KEYS TO THE ARTHROPODS NOXIOUS TO MAN 257-317
Crustacea.
Myriapoda, or centipedes and millipedes.
Arachnida (Orders of).
Acarina or ticks.
Hexapoda (Insecta).
Siphunculata and Hemiptera (lice and true bugs).
Diptera (mosquitoes, midges, and flies).
Siphonaptera (fleas).
APPENDIX
Hydrocyanic acid gas against household insects 318-320
Proportion of ingredients. A single room as an example.
Fumigating a large house. Precautions.
Lesions produced by the bite of the black-fly 321-326
BIBLIOGRAPHY 327-340
INDEX 341-348
CHAPTER I.
INTRODUCTION
EARLY SUGGESTIONS REGARDING THE TRANSMISSION OF DISEASE BY INSECTS
Until very recent years insects and their allies have been considered as
of economic importance merely in so far as they are an annoyance or
direct menace to man, or his flocks and herds, or are injurious to his
crops. It is only within the past fifteen years that there has sprung
into prominence the knowledge that in another and much more insiduous
manner, they may be the enemy of mankind, that they may be among the
most important of the disseminators of disease. In this brief period,
such knowledge has completely revolutionized our methods of control of
certain diseases, and has become an important weapon in the fight for
the conservation of health.
It is nowhere truer than in the case under consideration that however
abrupt may be their coming into prominence, great movements and great
discoveries do not arise suddenly. Centuries ago there was suggested the
possibility that insects were concerned with the spread of disease, and
from time to time there have appeared keen suggestions and logical
hypotheses along this line, that lead us to marvel that the
establishment of the truths should have been so long delayed.
One of the earliest of these references is by the Italian physician,
Mercurialis, who lived from 1530 to 1607, during a period when Europe
was being ravaged by the dread "black death", or plague. Concerning its
transmission he wrote: "There can be no doubt that flies feed on the
internal secretions of the diseased and dying, then, flying away, they
deposit their excretions on the food in neighboring dwellings, and
persons who eat of it are thus infected."
It would be difficult to formulate more clearly this aspect of the facts
as we know them to-day, though it must always be borne in mind that we
are prone to interpret such statements in the light of present-day
knowledge. Mercurialis had no conception of the animate nature of
contagion, and his statement was little more than a lucky guess.
Much more worthy of consideration is the approval which was given to his
view by the German Jesuit, Athanasius Kircher in 1658. One cannot read
carefully his works without believing that long before Leeuwenhook's
discovery, Kircher had seen the larger species of bacteria. Moreover, he
attributed the production of disease to these organisms and formulated,
vaguely, to be sure, a theory of the animate nature of contagion. It has
taken two and a half centuries to accumulate the facts to prove his
hypothesis.
The theory of Mercurialis was not wholly lost sight of, for in the
medical literature of the eighteenth century there are scattered
references to flies as carriers of disease. Such a view seems even to
have been more or less popularly accepted, in some cases. Gudger (1910),
has pointed out that, as far back as 1769, Edward Bancroft, in "An Essay
on the Natural History of Guiana in South America," wrote concerning the
contagious skin-disease known as "Yaws": "It is usually believed that
this disorder is communicated by the flies who have been feasting on a
diseased object, to those persons who have sores, or scratches, which
are uncovered; and from many observations, I think this is not
improbable, as none ever receive this disorder whose skins are whole."
Approaching more closely the present epoch, we find that in 1848, Dr.
Josiah Nott, of Mobile, Alabama, published a remarkable article on the
cause of yellow fever, in which he presented "reasons for supposing its
specific cause to exist in some form of insect life." As a matter of
fact, the bearing of Nott's work on present day ideas of the insect
transmission of disease has been very curiously overrated. The common
interpretation of his theory has been deduced from a few isolated
sentences, but his argument appears quite differently when the entire
article is studied. It must be remembered that he wrote at a period
before the epoch-making discoveries of Pasteur and before the
recognition of micro-organisms as factors in the cause of disease. His
article is a masterly refutation of the theory of "malarial" origin of
"all the fevers of hot climates," but he uses the term "insect" as
applicable to the lower forms of life, and specific references to
"mosquitoes," "aphids," "cotton-worms," and others, are merely in the
way of similes.
But, while Nott's ideas regarding the relation of insects to yellow
fever were vague and indefinite, it was almost contemporaneously that
the French physician, Louis Daniel Beauperthuy argued in the most
explicit possible manner, that yellow fever and various others are
transmitted by mosquitoes. In the light of the data which were available
when he wrote, in 1853, it is not surprising that he erred by thinking
that the source of the virus was decomposing matter which the mosquito
took up and accidentally inoculated into man. Beauperthuy not only
discussed the role of mosquitoes in the transmission of disease, but he
taught, less clearly, that house-flies scatter pathogenic organisms. It
seems that Boyce (1909) who quotes extensively from this pioneer work,
does not go too far when he says "It is Dr. Beauperthuy whom we must
regard as the father of the doctrine of insect-borne disease."
In this connection, mention must be made of the scholarly article by the
American physician, A. F. A. King who, in 1883, brought together an all
but conclusive mass of argument in support of his belief that malaria
was caused by mosquitoes. At about the same time, Finley, of Havana, was
forcefully presenting his view that the mosquito played the chief role
in the spread of yellow fever.
To enter more fully into the general historical discussion is beyond the
scope of this book. We shall have occasion to make more explicit
references in considering various insect-borne diseases. Enough has been
said here to emphasize that the recognition of insects as factors in the
spread of disease was long presaged, and that there were not wanting
keen thinkers who, with a background of present-day conceptions of the
nature of disease, might have been in the front rank of investigators
along these lines.
THE WAYS IN WHICH ARTHROPODS MAY AFFECT THE HEALTH OF MAN
When we consider the ways in which insects and their allies may affect
the health of man, we find that we may treat them under three main
groups:
A. They may be directly poisonous. Such, for example, are the scorpions,
certain spiders and mites, some of the predaceous bugs, and stinging
insects. Even such forms as the mosquito deserve some consideration from
this viewpoint.
B. They may be parasitic, living more or less permanently on or in the
body and deriving their sustenance from it.
Of the parasitic arthropods we may distinguish, first, the _true
parasites_, those which have adopted and become confirmed in the
parasitic habit. Such are the itch mites, the lice, fleas, and the
majority of the forms to be considered as parasitic.
In addition to these, we may distinguish a group of _accidental_, or
_facultative parasites_, species which are normally free-living, feeding
on decaying substances, but which when accidentally introduced into the
alimentary canal or other cavities of man, may exist there for a greater
or less period. For example, certain fly larvae, or maggots, normally
feeding in putrifying meat, have been known to occur as accidental or
facultative parasites in the stomach of man.
C. Finally, and most important, arthropods may be transmitters and
disseminators of disease. In this capacity they may function in one of
three ways; as _simple carriers_, as _direct inoculators_, or as
_essential hosts_ of disease germs.
As simple carriers, they may, in a wholly incidental manner, transport
from the diseased to the healthy, or from filth to food, pathogenic
germs which cling to their bodies or appendages. Such, for instance, is
the relation of the house-fly to the dissemination of typhoid.
As direct inoculators, biting or piercing species may take up from a
diseased man or animal, germs which, clinging to the mouth parts, are
inoculated directly into the blood of the insect's next victim. It it
thus that horse-flies may occasionally transmit anthrax. Similarly,
species of spiders and other forms which are ordinarily perfectly
harmless, may accidentally convey and inoculate pyogenic bacteria.
It is as essential hosts of disease germs that arthropods play their
most important role. In such cases an essential part of the life cycle
of the pathogenic organism is undergone in the insect. In other words,
without the arthropod host the disease-producing organism cannot
complete its development. As illustrations may be cited the relation of
the Anopheles mosquito to the malarial parasite, and the relation of the
cattle tick to Texas fever.
A little consideration will show that this is the most important of the
group. Typhoid fever is carried by water or by contaminated milk, and in
various other ways, as well as by the house-fly. Kill all the
house-flies and typhoid would still exist. On the other hand, malaria is
carried only by the mosquito, because an essential part of the
development of the malarial parasite is undergone in this insect.
Exterminate all of the mosquitoes of certain species and the
dissemination of human malaria is absolutely prevented.
Once an arthropod becomes an essential host for a given parasite it may
disseminate infection in three different ways:
1. By infecting man or animals who ingest it. It is thus, for example,
that man, dog, or cat, becomes infected with the double-pored dog
tapeworm, _Dipylidium caninum_. The cysticercoid stage occurs in the
dog louse, or in the dog or cat fleas, and by accidentally ingesting the
infested insect the vertebrate becomes infested. Similarly, _Hymenolepis
diminuta_, a common tapeworm of rats and mice, and occasional in man,
undergoes part of its life cycle in various meal-infesting insects, and
is accidentally taken up by its definitive host. It is very probable
that man becomes infested with _Dracunculus (Filaria) medinensis_
through swallowing in drinking water, the crustacean, _Cyclops_,
containing the larvae of this worm.
2. By infecting man or animals on whose skin or mucous membranes the
insect host may be crushed or may deposit its excrement. The pathogenic
organism may then actively penetrate, or may be inoculated by
scratching. The causative organism of typhus fever is thus transmitted
by the body louse.
3. By direct inoculation by its bite, the insect host may transfer the
parasite which has undergone development within it. The malarial
parasite is thus transferred by mosquitoes; the Texas fever parasite by
cattle ticks.
CHAPTER II.
ARTHROPODS WHICH ARE DIRECTLY POISONOUS
Of all the myriads of insects and related forms, a very few are of
direct use to man, some few others have forced his approbation on
account of their wonderful beauty, but the great hordes of them are
loathed or regarded as directly dangerous. As a matter of fact, only a
very small number are in the slightest degree poisonous to man or to the
higher animals. The result is that entomologists and lovers of nature,
intent upon dissipating the foolish dread of insects, are sometimes
inclined to go to the extreme of discrediting all statements of serious
injury from the bites or stings of any species.
Nevertheless, it must not be overlooked that poisonous forms do exist,
and they must receive attention in a consideration of the ways in which
arthropods may affect the health of man. Moreover, it must be recognized
that "what is one man's meat, is another man's poison," and that in
considering the possibilities of injury we must not ignore individual
idiosyncrasies. Just as certain individuals may be poisoned by what, for
others, are common articles of food, so some persons may be abnormally
susceptible to insect poison. Thus, the poison of a bee sting may be of
varying severity, but there are individuals who are made seriously sick
by a single sting, regardless of the point of entry. Some individuals
scarcely notice a mosquito bite, others find it very painful, and so
illustrations of this difference in individuals might be multiplied.
In considering the poisonous arthropods, we shall take them up by
groups. The reader who is unacquainted with the systematic relationship
of insects and their allies is referred to Chapter XII. No attempt will
be made to make the lists under the various headings exhaustive, but
typical forms will be discussed.
ARANEIDA OR SPIDERS
Of all the arthropods there are none which are more universally feared
than are the spiders. It is commonly supposed that the majority, if not
all the species are poisonous and that they are aggressive enemies of
man and the higher animals, as well as of lower forms.
That they really secrete a poison may be readily inferred from the
effect of their bite upon insects and other small forms. Moreover, the
presence of definite and well-developed poison glands can easily be
shown. They occur as a pair of pouches (fig. 1) lying within the
cephalothorax and connected by a delicate duct with a pore on the claw
of the chelicera, or so-called "mandible" on the convex surface of the
claw in such a position that it is not plugged and closed by the flesh
of the victim.
[Illustration: 1. Head of a spider showing poison gland (_c_) and its
relation to the chelicera (_a_).]
The glands may be demonstrated by slowly and carefully twisting off a
chelicera and pushing aside the stumps of muscles at its base. By
exercising care, the chitinous wall of the chelicera and its claw may be
broken away and the duct traced from the gland to its outlet. The inner
lining of the sac is constituted by a highly developed glandular
epithelium, supported by a basement membrane of connective tissue and
covered by a muscular layer, (fig. 2). The muscles, which are striated,
are spirally arranged (fig. 1), and are doubtless under control of the
spider, so that the amount of poison to be injected into a wound may be
varied.
[Illustration: 2. Section through a venom gland of Latrodectus
13-guttatus showing the peritoneal, muscular and epithelial layers.
After Bordas.]
The poison itself, according to Kobert (1901), is a clear, colorless
fluid, of oily consistency, acid reaction, and very bitter taste. After
the spider has bitten two or three times, its supply is exhausted and
therefore, as in the case of snakes, the poison of the bite decreases
quickly with use, until it is null. To what extent the content of the
poison sacs may contain blood serum or, at least, active principles of
serum, in addition to a specific poison formed by the poison glands
themselves, Kobert regards as an open question. He believes that the
acid part of the poison, if really present, is formed by the glands and
that, in the case of some spiders, the ferment-like, or better, active
toxine, comes from the blood.
[Illustration: 3. Chelicera of a spider.]
But there is a wide difference between a poison which may kill an insect
and one which is harmful to men. Certain it is that there is no lack of
popular belief and newspaper records of fatal cases, but the evidence
regarding the possibility of fatal or even very serious results for man
is most contradictory. For some years, we have attempted to trace the
more circumstantial newspaper accounts, which have come to our notice,
of injury by North American species. The results have served, mainly, to
emphasize the straits to which reporters are sometimes driven when there
is a dearth of news. The accounts are usually vague and lacking in any
definite clue for locating the supposed victim. In the comparatively few
cases where the patient, or his physician, could be located, there was
either no claim that the injury was due to spider venom, or there was no
evidence to support the belief. Rarely, there was evidence that a
secondary blood poisoning, such as might be brought about by the prick
of a pin, or by any mechanical injury, had followed the bite of a
spider. Such instances have no bearing on the question of the venomous
nature of these forms.
[Illustration: 4. The Italian tarantula (Lycosa tarantula). After
Kobert.]
The extreme to which unreasonable fear of the bites of spiders
influenced the popular mind was evidenced by the accepted explanation of
the remarkable dancing mania, or tarantism, of Italy during the Middle
Ages. This was a nervous disorder, supposed to be due to the bite of a
spider, the European tarantula (fig. 4), though it was also, at times,
attributed to the bite of the scorpion. In its typical form, it was
characterized by so great a sensibility to music that under its
influence the victims indulged in the wildest and most frenzied dancing,
until they sank to the ground utterly exhausted and almost lifeless. The
profuse perspiring resulting from these exertions was supposed to be the
only efficacious remedy for the disease. Certain forms of music were
regarded as of especial value in treating this tarantism, and hence the
name of "tarantella" was applied to them. Our frontispiece, taken from
Athanasius Kircher's _Magnes sive de Arte Magnetica_, 1643 ed.,
represents the most commonly implicated spider and illustrates some of
what Fabre has aptly designated as "medical choreography."
The disease was, in reality, a form of hysteria, spreading by sympathy
until whole communities were involved, and was paralleled by the
outbreaks of the so-called St. Vitus's or St. John's dance, which swept
Germany at about the same time (fig. 5). The evidence that the spider
was the cause of the first is about as conclusive as is that of the
demoniacal origin of the latter. The true explanation of the outbreaks
is doubtless to be found in the depleted physical and mental condition
of the people, resulting from the wars and the frightful plagues which
devastated all Europe previous to, and during these times. An
interesting discussion of these aspects of the question is to be found
in Hecker.
[Illustration: 5. Dancing mania. Illustration from Johann Ludwig
Gottfried's Chronik. 1632.]
So gross has been the exaggeration and so baseless the popular fear
regarding spiders that entomologists have been inclined to discredit all
accounts of serious injury from their bites. Not only have the most
circumstantial of newspaper accounts proved to be without foundation but
there are on record a number of cases where the bite of many of the
commoner species have been intentionally provoked and where the effect
has been insignificant. Some years ago the senior author personally
experimented with a number of the largest of our northern species, and
with unexpected results. The first surprise was that the spiders were
very unwilling to bite and that it required a considerable effort to get
them to attempt to do so. In the second place, most of those
experimented with were unable to pierce the skin of the palm or the back
of the hand, but had to be applied to the thin skin between the fingers
before they were able to draw blood. Unfortunately, no special attempt
was made to determine, at the time, the species experimented with, but
among them were _Theridion tepidariorum_, _Miranda aurantia_
(_Argiopa_), _Metargiope trifasciata_, _Marxia stellata_, _Aranea
trifolium_, _Misumena vatia_, and _Agelena naevia_. In no case was the
bite more severe than a pin prick and though in some cases the sensation
seemed to last longer, it was probably due to the fact that the mind was
intent upon the experiment.
[Illustration: 6. An American tarantula (Eurypelma hentzii). Natural
size. After Comstock.]
Similar experiments were carried out by Blackwell (1855), who believed
that in the case of insects bitten, death did not result any more
promptly than it would have from a purely mechanical injury of equal
extent. He was inclined to regard all accounts of serious injury to man
as baseless. The question cannot be so summarily dismissed, and we shall
now consider some of the groups which have been more explicitly
implicated.
THE TARANTULAS.--In popular usage, the term "tarantula" is loosely
applied to any one of a number of large spiders. The famous tarantulas
of southern Europe, whose bites were supposed to cause the dancing
mania, were Lycosidae, or wolf-spiders. Though various species of this
group were doubtless so designated, the one which seems to have been
most implicated was _Lycosa tarantula_ (L.), (fig. 4). On the other
hand, in this country, though there are many Lycosidae, the term
"tarantula" has been applied to members of the superfamily Avicularoidea
(fig. 6), including the bird-spiders.
Of the Old World Lycosidae there is no doubt that several species were
implicated as the supposed cause of the tarantism. In fact, as we have
already noted, the blame was sometimes attached to a scorpion. However,
there seems to be no doubt that most of the accounts refer to the spider
known as _Lycosa tarantula_.
There is no need to enter into further details here regarding the
supposed virulence of these forms, popular and the older medical
literature abound in circumstantial accounts of the terrible effects of
the bite. Fortunately, there is direct experimental evidence which bears
on the question.
Fabre induced a common south European wolf-spider, _Lycosa narbonensis_,
to bite the leg of a young sparrow, ready to leave the nest. The leg
seemed paralyzed as a result of the bite, and though the bird seemed
lively and clamored for food the next day, on the third day it died. A
mole, bitten on the nose, succumbed after thirty-six hours. From these
experiments Fabre seemed justified in his conclusion that the bite of
this spider is not an accident which man can afford to treat lightly.
Unfortunately, there is nothing in the experiments, or in the symptoms
detailed, to exclude the probability that the death of the animals was
the result of secondary infection.
As far back as 1693, as we learn from the valuable account of Kobert,
(1901), the Italian physician, Sanguinetti allowed himself to be bitten
on the arm by two tarantulas, in the presence of witnesses. The
sensation was equivalent to that from an ant or a mosquito bite and
there were no other phenomena the first day. On the second day the wound
was inflamed and there was slight ulceration. It is clear that these
later symptoms were due to a secondary infection. These experiments have
been repeated by various observers, among whom may be mentioned Leon
Dufour, Josef Erker and Heinzel, and with the similar conclusion that
the bite of the Italian tarantula ordinarily causes no severe symptoms.
In this conclusion, Kobert, though firmly convinced of the poisonous
nature of some spiders, coincides. He also believes that striking
symptoms may be simulated or artificially induced by patients in order
to attract interest, or because they have been assured that the bite,
under all circumstances, caused tarantism.
The so-called Russian tarantula, _Trochosa singoriensis_ (fig. 7), is
much larger than the Italian species, and is much feared. Kobert carried
out a series of careful experiments with this species and his results
have such an important bearing on the question of the venomous nature of
the tarantula that we quote his summary. Experimenting first on nearly a
hundred living specimens of _Trochosa singoriensis_ from Crimea he says
that:
"The tarantulas, no matter how often they were placed on the skin,
handled, and irritated, could not be induced to bite either myself, the
janitor, or the ordinary experimental animals. The objection that the
tarantulas were weak and indifferent cannot stand, for as soon as I
placed two of them on the shaved skin of a rabbit, instead of an attack
on the animal, there began a furious battle between the two spiders,
which did not cease until one of the two was killed."
[Illustration: 7. Trochosa singoriensis. After Kobert.]
"Since the spiders would not bite, I carefully ground up the fresh
animals in physiological salt solution, preparing an extract which must
have contained, in solution, all of the poisonous substance of their
bodies. While in the case of _Latrodectus_, as we shall see, less than
one specimen sufficed to yield an active extract, I have injected the
filtered extract of six fresh Russian tarantulas, of which each one was
much heavier than an average _Latrodectus_, subcutaneously and into the
jugular vein of various cats without the animals dying or showing any
special symptoms. On the basis of my experiments I can therefore only
say that the quantity of the poison soluble in physiological salt
solution, even when the spiders are perfectly fresh and well nourished,
is very insignificant. That the poison of the Russian tarantula is not
soluble in physiological salt solution, is exceedingly improbable.
Moreover, I have prepared alcoholic extracts and was unable to find them
active. Since the Russian spider exceeds the Italian in size and in
intensity of the bite, it seems very improbable to me that the
pharmacological test of the Italian tarantula would yield essentially
other results than those from the Russian species."
To the AVICULAROIDEA belong the largest and most formidable appearing of
the spiders and it is not strange that in the New World they have fallen
heir to the bad reputation, as well as to the name of the tarantula of
Europe. In this country they occur only in the South or in the far West,
but occasionally living specimens are brought to our northern ports in
shipments of bananas and other tropical produce, and are the source of
much alarm. It should be mentioned, however, that the large spider most
frequently found under such circumstances is not a tarantula at all, but
one of the Heteropodidae, or giant crab-spiders, (fig. 8).
[Illustration: 8. The giant crab-spider or banana spider (Heteropoda
venatoria). Natural size. After Comstock.]
In spite of their prominence and the fear which they arouse there are
few accurate data regarding these American tarantulas. It has often been
shown experimentally that they can kill small birds and mammals, though
it is doubtful if these form the normal prey of any of the species, as
has been claimed. There is no question but that the mere mechanical
injury which they may inflict, and the consequent chances of secondary
infection, justify, in part, their bad reputation. In addition to the
injury from their bite, it is claimed that the body hairs of several of
the South American species are readily detached and are urticating.
Recently, Phisalix (1912) has made a study of the physiological effects
of the venom of two Avicularoidea, _Phormictopus carcerides_ Pocock,
from Haiti and _Cteniza sauvagei_ Rossi, from Corsica. The glands were
removed aseptically and ground up with fine, sterilized sand in
distilled water. The resultant liquid was somewhat viscid, colorless,
and feebly alkaline. Injected into sparrows and mice the extract of
_Phormictopus_ proved very actively poisonous, that from a single spider
being sufficient to kill ten sparrows or twenty mice. It manifested
itself first and, above all, as a narcotic, slightly lowering the
temperature and paralyzing the respiration. Muscular and cardiac
weakening, loss of general sensibility, and the disappearance of
reflexes did not occur until near the end. The extract from _Cteniza_
was less active and, curiously enough, the comparative effect on
sparrows and on mice was just reversed.
SPIDERS OF THE GENUS LATRODECTUS.--While most of the popular accounts of
evil effects from the bites of spiders will not stand investigation, it
is a significant fact that, the world over, the best authenticated
records refer to a group of small and comparatively insignificant
spiders belonging to the genus _Latrodectus_, of the family Theridiidae.
The dread "Malmigniatte" of Corsica and South Europe, the "Karakurte" of
southeastern Russia, the "Katipo" of New Zealand, the "Mena-vodi" and
"Vancoho" of Madagascar, and our own _Latrodectus mactans_, all belong
to this genus, and concerning all of these the most circumstantial
accounts of their venomous nature are given. These accounts are not mere
fantastic stories by uneducated natives but in many cases are reports
from thoroughly trained medical men.
The symptoms produced are general, rather than local. As summarized by
Kobert (1901) from a study of twenty-two cases treated in 1888, in the
Kherson (Russia) Government Hospital and Berislaw (Kherson) District
Hospital the typical case, aside from complications, exhibits the
following symptoms. The victim suddenly feels the bite, like the sting
of a bee. Swelling of the barely reddened spot seldom follows. The
shooting pains, which quickly set in, are not manifested at the point of
injury but localized at the joints of the lower limb and in the region
of the hip. The severity of the pain forces the victim to the hospital,
in spite of the fact that they otherwise have a great abhorrence of it.
The patient is unable to reach the hospital afoot, or, at least, not
without help, for there is usually inability to walk. The patient, even
if he has ridden, reaches the hospital covered with cold sweat and
continues to perspire for a considerable period. His expression
indicates great suffering. The respiration may be somewhat dyspnoeic,
and a feeling of oppression in the region of the heart is common. There
is great aversion to solid food, but increasing thirst for milk and tea.
Retention of urine, and constipation occur. Cathartics and, at night,
strong narcotics are desired. Warm baths give great relief. After three
days, there is marked improvement and usually the patient is dismissed
after the fifth. This summary of symptoms agrees well with other
trustworthy records.
It would seem, then, that Riley and Howard (1889), who discussed a
number of accounts in the entomological literature, were fully justified
in their statement that "It must be admitted that certain spiders of the
genus _Latrodectus_ have the power to inflict poisonous bites, which may
(probably exceptionally and depending upon exceptional conditions) bring
about the death of a human being."
And yet, until recently the evidence bearing on the question has been
most conflicting. The eminent arachnologist, Lucas, (1843) states that
he himself, had been repeatedly bitten by the Malmigniatte without any
bad effects. Dr. Marx, in 1890, gave before the Entomological Society of
Washington, an account of a series of experiments to determine whether
the bite of _Latrodectus mactans_ is poisonous or not. He described the
poison glands as remarkably small[A] and stated that he had introduced
the poison in various ways into guinea-pigs and rabbits without
obtaining any satisfactory results. Obviously, carefully conducted
experiments with the supposed venom were needed and fortunately they
have been carried out in the greatest detail by Kobert (1901).
This investigator pointed out that there were two factors which might
account for the discrepancies in the earlier experiments. In the first
place, the poison of spiders, as of snakes, might be so exhausted after
two or three bites that further bites, following directly, might be
without visible effect. Secondly, the application of the poison by means
of the bite, is exceedingly inexact, since even after the most careful
selection of the point of application, the poison might in one instance
enter a little vein or lymph vessel, and in another case fail to do so.
Besides, there would always remain an incalculable and very large amount
externally, in the nonabsorptive epithelium. While all of these factors
enter into the question of the effect of the bite in specific instances,
they must be as nearly as possible obviated in considering the question
of whether the spiders really secrete a venom harmful to man.
Kobert therefore sought to prepare extracts which would contain the
active principles of the poison and which could be injected in definite
quantities directly into the blood of the experimental animal. For this
purpose various parts of the spiders were rubbed up in a mortar with
distilled water, or physiological salt solution, allowed to stand for an
hour, filtered, and then carefully washed, by adding water drop by drop
for twenty-four hours. The filtrate and the wash-water were then united,
well mixed and, if necessary, cleared by centrifuging or by exposure to
cold. The mixture was again filtered, measured, and used, in part, for
injection and, in part, for the determination of the organic materials.
Such an extract was prepared from the cephalothoraces of eight dried
specimens of the Russian _Latrodectus_ and three cubic centimeters of
this, containing 4.29 mg. of organic material, were injected into the
jugular vein of a cat weighing 2450 grams. The previously very active
animal was paralyzed and lay in whatever position it was placed. The
sensibility of the skin of the extremities and the rump was so reduced
that there was no reaction from cutting or sticking. There quickly
followed dyspnoea, convulsions, paralysis of the respiratory muscles
and of the heart. In twenty-eight minutes the cat was dead, after having
exhibited exactly the symptoms observed in severe cases of poisoning of
man from the bite of this spider.
These experiments were continued on cats, dogs, guinea pigs and various
other animals. Not only extracts from the cephalothorax, but from other
parts of the body, from newly hatched spiders, and from the eggs were
used and all showed a similar virulence. Every effort was made to avoid
sources of error and the experiments, conducted by such a recognized
authority in the field of toxicology, must be accepted as conclusively
showing that this spider and, presumably, other species of the genus
_Latrodectus_ against which the clinical evidence is quite parallel,
possess a poison which paralyzes the heart and central nervous system,
with or without preliminary stimulus of the motor center. If the
quantity of the poison which comes into direct contact with the blood is
large, there may occur haemolysis and thrombosis of the vessels.
On the other hand, check experiments were carried out, using similar
extracts of many common European spiders of the genera _Tegenaria_,
_Drassus_, _Agelena_, _Eucharia_ and _Argyroneta_, as well as the
Russian tarantula, _Lycosa singoriensis_. In no other case was the
effect on experimental animals comparable to the _Latrodectus_ extract.
Kobert concludes that in its chemical nature the poison is neither an
alkaloid, nor a glycoside, nor an acid, but a toxalbumen, or poisonous
enzyme which is very similar to certain other animal poisons, notably
that of the scorpion.
[Illustration: 9. Latrodectus mactans; (_a_) female, x3; (_b_) venter
of female; (_c_) dorsum of male. After Comstock.]
The genus _Latrodectus_ is represented in the United States by at least
two species, _L. mactans_ and _L. geometricus_. Concerning _L. mactans_
there are very circumstantial accounts of serious injury and even death
in man[B]. _Latrodectus mactans_ is coal black, marked with red or
yellow or both. It has eight eyes, which are dissimilar in color and
are distinctly in front of the middle of the thorax, the lateral eyes of
each side widely separate. The tarsus of the fourth pair of legs has a
number of curved setae in a single series. It has on the ventral side of
its abdomen an hour-glass shaped spot. The full-grown female is about
half an inch in length. Its globose abdomen is usually marked with one
or more red spots dorsally along the middle line. The male is about half
as long but has in addition to the dorsal spots, four pairs of stripes
along the sides. Immature females resemble the male in coloring (fig.
9).
Regarding the distribution of _Latrodectus mactans_, Comstock states
that: "Although it is essentially a Southern species, it occurs in
Indiana, Ohio, Pennsylvania, New Hampshire, and doubtless other of the
Northern States." _L. geometricus_ has been reported from California.
OTHER VENOMOUS SPIDERS--While conclusive evidence regarding the venomous
nature of spiders is meager and relates almost wholly to that of the
genus _Latrodectus_, the group is a large one and we are not justified
in dismissing arbitrarily, all accounts of injury from their bites.
Several species stand out as especially needing more detailed
investigation.
_Chiracanthium nutrix_ is a common European species of the family
Clubionidae, concerning which there is much conflicting testimony. Among
the reports are two by distinguished scientists whose accounts of
personal experiences cannot be ignored. A. Forel allowed a spider of
this species to bite him and not only was the pain extreme, but the
general symptoms were so severe that he had to be helped to his house.
The distinguished arachnologist, Bertkau reports that he, himself, was
bitten and that an extreme, burning pain spread almost instantaneously
over the arm and into the breast. There were slight chills the same day
and throbbing pain at the wound lasted for days. While this particular
species is not found in the United States, there are two other
representatives of the genus and it is possible that they possess the
same properties. We are unaware of any direct experimental work on the
poison.
_Epeira diadema_, of Europe, belongs to a wholly different group, that
of the orb-weavers, but has long been reputed venomous. Kobert was able
to prepare from it an extract whose effects were very similar to that
prepared from _Latrodectus_, though feebler in its action. Under
ordinary circumstances this spider is unable to pierce the skin of man
and though Kobert's results seem conclusive, the spider is little to be
feared.
_Phidippus audax_ (_P. tripunctatus_) is one of our largest Attids, or
jumping spiders. The late Dr. O. Lugger describes a case of severe
poisoning from the bite of this spider and though details are lacking,
it is quite possible that this and other large species of the same
group, which stalk their prey, may possess a more active poison than
that of web-building species.
SUMMARY--It is clearly established that our common spiders are not to be
feared and that the stories regarding their virulence are almost wholly
without foundation. On the other hand, the chances of secondary
infection from the bites of some of the more powerful species are not to
be ignored.
Probably all species possess a toxin secreted by the poison gland,
virulent for insects and other normal prey of the spiders, but with
little or no effect on man.
There are a very few species, notably of the genus _Latrodectus_, and
possibly including the European _Chiracanthium nutrix_ and _Epeira
diadema_, which possess, in addition, a toxalbumen derived from the
general body tissue, which is of great virulence and may even cause
death in man and the higher animals.
[Illustration: 10. A whip-scorpion (Mastigoproctus giganteus). Half
natural size. After Comstock.]
THE PEDIPALPIDA OR WHIP-SCORPIONS
The tailed whip-scorpions, belonging to the family Thelyphonidae, are
represented in the United States by the giant whip-scorpion
_Mastigoproctus giganteus_ (fig. 10), which is common in Florida, Texas
and some other parts of the South. In Florida, it is locally known as
the "grampus" or "mule-killer" and is very greatly feared. There is no
evidence that these fears have any foundation, and Dr. Marx states that
there is neither a poison gland nor a pore in the claw of the
chelicera.
THE SCORPIONIDA, OR TRUE SCORPIONS
The true scorpions are widely distributed throughout warm countries and
everywhere bear an evil reputation. According to Comstock (1912), about
a score of species occur in the Southern United States. These are
comparatively small forms but in the tropics members of this group may
reach a length of seven or eight inches. They are pre-eminently
predaceous forms, which lie hidden during the day and seek their prey by
night.
[Illustration: 11. A true scorpion. After Comstock.]
The scorpions (fig. 11) possess large pedipalpi, terminated by strongly
developed claws, or chelae. They may be distinguished from all other
Arachnids by the fact that the distinctly segmented abdomen is divided
into a broad basal region of seven segments and a terminal, slender,
tail-like division of five distinct segments.
The last segment of the abdomen, or telson, terminates in a
ventrally-directed, sharp spine, and contains a pair of highly developed
poison glands. These glands open by two small pores near the tip of the
spine. Most of the species when running carry the tip of the abdomen
bent upward over the back, and the prey, caught and held by the
pedipalpi, is stung by inserting the spine of the telson and allowing it
to remain for a time in the wound.
The glands themselves have been studied in _Prionurus citrinus_ by
Wilson (1904). He found that each gland is covered by a sheet of muscle
on its mesal and dorsal aspects, which may be described as the
_compressor muscle_. The muscle of each side is inserted by its edge
along the ventral inner surface of the chitinous wall of the telson,
close to the middle line, and by a broader insertion laterally. A layer
of fine connective tissue completely envelops each gland and forms the
basis upon which the secreting cells rest. The secreting epithelium is
columnar; and apparently of three different types of cells.
1. The most numerous have the appearance of mucous cells, resembling the
goblet cells of columnar mucous membranes. The nucleus, surrounded by a
small quantity of protoplasm staining with haematoxylin, lies close to
the base of the cell.
2. Cells present in considerable numbers, the peripheral portions of
which are filled with very numerous fine granules, staining with acid
dyes such as methyl orange.
3. Cells few in number, filled with very large granules, or irregular
masses of a substance staining with haematoxylin.
The poison, according to Kobert (1893), is a limpid, acid-reacting
fluid, soluble in water but insoluble in absolute alcohol and ether.
There are few data relative to its chemical nature. Wilson (1901) states
that a common Egyptian species, _Buthus quinquestriatus_, has a specific
gravity of 1.092, and contains 20.3% of solids and 8.4% ash.
The venom of different species appears to differ not only quantitatively
but qualitatively. The effects of the bite of the smaller species of the
Southern United States may be painful but there is no satisfactory
evidence that it is ever fatal. On the other hand, certain tropical
species are exceedingly virulent and cases of death of man from the bite
are common.
In the case of _Buthus quinquestriatus_, Wilson (1904) found the
symptoms in animals to be hypersecretion, salivation and lachrymation,
especially marked, convulsions followed by prolonged muscular spasm;
death from asphyxia. The temperature shows a slight, rarely
considerable, rise. Rapid and considerable increase of blood-pressure
(observed in dogs) is followed by a gradual fall with slowing of the
heart-beat. The coagulability of the blood is not affected.
An interesting phase of Wilson's work was the experiments on desert
mammals. The condition under which these animals exist must frequently
bring them in contact with scorpions, and he found that they possess a
degree of immunity to the venom sufficient at least to protect them from
the fatal effects of the sting.
As far as concerns its effect on man, Wilson found that much depended
upon the age. As high as 60 per cent of the cases of children under
five, resulted fatally. Caroroz (1865), states that in a Mexican state
of 15,000 inhabitants, the scorpions were so abundant and so much feared
that the authorities offered a bounty for their destruction. A result
was a large number of fatalities, over two hundred per year. Most of the
victims were children who had attempted to collect the scorpions.
The treatment usually employed in the case of bites by the more
poisonous forms is similar to that for the bite of venomous snakes.
First, a tight ligature is applied above the wound so as to stop the
flow of blood and lymph from that region. The wound is then freely
excised and treated with a strong solution of permanganate of potash, or
with lead and opium lotion.
In recent years there have been many attempts to prepare an antivenom,
or antiserum comparable to what has been used so effectively in the case
of snake bites. The most promising of these is that of Todd (1909),
produced by the immunization of suitable animals. This antivenom proved
capable of neutralizing the venom when mixed _in vitro_ and also acts
both prophylactically and curatively in animals. Employed curatively in
man, it appears to have a very marked effect on the intense pain
following the sting, and the evidence so far indicates that its prompt
use greatly reduces the chance of fatal results.
THE SOLPUGIDA, OR SOLPUGIDS
The SOLPUGIDA are peculiar spider-like forms which are distinguished
from nearly all other arachnids by the fact that they possess no true
cephalothorax, the last two leg-bearing segments being distinct,
resembling those of the abdomen in this respect. The first pair of legs
is not used in locomotion but seemingly functions as a second pair of
pedipalpi. Figure 12 illustrates the striking peculiarities of the
group. They are primarily desert forms and occur in the warm zones of
all countries. Of the two hundred or more species, Comstock lists twelve
as occurring in our fauna. These occur primarily in the southwest.
[Illustration: 12. A solpugid (Eremobates cinerea). After Comstock.]
The Solpugida have long borne a bad reputation and, regarding virulence,
have been classed with the scorpions. Among the effects of their bites
have been described painful swelling, gangrene, loss of speech, cramps,
delirium, unconsciousness and even death. Opposed to the numerous loose
accounts of poisoning, there are a number of careful records by
physicians and zoologists which indicate clearly that the effects are
local and though they may be severe, they show not the slightest symptom
of direct poisoning.
More important in the consideration of the question is the fact that
there are neither poison glands nor pores in the fangs for the exit of
any poisonous secretion. This is the testimony of a number of prominent
zoologists, among whom is Dr. A. Walter, who wrote to Kobert at length
on the subject and whose conclusions are presented by him.
However, it should be noted that the fangs are very powerful and are
used in such a manner that they may inflict especially severe wounds.
Thus, there may be more opportunity for secondary infection than is
usual in the case of insect wounds.
The treatment of the bite of the Solpugida is, therefore, a matter of
preventing infection. The wound should be allowed to bleed freely and
then washed out with a 1:3000 solution of corrosive sublimate, and, if
severe, a wet dressing of this should be applied. If infection takes
place, it should be treated in the usual manner, regardless of its
origin.
THE ACARINA, OR MITES AND TICKS
A number of the parasitic Acarina evidently secrete a specific poison,
presumably carried by the saliva, but in most cases its effect on man is
insignificant. There is an abundant literature dealing with the
poisonous effect of the bite of these forms, especially the ticks, but
until recently it has been confused by failure to recognize that various
species may transmit diseases of man, rather than produce injury through
direct poisoning. We shall therefore discuss the Acarina more especially
in subsequent chapters, dealing with parasitism and with disease
transmission.
Nevertheless, after the evidence is sifted, there can be no doubt that
the bites of certain ticks may occasionally be followed by a direct
poisoning, which may be either local or general in its effects. Nuttall
(1908) was unable to determine the cause of the toxic effect, for, in
_Argas persicus_, the species most often implicated, he failed to get
the slightest local or general effect on experimental animals, from the
injection of an emulsion prepared by crushing three of the ticks.
It seems clearly established that the bite of certain ticks may cause a
temporary paralysis, or even complete paralysis, involving the organs of
respiration or the heart, and causing death. In 1912, Dr. I. U. Temple,
of Pendleton, Oregon, reported several cases of what he called "acute
ascending paralysis" associated with the occurrence of ticks on the head
or the back of the neck. A typical severe case was that of a six year
old child, who had retired in her usual normal health. The following
morning upon arising she was unable to stand on her feet. She exhibited
paralysis extending to the knees, slight temperature, no pain, sensory
nerves normal, motor nerves completely paralyzed, reflexes absent. The
following day the paralysis had extended to the upper limbs, and before
night of the third day the nerves of the throat (hypoglossal) were
affected. The thorax and larynx were involved, breathing was labored,
she was unable to swallow liquids, phonation was impossible and she
could only make low, guttural sounds. At this stage, two ticks, fully
distended with blood, were found over the junction of the spinal column
with the occipital bones in the hollow depression. They were removed by
the application of undiluted creoline. Though the child's life was
despaired of, by the following morning she was very much improved. By
evening she was able to speak. The paralysis gradually receded,
remaining longest in the feet, and at the end of one week the patient
was able to go home.
There was some doubt as to the exact species of tick implicated in the
cases which Dr. Temple reported, although the evidence pointed strongly
to _Dermacentor venustus_.[C] Somewhat later, Hadwen (1913) reported
that "tick paralysis" occurs in British Columbia, where it affects not
only man, but sheep and probably other animals. It is caused by the
bites of _Dermacentor venustus_ and was experimentally produced in lambs
and a dog (Hadwen and Nuttall, 1913). It is only when the tick begins to
engorge or feed rapidly, some days after it has become attached, that
its saliva produces pathogenic effects.
Ulceration following tick bite is not uncommon. In many of the instances
it is due to the file-like hypostome, with its recurved teeth, being
left in the wound when the tick is forcibly pulled off.
THE MYRIAPODA, OR CENTIPEDES AND MILLIPEDES
The old class, Myriapoda includes the DIPLOPODA, or millipedes, and the
CHILOPODA, or centipedes. The present tendency is to raise these groups
to the rank of classes.
The Diplopoda
The DIPLOPODA, or millipedes (fig. 13), are characterized by the
presence of two pairs of legs to a segment. The largest of our local
myriapods belong to this group. They live in moist places, feeding
primarily on decaying vegetable matter, though a few species
occasionally attack growing plants.
[Illustration: 13. A millipede. After Comstock.]
The millipedes are inoffensive and harmless. _Julus terrestris_, and
related species, when irritated pour out over the entire body a
yellowish secretion which escapes from cutaneous glands. It is volatile,
with a pungent odor, and Phisalix (1900) has shown that it is an active
poison when injected into the blood of experimental animals. This,
however, does not entitle them to be considered as poisonous arthropods,
in the sense of this chapter, any more than the toad can be considered
poisonous to man because it secretes a venom from its cutaneous glands.
The Chilopoda
[Illustration: 14. Two common centipedes.
(_a_) Lithobius forficatus. After Comstock.
(_b_) Scutigera forceps. Natural size; after Howard.]
The CHILOPODA, or centipedes (fig. 14), unlike the millipedes, are
predaceous forms, and possess well developed poison glands for killing
their prey. These glands are at the base of the first pair of legs
(fig. 15), which are bent forward so as to be used in holding their
prey. The legs terminate in a powerful claw, at the tip of which is the
outlet of the poison glands.
The poison is a limpid, homogeneous, slightly acid fluid, which
precipitates in distilled water. Briot (1904) extracted it from the
glands of _Scolopendra morsitans_, a species common in central France,
and found that it was actively venomous for the ordinary experimental
animals. A rabbit of two kilograms weight received an injection of three
cubic centimeters in the vein of the ear and died in a minute. A white
rat, weighing forty-eight grams, received one and a half cubic
centimeters in the hind leg. There was an almost immediate paralysis of
the leg and marked necrosis of the tissues.
[Illustration: 15. Mandible of Scolopendra cingulata showing venom
gland. After Dubosq.]
As for the effect on man, there is little foundation for the fear with
which centipedes are regarded. Our native species produce, at most,
local symptoms,--sometimes severe local pain and swelling,--but there is
no authentic record of fatal results. In the tropics, some of the
species attain a large size, _Scolopendra gigantea_ reaching a length of
nearly a foot. These forms are justly feared, and there is good evidence
that death sometimes, though rarely, results from their bite.
One of the most careful accounts of death from the sting of the scorpion
is that of Linnell, (1914), which relates to a comparatively small
Malayan species, unfortunately undetermined. The patient, a coolie, aged
twenty, was admitted to a hospital after having been stung two days
previously on the left heel. For cure, the other coolies had made him
eat the head of the scorpion. On admission, the patient complained of
"things creeping all over the body". Temp. 102.8^o. On the fourth day he
had paralysis of the legs, and on the fifth day motor paralysis to the
umbilicus, sensation being unaltered. On the sixth day there was
retention of the urine and on the ninth day (first test after third day)
sugar was present. On the thirteenth day the patient became comatose,
but could be roused to eat and drink. The temperature on the following
day fell below 95^o and the patient was still comatose. Death fifteenth
day.
Examination of the spinal (lumbar) cord showed acute disseminated
myelitis. In one part there was an acute destruction of the anterior
horn and an infiltration of round cells. In another portion Clarke's
column had been destroyed. The perivascular sheaths were crowded with
small round cells and the meninges were congested. Some of the cells of
the anterior horn were swollen and the nuclei eccentric; chromatolysis
had occurred in many of them.
As for treatment, Castellani and Chalmers (1910), recommend bathing the
part well with a solution of ammonia (one in five, or one in ten). After
bathing, apply a dressing of the same alkali or, if there is much
swelling and redness, an ice-bag. If necessary, hypodermic injections of
morphine may be given to relieve the pain. At a later period
fomentations may be required to reduce the local inflammation.
THE HEXAPODA OR TRUE INSECTS
There are a number of HEXAPODA, or true insects, which are, in one way
or another, poisonous to man. These belong primarily to the orders
Hemiptera, or true bugs; Lepidoptera, or butterflies and moths (larval
forms); Diptera, or flies; Coleoptera, or beetles; and Hymenoptera, or
ants, bees, and wasps. There are various ways in which they may be
poisonous.
1. _Piercing_ or _biting_ forms may inject an irritating or poisonous
saliva into the wound caused by their mouth-parts.
2. _Stinging forms_ may inject a poison, from glands at the caudal end
of the abdomen, into wounds produced by a specially modified ovipositer,
the _sting_.
3. _Nettling_ properties may be possessed by the hairs of the insect.
4. _Vescicating_, or _poisonous blood plasma_, or _body fluids_ are
known to exist in a large number of species and may, under exceptional
circumstances, affect man.
For convenience of discussion, we shall consider poisonous insects under
these various headings. In this, as in the preceding discussion, no
attempt will be made to give an exhaustive list of the poisonous forms.
Typical instances will be selected and these will be chosen largely from
North American species.
PIERCING OR BITING INSECTS POISONOUS TO MAN
HEMIPTERA
Several families of the true bugs include forms which, while normally
inoffensive, are capable of inflicting painful wounds on man. In these,
as in all of the Hemiptera, the mouth-parts are modified to form an
organ for piercing and sucking. This is well shown by the accompanying
illustration (fig. 16).
The upper lip, or _labrum_, is much reduced and immovable, the lower
lip, or _labium_, is elongated to form a jointed sheath, within which
the lance-like mandibles and maxillae are enclosed. The mandibles are
more or less deeply serrate, depending on the species concerned.
[Illustration: 16. Beak of hemipteron.]
The poison is elaborated by the salivary glands, excepting, possibly, in
_Belostoma_ where Locy is inclined to believe that it is secreted by the
maxillary glands. The salivary glands of the Hemiptera have been the
subject of much study but the most recent, comprehensive work has been
done by Bugnion and Popoff, (1908 and 1910) to whose text the reader is
referred for details.
The Hemiptera have two pairs of salivary glands: the _primary gland_, of
which the efferent duct leads to the salivary syringe, and the
_accessory gland_, of which the very long and flexuous duct empties into
the primary duct at its point of insertion. Thus, when one observes the
isolated primary gland it appears as though it had efferent ducts
inserted at the same point. In _Nepa_ and the _Fulgoridae_ there are two
accessory glands and therefore apparently three ducts at the same point
on the primary gland. The _ensemble_ differs greatly in appearance in
different species but we shall show here Bugnion and Popoff's figure of
the apparatus of _Notonecta maculata_, a species capable of inflicting a
painful bite on man (fig. 17).
[Illustration: 17. Salivary glands of Notonecta maculata. After Bugnion
and Popoff.]
[Illustration: 18. Pharyngeal syringe or salivary pump of Fulgora
maculata. After Bugnion and Popoff.]
[Illustration: 19. Heteroptera, (_a_) Melanolestes picipes; (_b_)
Notonecta undulata; (_c_, _d_) Aradus robustus (_c_) adult, (_d_) nymph,
much enlarged; (_e_) Arilus cristatus; (_f_) Belostoma americana; (_g_)
Nabis (Coriscus) subcoleoptratus, enlarged; (_h_) Cimex lectularius,
(_i_) Oeciacus vicarius, much enlarged; (_j_) Lyctocoris fitchii, much
enlarged. After Lugger.]
Accessory to the salivary apparatus there is on the ventral side of the
head, underneath the pharynx, a peculiar organ which the Germans have
called the "Wanzenspritze," or syringe. The accompanying figure of the
structure in _Fulgora maculata_ (fig. 18) shows its relation to the
ducts of the salivary glands and to the beak. It is made up of a
dilatation forming the body of the pump, in which there is a chitinous
piston. Attached to the piston is a strong retractor muscle. The
function of the salivary pump is to suck up the saliva from the salivary
ducts and to force it out through the beak.
Of the Hemiptera reported as attacking man, we shall consider briefly
the forms most frequently noted.
The NOTONECTIDAE, or _back swimmers_, (fig. 19_b_) are small, aquatic
bugs that differ from all others in that they always swim on their
backs. They are predaceous; feeding on insects and other small forms.
When handled carelessly they are able to inflict a painful bite, which
is sometimes as severe as the sting of a bee. In fact, they are known in
Germany as "Wasserbienen."
The BELOSTOMATIDAE, or _giant water bugs_, (fig. 19_f_) include the
largest living Hemiptera. They are attracted to lights and on account of
the large numbers which swarm about the electric street lamps in some
localities they have received the popular name "electric light bugs."
Our largest representatives in the northern United States belong to the
two genera _Belostoma_ and _Banacus_, distinguished from each other by
the fact that _Belostoma_ has a groove on the under side of the femur of
the front leg, for the reception of the tibia.
The salivary glands of Belostoma were figured by Leidy (1847) and later
were studied in more detail by Locy (1884). There are two pairs of the
glands, those of one pair being long and extending back as far as the
beginning of the abdomen, while the others are about one-fourth as long.
They lie on either side of the oesophagus. On each side of the
oesophagus there is a slender tube with a sigmoid swelling which may
serve as a poison reservoir. In addition to this salivary system, there
is a pair of very prominent glands on the ventral side of the head,
opening just above the base of the beak. These Locy has called the
"cephalic glands" and he suggests that they are the source of the
poison. They are the homologues of the maxillary glands described for
other Hemiptera, and it is by no means clear that they are concerned
with the production of venom. It seems more probable that in
_Belostoma_, as in other Hemiptera, it is produced by the salivary
glands, though the question is an open one.
The Belostomatidae feed not only on insects, but on small frogs, fish,
salamanders and the like. Matheson (1907) has recorded the killing of a
good-sized bird by _Belostoma americana_. A woodpecker, or flicker, was
heard to utter cries of distress, and fluttered and fell from a tree. On
examination it was found that a bug of this species had inserted its
beak into the back part of the skull and was apparently busily engaged
in sucking the blood or brains of the bird. Various species of
_Belostoma_ have been cited as causing painful bites in man. We can
testify from personal experience that the bite of _Belostoma americana_
may almost immediately cause severe, shooting pains that may extend
throughout the arm and that they may be felt for several days.
[Illustration: 20. Reduvius (Opsicoetus) personatus. (x2).]
Relief from the pain may be obtained by the use of dilute ammonia, or a
menthol ointment. In the not uncommon case of secondary infection the
usual treatment for that should be adopted.
[Illustration: 21. (_a_) Reduvius personatus, nymph. Photograph by M. V.
S.]
The REDUVIIDAE, or _assassin-bugs_ are capable of inflicting very painful
wounds, as most collectors of Hemiptera know to their sorrow. Some
species are frequently to be found in houses and outhouses and Dr.
Howard suggests that many of the stories of painful spider bites relate
to the attack of these forms.
[Illustration: 21. (_b_) Reduvius personatus, adult (x2) Photograph by
M. V. S.]
An interesting psychological study was afforded in the summer of 1899,
by the "kissing-bug" scare which swept over the country. It was reported
in the daily papers that a new and deadly bug had made its appearance,
which had the unpleasant habit of choosing the lips or cheeks for its
point of attack on man. So widespread were the stories regarding this
supposedly new insect that station entomologists all over the country
began to receive suspected specimens for identification. At Cornell
there were received, among others, specimens of stone-flies, may-flies
and even small moths, with inquiries as to whether they were
"kissing-bugs."
[Illustration: 22. Rasahus biguttatus. (x2). After Howard.]
Dr. L. O. Howard has shown that the scare had its origin in newspaper
reports of some instances of bites by either _Melanolestes picipes_
(fig. 19a) or _Opsicoetes personatus_ (fig. 20), in the vicinity of
Washington, D. C. He then discusses in considerable detail the more
prominent of the Reduviidae which, with greater or less frequency pierce
the skin of human beings. These are _Opsicoetes personatus_,
_Melanolestes picipes_, _Coriscus subcoleoptratus_ (fig. 19_g_),
_Rasahus thoracicus_, _Rasahus biguttatus_ (fig. 22), _Conorhinus
sanguisugus_ (fig. 71), and _Melanolestes abdominalis_ (fig. 23).
[Illustration: 23. Melanolestes abdominalis (x2). After Marlatt.]
One of the most interesting of these species is _Reduvius personatus_,
(= _Opsicoetus personatus_), which is popularly known as the "masked
bed-bug hunter." It owes this name to the fact that the immature nymphs
(fig. 21) have their bodies and legs completely covered by dust and
lint, and that they are supposed to prey upon bed-bugs. LeConte is
quoted by Howard as stating that "This species is remarkable for the
intense pain caused by its bite. I do not know whether it ever willingly
plunges its rostrum into any person, but when caught, or unskilfully
handled it always stings. In this case the pain is almost equal to the
bite of a snake, and the swelling and irritation which result from it
will sometimes last for a week."
A species which very commonly attacks man is _Conorhinus sanguisugus_,
the so-called "big bed-bug" of the south and southern United States. It
is frequently found in houses and is known to inflict an exceedingly
painful bite. As in the case of a number of other predaceous Hemiptera,
the salivary glands of these forms are highly developed. The effect of
the bite on their prey and, as Marlatt has pointed out, the constant and
uniform character of the symptoms in nearly all cases of bites in man,
clearly indicate that their saliva contains a specific substance. No
satisfactory studies of the secretions have been made. On the other
hand, Dr. Howard is doubtless right in maintaining that the very serious
results which sometimes follow the bite are due to the introduction of
extraneous poison germs. This is borne out by the symptoms of most of
the cases cited in literature and also by the fact that treatment with
corrosive sublimate, locally applied to the wound, has yielded favorable
results.
OTHER HEMIPTERA REPORTED AS POISONOUS TO MAN--A large number of other
Hemiptera have been reported as attacking man. Of these, there are
several species of Lygaeidae, Coreidae, and Capsidae. Of the
latter, _Lygus pratensis_, the tarnished plant-bug, is reported by
Professor Crosby as sucking blood. _Orthotylus flavosparsus_ is another
Capsid which has been implicated. _Empoasca mali_ and _Platymetopius
acutus_ of the Jassidae have also been reported as having similar
habits.
Whenever the periodical cicada or "seventeen-year locust" becomes
abundant, the newspapers contain accounts of serious results from its
bites. The senior author has made scores of attempts to induce this
species to bite and only once successfully. At that time the bite was in
no wise more severe than a pin-prick. A student in our department
reports a similar experience. There is no case on record which bears
evidence of being worthy of any credence, whatsoever.
Under the heading of poisonous Hemiptera we might consider the bed-bugs
and the lice. These will be discussed later, as parasites and as
carriers of disease, and therefore need only be mentioned here.
DIPTERA
Several species of blood-sucking Diptera undoubtedly secrete a saliva
possessing poisonous properties. Chief among these are the Culicidae, or
mosquitoes, and the Simuliidae, or black-flies. As we shall consider
these forms in detail under the heading of parasitic species and
insects transmitting disease, we shall discuss here only the poison of
the mosquitoes.
It is well known that mosquitoes, when they bite, inject into the wound
a minute quantity of poison. The effect of this varies according to the
species of mosquito and also depends very much on the susceptibility of
the individual. Soon after the bite a sensation of itching is noticed
and often a wheal, or eminence, is produced on the skin, which may
increase to a considerable swelling. The scratching which is induced may
cause a secondary infection and thus lead to serious results. Some
people seem to acquire an immunity against the poison.
The purpose of this irritating fluid may be, as Reaumur suggested, to
prevent the coagulation of the blood and thus not only to cause it to
flow freely when the insect bites but to prevent its rapid coagulation
in the stomach. Obviously, it is not developed as a protective fluid,
and its presence subjects the group to the additional handicap of the
vengeance of man.
[Illustration: 24. Diagram of a longitudinal section of a mosquito.]
As to the origin of the poison, there has been little question, until
recent years, that it was a secretion from the salivary glands.
Macloskie (1888) showed that each gland is subdivided into three lobes,
the middle of which differs from the others in having evenly granulated
contents and staining more deeply than the others (fig. 24). This middle
lobe he regarded as the source of the poison. Bruck, (1911), by the use
of water, glycerine, chloroform, and other fluids, extracted from the
bodies of a large number of mosquitoes a toxine which he calls
_culicin_. This he assumes comes from the salivary glands. Animal
experimentation showed that this extract possessed hemolytic powers.
Inoculated into the experimenter's own skin it produced lesions which
behaved exactly as do those of mosquito bites.
Similarly, most writers on the subject have concurred with the view that
the salivary glands are the source of the poison. However, recent work,
especially that of Nuttall and Shipley (1903), and Schaudinn (1904), has
shown that the evidence is by no means conclusive. Nuttall dissected out
six sets (thirty-six acini) of glands from freshly killed _Culex
pipiens_ and placed them in a drop of salt solution. The drop was
allowed to dry, it being thought that the salt crystals would facilitate
the grinding up of the glands with the end of a small glass rod, this
being done under microscopic control. After grinding up, a small drop of
water was added of the size of the original drop of saline, and an equal
volume of human blood taken from the clean finger-tip was quickly mixed
therewith, and the whole drawn up into a capillary tube. Clotting was
not prevented and no hemolysis occurred. Salivary gland emulsion added
to a dilute suspension of corpuscles did not lead to hemolysis. This
experiment was repeated a number of times, with slight modification, but
with similar results. The data obtained from the series "do not support
the hypothesis that the salivary glands, at any rate in _Culex pipiens_,
contain a substance which prevents coagulation."
Much more detailed, and the more important experiments made along this
line, are those of Schaudinn (1904). The results of these experiments
were published in connection with a technical paper on the alternation
of generations and of hosts in _Trypanosoma_ and _Spirochaeta_, and for
this reason seem to have largely escaped the notice of entomologists.
They are so suggestive that we shall refer to them in some detail.
Schaudinn observed that the three oesophageal diverticula (commonly,
but incorrectly, known as the "sucking stomach") (fig. 24) usually
contain large bubbles of gas and in addition, he always found yeast
cells. On the ground of numerous observations, Schaudinn was convinced
that these yeast plants are normal and constant commensals of the
insect. He regarded them as the cause of the gas bubbles to be found in
diverticula. It was found that as the insect fed, from time to time the
abdomen underwent convulsive contractions which resulted in the emptying
of the oesophageal diverticula and the salivary glands through blood
pressure.
In order to test the supposed toxic action of the salivary glands,
Schaudinn repeatedly introduced them under his skin and that of his
assistant, in a drop of salt solution, and never obtained a suggestion
of the irritation following a bite of the insect, even though the glands
were carefully rubbed to fragments after their implantation. Like
Nuttall, he failed to get satisfactory evidence that the secretion of
the salivary glands retarded coagulation of the blood.
He then carefully removed the oesophageal diverticula with their
content of yeast and introduced them into an opening in the skin of the
hand. Within a few seconds there was noticeable the characteristic
itching irritation of the mosquito bite; and in a short time there
appeared reddening and typical swelling. This was usually much more
severe than after the usual mosquito bite, and the swelling persisted
and itched longer. This was because by the ordinary bite of the mosquito
most of the yeast cells are again sucked up, while in these experiments
they remained in the wound. These experiments were repeated a number of
times on himself, his assistant and others, and always with the same
result. From them Schaudinn decided that the poisonous action of the
mosquito bite is caused by an enzyme from a commensal fungus. These
conclusions have not, as yet, been satisfactorily tested.
Relief from the effect of the mosquito bite may be obtained by bathing
the swellings with weak ammonia or, according to Howard, by using moist
soap. The latter is to be rubbed gently on the puncture and is said to
speedily allay the irritation. Howard also quotes from the _Journal of
Tropical Medicine and Hygiene_ to the effect that a few drops of a
solution of thirty to forty grains of iodine to an ounce of saponated
petroleum rubbed into the mosquito bite, or wasp sting, allay the pain
instantaneously.
Methods of mosquito control will be discussed later, in considering
these insects as parasites and as carriers of disease.
STINGING INSECTS
The stinging insects all belong to the order HYMENOPTERA. In a number of
families of this group the ovipositor is modified to form a sting and is
connected with poison-secreting glands. We shall consider the apparatus
of the honey-bee and then make briefer reference to that of other forms.
APIS MELLIFICA, THE HONEY BEE--The sting of the worker honey-bee is
situated within a so-called sting chamber at the end of the abdomen.
This chamber is produced by the infolding of the greatly reduced and
modified eighth, ninth and tenth abdominal segments into the seventh.[D]
From it the dart-like sting can be quickly exserted.
The sting (fig. 25) is made up of a central shaft, ventro-laterad of
which are the paired _lancets_, or darts, which are provided with sharp,
recurved teeth. Still further laterad lie the paired whitish,
finger-like _sting palpi_. Comparative morphological as well as
embryological studies have clearly established that these three parts
correspond to the three pairs of gonopophyses of the ovipositor of more
generalized insects.
[Illustration: 25. Sting of a honey bee. _Psn Sc_, base of acid poison
gland; _B Gl_, alkaline poison gland; _Stn Plp_, sting palpi; _Sh B_,
bulb of sting; _Sh A_, basal arm; _Lct_, lancets or darts; _Sh s_, shaft
of sting. Modified from Snodgrass.]
[Illustration: 26. Poison apparatus of a honey bee. Modified from
Snodgrass.]
An examination of the internal structures (fig. 26) reveals two distinct
types of poison glands, the acid-secreting and the alkaline-secreting
glands, and a prominent poison reservoir. In addition, there is a small
pair of accessory structures which have been called lubricating glands,
on account of the supposed function of their product. The acid-secreting
gland empties into the distal end of the poison reservoir which in turn
pours the secretion into the muscular bulb-like enlargement at the base
of the shaft. The alkaline secreting gland empties into the bulb ventrad
of the narrow neck of the reservoir.
The poison is usually referred to as formic acid. That it is not so
easily explained has been repeatedly shown and is evidenced by the
presence of the two types of glands. Carlet maintains that the product
of either gland is in itself innocent,--it is only when they are
combined that the toxic properties appear.
The most detailed study of the poison of the honey-bee is that of Josef
Langer (1897), who in the course of his work used some 25,000 bees.
Various methods of obtaining the active poison for experimental purposes
were used. For obtaining the pure secretion, bees were held in the
fingers and compressed until the sting was exserted, when a clear drop
of the poison was visible at its tip. This was then taken up in a
capillary tube or dilute solutions obtained by dipping the tip of the
sting into a definite amount of distilled water.
An aqueous solution of the poison was more readily obtained by pulling
out the sting and poison sacs by means of forceps, and grinding them up
in water. The somewhat clouded fluid was then filtered one or more
times. For obtaining still greater quantities, advantage was taken of
the fact that while alcohol coagulates the poison, the active principle
remains soluble in water. Hence the stings with the annexed glands where
collected in 96 per cent alcohol, after filtering off of the alcohol
were dried at 40^o C., then rubbed to a fine powder and this was
repeatedly extracted with water. Through filtering of this aqueous
extract there was obtained a yellowish-brown fluid which produced the
typical reactions, according to concentration of the poison.
The freshly expelled drop of poison is limpid, of distinct acid
reaction, tastes bitter and has a delicate aromatic odor. On
evaporation, it leaves a sticky residue, which at 100 degrees becomes
fissured, and suggests dried gum arabic. The poison is readily soluble
in water and possesses a specific gravity of 1.1313. On drying at room
temperature, it leaves a residue of 30 per cent, which has not lost in
poisonous action or in solubility. In spite of extended experiments,
Langer was unable to determine the nature of the active principle. He
showed that it was not, as had been supposed, an albuminous body, but
rather an organic base.
The pure poison, or the two per cent aqueous solution, placed on the
uninjured skin showed absolutely no irritating effect, though it
produced a marked reaction on the mucus membrane of the nose or eye. A
single drop of one-tenth per cent aqueous solution of the poison brought
about a typical irritation in the conjunctiva of the rabbit's eye. On
the other hand, the application of a drop of the poison, or its
solution, to the slightest break in the skin, or by means of a needle
piercing the skin, produced typical effects. There is produced a local
necrosis, in the neighborhood of which there is infiltration of
lymphocytes, oedema, and hyperaemia.
The effect of the sting on man (fig. 27) is usually transitory but there
are some individuals who are made sick for hours, by a single sting.
Much depends, too, on the place struck. It is a common experience that
an angry bee will attempt to reach the eye of its victim and a sting on
the lid may result in severe and prolonged swelling. In the case of a
man stung on the cheek, Legiehn observed complete aphonia and a breaking
out of red blotches all over the body. A sting on the tongue has been
known to cause such collateral oedema as to endanger life through
suffocation. Cases of death of man from the attacks of bees are rare but
are not unknown. Such results are usually from a number of stings but,
rarely, death has been known to follow a single sting, entering a blood
vessel of a particularly susceptible individual.
[Illustration: Effect of bee stings. After Root.]
It is clearly established that partial immunity from the effects of the
poison may be acquired. By repeated injections of the venom, mice have
been rendered capable of bearing doses that certainly would have killed
them at first. It is a well-known fact that most bee-keepers become
gradually hardened to the stings, so that the irritation and the
swelling become less and less. Some individuals have found this immunity
a temporary one, to be reacquired each season. A striking case of
acquired immunity is related by the Roots in their "A B C and X Y Z of
Bee Culture." The evidence in the case is so clear that it should be
made more widely available and hence we quote it here.
A young man who was determined to become a bee-keeper, was so
susceptible to the poison that he was most seriously affected by a
single sting, his body breaking out with red blotches, breathing growing
difficult, and his heart action being painfully accelerated. "We finally
suggested taking a live bee and pressing it on the back of his hand
until it merely pierced his skin with the sting, then immediately
brushing off both bee and sting. This was done and since no serious
effect followed, it was repeated inside of four or five days. This was
continued for some three or four weeks, when the patient began to have a
sort of itching sensation all over his body. The hypodermic injections
of bee-sting poison were then discontinued. At the end of a month they
were repeated at intervals of four or five days. Again, after two or
three weeks the itching sensation came on, but it was less pronounced.
The patient was given a rest of about a month, when the doses were
repeated as before." By this course of treatment the young man became so
thoroughly immunized that neither unpleasant results nor swelling
followed the attacks of the insects and he is able to handle bees with
the same freedom that any experienced bee-keeper does.
In an interesting article in the _Entomological News_ for November,
1914, J. H. Lovell calls attention to the fact that "There has been a
widespread belief among apiarists that a beekeeper will receive more
stings when dressed in black than when wearing white clothing. A large
amount of evidence has been published in the various bee journals
showing beyond question that honey-bees under certain conditions
discriminate against black. A few instances may be cited in
illustration. Of a flock of twelve chickens running in a bee-yard seven
black ones were stung to death, while five light colored ones escaped
uninjured. A white dog ran among the bee-hives without attracting much
attention, while at the same time a black dog was furiously assailed by
the bees. Mr. J. D. Byer, a prominent Canadian beekeeper, relates that a
black and white cow, tethered about forty feet from an apiary, was one
afternoon attacked and badly stung by bees. On examination it was found
that the black spots had five or six stings to one on the white. All
noticed this fact, although no one was able to offer any explanation. A
white horse is in much less danger of being stung, when driven near an
apiary, than a black one. It has, indeed, been observed repeatedly that
domestic animals of all kinds, if wholly or partially black, are much
more liable to be attacked by bees, if they wander among the hives, than
those which are entirely white."
In order to test the matter experimentally, the following series of
experiments was performed. In the language of the investigator:
"On a clear, warm day in August I dressed wholly in white with the
exception of a black veil. Midway on the sleeve of my right arm there
was sewed a band of black cloth ten inches wide. I then entered the
bee-yard and, removing the cover from one of the hives, lifted a piece
of comb with both hands and gently shook it. Instantly many of the bees
flew to the black band, which they continued to attack as long as they
were disturbed. Not a single bee attempted to sting the left sleeve,
which was of course entirely white, and very few even alighted upon it."
"This experiment was repeated a second, third and fourth time; in each
instance with similar results. I estimated the number of bees on the
band of black cloth at various moments was from thirty to forty; it was
evident from their behavior that they were extremely irritable. To the
left white sleeve and other portions of my clothing they paid very
little attention; but the black veil was very frequently attacked."
"A few days later the experiments were repeated, but the band of black
cloth, ten inches wide, was sewed around my left arm instead of around
the right arm as before. When the bees were disturbed, after the hive
cover had been removed, they fiercely attacked the band of black cloth
as in the previous experiences; but the right white sleeve and the white
suit were scarcely noticed. At one time a part of the black cloth was
almost literally covered with furiously stinging bees, and the black
veil was assailed by hundreds. The bees behaved in a similar manner when
a second hive on the opposite side of the apiary was opened."
"A white veil which had been procured for this purpose, was next
substituted for the black veil. The result was most surprising, for,
whereas in the previous experiments hundreds of bees had attacked the
black veil, so few flew against the white veil as to cause me no
inconvenience. Undoubtedly beekeepers will find it greatly to their
advantage to wear white clothing when working among their colonies of
bees and manipulating the frames of the hives."
When a honey-bee stings, the tip of the abdomen, with the entire sting
apparatus, is torn off and remains in the wound. Here the muscles
continue to contract, for some minutes, forcing the barbs deeper and
deeper into the skin, and forcing out additional poison from the
reservoir.
Treatment, therefore, first consists in removing the sting without
squeezing out additional poison. This is accomplished by lifting and
scraping it out with a knife-blade or the fingernail instead of grasping
and pulling it out. Local application of alkalines, such as weak
ammonia, are often recommended on the assumption that the poison is an
acid to be neutralized on this manner, but these are of little or no
avail. They should certainly not be rubbed in, as that would only
accelerate the absorption of the poison. The use of cloths wrung out in
hot water and applied as hot as can be borne, affords much relief in the
case of severe stings. The application of wet clay, or of the end of a
freshly cut potato is sometimes helpful.
In extreme cases, where there is great susceptibility, or where there
may have been many stings, a physician should be called. He may find
strychnine injections or other treatment necessary, if general symptoms
develop.
[Illustration: 28. The poison apparatus of Formica. Wheeler, after
Forel.]
OTHER STINGING FORMS--Of the five thousand, or more, species of bees,
most possess a sting and poison apparatus and some of the larger forms
are capable of inflicting a much more painful sting than that of the
common honey-bee. In fact, some, like the bumble bees, possess the
advantage that they do not lose the sting from once using it, but are
capable of driving it in repeatedly. In the tropics there are found many
species of stingless bees but these are noted for their united efforts
to drive away intruders by biting. Certain species possess a very
irritating saliva which they inject into the wounds.
The ants are not ordinarily regarded as worthy of consideration under
the heading of "stinging insects" but as a matter of fact, most of them
possess well developed stings and some of them, especially in the
tropics, are very justly feared. Even those which lack the sting possess
well-developed poison glands and the parts of the entire stinging
apparatus, in so far as it is developed in the various species, may
readily be homologized with those of the honey-bee.
The ants lacking a sting are those of the subfamily CAMPONOTINAE, which
includes the largest of our local species. It is an interesting fact
that some of these species possess the largest poison glands and
reservoir (fig. 28) and it is found that when they attack an enemy they
bring the tip of the abdomen forward and spray the poison in such a way
that it is introduced into the wound made by the powerful mandibles.
[Illustration: 29. A harmless, but much feared larva, the "tomato worm."
Natural size. Photograph by M. V. S.]
More feared than any of the other Hymenoptera are the hornets and wasps.
Of these there are many species, some of which attain a large size and
are truly formidable. Phisalix (1897), has made a study of the venom of
the common hornet and finds that, like the poison of the honey-bee, it
is neither an albuminoid nor an alkaloid. Its toxic properties are
destroyed at 120^o C. Phisalix also says that the venom is soluble in
alcohol. If this be true, it differs in this respect from that of the
bee. An interesting phase of the work of Phisalix is that several of her
experiments go to show that the venom of hornets acts as a vaccine
against that of vipers.
NETTLING INSECTS
So far, we have considered insects which possess poison glands connected
with the mouth-parts or a special sting and which actively inject their
poison into man. There remain to be considered those insects which
possess poisonous hairs or body fluids which, under favorable
circumstances, may act as poisons. To the first of these belong
primarily the larvae of certain Lepidoptera.
LEPIDOPTERA
[Illustration: 30. Another innocent but much maligned caterpillar, the
larva of the Regal moth. Photograph by M. V. S.]
When we consider the reputedly poisonous larvae of moths and butterflies,
one of the first things to impress us is that we cannot judge by mere
appearance. Various species of Sphingid, or hawk-moth larvae, bear at the
end of the body a chitinous horn, which is often referred to as a
"sting" and regarded as capable of inflicting dangerous wounds. It would
seem unnecessary to refer to this absurd belief if it were not that each
summer the newspapers contain supposed accounts of injury from the
"tomato worm" (fig. 29) and others of this group. The grotesque, spiny
larva (fig. 30) of one of our largest moths, _Citheronia regalis_ is
much feared though perfectly harmless, and similar instances could be
multiplied.
[Illustration: 31. The flannel moth (Lagoa crispata). (_a_) Poisonous
larva.]
[Illustration: 31. (_b_) Adult. Enlarged. Photographs by M. V. S.]
But if the larvae are often misjudged on account of their ferocious
appearance, the reverse may be true. A group of most innocent looking
and attractive caterpillars is that of the flannel-moth larvae, of which
_Lagoa crispata_ may be taken as an example. Its larva (fig. 31) has a
very short and thick body, which is fleshy and completely covered and
hidden by long silken hairs of a tawny or brown color, giving a convex
form to the upper side. Interspersed among these long hairs are
numerous short spines connected with underlying hypodermal poison
glands. These hairs are capable of producing a marked nettling effect
when they come in contact with the skin. This species is found in our
Atlantic and Southern States. Satisfactory studies of its poisonous
hairs and their glands have not yet been made.
[Illustration: 32 The poisonous saddle back caterpillar. Empretia
(Sibine) stimulea. Photograph by M. V. S.]
[Illustration: 33_a_. Io moth larvae on willow. Photograph by M. V. S.]
_Sibine stimulea (Empretia stimulea)_, or the saddle-back caterpillar
(fig. 32), is another which possesses nettling hairs. This species
belongs to the group of Eucleidae, or slug caterpillars. It can be
readily recognized by its flattened form, lateral, bristling spines and
by the large green patch on the back resembling a saddle-cloth, while
the saddle is represented by an oval, purplish-brown spot. The small
spines are venomous and affect some persons very painfully. The larva
feeds on the leaves of a large variety of forest trees and also on
cherry, plum, and even corn leaves. It is to be found throughout the
Eastern and Southern United States.
[Illustration: 33_b_. Io moth. Full grown larva. Photograph by M. V. S.]
[Illustration: 34. Io moth. Adult. Photograph by M. V. S.]
_Automeris io_ is the best known of the nettling caterpillars. It is the
larva of the Io moth, one of the Saturniidae. The mature caterpillar,
(fig. 33), which reaches a length of two and one-half inches, is of a
beautiful pale green with sublateral stripes of cream and red color and
a few black spines among the green ones. The green radiating spines give
the body a mossy appearance. They are tipped with a slender chitinous
hair whose tip is readily broken off in the skin and whose poisonous
content causes great irritation. Some individuals are very susceptible
to the poison, while others are able to handle the larvae freely without
any discomfort. The larvae feed on a wide range of food plants. They are
most commonly encountered on corn and on willow, because of the
opportunities for coming in contact with them.
[Illustration: 35. Larva of brown-tail moth. (Natural size). Photograph
by M. V. S.]
The larvae of the brown-tail moth (_Euproctis chrysorrhoea_) (fig. 35
and 36), where they occur in this country, are, on account of their
great numbers, the most serious of all poisonous caterpillars. It is not
necessary here, to go into details regarding the introduction of this
species from Europe into the New England States. This is all available
in the literature from the United States Bureau of Entomology and from
that of the various states which are fighting the species. Suffice to
say, there is every prospect that the pest will continue to spread
throughout the Eastern United States and Canada and that wherever it
goes it will prove a direct pest to man as well as to his plants.
Very soon after the introduction of the species there occurred in the
region where it had gained a foothold, a mysterious dermatitis of man.
The breaking out which usually occurred on the neck or other exposed
part of the body was always accompanied by an intense itching. It was
soon found that this dermatitis was caused by certain short, barbed
hairs of the brown-tail caterpillars and that not only the caterpillars
but their cocoons and even the adult female moths might harbor these
nettling hairs and thus give rise to the irritation. In many cases the
hairs were wafted to clothing on the line and when this was worn it
might cause the same trouble. Still worse, it was found that very
serious internal injury was often caused by breathing or swallowing the
poisonous hairs.
[Illustration: 36. Browntail moths. One male and two females. Photograph
by M. V. S.]
The earlier studies seemed to indicate that the irritation was purely
mechanical in origin, the result of the minute barbed hairs working into
the skin in large numbers. Subsequently, however, Dr. Tyzzer (1907)
demonstrated beyond question that the trouble was due to a poison
contained in the hairs. In the first place, it is only the peculiar
short barbed hairs which will produce the dermatitis when rubbed on the
skin, although most of the other hairs are sharply barbed. Moreover, it
was found that in various ways the nettling properties could be
destroyed without modifying the structure of the hairs. This was
accomplished by baking for one hour at 110^o C, by warming to 60^o C in
distilled water, or by soaking in one per cent. or in one-tenth per
cent. of potassium hydrate or sodium hydrate. The most significant part
of his work was the demonstration of the fact that if the nettling
hairs are mingled with blood, they immediately produce a change in the
red corpuscles. These at once become coarsely crenated, and the roleaux
are broken up in the vicinity of the hair (fig. 37_b_). The corpuscles
decrease in size, the coarse crenations are transformed into slender
spines which rapidly disappear, leaving the corpuscles in the form of
spheres, the light refraction of which contrasts them sharply with the
normal corpuscles. The reaction always begins at the basal sharp point
of the hair. It could not be produced by purely mechanical means, such
as the mingling of minute particles of glass wool, the barbed hairs of a
tussock moth, or the other coarser hairs of the brown-tail, with the
blood.
[Illustration: 37. (_a_) Ordinary hairs and three poison hairs of
subdorsal and lateral tubercles of the larva of the browntail moth.
Drawing by Miss Kephart.]
The question of the source of the poison has been studied in our
laboratory by Miss Cornelia Kephart. She first confirmed Dr. Tyzzer's
general results and then studied carefully fixed specimens of the larvae
to determine the distribution of the hairs and their relation to the
underlying tissues.
[Illustration: 37. (_b_) Effect of the poison on the blood corpuscles of
man. After Tyzzer.]
The poison hairs (fig. 37), are found on the subdorsal and lateral
tubercles (fig. 38), in bunches of from three to twelve on the minute
papillae with which the tubercles are thickly covered. The underlying
hypodermis is very greatly thickened, the cells being three or four
times the length of the ordinary hypodermal cells and being closely
crowded together. Instead of a pore canal through the cuticula for each
individual hair, there is a single pore for each papillae on a tubercle,
all the hairs of the papilla being connected with the underlying cells
through the same pore canal, (figs. 39 and 40).
[Illustration: 38. Cross section of the larva of the browntail moth
showing the tubercles bearing the poison hairs. Drawing by Miss
Kephart.]
The hypodermis of this region is of two distinct types of cells. First,
there is a group of slender fusiform cells, one for each poison hair on
the papilla, which are the trichogen, or hair-formative cells. They are
crowded to one side and towards the basement membrane by a series of
much larger, and more prominent cells (fig. 40), of which there is a
single one for each papilla. These larger cells have a granular
protoplasm with large nuclei and are obviously actively secreting. They
are so characteristic in appearance as to leave no question but that
they are the true poison glands.
[Illustration: 39. Epithelium underlying poison hairs of the larva of
the browntail moth. Drawing by Miss Kephart.]
Poisonous larvae of many other species have been reported from Europe and
especially from the tropics but the above-mentioned species are the more
important of those occurring in the United States and will serve as
types. It should be noted in this connection that through some curious
misunderstanding Goeldi (1913) has featured the larva of _Orgyia
leucostigma_, the white-marked tussock moth, as the most important of
the poisonous caterpillars of this country. Though there are occasional
reports of irritation from its hairs such cases are rare and there is no
evidence that there is any poison present. Indeed, subcutaneous
implantation of the hairs leads to no poisoning, but merely to temporary
irritation.
[Illustration: 40. Same as figure 39, on larger scale.]
Occasionally, the hairs of certain species of caterpillars find
lodgement in the conjunctiva, cornea, or iris of the eye of man and give
rise to the condition known as _opthalmia nodosa_. The essential feature
of this trouble is a nodular conjunctivitis which simulates tuberculosis
of the conjunctiva and hence has been called _pseudo-tubercular_. It may
be distinguished microscopically by the presence of the hairs.
[Illustration: 41. (_a_) Nodular conjunctivitis in the eye of a child.
De Schweinitz and Shumway.]
Numerous cases of opthalmia nodosa are on record. Of those from this
country, one of the most interesting is reported by de Schweinitz and
Shumway (1904). It is that of a child of fifteen years whose eye had
become inflamed owing to the presence of some foreign body. Downward and
inward on the bulbar conjunctiva were a number of flattened,
grayish-yellow nodules, between which was a marked congestion of the
conjunctival and episcleral vessels (fig. 41_a_). Twenty-seven nodules
could be differentiated, those directly in the center of the collection
being somewhat confluent and assuming a crescentic and circular
appearance. The nodules were excised and, on sectioning, were found to
be composed of a layer of spindle cells and round cells, outside of
which the tissue was condensed into a capsule. The interior consisted of
epithelioid cells, between which was a considerable intercellular
substance. Directly in the center of a certain number of nodules was
found the section of a hair (fig. 41_b_). The evidence indicated that
the injury had resulted from playing with caterpillars of one of the
Arctiid moths, _Spilosoma virginica_. Other reported cases have been
caused by the hairs of larvae of _Lasiocampa rubi_, _L. pini_,
_Porthetria dispar_, _Psilura monacha_ and _Cnethocampa processionea_.
[Illustration: 41_b_. Section through one of the nodules showing the
caterpillar hair. De Schweinitz and Shumway.]
RELIEF FROM POISONING BY NETTLING LARVAE--The irritation from nettling
larvae is often severe and, especially in regions where the brown-tail
abounds, inquiries as to treatment arise. In general, it may be said
that cooling lotions afford relief, and that scratching, with the
possibilities of secondary infection, should be avoided, in so far as
possible.
Among the remedies usually at hand, weak solutions of ammonia, or a
paste of ordinary baking soda are helpful. Castellani and Chalmers
recommend cleaning away the hairs by bathing the region with an alkaline
lotion, such as two per cent solution of bicarbonate of soda, and then
applying an ointment of ichthyol (10%).
In the brown-tail district, there are many proprietary remedies of which
the best ones are essentially the following, as recommended by Kirkland
(1907):
Carbolic acid 1/2 drachm.
Zinc oxide 1/2 oz.
Lime water 8 oz.
Shake thoroughly and rub well into the affected parts.
In some cases, and especially where there is danger of secondary
infection, the use of a weak solution of creoline (one teaspoonful to a
quart of water), is to be advised.
VESCICATING INSECTS AND THOSE POSSESSING OTHER POISONS IN THEIR BLOOD
PLASMA
We have seen that certain forms, for example, the poisonous spiders, not
only secrete a toxine in their poison glands, but that such a substance
may be extracted from other parts of their body, or even their eggs.
There are many insects which likewise possess a poisonous blood plasma.
Such forms have been well designated by Taschenberg as _cryptotoxic_
([Greek: kryptos] = hidden). We shall consider a few representative
forms.
[Illustration: 42_a_. Blister beetle.]
[Illustration: 42_b_. An American blister beetle. Meloe angusticollis.
Photograph by M. V. S.]
THE BLISTER BEETLES--Foremost among the cryptotoxic insects are the
_Meloidae_ or "blister beetles," to which the well-known "Spanish fly"
(fig. 42_a_), formerly very generally used in medical practice, belongs.
The vescicating property is due to the presence in the blood plasma of a
peculiar, volatile, crystalline substance known as _cantharidin_, which
is especially abundant in the reproductive organs of the beetle.
According to Kobert, the amount of this varies in different species from
.4 or .5% to 2.57% of the dry weight of the beetle.
While blister beetles have been especially used for external
application, they are also at times used internally as a stimulant and a
diuretic. The powder or extract was formerly much in vogue as an
aphrodisiac, and formed the essential constituent of various philters,
or "love powders". It is now known that its effects on the reproductive
organs appear primarily after the kidneys have been affected to such an
extent as to endanger life, and that many cases of fatal poison have
been due to its ignorant use.
There are many cases on record of poisoning and death due to internal
use, and in some instances from merely external application. There are
not rarely cases of poisoning of cattle from feeding on herbage bearing
a large number of the beetles and authentic cases are known of human
beings who have been poisoned by eating the flesh of such cattle. Kobert
states that the beetles are not poisonous to birds but that the flesh of
birds which have fed on them is poisonous to man, and that if the flesh
of chickens or frogs which have fed on the cantharidin be fed to cats it
causes in them the same symptoms as does the cantharidin.
Treatment of cases of cantharidin poison is a matter for a skilled
physician. Until he can be obtained, emetics should be administered and
these should be followed by white of egg in water. Oils should be
avoided, as they hasten the absorption of the poison.
OTHER CRYPTOTOXIC INSECTS--Though the blister beetles are the best known
of the insects with poisonous blood plasma, various others have been
reported and we shall refer to a few of the best authenticated.
One of the most famous is the Chrysomelid beetle, _Diamphidia simplex_,
the body fluids of whose larvae are used by certain South African bushmen
as an arrow poison. Its action is due to the presence of a toxalbumin
which exerts a haemolytic action on the blood, and produces inflammation
of the subcutaneous connective tissue and mucous membranes. Death
results from general paralysis. Krause (1907) has surmised that the
active principle may be a bacterial toxin arising from decomposition of
the tissues of the larva, but he presents no support of this view and it
is opposed by all the available evidence.
In China, a bug, _Heuchis sanguinea_, belonging to the family Cicadidae,
is used like the Meloidae, to produce blistering, and often causes
poisoning. It has been assumed that its vescicating properties are due
to cantharidin, but the presence of this substance has not been
demonstrated.
Certain Aphididae contain a strongly irritating substance which produces,
not merely on mucous membranes but on outer skin, a characteristic
inflammation.
It has been frequently reported that the larvae of the European cabbage
butterfly, _Pieris brassicae_, accidentally eaten by cows, horses, ducks,
and other domestic animals, cause severe colic, attempts to vomit,
paralysis of the hind legs, salivation, and stomatitis. On _postmortem_
there are to be found haemorrhagic gastro-enteritis, splenitis, and
nephritis. Kobert has recently investigated the subject and has found a
poisonous substance in the blood of not only the larvae but also the
pupae.
FOOTNOTES:
[A] This is diametrically opposed to the findings of Bordas (1905) in
the case of the European _Latrodectus 13-guttatus_, whose glands are
"much larger than those of other spiders." From a considerable
comparative study, we should also unhesitatingly make this statement
regarding the glands of our American species, _L. mactans_.
[B] Dr. E. H. Coleman (Kellogg, 1915) has demonstrated its virulence by
a series of experiments comparable with those of Kobert.
[C] According to Stiles, the species occurring in the Northwest which is
commonly identified as _D. venustus_ should be called _D. andersoni_
(see footnote, chapter 12).
[D] It should be remembered that in all the higher Hymenoptera the first
abdominal segment is fused with the thorax and that what is apparently
the sixth segment is, in reality, the seventh.
CHAPTER III
PARASITIC ARTHROPODA AFFECTING MAN
The relation of insects to man as simple parasites has long been
studied, and until very recent years the bulk of the literature of
medical entomology referred to this phase of the subject. This is now
completely overshadowed by the fact that so many of these parasitic
forms are more than simple parasites, they are transmitters of other
microscopic parasites which are pathogenic to man. Yet the importance of
insects as parasites still remains and must be considered in a
discussion of the relation of insects to the health of man. In taking up
the subject we shall first consider some general features of the
phenomenon of animal parasitism.
Parasitism is an adaptation which has originated very often among living
organisms and in widely separated groups. It would seem simple to define
what is meant by a "parasite" but, in reality, the term is not easily
limited. It is often stated that a parasite is "An organism which lives
at the expense of another," but this definition is applicable to a
predatory species or, in its broadest sense, to all organisms. For our
purpose we may say with Braun: "A parasite is an organism which, for the
purpose of obtaining food, takes up its abode, temporarily or
permanently, on or within another living organism".
Thus, parasitism is a phase of the broad biological phenomenon of
_symbiosis_, or living together of organisms. It is distinguished from
_mutualism_, or symbiosis in the narrow sense, by the fact that only one
party to the arrangement obtains any advantage, while the other is to a
greater or less extent injured.
Of parasites we may distinguish on the basis of their location on or in
the host, _ecto-parasites_, which live outside of the body; and
_endo-parasites_, which live within the body. On account of their method
of breathing the parasitic arthropods belong almost exclusively to the
first of these groups.
On the basis of relation to their host, we find _temporary parasites_,
those which seek the host only occasionally, to obtain food; and the
_stationary_ or _permanent_ parasites which, at least during certain
stages, do not leave their host.
_Facultative parasites_ are forms which are not normally parasitic, but
which, when accidentally ingested, or otherwise brought into the body,
are able to exist for a greater or less period of time in their unusual
environment. These are generally called in the medical literature
"pseudoparasites" but the term is an unfortunate one.
We shall now take up the different groups of arthropods, discussing the
more important of the parasitic forms attacking man. The systematic
relationship of these forms, and key for determining important species
will be found in Chapter XII.
ACARINA OR MITES
The ACARINA, or _mites_, form a fairly natural group of arachnids,
characterized, in general, by a sac-like, unsegmented body which is
generally fused with the cephalothorax. The mouth-parts have been united
to form a beak or rostrum.
The representatives of this group undergo a marked metamorphosis.
Commonly, the larvae on hatching from the egg, possess but three pairs of
legs, and hence are called _hexapod larvae_. After a molt, they transform
into nymphs which, like the adult, have four pairs of legs and are
called _octopod nymphs_. These after a period of growth, molt one or
more times and, acquiring external sexual organs, become adult.
Most of the mites are free-living, but there are many parasitic species
and as these have originated in widely separated families, the Acarina
form an especially favorable group for study of the origin of
parasitism. Such a study has been made by Ewing (1911), who has reached
the following conclusions:
"We have strong evidence indicating that the parasitic habit has
originated independently at least eleven times in the phylogeny of the
Ararina. Among the zoophagous parasites, the parasitic habit has been
developed from three different types of free-living Acarina: (a)
predaceous forms, (b) scavengers, (c) forms living upon the juices of
plants."
Ewing also showed that among the living forms of Acarina we can trace
out all the stages of advancing parasitism, semiparasitism, facultative
parasitism, even to the fixed and permanent type, and finally to
endoparasitism.
Of the many parasitic forms, there are several species which are serious
parasites of man and we shall consider the more important of these.
Infestation by mites is technically known as _acariasis_.
[Illustration: 43. Effect of the harvest mites on the skin of man.
Photograph by J. C. Bradley.]
The Trombidiidae, or Harvest Mites
In many parts of this country it is impossible for a visitor to go into
the fields and, particularly, into berry patches and among tall weeds
and grass in the summer or early fall without being affected by an
intolerable itching, which is followed, later, by a breaking out of
wheals, or papules, surrounded by a bright red or violaceous aureola,
(fig. 43). It is often regarded as a urticaria or eczema, produced by
change of climate, an error in diet, or some condition of general
health.
Sooner or later, the victim finds that it is due to none of these, but
to the attacks of an almost microscopic red mite, usually called
"jigger" or "chigger" in this country. As the term "chigger" is applied
to one of the true fleas, _Dermatophilus penetrans_, of the tropics,
these forms are more correctly known as "harvest mites." Natives of an
infested region may be so immune or accustomed to its attacks as to be
unaware of its presence, though such immunity is by no means possessed
by all who have been long exposed to the annoyance.
[Illustration: 44. Harvest mites. (Larvae of Trombidium). After C. V.
Riley.]
The harvest mites, or chiggers, attacking man are larval forms,
possessing three pairs of legs (fig. 44). Their systematic position was
at first unknown and they were classed under a special genus _Leptus_, a
name which is very commonly still retained in the medical literature. It
is now known that they are the larval forms of various species of the
genus _Trombidium_, a group of predaceous forms, the adults of which
feed primarily on insects and their eggs. In this country the species
best known are those to be found late in summer, as larvae at the base of
the wings of houseflies or grasshoppers.
There is much uncertainty as to the species of the larvae attacking man
but it is clear that several are implicated. Bruyant has shown that in
France the larvae _Trombidium inapinatum_ and _Trombidium holosericeum_
are those most frequently found. The habit of attacking man is abnormal
and the larvae die after entering the skin. Normally they are parasitic
on various insects.
Most recent writers agree that, on man, they do not bore into the skin,
as is generally supposed, but enter a hair follicle or sebaceous gland
and from the bottom of this, pierce the cutis with their elongate
hypopharynx. According to Braun, there arises about the inserted
hypopharynx a fibrous secretion--the so-called "beak" which is, in
reality, a product of the host. Dr. J. C. Bradley, however, has made
careful observations on their method of attack, and he assures us that
the mite ordinarily remains for a long time feeding on the surface of
the skin, where it produces the erythema above described. During this
time it is not buried in the skin but is able to retreat rapidly into it
through a hair follicle or sweat gland. The irritation from the mites
ceases after a few days, but not infrequently the intolerable itching
leads to so much scratching that secondary infection follows.
Relief from the irritation may be afforded by taking a warm salt bath as
soon as possible after exposure or by killing the mites by application
of benzine, sulphur ointment or carbolized vaseline. When they are few
in number, they can be picked out with a sterile needle.
Much may be done in the way of warding off their attacks by wearing
gaiters or close-woven stockings extending from ankle to the knee. Still
more efficacious is the sprinkling of flowers of sulphur in the
stockings and the underclothes from a little above the knee, down. The
writers have known this to make it possible for persons who were
especially susceptible to work with perfect comfort in badly infested
regions. Powdered naphthalene is successfully used in the same way and
as Chittenden (1906) points out, is a safeguard against various forms of
man-infesting tropical insect pests.
The question of the destruction of the mites in the field is sometimes
an important one, and under some conditions, is feasible. Chittenden
states that much can be accomplished by keeping the grass, weeds, and
useless herbage mowed closely, so as to expose the mites to the sun. He
believes that in some cases good may be done by dusting the grass and
other plants, after cutting, with flowers of sulphur or by spraying with
dilute kerosene emulsion in which sulphur has been mixed. More recently
(1914) he calls attention to the value of cattle, and more especially
sheep, in destroying the pests by tramping on them and by keeping the
grass and herbage closely cropped.
IXODOIDEA OR TICKS
Until recently, the ticks attracted comparatively little attention from
entomologists. Since their importance as carriers of disease has been
established, interest in the group has been enormously stimulated and
now they rank second only to the mosquitoes in the amount of detailed
study that has been devoted to them.
[Illustration: 45_a_. Argus persicus. Capitulum of male. After Nuttall
and Warburton.]
The ticks are the largest of the Acarina. They are characterized by the
fact that the hypostome, or "tongue" (fig. 45) is large and file-like,
roughened by sharp teeth. They possess a breathing pore on each side of
the body, above the third or fourth coxae (fig. 45_b_).
[Illustration: 45_b_. Left spiracle of nymph of _Argas persicus_. After
Nuttall and Warburton.]
There are two distinct families of the IXODOIDEA, differing greatly in
structure, life-history and habits. These are the ARGASIDAE and the
IXODIDAE. We shall follow Nuttall (1908) in characterizing these two
families and in pointing out their biological differences, and shall
discuss briefly the more important species which attack man. The
consideration of the ticks as carriers of disease will be reserved for a
later chapter.
Argasidae
In the ticks belonging to the family ARGASIDAE, there is comparatively
little sexual dimorphism, while this is very marked in the Ixodidae. The
capitulum, or so-called "head" is ventral, instead of terminal; the
palpi are leg-like, with the segments subequal; the scutum, or dorsal
shield, is absent; eyes, when present, are lateral, on supracoxal folds.
The spiracles are very small; coxae unarmed; tarsi without ventral
spurs, and the pulvilli are absent or rudimentary.
In habits and life history the Argasidae present striking
characteristics. In the first place, they are long-lived, a factor which
counts for much in the maintenance of the species. They are intermittent
feeders, being comparable with the bed-bug in this respect. There are
two or more nymphal stages, and they may molt after attaining maturity.
The female lays comparatively few eggs in several small batches.
Nuttall (1911) concludes that "The Argasidae represent the relatively
primitive type of ticks because they are less constantly parasitic than
are the Ixodidae. Their nymphs and adults are rapid feeders and chiefly
infest the habitat of their hosts. * * * Owing to the Argasidae infesting
the habitats of their hosts, their resistance to prolonged starvation
and their rapid feeding habits, they do not need to bring forth a large
progeny, because there is less loss of life in the various stages, as
compared with the Ixodidae, prior to their attaining maturity."
[Illustration: 46. Argus persicus. Dorsal and ventral aspects. (x4).
After Hassell.]
Of the Argasidae, we have in the United States, several species which
have been reported as attacking man.
_Argas persicus_, the famous "Miana bug" (fig. 46), is a very widely
distributed species, being reported from Europe, Asia, Africa, and
Australia. It is everywhere preeminently a parasite of fowls. According
to Nuttall it is specifically identical with _Argas americanus_ Packard
or _Argas miniatus_ Koch, which is commonly found on fowls in the United
States, in the South and Southwest. Its habits are comparable to those
of the bed-bug. It feeds intermittently, primarily at night, and instead
of remaining on its host, it then retreats to cracks and crevices.
Hunter and Hooker (1908) record that they have found the larva to remain
attached for five or eight days before dropping. Unlike the Ixodidae, the
adults oviposit frequently.
[Illustration: 47. Otiobius (Ornithodoros) megnini, head of nymph. After
Stiles.]
[Illustration: 48. Otiobius (Ornithodoros) megnini, male. (_a_) dorsal,
(_b_) ventral aspect. After Nuttall and Warburton.]
The most remarkable feature of the biology of this species is the great
longevity, especially of the adult. Hunter and Hooker report keeping
larvae confined in summer in pill boxes immediately after hatching for
about two months while under similar conditions those of the Ixodid,
_Boophilus annulatus_ lived for but two or three days. Many writers have
recorded keeping adults for long periods without food. We have kept
specimens in a tin box for over a year and a half and at the end of that
time a number were still alive. Laboulliene kept unfed adults for over
three years. In view of the effectiveness of sulphur in warding off the
attacks of Trombidiidae, it is astonishing to find that Lounsbury has
kept adults of _Argas persicus_ for three months in a box nearly filled
with flowers of sulphur, with no apparent effect on them.
We have already called attention to the occasional serious effects of
the bites of this species. While such reports have been frequently
discredited there can be no doubt that they have foundation in fact. The
readiness with which this tick attacks man, and the extent to which old
huts may be infested makes it especially troublesome.
_Otiobius (Ornithodoros) megnini_, the "spinose ear-tick" (figs. 47,
48), first described from Mexico, as occurring in the ears of horses, is
a common species in our Southwestern States and is recorded by Banks as
occurring as far north as Iowa.
The species is remarkable for the great difference between the spiny
nymph stage and the adult. The life history has been worked out by
Hooker (1908). Seed ticks, having gained entrance to the ear, attach
deeply down in the folds, engorge, and in about five days, molt; as
nymphs with their spinose body they appear entirely unlike the larvae. As
nymphs they continue feeding sometimes for months. Finally the nymph
leaves the host, molts to form the unspined adult, and without further
feeding is fertilized and commences oviposition.
The common name is due to the fact that in the young stage the ticks
occur in the ear of their hosts, usually horses or cattle. Not
uncommonly it has been reported as occurring in the ear of man and
causing very severe pain. Stiles recommends that it be removed by
pouring some bland oil into the ear.
Banks (1908) reports three species of _Ornithodoros_--_O. turicata_,
_coriaceus_ and _talaje_--as occurring in the United States. All of these
attack man and are capable of inflicting very painful bites.
Ixodidae
The ticks belonging to the family IXODIDAE (figs. 49 and 50) exhibit a
marked sexual dimorphism. The capitulum is anterior, terminal, instead
of ventral as in the Argasidae; the palpi are relatively rigid (except in
the subfamily Ixodinae), with rudimentary fourth segment; scutum present;
eyes, when present, dorsal, on side of scutum. The spiracles are
generally large, situated well behind the fourth coxae; coxae generally
with spurs; pulvilli always present.
In habits and life history the typical Ixodidae differ greatly from
the Argasidae. They are relatively short-lived, though some recent
work indicates that their longevity has been considerably
under-estimated. Typically, they are permanent feeders, remaining on the
host, or hosts, during the greater part of their life. They molt twice
only, on leaving the larval and the nymphal stages. The adult female
deposits a single, large batch of eggs. Contrasting the habits of the
Ixodidae to those of the Argasidae, Nuttall (1911) emphasizes that
the Ixodidae are more highly specialized parasites. "The majority are
parasitic on hosts having no fixed habitat and consequently all stages,
as a rule, occur upon the host."
[Illustration: 49. Ixodes ricinus; male, ventral aspect. After Braun and
Luehe.]
As mere parasites of man, apart from their power to transmit disease,
the Ixodidae are much less important than the Argasidae. Many are
reported as occasionally attacking man and of these the following native
species may be mentioned.
[Illustration: 50. Ixodes ricinus, var. scapularis, female. Capitulum
and scutum; ventral aspect of capitulum; coxae; tarsus 4; spiracle;
genital and anal grooves. After Nuttall and Warburton.]
_Ixodes ricinus_, the European castor bean tick (figs. 49, 50), is a
species which has been often reported from this country but Banks (1908)
has shown that, though it does occur, practically all of the records
apply to _Ixodes scapularis_ or _Ixodes cookei_. In Europe, _Ixodes
ricinus_ is very abundant and very commonly attacks man. At the point
of penetration of the hypostome there is more or less inflammation but
serious injury does not occur unless there have been introduced
pathogenic bacteria or, unless the tick has been abruptly removed,
leaving the capitulum in the wound. Under the latter circumstances,
there may be an abscess formed about the foreign body and occasionally,
serious results have followed. Under certain conditions the tick, in
various stages, may penetrate under the skin and produce a tumor, within
which it may survive for a considerable period of time.
_Ixodes cookei_ is given by Banks as "common on mammals in the Eastern
States as far west as the Rockies." It is said to affect man severely.
_Amblyomma americanum_, (fig. 158c), the "lone star tick," is widely
distributed in the United States. Its common name is derived from the
single silvery spot on the scutum of the female. Hunter and Hooker
regard this species as, next to _Boophilus annulatus_, the most
important tick in the United States. Though more common on cattle, it
appears to attack mammals generally, and "in portions of Louisiana and
Texas it becomes a pest of considerable importance to moss gatherers and
other persons who spend much time in the forests."
_Amblyomma cajennense_, noted as a pest of man in central and tropical
America, is reported from various places in the south and southwestern
United States.
_Dermacentor variabilis_ is a common dog tick of the eastern United
States. It frequently attacks man, but the direct effects of its bite
are negligible.
The "Rocky Mountain spotted fever tick" (_Dermacentor andersoni_
according to Stiles, _D. venustus_ according to Banks) is, from the
viewpoint of its effects on man, the most important of the ticks of the
United States. This is because, as has been clearly established, it
transmits the so-called "spotted fever" of man in our northwestern
states. This phase of the subject will be discussed later and it need
merely be mentioned here, that this species has been reported as causing
painful injuries by its bites. Dr. Stiles states that he has seen cases
of rather severe lymphangitis and various sores and swellings developing
from this cause. In one case, of an individual bitten near the elbow,
the arm became very much swollen and the patient was confined in bed for
several days. The so-called tick paralysis produced by this species is
discussed in a preceding chapter.
There are many other records of various species of ticks attacking man,
but the above-mentioned will serve as typical and it is not necessary to
enter into greater detail.
TREATMENT OF TICK BITES--When a tick attaches to man the first thing to
be done is to remove it without leaving the hypostome in the wound to
fester and bring about secondary effects. This is best accomplished by
applying to the tick's body some substance which will cause it to more
readily loosen its hold. Gasoline or petroleum, oil or vaseline will
serve. For removing the spinose ear-tick, Stiles recommends pouring some
bland oil into the ear. Others have used effectively a pledget of cotton
soaked in chloroform.
In general, the treatment recommended by Wellman for the bites of
_Ornithodoros moubata_ will prove helpful. It consists of prolonged
bathing in very hot water, followed by the application of a strong
solution of bicarbonate of soda, which is allowed to dry upon the skin.
He states that this treatment is comforting. For severe itching he
advises smearing the bites with vaseline, which is slightly impregnated
with camphor or menthol. Medical aid should be sought when complications
arise.
The DERMANYSSIDAE are Gamasid mites which differ from others of the group
in that they are parasitic on vertebrates. None of the species normally
attack man, but certain of them, especially the poultry mite, may be
accidental annoyances.
[Illustration: 51. Dermanyssus gallinae, female. After Delafond.]
_Dermanyssus gallinae_ (fig. 51), the red mite of poultry, is an
exceedingly common and widespread parasite of fowls. During the day it
lives in cracks and crevices of poultry houses, under supports of
roosts, and in litter of the food and nests, coming out at night to
feed. They often attack people working in poultry houses or handling
and plucking infested fowls. They may cause an intense pruritis, but
they do not produce a true dermatosis, for they do not find conditions
favorable for multiplication on the skin of man.
Tarsonemidae
The representatives of the family TARSONEMIDAE are minute mites, with the
body divided into cephalothorax and abdomen. There is marked sexual
dimorphism. The females possess stigmata at the anterior part of the
body, at the base of the rostrum, and differ from all other mites in
having on each side, a prominent clavate organ between the first and
second legs. The larva, when it exists, is hexapodous and resembles the
adult. A number of the species are true parasites on insects, while
others attack plants. Several of them may be accidental parasites of
man.
[Illustration: 52. Pediculoides ventricosus, female. After Webster.]
[Illustration: 53. Pediculoides ventricosus, gravid female. (x80).
After Webster.]
_Pediculoides ventricosus_ (fig. 52 and 53) is, of all the
Tarsonemidae reported, the one which has proved most troublesome to
man. It is a predaceous species which attacks a large number of insects
but which has most commonly been met with by man through its fondness
for certain grain-infesting insects, notably the Angoumois grain moth,
_Sitotroga cerealella_, and the wheat straw-worm, _Isosoma grande_. In
recent years it has attracted much attention in the United States and
its distribution and habits have been the object of detailed study by
Webster (1901).
[Illustration: 54. Pediculoides ventricosus, male. After Braun.]
There is a very striking sexual dimorphism in this species. The
non-gravid female is elongate, about 200u by 70u (fig. 52), with the
abdomen slightly striated longitudinally. The gravid female (fig. 53)
has the abdomen enormously swollen, so that it is from twenty to a
hundred times greater than the rest of the body. The species is
viviparous and the larvae undergo their entire growth in the body of the
mother. They emerge as sexually mature males and females which soon
pair. The male (fig. 54) is much smaller, reaching a length of only 320u
but is relatively broad, 80u, and angular. Its abdomen is very greatly
reduced.
As far back as 1850 it was noted as causing serious outbreaks of
peculiar dermatitis among men handling infested grain. For some time the
true source of the difficulty was unknown and it was even believed that
the grain had been poisoned. Webster has shown that in this country (and
probably in Europe as well) its attacks have been mistaken for those of
the red bugs or "chiggers" (larval Trombiidae). More recently a number of
outbreaks of a mysterious "skin disease" were traced to the use of straw
mattresses, which were found to be swarming with these almost
microscopic forms which had turned their attentions to the occupants of
the beds. Other cases cited were those of farmers running wheat through
a fanning mill, and of thrashers engaged in feeding unthrashed grain
into the cylinder of the machine.
[Illustration: 55. Lesions produced by the attacks of Pediculoides
ventricosus. After Webster.]
The medical aspects of the question have been studied especially by
Schamberg and Goldberger and from the latter's summary (1910) we derive
the following data. Within twelve to sixteen hours after exposure,
itching appears and in severe cases, especially where exposure is
continued night after night by sleeping on an infested bed, the itching
may become almost intolerable. Simultaneously, there appears an eruption
which characteristically consists of wheals surrounded by a vesicle
(fig. 55). The vesicle as a rule does not exceed a pin head in size but
may become as large as a pea. Its contents rapidly become turbid and in
a few hours it is converted into a pustule. The eruption is most
abundant on the trunk, slight on the face and extremities and almost
absent on the feet and hands. In severe cases there may be
constitutional disturbances marked, at the outset, by chilliness,
nausea, and vomiting, followed for a few days by a slight elevation of
temperature, with the appearance of albumin in the urine. In some cases
the eruption may simulate that of chicken-pox or small-pox.
Treatment for the purpose of killing the mites is hardly necessary as
they attach feebly to the surface and are readily brushed off by
friction of the clothes. "Antipruritic treatment is always called for;
warm, mildly alkaline baths or some soothing ointment, such as zinc
oxide will be found to fulfil this indication." Of course, reinfestation
must be guarded against, by discarding, or thoroughly fumigating
infested mattresses, or by avoiding other sources. Goldberger suggests
that farm laborers who must work with infested wheat or straw might
protect themselves by anointing the body freely with some bland oil or
grease, followed by a change of clothes and bath as soon as their work
is done. We are not aware of any experiments to determine the effect of
flowers of sulphur, but their efficiency in the case of "red bugs"
suggests that they are worth a trial against _Pediculoides_.
Various species of TYROGLYPHIDAE (fig. 150_f_) may abound on dried fruits
and other products and attacking persons handling them, may cause a
severe dermatitis, comparable to that described above for _Pediculoides
ventricosus_. Many instances of their occurrence as such temporary
ectoparasites are on record. Thus, workers who handle vanilla pods are
subject to a severe dermatitis, known as vanillism, which is due to the
attacks of _Tyroglyphus siro_, or a closely related species. The
so-called "grocer's itch" is similarly caused by mites infesting various
products. Castellani has shown that in Ceylon, workers employed in the
copra mills, where dried cocoanut is ground up for export, are much
annoyed by mites, which produce the so-called "copra itch." The skin of
the hands, arms and legs, and sometimes of the whole body, except the
face, is covered by fairly numerous, very pruriginous papules, often
covered by small, bloody crusts due to scratching. The condition is
readily mistaken for scabies. It is due to the attacks of _Tyroglyphus
longior castellanii_ which occur in enormous numbers in some samples of
the copra.
Sarcoptidae
The SARCOPTIDAE are minute whitish mites, semi-globular in shape, with a
delicate transversely striated cuticula. They lack eyes and tracheae. The
mouth-parts are fused at the base to form a cone which is usually
designated as the head. The legs are short and stout, and composed of
five segments. The tarsi may or may not possess a claw and may terminate
in a pedunculated sucker, or simple long bristle, or both. The presence
or absence of these structures and their distribution are much used in
classification. The mites live on or under the skin of mammals and
birds, where they produce the disease known as scabies, mange, or itch.
Several species of the Sarcoptidae attack man but the most important of
these, and the one pre-eminent as the "itch mite" is _Sarcoptes
scabiei_.
The female of _Sarcoptes scabiei_, of man, is oval and yellowish white;
the male more rounded and of a somewhat reddish tinge, and much smaller.
The body is marked by transverse striae which are partly interrupted on
the back. There are transverse rows of scales, or pointed spines, and
scattered bristles on the dorsum.
[Illustration: 56_a_. Sarcoptes scabiei, male. (x100). After
Furstenberg.]
The male (fig. 56) which is from 200-240u in length, and 150-200u in
breadth, possesses pedunculated suckers on each pair of legs except the
third, which bears, instead, a long bristle. The female (fig. 56)
300-450u in length and 250-350u in breadth, has the pedunculated suckers
on the first and second pairs of legs, only, the third and fourth
terminating in bristles.
[Illustration: 56_b_. Sarcoptes scabiei, female. (x100.) After
Furstenberg.]
The mite lives in irregular galleries from a few millimeters to several
centimeters in length, which it excavates in the epidermis (fig. 57). It
works especially where the skin is thin, such as between the fingers, in
the bend of the elbows and knees, and in the groin, but it is by no
means restricted to these localities. The female, alone, tunnels into
the skin; the males remain under the superficial epidermal scales, and
seldom are found, as they die soon after mating.
As she burrows into the skin the female deposits her eggs, which measure
about 150 x 100u. Furstenberg says that each deposits an average of
twenty-two to twenty-four eggs, though Gudden reports a single burrow as
containing fifty-one. From these there develop after about seven days,
the hexapod larvae. These molt on the sixteenth day to form an octopod
nymph, which molts again the twenty-first day. At the end of the fourth
week the nymphs molt to form the sexually mature males and the so-called
pubescent females. These pair, the males die, and the females again cast
their skin, and become the oviparous females. Thus the life cycle is
completed in about twenty-eight days.
The external temperature exercises a great influence on the development
of the mites and thus, during the winter, the areas of infestation not
only do not spread, but they become restricted. As soon as the
temperature rises, the mites increase and the infestation becomes much
more extensive.
[Illustration: 57. Sarcoptes scabiei. Diagrammatic representation of the
course in the skin of man.]
In considering the possible sources of infestation, and the chances of
reinfestation after treatment, the question of the ability of the mite
to live apart from its host is a very important one. Unfortunately there
are few reliable data on this subject. Gerlach found that, exposed in
the dry, warm air of a room they became very inactive within twenty-four
hours, that after two days they showed only slight movement, and that
after three or four days they could not be revived by moisture and
warming. The important fact was brought out that in moist air, in folded
soiled underwear, they survived as long as ten days. Bourguignon found
that under the most favorable conditions the mites of _Sarcoptes scabiei
equi_ would live for sixteen days.
The disease designated the "itch" or "scabies," in man has been known
from time immemorial, but until within less than a hundred years it was
almost universally attributed to malnutrition, errors of diet, or "bad
blood." This was in spite of the fact that the mite was known to Mouffet
and that Bonomo had figured both the adult and the egg and had declared
the mite the sole cause of the disease. In 1834 the Corsican medical
student, Francis Renucci, demonstrated the mite before a clinic in Saint
Louis Hospital in Paris and soon thereafter there followed detailed
studies of the life history of the various itch mites of man and
animals.
[Illustration: 58. Scabies on the hand. From portfolio of Dermochromes
by permission of Rebman & Co., of New York. Publishers.]
The disease is a cosmopolitan one, being exceedingly abundant in some
localities. Its spread is much favored where large numbers of people are
crowded together under insanitary conditions and hence it increases
greatly during wars and is widely disseminated and abundant immediately
afterwards. Though more commonly to be met with among the lower classes,
it not infrequently appears among those of the most cleanly, careful
habits, and it is such cases that are most liable to wrong diagnosis by
the physician.
Infection occurs solely through the passage, direct or indirect, of the
young fertilized females to the skin of a healthy individual. The adult,
oviparous females do not quit their galleries and hence do not serve to
spread the disease. The young females move about more or less at night
and thus the principal source of infestation is through sleeping in the
same bed with an infested person, or indirectly through bedclothes, or
even towels or clothing. Diurnal infestation through contact or clothing
is exceptional. Many cases are known of the disease being contracted
from animals suffering from scabies, or mange.