The Animal Parasites of Man by Fantham, Braun, Stephens, and Theobald
1889. Stiles, in a work recently published, states that there were
25865 words | Chapter 17
8,491 cases of trichinosis with 513 deaths (6·04 per cent.) in Germany
from 1860 to 1880; and 6,329 cases and 318 deaths (5·02 per cent.)
between 1881–1898. Of these latter, 1881–1898, 3,822 (225 deaths)
occurred in Prussia, 1,634 (76 deaths) in Saxony, and 873 (17 deaths)
in the remaining states. There is, however, no doubt that many deaths
from trichinosis were not recognized, as proved by experience at
_post-mortems_.
_Prophylaxis._--The grave nature of the disease and the comparatively
high mortality relating to trichinosis led the authorities to adopt
certain preventive measures, which are the more necessary as national
customs cannot be altered in a short time. As the usual process of
pickling and smoking, even when long continued, does not certainly
ensure the death of the Trichinellæ contained in the meat, and also
because in roasting and boiling large pieces of pork a considerable
time is necessary to permit the temperature required to kill off the
parasites (62° to 70° C.) to penetrate to the middle of the joint,
it appeared to be most practical to have all pigs microscopically
examined for Trichinellæ before they, or parts of them, were placed
on the market, and all infected meat condemned, no matter whether the
Trichinellæ were present in large or small numbers, still undeveloped
or calcified. Since 1877 obligatory examination of pork has been
introduced in Prussia, though as yet it is not thoroughly carried
out; other states of North Germany as well as the larger towns of
South Germany soon followed; a complete army of trichina inspectors,
officially examined and periodically controlled by experts, and
whose number in Prussia amounted to 27,602 in 1896, this being even
increased to 28,224 in 1899, have the charge of examining pork on
certain lines laid down. These are at the present time uniformly
administered. The proceeding is usually that the trichina inspector
himself goes to the slaughterhouses, or special samplers take pieces
of the muscles that are known to be the favourite seats of the
parasite (pillars of the diaphragm, the costal part of the diaphragm,
muscles of the tongue and larynx, intercostal and abdominal muscles);
six small portions are separated from each piece, pressed between
slides or special compressors, and carefully gone through by
examining them with a low power of the microscope. The pigs free
from Trichinellæ are passed for commerce; trichinous pigs, on the
other hand, in Prussia, are only allowed to be used for industrial
purposes, _i.e._, the hide and bristles are used, the fat is allowed
to be melted down, or certain parts are used for the manufacture of
soap or glue. In Saxony, however, it is still permitted to place
trichinous flesh on the market, fully declaring its nature, and after
having been heated to its deepest strata at a temperature of 100° C.
in a suitable apparatus, and under the supervision of a veterinary
surgeon.
AS TO THE PROPORTION OF TRICHINOUS PIGS to healthy ones, the
following tables give the figures for Prussia:--
------+---------------+-----------------+------------
| Number of | Number of |
Year | pigs examined | trichinous pigs | Proportion
------+---------------+-----------------+------------
1878 | 2,524,105 | 1,222 | 1 : 2,065
1879 | 3,164,656 | 1,938 | 1 : 1,632
1881 | 3,118,780 | 1,695 | 1 : 1,839
1882 | 3,808,142 | 1,852 | 1 : 2,056
1883 | 4,248,767 | 2,199 | 1 : 1,932
1884 | 4,611,689 | 2,624 | 1 : 1,741
1885 | 4,421,208 | 2,387 | 1 : 1,852
1886 | 4,834,898 | 2,114 | 1 : 2,287
1887 | 5,486,416 | 2,776 | 1 : 1,988
1888 | 6,051,249 | 3,111 | 1 : 1,945
1889 | 5,500,678 | 3,026 | 1 : 1,818
1890 | 5,590,510 | 1,756 | 1 : 3,183
1891 | 6,550,182 | 2,187 | 1 : 2,996
1892 | 6,234,559 | 2,085 | 1 : 2,992
1896 | 8,759,490 | 1,877 | 1 : 4,666
1899 | 9,230,353 | 1,021 | 1 : 9,040
1902 | 9,093,210 | 725 | 1 : 12,397
------+---------------+-----------------+------------
The proportion, however, is not only subject to variation in separate
years, but differs according to the district; thus, in 1884, in the
state district of Minden there was one trichinous pig to 30,146
healthy animals, in Erfurt 1 to 14,563, in the district of Gnesen 1
to 101, in Schrimm 1 to 86, and in Schroda 1 to 68.
In Germany Trichinella is becoming LESS COMMON in pigs (Ostertag):--
(_a_) _Prussia._
Pigs found
Year to be trichinous
1878–1885 0·061–0·048 per cent.
1886–1892 0·033–0·043 "
1896 0·021 "
1899 0·014 "
1902 0·011 "
(_b_) Saxony.
Number of pigs
Year found to be trichinous
1891 0·014 per cent.
1892 0·011 "
1893 0·008 "
1894 0·007 "
1895 0·012 "
1896 0·0102 "
1899 0·004 "
1902 0·0056 "
(_c_) City of Berlin.
Number of pigs
Year found to be trichinous
1883–1893 0·035–0·064 per cent.
1893–1897 0·022–0·015 "
1902 0·006 "
There is no doubt that the excellent preventive measure of official
inspection for Trichinella has led to the avoidance of grave disasters;
its introduction has not yet caused an entire cessation of trichinosis
in man, because inspection of pork is not obligatory everywhere, so
that human beings may become infected by unexamined trichinous pigs
from their own country or from abroad, and also because an infection
may occasionally escape notice. For these reasons the meat imported
into Berlin from abroad as free from Trichinæ is examined again and not
always in vain; finally, also, gross negligence may at times occur, or
fatal errors may be made.
In addition _private prophylaxis_ must not be neglected, and its chief
aim should be directed to the suitable preparation of pork.
Family. *Dioctophymidæ.*
Genus. *Dioctophyme*, Collet-Megret, 1802.
Syn.: _Eustrongylus_, Dies., 1851.
Large worms. Anterior extremity unarmed; the mouth is surrounded by
six papillæ. One ovary. The vulva is in the anterior region of the
body.
*Dioctophyme gigas*, Rudolphi, 1802.
Syn.: _Dioctophyme renale_, Goeze, 1782; _Ascaris canis_ et _martis_,
Schrank, 1788; _Ascaris visceralis_ et _renalis_, Gmelin, 1789;
_Strongylus gigas_, Rud., 1802; _Eustrongylus gigas_, Dies., 1851;
_Strongylus renalis_, Moq. Tand., 1860; _Eustrongylus visceralis_,
Raill., 1885.
Colour blood-red; the anterior extremity somewhat slender; there is a
series of about 150 papillæ along the lateral lines; the sub-median
lines are strongly developed, and from them spring the radial muscles
for the intestine.
The males attain a length of 40 cm. and a diameter of 4 to 6 mm.; the
posterior extremity is transversely truncated; the anal orifice is
within the base of the collar-like bursa, the thickened edges of which
are beset with papillæ; the spicule measures 5 to 6 mm. in length.
The females attain a length of 100 cm. and a breadth of 12 mm. The anus
is crescent-shaped and terminal. The vulva is 50 to 70 mm. distant
from the anterior extremity. The eggs are oval and have a thick shell
presenting numerous depressions; the shell itself is brownish, but it
is colourless at the somewhat thickened poles; it measures 60 µ in
length by 40 µ in breadth. The larva measures 240 µ by 14 µ.
_Dioctophyme gigas_ lives in the pelvis of the kidney, more rarely in
the abdominal cavity of the seal, otter, dog, wolf, fox, horse, marten
and polecat, exceptionally also in human beings. It also occurs in
tumours of the mamma and perinæum. Most of the cases in which this
parasite has been reported as occurring in man may be traced back
to unrecognized _Ascaris lumbricoides_ or to clots of fibrin; seven
certain cases, eight more or less doubtful, however, remain.
[Illustration: FIG. 306.--_Dioctophyme gigas_, male. Natural size.
(After Railliet.)]
[Illustration: FIG. 307.--Eggs of _Dioctophyme gigas_; above seen from
the flat, below in optical section. 400/1. (After Railliet.)]
The source of infection is unknown, but according to Balbiani the
eggs develop an embryo in water or moist soil, and this embryo may
remain alive several years without hatching; the infection of dogs
with embryo-containing eggs did not succeed; an intermediate stage in
fishes is conjectured, but still the infection of cattle and horses is
unintelligible.
Family. *Strongylidæ.*
Sub-family. *Metastrongylinæ*, Leiper, 1908.
Buccal capsule absent or slightly developed, vagina elongate,
uteri convergent[307] and have a simple musculature. Parasitic in
the respiratory or circulatory system. Genera: Metastrongylus,
Synthetocaulus.
[307] Convergent: _i.e._, the uteri are parallel, converging from the
anterior part of body to the vagina, which is near the anus, this
position being associated with convergence of the uteri. Divergent:
Uteri run anterior and posterior, diverging from the vagina, which in
this case is near middle of body.
Genus. *Metastrongylus*, Molin, 1861.
Mouth with six lips, of which the two lateral are the largest. Postero-
and postero-external rays[308] of bursa thin, the rest thick. Only the
median ray double. Spicules very long and slender, striated. Vulva
immediately in front of anus. Eggs contain an embryo when laid.
[308] For nomenclature of rays _vide_ p. 449.
*Metastrongylus apri*, Gmelin, 1789.
Syn.: _Gordius pulmonalis apri_, Ebel, 1777; _Ascaris apri_, Gmelin,
1789; _Strongylus suis_, Rud., 1809; _Strongylus paradoxus_,
Mehlis, 1831; _Strongylus elongatus_, Duj., 1845; _Strongylus
longevaginatus_, Dies., 1851.
The male measures 12 to 25 mm. in length; the bursa is bilobed; there
are five rays in each lobe; the spicules are thin and up to 4 mm. in
length. The females measure 20 to 50 mm. in length, the anus is close
in front of the posterior extremity, which has a recurved, hook-like
process; the vulva is close in front of the anus. The eggs are
elliptical, 57 µ to 100 µ in length, 39 µ to 72 µ in breadth; when the
eggs are deposited the embryo is already formed, 220 µ to 350 µ by 10 µ
to 12 µ.
_Metastrongylus apri_ frequently lives in the bronchial tubes--usually
the smaller ones--of the pig[309] and wild boar; it is also found
occasionally in sheep and in man; in young pigs it is apt to set up a
bronchitis, which frequently causes death.
[309] The reports of the city inspection of meat in Berlin state that
_Strongylidæ_ in the lungs of pigs are by no means rare; therefore the
lungs of 1,941 pigs were condemned between 1885–1886, of 1,641 between
1886–1887, of 3,237 between 1887–1888, of 4,855 between 1888–1889, of
7,197 between 1889–1890, and of 5,574 pigs between 1890–1891, etc.
Ostertag found _Strongylus apri_ in 60 per cent. of the pigs examined
in the Berlin abattoir; Meyer, in Leipzig, found the parasite in 15 per
cent. of the native pigs and in 52 per cent. of the Hungarian pigs.
The first communication as to the occurrence of this species in man
was that of Diesing, who, in 1845, in Klausenburg, had the opportunity
of examining _Strongylidæ_ found by Jortsits in the lung of a little
boy, aged 6, in Transylvania; probably also the Nematodes found in the
trachea and larynx of man, and described by Rainey and Bristowe as
specimens of _Filaria trachealis_, belong to this group; according to
Chatin, _Metastrongylus apri_ may also occur in the intestine of man;
this occurrence, however, may in all probability have been due to an
accidental introduction of adult worms into the intestine, and should
not be attributed to an infection by the larval stage.
[Illustration: FIG. 308.--_Metastrongylus apri_: one side of bursa.
_a._, anterior; _a.e._, antero-external; _a.m._, antero-median; _p.m._,
postero-median; _p.e._, postero-external; _p._, one division of
posterior ray. (Stephens.)]
No experiments to induce infection have been made; it is probable,
however, that infection is direct and without the aid of an
intermediate host.
Sub-family. *Trichostrongylinæ*, Leiper, 1908.
Strongylidæ with buccal capsule absent or slightly developed, vagina
short, uteri divergent (_i.e._, anterior and posterior), ovejectors
differentiated. Parasitic in the alimentary canal. Contains the genera
Trichostrongylus, Hæmonchus, Ostertagia, Nematodirus, Cooperia,
Dictyocaulus.[310]
[310] _Dictyocaulus_ is parasitic in the bronchi.
Genus. *Trichostrongylus*, Looss, 1905.
Very small _Strongylidæ_. Mouth with three small lips and nodular or
punctiform papillæ. Cervical papillæ absent. Bursa entirely closed,
with large lateral lobes, and median lobe not distinctly defined.
Anterior[311] rays double, the branches widely divergent, one thin, the
other thick, and close to the antero-median. The postero-median ray
is thin and close to the postero-external. Posterior ray bifurcate,
each branch bifid at the tip (fig. 311). Spicules short, spoon or
spatula-like, with on the broad anterior end a lateral knob or disc
and in front of the point an angular projection. Gubernaculum of a
peculiar canoe or shoe shape in profile. Vulva in the hinder half of
the body. Tail with two minute papillæ just in front of tip. Eggs thin
shelled; when laid they show eight to thirty-two segments. Parasitic in
duodenum, seldom in the stomach of herbivora.
[311] When the anterior ray is double, the branches of it are called
antero-anterior and latero-anterior.
[Illustration: FIG. 309.--_Trichostrongylus instabilis_: left,
posterior end of male; right, spicule and gubernaculum, side view.
_Cf._ fig. 311. Magnified. (After Looss.)]
[Illustration: FIG. 310.--_Trichostrongylus instabilis_: posterior end
of female. Magnified. (After Looss.)]
*Trichostrongylus instabilis*,[312] Railliet, 1893.
[312] Identical with _T. colubriformis_ of the sheep according to
Leiper. If so, this latter name has priority.
Syn.: _Strongylus instabilis_, Railliet, 1893; _Strongylus subtilis_,
Looss, 1895.
Male 4 to 5·5 mm. long, 0·08 mm. thick in front of bursa. Spicule 0·135
to 0·145 mm. long, accessory piece (gubernaculum) 0·07 mm. thick.
Antero-external ray usually thickest of all, occasionally only as
thick as the antero-median; postero-median far more slender than the
antero-external and antero-median and nearer to the postero-external
than to the antero-median. Female 5 to 6 mm. long, vulva 1·05 to
1·2 mm. distant from the tip of the tail, placed _longitudinally_,
50 µ to 55 µ long, always shorter than the unpaired portion of the
canal formed by the union of the two ovejectors; anus 0·055 to 0·07 mm.
distant from tip of the tail; ova 73 µ to 80 µ by 40 µ to 43 µ.
This species lives in the duodenum, exceptionally also in the
stomach of _Ovis aries_, _O. laticauda_, _Antilope dorcas_, _Camelus
dromedarius_ (Egypt), _Cynocephalus hamadryas_ (North Africa), sheep
and goats (France), and has been found by Looss in bodies of fellaheen
at Alexandria and in the stomach of a Japanese female by Ijima.
*Trichostrongylus probolurus*, Railliet, 1896.
Syn.: _Strongylus probolurus_, Railliet, 1896.
Male 4·5 to 5·5 mm. long, in front of bursa 0·08 mm. thick; spicule
0·126 to 0·134 mm. long, gubernaculum 0·075 to 0·08 mm. long.
Bursa: latero-anterior rib thickest; antero-external thicker than
antero-median, postero-median and postero-external very short and close
together. Female 4·5 to 6 mm. long, vulval opening 1·08 to 1·25 mm.
from tip of tail, placed _longitudinally_, and slightly curved, 76 µ
long, always longer than the unpaired portion of the ovejector; anus
0·040 to 0·05 mm. distant from tip of tail. Posterior end thick, point
of tail short. Ova 76 µ to 80 µ by 43 µ to 46 µ.
[Illustration: FIG. 311.--_Trichostrongylus probolurus_: tail of male
from left side. _d._, posterior; _e.d._, postero-external; _p.l._,
postero-median; _m.l._, antero-median; _e.l._, antero-external;
_l.v._, latero-anterior; _v.v._, antero-anterior; _gub._, portion of
gubernaculum; _sp._, portion of spicules. × c. 300. (After Looss.)]
[Illustration: FIG. 312.--_Trichostrongylus probolurus_: spicules and
gubernaculum of male; on left, ventral view; on right, lateral view. ×
c. 300. (After Looss.)]
_Habitat._--In the duodenum of _Ovis aries_, _O. laticauda_, _Antilope
dorcas_, _Camelus dromedarius_ (Egypt) and occasionally also in man
(Egypt).
*Trichostrongylus vitrinus*, Looss, 1905.
Male 4 to 5·5 mm. long, in front of bursa 0·085 mm. thick. Bursa
larger than in the other two species, antero-external rib thickest,
antero-anterior and postero-median equally thick, straight. Spicule
0·16 to 0·17 mm. long, gubernaculum 0·085 to 0·095 mm. long. Female 5
to 6·5 mm. long, vulval opening 1·15 to 1·25 mm. distant from tip of
tail, crescent shaped, _oblique_ to body axis, and around it irregular
thickenings. Ova 84 µ to 90 µ by 46 µ to 50 µ.
In duodenum of _Ovis aries_, _O. laticauda_, occasionally in _Camelus
dromedarius_ and in man (Egypt).
[Illustration: FIG. 313.--_Trichostrongylus vitrinus_: tail of male
from left side. _d._, posterior; _e.d._, postero-external; _p.l._,
postero-median; _m.l._, antero-median; _e.l._, antero-external;
_l.v._, latero-anterior; _v.v._, antero-anterior; _gub._, portion of
gubernaculum; _sp._, portion of spicule. × c. 300. (After Looss.)]
[Illustration: FIG. 314.--_Trichostrongylus vitrinus_: spicules and
gubernaculum; on left, ventral view; on right, lateral view. × c. 300.
(After Looss.)]
Genus. *Hæmonchus*, Cobb., 1898.
Small mouth cavity contains a “tooth” or “lancet” arising from the
dorsal side. Cuticle of head and neck not inflated. Cervical papillæ
well marked. Bursa bilateral, with large lateral lobes and a small
dorsal lobe _not median_, but lateral, attached to the base of one of
the lateral lobes (fig. 316). Posterior ray bifurcate, each branch
bifid apically. Each lateral lobe six rays. Anterior rays separated
distally, curving forward. Antero-median and postero-median rays
distally curve away from the antero-external. Postero-external ray long
and slender. Spicules less than 1 mm. Gubernaculum present. Vulva in
posterior part of body covered by a prominent tongue-like flap. Eggs
ellipsoidal.
*Hæmonchus contortus*, Rudolphi, 1803; Cobb., 1898.
Dorsal “tooth” or “lancet” 10 µ to 15 µ long. Cervical papillæ 0·3 mm.
from head.
Male 20 mm. long by 400 µ thick (maximum). Asymmetrical lobe of bursa
150 µ by 125 µ attached to left lateral lobe. Posterior ray bifurcate;
each branch bifid. Stem of ray less than twice as long as its branches.
Spicules 300 µ to 500 µ with knobbed tips, and the left spicule with
a barb 20 µ from the tip, right spicule with a barb 40 µ from tip.
Gubernaculum 200 µ by 25 µ to 35 µ, fusiform with thickened edges.
Female 18 to 30 mm. by 500 µ (maximum). Vulva 3 to 4·5 mm. from tip.
Linguiform flap 0·5 mm. (a second one exists, according to Brumpt).
Anus 400 µ to 630 µ from tip. Tail acutely pointed. Eggs 75 µ to 95 µ
by 40 µ to 50 µ.
[Illustration: FIG. 315.--_Hæmonchus contortus_: vulval region of
female viewed from left side. _int._, intestine; _lab._, linguiform
process covering vulva; _ov._, ovary; _ovij._, ovejector; _ut._,
uterus; _vag._, vagina; _vul._, vulva. × 75. (After Ransom.)]
[Illustration: FIG. 316.--_Hæmonchus contortus_: tail of male,
dorsal view, _d._, posterior ray of the asymmetrically placed posterior
lobe; _e.d._, postero-external; _p.l._, postero-median; _m.l._,
antero-median; _e.l._, antero-external; _l.v._, latero-anterior;
_v.v._, antero-anterior; _gub._, gubernaculum; _sp._, spicule. × 75.
(After Ransom.)]
_Habitat._--Fourth stomach of cattle, sheep, antelope.
_Distribution._--Europe, America, Africa, Asia, Australia, New Zealand.
Once in man in South America by de Magalhães.
_Pathology._--Produces anæmia, emaciation, dropsy in sheep; and in the
human case the symptoms were mistaken for those of ancylostomiasis.
_Life-history._--Rhabditic embryos easily hatch in water, then moult
several times, becoming eventually “filariform” larvæ enclosed in the
moulted skin. These crawl up blades of grass and are swallowed by
sheep, etc.
Genus. *Nematodirus*, Ransom, 1907, emend. Railliet, 1912.
Head over 50 µ in diameter. Cuticle may be slightly inflated and often
transversely striated. Cuticle with eighteen distinct longitudinal
ridges. Cervical papillæ absent (?). Posterior lobe of bursa reduced to
short lobules each with a dorsal ray. Antero-anterior + latero-anterior
(= anterior double) rays close together, parallel; antero-external
ray diverges widely from antero- and postero-median, which are close
together and parallel. Postero-external ray slender. Spicules more
than 0·5 mm. long, at most one-twelfth of body, united by a membrane
throughout their length or only distally. Gubernaculum absent. Vulva
behind middle of body. Eggs ellipsoidal, shell rather thick.
_Habitat._--Duodenum of ruminants.
Sub-genus. *Mecistocirrus*, Railliet, 1912.
Head slightly inflated, with transverse striations. Skin with eighteen
longitudinal ridges, but little apparent; cervical papillæ distinct.
Bursa bilobed; median ray double (= postero-median + antero-median);
very large antero-external at the edge, close to the anterior. Spicules
very long, slender, one-sixth length of body (3·5 mm.); tail pointed.
Vulva immediately in front of anus.
_Habitat._--Stomach of ruminants.
*Mecistocirrus fordi*, Daniels, 1908.
Syn.: _Strongylus fordi_, Daniels, 1908; _Strongylus gibsoni_,
Stephens, 1909; _Nematodirus fordi_, Leiper, 1911.
Male 21 mm. long by 0·4 mm. thick. Cervical papillæ 0·45 mm. behind the
head. Spicules about 7 mm. long, _i.e._, one-third of the body length.
At the level of the postero-external rays of the bursa, the bursa has a
projecting lobule.
Female 25 mm. long. Anus 0·2 mm., vulva 0·5 mm. from the tip of tail.
Eggs 100 µ by 53 µ.
Sub-family. *Ancylostominæ*, Railliet, 1909.
_Strongylidæ_ with buccal capsule, well developed. Uteri divergent.
Parasitic in the alimentary canal, exceptionally in the respiratory
system.
Group. *Œsophagostomeæ*, Railliet and Henry, 1909.
Bursa with anterior and median ray cleft (not double), postero- and
postero-external arising from a common trunk, posterior bifurcated,
each limb bidigitate.
Contains at present four genera: (1) Ternidens, (2) Chabertia, (3)
Œsophagostomum, (4) Agriostomum.
[Illustration: FIG. 317.--Mecistocirrus fordi: bursa of male, dorsal
view. The rays are (1) postero-external, (2) median (= postero-median
+ antero-median), (3) antero-external, (4) latero-anterior, (5)
antero-anterior. These two latter are parallel. The posterior ray is
absent. (After Stephens.)]
Genus. *Ternidens*, Railliet, 1909.
Buccal capsule sub-globular, opening obliquely in the dorsal surface,
and having at the bottom three complex teeth resembling those of
Triodontophorus.[313] Two crowns of leaflets; peristomic collar
moderate, edge of bursa slightly toothed.
[313] Triodontophorus belongs to the group _Cylicostomeæ_, which has
the following bursal formula: (1) anterior cleft, (2) median double,
(3) postero-external and posterior arising _separately_, (4) posterior
double, each branch giving off two lateral branches.
_Type._--_T. deminutus_, Railliet and Henry.
*Ternidens deminutus*, Raill. and Henry, 1905.
Syn.: _Triodontophorus deminutus_, Raill. and Henry, 1905.
Body relatively thick. Cervical papillæ 0·5 mm. behind the head. Buccal
capsule 40 µ deep. Teeth 40 µ long.
[Illustration: FIG. 318.--_Ternidens deminutus._ *A*, head end, ventral
view: c, crown of leaflets; v.o., buccal cavity; d, pharyngeal plates;
ph., pharynx; n., valve. *B*, lateral view. *C*, tail of female. *D*,
bursa of male: a., anterior ray; a.e., antero-external; m., median;
p.e., postero-external; p., posterior. *E*, pharyngeal plate. Enlarged.
(After Railliet and Henry.)]
Male 9·5 mm. long by 560 µ thick. Œsophagus 660 µ long. Bursa broader
than long, the lateral lobes united by a small posterior lobe with
slightly sinuous margin; edge of bursa finely toothed. Spicules about
900 µ long.
Female 12 to 16 mm. long by 650 µ to 730 µ thick. Œsophagus 860 µ long.
Vulva forms a distinct projection 480 µ from tip of tail. Anus 240 µ to
270 µ from tip. Eggs 60 µ to 80 µ by 40 µ.
_Habitat._--Large intestine of a negro (Comoro Islands) and in the
natives of Nyasaland and Portuguese East Africa. Also in large
intestine of _Macacus sinensis_ and _Macacus cynomolgus_.
Genus. *Œsophagostomum*, Molin, 1861.
No teeth. Cuticle around the mouth dilated to form a narrow cuticular
“peristomic collar.” Separated by a constriction from this is a much
more extensive inflation, the “cephalic vesicle,” bounded abruptly
behind on the ventral side by a transverse groove, the “ventral cleft,”
which is always present even in absence of the vesicle. Buccal cavity
of slight depth with a short dorsal tunnel. Internal margin of the
mouth armed with chitinous leaflets (“external crown”); internal border
of the buccal capsule armed with short tongue-like leaflets (internal
crown). Lateral membranous wings may extend backwards from the
ventral cleft. Cervical papillæ present. Bursa with two lateral lobes
united by a smaller median lobe. Spicules over 5 mm. long, slender;
gubernaculum inconspicuous. Vulva in front of anus. Adults usually in
large intestine of ruminants, suidæ, tapirs, edentates and apes. Larvæ
sometimes in nodules in intestinal wall.
*Œsophagostomum brumpti*, Railliet and Henry, 1905.
Female immature, 8·5 to 10·2 mm. long, 0·295 to 0·325 mm. thick.
Cuticle transversely striated. The cephalic vesicle immediately
behind the vestibulum oris, embracing the anterior two-fifths of the
œsophagus, extending ventrally, however, towards its posterior end.
Vestibulum oris formed by a cuticular band provided with a crown of
twelve apical leaflets directed forward and inwards; six cephalic
papillæ (two lateral, four submedian); buccal capsule in front of
cervical swelling not delineated circularly behind, but provided with
three wide incisions (one dorsal, two sub-ventral). Œsophagus, 0·470
to 0·500 mm. long, two cervical papillæ at five-eighths of its length.
Vulva 0·350 to 0·475 mm., anus 0·170 to 0·200 mm., before tip of tail.
_Habitat._--Found by Brumpt in tumours of the cæcum and colon of a
native of the River Omo (Lake Rudolph), East Africa. Immature forms
only were present. Adults have been found in similar tumours in monkeys.
[Illustration: FIG. 319.--_Œsophagostomum stephanostomum_ var.
_thomasi_. 1, male, natural size; 2, female, natural size; 3, head
of female, ventral view, showing cephalic vesicle and ventral cleft
limiting it behind, × 55; 4, head of female, dorsal view, × 225; 5,
head of male, end view, showing external and internal leaf crowns,
× 225; 6, tail of male, lateral view (_cf._ fig. 318, D), × 20; 7,
tail of female, lateral view, × 20; 8, _Œs. thomasi_, posterior ray of
bursa, × 150; 9, _Œs. dentigerum_, from chimpanzee, posterior ray of
bursa, × 150; 10, _Œs. stephanostomum_, from gorilla, posterior ray of
bursa, × 150.]
_Pathology._--They occur in hæmorrhagic cysts in the submucosa or
muscularis mucosæ of the gut wall. The cysts project internally and
externally, and contain immature adults, which eventually escape into
the lumen of the gut.
[Illustration: FIG. 320.--_Œsophagostomum stephanostomum_ var.
_thomasi_: cæcum and ascending colon. Subperitoneal cysts are seen on
the top right hand, and in the lumen of the gut numerous cysts arranged
transversely. The small roundish patches are areas of necrosis in the
cyst walls. (After Thomas.)]
*Œsophagostomum stephanostomum* var. *thomasi*, Raill. and Henry, 1909.
Body thick, pointed only at the ends. Buccal capsule much reduced.
External crown of thirty-eight leaflets (the “crown” nearest the centre
of fig. 319, 5). Male 17 to 22 mm. long by 750 µ thick. Spicule 1·380
to 1·475 mm., slightly curved at the tip. Female, immature, 16 to
20 mm. long by 900 µ thick, tail ending in a little conical appendage.
Anus 230 µ, vulva 500 µ to 525 µ from tip. Ovejectors close together.
Uteri very short in form of oblong pouch.
_Œs. stephanostomum_, Stossich, 1904, in the large intestine of
gorilla. _Œs. stephanostomum_ var. _dentigera_, Raill. and Henry, 1909,
in the chimpanzee.
_Habitat._--In large and small intestine of man, Brazil.
_Pathology._--Nodules occur in the gut wall; 187 were found by Thomas
in his, the sole case. The tumours contain each a single worm.
[Illustration: FIG. 321.--_Œsophagostomum stephanostomum_ var.
_thomasi_: portion of the ileum, showing a cyst with protruding worm.
× 8. (After Thomas.)]
[Illustration: FIG. 322.--_Œsophagostomum stephanostomum_ var.
_thomasi_: colon with œsophagostome withdrawn from its cyst cavity.
× 20. (After Thomas.)]
*Œsophagostomum apiostomum*, Willach, 1891.
According to Leiper, _Œs. brumpti_ is identical with, and hence a
synonym of, this species. Parasitic in large intestines of monkeys,
producing dysentery, and in man (Northern Nigeria).
According to Walker this species is common in Philippine monkeys. Ova
are scanty in the fæces. They measure 73 µ to 84 µ by 44 µ to 57 µ and
are in the morula stage. They are easily cultivated. The rhabditiform
larva is 340 µ by 16 µ and has a long filiform tail. It moults twice,
and at the second moult becomes a filariform larva retaining the skin
of this moult, this stage being that of the mature larva. It now
measures 9 mm. long by 30 µ thick. Walker suggests that the mode of
infection is similar to that of ancylostomes.
Group. *Ancylostomeæ*, Railliet and Henry, 1909.
Bursa with anterior ray cleft, median double,[314] postero- and
postero-external arising from a common trunk, posterior bifurcate,
each limb being tridigitate. Vulva in posterior third of body. Uteri
divergent.
[314] _I.e._, with a distinct space between the limbs.
Contains the following genera: (1) Strongylus,[315] (2) Ancylostoma,
(3) Uncinaria, (4) Characostomum, etc.
[315] Strongylus (Syn.: Sclerostomum) differs slightly in its posterior
ray from the other genera of the group. Each bifurcation is trifurcate
rather than tridigitate.
Genus. *Ancylostoma*, Dubini, 1843, emend. Looss, 1905.
Ventral margin of mouth capsule armed with teeth, the “roots” of which
are continued backwards and appear on the _external_ surface of capsule
as rib-like thickenings. Terminal third of dorsal ray cleft. Genital
tubes very long, with short, closely packed diagonal convolutions.
*Ancylostoma duodenale*, Dubini, 1843.
Male 9 mm. long by 0·45 mm. thick, female 12 mm. long by 0·6 mm. thick.
Pale flesh colour, or an intense red in posterior third. Anteriorly may
be more or less black due to (blood) pigment in the cells of the chyle
intestine (= stomach + small intestine). The worm is about the same
thickness all through and is plump and rigid. Cuticle striated. The
body has a marked torsion, so that if the ventral side of the head is
upwards the anus appears to open laterally and _vice versâ_. The dorsal
curve of the head end is only slight and the œsophagus is roughly
cylindrical.
_Buccal Capsule._--The buccal capsule is bent dorsally, 0·21 mm. long,
0·19 mm. broad. If a worm is rolled under the cover-glass so that the
dorsal side is upwards, we observe the following features (fig. 325):
In the dorsal edge of the chitinous capsule there is a gap as if a
*U*-piece had been punched out. This is the “dorsal gap or incision.”
The so-called “dorsal teeth” are simply the rounded edges of the tips
of this gap. They project _beyond_ the skin which covers the capsule
externally. Below this gap is seen a curved line which, if followed
along the sides of the capsule on each side, merges into the base of
the most posterior ventral tooth. This line is the optical expression
of a very shallow groove on the _inside_ of the capsule. The skin
on the outside of the capsule, which is reflected over the edge of
and into the capsule, dips into this groove, which gives it a firm
attachment. Below the middle (dorsally) of this curved line there is
a thickening in the capsule wall, which is perforated by the opening
of the dorsal œsophageal gland. This is the “dorsal ridge”; in optical
section it has a conical appearance with a lumen (of the duct).
[Illustration: FIG. 323.--_Ancylostoma duodenale_, male. _B_, bursa;
_Bm_, bursal muscles; _Cdr_, cement gland surrounding the ejaculatory
duct; _Glc_, cervical glands; _N_, nucleus of cephalic gland; _Nr_,
nerve ring; _T_, testes; _Sp_, spicule; _Vs_, vesicula seminalis. × 15.
(After Looss.)]
[Illustration: FIG. 324.--_Ancylostoma duodenale_, female. _A_, anus;
_Gcph_, cephalic gland; _N_, nucleus of cephalic gland; _Glc_, cervical
gland; _Ov_, ovary; _Pex_, excretory pore; _Rs_, receptaculum seminis;
_Ut_, uterus; _V_, vagina. × 15. (After Looss.)]
On the ventral wall one sees the two pairs of strong teeth, their
points being directed somewhat backwards. They are covered by cuticle
above and below, but their points are free, piercing the cuticle. The
“roots” of these teeth followed backwards appear as two thickenings
or ribs on the _outside_ of the capsule wall, so that the outside
wall is not smooth--a characteristic of the genus Ancylostoma. In the
space between these ribs lies the ventral nerve papilla, and lying
against the outside of the outer root the lateral nerve papilla. The
nerve papillæ are thus, as it were, concealed by these roots, and not
conspicuous as they are in Necator. Following the ventral curve of the
capsule on the inside, posteriorly we next find two triangular ventral
lancets.[316] These stand straight up into the capsule on either side
of the longitudinal axis, converging at their summits. So that to sum
up, the cutting apparatus is entirely ventral, consisting of two pairs
of cutting teeth and a pair of lancets.
[316] The ventral lancet (of one side) of Necator is seen in fig. 335.
_Cervical Papillæ._--Two, one on each side behind the head at the level
of the excretory pore. They consist of “pulp,” _i.e._, extensions of
the substance of the lateral bands covered by cuticle and supplied with
a nerve (fig. 326).
[Illustration: FIG. 325.--_Ancylostoma duodenale_: showing ventral
teeth, dorsal cleft, and behind it the dorsal ridge with duct of dorsal
œsophageal gland. × c. 200. (After Looss.)]
_Œsophageal Glands_ (3).--The chitin of the triradiate œsophagus is
continuous with that of the buccal capsule. In its muscular walls are
three glands--one dorsal, two sub-ventral. The dorsal gland opens into
the buccal cavity through the dorsal ridge; the two others into the
lumen of the œsophagus at the nerve ring. They branch freely amidst the
muscles. They are probably digestive in function.
_Cephalic Glands_ (2).--Lie in the lateral lines or bands on either
side. They begin about the middle line of the body, and their ducts
open at the base of the outer ventral tooth on the surface of the skin
on each side. Each is 0·15 mm. thick in the middle, and has a single
nucleus about as big as an ancylostome egg (_N_, fig. 323). They
probably function as poison glands.
_Excretory System and Cervical Glands_ (2).--The excretory pore lies
in the mid line ventrally behind the œsophageal nerve ring (figs. 324
and 326). It opens into the excretory vesicle, a cavity in a large
cell with lateral appendages which fuse with the lateral lines, this
cell thus forming the “bridge” of the excretory system. Adhering to
this (bridge) cell are the spindle-shaped cervical glands (_Glc_,
fig. 324), and branches from the excretory vesicle enter the glands,
which are excretory in function; the vesicle also receives branches
from the lateral excretory canals (fig. 326) running in the lateral
lines or bands. The cervical glands are swollen anteriorly, forming the
so-called ampullæ just in front of the bridge. They extend backwards a
little beyond the anterior loop of the testis.
[Illustration: FIG. 326.--_Ancylostoma duodenale_: diagrammatic
representation of excretory system. _ex.p._, excretory pore; _e.c.g._,
excretory cervical gland; _Ex. ves._, excretory vesicle in _B.c._,
bridge cell, which is connected with _c.g._, cervical gland, and
_l.l._, lateral lines; _ceph.g._, cephalic gland; _l.ex.c._, lateral
excretory canal passing into the bridge cell; l.l., lateral line
containing excretory canal and cephalic gland; _c.p._, cervical
papilla; _n._, nuclei of bridge cell. (After a drawing of Looss.)]
_Lateral Lines._--(1) Are broad elevations of the subcuticle, in which,
here and there, a nucleus occurs. (2) Near the bursa in the male they
increase in volume, and finally divide into branches which form the
“pulp” of the different rays. (3) In addition to the lateral lines or
bands, there is also a dorsal and ventral band. (4) The ventral band
is well developed caudally, forming a large pad dorsal to the cloaca,
“pulvillus post-analis.”
The bursal rays are outgrowths of the lateral lines. Beside this “pulp”
they contain a nerve, and at their bases complex muscles.
_The Bursa_ is closed on all sides with a short median (ventral) lobe,
which may be tucked inwards. It is an outgrowth of the inner layer of
the skin pushing the outer layer before it, so that it consists of
three layers, not four, as it would be if it were a fold. The bursa
is twice as broad as long. It is supported by a variety of rays, the
arrangement of which is best followed from the figure (fig. 327).
The different terminology for these rays as used by various authors
should be noted: Ventral = anterior; externo-lateral = antero-external;
medio-lateral + postero-lateral or antero-median + postero-median
= median (doubled); externo-dorsal = postero-external; dorsal =
posterior. All the rays end in tactile papillæ, seven, on each side;
the postero-external and antero-external on the _outer_ surface of the
bursa, the five others on the _inner_ surface.[317] Of the six terminal
digitations of the _dorsal_ ray, only the external two contain tactile
papillæ.
[317] This also occurs in other _Strongylidæ_, _e.g._, in the genus
Strongylus (Syn.: Sclerostomum).
In the male there are prebursal papillæ and minute caudal papillæ in
the female.
In the female the inner layer of the cuticle projects at the posterior
end as a sharp spike, 20 µ long, which may sometimes be broken off.
[Illustration: FIG. 327.--_Ancylostoma duodenale_: bursa enlarged.
_Ca_, anterior ray cleft; _cle_, antero-external; _cls_, antero-median;
_clp_, postero-median; _Cde_, postero external; _Cd_, posterior
bifurcated, each bifurcation tridigitate. (After Railliet.)]
_Ovaries._--The anterior tube runs from the cephalic to the posterior
end and back again. The posterior tube begins anteriorly, runs to
the posterior end of the body, and then back to the cephalic end,
forming a vulval loop before ending. The ovaries on the whole run in
oblique coils. The uterus is the thicker portion of the tube, 5 mm.
long. A short tube connecting the ovary and uterus is the oviduct.
The two uteri unite to form a single duct, the vagina, opening 1 mm.
_behind_ the middle line. The portion of the uterus next to the oviduct
functions as a seminal receptacle, whereas the part next the vagina
functions as an ovejector.
_Testis._--The blind end begins a little behind the beginning of the
_cement gland_. The transverse coils occupy the middle third of the
body. About the middle of the body it passes into the spindle-shaped
seminal vesicle, which, with the spicular canal and rectum, opens
into the cloaca. An anterior longitudinal coil pushing in between the
cervical glands is characteristic of Ancylostoma. The cement gland
surrounds the ejaculatory duct for practically its whole course, and it
occupies nearly the posterior half of the body and secretes a brown or
black cement. The spermatozoa are curved rods about 2 µ long.
_Spicules_ are 2 mm. long, ending in a fine point. They are moved by
exsertor and retractor muscles. At first they lie free in the body
cavity; next in a groove in the dorsal wall of the cloaca; then in an
isolated canal, and finally in two canals. Anteriorly each has two
longitudinal crests on its inner surface. These meet the corresponding
crests of the other spicule, and so form a canal along which the sperm
passes into the female. The gubernaculum is a thickening of the dorsal
wall of the cloaca. It is not a free piece, but is moved by various
muscles.
[Illustration: FIG. 328.--_Ancylostoma duodenale_: bursa of male. The
rays from left to right are: (1) anterior cleft; (2) antero-external;
(3) and (4) median doubled, _i.e._, antero-median and postero-median;
(5) postero-external arising from a common trunk with the posterior. ×
c. 120. (After Looss.)]
_Genital Cone_ is a prominence on the floor of the bursa on the ventral
side of the body, on which the genito-anal orifice opens. The cone
is only slightly marked in _Ancylostoma duodenale_, but is much more
prominent in _Necator americanus_.
_Distribution._--Africa, Egypt, Europe, Japan, China (mainly), but in
association with _Necator americanus_ in Southern States of America,
British India, Assam, Burma, Hongkong, Liberia, Jamaica, Martinique,
Costa Rica, Colombia, Antigua, Guadeloupe.
_Habitat._--The worms live in the jejunum, less frequently in the
duodenum, of man only.
_Food._--The worms feed on the mucous membrane of the gut, attaching
themselves to the base of the villi, sucking these in; and when these
are destroyed they attack further the submucosa. As a rule the worms
have no blood in the gut, but in their attack on the submucosa a
blood-vessel may be eroded, and so the gut of the worm filled with
blood.
_Development._--The eggs are oval with broadly rounded poles, 56 µ
to 61 µ by 34 µ to 38 µ. In _fresh_ fæces they contain four granular
nucleated segmentation masses of the ovum (fig. 329) separated by a
clear space from the shell.
_Egg of Ancylostome_ appears to have a single contour. Under high
powers this appears double, but they are the outer and inner surface
of the true (chitinous) egg-shell. Internal to this is the extremely
delicate yolk-envelope, a kind of skin secreted by the egg cell around
itself for protection. The function of this is probably to absorb water
to swell and burst the outer chitinous shell. The embryos when hatched
are termed larvæ.
_Embryos_ which are ready to hatch have their bodies almost free from
granules; others, though apparently mature, that have granules will not
hatch.[318]
[318] TABLE OF DIFFERENCES BETWEEN LARVÆ OF _A. duodenale_ AND _S.
stercoralis_.
--------------+----------------------+------------------------------
| _A. duodenale_ | _S. stercoralis_
--------------+----------------------+------------------------------
(1) Vestibulum| | }
oris |1·8 µ broad |3 µ }Rhabditiform.
(2) Genital | | }
rudiment|3 µ to 5 µ long |25 µ to 33 µ }
| |
(3) Thickness |Thicker | -- }
(4) Œsophagus |One-fourth body length|Half body length}
(5) Tail |Pointed |Two fine points }Filariform.
(6) Motion |Less active than | -- }
(7) Gut |Soon fills with dark | }
| granules | -- }
--------------+----------------------+------------------------------
[Illustration: FIG. 329.--_Ancylostoma duodenale_: eggs in different
stages of development. _a_ to _c_, in fresh fæces; _d_, containing a
larva, only in old fæces. × 336. (After Looss.)]
_Larva._--_Stage I_: Average length, 25 mm. Maximum thickness in
œsophageal region, 17 µ. Head end fairly blunt, from behind the
anus (the tail) tapering in an uniform manner. Buccal cavity is
characteristic, 10 µ to 12 µ by 1 µ to 8 µ, longer and narrower than
the corresponding larvæ of _Strongyloides stercoralis_. Œsophagus
“rhabditic” in character, _i.e._, it has three sections, but they are
not so clearly marked off as in larvæ of the genus Rhabditis. The
posterior bulb has a *Y*-shaped valve, the function of which, according
to Looss, is to prevent regurgitation of food. The granules of the gut
serve as a reserve of food, and are used up if the larvæ are starved.
The _genital rudiment_ consists of two cells half-way between the end
of the œsophagus and the anus in the mid-ventral line. The larva lives
on fæcal matter and grows to about 0·4 mm., then moult[319] I takes
place in two days or more, the skin being ruptured by the activity of
the larva.
[319] Moults take place by the formation of a new skin below the old
one, the two being in close apposition at first.
[Illustration: FIG. 330.--_Ancylostoma duodenale_ larva on fourth day
of culture on right; _Strongyloides stercoralis_ larva on left. (After
Leichtenstern.)]
_Stage II_: The larva is now in this stage, which does not differ much
from the previous one. It grows to 0·5 mm. The mouth opening closes.
The œsophagus elongates, becoming cylindrical or “filariform”; a new
skin is formed underneath the old one, and in about a week moult II
takes place.
_Stage III_: The _mature larva_ remains enclosed in the old skin. Its
movements are now much more active and of a boring character. Length is
now 0·6 mm. This mature stage has been erroneously called the encysted
larva, because there is no cyst _secreted_ from its surface by the
larva, but it is simply the old skin, which is not cast off, but is
retained for purposes of protection, as the larva is free living, but
casts it as soon as it assumes parasitic life again. From the egg to
this mature stage is thus six to ten days.
[Illustration: FIG. 331.--_Ancylostoma duodenale_: left, four days
after transmission into dog, 190/1; in the centre, at the commencement
of the second stage of development (five to six days), 105/1; on the
right, fourteen to fifteen days after transmission. 42/1. (After
Looss.)]
_Bionomics of Development._--_Air_: Eggs can develop when shut off from
the air for a “comparatively long” time.
_Temperature_: Hatching takes from eight hours upwards. Eggs develop
best at 25° to 30° C., but will not develop below 8° to 10° C. The
larvæ, however, will stand freezing.
_Moisture_: Eggs and larvæ do not live long under water, because they
suffocate or starve, but _mature_ larvæ will live for months (six to
twelve) in water; they require no food--in fact, can take none in--but
live on their reserve granules, and in course of time become as clear
as glass.
_Thigmotropism_: The mature larvæ, after casting their skin, will
penetrate pith, climb up stems, stalks, etc., and creep into any pore.
It is important to recognize that this third stage of the _mature
larva_ is the only infective one.
_Mode of Entry into the Body._--Infection is effected through the mouth
(Leichtenstern and others), and also through the skin, as was first
discovered by Looss and afterwards confirmed from the most diverse
quarters, partly in the case of _Ancylostoma duodenale_, partly in
that of _A. caninum_ in dog, man, and monkey. The larvæ that gain
access to the intestine partly through contaminated food, or through
unwashed hands, or under some circumstances through water, first throw
off their “sheath”--that is, they complete moult II. Moult III takes
place four to five days after they have reached the gut, and they now
have a mouth capsule supplied with four small teeth arranged crosswise,
enabling them to fasten on to the intestinal epithelium, upon which
they feed. On this food the worms grow in four to six days to 3 to
5 mm. in length, and now moult IV. takes place, thus attaining their
definite shape and distinctive character. About eight days later the
sexual organs commence to function; at this time the first copulation
should be taking place--it will later be frequently repeated--and a few
days later the first ova are laid, first in less and later in larger
numbers, so that they appear in the fæces about four to five weeks
after the infection.[320]
[320] From the number of eggs present in a given quantity of fæces, the
number of female Ancylostomes present in the gut can be reckoned by
a formula of Leichtenstern’s (x = _a_/47, in which _a_ signifies the
number of eggs counted in a single gramme of fæces).
_Infection by the Skin._--Mature larvæ, which are placed on the skin of
man or suitable animals, cast their “sheath” and bore their way through
delicate fissures either horizontal in the superficial scales of the
epidermis, or through vertical fissures into hair follicles where these
exist, and then they invade the cutis. Now according as they migrate
further into the lymphatic vessels or the small vesicles, the final
path to the gut differs to some extent. The blood path leads to the
right heart, and from there into the lungs; here the larvæ leave the
blood stream and enter the air passages, over the mucosa of which they
travel further headwards, through the bronchi into the trachea and
larynx, and from hence through the œsophagus to the stomach; in some
cases also they are swallowed. The lymphatic path leads finally also
into the blood stream, but the lymphatic glands must first be passed,
and in these many larvæ are retained and perish. In the cutaneous
infection seven to ten weeks elapse till the time of appearance of the
first ova in the fæces.
The penetration of the skin by the larvæ also in man causes reddening
and burning at the affected points, and this is followed in a
few days by transitory swelling in the subcutaneous connective
tissue. Skin affections can also be set up by such Ancylostoma
(and Strongyloides) larvæ as do not gain access to the blood or
lymphatic vessels or gut; such larvæ apparently wander further in the
connective tissue, and, as Looss has in his own person observed, gain
access to the cutis at different points, thus causing progressive
swellings (accompanied by intense itching), which cease when the
worm again penetrates into the deep tissues. Skin affections such
as “ground-itch” or “pani-ghao” occurring in the tropics and only
attacking the feet, or other affections (_e.g._, sump bunches) are
now well recognized as being due to the invasion of Ancylostoma larvæ.
Other names for these skin affections are water-sore, sore feet of
coolies, maza-morra, bunches, botches, quaddeln, krätze, ampoules,
gourmes, taons, pitirr. Whether oral or dermal infection is the more
important one further observation must decide.
The duration of life of _Ancylostoma duodenale_, which is a specific
parasite of man and has not been observed in other mammals, amounts to
about five years, as strayed larvæ according to Looss wander for this
extent of time in the body.
_Cultivation of Larvæ._--(1) Mix the fæces (free from drugs such as
salines or thymol) with animal charcoal, adding water if necessary till
a consistence of porridge is obtained. If the stools are very fluid,
allow to sediment first and pour off the fluid. The best charcoal is
that made from bones, and should not have an acid reaction. Charcoal
is necessary in order to prevent fermentation, which kills the larvæ.
Spread in layers 2 to 3 mm. thick in Petri dishes. Incubate at room
temperature. To extract the larvæ from the culture allow the surface
thoroughly to dry, then pour on water; the larvæ wander out and are
poured off and subsequently further purified by sedimentation or
filtering through blotting paper, the larvæ passing through.
(2) A funnel is plugged with cotton wool, then filled with washed sand
to within a centimetre or two of the rim. Stand this in a jar of water
so that the level of the water is slightly below that of the sand. On
the surface of the wet sand now place layers of blotting paper, and
spread the fæces, diluted if necessary, on this in layers of a few
millimetres thick (_vide_ p. 474).
_Detection of Eggs._--_Vide_ p. 473.
_Dermal Infection of Dogs._--Infection with larvæ of _A. caninum_. In
two hours most of the larvæ are free in the cutis and in four hours in
the subcutaneous tissue. By scraping a few days later the mucosa of the
trachea large numbers of larvæ are found there.
*Ancylostoma ceylanicum*, Looss, 1911.
[Illustration: FIG. 332.--_Ancylostoma ceylanicum_: head end, two teeth
on each side, the inner almost concealed by the outer. × c. 200. (After
Looss.)]
At the anterior edge of mouth capsule one large tooth; below or behind
this towards the middle line a very small tooth, the tip only of which
is seen. Male 5 mm. average. Lobes of bursa almost as long as broad,
strongly projecting towards the ventral side. Rays short and relatively
thick. Female 7 mm.
_Habitat._--Intestine civet cat (_Viverricula malacensis_), Ceylon, and
man in Bengal according to Clayton-Lane.
Other species are: _A. caninum_ (Ercolani), in cat and dog, Europe
and Africa; _A. malayanum_ (Alessandrini), 1905, in the Malay bear
(_Helaretos malayanus_); _A. pluridentatum_ (Alessandrini), 1905, in
_Felis mitis_, Brazil.
*Ancylostoma braziliense*, Gomez de Faria, 1910.
In cats (and dog), Brazil. Female 8·5 mm., male 7·5 mm. long. Eggs 65 µ
by 32 µ. Leiper considers it to be identical with _A. ceylanicum_.
[Illustration: FIG. 333.--_Ancylostoma braziliense_: bursa of male.
(After Gomez de Faria.)]
Group. *Bunostomeæ*, Railliet and Henry, 1909.
Bursa with median double, postero- and postero-external arising from a
common trunk, posterior bifurcated, each limb bidigitate (fig. 336).
Vulva in middle of body or a little in front. Uteri divergent.
Contains the following genera: (1) Bunostomum (= Monodontus); (2)
Necator; (3) Bathmostomum; (4) Gaigeria.
Genus. *Necator*, Stiles, 1903.
Mouth capsule small, narrowed anteriorly (ventrally) by chitinous
plates, as in Uncinaria. On each side of the base of the dorsal cone
a lateral chitinous plate or lancet with smooth edge (not serrated),
ventral lancets as in Ancylostoma. No ridges on outside of ventral
wall. Aperture of dorsal œsophageal gland on tip of a cone projecting
freely into the buccal capsule. Bursa closed. Posterior ray cleft to
its root.
*Necator americanus*, Stiles, 1902.
Syn.: _N. africanus_, Harrison, 1910.
[Illustration: FIG. 334.--_Necator americanus._ Showing cutting plates
and the projecting dorsal ridge, and deep in the cavity the edges of
the ventral lancets. × c. 475. (After Looss.)]
Male 8 mm. long, female 10 mm. The head is strongly bent dorsalwards
so that almost by this character alone it can be distinguished from
_Ancylostoma duodenale_. The buccal capsule is markedly small--in
the male, 0·093 by 0·084 mm., in the female 0·11 by 0·097 mm. There
are no teeth anteriorly on the ventral side of the capsule, but
instead there are two cutting chitinous plates, the anterior portions
of which are prominent and angular, and meet in the middle line
in front. Posteriorly on each side the plate projects less, while
between the anterior and posterior parts there is a deep angle. The
inner (posterior) ventral lancets which also occur in _A. duodenale_
are large, and project far into the lumen, the tips of these, of the
lateral lancets, and of the dorsal cone almost meeting in the centre of
the lumen. As already stated in the definition of the genus Necator,
there are also lateral lancets which start from the base of the dorsal
cone. This dorsal ridge, or rather in this case cone, is a striking
object in the mouth, and projects right out into the cavity, and on
its summit opens the dorsal œsophageal gland.
[Illustration: FIG. 335.--_Necator americanus_: lateral view, showing
the dorsal ridge perforated by the duct of the dorsal œsophageal gland,
the lateral lancet and ventral lancet and the nerve papillæ. × c. 475.
(After Looss.)]
The bursa is about as long as broad, but has the lateral lobes
strikingly lengthened, giving a trilobed appearance (fig. 336), but
as in _Ancylostoma duodenale_ it is closed on the ventral side. The
distribution of the rays is best understood from the figure. The
genital aperture lies on a marked conical protuberance; the cement
gland is bilobed in transverse section. In the female the opening of
the vulva is in front of the middle line, in _A. duodenale_ it is
behind.
The spicules, 0·92 mm. long are hooked at the extremity.
[Illustration: FIG. 336.--_Necator americanus_: bursa of male. The rays
from right (top) to left are: (1) posterior, (2) postero-external,
(3) and (4) median doubled, _i.e._, postero-median and antero-median,
(5) antero-external, (6) anterior (cleft), and above it on left a
pre-bursal ray. × c. 120. (After Looss.)]
Eggs more pointed at the poles than those of _A. duodenale_, 64 µ to
72 µ by 36 µ, so that it may not be possible to distinguish single eggs
owing to individual variations, yet on comparing a number they can be
distinguished.
_Geographical Distribution._--Brazil, Porto Rico, Cuba, Central Africa,
East Africa, Victoria Nyanza, Gold Coast, Uganda, North-Western
Rhodesia, Ceylon, Mysore. For other localities where _A. duodenale_ is
also found see p. 450.
_Habitat._--In small intestine of man and gorilla (_Troglodytes
gorilla_).
*Necator exilidens*, Cummins, 1912.
Syn.: _N. africanus_, Looss, 1911.
Male 7 mm., female 9 mm. long. The edges of the cutting plates are
rounded, not angular, and do not meet in the middle line. Inner
(posterior) ventral lancets very small. Lateral lobes of bursa broader
than long. Rays thick and plump.
_Habitat._--In the chimpanzee (_Anthropopithecus troglodytes_).
ANCYLOSTOMIASIS.
_Morbid Anatomy._--Organs pale and bloodless. Abdominal organs sodden,
and there is fluid in the serous cavities. Lungs: œdema. Kidneys: fatty
changes, especially large pale kidney. Liver and heart also show fatty
changes--there is much hæmosiderin in the liver cells. Blood: early
stages, a leucocytosis 20,000 upwards, and eosinophilia 50 per cent.
Later, anæmia (hydræmia). The number of worms found varies from ten to
1,000. They are rare in the duodenum, but occur as far as 6 ft. from
the pylorus.
Group. *Syngameæ*, Railliet and Henry, 1909.
Bursa with anterior and median ray cleft; antero-external, close to
median; postero-external, arising separately from posterior; posterior
bifurcate to base, each branch bifurcate or trifurcate. Vulva in the
anterior fourth of body. Uteri divergent.
Genus. *Syngamus*, von Siebold, 1836.
Head thickened, not tapering; broad mouth with gaping buccal capsule.
Male and female often in permanent copulâ.
Parasitic in respiratory passages of birds and mammals.
_Habitat._--_S. trachealis_ in poultry; _S. bronchialis_ in goose; _S.
laryngeus_ in cattle; _S. vasicola_ in goats, etc.
*Syngamus kingi*, Leiper, 1913.
Buccal capsules of male and female on same level. In _S. trachealis_
and _S. laryngeus_, that of male in front of that of female. In
_S. dispar_, that of male behind that of female. Œsophagus of male
one-sixth, that of female one-ninth of total length. Mouth capsule
in male and female terminal; it is dorsal in _S. trachealis_ and in
mammalian species. Tail of female bluntly pointed. Ovary reaches to
anus. Excretory pore opposite the middle of the bulb of œsophagus. In
_S. trachealis_ it is opposite the œsophageal valves.
_Habitat._--Found in sputum of patient by King in St. Lucia. Normal
host probably a carnivore.
[Illustration: FIG. 337.--_Syngamus kingi_: anterior end of male.
(After Leiper.)]
[Illustration: FIG. 338.--_Syngamus kingi_: anterior end of female.
(After Leiper.)]
Family. *Physalopteridæ.*
Genus. *Physaloptera*, Rudolphi, 1819.
Mouth surrounded by _two_ large lateral lips bounded posteriorly by
a cuticular band projecting anteriorly, forming a collar. Each lip
bears anteriorly and inwardly a cuticular appendage, the external
tooth. Immediately below and internal to the external teeth the
internal teeth, one on each lip. Each lip bears two large submedian
papillæ. Tail of male with four pairs of pedunculated papillæ in a
row on each side external to the six pairs of unpedunculated papillæ.
Spicules unequal. Vulva in the anterior region of the body. Eggs with a
characteristic thick smooth shell.
Parasitic in the intestine, more especially the stomach, of mammals
(twenty species), birds (twelve species), reptiles (fourteen species).
*Physaloptera caucasica*, v. Linstow, 1902.
The male measures 14·2 mm. in length and 0·71 mm. in breadth; the bursa
is broad, rounded off in front and narrower at the back; the right
spicule measures 0·62 mm. in length, the left spicule 1·76 mm.; there
are two papillæ in front of the orifice of the cloaca, four behind
it and six unpedunculated on the tail. The female measures 27 mm. in
length, 1·14 mm. in breadth; the caudal extremity is rounded off; the
vulva is on the border of the first and second sixth of the length of
the body; the eggs have thick shells, and measure 57 µ by 39 µ. It has
hitherto only been observed once, by Ménétriés in the intestine of man
(Caucasus).
[Illustration: FIG. 339.--Bursa of _Syngamus trachealis_. _a._,
anterior ray cleft; _a.e._, antero-external; _m.a._, antero-median;
_m.l._, postero-median; _p.e._, postero-external; _p._, one branch of
posterior (trifurcate). (Stephens.)]
*Physaloptera mordens*, Leiper, 1907.
Large worms resembling an immature _Ascaris lumbricoides_.
The inner lancet-shaped teeth have a sharp cutting edge towards the
lumen. Below each is a cuticular boss projecting into the mouth
(fig. 340).
Male 30 to 50 mm., bursa with four pairs of pedunculated papillæ, the
second and third lying external to the first and fourth on each side.
Spicules unequal, one slender (4·6 mm.), the other stouter (6 mm.).
Female 40 to 55 mm. Tail sharp. Vulva opens between the anterior fourth
and fifth of the body. Eggs 43·6 µ by 35·3 µ with a thick smooth shell.
_Habitat._--Œsophagus, stomach, small intestine of man (several cases).
Nyasaland and Portuguese East Africa.
Family. *Ascaridæ*, Cobbold, 1864.
Sub-family. *Ascarinæ.*
Without œsophageal or intestinal diverticula; spicules without flanges.
Genus. *Ascaris*, L., 1758.
Intermediate lips and auricles absent. Lips edged with fine teeth.
Lips triangular in cross section. Not grooved on internal surface.
[Illustration: FIG. 340.--_Physaloptera mordens_, Leiper, 1907. (*1*)
adult male: _o.e._, œsophagus; _ch.i._, chyle intestine; _t.c._,
testicular coils; _ves. sem._, vesicula seminalis; _sp._ 1, long
spicule; _sp._ 2, short spicule; _B._, bursa. (*2*) Mouth parts:
_c._, cuticular collar embracing the two lips posteriorly; _c.b._,
cuticular bosses guarding the mouth laterally; _e.d._, external tooth;
_i.d._, internal tooth; _sm.p._, submedian papillæ. (*3*) egg of _P.
caucasica_. (*4*) egg of _P. mordens_. (*5*) bursa enlarged: _ped.p._,
pedunculated papillæ; _ses.p._, sessile papillæ. (After Leiper.)]
*Ascaris lumbricoides*, L., 1758.
The colouring, in the fresh condition, is reddish-yellow or
greyish-yellow; the body is of an elongated spindle shape. The oral
papillæ are finely toothed. The dorsal papilla carries two sensory
papillæ, the two ventral papillæ each one sensory papilla. The
male measures from 15 to 17 to 25 cm. in length, and about 3 mm.
in diameter; the posterior extremity is conical and bent hook-like
ventrally; the spicules measure 2 mm. in length, are curved, and
somewhat broadened at their free end; on each side around the orifice
of the cloaca there are seventy to seventy-five papillæ, of which
seven pairs are post-anal. The testicular tube is much folded, showing
through the body integument, and is about eight times the length of
the body. The female measures 20 to 25 to 40 cm. in length and about
5 mm. in diameter; the posterior extremity is conical and straight.
The vulva is at the junction of the anterior and middle thirds of the
body, which, at this point, has a slight ring-like constriction; the
convoluted ovaries measure ten times the length of the body.
[Illustration: FIG. 341.--_Ascaris lumbricoides._ _a_, posterior
extremity of the male with the spicules protruding from the orifice of
the cloaca (_Sp._); _b_, anterior extremity from the dorsal surface,
the two lobes of the pulp of the lip separated by the “saddle”; _c_,
anterior extremity from the ventral surface; _P._, excretory pore.
(From Claus.)]
[Illustration: FIG. 342.--Ovum of _Ascaris lumbricoides_, with shell
and albuminous envelope. 400/1.]
The ova are elliptical with a thick (4 µ) transparent shell (fig. 342)
and an external albuminous coating which forms protuberances; the ova
measure 50 µ to 70 µ in length, 40 µ to 50 µ in breadth; they are
deposited _before_ segmentation; the albuminous coating is stained
yellow by the colouring matter of the fæces, but is sometimes absent.
The egg cell is unsegmented, it almost completely fills the shell, and
its nucleus is concealed by the large amount of coarse yolk granules.
Abnormal or unfertilized eggs also occur in fæces. They are
distinguished by their elongated form (80 µ by 45 µ), irregularly
cylindrical, its contents consisting of refractive granules.
_Ascaris lumbricoides_ is one of the most frequent parasites of man; it
is distributed all over the inhabited parts of the world, and though
it is particularly frequent in the warmer regions, yet it also occurs
in Finland, Greenland, etc. In temperate climates _A. lumbricoides_
occurs most frequently in young children; it is, moreover, more common
amongst country dwellers than amongst the inhabitants of towns, but is
not lacking in infants, adults and aged persons. As a rule only a few
specimens are present in the intestine, but many cases are known in
temperate zones in which several hundreds of worms have been found in
the same patient. This species is particularly numerous in the negroes
of Africa and America. It occurs also in the monkey, dog and pig (? _A.
suilla_).
The parasite was known in ancient times; the Greeks called it ἐλμινς
στρογγύλη, Plinius termed it _Tinea rotunda_, later on it was named
_Lumbricus teres_. The ἄσκαρις of the Greeks is our Oxyuris.
The small intestine is the normal habitat of _Ascaris lumbricoides_;
the worms, however, often leave this part of the intestine and wander
into the stomach, whence they are frequently evacuated by vomiting,
or they may creep through the œsophagus into the pharynx and crawl
out through the nose or mouth; very rarely they may find their way
into the Eustachian tube or into the naso-lachrymal duct, or into
the excretory ducts of the liver and pancreas; exceptionally they
may gain the trachea, and they have also been found in the abdominal
cavity. They may bore through adhesions between the intestinal wall
and the omentum (worm abscess); they occasionally penetrate the
urinary apparatus and are passed with the urine; in febrile diseases
_A. lumbricoides_ usually leaves the intestine spontaneously. It is
obvious that these wanderings may be accompanied by the most serious
symptoms, but in sensitive persons the invasion of even only a few
intestinal Ascarides gives rise to a series of almost inexplicable
symptoms (hysterical, epileptiform attacks, cerebral congestion,
aphonia, etc.), which cease with the expulsion of the worms, so
that many authors are driven to the conclusion that these Ascarides
secrete a toxin. Fortunately, the presence of _A. lumbricoides_ in
the intestine is easily demonstrated by the microscopical examination
of the fæces.
_Development._--Several authors (Gros, Schubart, Richter, Leuckart and
Davaine) have demonstrated that the ova of Ascaris develop in water or
moist earth after a long period of incubation. Freezing and desiccation
(if not too long) do not injure their powers of development; the
duration of the development depends on the degree of the surrounding
temperature. At a medium temperature, after a varying period of
incubation, it takes from thirty to forty days for the embryo to become
formed. The spirally rolled up embryo, with its so-called “tooth,”
formed by three papillæ close together, never leaves the egg-shell
in the open, even if the eggs are kept for years under favourable
conditions. Davaine proved that the larvæ hatch out in the intestine of
the rat, but are again expelled with the fæces; he therefore concluded
that the hatching likewise takes place within the intestine of man,
but is followed by the invasion of the larvæ. In the meantime Leuckart
had sought to infect himself by swallowing embryo-containing eggs,
but without results; he therefore conjectured that there must be an
intermediary host, and v. Linstow thought he had found it in myriapods
(_Julus guttulatus_). Subsequently, Davaine’s opinion proved correct.
First of all Grassi succeeded in infecting himself by swallowing 100
embryo-containing eggs of _Ascaris lumbricoides_; five weeks after
ingestion the worms had attained maturity and their ova appeared in
the fæces. Calandruccio also sought to infect himself, but failed,
yet he succeeded in infecting a little boy aged 7. Lutz also reports
a successful experiment which must have been positive, as young worms
5·5 to 18 mm. long were expelled. Lutz proved that the eggs lost their
albuminous shell by long lying in water and then died when introduced
into the stomach; this would explain the failure of Leuckart’s
experiment; in moist earth the albuminous shell is retained. Finally,
Epstein conducted unimpeachable experiments on three children which
place direct infection with embryo-containing eggs beyond doubt; he,
moreover, proved that the development of the eggs takes place more
rapidly in the fæces when there is free admission of air, sun, and a
sufficiency of moisture.
Accordingly, infection occurs partly through water, but principally
direct from the soil.
*Ascaris*, sp.
Wellmann states that yet another species of Ascaris in man occurs in
the highlands of Angola: up to the present nothing certain is known
about it (Welland, “Critical Notes on Tropical Diseases of the Angola
Highlands,” _New York Med. Journ. and Philadelphia Med. Journ._, August
12 to September 2, 1905.)
*Ascaris texana*, Smith et Goeth, 1914.
Female alone known; 58 to 60 mm. and upwards in length; characterized
by the serration of the anterior border of the lip and by the
appearance of interlabia. Evacuated by a white settler in Texas.
Position of this worm doubtful.
*Ascaris maritima*, Leuckart, 1876.
Only one immature specimen, a female (43 mm. in length and 1 mm. in
breadth), has hitherto been described, and it was vomited by a child in
North Greenland in 1865. (R. Leuckart, “Die menschlichen Parasiten,”
1876, edition 2, i, p. 877.)
Genus. *Toxascaris* (τόξον, an arrow), Leiper, 1907.
Body anteriorly bent dorsally, cuticle finely striated. Œsophagus
without a distinct bulb. Tail of male tapers to a point. Testis in
anterior portion of posterior half of body. Vulva about middle of body.
Eggs oval and smooth.
*Toxascaris limbata*, Railliet and Henry, 1911.
Syn.: _Lumbricus canis_, Werner, 1782; _Ascaris teres_, Goeze, 1782;
_Ascaris cati_ et _caniculæ_, Schrank, 1788; _Ascaris canis_ et
_felis_, Gmelin, 1789; _Ascaris tricuspidata_ et _felis_, Bruguiere,
1791; _Ascaris werneri_, Rud., 1793; _Fusaria mystax_, Zeder, 1800;
_Ascaris marginata_ et _mystax_, Rud., 1802; _Ascaris alata_,
Bellingham, 1839.
Striations 6 µ to 12 µ apart. Cephalic wings long, narrow,
semi-lanceolate. Male, 4 to 6 cm. Spicules, 1,002 µ and 1,005 µ.
Female, 0·5 to 10 cm. Eggs, 75 µ to 85 µ, shell thick and smooth. Host:
dog, occasionally man.
[Illustration: FIG. 343.--Ovum of _Toxascaris limbata_, with thin
albuminous envelope. Magnified.]
[Illustration: FIG. 344.--Transverse section through the head part of
_Belascaris cati_ from the cat, with the lateral wings. In addition,
one may note the four fields of muscles, the longitudinal lines with
the œsophagus in the centre. Magnified. (After Leuckart.)]
Genus. *Belascaris* (Βέλος, an arrow), Leiper, 1907.
Body anteriorly bent ventrally, cuticle coarsely striated. Œsophagus
with a distinct bulb. Tail of male conical. A papillæ-bearing
protuberance behind the anus. Testis in anterior half of body. Vulva in
anterior part of body. Eggs corrugated.
*Belascaris cati*, Schrank, 1788.
Syn.: _Belascaris mystax_, Leiper, 1907; _Ascaris mystax_.
Striations 12 µ to 16 µ apart. Cephalic wings lanceolate. Male 3 to
6 cm. Spicules 17 to 1·9 mm. Female 4 to 10 cm. Eggs, 65 µ to 75 µ in
diameter, surface finely honeycombed. Host: domestic cat, and man,
eight or nine cases.
*Belascaris marginata*, Rudolphi, 1802.
Striations 16 µ to 22 µ apart. Cephalic wings long, narrow,
semi-lanceolate. Male, 5 to 10 cm. Spicules, 750 µ and 950 µ. Female, 9
to 18 cm. Eggs, 75 µ to 80 µ. Shell finely honeycombed. Host: dog.
Genus. *Lagocheilascaris*, Leiper, 1909.
Thick lips separated by a furrow from the body; between the lips
small intermediate lips without “pulp.” The cutting angle of each lip
bifurcated. Along each lateral line a cuticular wing extending the
whole length of the body. Eggs, thick shell with a mosaic pattern.
*Lagocheilascaris minor*, Leiper, 1909.
Male, 9 mm., tail sharply curved. Spicules colourless, 3·5 and 4 mm.
long. More than twenty-four pairs of pre-anal papillæ, at least five
pairs of post-anal. Female, 15 mm. Straight posteriorly. Vulva 6 mm.
from head with two lips. Eggs, 65 µ in diameter. Host: man, cutaneous
abscesses. Trinidad.
Family. *Oxyuridæ.*
Genus. *Oxyuris*, Rudolphi, 1803.
Mouth unarmed. The three labial papillæ are only slightly
protuberant, the œsophagus is long and presents two well-marked
bulbs. The vulva is in the anterior part of the body.
*Oxyuris vermicularis*, Linnæus, 1767.
Syn.: _Ascaris vermicularis_, L.; _Fusaria_, Zeder, 1803.
Colour white, the striated cuticle forms projections at the anterior
end which extend some distance back along the middle of the ventral
and dorsal surfaces; the longitudinal lateral flanges of the skin
corresponding to the lateral lines are well seen in transverse
sections; there are three small retractile labial papillæ around the
mouth. The male measures 3 to 5 mm. in length, and shortens on death;
the posterior extremity of the body is curved ventrally and presents
six papillæ. Spicule 70 µ long, hook-like. The female is 10 mm. in
length and 0·6 mm. in diameter; the anus is about 2 mm. in front of
the tip of the tail; the vulva is in the anterior third of the body;
the eggs are oval, asymmetrical, with double-contoured shells, and
measure 50 µ to 55 µ by 16 µ to 25 µ; they are deposited with clear,
non-granular tadpole-like embryos already developed.
_Habitat._--Adults in large intestine of man. Young forms in small
intestine and often in the appendix.
The worm lives in the lower part of the small intestine, cæcum and
vermiform appendix, and before becoming adult undergoes two or three
moults (Heller). According to Wagener the worms at times live in the
gut wall, giving rise to calcareous nodules. When the uterus of the
fertilized females begins to fill with eggs they leave the cæcum and
travel through the colon to the rectum. The uterus is now packed with
eggs which contain a tadpole-shaped embryo. Egg-laying now takes
place, partly in the rectum, partly outside, the mode of exit being
not only passive through defecation but also an active one on the
part of the worms when the patient is in bed. In this case the worms
crawl out of the anus, producing a most intolerable itching as they
scatter their eggs between the nates and the perinæum. From here
in the case of girls they may get occasionally into the vulva and
vagina, and even into the oviducts and so into the body cavity. The
worms also may wander through the alimentary canal in the opposite
direction, getting out occasionally through the mouth. Recently a
_rôle_ has been assigned to them, as to other gut parasites, in
appendicitis and typhlitis.
It is stated that the males die after fertilizing the females,
thus explaining why they are so rarely met with in fæces [but
it is probable that they often escape notice from their small
size.--J. W. W. S.].
[Illustration: FIG. 345.--_A_, male, and _B_, female, of _Oxyuris
vermicularis_. 5/1.]
[Illustration: FIG. 346.--On the left, female; on the right, male. _A_,
anus; _M_, mouth; _V_, vulva. Greatly enlarged. (After Claus.)]
[Illustration: FIG. 347.--_Oxyuris vermicularis_: egg freshly
deposited, with tadpole-like embryo. × 640.]
[Illustration: FIG. 348.--_Oxyuris vermicularis_: egg twelve hours
after deposition, with nematode-like embryo. × 640.]
_Development._--The eggs, which often adhere together, contain a
tadpole-like embryo, the thin tail of which is bent upwards ventrally;
the embryo in a short time, given a sufficiently high temperature,
passes into a second folded nematode-like embryonal stage, lying in the
egg-shell, either in the fæces, with which also numerous females pass
out, or in the moisture of the groove between the buttocks, and they
there await the opportunity of being reintroduced into man _per os_. It
is very improbable that infection takes place directly in the large
intestine, as is occasionally stated, because although the harbourers
of Oxyuris are frequently liable to auto-infection, this takes place
exclusively through the mouth, and is conveyed by the fingers, on which
the ova of Oxyuris, and occasionally the female worms, have clung.
The opportunity for this is afforded every evening, as naturally the
troublesome itching caused by the wandering of the worms is met by
scratching and rubbing with the fingers. It is therefore possible
that the eggs may even thus be introduced into the nose, where the
young Oxyuris are perhaps hatched out, if they get high enough up on
the moist pituitary mucous membrane. As a matter of fact, the larvæ
of Oxyuris have been found in the nose. Moreover, one can understand
that the eggs of Oxyuris are transferred from person to person by
the hand, directly or indirectly. This again explains the wholesale
infections which occur in collective dwellings, after a person
harbouring Oxyuris has been admitted into boarding-houses, etc. The
primary infection may be also caused in other ways--by foods, fruits,
vegetables and other articles that are eaten raw, and are polluted
with the ova. Perhaps also flies or their excrement play a part in
the distribution of the parasite, similar to that demonstrated by
Grassi as taking place in the spread of the ova of Trichocephalus and
Tænia.
The assumption of a _direct development_ without an intermediary host
was first substantiated by Leuckart by experiments on himself and
three of his students; about fourteen days after swallowing the eggs
the Oxyuris has attained 6 to 7 mm. in length; Grassi, and later on
Calandruccio, infected themselves by swallowing adult female Oxyuris,
with the same results. Heller found worms in the gut (appendix
vermiformis) of a male child five weeks old.
Other species are: _O. compar_ in the cat; _O. curvula_ and _O.
mastigodes_ in horse, ass, mule; _O. ambigua_ in the rabbit; _O.
poculum_ in the horse; _O. tenuicauda_ in the horse. Many species
occur in insects, especially in _Blattidæ_ and _Hydrophilidæ_
(aquatic beetles).
Family. *Mermithidæ.*
Genus. *Mermis*, Dujardin, 1845.
With characters of the family.
*Mermis hominis oris*, Leidy, 1850.
Fourteen centimetres in length, 0·16 mm. in breadth; mouth terminal;
posterior extremity obtuse and provided with a recurved hook 50 µ long.
The parasite was “obtained from the mouth of a child.” Stiles
considers it to be probably a Mermis, possibly swallowed in an apple.
*Agamomermis*, Stiles, 1903.
Group name for immature _Mermithidæ_.
*Agamomermis restiformis*, Leidy, 1880.
This worm measures 65 cm. in length, pointed anteriorly, the posterior
extremity broadened and rounded off (1·5 mm. in breadth); the mouth is
terminal, without lips. Behind the mouth six papillæ; the œsophagus
measures 1·125 mm. in length; the intestine appears to terminate
blindly.
This parasite was obtained in West Virginia from the urethra of a
young man, aged 20, who for a few days previous to expelling the worm
passed turbid and bloody urine.
TECHNIQUE.
PRESERVATION AND EXAMINATION OF FLUKES.
_Fixation._--(Method A.) (1) Place the flukes in a test tube or small
bottle a quarter full of normal saline. Shake the contents _as hard as
possible_ (the object of this is to _extend_ the flukes) for half a
minute.
(2) Add _immediately_ an equal bulk of saturated aqueous solution of
corrosive sublimate and shake again as vigorously as possible for a few
minutes.
(3) Transfer when convenient to 70 per cent. alcohol. (Before staining
and mounting remove the sublimate with tincture of iodine.)
(Method B.) In case of large flukes, _e.g._, _Fasciola hepatica_,
_Fasciolopsis buski_, compress the flukes between two glass slides with
rubber bands or thread. Fix in sublimate or in absolute alcohol, or in
10 per cent. formalin.
(Method C.) Place the flukes in 10 per cent. formalin solution.
_Staining_ is successfully effected by using quite _dilute_ solutions
of carmine or hæmatein overnight. This is far preferable to using
strong solutions, as it may be almost impossible to remove a too
intense stain. Almost any dilute carmine solution suffices. One of the
best is acetic-alum carmine (boil excess of carmine in a saturated
aqueous solution of potash-alum for about fifteen minutes; add glacial
acetic acid to the extent of 10 per cent.; let it stand for a week;
filter). For use, dilute about thirty times with water. Place the
flukes directly in the stain. Stain overnight or longer.
_Differentiation._--In order to get the sharpest picture, it is best
now to differentiate (but this may often be omitted) with acid alcohol
(70 per cent. alcohol 100 parts, HCl 5 drops). Allow to act from one
to twenty-four hours, according to the appearance of the flukes.
Similarly, in staining with _hæmatoxylin_ solution, dilute twenty
to thirty times so that the water is merely tinged with the stain.
Differentiate as before. After staining, dehydrate, clear, and mount in
balsam if required.
_Clearing and Mounting._--(1) _Carbolic acid_ (carbolic acid 94,
water 6) is a very convenient clearing agent. It may be used for
stained or unstained specimens. It will clear rapidly without previous
dehydration. If it is required to mount a specimen permanently,
transfer from carbolic to alcohol, then cedar-wood oil (or xylol,
etc.), then balsam.
(2) _Creasote._--Dehydrate the specimen, stained or unstained, transfer
to creasote. If it is desired to mount permanently, transfer back to
alcohol, then cedar-wood oil, then balsam.
(3) _Cedar-wood Oil._--Preferable to xylol or oil of cloves. Dehydrate
the specimen in alcohol. To mount permanently, transfer to balsam.
(4) _Glycerine._--_Vide_ under methods of preservation of ova; to mount
permanently, transfer to glycerine jelly; subsequently to harden the
jelly, expose to formalin vapour.
Of these media, carbolic acid has the greatest refractive index
excepting that of balsam. The latter may, in some cases, render
structures too transparent, and it may be advisable to use only
glycerine or glycerine jelly.
PRESERVATION OF OVA IN FÆCES, URINE, BILE, ETC.
Heat some 70 per cent. alcohol in a basin to about 60 to 70° C. (until
bubbles begin to appear). Add the fæces, etc., in the proportion of one
part to about nine of fixative; keep stirring. Allow the sediment to
settle. Transfer to a bottle with some fresh 70 per cent. alcohol.
_Transference to Glycerine._--Prepare 5 per cent., 10 per cent., 20
per cent. solutions of glycerine in 70 per cent. alcohol. Pour off
the alcohol in the bottle of fæces, etc., and replace by 5 per cent.
glycerine solution. Allow to stand an hour or so. Then in the same way
replace the 5 per cent. by a 10 per cent. glycerine, and finally by a
20 per cent. glycerine solution. When in this latter expose freely to
the air (protecting from dust), so as to allow the alcohol and water
to evaporate. Add a few drops of glycerine from time to time till
eventually the ova are in pure glycerine. (In a very moist climate it
may be necessary to use lime or calcium chloride to dry the air.) To
mount permanently transfer some of the sediment to glycerine jelly.
PRESERVATION AND EXAMINATION OF CESTODES.
_Fixation._--(1) _Saturated aqueous corrosive sublimate._--Add to this
glacial acetic acid to the extent of 1 per cent. (Note this fixative
will dissolve the “calcareous corpuscles”; 10 grammes of sublimate to
160 c.c. of water will give a saturated solution.) Warm the fixative
to 70° to 80° C. (Avoid the use of needles.) Use plenty of fixative.
Allow to act for a quarter of an hour or so. (_a_) Transfer to 70 per
cent. alcohol. (It is advisable to remove the sublimate by the use of
Lugol’s solution, or a solution containing tincture of iodine, adding
this until the iodine colour is permanent.) Or (_b_) transfer for
preservation to 10 per cent. formalin.
Or (2) 10 _per cent. formalin_.--In order to prevent contraction it is
advisable to extend the tapeworm and keep it fixed by glass plates, or
wind the worm around a wide glass tube or bottle, and then fix it.
Or (3) _fix in hot alcohol_.
_Staining._--As under flukes. It is necessary to sacrifice portions of
the tapeworm for this purpose, cutting out, _e.g._, mature segments, so
as to study the topography of the genitalia.
_Clearing._--As under flukes. To examine the hooks satisfactorily it is
best to cut off the head with a sharp knife and mount. A certain amount
of pressure is then advisable in order to view the hooks completely so
as to measure them.
PRESERVATION OF OVA IN FÆCES, ETC.
As under flukes.
PRESERVATION AND EXAMINATION OF NEMATODES.
_Fixation._--(1) Thoroughly wash the worms to get rid of mucus, etc.,
by shaking up in warm saline (or water) till the washings are clean.
Then transfer to 70 per cent. alcohol _heated_ to about 70° C. It is
absolutely necessary to use _hot_ fixatives in order to _extend_ the
worms. If no alcohol or spirit is immediately available, drop the worms
into _hot_ water, or saline, and transfer later to 70 per cent. alcohol.
(2) Drop into hot 10 per cent. formalin.
_Clearing._--(1) Carbolic acid, _vide_ p. 471.
(2) Creasote, _vide_ p. 471.
(3) Glycerine, _vide_ p. 472.
_Staining._--In case of quite small Nematodes, _e.g._, _Anguillulidæ_,
carmine may be used, but as a rule staining is not advantageous.
_Rolling._--In order to study the mouth parts, or bursa, etc., it is
necessary to place the worm in any desired position. This is done as
one would roll a penholder along the table by one’s finger placed on
top of it. In the case of a worm, one edge of the cover-glass is placed
over the worm, the other is supported by a strip of cardboard. By
tapping the cover-glass the worm will now revolve as much as required
provided it is _round_ and _straight_. In certain cases it may be
necessary for this purpose to cut off the head or tail. Roll these
separately.
When a suitable position is got, the worm may be fixed in this position
by pressure on the cover-glass, so as slightly to flatten it.
_Mounting the Head._--If it is required to get an end view, it is
necessary to cut off the head transversely as near the end as possible,
and then mount.
_Detection of Eggs_ (Bass and Hall).--Mix the fæces thoroughly with
ten times the volume of water. Filter through gauze. Centrifugalize
the filtrate. Wash the sediment and centrifugalize. Repeat twice.
To sediment add CaCl_{2} solution, sp. gr. 1250. The eggs float
to surface. Pour off surface fluid. Dilute to sp. gr. 1050.
Centrifugalize. Examine sediment, which contains practically all the
eggs in the stool.
_Detection of Small Nematodes._--Mix the fæces thoroughly with water.
Allow to settle for five minutes. Carefully decant off, or better,
syphon off the fluid. Mix the sediment again with water. Allow to
settle. Remove the fluid. Repeat several times. Examine the sediment
in a Petri dish. As the fluid is poured off, the worms will be seen
collected in the backwater. Remove them with a brush. Fix in hot 70 per
cent. alcohol.
CULTIVATION OF LARVAL FORMS OF ANCYLOSTOMA AND STRONGYLOIDES.
A modification of the second method of Looss (p. 455) is that of
Fülleborn. A glass filter funnel is lined with linen or with cotton
wool dyed black with iron-tannin. On this is placed a layer of sterile
sand, and on top of this the fæces. The whole is moistened. The larvæ
hatch out and wander through the meshes of the wool, appearing on the
edges of the same as white threads visible to the naked eye. With a
platinum needle these can be easily removed. The glass filter can
be placed on a glass cylinder, and this in another large stoppered
cylinder containing caustic potash solution at the bottom, so that any
larvæ escaping from the funnel are killed.
D. *ACANTHOCEPHALA*, Rud.
Gutless, nematode-like worms that carry at their anterior end a
retractile rostrum beset with hooks. In their adult condition they
only live in vertebrate animals. During their larval stage they are
often parasitic in invertebrate animals.
The _Acanthocephala_ are elongated cylindrical worms, with a
rounded posterior end. In some species an annulation is distinctly
recognizable; they are, however, not segmented. The size varies
according to the species, between about 5 to 10 mm. and 40 to 50 cm.;
in general, however, there is a preponderance of the small species.
The sexes are separate, and the males can easily be distinguished
from the females without examination of the genitalia, as the females
are both larger and thicker.
The body wall of Echinorhynchus is limited by a thin cuticle, which
is attached inwardly to the hypodermis. In only exceptional cases
a syncytium with large nuclei, even in the adult condition, is
represented by the hypodermis; and in it fibre systems, the elements
of which run in layers in various directions, appear, and it is only
towards the interior from these strata of fibres that the nuclei of
the hypodermis are found. As a rule, these fibres, at all events
the radiary fibres, are regarded as muscles. Hamann describes them
as elastic fibres, which lie in a viscid gelatinous connective
substance (transformed protoplasm?); a lacune system filled with
a granular fluid, the central part of which are two longitudinal
lacunes lying at the sides, also belongs to the cutaneous strata, as
do the so-called lemnisci, two short, flat organs suspended in the
body cavity, and the pedicles of which are attached anteriorly at the
border between the rostrum and body; their structure as well as their
origin permit them to be traced to the skin (fig. 348A).
Finally, inwardly below the skin there follows a layer of annular,
and after these a layer of longitudinal muscles, the structure
cells of which remain present in the residues, carrying nuclei. The
motor apparatus of the rostrum, the sheath of the rostrum, and the
lemnisci also belong to the muscular system. The rostrum represents a
finger-shaped hollow process of the cutaneous layer; but, according
to Hamann, it originates from the entoderm and passes through the
skin secondarily. It is covered by a thin cuticle, and as a rule
contains a large number of regularly placed chitinous hooks that
adjoin a granular formation tissue. From the base of the rostrum
springs a tubular hollow muscle extending into the body cavity; this
is the RECEPTACULUM PROBOSCIDIS, from the base of which again bundles
of longitudinal muscles originate, which pass along its axis and
that of the rostrum itself, and are inserted at the inner surface of
its anterior end (RETRACTOR PROBOSCIDIS). These muscles when they
contract invaginate the proboscis and draw it into the receptaculum;
when reversed they act again as PROTRUSOR PROBOSCIDIS. The whole of
the anterior body, however, can be invaginated, and for this purpose
there is a muscle that originates from the body wall at a variable
distance back, and which is joined to the receptaculum (RETRACTOR
RECEPTACULI); there is also a bell-shaped muscle which springs from
the body wall behind the lemnisci in rings, and passes forward to the
spot of attachment of the lemnisci.
The nervous system consists of a cluster of ganglia situated at the
base of the rostrum, from which three nerves pass towards the front
and two towards the back. No sensory organs are known.
The excretory organs, according to Kaiser, lie at the upper border of
the ductus ejaculatorius in the male and at the so-called bell in the
female. Here they represent the long-known villous tufts, placed on
disc-like cushions. In each of the cylindrical villi--which terminate
blindly towards the body cavity--there lies a cilium, which springs
from the membrane lining the villus, and which lies in a space cavity
of the villus, which ultimately proceeds as a little canal. There are
three canals discharging into the uterus that serve to conduct the
excretory materials from the body cavity; special glandular cells
corresponding to the terminal cells of the Platyhelminths, at the
commencement of the system, are not present in the Acanthocephala.
SEXUAL ORGANS.
(_a_) _Male Organs._--The greatest part of the male genital apparatus
is contained in a muscular sheath--the ligament--which originates
at the posterior end of the receptaculum proboscidis, passes along
longitudinally through the body cavity, and is inserted at the
posterior end of the worm. The two oval testicles usually lie one
behind the other; their vasa efferentia unite sooner or later into a
vas deferens which passes backwards, and finally terminates in the
penis; the terminal portion of the conducting apparatus is surrounded
by six large glandular cells (prostatic glands) the excretory ducts
of which open into the vas deferens. The penis itself is placed at
the base of a bell-shaped invagination of the posterior end, the
bursa, which is everted during copulation.
[Illustration: FIG. 348A.--The male of _Echinorhynchus augustatus_.
_L._, lemnisci; _T._, testicles; _P._, prostatic glands; _P.r._, sheath
of proboscis, with ganglion; _R.r._, retractor of sheath of proboscis.
25/1.]
[Illustration: FIG. 348B.--Anterior portion of the female apparatus of
_Echinorhynchus acus_. On the left seen from behind, on the right seen
from the front. _F_, inferior orifice of the bell; _B_, bell; _Lig_,
ligament; _M_, mouth of bell; _Ut_, uterus. Magnified. (After Wagener.)]
(_b_) _Female Organs._--There are only two ovaries present in the
ligament during the larval stage. During the course of growth they
divide into accumulations of cells (placentulæ, loose or floating
ovaries), which finally cause the ligament to burst and they thus
attain the body cavity. Thence a peculiarly constructed apparatus
finally conveys the eggs out. This apparatus consists of the uterine
bell and vagina, the latter discharging at the posterior extremity
of the body. The bell is a muscular canal provided with apertures at
both the anterior and posterior extremities. Its interior space is in
direct communication with the body cavity, and the anterior orifice
takes up all materials floating in the cavity--egg-balls, mature and
immature eggs--and pushes these further backwards. The continuation
of the bell lumen is now narrowed by a number of large cells in such
a manner that only bodies of a certain form can pass through this
tract and attain the uterus; everything else is conveyed back into
the body cavity through the posterior opening of the bell.
The eggs are already fertilized in the body cavity, and in this
position go through their development to the formation of the embryo.
Completely developed eggs are surrounded by three shells, and are
generally fusiform. The eggs agglomerate in masses in the uterus
until they are finally deposited through the vagina and vulva.
For the further development, the transmission of the eggs into an
intermediary host--usually a crustacean or an insect--is necessary;
the metamorphosis is very complicated; but this transmission may be
very easily effected artificially by feeding suitable crustaceans
(_Asellus_, _Gammarus_, etc.) with the eggs of _Acanthocephala_; this
being the only method of inducing the larva to hatch out so that
its structure may be studied. The larva appears in the form of an
elongated, somewhat bent body, at the stumpy anterior end of which
there is a crown of hooks or spines, whereas the posterior end is
pointed. Especial retractors draw in the hook-beset anterior surface,
and an elastic cushion beneath them jerks them forward again when
required. In the middle of the body a roundish heap of small cells is
seen, from which the entire body of the Echinorhynchus originates,
even to the cutaneous layer; the latter is also the larval skin in
which the small Echinorhynchus gradually grows. The development of
all the organs takes place within the intermediary host, and the
parasite only needs to be imported into the terminal host to attain
the adult stage after a certain growth. In some cases, however, a
second intermediary host is utilized.
Species of _Acanthocephala_ only occur exceptionally in human beings.
*Echinorhynchus gigas*, Goeze, 1782.
Syn.: _Tænia hirudinacea_, Pallas, 1781.
[Illustration: FIG. 348C.--Egg of _Echinorhynchus gigas_. 300/1. (After
Leuckart.)]
The body is elongated, gradually decreasing in thickness towards the
back. The rostrum is almost spherical, and is beset with five or six
rows of recurved hooks. The males measure 10 to 15 cm. in length,
the females 30 to 50 cm.; the eggs are provided with three shells,
of which the middle one is the thickest. The eggs measure 0·08 to
0.1 mm. in length. The GIANT ECHINORHYNCHUS occurs especially in the
intestinal canal of the domestic pig; it is less common in other
mammals. It bores deep into the mucous membrane with its rostrum,
and causes an annular proliferation around the perforated spot;
occasionally also it causes perforation of the intestine.
It is doubtful whether the giant Echinorhynchus occurs in man.
Leuckart admitted that there were a few positive cases. According
to Lindemann, _Ech. gigas_ occurs in human beings in South Russia,
and its presence is not rare. This statement, however, has not been
confirmed. Its presence in man is by no means impossible, as its
intermediary host, the cankerworm, or cock-chafer (_Melolontha_),
is, according to Schneider, occasionally eaten raw by human
beings. According to Kaiser, the golden beetle (_Cetonia aurata_)
and, according to Stiles, another beetle in America (_Lachnosterna
arcuata_) are also intermediary hosts.
*Echinorhynchus hominis*, Lambl, 1859.
This term is applied to an ECHINORHYNCHUS found by Lambl in the
intestine of a boy who had died of _leucæmia_; the worm was 5·6 mm.
in length, and the almost spherical head was beset with twelve
transverse rows of hooks.
*Echinorhynchus moniliformis*, Bremser, 1819.
The male is 4 cm. in length, the female 8 cm. long. This species
lives in the intestine of field-mice, rats, marmots and _Myoxus
quercinius_. A beetle (_Blaps mucronata_) is the intermediary host.
This species has also once been artificially cultivated in man
(Grassi and Calandruccio).
E. *GORDIIDAE.*
Very long thin worms similar to Filariæ, which, in their adult
condition, live free in brooks, pools and springs; the mouth and the
commencement of the intestine are obliterated; there are no lateral
ridges, and the muscular system presents a structure different to
that of the _Nematoda_. The posterior end of the male is split, and
spicules are lacking; there are two testicles. In both sexes the
genitalia discharge through the terminal gut.
The larvæ, which carry a rostrum beset with hooks, force themselves
into the larvæ of water-insects; more rarely they invade molluscs,
and they then become encysted within the body of the host. According
to Villot, at least a part of them attain the intestine of fishes,
where they again become encysted, and after a period of rest they
travel into the tissues of their hosts, and finally again reach the
exterior by way of the intestine, where they then become adult. In
most cases, however, the gordius larvæ are taken up by predacious
water insects; they live for a while in the body cavity of these
insects, undergo a metamorphosis, and finally wander into the water.
A few species invade man accidentally with water, in which case they
are usually vomited up:--
_Gordius aquaticus_, Dujardin, 30 to 90 cm. in length (Aldrovandi,
Degland, Siebold, Patruban).
_Gordius tolosanus_, Duj., 11 to 13 cm. in length (Fiori).
_Gordius varius_, Leidy, 10 to 16 cm., female, up to 30 cm. in length
(Diesing).
_Gordius chilensis_, Blanch. (Guy). _Gordius villoti_, Rosa
(Bercutti, Camerano); _Gordius tricuspidatus_, L. Def. (R.
Blanchard), _Gordius violaceus_, Baird (Topsent), and _Gordius
pustulosus_, Baird (Parona).
F. *HIRUDINEA s. DISCOPHORA* (Leech).
The _Hirudinea_, which have been appropriately included amongst the
Annelida, differ in many respects from the typical members of the
group; their body is long and flat, it lacks the parapodia that
are characteristic to all forms of Annelida; but, on the other
hand, possesses a terminal posterior sucker, and in many species
there is also an anterior sucker. The mouth is terminal at the
anterior end, the anus lies dorsally above the posterior sucker
(fig. 348D). The body is segmented, but this is less manifest in
the body covering than it is in the arrangement of the internal
organs; the segmentation, nevertheless, is also indicated exteriorly
by the appearance of the cutaneous sensory organs which correspond
to the segments. This shows what the condition of the ganglia in
the abdominal ganglion chain has taught us, that the anterior and
the most posterior segments are considerably abbreviated--a part of
the latter taking part in the formation of the suctorial organs.
In a great many species the skin is distinctly annulated, four or
five of such rings, at least in the central region of the body,
appearing on one segment of the body. The condition of their body
cavity is another peculiarity of the _Hirudinea_; it is narrowed by
the powerful development of the connective tissue and the muscular
system into four tubular sinuses, which have the appearance of
blood-vessels. There are usually one dorsal and one ventral median
trunks, as well as two lateral trunks; in addition, a particular
blood-vessel system exists.
[Illustration: FIG. 348D.--The internal organs of the leech. The
creature has been opened from the dorsal surface, and part of the
intestine has been removed. The testicles, with vas deferens, may be
seen between the blind ducts of the intestine; beyond these on either
side the segmental organs. The female genital organs are in front of
the most anterior pair of testicles. (After Kennel.)]
The skin consists of a very thin cuticle that is cast off from time
to time; it is secreted by the underlying cylindrical epithelium,
which contains numerous goblet cells. The muscular system is
strongly developed; it consists of long tubular fibres, which run
circularly, longitudinally and in the dorso-ventral direction; the
muscular system is subject to a particular expansion in the clinging
organs and at the commencement of the intestine. On the whole, the
alimentary canal represents a tube running straight from the mouth to
the anus, which possesses a number of blind sac-like protuberances
at the sides varying according to the species. The most anterior
section, the pharynx, in the leeches with maxillæ carries three
chitinous, semicircular plates furnished with teeth--the jaws--which
serve to tear up the epidermis in order to open the blood-vessels;
in the leeches with rostra a long protractile proboscis rises from
the base of the elongated pharynx. Numerous salivary glands, the
secretion from which possesses toxic properties, discharge into
the pharynx. The œsophagus, which follows the pharynx, and to the
exterior of which numerous radiary muscles are fixed, is a suctorial
organ in its entire structure. The nutriment in the larger species
consists of the blood of vertebrate animals, in smaller species and
in the young stages the food consists of small invertebrate animals.
The NERVOUS SYSTEM exhibits the typical structure of other segmented
worms; the sensory organs consist of the previously mentioned
goblet-shaped cutaneous sensory organs, of the organs of taste, and
of eyes, the latter frequently being present in large numbers.
The EXCRETORY or segmental organs exhibit many peculiarities, which
cannot, however, be detailed here. They commence with funnels in the
lacunes of the body cavity, and usually discharge on the ventral
surface.
Almost all the _Hirudinea_ are hermaphrodite and copulate
reciprocally. The two ovaries are very small, and the oviducts that
proceed from them soon unite into a common duct, which then passes
into the uterus and discharges through the short vagina in the
median line of the ventral surface behind the male organs into the
so-called clitellar region. The male sexual apparatus consists of
symmetrically arranged testicles, varying in number according to the
species, the short vasa efferentia of which, one by one, run into the
vas deferens, passing towards the front on each side. In front, at
about the level, or a little in front, of the female genitalia, the
two vessels pass into a convoluted mass of tubes to the so-called
epididymis, and then discharge into the single protractile penis
(fig. 348D).
All leeches deposit so-called COCOONS. These are small barrel-shaped
or pouch-like bodies, which are surrounded by a thicker shell and
contain a number of eggs in a large mass of albumen; the albumen
originates from glands of the generative organs, the shell substance
from cutaneous glands of the clitellar region.
Family. *Gnathobdellidæ* (Leeches with Jaws).
These are distinguished by the possession of usually three jaws in
the pharynx; the body consists of twenty-six segments. The posterior
sucker is large and flat; the anterior sucker is smaller. The
_Hirudinea_ have five pairs of eyes, the _Nephelinæ_ have four pairs.
Genus. *Hirudo*, L., 1758.
The entire body consists of 102 annulations, five appearing on one
segment in the central region of the body. The pharynx has three
semicircular jaws, the arched border of which is beset with numerous
teeth (50 to 100). The male sexual orifice lies between the thirtieth
and thirty-first rings, the female orifice between the thirty-fifth
and thirty-sixth. There are numerous species, some of which are
utilized for medicinal purposes.
*Hirudo medicinalis*, L., 1758.
[Illustration: FIG. 348E.--_Hirudo medicinalis._ _a_, anterior end,
with open buccal cavity, with the jaws, _J_, at the; _b_, one jaw
isolated. (After Claus.)]
It occurs in numerous colour varieties, one of which has been
designated _Hirudo officinalis_, Moq.-Tandon. Usually the dorsal
surface is greyish-green and is marked with six rusty-red
longitudinal stripes. The ventral surface is olive-green, more
or less spotted with black, and marked at the sides with a black
longitudinal line. The length averages 8 to 12 to 20 cm. This leech
lives in swamps, ponds and brooks, overgrown with plants and having
a muddy bed. The cocoons are deposited in the soil at the sides.
Europe, as well as North Africa, is its home. At the present day it
has been exterminated from most parts of Central Europe, but it is
still very common in Hungary. Its use for medicinal purposes is well
known. A large leech can suck about 15 grs. of blood, and about the
same amount is lost through secondary hæmorrhage.
*Hirudo troctina*, Johnston, 1816.
Syn.: _Hirudo interrupta_, Moq.-Tandon, 1826.
This species measures 8 to 10 cm. in length. The back is greenish,
with six rows of black spots surrounded by red; the lateral borders
are orange-coloured; the abdomen spotted or unspotted. Its habitat is
in North Africa and Sardinia. It is applied medicinally in England,
Spain, France, Algeria, etc.
Genus. *Limnatis*, Moq.-Tandon, 1826.
Nearly related to Hirudo, but is differentiated by a longitudinal
groove on the inner surface of the upper lip of the anterior sucker.
The jaws are furnished with over 100 very sharp toothlets.
*Limnatis nilotica*, Savigny, 1820.
Syn.: _Bdella nilotica_, Sav.; _L. nilotica_, Moq.-Tandon; _Hæmopis_
(_vorax_), Moq.-Tandon, 1826, _p. p._; _Hæmopis sanguisuga_,
Moq.-Tandon, 1846 (_nec Hir. sanguis_, Bergm., 1757).
This species measures 8 to 10 cm. in length, and becomes gradually
more pointed towards the front; the body is always soft. The back is
brown or greenish, and has usually six longitudinal rows (rarely only
two or four) of black dots. The abdomen is dark; but numerous colour
variations occur.
The native place is North Africa, especially the coastal regions; it
is also found in the Canaries, the Azores, Syria, Armenia, Turkestan,
perhaps also Southern Europe. It is taken into the mouth with
drinking water, and may settle in the pharynx, larynx, œsophagus, and
nasal cavities of human beings. This species has also been observed
in the vagina and on the conjunctiva. It is equally fond of attacking
domestic animals.
_Hirudo mysomelas_ (Senegambia) and _Hirudo granulosa_ (India) are
placed with this genus, and, like our leech, are also used for
medicinal purposes.
Genus. *Hæmadipsa*, Tennent, 1861.
These leeches live on land, and measure 2 to 3 cm. in length. About a
dozen species are known. They are a veritable scourge to persons in
the tropics (Asia, South America), as they attack them to suck their
blood. They are able to force their way even under close-fitting
garments, so that it is difficult to protect oneself from their
assaults (_Hæmadipsa ceylonica_, Bl., and other species).
Family. *Rhynchobdellidæ* (Leeches with Rostrum).
These are furnished with a proboscis in lieu of the jaws; the segment
consists of three annulations.
Genus. *Hæmentaria*, de Filippi, 1849.
*Hæmentaria officinalis*, de Fil.
Inhabit Mexico, where they are used for medicinal purposes.
Genus. *Placobdella*, R. Blanch.
*Placobdella catenigera*, Moq.-Tandon.
Indigenous to South Russia, Hungary, Italy and South France. It is a
parasite of the swamp turtle, but frequently attacks human beings.
G. *ARTHROPODA* (Jointed-limbed Animals).
BY
FRED. V. THEOBALD, M.A.
BILATERALLY symmetrical segmented animals which are covered with a
thick cuticle that is frequently calcareous (_Crustacea_), but always
thinner between the segments; they carry (primitively) a pair of
jointed appendages on every segment.[321] The segments of the body
are uniform in certain regions, but differ from those of contiguous
regions, so that it is easy to distinguish three parts (head, thorax
and abdomen), each composed of segments. The cephalic segments are
always formed into a uniform head, the segmentation being scarcely
recognizable at either end; the thoracic segments may also fuse, or
part or all of them may coalesce with the head; the abdomen, as a
rule, retains its segmentation, but this may possibly also be lost,
in which case it is [sometimes] united to the cephalothorax. The
structure of the three regions depends mostly on the varying form
and function of the appendages: those on the head are primitively
locomotive organs (and frequently are still so in the early stages),
but they become transformed into feelers and mouth-parts (mandibles,
maxillæ); the limbs of the thorax, however, usually retain their
ambulatory functions, as frequently do those of the abdomen;
sometimes, however, the abdominal limbs disappear, entirely or
partly; in the latter case they are then utilized for other purposes.
[321] [In most _Arthropoda_ the skin is hardened by a deposit of chitin
(_Hexapoda_, etc.).--F. V. T.]
In their organization the _Arthropoda_ approach the segmented worms.
The _Arthropoda_ are generally divided into five groups
(_Crustacea_,[322] _Protracheata_, _Arachnoidea_, _Myriapoda_,[323]
and _Insecta_ or _Hexapoda_), of which only the _Arachnoidea_ and the
_Hexapoda_ interest us here.
[322] Parasitic or free-living Crustaceans may now and then invade man
abnormally. Thus, according to Betten, _Caligus curtus_ invade the
cornea (Betten, R. A., “Par. Crust. as a Foreign Body on the Cornea,”
_Lancet_, 1900, i, p. 1002; and _Centralbl. f. Bakt. u. Par._, xxix,
p. 506). According to Laboulbène, also _Gammarus pulex_ (Laboulbène,
A., “Obs. d’accid. caus. par le _G. pul._ apport. avec l’eau de boison
dans l’estomac d’un homme,” _Bull. Acad. méd._, 1898, p. 21).
[323] R. Blanchard has compiled thirty-five cases in which _Myriapoda_
have been observed in the intestine as well as in the nose of human
beings (“Sur le pseudopar. d. myriap. chez l’homme,” _Arch. de Par._,
1898, i, p. 452). E. Munoz Ramos reports an additional case (_ibid._,
p. 491). A few years ago a doctor in East Prussia sent me a rain
worm out of a lady’s nose (_cf._ Hanau, A., “Wahrsch. Pseudo-paras.
v. Schmeissfliegenlarv. u. angebl. Paras. v. Regenwürmern b. einer
Hysterischen,” _Arch. de Par._, 1899, ii, p. 23).
_A._ *ARACHNOIDEA* (Spiders, Mites, etc.).
The head and thorax are always united together; the abdomen is either
segmented or without exterior segmentation, in which case it is united
with the cephalothorax.[324] The number of pairs of appendages amount
to six, of which the two front pairs, the cheliceræ and the pedipalpi,
are attached to the head region and the four remaining pairs to the
thoracic region.[325] The abdomen in the adult condition has no
appendages. The Arachnoids are air-breathers, and for this purpose
are either provided with tracheæ or with so-called lung-sacs, or they
breathe through the surface of the body. Some aquatic forms breathe by
gills.
[324] [This is only so in the _Acarina_ or mites, not in the _Araneida_
or spiders.--F. V. T.]
[325] [The true character of the _Arachnoidea_ is the presence of
four pairs of ambulatory appendages. This number is reduced to two
pairs in the gall-making _Phytoptidæ_, and they differ from all other
_Arthropoda_ in having no antennæ.--F. V. T.]
There are eight or ten orders of Arachnoids,[326] of which, however,
only two, the _Acarina_ and the _Linguatulida_, have to be considered
here.[327]
[326] Twelve orders are now recognized, as follows: _Pentastomida_
or Linguatulids; _Tardigrada_ or bear-animalcules; _Phalangidæ_ or
harvest-men; _Acarina_ or ticks and mites; _Palpigradi_; _Solifugæ_;
_Pseudoscorpionidea_ or book mites; _Pedipalpi_ or false scorpions;
_Scorpionidea_ or true scorpions; _Araneida_ or spiders; _Xiphosura_ or
king crabs; and _Pycnogonida_, marine Arachnoids.
[327] _Chelifer cancroides_ has also been observed as a pseudoparasite
in man (Arnault de Very, S., “Pseudopar. du. _Chel. cancr._ chez
l’homme,” _Compt. rend. Soc. de Biol._, 1901, liii, p. 105).
Order. *Acarina* (Mites).
Small Arachnoids, the three parts of the body of which are, as a
rule, coalesced; it is only rarely that a faint line indicates the
division between a cephalothorax and abdomen. The two appendages
on the head are designed for biting or puncturing and sucking, and
vary according to their use. The cheliceræ[328] are fang-like jaws
or puncturing bristles forming a kind of rostrum, the pedipalpi are
claw-like or shear-shaped, or form a suctorial proboscis.[329] The
four pairs of legs are usually well developed, more rarely they
are rudimentary or have partly vanished; many parasitic forms are
provided with pedunculated suckers [ambulacra--F. V. T.]. Respiratory
organs (tracheal tufts) may be present or absent. The nervous system
is reduced to a minimum, eyes are usually lacking. The intestine,
situated in the central part, generally has three blind appendages;
the anus is situated on the venter above the posterior end. Sexes
separated; nearly all the species deposit eggs, from which six-legged
larvæ hatch. The _Acarina_ live either free in the water or in moist
soil, or they are parasitic on plants and animals.[330]
[328] [The cheliceræ are sometimes regarded as modified antennæ, but it
is more natural to regard them as the morphological equivalent of the
mandibles of _Hexapoda_.--F. V. T.]
[329] [The pedipalpi, or second pair of jaws, consist of a stout
basal segment and a palp, which may have the appearance of a leg in
_Arachnida_; this may end with or without a claw, or with a chela
(scorpions); they may also form a tube enclosing the styliform
cheliceræ (mites).--F. V. T.]
[330] [_Acarina_ are also found living upon trees, feeding upon other
Arthropods and also upon spores of lichen and fungi (_Oribatidæ_ or
beetle mites); they also swarm indoors amongst stores and provisions
(_Tyroglyphidæ_ and _Glyciphagi_, household, sugar and cheese mites).
This order is very important, as many are parasites upon man, his
domestic animals and his cultivated plants, and attack his provisions
and stores. Some live on blood, and in some of the ticks distribute
various protozoal and other blood parasites and germs.--F. V. T.]
Family. *Trombidiidæ* (Running Mites).
Soft-skinned _Acarina_ with tracheæ and with two eyes, usually
pedunculated; they are often brightly coloured; cheliceræ lancet- or
claw-shaped; pedipalpi claw-like; legs composed of six segments, with
suctorial discs between the terminal ungues.[331] Larvæ six-legged.
To the latter belong the larvæ of several species of Trombidium such
as:--
[331] [Some have seven segments to the legs.--F. V. T.]
Genus. *Trombidium*, Latreille (and *Leptus*).
*Leptus autumnalis*, Shaw, 1790.
Leptus occur as parasites in the human skin and cause a cutaneous
disease known as autumn erythema, and produce a very unpleasant
sensation on account of the troublesome itching; in children it is very
often accompanied by fever.[332]
[332] [This minute parasite is especially obnoxious in barley fields.
In walking over barley stubble one is sure to be attacked by this
Acarus in many districts. Trombidium is often very prevalent in
gardens, especially along rows of peas, and in spring they may be found
on fruit trees and bushes. Nut-pickers are frequently attacked by
Leptus, and also pickers in other fruit plantations. It is often called
the harvest mite.--F. V. T.]
[Illustration: FIG. 349.--_Leptus autumnalis_, with so-called sucking
proboscis. Enlarged. (After Gudden.)]
[Illustration: FIG. 350.--_Leptus autumnalis_: the so-called proboscis
is formed around the hypopharynx sunk into the skin. 100/1. (After
Trouessart.)]
Formerly these mites were considered adult forms, but when they were
recognized as mite larvæ they were taken for those of the spider-mite
(_Tetranychus telarius_); the investigations of Hanstein, however,
showed this to be a mistake. When Henking first investigated the
development of _Trombidium fuliginosum_, parasitic in the larval
stage on vine-fretters, he demonstrated the occurrence of a form
very similar to _Leptus autumnalis_, and the “autumn, grass, or
gooseberry” louse was commonly designated the Trombidium larva. Even
before Henking’s work it had been described by Mégnin as the larva
of _Trombidium holosericeum_, a red-coloured species, frequently
occurring in spring and summer on the ground, trees, etc. This
assumption, however, as Moniez was the first to explain, is not
correct; indeed, as many as three species come under consideration:
_T. gymnopterosum_, L., _T. fuliginosum_, Herm. (according to
Brucker), and two species known hitherto only in the early stage, _T.
striaticeps_, Helm. et Oudem., and _T. poriceps_, Helm. et Oudem.,
which are not only parasitic on mammals, but on birds, on Arthropods
and especially on insects. Arthropods appear to be the normal hosts
for the larvæ.
The above-mentioned forms invade the skin of man by means of their
oral apparatus, by preference invading the orifices of the sebaceous
glands so as to suck the blood; around the point attacked there arises
a wheal about the size of a lentil, and around the inserted hypopharynx
a fibrinous secretion, the “proboscis,” which, however, is a product
of the host, just as chitinous secretions are provoked by Trombidia
parasitic on Arthropods.
Further species, analogous in habit to _Leptus autumnalis_, are
described by Riley from Central and South America as _L. americanus_
and _L. irritans_.
[_L. autumnalis_ attacks small mammals by preference, such as moles
and hares, which are often literally covered with them. Dogs are also
subject to their attack, and cats suffer similarly. This mite also
frequently appears in colonies on cows; cavalry horses after autumn
manœuvres often suffer from an erythematous affection about the hocks
and knees due to this pest.
[A number of Leptus, so far undescribed, occur abroad which attack man
in the same way as _L. autumnalis_ in Europe. Dr. Durham has brought
me specimens from British Guiana called _bête rouge_; this species
works under the skin much as does our European species, but it is very
distinct, being considerably larger.--F. V. T.]
*Trombidium tlalsahuate*, Lemaire, 1867.
_T. tlalsahuate_ occurs in Mexico under conditions similar to those of
Leptus here. It also frequently attacks men, and especially fastens
itself on to the eyelids, in the axillae, navel, or on the prepuce; it
induces itching and swelling of the parts affected, and sometimes even
causes suppuration; the symptoms, however, generally disappear after a
week and remain localized.[333]
[333] Lemaire, “Import. en France du tlalsahuate,” _Compt. rend. Acad.
Sci._, Paris, 1867, lxv, p. 215.
Other species of mites which attack man are reported, mostly by
travellers, from various other places; zoologically, however, there
is little known about them. The pou d’agouti in Guiana, niaibi in New
Granada, colorada in Cuba, mouqui in Para, and the buschmucker in New
Guinea represent a few of these.
*Akamushi* or *Kedani*.
In a few districts of Japan there occurs a serious illness, with a
mortality of 40 to 70 per cent. It is called river or flood fever, and
the Japanese doctors have connected it with a small mite (akamushi,
kedani). Baelz has opposed this opinion on the grounds that he has
repeatedly observed the same species of mite in his dwelling without
any subsequent illness occurring. According to Keïsuke Tanaka, however,
a connection certainly does exist, inasmuch as the akamushi, like
Leptus, attacks persons to suck blood. If the mite is not removed,
or if the spot attacked is injured by scratching, etc., a papule
surrounded by a red area forms, and a pustule ensues; and finally a
black scab covers the seat of injury. The lesion becomes the point of
entrance of bacteria, especially a species of _proteus_ which produces
river fever. If the mites are carefully removed no general illness
takes place.
[Illustration: FIG. 351.--The kedani mite. Enlarged. (After Tanaka.)]
The orange-red mites, which we only know in their larval condition,
measure 0·16 to 0·38 mm. in length by 0·10 to 0·24 mm. in breadth. They
have leg-like palpi with three joints, hirsute bodies, and very hairy
legs composed of five segments, terminating with three ungues.
Family. *Tetranychidæ* (Spinning Mites).
These have tracheæ and eyes; the palpi are composed of four segments,
of which the last but one has a powerful claw. The legs have six
segments with sucker discs between the claws.
[The red spiders or spinning mites (Tetranychi) are usually placed in
the family _Trombidiidæ_.--F. V. T.]
Genus. *Tetranychus*, Dufour.
*Tetranychus molestissimus*, Weyenbergh, 1886.[334]
[334] [This species is also known as _Bicho colorado_. It spins a web
under the lower surface of the leaves, and it is only from December to
February that it attacks warm-blooded animals and man.--F. V. T.]
Found in Argentine and Uruguay on the under surface of the leaves of
_Xanthium macrocarpum_; it attacks mammals and men, producing severe
itching, accompanied by fever in the latter.
It has been asserted by Haller that the CAPE AILMENT (Port Natal
sickness) is caused by mites, but this statement has been contested.
*Tetranychus telarius*, L., 1758,[335] var. *russeolus*, Koch.
[335] [There is something wrong here, probably in the identification.
_T. telarius_ is purely a plant-feeder, and it is extremely unlikely
a variety would attack man. Anyhow, it will not do so in Great
Britain.--F. V. T.]
[Illustration: FIG. 352.--_Tetranychus telarius_ var. _russeolus_,
Koch. Enlarged. (After Artault.)]
This common spinning mite likewise attacks human beings, but the
papules produced by it very soon disappear.
Family. *Tarsonemidæ.*
A family distinguished by complete sexual dimorphism, the species of
which are provided with tracheæ; the legs have five segments; the
terminal segments of the front pair of legs of both sexes possess a
claw; the terminal segment of the posterior pair of legs of the male
likewise has a claw. In the female this pair of legs, like the second
and third pairs of both sexes, is provided with two hooklets and a
sucking disc. The cuticle of the body on the back is “annulated.”
[This family of small transparent mites live normally as plant
parasites. The last two pairs of legs are widely separated from the two
front pairs.--F. V. T.]
Genus. *Pediculoides.*
*Pediculoides ventricosus*, Newport, 1850.
Syn.: _Heteropus ventricosus_, Newport, 1850; _Acarus tritici_,
Lagrèze-Fossot, 1851; _Physogaster larvarum_, Lichtenstein, 1868;
_Sphærogyna ventricosa_, Laboulbène and Mégnin, 1885.
Males are oval in shape, 0·12 mm. in length and 0·08 mm. in breadth,
flattened. There are six pairs of chitinous hairs on the dorsal surface
and a lyre-shaped lamella on the posterior part. The female in the
non-gravid state is cylindrical in form, 0·2 mm. in length and 0·07 mm.
in breadth; when gravid the posterior part of the body becomes enlarged
into a ball, which may attain 1·5 mm. in size, as in the case of _Pulex
penetrans_ and of the female Termites. On emerging the young are
already provided with four pairs of legs and copulate soon after birth.
[Illustration: FIG. 353.--_Pediculoides ventricosus._ _a_, male; _b_,
young female; _c_, gravid female. Enlarged. (After Laboulbène and
Mégnin.)]
These animals live on the stalks of cereals, and feed on vegetable
and animal juices; they are also found on corn-infesting insects.
They invade the barns and seek out the insects living in the dry
grains of corn, or wait for an opportunity of obtaining food. They
have been repeatedly observed on human beings, particularly labourers
occupied in handling grain; their bite causes severe irritation,
local elevation and reddening of the epidermis, as well as fever. It
cannot be positively asserted that all cases of the occurrence of
cereal mites on man relate to _P. ventricosus_, as the descriptions
are often insufficient. Geber states that one form is _Chrithoptes
monunguiculosus_, or _Acarus hordei_; Flemming mentions _Tarsonemus
uncinatus_; Koller _Oribates_ sp.; and Karpelles _Tarsonemus
intectus_.
[The pregnant female Pediculoides has a large round inflated abdomen,
in which the ova hatch and the young mature. Later they escape from
the parent as adults.--F. V. T.]
Genus. *Nephrophages.*
*Nephrophages sanguinarius*, Miyake and Scriba, 1893.
Males measure 0·117 mm. in length and 0·079 mm. in breadth; females up
to 0·360 mm. in length by 0·120 mm. in breadth. The head is provided
with two very large scissors-like jaws and two large round eyes. The
legs are composed of five segments and are all of equal length; the
three anterior pairs of legs have pedunculated ambulacra, the posterior
ones terminate in a claw. The cuticle on the back is thickened in
three places, shield-like; the abdominal surface without scutellum
is longitudinally striped and is beset with chitinous hairs. Colour
greenish to brownish-yellow. Eggs 0·046 to 0·040 mm.
[Illustration: FIG. 354.--_Nephrophages sanguinarius_: male, ventral
surface. Enlarged. (After Miyake and Scriba.)]
[Illustration: FIG. 355.--_Nephrophages sanguinarius_: female, dorsal
aspect. Enlarged. (After Miyake and Scriba.)]
The authors discovered these mites, but always dead, in the urine
of a Japanese suffering from fibrinuria complicated with chyluria
and hæmaturia. They surmised that they were endoparasites, probably
situated in the kidney; but this view is not convincing, though they
also report that for a week, day after day, the mites were found in
the patient’s urine, as well as in urine drawn off by means of a
catheter, and in the water used to wash out the bladder (one or two
specimens and an egg). The statement that these mites have large eyes
makes the discovery suspicious, to say the least. The significance of
the discovery is not supported by the further statement that Disse
is supposed to have found an encapsuled mite closely related to the
Tyroglyphides on the wall of the vena cava.
In the case of Marpmann, who found a dead Acarid in the urine of a
man suffering from chronic nephritis, and in whom later examinations
proved negative, the author himself was of opinion that the mite had
reached the urine from outside.
We are certainly acquainted with mites living endoparasitically,
namely, the _Cysticolæ_, _Analgesinæ_, of which _Laminosioptes
gallinarum_ live in the intramuscular and subcutaneous connective
tissue of fowls, and _Cytoleichus sarcoptoïdes_ in their air sacs.
Another kind of mite (_Halarachne halichœri_) is occasionally found
in the nasal mucous membrane of the seal (_Halichœrus grypus_), and,
quite recently, _Pneumonyssus simicola_, which is more nearly related
to Halarachne, has been found in the lung of _Cynocephalus_ sp. It is
therefore not improbable that endoparasitic mites are found in man;
but no definite discovery has yet been made.
Family. *Eupodidæ.*
Small tracheate mites, with moderately long or short pedipalpi,
composed of four segments, of which the last segments bend; cheliceræ
forceps-shaped, with serrated edges; legs with two claws, more rarely
with one, and terminating in a tuft ornamented with fine hairs;
genital orifices on the abdomen, surrounded by a circle of little
hairs. Most species live free--one lives parasitically on the bodies
of slugs.
Genus. *Tydeus*, Koch.
*Tydeus molestus*, Moniez, 1889.
[Illustration: FIG. 356.--_Tydeus molestus_: seen in profile. Enlarged.
(After Moniez.)]
Male, 0·2 mm. in length, 0·125 mm. in breadth. Females, 0·225 mm. in
length, 0·135 mm. in breadth; gravid female 0·315 to 0·360 mm. in
length and 0·180 mm. in breadth. They were observed by Moniez on an
estate in Belgium, whither the creature had apparently been imported
twenty-five years previously with Peruvian guano; they appeared
regularly in the summer and remained until the first frost set in;
they were found on grass-plots, on trees and bushes in masses; they
regularly attacked human beings, mammals and birds, tormenting their
hosts in a terrible manner.
Family. *Gamasidæ* (Coleopterous or Insect Mites).
Cheliceræ chelate or styliform; pedipalpi filiform; the legs are
composed of six segments with two terminal ungues and a bladder-like
sucking disc [caruncle--F. V. T.]. Stigmata situated between the
third and fourth pairs of legs; the cuticle thickened, leather-like;
no eyes; the larvæ have six legs.
The _Gamasidæ_ are predaceous on small insects and other mites; some
are parasitic on insects, and one is noticeable as a pest on birds,
etc.
Genus. *Dermanyssus*, Dugés.
*Dermanyssus gallinæ*, de Geer, 1778.
Syn.: _Pulex gallinæ_, Redi, 1674; _Atarus gallinæ_, de Geer, 1778;
_Dermanyssus avium_, Dugés, 1834.
The male measures 0·6 mm. in length by 0·32 mm. in breadth; the female
0·7 to 0·75 mm. in length by 0·4 mm. in breadth. The body is somewhat
pear-shaped; the colour whitish, reddish, or reddish-black, according
to the contents of the intestine. The legs are fairly short and strong.
During the day they live concealed in the nests, cracks, etc., of the
hen-house, and at night attack the inmates in order to suck their
blood; they rarely remain long on the birds. They have been repeatedly
found on persons, on whose skin they produce an itching eruption.
*Dermanyssus hirundinis*, Hermann, 1804.
Syn.: _Acarus hirundinis_, Herm., 1804.
Of a brownish colour, 1·2 or 1·4 mm. in length; lives in the nests of
swallows and is occasionally found on man.
[Illustration: FIG. 357.--_Dermanyssus gallinæ._ Enlarged. (After
Berlese.)]
[Illustration: FIG. 358.--_Dermanyssus hirundinis._ 40/1. (After
Delafond.)]
[The red hen mite (_Dermanyssus gallinæ_) not only attacks poultry
and man, as stated above, but is found on all birds and many mammals.
The _D. gallinæ_ is the same as _D. avium_. The species found in
swallows’ nests is also said to be the same. This mite can remain for
weeks without any food from its normal host. They only attack man
when entering or cleaning dirty and neglected fowl-houses; they do
not produce a true dermatosis. They chiefly attack the backs of the
hands and forearms of those who constantly attend poultry and give
rise to symptoms similar to the papular eczema of scabies. That they
may remain some time upon the human body we know from the following
cases out of many recorded: Geber observed that the Dermanyssus had
caused a diffused eczema on a woman, which lasted four weeks and then
disappeared. The _tique_ of F. V. Raspail is the bird Dermanyssus;
he records children and adults being attacked not only when handling
pigeons, but even when walking in a garden manured with pigeons’ dung.
The affection soon disappeared when the pigeons were destroyed and the
excreta buried. I have frequently heard of poultrymen being seriously
attacked by this pest.--F. V. T.]
Genus. *Holothyrus.*
*Holothyrus coccinella*, Gervais, 1842.
Measures 5 mm. in size; lives on birds in the Island of Mauritius;
ducks and geese frequently fall victims to its bite; it also attacks
human beings, on whose skin it causes severe burning and swelling, but
no reddening; it may be dangerous to children, especially by settling
in the oral cavity.
Other Gamasides occasionally occur in man, for instance, according
to Moniez, _Leignathus sylviarum_, Canestr. et Fanzago; according to
Neumann _Lælaps stabularis_. The former live normally in the nests of
various species of _Sylvia_, Lælaps on dried vegetable substances,
also in houses.
[Marchoux and Conoy (_Bull. Soc. Path. exot._, 1912, v, No. 10, pp.
796–798) found Leishman granules in _Lælaps echidninus_. It is assumed
that Leishman granules may be found in most Arachnoids, and have no
connection with Spirochæta.--F. V. T.]
Family. *Ixodidæ* (Ticks).
Comparatively large Acarines with a leathery skin; they are flattened
in form, but after sucking blood the abdomen becomes spherical;
the cheliceræ are rod-like and possess a serrated terminal joint,
bent hook-like; the median parts of the pedipalpi (maxillæ) form a
rostrum furnished with barbed hooks (fig. 359); the maxillary palpi
themselves are club-like or rounded; the legs are composed of six
segments with two terminal ungues, often also with “sucking discs”;
the stigmata are at the sides of the body, posterior to the fourth or
third pair of legs. The larvæ are six-legged.
[The true ticks (_Ixodidæ_) are all blood-suckers, and as far as
is known they do not take vegetable food at all. Not only are the
_Ixodidæ_ important as actual parasites, but they are most so on
account of the fact that they are the active agents in carrying various
diseases in animals and apparently in man. It has been conclusively
proved that the bont tick (_Amblyomma hebræum_) is the carrier of the
fatal “heart-water fever” so rife amongst sheep in South Africa, that
the dog tick (_Hæmaphysalis leachi_) is the agent by which the protozoa
that cause malignant jaundice in dogs are distributed, that Texas
fever in cattle is spread by _Rhipicephalus annulatus_, and Coast or
Rhodesian fever by _R. appendiculatus_ and _R. simus_. Their importance
as disease carriers amongst mammals is therefore considerable, and
it may prove to be so for man.[336] They frequently attack man, but
chiefly, according to my observations, in their early stages in Europe;
this is not so, however, abroad. The life-history of a number of ticks
has been clearly demonstrated. Mr. Wheler has shown that in _Ixodes
reduvius_ it is as follows: the female deposits her eggs in masses upon
the ground, gradually reducing in size as the eggs pass out, until she
finally remains a mere shrivelled empty bag and then dies. The eggs are
oval, golden brown in colour and smooth; in length they are 0·59 mm.;
as in all _Ixodidæ_ they are covered with a glutinous secretion, by
means of which they adhere together in masses. These egg masses may be
deposited anywhere on the ground, but amongst rough, coarse herbage
seems to be the favourite place. The egg stage may last as long as
twenty-two weeks, or it may only take eight weeks. In the case of the
bont tick a single female may deposit 15,000 or more eggs. The process
of egg-laying is described as follows by Mr. Wheler: “When egg-laying
is about to take place, the head is further depressed till it rests
close against the under side of the body. In this attitude the end of
the rostrum actually touches the genital orifice, the palpi being at
the same time widely opened out. Behind the head and from beneath the
shield, at what for the purposes of explanation may be described as the
back of the neck, a white, perfectly transparent, delicate gelatinous
membrane is brought down through inflation, either with air or with a
transparent fluid, above the head, which it temporarily conceals. The
end of this membrane terminates in two conical points which appear
to be covered with a glutinous secretion, and at the same time an
ovipositor of a somewhat similar character, but only semi-transparent,
is pushed forward from the genital orifice. This latter is a tube,
within which is the egg. As the ovipositor projects it turns itself
inside out, like the finger of a glove, leaving the egg protruded at
the end and lying between the two finger-like points of the membrane.
The membrane and the ovipositor are then withdrawn each from the other.
The egg adheres to the former, which collapses through the withdrawal
of its contents, dragging the head forward and depositing it on the top
of the head. Neither legs, palpi, nor the organs of the mouth take any
part in oviposition, but after the collapse of the membrane the palpi
are closed and the head is raised, by which action the egg is pushed
forward to the front edge of the shield, forming in time an adherent
mass of eggs, which are deposited in front of the tick.”
[336] This has been proved in Uganda--so-called tick fever in man.
[The egg gives rise to the larval form, the so-called “seed-tick”
stage. At first these minute specks are pallid and soft, but they soon
harden and darken in colour. These larvæ are six-legged and crawl up
grasses and various plants, and there await a passing host, waving
their two front legs in the air and becoming attached by this means.
The larval ticks feed upon the blood of the host, and when replete
fall to the ground, the body becoming inflated in the meanwhile. These
larvæ may remain on the host only two days, or they may remain much
longer. Eventually they moult on the ground and change to the nymph or
pupal stage, which has four pairs of legs. This pupa acts just as the
larva, crawls up plants and waits to regain the host. After a time the
nymphs, having gorged themselves with blood, fall off and remain on the
ground for nearly three months; they then moult and become adult males
and females. In about ten days they assume their normal colour and
regain the host afresh; the female gradually swells until she attains
that large inflated form so characteristic of ticks. The male does not
swell, but nevertheless feeds upon the host and fertilizes the female.
[The act of coitus is strange: the male tick inserts its rostrum and
other mouth organs into the sexual orifice of the female, between the
base of the posterior pair of legs. The males then die and the females
fall to the ground and deposit the ova. There are variations in the
different species, of course, from those given above, which apply
solely to _Ixodes reduvius_. The larvæ and nymphs seem to attack most
animals, but the adults mainly keep to the same host. The periods in
the life-cycle of ticks not only vary in the different species, but in
each species according to climatic conditions; for instance, in the
bont tick (_Amblyomma hebræum_, Koch), Lounsbury has shown that the
development is rapid in summer, slow in winter. The period from the
time that the female drops to the time she commences to lay eggs varied
in specimens observed by him from twelve days in summer to twelve weeks
in winter, and the complete period from the dropping of the female to
the hatching of the eggs, from eleven weeks in summer to thirty-six
weeks through the winter. Other stages vary in a similar manner.
[Ticks may live a long time away from the host provided they are
supplied with a certain amount of moisture. Mr. Wheler kept dog ticks
(_Ixodes plumbeus_) in the larval stage for ten months; the pupæ, male
and female, of _I. reduvius_ for six months.
[I have kept _Ornithodorus moubata_ alive for eighteen months without
food.
[In many species moulting takes place off the host, but in _I. bovis_,
now known as _Rhipicephalus annulatus_, Say (the carrier of Texas
fever), moulting takes place on the host, and in many other species
also.[337] Some species of ticks leave their host on its death (as the
dog tick, _Hæmaphysalis leachi_), but others die with the host (bont
tick, _Amblyomma hebræum_).
[337] Some ticks require only one (_R. decoloratus_), others two (_R.
evertsi_), and some three hosts (_R. appendiculatus_) in order to reach
maturity.
[Two species are of special importance, namely _Ornithodorus moubata_,
Murray, which may infect human beings with the spirillum of African
tick fever, and _Dermatocentor reticulatus_ var. _occidentalis_, which
is said to be the carrier of Rocky Mountain spotted fever.
CLASSIFICATION OF _Ixodidæ_.
[The ticks, or _Ixodidæ_, are divided into two groups, known as (1)
_Argantinæ_, (2) _Ixodinæ_. The _Argantinæ_ are told from the _Ixodinæ_
by the absence of dorsal or ventral shields in both sexes, and also by
the rostrum being placed beneath the cephalothorax, which covers it
over: except in the larval stage, in which it is subterminal, and in
the pupal, when it partly projects. Legs nearly equal in length. The
sexual orifice is situated between the two first pairs of legs. The
males usually smaller than the females.
[The _Ixodinæ_ have the legs unequal, of six segments with two false
segments, making them look as if composed of eight segments. The
rostrum is terminal and never hidden beneath the body. The sexual
orifice is situated between the bases of the first three pairs of
legs. In the males the orifice is obsolete or very rudimentary, sexual
intercourse being effected by the rostrum. The males are smaller
than the females. The shield in the females never covers so much as
one-half of the body even when fasting, also in the larvæ and nymphs;
but in the males, which do not distend, the shield covers the body
entirely, or all but a narrow margin. The _Ixodinæ_ are divided into
two groups: (i) the _Ixodæ_, and (ii) the _Rhipicephalæ_. The former
have a long proboscis reaching nearly to the end of the palpi or even
a little longer than the palpi. The palpi are longer than broad. The
_Rhipicephalæ_ have short palpi, nearly or quite as broad as long,
more or less conical or subtriangular. They were called _Conipalpi_ by
Canestrini.
SYNOPSIS OF GENERA.
[A. _Argantinæ_: Rostrum concealed in adult, partly
exposed in larvæ and nymphs. No dorsal and
ventral shields.
Body flat with thin edges, finely shagreened and
punctate _Argas_.
Body with numerous small round granules and with
thick sides _Ornithodorus_.
[B. _Ixodinæ_: Rostrum terminal. Body with dorsal
shield over some part of it.
I. Rostrum and palpi longer than broad (_Ixodæ_).
α. A groove around anus in front.
Palpi caniculated in both sexes _Ixodes_.
Palpi claviform, not caniculated in the
male; legs very long _Eschatocephalus_.
Palpi claviform, not caniculated in the
male; _anal groove absent in the female_ _Ceratixodes_.
β. A groove around the anus behind.
No eyes; adanal shields _Aponomma_.
Eyes present.
Males with no adanal shields _Amblyomma_.
Males with adanal shields _Hyalomma_.
II. Labium and palpi short; palpi not, or very
little, longer than broad (_Rhipicephalæ_).
α. No eyes.
Rostrum rectangular; no ventral shields in
the male _Hæmaphysalis_.
β. Eyes present.
No adanal shields, but usually with greatly
developed coxæ on fourth pair of legs.
Capitulum quadrangular _Dermacentor_.
Capitulum hexagonal _Rhipicentor_.
Adanal shields in male. Stigmata comma-
shaped _Rhipicephalus_.
Stigmata oval or round; legs normal _Boophilus_.
Segments of legs expanded _Margaropus_.
[The genus Ceratixodes of Neumann, 1902, occurs on birds.
[The genus Eschatocephalus of Frauenfeld, 1853, of which seven
species are known, is mostly parasitic on bats, and is found
in holes, caves, and church towers.
[The genus Aponomma of Neumann, 1899, is exotic, and almost
entirely confined to snakes and saurians.
[The following are synonyms of the different genera:--
_Argas_, Latreille, 1796 (_Rhynchoprion_, Hermann, 1804).
_Ixodes_, Latreille, 1795 (_Acarus_, Linnæus, 1758;
_Cynorhæstes_, Hermann, 1804; _Crotonus_, Dumeril, 1822).
_Ceratixodes_, Neumann, 1902 (_Ixodes_, Cambridge, 1879;
_Hyalomma_, Cambridge, 1879).
_Eschatocephalus_, Frauenfeld, 1853 (_Sarconyssus_, Kolenati,
1857).
_Amblyomma_, Koch, 1844 (_Ixodes_, Latreille, 1795).
_Hæmaphysalis_, Koch, 1844 (_Rhipistoma_, Koch, 1844;
_Gonixodes_, Dugès, 1888; _Opitodon_, Canestrini, 1897).
_Rhipicephalus_, Koch, 1844 (_Acarus_, Linnæus, 1758;
_Ixodes_, Latreille, 1795; _Phanloixodes_, Berlese, 1889;
_Boophilus_, Curtice, 1890).
_Dermacentor_, Koch, 1844 (_Ixodes_, Latreille, 1795;
_Pseudixodes_, Haller, 1882).--F. V. T.]
Genus. *Ixodes*, Latreille.
*Ixodes reduvius*, L., 1758.[338]
[338] _Ixodes reduvius_ and _I. ricinus_ are synonymous. [The above
should read _Ixodes ricinus_, Latreille, 1806.--F. V. T.]
Syn.: _Acarus reduvius_ and _ricinus_, L.; _Ixodes ricinus_,
Latreille, 1806.
The males are oval; their length 1·2 to 2 mm.; they are brownish-red
or black in colour; the females are yellowish-red, 4 mm. long; when
gorged they are lead-coloured, and may attain 12 mm. in length by 6 to
7 mm. in breadth.
The dog tick (fig. 360) lives in thickets on leaves, etc., and
attacks sheep and oxen, and more rarely dogs, horses, and human
beings, into the skin of which the female bores with the rostrum in
order to suck blood; the bite is not dangerous, and sometimes is
not even felt. Inflammation, however, is set up if the creatures
are forcibly removed from the wound, as the rostrum as a rule
is torn off in the process. If left alone or smeared over with
some grease--vaseline, oil, butter, etc.--the creatures drop off
spontaneously. Sometimes the entire tick bores itself into the skin;
they also appear to be permanent inmates of kennels.
[Illustration: FIG. 359.--A., the rostrum of _Ixodes ricinus_ (male);
B., the terminal joint of the maxillary palpi of the female. Enlarged.
(After Pagenstecher.)]
[Illustration: FIG. 360.--Female of _Ixodes ricinus_, gorged full,
dorsal and ventral surfaces. 2/1. (After Pagenstecher.)]
[The species _I. reduvius_ is the same as _I. ricinus_, Latreille.
The male is 2·35 to 2·80 mm. long; the body is dark brown, almost
black, with a pale, almost white, margin; there are also traces of
reddish mottling. Coxæ of the first pair of legs with a short spine.
Rostrum much shorter than that of the female; shield oval; anal shield
small, about one-third the length of ventral shield. The adult female
varies from 2·80 to 3·5 mm. when not distended, but when gorged may
reach 10 mm. long. The shield and legs are dark blackish-brown, body
deep orange-red with four dark longitudinal lines, paler beneath and
light grey in front. When distending it is pale red to grey or white;
when fully gorged olive-green, or dark red to black, with irregular
yellow streaks on the back and sides just before egg-laying. Sexual
orifice opposite fourth pair of legs. The nymph varies from 1·60 to
1·70 mm. long when fasting; the body is olive-white, opaque, with four
distinct brown posterior markings and similar anterior ones, leaving
a pale centre to the shield. When fully gorged it is 3 mm. long. As
the nymph distends, it changes from opaque white to blue-black, and
finally black. The little larva is 0·80 to 1·50 mm. long, transparent
with olive-green intestinal markings; as it becomes inflated it
changes to blue-black, and then black. There are no eyes. It is widely
distributed, and chiefly attacks sheep; sometimes it occurs on dogs and
also attacks man. Mégnin records it from horses in the nymph stage.
Amongst its other numerous hosts are goats, cattle, deer, hedgehogs,
moles, bats, birds, and lizards. It is usually known as the grass tick
and bottle-nosed tick. This species occurs in Europe, Asia, North
Africa, and North America.
[_Synonyms._--Considerable confusion exists over the name of this and
other common ticks, owing to the same species having been described
under a great many names. Observers have taken the same species on
different animals and in various stages to be distinct, and have
described them accordingly.
[The name _Ixodes reduvius_, Leach, does not stand, as Leach was
describing quite a different parasite. The name _I. ricinus_,
Latreille, 1806, is now substituted by Neumann and Wheler.
[The synonyms given by Wheler are as follow: _Reduvius_, Charleton,
1668; _Ricinus caninus_, Ray, 1710; _Acarus ricinoides_, de Geer, 1778;
_Acarus ricinus_, Linnæus, 1788; _Cynorhæstes reduvius_, Hermann, 1804;
_Cynorhæstes ricinus_, Hermann, 1804; _Ixodes megathyreus_, Leach,
1815; _Ixodes bipunctatus_, Risso, 1826; _Cynorhæstes hermanni_, Risso,
1826; _Crotonus ricinus_, Dumeril, 1829; _Ixodes trabeatus_, Audouin,
1832; _Ixodes plumbeus_, Dugés, 1834; _Ixodes reduvius_, Hahn, 1834;
_Ixodes fuscus_, Koch, 1835 (?); _Ixodes lacertæ_, Koch, 1835 (?);
_Ixodes pustularum_, Lucas, 1866; _Ixodes fodiens_, Murray, 1877;
_Ixodes rufus_, _Ixodes sulcatus_, and _Ixodes sciuri_, Koch.--F. V. T.]
*Ixodes holocyclus*, Neumann, 1899.
[Under the name _I. holocyclus_, Cleland (_Journ. Trop. Med. and Hyg._,
1913, xvi, No. 3, pp. 43–45) says that: “This tick is common in man
where there is dense scrub and tropical jungle along the east coast of
Australia at certain times of the year. It may cause severe symptoms in
children resulting in death.” He records a child being attacked in 1884
which died, and another case from which 200 ticks were removed, the
symptoms being weak heart, collapse, syncope, but the patient recovered
under treatment; again, in the same journal (pp. 188, 189), the case
of a 4-1/2-year-old girl who was bitten showed widespread muscular
paralysis, and other cases resembling conium poison.
[Taylor (_Rep. Ent. Aust. Inst. Trop. Med._, 1911, p. 21, 1913) refers
to this species as the scrub tick of New South Wales. The partially
fed female has a dark reddish-yellow scutum and is almost as broad as
long, punctations very numerous, equal and confluent in places, long
white hairs on the lower half of each coxa. He records it as attacking
man commonly, mentioning Kamerunga, Cairns district, Queensland, and
Sydney, N.S.W., as localities.--F. V. T.]
*Ixodes hexagonus*, Leach, 1815.
Syn.: _Ixodes sexpunctatus_, Koch, 1897; _I. vulpis_, Pagenstecher,
1861.
Lives in the same manner as the foregoing; especially attacks hounds,
but also other mammals and even birds. The difference consists in the
shape of the legs, the shorter rostrum, and the larger size of the
male. It also occasionally attacks man, but is usually confused with
the previously mentioned species.
[The synonyms of this species are as follow:[339] _Ixodes autumnalis_,
Leach, 1815; _I. erinacei_, Audouin, 1832; _I. reduvius_, Audouin,
1832; _I. crenulatus_, Koch; _I. erinaceus_, Murray, 1877; _I.
ricinus_, Mégnin, 1880.[339] Two other synonyms are given above by
Braun.
[339] Neumann, G. L., “Rev. de la fam. des Ixodides,” _III Mém. Soc.
Zool. France_, 1899, xii, p. 129.
[The female when fully replete is 11 mm. long, when fasting 3·86 mm.;
the shield is heart-shaped and punctate, body finely hairy; palpi short
and broad; labium shorter, and tarsi of all the legs more truncate than
in _I. ricinus_. The colour of the distended body is drab and somewhat
waxy; rostrum, shield and legs light testaceous. The male varies from
3·5 to 4·0 mm. long, and is reddish-brown in colour with lighter legs;
the shield is punctate and leaves a narrow margin around the body;
the body is elliptical, almost as large in front as behind. There is
a spine on the coxæ of the first pair of legs, which is shorter than
in the male _I. ricinus_ and longer than in the female. The genital
orifice is opposite the interval between the second and third pair of
legs. The fasting _nymph_ is 1·76 mm. long, light bluish-grey, margined
and transparent, with four large posterior intestinal marks joined
together behind the shield and smaller ones extending to the front and
sides. When fully distended it is uniformly brownish-white; shield,
legs and rostrum pale testaceous. The _larva_ varies from 0·88 mm.
when fasting to 1·76 mm. when gorged. Its body is light, but gradually
becomes darker, with similar intestinal marks to _ricinus_.
[This tick is very common, especially on ferrets, stoats and hedgehogs.
It is also found on sheep, cattle, etc. The males do not generally
occur in company with the females on the host. Pairing probably takes
place on the ground.--F. V. T.]
Genus. *Amblyomma*, Koch.
*Amblyomma cayennense*, Koch, 1844.
Syn.: _Amblyomma mixtum_, Koch, 1844; _Ixodes herreræ_, Dugés, 1887;
_Amblyomma sculptum_, Berlese, 1888.
Characterized by the possession of eyes. The male measures 3·8 mm.
in length by 3 mm. in breadth; the female 4 mm. in length by 3 mm.
in breadth, but when full of blood may become 13 mm. in length and
11 mm. in breadth. They are common in the whole of Central America
(Carrapatas), and attack mammals, amphibious animals and man.[340]
[340] Neumann, G. L., _loc. cit._, p. 205.
[This species was described by Fabricius. It occurs in Cayenne, Guiana,
in Southern Texas, Florida, California, Mexico, Guatemala, Honduras,
Nicaragua, Costa Rica, Panama, Bermuda, Cuba, Jamaica, Trinidad,
Colombia, Venezuela, French Guiana, Brazil, Paraguay and the Argentine.
It is called the silver tick. It frequently attacks man. Schwarz and
Bishopp (_Bull._ 105, U.S. Dept. Agric., p. 158) heard of one man whose
legs were well covered with suppurating sores and who was ill from
the attack of these ticks and the wounds produced by scratching, and
records other cases of their swarming on man. Newstead (_Ann. Trop.
Med. and Par._, 1909, iii, No. 4, p. 442) records it as the worst pest
to man in Jamaica.--F. V. T.J
*Amblyomma americana*, Linnæus.
The so-called long star tick, from the silvery spot on the apex of the
scutum of the female. It will attack any mammal and even birds and
also man. It occurs in North America, and also in Brazil, Guiana and
Guatemala. Its punctures frequently end in suppuration. In the Eastern
and Southern States man is more frequently attacked by this species
than any other. Moss-gatherers in Louisiana are badly attacked by
it.[341] It also attacks the milkers in dairies. Attempts to transmit
Texas fever failed with this species.
[341] Morgan, “Ticks and Texas Fever,” _Louisiana State Bull._ 55,
pp. 134, 135, pl. 59.
*Amblyomma maculatum*, Koch.
The so-called Gulf Coast tick, of the Gulf Coast, occurs on birds,
mammals and man, especially cattle, and attacks the ears.
Genus. *Hyalomma*, Koch.
*Hyalomma ægyptium*, L., 1758.
Syn.: _Acarus ægyptius_, L., 1758; _Ixodes camelinus_, Fischer, 1823.
A species frequently found in Africa, particularly in Egypt and
Algeria, and which also occurs in France and Italy, as well as in Asia.
Male 8 mm. in length, 4·5 mm. in breadth. Female up to 24 mm. in length
and 15 mm. in breadth. It infests large and small animals as well as
human beings.[342]
[342] Neumann, G. L., _loc. cit._, p. 285; Ronsisvalle, “Sui fenomeni
morb. prodotti nel uomo da un Ixodide denominato _Hyal. æg._,” _Boll.
Acc. Gioenia sci. nat._, 1891, xvii.
[This is one of the largest ticks, nearly reaching the size of the bont
tick. It is known in Africa as the bont leg-tick; all farm stock is
attacked, but sheep and goats suffer most. Only one generation appears
to occur each year. The male is almost black with a pale marginal
stripe; the replete female brown with irregular light blue stripes. It
is abundant in parts of South Africa.]
Genus. *Hæmaphysalis*, Koch.
*Hæmaphysalis punctata*, Canestrini and Fanzago, 1877–1878.
Syn.: _Hæmaphysalis sulcata_, Canestrini and Fanzago, 1877–1878;
_Rhicocephalus expositicius_, Koch, 1877; _Hæmaphysalis peregrinus_,
Cambridge, 1889; _Herpetobia sulcata_, Canestrini, 1890.
[This species does not appear to be common. It occurs on sheep, goats,
horses and cattle. I have seen a female taken from man in Britain. The
female when fasting is 3·44 mm. long, when gorged 12 mm. long. Colour,
reddish-brown, leaden-grey when gorged; dorsal shield deeply indented
in front; rostrum, shield and legs brownish; body finely punctate, both
above and below; sexual opening opposite the coxæ of the second pair
of legs in both sexes. Palpi a little longer than the labium; first
segment short and narrow, second and third widened on the dorsal face.
Coxæ with a short, broad blunt spine; tarsi short, terminated with a
spur on the first pair. The male is 3·10 mm. long. Body rather narrow,
yellowish to reddish-brown; dorsal shield nearly covers the whole body;
numerous punctures over the whole surface. Eleven indentations on the
posterior margin of the body; peritremes lighter in colour, large and
comma-shaped. The three anterior pairs of legs with a short spine on
the haunches, the fourth with a very long one directed backwards. The
nymph varies from 2·5 to 3·0 mm., is oval, and light yellow to dark
red in colour. Dorsal shield rounded with few punctations. No spur on
tarsi, and sexual orifice nearly obsolete. Larva short and oval. Length
1·20 mm.--F. V. T.]
Genus. *Dermacentor*, Koch.
*Dermacentor reticulatus*, Fabricius, 1794.
Syn.: _Acarus reticulatus_, Fabr., 1794; _Ixodes reticulatus_,
Latreille, 1806; _I. marmoratus_, Risso, 1826.
This tick is provided with eyes, but it is distinguished from Ixodes
and analogous genera by the lack of the abdominal plastron in the male,
which measures 5 to 6 mm. in length by 3·5 mm. in breadth. The female
may attain 16 mm. in length and 10 mm. in breadth. It is found in the
South of Europe, in Asia, and in America; it attacks chiefly oxen,
sheep and goats, and occasionally man.[343]
[343] Neumann, G. L., “Rev. de la fam. des Ixodides,” _Mém. Soc. Zool.
France_, 1897, x, p. 360.
[This tick sometimes causes much annoyance to human beings. It was once
most troublesome at Revelstoke. Specimens have recently been found on
fowls, turkeys and pheasants in Kent.
[Other synonyms are as follows: _Cynorhæstes pictus_, Hermann, 1804;
_Crotonus variegatus_, Dumeril, 1829; _I. pictus_, Gervais, 1844;
_Dermacentor albicollis_, Koch, 1844–1847; _D. pardalinus_, Koch,
1844–1847; _D. ferrugineus_, Koch, 1844–1847; _Ixodes holsatus_,
Kolenati, 1857; _Pseudixodes-holsatus_, Haller, 1882; _Hæmaphysalis
marmorata_, Berlese, 1887.
[The _female_ when fasting is 3·86 mm. long by 2 mm. wide. The body is
depressed, larger behind and reddish-brown in colour. The shield is
very large and extends to the level of the third pair of legs, with
a few large and many small punctations, milky white, variegated with
reddish-brown. Sexual orifice opposite the coxæ of the second pair
of legs. Coxæ of the front legs are deeply bifid, the others with a
moderate spine. When gorged light brown, and may reach 16 mm. When
depositing eggs the female is mottled with dark brown above and below.
The _male_ is like the female. The shield is reddish-brown, variegated
with a milky white pattern. Coxæ of the fourth pair of legs three times
the size of the third. There is a sharp backwardly pointing spine on
the second palpal segment, also seen (but smaller) in the female.
Length 4·20 mm.
[According to Mr. Wheler this is a very variable species both in size
and colour. It occurs in England on sheep, but not commonly. It has
probably been introduced into Britain. Besides the animals mentioned
above it is also found on deer.--F. V. T.]
*Dermacentor venustus*, Banks.
[The Rocky Mountain tick fever tick. This species has been wrongly
called _Dermatocentor reticulatus_ var. _occidentalis_. The correct
name of the carrier of Rocky Mountain tick fever is _Dermacentor
venustus_, Banks (Hooker, Bishopp and Wood, _Bull._ 106, U.S. Dept.
Agric., Div. Ent., p. 165).
[The female is from 13·8 by 10 by 6·4 mm. to 16·5 by 11·4 by 6·9 mm.
when gorged; the male from 2·1 by 1·5 mm. to 6 by 3·7 by 1·4 mm. The
male reddish-brown; scutum with an extensive pattern of white lines,
usually but little white on the mid-posterior region, legs slightly
lighter than scutum, joints tipped with white. Female with scutum
mostly covered with white, abdomen reddish-brown, legs as in male. The
nymph when unengorged reddish-brown, when gorged dark bluish-grey; the
larva is yellowish-brown when unengorged, slate blue when engorged. The
ova light brown, shiny and smooth.
[The chief wild hosts are the brown bear, coyote, woodchuck, rabbit,
wild cat, badger and mountain goat for the larvæ; practically all
small mammals act as hosts for larvæ and nymphæ, whilst the adults are
seldom found on other than large domestic animals; horses and cattle
are preferred. It occurs in British Columbia, southward to Northern
New Mexico, and from the foothills of the Rocky Mountains in Colorado
to the base of the Cascade Range in Oregon and California; abundant in
Western Montana, Idaho, Eastern Washington, Oregon, North Utah, West
Wyoming and North-west Colorado.
[Of great importance in the Bitter Root Valley of Montana, where a
number of cases of fever occur each year, with a mortality of about 70
per cent. In British Columbia this tick causes tick paralysis in man
and sheep. Only the adults seem to attack man and animals there (Hadwen
and Nuttall, _Parasitology_, 1913, vi, No. 3, pp. 288–297 and 298–301)
according to the _Canadian Medical Association Journal_, December,
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