderlings
in the crop. It may be said in brief that all the larger animals with

‘Season
Changes.

Fic. 321. A moulting lizard eating a spider.

Animals.

i i es

380 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

insectivorous habits embrace the aranead nations also in their menu. Mon-
keys eat them; Hentz discovered a rat eating Oxyopes viridans; * and we
have an account, which may pass for what it is worth, of even sheep
upon the Steppes of Russia devouring certain ground spiders.?

Many spiders while yet immature fall victims to the voracity of their
own species. I have elsewhere considered the charge of cannibalism as
lodged against spiderlings while yet within the cocoon, show-
ing that as a rule their cradle life and earliest babyhood are
largely exempt from the perils of internecine hunger. But when
once the solitary habit of the race has compelled the individuals to sep-
arate and dwell apart, Nature relaxes her restraining influence and hunger
converts all available objects into legitimate prey. After this period it is
not possible to defend our aranead friends against the charge of canni-

balism, even of the most revolting
kind. Brothers and sisters eat each
other up without hesitation, and since,
naturally, fellow broodlings are likely
to pitch their tents and spin their
snares in closest contiguity, it comes
to pass that many of every brood are
devoted as sacrifices to the growth
and development of the few surviv-
ors to whom Nature has committed
the perpetuity of the species. Out-
side of these limits, everywhere, spi-
ders will prey upon their kind as
opportunity allows, even the hours
allotted to courtship and amatory em-
brace not being wholly exempt from
the perils of this general tendency.

Canni-
balism.

LiL

Perhaps the most persistent and
: destructive natural enemies of spiders
Fic. 822, A wasp plucking an Orbweayer from are certain hymenopterous insects be-
ee longing to the large family of wasps

known popularly 4s mud daubers and diggers.
It has often been remarked by ordinary observers that wasps can visit
a spider’s web not only with impunity, but as a successful assailant of
the occupant thereof. This fact has crept into literature, and is embalmed
by Goethe in a striking allusion to his father. ‘“Willingly,” he writes in
his autobiography, “as I have made myself familiar with all sorts of

1 Spiders of the U. S., page 46. * Walckenaer, Apt., Vol. I., page 172.

:
‘ |

ENEMIES AND THEIR INFLUENCE. 381

conditions, and many as had been my inducements to do so, an excessive
aversion from all Inns had, nevertheless, been instilled into me by my
father. This feeling had rooted itself firmly in him on his trav-

Goethe els through Italy, France, and Germany. Although he seldom
ge ronbar. poke in images, and only called them to his aid when he was
Wasps. ‘I Bes, y sie eae

very cheerful, yet he used often to repeat that he always fancied
he saw a great cobweb spun across the gate of an Inn so ingeniously
that the insects could indeed fly in, but that even the privileged wasp
could not fly out again unplucked.” But the number of those who, hay-
ing observed the scathless incursions of “the privileged wasp” into cob-
web domains, also know the purpose thereof, is exceedingly small. Yet it
is inspired by one of the most common and interesting instincts in the
insect world.

If we follow the wasp a little space backward from her cobweb raid, we
shall see her fluttering over the muddy margin of pond, puddle, or stream.
She is seeking mortar, which, gathered between her mandibles, she carries
away through the air. Following her flight, we find her engaged upon the
broken face of a cliff, the rugose surface of a wall, or the rough boards
or beams in angle or cornice of some house, stable, or
outbuilding. She carefully spreads her mortar, smooths
it, rounds and arches it, until, after many successive
visits to the mud bed, she has built a cell about an
inch long and three-eighths to half an inch thick.
(Fig. 328.) The middle of this cell is a hollow cylinder,
within which the mother wasp, for such the little ma-
son is, deposits a single egg. It is at this point that
the raids upon spider webs begin. The egg in course
of time is to become a ravenous, flesh eating worm, an
arachnophagous larva; a soft, legless, whitish maggot,
with a somewhat horny head and a strong pair of
jaws, but no other weapons whatever. The food which
Nature foreordains for it is living spiders, and those spiders the mother
proceeds to capture and entomb within- her mud daub nursery. On this
errand she may be seen hawking over and near cobwebs of various sorts,
venturing within the meshed and beaded snares that prove fatal to most
incomers, and sometimes even to herself. She rarely fails in her errand.
If the aranead occupant, expectant of prey, sallies forth to seize the in-
truder, it finds itself a captive, not a captor. For the wasp shakes the
silken filaments from feet and wings, turns upon the spider, seizes and
stings it, bears it to her cell, and thrusts it therein.

She does not Jimit her hawking to cobwebs, but flutters over flowers,
burrows among leaves, creeps with nervous, twitching tread along branches
of trees, wherever spiders dwell or hunt, and with relentless cunning, zeal,
and ferocity snatches those creatures away to add to the growing store

Fic. 323. The nidus of
the mud dauber wasp.

382 AMBRICAN SPIDERS AND THEIR SPINNINGWORK.

within her egg nest. At last the cavity is filled, the circular opening
sealed up, and the spiders left literally entombed alive within that clay
sarcophagus.

If one at this stage should break open the mud dauber’s cell, he might
dispute the statement that the imprisoned spiders are alive. To all ap-
pearances they are dead. In point of fact they are simply paralyzed. The
effect of the poison injected by the wasp’s sting within the tissues of her
victim is such that all activity is at once and completely suspended, with-
out destroying life. Thus, when the larval waspkin awakes to the pangs
of hunger, it finds itself in the midst of a generous supply of the very
food which Nature intended for it. The mother whom it is never to
know, and who already perhaps has paid the last debt to Nature, had con-
sumed her closing days in providing for the offspring which she was
never to see. I have found these larve, fat, white grubs, in the midst
of their “preserved meats,” feasting thereon, and have wondered at their
enormous appetite and the greedy vigor with
which it was satisfied. (Fig. 324.)

Thus, before the era of man, Nature, in the
J person of a wasp, had solved the problem of
preserving animal flesh without impairing its
value as food. A like discovery by the human
species, with due application to the edible do-
Fic. 824. A athe feeding upon mestic animals, would solve an important prob-

lem in commercial economy which has only
been distantly approached by the ice chambers within which great trans-
portation lines convey butchers’ meats.

It would be interesting to know the nature of the poison which pro-
duces such remarkable effects, but one cannot hope that it will ever be
procured in sufficient quantity to permit analysis. How long
the virus may preserve its peculiar effect before death results,
or whether a spider once stung can recover health, and to what
extent sensation is retained, have been points of inquiry and of some ex-
periment, On two occasions I kept under observation spiders rescued
from the jaws of wasps. One specimen was a species of 'Tubeweaver,
which I took from a blue wasp; it lived about twa weeks. The other
species was a large female Wolf spider, taken by a friend and sent to me
October 5th, 1875. It lived until the 17th; twelve days. During this
period the creature remained entirely motionless and the limbs retained
any position in which they were placed. These examples indicate that
there is no recovery from the poison, and that death is suspended for
about two weeks.

I do not know the exact period required for the deyelopment of the
wasp egg to a feeding larva, but it is something longer than two weeks.
Tn some cases I have found the spiders within the wasp’s nidus dead and

Wasps
Poison.

ENEMIES AND THEIR INFLUENCE. 383

shriveled, the egg probably having proved infertile. Again, a few spiders
would be dried up, while others were plump and edible, a condition in
which more frequently most of them are found. It is certainly one of
the unhappy possibilities in the destiny of the spider that it may be con-
strained to abide in a living death within this dark vault awaiting the
awakening appetite of a voracious worm. It is to be hoped that a kindly
Nature has so far tempered this hard doom as to deprive the entombed
creature of all consciousness of her condition and consequent suffering
therein. Indeed the evidence is well nigh conclusive that sensation is
wholly suspended at the prick of the insect’s sting.

ITI.

With the single exception, perhaps, of one small order, Neuroptera, no
order of insects is exempt from the attacks of the all devouring wasps.
Some provision their nests with grasshoppers, some with cockroaches, some
with snoutbeetles of various kinds,
some with ants and bees, a few with
different kinds of bugs, frog spittle,
insects, and plant lice; a great num-
ber of them with various kinds of
two winged flies, and a still greater
number, perhaps, with the larve of
various moths.! Most observing coun-
try lads have noticed the assault of
the handsome digger wasp, Sphecius
speciosus Drury (Sphex), (Fig. 325),
upon the so called “locust,” the cicada or harvest fly, and I have dug
that insect, Cicada pruinosa, out of a burrow of this wasp in the terrace
of a West Philadelphia yard.

Those wasps which prey upon spiders comprise many distinct species
belonging to widely separated genera. Some of these gather many spi-
ders into one cell, others only one. The insects heretofore noticed are of
the former class, the species most destructive in this region being prob-
ably the common indigo blue mud dauber, Chalybion ceruleum Linn.
(Sphex). (Plate V., Fig. 7, natural size.) The larval cells of the blue
mud dauber are commonly laid in small masses, one on top of another.
(Fig. 323.) The cells of the common mud dauber are composed of one
or more layers or tiers of clay tubes, arranged one above another or side
by side like a set of Pan’s pipes, and cemented to some surface protected
from the weather. One such specimen, collected in the autumn (Fig. 326),
I kept in my cabinet, and about the beginning of July following, a num-
ber of black digger wasps, T'rypoxylon politum Say, escaped therefrom.

Fig. 325. The Cicada wasp (Sphecius speciosus.)

* Walsh, American Entomologist, Vol. I., No. 7, 1869, page 126.

384 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

I obtained no other species from these nests, but cannot affirm that no
other escaped.! It may be a question, perhaps, whether the mud daubs
were made by Chalybion or Trypoxylon; but we have the great authority
of the late Benjamin D. Walsh that the latter species is really a guest wasp,
not building and provisioning any nest for itself, but laying its eggs
in the nest built and provisioned by the former, thus appropriating
for its own future progeny the spider store laid up by the industrious
Chalybion for its young.? It is curious and suggestive to trace this use
and wont from the guest wasp and the cuckoo up to the human species
as represented alike by the imperial “annexers” of Europe, Africa, and
the Orient, and the “land grabbers” of the Indian Territory, the “squat-
ter sovereigns” of the border, and the
“claim jumpers” of Rocky Mountain
mining districts.

Among the wasps that provision
their nests with single spiders is the
common blue digger wasp, Chlorion
veruleum Drury (Sphex), which, un-
like species hitherto alluded to, bur-
rows in the earth. It excavates its
egg nest in an incredibly short time,
sometimes consuming not more than
a minute or a half minute, and then
places therein a single egg together
with a spider, which is generally a
large one. With its front pair of feet
it then scrapes back the dirt which
it had withdrawn, frequently stopping
to pat it down with its abdomen.
When the hole is filled the surface
is smoothed to the level of the sur-
rounding soil. The large and beauti-
ful Elis 4-notata Fabr. (Scolia), (Plate
Po akin ot ips te a ae as, i a) dna NR

Lycosids, especially Lycosa tigrina,
and the small Priocnemus pomilius Cresson has been taken while carrying
a Laterigrade, a species of Xysticus, in its jaws.

Another example of wasps that store single spiders is the large and
beautiful Pepsis formosa Say (Pompilus), an inhabitant of the
Southwestern States of North America, where it is popularly
known as the “tarantula killer.” (Plate V., Fig. 2.) This
name is given because of its habit of storing its burrow with that most

Tarantula
Killer.

1 The figure here given (Fig. 326) was drawn from a series sawed out of a shed at Bell-
wood, Pennsylvania. * American Entomologist, Vol. I., page 183.

—_—s l

ENEMIES AND THEIR INFLUENCE. 385

formidable of our spider fauna, Eurypelma hentzii. (Plate V., Fig. 1.) I
have seen this insect in Texas hawking for its gigantic victim, which
showed by its hurried and excited action full consciousness of its peril as
it fled with eager and trembling speed before its pursuer. The late Pro-
fessor Buckley, of Austin, describes an encounter on Texas soil between
these two formidable creatures. ,

. The tarantula killer is a bustling, unquiet insect, always in motion, fly-
ing now here, now there, and when running on the ground its wings are in
a state of constant vibration. Should it discover a tarantula it begins in-
stantly to fly in circles in the air around its victim. The spider, as if
knowing its fate, trembles violently, standing up and making a show of
fight, but the resistance is feeble and of no avail. The spider’s foe soon
discovers a favorable moment and darts upon the tarantula, whom it
wounds with its sting, and again commences flying in circles. The injured
spider is thrown into a tremor, and often becomes at once paralyzed,
though the influence of a second, and even a third, wound is sometimes
necessary. Sooner or later the spider becomes powerless when the victor
approaches, carefully feeling its way to see if its work has been effectually
done. It then begins to drag the tarantula into a hole which it has pre-
viously dug in the ground, wherein it is covered up after the deposition
of an egg.'| The courage and address thus shown in assault upon so for-
midable an animal, and the strength and perseverance required for its sub-
sequent entombment, are of the highest order and surely evoke admiration,
however much we may pity a foe doomed to so hard a fate as to be par-
alyzed, buried alive, and afterward devoured by a greedy grub.

In estimating the ravages wrought among spiders’ by the various tribes
of wasps, it must be remembered that in the above and all like cases, the
mother wasp, although depositing but one egg in each nidus, has a num-
ber of eggs, more or less, to dispose of. As she never ceases her’ work
until every egg is duly deposited and its future offspring provided for, the
vast destruction carried into the aranead hosts during the period of ma-
ternal activity may better be imagined than expressed.

IV.
The thought had occurred to me while examining the contents of mud

daubs, that certain species of spiders were preferred by the wasps as pro-
vision, and that possibly certain species of wasps affect certain

alee spider species. In case of the true Diggers, who store but a sin-
ae Shes gle individual, there is no doubt a narrower range for selection,

and even a specific choice, as with the Tarantula killer and the
ee apaeyet aaa But not so with mud daubers. I have found avery

' Proceedings Amer. Ento. Soc. (Philadelphia) Vol. I., page 188. See also an account by
Dr. G, Lincecum, Amer, Entomologist, Vol. I., No. 6, page 111.

386 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

ereat group of our indigenous spider fauna represented in broken cells.
The most numerous are Orbweavers, many species of which I have col-
lected from mud dau

It is, however, true that in the gatherings of any individual wasp there
is apt to be a preponderance of a single species of spider. Thus it would
seem that a wasp starting out with Epeira strix, for example, is apt to
devote herself chiefly to collecting that particular species. So with other
species. Can it be that the mode of capturing her first victim, itself some-
what a matter of chance, so impresses itself upon the “brain” of the wasp
that she almost mechanically drops into the same mode for subsequent
capture, and thus finds herself habitually hunting along the trail of the
domicile and hiding place uncovered in her first capture ?

Next to Orbweavers, Laterigrades perhaps have been oftenest found
by me in mud daubs. The species most frequently seen is the large yel-
low, white, or variegated species, Misumena vatia, which lurks for its prey
on flowers, the mimicry of whose colors (see Chapter XII.) seems, in this
case, not to be a “protective ” resemblance. Here again, where this spider
is found it usually predominates, as though the wasp, making her original
capture upon the habitual feeding grounds of the species, had gained
what might be called an “experience” and followed in the lines of her
first finds.

I find some confirmation of my own impression in the opinion ex-
pressed by Mr. John Abbot, who observed the habits of American spiders
in Georgia as early as 1792. He says that wasps generally confine their
hawking to one species, when in search of spiders with which to store
their mud daub nests.1

If we now turn from the more arboreal species to those which during
the day will be found chiefly upon their webs, we shall observe a strong
tendency in the same general line of habit. That vigorous and destructive
Retitelarian, Theridium tepidariorum, so common in our outhouses, when
found within a mud daub will be the prevailing species. Such Orbweayers
as Argiope argyraspis, Tetragnatha extensa, Epeira labyrinthea, or Argy-
roepeira hortorum, which habitually hang upon their webs and must be
thence seized by the raiding wasp, I have found subject to the same gen-
eral tendency. It is needless further to multiply examples. I disclaim
the purpose of indicating here an inflexible conclusion, or even one sus-
tained by’ satisfactory evidence. But the facts which have fallen under
my notice do justify one, if not in inferring, at least in suggesting, what
future observers may find worthy of careful study. The line of inquiry
certainly points along fields full of interest.

A brief reference to some of the special characteristics of a few of the
spider species preyed upon by the mud dauber wasps will give a better

1 Walckenaer, Hist. Nat. des Insectes, Aptéres, Vol. I., page 174.

ENEMIES AND THEIR INFLUENCE. 387

idea of the skill and acumen of these creatures in their raids. For ex-
ample, there is no species with stronger secretive tendencies than Epeira
strix. Its ordinary hiding place in a rolled leaf is so carefully
Charac- selected and separated from its snare that I am continually
teristics thwarted in search for it. Yet the mud dauber finds it. So with
of Cap- 5 i ’ : eke
Fives: the Laterigrade spider Misumena vatia. Its mimicry of the va-
rious colors of the particular flowers upon which it lurks, is sur-
prisingly exact, although for the most part it affects yellow and pinkish
white colors, Yet it is precisely this species which the wasp, in her in-
dustrious quest among leaves and blossoms, most frequently falls upon.
I confess myself equally puzzled and interested at the facts which here
present themselves. If one were at liberty to do so, he might fancy that
this curious hymenopter feels some trace of that noble rage which inspires
the breast of the huntsman, and, scorning more inglorious game, devotes
herself to that which most excites her enterprise and evokes her skill. I
have admired the intensity of action shown by the blue mud dauber when
hunting spiders among bushes. It fairly jerks itself along from leaf to
leaf and from stem to stem, prying under every corner and thrusting its
antenne beneath leaves, peeking into every cranny, angle, and nook where
a spider could possibly be reposing. I do not wonder, after watching one
of these creatures stalking its prey, that even the most secretive of our
araneads fails to escape the detective skill and quenchless ardor of the
remorseless insect.

The solitary wasps, diggers, and mud daubers are not the only ones
whom maternal instinct makes hostile to spiders. The social or paper
making wasps may be included in the same list. The digger
wasps appear to feed upon vegetable matter exclusively, although
they provide animal food for their larve. It is difficult to ac-
count for the development of such a habit and such a taste. How could
the insectivorous habit have come to a larva! by heredity from a nectar
feeding ancestry? On what principle can one explain why a mother with
such a taste should provide for a sarcophagous offspring? Evolutionism
has here a series of facts that lay formidable obstacles in its path.

If, now, we could show in the digger wasps some such facts as appear
in the life of the social wasps, we might, perhaps, escape the difficulty.
These insects also feed upon the honey and pollen of flowers, but the
opportunity to acquire a taste for animal food is sufficient, for they di-
rectly feed their larvee as do bees and ants, not leaving them to serve
themselves as do the young of the mud daubers. That food consists
chiefly of desiccated insects, but spiders contribute a portion to the larval
bill of fare. The assaults of hornets upon the flies swarming in country
kitchens are well known to American housewives; the webs of spiders
are raided for the same purpose. ‘These captives are chewed into juicy
pulp and fed by mouth to the white worms that occupy the regular cells

Social
Wasps.

388 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of the beautiful paper nest. Now, in the act of reducing spider flesh to
pulp it is natural to suppose that a taste for such food might be ac-
quired (and, perhaps, it is even gratified) in sufficient strength to lay
the foundation, at least, for an insectivorous habit in the progeny.

But our mud dauber does not feed her own larve at all; the far away
originals of her species could have had no reasonable origin for a faintest
suggestion of arachnophagous necessity in her progeny, and how then did
she begin her persistent harvesting of spiders? It is, perhaps, possible
to conceive that it may have come by the long, roundabout way of an
insect chewing hornet or rust red wasp, but whether it is worth while to
go so far to get so little, the advocates of the development theory must
consider. The point in which the author is here specially interested is
that the social wasps also are to be ranked with the enemies of spiders.

Ni;

It has been stated that all spiders are addicted to cannibalism, no
species scrupling to prey upon individuals of its own order. As a rule,
however, every species takes aranead prey, as it does insects,
Nest Par- otter its own characteristic modes, But the habit of cannibal-
asitism. . : : , F 5
ism has a peculiar manifestation in the case of several species
belonging to the Retitelaria. One of these is a beautiful California spi-
der, first sent to me by Mr. ©. R. Orcutt, which is conspicuous by bright
metallic silver markings upon a black triangular body. It is a small
creature, but is evidently possessed of unusual cunning and ferocity. My
knowledge of its habits is received from Mrs. Eigenmann, who forwarded
to me living examples of both sexes. I have named the species (in lit-
teris) Argyrodes piraticum.! The spiders were established on what seemed
to be foundation lines of their own, which were attached to the broad
foundation lines of a large orbweb of a species of Epeira. In one case
an Argyrodes was found in the act of preying upon a large Orbweaver
which she had encased within a silken enswathment and trussed up on
its own web. It seemed remarkable to the observer that the
The little silvery spider could slay and eat a creature so much larger
Pirate : : : ;
Spidey. than itself, and, indeed, nothing short of actual observation would
justify belief. My informant has found this pirate spider upon
the snares of Gasteracantha, Argiope argenteola, and Zilla x-notata, as
well as upon orbs of various species of Epeira. It takes its station quite
habitually upon the outskirts of the snares of these Orbweavers, from
which point it makes its raid upon the lawful owner of the web, and
perhaps, also, as Mrs. Eigenmann thinks, feeds upon the excess of insects
which may often be found adhering in considerable numbers to the viscid
portions of the orb.

1 Tt has probably been described, but I cannot identify it.

———

ENEMIES AND THEIR INFLUENCE. 389

This genus is represented by a species, Argyrodes trigonum,! common in
the neighborhood of Philadelphia and throughout the Eastern States. It
makes the usual nest of crossed and netted lines common to its

ee . tribe, but is also parasitic in its nesting habits, for I have found
eitnet it on the retitelarian section of the Labyrinth spider’s web,

where it had made itself very much at home. I have also found
it upon the webs of other species, as Linyphia communis and a small The-
ridium, and the upper intersecting supports and lines of Agalena neevia.
Mr. Emerton? has observed the same tendency to nest parasitism, having
found 'Trigonum in the upper part of the web of Linyphia scripta and
also among the upper cross threads of Agalena nevia. There is no record,
however, as far as I know, of this species actually preying upon its hosts,
and the creature must be endowed with unusual cunning if it really suc-
ceeds in doing so.

The most decided example of this particular habit is found in a Line-
weaving spider described by Professor Hentz as Mimetus interfector. Hentz
found the species in Alabama; I have found it in Pennsylvania,
Ohio, New York, and elsewhere; and Emerton has collected it in
Massachusetts and Connecticut. According to Mr. Simon the
species also occurs in southern Europe. Hentz says that Inter-
fector spins a web resembling that of Theridium, but prefers prowling in
the dark and taking possession of the snares of Epeira and Theridium after
murdering the proprietor.

This singular depredator is not rare, and was usually found by its dis-
coverer in houses, which enabled him to make many observations upon
its manners. The first specimen observed was a female, which had made
two cocoons under a table in his study, near and among the webs of sey-
eral individuals of Theridium tepidariorum. The cocoon of Mimetus is
oblong, and tapers equally at both ends, which are secured by many
threads connected with a retitelarian web. The mother was watching the
young, which were issuing from the lower one of her two cocoons. Thus
she appears to possess in a strong degree that maternal solicitude which
marks so many of her order.

A second observation discovered a very different state of feeling as to
the young of other species, for she was observed devouring the eggs of

: Theridium tepidariorum, most probably after having eaten the
ee mother. A third specimen was found dead in the web of some
species of Theridium, which no doubt had killed it, an illustra-

tion of the fact that sometimes in her predatory expeditions she man-
ages, like human robbers, to “catch a Tartar.’ A fourth Interfector was

Mimetus
inter-
fector.

' Argyrodes argyrodes Wlck.

* Notes in Hentz’s Spiders U. S., page 153.

5’ New England Spiders, Family Theridide, page 17.
* Arachnides de France, Vol. V., page 29.

390 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

found eating the same Theridium that had devoured her predecessor.
Such observations indicate a marked hostility existing between these two
powerful examples of their family—a sort of aranead feud.

Professor Hentz sometimes enclosed specimens of these spider eaters
with other araneads in a glass jar, in order to watch their motions. ‘The
moment another spider was thrown in, Interfector showed by its
attitude that it was conscious of the presence of an enemy. For
a moment it moved its first and second pairs of legs up and
down, and then slowly approached its victim, and generally killed it. A
Theridium tepidariorum thrown into the jar manifested great terror, but
in a little while, or, as Professor Hentz puts it, “after some seeming reflec-
tions upon fortitude and necessity,” it prepared for the mortal combat, and
cautiously advanced towards the Mimetus, which began to move more
slowly. Theridium, when near her adversary, threw out a long thread on
which were several globules of transparent fluid. This partially succeeded,
for Mimetus was caught by one leg, and while Theridium retreated for ob-
servation it was dragged about for a long time before it succeeded in free-
ing itself. The battle presently was renewed, and this time Theridium was
conquered and eaten.!

From these interesting observations it appears that this formidable spe-
cies of Theridium is a favorite object of attack with Mimetus. It is one
of the most powerful and ferocious of its kind, being able to overcome and
destroy the largest insects, and, as we have shown (Vol. I., Chapter XIII),
even to entrap and destroy a small snake and a half grown mouse. The
courage, ferocity, and combative skill of the adversary which is able suc-
cessfully to meet and vanquish it are thus at once manifest. I have
found Mimetus interfector parasitic upon the snares of other spiders in
the neighborhood of Philadelphia, but have never witnessed an actual at-
tack by her. On one occasion I found her ensconced upon the snare of
an Orbweaver, having evidently destroyed the occupant.

Mimetus syllepsicus, according to Hentz,? has the same piratical habit
as her congener Interfector. This spider was found in the tent of Epeira
labyrinthea, which it had no doubt killed. The webs and cocoons of its
victim were untouched, and the squatter seemed perfectly at home in its
new domicile. It strikes one as a remarkable development of instinct
which has formed within a tribe and families having fixed sedentary
habits a disposition to leave the snare and go, like the Wandering groups,
to seek prey, and especially to raid the nests of fellow araneads therefor.
But it may be noted that the piratical destroyer confines its raids
to species whose nesting habits, in whole or in part, are sim-
ilar to its own. That is to say, Mimetus finds its best pre-
serves and hunting grounds upon the netted cross lines of Theridium and

Spider
Duels.

Origin of
the Habit.

1Spiders of the U. 8., page 138. 2 Spid. U. S., page 140.

. by Mai
wae lS

ENEMIES AND THEIR INFLUENCE. 391

Agalena and the mazy annex of Labyrinthea, or the foundation lines of
Orbweavers, which somewhat resemble her own retitelarian forms. The
same remark applies to Argyrodes piraticum. It is found upon the com-
pound snare of Zilla, which combines the orbweb with the retitelarian,
and stations itself for its piratical raids upon the netted outlying and
foundation lines of other Orbweavers. The same observation holds good
as to Argyrodes trigonum,as far as its nesting parasitism is concerned,
for she finds congenial raiding grounds within the netted lines that over-
hang the funneled sheet of Agalena and the orb of Labyrinthea. Un-
doubtedly she feels more courage, confidence, and vigor within an environ-
ment which gives her the sense of being upon her native snare.

It is possible for us to conceive that the habit may have originated
from the facility thus presented for obtaining a foothold and home upon
the webs of its neighbors. Having done this, it only needed a favoring
opportunity, many of which must have presented, to throw the host of
the web into the power of its guest. An act so advantageous would be
likely to be repeated and persisted in; we have thus an easy way to the
development of a fixed habit from what at first may have been an acci-
dental feature in the life of ancestors. Even if this conjecture should be
adjudged plausible, I cannot free myself from the wonder that so mani-
festly convenient a mode of securing food should have been fixed upon by
so few of all the numerous species and innumerable individuals of the
great Retitelarian tribe.

We pass now to note the parasitic enemies of Orbweavers and others
of their order. Mr. Blackwall has shown that immature spiders, Epei-
roids and others, are infested by the larvee of Polysphincta car-
bonaria, a hymenopterous insect belonging to the Ichneumonide.
This parasite is always attached to the upper part of the abdo-
men, near its union with the cephalothorax, and, although it proves a
source of constant irritation, is secured by its position from every attempt
of the spider to displace it. Being without feet, it appears to retain its
hold upon its victim solely by the instrumentality of the mouth and a
viscid secretion emitted from its caudal extremity. But one larva has ever
been seen upon a single spider.

The ichneumon probably deposits its eggs on the spiders in the au-
tumn, attaching one egg to each individual. In the spring, towards the
end of May, having gone through its final moult and increased consider-
ably in size, the larva becomes restless and rapidly destroys the spider,
which it abandons after having reduced it to a mere corrugated skin. It
then attaches itself to some convenient point, the cork of a bottle if it be
in confinement, and begins to spin its cocoon, which it completes in a day
or two. This cocoon is of yellowish white silk of compact texture, and

Body
Parasites.

tee | ze

392 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

measures one-third of an inch in length and one-tenth in diameter. It
is of an oblong quadrilateral figure tapering to its extremities, one of
which is more pointed than the other. It is lashed to its site by numer-
ous fine silken lines. In about one month the perfect insect appears.
Blackwall found this parasite on four species of Epeira and two of Liny-
phia.'| The same author figures Hermeteles fasciatus and H. formosus,
ichneumon parasites on Agalena brunnea.?

A correspondent of “Science Gossip” gives an interesting note with draw-
ings. of an ichneumon wasp larva that preys upon a small spider in Cey-
lon, India. The spider usually attacked is a small black ani-
mal, with globose abdomen, that spins a loose irregular web on
the under surface of leaves. The ichneumon wasp appears to
oviposit upon female spiders only, the males being much smaller and
unable to support the wasp grub. The egg is fixed to the abdomen of the
spider close to its junction with the cephalothorax. The newly hatched
larva immediately pierces the skin, and commences to absorb the juices
of its host. The spider continues to feed, and re-
mains apparently in good health until the parasite
is full grown, when the latter destroys its victim,
leaving nothing but the empty skin. The larva then
commences to spin a flask shaped silken cocoon, at-
tached generally to the under side of a cinchona
leaf. It builds up the cocoon gradually, complet-
7 ne Pochette ing the walls as it proceeds, forming first a cup

shaped receptacle, which is lengthened by regular
additions to the open edge, and finally closed. A specimen under observa-
tion completed its work in forty-eight hours.*

It is an interesting fact, to which Blackwall has called attention, that
immature spiders infested by the larva of Polysphincta carbonaria do not
change their skins. In what way the parasite can affect the animal thus
to cause a suspension of so ordinary a function is not known, but the .
economy of the fact is apparent. If the moulting were to proceed, the
parasitic larva would probably be cast off with the skin and would inevi-
tably perish, thus causing a failure of its manifest end in Nature, which is
to conserve the life of its kind both directly and indirectly—directly by
feeding upon the body of the spider, and indirectly by checking the undue
increase of that deadly enemy of insect tribes.

Spiders are also attacked by parasites within the body. A full grown
specimen of Epeira cinerea,® sent me by Mrs. Treat, had been dropped into
alcohol to kill it. Immediately there issued from the abdomen a white

An India
Parasite.

‘Ann. and Mag. Nat. Hist., Vol. XI., 1843, pages 1 and 2, and Spiders Gt. Br. & In,
page 352. 2 Spiders Gt. Br. & Ir., pl. xii, A A, BB.

* KE. Ernest Green, Science Gossip, July, 1888, pages 159, 160.

* Ann. and Mag. Nat. Hist., Vol. XI., 1843, page 4. ® Epeira cayatica Keys.

’ Larve.

ENEMIES AND THEIR INFLUENCE, 393

ichneumonid larva one-half inch long. The spirit bath had evidently dis-
turbed the creature when near the period of emerging, and being affected
; by it, it at once cut a way through the skin of its host, and
Parasites wriggled out of the body into the alcohol, where, of course, it was
Within destroyed. ‘The site of this lar its host corresponded
the Body, “stroyed. e site of this larva upon its hos p
closely with that of the parasitic guest described by Blackwall,
on the upper part of the abdomen. In size the two larve are nearly equal,
and they probably belong to at least the same genus,

Menge has added to our knowledge of the parasites infesting spiders,
and I present a brief abstract of his observations.! Micriphantes, The-
ridium, Bolyphantes, and other species found crawling on the
ground are infested with a cinnabar red species of Dermanyssa.
But one of these is usually found on an individual, seldom two,
and hardly ever three. It is fatal to the smaller varieties alone, and only
infests the larger varieties while young. He found Mermis allicans on a
Water spider (Argyroneta aquatica), and saw it, escape from the body and
tumble about in the water. During the latter part of June he took a fe-
male of Clubiona putris within its little silken sac on a stem of heath, and
confined it for observation. A week thereafter it had disappeared, while
within the sac, on a few horizontal threads suspended in the centre, lay
a yellowish white pupa about five millimetres long, which had eaten the
spider except the legs and a small part of the skin. In another week a
winged insect, probably Henops marginatus or Oncodes pallipes, emerged
from the pupa. During the brief adult life of this insect it takes no
nourishment, but soon finds its mate and deposits its eggs upon the spider.
Immediately after hatching, the maggot makes its way into its host’s body,
probably through the rima prudendi.

The same author describes two other parasitic larvee which he failed
to bring to maturity, but which probably also belong to the Hymenoptera.
The first was found August 27th, upon the posterior part of a
full grown female Arctosa cinerea, taken in the sand under
fallen leaves; it was naked, reddish white, without feet, two mil-
limetres long. It astonished the observer to note that the large spider re-
mained perfectly quiet while the larva nibbled its way into the body,
when a movement with the legs would have removed it. On the second
day the spider was quite dead, and the larva was then four millimetres
long. Eight days thereafter the larva had devoured the entire abdomen,
the inner cephalothorax, and the thigh of one hind leg; it had increased
about one-half in size, had satisfied its hunger, but was very uneasy. It
was placed in sand to mature, and there overspun itself, but never further
developed.

Another larva was found, July 28th, on an immature male of Miranda

Menge on
Parasites.

Parasitic

1 Menge, Preussische Spinnen, under Parasites of the Spider.

394 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

adianta, a species of Orbweaver. The parasite was two millimetres long,
a yellowish green color, and smooth, except slight warts upon the back.
At first the spider seemed to feel little inconvenience from its guest, but
on the fourth diy it sat motionless, and on the fifth it had been de-
voured, only a small bit of skin remaining, while the caterpillar lay
curled up in a half circle, grown to twelve millimetres in length and two
millimetres in width across the middle of the body. Subsequently it made
a cocoon on the heath, but developed no further. None of the above
larvee moulted, which, according to Menge, is characteristic of parasites
within the body, a habit that shows quite as striking adaptation as does
freedom from moulting in an infested spider.

One of the most common superstitions heard among persons unin-
formed in natural history, is that a horse hair, when placed in the water
under certain conditions, will turn into a snake. I have heard
this fact averred by eye witnesses, who believed confidently that
they had seen the hair: suddenly come to life and wiggle off through the
water. This astonishing statement is explained by that
most interesting and least enjoyable of natural facts,
parasitism. Crickets, grasshoppers, and spiders are
known to be the hosts of a species of our common
Gordius, the same probably as that described a num-

in oi!” ~ piesa: “2 DET OF years.ago by Prof. Joseph Leidy as Gordius
Parasitic Gordius (Fra.328) aquaticus.! From this eminent naturalist I have re-
rere Pet seutu- ceived a specimen of Lycosa scutulata, from which a

eng eye Gordius was taken. I have figured the spider just as
I received it, it being very much damaged when it came to my hand;
the parasite is also drawn, both figures natural size. (Figs. 328 and 329.)

Gordius.

VII.

We have thus far considered the foes which assail the life of the
spider after it has escaped from the cocoon. This does not complete the
doleful record. Her cradle life is beset by even more formida-
ble perils. The maternal instinct which, in the spider mother,
prompts to cunning protection of her eggs in admirably wrought
cocoons, inspires the Ichneumon fly to penetrate the silken bars and wards,
and place the eggs of her parasitic young upon the spider’s eggs. Our
knowledge of the parasitic Hymenoptera preying upon cocoons of spiders
has yet to be much enlarged, but we know that the genus Pezomachus is
one of the most persistent guests, and that she carries vast ravages into
the aranead ranks. A few notes will be given, culled from many observa-
tions upon the destructive habits of this genus.

Parasites
on Eggs.

1 Proceed. Acad. Nat. Sci., 1850-51, page 98.

| '
—— ll

——s Tl eon

ENEMIES AND THEIR INFLUENCE. 395

The outer case of a cocoon of Argiope cophinaria was taken at Atlantic
City during winter (1883), and opened June 6th following. It was then
pierced with several round holes. The lower part of the flask

sae was occupied by a number of white cocoons of a parasitic hy-
arasit- : = :
ian menopterous insect. They were each three-eighths of an inch

long, were grouped in a bowl shaped*mass quite around the

bottom of the egg sac, and were covered with a delicate white silken floss.
Many of them were pierced at one end with a hole corresponding with
the one on the outside of the spider’s cocoon. (Fig. 330.) From these
holes the insects, probably a species of Pezomachus, had made their escape,
leaving their mahogany colored shells within their white pupa cases. Some
of the Ichneumonid cocoons were without the single large opening, but
had minute punctures not much larger
than pin holes. These were doubtless the
exit holes of a species of Chalcidian.!

Thus the larve of the parasitic Ich-
neumons were themselves preyed upon by
a parasite. However, in each case some
individuals of the original host escaped the
parasitic destroyer. The Chalcidians did
not destroy all the Ichneumons, as the exit
holes attested; and, notwithstanding the
entire lower part of the spider’s egg sac
was occupied by the hymenopterous en-
campment, whose white tents pushed up
against the brown wadding spun by the
mother spider, a large number of young
spiderlings occupied the field. They were
active and apparently healthy, scrambling ye. 330, cocoon of Argiope cophinaria,
among the woolly fibres of their home opened to showthe pupa cases of a brood

i Sor of parasitic Ichneumon flies.
quite down to the cocoons of their invaders.
Whatever ravages the Ichneumons may have made among the spiderlings,
there were certainly enough of them still left. I have found other cocoons
of Cophinaria similarly occupied with some of the perfect Chalcids entan-
gled in the spider silk.

These are, not the only examples of peaceful occupation of a cocoon
by the Orbweaver’s young along with various “squatter sovereigns” of the

parasitic tribes. I received from Mrs. Eigenmann, San Fran-

Treble cisco (August, 1883), a specimen of Epeira accompanied by what
Parasit- : ry.
ian was supposed to be, and probably is, the cocoon. The adult

spider, a female, is black, with faint dorsal foliated marks, and

Science, 1873, page 258.

a Oe

396 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of these were alive within the cocoon, a flossy ball of blackish colored
silk different from any I have ever seen. In the centre was a small agglu-
tinated mass of white parasitic cells, similar to those above described, to
which a number of infertile spider eggs still clung. ‘These cells were
probably those of an Iechneumon, Pezomachus dimidiatus Cresson (Plate V.,
Fig. 5, female, x 4), an example of which was found, dead, clinging to the
padding of the spider’s egg sac. They had evidently burrowed within
the mass of Epeiroid eggs, destroying a number but leaving still many
to hatch out. Within these parasitic cocoons, which were all empty of
their original occupants, were several Dermestid larvee of various sizes.
They were in an intense state of activity, but I did not observe that they
preyed upon the living spiders. Besides these I found in the box two
specimens of a minute Chalcidian, and
several living specimens of a small spe-
cies of ant, apparently an undescribed
Solenopsis. Thus this family consisted
of the original spider hosts, their proper
parasitic guests Pezomachus, the para-

Fic. 331.
FiG. 331. Cocoons of Pezomachus gracilis ina site’s parasite Chaleidian, the universal

Laterigrade spider’s cocoon. Fic. 332. Spi-
der cocoons; one healthy, one infested.

destroyer the Dermestid larvae, and that
inquisitive interloper the ant. It has not
been my fortune to see a more miscellaneous natural combination than this.
Pezomachus does not limit herself to any single group of spiders, but
apparently preys upon all. I succeeded in hatching several, both male and
female, of Pezomachus gracilis Cresson (Plate V., Fig. 6, female,

eis x 4) from cocoons of.a Laterigrade spider. Two cocoons found
uae on the banks of the Schuylkill, attached to the inner side of the

bark of a tree, were joined together as at Fip. 332. One of these
was completely occupied by Ichneumon cocoons. The spider’s cocoons are
made of very stiff silk, and are covered more or less thickly with minute
daubs of mud. Through this covering Pezomachus had penetrated and
lodged her eggs upon the spider eggs within. In due time they were
hatched, devoured the eggs, the shells of which were within. Five pupa
cases of the parasite occupied the interior. (Fig. 331.) In the adjoining
cocoon were healthy young spiderlings and a few eggs.
The cocoon of Epeira apoclisa of England is spun of yellowish silk, of
a loose texture, about half an inch in diameter, and contains about two
hundred and twenty spherical eggs. From this cocoon, on the
18th of July, Blackwall took both sexes of a small Ichneumon
fly, the female of which is apterous, and on another occasion
he obtained specimens of the same insect from the cocoon of Epeira um-
bratica. !

English
Parasites.

1 Spiders Gt. Br. & Ir., page 327.

ENEMIES AND THEIR INFLUENCE. 397

October, 1884, Mr. F. M. Webster sent me from Oxford, Indiana, a
parasitized cocoon, evidently of some Saltigrade species, which appeared to
be that of Phidippus morsitans. The cocoon contained within the outer
flossy case about eighty cells or pupa cases and a num-
ber of mature black hymenopterous insects about one-
eighth inch long. (Fig. 334.) ‘The cells were ovoid,
gray, blackish at the closed
end, probably from excre-
tions from the enclosed lar-
ve, One end was cut open,
showing where the insects
had escaped. (Fig. 333.)

With the exception of a few . b>
F Fic. 333. Magnified cell of parasitic hymenopter, probably Aco-
hard, dry, yellowish brown loides saitidis. Fra. 334. Saltigrade cocoon, with parasitic

examples all the eggs of cells enclosed, somewhat magnified. The fly on the edge is
’

about natural size.
the spiders had disappeared.
The specimens were sent to Mr. L. O. Howard, who thought them to be
Proctotrupids, belonging to the subfamily Scelionine, and seeming to form
an entirely new genus.?
This gentleman has lately published * a description of a hymenopterous
parasite on spiders sent to him by Mr. L. Bruner, of Lincoln, Nebraska,
: which was collected from the eggs of a Saltigrade, Saitis pulex.
igen The eggs of this spider are a little more than a millimetre in
Guests. Circumference, and each egg harbors but one parasite, which
issues by splitting the egg case open, rather than by gnawing a
hole. This insect belongs to the same family and subfamily and is prob-
ably the same species as that col-
lected by Mr. Webster. Mr. How-
ard has named it Acoloides saitidis,
and a copy of the drawing of the
insect is given at Fig. 335. In the
same connection Mr. Howard de-
scribes, under the name of Brus
americanus, a new species, a mi-

Exes 5b: FIG. 836. nute wingless Scelioninz, from spec-
Fic, 385. Acoloides saitidis, a hymenopterous parasite +; ae <e0e
on the Saltigrade, Saitis pulex. Fie. 386. Breus imens sent him ten ¥ ears ago by

americanus, (After Howard.) Natural sizes shown Dr, Marx, who appears to have re-
SR abe ceived them from Col. Nicholas
Pike, of Brooklyn, New York. ‘They are labeled: “Parasites in spider’s
eggs in an orange cocoon.” Dr. Marx, after examining the eggs, expressed
the opinion that the host from which these little parasites were established

FIG. 333. Fic. 334,

' Bureau of Entomology, Department of Agriculture, Washington, D. C.
2 Proceed. Acad. Nat. Sci., Philadelphia, 1884, page 294.
®’“Tnsect Life,’ Vol. II, No. 9, 1890, page 269.

.. pt ieeeie eke eg = *

— ss i ae

398 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

belonged to the Orbweayers. (See Fig. 336.) The natural size of the insect
is given within the circle at the side of the drawing. The insect may not
have been directly parasitic on the spider’s eggs, but on the larve of
Pezomachus or some other spider parasite.

The above examples are sufficiently representative, and they indicate the

mode in which the Hymenoptera wage war upon the Aranez. As to the
extent of that war I may simply say that I find a large propor-
Extent tion of Epeiroid cocoons infested by parasites. In going among
of Par- ; : ame ,
asitism, ‘2 old cocoons in the spring it is often more usual to find them
occupied by cocoons of Pezomachus than full of healthy eggs.
Of five large egg nests of Epeira cinerea now before me, not one is free
from parasitic cocoons. Often the spider brood will be in part preserved;
frequently the parasites have full possession; and, again, the traces of
a full and healthy brood are shown by the shells or first moults within
the central bag. Of course the ratio
of destruction varies at different times
and places.

There appears to be no special
proclivity on the part of parasitic hy-
menopters to confine their operations
to any species of spider. They appear
to choose their host indiscriminately
a from among the cocoons in which the
"inet token the nfceion iaeneeae finer eggs are swathed. Pezomachus gra-

cilis will attack the hard, stiff, and
compact cocoon case of Argiope cophinaria, or will choose as a host the
eggs of a species of Epeira, or indeed of other tribes.

This point needs further investigation, and would be a matter of some
importance to solve with absolute certainty. It would greatly add to our

__. respect for the discriminating powers of these strange insects if
cone we were to establish the fact that they can select a spider's eggs
creed even when they are hidden away under protecting cases so widely

differing in appearance, construction, and location. The question
would then rise, in what manner do the ichneumons determine the pres-
ence of the eggs? Do they watch the spiders themselves? Are they able
to detect the presence of spiders’ eggs through the enclosing enswathment
by some sense so delicate that it cannot be appreciated by human beings ?
The wingless condition of the females doubtless greatly favors them in their
search for objects hidden away as spider’s cocoons commonly are.

VIII.

These are not all the hostile agents arrayed against the embryo life of
spiders. Parasitic plants as well as parasitic insects assail them, for not

ENEMIES AND THEIR INFLUENCE. 399

unfrequently the eggs are destroyed by vegetable mold. According to the
observations of Homburg, house spiders in the kingdom of Naples are sub-
ject to a malady which makes them appear hideous. Their body
Mold, becomes covered over with scales, bristling one above the other,
Flies, : , ; ; . ;
Birds. among which numbers of a species of mites are discovered. When
the spider walks, it shakes itself and throws off part of the scales
and some of the parasites.! :
One day I was dissecting a cocoon of Epeira sclopetaria, and had just
turned back the white sheeting of the interior sac, thus quite exposing
the eggs, when a house fly lit upon the mass, and instantly thrust her
proboscis into and sucked out the contents of an egg. I permitted the
insect to continue its feast long enough to show that the innumerable com-
pany of common flies only require an opportunity to wholly cut off and
exterminate their hereditary foes at the very fountain head of life.
Spiders themselves enjoy a meal of spiders’ eggs; for example, Staveley
speaks of two species of Clubiona feeding upon the eggs of other species. ?
Birds have already been alluded to, in the chapter on Aeronautic Habits,
as utilizing spider cocoonery in the construction of their nests. Among
those addicted to this habit are the pewit,® the
wren,* and the vireo. I have several specimens
of nests made by a species of the last named bird,
probably Vireo noveborocensis, collected in Fairmount Park,
in all of which cobwebs have been used more or less freely.
(Fig. 339.) I am told that this is habitual with that bird.
The texture of the spinningwork shows, evidently, that it
had been plucked from cocoons; and if this were done
before abandonment by the brood, at least before hatch-
ing, the destruction of the contents must have followed. jc. 338, 4 cocoon
It illustrates the catholicity of habit among the animal — nest of Epeira,
. sy re Sea : rifled of its eggs.
races, that Dr. David Livingstone, the distinguished mis-
sionary explorer, found a like habit in Africa among the sunbirds.°
Mr. Carl Voelker has a specimen of the nest of a hummingbird, which
is composed in considerable part of various portions of spinningwork taken
from the snares and cocoons of spiders. He has seen our common red
throat hummingbird, Trojilus rubicolis, darting at the webs of spiders
and gathering the threads in its bill for nesting purposes. He has also
found minute spiders in the throats of birds of this species, and believes,
therefore, that it is their habit to feed upon spiders. The Blue Gray Gnat-
catchers also use spider webs for the construction of their nests, and thus

Bird
Enemies.

2 British Spiders, page 101.
8’ Mr. Thomas Meehan, the botanist, is my authority for this statement.
* Mrs. Treat, “My Garden Pets.” ® Livingstone’s Last Journals, page 453.

400 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

The above facts, uncovering as they do so hard a destiny impending
over every stage of aranead life, might well awake sympathy in the breast
of the most pronounced spider hater. To those who know the usefulness
to man of the much enduring race, and yiew its destruction from the
standpoint of human disadvantage, the facts are melancholy enough. But
after all there seems a judicial fitness in the order of things which ap-
points avengers from the midst of the insect world against the chief de-
stroyer of the insect hosts. Seeing, therefore, that some check is required’
upon the excessive increase of spiders, we may regard their relation to
the Hymenoptera with some complacency from the view point of natural

justice,

Fic, 339. Nest of Vireo noveborocensis woven together, with bands and threads
of plundered spider webs.

IX.

In speaking of the enemies of the spider we have thus far omitted
one of the most determined and destructive—man himself. But it will be
observed that I have been writing of the natural enemies of
Foolishly spiders, and in my opinion man cannot reasonably be classed
Hostile : rie ; Ra j
stan among these. His hostility to the various families of the spider
world is without reason not only, but is against reason. It is an
example of indulgence in a prejudice which has been long fostered by

‘eS = =—§s a

—— vor

ENEMIES AND THEIR INFLUENCE. 401

ignorance, and which, I am thankful to record, is yielding before the
light of modern science. In truth, the spider is not only a harmless creat-
ure as far as man is concerned, but is, on the contrary, a most helpful
one to him in many respects. She is one of those checks established in
the economy of Nature against the increase of insects whose presence
would make the world well nigh uninhabitable by the human species.
Some idea of the destruction wrought in the insect world by the cun-
ningly devised snares of Orbweavers may be had from the following facts:
I have counted nearly two hundred and fifty insects, small and great, hang-
ing entangled in one web. In another net, in Fairmount Park, I counted
thirty-eight mosquitoes ; in another, hung under a bridge at Asbury Park,
and out of reach, there must have been tivo or three times as many.
Greenhead flies by the legion have been seen in the snares that fairly en-
lace the boat houses at Atlantic City and Cape May. Very small spiders
prey upon microscopic insects, like gnats, and devour myriads, A glance
at the fields, bushes, and trees on a dewy morning in September will reveal
an innumerable multitude of webs spread over the landscape, all occupied
by spiders of various ages, sizes, and families, and all busy destroying the
insect pests of man. These webmaking spiders thus revealed are only a part
of the numberless hosts engaged in this friendly service. On the ground,
in crevices and crannies into which man never looks; lurking on flowers,
on leaves, on limbs and twigs of trees, shrubs, grasses; in barns, cellars,
outhouses; everywhere, indeed, upon the face of Nature, one who will take
the pains to look will find legions of spiders carrying forward day and
night without cessation the same vigilant and unrelenting warfare upon
the insect world. They are of all sizes and of various forms; in all stages
of their life, from the spiderling upon its tyro web to the grizzly veteran
just ready to give up its life as the frosts of autumn fall. If one stops
to consider that all these creatures must find food, and do find food, and
that the chief supply is furnished from the realms of insect life, he may
form some faint conception of the destruction which is wrought, and, by
consequence, the service done to man.
To the testimony thus gleaned from field observations we may add the
evidence of postmortem examination made by a careful and learned stu-
dent with the aid of the microscope. Dr. ©. Keller, of Zurich,
Arachne glaims that spiders perform an important part in the preserva-
Fieiak tion of forests, by defending the trees against the depredations
Keeper. °f aphides and insects. He has examined a great many spiders,
both in their viscera and by feeding them in captivity, and has
found them to be voracious destroyers of these pests; and he believes that
the spiders in a particular forest do more effective work of this kind than
all the insect eating birds that inhabit it. He has verified his views by
observations on coniferous trees, a few broad leaved trees, and apple trees,
An important feature of the spiders’ operations is that they prefer shaded

ee a es

402 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

spots, and therefore work most in the places which vermin most infest,
but which are likely to be passed by other destroying agents.

We thus see that man is not only indebted to Arachne for protection to
his own personal safety and comfort, but also for the protection of his for-
ests, fruit orchards, gardens, and fields. Indeed, the whole vegetable world
may well join with man in a tribute of gratitude to a creature whose
service is so eminently useful.

Surely, in view of these facts, we need not hesitate, through fear of
being charged with undue enthusiasm, to declare the spider a universal

philanthropist. She labors unceasingly to check the increase of a
Arachne horde of tiny insect enemies which else would banish the hu-
ohare man species from many parts of the earth. Nor does she make

ropist. ee : ¥ e

reprisals of any sort for all this service. She never attacks fields,
harvests, vineyards, and orchards, like beetles, grasshoppers, and various
other insects in the perfect and larval state; she never forages upon the
goodies in ladies’ kitchens and pantries, as do roaches and ants; she does not
torment and afflict by cutting, piercing, sawing, and pumping, by buzzing,
humming, and blowing, like the mosquito and house fly, to say nothing of
less desirable denizens of the entomological kingdom. An occasional (and
doubtful) “spider bite” one does hear of at rare intervals; a harmless cob-
web here and there in a cranny or corner of one’s house, that is all that
can be charged against her. Yet this useful aranead is despised, abhorred,
persecuted, and slain with a zest that is hardly shown against any other
creature, except the snake. No; man is not a “natural” enemy of the
spider, but an enemy by a culture most unnatural and unreasonable. What
stupid ingrates men are often found! ‘What fools these mortals be!”

This subject could not be held complete without reference to the rela-
tion which undoubtedly exists between the facts which the chapter un-
covers and many of those interesting habits described in this
Influence. yolume and elsewhere. ‘That the instincts and industry of spiders

cme are in a large degree protective will not admit of dispute. It is
Industry. Certainly a reasonable theory that they have in part, at least, arisen

or been modified by that fact. That is to say, the original en-
dowments of the creature have been enlarged and varied by the peculiar
perils with which successive generations have had to contend.

We have already anticipated the influence of enemies in developing the
industrial habits of spiders, in the chapter on Nesting Habits and Protective
Architecture. (Vol. I., Chapter XVII., pages 807-309.) By referring to the
summary of the various forms of tents there described, the importance of
this influence, particularly in the case of Orbweavers, will be noted, It
appears that Orbweavers live continually in dread of enemies, and that

ENEMIES AND THEIR INFLUENCE. 403

— st ——— i ——— = ——— : —S

their whole life is spent in a defensive industrial warfare for the protec-
tion of their persons. In order to illustrate this truth a little more fully
I propose to consider the habits of those spiders which make burrows in
the ground, namely, the Citigrades and Tunnelweavers. Both of these
tribes, but particularly the latter, are noted for the admirable dwelling
places which they construct. I hope to show that these ingenious homes
are largely defensive.

Beginning with the Citigrades, we find, in the first place, that during
the period of moulting, when the spider is conscious of its de-
fenseless condition by reason of physical weakness, it is in the
habit of protecting itself by covering the mouth of its den with
a silken sheet, or by spinning a special cell in which to shed its skin.

Again, I have observed Lycosa, when about to make her cocoon, con-
struct a cell in the earth and carefully cover the entrance thereto with a
silken curtain, She was animated, as I conjectured, by the wish to pre-
serve herself from enemies during this crisis period of her life. I have
also shown that it is the universal habit of these Lycosids to construct co-

cooning caves or nests underneath stones, logs, and like situa-
Ste: tions, which are carefully plastered and enclosed on all sides,
Caves, leaving an entrance which is usually well protected from assaults

of ordinary enemies. This appears to be a cosmopolitan habit;
at least, Mr. Campbell tells us that some English Lycosids dig an irregular
oval cavity about one inch by a half inch in diameter, close it with a con-
glomerate of silken threads and earth, and remain therein with the
cocoon. He kept one Lycosid in confinement and twice destroyed her
retreat, only to find another made the following morning. The top was
covered with granular pieces of soil, such as might have been raked over
the silken lining with her feet. In both the above conditions, namely,
while moulting and while cocooning, these Citigrades appear to be driven
to special industrial provisions by the impulse of self protection.

That the same habit prevails as a protection against the destructive
influences of climate is well attested. Lycosids everywhere appear to seal

j up the openings to their cylindrical burrows at the advent of
a winter. The advantage of such artificial closure, as a protec-
Covers, tion against winter cold, was well demonstrated by Mrs. Treat.

A large example of Lycosa carolinensis, which makes a beau-
tiful nest (Fig. 291, Vol. I., page 316) was brought from New Hampshire
and domiciled in Mrs. Treat’s ground. Its burrow was only eight inches
deep, yet there was a marked increase of temperature above that at the
surface. This could be discerned by placing the hand at the mouth of
the tube. One cold morning when the thermometer stood twenty degrees
above zero (Fahrenheit) a thermometer was introduced, It ran up to
forty degrees, making a difference of twenty degrees in temperature be-
tween that prevailing at the surface and that within the burrow. Soon

1. Moult-
ing Tents.

OO a ae i mill
4 La Sees “3 =

404 AMERICAN SPIDERS AND THETR SPINNINGWORK.

after this test was made, late in November, the spider closed its burrow
with a canopy of thick web, over which were drawn a few sticks and
straws. It cannot be doubted that’ the advantage thus secured by the bur-
rowing habit, against the exigencies of a severe winter, strongly tend to
protect the life of Lycosids.

I have also called attention to the fact, while treating upon the cocooning
industry and maternal instincts of spiders, that the industrial product of
such skillful architects as certain Lycosids and the Trapdoor spiders, is
probably influenced by the mother’s wish to protect her eggs. I refer the
reader to what is said Chapter III., page 64, merely remarking that it
would indeed be strange were not the strongest feelings in animal nature
to leave their impress in some form upon the industrial life of animals.
I have also suggested that even the sexual excitement of the male reacts
upon his industrial energies and tends to the development of a higher skill,
at least in certain species. (See page 65.)

I now proceed to show that the necessity of self protection against their
most persistent and formidable enemies, the wasps, has led certain Lycosids

to adopt a special and interesting form of ptotective architecture.
4. Self We fortunately have abundant facts in the case of Lycosa tigri-
eee en na,! as carefully observed and recorded by Mrs. Mary Treat.?
dustry. /igrina’s method of working, as observed from a large female
specimen in confinement, is as follows: She first spins a canopy
of web over her tunnel, leaving a place of exit on one side. She next goes
out and carefully moves over the canopy as if to see whether it is strong
and secure. Satisfied that it is all right, she steps down, just letting her
fore feet touch the web, while with her hind legs she feels and apparently
examines the material. Finally, she selects a dry oak leaf about two inches
broad and three in length, lays it over the canopy, and proceeds to fasten
it down all around except at the entrance. After the leaf is made secure
she reaches up and pulls down blades of grass, lays them over the leaf and
dexterously fastens them down with webs. This makes a strong roof for
her domicile. Then she goes within and puts the finishing touches on the
inside. This done, she stands in the door of her neat apartment awaiting
At insects that may chance to come within her range. If a beetle,
ae for example, approaches, she rushes upon it and bears it into
Hone her den, A few days after the work is thus begun, Tigrina com-
pletely closes the entrance to her domicile, and the observer avers
that if she had not known the spot in which it was located, she would
not have been able to find it.

The burrowing habit of this species is there for the first time fully described by me, and a
brief deseription given of the female. Emerton (New England Spiders of the Family Lyco-
sidee, 1885) describes the species as Lycosa vulpina, I judge that my own name has pre-
cedence. 2 Home Studies in Nature, Harper’s Magazine, 1880, page 710,

idl pele ied he.

SEE = - . e.CLClh wine —— =

ENEMIES AND THEIR INFLUENCE, 405

At one time twenty-eight of these spiders were kept under observation
in the obseryer’s grounds. She visited them all frequently and found that
more than half the number, both males and females, had closed their doors
firmly. Some of the burrows were situated in bits of moss, and the moss
was so cunningly arranged over them that the most expert naturalist would
have found it difficult to tell where they were located. Mrs. Treat had
often tested the matter with her friends, to see if they could find one
of these concealed burrows, limiting their search to afew square inches
of space. But they rarely Fit upon the right spot. Mrs. Treat kindly con-#
tributed some of the coverings of these Tigrina burrows to my collection

Owe: sity
fl co “ Wy

uA as
fi , Mi

Fic. 340. A vestibule and dome of vegetable débris reared over the burrow of Lycosa tigrina.
Natural size, (From a cabinet specimen.)

of aranead architecture. They are masses of vegetable débris, moss, grass
blades, chippage, dry twigs, the shell of an acorn, etc., arranged in an
irregular dome several inches in diameter across the base and an inch or
more in height. (See Fig. 340 and Figs. 341, 342.)

What is the explanation of this care shown by Tigrina in the covering
of its den in midsummer? The closure in winter is explained
by the natural desire to protect the burrow from frost and
snow. ‘The closure during or previous to the moulting season is
explained by the fact that the spider requires especial protection at a period

The Eix-
planation.

406 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

of special weakness and inability for defense. The closure of the domicile
just before cocooning is explained by the maternal instinct which drives
mothers into hiding at that crisis period of life. But August is the time
when insects are most plentiful in our climate, and one would think that then
Tigrina would keep her burrow, open in order more freely to prey upon
insects.

The explanation is certainly found in the presence of a species of Digger
wasp, Elis 4-notata Fabr. (Scolia). (Plate V., Fig. 3.) She belongs to the
arachnophagous species, and, although the mother appears to feed
upon nectar and honey, the grub feeds upon the juices of spi-
ders, and the particular species affected by the wasp larva is
Lycosa tigrina. The wasp is large and strong, has smoky brown wings
with a strong purplish blue iridescence, and two bright orange spots on
either side of the abdomen. She runs over the ground swiftly, peering
here and there into various recesses, until she alights upon an open bur-
row of Tigrina. Down into this she plunges, and soon returns dragging
up the inhabitant, which she has already paralyzed with her powerful
sting. Woe now to all spiders with unclosed doors, for Elis is sure to
find them !

Sometimes two wasps are hunting in the same vicinity, and when one
finds a spider the other tries to wrest it from her. A furious battle en-
sues. The combatants drop the prey and clingh in conflict, seemingly try-
ing to stab each other with their stings. The victorious party returns to
the spider, which is often heavier than herself, and proceeds to drag it to
her nest. She moves backward for a time, drawing it over the ground,
then tries flying a short distance, but the burden is so heavy that she soon
comes to the ground again. She is so active and quick in her movements
that one has to walk quite fast to keep abreast with her. She carries the
spider several rods from where she obtained it, lays it down on the gravel
walk, and hunts over the ground. Presently she finds a burrow which she
had previously dug, takes up her spider, and disappears within.
She comes out empty handed and proceeds to fill up the hole
with the earth she had thrown up. She works so rapidly that
one can scarcely tell which feet she uses most. She seems to dig with
her fore feet and rake the earth in backward with her hind feet. Soon
the hole is full, And now she makes a battering ram of herself by rapidly
striking her body on the ground, as if to pound the earth down. This
done, she rakes the ground all over and around the place to make it level,
then collects small pebbles in her mandibles and lays them over the spot,
until it looks little different from the surrounding ground. Elis also
knows how to practice local mimicry!

When one of these mother wasps had retired, the observer dug up
the paralyzed spider, which was about four inches below the surface, and
found an egg sticking in the body. This egg hatches into a white grub

Elis the
Invader.

The
Burial.

a
a ae ° ; st = =n ;

ENEMIES AND THEIR INFLUENCE. 407

in about six weeks, when it at once begins to feed upon the stored spider.
When full grown it passes into a chrysalis state, in which it remains until
the following summer, when it emerges, a mature insect, and, like its
mother, begins a remorseless raid upon the Tiger spiders. The wasps con-
tinue their raids for two or three weeks, only the spiders with closed
doors escaping. Sometimes a spider will keep herself shut up for two
weeks and then timidly open her door and look out.

After the end of August the maternal rage had exhausted itself, the
wasps disappeared, and then Mrs. Treat found that out of twenty-eight
spiders only five were left! ‘These survivors soon opened their doors, and
occasionally one would cut the threads of the spinningwork which unites
the thatching material in such a manner as to make a sort of trapdoor,
leaving hinge on'one side. (See Figs. 341 and 342.) But more commonly
a hole was left in one end of the oven shaped cover, which the spider
would readily close by drawing the material together and fastening it with
threads.

It seems impossible, in view of such a careful and intelligent record
as this, to doubt the fact that the instinct by which Lycosa tigrina is im-

pelled to construct the vestibule roof and door to her den, has

a eel been vitalized by, or at least associated with, the natural in-
=i dustry, stinct of self protection against the raids of its formidable enemy,

the Digger wasp. This is further emphasized by the fact that
the partly grown Tigrinas, who are not molested by the wasps, have never
been observed to conceal their burrows in the manner of the adult. Tigri-
na’s rude architecture is a product of her peril.

The defensive motive in the architecture of Lycosa tigrina, which is
thus clearly demonstrated, serves as a key to the purpose underlying all
the architecture, not only of the Lycoside, but of other burrowing spe-
cies. The watch tower which guards the nest of the Turret spider (Vol-
ume I., Fig. 289, page 314); the interesting structure so closely resembling
a bird’s nest, reared above her nest by Lycosa carolinensis (Volume L.,
Fig. 291); and other forms of industry of which these are types, may all
be considered as in part, at least, the result of protective industry.

According to the account of Dufour, the habits of Lycosa tigrina and
Lycosa arenicola are quite similar to those of the famous Lycosa tarentula
of Italy, the heroine of the “'Tarentula dance.” This aranead
forms a cylindrical burrow in the earth, often more than a foot
long, and about one inch in diameter. At about four or five
inches below the surface the perpendicular tube is bent horizontally, and
it is at this angle that Tarentula watches for the approach of enemies or
prey. The external office of the burrow is ordinarily surmounted by a
separately constructed tube. This tube and outer piece of architecture
rises about an inch above the surface of the ground, and is sometimes as
much as two inches in diameter, being thus even larger than the burrow

Lycosa
tarentula.

——E—————————

408 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

FiG. 341 (upper figure). Vestibule of Tigrina’s burrow, with door open, FiG. 342 (lower figure),
The same, with door closed. Natural size. (Drawn from cabinet specimens.)

—-- =

PNEMIES AND THEIR INFLUENCE. 409

itself. The tube is composed of fragments of white silk fastened together
with particles of clay, etc., and so artistically disposed, one above the
other, that they form a scaffold, having the form of an upright column,
of which the interior is a hollow cylinder. The tube is lined with silk
throughout its whole length.?

aol

I have no hesitation in also applying the key thus furnished by the
habits of Tigrina to interpret the motive of Trapdoor spiders in
their remarkable industry. With this in mind, and aided by
other facts and conclusions drawn from a direct study of this
form of nest, we may venture to approach the subject.

M. Eugene Simon has contributed largely to our knowledge of Tunnel-
weaving spiders in a recent paper presented in the Annals of the Entomo-
logical Society of France? and in the Acts of the Linnean Society of
Bordeaux.’ His descriptions are accompanied by notes upon the habits
and architecture of the spiders, with admirable illustrations, which greatly
enlarge our knowledge of the nesting habits of these interesting creatures,
who rank among the most skillful artificers of the animal world. Mr.
Simon’s notes were made from specimens obtained in Venezuela and North
Africa, the Venezuelan specimens being studied personally during a visit
to that country. A comparative study of his papers enables us to trace
the progressive development of the nesting architecture of Tunnelweavers
from the simplest tube in the ground to the hinged doors or trapdoors
which close the silk lined burrows, and which are so well known for
their mechanical perfection.

The simplest form of burrow is that of the Tarantulas, which represent
the largest known spiders. These huge araneads appear to depend wholly

upon their size to resist the assaults of enemies who invade their
1. Taran- den, At least I have not found satisfactory evidence that they
ae erect any artificial barrier over the entrance to their tunnels.
Burrow: A more complicated burrow and one better serving for defense

is that of Leptopelma cavicula of northern Africa. The drawing
(Fig. 343) shows a section view of the upper part of the burrow, the en-
trance to which is without any door or other defense as in the case of the
tarantulas. The burrow descends perpendicularly for a little ways, but at
the top a special branch diverges laterally, which curves and again descends
perpendicularly for a considerable distance. At the summit of this second
and parallel perpendicular tube another branch issues, inclining upwards
towards the surface. A glance at this structure, if we suppose it to be

Trapdoor
Spiders.

' Lucas, Hist. Nat. des Animaux Crust. et Arach., page 357.

2 Extrait des Annales de la Société Entomologique de France, “Arachnides du Vene-
zuela,” December, 1887, April, 1888, pages 170-220, plates i., ii., iii.

® Actes de la Société Linnéenne de Bordeaux, Vol. XLIL., 1888.

a ee ees a RO Oe a ee eee

410 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

| Fie, 844,

4
Sign i

Fic, 346.

| FiG. 343. Burrow of Leptopelma cavicula; section view of upper part. Fic. 344, Lily shaped

tube of Leptopelma elongata. Fic. 345. Turret of Dolichoscaptus inops. (Natural size.)
Fic. 346. Turret, with trapdoor, of Dolichoscaptus latastei, supported on a plant. Four
inches high. (After Simon.)

ENEMIES AND THEIR INFLUENCE. 411

characteristic of the species and not an accidental formation, will show that
it makes an admirable protection against heavy rains, which sink away into
the first burrow as a kind of reservoir, enabling the spider to escape
2. Lepto- by the diverging branch. Against enemies who pursue it into its
eaeates den this structure also presents an effectual defense, for, while an
Burrow, ¢Cnemy naturally would rush downward into the first direct pass-
age, the spider may escape by the lateral branch. Supposing that
the enemy, observing the mistake, ascends and follows along the branches,
the spider has the opportunity to push up into the second branch while the
pursuer, again following its natural instinct, would rush down the second
perpendicular tube. I am here in the region of conjecture, but perhaps no
better explanation presents itself.
A third stage in the development of this defensive industry is repre-
sented at Fig. 344, which shows the external tube of Leptopelma elongata. !
This is simply a lily shaped tube of pure white spinningwork,
8. Silken rising directly above the burrow, and supported by surrounding
ete foliage. The purpose of this structure has not been positivel
Tower, oliage. e purpose not been positively
determined. As able a naturalist as A. R. Wallace has conjectured
that it may be deceptive in its uses, its resemblance to a flower attracting
to it insects, which are thus preyed upon by the proprietor. Such elevated
objects are certainly apt to attract insects, who are disposed to alight upon
them even without regard to their promise of providing food. But I am
inclined to believe that Leptopelma’s silken lily serves as a watch tower
from which she can observe the approach of enemies and make good her
escape in time. Moreover, I believe that it is possible for her to pull
together the sides of the sheeted turret and thus erect a barrier between
herself and some of her feebler pursuers.
Another form of defensive industry is presented at Fig. 345, which is
the exterior part of the turret tube of Dolichoscaptus inops Simon, This
is about an inch in height, and is composed of mingled chippage

. Con- : and mud, a sort of: débris of chopped straw and soil.
Pawan * © A still further stage is shown at Fig. 346, which represents a

Doorless, COlumnar turret of Dolichoscaptus latastei several inches high.
This resembles the tower of the preceding species, but adds thereto
a hinged covering after the manner of the trapdoor. This turret is also
composed of chippage and débris of various sorts gathered from the neigh-
borhood, and is supported upon the surrounding foliage, which in

: Trap- the drawing is a plant of Lavandula dentata. All the uses to
Tones which such an elevated structure can be put are served by this

ingenious structure, and, in addition, the trapdoor is manifestly
intended to defend the inmate from the assault of enemies.
We come now to the trapdoor nests of Nemesia meredionalis, and other

1 Cyrtauchenius elongatus; see Volume I., page 322, Fig. 304. 7

412 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

species making traps of the wafer type, as so fully described by Moggridge.
Here we have simply a dropping away of the turret of Dolichoscaptus
and the use of the burrow independently of the same, but with the trapdoor
retained. In the species studied by Moggridge a single burrow is the or-
dinary rule, but there are many variations, some of which are manifestly
characteristic of species, and others which are probably occasional and ac-
cidental.
A variation described by Mr. Simon is shown at Fig. 349, the nest of
Stothis astuta, which inhabits the forest of Cartuche, near Caracas, South
America. The drawing shows a section of the burrow, indicating
6. Bur- the curved course, and also the two wafer like trapdoors habit-
eh ually placed at either end. That this peculiar industry is defensive
Dace is probable, for we can readily imagine the spider disappearing
within its den at one door, and, if its pursuer should succeed in
entering the same, escaping at the other. We might, without much stress
of imagination, carry the conception a little further, and suppose again the
enemy making its exit from one door and the spider again descending

cl
iS ai 4

if ie

LNs das

Bik Ky
f SN f

Wt fae ate
aN

Fic, 347. Fic. 348.

Fic. 347. Silk lined case of Stothis astuta, with two doors.
Fic, 348. Front view of a door.

into its burrow by the other. This game of bo-peep might evidently be
played to the great advantage of the Trapdoor spider and manifest dis-
concerting of its enemy.

Simon gives an interesting example of the ability of a spider of this
species to change its habit and adapt its industry to unexpected sur-
roundings. The species commonly seeks dark and damp localities, and
digs in vegetable earth a burrow not very deep. The nest, which is
drawn in side view at Fig. 347, and a front view of the door shown at
Fig. 348, was begun underneath a stone in soil which was so rocky as to
be impenetrable. Not wishing to change its site, and not to be cheated
out of its proposed domicile, Stothis proceeded to erect a cylindrical case
about two inches long, composed of a conglomerate gathered from surround-
ing particles of soil and vegetable chippage. ‘These were cunningly wrought
together, the whole structure silk lined, and the ¢haracteristic trapdoors
hung, one at either end. Thus, while varying her habit in so far as to
build a surface tunnel instead of a subterranean one, Stothis preserved her
defensive habit of erecting for herself a back door by which she could
retreat in case of invasion at the front door.

. Fig. 350 represents the burrow of Stothis cenobita Simon, which is

See eer .”.h Che

ENEMIES AND THEIR INFLUENCE. 413

Fie, 350.

Fig. 351.

Fra. 349. Section view of curved burrow of Stothis astuta, showing double trapdoor entrance.
Fig. 350. The globular burrow, with trapdoor of Stothis cenobita. (Section view.) FuG. 351.
Front view of trapdoor. (Natural size.)

ee ee a

414 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

simply a rounded chamber underneath the surface and closed by a trap-
door, Fig. 352, which differs in no particular, as far as I can observe, from
the ordinary trapdoor of the American Cteniza californica, (See
Bate Fig. 240 B, page 183.) ‘
“Cork” It is difficult to say what may be the enemies of the Trapdoor
nor spider against which such ingenious architecture has been reared
and such vigilant watch is exercised. But the quite general testi-
mony is that these spiders leave their tubes at night and go forth in search
of prey; or, as in other cases, open the lids of their tunnels and spread strag-
gling lines near by, upon which passing insects are entangled and delayed
long enough to allow the spiders to pounce upon them from their open
caves. If we credit these accounts we might infer that the enemies which
the Trapdoor spiders most dread are not such as are abroad at night.
Evidently the creatures are fearless at that time, a state of mind which
doubtless results from their knowledge that they are comparatively free
from their worst enemies. The
enemies which they most dread
may therefore be reasonably look-
ed for among diurnal creatures,
and not among those of nocturnal
habits.

Among these foes at least one
of the most formidable and irre-
sistible is a diurnal insect, the fe-
male of the terrible Digger wasp,
which I do not doubt will be
found to store Trapdoor spiders,
as well as Tarantulas and Lyco-
sids. There is no evidence known
to me that Pepsis formosa in-
vades the tunnel of the Myga-
lidee in order to dig them out.
sei Such an act is not indeed be-

Fs 22, Traptoor af Cte eerie ator sie, yond her powers, and, reasoning
s a on the silk lining. . ‘
from the conduct of Elis 4-notata,
it is highly probable. But we aré not yet warranted in attributing the
habit to her. Some lizard or mammal that might pull open the trap
with its claws may be looked for as also a probable enemy against which
Trapdoor spiders erect and defend their ingenious barrier,

At all events, the spider herself is well aware of these enemies. Abbe
Sauvages invariably found, when he attempted to open the door of the
nest of “the Mason spider” (Nemesia and Cteniza), that the mother was
on guard, holding down the lid of her tunnel with great force. In his
efforts to pull the trapdoor up the spider would jerk it down again, and

me ’
a ee

——s eee,

ENEMIES AND THEIR INFLUENCE. 415

there would be an alternate opening and shutting of the nest until his
purpose was accomplished.!

It is the habit, according to Moggridge, Simon, and all observers who
have noted the point at all, for these animals to hang back downward
upon the inner surface of the door. In many nests which I have seen
there are holes along the outer or free edge of the door—the part directly
opposite the hinge—which mark the points at which, probably, the fangs
of the spider had been fixed, in order to give it a strong purchase against
intruders. ‘These holes are usually three, and show with great distinct-
ness, even in the photograph from which Fig. 352 was drawn, which is
the upper part of the nest of our California Trapdoor spider, Cteniza cali-
fornica.

\ XII.

One of the most remarkable developments of industrial skill under
hostile influences is that described by Moggridge in the inner door of
Nemesia congener.” The general character
of the nest is that of a tube ten or twelve
inches long, drilled horizontally into the side
of a slope, and closed outside with a wafer
door. This horizontal burrow bends abrupt-
ly, and is continued perpendicularly down-
ward, with a short branch extending upwards
towards the surface. At the juncture or
bending is suspended a wedge shaped double
door, which tapers from below upwards to a
hinge. The door has two crowns, separated
from each other by the gusset like web of
silk that connects the door on either side with
the lining of the main tube. One of these
crowns fits into and closes the main tube,
while the other fits into the aperture of the
upward branch. This swinging door acts very
much in the fashion of a valve, dropping
down to close the entrance from the first to
the second tube, and swinging upward in such
a Way as to protect the spider when it is Fig. 358. Cyclocosmia truncata. Fic.
cowering within the upper branch of the per- — ** Site view of same (After Hentz.)
pendicular tube. The wedge like structure of this door is seen not only
in the adult spider’s nest, but in even more exaggerated shape in the nests
of the younger ones.

‘Cuvier, Animal Kingdom, Lond. ed., Vol. XIII., Supplement, page 465.
* Trapdoor Spiders, Supplement, page 228 and pl. xvi.

416 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Mr. George I’. Atkinson, who has given some attention to the Ameri-
can Territelarie, dissents from Mr. Moggridge’s supposition that this and
other inside doors affords the Trapdoor spider a means of escape when
pursued by an enemy, the view which I have favored in the foregoing
pages. Mr. Atkinson, on the contrary, found indications that the main
tube of Myrmeciaphila foliata! is constructed to serve as a gallery for the
passage of ants or other insects, and that the branch so constructed is a
real trap in which the spider awaits the passing of an ant, when it opens
the door and catches the insect. In support of this opinion, he states that
he found that the trapdoor nests were all made in places where ants had
underground passages; that the main tube connected directly with some of
the ant galleries; that the trapdoor at the surface of the ground had the
appearance of being little used; and, finally, that one nest had only one
door, which led into a short tube that opened into the floor of a broad
hall in an ant’s nest leading into several galleries. This hall was the
gangway to the surface of the ground made by the ants, and through this
the spider probably entered the hall to construct her branch tube into
the floor.

Again, in May, at Chapel Hill, North Carolina, Mr. Atkinson found a
nest of the same species under conditions which seemed to give conclusive
evidence that the main tube is intended to entrap un-
wary insects as they pass the door of the branch where
the spider lurks. This nest was made in a broad foot
path where the clay soil was very hard. It was discovered
by seeing the open door. The following day Atkinson
visited the place with trowel in hand, to take up the spi-
der. He found the door still open. The main tube was
about nine inches long; the branch about one inch long,

this the spider was found. The door to the branch was
a “cork” door, that is, a thick beveled one, while that at
Foe Mtksanttesecr ts the surface of the ground was a “wafer” or thin door.
view of Truncata,clos- It appears in cases where the spider’s tube is not made
ne pit with in the ant’s nest, that the outer door is set open, thus
offering an attractive place for insects crawling on the sur-

face of the ground in search of food. They enter the main tube and, as
they pass the branch, the door is suddenly thrown open, and to their sur-
prise they are taken captive and made a meal of by the cunning spider.?
There is no doubt that Mr. Atkinson is correct in so far as the burrow

of the Trapdoor spider does serve as a true trap for the capture of ants.
Mr. Moggridge shows this in the case of Nemesia coomentaria. (See above,

1 Entomologia Americana, October and November, 1886.
* Psyche, Cambridge Entomological Club, Vol. V., July-August, 1888, page 89.

situated six inches from the surface of the ground. In -

Le hCUrl
Me ee a ee oe a
5 > 7 - T ,

ENEMIES AND THEIR INFLUENCE. 417

Chapter XII., page 355.) Mr. Simon also attributes this use of the tube
to the ants of northern Africa, particularly to Dolichoscaptus vittatus,
which drives into the earth a horizontal burrow that is considerably con-
tracted and bent towards the end in the form of a little cul de sac, which
is full of the débris of insects upon which the spider has fed. In one
locality this insect débris consisted almost wholly of the remains of Atta
barbara, the well known harvesting ant of Palestine and the Mediterra-
nean shores.’ I do not see that the two facts are at all contradictory.
The use of the silk lined burrow as a decoy for curious insects in no-
wise hinders it from being also useful as a defense against enemies of
various sorts. x

One of the most curious examples of relation of structure to enemies,
or perhaps of the reaction of hostile environment and agents upon struct-
ure is found in a Territelarian spider, Cyclocosmia trun-
cata.2 This aranead, according to Hentz, dwells like
others of its kind in cylindrical cavities in the earth.
Though many specimens were found, he never saw any
lid or closure to the aperture of its dwelling. The very
singular formation of its abdomen, which is as hard
as leather behind and is truncated to form a perfect
circle, induced Hentz to believe that when in danger
it closes its dwelling with that part of its body instead
of with a trapdoor or lid. This conjecture, of course,
needs confirmation, though it seems not im-
probable; and one may imagine the intellectual
confusion of a pursuing enemy, which finds its
prey suddenly disappearing within a hole in
the ground, but which, when investigated, pre-
sents nothing but a level surface where cer-
tainly a hole ought to have been! The dorsal
view of the spider is given at Fig. 353; the side
view at Fig. 354; and a diagramatic section ane ;

Fic. 356. Theridium differens, much
view of the accuutns is drawn at Pig. 355, a8 16 © enlarged, grasping her cocoon when
probably would appear when closing up the riage oe BUCGAH ean GE
opening to its burrow.

Another mode in which enemies may influence the formation of habit
in spiders has been suggested by observing the manner in which certain
Theridioids defend their cocoons. Theridium differens, a pretty little spi-
der inhabiting leaves and foliage (Fig. 857), which makes a flossy round
cocoon somewhat larger than itself (about a quarter of an inch in diameter),
will grasp her cocoon in her mouth when annoyed by one’s finger or by

FIG. 356, Fig, 357.

1“Ttude sur les ee de la Famille des Avic larides an of North Athen, Botes de la
Soc, Linn. de Bordeaux, Vol. XLII, 1888, page 11.
* Mygale truncata, Hentz, Spiders U.S., page 16, pl. i, Fig. 1

418 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

disturbance of the surrounding foliage. (See Fig. 356.) Her first impulse
at the approach of what she regards as danger to her offspring, is to
seize the little ball and carry it away to another part of her snare, or
simply to interpose her own person between it and threatened peril, or at
least to secure it by her own personal possession.

Now, it has already been shown (see pages 119, 120) that certain spe-
cies of Lineweavers have acquired the habit of permanently carrying
about their cocoons in their jaws and under their legs. This is their
method of protecting their offspring from assail of enemies. The same
habit, with varying methods, prevails with certain Laterigrades, with most
Citigrades, and perhaps also with some Tunnelweavers. May it not be
that this fixed habit of protecting cocoons by personal possession may
have originated from such occasional acts as that common with Theridium
differens, and which, by transmission and gradual growth, have come at
last to be characteristic ?

CHAPTER XIV.

DEATH AND ITS DISGUISES, HIBERNATION AND
DEATH FEIGNING.

As one passes through the fields in the latter part of September or
early in October he marks the cessation of activity on the part of Ar-
giope cophinaria. The splendid creatures, whose restless vigor
The De- in spinningwork and ferocious activity in capturing prey were so
eae oe apparent a few weeks before, have nearly all disappeared. The
rgiope. ;
males have gone weeks before. Not one of the courtiers that
were seen hanging around the outer courts of their lady loves’ snares has
survived the mating season. Occasionally one notes a female, a shrunken
remnant of her former self, suspended in listless mood upon a tattered
web, or crawling sluggishly around the circle of her orb, weaving in her
spirals as though spreading a table for the last banquet that life affords.

A little further on one will see the dead forms of other individuals hang-
ing in various postures from broken snares, or from tattered remnants of the
silken shield, or from snatches of cross lines dangling from leaves and
bowers. Still further, as one moves on, he sees fragments of the once beau-
tiful shares stretched out at various points between the grasses and branches
of low lying shrubbery. he strands flutter in the breeze. The great cen-
tral patch of white silk flaunts like a tattered banner after a battle. The
radii are snapped, the spirals have lost their viscidity, or have only retained
them to capture hapless insects that expire without even the poor satisfac-
tion of helping rejuvenate exhausted Nature by rendering their lives an of-
fering to the vigor of another creature. The race of Argiope is gone for
the current year.

Where are these noble araneads that so lately brightened and enlivened
the landscape? They have crawled away into various nooks beneath em-
bowering leaves or other cozy retreats, and there have woven the beautiful
basket like cocoons which characterize the species. The last force of life
has been expended in this act and, somewhere near, the dry and shriveled
corpse of Cophinaria may be found hanging, after a little while, to the
threads on which:she perished, soon to be washed down by the rains of
autumn and mingled with the dust beneath. When the warmth of spring
has once more revived the earth, another generation will issue from these
cocoons and go forth to follow the life round of the race that has now
passed away. This record of the decline and fall of Argiope is a picture

(419)

i ee ee
° .

420 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

¥ mn
| "t

Fic. 358. The fashion of death. Argiope cophinaria hanging dead in her snare,
on an ampelopsis vine against a wall,

DEATH AND ITS DISGUISES. 421

of what one may everywhere see of other species during autumn days,
and, indeed, at other seasons also, for the limit of life with some species
is reached before the fall.

I have heretofore remarked that the most natural death of the spider

is, perhaps, a violent one. To feed the hungry maw of a stronger, more

skillful, or more fortunate fellow aranead; to be paralyzed and

The Fash- entombed within a clay sarcophagus by a mother wasp and serve

ee as food for a growing waspling worm; to be snapped up as a

delicate tidbit by birds, toads, and all the other creatures. that

prey upon her—these are some of the modes by which, in the appoint-

ments of Nature, the spider meets that doom which must befall all the
living. And a painless doom it doubtless is, even thus.

But there are some that end their life by what we commonly call a
natural death; that is to say, they do not perish through violence, but
cease to live because of the natural exhaustion of vital forces. It is one.
of the most difficult matters, among the many difficult ones in the study
of spider life, to find an unin-
terrupted opportunity for consec-
utive observations of a spider
while undergoing this fashion of
death. But I have been fortu-
nate enough to create opportu-
nities which have afforded me
satisfactory results. Most of the
examples studied were females of
Argiope cophinaria, which I had
colonized upon the vines in my
manse yard. One of these, called
for convenience Prima, had oe-
cupied a position upon a honey-
suckle vine for several weeks,
and highly enjoyed herself cap-
turing and deyouring numerous
flies attracted to the spot from te. 359, The death fashion of Argiope. Position of
a neighboring stable. The first ee La PRE Aas
stage of mortality was simply a condition of inactivity.

In my daily rounds among my pets, I noticed nothing peculiar in Prima
except that she seemed to be hanging inactive behind her central shield of
white silk. But as this is not an unusual circumstance, it attracted no spe-
cial attention until September 24th, when I found her hanging in a position
that at once indicated disaster. I touched her and tried her sensibilities
in various ways; but» she was dead. She had, indeed, evidently died

—_— i > -- °°

422 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

upon her web unnoticed by me. Now she hung behind her orb in the
position represented by Fig. 359. Only a scant patch of the central shield

remained. One hindermost foot was extended upward almost
First straight from the cephalothorax, and grasped the ragged edge
iain of this patch. The corresponding last leg on the opposite side of
tality. the spider was outstretched in a like position and held to the op-

posite portion of the ragged shield. The third legs on both sides
were holding to straggling threads. The second legs were curled forward
towards the snare behind which she hung, and the claws held to cross
lines of the notched zone. The first pair of legs hung free, and were
projected through the meshes of the snare. Thus, even in death was
maintained the habitual position of the creature when watching for a
victim or an enemy, the first pair of legs being kept free in order to feel
towards and find contact with an object of desire or dread. Between the
patch of silken shield and the parts of the snare immediately beneath was
a great gap, the size of the spider’s body, which had evidently been pro-
duced by the weight of the creature as she hung downward. ‘The whole
web, indeed, was relaxed in all its remaining parts. The abdomen of the
spider hung downward at an inclination from the cephalothorax of perhaps
forty-five degrees, in which position it was held by a trapline attached to
the spinnerets and at the opposite end of the ragged shield.

The next day, September 25th, the position of the spider was sub-
stantially the same. September 26th the first and second legs were bent
over towards the body, and the claws seemed to have entangled with portions
of the cross lines. The palps were as at the time of the first observation,
one stretched out holding a thread by the palpal claw, and the other bent
over a line, as one would bend his arm across a rope by which it was
supported at the bended joint. By September 28th the web which had
been gradually sinking was quite sagged down behind the vines, and but
few portions of it remained; but upon lifting the leaves, the spider was
seen, hanging, but with the legs considerably more procurved. One fore leg,
however, was still stretched out straight, and held to the drooping line by
the clasped claws.

There is little to detail concerning the subsequent history of the deceased
Prima. I watched during the first week of October, and saw the various
fragments of the orb frayed away by the winds and rains, which
were quite severe. But the corpse hung in the position last de-
scribed, the one outstretched leg strained to high tension and supporting
the whole weight of the body. When last seen, the remaining legs were
rigidly bent at the joints and clustered together over and around the head.
October 8th the spider had disappeared, dropped down or washed down by
the rains into the mass of leaves and tendrils below, no doubt, although I
could not find it. Thus the chapter ended; a seemingly quiet, gradual,
painless death; a winding sheet among the leaves like an ancient Egyptian

“Finis!”

a ee ee Loe VTe=—

DEATH AND ITS DISGUISES. 423

mummy, and a sepulchre amid the tangled tendrils of fragrant honeysuckle.
Not an undesirable kind of death and burial.

The second example, the Secunda of my notes, hung upon an ampe-
lopsis vine against the chapel wall. I quote my journal: “. . . . For
two days, September 28th, she has hung y
absolutely inactive. Yesterday I touched
her, and she only slightly moved her fore
legs, then sank back into position, She is
entirely natural in her appearance, and no
one observing her would suspect that any-
thing is the matter with her. This morn-
ing I put a vibrating tuning fork to one
of her legs, and the only sign of anima-
tion she gave was slowly but slightly draw-
ing the legs towards her. Under ordinary circumstances this act would
have produced the wildest excitement. At four of the afternoon I repeated
the test, and action seemed to be a little more decided. The fore legs
were curyed inward, and an hour afterward were not relaxed again. I
then touched the spider with my finger, and she drew her legs up a little
closer, making no further sign... . . Sep-
tember 29th. Secunda has left her position
on the shield, crawled along the stem of an
adjoining leaf, and is hanging with her back
downward and her feet clasped around the
stem close up against the wall.” (Fig. 360.)

For a week thereafter the record contin-
ued with little variation, except that Secunda
would shift her situation a little, several
inches to one side, and above or below. Once
after long hunting I discovered her by see-
ing her swing down by a thread between my
hands. She dropped six or seven inches,
climbed up the thread sluggishly, and re-
sumed position with her feet clasped above
the stem. I never could find her again. She
had doubtless nestled out of sight and died
in the fashion she had habitually maintained
during the few days immediately preceding
her disappearance.

Fic. 361, A dead Argiope hanging in her Another example may be cited from my
snare, (Sexta.)

journal. It was followed up so closely and

continuously that it well illustrates the manner in which spiders pass away

from life. The animal was sixth of my series, and noted under the name

of Sexta. She was transferred from the banks of the Schuylkill River to

Fic, 860. Death fashion of Secunda.

424 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

an ampelopsis vine upon the outer wall of my manse. [or several weeks
she hung in the normal position of her species, frequently changed her

web, and occupied herself in the usual manner of spiders. Dur-
Sexta’s ing cold weather that occurred in the early part of October,
nape} 1888, she appeared to be a little torpid, at least was decidedly

inactive. Throughout those days she would move her legs when
touched by my finger or by a vibrating tuning fork, but showed little
excitement. The last evidence of activity which she gave before passing
into a lethargic condition was to move slowly to one end of her snare
from the centre. The next day, October 4th, I found her with legs
doubled quite over certain lines of her web, and rigid. There appeared
to be a little life in her, but on the following morning, October 5th, she
was hanging in the same position dead. (See Fig. 361.) The axis of her
body was at right angles with the position in which the
spider usually hangs; that is to say, she was stretched
crosswise of her web. ‘The limbs were all bent at the
middle joints, in the angles of which the linework was
looped. The spider appeared to be chiefly sustained in
this way, although some of the feet were still attached
to parts of the snare.

I continue the description by extracts from my note
book : “October 6th. Sexta hangs in the same position.
October 7th. Ditto. October 8th. Certain lines in the
web have given way so that the fore part of the body
has dropped downward, causing the spider to hang now
in her natural position. (See Fig. 362.) The two pairs
of hind legs are stretched out to their utmost extent, and
the feet hold on mechanically to threads. The fore legs
are bunched and bent, as in the first position, with very
little change. The weight of the spider’s body has evi-
Fic, 362, Sextain dying dently drawn out the two hind legs by which she is sus-

eae tained, October 9th, 9 A. M. Sexta still hangs in the
above position. No change observable. A cold morning and clear. Octo-
ber 9th, 6 P. M. The position of Sexta is now changed. The left hind
leg is loose, the thread to which it held having been broken by the wind
or by a dropping leaf. The claw still holds to the line, a broken frag-
ment of which floats out from one side. The body has swayed quite over,
and the abdomen is twisted into a position at an angle nearly forty-five
degrees to the perpendicular. The other legs remain about the same, ex-
cept that the fore part of the body is swung upward and to one side. Its
weight is largely supported on the one outstretched hind leg.” (See
Fig. 363.)

From this date and up to October 22d, Sexta was observed every day,
morning, and afternoon. Although high winds and heayy rains prevailed

a Ne
. r >

DEATH AND ITS DISGUISES. 425

during this period, and the leaves were continually dropping from the
vine, no change at all of any decided character occurred in the position.
The lines maintained their strength and tension. The next day,
however, showed a change. ‘‘October 22d. This morning the
threads of Sexta’s web haye relaxed and broken, and the position is quite
changed. The abdomen is shrunken up, a mere hard, dry shell. . .
October 24th. Fearing that Sexta would be carried off by the high wind
and falling leaves, and wishing to preserve the body, I removed it from
its lines, and the shriveled corpse now rests in my collection at the Phil-
adelphia Academy of Natural Sciences.

I have observed something of the same sort in spiders kept in confine-
ment within my breeding boxes, where I had placed them for various
observations, particularly to secure cocoons.
I was sure to find them some morning lying
upon the bottom of the box, quite shriveled
up and dead. In the case of spiders after
the act of cocooning, the process is
very much as above described, so
far at least as I can judge from
disjointed observations upon various species.
Of course, those spiders which make several
cocoons remained active until the last cocoon
has been spun; but with those who make
but one the forces of life seem to be entirely
or largely expended in the act of maternity.
After a little while the creature hangs to the
maze of lines within which her cocoon is
usually suspended, or to some bit of web ad-
joining, and then simply drops off dead. As
she lies in this attitude the legs are usually
bent beneath the body and towards the mouth
parts. Sometimes they will be found clus-
tered close together just beneath the mouth
or some part of the sternum. The abdomen
frequently shows gaunt and shriveled. In the ease of the spiders above
described, who had not made cocoons, the abdomen immediately after
death was sufficiently plump, at least not shriveled. Some spiders, after
the act of cocooning, have enough energy remaining to spin a web and
even capture prey, but with the Orbweavers which make only one cocoon,
this is the exception and not the rule.

“Finis!”

After Co-
cooning.

Fic. 363. Sexta after death.

Until lately little has been known concerning the possibilities of pro-
longed life among the lower orders of animals. The waste of life is great

426 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

in the natural conditions surrounding most inferior creatures, so that the
immense fecundity of insects and araneads, for example, is abundantly
he checked. I have counted over eleven hundred eggs and young
a ee spiders in the single cocoon of Argiope cophinaria; yet, though
a score of cocoons may hang in a field, one will scarcely

find as many spiders as cocoons the next summer. In efforts to breed
spiders from cocoons, I have at various times seen colonies numbering from

Fic. 364. Argiope cophinaria hanging dead beside her cocoon.

one and two to ten hundred dispersed from the maternal egg nest to sur-
rounding foliage, of which during the year not a single survivor could be
traced.

Bee keepers are well aware of the great mortality among working bees,
caused not only by disease and accidents, but especially by those enemies
which prey upon them. Ants are quite as much, perhaps even more
exposed to loss from accidents, the exigencies of weather, and the appetites

es eo

ee eee

DEATH AND ITS DISGUISES. 427

of various insectivorous animals. There is, therefore, abundant occasion
for the seemingly exhaustless fertility of the queen mothers of formicaries.
These queens probably have a longer life than the workers. They are
larger in size and apparently organized for more vigorous resistance of the
influences which work for their destruction. Moreover, the instinct of
the workers has provided a system of preservation by surrounding the
queen with a guard of attendants which never leave her unprotected, which
care for all her wants, and vigilantly separate her, by a regular system of
seclusion within the portals of the formicary, from many influences which
would prove hostile to health and fatal to life.

How long the ant queen may live in an entirely natural habitat is un-
known, and perhaps cannot be determined. But recently, through the

patience and ingenuity of Sir John Lubbock, we have learned
Lub- that under artificial protection both workers and queens of certain
bock’s : ‘ Ri
Old Ant “Pecies may attain a great age. Some eight years ago I had the
Queens. Privilege of visiting this distinguished naturalist at his country

seat, High Elms, Kent, and examining under his personal direction
his artificial formicaries, and the mode in which they are preserved. At that
time I saw a queen of the Fuscous ant, Formica fusca, which was nearly
eight years old. On the last day of July, 1887, I again visited Sir John
at his house in London, and on inquiry after the aged queen, which I sup-
posed to be still alive, was informed that it had died the evening before,
having at the time reached the wonderful age of thirteen years.

I was permitted to see this venerable queen as she lay in death on the
floor of one of the wide chambers which the workers had excavated in the
soil compacted between glass plates that bounded their formicary. She was
still attended by a circle of the “courtiers” which, according to my pub-
lished observations,’ are in the habit of watching continually upon ant
queens. Some of these attendants were licking the dead queen, or touching
her with their antenne, and making other demonstrations, as though solicit-
ing her attention or wishing to wake her out of sleep. “They do not
appear to have discovered that she is really dead,’ remarked Sir John.
And so, indeed, it seemed. It was certainly a touching sight to witness
these faithful attendants, surrounding the dead body of one who had so
long presided over the maternal destinies of the colony, and seeking by their
caresses to evoke the attention which never again could respond to their
solicitations,

In answer to a letter of inquiry concerning the life of this queen and
her companion, Sir John wrote me? as follows: “As they had lived with
me since December, 1874, they must have been born in the spring of that
year. One of these ‘asd after ype for some she died on the 30th of

1 Honey and Occident. Ants, Chapter IV., page 41, plate vi., Fig. 29.
2 Under date of May 10th, 1890.

428 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

July, 1887. She must then have been more than thirteen years old. I was
at first afraid that the other one might be affected by the death of her com-
panion. She lived, however, until the 8th of August, 1888, when she must
have been nearly fifteen years old.”

This ‘longevity is, as far as I know, unparalleled in the history of in-
vertebrate animals. Such experiments as the above clearly indicate that
artificial environment may have a beneficial influence upon insects as well
as domestic animals, and that the interference of human intelligence may
be a preservative factor, as well as a destructive one in the lives ae even
our most lowly organized fellow creatures.

Early in the year 1882 I received from Dr. Joseph Leidy a specimen
of our common tarantula, Eurypelma hentzii.! As the individual seemed

to be in good health, I preserved its life in order to gain infor-
The mation as to its habits and vital endurance. It was first placed
alder in a large glass globe on a bed of earth, where it was kept for

more than a year. It was then transferred to a wooden box
made with glazed sides and a sliding glass door at the top, the whole be-
ing eighteen inches long, twelve inches wide, and ten high. One end was
filled with dry soil, which was slightly compacted and heaped up; the
other end was sparsely covered with earth. There was thus presented a bit
of level space for the spider to burrow should it be inclined to its natural
tastes. I last saw it early in July, just prior to my departure for Eng-
land. On June 22d, 1887, I made this note: “This spider, which has
been kept ever since 1882, is to-day in good health. It is on the outside of
the earth moundlet in its box, looking hearty after the winter’s fast. It
has had nothing to eat since October last—at least eight months—but has
had water freely. Some flies have been put into the box lately, but I do
not know that they have been eaten.” The spider was then left in the care
of Professor Fronani, who for several summers, while at work in the
library hall of the Academy, had kindly cared for it during my absence,
giving it water and feeding it with insects, particularly grasshoppers or
locusts.

On my return from abroad I was met at the Academy by the intelli-
gence that my tarantula was dead. About the close of July it had de-
scended into the burrow which for several years it had maintained close to
the side of the box, and since then had not come up. Looking into the
box I could see against the glass the fragments of a moulted skin on
one side of the cayity, and on the other side the outlines of the creat-
ure’s dead body. It had evidently died shortly after moulting.

Reckoning its death: as having occurred at the close or July, 1B, the

‘Tt was captured about the beginning of April, 1882, at Hills Tene, Stanislaus County:
California, was kept in a bottle without food for two weeks, then sent to Professor G. 1. TH.
Weaver, at Media, then a student in Swarthmore College. Mr. Weaver fed it on beefsteak,
which it took readily.

TE a CC CCl =
az re bs
'

DEATH AND ITS DISGUISES.

“es SS eel

429

spider was five years and three months in my possession. I have not
sufficient data to estimate accurately the rapidity of growth in this spe-

cies, but judging from such facts and. indications as I have ob-
iaceee. served the animal must have been from eighteen months to two

years old when I received it from Dr. Leidy. At the period of
its death, therefore, it must have been at least seven years old, and may
have been eight or more. It thus attained the distinction of having
reached the greatest age of any spider known to science. How long this
species and members of the Theraphoside generally live in their natural
habitat is of course unknown. I have little doubt that they live much
longer than other tribes, but am inclined to think that it is not usual for
them to reach such an age as my tarantula “Leidy.” In its case, as in
that of Sir John Lubbock’s queen ant, human protection probably pro-
longed life.

Other observations on the age of spiders fall in with this indication
of their vital endurance from the tarantula’s prolonged age. Blackwall
kept spiders of the species Tegenaria domestica and 'T. civilis to the age
of four years.1 Moggridge made a calculation of the age of Trapdoor
spiders, based on average growth in nests of the young; for he established
the fact, which has subsequently been confirmed, that a young spider, in-
stead of abandoning its nest, enlarges it with its growth. He concluded
that it took at least four years to produce a full size trapdoor nest, and, of

course, the architect must be at least that old.? . The most recent

eer information on this point is from Mr. Frederick Enock.* This
eo : ; : pee
eect observer, in an extended and interesting communication on the

habits of the British Atypus, speaks of one individual which
he kept over three years, and which, judging from its size when first
captured, he puts at the age of six years. Other examples, under observa-
tion for more than two years, were well grown when first transferred to
his artificial colony, and at the date of his paper, June, 1885, were still
in good health. He ventures the inference that Atypus is about four years
in reaching maturity, then retains her young for eighteen months under
her care before turning them out to shift for themselves, and after that
lives in vigorous health for a period which he believes may sometimes
reach the advanced age of ten years. Thus, a spider’s life may vary in
length, according to organization and surroundings, from a single season
to two, four, and even eight or ten years.

I may add here, as in the same line of research, that Dr. George H.
Horn, a distinguished authority in the Coleoptera, has called my attention
to the fact that a female of Cybister roeselii was preserved for eight years
of continuous life by Dr. David Sharp.

1 Spiders Gr. Bt. & Iv, page 8. 2 “Warvesting Ants and Trapdoor Spiders,” page 127.
8“The Life History of Atypus piceus Sulz.,” Trans. Entom. Soc., London, 1889,
page 416.

r \ = QE) a ON
: 2 7 a

430 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

III.

If one will go to any woodside or other spot where the foliage of trees
and vines has been amassed, and examine one after another the withered
leaves, he will have opened to him a new and interesting chap-
ter in the life of spiders. This process is familiarly known as
“sifting,” and it is an admirable method of collecting in the
late autumn, winter, or the early spring. Clear away the mass of snow
overlying that windrow of withered leaves, fill a generous basket full, and
carry them into your study. Open carefully the curled leaves one after
another, and you will find a number of spiders of various species, that
have found their winter refuge and rest within these seemingly inade-
quate receptacles.

Here are Saltigrades, nested within their white, thick, silken cells.
Here are all sorts of Tubeweavers, Disderids, Drassids, Agalena, Tegenaria,
Dictyna. Some of them are underneath silken cells, others clinging to
simple strings of intersecting lines. If the weather be very cold, most of
them will be found quite torpid; but in the warm atmosphere of the
room they will soon renew their vitality and freely creep about. If the
temperature be mild, or if the sifting be made at that part of the year
which lies just between winter and spring, the spiders will have recovered
from their hibernation, but many of them will be certain, as soon as they
feel the touch of the inquisitive observer, to double themselyes up in that
strange mimicry of death which marks so many species.

Such an examination as the above has increased my surprise at the
immense host of spiders that must be preserved throughout the winter by

nestling under leaves and forest mold. The autumn broods of
Hibernat- younglings here find refuge in numbers, and when the snows
ean: have been melted away by the south wind and the increasing

heat of the sun, they creep forth from their leafy lairs and
enter upon the active duties of their lives. Nearly all species in all the
several tribes thus find winter homes in such places. This is not only
true of the woods and wild fields, but of the lawns, groves,.and parks sur-
rounding suburban and city homes. When the bright, soft days of April
come, and the gardener begins his annual task of raking withered leaves
together and burning them, my heart has many a spasm of pity at the
reflection that this seemingly harmless and necessary work is the holocaust
of millions of hapless spiders. Thus, even in the discharge of ordinary
duties, man is unconsciously one of the most destructive enemies of the
children of Arachne.

A good time to uncover the winter habits of spiders in the latitude
of Philadelphia is the early' or middle part of April. Frequently there
will come a few successive days of warm sunshine, particularly if the
preceding winter has been mild, that invite the Sedentary spiders from

Winter
Habits.

a a a ee ey ee

DEATH AND, ITS DISGUISES. 431

their lairs, and tempt them to spin their first webs. These webs betray
their winter quarters. Here, for example, along these hedge rows of arbor

vitae are a number of round webs whose proportions indicate ma-
Epeira’s ture weavers, and whose construction gives the experienced eye a
aa token that Epeira strix has probably spun the snare. She is not

en. : : Bee y

upon her orb at this hour of the day, and is doubtless resting in
some secluded spot near by, which spot, considering the season of the year,
is almost certain to be the den within which she hibernated.

The tyro spider hunter would vainly search along the hedge row for
this refuge, but certain signs which experience has taught lead one to a
particular point, where a larger concentration of threads, diverging from
the foundation lines of the orb, form a sort of guide board to the desired
haven. Follow this clue, gently separate the sprigs of foliage, and push
aside the twigs, and one will see a few inches below the surface, at a point
where the branches diverge, a mass of rubbish. It is accumulated between
the forks of the twigs, and has been retained in its position by the same.
These leaves have dropped from the bush above, and have drifted in from
surrounding plants. It is possible, also, though I cannot affirm it, some
of the material may have been collected by the spider and added to the
nucleus which accident furnished. At all events, here is a lump of rub-
bish as large asa hen’s egg. The whole is lashed together by scant threads
of spinningwork, which assist the office of the encompassing twigs, and
brace it in its place at the point of juncture.

If one thrusts a finger beneath the mass, a slight opening will be
found, which is manifestly the door of the den. Now, with fingers or scis-
sors, separate the ball of rubbish, and lo! inside, snugly ensconced in the
very heart of the heap, is the weaver of the web and proprietor of the
den, our interesting friend Epeira strix. Here she has lived throughout
the winter, and, as she is entirely mature, she must have been well grown
when she first went into winter quarters. One day (April 14th), while
walking with my secretary, we found a number of these nests within a
short space in the precincts of Woodland Cemetery, on the banks of the
Schuylkill River. Several mature females and one mature male were dis-
covered, all of them occupying some sort of a den of miscellaneous rub-
bish, gathered together with varying degrees of efficiency. With the com-

* pact roof of evergreen leaves, which forms the outer surface of the plant,
stretched above the den and serving as a screen from snow and frosts, it
is evident that this winter nest is a safe or at least sufficient refuge for
the Orbweaver.

Another favorite winter resort for spiders is the stump of an old tree;
another, the hump of earth and roots which marks the spot where a tree
has fallen. Every hollow and cranny forms a refuge for some species.
Favorite spots are the tubes or “casts” beneath the soil formed by decayed
roots. ‘Tearing away the earth at this point, for example, one finds the

432 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

soil penetrated by a well rounded pipe, whose walls are protected by the
outer bark of a root. Only that remains. All the rest has per-
Other ished, leaving a long tube in the earth» wherever the root had
Winter yun. Here many spiders have found winter refuge, and from
Resorts.
the mouth of one of these natural dens a full grown specimen
of Tegenaria medicinalis is unearthed.
Young spiders survive the winter in the admirably arranged cocoons pro-
vided by the maternal instinct. But early in spring many adults of both
sexes are found nearly full grown, that have also safely weathered
Winter the cold months. I have at various times in the winter col-
reece lected Epeira strix,and have found the species adult in spring.
Specimens of Strix may be frequently taken during winter
months from rolled leayes within which they have withstood our hard
frosts. These rolled leaves also serve for nests during summer. Dr. George
Marx has informed me that on the capacious Government grounds in
Washington City he often sees
such curled leaves suspended,
conspicuous amid the verdure-
less branches, and has learned
to recognize them easily as the
winter quarters of this species.
It of course follows that, either
from purpose or by the acci-
dental enwrapping of threads
during continual journeys back
and forth, trailing her dragline behind her, she secures the leaf from falling.
A vast colony of Epeira sclopetaria inhabits the boat houses grouped
around the inlet at Atlantic City.' I visited this colony in the latter part
of May (1882), when the season had been remarkably backward, cold, and
rainy. The trees on the island had not yet leaved; insect life had
scarcely appeared; in short, the season had adyanced little further than
the first of May in ordinary years. The inlet colony, however, had already
appeared in large numbers, and had swung their orbs between the timbers
of the houses and the piles which supported them. These were of various
sizes, full grown, half grown, and young several weeks out of the cocoon.
All the cocoons, which were thickly laid along the angles of the joists and -
cornices, were empty. The number of young spiders was, however, remark-
ably small, a fact which I could account for only on the supposition that
in the absence of the usual insect food supply the adults had been
driven to prey upon the young, and the young upon each other to an un-
usual degree,
Many of these spiders were hanging in the centre of thei* round snares.

Fic. 365. Arched shelter tent for winter service.

‘See page 232 and Fig. 256.

er ere ee UCU - aii-
— =e - “ve li dia -

DEATH AND ITS DISGUISES. 433

Others, the greater part indeed, were covered within a thick tubular or
rather arched screen (Fig. 365), open at both ends, which was bent in the
angles of the woodwork, or were sheltered beneath an irregular rectangular
silken patch (Fig. 366) stretched across a corner. Many others were bur-
rowed behind cocoons quite covered up by their thick, flossy fibre, in
which condition they had undoubtedly spent the winter. I have found
examples of Epeira strix blanketed in precisely the same way during the
winter months. I asked some of the young boatmen what the spiders did
in the wintertime. ‘They crawl into their bags,” one answered, referring
to the screens and tubes above described (Figs. 365 and 366), “and stay
there. They came out about a month ago (the last of April), and then
shed. A couple of weeks ago the sides of the houses were all covered
with these sheds.” “Shed,” it should be understood, is vernacular for

“moulted” or “ moult.”
English spiders have like habits. Epeira apoclisa frequents gorse, heath,
and rank herbage growing near marshes, lakes, pools, and brooks, or
other damp situations, among which it constructs a dome shaped

onal cell of white silk of compact texture. In this cell, after distrib-
meee tl uting upon its exterior surface the withered leayes of plants,

and enclosing its entrance with a tissue of silk, the spider passes
the winter in a state of tor-
pidity.! It is said that Apo-
clisa possesses the power of
closing the door of her nest
against intruders by seizing
the sides with its claws. The
eggs are placed in her cell, en-
closed in several slight, round-
ish, yellow cocoons about half
an inch in diameter, Simi-
lar nests attributed to Epeira
quadrata, although Staveley?
thinks the deserted nests of
Kpeira apoclisa are alluded to,
are selected by the dormouse,
according to Rennie, as a ready
made roof for its nest of dried grass. That the old spider dens are not
accidentally chosen by the mouse appeared from the fact that out of
about a dozen mouse nests of this sort found during winter in a copse
in Kent, England, every second or third one was furnished with such a
roof.®

Fic. 366. A winter bivouac tent of Epeira sclopetaria.

! Blackwall, Spiders Gt. Br. & Ty, page 821. 2 British Spiders, page 230,
* Rennie, “Insect: Architecture,” page 109.

434 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

The winter habits of the Retitelarize are various. Many of them creep
into hollow trees, into holes and crevices of divers sorts, or spend the
‘winter under fallen leaves and in vegetable mold. Many of
Line- them have great powers of endurance, and in the neighbor-
Tea ee hood of houses, barns, and outhouses may be seen in the coldest
Tribes, Weather simply snugged up against a corner or angle of wall or
woodwork, with a few straggling lines beneath them, taking the
cold weather without any protection or attempt to secure such. Tube-
weavers, with few exceptions, find their winter homes among leaves and
forest mold, in hollow roots, deep crannies, in rocks, underneath stones,
and like positions. These are also common winter resorts of Laterigrades
and Saltigrades, although they prefer lodgings underneath old bark. But
the latter always spin around themselves a tube of thick silk, which serves
them as a blanket. On a warm day in winter these vigorous little creat-
ures may sometimes be seen jumping about upon the surface of the snow,
having been attracted by a patch of sunlight pouring upon their winter
dens to try their fortunes at winter hunting.

The Lycosids, without exception, as far as I know, spend the winter in
earth burrows. One example of Lycosa tigrina observed by Mrs. Treat
made but little change in the dome shaped covering of her burrow until
November, when it was cut down level with the ground, perfectly con-
cealed with leaves and moss, and held firmly down with strong webs. This
covering remained until the following April. In spring, the gardener, not
knowing that this spot was set apart for special study, raked away the
leaves and rubbish, preventing observation of the manner in which Tigrina
herself would have removed her winter covering; but in a few days there-
after she had made another cover, not like the flat winter thatch, but more
like a little room.!

There is much to learn concerning the life history of Lycosids,
and it may be yet found that their winter life shows a greater activity
and variety of habit than has generally been supposed. It is
possible that they may not remain enclosed within their bur-
rows during the entire winter in a state of semihibernation, Dr, Allen
Gentry, a careful observer, informed me that he observed this incident
while visiting a frozen pond in the vicinity of Philadelphia. He cut a
slab from the ice, about eight or ten feet from the bank, and was sur-
prised to see spiders running about in the water. They were passing from
point to point by silken lines stretched underneath the surface between
certain water plants. Several specimens were collected, but unfortunately
were not preserved. They were supposed to be Lycosids, and, from Mr.
Gentry’s description of the eyes, his supposition is evidently correct. It
is a remarkable and notable fact that these creatures can thus live in full

Lycosids.

‘Am. Naturalist, August, 1879, page 488.

DEATH AND ITS DISGUISES. 435

health and activity, within the waters of a frozen pond, in midwinter,
and so far from the bank in which their burrows are commonly found.
It has been believed heretofore, and doubtless it is generally true, that
Lycosids winter in deep burrows in the ground, sealed up tightly to main-
tain a higher temperature. But the above observation opens a new and
strange chapter in the winter behavior of these araneads, as well as in the
amphibious nature of their habits.!

TY.

The effect of low temperature upon spiders was observed in several
young specimens of Theridium tepidariorum. ‘They hung on a few short
lines to the plastered wall of a brick building on my premises,
the plaster being laid directly on the brick, forming a cold sur-
face. The spiders were protected from wind and snow, but
wholly exposed to the frost. On January 14th (1885), with thermometer
ranging from 20° to 25° above zero (Fahrenheit) the spiders were hang- |
ing motionless. When touched by the tip of a pencil they dropped down
in the usual manner of their kind, holding on by the outspun threads,
which reached a length of over one foot. They ascended to their perch
afterward, and crawled over the roof for a short distance.

At a temperature of 18.6° they again were able to drop from the perch.
January 19th, with the thermometer ranging from 17.5° to 20°, they seemed
less active; one individual, when touched, dropped about one inch, another
six inches. Four hours thereafter they were suspended in the same posi-
tion and place. As the natural habit of the creature is to ascend in a
moment or two after disturbance, this shows that the frost had affected
the normal energy. One of the specimens, however, on being gently lifted
upon my finger, moved its legs and very slowly began to ascend. Five
hours thereafter it was at its perch against the roof. These spiders, at
this temperature with some variations (January 21st), moved their position,
one passing along the angle of the roof a distance of four feet. This
change of site was probably caused by the annoyance which my experi-
ments produced.

February 11th the thermometer stood at zero; in West Philadelphia,
where my observations were made, the temperature was lower. On the

12th the Signal Service reported 1° above zero, at my house it

Hiberna-
tion.

Sudden was below zero. On this day I removed from its position one
Resusci- P eh ;
aol ain of the specimens, a young female about two-thirds grown, and

placed it in my library, where the temperature was summer
heat. She was put upon a table in the sunshine, at which moment her
legs were drawn up around the cephalothorax in the usual hunched way
when torpid or feigning death. There was a slight and regular pulsation

"Proceed. Acad. Nat. Sci., Philadelphia, 1884, page 140.

436 AMERICAN SPIDERS AND THEIR SPINNINGWORK,

of the fect. In less than ten minutes, upon being touched, she stretched
forth her legs and began to move slowly over the paper upon which she
had been placed. When touched, her motion was much accelerated, and
she began vigorously to perambulate her bounds, meanwhile anchored to
and pulling out after her the usual dragline. When lifted up on the tip
_of a pencil, she spun out a long thread, to the end of which she hung in
the little foot basket of silken cords which I have elsewhere described.
Indeed, her action was in every respect normal, and showed a remarkable,
sudden, and complete renewal of activity after so long an exposure to such
extreme cold.

February 26th, a younger specimen, about one-third grown, hanging
in a crevice in the site above described, when touched and lightly pressed
down, slowly moved its legs and began to struggle back to its perch. The
thermometer ranged from 20° to 25° above zero; on the day before the
range was from 21° at 7 A. M. to 28° at 11 A. M. During the six weeks
over. which these observations extended the temperature was unusually low
for this vicinity; for a great part of the time the thermometer stood below
zero. The month of March following was unusually severe, the thermom-
eter frequently reaching winter temperature. On the Ist of April, how-
ever, the above named spiders and others of a younger brood were in their
webs, hale and active, having been drawn out by the first soft days of

spring. It would seem, therefore, first, that the hibernation of
pe spiders, of this species at least, is not accompanied with a great

degree of torpidity; second, that they preserve their activity and
spinning habit while exposed to cold ranging from freezing point to zero
(Fahrenheit); third, that after long and severe exposure the recovery of
complete activity, when brought into a warm temperature, is very rapid,
almost immediate; and, fourth, that on the return of spring, even after a
prolonged and severe winter, they at once resume the habits of their
kind. The above experiments were made upon Theridioids, but I have
made like trials with other species, as Epeira strix, Dictyna philoteichous,
and Eurypelma hentzii, and the results vary in no essential particular.

Tn all the above specimens the abdomens were full, indicating perfect
health. Other spiders hung upon their webs with shriveled abdomens,
quite dead, among them one of my specimens, a male who died during
the course of observations. A Pholeus phalangioides hung thus dried up,
holding with a death grip to her web by the two fore pairs of legs, which
supported the cephalothorax in a position parallel to. the plane of the
horizon, while the long abdomen hung down at right angles thereto, and
the third and fourth pairs. of legs were drooped downward and backward.
I could only conjecture that this and other spiders perished by the cold.
The living individuals were all characterized by the plump abdomen, as
though there had been little or no absorption of tissues for nourishment
of life. There appeared to be no growth during hibernation.

OOO ——— eS er a ==

DEATH AND ITS DISGUISES. 437

Wes

Many spiders have the habit, which belongs to certain insects and pre-
vails eyen among the vertebrates, of feigning death. This habit, which
is common among many species of spiders, appears to be par-
ticularly developed in the Orbweayers. One who touches an
Orbweaver when hanging upon its web will often be surprised
to see it suddenly cast itself from the snare, or appear to drop from it, as
though shot off by some unseen force. Unless he understands the nature of
the creature, he will be utterly at a loss to know what has become of it.
In truth, it has simply dropped upon the ground by a long thread which
had been instantaneously emitted, and had sustained the aranead in its
remarkable exit, so that its fall was not only harmless but its return to
the web assured. If the creature be now examined it will be found mo-
tionless. Its legs are drawn up around the body, and to the inexperienced
eye it has the external semblance of death. In this condition it may be
handled, it may be turned over, it may be picked up, and, for a little
while at least, will retain its death like appearance.

It has been conjectured that this behavior is simply the result of fear,
and is largely beyond the volition of the spider. It is a case, in other
words, of what has been called kataplexy, or fear paralysis. It is, per-
haps, difficult to disprove this theory, but I can by no means accept it. To
me it scems a case of genuine “’possuming,” if I may use a term which
was commonly applied in the West, during my boyhood, to characterize
all shamming or feigning among our associates.

Dr. Preyer, of Jena, has published his experiments on animals while
under the influence of sudden fright,! the general trend of which is to

show that unconsciousness is the resulting state. I do not know

Death
Feigning.

? . . .
eee what value these experiments may have with experts in that

ata- ‘ :
uleer general field of research, but I cannot accept the conclusion as

to spiders. Mr. Campbell, however, appears to favor the theory.”
Examples will occur to every one, of wild animals met in walks through
the woods or fields, suddenly pausing as though the first appearance
of a supposed enemy had shocked them into inaction, A child falls, and,
though not hurt, loses himself for a few seconds. When regaining con-
sciousness he bursts out into a loud roar. Many moths never attempt to
fly when touched, and the white ermine, the satin, the swallow tail, and
the male ghost moth will fall as if paralyzed when a net is swept under
them at night while on the wing. The clouded yellow butterfly will drop
as if lifeless when closely pursued. Many species of beetles are inimitable
death feigners, as most boys know who have any knowledge of field life.

1 Samlung physiologischer Abhandlungen, Zweite Reihe, Erster Heft, 1878.
* Observations on Spiders, page 46.

i i

438 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

But it is needless to multiply examples; let us turn to the animals with
which we are here chiefly concerned.

I have frequently watched spiders in this condition, to determine the
point in question, and their behavior always impressed me as being a
genuine feigning of death, and therefore was entirely
within their volition. The evidence is of such an
indefinite nature that one can hardly venture to
give it visible expression, but my conviction is none
the less decided. I may say, however, that my ob-
aS S\ servations indicate that the spiders remained in this
Fic. 367. Dorsal view of condition as long as there seemed to be any threat-

Kabyrinth spider in actof ened danger; now and again the legs would be re-
eath feigning.

laxed slightly, as though the creature were about
getting ready to resume its normal condition, but at the slightest alarm
withheld its purpose and relapsed into rigidity. The slight un-
Not Fear Glasping of the legs, the faint quivering indications of a purpose
ee to come to life, and then the instant suppression of the purpose
untary. Were so many evidences that the power of volition was retained,
and that the aranead might have at once recovered if it had

been disposed to do so.

Again, I think that I have never noticed anything like that gradual
emergence from the kataplectic condition which one would naturally expect
if the act were not a voluntary one. On the contrary, the spider inva-
riably recovered, immediately sprang upon its legs, and hoisted itself to its
snare, or ran vigorously away among the grasses.

Two positions of the Labyrinth spider while in the act of death feign-
ing are here presented, from a number of sketches made from Nature,
Fig. 367 represents her from a front view, leaning slightly upon one side.
The two fore legs on the further side are doubled under the face, and the
feet may be seen projecting on the opposite side;
that is, nearest the observer. The other pair of fore
legs are doubled under the side, the feet almost
touching the third and fourth legs, which are bent
in a position quite like that commonly assumed
when the spider is sitting at her natural rest. The
whole attitude appears to the familiar observer en-
tirely different from any posture during death, and
this may perhaps be seen by comparing these draw-

see
ings with the death fashions shown in the first pages jg. 39g, ventral view of
of this chapter. Labyrinth spider while

ee F pi ie death feigning.
The second position is shown at Fig. 368, giving

a view of the same spider from the lower part of her body while resting
upon her back. The fore legs are all bent and doubled over around the
mouth parts, the feet extending almost to the lower end of the sternum.

EL ———— ee ee a
nr " - 1s . - : me. ‘

DEATH AND ITS DISGUISES. 439

The third and fourth pairs of legs are folded in an easy position upon
the venter. Applying a magnifying lens to these legs, one can see that
they are connected by threads, which are attached to the spinnerets in the
ordinary way, showing that the aranead is by no means unconscious of

and indifferent to her usual methods of escape.
I give three other drawings which present in natural size three positions
assumed by Epeira trifolium while death feigning. Two of these (Figs. 369
and 370) represent her lying upon her back with her claws doub-

Sey es led up in the manner previously described of the Labyrinth spider,
eataid ~ and showing the same readiness to immediately relax the limbs

and assume the ordinary position. ‘The third drawing (Fig. 371)
shows the spinnerets and two fourth pairs of legs holding on to the apex
of little pyramids of threads which had been instantly thrown out just as
the spider passed into its death feigning condition. One third leg may also
be noticed, reaching downward to the spinnerets, and holding on to a line
which had been outspun at the same time.

This action in itself seemed to me sufficient indication that the spider
retained entire control of her faculties. Instead of falling at once into

FIG. 369, Fic. 370.
Fic. 871.

Death feigning attitudes of the Shamrock spider.

fright paralysis without any preparatory efforts at protecting herself, she
secured herself by her spinningwork anchorages from being carried away
without her knowledge, and gave herself the means of recovery to natural
position. Of course the period of time between this act of self protection
and the assuming of the death feigning attitude could scarcely be caleu-
lated; yet the whole behavior showed that after the act which is supposed
to have induced kataplexy, and between it and the kataplectic attitude, there
was this deliberate effort to secure herself in the ordinary and natural
manner. This was so manifest that, on observing it time and again, I was
extremely amused by the manifest stage effects of the aranead actress, and
could not help comparing it with the alleged method of certain women ad-
dicted to voluntary “fainting,’ who are said deliberately to pick out the
most agreeable spot on which to fall, and to adjust their limbs and drapery in
the most graceful and convenient manner before the faint! In the case of
the Shamrock spider also, as with Labyrinthea, the recovery from the death
feigning attitude into natural posture showed, in the transition, the marks

440 AMERICAN SPIDERS AND THEIR SPINNINGWORK.,

of entire self possession, as though consciousness had never for a moment
been lost.
My conclusions in this matter are substantially supported by the ex-
periments made by the Peckhams, which are recorded quite at length!
These naturalists made two hundred and ten experiments on this
aac subject upon spiders taken from fifteen different genera, and their
Studies. resulting conclusion is that no spider under their observation
ever fell into a kataplectic condition. A few of their experi-
ments may be quoted, with the remark that, as far as they cover the spe-
cies experimented upon by myself, they are confirmatory of my results.
One of the best death feigners is Epeira bombicinaria, a species iden-
tical with Epeira parvula. A pretty little female was softly touched as
she hung in her web. She fell two feet, and then swung to a neighbor-
ing branch, where she crouched motionless for three minutes. Being again
gently touched, she fell to the ground with her legs outstretched, and then,
quickly drawing them in, remained clinging, in a very inconspicuous heap,
to a blade of grass. Here she stayed motionless for one hour, when she
was placed in a bottle, carried into the house, and, still keeping perfectly
quiet, was shaken out on a table. After two hours she was pushed by
the end of a brass rod. Then her legs were lifted one by one with a
needle. She seemed so lifeless that they began to wonder if
they had been watching a dead spider after all. They finally
touched her with the point of a needle, but at the first sugges-
tion of a prick she ran rapidly away. She was knocked over as
she ran and remained motionless just as she fell, resting on the cephalo-
thorax with all the legs drawn closely in, excepting one which was slightly
extended, She did not look like a live spider, nor like a dead one, nor
like anything except a bit of bark or lump of dirt. She laid thus with-
out a perceptible quiver for more than two hours and a half, and then
suddenly ran away. She was reduced to quiet several times after this,
but was less patient and endured no more handling. She did not usually
lie still just as she fell, but deliberately gathered up her legs in such a
way that they were indistinguishable from each other and from her body.
Another example of remarkable death feigning was a large female
Kpeira infumata, She was put into a tumbler and left until the follow-
ing morning, when one of the Peckhams, on looking at her, exclaimed
that she was dead! Her legs were drawn up and bent, and
Infu- :
cine she looked stiff and dry. She was handed from one to an-
Feigning. Other of those present. Her demise was duly regretted, and her
wonderful protective coloring was remarked upon. She was then
put back into the tumbler. An hour later, much to their astonishment,
she was found moving about, alive and well. As they were experimenting

Parvula’s
Sham-
ming.

* Journal of Morphology, Vol. I., No. 2, 1887, page 408, sq.

DEATH AND ITS DISGUISES. 44]

at this time upon the color sense, Infumata was placed in one of the boxes
of colored glass described in a preceding chapter, and, at intervals of one
hour during the day, was moved from the section in which she had set-
tled to another. Every time this removal was made she fell stiffly on
her side, drawing her legs in and remaining thus for about three minutes.
In experiments with her afterwards, the Peckhams found that, when
knocked about on a table, she would stay in the position in which she
fell, although this was often an uncomfortable one. She showed no sign
of life when rolled about, but jumped up at the least prick of a needle.
She never remained quiet for more than twenty-seven minutes, and never
absolutely motionless for so long a time as this, there being slight quiv-
ering movements of the legs and palps at intervals of three or four
minutes, !

The Peckhams had found no spider that would endure bad treatment
without showing signs of life, until they experimented on the Insular spi-

p der. When put on a table, Insularis acted much as Infumata
aa hae had done, but had no such rigid, lifeless appearance. When
she was knocked or touched with the point of a needle, there
was a convulsive twitch of the legs, though she seemed to be trying to
keep quiet. The first time she was pricked so as to puncture the skin
she remained motionless. But at the second puncture she ran. After-
wards, from both males and females of this species, the observers obtained
similar results, once finding an individual that did not run until the skin
had been punctured five times. When the needle entered the skin, there
was usually a twitching of the legs, which seemed to show that sensation
was present. Outside of this species the Peckhams found no spider that
would endure a puncture of the skin without running away, and_ they
rarely found one that would keep quiet while being handled.”

Such a remarkable instinct as that of death feigning did not, of course,
escape the observation of such an incomparable naturalist as Darwin. He
tells us*® that he carefully noted the simulated positions of sey-
enteen different kinds of insects, including among them a spider.
These belonged to the most distinct genera, both poor and first
rate shammers. He afterwards procured naturally dead specimens of some
of these insects, and others he killed with camphor by an easy, slow death.
The result was that in no one instance was the attitude exactly the same,
and in several instances the attitude of the feigners and the really dead
were as unlike as they possibly could be.

The Peckhams in the course of their experiments received the impres-
sion that the habit of keeping still after dropping must not only help the
spider to avoid detection, but must also make it more certain of finding

Darwin's
View.

! Thid., page 410. * Tbid., page 410.
* Essay on Instinct, Appendix to Mental Evolution of Animals, by G. J. Romanes,
page 363, ,

442 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

its way home after the danger is over. There would thus be a double

advantage in absolute quiet.! It will be remembered that as a spider

drops from a web or other roosting place it spins a line, which forms a

straight path backward from the starting point to the stopping

Origin of point. It can thus easily return to its snare or roost by means

cae of this trapline, provided it remains quiet at the first point of

‘stoppage. But, if the spider moves, its trapline becomes a drag-

line, the end of which adheres to the first point of stoppage; after another

short interval this is attached to another point, and so on, to another
and another as the spider moves.

Now, the Peckhams think that this last action tends to confuse a spi-
der, and make its path homeward indirect. In this view they seem to
think themselves justified by some experiments made with the Labyrinth
spider, and to some extent I have no doubt they are correct. But I hardly
agree with them in the importance which they give to this fact as em-
phasizing the theory that it would be of great value to the spider to re-
main quiet at the point first reached after dropping from its web. It
seems to me that it is not difficult for a spider to return by its dragline
to the point where it might reach its dropline, unless, indeed, it should
wander far into the mazes of leaves, or by any misfortune its dragline
should be broken and thus lose its trail. The truth or falsity of this
view is interesting, because of the opinion of the Peckhams that the pos-
sibility of losing itself makes it much more to the interest of the spider
to remain quiet at the place it first reached when dropping from its snare ;
and, further, that this usefulness of the quiet attitude may have been the
starting point from whjch, by natural selection or otherwise, the death
feigning habit may have been developed.

The matter seems to me to require further test before one can posi-
tively decide. At all events, the Peckhams accept Darwin’s explanation
of the habit of lying motionless as the result of natural selection, and
that it has been acquired by different species in different degrees accord-
ing to its usefulness in their various modes of life. Thus we find it in its
greatest development among the comparatively sluggish Epeirids, whereas
it is badly developed or lacking in the running or jumping spiders which
are able, as any one who has pursued them will testify, to move with as-
tonishing rapidity.”

In connection with this subject the question naturally arises as to
whether insects show any sign of fear in the presence of spiders. Camp-
bell only once observed an attitude in a fly which might be taken
as coincident with fright paralysis. The fly was about one and
a half inch from Tegenaria domestica, was busy cleaning itself,
when suddenly it stood motionless in the very act of rubbing its claws

Fearless
Flies.

1 Op. cit., page 413. * Ibid., page 417.

eee = i — -
. .

DEATH AND ITS DISGUISES. 443

together, until it was shortly afterward seized.1_ Mr. Belt states that he has
seen cockroaches retreat in full haste when they had unexpectedly ap-
proached a large spider.2 I have already, when speaking of warning
coloration (page 340), expressed my lack of faith in the supposed paralyzing
influence of spider enemies, at least on insects. Of a vast number of in-
sects, especially flies and grasshoppers, fed to spiders of various species in
captivity, I have not noted and do not remember a single individual that
showed the least evidence of fear or disturbance of any sort. Even when
placed in the box with the giant of the order, the huge Tarantula, insects
appeared unaffected. *

These observations of the apparent fearlessness of flies in the presence
of their natural enemy is confirmed by Moggridge.4 He habitually fed his
captive spiders with common house flies, and remarks that it was curious
to see how entirely the latter were wanting in any instinctive fear of even
the largest spiders. They would creep between a spider’s legs, causing it to
start as if electrified; and frequently it was not until a fly, after repeating
this annoyance several times, actually walked up to and almost touched
the fangs of the spider that it was punished. Certainly such facts indi-
eate a lack of anything like fear paralysis or consciousness of danger on
the part of flies in the presence of spiders.

There is no doubt that some higher animals possess the power of vol-
untarily assuming the external form of death. Numbers of well authen-

ticated examples of this power are recorded among men. Dr.
Feigned § Weir Mitchell related to me an example which occurred under
Death his father’s observation many years ago in a Chinese port. A
Among ‘18 father’s observ yy go port.
‘Neu: Chinaman came on board for purposes of barter, and made him-

self so disagreeable by his importunity that he was finally or-
dered to be put off the vessel into his boat. He resisted the order, and
as it was being enforced fell down apparently dead. All efforts to restore
him were fruitless. He was taken ashore amid great lamentations on the
part of his countrymen and friends, and of course damages were assessed
upon the shipmen. The officers refused to do anything unless the body
were brought on board, and it was accordingly carried to the ship in a boat,
and laid down upon the deck, still maintaining every appearance of actual
death. The officers, whose suspicions were thoroughly aroused by this time,
still refused to pay the demanded recompense until they had made some
last, severe, and satisfactory test that the man was actually dead. As they
were about to proceed with this test the supposed dead man rose to his
feet, and, with grumbling and maledictions, which were echoed by his dis-
comfited associates, descended to the boat and pulled ashore. The self hyp-
notism in this case was so decided that the shrewd American observers
were for a long time thoroughly deceived.

' Observations, page 47. 2 The Naturalist in Nicaragua, page 110.
* See also Vol. I., page 256. * Trapdoor Spiders, page 246.

~~ Ne

444 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Well authenticated cases of self hypnotism, far more striking than
these, are recorded as occurring in India. One was seen by Captain Os-
born—a fakir who buried himself alive at the court of Runjeet
Sonny “Singh for six weeks. Another was observed by Sir C. KE. Trev-
notism. ‘ ens : ‘
elyan, of a fakir who buried himself for ten days.’ In these
cases transition from the normal condition of life to the appearance of
death was gradual, but was undoubtedly voluntary.
Most persons are acquainted with the paralyzing influence of fear, or
the sudden excitation of emotions of any kind, whether sorrowful or joyful,
upon the human mind. The most quick witted are subject there-
Symp- to, and persons with more sluggish intellect are even more af-
aoe of fected thereby; but the symptoms of this action, so far as I
Paralysis, have been able to study them in human subjects, are wholly
different from those which I have observed in the case of spi-
ders. Of course, the vast difference of grade between the two creatures
in the order of organization and intellect may be considered quite suffi-
cient reason for this; yet one might be excused for expecting that there
would at least be so much of analogy between the two as to form a basis
for judgment. In the absence of such analogy I must adhere to my opin-
ion that the behavior of the spider is a genuine case of voluntary death
feigning. One who has observed the process will come to the conclusion
that Arachne’s histrionic abilities are of no mean ‘grade.
The purpose of this habit is undoubtedly protective. I have often
been deprived of coveted specimens by their sudden exit from the web
just as I was about to seize them, and, although I knew thor-
See oughly their habit, and just where to look for them in the grass
eae beneath the snare, I have frequently been disappointed in my
search. One may, therefore, well conceive the amazement, or
whatever emotion there may be akin to that within the minds of lower
animals, that seizes upon the raiding bird or wasp which darts, with
seeming good aim, at the plump prize in the centre of the snare, and finds
beak and claws grasping empty air or seizing only strands of the still
quivering web. One might carry the fancy still further, and imagine
how the spider from its grassy shelter must laugh, or go through what-
ever intellectual process may be analogous to that action in the spider
brain, when it thinks of the discomfiture of its enemy as it flies empty
away !
When we come to think of the origin of this habit, perhaps, we may
be justified in giving some place to the theory of fright paralysis.? Pos-
sibly the success in escaping hostile attacks experienced by some remote

‘See observations on Trance or Human Hibernation, by James Baird, M. R., C. 8. E.,
C. M. W. 8., London, 1850.
2 Romanes, Mental Kyolution in Animals, page 308.

OO ee eee eS —— |

DEATH AND ITS DISGUISES. 445

spider ancestor may have been a feeble beginning of the habit, which
£: gradually was developed into the fixed characters which we now
Origin observe. A supposition of this sort, it is true, has no facts to
er support it, but is in accordance with prevailing ideas as to
‘the evolution of many, if not all the interesting traits in ani-

mal behavior.

In this connection one may perhaps allude to the remarkable sem-
blance of death into which the spider involuntarily falls when pricked
with the sting of the digger wasp. I have referred to this in the preced-
ing chapter, and quote here in confirmation a remark of Mr. Fabre, de-
scriptive of the condition of Lycosa narbonensis of France, after being
paralyzed by Pompilus annulatus. The spider is immobile, lithe
as when living, without the slightest trace of a wound. It is
life, in fact, minus movement. Viewed from a distance, the tip
of the feet tremble a little; and that is all. One specimen disentombed
from a wasp’s burrow was placed in a box, where it kept fresh, presery-
ing the flexibility of life from the 2d of August to the 20th of September,
a space of seven weeks.!_ With spiders in such condition there is really
no appearance of death. They are unconscious though living, and there-

Death
Paralysis.

Habits of Insects, page 210, 1882.

PART VI.—FOSSIL SPIDERS.

CHAPTER 2;
ANCESTRAL SPIDERS AND THEIR HABITS.

Tur interest which attaches to the spider fauna of the present era
naturally reaches backward to those of geologic time. I therefore under-
take a sketch of the fossil remains of spiders, with particular view to
gaining, if possible, some key to ancestral habits. The material for our
review is not abundant, but fortunately we have sufficient data to give
our inquiry an intelligent interest.

According to Scudder,! one hundred and ninety species of spiders have
been discovered from the Tertiary deposits of Europe. Mr. Seudder de-
scribes thirty-two from America in his work on the Tertiary
Insects, of which fourteen are Orbweavers, being forty-four per
cent of the whole number of species. The proportion of known
fossil Orbweavers in America is much greater than in Europe.

A notable addition to our knowledge of the spiders of Tertiary Europe
has been made by Gourret in a paper on those of Aix, in which, among
others, eighteen species are described, including two of Erisoide, two of
Lycosoidee, one of Theraphosoide, one of Dysderides, two species of Her-
sillioidee, two species of Erocteroide, one Enyoide, none of which families
had been before found in European rocks, and the last two named not
even in amber.’

Of the fossil spiders of Europe, one hundred and sixty-eight are de-
scribed from enclosures within amber, forty-one only from the rocks. It
will thus be seen that while Europe is much richer in spiders when the
amber fossils are included, America has yielded more than three-fourths as
many from the Tertiary rocks and one from the Carboniferous.

The fossil spiders found in America are distributed as follows: Salti-
grades, three species of Attids; Laterigrades, three species of Thomisids.
Tubitelarie : Dysderides, one species; Drassides, five species; Agalenades,
two species. Retitelarize: Theridides, four species. Orbitelarie: Epeirids,
fourteen species.* Arthrolycosa antiqua is probably a Territelarian.

Fossil
Spiders.

‘ Tertiary Insects of North America, pages 48-90, U. 8. Geol. Survey of the Territories,
Vol. XIII. * Thid., page 52. 5 Thid., page 49.
(446)

ANCESTRAL SPIDERS AND THEIR HABITS. 447

The discovery of fossil spiders on this continent is confined chiefly to
a single point, Florissant, Colorado, although Green River, Wyoming, and
: Quesnel, British Columbia, have contributed some specimens.
Biapes The remains occur in a series of lacustrine deposits formed
Sites. p
within an ancient lake basin which lies in the valley of the
present South Fork of Twin Creek, and of the upper half of the same
after the South Fork has joined it. This ancient Florissant Lake basin
lies among a series of low wooded hills and ravines marked by an irreg-
ular L-shaped grassy meadow. At the period of the Oligocene this ele-
vated lake must have been a beautiful shallow sheet of fresh water. It
was hemmed in on all sides by granitic hills, whose wooded slopes came
to the water’s edge, sometimes, especially on the wooded sides, rising
abruptly, at others gradually sloping, so that reeds and flags grew in the
shallow waters by the shore. The waters of the lake penetrated in deep
inlets between the hills, giving it a varied and tortuous outline. Steep
promontories projected abruptly into the lake from either side, dividing
it into a chain of three or four unequal and irregular ponds united by a
narrow channel to a larger and less indented sheet, dotted with numerous
long and narrow wooded islets just rising above the surface. Along these
wooded islands and indented shores, a most congenial habitat, the spiders
of the Tertiary had their homes. The Orbweavers and other Sedentary
groups hung their snares among the branches of young hickories, oaks,
birches, poplars, willows, elms, wild roses, sumac, alder, ferns, catalpa, and
bignonia, precisely as in our own woods; or spread them among the blos-
soms of water lilies and clumps of grasses, reeds, and iris that thrust their
stalks out of the shallow waters, as one may see to-day in the ponds of
New Jersey and the lagoons of the South.!
The promontories projecting into this lake bed on either side are
formed of trachite or other volcanic lavas; masses of the same occur at
many different points along the ancient shore. They seem to
Cause of be confined to the edges, for the most part, but some of the
Entomb- : : : ;
ieee mesas, or ancient islands, have trachite flows over them, and
their slopes covered with quantities of vesicular scorie. We
haye thus pointed out the principal cause of the fossil strata whose ex-
ploration has uncovered for us these pages in the life of the spiders of
the Tertiary. The shales of the lake in which the myriad of plants and
insects are entombed are wholly composed of volcanic ash and sand, which
he fifteen feet thick or more in alternating layers of coarser and finer
material. ?

‘ Lesquereux identifies these as among the plants found in the fossil yielding strata.
The genera are identical with the corresponding existing plants. U. 8. Geolog. Surv. Terr.,
Vol. VII., Tertiary Flora, 1878. Insects and spiders are usually found in the same shales
that yield the plants,

2 Paleontology of Florissant, 8. H. Scudder.

_ a —— 2 "=e" a
.

448 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

These insect bearing strata, as described by Dr. M. E. Wadsworth,' are
brownish and grayish brown shales, being simply the finer material of the
tufas laid down in lamine of varying thickness and coarseness. ‘This yol-
canic material has evidently been worked over by water; so far, however, |
as can be judged by microscopic examination, when the water commenced
its work the material was in loose, unconsolidated deposits, That it was
thrown out as an ash, or rather deposited as a moya or mud flow near its
present location, is the most probable supposition. The deposition ap-
pears to have been gentle but comparatively rapid, for there is no sign of
violence or even of such decomposition as one should expect in slow dep-
osition; and showers of ashes falling on still water or a lake, acting on
an unconsolidated tufa bank, answer best the conditions called for here.

Il.

According to Lesquereux the numerous leaves of some of the species
of plants are not crumpled, folded, or rolled as if driven by currents, but
flat as if they had been imbedded in the muddy surface of the

Manner }ottom when falling from trees or shrubs along the border of

pesca ~ the lake. As leaves, seeds, and other parts of a plant are al-
ment. ways intermingled with the fossil insects and araneads, we may

conclude that their entombment resulted from dropping along
with the leaves into the water. It will be remembered that many spiders
make their snares permanently among leayes, or within the inner sur-
faces of leaves, so that, when they are stripped from their stems by
violence or natural decay, they must often drift from the banks into
streams, and if overhanging the water drop directly therein. Others, like
our Insular and Shamrock spiders, dwell within nests of curled leaves,
and these would meet the same fate under like circumstances. It is not
now uncommon to see such nests overhanging the borders of streams or
woven among the foliage of plants in the immediate vicinity. Supposing,
as we have a right to do, the same habits prevailing in the Oligocene
period as the present, all these leaf dwelling species would haye been ex-
posed to submergence in the ancient Florissant Lake, and, being imbedded
in the mud, some of them, at least, might be preserved.

Lesquereux further believes that the deposition of the vegetable ma-
terials took place in the springtime and that the lake gradually dried
during summer, He bases this inference on the complete absence of hard
fruits, together with the presence of flowers, of unripe carpels of elm and
maple, and of well preserved branches of taxodium, which, in the living
species, are mostly detached and thrown upon the ground in wintertime
or early spring. If this were so, there would have been far fewer mature
spiders at that season, and the very young would be less likely to fossilize.

1 Scudder, Paleon. Floriss.

LL ee ee ee

ANCESTRAL SPIDERS AND THEIR HABITS. 449

Scudder thinks that the structure of the rocks indicates a quiet depo-
sition of the materials in an unruffled lake through long periods, inter-
rupted at intervals by the influx of new lava flows, or the bury-
ing of the bottom sediments beneath heavy showers of volcanic
ashes.' That many insects and spiders were beaten down by
these showers, destroyed, and buried, is at least probable. Certainly we
shall not go far astray in picturing such an exigency in the life history
of the disentombed fossil spiders in our possession. Thus the story of
Pompeii was enacted among the aranead inhabitants of this upland lake
shore in the distant Tertiary.

In this case, every season must have added contributions to the im-
bedded forms. After the final act of maternity female spiders soon die.
They may often be found, dried up, quite dead, hanging to grass or foli-
age, whence they drop off with the leaves. It was not different with the
fossils of Lake Florissant; they dropped to the ground and were carried
into the water, or dropped directly into the lake, and sank into the muddy
sediment, and were buried under the voleanic mud flow.

By a process somewhat similar the spiders of the Swiss Miocene ap-
pear to have been entombed, these soft animals being preserved only in

Volcanic
Showers.

aa the calcareous marl of the lower Oeningen quarry. Twenty-
eningen _. . ; aimee
Spi mc eight species have been uncovered, of which one, Epeira molas-

sica Heer, is an Orbweaver.? These fossils are, for the most
part, small, delicate creatures, belonging, with one exception, to genera
widely represented among living fauna. Eleven species are figured but not
described by Heer, whose figures are repeated by Heywood.

Of the insects which fell into the water of the ancient Lake Oeningen,
only those have been preserved which were quickly covered by the mud,
and thus saved from destruction. Aquatic insects are numerous, and are
found in all stages as larvee, pup, and imagines. Many were so rapidly
enveloped by the fine calcareous deposit that they have not merely pro-
duced an impression in it, but even the organic substance has been pre-
served. By this rapid covering the softest midgets are so admirably pre-
served that, under the microscope, the hairs of their legs and wings can be
recognized, and the color of the land bugs can still be ascertained. Thus,
in Europe as in America, we can picture the local conditions under which
the ancient spiders lived as not very different from some of our littoral
Atlantic lakes, as Deal Lake, for example, or those of Florida, and the
lagoons and bayous of the Southwest Mississippi. The general aspect of
the landscape; the forms and foliage of plants; the flowers and the insects
that visited them, like the spiders that made them their prey, must have
given a familiar face to the scenery.

' Paleon. Florissant, page 298.
* The Primeval World of Switzerland. By Professor Heer, Heywood’s English transla-
tion, Vol. II., page 10, 1876.

450 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

Il.

The climatic conditions under which the Florissant spiders lived and

died are established by a testimony which is quite accordant, both from

the fossil flora and fauna. According to Lesquereux, the plants

Climatic jndicate a climate like that of the northern shores of the Gulf

noaes of Mexico at the present epoch, and have a general aspect which

; recalls that of the vegetation of uplands or valleys of mountains.

The fishes, according to Professor Cope (quoted by Scudder), indicate a

climate like that at present found in latitude thirty-five degrees in the
United States.

Professor Scudder thinks that the insects from. their general ensemble
prove a somewhat warmer climate. He refers especially to the presence of
a great number of white ants imbedded in the shales, a testimony that is con-
firmed by a study of other insects, which are found to be largely tropical or
subtropical in their nature.! In a subsequent brief and interesting review
of thé Florissant spiders, and comparison with those from the European
beds,?2 Mr. Scudder repeats this opinion. He considers that the present
distribution of the allies of the fossil spiders points to a climate like that
of the middle zone of our Southern States, or the two shores of the
Mediterranean in Europe. However, of the genera which he cites in proof
of this, only one, Nephila, seems to me in point. This spider is undoubt-
edly tropical. I have a number of species from Africa, Zululand, Mada-
gascar, Liberia, etc., where they have an enormous development.

Our Gulf States have one species, Nephila plumipes, which is abundant
in many parts thereof, and is even more characteristic of the spider fauna of
Central America and’ the West Indies. As it has never yet been found
in the United States outside the southern belt of the Southern States, the
presence of a closely related species in the Florissant shales would
seem to be conclusive as to the nature of the climate during the
period at which their fossils were imbedded. The specimen pub-
lished by Professor Scudder, which I reproduce (Fig. 372), is well enough
preserved to prevent any doubt as to its generic identity.

According to Scudder® it is a much smaller species than Nephila plu-
mipes Koch, if the fossil be fully grown, and differs from it in some
striking points. The eyes differ considerably, although the position of only
two of those of the fossil species is known. ‘The corselet is squarer in the
fossil, and per contra the abdomen is oval and not quadrate, while the tarsi
are unusually long in proportion to the whole leg. The tufts of hairs occur
only on the extremity of the tibia. I have not seen the fossil, but judging

Fossil
Nephila.

' Paleontology of Florissant, page 299.

2 Fossil Spiders, Harvard University Bulletin, No. 21, page 303; see also Tertiary Insects,
page 51.

’ Tertiary Insects of North America, page 90, pl. xi, Fig. 12.

ee ee ee

ANCESTRAL SPIDERS AND THEIR HABITS. 451

from its general aspect, as displayed in the figure alone, I would suspect
it to be a young female Nephila plumipes. I have specimens of a species
collected by Mr. C. H. Townsend, at Swan Island, Caribbean Sea, which in
size and general appearance more closely resembles Scudder’s description of
Pennatipes than the modern Plumipes. The femoral brush is lacking
in these specimens, as it is in Scudder’s fossil (although there they may
have simply been worn away), and the shape of the abdomen is also cylin-
drical, as with Nephila pennatipes, instead of being quadrate as with our
species, We have thus a living Orbweaver which, as far as it is possible
to judge, differs little from this ancient aranead.

On the presence of this fossil species alone I would assimilate the climate
of the ancient Florissant Lake to
that of a region even
further south than that
assigned by Mr. Seudder. e ro

Scudder? describes a fossil
Tetragnatha, T. tertiaria, which
he thinks does not appear to
have any special affinity with
the American species with which
he has been able to compare it,
being stouter bodied than they.
Tlis conjecture, however, is hard-
ly a true one, that the presence
of this genus in the neighbor-
hood of the lake deposits of
Florissant indicates a warmer cli-
mate than the present. Tetrag-
natha, in several species, has a
range over the whole of the Fig. 372. The ype gti hee pennatipes.
United States, and I have fine
specimens from as far north as the borders of Alaska. They are extremely
numerous in such a climate as Philadelphia, for example, where we have
the European species Tetragnatha extensa; and along the margins of our
ponds and waters are seen immense numbers of large examples of the Stilt
spider of Hentz, Tetragnatha grallator, which is probably identical with
Tetragnatha elongata of Walckenaer.

A study of the spider fauna also justifies the inference that the climate
of the Tertiary period in Europe was essentially the same as that of
Florissant. ‘This is especially strengthened by a view of the recovered
insect forms of the two continents.? Of the insects in amber Mr. Hope

Pd
Climate.

1 Ter. Ins. N. A., page 77.
* Recherches sur les Insectes Fossiles des Terrains Tertiaires de la France. Par M.
Oustalet, pages 6 and 38. Bibliothéque de Ecole des Hautes Etudes, 1874.

7S | ies _ eS _ ee

452 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

states that they are extra-European, many belonging to tropical and tem-
perate climes. Dr. G. Mayr thinks the amber ants have few relations
with ants of tropical Africa and America.

IV.

It may be remarked, in this connection, that a comparison of the fossil
spiders of Europe with those of Florissant shows, on the whole, a general
correspondence between the two fauna. The same families are
Europe represented in the stratified deposits of Europe and America; and
aaa the correspondence holds good, to a considerable extent, as to the
amber species. Among Orbweavers this correspondence is not so
close, but obtains if we confine the comparison to families, and is true in
a measure of the genus Epeira and its near allies. Of the Oeningen spi-
ders one is gn Epeira. From the Brown-coal the Gea of Von Heyden!
is an Epeira also, according to Thorell.2 Of the Amber species,* Grea
Thor. (Gea Koch and Berendt), and Antopia (Menge) are near Epeira; Siga
(Menge) is near Zilla. All of
these belong with the family
Epeirine. Androgeus (Koch
and Ber.) alone probably be-
longs to another family, the
Uloborinee. Scudder divides
the Orbweaving species of Flor-
issant among four genera, Epe-
Fic. 378. Fic. 374. ira, Tethneus (new), Nephila,
_Fosall spiders from the amber. (After Berendt.*) baal Tetr agnatha, all Epeirine.
Fig. 372. Gea epeiroidea. Fic, 373, Androgeus militaris; male. __,, :
Thus all the Orbweavers in
both continents, with the exception of Androgeus (if Androgeus be, indeed,
an Orbweayer), belong to the same family Epeirinee, and most of them to
Epeira and closely related genera.

The above comparison also shows a close resemblance between existing
spider fauna and that of the Tertiary both of Europe and America, Tor
: example, the Orbweaving genera Epeira, Zilla, Tetragnatha, and
sete) Nephila are now common to both hemispheres, are all found in
Fauna, the United States, and the first three abundant. We should con-
sider, moreover, how closely related the remaining fossil genera
are to these and other existing ones. Tethneus, Gea, Groea, and Antopia
(Epeira), Siga (Zilla), and Androgeus (Uloborus) can, in this view, scarcely
be said with confidence to differ from existing Orbweaving genera. The

1 Paleontographica, Beitrage zur Naturgeschichte der Vorwelt, Band VIII. “ Fossile
Insekten aus der Rheinischen Braun-kéhle,” yon C. yon Heyden. Taf. I, Fig. 11, page 2.
Gea krantzi Heyd. Fundort: Rott, Sammlung Krantz, 2 Huropean Spiders, page 223.

® Tbid. 4 Op. cit. below, Tab. IIL, Figs. 12, 17.

OO SL =~ = — vr ~. * ¥ = ——

ANCESTRAL SPIDERS AND THEIR HABITS. 453

fact may be readily seen by comparing Berendt’s numerous figures of the
well preserved amber spiders with examples from corresponding genera,
(See Fig. 873, compared with Figs. 8376 and 877, and the full page cut
further on.)

The Florissant fossils are of course not so well preserved, but some of
the specimens retain their characteristics with sufficient distinctness to
compel the same conclusion. Scudder’s figures, as they are displayed upon
his plate, might well stand for good drawings of a miscellaneous collec-
tion of damaged specimens of our living spiders. Compare his figure of
the fossil Orbweaver Epeira meekii, for example (Fig. 375), with our familiar
Epeira strix (Fig. 376) or Epeira insularis (Fig. 377).

FG. 375. FiG. 376. FIG. 377.

Fie. 375. Fossil spider of Florissant, Epeira meekii. (After Scudder.) Fic. 376. Existing spider
Epeira strix; male. Fic, 377. Existing spider Epeira insularis; male.

Turning to the oldest known fossil aranead, Protolycosa anthrocophila
Romer, we are brought face to face with a species closely related to exist-
ing fauna. (Fig. 378.) Protolycosa belongs to the Carbonifer-

Stes ous, being found in the argillaceous slate of Kattowitz, upper
Spider’ Silesia.! Fig. 379 is an enlarged drawing, and Fig. 380 is an

Fossil, Outline restoration by the author. Rémer placed the fossil near
the genus Lycosa of the Citigrades, which rank among the highest
of the araneads. Thorell, on the ground of the extremely coarse and short,
strong legs and palps, assigns it to the Territelarie, which puts it within
a closely related group, in which we have found the largest existing spiders,
Theraphosoide, the Tarantulas, and such also as possess the highest me-
chanical instincts, as Trapdoor spiders. The first apparition of the spider
is therefore by no means that of a low example, but one rather which
presents a plenitude of faunal characteristics, and gives the possibility of
high industrial skill.
Moreover, Protolycosa is nearly related to a living species. Thorell
points out its marked resemblance to Schiodte’s wonderful East India genus

1 Neues Jahrbuch fiir Mihendlogia, Genlagts — Palae: sntativgtoy- Tah: 1866, pages 136-
143, Taf. III., Figs. 1-3.

fide ee

454 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

| FG. 380.

Fic. 378. Fic. 379.

The oldest fossil spider, Protolycosa anthrocophila. (After Rémer.)

Fic, 378. The spider in site. (Natural size.) Fig. 379. An enlarged drawing of the spider.
Fic. 380. A restored figure.

ANCESTRAL SPIDERS AND THEIR HABITS, 455

Liphistius,! and for this reason assigns it to his own family of Liphis-
tioidee. ” ,

Among the oldest fossil spiders, and probably the oldest in America, is
Arthrolycosa antiqua, from the Coal measures of Illinois, described by
Soeak Mr. Harger.* As the horizon in which this fossil was taken
Raa is synchronous with that of the argillaceous slate of upper
Spider, Silesia in which Protolycosa was discovered, this American spi-

der has the distinction of being one of the oldest as yet known
to science. Its true determination is therefore a matter of great impor-
tance. The aranead has generally been recognized as a type of a new
family, Arthrolycoside, as first established by Mr. Harger. Professor
Scudder placed the family at the beginning of the order Anthracomarti.
The supposed forcipulate character of the mandibles was considered suffi-
cient evidence to overcome the otherwise general resemblance to the Ter-
ritelariz, but this characteristic now ap-
pears to be without sufficient warrant.

Professor Scudder made a reéxamina-
tion of the type in 1884, and decided
against the forcipulate character of the
palps as described by Harger. More re-
cently Prof. Charles E. Beecher has made
a thorough study of the type specimen,
after cleaning it and exposing the append-
ages by removal of the superincumbent
matrix. After noting the differences which
the study of the specimens under these con- a
ditions developed, Professor Beecher con- Sa
cluded that on account of these important yy¢. 3g1. possil spider Arthrolycosa antiqua.
mein. differences it seems necessary to caicemaatens Meare a Gantt teneny
eddy exclude the genus from the order ;
antiqua. Anthracomarti.6 The marked resemblance between the ancient

Avicularidee and the fossil seemed to suggest that Arthrolycosa
is entitled to a place in the suborder Tetrapneumones, among the Terri-

FiG, 381.

1 “Om en afvigende Slaegt af Spindlernes rie J.C. Schiodte. Natur historisk Anden
Raeckes andet Bind Tidskrift, 1846-9, Bd. I1., Rak. 2, page 617, sq. Thorell’s reference is wrong,
a typographical error doubtless, making pages 6-7 for 617. As I count, he also errs in the
order of length of legs, which is 4, 8, 2, 1, instead of 4, 2, 8,1. There is, however, but a frae-
tional difference between third (172% lin.) and second (17%), and this does not change the
force of the inference. The species is Liphistius desultor; female. Habitat, Pinang Island.

2 Huropean Spiders, page 222.

8 American Journal of Science, 1874, Vol. VII., pages 219-223.

4 Proc. Amer. Acad. Arts and Sci., Vol. XX., 1884, page 15, “A Contribution to our
Knowledge of Paleozoic Arachnida.” :

5 Note on the Fossil Spider Arthrolycosa antiqua Harger, by Charles E. Beecher, Amer,
Journ. of Science, Vol. XXX VIII., 1889, page 219.

456 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

telarie. I insert a fac simile copy of the figure published by Professor
Beecher (Fig. 381), representing a dorsal view of the fossil, and (Fig. 382)
a bare outline when viewed directly in front. From the figure and profile
it is seen that all the limbs of the spider are in nearly their natural posi-
tion, having undergone but slight displacement and decay, while its per-
fection indicates that it is not a shed skin which is preserved, but that
the actual animal was entombed. It throws an interesting side light upon
the life habits of this creature, to learn that in the same coneretion which
contains the fossil are fragments of the broad leaves of a rush like plant
which, as Professor Beecher thinks, probably furnished a float by which
the spider was carried out from land, so that its remains are found min-
gled in the same bed with marine organisms.

In this connection I may call attention to another fossil spider which
has been supposed also to belong to the Territelaria. While visiting the
British Museum of Natural History at South
Kensington, London, in the summer of 1887,
my attention was called to some fossil spiders
by Dr. Henry Woodward, Keeper of the Geo-
logical Department. Among these I observed
one which seemed new to science, and closely
related to the genus Atypus. After my return
to America, Dr. Woodward sent me casts both
in wax and plaster, from which a description
of the species was made, and the name Ko-
atypus woodwardii suggested.! The fossil is
simply an impression in the shale, which, how-
ever, is tolerably well preserved, but exhibits
few features necessary to classification. The
eyes are not defined, and nothing but a little

ING. 384. roughened elevation in the centre of the caput,

FiG. 383. Fossil spider Eoatypus wood- : ‘
wardii McCook. Dorsal view. x3, Which may or may not be an organic cast,
sigs) algae woodwardii. Side gives any suggestion of the eye space. As far
as it goes, this appears to follow the charac-
teristics of Atypus and the Territelarie generally. ‘The appearance of the
mandibles also suggests this relation, and the general facies of the fossil is
to the same effect. The drawings have been made from a plaster cast,
Fig. 383 representing the dorsal view, and Fig. 384 the same in outline,
both magnified three times natural size.2

"Proc. Acad. Nat. Sci., Phila., 1888, page 200, for full description of the species.

*I hesitated much as to whether this fossil should be assigned to the Lycosidee, the
Attidee, or to Atypinse. On the whole, I decided, though not positively, as above, and on
the above named grounds. It seemed impossible, in the absence of the characteristic eyes
and long jointed superior spinners to relegate the species positively to the genus Atypus.
Besides expressing the general facies of the fossil as above described, the generic yalue of
the name Koatypus consists largely in assigning the specimen rank as a fossil spider.

ANCESTRAL SPIDERS AND THEIR HABITS. 457

The horizon from which this fossil was obtained is the Eocene Ter-
tiary, Garnet Bay, Isle of Wight. It is, therefore, probably somewhat older
than most’ Kuropean and American aranead fossils.

According to Scudder, more than one-half the genera of known fossil
spiders to which species have been referred have been described as new
and peculiar to Tertiary times. ‘These genera include about two-fifths of
the species. Among the genera are some remarkable forms, such as Archea
and Mizalia, each of which is considered by Thorell and others as repre-
senting distinct families! Further on I reproduce Berendt’s drawings of
Archea paradoxa, to illustrate these peculiar forms.

Fic. 385. Fic. 386.

Fics. 385 and 386. Views of Palpipes priscus, a fossil crustacean larva. (After Von Meyer.)

Two genera only of the thirteen to which the American species are
referred are described as new, and ‘to them are referred seven of the thirty-
two species. Other genera not before recognized in a fossil state, but here
recorded from American strata, are Titanceca, Tetragnatha, and Nephila.
To enter into details, seventy-one genera of spiders have been described
from the Tertiaries, sixty-six from Europe, and thirteen by Scudder from
America, eight genera being common to both. Of these seventy-one gen-
era, thirty-seven are counted extinct, thirty-five from Europe, and two
from America, none of these extinct species being found in both countries.
The European genera are, as may be supposed, largely composed of amber
species, no less than fifty-two, including thirty-two distinct genera, being
confined to amber deposits, besides others which they possess in common
with the stratified beds.?

Palpipes priscus® has been so long regarded as a Jurassic spider that I
have alluded to it in this chapter, but that it is not a true spider, but

1 Thorell, European Spiders, pages 223-233.
* Scudder, Tertiary Insects of N. A., page 51. I do not here include Eoatypus.
®’ Von Meyer, Palseontographica, Bd. X., pages 299-304, Taf. L., Figs. 14, Cassel, 1863.

‘a. - Ee

458 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

a crustacean larva appears to me to be yery clear from an examination
of the figures which I reproduce, Figs. 385 and 386, and, indeed, this
has already been shown by Seebach.!

It remains to notice a little more definitely the geological position of
the fossil spiders of America. Professor Cope, in view of the character
of the fish fauna, relegates the Florissant deposits to the later
Geological Rocene or early Miocene.? Lesquereux, judging from the plants,
ile oy of refers this deposit to the lower Miocene or Oligocene.* ‘This
orissant ; : Pie
Spiders, Would place the spiders and the insects of these beds within
the same horizon, substantially, as those of the amber and the
Oeningen and other Tertiary strata of Europe. Or, as Scudder has ex-
pressed it, “We may therefore provisionally conclude, from the evidence
afforded by the plants and vertebrates, that the Florissant beds belong in
or near the Oligocene.” The evidence derived from insects and spiders is
thus in harmony with that from vegetables and higher animals.
I have attempted, by the following tabulated statement, to express ap-
proximately the relations of the Florissant spider bearing deposits with
those of Europe in which spiders have also been found. ;

TERTIARY.

PLIOCENE.
Upper. 1. Fresh water formations, Oeningen, Switzerland,

Miocene. | Middle. 2. Sulphur impregnated strata, Radoboj, Croatia.
| Lower. 3. Brown-coal strata of the Siebengebirge, Rhine.

( 4. Florissant Basin, Florissant, Colorado, U. 8.
OLIGOCENE. 5. Amber, Prussian Baltic.
| 6. Fresh water formations,t Aix, Proyence.°
Eocene. 7. Garnet Bay, Isle of Wight (Eoatypus woodwardii).
CRETACEOUS.
JURASSIC. 8. Lithographic limestone, Solenhofen, Bayaria (Palpipes priscus).°
TRIASSIC.
PERMIAN,
. 9. Argillaceous slate, Kattowitz, Upper Silesia (Protolycosa anthro-
CARBONIFEROUS. | cophila).

10. Coal measures of Illinois (Arthrolycosa antiqua).

1 Zeitschr. deutsch geol. Gesellsch, X XIII., page 340.
2 Bull. U. S. Geological Survey Territories, 2d series, No. 1, 1875.

3 Report U. S. Geological Survey Territories, Vol. 7, 1878. American Journal Science,
XVII. page 279.

* Oustalet, Recherches sur les Insectes Fossiles des Terrains Tertiaires de la France,
page 36. Oustalet presents the various views of geologists as to the position of this forma-
tion, from which I have placed it as here. . :

5 A well preserved Theridioid spider from Aix may be seen in “Geology and Mineral-
ogy,” Bridgewater Treatise, by Rey. Wm. Buckland, D.D., Vol. II., page 79, and plate 46,
Fig. 12, Theridium bucklandii Thorell. Gourret has recently described about eighteen Oli-
gocene species from Aix. Rec. Zool. Suisse, 1V., page 431, 1887.

5 A crustacean larva, see above, page 457.

wet OE a hhh ee ee ——- Tl diets 7 ——————

ANCESTRAL SPIDERS AND THEIR HABITS. 459

The fragile nature of the spider’s spinningwork has passed into a proy-
erb expressive of utter weakness and ephemeral age. Yet Mr. Scudder has
uncovered for us a fossil cocoon, about one-fifth of an inch long,

ae that dates from the distant period of the Oligocene, and which
pee ee he describes under the name of Aranea columbie.! This co-

coon has been found at widely separated points—Florissant, Green
River, Wyoming, and British Columbia—and thus appears to have had
some fayored environment or especial qualities inducing preservation. - One
might suppose that the large cocoons of Orbweavyers, especially those with
tough encasements, like Argiope and Cyrtarachne, or the large flossy silken
ball of Nephila, might easily have been fossilized under circumstances that
allowed the preservation of the araneads themselves. None of these, how-
ever, have yet been discovered, and the little Aranea columbiz cocoons
are the sole representatives of the spinningwork of the aranead weavers
of the Tertiary. Eleven of these in all have been found, and the survival
of this minute bit of cocooning spinningwork is so interesting and im-
portant that I give a full abstract of Scudder’s description thereof.?
Among the stones obtained by Dr. George M. Dawson in British Co-
lumbia are several containing the flattened remains of the egg cocoons of
spiders. There are no _ less
than eight of them, occurring
by pairs, none of them re-
verses of others. They vary slightly in
size, and more in shape, owing, no
doubt, to their varying position when
crushed; probably they were globular,
or possibly slightly oval in shape; ay-
eraging about five millimetres in the

Fossil
Cocoons.

longer and four millimetres in the short- FXG, 387. Fr, 388.
er diameter ; of a firm structure : testa- The fossil spider cocoon, Aranea columbiz.
- Fic. 887. With the pedicle by which it was
ceous in color, and hung by a slender anayentied,, Sia, eas. Muah elonpated by
thread, less or much less than quarter presiure. “Both ‘fgares are enlarged be-

tween five and six times. (After Scudder.)

the length of the egg cocoon (averag-
ing, perhaps, one millimetre in length), to a thickened mass of web, at-
tached to some object or to the mother’s web.

That they have been preserved by pairs upon the stones has no signifi-
cance, and, indeed, may be due simply to the way the stones were broken,
for they lie at varying distances apart, with no sign of connection, and
placed with no definite relations to each other.’ Two of them show no

' First described in the Report of the Geological Survey of Canada for 1876-77, pages
463, 464. 2 See Tertiary Insects of N. A.
% Many spiders make two or more cocoons, which sufticiently accounts for the above fact.

460 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

sign of a pedicle, but this may be due to poor preservation; and a single
one not only has no pedicle, but appears to be formed of a lighter, flim-
sier tissue, and may belong to a different species.

The egg cocoon of a spider of exactly the same size, shape, and gen-
eral appearance as those described above, excepting that from a break in
the stone there is no trace of a pedicle, was found by Seudder in the shales
at Green River, Wyoming. A single specimen was also found at Floris-
sant, Colorado, having the same general appearance, but with no trace of
a pedicle and slightly larger than any of the others, being six millimetres
long*’and four millimetres broad. It is, of course, impossible to say that
it is the same species. Still another was brought by the Princeton ex-
pedition from Florissant, different in the opposite direction, being con-
siderably smaller and so preserved as to appear broader than long. It is
provided with a pedicle one and four-tenths millimetres long, but is itself
only two millimetres long and two and a half broad.

If the reader will turn back to pages 114 and 115, in the chapter on
General Cocooning Habits, he will see examples of cocoons which correspond,

both in size and general character, to these fossil cocoons of the
oa Tertiary. Cocoons of Ero thoracica, for example (Figs. 111 and
ypes. 2 : ‘ 5
116), are represented in my drawings about twice natural size;
that is, they are about one-eighth inch long, or a little over three milli-
metres. They are suspended by a thread, from various objects, in a man-
ner which is suggested by the character of Aranea columbie.

Another cocoon represented among these drawings (Figs. 112, 113) I
there attribute to Theridium frondeum on the authority of Dr. Marx. A
number of observations made since those pages were printed, both by my-
self and my secretary, have led me seriously to doubt the identification,
and to believe that this little orange colored hanging cocoon, which has
so long puzzled me to identify, is probably the cocoon of Theridiosoma
radiosum. We have found it a number of times hanging close by the
snares of females of that species in Belmont Glen and other ravines of
Fairmount Park, and in the country surrounding Philadelphia; and no
other species was found in the neighborhood to which such a cocoon
could be attributed. I am therefore inclined at the present date to be-
lieve that the Ray spider is responsible for this pretty little egg sac. In
addition to this, I have examined young specimens raised from the cocoon,
and although the determination of a species by just hatched spiderlings is

_.. Well known to be extremely uncertain, yet this examination has
ia confirmed me in the above opinion, The shape of cephalo-
Cocoon. *horax and abdomen, arrangement of eyes, proportion of legs,

and general ensemble of the younglings lead me to conclude

that, if they are not Theridiosoma, they belong to no species with which
I am acquainted.

In further confirmation I may add that Dr. L. Koch says of the cocoon

_—

re ee SS ee oer rth OO eee — ——

ANCESTRAL SPIDERS AND THEIR HABITS. 461

of Theridiosoma gemmosum that it is pyriform, pediculated, of yellow
brown color, with pedicle white, and that the female makes her cocoon at
the end of June.! This description well agrees with the cocoon under
question. Simon himself says that Theridiosoma gemmosum is found
along the borders of waters, making its snares upon aquatic plants. Its
cocoon is in the form of a balloon, with a pedicle like that of Ero.? As
Theridiosoma gemmosum and 'T. radiosum are probably identical, or at least
closely related, this evidence appears to be almost conclusive.

I have measured many of these Theridiosoma cocoons, and they aver-
age in length about one-eighth inch, or, more accurately, three and one-
half millimetres. Their width is a little less. In other words, the cocoon
is almost spherical, but the addition of the pedicle or stalk makes it seem
longer. I have seen some cocoons which were five millimetres long. ‘The-
ridiosoma’s cocoons are closely woven and of tough fibre, well fitted for
preservation. If now we compare the above named structures with Scud-
der’s fossil cocoons, we shall find a close resemblance. We may therefore
have little hesitation in relegating Aranea columbie to some such The-
ridioid genus as Ero or Theridium, or perhaps to the ancestors of Therid-
iosoma, The Ray spider has evident relationship to Theridium, as appears
from the fact that such accomplished araneologists as Cambridge, Simon,
and the late Count Keyserling have classed it with the Retitelaria. One
might therefore venture to attribute to it an ancient lineage, and even to
risk the conjecture that a species of Theridiosoma may have been the au-
thor of some of Scudder’s fossil cocoons.

The preservation of any spinningwork through so vast a period is
greatly interesting; but I find the chief value of the fact in the inference

_ that the general habits of spiders have followed even more
Unmodi- ¢losely the law of unmodified survival that appears to mark
fied In-
duatiy. the general structure of araneads. Indeed, I am not able here

to note any difference. Precisely the same industry that we see
everywhere exemplified in the pretty hanging basket cocoonery of our
modern Ero, Theridium, or Theridiosoma, characterized the fossil Aranea
columbiz that wrought her spinningwork along the shores of Lake Flor-
issant in the early period of the Tertiary. It is certainly not an unwar-
ranted inference that the spinning organs by which these cocoons were
produced differed in no essential particular from those possessed by mod-
ern spiders. This likeness implies structural similarity in other vital
organs, and hence, reasoning from industrial product to function, from
function to organ, from special organs to general structure, we arrive at
the same conclusion that seems justified by a study of Scudder’s Ameri-
can fossils, that many spiders of the Tertiary were not widely different

1 Simon, Arach. de France, Vol. V., page 27. 2 Tbid., page 25,
® See Vol. 1, Chapter II.

i ~~’? = |= Oe a

462 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

generically, and some probably even specifically, from the spiders which
now inhabit our continent.

VII.

Since most fossil spiders known to us are preserved enclosed in amber,
it is important in our study of the life of ancestral araneads to know

something of the history and character of this important sub-
~ stance. Amber is a product of the prehistoric world, a hard-
‘ened resin which issued from the bark of certain trees. The
chief geographical source of the amber wood is in the bottom of the
Baltic Sea in the neighborhood of what is now called Samland, near Pil-
lau. The amber tree is known as Pinites succinifer Gépp. and Ber., and
has been described from various vegetable inclusions—wood, blossom, fruit,
and needle leaves—along with various insects and araneads. ‘The species
Succinifer rightly belongs to the genus Pinus, although that name is really
a collective name, inasmuch as included needle leaves and other vegeta-
ble formations show there must have been at least four species of pine
in the amber fields. Since it cannot be determined which one of these
actually secreted the resin, the specific name must be a comprehensive
one. The trees which produce the amber are not now known to exist,
but Berendt says that the Balsamea most closely resembles it.!

Every gale from the north still throws up, as for unknown ages it has
done, masses of amber on the shore of the Baltic Sea, and each point of
the coast is said to receive a particular kind so peculiar that practiced
cutters are able, when looking at a rough piece, to decide whether it came
from a quarter to the east of Danzig or from the west on the coast of
Pomerania; they are therefore probably the product of different trees.

The sources of amber are submarine forests which, in the middle epoch
of the Brown-coal, as Berendt conceives, covered the shores of an island

continent that occupied the northern portion of the great Ter-
Sources tiary sea that covered most of Germany. This island, or group
of Amber:
Samland, © islands, had its geographical centre in the southeastern part

of the present sea, basin, under the fifty-fifth degree of latitude,
and its northwestern border extending higher than the present north-
western point of Samland.

The name Samland will not be found upon many maps, and it may,
therefore, be defined as distinguishing that part of Prussia bounded on the
west by the Baltic Sea; on the north in part by the same sea, the Ku-
rische Nehrung, and Kurische Haff. The southern boundary is the river
Pregel and the Frische Haff; while the eastern ee | is an arm of

' Berendt, G. K., Die im Bernstein befindlichen Oneiieowen Reste der Vorwelt gesam-
melt in Verbindung. mit Mehreren bearbeitet und herausgegeben, von C. L. Koch und Dr.
Georg Karl Berendt. Band I., Abth. II., page 28, Berlin, 1854 (1845).

ee Se lee ee ——_ | a ——

ANCESTRAL SPIDERS AND THEIR HABITS. 463

the Pregel, the Deima. It is hilly towards the northwest, the ground
rising to heights of two and three hundred feet, and becoming flat towards
the northeast and east, and gradually sinking down towards the north-
eastern angle. In the elevated northwestern coast Tertiary beds are con-
spicuous at a height from eighty to one hundred and twenty-five feet
above the sea level, in which amber deposits are found.

Zaddach! defines the site of the amber forests as a bay whose bed in-
cluded the whole of West Prussia, a neighboring portion of Pomerania, and
the western half of East Prussia, and which was connected in the
southwest with the great Tertiary sea that covered the larger por-
tion of Germany. ‘The northern boundary of this bay left Sam-

land at some distance, and was continued westward with some irregularity to
Ruckshéft (Rixhéft), which lies at the foot of the peninsula of Hela, and
where thick Brown-coal beds crop out on the coast of the Baltic. The
bay was a basin in the Cretaceous formation, and was bordered by widely
extended flat coasts, which mark the last upheaval of the district. Number-
less rivulets with small discharge emptied themselves into the bay and
carried solid matter into it, and another stream from the northwest, which
flowed from the southern portion of the Cretaceous land, also discharged
itself here.
The coasts of this bay were covered with luxuriant plant growths, a
flora whose delicate structure is still preserved to us in the amber and
‘coal, The forests which covered the shores of this bay and oc-
mt cupied the group of islands or insular continent beyond, were,
unning : , :
Amber, according to Zaddach, the native home of the amber. This
amber resin issued from the trees as pitch issues from pine trees,
and gum from our cherry and plum trees. In the Adirondack forests I
have seen guides and visitors collecting vials full of the aromatic resin
which issues from the fragrant balsam tree. Certain resins and gums of
commerce, as copal, anime, benzoe resin, mastix, and balsam, are collected
by making slits in the bark of trees so that the resin runs down in chan-
nels to the ground, where it hardens and is collected for transportation.
Copal perhaps affords the best analogy between modern resins and the
ancient amber, because it comes nearest it, and, indeed, according to
Berendt, may be considered its modern representative. One species of
copal belongs to the prehistoric world, but Berendt thinks that it did not
grow in the same native home with the amber tree, because the organic
inclusions of the two resins show no identity.
The great amount of amber already collected gives but slight indica-
tion of the incalculable quantity that must have been secreted by the
amber pines of the Tertiary. ‘The sunken storehouse thereof, the former

Amber
Bay.

1 Amber: Its Origin and istry. as jiigeumted = ihe Gace of Gamlaxic, ‘by Dn G,
Zaddach, Professor in the University of Kénigsberg. Quarterly Journal of Sciences, London,
1868, page 167.

464 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

soil of the forests, seems to be full of it. Although storms and floods
during thousands and thousands of years have been tearing up and wash-
ing away these stores, the quantity seems to have been lessened

bake to only a trifling degree. All the Baltic shores which lie clos-
Stores. st to this supposed sunken continent, also the west shore of

Samland and the north shore of the Frische Nehrung, have
always received and still receive a large quantity of amber. However,
the storms from the west and west northwest bring up the amber most
abundantly.

The temperature was then much higher than now, and the flora of
the amber continent contained certain northern forms associated with
plants in temperate climate, and others whose nearest allies now live in
much more southern regions. Thus camphor trees (Cinnamomum poly-
morphum Heer) occur with willows, beeches, and numerous oaks. Among
the conifers, the most abundant tree was a Thuja, very similar to the Thuja
occidentalis now living in America, next to which abounded Widdringtonia,
pines and firs in great variety, and among them the amber pine. Many of
the last already had perished, and, while the wood decayed, the resin with
which the stem and branches were stored might have accumulated in
large quantities in bogs and lakes in the soil of the forest.

In order to explain, however, that this accumulation of amber could
be suddenly broken up, floated away, and scattered, Zaddach assumes that

the coast of the district was on the point of sinking. Alternate
Breaking ypheavals and depressions of the country may be positively
ea proved to have occurred in the immediately succeeding period.
house, /f at that time the coast sank but slowly, in the lapse of a few

centuries, or even a shorter time, a great portion of the flat
coast terraces might have been covered by the sea. The forest earth was
washed up by the waves, and the amber carried into the sea. The greater
portion being probably still attached to the wood, with all its animal en-
closures, it could float about in the water for some time before sinking,
The forest of the inundated coast was also destroyed, but the stems of
the trees which floated out into the open sea were scattered about, only
those pieces of wood imbedded in the amber charged earth sinking with
it to the bottom. Thus perished the amber forests; in great part, at least,
for one need not assume that they were then all destroyed, as it is prob-
able that in the higher districts of the country there still remained many
forests which also were rich in amber trees.!

At last, after alternate upheavals and depressions, the land gradually
rose to its present height. And now, when lashed by storms, the sea tears
up the amber out of the deep lying beds of amber earth. By the help
of sea weeds turned up at the same time, the resin is heaved upwards and

1 Zaddach, op. cit.

ails it er a

a xg2E————< - eae ee . a e. ~~

carried on the surface of the water; and when the storms abate and the
sea becomes calm it carries the amber, together with pieces of older Brown-

coal and fresh marine plants, on to the beach, where a hundred
Collect- hands are waiting to intercept it with nets. hat is the “amber
Metice, drawing,” a trying occupation, which demands a strong and

hearty frame, for the cold winter storms yield the richest booty.
But many pieces of amber, nevertheless, do not reach the shore, for the
largest and heaviest pieces have already sunk to the bottom, and lie be- ’
tween the large boulders which cover the sea bed. ‘Therefore, in calm
weather, the inhabitants of the coast take boats and turn the stones with
hooks fastened to long poles, dislodge the amber in the interspaces, and
draw it up with small nets. This is called “striking for amber” (Bern-
stein strechen).

Amber is occasionally met with in the gravel beds near London. At
Alborough, on the coast of Suffolk, after a wrecking tide, it is thrown
on the beach in considerable quantities along with masses of jet, and if
not torn from the bed of the sea may have been washed from the Bal-
tic. There are regular mines of amber in Spain, and it is also abundant
on the shores of Sicily and the Adriatic Sea.

According to Mr. Hope, who speaks as an entomologist, many of the
insects recognized in amber indicate a tropical climate, and evince a South

American relationship ; yet the Blattidee and some of the Hymen-
Climate optera resemble closely oriental species. The presence of many
of Amber panes : :
Teme: other genera indicates a northern climate. From the above dis-

crepancies, it may be adduced that the climate and temperature
of Europe have undergone considerable change. The examples of tropical
insects sufficiently testify that the amber tree did not flourish in a climate
such as Prussia now enjoys, but in a warmer region.!

VL:

One who reads a list of Succinie Insects, as, for example, that pub-
lished by Mr. Hope,? will find represented the orders of insects with which
we are now familiar, These must have formed the food sup-

Insect plies of the amber spiders, A large proportion of our com-
og mon families are therein represented, and underneath these fam-
Spiders, ilies numerous genera of prevalent insects appear. It would thus
seem that the generic aspects of the insect fauna of the amber

period resembled that of the present time; indeed, Mr. Hope has said, per-
haps somewhat too strongly, “the major part exhibit a close resemblance

1 Rey. F. W. Hope, F. R.S., President Entomological Society. “Observations on Succinic —
Insects.” Transactions Entomological Society of London, Vol. I., 1836, page 133, sq.
* Tbid., pages 189, 147. ;

a ee ns a es

466 AMBPRICAN SPIDERS AND THEIR SPINNINGWORK.

to existing species, and can be satisfactorily placed under the published
genera.” However, we have not been able to trace specific identity. In
this antediluvian and amber forest now lying beneath the North Sea waves,
and along the shores of this Tertiary Amber Bay, we can readily picture to
ourselves vast numbers of Coleoptera burrowing in the ground, boring in
trees, flying among the branches, pursuing the same round of habits with
which we are to-day familiar. The Homoptera are represented by the
* Cicada, who doubtless then as now filled the forest with his piping notes.
Dragon-flies hunted their insect prey, and Libellula and Agrion carried
havoc among the entomological hosts, as they do to-day in the neighbor-
hood of Philadelphia. Ichneumon flies doubtless exercised their parasitic
habits upon victims like their modern hosts. Wasps of various sorts
dragged numberless spiders, flies, and other insects into their mud daub
nests to feed their voracious grubs.

Ants and bees were present in great numbers. Among the Orthoptera,
cockroaches, locusts, grasshoppers, and many other genera were represented.
Among the Lepidoptera such well known genera as Papilio, Tinea, and
Sphinx might have been seen; and minute Diptera, some of which, at
least, were similar to those of modern Europe, everywhere abounded in field
and forest. We may, therefore, conclude that the picture of this submarine
antediluvian amber forest, which we can draw from the facts presented
to us by the entomologist, botanist, and geologist, would not largely differ
from that of the midsummer aspect of the forests of the Adirondack Mount-
ains in New York, where various sorts of pine trees reach immense pro-
portions, and the balsam especially abounds, forming the fragrant upholstery
for the beds of those who bivouac or camp along the lakes and rivers of
that favorite region of American summer tourists. In the midst some
such scenes, and surrounded by similar insect hordes, the aranead ancestors
of our existing spiders dwelt. The reader may know just how they looked.
They are embalmed for. us in the liquid resin secreted in the forests of
Amber Island and Amber Bay.

On the accompanying full page engraving I have presented a few selec-
tions from the figures of amber spiders, as given in Berendt’s noble work.
Figs. 389 and 390 represent Orbweavers of the genus Zilla. The
Lineweavers are represented by Figs. 395 and 398, Ero and 'The-
ridium. The Tubeweayvers by Figs. 393 and 397, Segestria and
Clubiona. The Saltigrades, by the unmistakable Eresus at Fig. 394, and
the Laterigrades by Philodromus and Syphax, Figs. 396 and 399. It is at
once manifest by a glance at these drawings that in their general facies not
only, but in their detailed characteristics, they show a close resemblance to
corresponding genera as they are known to-day.

This resemblance, however, to existing genera (as far as now known) is
not always so apparent from the figures presented by Berendt, For ex-
ample, Archea paradoxa, which is represented much enlarged in both the

Amber
Spiders.

ANCESTRAL SPIDERS AND THEIR HABITS.

Fossil spiders of the amber. (After Berendt.)

Fig, 389. Zilla porrecta, female. Fic. 890. Zilla gracilis, female. Fic. 891. Phidippus fre-
natus, female. Fic. 392. Phidippus frenatus, male. Fic. 393. Segestria nana, female.
Fig. 394. Eresus monachus, female. FiG. 395. Ero setulosa, female. Fic. 396. Philo-
dromus microcephalus, male. FiG, 397. Clubiona attenuata, female. Fie. 398. The-
ridium hirtum, female. Fic. 399. Syphax megacephalus, female.

——— Eee

— ee a ee ee aT eee ne

468 AMERICAN SPIDERS AND THEIR SPINNINGWORK.

female form (Fig. 400) and the male form (Fig. 401), shows a wide diverg-
ence from any spider with which I am acquainted. Modern spiders cer-
tainly present some forms that are equally remarkable in their divergence
from the typical spider facies. But this genus appears to stand by itself,
without any modern representative, and is probably extinct.

As the climate of the amber forests covering the shores of the Tertiary
Amber Bay, and the islands grouped within it, was that of a semitropical

rather than of a temperate zone, we may conceive these endless
Embalm- woods of amber pine exuding streams of resin under a hot sum-
Faee mer sun, The liquid product freely flowed down the trunks of
Insects. the trees and accumulated in great lumps around the roots, mass

mingling with mass, as the trees stood close together in the for-
ests, until in the course of time the soil was surcharged with solidified
resin. In that period, as
now, insects frequented
trees, and were continu-
ally hovering around the
trunks, alighting thereon,
creeping along the bark.
Then, too, any aromatic
substance dropping from
the branches upon the
ground must have at-
tracted swarms of them,
as I have often seen in —
American forests.

We have thus the
conditions under which
the amber fossils were
entombed ; for a™single
touch of an insect upon
the liquid resin would at
once arrest its flight, and the soft, flowing stream would instantly imbed
it. For the most part this enclosure seems to have been painless; at least,
the attitude of the included insects and spiders is such as to suggest the
absence of all violent struggle. At any rate, their limbs soon sank into a
position of repose, and they are thus preserved to us.

Where insects are, there spiders resort in search of their natural food.
Lurking upon the branches, crouching, walking, jumping upon the trunks,

: spinning their webs in the grasses at the foot of trees, and
as ig stringing them from bough to bough, it is not strange that, in
balmed: the ordinary course of life, they too found sepulture within the

liquid runlets and masses of resin, and thus have been preserved
to us, along with the insects whose lives they sought, imbedded in amber.

Fic, 400. The fossil spider Archea paradoxa; female. (After
Berendt.) Natural size shown in the circle.

eee ee EE ——

ANCESTRAL SPIDERS AND THEIR HABITS. 469

One might well be excused for giving his imagination some play in
depicting the strange mutations of these creatures of the amber forests.
But the simple truth seems fanciful enough. Their life in those fragrant
woods of the Tertiary, along the islands and shores of the ancient Tertiary
sea; their swift entombment within the aromatic balsam; their long repose
within the soil of the ancient forests; the convulsions by which they were
sunk deep within the sea, and their recovery again to the surface; their final
repose in the deep bed of the Baltic Sea, after the recurring depressions
and elevations had ceased; in some cases, at teast, their settlement and
subsidence, after drifting here and there,
attached to broken and decayed trunks and
roots, the sport of waves and currents of
the ocean; their long, long sleep under-
neath deep sea waves, while the marvelous
changes that have made our present world
were being wrought out; their rupture from
their rest of milleniads by the grinding
force of winter storms; their drifting be-
fore the force of breaking waves upon the
shores of Samland; their capture by the
fishermen and amber strikers of Germany ; Fic. 401. Arachea paradoxa; male.

5 ‘i A ie, ‘ (After Berendt.)
their cutting, shaping, and polishing in
the hands of lapidaries; their transit from hand to hand among venders
and merchants; their resting place in cabinets of entomologists, collectors,
and scientific societies; their voyage from country to country,
and once more upon the sea; their lodgment here beneath the
curious eye and lens of the writer, who studies them and depicts
their forms for science as they rest embalmed in their amber sarcophagus—
- all this is certainly a picture upon which fancy might fondly dwell. It
reads like a romancer’s tale; yet the story, nevertheless, presents no merely
fanciful features, but, in good sooth, is all within the realm of sober facts
which naturalists have disclosed.

Resur-
gam.

THE END.

INDEX OF VOLUME II.

Abbot, John, 138, 386.

Abdomen, curious use of, 417.

Acoloides saitidis, 397.

Acrosoma, 339.

Acrosoma rugosa, 285, 289, 375.

Activity of female, 70.

Adaptation, 412.

Adirondack Mountains, 463, 466.

Aeronautic habit, 15, Chapter IX., 256, 399 ;
flight, 179.

African spiders, 208, 399, 450.

Agalena brunnea, 124, 292.

Agalena labyrinthea, 29, 45, 122, 189.

~Agalena neevia, 33, 36, 44, 85, 165, 166, 187, 189,
208, 211, 236, 251, 252, 253, 288, 334, 337, 347,
389; cocoons of, 121; upholstering of co-
coon, 122.

Agamic reproduction, 73.

Age of spiders, 3841; influencing color, 331,
428; of ants, 427.

Agroeca brunnea, 126, 132.

Agreeca proxima, 126,

Alaska, spider fauna, 97.

“Albatross,” Fish Commission steamer, 334.

Alexandria Bay, New York, 131.

Amber bay, 466; collecting, 465; embalming,
468 ; island, 466; stores of, 464; tree, 462;
sources of, 462.

Andover Review, 280.

Androgeus, 452.

Animals, senses of, 345.

Annisquam, 836.

Anthrobia mammouthia, 154, 156, 189, 286,
291, 335.

Anthrocophila antiqua, 455.

Ant formed spiders, 357, 369.

Ant thrush, 363.

Ants, 427; cutting ants, 353; destroyed by
birds, 362; driver ants, 863; eat spiders,
364.

Arachnophagous wasps, 388.

Aranea columbiee, 459, 461.

Archea paradoxa, 457, 466, 468, 469.

Architecture, 355; influenced by motherhood,
185; inspired by motherhood, 64,

Arctosa cinerea, 393.

Argiope argenteola, 83, 84; construction of
cocoon, 84, 388.

Argiope argyraspis, 21, 71, 289, 334, 338, 347,
349, 386; cocoon of, manner of suspen-
sion, 82; hung among wild flowers, 83.

Argiope aurelia, life of young, 228, 229.

Argiope cophinaria, 20, 21, 22, 26, 71, 188, 189,
201, 203, 207, 209, 210, 288, 300, 329, 334, 338,
346, 348, 350, 395, 398, 426; cocoon, con-
struction of, 159, Chapter IT.; sites of, 75;
methods of supension, 76; among grasses
and wild flowers, 77; stability of poise,
78; hung to a curtain, 79; internal struct-
ure of, 80; variation in structure, 81; ovi-
positing, 160; spinning the brown pad-
ding of her cocoon, 160; weaving the co-
coon case, 161, 162; winding the thread,
163; mechanical ingenuity in weaving,
163; weaving, Argiope’s method of, 163;
decline and death of, 419; male, courtship
of, 18; maternity, 19; pairing of, 37; sey-
eral cocoons, 108, 110.

Argiope fenestrinus, 84.

Argiope multiconcha, 108, 188.

Argyrodes argyrodes, see Argyrodes trigonum,

Argyrodes piraticum, 388, 390.

Argyrodes trigonum, 113, 114, 376, 389; see
Argyrodes argyrodes, 389.

Argyroepeira hortorum, 39, 299.

Argyroneta aquatica, 24, 29, 65, 125, 126, 188,
239, 393; cell and eggs of, 125; male and
female, 23; pairing, 45.

Ariadne bicolor, 134.

Arthropoda, 314.

Astia vittata, 262, 295; dance, 53, 54.

Atkinson, George T., 416.

Atrophy of eyes, 292.

Atta fervens, 357.

Attidee, 69, 333, 359, 864; colors of, 827; pair-
ing of, 50; attitudes in courtship, 59,

(470)

ss Se i —

INDEX.

Attus nubilis, 196.

Attus terebratus, 314.

Atypus, 169.

Atypus abbotii, 138; see Purseweb spider.
Atypus piceus, 29, 137, 138, 245, 246, 429.
Audebert, 73.

Auditory organs, 300; hairs, 309, 312.
Ausserer, Prof. Ant., 73.

Azara, Don Felix de, 230.

Babyhood of spiders, 206.

Beeus americanus, 397.

Baird, James, M. D., 444.

Ballooning, 109; among Orbweavers, 266; at-
titude during flight, 260, 261; circumnayi-
gation by, 268, 269, 270; habit, 256; height
attained, 264; modes of, 268; process of,
264; -to distribute species, 272; species, 280 ;
spiders at sea, 273; spiders, 262, 263, 379.

Balloons, description of, 267.

Balsam, 463.

Balsamea, 462.

Baltic Sea, 462.

Banks, Mr. Isaac, 94.

Bat Cave, 292.

Bates, Mr., 324, 359.

Beaded hair, 310.

Beauty of spiders, 325.

Beauvois, Palissot de, 142.

Beecher, Prof. Charles E., 455, 456.

Beethoven, Ludwig van, 307.

Belt, Mr., 443.

Berendt, Dr. G. K., 453, 457, 462, 463, 466, 469.

Bert, Prof. Paul, 342.

Bertkau, Prof. Philip, 78, 74.

Birds, 399; eat ants, 361; love dances of, 56;
of Paradise, courting display of males, 56 ;
opening cocoons, 210; sight of, 360.

Blackwall, John, 16, 30, 94, 111, 118, 119, 190,
261, 267, 268, 277, 278, 279, 280, 292, 297,
298, 347, 392, 396, 433.

Blue wasp, 382; mud dauber wasps, 383.

Blanchard, Prof. Emile, 16.

Boys, Mr. C. V., 305.

Bridge building, by spiderlings, 226.

British spiders, 194, 297, 358; see English.

Brooding cocoons, 171, 191.

Bruner, Mr. L., 397.

Buckland, Rey. Wm.,

Buckley, Prof., 385.

Butler, Mr. A. G., 334, 871.

D. D,, 458.

California, spiders of, 83, 98,
187, 209, 225, 242,

98, 135, 147, 149,
329, 333, 388, 414, 428.

AT1

Jambridge, Rey. O. Pickard, 26, 29, 66, 67, 116,
138, 206, 287, 328, 358, 364, 369, 370, 461.
Campbell, Mr. F. M., 16, 24, 47, 48, 74, 182, 202,

284, 308, 317, 318, 319, 403, 437.
Camponotus pennsylyanicus, 361.
Jannibalism, 209, 380.
Caracas, South America, 140.
Caressing, sexes of Water spider, 46, 47.
Caye fauna, origin of, 156, 157; life, effects of,
157; spiders, 154, 286, 291, 330.
Cayenne, spiders of, 142.
Central America, spiders of, 148.
Cephalothorax, color of, 349.
Chalcidians, 395, 396.
Chalybion czeruleum, 383, 384.
Chlorion czertleum, 384.
Chromatophores, 349, 350, 351.

| Citigradse, cocoons of, 143, 403; colors of, 324.

Cicada, 466.

Cicada pruinosa, 383.

Cicada septendecim, 314.

Cicada wasp, 383.

Ciniflo atrox, 280.

Clerck, Carolii, the Swedish naturalist, 29, 125.

Climate, influence on color, 333; covers, 403 ;
of Tertiary, 451.

Clubshaped hair, 311.

Clubiona, male, pairing, 45.

Clubiona attenuata, 467.

Clubiona erratica, 152, 153.

Clubiona hollosericea, 132, 194.

Clubiona pallens, 182, 288.

Clubiona putris, 393.

Clubiona tranquilla, 126, 127.

Cock-of-the-rock, 56,

Cockerell, Mr. T. B., 362.

Cocoons, 417 ; brooding of, 191; colors, 347, 348;
egress from, 210; fossil, 459; life in, 206,
207; mimicry, 372, 376; of Water spider,
46; of Wandering spiders, 433; shape of,
185; of Theridiosoma, 460; simplicity and
complexity of, 186; tent, 294; vigils of,
186; weaving of, 203; carrying, 418; har-
borage of, 171; position of, 168; methods
of production, 170; of British spiders, 194;
of Orbweavers, Chapter IV.; of Trapdoor
spiders, 64; parasitized, 397; protecting by
portage, 172; by suspensory lines, 172; re-
lation of aati 346; secreting of, 179; sev-
eral, 95; variety and complexity of, 174.

Cocooning, 63; at night, 180; caves, 403; habit,
460; in the dark, 285; multifold, 187; pe-
riod of, 229.

Ccelotes saxatilis, 125.

Coftin’s Beach, 336.

472 INDEX.

Coleoptera, 466; long life, 429.

Colonies of Medicinal spider, 237, 238; on
adult webs, 234, 2835; over water, 233; of
South American Epeiras, 231; of Epeira
triaranea, 231.

Color and color sense, Chapter XT.

Color, causes that modify, 841; consciousness
of, 338, 341; development, 60; hairs, 351;
mimicry, 867; normal, 325; of Cave spi-
ders, 385: patterns, 348; utility of, 337;
sense of, 346; structural causes of, 349; ex-
hibited by males in wooing, 52; preference
for, 3483; relation to cocoons, 347.

Colorado, Gray’s Peak, 128.:

Colors and sex, 328.

Combativeness in females, 33; in? males, 32, 33.

Communication by sound, 314.

Communities of spiders, 230.

Jonflicts of males, 30.

Conifers of the Tertiary, 464.

Consciousness of color, 333, 842.

Copal, 463.

Cope, Prof. Hdward, 450, 458.

Copulation, 73.

Coreopsis, 367.

Cornwallis, Mr. E. C., 362.

Courtship, 333, 335; attitude in, 62; first stages
of, 20.

Cowan, Edward, 307.

Crab, male, dances of, 55.

Cresson, Ezra T., 131, 396.

Cretaceous formations, 463.

Cricket, 314, 315.

Crum Creek, 368.

Crustaceans, 342; sound making, 315.

Cteniza ariana, 250.

Cteniza californica, 169, 414, 415; eggs of, 182,
183.

Cteniza fodiens, 249.

Ctenus, 148; young of, 226, 227.

Cutting ants, 357,

Cuvier, Baron, 415.

Cyclocosmia truncata, 415, 416, 417.

Cyclosa bifurca, 189, 372, 376.

Cyclosa caudata, 102, 104, 169, 191, 204, 232,
301, 372, 373, 376.

Cyclosa conica, 374.

Cynips, 142; parasitic on spiders, 246, 247.

Cyrtarachne, cocoons of, 95, 98, 97.

Cyrtarachne cornigera, 98, 99.

Cyrtophora bifurca, 95.

Dahl, Herr, 309, 312.
Dances, love, of males, 50.

Danger, influence of, 341; influences indus-

try, 407; insects unconscious of, 340.

Daphnia pulex, 342.

Darwin, Charles, 16, 80, 56, 280, 282, 273, 315,
319, 358, 359, 441, 442,

Dawson, Dr. G, M., 459.

Death, 378; feigning, 255, 488, 442; of spiders,
Chapter XTY., 419.

De Geer, Baron, 16, 27, 29, 50, 183, 150.

De Lignac, 45.

Dendryphantes capitatus, 31, 52.

Dendryphantes elegans, 33.

Dens, spider, 433.

Dermestidie, 87, 396.

Development, effects of, 208.

Diadem spider, 22; see Epeira diademata.

Digger wasp, 383, 406.

Diptera, 340,

Dispersion of young, 220.

Dissimulation of insects, 359.

Distribution of species, 97, 272, 274; vertical,
of Orbweavers, 178.

Diurnal eyes, 290.

Dodge, Capt. George H., 273.

Dolichoscaptus imops, 410, 411.

Dolichoscaptus latastei, 410.

Dolichoscaptus vittatus, 417.

Dolomedes albinius, 193.

Dolomedes lanceolatus, 194.

Dolomedes mirabilis, 27, 146, 147, 189.

Dolomedes scriptus, 195.

Dolomedes sexpunctatus, 145, 146.

Dolomedes tenebrosus, 192, 201, 299, 301.

Dolomedes, young of, 240, 241.

Domesticity of spiders, 27, 28, 39, 63.

Domicile spider, 302.

Doors of Lycosids, 405.

Dragon flies, 466.

Drassids, cocoon and nest of, 132; cocoons of,
125, 210; of England, 133; ovipositing, 181.

Drassus ater, 133, 194.

Drassus lapidicolens, 133, 194.

Drassus nitens, 133.

Drassus sylvestris, 133.

Dufour, M., 407.

Dwight, Dr. Sereno F., 280.

Dyctina, nests and cocoons of, 136, 137.

Dying, manner of, 419.

Dysdera bicolor, 134, 135, 189.

Dysdera hombergii, 189.

Eating, 428. .

Edgerton, Lord, of Tatton Hall, 290.

Edwards, Dr. Jonathan, as a naturalist, 280,
281.

——

INDEX,

Eggs, spider, 75, 88, 193, 199, 201, 202, 396;
deyoured by mother, 182; fecundity of,
177 ; irregular oviposition, 184; number of,
90, 188; parasites on, 394; within the ab-
domen, 180.

Egress from cocoon, 212.

Eigenmann, Mrs. Rosa, 83, 93, 135, 225, 329,
388, 395.

Electricity, influencing ballooning, 279.

Elis 4-notata, 384, 406, 414. :

Emerton, J. H., 21, 27, 36, 44, 49, 95, 109, 120,
121, 128, 185, 154, 155, 165, 181, 182, 187,
212, 261, 291, 292, 335, 351, 389, 404.

Enemies and their influence, Chapter XIIL.,
378.

Enemies of spiders, 191, 363, 875, 399, 411.

English spiders, 280, 284, 289, 290, 292, 328,
330, 348, 366, 369, 374, 403, 429, 433; Dras-
sids, 133; Atypus, 187; gossamer showers
of, 276.

Enock, Mr. Frederick, 29, 137, 244, 245, 249,
429,

Enyironment, 352; influence of, 412; influ-
ence on color, 334.

Entombment, manner of, in amber, 468; of
fossils, 447.

Ko thoracica, 460.

Kotypus woodwardii, 456.

Epeira apoclisa, 27, 36, 38, 187, 433.

Epeira basilica, 169, 191; cocoon string of, 106.

Kpeira bicentennaria, 330.

Epeira bifurca, 95, 169; cocoon string, 104;
see Cyclosa bifurca.

Epeira bombicinaria, 440.

Kpeira cayatica, see Epeira cinerea.

Epeira cinerea, 89, 190, 224, 398.

Epeira cooksonii, 334.

Epeira cornigera, 207.

Kpeira cornuta, 290.

Epeira cucurbitina, 330, 370.

Epeira diademata, 21, 110, 187, 188, 189, 190,
224, 283, 284, 289, 328; pairing of, 34.

Epeira domiciliorum, 224, 384; cocoon, 86, 87;
time of cocooning, 88.

Epeira epiblemum, 207.

Epeira fusca, 27; see Meta menardii.

‘Epeira gemma, 330.

Epeira inclinata, 24, 292.

Epeira infumata, 440, 441.

Epeira insularis, 208, 214, 289, 441, 453; co-
coon of, 86, 87; male, 20.

Epeira labyrinthea, 62, 99, 168, 187, 191, 222,
289, 305, 3833, 386, 390; cocoons strung in
site, 100; string of, 101; suspension of, 102;
courtship, 21.

473

Kpeira marmorea, 34; see Epeira insularis.

Epeira meekii, 453.

Kpeira parvula, 328, 371, 440.

Epeira patagiata, 327.

Epeira quadrata, 28, 189, 289, 433.

BEpeira sclopetaria, 36, 62, 207, 234, 254, 289,
293, 399, 432; time of cocooning, 88.

Epeira strix, 24, 26, 164, 165, 181, 288, 285, 329,
388, 341, 386, 431, 433, 453.

Bpeira thaddeus, 331; cocoon, 90.

Epeira triaranea, 89, 90, 195, 208, 222, 231, 338,
339.

Epeira trifolium, 289, 331, 439.

EKpeira umbratica, 179, 290, 396.

Epeira vertebrata, 26, 334.

Epiblemum scenicum, 30, 50, 57, 286.

Erigone, cocoon of, 118.

Erigonum, 147.

Erber, Mr., 250.

Eresus monarchus, 467.

Ergatis benigna, 27.

Ero, 466.

Ero setulosa, 467.

Ero thoracica, 114, 115.

Ero yariegata, 114, 115.

Eumenes, genus of wasps, 150, 131.

European spiders, 95, 289, 452.

Eurypelma hentzii, 140, 249, 321, 385, 428.

Evolution, 60, 360, 378.

Eye tubercles, 298.

Eye turrets, 297, 298.

Eyes of spiders, 283; atrophy of, 292; color of,
287; night and day, 288; structure of, 284.

Fabre, J. H., 445.

Fakir, 444.

Fairmount Park, 399.

Faithfulness, maternal, 199.

Fear paralysis, 438.

Fecundity of female spiders, 177.

Feigning death, 195, 255.

Female spiders, quiescent during courtship,
57; relative activity, 70.

Ferocity, 66.

Fertility of spiders, 189.

Fertilization, 72.

Feud among spiders, 389.

Fighting of females, 33; of males, 30, 31.

| Fingal’s Cave, spiders in, 179, 290.

Flies, 387, 442.

Florida spiders, 91, 201, 356.

Florissant spiders, 447, 458.

Flowers, attracting insects and spiders, 346;
mimicked, 368.

' Flying spiders, 256.

474

Food of cave spiders, 156; of young spiders,
213, 248.

Forel, Prof. Auguste, M. D., 295, 345.

Foreordination in Nature, 88.

Forests protected by spiders, 401.

Forethought, 202, 204.

Form mimicry, 357.

Formica exsecta, 362.

Formica fusca, 362, 427.

Formica integra, 362.

Formica rufa, 362, 364.

Formicariidie, 363.

Fossil spiders, 446; life of, 469.

Fountain Cave spiders, 291, 292.

Four spotted Elis, 385.

Franklin, Clarence P., 155.

Fraternity among broodlings, 225, 255.

Fright, 341.

Fronani, Prof., 428.

Furrow spider; see Epeira strix.

Gasteracantha, 93, 329, 340, 388.

Gasteracantha bourbonica, 93, 208.

Gentry, Dr. Allan, 434.

Geology, 458.

Geotrachea crocata, 351.

Germany, Tertiary Sea, 462; Tertiary bound-
aries of, 463.

Gerstaecker, Herr, 271, 272.

Gestation, 341; influence on color, 331.

Gnaphosa, cocoons of, 128.

Gnatcatcher, blue gray, 399.

Goeppert, Prof. Dr. H. R., 462.

Goethe on wasps, 380, 381.

God, presence of, in Nature, 204.

Golden rod, 367.

Gordius, 394.

Gossamer, floating, 259, 265, 274; how formed,
277; showers, origin of, 274, 275, 276, 278.

Gourret, M., 446, 458.

Graber, Mr., 345.

Grecian Archipelago, Trapdoor spiders of, 250.

Green, Mr. E. Ernest, 392.

Gregarious habit, 216, 217, 280.

Grenacher, H., 283.

Gretrey, M., 307.

Guerin, M., 142.

Guest wasps, 384.

Habit influencing industry, 406; modification
of, 412; value of, 61.

Habitat of spiders, 401.

Habrocestus splendens, 52, 333.

Hairs, auditory, 309; colored, 350, 351.

INDEX.

Harger, Mr., 455.

Harris, Dr. T. W., 194.

Hatching of young, period of, 207, 294.

Hawkins, Sir John, 307.

Hearing, organs of, 301.

Heer, Prof., 449.

Tienops marginatus, 393.

Hentz, Prof. Marcellus, 102, 107, 147, 192, 198,
357, 390, 417, 451.

Herman, Mr. Otto, 236, 237.

Hermeteles fasciatus, 392.

Hermeteles formosus, 392.

Herpyllus aureata, 127, 128.

Herpyllus ecclesiasticus, 191, 299, 501.

Heterapoda venatoria, 109, 153, 272, 273.

Heywood, Mr., 449.

Hibernating, 4380, 435.

Holden, Mr. William, 271.

Hope, Rev. F. W., 452, 465,

Horn, Dr. George H., 429.

Hornets, 387.

Howard, Mr. L. O., 397.

Huntsman spider, 153, 268.

Hummingbirds, 210, 399.

Hymenoptera, parasitic, 394, 397.

Hypnotism, voluntary, 444.

Ichneumon flies, 129, 189, 338, 392, 395, 466.

Icius mitratus, 54.

Illinois spiders of, 128.

Impregnation of female spider, 49, 74.

India, spiders of, 392.

Industrial mimiery, 352.

Industrial skill, 415; intuitive, 202, 338; influ-
enced by enemies, 402; by maternity, 64.

Industry, maternal, 75; unmodified, 461.

Insects, 314, 335, 340; color sense, 348; stored
by spiders, 383; fossil, 452; succinic or
amber, 465.

Instinct, 201, 202; manifest in young, 250, 251;
maternal, 75, 193, 196, 199, 200.

Insular spider, 28, 338; see Epeira insularis.

Intelligence, maternal, 185.

Internal structure, 108.

Tridescence, 335, 351.

Ithomia, 359.

Joannés, Moreau de, 142.
Jones, Rey. P. L., 186.

Keller, Dr. C., 401.

Kent, J. Sackville, 315.
Kirby and Spence, 375.
Koch, Dr. L., 84, 151, 271.

Gee, ine oe eee Es

INDEX.

475

Labyrinth spider; see Hpeira labyrinthea.

Laterigrades, 69, 180, 434, 466 ; cocoon of, 151 ;
colors of, 324, 369.

Latreille, 281.

Lathrodectus mactans, 112.

Lebert, Mr., 288.

Leidy, Prof. Joseph, 154, 336, 394, 428, 429.

Legs, 313; restored when lost, 229; color of,
349; relative length of male and female, 26.

Lepidoptera, 466.

Leptalis, 359.

Leptopelma cavicula, 409.

Leptopelma elongata, 411.

Lesquereux, 447, 450, 458.

Life, prolonged, 425.

Lignac, Abbe de, 28.

Lincecum, Dr. G., 264, 267, 385.

Linton Park, England, 362.

Linyphia communis, 341, 389.

Linyphia costata, 27; see Linyphia phry-
giana.

Linyphia erypticolens, 119.

Linyphia inserta, 292.

Linyphia marginata, 21, 29, 33, 36,.78, 119;
pairing of, 41, 42.

Linyphia montana, 16, 119.

Linyphia phrygiana = L. costata, 27.

Linyphia seripta, 389.

Linyphia subterranea, 335.

Linyphia tenebricola, 318.

Linyphia weyerii, 154, 289, 290.

Liphistius, 455.

Lister, Dr. Martin, 35, 95, 187, 264, 279, 374,
375.

Livingstone, Dr. David, 399.

Lizards, eating spiders, 379.

Local mimicry, 365.

Locy, William H., 284.

Love, a, bower, 57; call, 315; dances of male
spiders, 51; maternal, 205; signals, 21, 59.

Lubbock, Sir John, 201, 288, 296, 342, 343, 344,
345, 427, 429.

Lucas, M. H., 252.

Luray Cave, 289, 290, 336.

Lycosids, 72, 301, 334, 344, 364, 382, 403, 434;
cocoon of, 143; cocoon making, 166; effects
of music on, 301; maternal feeling of, 198 ;
maternal instincts of, 198; sight of, 295;
young of, 240, 242.

Lycosa agrestis, 189.

Lycosa arenicola, 336.

Lycosa carolinensis, 403, 407.

Lycosa herbigrada, 370.

Lycosa lenta, 193.

Lycosa narbonensis, 189, 445.

Lycosa riparia, 143.

Lycosa scutulata, 394.

Lycosa tarentula, 407.

Lycosa tigrina, 244, 384, 404, 407.
Lycosa saceata, 144, 280, 296, 314.

Magellan, Straits of, 333.

Males, 206, 333; amorous solicitations, 63; at-
titude of, 58; before mating, 18; dwelling
with females, 27; fights of, 30; love call,
315; love dances of, 50, 51,52; displays to
attract females, 57; interruptions during
pairing, 48; office of, 15; peril of, 22; po-
sition when mating, 59; pugnacity of, 32;
relative activity, 70; relative number, 16;
relative size of, 68; revelry and quarrel-
someness, 63; sluggishness of, 71; snare,
19; immature web of, 19.

Mammoth Cave, 386, 387; spiders of, 154.

Mandibles, 322.

Manifold cocooning, 95.

Marptusa familiaris, 51, 58.

Martindale, Isaac H., 98.

Marx, Dr. George, 93, 95, 98, 106, 107, 151, 334,
377, 397, 432, 460.

Mason, Prof. Wood, 322.

Maternal instincts, motherhood, Chapter VII.,
178, 190, 195, 198, 202, 212, 417, 418; and
industry, Chapter IV., 75; influence on
industry, 64.

Mating, 337.

Mating habits, comparative views of, 61.

Mechanical skill of spiders, 129, 203.

Medicinal spider, 123, 288; see Tegenaria me-
dicinalis.

Meehan, Thomas, 210, 399.

Memory, 199.

Menge, Herr A., 22, 28, 34, 73, 182, 190, 212,
218, 272, 332, 393, 394, 452.

Merejkowski, M., 342.

Merian, Madame, 142.

Mermis allicans, a “hair snake,” 393.

Meta, 178.

Meta menardii, 94, 288.

Meta segmentata, 29.

Metallic colors, 325, 349, 371.

Metatarsus, hairs on, 313.

Micaria aureata, 127, 128.

Micaria limicune, 129, 180, 203, 204.

Micaria longipes, 351.

Micaria scintilans, 358.

Micromata marmorata, 194.

Micromata ornata, 332.

Mimetic harmonies, 335, 837 ; resemblance, 353.

Mimetus interfector, 889, 390.

Mimetus syllepsicus, 390.

Mimicry, 152, 190, 337, 345; of animal forms,
357; of colors, 367; of cocoons, 372; of en-
vironment, 365; knots and buds, 366 ;
ground, 369.

Miranda adianta, 394.

Missouri spider fauna, 108.

Mistakes of mothers, 200. S

Misumena vatia, 152, 192, 324, 344, 346, 367,
369, 371, 386.

Mitchell, Dr. S. Weir, 443.

Mogeridge, T. Traherne, 182, 184, 247, 248, 249,
250, 352, 354, 355, 356, 412, 415, 416, 429, 443,

Morgan, T. H., 55.

Mortality among spiders, 222, 228; first stages
of, 422.

Motherhood, 72, 178, 186, 192, 193, 194, 197,
200, 205.

Moulting, 207, 208, 220, 229, 341; dangers of,
428; influence on color, 331; of Argiope,
22, 23; tents, 403.

Mud cocoons, 129, 203.

Mud dauber wasps, 364, 381, 382, 383, 387.

Miiller, Herr (Alpen Bliimen), 346.

Multifold cocoons, 108.

Murray, John, 276.

Muscular action, 332, 341.

Music, effects on spiders, 300, 305, 306, 307,
1309.

Mygale avicularia, 142.

Mygale blondii, 142, 189,

Mygale stridulans, 319.

Mygale truncata, see Cyclocosmia truncata.

Mygalidee, 141, 142, 169, 316, 321.

Natural selection, 363, 370, 442, 445,

Navigating spiders, 268, 269,

Nemesia coementaria, 249, 353, 355, 416.

Nemesia congener, 248, 415,

Nemesia eleanora, 248.

Nemesia manderstjernie, 356.

Nemesia meridionalis, 353, 355, 411; nest of
young, 250.

Nemesia moggridgii, 248, 356.

Nephila chrysogaster, 24.

Nephila cocoons, 92; fossil, 450.

Nephila inaurata, 93, 235,

Nephila nigra, 25, 26, 66, 235.

Nephila pennatipes, 451.

Nephila plumipes (wilderi), 66, 91, 189, 450.

Neriene dentipalpis, 277.

Nests, 339; building, 355; cocooning of Dras-
sids, 184, 135; development of trapdoor,
248; parasitism.’ 235, 888; repairing of,
196; winter, 431, 432.

INDEX.

Nesting habits, 70, 402; of Argyroneta, 45, 46;
influence of habit, 67; Trapdoor spiders, 64.

Nesticus pallidus, 154, 156, 189.

Nesticus speluncarum, 291,

New England spider fauna, 90.

New Lisbon, Ohio, spider fauna, 89.

Night eyes, 288; habits, 180, 287, 308.

Niantic, Connecticut, 20.

Nocturnal eyes, 290.

Numerical proportion, 69.

Ocellus, 283.

Odors, effects on spiders, 299.

Oeningen spiders, 452.

Ogontz Seminary, 218,

Olfactory organs, 300.

Oligocene spiders, 447, 458.

Olivet, Abbe, 307.

Oncodes pallipes, 393.

Oonops pulcher, 189.

Orbweavers, 65, 71; difference between sexes,
62; position in pairing, 62; favorite sites
of, 178; stored by wasps, 386; fossil, 466.

Orchard spider, 339, 350, 366.

Oreutt, C. R., 88, 388.

Organs of hearing, 301 302,

Ornamentation, 333. \

Orthoptera, 466. \;

Osborn, Capt., 444. ‘

Oustalet, M., 451, 458. \

Ovaries, 180. }

Oviposition, 181, 184.

Oxyopes salticus, 147.

Oxyopes viridens, 147, 193, 380.

Pachygnatha, 27.

Packard, Prof. A. 8., Jr., 128, 154, 155, 156, 286,
335.

Pairing of spiders, Chapter II., 41.

Palmer, L. Chalkley, 368.

Palpal bulb, use of, 42.

Palpipes priseus, 457.

Palps, 48, 72, 302.

Paralysis, fear, 438.

Paralyzed spiders, 383, 406.

Parasites, 129, 142, 393, 394; of body, 391; veg-
etable, 399.

Parasitic larvee, 393; spiders, 235.

Parasitism, 395, 398.

Parson spider, 127; see Herpyllus ecclesias-
ticus.

Parthenogenesis, 74.

Partridges eat ants, 362.

Patterns, dorsal, 348.

Pavesi, Sig. Prof., 291.

SS |

Peal, Mr. §, E., 319, 321.

Peckham, Prof. George W. and Elizabeth G.,
16, 21, 31, 33, 50, 51, 54, 60, 151, 187, 188,
189, 198, 199, 201, 295, 299, 301, 304, 305,
323, 328, 382, 333, 336, 337, 343, 357, 358,
365, 371, 372, 376, 440, 441, 442.

Pellisson, M., 307.

Penny, Rey. C. W., 369.

Pepsis formosa, 384, 414.

Perils of spiders, 378.

Pezomachus, 395, 398.

Pezomachus dimidiatus, 396.

Pezomachus gracilis, 396.

Pezomachus meabilis, 131.

Phidippus frenatus, 467.

Phidippus galathea, 182.

Phidippus jonsonii, 331.

Phidippus morsitans, 33, 59, 148, 189, 190, 295,
335, 350, 351, 397.

Phidippus opifex, 149, 150.

Phidippus rufus, 33, 59, 167.

Philanthropy, spider, 400.

Phileus militaris, 31, 32, 53, 57, 832.

Philodromin, 147.

Philodromus fallax, 370.

Philodromus microcephalus, 467.

Philodromus mollitor, 151.

Pholeus phalangioides, 120, 186, 222, 236.

Physical vigor, 63,

Pigment, 341, 349, 351.

Pike, Col, Nicholas, 397.

Pinites succinifer, 462.

Pirata piraticus, 198.

Pittidee, 362.

Pliny on gossamer showers, 274.

Poison of wasps, 382.

Pollock, Frederick, Esq., 188, 209, 228.

Polysphincta carbonaria, 391.

Pompilus annulatus, 445.

Pompilus formosus; see Pepsis formosa.

Portage of cocoon, 119, 120, 153.

Presbyterian and Reformed Review, 282.

Prey, capturing, 70, 286, 368.

Preyer, Dr., 487.

Priocnemus pomilius, 384.

Proctotrupids, 397.

Prosthesima eecclesiastica; see Herpyllus ec-
clesiasticus.

Protection, 192; by colors, 3388; of female
during courtship, 57.

Protective architecture, 402, 409; forms, 358 ;
habits, 878; resemblance, $75.

Protolycosa anthrocophila, 450, 454.

Psalistops melanophygia, 140.

Pseudidiops opifex, 356.

ATT

Pucetia aurora, 147, 149, 241.
Pugnacity of males, 32.
Purseweb spiders, 356; see Atypus abbotii.

Quails eat ants, 362.
Queen of ants, 427.

Rats eat spiders, 380.

Ray spider, 288, 460; see Theridiosoma radi-
osum.

Rearing spiders artificially, 213.

Red color preferred by spiders, 344.

Rennie, Mr., 433.

Resins, 463.

Retitelarize, 434.

Rhytidicolus structor, 139.

Riley, Dr. C. V., 362.

Romer, Mr., 453.

Romanes, G. J., 441, 444.

Russian spiders, 289.

Saitis pulex, 51, 397.

Saltigrades, 30, 180, 434, 466; brilliant eyes of,
287; cocoons of, 148; cocoon making, 167;
sense of smell, 299; sight of, 295.

Samland, 462, 463, 469.

Sanborn, F. G., 154.

San Domingo, 142.

Sauvages, Abbe, 64, 414.

Seales of colors, 351.

Scelionine, 397.

Schindler, Anthon, 307.

Schiodte, Mr. J. C., 438.

Scorpions, 315; stridulating, 316.

Scudder, Prof. 8. H., 148, 226, 315, 446, 447, 448,
450, 452, 455, 457, 458, 459.

Scytodes thoracica, 120,

Season, influences of, 209.

Secretiveness, 355.

Seebach, Herr, 458.

Segestria, 466.

Segestria canities, 185, 136,

Segestria nana, 467.

Segmentation, 349.

Self protection, 353, 354, 404.

| Senses of spiders, 288, 314.

Sex, influence on color, 328, 332.

Sexes, numerical proportion of, 69; relative
size of,.24.

Shaler, Prof., 154,

Shamming death, 440.

Shamrock spider, 17, 28; see Epeira trifolium.

Sharp, Dr. Dayid, 429.

Sheep eat spiders, 380.

Showers of gossamer, 274, 275.

478

INDEX,

Shrilling of insects, 314.

Sight of spiders, accuracy of, 286; limited,
295, 296.

Silk, spider, colors of, 348.

Silliman, Prof. Benjamin, 281, 282.

Simon, M. Eugene, 69, 188, 189, 140, 169, 184,
287, 288, 297, 332, 347, 356, 389, 409, 411,
412, 415, 417, 461.

Simonella americana, 357.

Sites, cocooning, 75.

Size, spiders’, difference in, 67; disparity in,
62; influencing development, 66; varia-
tion in, 26,

Smell, organs of, 800; sense of, 299.

Smyth, Prof. Egbert C., 280.

Solitary habit, 63; wasps, 387.

Sounds, effects of, on spiders, 301; uses of, 314.

South American spiders, 333, 358, 412.

Sparrows eat ants, 362.

Speckled Agalena, see Agalena neevia.

Spermatozoa, 72.

Sphecius speciosus, 383.

Spheroma, 515.

Spiderlings, life of, 197.

Spines, abdominal, 330; protective, 339.

Spinning, habit influencing courtship, 62; in-
dustry defective in male, 63.

Skill, mechanical, of spiders, 202, 203, 404.

Skinner, Miss C., 218.

Staveley, E. F., 28, 118, 119, 132, 188, 189, 194,
347, 399.

Steatoda bipunctata, 318.

Steatoda borealis, 27, 169) 186; pairing, 44.

Steatoda guttata, 317.

Steatoda maculata, 120.

Stothis astuta, 412, 413.

Stothis cenobita, 412, 413.

Stridulating apparatus, 316; Mygale, 319.

Stridulation, of spiders, 317, 318; organs of,
317, 318; uses of, 319.

Structure and color, 351.

Sunbirds, 399.

Survival of the fittest, 370.

Swallows eat spiders, 379.

Swedish spiders, 316.

Swifts, 379.

Swinging basket, 254, 281.

Synagales picata, 33, 189; love dance, 51.

Synemosyna americana, 359.

Synemosyna formica, 357, 358.

Syphax megacephalus, 467.

Sytodes cameratus, 120, 121.

Tactile hairs, 310, 313.
Tailed spider, 102; see Cyclosa caudata,

Tarantula, 140, 166, 800, 885, 409, 443; age of,
429; cocoon of, 141; striking, 320, 322.

Tarantula killer, 384, 385.

Tarentula tigrina, see Lycosa tigrina.

Tatton Hall, 290.

Tegenaria agrestis, 181, 187.

Tegenaria ciyilis, 280, 429.

Tegenaria derhamii, 123.

Tegenaria domestica, 189, 308, 429.

Tegenaria emaciata, 131.

Tegenaria guyonii, 16, 47, 74, 202; courtship
of, 24.

Tegenaria medicinalis, 128, 124, 169, 189, 286,
330, 334, 387, 347.

Tegenaria persica, 123.

Tents, cocooning, 86, 87, 294.

Termeyer, Raymond, 21, 22, 33, 110, 142, 187.

Territelarix, 455; cocoons of, 137, 148; mak-
ing cocoons, 166.

Tertiary spiders, 446; trees, 464.

Tethneus, 452.

Tetragnatha, 379; mimicry of, 147.

Tetragnatha elongata, 365.

Tetragnatha extensa, 365, 366, 386, 451; co-
coon. of, 94, 96; pairing of, 34.

Tetragnatha grallator, 365, 451.

Tetragnatha tertiaria, 451.

Teutana triangulosa, 377; see Theridium ser-
pentinum.

Texas, Gossamer spider of, 267.

Textris lycosina, 125.

Theraphosoidee, 140,

Theridioids, colors of, 324.

Theridiosoma gemmosum, 461,

Theridiosoma radiosum, cocoon of, 461.

Theridium, 466.

Theridium carolinum, 120.

Theridium differens, 116, 417.

Theridium frondeum, 114, 115, 460; cocoon,
see Theridiosoma radiosum, 461.

Theridium globosum, 199.

Theridium hirtum, 467.

Theridium lineatum, 116.

Theridium maculatum, 120.

Theridium neryosum, 119.

Theridium pallens, 116.

Theridium riparium, 65.

Theridium serpentinum, see Teutana triangu-
losa, 112, 877.

Theridium studiosum, 169, 193. «

Theridium tepidariorum, 27, 111, 112, 164, 165,
169, 222, 237, 334, 876, 386; 389, 435.

Theridium varians, 118.

Theridium variegatum, 189.

Theridium zelotypum, 64, 119.

OE ee eae Ne eee ee ee

Thomisoids, 364, “4

Thomisus cristatus, 151, 280.

Thorell, Prof. Tamerlan, 147, 272, 452, 453, 457.

Tibellus oblongus, 365.

Tibia, hairs on, 313.

Tiger spider; see Lycosa tigrina.

Tigrina, Lycosa, 408.

Touch, sense of, 200, 285, 303.

Tower, trapdoor, 411.

Townsend, Charles H., 451.

Trapdoor spiders, 64, 169, 183, 184, 247, 354,
355, 356, 404, 409, 411, 414, 429 ;. cocoons of,
139, 140.

Treat, Mrs. Mary, 91, 107, 108, 186, 187, 188,
190, 192, 193, 195, 196, 197, 201, 211, 243, 367,
375, 403, 404, 405, 434.

Tree, trapdoor, 356,

Trees of Tertiary, 464.

Trevelyan, Sir C. E., 444.

Trifolium, Epeira, male of, 326; varieties of
color, 325, 326,

Trypoxylon politum, 383, 384.

Tubercles of eyes, 298.

Tubeweayers, 179; colors of, 324; sexual size
in, 67.

Tuning fork experiments, 302. |

Tunnel weavers, 179, 324, 364, 409; industry, 64. |

Turret spider, 198, 242, 243, 337, 407.

Turrets on eyes, 297.

Uloborus, 106, 866, 376.

Uloborus mammeatus, 107, 192.
Uloborus plumipes, 108, 109, 285, 376.
Uloborus riparia, 95.

Uloborus waleckenaerius, 107.
Unmodified industry, 461.
Upholstering cocoons, 131.

Variation, 359.

Venezuela, spiders of, 139, 409.
Vigils, maternal, 186, 190.
Vinson, Dr., 24, 93, 235, 287, 329.
Vireo noveborocensis, 211, 399.
Virey, M., 279.

Vision in spiders, 285, 295, 296.
Voelker, Mr. Carl, 361, 399.
Volcanic showers, 449.

Von Heyden, C., 452.

Von Meyer, Herr, 457.

Wafer trapdoor, 412.

Wagner, Mr. Waldemar, 289, 310, 311, 312,
313, 314.

Walckenaer, Baron, 24, 27, 28, 35, 36, 38, 45, 47,
95, 181, 132, 142, 188, 189, 230, 246, 249, 307,

346,

— =

479

Walckenaera acuminata, 121, 297.

Wallace, Alfred Russell, 56, 251, 324, 326, 343,
363, 411.

Walsh, Benj. D., 384.

Wanderers, cocoon site, 179.

Warning coloration, 335, 340.

Wasps, 466; as mimics, 363; mud daubs, 338;
mud nests of, 130; solitary, diggers, mud
daubers, social, 387.

Water spider, young of, 239; see Argyroneta
aquatica,

Weather, 404.

Weaver, Prof. G. E. H., 428.

Weaving, in darkness, 286; process of, ital:
161, 163.

Webster, F. M., 128, 129, 397.

Westring, Prof. Nicolas, 189, 315, 316, 317, 319.

White, Rey. Gilbert, 275. ‘

Wilder, Prof. Burt G., 34, 91, 142, 189, 207, 209,
210, 212, 285.

Winds, Trade, carrying spiders, 268.

Winter, effects, 213; habits, 430.

Wisconsin, spiders of, 371.

Wittfield, Miss Anna, 104.

Wood, Rev. J. H., 93.

Wood-Mason, Prof. James, 315, 316, 319, 321.

Woodpecker, pileated, 361.

Woodward, Dr. Henry, 456.

Wooing and mating habits, Chapter I., 15, 63.

Workman, Mr. Thomas, 289.

Wright, Mr. W. G., 135, 147, 149, 242.

Wright, Rev. A., 362.

Xenarchus, 315.
Xysticus audax, 151.
Xysticus ferox, 33.

| Xysticus gulosus, 33.

Xysticus sabulosis, 370.
Xysticus trivittata, pairing of, 49.

Young spiders, 193, 197, 314, 370, 375, 376; of
Agalena, 251, 252; of Gasteracantha, 340;
of Lycosids, 72, 240; of Dolomedes, 241; of
Trapdoor spiders, 247; ballooning habit of,
256; development of color, 327; feeding of,
195; first movements, 217, 218; mimics, 374 ;
escape from cocoon, 211, 214; no metamor-
phosis, 206; sensitive to light, 292, 293.

Zaddach, Prof. G., 463, 464.
Zilla, 91, 209, 466,

Zilla callophila, 39.

Zilla gracilis, fossil, 467.

Zilla porrecta, 467.

Zilla x-notata, 225, 292, 328, 388,
Zygoballus bettini, 31.

SUPPLEMENTARY LIST OF SUBSORIBERS.*

Prof. Aug. Weismann, Freiburg, Baden, Germany.

Isaac C. Martindale, Philadelphia,

Public. Library, Bangor, Maine. (Mrs. M. H. Curran.)

Hon. J. K. Valentine, Philadelphia.

Lieut. Col. Linley Blathwayt, Bath, England. (William Wesley & Son, London.)

Feret & Fils, Bordeaux, France. ?

Prof. A. Agassiz, Cambridge, Massachusetts.

R. C. MeMurtrie, Philadelphia.

Alfred C. Harrison, Philadelphia.

Col. A. K. McClure, Philadelphia.

George C. Thomas, Philadelphia.

Carl Edelheim, Philadelphia.

Rey. W. C. Cattell, D. D., LL. D., Philadelphia.

Gen. I. J. Wistar, Philadelphia.

Dr. J. B. Brinton, Philadelphia.

Koenigliche Bibliothek, Berlin, Germany.

Kaiserliche Universitiits und Landesbibliothek, Strassburg, Germany. (Otto Harrassowitz,
Leipzig.) :

State Laboratory of Natural History, Champaign, Illinois.

Conyers Button, Philadelphia. (Two copies.)

The Signet Library, Edinburgh, Scotland. (Thomas G. Law.)

W. B. Clark & Co., Boston, Massachusetts. (Two copies.) :

Dartmouth College, Hanover, New Hampshire. (Charles F, Richardson.)

Charles Schaffer, Philadelphia.

Rey. F. O. Pickard-Cambridge, Carlisle, England. (William Wesley & Son, London.)

Enoch Pratt Library, Baltimore. (Lewis H. Steiner.)

Justus C. Strawbridge, Philadelphia. (Two copies.)

Haverford College, Haverford, Pennsylyania.?

Librairie, H. Georg, Geneve, Switzerland.

Edinburgh Public Library, Scotland. (Hodder & Stoughton, London.)

Mrs. Henry Draper, New York City.

J. B. Lippincott Co., Philadelphia.

Proyidence Atheneum, Providence, Rhode Island. (Daniel Beckwith.)

University of Minnesota, Minneapolis. (Prof. Henry F. Nachtuit.)

State Library of Pennsylvania, Harrisburg. (Porter & Coates, Philadelphia.)

Mrs. Emma W. Bucknell, Philadelphia.

John MacFarlane, Detroit, Michigan, _

James Kelly, New York City. —s

Entomological Society of Ontario, London, Ontario, Canada.

University of California, Berkeley, California. (Methodist Book Depository, San Francisco.)

Kegan Paul, Trench, Tribner & Co., London, England.

A. 8. Bertolet, Cummings, Illinois. (A. C. McClurg & Co., Chicago.)

1This list includes all subseriptions made since November Ist, 1889. All before that
date are published in Volume I, The author wishes to print a complete and corrected list
at the close of Volume III., and requests that any mistakes heretofore made may be com-
municated to him. He also requests all subscribers to send their names and addresses in
full, with proper titles of individuals and full names of societies, libraries, ete. )
2 Presented by Mr. Justus C. Strawbridge. *

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Full text of "American spiders and their spinning work. A natural history of the orbweaving spiders of the United States, with special regard to their industry and habits"

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