TRANSCRIBERS' NOTES
Barring some obvious typos, the text has been left as printed. Discrepancies identified are listed at the end of the text. Most images are linked to a larger image of the same picture.
HANDBOOK OF MEDICAL ENTOMOLOGY
WM. A. RILEY, Ph.D.
Professor of Insect Morphology and Parasitology, Cornell University
and
O. A. JOHANNSEN, Ph.D.
Professor of Biology, Cornell University
ITHACA, NEW YORK
THE COMSTOCK PUBLISHING COMPANY
1915
COPYRIGHT, 1915
BY THE COMSTOCK PUBLISHING COMPANY,
ITHACA, N. Y.
Press of W. F. Humphrey
Geneva, N. Y.
PREFACE
The Handbook of Medical Entomology is the outgrowth of a course of lectures along the lines of insect transmission and dissemination of diseases of man given by the senior author in the Department of Entomology of Cornell University during the past six years. More specifically it is an illustrated revision and elaboration of his "Notes on the Relation of Insects to Disease" published January, 1912.
Its object is to afford a general survey of the field, and primarily to put the student of medicine and entomology in touch with the discoveries and theories which underlie some of the most important modern work in preventive medicine. At the same time the older phases of the subject—the consideration of poisonous and parasitic forms—have not been ignored.
Considering the rapid shifts in viewpoint, and the development of the subject within recent years, the authors do not indulge in any hopes that the present text will exactly meet the needs of every one specializing in the field,—still less do they regard it as complete or final. The fact that the enormous literature of isolated articles is to be found principally in foreign periodicals and is therefore difficult of access to many American workers, has led the authors to hope that a summary of the important advances, in the form of a reference book may not prove unwelcome to physicians, sanitarians and working entomologists, and to teachers as a text supplementing lecture work in the subject.
Lengthy as is the bibliography, it covers but a very small fraction of the important contributions to the subject. It will serve only to put those interested in touch with original sources and to open up the field. Of the more general works, special acknowledgment should be made to those of Banks, Brumpt, Castellani and Chalmers, Comstock, Hewitt, Howard, Manson, Mense, Neveau-Lemaire, Nuttall, and Stiles.
To the many who have aided the authors in the years past, by suggestions and by sending specimens and other materials, sincerest thanks is tendered. This is especially due to their colleagues in the Department of Entomology of Cornell University, and to Professor Charles W. Howard, Dr. John Uri Lloyd, Mr. A. H. Ritchie, Dr. I. M. Unger, and Dr. Luzerne Coville.
They wish to express indebtedness to the authors and publishers who have so willingly given permission to use certain illustrations. Especially is this acknowledgment due to Professor John Henry Comstock, Dr. L. O. Howard, Dr. Graham-Smith, and Professor G. H. T. Nuttall. Professor Comstock not only authorized the use of departmental negatives by the late Professor M. V. Slingerland (credited as M. V. S.), but generously put at their disposal the illustrations from the Manual for the Study of Insects and from the Spider Book. Figures [5] and [111] are from Peter's "Der Arzt und die [Heilkunft] in der deutschen Vergangenheit." It should be noted that on examining the original, it is found that Gottfried's figure relates to an event antedating the typical epidemic of dancing mania.
Wm. A. Riley.
O. A. Johannsen.
Cornell University,
January, 1915.
ADDITIONS AND CORRECTIONS
vi [line 11], for Heilkunft read Heilkunst.
18 [line 2], for tarsi read tarsus.
32 [line 21], and legend under [fig. 23], for C. (Conorhinus) abdominalis read Melanolestes abdominalis.
47 legend under [figure] for 33c read 34.
92 line [22] and [25], for sangiusugus read sanguisugus.
116 legend under [fig. 83], for Graham-Smith read Manson.
136 [line 10, from bottom], insert "ring" after "chitin".
137 [line 3], for meditatunda read meditabunda.
145 [line 7, from bottom], for Rs read R5.
158 [line 20], for have read has.
212 after the [chapter heading] insert "continued".
219 [line 10, from bottom], for Cornohinus read Conorhinus.
266 [line 1], [fig. 158j] refers to the female.
272 [line 5], insert "palpus" before "and leg".
281 [line 6], for discodial read discoidal.
281 [last line], insert "from" before "the".
284 [line 5], for "tubercle of" read "tubercle or".
305 lines [19], [28], [44], page 306 lines [1], [9], [22], [27], [30], page 307 [line 7], page 309 lines [8], [11], for R4+5 read M1+2.
309 legend under [fig. 168] add Bureau of Entomology.
312 [line 36], for "near apex" read "of M1+2".
313 running head, for Muscidæ read Muscoidea.
314 [line 29], for "distal section" read "distally M1+2".
315 legend under [fig. 172], for Pseudopyrellia read Orthellia, for Lyperosia read Hæmatobia, for Umbana read urbana.
[323] and [325] legends under the figures, add "After Dr. J. H. Stokes".
328 [line 7 from bottom] for Apiochæta read Aphiochæta.
CONTENTS
[CHAPTER I]
[INTRODUCTION] [1-5]
[Early suggestions regarding the transmission of disease by] [insects.] [The ways in which arthropods may affect the health of man.]
[CHAPTER II]
[ARTHROPODS WHICH ARE DIRECTLY POISONOUS] [6-56]
[The Araneida, or Spiders.] [The tarantulas.] [Bird spiders.] [Spiders of the genus] [Latrodectus.] [Other venomous spiders.] [Summary.] [The Pedipalpida, or whip-scorpions.] [The Scorpionida, or true scorpions.] [The Solpugida, or solpugids.] [The Acarina, or mites and ticks.] [The Myriapoda, or centipedes and millipedes.] [The Hexapoda, or true insects.] [Piercing or biting insects poisonous to man.] [Hemiptera, or true bugs.] [The Notonectidæ or back-swimmers.] [Belostomidæ or giant] [water-bugs.] [Reduviidæ, or assassin bugs.] [Other] [Hemiptera reported as poisonous to man.] [Diptera; the midges, mosquitoes and flies.] [Stinging insects.] [Apis mellifica, the honey bee.] [Other stinging forms.] [Nettling insects.] [Lepidoptera, or butterflies and moths.] [Relief from] [poisoning by nettling larvæ.] [Vescicating insects and those possessing other poisons] [in their blood plasma.] [The blister beetles.] [Other] [cryptotoxic insects.]
[CHAPTER III]
[PARASITIC ARTHROPODS AFFECTING MAN] [57-130]
[Acarina, or mites.] [The Trombidiidæ, or harvest mites.] [The Ixodoidea, or ticks.] [Argasidæ.] [Ixodidæ.] [Treatment of tick bites.] [The mites.] [Dermanyssidæ.] [Tarsonemidæ.] [Sarcoptidæ, the itch mites.] [Demodecidæ, the follicle mites.] [Hexapoda, or true insects.] [Siphunculata, or sucking lice.] [Hemiptera.] [The bed-bug.] [Other bed-bugs.] [Parasitic Diptera, or flies.] [Psychodidæ, or moth flies.] [Phlebotominæ.] [Culicidæ, or] [mosquitoes.] [Simuliidæ, or black-flies.] [Chironomidæ, or] [midges.] [Tabanidæ, or horse-flies.] [Leptidæ or] [snipe-flies.] [Oestridæ, or bot-flies.] [Muscidæ, the] [stable-fly and others.] [Siphonaptera, or fleas.] [The fleas affecting man, the dog, cat, and rat.] [The true chiggers, or chigoes.]
[CHAPTER IV]
[ACCIDENTAL OR FACULTATIVE PARASITES] [131-143]
[Acarina, or mites.] [Myriapoda, or centipedes and millipedes.] [Lepidopterous larvæ.] [Coleoptera, or beetles.] [Dipterous larvæ causing myiasis.] [Piophila casei, the cheese skipper.] [Chrysomyia macellaria,] [the screw-worm fly.] [Calliphorinæ, the bluebottles.] [Muscinæ, the house or typhoid fly, and others.] [Anthomyiidæ, the lesser house-fly and others.] [Sarcophagidæ, the flesh-flies.]
[CHAPTER V]
[ARTHROPODS AS SIMPLE CARRIERS OF DISEASE] [144-163]
[The house or typhoid fly as a carrier of disease.] [Stomoxys calcitrans, the stable-fly.] [Other arthropods which may serve as simple carriers of] [pathogenic organisms.]
[CHAPTER VI]
[ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS] [164-174]
[Some illustrations of direct inoculations of disease germs] [by arthropods.] [The rôle of fleas in the transmission of the plague.]
[CHAPTER VII]
[ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC ORGANISMS] [175-185]
[Insects as intermediate hosts of tape-worms.] [Arthropods as intermediate hosts of nematode worms.] [Filariasis and mosquitoes.] [Other nematode parasites of man and animals.]
[CHAPTER VIII]
[ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA] [186-211]
[Mosquitoes and malaria.] [Mosquitoes and yellow fever.]
[CHAPTER IX]
[ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA] [212-229]
[Insects and trypanosomiases.] [Fleas and lice as carriers of Trypanosoma lewisi.] [Tsetse-flies and nagana.] [Tsetse-flies and sleeping sickness in man.] [South American trypanosomiasis.] [Leishmanioses and insects.] [Ticks and diseases of man and animals.] [Cattle tick and Texas fever.] [Ticks and Rocky Mountain Spotted fever of man.]
[CHAPTER X]
[ARTHROPODS AS ESSENTIAL HOSTS OF PATHOGENIC PROTOZOA][(Continued)] [230-240]
[Arthropods and Spirochætoses of man and animals.] [African relapsing fever of man.] [European relapsing fever.] [North African relapsing fever of man.] [Other types of relapsing fever of man.] [Spirochætosis of fowls.] [Other spirochæte diseases of animals.] [Typhus fever and lice.]
[CHAPTER XI]
[SOME POSSIBLE, BUT IMPERFECTLY KNOWN CASES OF][ARTHROPOD TRANSMISSION OF DISEASE] [241-256]
[Infantile paralysis, or acute anterior poliomyelitis.] [Pellagra.] [Leprosy.] [Verruga peruviana.] [Cancer.]
[CHAPTER XII]
[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 rôle 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 rôle 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 larvæ, 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 rôle. 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 larvæ 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.
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.
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.
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.
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.
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 nævia. 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.
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 Lycosidæ, 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 Lycosidæ, the term "tarantula" has been applied to members of the superfamily Avicularoidea ([fig. 6]), including the bird-spiders.
Of the Old World Lycosidæ 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."
"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 Heteropodidæ, or giant crab-spiders, ([fig. 8]).
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 Theridiidæ. 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 dyspnœic, 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 dyspnœa, 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 hæmolysis 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.
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 tarsi of the fourth pair of legs has a number of curved setæ 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 Clubionidæ, 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.
THE PEDIPALPIDA OR WHIP-SCORPIONS
The tailed whip-scorpions, belonging to the family Thelyphonidæ, 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.
The scorpions ([fig. 11]) possess large pedipalpi, terminated by strongly developed claws, or chelæ. 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 hæmatoxylin, 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 hæmatoxylin.
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.
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 zoölogists 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 zoölogists, 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.
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
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.
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°. 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° 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 maxillæ are enclosed. The mandibles are more or less deeply serrate, depending on the species concerned.
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 Fulgoridæ 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]).
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 Notonectidæ, or back swimmers, ([fig. 19b]) 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 Belostomatidæ, or giant water bugs, ([fig. 19f]) 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 œsophagus. On each side of the œsophagus 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 Belostomatidæ 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.
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.
The Reduviidæ, 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.
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."
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 Reduviidæ which, with greater or less frequency pierce the skin of human beings. These are Opsicoetes personatus, Melanolestes picipes, Coriscus subcoleoptratus ([fig. 19g]), Rasahus thoracicus, Rasahus biguttatus ([fig. 22]), Conorhinus sanguisugus ([fig. 71]), and C. abdominalis ([fig. 23]).
One of the most interesting of these species is Reduvius personatus, (= Opsicœtus 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 Lygæidæ, Coreidæ, and Capsidæ. 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 Jassidæ 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 Culicidæ, or mosquitoes, and the Simuliidæ, 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.
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 Spirochæta, 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 œsophageal 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 œsophageal 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 œsophageal 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.
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° 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, œdema, and hyperæmia.
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 œdema 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.
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.
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 Camponotinæ, 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.
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° 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 larvæ of certain Lepidoptera.
LEPIDOPTERA
When we consider the reputedly poisonous larvæ 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 larvæ, 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.
But if the larvæ 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 larvæ, 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.
Sibine stimulea (Empretia stimulea), or the saddle-back caterpillar ([fig. 32]), is another which possesses nettling hairs. This species belongs to the group of Eucleidæ, 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.
Automeris io is the best known of the nettling caterpillars. It is the larva of the Io moth, one of the Saturniidæ. 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 larvæ freely without any discomfort. The larvæ 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.
The larvæ of the brown-tail moth (Euproctis chrysorrhœa) (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.
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° C, by warming to 60° 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. 37b]). 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.
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 larvæ to determine the distribution of the hairs and their relation to the underlying tissues.
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 papillæ 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 papillæ on a tubercle, all the hairs of the papilla being connected with the underlying cells through the same pore canal, (figs. [39] and [40]).
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.
Poisonous larvæ 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 Gœldi (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.
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.
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. 41a]). 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. 41b]). 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 larvæ of Lasiocampa rubi, L. pini, Porthetria dispar, Psilura monacha and Cnethocampa processionea.
Relief from Poisoning by Nettling Larvæ—The irritation from nettling larvæ 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 | ½ drachm. |
| Zinc oxide | ½ 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 (κρυπτος = hidden). We shall consider a few representative forms.
The Blister Beetles—Foremost among the cryptotoxic insects are the Meloidæ or "blister beetles," to which the well-known "Spanish fly" ([fig. 42a]), 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 larvæ are used by certain South African bushmen as an arrow poison. Its action is due to the presence of a toxalbumin which exerts a hæmolytic 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 Cicadidæ, is used like the Meloidæ, 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 Aphididæ 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 larvæ of the European cabbage butterfly, Pieris brassicæ, 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 hæmorrhagic gastro-enteritis, splenitis, and nephritis. Kobert has recently investigated the subject and has found a poisonous substance in the blood of not only the larvæ but also the pupæ.
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 larvæ on hatching from the egg, possess but three pairs of legs, and hence are called hexapod larvæ. 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.
The Trombidiidæ, 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.
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 larvæ at the base of the wings of houseflies or grasshoppers.
There is much uncertainty as to the species of the larvæ attacking man but it is clear that several are implicated. Bruyant has shown that in France the larvæ Trombidium inapinatum and Trombidium holosericeum are those most frequently found. The habit of attacking man is abnormal and the larvæ 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.
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 coxæ ([fig. 45b]).
There are two distinct families of the Ixodoidea, differing greatly in structure, life-history and habits. These are the Argasidæ and the Ixodidæ. 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.
Argasidæ
In the ticks belonging to the family Argasidæ, there is comparatively little sexual dimorphism, while this is very marked in the Ixodidæ. 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; coxæ unarmed; tarsi without ventral spurs, and the pulvilli are absent or rudimentary.
In habits and life history the Argasidæ 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 Argasidæ represent the relatively primitive type of ticks because they are less constantly parasitic than are the Ixodidæ. Their nymphs and adults are rapid feeders and chiefly infest the habitat of their hosts. * * * Owing to the Argasidæ 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 Ixodidæ, prior to their attaining maturity."
Of the Argasidæ, 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 Ixodidæ, the adults oviposit frequently.
The most remarkable feature of the biology of this species is the great longevity, especially of the adult. Hunter and Hooker report keeping larvæ 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 Trombidiidæ, 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 larvæ. 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.
Ixodidæ
The ticks belonging to the family Ixodidæ (figs. [49] and [50]) exhibit a marked sexual dimorphism. The capitulum is anterior, terminal, instead of ventral as in the Argasidæ; the palpi are relatively rigid (except in the subfamily Ixodinæ), with rudimentary fourth segment; scutum present; eyes, when present, dorsal, on side of scutum. The spiracles are generally large, situated well behind the fourth coxæ; coxæ generally with spurs; pulvilli always present.
In habits and life history the typical Ixodidæ differ greatly from the Argasidæ. 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 Ixodidæ to those of the Argasidæ, Nuttall (1911) emphasizes that the Ixodidæ 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."
As mere parasites of man, apart from their power to transmit disease, the Ixodidæ are much less important than the Argasidæ. Many are reported as occasionally attacking man and of these the following native species may be mentioned.
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 Dermanyssidæ 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.
Dermanyssus gallinæ ([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.
Tarsonemidæ
The representatives of the family Tarsonemidæ 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.
Pediculoides ventricosus (fig. [52] and [53]) is, of all the Tarsonemidæ 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).
There is a very striking sexual dimorphism in this species. The non-gravid female is elongate, about 200µ by 70µ ([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 larvæ 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 320µ but is relatively broad, 80µ, 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 Trombiidæ). 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.
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 Tyroglyphidæ ([fig. 150f]) 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.
Sarcoptidæ
The Sarcoptidæ are minute whitish mites, semi-globular in shape, with a delicate transversely striated cuticula. They lack eyes and tracheæ. 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 Sarcoptidæ 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 striæ which are partly interrupted on the back. There are transverse rows of scales, or pointed spines, and scattered bristles on the dorsum.
The male ([fig. 56]) which is from 200-240µ in length, and 150-200µ in breadth, possesses pedunculated suckers on each pair of legs except the third, which bears, instead, a long bristle. The female ([fig. 56]) 300-450µ in length and 250-350µ in breadth, has the pedunculated suckers on the first and second pairs of legs, only, the third and fourth terminating in bristles.
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 × 100µ. Fürstenberg 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 larvæ. 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.
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.
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.
When a person is exposed to infestation, the trouble manifests itself after eight or ten days, though there usually elapses a period of twenty to thirty days before there is a suspicion of anything serious. The first symptom is an intense itching which increases when the patient is in bed. When the point of irritation is examined the galleries may usually be seen as characteristic sinuous lines, at first whitish in color but soon becoming blackish because of the contained eggs and excrement. The galleries, which may not be very distinct in some cases, may measure as much as four centimeters in length. Little vesicles, of the size of a pin head are produced by the secretions of the feeding mite; they are firm, and projecting, and contain a limpid fluid. Figures [58] and [59] show the typical appearance of scabies on the hands, while [figure 60] shows a severe general infestation. The intolerable itching induces scratching and through this various complications may arise. The lesions are not normally found on the face and scalp, and are rare on the back.
Formerly, scabies was considered a very serious disease, for its cause and method of treatment were unknown, and potentially it may continue indefinitely. Generation after generation of the mites may develop and finally their number become so great that the general health of the individual is seriously affected. Now that the true cause of the disease is known, it is easily controlled.
Treatment usually consists in softening the skin by friction with soap and warm water, followed by a warm bath, and then applying some substance to kill the mites. Stiles gives the following directions, modified from Bourguignon's, as "a rather radical guide, to be modified according to facilities and according to the delicacy of the skin or condition of the patient":
1. The patient, stripped naked, is energetically rubbed all over (except the head) for twenty minutes, with green soap and warm water. 2. He is then placed in a warm bath for thirty minutes, during which time the rubbing is continued. 3. The parasiticide is next rubbed in for twenty minutes and is allowed to remain on the body for four or five hours; in the meantime the patient's clothes are sterilized, to kill the eggs or mites attached to them. 4. A final bath is taken to remove the parasiticide.
The parasiticide usually relied on is the officinal sulphur ointment of the United States pharmacopœia. When infestation is severe it is necessary to repeat treatment after three or four days in order to kill mites which have hatched from the eggs.
The above treatment is too severe for some individuals and may, of itself, produce a troublesome dermatitis. We have seen cases where the treatment was persisted in and aggravated the condition because it was supposed to be due to the parasite. For delicate-skinned patients the use of balsam of Peru is very satisfactory, and usually causes no irritation whatever. Of course, sources of reinfection should be carefully guarded against.
Sarcoptes scabiei crustosæ, which is a distinct variety, if not species, of the human itch mite, is the cause of so-called Norwegian itch. This disease is very contagious, and is much more resistant than the ordinary scabies. Unlike the latter, it may occur on the face and scalp.
Sarcoptes scabiei not only attacks man but also occurs on a large number of mammals. Many species, based on choice of host, and minute differences in size and secondary characters, have been established, but most students of the subject relegate these to varietal rank. Many of them readily attack man, but they have become sufficiently adapted to their normal host so that they are usually less persistent on man.
Notoedres cati (usually known as Sarcoptes minor) is a species of itch mites which produce an often fatal disease of cats. The body is rounded and it is considerably smaller than Sarcoptes scabiei, the female ([fig. 61]) measuring 215-230µ long and 165-175µ wide; the males 145-150µ by 120-125µ. The most important character separating Notoedres from Sarcoptes is the position of the anus, which is dorsal instead of terminal. The mite readily transfers to man but does not persist, the infestation usually disappearing spontaneously in about two weeks. Infested cats are very difficult to cure, unless treatment is begun at the very inception of the outbreak, and under ordinary circumstances it is better to kill them promptly, to avoid spread of the disease to children and others who may be exposed.
Demodecidæ
The Demodecidæ are small, elongate, vermiform mites which live in the hair follicles of mammals. The family characteristics will be brought out in the discussion of the species infesting man, Demodex folliculorum.
Demodex folliculorum ([fig. 62]) is to be found very commonly in the hair follicles and sebaceous glands of man. It is vermiform in appearance, and with the elongate abdomen transversely striated so as to give it the appearance of segmentation. The female is 380-400µ long by 45µ; the male 300µ by 40µ. The three-jointed legs, eight in number, are reduced to mere stubs in the adult. The larval form is hexopod. These mites thus show in their form a striking adaptation to their environment. In the sebaceous glands and hair follicles they lie with their heads down ([fig. 63]). Usually there are only a few in a gland, but Gruby has counted as many as two hundred.
The frequency with which they occur in man is surprising. According to European statistics they are found in 50 per cent to 60 per cent or even more. Gruby found them in forty out of sixty persons examined. These figures are very commonly quoted, but reliable data for the United States seem to be lacking. Our studies indicate that it is very much less common in this country than is generally assumed.
The Demodex in man does not, as a rule, cause the slightest inconvenience to its host. It is often stated that they give rise to comedons or "black-heads" but there is no clear evidence that they are ever implicated. Certain it is that they are not the usual cause. A variety of the same, or a very closely related species of Demodex, on the dog gives rise to the very resistant and often fatal follicular mange.
Hexapoda or True Insects
The Hexapoda, or true insects, are characterized by the fact that the adult possesses three pairs of legs. The body is distinctly segmented and is divided into head, thorax, and abdomen.
The mouth-parts in a generalized form, consist of an upper lip, or labrum, which is a part of the head capsule, and a central unpaired hypopharynx, two mandibles, two maxillæ and a lower lip, or labium, made up of the fused pair of second maxillæ. These parts may be greatly modified, dependent upon whether they are used for biting, sucking, piercing and sucking, or a combination of biting and sucking.
Roughly speaking, insects may be grouped into those which undergo complete metamorphosis and those which have incomplete metamorphosis. They are said to undergo complete metamorphosis when the young form, as it leaves the egg, bears no resemblance to the adult. For example, the maggot changes to a quiescent pupa and from this emerges the winged active fly. They undergo incomplete metamorphosis, when the young insect, as it leaves the egg, resembles the adult to a greater or less extent, and after undergoing a certain number of molts becomes sexually mature.
Representatives of several orders have been reported as accidental or faculative parasites of man, but the true parasites are restricted to four orders. These are the Siphunculata; the Hemiptera, the Diptera and the Siphonaptera.
Siphunculata
The order Siphunculata was established by Meinert to include the true sucking lice. These are small wingless insects, with reduced mouth-parts, adapted for sucking; thorax apparently a single piece due to indistinct separation of its three segments: the compound eyes reduced to a single ommatidium on each side. The short, powerful legs are terminated by a single long claw. Metamorphosis incomplete.
There has been a great deal of discussion regarding the structure of the mouth-parts, and the relationships of the sucking lice, and the questions cannot yet be regarded as settled. The conflicting views are well represented by Cholodkovsky (1904 and 1905) and by Enderlein (1904).
Following Graber, it is generally stated that the mouth-parts consist of a short tube furnished with hooks in front, which constitutes the lower lip, and that within this is a delicate sucking tube derived from the fusion of the labrum and the mandibles. Opposed to this, Cholodkovsky and, more recently, Pawlowsky, (1906), have shown that the piercing apparatus lies in a blind sac under the pharynx and opening into the mouth cavity ([fig. 64]). It does not form a true tube but a furrow with its open surface uppermost. Eysell has shown that, in addition, there is a pair of chitinous rods which he regards as the homologues of the maxillæ.
When the louse feeds, it everts the anterior part of the mouth cavity, with its circle of hooks. The latter serve for anchoring the bug, and the piercing apparatus is then pushed out.
Most writers have classed the sucking lice as a sub-order of the Hemiptera, but the more recent anatomical and developmental studies render this grouping untenable. An important fact, bearing on the question, is that, as shown by Gross, (1905), the structure of the ovaries is radically different from that of the Hemiptera.
Lice infestation and its effects are known medically as pediculosis. Though their continued presence is the result of the grossest neglect and filthiness, the original infestation may be innocently obtained and by people of the most careful habits.
Three species commonly attack man. Strangely enough, there are very few accurate data regarding their life history.
Pediculus humanus ([fig. 65]), the head louse, is the most widely distributed. It is usually referred to in medical literature as Pediculus capitis, but the Linnean specific name has priority. In color it is of a pale gray, blackish on the margins. It is claimed by some authors that the color varies according to the color of the skin of the host. The abdomen is composed of seven distinct segments, bearing spiracles laterally. There is considerable variation in size. The males average 1.8 mm. and the females 2.7 mm. in length.
The eggs, fifty to sixty in number, stick firmly to the hairs of the host and are known as nits. They are large and conspicuous, especially on dark hair and are provided with an operculum, or cap, at the free end, where the nymphs emerge. They hatch in about six days and about the eighteenth day the young lice are sexually mature.
The head lice live by preference on the scalp of their host but occasionally they are found on the eyelashes and beard, or in the pubic region. They may also occur elsewhere on the body. The penetration of the rostrum into the skin and the discharge of an irritating saliva produce a severe itching, accompanied by the formation of an eczema-like eruption ([fig. 66]). When the infestation is severe, the discharge from the pustules mats down the hair, and scabs are formed, under which the insects swarm. "If allowed to run, a regular carapace may form, called trichoma, and the head exudes a fœtid odor. Various low plants may grow in the trichoma, the whole being known as plica palonica."—Stiles.
Sources of infestation are various. School children may obtain the lice from seatmates, by wearing the hats or caps of infested mates, or by the use, in common, of brushes and combs. They may be obtained from infested beds or sleeper berths. Stiles reports an instance in which a large number of girls in a fashionable boarding school developed lousiness a short time after traveling in a sleeping car.
Treatment is simple, for the parasites may readily be controlled by cleanliness and washing the head with a two per cent solution of carbolic acid or even kerosene. The latter is better used mixed with equal parts of olive oil, to avoid irritation. The treatment should be applied at night and followed the next morning by a shampoo with soap and warm water. It is necessary to repeat the operation in a few days. Xylol, used pure, or with the addition of five per cent of vaseline, is also very efficacious. Of course, the patient must be cautioned to stay away from a lighted lamp or fire while using either the kerosene or xylol. While these treatments will kill the eggs or nits, they will not remove them from the hairs. Pusey recommends repeated washings with vinegar or 25 per cent of acetic acid in water, for the purpose of loosening and removing the nits.
Treatment of severe infestations in females is often troublesome on account of long hair. For such cases the following method recommended by Whitfield (1912) is especially applicable:
The patient is laid on her back on the bed with her head over the edge, and beneath the head is placed a basin on a chair so that the hair lies in the basin. A solution of 1 in 40 carbolic acid is then poured over the hair into the basin and sluiced backwards and forwards until the whole of the hair is thoroughly soaked with it. It is especially necessary that care should be taken to secure thorough saturation of the hair over the ears and at the nape of the neck, since these parts are not only the sites of predilection of the parasites but they are apt to escape the solution. This sluicing is carried out for ten minutes by the clock. At the end of ten minutes the hair is lifted from the basin and allowed to drain, but is not dried or even thoroughly wrung out. The whole head is then swathed with a thick towel or better, a large piece of common house flannel, which is fastened up to form a sort of turban, and is allowed to remain thus for an hour. It can then be washed or simply allowed to dry, as the carbolic quickly disperses. At the end of this period every pediculus and what is better, every ovum is dead and no relapse will occur unless there is exposure to fresh contagion. Whitfield states that there seem to be no disadvantages in this method, which he has used for years. He has never seen carboluria result from it, but would advise first cutting the hair of children under five years of age.
Pediculus corporis (= P. vestimenti) the body louse, is larger than the preceding species, the female measuring 3.3 mm., and the male 3 mm. in length. The color is a dirty white, or grayish. P. corporis has been regarded by some authorities as merely a variety of P. humanus but Piaget maintains there are good characters separating the two species.
The body louse lives in the folds and seams of the clothing of its host, passing to the skin only when it wishes to feed. Brumpt states that he has found enormous numbers of them in the collars of glass-ware or grains worn by certain naked tribes in Africa.
Exact data regarding the life-history of this species have been supplied, in part, by the work of Warburton (1910), cited by Nuttall. He found that Pediculus corporis lives longer than P. humanus under adverse conditions. This is doubtless due to its living habitually on the clothing, whereas humanus lives upon the head, where it has more frequent opportunities of feeding. He reared a single female upon his own person, keeping the louse enclosed in a cotton-plugged tube with a particle of cloth to which it could cling. The tube was kept next to his body, thus simulating the natural conditions of warmth and moisture under which the lice thrive. The specimen was fed twice daily, while it clung to the cloth upon which it rested. Under these conditions she lived for one month. Copulation commenced five days after the female had hatched and was repeated a number of times, sexual union lasting for hours. The female laid one hundred and twenty-four eggs within twenty-five days.
The eggs hatched after eight days, under favorable conditions, such as those under which the female was kept. They did not hatch in the cold. Eggs kept near the person during the day and hung in clothing by the bedside at night, during the winter, in a cold room, did not hatch until the thirty-fifth day. When the nymphs emerge from the eggs, they feed at once, if given a chance to do so. They are prone to scatter about the person and abandon the fragment of cloth to which the adult clings.
The adult stage is reached on the eleventh day, after three molts, about four days apart. Adults enter into copulation about the fifth day and as the eggs require eight days for development, the total cycle, under favorable conditions, is about twenty-four days. Warburton's data differ considerably from those commonly quoted and serve to emphasize the necessity for detailed studies of some of the commonest of parasitic insects.
Body lice are voracious feeders, producing by their bites and the irritating saliva which they inject, rosy elevations and papules which become covered with a brownish crust. The intense itching provokes scratching, and characteristic white scars ([fig. 67]) surrounded by brownish pigment ([fig. 68]) are formed. The skin may become thickened and take on a bronze tinge. This melanoderma is especially marked in the region between the shoulders but it may become generalized, a prominent characteristic of "vagabond's disease." According to Dubre and Beille, this melanoderma is due to a toxic substance secreted by the lice, which indirectly provokes the formation of pigment.
Control measures, in the case of the body louse, consist in boiling or steaming the clothes or in some cases, sterilizing by dry heat. The dermatitis may be relieved by the use of zinc-oxide ointment, to which Pusey recommends that there be added, on account of their parasiticidal properties, sulphur and balsam of Peru, equal parts, 15 to 30 grains to the ounce.
Phthirius pubis (= P. inguinalis), the pubic louse, or so-called "crab louse," differs greatly from the preceding in appearance. It is characterized by its relatively short head which fits into a broad depression in the thorax. The latter is broad and flat and merges into the abdomen. The first pair of legs is slender and terminated by a straight claw. The second and third pairs of legs are thicker and are provided with powerful claws fitted for clinging to hairs. The females ([fig. 69]) measure 1.5 to 2 mm. in length by 1.5 mm. in breadth. The male averages a little over half as large. The eggs, or nits, are fixed at the base of the hairs. Only a few, ten to fifteen are deposited by a single female, and they hatch in about a week's time. The young lice mature in two weeks.
The pubic louse usually infests the hairs of the pubis and the perineal region. It may pass to the arm pits or even to the beard or moustache. Rarely, it occurs on the eyelids, and it has even been found, in a very few instances, occurring in all stages, on the scalp. Infestation may be contracted from beds or even from badly infested persons in a crowd. We have seen several cases which undoubtedly were due to the use of public water closets. It produces papular eruption and an intense pruritis. When abundant, there occurs a grayish discoloration of the skin which Duguet has shown is due to a poisonous saliva injected by the louse, as is the melanoderma caused by the body louse.
The pubic louse may be exterminated by the measures recommended for the head louse, or by the use of officinal mercurial ointment.
Hemiptera
Several species of Hemiptera-Heteroptera are habitual parasites of man, and others occur as occasional or accidental parasites. Of all these, the most important and widespread are the bed-bugs, belonging to the genus Cimex (= Acanthia).
The Bed-bugs—The bed-bugs are characterized by a much flattened oval body, with the short, broad head unconstricted behind, and fitting into the strongly excavated anterior margin of the thorax. The compound eyes are prominent, simple eyes lacking. Antennæ four-jointed, the first segment short, the second long and thick, and the third and fourth slender. The tarsi are short and three segmented.
It is often assumed in the literature of the subject that there is but a single species of Cimex attacking man, but several such species are to be recognized. These are distinguishable by the characters given in Chapter XII. We shall consider especially Cimex lectularius, the most common and widespread species.
Cimex lectularius (= Acanthia lectularia, Clinocoris lectularius), is one of the most cosmopolitan of human parasites but, like the lice, it has been comparatively little studied until recent years, when the possibility that it may be concerned with the transmission of various diseases has awakened interest in the details of its life-history and habits.
The adult insect ([fig. 70]) is 4-5 mm. long by 3 mm. broad, reddish brown in color, with the beak and body appendages lighter in color. The short, broad and somewhat rectangular head has no neck-like constriction but fits into the broadly semilunar prothorax. The four segmented labium or proboscis encloses the lancet-like maxillæ and mandibles. The distal of the four antennal segments is slightly club-shaped. The prothorax is characteristic of the species, being deeply incised anteriorly and with its thin lateral margins somewhat turned up. The mesothorax is triangular, with the apex posteriorly, and bears the greatly atrophied first pair of wings. There is no trace of the metathoracic pair. The greatly flattened abdomen has eight visible segments, though in reality the first is greatly reduced and has been disregarded by most writers. The body is densely covered with short bristles and hairs, the former being peculiarly saber-shaped structures sharply toothed at the apex and along the convex side ([fig. 159b]).
The peculiar disagreeable odor of the adult bed-bug is due to the secretion of the stink glands which lie on the inner surface of the mesosternum and open by a pair of orifices in front of the metacoxæ, near the middle line. In the nymphs, the thoracic glands are not developed but in the abdomen there are to be found three unpaired dorsal stink glands, which persist until the fifth molt, when they become atrophied and replaced by the thoracic glands. The nymphal glands occupy the median dorsal portion of the abdomen, opening by paired pores at the anterior margin of the fourth, fifth and sixth segments. The secretion is a clear, oily, volatile fluid, strongly acid in reaction. Similar glands are to be found in most of the Hemiptera-Heteroptera and their secretion is doubtless protective, through being disagreeable to the birds. In the bed-bug, as Marlatt points out, "it is probably an illustration of a very common phenomenon among animals, i.e., the persistence of a characteristic which is no longer of any special value to the possessor." In fact, its possession is a distinct disadvantage to the bed-bug, as the odor frequently reveals the presence of the bugs, before they are seen.
The eggs of the bed-bug ([fig. 70]) are pearly white, oval in outline, about a millimeter long, and possess a small operculum or cap at one end, which is pushed off when the young hatches. They are laid intermittently, for a long period, in cracks and crevices of beds and furniture, under seams of mattresses, under loose wall paper, and similar places of concealment of the adult bugs. Girault (1905) observed a well-fed female deposit one hundred and eleven eggs during the sixty-one days that she was kept in captivity. She had apparently deposited some of her eggs before being captured.
The eggs hatch in six to ten days, the newly emerged nymphs being about 1.5 mm. in length and of a pale yellowish white color. They grow slowly, molting five times. At the last molt the mesathoracic wing pads appear, characteristic of the adult. The total length of the nymphal stage varies greatly, depending upon conditions of food supply, temperature and possibly other factors. Marlatt (1907) found under most favorable conditions a period averaging eight days between molting which, added to an equal egg period, gave a total of about seven weeks from egg to adult insect. Girault (1912) found the postembryonic period as low as twenty-nine days and as high as seventy days under apparently similar and normal conditions of food supply. Under optimum and normal conditions of food supply, beginning August 27, the average nymphal life was 69.9 days; average number of meals 8.75 and the molts 5. Under conditions allowing about half the normal food supply the average nymphal life was from 116.9 to 139 days. Nymphs starved from birth lived up to 42 days. We have kept unfed nymphs, of the first stage, alive in a bottle for 75 days. The interesting fact was brought out that under these conditions of minimum food supply there were sometimes six molts instead of the normal number.
The adults are remarkable for their longevity, a factor which is of importance in considering the spread of the insect and methods of control. Dufour (1833) (not De Geer, as often stated) kept specimens for a year, in a closed vial, without food. This ability, coupled with their willingness to feed upon mice, bats, and other small mammals, and even upon birds, accounts for the long periods that deserted houses and camps may remain infested. There is no evidence that under such conditions they are able to subsist on the starch of the wall paper, juices of moistened wood, or the moisture in the accumulations of dust, as is often stated.
There are three or four generations a year, as Girault's breeding experiments have conclusively shown. He found that the bed-bug does not hibernate where the conditions are such as to allow it to breed and that breeding is continuous unless interrupted by the lack of food or, during the winter, by low temperature.
Bed-bugs ordinarily crawl from their hiding places and attack the face and neck or uncovered parts of the legs and arms of their victims. If undisturbed, they will feed to repletion. We have found that the young nymph would glut itself in about six minutes, though some individuals fed continuously for nine minutes, while the adult required ten to fifteen minutes for a full meal. When gorged, it quickly retreats to a crack or crevice to digest its meal, a process which requires two or three days. The effect of the bite depends very greatly on the susceptibility of the individual attacked. Some persons are so little affected that they may be wholly ignorant of the presence of a large number of bugs. Usually the bite produces a small hard swelling, or wheal, whitish in color. It may even be accompanied by an edema and a disagreeable inflammation, and in such susceptible individuals the restlessness and loss of sleep due to the presence of the insects may be a matter of considerable importance. Stiles (1907) records the case of a young man who underwent treatment for neurasthenia, the diagnosis being agreed upon by several prominent physicians; all symptoms promptly disappeared, however, immediately following a thorough fumigation of his rooms, where nearly a pint of bed-bugs were collected.
It is natural to suppose that an insect which throughout its whole life is in such intimate relationship with man should play an important rôle in the transmission of disease. Yet comparatively little is definitely known regarding the importance of the bed-bug in this respect. It has been shown that it is capable of transmitting the bubonic plague, and South American trypanosomiasis. Nuttall succeeded in transmitting European relapsing fever from mouse to mouse by its bite. It has been claimed that Oriental sore, tuberculosis, and even syphilis may be so carried. These phases of the subject will be considered later.
The sources of infestation are many, and the invasion of a house is not necessarily due to neglect, though the continued presence of the pests is quite another matter. In apartments and closely placed houses they are known to invade new quarters by migration. They are frequently to be met with in boat and sleeper berths, and even the plush seats of day coaches, whence a nucleus may be carried in baggage to residences. They may be brought in the laundry or in clothes of servants.
Usually they are a great scourge in frontier settlements and it is generally believed that they live in nature under the bark of trees, in lumber, and under similar conditions. This belief is founded upon the common occurrence of bugs resembling the bed-bug, in such places. As a matter of fact, they are no relation to bed-bugs but belong to plant-feeding forms alone ([fig. 19] c, d).
It is also often stated that bed-bugs live in poultry houses, in swallows nests, and on bats, and that it is from these sources that they gain access to dwellings. These bugs are specifically distinct from the true bed-bug, but any of them may, rarely, invade houses. Moreover, chicken houses are sometimes thoroughly infested with the true Cimex lectularius.
Control measures consist in the use of iron bedsteads and the reduction of hiding places for the bugs. If the infestation is slight they may be exterminated by a vigilant and systematic hunt, and by squirting gasoline or alcohol into cracks and crevices of the beds, and furniture. Fumigation must be resorted to in more general infestations.
The simplest and safest method of fumigation is by the use of flowers of sulphur at the rate of two pounds to each one thousand cubic feet of room space. The sulphur should be placed in a pan, a well made in the top of the pile and a little alcohol poured in, to facilitate burning. The whole should be placed in a larger pan and surrounded by water so as to avoid all danger of fire. Windows should be tightly closed, beds, closets and drawers opened, and bedding spread out over chairs in order to expose them fully to the fumes. As metal is tarnished by the sulphur fumes, ornaments, clocks, instruments, and the like should be removed. When all is ready the sulphur should be fired, the room tightly closed and left for twelve to twenty-four hours. Still more efficient in large houses, or where many hiding places favor the bugs, is fumigation with hydrocyanic acid gas. This is a deadly poison and must be used under rigid precautions. Through the courtesy of Professor Herrick, who has had much experience with this method, we give in the Appendix, the clear and detailed directions taken from his bulletin on "Household Insects."
Fumigation with formaldehyde gas, either from the liquid or "solid" formalin, so efficient in the case of contagious diseases, is useless against bed-bugs and most other insects.
Other Bed-bugs—Cimex hemipterus (= C. rotundatus) is a tropical and subtropical species, occurring in both the old and new world. Patton and Cragg state that it is distributed throughout India, Burma, Assam, the Malay Peninsula, Aden, the Island of Mauritius, Reunion, St. Vincent and Porto Rico. "It is widely distributed in Africa, and is probably the common species associated there with man." Brumpt also records it for Cuba, the Antilles, Brazil, and Venezuela.
This species, which is sometimes called the Indian bed-bug, differs from C. lectularius in being darker and in having a more elongate abdomen. The head also is shorter and narrower, and the prothorax has rounded borders.
It has the same habits and practically the same life cycle as Cimex lectularius. Mackie, in India, has found that it is capable of transmitting the Asiatic type of recurrent fever. Roger suggested that it was also capable of transmitting Kala-azar and Patton has described in detail the developmental stages of Leishmania, the causative organism of Kala-azar, in the stomach of this bug, but Brumpt declares that the forms described are those of a common, non-pathogenic flagellate to be found in the bug, and have nothing to do with the human disease. Brumpt has shown experimentally that Cimex hemipterus may transmit Trypanosoma cruzi in its excrement.
Cimex boueti, occurring in French Guinea, is another species attacking man. Its habits and general life history are the same as for the above species. It is 3 to 4.5 mm. in length, has vestigial elytra, and much elongated antennæ and legs. The extended hind legs are about as long as the body.
Cimex columbarius, a widely distributed species normally living in poultry houses and dove cotes, C. inodorus, infesting poultry in Mexico, C. hirundinis, occurring in the nests of swallows in Europe and Oeciacus vicarius ([fig. 19i]) occurring in swallows' nests in this country, are species which occasionally infest houses and attack man.
Conorhinus sanguisugus, the cone-nosed bed-bug. We have seen in our consideration of poisonous insects, that various species of Reduviid bugs readily attack man. Certain of these are nocturnal and are so commonly found in houses that they have gained the name, of "big bed-bugs." The most noted of these, in the United States, is Conorhinus sangiusugus ([fig. 71]), which is widely distributed in our Southern States.
Like its near relatives, Conorhinus sangiusugus is carnivorous in habit and feeds upon insects as well as upon mammalian and human blood. It is reported as often occurring in poultry houses and as attacking horses in barns. The life history has been worked out in considerable detail by Marlatt, (1902), from whose account we extract the following.
The eggs are white, changing to yellow and pink before hatching. The young hatch within twenty days and there are four nymphal stages. In all these stages the insect is active and predaceous, the mouth-parts ([fig. 72]) being powerfully developed. The eggs are normally deposited, and the early stages are undoubtedly passed, out of doors, the food of the immature forms being other insects. Immature specimens are rarely found indoors. It winters both in the partly grown and adult stage, often under the bark of trees or in any similar protection, and only in its nocturnal spring and early summer flights does it attack men. Marlatt states that this insect seems to be decidedly on the increase in the region which it particularly infests,—the plains region from Texas northward and westward. In California a closely related species of similar habits is known locally as the "monitor bug."
The effect of the bite of the giant bed-bug on man is often very severe, a poisonous saliva apparently being injected into the wound. We have discussed this phase of the subject more fully under the head of poisonous insects.
Conorhinus megistus is a Brazilian species very commonly attacking man, and of special interest since Chagas has shown that it is the carrier of a trypanosomiasis of man. Its habits and life history have been studied in detail by Neiva, (1910).
This species is now pre-eminently a household insect, depositing its eggs in cracks and crevices in houses, though this is a relatively recent adaptation. The nymphs emerge in from twenty to forty days, depending upon the temperature. There are five nymphal stages, and as in the case of true bed-bugs, the duration of these is very greatly influenced by the availability of food and by temperature. Neiva reckons the entire life cycle, from egg to egg, as requiring a minimum of three hundred and twenty-four days.
The nymphs begin to suck blood in three to five days after hatching. They usually feed at night and in the dark, attacking especially the face of sleeping individuals. The bite occasions but little pain. The immature insects live in cracks and crevices in houses and invade the beds which are in contact with walls, but the adults are active flyers and attack people sleeping in hammocks. The males as well as the females are blood suckers.
Like many blood-sucking forms, Conorhinus megistus can endure for long periods without food. Neiva received a female specimen which had been for fifty-seven days alive in a tightly closed box. They rarely feed on two consecutive days, even on small quantities of blood, and were never seen to feed on three consecutive days.
Methods of control consist in screening against the adult bugs, and the elimination of crevices and such hiding places of the nymphs. Where the infestation is considerable, fumigation with sulphur is advisable.
Parasitic Diptera or Flies
Of the Diptera or two-winged flies, many species occasionally attack man. Of these, a few are outstanding pests, many of them may also serve to disseminate disease, a phase of our subject which will be considered later. We shall now consider the most important of the group from the viewpoint of their direct attacks on man.
Psychodidæ or Moth-Flies
The Psychodidæ or Moth-flies, include a few species which attack man, and at least one species, Phlebotomus papatasii, is known to transmit the so-called "three-day fever" of man. Another species is supposed to be the vector of Peruvian verruga.
The family is made up of small, sometimes very small, nematocerous Diptera, which are densely covered with hairs, giving them a moth-like appearance. The wings are relatively large, oval or lanceolate in shape, and when at rest are held in a sloping manner over the abdomen, or are held horizontally in such a way as to give the insect a triangular outline. Not only is the moth-like appearance characteristic, but the venation of the wings ([fig. 163, d]) is very peculiar and, according to Comstock, presents an extremely generalized form. All of the longitudinal veins separate near the base of the wing except veins R2 and R3 and veins M1 and M2. Cross veins are wanting in most cases.
Comparatively little is known regarding the life-history and habits of the Psychodidæ, but one genus, Phlebotomus, contains minute, blood-sucking species, commonly known as sand-flies. The family is divided into two subfamilies, the Psychodinæ and the Phlebotominæ. The second of these, the Phlebotominæ, is of interest to us.
The Phlebotominæ—The Phlebotominæ differ from the Psychodinæ in that the radical sector branches well out into the wing rather than at the base of the wing. They are usually less hairy than the Psychodinæ. The ovipositor is hidden and less strongly chitinized. The species attacking man belong to the genus Phlebotomus, small forms with relatively large, hairy wings which are held upright, and with elongate proboscis. The mandibles and maxillæ are serrated and fitted for biting.
According to Miss Summers (1913) there are twenty-nine known species of the genus Phlebotomus, five European, eleven Asiatic, seven African and six American. One species only, Phlebotomus vexator, has been reported for the United States. This was described by Coquillett, (1907), from species taken on Plummer's Island, Maryland. It measures only 1.5 mm. in length. As it is very probable that this species is much more widely distributed, and that other species of these minute flies will be found to occur in our fauna, we quote Coquillett's description.
Phlebotomus vexator, Coq.: Yellow, the mesonotum brown, hairs chiefly brown; legs in certain lights appear brown, but are covered with a white tomentum; wings hyaline, unmarked; the first vein (R1) terminates opposite one-fifth of the length of the first submarginal cell (cell R2); this cell is slightly over twice as long as its petiole; terminal, horny portion of male claspers slender, bearing many long hairs; the apex terminated by two curved spines which are more than one-half as long as the preceding part, and just in front of these are two similar spines, while near the middle of the length of this portion is a fifth spine similar to the others. Length 1.5 mm.
The life-history of the Phlebotomus flies has been best worked out for the European Phlebotomus papatasii and we shall briefly summarize the account of Dœrr and Russ (1913) based primarily on work on this species. The European Phlebotomus flies appear at the beginning of the warm season, a few weeks after the cessation of the heavy rains and storms of springtime. They gradually become more abundant until they reach their first maximum, which in Italy is near the end of July (Grassi). They then become scarcer but reach a second maximum in September. At the beginning of winter they vanish completely, hibernating individuals not being found.
After fertilization there is a period of eight to ten days before oviposition. The eggs are then deposited, the majority in a single mass covered by a slimy secretion from the sebaceous glands. The larvæ emerge in fourteen to twenty days. There is uncertainty as to the length of larval life, specimens kept in captivity remaining fifty or more days without transforming. Growth may be much more rapid in nature. The larvæ do not live in fluid media but in moist detritus in dark places. Marett believes that they live chiefly on the excrement of pill-bugs (Oniscidæ) and lizards. Pupation always occurs during the night. The remnants of the larval skin remain attached to the last two segments of the quiescent pupa and serve to attach it to the stone on which it lives. The pupal stage lasts eleven to sixteen days, the adult escaping at night.
Only the females suck blood. They attack not only man but all warm-blooded animals and, according to recent workers, also cold-blooded forms, such as frogs, lizards, and larvæ. Indeed, Townsend (1914) believes that there is an intimate relation between Phlebotomus and lizards, or other reptiles the world over. The Phlebotomus passes the daylight hours within the darkened recesses of the loose stone walls and piles of rock in order to escape wind and strong light. Lizards inhabit the same places, and the flies, always ready to suck blood in the absence of light and wind, have been found more prone to suck reptilian than mammalian blood.
On hot summer nights, when the wind is not stirring, the Phlebotomus flies, or sand-flies, as they are popularly called, invade houses and sleeping rooms in swarms and attack the inmates. As soon as light begins to break the flies either escape to the breeding places, or cool, dark places protected from the wind, or a part of them remain in the rooms, hiding behind pictures, under garments, and in similar places. Wherever the Phlebotomus flies occur they are an intolerable nuisance. On account of their small size they can easily pass through the meshes of ordinary screens and mosquito curtains. They attack silently and inflict a very painful, stinging bite, followed by itching. The ankles, dorsum of the feet, wrists, inner elbow, knee joint and similar places are favorite places of attack, possibly on account of their more delicate skin.
Special interest has been attracted to these little pests in recent years, since it has been shown that they transmit the European "pappatici fever" or "three day fever." More recently yet, it appears that they are the carriers of the virus of the Peruvian "verruga." This phase of the subject will be discussed later.
Control measures have not been worked out. As Newstead says, "In consideration of the facts which have so far been brought to light regarding the economy of Phlebotomus, it is clearly evident that the task of suppressing these insects is an almost insurmountable one. Had we to deal with insects as large and as accessible as mosquitoes, the adoption of prophylactic measures would be comparatively easy, but owing to the extremely minute size and almost flea-like habits of the adult insects, and the enormous area over which the breeding-places may occur, we are faced with a problem which is most difficult of solution." For these reasons, Newstead considers that the only really prophylactic measures which can at present be taken, are those which are considered as precautionary against the bites of the insects.
Of repellents, he cites as one of the best a salve composed of the following:
| Ol. Anisi | 3 grs. |
| Ol. Eucalypti | 3 grs. |
| Ol. Terebenth | 3 grs. |
| Unq. Acid Borac. |
Of sprays he recommends as the least objectionable and at the same time one of the most effective, formalin. "The dark portions and angles of sleeping apartments should be sprayed with a one per cent. solution of this substance every day during the season in which the flies are prevalent. A fine spraying apparatus is necessary for its application and an excessive amount must not be applied. It is considered an excellent plan also to spray the mosquito curtains regularly every day towards sunset; nets thus treated are claimed to repel the attacks of these insects." This effectiveness of formalin is very surprising for, as we have seen, it is almost wholly ineffective against bed-bugs, mosquitoes, house flies and other insects, where it has been tried.
A measure which promises to be very effective, where it can be adopted, is the use of electric fans so placed as to produce a current of air in the direction of the windows of sleeping apartments. On account of the inability of the Phlebotomus flies to withstand even slight breezes, it seems very probable that they would be unable to enter a room so protected.
Culicidæ or Mosquitoes
From the medical viewpoint, probably the most interesting and important of the blood-sucking insects are the mosquitoes. Certainly this is true of temperate zones, such as those of the United States. The result is that no other group of insects has aroused such widespread interest, or has been subjected to more detailed study than have the mosquitoes, since their rôle as carriers of disease was made known. There is an enormous literature dealing with the group, but fortunately for the general student, this has been well summarized by a number of workers. The most important and helpful of the general works are those of Howard (1901), Smith (1904), Blanchard (1905), Mitchell (1907), and especially of Howard, Dyar, and Knab, whose magnificent monograph is still in course of publication.
Aside from their importance as carriers of disease, mosquitoes are notorious as pests of man, and the earlier literature on the group is largely devoted to references to their enormous numbers and their blood-thirstiness in certain regions. They are to be found in all parts of the world, from the equator to the Arctic and Antarctic regions. Linnæus, in the "Flora Lapponica," according to Howard, Dyar and Knab, "dwells at some length upon the great abundance of mosquitoes in Lapland and the torments they inflicted upon man and beast. He states that he believes that nowhere else on earth are they found in such abundance and he compares their numbers to the dust of the earth. Even in the open, you cannot draw your breath without having your mouth and nostrils filled with them; and ointments of tar and cream or of fish grease are scarcely sufficient to protect even the case-hardened cuticle of the Laplander from their bite. Even in their cabins, the natives cannot take a mouthful of food or lie down to sleep unless they are fumigated almost to suffocation." In some parts of the Northwestern and Southwestern United States it is necessary to protect horses working in the fields by the use of sheets or burlaps, against the ferocious attacks of these insects. It is a surprising fact that even in the dry deserts of the western United States they sometimes occur in enormous numbers.
Until comparatively recent years, but few species of mosquitoes were known and most of the statements regarding their life-history were based upon the classic work of Reaumur (1738) on the biology of the rain barrel mosquito, Culex pipiens. In 1896, Dr. Howard refers to twenty-one species in the United States, now over fifty are known; Giles, in 1900, gives a total of two hundred and forty-two for the world fauna, now over seven hundred species are known. We have found eighteen species at Ithaca, N. Y.
All of the known species of mosquitoes are aquatic in the larval stage, but in their life-histories and habits such great differences occur that we now know that it is not possible to select any one species as typical of the group. For our present purpose we shall first discuss the general characteristics and structure of mosquitoes, and shall then give the life-history of a common species, following this by a brief consideration of some of the more striking departures from what have been supposed to be the typical condition.
The Culicidæ are slender, nematocerous Diptera with narrow wings, antennæ plumose in the males, and usually with the proboscis much longer than the head, slender, firm and adapted for piercing in the female. The most characteristic feature is that the margins of the wings and, in most cases, the wing veins possess a fringe of scale-like hairs. These may also cover in part, or entirely, the head, thorax, abdomen and legs. The females, only, suck blood.
On account of the importance of the group in this country and the desirability of the student being able to determine material in various stages, we show in the accompanying figures the characters most used in classification.
The larvæ ([fig. 73]) are elongate, with the head and thorax sharply distinct. The larval antennæ are prominent, consisting of a single cylindrical and sometimes curved segment. The outer third is often narrower and bears at its base a fan-shaped tuft of hairs, the arrangement and abundance of which is of systematic importance. About the mouth are the so-called rotary mouth brushes, dense masses of long hairs borne by the labrum and having the function of sweeping food into the mouth. The form and arrangement of thoracic, abdominal, and anal tufts of hair vary in different species and present characteristics of value. On either side of the eighth abdominal segment is a patch of scales varying greatly in arrangement and number and of much value in separating species. Respiration is by means of tracheæ which open at the apex of the so-called anal siphon, when it is present. In addition, there are also one or two pairs of tracheal gills which vary much in appearance in different species. On the ventral side of the anal siphon is a double row of flattened, toothed spines whose number and shape are likewise of some value in separating species. They constitute the comb or pecten.
The pupa ([fig. 139, b]) unlike that of most insects, is active, though it takes no food. The head and thorax are not distinctly separated, but the slender flexible abdomen in sharply marked off. The antennæ, mouth-parts, legs, and wings of the future adult are now external, but enclosed in chitinous cases. On the upper surface, near the base of the wings are two trumpets, or breathing tubes, for the pupal spiracles are towards the anterior end instead of at the caudal end, as in the larva. At the tip of the abdomen is a pair of large chitinous swimming paddles.
As illustrative of the life cycle of a mosquito we shall discuss the development of a common house mosquito, Culex pipiens, often referred to in the Northern United States as the rain barrel mosquito. Its life cycle is often given as typical for the entire group, but, as we have already emphasized, no one species can serve this purpose.
The adults of Culex pipiens hibernate throughout the winter in cellars, buildings, hollow trees, or similar dark shelters. Early in the spring they emerge and deposit their eggs in a raft-like mass. The number of eggs in a single mass is in the neighborhood of two hundred, recorded counts varying considerably. A single female may deposit several masses during her life time. The duration of the egg stage is dependent upon temperature. In the warm summer time the larvæ may emerge within a day. The larvæ undergo four molts and under optimum conditions may transform into pupæ in about a week's time. Under the same favorable conditions, the pupal stage may be completed in a day's time. The total life cycle of Culex pipiens, under optimum conditions, may thus be completed in a week to ten days. This period may be considerably extended under less favorable conditions of temperature and food supply.
Culex pipiens breeds continuously throughout the summer, developing in rain barrels, horse troughs, tin cans, or indeed in any standing water about houses, which lasts for a week or more. The catch basins of sewers furnish an abundant supply of the pests under some conditions. Such places, the tin gutters on residences, and all possible breeding places must be considered in attempts to exterminate this species.
Other species of mosquitoes may exhibit radical departures from Culex pipiens in life-history and habits. To control them it is essential that the biological details be thoroughly worked out for, as Howard, Dyar, and Knab have emphasized, "much useless labor and expense can be avoided by an accurate knowledge of the habits of the species." For a critical discussion of the known facts the reader is referred to their monograph. We shall confine ourselves to a few illustrations.
The majority of mosquitoes in temperate climates hibernate in the egg stage, hatching in the spring or even mild winter days in water from melting snow. It is such single-brooded species which appear in astounding numbers in the far North. Similarly, in dry regions the eggs may stand thorough dessication, and yet hatch out with great promptness when submerged by the rains. "Another provision to insure the species against destruction in such a case, exists in the fact * * * that not all the eggs hatch, a part of them lying over until again submerged by subsequent rains." In temperate North America, a few species pass the winter in the larval state. An interesting illustration of this is afforded by Wyeomia smithii, whose larvæ live in pitcher plants and are to be found on the coldest winter days imbedded in the solid ice. Late in the spring, the adults emerge and produce several broods during the summer.
In the United States, one of the most important facts which has been brought out by the intensive studies of recent years is that certain species are migratory and that they can travel long distances and become an intolerable pest many miles from their breeding places. This was forcibly emphasized in Dr. Smith's work in New Jersey, when he found that migratory mosquitoes, developing in the salt marshes along the coast, are the dominant species largely responsible for the fame of the New Jersey mosquito. The species concerned are Aedes sollicitans, A. cantator and A. tæniorhynchus. Dr. Smith decided that the first of these might migrate at least forty miles inland. It is obvious that where such species are the dominant pest, local control measures are a useless waste of time and money. Such migratory habits are rare, however, and it is probable that the majority of mosquitoes do not fly any great distance from their breeding places.
While mosquitoes are thought of primarily as a pest of man, there are many species which have never been known to feed upon human or mammalian blood, no matter how favorable the opportunity. According to Howard, Dyar, and Knab, this is true of Culex territans, one of the common mosquitoes in the summer months in the Northern United States. There are some species, probably many, in which the females, like the males, are plant feeders. In experimental work, both sexes are often kept alive for long periods by feeding them upon ripe banana, dried fig, raisins, and the like, and in spite of sweeping assertions that mosquitoes must have a meal of blood in order to stimulate the ovaries to development, some of the common blood-sucking species, notably Culex pipiens, have been bred repeatedly without opportunity to feed upon blood.
The effect of the bite varies greatly with different species and depends upon the susceptibility of the individual bitten. Some persons are driven almost frantic by the attacks of the pests when their companions seem almost unconscious of any inconvenience. Usually, irritation and some degree of inflammation appear shortly following the bite. Not infrequently a hardened wheal or even a nodule forms, and sometimes scratching leads to secondary infection and serious results.
The source of the poison is usually supposed to be the salivary glands of the insect. As we have already pointed out, ([p. 34]), Macloskie believed that one lobe of the gland, on each side, was specialized for forming the poison, while a radically different view is that of Schaudinn, who believed that the irritation is due to the expelled contents of the œsophageal diverticula, which contain a gas and a peculiar type of fungi or bacteria. In numerous attempts, Schaudinn was unable to produce any irritation by applying the triturated salivary glands to a wound, but obtained the typical result when he used the isolated diverticula.
The irritation of the bite may be relieved to some extent by using ammonia water, a one per cent. alcoholic solution of menthol, or preparations of cresol, or carbolic acid. Dr. Howard recommends rubbing the bite gently with a piece of moist toilet soap. Castellani and Chalmers recommend cleansing inflamed bites with one in forty carbolic lotion, followed by dressing with boracic ointment. Of course, scratching should be avoided as much as possible.
Repellents of various kinds are used, for warding off the attacks of the insects. We have often used a mixture of equal parts of oil of pennyroyal and kerosene, applied to the hands and face. Oil of citronella is much used and is less objectionable to some persons. A recommended formula is, oil of citronella one ounce, spirits of camphor one ounce, oil of cedar one-half ounce. A last resort would seem to be the following mixture recommended by Howard, Dyar, and Knab for use by hunters and fishermen in badly infested regions, against mosquitoes and blackflies.
Take 2¼ lbs. of mutton tallow and strain it. While still hot add ½ lb. black tar (Canadian tar). Stir thoroughly and pour into the receptacle in which it is to be contained. When nearly cool stir in three ounces of oil of citronella and 1¼ oz. of pennyroyal.
At night the surest protection is a good bed net. There are many types of these in use, but in order to be serviceable and at the same time comfortable it should be roomy and hung in such a way as to be stretched tightly in every direction. We prefer one suspended from a broad, square frame, supported by a right-angled standard which is fastened to the head of the bed. It must be absolutely free from rents or holes and tucked in securely under the mattress or it will serve merely as a convenient cage to retain mosquitoes which gain an entrance. While such nets are a convenience in any mosquito ridden community, they are essential in regions where disease-carrying species abound. Screening of doors, windows and porches, against the pests is so commonly practiced in this country that its importance and convenience need hardly be urged.
Destruction of mosquitoes and prevention of breeding are of fundamental importance. Such measures demand first, as we have seen, the correct determination of the species which is to be dealt with, and a knowledge of its life-history and habits. If it prove to be one of the migratory forms, it is beyond mere local effort and becomes a problem demanding careful organization and state control. An excellent illustration of the importance and effectiveness of work along these lines is afforded by that in New Jersey, begun by the late Dr. John B. Smith and being pushed with vigor by his successor, Dr. Headlee.
In any case, there is necessity for community action. Even near the coast, where the migratory species are dominant, there are the local species which demand attention and which cannot be reached by any measures directed against the species of the salt marshes. The most important of local measures consist in the destruction of breeding places by filling or draining ponds and pools, clearing up of more temporary breeding places, such as cans, pails, water barrels and the like. Under conditions where complete drainage of swamps is impracticable or undesirable, judicious dredging may result in a pool or series of steep-sided pools deep enough to maintain a supply of fish, which will keep down the mosquito larvæ. Where water receptacles are needed for storage of rain water, they should be protected by careful screening or a film of kerosene over the top of the water, renewed every two weeks or so, so as to prevent mosquitoes from depositing their eggs. When kerosene is used, Water drawn from the bottom of the receptacle will not be contaminated by it to any injurious extent. Where ponds cannot be drained much good will be accomplished by spraying kerosene oil on the surface of the water, or by the introduction of fish which will feed on the larvæ.
Detailed consideration of the most efficient measures for controlling mosquitoes is to be found in Dr. Howard's Bulletin No. 88 of the Bureau of Entomology, "Preventive and remedial work against mosquitoes" or, in more summarized form, in Farmers' Bulletin No. 444. One of these should be obtained by any person interested in the problems of mosquito control and public health.
The Simuliidæ, or Black Flies
The Simuliidæ, or black flies, are small, dark, or black flies, with a stout body and a hump-back appearance. The antennæ are short but eleven-segmented, the wings broad, without scales or hairs, and with the anterior veins stout but the others very weak. The mouth-parts ([fig. 74]) are fitted for biting.
The larvæ of the Simuliidæ ([fig. 75]) are aquatic and, unlike those of mosquitoes, require a well ærated, or swiftly running water. Here they attach to stones, logs, or vegetation and feed upon various micro-organisms. They pupate in silken cocoons open at the top. Detailed life-histories have not been worked out for most of the species. We shall consider as typical that of Simulium pictipes, an inoffensive species widely distributed in the Eastern United States, which has been studied especially at Ithaca, N.Y. (Johannsen, 1903).
The eggs are deposited in a compact yellowish layer on the surface of rock, on the brinks of falls and rapids where the water is flowing swiftly. They are elongate ellipsoidal in shape, about .4 by .18 mm. As myriads of females deposit in the same place the egg patches may be conspicuous coatings of a foot or much more in diameter. When first laid they are enveloped in a yellowish white slime, which becomes darker, until finally it becomes black just before the emerging of the larvæ. The egg stage lasts a week.
The larvæ ([fig. 75]) are black, soft skinned, somewhat cylindrical in shape, enlarged at both ends and attenuated in the middle. The posterior half is much stouter than the anterior part and almost club-shaped. The head bears two large fan-shaped organs which aid in procuring food. Respiration is accomplished by means of three so-called blood gills which are pushed out from the dorsal part of the rectum. The larvæ occur in enormous numbers, in moss-like patches. If removed from their natural habitat and placed in quiet water they die within three or four hours. Fastened to the rock by means of a disk-like sucker at the caudal end of the body, they ordinarily assume an erect position. They move about on the surface of the rocks, to a limited extent, with a looping gait similar to that of a measuring worm, and a web is secreted which prevents their being washed away by the swiftly flowing water. They feed chiefly upon algæ and diatoms.
The complete larval stage during the summer months occupies about four weeks, varying somewhat with the temperature and velocity of the water. At the end of this period they spin from cephalic glands, boot-shaped silken cocoons within which they pupate. The cocoon when spun is firmly attached to the rock and also to adjacent cocoons. Clustered continuously over a large area and sometimes one above another, they form a compact, carpet-like covering on the rocks, the reddish-brown color of which is easily distinguishable from the jet-black appearance of the larvæ. The pupal stage lasts about three weeks. The adult fly, surrounded by a bubble of air, quickly rises to the surface of the water and escapes. The adults ([fig. 76]) are apparently short lived and thus the entire life cycle, from egg to egg is completed in approximately eight weeks.
In the case of Simulium pictipes at Ithaca, N. Y., the first brood of adults emerges early in May and successive generations are produced throughout the summer and early autumn. This species winters in the larval condition. Most of the other species of Simulium which have been studied seem to be single brooded.
While Simulium pictipes does not attack man, there are a number of the species which are blood-sucking and in some regions they are a veritable scourge. In recent years the greatest interest in the group has been aroused by Sambon's hypothesis that they transmit pellagra from man to man. This has not been established, and, indeed, seems very doubtful, but the importance of these insects as pests and the possibility that they may carry disease make it urgent that detailed life-histories of the hominoxious species be worked out.
As pests a vivid account of their attacks is in Agassiz's "Lake Superior" (p. 61), quoted by Forbes (1912).
"Neither the love of the picturesque, however, nor the interests of science, could tempt us into the woods, so terrible were the black flies. This pest of flies which all the way hither had confined our ramblings on shore pretty closely to the rocks and the beach, and had been growing constantly worse, here reached its climax. Although detained nearly two days, * * * we could only sit with folded hands, or employ ourselves in arranging specimens, and such other operations as could be pursued in camp, and under the protection of a 'smudge.' One, whom scientific ardor tempted a little way up the river in a canoe, after water plants, came back a frightful spectacle, with blood-red rings round his eyes, his face bloody, and covered with punctures. The next morning his head and neck were swollen as if from an attack of erysipelas."
There are even well authenticated accounts on record of death of humans from the attacks of large swarms of these gnats. In some regions, and especially in the Mississippi Valley in this country, certain species of black flies have been the cause of enormous losses to farmers and stockmen, through their attacks on poultry and domestic animals. C. V. Riley states that in 1874 the loss occasioned in one county in Tennessee was estimated at $500,000.
The measures of prevention and protection against these insects have been well summarized by Forbes (1912). They are of two kinds: "the use of repellents intended to drive away the winged flies, and measures for the local destruction of the aquatic larvæ. The repellents used are either smudges, or surface applications made to keep the flies from biting. The black-fly will not endure a dense smoke, and the well-known mosquito smudge seems to be ordinarily sufficient for the protection of man. In the South, leather, cloth, and other materials which will make the densest and most stifling smoke, are often preserved for this use in the spring. Smudges are built in pastures for the protection of stock, and are kept burning before the doors of barns and stables. As the black-flies do not readily enter a dark room, light is excluded from stables as much as possible during the gnat season. If teams must be used in the open field while gnats are abroad, they may be protected against the attacks of the gnats by applying cotton-seed oil or axle grease to the surface, especially to the less hairy parts of the animals, at least twice a day. A mixture of oil and tar and, indeed, several other preventives, are of practical use in badly infested regions; but no definite test or exact comparison has been made with any them in a way to give a record of the precise results."
"It is easy to drive the flies from houses or tents by burning pyrethrum powder inside; this either kills the flies or stupifies them so that they do not bite for some time thereafter." * * * "Oil of tar is commonly applied to the exposed parts of the body for the purpose of repelling the insects, and this preparation is supplied by the Hudson Bay Company to its employees. Minnesota fishermen frequently grease their faces and hands with a mixture of kerosene and mutton tallow for the same purpose." We have found a mixture of equal parts of kerosene and oil of pennyroyal efficient.
Under most circumstances very little can be done to destroy this insect in its early stage, but occasionally conditions are such that a larvicide can be used effectively. Weed (1904), and Sanderson (1910) both report excellent results from the use of phinotas oil, a proprietary compound. The first-mentioned also found that in some places the larvæ could be removed by sweeping them loose in masses with stiff stable brooms and then catching them downstream on wire netting stretched in the water.
Chironomidæ or Midges
The flies of this family, commonly known as midges, resemble mosquitoes in form and size but are usually more delicate, and the wing-veins, though sometimes hairy, are not fringed with scales. The venation is simpler than in the mosquitoes and the veins are usually less distinct.
These midges, especially in spring or autumn, are often seen in immense swarms arising like smoke over swamps and producing a humming noise which can be heard for a considerable distance. At these seasons they are frequently to be found upon the windows of dwellings, where they are often mistaken for mosquitoes.
The larvæ are worm-like, but vary somewhat in form in the different genera. Most of them are aquatic, but a few live in the earth, in manure, decaying wood, under bark, or in the sap of trees, especially in the sap which collects in wounds.
Of the many species of Chironomidæ, (over eight hundred known), the vast majority are inoffensive. The sub-family Ceratopogoninæ, however, forms an exception, for some of the members of this group, known as sandflies, or punkies, suck blood and are particularly troublesome in the mountains, along streams, and at the seashore. Most of these have been classed under the genus Ceratopogon, but the group has been broken up into a number of genera and Ceratopogon, in the strict sense, is not known to contain any species which sucks the blood of vertebrates.
The Ceratopogoninæ—The Ceratopogoninæ are among the smallest of the Diptera, many of them being hardly a millimeter long and some not even so large. They are Chironomidæ in which the thorax is not prolonged over the head. The antennæ are filiform with fourteen (rarely thirteen) segments in both sexes, those of the male being brush-like. The basal segment is enlarged, the last segment never longer than the two preceding combined, while the last five are sub-equal to, or longer than the preceding segment. The legs are relatively stouter than in the other Chironomidæ. The following three genera of this subfamily are best known as blood suckers in this country.
Of the genus Culicoides there are many species occurring in various parts of the world. A number are known to bite man and animals and it is probable that all are capable of inflicting injury. In some localities they are called punkies, in others, sand-flies, a name sometimes also applied to the species of Simulium and Phlebotomus. Owing to their very small size they are known by some tribes of Indians as No-see-ums. The larvæ are found in ponds, pools, water standing in hollow tree stumps, and the like. Though probably living chiefly in fresh water, we have found a species occurring in salt water. The larvæ are small, slender, legless, worm-like creatures ([fig. 77c]) with small brown head and twelve body segments. The pupæ ([fig. 77e]) are slender, more swollen at the anterior end and terminating in a forked process. They float nearly motionless in a vertical position, the respiratory tubes in contact with the surface film. The adults are all small, rarely exceeding 2¼ mm. in length. The wings are more or less covered with erect setulæ or hairs and in many species variously spotted and marked with iridescent blotches. The antennæ have fourteen segments, the palpi usually five. The wing venation and mouth-parts are shown in figures [77] and [78]. Of the twenty or more species of this genus occurring in the United States the following are known to bite: C. cinctus, C. guttipennis, C. sanguisuga, C. stellifer, C. variipennis, C. unicolor.
One of the most widely distributed and commonest species in the Eastern States is C. guttipennis ([fig. 77a]). It is black with brown legs, a whitish ring before the apex of each femur and both ends of each tibia; tarsi yellow, knobs of halteres yellow. Mesonotum opaque, brown, two vittæ in the middle, enlarging into a large spot on the posterior half, also a curved row of three spots in front of each wing, and the narrow lateral margins, light gray pruinose. Wings nearly wholly covered with brown hairs, gray, with markings as shown in the figure. Length one mm.
Johannseniella Will. is a wide-spread genus related to the foregoing. Its mouth-parts are well adapted for piercing and it is said to be a persistent blood sucker, particularly in Greenland. This genus is distinguished from Culicoides by its bare wings, the venation ([fig. 163], c), and the longer tarsal claws. There are over twenty North American species.
In the Southwestern United States, Tersesthes torrens Towns. occurs, a little gnat which annoys horses, and perhaps man also, by its bite. It is related to Culicoides but differs in the number of antennal segments and in its wing venation ([fig. 163], e). The fly measures but two mm. in length and is blackish in color. The antennæ of the female have thirteen segments, the palpi but three, of which the second is enlarged and swollen.
Tabanidæ or Horse-Flies
The Tabanidæ,—horse-flies, ear-flies, and deer-flies,—are well-known pests of cattle and horses and are often extremely annoying to man. The characteristics of the family and of the principal North American genera are given in the keys of Chapter XII. There are over 2500 recorded species. As in the mosquitoes, the females alone are blood suckers. The males are flower feeders or live on plant juices. This is apparently true also of the females of some of the genera.
The eggs are deposited in masses on water plants or grasses and sedges growing in marshy or wet ground. Those of a common species of Tabanus are illustrated in [figure 80, a]. They are placed in masses of several hundred, light colored when first deposited but turning black. In a week or so the cylindrical larvæ, tapering at both ends ([fig. 80, b]), escape to the water, or damp earth, and lead an active, carnivorous life, feeding mainly on insect larvæ, and worms. In the forms which have been best studied the larval life is a long one, lasting for months or even for more than a year. Until recently, little was known concerning the life-histories of this group, but the studies of Hart (1895), and Hine (1903 +) have added greatly to the knowledge concerning North American forms.
Many of the species attack man with avidity and are able to inflict painful bites, which may smart for hours. In some instances the wound is so considerable that blood will continue to flow after the fly has left. We have seen several cases of secondary infection following such bites.
The horse-flies have been definitely convicted of transferring the trypanosome of surra from diseased to healthy animals and there is good evidence that they transfer anthrax. The possibility of their being important agents in the conveyal of human diseases should not be overlooked. Indeed, Leiper has recently determined that a species of Chrysops transfers the blood parasite Filaria diurna.
Leptidæ or Snipe-Flies
The family Leptidæ is made up of moderate or large sized flies, predaceous in habit. They are sufficiently characterized in the keys of Chapter XII. Four blood-sucking species belonging to three genera have been reported. Of these Symphoromyia pachyceras is a western species. Dr. J. C. Bradley, from personal experience, reports it as a vicious biter.
Oestridæ or Bot-flies
To the family Oestridæ belong the bot and warble-flies so frequently injurious to animals. The adults are large, or of medium size, heavy bodied, rather hairy, and usually resemble bees in appearance.
The larvæ live parasitically in various parts of the body of mammals, such as the stomach (horse bot-fly), the subcutaneous connective tissue (warble-fly of cattle), or the nasal passage (sheep bot-fly or head maggot).
There are on record many cases of the occurrence of the larvæ of Oestridæ as occasional parasites of man. A number of these have been collected and reviewed in a thesis by Mme. Pètrovskaia (1910). The majority of them relate to the following species.
Gastrophilus hæmorrhoidalis, the red tailed bot-fly, is one of the species whose larvæ are most commonly found in the stomach of the horse. Schoch (1877) cites the case of a woman who suffered from a severe case of chronic catarrh of the stomach, and who vomited, and also passed from the anus, larvæ which apparently belonged to this species. Such cases are exceedingly rare but instances of subcutaneous infestation are fairly numerous. In the latter type these larvæ are sometimes the cause of the peculiar "creeping myasis." This is characterized at its beginning by a very painful swelling which gradually migrates, producing a narrow raised line four to twenty-five millimeters broad. When the larva is mature, sometimes after several months, it becomes stationary and a tumor is formed which opens and discharges the larva along with pus and serum.
Gastrophilus equi is the most widespread and common of the horse bot-flies. Portschinsky reports it as commonly causing subcutaneous myasis of man in Russia.
Hypoderma bovis (= Oestrus bovis), and Hypoderma lineata are the so-called warble-flies of cattle. The latter species is the more common in North America but Dr. C. G. Hewitt has recently shown that H. bovis also occurs. Though warbles are very common in cattle in this country, the adult flies are very rarely seen. They are about half an inch in length, very hairy, dark, and closely resemble common honey-bees.
They deposit their eggs on the hairs of cattle and the animals in licking themselves take in the young larvæ. These pass out through the walls of the œsophagus and migrate through the tissues of the animal, to finally settle down in the subcutaneous tissue of the back. The possibility of their entering directly through the skin, especially in case of infestation of man, is not absolutely precluded, although it is doubtful.
For both species of Hypoderma there are numerous cases on record of their occurrence in man. Hamilton (1893) saw a boy, six years of age, who had been suffering for some months from the glands on one side of his neck being swollen and from a fetid ulceration around the back teeth of the lower jaw of the same side. Three months' treatment was of no avail and the end seemed near; one day a white object, which was seen to move, was observed in the ulcer at the root of the tongue, and on being extracted was recognized as a full grown larva of Hypoderma. It was of usual tawny color, about half an inch long when contracted, about one third that thickness, and quite lively. The case resulted fatally. The boy had been on a dairy farm the previous fall, where probably the egg (or larva) was in some way taken into his mouth, and the larva found between the base of the tongue and the jaw suitable tissue in which to develop.
Topsent (1901) reports a case of "creeping myasis" caused by H. lineata in the skin of the neck and shoulders of a girl eight years of age. The larva travelled a distance of nearly six and a half inches. The little patient suffered excruciating pain in the place occupied by the larva.
Hypoderma diana infests deer, and has been known to occur in man.
Oestris ovis, the sheep bot-fly, or head maggot, is widely distributed in all parts of the world. In mid-summer the flies deposit living maggots in the nostrils of sheep. These larvæ promptly pass up the nasal passages into the frontal and maxillary sinuses, where they feed on the mucous to be found there. In their migrations they cause great irritation to their host, and when present in numbers may cause vertigo, paroxysms, and even death. Portschinsky in an important monograph on this species, has discussed in detail its relation to man. He shows that it is not uncommon for the fly to attack man and that the minute living larvæ are deposited in the eyes, nostrils, lips, or mouth. A typical case in which the larvæ were deposited in the eye was described by a German oculist Kayser, in 1905. A woman brought her six year old daughter to him and said that the day before, about noontime, a flying insect struck the eye of the child and that since then she had felt a pain which increased towards evening. In the morning the pain ceased but the eye was very red. She was examined at about noon, at which time she was quiet and felt no pain. She was not sensitive to light, and the only thing noticed was a slight congestion and accumulation of secretion in the corner of the right eye. A careful examination of the eye disclosed small, active, white larvæ that crawled out from the folds of the conjunctiva and then back and disappeared. Five of these larvæ were removed and although an uncomfortable feeling persisted for a while, the eye became normal in about three weeks.
Some of the other recorded cases have not resulted so favorably, for the eyesight has been seriously affected or even lost.
According to Edmund and Etienne Sergent (1907), myasis caused by the larvæ of Oestris ovis is very common among the shepherds in Algeria. The natives say that the fly deposits its larvæ quickly, while on the wing, without pause. The greatest pain is caused when these larvæ establish themselves in the nasal cavities. They then produce severe frontal headaches, making sleep impossible. This is accompanied by continuous secretion from the nasal cavities and itching pains in the sinuses. If the larvæ happen to get into the mouth, the throat becomes inflamed, swallowing is painful, and sometimes vomiting results. The diseased condition may last for from three to ten days or in the case of nasal infection, longer, but recovery always follows. The natives remove the larvæ from the eye mechanically by means of a small rag. When the nose is infested, tobacco fumigations are applied, and in case of throat infestation gargles of pepper, onion, or garlic extracts are used.
Rhinœstrus nasalis, the Russian gad-fly, parasitizes the nasopharyngeal region of the horse. According to Portschinsky, it not infrequently attacks man and then, in all the known cases deposits its larvæ in the eye, only. This is generally done while the person is quiet, but not during sleep. The fly strikes without stopping and deposits its larva instantaneously. Immediately after, the victim experiences lancinating pains which without intermission increase in violence. There is an intense conjunctivitis and if the larvæ are not removed promptly the envelopes of the eye are gradually destroyed and the organ lost.
Dermatobia cyaniventris—This fly ([fig. 83]) is widely distributed throughout tropical America, and in its larval stage is well known as a parasite of man. The larvæ (figs. [81] and [82]) which are known as the "ver macaque," "torcel," "ver moyocuil" or by several other local names, enter the skin and give rise to a boil-like swelling, open at the top, and comparable with the swelling produced by the warble fly larvæ, in cattle. They cause itching and occasional excruciating pain. When mature, nearly an inch in length, they voluntarily leave their host, drop to the ground and complete their development. The adult female is about 12 mm. in length. The face is yellow, the frons black with a grayish bloom; antennæ yellow, the third segment four times as long as the second, the arista pectinate. The thorax is bluish black with grayish bloom; the abdomen depressed, brilliant metallescent blue with violet tinge. The legs are yellowish, the squamæ and wings brownish.
The different types of larvæ represented in [figure 81] were formerly supposed to belong to different species but Blanchard regards them as merely various stages of the same species. It is only very recently that the early stage and the method by which man becomes infested were made known.
About 1900, Blanchard observed the presence of packets of large-sized eggs under the abdomen of certain mosquitoes from Central America; and in 1910, Dr. Moralès, of Costa Rica, declared that the Dermatobia deposited its eggs directly under the abdomen of the mosquito and that they were thus carried to vertebrates. Dr. Nunez Tovar observed the mosquito carriers of the eggs and placing larvæ from this source on animals, produced typical tumors and reared the adult flies. It remained for Surcouf (1913) to work out the full details. He found that the Dermatobia deposits its eggs in packets covered by a very viscid substance, on leaves. These become attached to mosquitoes of the species Janthinosoma lutzi ([fig. 84]) which walk over the leaves. The eggs which adhere to the abdomen, remain attached and are thus transported. The embryo develops, but the young larva ([fig. 82]) remains in the egg until it has opportunity to drop upon a vertebrate fed upon by the mosquito.
Muscidæ
The following Muscidæ, characterized elsewhere, deserve special mention under our present grouping of parasitic species. Other important species will be considered as facultative parasites.
Stomoxys calcitrans, the stable-fly, or the biting house-fly, is often confused with Musca domestica and therefore is discussed especially in our consideration of the latter species as an accidental carrier of disease. Its possible relation to the spread of infantile paralysis is also considered later.
The tsetse flies, belonging to the genus Glossina, are African species of blood-sucking Muscidæ which have attracted much attention because of their rôle in transmitting various trypanosome diseases of man and animals. They are characterized in Chapter XII and are also discussed in connection with the diseases which they convey.
Chrysomyia macellaria, (= Compsomyia), the "screw worm"-fly is one of the most important species of flies directly affecting man, in North America. It is not normally parasitic, however, and hence will be considered with other facultative parasites in Chapter IV.
Auchmeromyia luteola, the Congo floor maggot. This is a muscid of grewsome habits, which has a wide distribution throughout Africa. The fly ([fig. 86]) deposits its eggs on the ground of the huts of the natives. The whitish larvæ ([fig. 85]) on hatching are slightly flattened ventrally, and each segment bears posteriorly three foot-pads transversely arranged. At night the larvæ find their way into the low beds or couches of the natives and suck their blood. The adult flies do not bite man and, as far as known, the larvæ do not play any rôle in the transmission of sleeping sickness or other diseases.
This habit of blood-sucking by muscid larvæ is usually referred to as peculiar to Auchmeromyia luteola but it should be noted that the larvæ of Protocalliphora frequent the nests of birds and feed upon the young. Mr. A. F. Coutant has studied especially the life-history and habits of P. azurea, whose larvæ he found attacking young crows at Ithaca, N.Y. He was unable to induce the larvæ to feed on man.
Cordylobia anthropophaga, (Ochromyia anthropophaga), or Tumbu-fly ([fig. 87]) is an African species whose larvæ affect man much as do those of Dermatobia cyniventris, of Central and South America. The larva ([fig. 88]), which is known as "ver du Cayor" because it was first observed in Cayor, in Senegambia, develops in the skin of man and of various animals, such as dogs, cats, and monkeys. It is about 12 mm. in length, and of the form of the larvæ of other muscids. Upon the intermediate segments are minute, brownish recurved spines which give to the larva its characteristic appearance. The life-history is not satisfactorily worked out, but Fuller (1914), after reviewing the evidence believes that, as a rule, it deposits its young in the sleeping places of man and animals, whether such be a bed, a board, the floor, or the bare ground. In the case of babies, the maggots may be deposited on the scalp. The minute maggots bore their way painlessly into the skin. As many as forty parasites have been found in one individual and one author has reported finding more than three hundred in a spaniel puppy. Though their attacks are at times extremely painful, it is seldom that any serious results follow.
The Siphonaptera or Fleas
The Siphonaptera, or fleas ([fig. 89]) are wingless insects, with highly chitinized and laterally compressed bodies. The mouth-parts are formed for piercing and sucking. Compound eyes are lacking but some species possess ocelli. The metamorphosis is complete.
This group of parasites, concerning which little was known until recently, has assumed a very great importance since it was learned that fleas are the carriers of bubonic plague. Now over four hundred species are known. Of these, several species commonly attack man. The most common hominoxious species are Pulex irritans, Xenopsylla cheopis, Ctenocephalus canis, Ctenocephalus felis, Ceratophyllus fasciatus and Dermatophilus penetrans, but many others will feed readily on human blood if occasion arises.
We shall treat in this place of the general biology and habits of the hominoxious forms and reserve for the systematic section the discussion of the characteristics of the different genera.
The most common fleas infesting houses in the Eastern United States are the cosmopolitan dog and cat fleas, Ctenocephalus canis ([fig. 90]) and C. felis. Their life cycles will serve as typical. These two species have until recently been considered as one, under the name Pulex serraticeps. See [figure 92].
The eggs are oval, slightly translucent or pearly white, and measure about .5 mm. in their long diameter. They are deposited loosely in the hairs of the host and readily drop off as the animal moves around. Howard found that these eggs hatch in one to two days. The larvæ are elongate, legless, white, worm-like creatures. They are exceedingly active, and avoid the light in every way possible. They cast their first skin in from three to seven days and their second in from three to four days. They commenced spinning in from seven to fourteen days after hatching and the imago appeared five days later. Thus in summer, at Washington, the entire life cycle may be completed in about two weeks. (cf. fig. [91], [92]).
Strickland's (1914) studies on the biology of the rat flea, Ceratophyllus fasciatus, have so important a general bearing that we shall cite them in considerable detail.
He found, to begin with, that there is a marked inherent range in the rate of development. Thus, of a batch of seventy-three eggs, all laid in the same day and kept together under the same conditions, one hatched in ten days; four in eleven days; twenty-five in twelve days; thirty-one in thirteen days; ten in fourteen days; one in fifteen days; and one in sixteen days. Within these limits the duration of the egg period seems to depend mainly on the degree of humidity. The incubation period is never abnormally prolonged as in the case of lice, (Warburton) and varying conditions of temperature and humidity have practically no effect on the percentage of eggs which ultimately hatch.
The same investigator found that the most favorable condition for the larva is a low temperature, combined with a high degree of humidity; and that the presence of rubbish in which the larva may bury itself is essential to its successful development. When larvæ are placed in a bottle containing either wood-wool soiled by excrement, or with feathers or filter paper covered with dried blood they will thrive readily and pupate. They seem to have no choice between dried blood and powdered rat feces for food, and also feed readily on flea excrement. They possess the curious habit of always devouring their molted skins.
An important part of Strickland's experiments dealt with the question of duration of the pupal stage under the influence of temperature and with the longevity and habits of the adult. In October, he placed a batch of freshly formed cocoons in a small dish that was kept near a white rat in a deep glass jar in the laboratory. Two months later one small and feeble flea had emerged, but no more until February, four months after the beginning of the experiment. Eight cocoons were then dissected and seven more found to contain the imago fully formed but in a resting state. The remainder of the batch was then placed at 70° F. for one night, near a white rat. The next day all the cocoons were empty and the fleas were found on the white rat.
Thus, temperature greatly influences the duration of the pupal period, which in Ceratophyllus fasciatus averages seventeen days. Moreover, when metamorphosis is complete a low temperature will cause the imago to remain within the cocoon.
Sexually mature and ovipositing fleas, he fed at intervals and kept alive for two months, when the experiment was discontinued. In the presence of rubbish in which they could bury themselves, unfed rat fleas were kept alive for many months, whereas in the absence of any such substratum they rarely lived a month. In the former case, it was found that the length of life is influenced to some degree by the temperature and humidity. In an experiment carried out at 70° F. and 45 per cent humidity, the fleas did not live for more than four months, while in an experiment at 60° F. and 70 per cent humidity they lived for at least seventeen months. There was no indication that fleas kept under these conditions sucked moisture from surrounding objects, and those kept in bell jars, with an extract of flea-rubbish on filter paper, did not live any longer than those which were not so supplied.
Curiously enough, although the rat is the normal host of Ceratophyllus fasciatus, it was found that when given the choice these fleas would feed upon man in preference to rats. However, none of the fleas laid eggs unless they fed on rat blood.
The experiments of Strickland on copulation and oviposition in the rat flea showed that fleas do not copulate until they are sexually mature and that, at least in the case of Ceratophyllus fasciatus, the reproductive organs are imperfectly developed for some time (more than a week) after emerging from the pupa. When mature, copulation takes place soon after the fleas have fed on their true host—the rat—but not if they have fed on a facultative host only, such as man. Copulation is always followed by oviposition within a very short time.
The effect of the rat's blood on the female with regard to egg-laying, Strickland concludes, is stimulating rather than nutritive, as fleas that were without food for many months were observed to lay eggs immediately after one feed. Similarly, the male requires the stimulus of a meal of rat's blood before it displays any copulatory activity.
Mitzmain (1910) has described in detail the act of biting on man, as observed in the squirrel flea, Ceratophyllus acutus. "The flea when permitted to walk freely on the arm selects a suitable hairy space where it ceases abruptly in its locomotion, takes a firm hold with the tarsi, projects its proboscis, and prepares to puncture the skin. A puncture is drilled by the pricking epipharynx, the saw-tooth mandibles supplementing the movement by lacerating the cavity formed. The two organs of the rostrum work alternately, the middle piece boring, while the two lateral elements execute a sawing movement. The mandibles, owing to their basal attachments, are, as is expressed by the advisory committee on plague investigations in India (Journal of Hygiene, vol. 6, No. 4, p. 499), 'capable of independent action, sliding up and down but maintaining their relative positions and preserving the lumen of the aspiratory channel.' The labium doubles back, the V-shaped groove of this organ guiding the mandibles on either side."
"The action of the proboscis is executed with a forward movement of the head and a lateral and downward thrust of the entire body. As the mouth-parts are sharply inserted, the abdomen rises simultaneously. The hind and middle legs are elevated, resembling oars. The forelegs are doubled under the thorax, the tibia and tarsi resting firmly on the epidermis serve as a support for the body during the feeding. The maxillary palpi are retracted beneath the head and thorax. The labium continues to bend, at first acting as a sheath for the sawing mandibles, and as these are more deeply inserted, it bends beneath the head with the elasticity of a bow, forcing the mandibles into the wound until the maxillæ are embedded in the skin of the victim. When the proboscis is fully inserted, the abdomen ceases for a time its lateral swinging."
"The acute pain of biting is first felt when the mandibles have not quite penetrated and subsequently during each distinct movement of the abdomen. The swinging of the abdomen gradually ceases as it becomes filled with blood. The sting of the biting gradually becomes duller and less sensitive as feeding progresses. The movements of the elevated abdomen grow noticeably feebler as the downward thrusts of the springy bow-like labium becomes less frequent."
"As the feeding process advances one can discern through the translucent walls of the abdomen a constant flow of blood, caudally from the pharynx, accompanied by a peristaltic movement. The end of the meal is signified in an abrupt manner. The flea shakes its entire body, and gradually withdraws its proboscis by lowering the abdomen and legs and violently twisting the head."
"When starved for several days the feeding of the rat fleas is conducted in a rather vigorous manner. As soon as the proboscis is buried to the full length the abdomen is raised and there ensues a gradual lateral swaying motion, increasing the altitude of the raised end of the abdomen until it assumes the perpendicular. The flea is observed at this point to gain a better foothold by advancing the fore tarsi, and then, gradually doubling back the abdomen, it turns with extreme agility, nearly touching with its dorsal side the skin of the hand upon which it is feeding. Meanwhile, the hungry parasite feeds ravenously."
"It is interesting to note the peculiar nervous action which the rodent fleas exhibit immediately when the feeding process is completed or when disturbed during the biting. Even while the rostrum is inserted to the fullest the parasite shakes its head spasmodically; in a twinkling the mouth is withdrawn and then the flea hops away."
A habit of fleas which we shall see is of significance in considering their agency in the spread of bubonic plague, is that of ejecting blood from the anus as they feed.
Fleas are famous for their jumping powers, and in control measures it is of importance to determine their ability along this line. It is often stated that they can jump about four inches, or, according to the Indian Plague Commission Xenopsylla cheopis cannot hop farther than five inches. Mitzmain (1910) conducted some careful experiments in which he found that the human flea, Pulex irritans, was able to jump as far as thirteen inches on a horizontal plane. The mean average of five specimens permitted to jump at will was seven and three-tenths inches. The same species was observed to jump perpendicularly to a height of at least seven and three-fourths inches. Other species were not able to equal this record.
The effect of the bite of fleas on man varies considerably according to the individual susceptibility. According to Patton and Cragg, this was borne out in a curious manner by the experiments of Chick and Martin. "In these, eight human hosts were tried; in seven, little or no irritation was produced, while in one quite severe inflammation was set up around each bite." Of two individuals, equally accustomed to the insects, going into an infested room, one may be literally tormented by them while the other will not notice them. Indeed it is not altogether a question of susceptibility, for fleas seem to have a special predilection for certain individuals. The typical itching wheals produced by the bites are sometimes followed, especially after scratching, by inflammatory papules.
The itching can be relieved by the use of lotions of carbolic acid (2-3 per cent), camphor, menthol lotion, or carbolated vaseline. If forced to sleep in an infested room, protection from attacks can be in a large measure gained by sprinkling pyrethrum, bubach, or California insect powder between the sheets. The use of camphor, menthol, or oil of eucalyptus, or oil of pennyroyal is also said to afford protection to a certain extent.
In the Eastern United States the occurrence of fleas as household pests is usually due to infested cats and dogs which have the run of the house. We have seen that the eggs are not attached to the host but drop to the floor when they are laid. Verrill, cited by Osborn, states that on one occasion he was able to collect fully a teaspoonful of eggs from the dress of a lady in whose lap a half-grown kitten had been held for a short time. Patton and Cragg record seeing the inside of a hat in which a kitten had spent the night, so covered with flea eggs that it looked "as if it had been sprinkled with sugar from a sifter." It is no wonder that houses in which pets live become overrun with the fleas.
One of the first control measures, then, consists in keeping such animals out of the house or in rigorously keeping them free from fleas. The latter can best be accomplished by the use of strong tar soap or Armour's "Flesope," which may be obtained from most druggists. The use of a three per cent solution of creolin, approximately four teaspoonfuls to a quart of warm water, has also been recommended. While this is satisfactory in the case of dogs, it is liable to sicken cats, who will lick their fur in an effort to dry themselves. Howard recommends thoroughly rubbing into the fur a quantity of pyrethrum powder. This partially stupifies the fleas which should be promptly swept up and burned.
He also recommends providing a rug for the dog or cat to sleep on and giving this rug a frequent shaking and brushing, afterwards sweeping up and burning the dust thus removed.
Since the larvæ of fleas are very susceptible to exposure, the use of bare floors, with few rugs, instead of carpets or matting, is to be recommended. Thorough sweeping, so as to allow no accumulation of dust in cracks and crevices will prove efficient. If a house is once infested it may be necessary to thoroughly scrub the floors with hot soapsuds, or to spray them with gasoline. If the latter method is adopted, care must be taken to avoid the possibility of fire.
To clear a house of fleas Skinner recommends the use of flake naphthalene. In a badly infested house he took one room at a time, scattering on the floor five pounds of flake naphthalene, and closed it for twenty-four hours. It proved to be a perfect and effectual remedy and very inexpensive, as the naphthalene could be swept up and transferred to other rooms. Dr. Skinner adds, "so far as I am concerned, the flea question is solved and if I have further trouble I know the remedy. I intend to keep the dog and cat."
The late Professor Slingerland very effectively used hydrocyanic acid gas fumigation in exterminating fleas in houses. In one case, where failure was reported, he found on investigation that the house had become thoroughly reinfested from pet cats, which had been left untreated. Fumigation with sulphur is likewise efficient.
The fact that adult fleas are usually to be found on the floor, when not on their hosts, was ingeniously taken advantage of by Professor S. H. Gage in ridding an animal room at Cornell University of the pests. He swathed the legs of a janitor with sticky fly-paper and had him walk back and forth in the room. Large numbers of the fleas were collected in this manner.
In some parts of the southern United States hogs are commonly infested and in turn infest sheds, barns and even houses. Mr. H. E. Vick informs us that it is a common practice to turn sheep into barn-lots and sheds in the spring of the year to collect in their wool, the fleas which abound in these places after the hogs have been turned out.
It is a common belief that adult fleas are attracted to fresh meat and that advantage of this can be taken in trapping them. Various workers, notably Mitzman (1910), have shown that there is no basis for such a belief.
The true chiggers—The chigoes, or true chiggers, are the most completely parasitic of any of the fleas. Of the dozen or more known species, one commonly attacks man. This is Dermatophilus penetrans, more commonly known as Sarcopsylla penetrans or Pulex penetrans.
This species occurs in Mexico, the West Indies, Central and South America. There are no authentic records of its occurrence in the United States although, as Baker has pointed out, there is no reason why it should not become established in Florida and Texas. It is usually believed that Brazil was its original home. Sometime about the middle of the nineteenth century it was introduced into West Africa and has spread across that continent.
The males and the immature females of Dermatophilus penetrans ([fig. 93]) closely resemble those of other fleas. They are very active little brown insects about 1-1.2 mm. in size, which live in the dust of native huts and stables, and in dry, sandy soil. In such places they often occur in enormous numbers and become a veritable plague.
They attack not only man but various animals. According to Castellani and Chalmers, "Perhaps the most noted feature is the way in which it attacks pigs. On the Gold Coast it appeared to be largely kept in existence by these animals. It is very easily captured in the free state by taking a little pig with a pale abdomen, and placing it on its back on the ground on which infected pigs are living. After watching a few moments, a black speck will appear on the pig's abdomen, and quickly another and another. These black specks are jiggers which can easily be transferred to a test tube. On examination they will be found to be males and females in about equal numbers."
Both the males and females suck blood. That which characterizes this species as distinguished from other fleas attacking man is that when the impregnated female attacks she burrows into the skin and there swells until in a few days she has the size and appearance of a small pea ([fig. 94]). Where they are abundant, hundreds of the pests may attack a single individual ([fig. 95]). Here they lie with the apex of the abdomen blocking the opening. According to Fülleborn (1908) they do not penetrate beneath the epidermis. The eggs are not laid in the flesh of the victim, as is sometimes stated, but are expelled through this opening. The female then dies, withers and falls away or is expelled by ulceration. According to Brumpt, she first quits the skin and then, falling to the ground, deposits her eggs. The subsequent development in so far as known, is like that of other fleas.
The chigoe usually enters between the toes, the skin about the roots of the nails, or the soles of the feet, although it may attack other parts of the body. Mense records the occurrence in folds of the epidermis, as in the neighborhood of the anus. They give rise to irritation and unless promptly and aseptically removed there often occurs pus formation and the development of a more or less serious abscess. Gangrene and even tetanus may ensue.
Treatment consists in the careful removal of the insect, an operation more easily accomplished a day or two after its entrance, than at first, when it is unswollen. The ulcerated point should then be treated with weak carbolic acid, or tincture of iodine, or dusted thoroughly with an antiseptic powder.
Castellani and Chalmers recommend as prophylactic measures, keeping the house clean and keeping pigs, poultry, and cattle away therefrom. "High boots should be used, and especial care should be taken not to go to a ground floor bathroom with bare feet. The feet, especially the toes, and under the nails, should be carefully examined every morning to see if any black dots can be discovered, when the jigger should be at once removed, and in this way suppuration will be prevented. It is advisable, also, to sprinkle the floors with carbolic lotion, Jeyes' fluid, or with pyrethrum powder, or with a strong infusion of native tobacco, as recommended by Law and Castellani."
Echidnophaga gallinacea ([fig. 96]) is a widely distributed Hectopsyllid attacking poultry ([fig. 97]). It occurs in the Southern and Southwestern United States and has been occasionally reported as attacking man, especially children. It is less highly specialized than Dermatophilus penetrans, and does not ordinarily cause serious trouble in man.
CHAPTER IV
ACCIDENTAL OR FACULTATIVE PARASITES
In addition to the many species of Arthropods which are normally parasitic on man and animals, there is a considerable number of those which may be classed as accidental or facultative parasites.
Accidental or facultative parasites are species which are normally free-living, but which are able to exist as parasites when accidentally introduced into the body of man or other animal. A wide range of forms is included under this grouping.
Acarina
A considerable number of mites have been reported as accidental or even normal, endoparasites of man, but the authentic cases are comparatively few.
In considering such reports it is well to keep in mind von Siebold's warning that in view of the universal distribution of mites one should be on his guard. In vessels in which animal and other organic fluids and moist substances gradually dry out, mites are very abundantly found. If such vessels are used without very careful preliminary cleaning, for the reception of evacuations of the sick, or for the reception of parts removed from the body, such things may be readily contaminated by mites, which have no other relation whatever to them.
Nevertheless, there is no doubt but that certain mites, normally free-living, have occurred as accidental parasites of man. Of these the most commonly met with is Tyroglyphus siro, the cheese-mite.
Tyroglyphus siro is a small mite of a whitish color. The male measures about 500µ long by 250µ wide, the female slightly larger. They live in cheese of almost any kind, especially such as is a little decayed. "The individuals gather together in winter in groups or heaps in the hollows and chinks of the cheese and there remain motionless. As soon as the temperature rises a little, they gnaw away at the cheese and reduce it to a powder. The powder is composed of excrement having the appearance of little grayish microscopic balls; eggs, old and new, cracked and empty; larvæ, nymphs, and perfect mites, cast skins and fragments of cheese, to which must be added numerous spores of microscopic fungi."—Murray.
Tyroglyphus siro, and related species, have been found many times in human feces, under conditions which preclude the explanation that the contamination occurred outside of the body. They have been supposed to be the cause of dysentery, or diarrhœa, and it is probable that the Acarus dysenteriæ of Linnæus, and Latreille, was this species. However, there is little evidence that the mites cause any noteworthy symptoms, even when taken into the body in large numbers.
Histiogaster spermaticus ([fig. 152]) is a Tyroglyphid mite which was reported by Trouessart (1902) as having been found in a cyst in the groin, adherent to the testis. When the cyst was punctured, it yielded about two ounces of opalescent fluid containing spermatozoa and numerous mites in all stages of development. The evidence indicated that a fecundated female mite had been introduced into the urethra by means of an unclean catheter. Though Trouessart reported the case as that of a Sarcoptid, Banks places the genus Histiogaster with the Tyroglyphidæ. He states that our species feeds on the oyster-shell bark louse, possibly only after the latter is dead, and that in England a species feeds within decaying reeds.
Nephrophages sanguinarius is a peculiarly-shaped, angular mite which was found by Miyake and Scriba (1893) for eight successive days in the urine of a Japanese suffering from fibrinuria. Males, .117 mm. long by .079 mm. wide, females .36 mm. by. 12 mm., and eggs were found both in the spontaneously emitted urine and in that drawn by means of a catheter. All the mites found were dead. The describers regarded this mite as a true endoparasite, but it is more probable that it should be classed as an accidental parasite.
Myriapoda
There are on record a number of cases of myriapods occurring as accidental parasites of man. The subject has been treated in detail by Blanchard (1898 and 1902), who discussed forty cases. Since then at least eight additions have been made to the list.
Neveau-Lamaire (1908) lists thirteen species implicated, representing eight different genera. Of the Chilognatha there are three, Julus terrestris, J. londinensis and Polydesmus complanatus. The remainder are Chilopoda, namely, Lithobius forficatus, L. malenops, Geophilus carpophagus, G. electricus, G. similis, G. cephalicus, Scutigera coleoptrata, Himantarium gervaisi, Chætechelyne vesuviana and Stigmatogaster subterraneus.
The majority of the cases relate to infestation of the nasal fossæ, or the frontal sinus, but intestinal infestation also occurs and there is one recorded case of the presence of a species in Julus ([fig. 13]) in the auditory canal of a child.
In the nose, the myriapods have been known to live for months and according to some records, even for years. The symptoms caused by their presence are inflammation, with or without increased flow of mucus, itching, more or less intense headache, and at times general symptoms such as vertigo, delirium, convulsions, and the like. These symptoms disappear suddenly when the parasites are expelled.
In the intestine of man, myriapods give rise to obscure symptoms suggestive of infestation by parasitic worms. In a case reported by Verdun and Bruyant (1912), a child twenty months of age had been affected for fifteen days by digestive disturbances characterized by loss of appetite, nausea and vomiting. The latter had been particularly pronounced for three days, when there was discovered in the midst of the material expelled a living myriapod of the species Chætechelyne vesuviana. Anthelminthics had been administered without result. In some of the other cases, the administration of such drugs had resulted in the expulsion of the parasite through the anus.
One of the extreme cases on record is that reported by Shipley (1914). Specimens of Geophilus gorizensis (= G. subterraneus) "were vomited and passed by a woman of 68 years of age. Some of the centipedes emerged through the patient's nose, and it must be mentioned that she was also suffering from a round worm. One of her doctors was of the opinion that the centipedes were certainly breeding inside the lady's intestines, and as many as seven or eight, sometimes more, were daily leaving the alimentary canal."
"According to her attendant's statements those centipedes had left the body in some hundreds during a period of twelve or eighteen months. Their presence produced vomiting and some hæmatemesis, and treatment with thymol, male-fern and turpentine had no effect in removing the creatures."
The clinical details, as supplied by Dr. Theodore Thompson were as follows:
"Examined by me July, 1912, her tongue was dry and glazed. There was bleeding taking place from the nose and I saw a living centipede she had just extracted from her nostril. Her heart, lungs and abdomen appeared normal. She was not very wasted, and did not think she had lost much flesh, nor was there any marked degree of anemia."
Shipley gives the following reasons for believing it impossible that these centipedes could have multiplied in the patient's intestine. "The breeding habits of the genus Geophilus are peculiar, and ill adapted for reproducing in such a habitat. The male builds a small web or nest, in which he places his sperm, and the female fertilizes herself from this nest or web, and when the eggs are fertilized they are again laid in a nest or web in which they incubate and in two or three weeks hatch out. The young Geophilus differ but very little from the adult, except in size. It is just possible, but improbable, that a clutch of eggs had been swallowed by the host when eating some vegetables or fruit, but against this is the fact that the Geophilus does not lay its eggs upon vegetables or fruit, but upon dry wood or earth. The egg-shell is very tough and if the eggs had been swallowed the egg-shells could certainly have been detected if the dejecta were examined. The specimens of the centipede showed very little signs of being digested, and it is almost impossible to reconcile the story of the patient with what one knows of the habits of the centipedes."
In none of the observed cases have there been any clear indications as to the manner of infestation. It is possible that the myriapods have been taken up in uncooked fruit or vegetables.
Lepidopterous Larvæ
Scholeciasis—Hope (1837) brought together six records of infestation of man by lepidopterous larvæ and proposed to apply the name scholeciasis to this type of parasitism. The clearest case was that of a young boy who had repeatedly eaten raw cabbage and who vomited larvæ of the cabbage butterfly, Pieris brassicæ. Such cases are extremely rare, and there are few reliable data relative to the subject. In this connection it may be noted that Spuler (1906) has described a moth whose larvæ live as ectoparasites of the sloth.
Coleoptera
Canthariasis—By this term Hope designated instances of accidental parasitism by the larvæ or adults of beetles. Reports of such cases are usually scouted by parasitologists but there seems no good basis for wholly rejecting them. Cobbold refers to a half dozen cases of accidental parasitism by the larvæ of Blaps mortisaga. In one of these cases upwards of 1200 larvæ and several perfect insects were said to have been passed per annum. French (1905) reports the case of a man who for a considerable period voided adult living beetles of the species Nitidula bipustulata. Most of the other cases on record relate to the larvæ of Dermestidæ (larder beetles et al.) or Tenebrionidæ (meal infesting species). Infestation probably occurs through eating raw or imperfectly cooked foods containing eggs or minute larvæ of these insects.
Brumpt cites a curious case of accidental parasitism by a coleopterous larva belonging to the genus Necrobia. This larva was extracted from a small tumor, several millimeters long, on the surface of the conjunctiva of the eye. The larvæ of this genus ordinarily live in decomposing flesh and cadavers.
Dipterous Larvæ
Myasis—By this term (spelled also myiasis, and myiosis), is meant parasitism by dipterous larvæ. Such parasitism may be normal, as in the cases already described under the heading parasitic Diptera, or it may be facultative, due to free-living larvæ being accidentally introduced into wounds or the body-cavities of man. Of this latter type, there is a multitude of cases on record, relating to comparatively few species. The literature of the subject, like that relating to facultative parasitism in general, is unsatisfactory, for most of the determinations of species have been very loose. Indeed, so little has been known regarding the characteristics of the larvæ concerned that in many instances they could not be exactly determined. Fortunately, several workers have undertaken comparative studies along this line. The most comprehensive publication is that of Banks (1912), entitled "The structure of certain dipterous larvæ, with particular reference to those in human food."
Without attempting an exhaustive list, we shall discuss here the more important species of Diptera whose larvæ are known to cause myasis, either external or internal. The following key will serve to determine those most likely to be encountered. The writers would be glad to examine specimens not readily identifiable, if accompanied by exact data relative to occurrence.
a. Body more or less flattened, depressed; broadest in the middle, each segment with dorsal, lateral, and ventral fleshy processes, of which the laterals, at least, are more or less spiniferous ([fig. 101]). Fannia (= Homalomyia).
In F. canicularis the dorsal processes are nearly as long as the laterals; in F. scalaris the dorsal processes are short spinose tubercles.
aa. Body cylindrical, or slender conical tapering toward the head; without fleshy lateral processes ([fig. 105]).
b. With the posterior stigmata at the end of shorter or longer tubercles, or if not placed upon tubercles, then not in pit; usually without a "marginal button" and without a chitinous ring surrounding the three slits; the slits narrowly or broadly oval, not bent ([fig. 171 i]). Acalyptrate muscidæ and some species of Anthomyiidæ. To this group belong the cheese skipper (Piophila casei, figs. [98], [99]), the pomace-fly (Drosophila ampelophila), the apple maggot (Rhagoletis pomonella), the cherry fruit fly (Rhagoletis cingulata), the small dung fly (Sepsis violacea, [fig. 170]), the beet leaf-miner (Pegomyia vicina, [fig. 171 i]), the cabbage, bean and onion maggots (Phorbia spp.) et. al.
bb. Posterior stigmata of various forms, if the slits are narrowly oval ([fig. 171]) then they are surrounded by a chitin ring which may be open ventro-mesally.
c. Integument leathery and usually strongly spinulose; larvæ hypodermatic or endoparasitic. Bot flies ([fig. 171, f, g, k]).—Oestridæ
cc. Integument not leathery and (except in Protocalliphora) spinulæ restricted to transverse patches near the incisures of the segments.
d. The stigmal plates in a pit; the lip-like margin of the pit with a number of fleshy tubercles; chitin of the stigma not complete; open ventro-mesally, button absent ([fig. 171 e]). Flesh flies.—Sarcophaga
dd. Stigmata not in a pit.
e. The chitin ring open ventra-mesally; button absent ([fig. 171 c]). Screw-worm fly. Chrysomyia
ee. The chitin ring closed.
f. Slits of the posterior stigmata straight; marginal "button" present ([fig. 171 b]); two distinct mouth hooks, fleshy tubercles around the anal area. Phormia ([fig. 171 f]), Lucilia and Calliphora ([fig. 172, a, b]), Protocalliphora ([fig. 171, j]), Cynomyia ([fig. 171, a]). Blow flies, bluebottle flies. Calliphorinæ
ff. Slits of the posterior stigmata sinuous or bent. Subfamily Muscinæ.
g. Slits of the posterior stigmata bent; usually two mouth hooks. Muscina stabulans ([fig. 171, l]), Muscina similis, Myiospila meditatunda ([fig. 172, i]), and some of the higher Anthomyiidæ.
gg. Slits of the posterior stigmata sinuous; mouth hooks usually consolidated into one. The house-fly (Musca domestica [fig. 171, d]), the stable fly (Stomoxys calcitrans), the horn fly (Lyperosia irritans), Pyrellia, Pseudopyrellia, Morellia, Mesembrina. Polietes, et. al. ([fig. 172] in part).
Eristalis—The larvæ of Eristalis are the so-called rat-tailed maggots, which develop in foul water. In a few instances these larvæ have been known to pass through the human alimentary canal uninjured. Hall and Muir (1913) report the case of a boy five years of age, who had been ailing for ten weeks and who was under treatment for indigestion and chronic constipation. For some time he had vomited everything he ate. On administration of a vermifuge he voided one of the rat-tailed maggots of Eristalis. He admitted having drunk water from a ditch full of all manner of rotting matter. It was doubtless through this that he became infested. It is worth noting that the above described symptoms may have been due to other organisms or substances in the filthy water.
Piophila casei, the cheese-fly ([fig. 99]), deposits its eggs not only in old cheeses, but on ham, bacon, and other fats. The larvæ ([fig. 98]) are the well-known cheese skippers, which sometimes occur in great abundance on certain kinds of cheese. Indeed, some people have a comfortable theory that such infested cheese is especially good. Such being the case, it is small wonder that this species has been repeatedly reported as causing intestinal myasis. Thebault (1901) describes the case of a girl who, shortly after consuming a large piece of badly infested cheese, became ill and experienced severe pains in the region of the navel. Later these extended through the entire alimentary canal, the excrement was mixed with blood and she suffered from vertigo and severe headaches. During the four following days the girl felt no change, although the excretion of the blood gradually diminished and stopped. On the fourth day she voided two half-digested larvæ and, later, seven or eight, of which two were alive and moving.
That these symptoms may be directly attributed to the larvæ, or "skippers," has been abundantly shown by experimental evidence. Portschinsky cites the case of a dog fed on cheese containing the larvæ. The animal suffered much pain and its excrement contained blood. On post mortem it was found that the small intestine throughout almost its entire length was marked by bloody bruises. The papillæ on these places were destroyed, although the walls were not entirely perforated. In the appendix were found two or three dead larvæ. Alessandri (1910) has likewise shown that the larvæ cause intestinal lesions.
According to Graham-Smith, Austen (1912) has recorded a case of myasis of the nose, attended with a profuse watery discharge of several weeks duration and pain, due to the larvæ of Piophila casei.
Anthyomyiidæ—The characteristic larvæ of two species of Fannia (= Homalomyia or Anthomyia, in part) ([fig. 101]) are the most commonly reported of dipterous larvæ causing intestinal myasis. Hewitt (1912) has presented a valuable study of the bionomics and of the larvæ of these flies, a type of what is needed for all the species concerned in myasis. We have seen two cases of their having been passed in stools, without having caused any special symptoms. In other instances their presence in the alimentary canal has given rise to symptoms vaguely described as those of tapeworm infestation, or helminthiasis. More specifically, they have been described as causing vertigo, severe headache, nausea and vomiting, severe abdominal pains, and in some instances, bloody diarrhœa.
One of the most striking cases is that reported by Blankmeyer (1914), of a woman whose illness began fourteen years previously with nausea and vomiting. After several months of illness she began passing larvæ and was compelled to resort to enemas. Three years previous to the report, she noticed frequent shooting pains in the rectal region and at times abdominal tenderness was marked. There was much mucus in the stools and she "experienced the sensation of larvæ crawling in the intestine." Occipital headaches were marked, with remissions, and constipation became chronic. The appetite was variable, there was a bad taste in the mouth, tongue furred and ridged, and red at the edges. Her complexion was sallow, and general nervousness was marked. As treatment, there were given doses of magnesium sulphate before breakfast and at 4 P. M., with five grain doses of salol four times a day. The customary parasiticides yielded no marked benefit. At the time of the report the patient passed from four to fifty larvæ per day, and was showing some signs of improvement. The nausea had disappeared, her nervousness was less evident, and there was a slight gain in weight.
The case was complicated by various other disorders, but the symptoms given above seem to be in large part attributable to the myasis. There is nothing in the case to justify the assumption that larvæ were continuously present, for years. It seems more reasonable to suppose that something in the habits of the patient favored repeated infestation. Nevertheless, a study of the various cases of intestinal myasis caused by these and other species of dipterous larvæ seems to indicate that the normal life cycle may be considerably prolonged under the unusual conditions.
The best authenticated cases of myasis of the urinary passage have been due to larvæ of Fannia. Chevril (1909) collected and described twenty cases, of which seven seemed beyond doubt. One of these was that of a woman of fifty-five who suffered from albuminuria, and urinated with much difficulty, and finally passed thirty to forty larvæ of Fannia canicularis.
It is probable that infestation usually occurs through eating partially decayed fruit or vegetables on which the flies have deposited their eggs. Wellman points out that the flies may deposit their eggs in or about the anus of persons using outside privies and Hewitt believes that this latter method of infection is probably the common one in the case of infants belonging to careless mothers. "Such infants are sometimes left about in an exposed and not very clean condition, in consequence of which flies are readily attracted to them and deposit their eggs."
Muscinæ—The larvæ of the common house-fly, Musca domestica, are occasionally recorded as having been passed with the feces or vomit of man. While such cases may occur, it is probable that in most instances similar appearing larvæ of other insects have been mistakenly identified.
Muscina stabulans is regarded by Portschinsky (1913) as responsible for many of the cases of intestinal myasis attributed to other species. He records the case of a peasant who suffered from pains in the lower part of the breast and intestines, and whose stools were mixed with blood. From November until March he had felt particularly ill, being troubled with nausea and vomiting in addition to the pain in his intestines. In March, his physician prescribed injections of a concentrated solution of tannin, which resulted in the expulsion of fifty living larvæ of Muscina stabulans. Thereafter the patient felt much better, although he suffered from intestinal catarrh in a less severe form.
Calliphorinæ—Closely related to the Sarcophagidæ are the Calliphorinæ, to which group belong many of the so-called "blue bottle" flies. Their larvæ feed upon dead animals, and upon fresh and cooked meat. Those of Protocalliphora, already mentioned, are ectoparasitic on living nestling birds. Larva of Lucilia, we have taken from tumors on living turtles. To this sub-family belongs also Auchmeromyia luteola, the Congo floor maggot. Some of these, and at least the last mentioned, are confirmed, rather than faculative parasites. Various species of Calliphorinæ are occasionally met with as facultative parasites of man.
Chrysomyia macellaria, the screw worm fly ([fig. 107]), is the fly which is responsible for the most serious cases of human myasis in the United States. It is widely distributed in the United States but is especially abundant in the south. While the larvæ breed in decaying matter in general, they so commonly breed in the living flesh of animals that they merit rank as true parasites. The females are attracted to open wounds of all kinds on cattle and other animals and quickly deposit large numbers of eggs. Animals which have been recently castrated, dehorned, or branded, injured by barbed wire, or even by the attacks of ticks are promptly attacked in the regions where the fly abounds. Even the navel of young calves or discharges from the vulva of cows may attract the insect.
Not infrequently the fly attacks man, being attracted by an offensive breath, a chronic catarrh, or a purulent discharge from the ears. Most common are the cases where the eggs are deposited in the nostrils. The larvæ, which are hatched in a day or two, are provided with strong spines and proceed to bore into the tissues of the nose, even down into or through the bone, into the frontal sinus, the pharynx, larynx, and neighboring parts.
Osborn (1896) quotes a number of detailed accounts of the attacks of the Chrysomyia on man. A vivid picture of the symptomology of rhinal myasis caused by the larvæ of this fly is given by Castellani and Chalmers: "Some couple of days after a person suffering from a chronic catarrh, foul breath, or ozæna, has slept in the open or has been attacked by a fly when riding or driving,—i.e., when the hands are engaged—signs of severe catarrh appear, accompanied with inordinate sneezing and severe pain over the root of the nose or the frontal bone. Quickly the nose becomes swollen, and later the face also may swell, while examination of the nose may show the presence of the larvæ. Left untreated, the patient rapidly becomes worse, and pus and blood are discharged from the nose, from which an offensive odor issues. Cough appears as well as fever, and often some delirium. If the patient lives long enough, the septum of the nose may fall in, the soft and hard palates may be pierced, the wall of the pharynx may be destroyed. By this time, however, the course of the disease will have become quite evident by the larvæ dropping out of the nose, and if the patient continues to live all the larvæ may come away naturally."
For treatment of rhinal myasis these writers recommend douching the nose with chloroform water or a solution of chloroform in sweet milk (10-20 per cent), followed by douches of mild antiseptics. Surgical treatment may be necessary.
Sarcophagidæ—The larvæ ([fig. 105]) of flies of this family usually feed upon meats, but have been found in cheese, oleomargerine, pickled herring, dead and living insects, cow dung and human feces. Certain species are parasitic in insects. Higgins (1890) reported an instance of "hundreds" of larvæ of Sarcophaga being vomited by a child eighteen months of age. There was no doubt as to their origin for they were voided while the physician was in the room. There are many other reports of their occurrence in the alimentary canal. We have recorded elsewhere (Riley, 1906) a case in which some ten or twelve larvæ of Sarcophaga were found feeding on the diseased tissues of a malignant tumor. The tumor, a melanotic sarcoma, was about the size of a small walnut, and located in the small of the back of an elderly lady. Although they had irritated and caused a slight hæmorrhage, neither the patient nor others of the family knew of their presence. Any discomfort which they had caused had been attributed to the sarcomatous growth. The infestation occurred in mid-summer. It is probable that the adult was attracted by the odor of the discharges and deposited the living maggots upon the diseased tissues.
According to Küchenmeister, Sarcophaga carnaria ([fig. 106]), attracted by the odor, deposits its eggs and larvæ in the vagina of girls and women when they lie naked in hot summer days upon dirty clothes, or when they have a discharge from the vagina. In malignant inflammations of the eyes the larvæ even nestle under the eyelids and in Egypt, for example, produce a very serious addition to the effects of small-pox upon the cornea, as according to Pruner, in such cases perforation of the cornea usually takes place.
Wohlfartia magnifica is another Sarcophagid which commonly infests man in the regions where it is abundant. It is found in all Europe but is especially common in Russia, where Portschinsky has devoted much attention to its ravages. It deposits living larvæ in wounds, the nasal fossæ, the ears and the eyes, causing injuries even more revolting than those described for Chrysomyia.
CHAPTER V
ARTHROPODS AS SIMPLE CARRIERS OF DISEASE
The fact that certain arthropods are poisonous, or may affect the health of man as direct parasites has always received attention in the medical literature. We come now to the more modern aspect of our subject,—the consideration of insects and other arthropods as transmitters and disseminators of disease.
The simplest way in which arthropods may function in this capacity is as simple carriers of pathogenic organisms. It is conceivable that any insect which has access to, and comes in contact with such organisms and then passes to the food, or drink, or to the body of man, may in a wholly accidental and incidental manner convey infection. That this occurs is abundantly proved by the work of recent years. We shall consider as typical the case against the house-fly, which has attracted so much attention, both popular and scientific. The excellent general treatises of Hewitt (1910), Howard (1911), and Graham-Smith (1913), and the flood of bulletins and popular literature render it unnecessary to consider the topic in any great detail.
The House-fly As a Carrier of Disease
Up to the past decade the house-fly has usually been regarded as a mere pest. Repeatedly, however, it had been suggested that it might disseminate disease. We have seen that as far back as the sixteenth century, Mercurialis suggested that it was the agent in the spread of bubonic plague, and in 1658, Kircher reiterated this view. In 1871, Leidy expressed the opinion that flies were probably a means of communicating contagious diseases to a greater degree than was generally suspected. From what he had observed regarding gangrene in hospitals, he thought flies should be carefully excluded from wounds. In the same year, the editor of the London Lancet, referring to the belief that they play a useful rôle in purifying the air said, "Far from looking upon them as dipterous angels dancing attendance on Hygeia, regard them rather in the light of winged sponges spreading hither and thither to carry out the foul behests of Contagion."
These suggestions attracted little attention from medical men, for it is only within very recent years that the charges have been supported by direct evidence. Before considering this evidence, it is necessary that we define what is meant by "house-fly" and that we then consider the life-history of the insect.
There are many flies which are occasionally to be found in houses, but according to various counts, from 95 per cent to 99 per cent of these in warm weather in the Eastern United States belong to the one species Musca domestica ([fig. 108]). This is the dominant house-fly the world over and is the one which merits the name. It has been well characterized by Schiner (1864), whose description has been freely translated by Hewitt, as follows:
"Frons of male occupying a fourth part of the breadth of the head. Frontal stripe of female narrow in front, so broad behind that it entirely fills up the width of the frons. The dorsal region of the thorax dusty grey in color with four equally broad longitudinal stripes. Scutellum gray with black sides. The light regions of the abdomen yellowish, transparent, the darkest parts at least at the base of the ventral side yellow. The last segment and a dorsal line blackish brown. Seen from behind and against the light, the whole abdomen shimmering yellow, and only on each side of the dorsal line on each segment a dull transverse band. The lower part of the face silky yellow, shot with blackish brown. Median stripe velvety black. Antennæ brown. Palpi black. Legs blackish brown. Wings tinged with pale gray with yellowish base. The female has a broad velvety back, often reddishly shimmering frontal stripe, which is not broader at the anterior end than at the bases of the antennæ, but become so very much broader above that the light dustiness of the sides is entirely obliterated. The abdomen gradually becoming darker. The shimmering areas on the separate segments generally brownish. All the other parts are the same as in the male."
The other species of flies found in houses in the Eastern United States which are frequently mistaken for the house or typhoid fly may readily be distinguished by the characters of the following key:
a. Apical cell (Rs) of the wide wing open, i.e., the bounding veins parallel or divergent ([fig. 100]). Their larvæ are flattened, the intermediate body segments each fringed with fleshy, more or less spinose, processes. Fannia
b. Male with the sides of the second and third abdominal segments translucent yellowish. The larva with three pairs of nearly equal spiniferous appendages on each segment, arranged in a longitudinal series and in addition two pairs of series of smaller processes ([fig. 100]) F. canicularis
bb. Male with blackish abdomen, middle tibia with a tubercle beyond the middle. The larva with spiniferous appendages of which the dorsal and ventral series are short, the lateral series long and feathered ([fig. 101]) F. scalaris
aa. Apical cell (R) of the wing more or less narrowed in the margin; i. e., the bounding veins more or less converging ([fig. 108]).
b. The mouth-parts produced and pointed, fitted for piercing.
c. Palpi much shorter than the proboscis; a brownish gray fly, its thorax with three rather broad whitish stripes; on each border of the middle stripe and on the mesal borders of the lateral stripes is a blackish brown line. Abdomen yellowish brown; on the second, third and fourth segments are three brown spots which may be faint or even absent. The larvæ live in dung. The stable-fly ([fig. 110]) Stomoxys calcitrans
cc. Palpi nearly as long as the proboscis. Smaller species than the house-fly. The horn-fly ([fig. 167]) Hæmatobia irritans
bb. Mouth-parts blunt, fitted for lapping.
c. Thorax, particularly on the sides and near the base of the wings with soft golden yellow hairs among the bristles. This fly is often found in the house in very early spring or even in the winter. The cluster-fly, Pollenia rudis
cc. Thorax without golden yellow hairs among the bristles.
d. The last segment of the vein M with an abrupt angle. ([fig. 108]). The larvæ live in manure, etc. House-fly, Musca domestica
dd. The last segment of vein M with a broad, gentle curve ([fig. 102]).
e. Eyes microscopically hairy; each abdominal segment with two spots. Larvæ in dung. Myiospila meditabunda
ee. Eyes bare; abdomen gray and brown marbled. Muscina
f. With black legs and palpi. M. assimilis
ff. With legs more or less yellowish; palpi yellow. Larvæ in decaying vegetable substances, dung, etc. M. stabulans
It is almost universally believed that the adults of Musca domestica hibernate, remaining dormant throughout the winter in attics, around chimneys, and in sheltered but cold situations. This belief has been challenged by Skinner (1913), who maintains that all the adult flies die off during the fall and early winter and that the species is carried over in the pupal stage, and in no other way. The cluster-fly, Pollenia rudis, undoubtedly does hibernate in attics and similar situations and is often mistaken for the house-fly. In so far as concerns Musca domestica, the important question as to hibernation in the adult stage is an open one. Many observations by one of the writers (Johannsen) tend to confirm Dr. Skinner's conclusion, in so far as it applies to conditions in the latitude of New York State. Opposed, is the fact that various experimenters, notably Hewitt (1910) and Jepson (1909) wholly failed to carry pupæ through the winter.
The house-fly breeds by preference in horse manure. Indeed, Dr. Howard, whose extensive studies of the species especially qualify him for expressing an opinion on the subject, has estimated that under ordinary city and town conditions, more than ninety per cent of the flies present in houses have come from horse stables or their vicinity. They are not limited to such localities, by any means, for it has been found that they would develop in almost any fermenting organic substance. Thus, they have been bred from pig, chicken, and cow manure, dirty waste paper, decaying vegetation, decaying meat, slaughter-house refuse, sawdust-sweepings, and many other sources. A fact which makes them especially dangerous as disease-carriers is that they breed readily in human excrement.
The eggs are pure white, elongate ovoid, somewhat broader at the anterior end. They measure about one millimeter (1-25 inch) in length. They are deposited in small, irregular clusters, one hundred and twenty to one hundred and fifty from a single fly. A female may deposit as many as four batches in her life time. The eggs hatch in from eight to twenty-four hours.
The newly hatched larva, or maggot ([fig. 108]), measures about two millimeters (1-12 inch) in length. It is pointed at the head end and blunt at the opposite end, where the spiracular openings are borne. It grows rapidly, molts three times and reaches maturity in from six to seven days, under favorable conditions.
The pupal stage, like that of related flies, is passed in the old larval skin which, instead of being molted, becomes contracted and heavily chitinized, forming the so-called puparium ([fig. 108]). The pupal stage may be completed in from three to six days.
Thus during the warm summer months a generation of flies may be produced in ten to twelve days. Hewitt at Manchester, England, found the minimum to be eight days but states that larvæ bred in the open air in horse manure which had an average daily temperature of 22.5° C., occupied fourteen to twenty days in their development, according to the air temperature.
After emergence, a period of time must elapse before the fly is capable of depositing eggs. This period has been tuned the preoviposition period. Unfortunately we have few exact data regarding this period. Hewitt found that the flies became sexually mature in ten to fourteen days after their emergence from the pupal state and four days after copulation they began to deposit their eggs; in other words the preoviposition stage was fourteen days or longer. Griffith (1908) found this period to be ten days. Dr. Howard believes that the time "must surely be shorter, and perhaps much shorter, under midsummer conditions, and in the freedom of the open air." He emphasizes that the point is of great practical importance, since it is during this period that the trapping and other methods of destroying the adult flies, will prove most useful.
Howard estimates that there may be nine generations of flies a year under outdoor conditions in places comparable in climate to Washington. The number may be considerably increased in warmer climates.
The rate at which flies may increase under favorable conditions is astounding. Various writers have given estimates of the numbers of flies which may develop as the progeny of a single individual, providing all the eggs and all the individual flies survived. Thus, Howard estimates that from a single female, depositing one hundred and twenty eggs on April 15th, there may be by September 10th, 5,598,720,000,000 adults. Fortunately, living forms do not produce in any such mathematical manner and the chief value of the figures is to illustrate the enormous struggle for existence which is constantly taking place in nature.
Flies may travel for a considerable distance to reach food and shelter, though normally they pass to dwellings and other sources of food supply in the immediate neighborhood of their breeding places. Copeman, Howlett and Merriman (1911) marked flies by shaking them in a bag containing colored chalk. Such flies were repeatedly recovered at distances of eight to one thousand yards and even at a distance of seventeen hundred yards, nearly a mile.
Hindle and Merriman (1914) continued these experiments on a large scale at Cambridge, England. They "do not think it likely that, as a rule, flies travel more than a quarter of a mile in thickly-housed areas." In one case a single fly was recovered at a distance of 770 yards but a part of this distance was across open fen-land. The surprising fact was brought out that flies tend to travel either against or across the wind. The actual direction followed may be determined either directly by the action of the wind (positive anemotropism), or indirectly owing to the flies being attracted by any odor that it may convey from a source of food. They conclude that it is likely that the chief conditions favoring the disposal of flies are fine weather and a warm temperature. The nature of the locality is another considerable factor. Hodge (1913) has shown that when aided by the wind they may fly to much greater distances over the water. He reports that at Cleveland, Ohio, the cribs of the water works, situated a mile and a quarter, five miles, and six miles out in Lake Erie are invaded by a regular plague of flies when the wind blows from the city. Investigation showed that there was absolutely nothing of any kind in which flies could breed on the crib.
The omnivorous habits of the house-fly are matters of everyday observation. From our view point, it is sufficient to emphasize that from feeding on excrement, on sputum, on open sores, or on putrifying matter, the flies may pass to the food or milk upon the table or to healthy mucous membranes, or uncontaminated wounds. There is nothing in its appearance to tell whether the fly that comes blithely to sup with you is merely unclean, or whether it has just finished feeding upon dejecta teeming with typhoid bacilli.
The method of feeding of the house-fly has an important bearing on the question of its ability to transmit pathogenic organisms. Graham-Smith (1910) has shown that when feeding, flies frequently moisten soluble substances with "vomit" which is regurgitated from the crop. This is, of course, loaded with bacteria from previous food. When not sucked up again these drops of liquid dry, and produce round marks with an opaque center and rim and an intervening less opaque area. Fly-specks, then, consist of both vomit spots and feces. Graham-Smith shows a photograph of a cupboard window where, on an area six inches square, there were counted eleven hundred and two vomit marks and nine fecal deposits.
From a bacteriologist's viewpoint a discussion of the possibility of a fly's carrying bacteria would seem superfluous. Any exposed object, animate or inanimate, is contaminated by bacteria and will transfer them if brought into contact with suitable culture media, whether such substance be food, or drink, open wounds, or the sterile culture media of the laboratory. A needle point may convey enough germs to produce disease. Much more readily may the house-fly with its covering of hairs and its sponge-like pulvilli ([fig. 109]) pick up and transfer bits of filth and other contaminated material.
For popular instruction this inevitable transfer of germs by the house-fly is strikingly demonstrated by the oft copied illustration of the tracks of a fly on a sterile culture plate. Two plates of gelatine or, better, agar medium are prepared. Over one of these a fly (with wings clipped) is allowed to walk, the other is kept as a check. Both are put aside at room temperature, to be examined after twenty-four to forty-eight hours. At the end of that time, the check plate is as clear as ever, the one which the fly has walked is dotted with colonies of bacteria and fungi. The value in the experiment consists in emphasizing that by this method we merely render visible what is constantly occurring in nature.
A comparable experiment which we use in our elementary laboratory work is to take three samples of clean (preferably, sterile) fresh milk in sterile bottles. One of them is plugged with a pledget of cotton, into the second is dropped a fly from the laboratory and into the third is dropped a fly which has been caught feeding upon garbage or other filth. After a minute or two the flies are removed and the vials plugged as was number one. The three are then set aside at room temperature. When examined after twenty-four hours the milk in the first vial is either still sweet or has a "clean" sour odor; that of the remaining two is very different, for it has a putrid odor, which is usually more pronounced in the case of sample number three.
Several workers have carried out experiments to determine the number of bacteria carried by flies under natural conditions. One of the most extended and best known of these is the series by Esten and Mason (1908). These workers caught flies from various sources in a sterilized net, placed them in a sterile bottle and poured over them a known quantity of sterilized water, in which they were shaken so as to wash the bacteria from their bodies. They found the number of bacteria on a single fly to range from 550 to 6,600,000. Early in the fly season the numbers of bacteria on flies are comparatively small, while later the numbers are comparatively very large. The place where flies live also determines largely the numbers that they carry. The lowest number, 550, was from a fly caught in the bacteriological laboratory, the highest number, 6,600,000 was the average from eighteen swill-barrel flies. Torrey (1912) made examination of "wild" flies from a tenement house district of New York City. He found "that the surface contamination of these 'wild' flies may vary from 570 to 4,400,000 bacteria per insect, and the intestinal bacterial content from 16,000 to 28,000,000."
Less well known in this country is the work of Cox, Lewis, and Glynn (1912). They examined over four hundred and fifty naturally infected house-flies in Liverpool during September and early October. Instead of washing the flies they were allowed to swim on the surface of sterile water for five, fifteen, or thirty minutes, thus giving natural conditions, where infection occurs from vomit and dejecta of the flies, as well as from their bodies. They found, as might be expected, that flies from either insanitary or congested areas of the city contain far more bacteria than those from the more sanitary, less congested, or suburban areas. The number of aerobic bacteria from the former varied from 800,000 to 500,000,000 per fly and from the latter from 21,000 to 100,000. The number of intestinal forms conveyed by flies from insanitary or congested areas was from 10,000 to 333,000,000 as compared with from 100 to 10,000 carried by flies from the more sanitary areas.
Pathogenic bacteria and those allied to the food poisoning group were only obtained from the congested or moderately congested areas and not from the suburban areas, where the chances of infestation were less.
The interesting fact was brought out that flies caught in milk shops apparently carry and obtain more bacteria than those from other shops with exposed food in a similar neighborhood. The writers explained this as probably due to the fact that milk when accessible, especially during the summer months, is suitable culture medium for bacteria, and the flies first inoculate the milk and later reinoculate themselves, and then more of the milk, so establishing a vicious circle.
They conclude that in cities where food is plentiful flies rarely migrate from the locality in which they are bred, and consequently the number of bacteria which they carry depends upon the general standard of cleanliness in that locality. Flies caught in a street of modern, fairly high class, workmen's dwellings forming a sanitary oasis in the midst of a slum area, carried far less bacteria than those caught in the adjacent neighborhood.
Thus, as the amount of dirt carried by flies in any particular locality, measured in the terms of bacteria, bears a definite relation to the habits of the people and to the state of the streets, it demonstrates the necessity of efficient municipal and domestic cleanliness, if the food of the inhabitants is to escape pollution, not only with harmless but also with occasional pathogenic bacteria.
The above cited work is of a general nature, but, especially in recent years, many attempts have been made to determine more specifically the ability of flies to transmit pathogenic organisms. The critical reviews of Nuttall and Jepson (1909), Howard (1911), and Graham-Smith (1913) should be consulted by the student of the subject. We can only cite here a few of the more striking experiments.
Celli (1888) fed flies on pure cultures of Bacillus typhosus and declared that he was able to recover these organisms from the intestinal contents and excrement.
Firth and Horrocks (1902), cited by Nuttall and Jepson, "kept Musca domestica (also bluebottles) in a large box measuring 4 × 3 × 3 feet, with one side made of glass. They were fed on material contaminated with cultures of B. typhosus. Agar plates, litmus, glucose broth and a sheet of clean paper were at the same time exposed in the box. After a few days the plates and broth were removed and incubated with a positive result." Graham-Smith (1910) "carried out experiments with large numbers of flies kept in gauze cages and fed for eight hours on emulsions of B. typhosus in syrup. After that time the infested syrup was removed and the flies were fed on plain syrup. B. typhosus was isolated up to 48 hours (but not later) from emulsions of their feces and from plates over which they walked."
Several other workers, notably Hamilton (1903), Ficker (1903), Bertarelli (1910) Faichnie (1909), and Cochrane (1912), have isolated B. typhosus from "wild" flies, naturally infected. The papers of Faichnie and of Cochrane we have not seen, but they are quoted in extenso by Graham-Smith (1913).
On the whole, the evidence is conclusive that typhoid germs not only may be accidentally carried on the bodies of house-flies but may pass through their bodies and be scattered in a viable condition in the feces of the fly for at least two days after feeding. Similar, results have been reached in experiments with cholera, tuberculosis and yaws, the last-mentioned being a spirochæte disease. Darling (1913) has shown that murrina, a trypanosome disease of horses and mules in the Canal zone is transmitted by house-flies which feed upon excoriated patches of diseased animals and then pass to cuts and galls of healthy animals.
Since it is clear that flies are abundantly able to disseminate viable pathogenic bacteria, it is important to consider whether they have access to such organisms in nature. A consideration of the method of spread of typhoid will serve to illustrate the way in which flies may play an important rôle.
Typhoid fever is a specific disease caused by Bacillus typhosus, and by it alone. The causative organism is to be found in the excrement and urine of patients suffering from the disease. More than that, it is often present in the dejecta for days, weeks, or even months and years, after the individual has recovered from the disease. Individuals so infested are known as "typhoid carriers" and they, together with those suffering from mild cases, or "walking typhoid," are a constant menace to the health of the community in which they are found.
Human excrement is greedily visited by flies, both for feeding and for ovipositing. The discharges of typhoid patients, or of chronic "carriers," when passed in the open, in box privies, or camp latrines, or the like, serve to contaminate myriads of the insects which may then spread the germ to human food and drink. Other intestinal diseases may be similarly spread. There is abundant epidæmiological evidence that infantile diarrhœa, dysentery, and cholera may be so spread.
Stiles and Keister (1913) have shown that spores of Lamblia intestinalis, a flagellate protozoan living in the human intestine, may be carried by house-flies. Though this species is not normally pathogenic, one or more species of Entamœba are the cause of a type of a highly fatal tropical dysentery. Concerning it, and another protozoan parasite of man, they say, "If flies can carry Lamblia spores measuring 10 to 7µ, and bacteria that are much smaller, and particles of lime that are much larger, there is no ground to assume that flies may not carry Entamœba and Trichomonas spores."
Tuberculosis is one of the diseases which it is quite conceivable may be carried occasionally. The sputum of tubercular patients is very attractive to flies, and various workers, notably Graham-Smith, have found that Musca domestica may distribute the bacillus for several days after feeding on infected material.
A type of purulent opthalmia which is very prevalent in Egypt is often said to be carried by flies. Nuttall and Jepson (1909) consider that the evidence regarding the spread of this disease by flies is conclusive and that the possibility of gonorrhœal secretions being likewise conveyed cannot be denied.
Many studies have been published, showing a marked agreement between the occurrence of typhoid and other intestinal diseases and the prevalence of house-flies. The most clear-cut of these are the studies of the Army Commission appointed to investigate the cause of epidemics of enteric fever in the volunteer camps in the Southern United States during the Spanish-American War. Though their findings as presented by Vaughan (1909), have been quoted very many times, they are so germane to our discussion that they will bear repetition:
"Flies swarmed over infected fecal matter in the pits and fed upon the food prepared for the soldiers in the mess tents. In some instances where lime had recently been sprinkled over the contents of the pits, flies with their feet whitened with lime were seen walking over the food." Under such conditions it is no wonder that "These pests had inflicted greater loss upon American soldiers than the arms of Spain."
Similar conditions prevailed in South Africa during the Boer War. Seamon believes that very much of the success of the Japanese in their fight against Russia was due to the rigid precautions taken to prevent the spread of disease by these insects and other means.
Veeder has pointed out that the characteristics of a typical fly-borne epidemic of typhoid are that it occurs in little neighborhood epidemics, extending by short leaps from house to house, without regard to water supply or anything else in common. It tends to follow the direction of prevailing winds (cf. the conclusions of Hindle and Merriman). It occurs during warm weather. Of course, when the epidemic is once well under way, other factors enter into its spread.
In general, flies may be said to be the chief agency in the spread of typhoid in villages and camps. In cities with modern sewer systems they are less important, though even under the best of such conditions, they are important factors. Howard has emphasized that in such cities there are still many uncared-for box privies and that, in addition, the deposition of feces overnight in uncared-for waste lots and alleys is common.
Not only unicellular organisms, such as bacteria and protozoa, but also the eggs, embryos and larvæ of parasitic worms have been found to be transported by house-flies. Ransom (1911) has found that Habronema muscæ, a nematode worm often found in adult flies, is the immature stage of a parasite occurring in the stomach of the horse. The eggs or embryos passing out with the feces of the horse, are taken up by fly larvæ and carried over to the imago stage.
Grassi (1883), Stiles (1889), Calandruccio (1906), and especially Nicoll (1911), have been the chief investigators of the ability of house-flies to carry the ova and embryos of human intestinal parasites. Graham-Smith (1913) summarizes the work along this line as follows:
"It is evident from the investigations that have been quoted that house-flies and other species are greatly attracted to the ova of parasitic worms contained in feces and other materials, and make great efforts to ingest them. Unless the ova are too large they often succeed, and the eggs are deposited uninjured in their feces, in some cases up to the third day at least. The eggs may also be carried on their legs or bodies. Under suitable conditions, food and fluids may be contaminated with the eggs of various parasitic worms by flies, and in one case infection of the human subject has been observed. Feces containing tape-worm segments may continue to be a source of infection for as long as a fortnight. Up to the present, however, there is no evidence to show what part flies play in the dissemination of parasitic worms under natural conditions."
Enough has been said to show that the house-fly must be dealt with as a direct menace to public health. Control measures are not merely matters of convenience but are of vital importance.
Under present conditions the speedy elimination of the house-fly is impossible and the first thing to be considered is methods of protecting food and drink from contamination. The first of these methods is the thorough screening of doors and windows to prevent the entrance of flies. In the case of kitchen doors, the flies, attracted by odors, are likely to swarm onto the screen and improve the first opportunity for gaining an entrance. This difficulty can be largely avoided by screening-in the back porch and placing the screen door at one end rather than directly before the door.
The use of sticky fly paper to catch the pests that gain entrance to the house is preferable to the various poisons often used. Of the latter, formalin (40 per cent formaldehyde) in the proportion of two tablespoonfuls to a pint of water is very efficient, if all other liquids are removed or covered, so that the flies must depend on the formalin for drink. The mixture is said to be made more attractive by the addition of sugar or milk, though we have found the plain solution wholly satisfactory, under proper conditions. It should be emphasized that this formalin mixture is not perfectly harmless, as so often stated. There are on record cases of severe and even fatal poisoning from the accidental drinking of solutions.
When flies are very abundant in a room they can be most readily gotten rid of by fumigation with sulphur, or by the use of pure pyrethrum powder either burned or puffed into the air. Herrick (1913) recommends the following method: "At night all the doors and windows of the kitchen should be closed; fresh powder should be sprinkled over the stove, on the window ledges, tables, and in the air. In the morning flies will be found lying around dead or stupified. They may then be swept up and burned." This method has proved very efficaceous in some of the large dining halls in Ithaca.
The writers have had little success in fumigating with the vapors of carbolic acid, or carbolic acid and gum camphor, although these methods will aid in driving flies from a darkened room.
All of these methods are but makeshifts. As Howard has so well put it, "the truest and simplest way of attacking the fly problem is to prevent them from breeding, by the treatment or abolition of all places in which they can breed. To permit them to breed undisturbed and in countless numbers, and to devote all our energy to the problem of keeping them out of our dwellings, or to destroy them after they have once entered in spite of all obstacles, seems the wrong way to go about it."
We have already seen that Musca domestica breeds in almost any fermenting organic material. While it prefers horse manure, it breeds also in human feces, cow dung and that of other animals, and in refuse of many kinds. To efficiently combat the insect, these breeding places must be removed or must be treated in some such way as to render them unsuitable for the development of the larvæ. Under some conditions individual work may prove effective, but to be truly efficient there must be extensive and thorough coöperative efforts.
Manure, garbage, and the like should be stored in tight receptacles and carted away at least once a week. The manure may be carted to the fields and spread. Even in spread manure the larvæ may continue their development. Howard points out that "it often happens that after a lawn has been heavily manured in early summer the occupants of the house will be pestered with flies for a time, but finding no available breeding place these disappear sooner or later. Another generation will not breed in the spread manure."
Hutchinson (1914) has emphasized that the larvæ of houseflies have deeply engrained the habit of migrating in the prepupal stage and has shown that this offers an important point of attack in attempts to control the pest. He has suggested that maggot traps might be developed into an efficient weapon in the warfare against the house-fly. Certain it is that the habit greatly simplifies the problem of treating the manure for the purpose of killing the larvæ.
There have been many attempts to find some cheap chemical which would destroy fly larvæ in horse manure without injuring the bacteria or reducing the fertilizing values of the manure. The literature abounds in recommendations of kerosene, lime, chloride of lime, iron sulphate, and other substances, but none of them have met the situation. The whole question has been gone into thoroughly by Cook, Hutchinson and Scales (1914), who tested practically all of the substances which have been recommended. They find that by far the most effective, economical, and practical of the substances is borax in the commercial form in which it is available throughout the country.
"Borax increases the water-soluble nitrogen, ammonia and alkalinity of manure and apparently does not permanently injure the bacterial flora. The application of manure treated with borax at the rate of 0.62 pound per eight bushels (10 cubic feet) to soil does not injure the plants thus far tested, although its cumulative effect, if any, has not been determined."
As their results clearly show that the substances so often recommended are inferior to borax, we shall quote in detail their directions for treating manure so as to kill fly eggs and maggots.
"Apply 0.62 pound borax or 0.75 pound calcined colemanite to every 10 cubic feet (8 bushels) of manure immediately on its removal from the barn. Apply the borax particularly around the outer edges of the pile with a flour sifter or any fine sieve, and sprinkle two or three gallons of water over the borax-treated manure.
"The reason for applying the borax to the fresh manure immediately after its removal from the stable is that the flies lay their eggs on the fresh manure, and borax, when it comes in contact with the eggs, prevents their hatching. As the maggots congregate at the outer edge of the pile, most of the borax should be applied there. The treatment should be repeated with each addition of fresh manure, but when the manure is kept in closed boxes, less frequent applications will be sufficient. When the calcined colemanite is available, it may be used at the rate of 0.75 pound per 10 cubic feet of manure, and is a cheaper means of killing the maggots. In addition to the application of borax to horse manure to kill fly larvæ, it may be applied in the same proportion to other manures, as well as to refuse and garbage. Borax may also be applied to the floors and crevices in barns, stables, markets, etc., as well as to street sweepings, and water should be added as in the treatment of horse manure. After estimating the amount of material to be treated and weighing the necessary amount of borax, a measure may be used which will hold the proper amount, thus avoiding the subsequent weighings.
"While it can be safely stated that no injurious action will follow the application of manure treated with borax at the rate of 0.62 pound for eight bushels, or even larger amounts in the case of some plants, nevertheless the borax-treated manure has not been studied in connection with the growth of all crops, nor has its cumulative effect been determined. It is therefore recommended that not more than 15 tons per acre of the borax-treated manure should be applied to the field. As truckmen use considerably more than this amount, it is suggested that all cars containing borax-treated manure be so marked, and that public-health officials stipulate in their directions for this treatment that not over 0.62 pound for eight bushels of manure be used, as it has been shown that larger amounts of borax will injure most plants. It is also recommended that all public-health officials and others, in recommending the borax treatment for killing fly eggs and maggots in manure, warn the public against the injurious effects of large amounts of borax on the growth of plants."
"The amount of manure from a horse varies with the straw or other bedding used, but 12 or 15 bushels per week represent the approximate amount obtained. As borax costs from five to six cents per pound in 100-pound lots in Washington, it will make the cost of the borax practically one cent per horse, per day. And if calcined colemanite is purchased in large shipments the cost should be considerably less."
Hodge (1910) has approached the problem of fly extermination from another viewpoint. He believes that it is practical to trap flies out of doors during the preoviposition period, when they are sexually immature, and to destroy such numbers of them that the comparatively few which survive will not be able to lay eggs in sufficient numbers to make the next generation a nuisance. To the end of capturing them in enormous numbers he has devised traps to be fitted over garbage cans, into stable windows, and connected with the kitchen window screens. Under some conditions this method of attack has proved very satisfactory.
One of the most important measures for preventing the spread of disease by flies is the abolition of the common box privy. In villages and rural districts this is today almost the only type to be found. It is the chief factor in the spread of typhoid and other intestinal diseases, as well as intestinal parasites. Open and exposed to myriads of flies which not only breed there but which feed upon the excrement, they furnish ideal conditions for spreading contamination. Even where efforts are made to cover the contents with dust, or ashes, or lime, flies may continue to breed unchecked. Stiles and Gardner have shown that house-flies buried in a screened stand-pipe forty-eight inches under sterile sand came to the surface. Other flies of undetermined species struggled up through seventy-two inches of sand.
So great is the menace of the ordinary box privy that a number of inexpensive and simple sanitary privies have been designed for use where there are not modern sewer systems. Stiles and Lumsden (1911) have given minute directions for the construction of one of the best types, and their bulletin should be obtained by those interested.
Another precaution which is of fundamental importance in preventing the spread of typhoid, is that of disinfecting all discharges from patients suffering with the disease. For this purpose, quick-lime is the cheapest and is wholly satisfactory. In chamber vessels it should be used in a quantity equal to that of the discharge to be treated. It should be allowed to act for two hours. Air-slaked lime is of no value whatever. Chloride of lime, carbolic acid, or formalin may be used, but are more expensive. Other intestinal diseases demand similar precautions.
Stomoxys calcitrans, the stable-fly—It is a popular belief that house-flies bite more viciously just before a rain. As a matter of fact, the true house-flies never bite, for their mouth-parts are not fitted for piercing. The basis of the misconception is the fact that a true biting fly, Stomoxys calcitrans ([fig. 110]), closely resembling the house-fly, is frequently found in houses and may be driven in in greater numbers by muggy weather. From its usual habitat this fly is known as the "stable-fly" or, sometimes as the "biting house-fly."
Stomoxys calcitrans may be separated from the house-fly by the use of the key on p. 145. It may be more fully characterized as follows:
The eyes of the male are separated by a distance equal to one-fourth of the diameter of the head, in the female by one-third. The frontal stripe is black, the cheeks and margins of the orbits silvery-white. The antennæ are black, the arista feathered on the upper side only. The proboscis is black, slender, fitted for piercing and projects forward in front of the head. The thorax is grayish, marked by four conspicuous, more or less complete black longitudinal stripes; the scutellum is paler; the macrochætæ are black. The abdomen is gray, dorsally with three brown spots on the second and third segments and a median spot on the fourth. These spots are more pronounced in the female. The legs are black, the pulvilli distinct. The wings are hyaline, the vein M1+2 less sharply curved than in the house-fly, the apical cell being thus more widely open (cf. [fig. 110]). Length 7 mm.
This fly is widely distributed, being found the world over. It was probably introduced into the United States, but has spread to all parts of the country. Bishopp (1913) regards it as of much more importance as a pest of domestic animals in the grain belt than elsewhere in the United States. The life-history and habits of this species have assumed a new significance since it has been suggested that it may transmit the human diseases, infantile paralysis and pellagra. In this country, the most detailed study of the fly is that of Bishopp (1913) whose data regarding the life cycle are as follows:
The eggs like those of the house-fly, are about one mm. in length. Under a magnifying glass they show a distinct furrow along one side. When placed on any moist substance they hatch in from one to three days after being deposited.
The larva or maggots ([fig. 110]) have the typical shape and actions of most maggots of the Muscid group. They can be distinguished from those of the house-fly as the stigma-plates are smaller, much further apart, with the slits less sinuous. Development takes place fairly rapidly when the proper food conditions are available and the growth is completed within eleven to thirty or more days.
The pupa ([fig. 110]), like that of related flies, undergoes its development within the contracted and hardened last larval skin, or puparium. This is elongate oval, slightly thicker towards the head end, and one-sixth to one-fourth of an inch in length. The pupal stage requires six to twenty days, or in cool weather considerably longer.
The life-cycle of the stable-fly is therefore considerably longer than that of Musca domestica. Bishopp found that complete development might be undergone in nineteen days, but that the average period was somewhat longer, ranging from twenty-one to twenty-five days, where conditions are very favorable. The longest period which he observed was forty-three days, though his finding of full grown larvæ and pupæ in straw during the latter part of March, in Northern Texas, showed that development may require about three months, as he considered that these stages almost certainly developed from eggs deposited the previous December.
The favorite breeding place, where available, seems to be straw or manure mixed with straw. It also breeds in great numbers in horse-manure, in company with Musca domestica.
Newstead considers that in England the stable-fly hibernates in the pupal stage. Bishopp finds that in the southern part of the United States there is no true hibernation, as the adults have been found to emerge at various times during the winter. He believes that in the northern United States the winter is normally passed in the larval and pupal stages, and that the adults which have been observed in heated stables in the dead of winter were bred out in refuse within the warm barns and were not hibernating adults.
Graham-Smith (1913) states that although the stable-fly frequents stable manure, it is probably not an important agent in distributing the organisms of intestinal diseases. Bishopp makes the important observation that "it has never been found breeding in human excrement and does not frequent malodorous places, which are so attractive to the house-fly. Hence it is much less likely to carry typhoid and other germs which may be found in such places."
Questions of the possible agency of Stomoxys calcitrans in the transmission of infantile paralysis and of pellagra, we shall consider later.
Other arthropods which may serve as simple carriers of pathogenic organisms—It should be again emphasized that any insect which has access to, and comes in contact with, pathogenic organisms and then passes to the food, or drink, or the body of man, may serve as a simple carrier of disease. In addition to the more obvious illustrations, an interesting one is the previously cited case of the transfer of Dermatobia cyaniventris by a mosquito ([fig. 81-84]). Darling (1913) has shown that in the tropics, the omnipresent ants may be important factors in the spread of disease.
CHAPTER VI
ARTHROPODS AS DIRECT INOCULATORS OF DISEASE GERMS
We have seen that any insect which, like the house-fly, has access to disease germs and then comes into contact with the food or drink of man, may serve to disseminate disease. Moreover, it has been clearly established that a contaminated insect, alighting upon wounded or abraded surfaces, may infect them. These are instances of mere accidental, mechanical transfer of pathogenic organisms.
Closely related are the instances of direct inoculation of disease germs by insects and other arthropods. In this type, a blood-sucking species not only takes up the germs but, passing to a healthy individual, it inserts its contaminated mouth-parts and thus directly inoculates its victim. In other words, the disease is transferred just as blood poisoning may be induced by the prick of a contaminated needle, or as the laboratory worker may inoculate an experimental animal.
Formerly, it was supposed that this method of the transfer of disease by arthropods was a very common one and many instances are cited in the earlier literature of the subject. It is, however, difficult to draw a sharp line between such cases and those in which, on the one hand, the arthropod serves as a mere passive carrier or, on the other hand, serves as an essential host of the pathogenic organism. More critical study of the subject has led to the belief that the importance of the rôle of arthropods as direct inoculators has been much overestimated.
The principal reason for regarding this phase of the subject as relatively unimportant, is derived from a study of the habits of the blood-sucking species. It is found that, in general, they are intermittent feeders, visiting their hosts at intervals and then abstaining from feeding for a more or less extended period, while digesting their meal. In the meantime, most species of bacteria or of protozoan parasites with which they might have contaminated their mouth-parts, would have perished, through inability to withstand drying.
In spite of this, it must be recognized that this method of transfer does occur and must be reckoned with in any consideration of the relations of insects to disease. We shall first cite some general illustrations and shall then discuss the rôle of fleas in the spreading of bubonic plague, an illustration which cannot be regarded as typical, since it involves more than mere passive carriage.
Some Illustrations of Direct Inoculation of Disease Germs by Arthropods
In discussing poisonous arthropods, we have already emphasized that species which are of themselves innocuous to man, may occasionally introduce bacteria by their bite or sting and thus cause more or less severe secondary symptoms. That such cases should occur, is no more than is to be expected. The mouth-parts or the sting of the insect are not sterile and the chances of their carrying pyogenic organisms are always present.
More strictly falling in the category of transmission of disease germs by direct inoculation are the instances where the insect, or related form, feeds upon a diseased animal and passes promptly to a healthy individual which it infects. Of such a nature are the following:
Various species of biting flies are factors in the dissemination of anthrax, an infectious and usually fatal disease of animals and, occasionally, of man. That the bacteria with which the blood of diseased animals teem shortly before death might be transmitted by such insects has long been contended, but the evidence in support of the view has been unsatisfactory. Recently, Mitzmain (1914) has reported a series of experiments which show conclusively that the disease may be so conveyed by a horse-fly, Tabanus striatus, and by the stable-fly, Stomoxys calcitrans.
Mitzmain's experiments were tried with an artificially infected guinea pig, which died of the disease upon the third day. The flies were applied two and one-half hours, to a few minutes, before the death of the animal. With both species the infection was successfully transferred to healthy guinea pigs by the direct method, in which the flies were interrupted while feeding on the sick animal. The evidence at hand does not warrant the conclusion that insect transmission is the rule in the case of this disease.
The nagana, or tsetse-fly disease of cattle is the most virulent disease of domestic animals in certain parts of Africa. It is caused by a protozoan blood parasite, Trypanosoma brucei, which is conveyed to healthy animals by the bite of Glossina morsitans and possibly other species of tsetse-flies. The flies remain infective for forty-eight hours after feeding on a diseased animal. The insect also serves as an essential host of the parasite.
Surra, a similar trypanosomiasis affecting especially horses and mules, occurs in southern Asia, Malaysia, and the Philippines where the tsetse-flies are not to be found. It is thought to be spread by various species of blood-sucking flies belonging to the genera Stomoxys, Hæmatobia, and Tabanus. Mitzmain (1913) demonstrated that in the Philippines it is conveyed mechanically by Tabanus striatus.
The sleeping sickness of man, in Africa, has also been supposed to be directly inoculated by one, or several, species of tsetse-flies. It is now known that the fly may convey the disease for a short time after feeding, but that there is then a latent period of from fourteen to twenty-one days, after which it again becomes infectious. This indicates that in the meantime the parasite has been undergoing some phase of its life-cycle and that the fly serves as an intermediate host. We shall therefore consider it more fully under that grouping.