MORPHOLOGICAL

CHARACTERISTICS OF
METAZOAN PARASITES IN TISSUE
SECTION

Metazoan parasites can be found in all organ systems of
animals. During necropsy or biopsy procedures these
parasites can be identified grossly since they usually 1)
look different than the tissue and 2) move. Their presence
in histological section, however, is less common. Many
organs are hollow, e.g., intestines, and many parasites do
not have the ability to attach to those tissues. Thus,
parasites "drop" out of tissue during histological process-
ing. It is important to correlate what is seen grossly with
what is seen histologically. This has been done in many of
the species presented in this Atlas.
Many sections of parasites do not allow identification
since salient features, needed in the identification process,
are absent. If possible prepare more sections. Sometimes,
serial sections are called for. Cross sections are usually
the "cut of choice" for study. It is important to study the
characteristics presented in th is Atlas and extrapolate
features in your section to these.
There are six groups of metazoan parasites: nema-
todes, acanthocephalans, trematodes, cestodes,
arthropods, and pentastomes. The table below is a
"working" key that is to be used in identification of meta-
zoan parasites in tissue section . The key consists of
characteristics of adult parasites; however, some charac-
teristics can also be used for the identification of larval
forms. All groups of parasites are not included (e.g.,
leeches) since few of those omitted are seen by parasitolo-
gists.
The body cavity may be present or absent in parasites.
If it is lacking the parasite is said to be acoelomate. If it is
present, it may be a coelom or a pseudocoelom (really
technical terms that can't be differentiated in histological
section). To identify a body cavity look for muscles, those
structures that stain eosinophilic with hematoxylin and
eosin. If muscles are pesent they usually support a body
wall. Thus, if there is a body wall, there should be an
associated cavity as well. Study the manner in which the
muscles are associated with the outside covering of the
body wall. In arth ropods and pentastomids, the muscles
are attached to the covering, i.e., cuticle. In the nematodes
and acanthocephalans, the muscles are associated with a
hypodermal layer that, in combination with the cuticle, form
the wall. Acanthocephalans have relatively more hypoder-
mis than muscles; nematodes usually have more muscle
than hypodermis. In trematodes and cestodes, the muscles
are found dispersed through the spongy parenchyma.
Digestive tracts are present in all groups except the
cestodes and acanthocephalans. When present, the tracts
appear to be tubes com posed of cells.
Some groups of parasites have unique features that
only that group possesses. Calcareous corpuscles are
found only in cestodes; sclerotized openings in the cuticle
and adidophilic digestive glands are found only in
pentastomes; lemnisci are found only in the acanthoceph-
alans, and tracheal ring respiratory systems are found only
in the arthropods. Each group of parasites has a "set" of
characteristics, that when taken in unison, allows identifi-
cation.
When describing or studying a parasite start your
discussion from the periphery and work your way to the
interior. Thus, describe the body covering and body wall
initially. Does it have a cuticle, how th ick is the cuticle, is
it supported by hypodermis, muscles, other structures?
Determine the presence or absence of a body cavity.
Determine what type of muscle, if any, is present (i.e., is
muscle striated or nonstriated). What type, if any, of
intestine is present? And finally, does the organism have
both sexes or only one sex? What is the sex?
The following sections discuss all features found in the
major groups of parasites. Study features as groups or
sets of features. It sounds difficult-but it really isn't!

QUICK KEY
1. No body cavity; solid with "holes" for structures; body usually flattened ....................... ............ ...... ............................. .... 5
Body cavity; coelom or pseudocoelom; body usually cylindrical ........................................................................................ 2
2. Anterior end armed with proboscis; no digestive tract .................................................. .................... ACANTHOCEPHALA
No proboscic; digestive tract present .............................................. ............................ ....................................... ...... ....... ..... 3
3. Cuticle with sclerotized openings; cephalic hooks present; head and acitophilic glands present ........ PENTASTOMIDA
None of the above; ............................................................................................ .................................................................... 4
4. Muscles striated ; jointed appendages; exoskeleton of chitin ........ ............................................................. ARTHROPODA
None of the above ............................................................................................................................................... NEMATODA
5. No digestive tract; presence of calcareous corpuscles; segmented body ...... ........ .. .. .. ........ ... .......................... CESTODA
Digestive tract; no segmentation or calcareous corpuscles .......................................................................... TREMATODA
MORPHOLOGICAL
CHARACTERISTICS OF
NEMATODES IN TISSUE SECTION
CUTICLE
The external cuticle of nematodes ranges in its thick-
ness. Many times the cuticle is very thick (Figs. 1.1, 4.1,
4.2), but often it is so thin it is not discernable (Figs. 1.2,
4.4). In some nematodes the cuticle is ornamented, con-
taining bumps, ridges, winglike structures, etc. It is
important to identify these features in histologic section
since their presence may lead to a more specific diagno-
sis.
The anterior end of the nematode may bear cuticular
ornamen tations. A bulb (Fig. 1.3) or co llar (Fig. 1.4) may
surround the mouth. These structures, of course, are
much more easily seen in gross specimens but may be
identified with fortuitous sectioning. The cuticle on some
nematodes has blister-like structures; these are called
bosses (Fig. 1.5). Serpentine cuticular ridges are present
on the anterior end in several groups of nematodes . The
ridges usually do not extend far posteriad and are called
cordons (Fig. 1.6). Alae are winglike extensions of the
cuticle. Alae are common on the anterior and posterior
ends where they are referred to as cephalic and caudal,
respectively. Cephalic alae are usually located laterally
(Figs. 1.5, 2.1) but in some nematodes they have "mi-
grated" to a sublateral position (Fig. 2.2). Caudal alae are
usually present on the posterior end of male worms and
are associated with the genital opening. Caudal alae
commonly bear papillae, bumps, ridges and/or spine-like
ornamentations (Fig. 9.6).
Regularly spaced, external ridges are present in
numerous genera of nematodes. These ridges are most
easily seen in cross or oblique sections (Figs. 2.4-2.6).
Sometimes generic identification can made just by
counting the ridges. Ridges can be equal in size and
shape (Figs. 2.4- 2.6), but in some instances the size,
shape, and distance from one to another varies greatly
(Fig. 3.1 ). Sometimes external ridges are found as rings
or annulations around the body of the nematode. In these
instances the ridges are not visible in cross section but
can only be identified in longitudmal section (Fig. 3.2).
Internally the cuticle may expand laterally to form
ridges (Figs. 3.3, 3.4). These are appropriately called
latera l internal ridges, are associated with the lateral
chords, and are common in some strongyles and most
filarial nematodes. The mouth or buccal cavity has a
cuticular li ning and in some nematodes, e.g ., the
strongyles, this cuticle is often quite thick (Fig. 3.5). Other
nematode groups, however, can also have strongly
cuticularized cavities (Fig . 3.6).
HYPODERMIS
Internal to the cuticle is a layer of hypodermis. In
phasmid nematodes the hypodermis divides the somatic
musculature into sections as it projects into the
pseudocoelom (i.e., the body cavity) of the worm at the
lateral areas. These extensions of the hypodermis are
called lateral chords (Figs. 4.1-4.4). In addition, nerve
chords are found associated with the hypodermis. Since
the nerve chords protrude in the ven tral and dorsal areas,
they, along with the lateral chords, divide the musculature
into quadrants. Th e ventral nerve cho rd is larger than the
dorsal (Figs. 33.3, 33.5).
In phasmid nematodes nuclei are sparse in the
hypodermis except in the lateral chords. The number and
arrangement of nuclei in the chords is often of diagnostic
importance. Th e lateral chords may protrude dramatically
into the pseudocoelom of the worm (Figs. 4.1 , 4.2) or
may be "low" in nature (Fig. 4.3). The chords may take
any number of forms, e.g., stalked (Fig. 4.1) or shaped
liked the wings of a butterfly (Fig. 14.5). The chords are
divided into sublaterals. In the an terior end of the
nematode an excretory cell may sometimes be associ-
ated with one or both of th e chords (Fig. 6.6).
Some nematodes, i.e., the metastrongyles, have
accessory hypodermal chords th at further divide the
quadrant of the worm (Fig. 4.5) . This feature is often
quite useful in their diagnosis.
The aphasmids do not have chords as described
above; instead they have hypodermal bacillary bands
that possess many nuclei (Figs. 4.6, 31.4, 31.5, 32 .3).
MUSCULATURE
Muscle cells are composed of a cytoplasmic and a
contractile portion. In hematoxylin and eosin (H&E)
stained sections , the cytoplasmic portion is usually clear,
grey, or light pink; the contractile portion is bright red.
Th e contractile portion is composed of numerous con-
tractile fibers that may be arranged perpendicular to the
cuticle (when th e worm is studied in cross section) or in a
parallel fashion.
Muscles are described as being coelomyarian (Figs.
5.2, 5.4, 5.6) or platymyarian (Figs. 5.1 , 5.3, 5.5).
Coelomyarian muscles project into the pseudocoelom of
the nematode in a cylinder-li ke fashion. Muscle cells
may project into the pseudocoelom so greatly that the
body cavity is almost entirely obscured (Fig. 5.4). At
other times, they are "low" in nature (Fig. 5.2). The
coelomyarian arrangement allows for many muscle cells
to be present in one section of worm . Thus , this muscu-
lature is also called polymyarian. Contractile fibers run
up the side of the cells and thus seem to be parallel to
the cuticle (Fig. 5).
Platymyarian muscles , on the other hand , do not often
extend into the pseudocoelom but lie flat on the hypoder-
mis (Figs. 5.1 , 5.3, 5.5). Platymyarian cells are usually
large and few, i.e. , meromyarian, in arrangement. The
contractile fibers of the cells are arranged perpendicular
to the hypodermis and cuticle when studied in cross
sections of worms (Figs. 5. 1- 5.6) .
In addition to somatic or body musculature, nematodes
have muscles associated with such structures as the
esophagus (Fig. 6.2), vagina (Fig. 29.3), and spicules (Fig.
31 .2) . These muscle ce lls are called ciromyarian since
they "circle" the structure.
DIGESTIVE TRACT
The digestive tract is composed of the mouth, buccal
cavity, esophagus, intestine, and anus. The shape of the
esophagus can be useful in the identification of nema-
todes. Some nematodes have an esophagus that is
uniform in diameter (Fig. 6.2) ; others contain narrowings,
bulbs, etc. (Figs. 6.1, 11.3, 13.5) . The lumen of the
esophagus, especially the anterior portion, is lined with
cuticle (as was the buccal cavity) and is tri-radiate in
shape (Figs. 6.3, 6.4 , 6.6) . In most nematodes the anterior
portion of the esophagus is muscu lar and often contains
ducts for esophageal glands (Fig. 6.3).
The esophagus gives rise to the intestine (Fig. 6.5 ).
The shape, si ze, and co mposition of the intestine varies
depend ing upon the group of nematode. Intestines are
described as being large (Figs. 7. 1, 7.3, 7.5) , medium , or
small (Figs. 7.2 , 7.4), relative to the diameter of the
nematode. Intestines can be composed of cuboidal to
columnar cells (Fi g. 7.5) or sometimes multinucleated
cells that are few in number (Figs. 7 .1, 7 .3, 7.6). Ami-
crovillar "brush" border may be present (Figs. 25.3, 25.4).
Yellow to black pigment, perhaps of hemosiderin origin,
may be contained within intestinal ce lls (Figs. 7 .2, 7.6,
12.2, 17.4) .
REPRODUCTIVE TRACTS
With the exception of some members of the Rhaditoids,
both sexes are present when nematodes are parasitic.
Females are usually larger than males. In the phasmid
nematodes , the female has at least two reproductive tracts
(Fig. 8.1 ). In cont rast, fema le aphasmids have one genital
tract (Figs . 31.3, 32.3). Ovaries give rise to oviducts
which empty into uteri. T he uteri contain developing eggs
or embryos (Figs. 8.3- 8.6, 9.2 ), which are liberated into
the environment via a vagina. Eggs vary among groups
and even within groups. It is importan t to remember the
type of egg in each nematode group (and exceptions) so
that these characteri stics can be used to identify the type
of parasite present. Eggs can contain a zygote or larva
(i.e., be embryonated) (Figs. 8.5, 8.6, 9.2), be thin - (Fig.
17.6) or thick-shelled (Figs. 8.3, 8.5, 8.6) , or the egg shell
can be so thin t hat it is not discernable, as seen with
microfi lariae (Fig. 10.3) and metastrongyle larvae (Figs.
8.4, 10. 4).
Adu lt males have only one gen ital tract (Figs. 8.2 ,
9.3). The testis produces sperm which, in turn , is found in
the vas deferens. Nematode sperm , when stained with
hematoxylin and eosin, is described as being smal l, oval ,
eosinophilic, and containing a dark, basophilic nucl eus
(Figs. 9.3, 13.1, 20 .2). Sperm can be found within both
the male and female since the distal part of the uteri in
the female , referred to as the seminal receptacle, stores
sperm from previous copulations (Fig. 9.4). Males may
possess copulatory bursas, caudal alae, and spicules as
reproductive structures (Figs . 9.5, 9.6).
Mature rep roductive cells, i.e., ova and sperm , are
produced differently in the phasmids and aphasmids. In
the phasmids the germ cells cleavage is referred to as
telegonic. In this type there is a single larg e cap cell at
the proximal end of the ovary or testis. This cell under-
goes cleavage and results in more mature germ cells
closer to the vagina or cloaca, respectively. Therefore ,
when th e reproductive t ract is histologically sectioned in
cross section the germ cells are all at t he same stage of
developmen t (Fig. 10.1 ). In the aphasmids, however, the
cleavage is much different. T his type of division is
referred to as hologonic. The germ cell is found along th e
ent ire "body" of t he ovary or testis and bu ds off germ cell s
throughout its length. Therefore, on histological section,
all stages of germ cell development can be seen in one
section (Fig. 10.2).
Alt hough most nematodes produce eggs, som e do
produce larvae. Common larvae found in histologic
section include Strongyloides larvae, microfilariae,
metastrongyle larvae, and Trichinella larvae. Microfi-
lariae, on light microscopy, appear to lack musculature
and resemble a cuticular-limited bag of nuclei (Fig. 10.3).
Intestines, lateral chords, and other features can not be
discerned. T he metastrongyle larvae have more structu re
to them and have a primitive enteron and are usually
much larger than the microfilariae (Fig. 10.4). Trichinella
larvae possess well-defined musculature , enterons, and
can be differentiated from other phasmid larvae by th e
presence of a stichosome (Fig. 10.5). This structure will
be discussed in more deta il in the aphasmid section.
There are numerous other types of nematode larvae
that may be foun d in histologic section . These differ f rom
adults in the fact that they are smaller and lack ova,
sperm , or zygotes (Fig. 10.6).

1.3
1.5
=igures 1.1-1.6. Cuticular manifestations in
1ematodes. 1.1. Larva of Contracaecum sp. in the
;tomach wall of a bullfrog. Note the thick cuticle.
<75. 1.2. Metastrongyle in lung of pig. Note the
hin, almost indiscernible, cuticle. X 150. 1.3.
.ongitudinal section of the anterior end of
'3nathostoma procyonis in the stomach of a
accoon. Note the cephalic bulb (b) that bears
;pines. The body is spined as well. X 75. 1.4.
.ongitudinal section of the anterior end of
1.2
1.4
1.6
Physaloptera procyonis in the stomach of a
raccoon. Note that the anterior end is surrounded
by a cuticular collar (c). X 750. 1.5. Cross
sections through the anterior end of
Gonglyonema sp. in the esophageal epithelium of
a deer. Note the cuticular bosses (b) and lateral
alae (a). X 300. 1.6. Cross section th rough the
anterior end of Echiura sp. in the stomach of a
swan. Note the cuticular cordons (c). X 120 .

2.5
Figures 2.1-2.6. Cuticular manifestations in
nematodes. 2.1. Cross section of Enterobius sp. in
the intestinal tissue of a chimpanzee. Note the
lateral alae (a). X 280. 2.2. Cross section
through the anterior end of Pterygodermatites
nycticebi in the intestine of a marmoset. Note that
the lateral alae (a) are sublateral in position. X
240. 2.3. Dirofilaria tmmitis in the heart of a dog.
Note the smooth external cuticle and the lateral
internal cuticular ridges (r). X 30. 2.4. Dirofilaria
r
2.4
2.6
ursi from the diaphragm of a bear. The external
cuticle has evenly spaced longitudinal ridges (r). X
70. 2.5. Cooperioides hepaticae in the bile duct of
an impala. The external cuticle has evenly spaced
longitudinal ridges (r). X 300. 2.6. Nochtia
nochtia in the intestine of a rhesus monkey. Note
how the longitudinal culicular ridges are flared at
the area of the lateral chords (1), contrasted to the
other cuticu lar ridges (r). X 750.
3.1
Figures 3.1-3.6. Cuticular manifestations in
nematodes. 3 .1. Nippostrongylus sp. in the
intestine of a rat. Note the external cuticular ridges
are not equal in size, nor are they evenly spaced.
X 300. 3.2. Longitudinal section of Onchocerca sp.
in periocular tissue of a dog. Note the evenly
spaced cuticular annulations (a). X 300. 3.3.
Dirofilaria immitis in the heart of a dog. Note the
lateral internal cuticular ridges (r). X 30.
3.4. Cross section of Ancylostoma caninum in the
3.4
intestine of a dog. Note the lateral internal
cuticular ridge (r). X 300. 3 .5. Longitudinal section
of larva of Strongylus vulgaris in mesenteric artery
of horse. Note the highly cuticularized buccal
cav1ty (c). X 120. 3.6. Longitudinal section of th e
ante nor end of Draschia megastoma from the
stomach of a horse. Note the highly cuticularized
buccal capsule (b). X 150.
4.1
Figures 4.1-4.6. Hypodermis in nematodes . 4.1.
Physaloptera sp . in the stomach of an opossum .
The hypodermis expands laterally to form two large
lateral chords (c). X 40. 4.2. Higher magnification
of the lateral chord in Physaloptera sp. This type
of lateral chord is described as being "stalked." X
100. 4.3. Cordophilus sagittus in the lung of a
kudu. Note the low lateral chord (c) and the lateral
internal cuticu lar ridge (r). X 300. 4.4. Spirurid
larva in the mesentery of a boa constrictor. Note
4.2
4.4
the huge lateral chords (c) filling the entire
pseudocoelom . X 300. 4.5. Metastrongyle in lung
of doll porpoise. Note the lateral chords (lc) and
the accessory hypodermal chords (c). X 300.
4.6. Trichuris vulpis in the intestine of a dog. This
worm lacks lateral hypodermal chords but has a
large bacillary band (b) throughout the length of its
anterior end. X 400.
 .· ·). .'(.-j
(

•
.

5.2

Figures 5.1-5.6. Musculature of nematodes. 5.1 .
Larva of Stephanurus dentatus in liver of pig. The
musculature is platymyarian. Note how the muscle
cells (m) are flat against the cuticle. X 120. 5.2.
Elaephora schneideri in the carotid artery of an elk.
The musculature is cylindrical in shaped and thus
referred to as coelomyarian. X 120. 5.3. Larva of
Stronqylus vulgaris in mesenteric artery of a horse.
The musculature (m) is platymyarian. X 300. 5.4.
Larva of Habronema sp. in the skin of a horse.
5.6
Note the coelomyarian musculature fills the
pseudocoelom. X 480. 5.5. Larva of
Oesophagostomum sp. in the intestine of a
chimpanzee. Note the platymyarian muscles (m).
X 120. 5.6. Dirofilaria ursi from the diaphragm of a
bear. Note the coelomyarian muscles (m). X 300.

Figures 6.1-6.6. Esophagi of nematodes. 6.1.
Longitudinal section of anterior end of Enterobius
sp. in the intestinal tissue of a chimpanzee. Note
the corpus (c), isthmus (i), and bulb (b) of the
rhabditiform esophagus. X 300. 6.2. Longitudinal
section of the anterior end of Draschia megastoma
in the stomach of a horse . Note that the esophagus
is uniform in shape and composed of an anterior
muscular portion (m) and a posterior glandular
portion (p). X 150. 6.3. Draschia megastoma in
6.6
'st •
the stomach of a horse. Note that the esophagus (e)
has a triradiate lumen (I) lined with cuticle . X 200.
6.4. Hastospiculum varani in the lung of a monitor
lizard. Note the triradiate lumen (I) of the esophagus
is not cuticularized and is very large. X 200. 6.5.
Draschia megastoma in stomach of horse. Note the
esophageal-intestinal junction (j). X 150. 6.6. Larva
of Contracaecum sp. in the stomach of a bullfrog.
Note the intestinal diverticu lum (d) that is situated
along side of the esophagus (e). X 200.
7.1

7.3

7.5
Figures 7.1-7.6. Intestines of nematodes. 7.1. intestine (i). X 120. 7.5. Physaloptera sp. in the
Angiostrongylus costaricensis 1n the intestine of a intestine of a monkey. Note the large intestine
monkey. Note the large intestine (i) composed of composed of uninucleate columnar cel ls. X 100.
few multinucleated cells. X 240. 7.2. Elaephora 7.6. Angiostrongylus cantonensis in the brain of a
poeli in the aorta of a water buffalo. Note the small gibbon. Note the large mtestine (i). X 300.
intestine (i) and that it contains black pigment. X
300. 7.3. Graphidium sp. in the intestine of a
rabbit. Note the large Intestine (i) composed of few
multinucleated cells. X 200. 7.4. Cordophilus
saggitus in the lung of a kudu. Note the very small

8. 1
Figures 8.1-8 .6 . Reproduction in nematodes.
8.1. Adult female Ascaris suum in the intestine of a
pig . Note the numerous sections of ovaries (o),
the two sections of uteri (u), and the single section
of Intestine (i). X 20. 8.2. Three sections of adult
male Elaephora schneideri in the carotid artery of
an elk. Note the single genital tract, i.e., vas
deferens, and the small intestine (i). X 60. 8.3.
Uterus of Ascaris suum. Note the developing eggs
limited by mammillated eggshells (s) with abundant
~­
~k~.
8.2
8.6
eosinophilic yolk gland within . X 75.
8.4. Parafilaroides sp. in the lung of a sea lion.
Note the developing larvae (1) , ova (o) , and sperm
(s) within these sections of an adult female parasite.
X 240. 8.5. Adult female oxyund in the intestine of a
mouse. Note the larvae (I) within the eggs and the
eggshells are longitudinally ridged (r) . X 240. 8 .6.
Eggs of Physaloptera sp. in the intestine of an
opossum . The eggs are described as being small,
thick-shelled ovals containing an embryo. X 300.
9.3
s
9.5
Figures 9.1-9.6. Reproduction in nematodes.
9.1. Egg of Anatrichosoma sp. in the skin of a
monkey. Note the bipolar plugs (p) of the
embryonated egg. X 750. 9.2. Tetrameres sp. in
the proventriculus of a woodcock. Note the
characteristic spirurid egg and that the egg has
polar filaments (f). X 300. 9.3. Adult male
lungworm in the lung of a pig. Note the sperm (s)
within the vas deferens. X 400. 9.4. Adult female
Angiostrongylus costaricensis in the intestine of a
9.4
monkey. Note that the female contains both ova
(o) and sperm (s) within the uterus. X 200. 9.5.
Adult male Graphidium sp. in the intestine of a
rabbit. Note the two highly cuticularized spicu les
(s). X 300. 9.6. Posterior end of adult male
Draschia megastoma in the stomach of a horse.
Note the highly cuticularized spicule (s) and the
caudal alae (c) with associated papillae (p). X
300.
Figures 10.1 - 10.6. Reproduction in nematodes. 1 0. 1. Adult
female Stephanurus dentatus from the peri renal region of a pig.
Note the three sections of ovary and that the ova (o) are attached
to a rachis (r) whi le developing. Each section of ovary have ova
in the same stage of development and therefore are undergoing
hologonic cleavage. X 200. 10.2 . Adult female Trichuris vulpis in
the intestine of a dog. The section of ovary (o) is undergoing
hologonic cleavage. All stages of the development of the ova ca n
be seen within thi s one section , from smaller less mature ova (s)
to the more mature, larger ova (I). X 300. 1 0.3. Microfilariae of
Stephanofilaria sp. in the sk1n of a cow. Note that the
microfilariae (m) resemble a cuticular-bound bag of nuclei. X
990. 10.4. Larva of Angiostrongylus costaricensis in the
intestine of a monkey. Note that this larva is more developed
than a microfilaria. X 480. 10.5. Larva of Trichinella spiralis in
the muscle of a pig. Note that this larva is very developed.
having visible musculature (m), intestine (i), and developing
gonad (g). X 300. 1 0.6. Larva of Contracaecum sp. in the
sto mach wall of a bullfrog. Note that th is larva resembles an
adu lt nematode but lacks a mature gonad. X 120.
 NEMATODE GROUPS

RHABDITOIDS
The rhabditoids are small nematodes. The most
common genera in this group include Strongyloides,
Rhabdias, Pelodera and Halicephalobus. Although these
worms have platymyarian musculature (Figs. 12.4, 12.6)
and an intestine that in composed of uninucleate cells
(Figs. 11.4, 12.1, 12.4), their small size often makes it
difficult to delineate these structures in histological
section .
Halicephalobus (pyeviously called Micronema) and
Strongyloides are female when parasitic. Halicephalobus
has one gential tract (Figs. ·11.4, 11.5) and therefore one
ova/egg can be seen in section. Strongyloides, on the
other hand, has a paired geni tal tract (Fig. 12.1) . Both lay
uninucleate eggs which embryonate in tissue in most

species (Fig. 11.6). Halicephalobus contains an esopha-

gus composed of a corpus, isthmus, and bulb in all
stages (Fig. 11.3). This characteristic rhabditiform
esophagus can be seen in fortuitous sections.
Rhabdias is found in the lung of the host. The
musculature is usually inapparent but the intestine is
quite characteristic. It commonly contains large amounts
of yellow to black pigment (Figs. 12.2-12.4). The lateral
chords in Rhabdias are unique in their vacuolated
appearance (Figs. 12.3, 12.4). Rhabdias females contain
developing larvae in the uterus (Fig. 12.3).
The atractids are another group of rhabditoids that
occur in large numbers in their host. The females may
contain large, well-developed larvae (Fig. 12.5).

11.5
Figures11.1-11.6. Rhabditoids . 11.1. Skinofa
dog with numerous Pelodera sp. in hair follicles.
Note the small size of the parasites (p). X 75. 11.2.
Higher magnification of Pelodera from dog hair
lollicle. Note the paired lateral alae (a) and
platymyarian musculature (m). The Intestine (i) is
indistinct. X 1500. 11 .3. Longitudinal section of
Ha/icephalobus sp. 1n brain of horse. Note the
characteristic esophagus with the corpus (c),
Isthmus (i), and bulb (b). X 990. 11.4. Cross
-/
~-a
11 .2
section of Halicephalobus in the brain of a horse.
Note the small size of the parasite , the intestine (i) ,
and the large, single genital tract containing an egg
(e) . X 1000. 11 .5. Longitudinal section of
Halicephalobus in bra1n of horse. Note the dorsal
reflection of the ovary (o) and the large, uninucleate
egg (e) . X 990. 11.6. A developing zygote of
Halicephalobus in the brain of a horse. X 990.
Figures 12.1-12.6. Rhabditoids. 12.1.
Strongyloides ratti in the small intestine of a rat.
Note the small size of the parasite, the relatively
large intestine (i) , and the two gential tracts (g). X
750. 12.2. Rhabdias sp. in the lung of a bullfrog.
Note the large intestine (i) whose cells contain
pigment and the developing larvae (I) within the
uteri. X 100. 12.3. Higher magnification of
Rhabdias sp. in bullfrog lung. Note the vacuolated
lateral chords (c) , the intestine (i) filled with pigment,
and the developing larvae. X 150. 12.4. C ross
section through the posterior end of a Rhabdtas.
Note the vacuolated lateral chords (c). X 240.
12.5. Cross section of an atractid in the intestine
of a tortoise. Note the well developed larvae (I)
within the uterus of the female. X 395. 12.6.
Adult male atractid in the Intestine of a tortoise.
Note the platymyarian musculature (m) , lateral
alae (a) , and the large lateral chords (c). X 420.
OXYURIDS

Oxyurids, the pinworms, are found in almost every

group of animals. They are small and are usually in large
numbers in their hosts. Most of these parasite s have a
rhabditoid esophagus, comprised of a corpus, isthmus,
and bulb (Figs. 13.3, 13.5). Lateral alae are common in
most groups (Figs. 13.2, 13.6). Musculature is
platymyarian (Figs. 13.1, 13.2, 13.4, 13.6). Eggs are
usually thick-shelled and contain an embryo (Fig. 13.4).
The intestine is composed of uninucleate cuboidal to
columnar cells (Figs. 13.1 , 13.2).
~-
13.1

\
.....
13.4~----

•
-., );: . ~ ......
-,.
...
. ---;;
·' ~ - ~~
~··,..,¥
ll.tf':' ~
13.5 13.6
Figures 13.1 - 13.6. Oxyurids. 13.1 . Enterobius esophagus. X 300. 13.4. Oxyurid in the intestine
sp. in the intestinal tissue of a chimpanzee. Note of a mouse. Note the larvae (I) within the eggs
the platymyarian musculature and lateral alae (a). and that the eggshells are longitudinally ridged (r) .
The worm is a male with one genital tract that X 240. 13.5. Longitudinal section of Oxyuris sp. in
contains sperm (s). X 280. 13.2. Enterobius sp. the intestine of a rock hyrax. Note the
in the intestine of a gorilla. Note the lateral alae, characteristic esophagus with corpus (c), isthmus
platymarian musculature, and intestine (i). X 240. (i), and bulb (b). X 150. 13.6. Passalurus sp. in
13.3. Longitudinal section of the anterior end of the intestine of a rabbit. Note the platymyarian
Enterobius sp. in intestinal tissue of a chimpanzee. musculature and that the lateral alae (a) are
Note the corpus (c), isthmus (i), and bulb (b) of the double. X 480.
ASCARIDS

Ascarids are large worms that are found, as adults, in

the intestines of their hosts. Larval ascarids, however,
may be found in any number of other tissues in the final
and/or intermediate host. Larval ascarids of mammals
commonly have lateral alae (Figs. 15.1-15.4) while those
larvae in birds and reptiles usually do not (Figs. 15.5,
15.6). Ascarids have coelomyarian musculature (Figs.
14.1- 15.6). Muscle cells may extend for some distance
into the pseudocoelom (Fig. 15.2) or may be "low" in
nature (Figs. 14.3, 14.5). Lateral chords in adult ascarids
usually do not extend farther into the pseudocoelom than
the muscle cells (Figs. 14.4, 14.5), but in larval ascarids
these chords may fill the pseudocoelom (Figs. 15.1, 15.3).
Many larval ascarids have an excretory cell in the anterior
end that is associated with one of the lateral chords (Figs.
15.2, 15.5, 15.6). The esophagus is simple and the
intestine is large and composed of uninucleate cuboidal to
columnar cells with a low brush border (Figs. 14.1, 14.4).
In some genera, e.g., Contracaecum, the intestine may
extend a caecum anteriad into the area of the esophagus
(Fig. 6.6). The ascarid adult female produces eggs which
contain a uninucleate zygote covered by a thick shell
(Figs. 8.3, 14.3, 14.4).
14.1 14.2

14.5
Figures 14.1-14.6. Adult ascarids. 14.1. procyonis in the intestine of a raccoon. Note the
Heterakis sp. in the intestine of a pheasant. Note "low" lateral chord (c) and the intestine (i) composed
the lateral alae (a) and characteristic lateral chords of columnar cells with a brush border (b) . X 150.
(c). The intestine is composed of uninucleate 14.5. Anasakis sp. in the intestine of a dolphin.
columnar cells and the musculature is coelomyarian. Note the butterfly-shaped lateral chord (c). X 75.
X 150. 14.2. Ascaridia sp. in the intestine of a 14.6. Porrocaecum sp. in the intestine of a hawk.
pigeon. Note the "low" lateral chord (c) and the Note the coelomyarian musculature, the large
coelomyarian musculature (m). X 150. 14.3. intestine (i), and the thick-shelled eggs (e). X 100.
Toxocara canis in the intestine of a dog. Note the
uteri filled with eggs (e). X 20. 14.4. Baylisascaris
Figures 15.1-15.6. Larval ascarids. 15.1. Coiled
larva of Ascaris suum in the lung of a pig. Note the
lateral alae (a) and large lateral chords (c) that fill
the pseudocoelom. X 580. 15.2. Larva of
Polydelphis sp. in the mesentery of a monkey. Note
the lateral alae (a), prominent coelomyarian
musculature, and the excretory cells (e) associated
with the large lateral chords. Polychrome stain. X
450. 15.3. Cross section of larva of Baylisascaris
procyonis in brain of woodchuck. Note the large
lateral chords (c) and lateral alae. X 480. 15.4.
Ascarid larva in mesentery of nine-banded
armadillo. Note coelomyarian musculature and
lateral alae (a). X 395. 15.5. Contracaecum sp. in
stomach of a frog. Note the large eosinophilic
gland (g) associated with one of the lateral chords
(c). X 75. 15.6. Larva of Brevimulticaecum sp. in
peritoneum of toad. Note the eosinophil ic gland
cells (e) associated with the lateral chords. X 200.
STRONGYLE$

There are three subgroups within the strongyles, i.e., the
true strongyles, the trichostrongyles, and the
metastrongyles. Parasites in all three groups have the
same type of intestine. The intestine for a strongyle is
described as being large, and composed of few multi-
nucleated cells (Figs. 16.5, 17.5, 18.5, 19.4, 20.3, 21.2).
Musculature and cuticular modifications vary by group.
True Strongyles
The true strongyles have platymyarian musculature
(Figs. 16.2, 16.5, 17.1, 17.3, 17.5). These muscles cells
are usually very large. The mouth usually has a chitinized
buccal cavity, i.e. capsule (Figs. 16.4, 16.6), and some
members have a lateral internal cuticular ridge (Fig. 16.2).
The cuticle is usually thick and smooth (Fig. 16.1 ). The
lateral chords often appear vacuolated (Figs. 16.2, 17.1,
17.3 , 17.5). The intestine often has a prominent brush
border (Figs. 16.1- 16.6, 17.1-17.6). Eggs are usually
thin-shelled and contain a morula at deposition (Fig.
17.6). Some of the members, e.g. Syngamus, produce
thick shelled, operculated eggs.
Metastrongyles
Metastrongyles, unlike the other members of the
strongyles, have coelomyarian muscles (Figs. 20.1-22.4).
The external cuticle is usually smooth but some genera
have ridges due to the presence of a loose cuticular
sheath (Figs. 20.3, 20.4). Accessory hypodermal chords
are usually present and are characteristic of the group
(Figs. 20.3, 21.2, 21.3, 22.3}. Adult females produce
either eggs or fully developed embryos (Figs. 20.1, 20.2,
20.4, 20.5, 22.1 ). If eggs are produced they are deposited
in host tissues where they embryonate (Fig. 22.5). The
first stage larvae can thus be found in the tissue of the
host and in its feces. Most metastrongyle larvae have the
tip of their tail excentric to the body, thus forming a finger-
like tip (Fig. 22.6). Additionally, some metastrongyle
larvae have caudal spines, the presence and shape of
which can aid in identification of the larvae.

Trichostrongyles
Trichostrongyles have platymyarian musculature.
Characteristically, the external cuticle bears either evenly
spaced (Figs. 18.1-18.6) or irregular longitudinal ridges
(Fig. 19.6). The number and shape of these ridges can
be used in the identification of the parasite. The buccal
cavity in this group is not as pronounced as that of the
true strongyles. Eggs are usually thin-shelled and contain
a morula at deposition.

~~
16.1
Figures 16.1-16.6. True strongyles. 16.1.
Ancylostoma caninum in intestine of a dog. Note
the thick cuticle (c), platymyarian muscles (m), and
large intestine (i) composed of few multinucleated
cells with a brush border. X 150. 16.2. Higher
magnification of Ancylostoma caninum in the
intestine of a dog. Note the low, vacuolated lateral
chords (c), platymarian muscles, and lateral internal
cuticular ridge (r). X 300. 16.3. Uncinaria sp. in
intestine of sea lion. Note the characteristic
c .~
16.2
f'l'
strongyle intestine (i) and the platymyarian muscles.
X 150. 16.4. Longitudinal section of Uncinaria sp. in
intestine of sea lion. Note the highly cuticularized buccal
capsule (c) that gives rise to a muscular esophagus (e).
X 150. 16.5. Larvae of Strongylus vulgaris in
mesenteric artery of horse. Note the platymyarian
muscles (m) and strongyle intestine (i). X 300.
16.6. Longitudial section of larvae of Strongylus vulgaris
in mesenteric artery of horse. Note the highly
cuticularized buccal capsule (c). X 120.
•I"
1~-·
17.1
-
17.5
Figures 17.1-17.6. True strongyles. 17.1. Larva
of Strongylus equinus in pancreas of horse. Note
the musculature, vacuolated lateral chords (c), and
intestine (i). X 320. 17.2. Larva of Oesophago-
tomum sp. in intestine of cow. Note the
characteristic strongyle intestine (i),with brush
border. X 595. 17.3. Larva of Oesophagostomum
sp. in intestine of chimpanzee. Note the
platymyarian musculature, the strongyle intestine,
and the vacuolated lateral chords (c). X 100.
17.4. Larva of Oesophagostomum sp. in intestine of
chimpanzee. Note the iron pigment within the
intestinal cells (i) and the vacuolated lateral chord
(c). X 240. 17.5. Larva of Stephanurus dentatus in
liver of pig. Note the platymyarian musculature (m)
and vacuolated lateral chords (c). X 120. 17.6.
Stephanurus dentatus in perirenal tissue of pig. Note
the lateral chords (c), musculature, intestine (i), and
uterus filled with thin shelled eggs (e). X 30.
18.3
18.5
Figures 18.1-18.6. Trichostrongyles. 18.1 .
Mo/ineus sp. in intestine of a monkey. Note
strongyle intestine (i), platymyarian muscles and
regularly spaced external longitudinal cuticular
ridges (r). X 480. 18.2. Nochtia nochtia in the
intestine of a rhesus monkey. Note how the
longitudinal cuticular ridges are flared at the area
of the lateral chords (1), contrasted to the other
cuticular ridges (r). X 750. 18.3. Cooperioides
hepaticae from the bile duct of an impala. Note the
r
18.6
large intestine and regularly spaced external
longitudinal cuticular ridges (r). X 300. 18.4.
Higher magnification of Cooperioides hepaticae.
X 750. 18.5. Graphidium sp. in intestine of rabbit.
Note the large intestine (i) composed of few
multinucleated cells and the regularly spaced
longitudinal cuticular ridges (r) . X 200. 18.6. Adult
male Graphidium sp. in the intestine of a rabbit.
Note the regularly spaced cuticular ridges (r) and
the brown chitinized spicules (s). X 300.

.-..
19.5
Figures 19.1-19.6. Trichostrongyles. 19. 1.
Cross sections of Longistrongylus sp. in the
abomassum of an impala. Note the large intestine
(i) composed of few multinucleated cells and the
tiny evenly spaced , external longitudinal cuticular
ridges (r) X 900. 19.2. Ostertagia sp. in the
abomassum of a cow. Note the external cuticular
ridges (r). X 470. 19.3. 0/lulanus tricuspis in the
stomach of a cat. Note the evenly spaced cuticular
ridges (r). X 900. 19.4. Haemonchus sp. in the
19.4
19.6
abomasum of a gazelle. Note the external
cuticular ridges (r) , the large intestine (i) composed
of few, multinucleated cells with a well-defined
brush border, and sections of ovary (o). X 240.
19.5. Nippostrongylus sp. in the intestine of a rat.
Note that the external cuticular ridges are not
evenly spaced , nor are they of equal size. X 400.
19.6 . Nippostrongylus sp. in the intestine of a rat.
Note the cuticular ridges (r) are not equal in size
nor are they evenly spaced. X 740.
20.5
.chs
Figures 20.1-20.6. Metastrongyles. 20.1. Filaroides milksi
in lung of dog. Note the large intestine (i) , the coelomyarian
musculature, and the developing larvae (I) within the female.
X 300. 20.2. Parafilaroides sp. in the lung of a sea lion.
Note the developing larvae (1) , the ova (o), and sperm (s)
within the female. X 240. 20.3. Metastrongyle (lungworm)
in lung of doll porpoise. Note the coelomyarian muscles, the
large intestine (i) composed of few multinucleated cells, the
accessory hypodermal chords (c) , the lateral chords (lc) , and
the loose cuticular sheath (s) surrounding the worm. X 300.
*' 20.6
20.4. Filariopsis sp. in lung of a monkey. Note the loose
cuticular sheath forming irregular cuticular ridging (r). X 395.
20.5. Lungworm in lung of pig. Note the difference of size in
the male (M) and female (F) worm. The intestine (i) is large
in both sexes. The male contains sperm (s) in its genital
tract; the female contains developing larva (I) within eggs.
X 150. 20.6. Higher magnification of male lungworm from
lung of pig. Note the coelomyarian musculature. The sperm
(s) are described as eosinophilic ovals with a dark, basoph ilic
nucleus. X 395.
21.5
Figures 21.1-21.6. Metastrongyles. 21.1.
Angiostrongy/us cantonensis in the lung of a rat.
Note the very large intestine (i), the coelomyarian
muscles, and the uteri with developing eggs (e).
X 150. 21.2. Angiostrongylus cantonensis in the
brain of a gibbon. Note the large intestine (i), the
accessory hypodermal chords (c), and the lateral
chords (lc). X 300. 21.3. Angiostrongylus
cantonensis in the brain of a rat. Note the large
intestine (i) and the accessory hypodermal chords
21.6
(c). X 300. 21.4. Angiostrongy/us costaricensis in
the intestine of a monkey. Note the large intestine
(i). X 395. 21 .5. Angiostrongytus costaricensis in
the intestine of a monkey. Note the large intestine
(i), the coelomyarian muscles, and the multiple
sections of ovary (o). X 240. 21.6. Angiostrongy/us
costaricensis in the intestine of a monkey. Note the
large intestine (i) and the ova (o) and sperm (s)
within the uterus of an adult female worm.
X 200.

Figures 22.1-22.6. Metastrongyles. 22.1.
Dictyocaulus viviparus from the lung of a deer. Note
the large intestine (i) and the developing larva (I)
within the uterus of an adult female lungworm. X
200. 22.2. Dictyocaulus viviparus from the lung of
springbok. Note the large intestine (i). X 450. 22.3.
Cystocaulus sp. from the lung of a sheep. Note the
large intestine (i), the two genital tracts (g), the
accessory hypodermal chords (c), and the lateral
chords {lc). X 450. 22.4. Parelaphostrongy/us sp.
22.6
in the brain of a mule deer. Note the large intestine
(i) composed of few, multinucleated cells and the
accessory hypodermal chords (c). X 450. 22.5.
Ae/urostrongylus sp. in the lung of a cat. Note that
the adult female (F) lays eggs (e) which develop
within the lung parenchyma. X 20. 22.6.
Pneumostrongylus sp. in lung of an impala. Note
that the posterior end of this mature larva contains
an eccentric tip (t). X 900.
SPIRURIDS

Members of this group differ greatly from one to another.

Many have cuticular ornamentations around the buccal
cavity in the shape of spines, cordons, bumps, or collars
(Figs. 24.1 , 24.2, 24.4, 25.2). The musculature in the
group is coelomyarian Figs. 23.1-23.3, 23.5, 24.4-24.6,
25.1-25.3, 25.5, 25.6, 26.1 - 26.4). The cytoplasmic
portion of some muscle cells is sometimes quite reduced
(Fig. 23.1 ). Intestines are usually large and composed of
uninucleate cells that are cuboidal to columnar in shape
(Figs. 23.3, 23.4, 25.3, 25.4). Often these cells have
microvilli that form a brush border (Fig. 25.4). Many
members have a eosinophilic fluid found in the
pseudocoelom (Figs. 24.4, 24.5, 25.3, 26.6). The lateral
chords may be enormous (Figs. 23.1, 23.5) and can often
have vacuoles (Fig. 26.3) .
Most adult females produce small, oval, thick-shelled
eggs that are embryonated (Figs. 23.6, 24.4, 26.6). This
is the characteristic spirurid egg. There are some
spirurids, however, that produce embryos without shells,
such as found in Draschia and Thelazia (Figs. 25.5, 25.6).
23.1
Figures 23.1-23.6. Spirurids. 23.1. Physaloptera
sp. in the stomach of an opossum. Note the thick
cuticule, coelomyarian musculature, large lateral
chords (c) that expand into the pseudocoelom, the
large intestine (i), and the numerous eggs within the
uterus. X 40. 23.2. Higher magnification of
Physaloptera in the stomach of an opossum. Note
the coelomyarian musculature and the eggs. X 300.
23.3. Physaloptera sp. in the intestine of a monkey.
Note the large intestine composed of uninucleate
23.2
columnar cells. X 100. 23.4. Higher magnification
of the intestine of Physaloptera. Note the
uninucleate columnar cells. X 300. 23.5.
Physaloptera sp. in the stomach of an opossum.
Note the large lateral chord that is described as
being "stalked." X 100. 23.6. Eggs of Physaloptera.
They are described as being small, thick-shelled,
oval, and containing an embryo. X 300.
24.1
&i'!1
'lj;.:':;,l_··
~
24.5
Figures 24.1-24.6. Spirurids. 24.1. Longitudinal section of the
anterior end of Gnathostoma procyonis in the stomach of a
raccoon. Note the cephalic bulb that contains spines (s) and that
the anterior portion of the body has even larger spines (S). X 75.
24.2. Longitudinal section of the anterior end of l 'hysa/optera
procyonis in the stomach of a raccoon. Note that the anterior end
is surrounded by a cuticular collar (c). X 750. 24.3. Longitudinal
section of the anterior end of Draschia megastoma in the stomach
of a horse. Note that the highly cuticularized buccal capsule (b)
that gives rise to an esophagus with muscular (m) and glandular
(g) portions. X 150. 24.4. Cross section of Echiura sp. in the
stomach of a swan. Note the external cuticular cordons (c), sections
of uteri filled with the typical spirurid eggs (e), eosinophilic fluid in the
pseudocoelom (x), and the large intestine (i). X 100. 24.5. Draschia
megastoma in the stomach of a horse. Note the lateral cephalic alae
(a), the coelomyarian musculature, eosinophilic fluid in the
pseudocoelom (x), and the esophagus (e) with a triradiate lumen (I)
lined by cuticle. X 200. 24.6. Cross section through the anterior end
of Pterygodermatites nycticebi in the intestine of a marmoset. Note
that the lateral alae (a) are sublateral in position. X 240.
25.5
Figures 25.1-25.6. Spirurids. 25.1. Gonglyonema
sp. in the esophageal epithelium of a rat. Note the
coelomyarian muscles and characteristic eggs
(arrows) . X 400. 25.2. Cross sections through the
anterior end of Gonglyonema sp. in the esophageal
epithelium of a deer. Note the cuticular bosses (b)
and lateral alae (a). X 300. 25.3. Spirocerca Jupi in
the esophagus of a dog. Note the coelomyarian
musculature (m) , the intestine (i) with prominent
brush border, the sections of uteri filled with eggs,
the lateral chords (c), and the eosinophilic material
(x) in the pseudocoelom. X 60. 25.4. Higher
magnification of the intestine of Spirocerca Jupi. The
intestine has a prominent brush border (b) . X 400.
25.5. Thelazia sp. in the eye of a horse. Note that
this female produces larvae (1) and not the
characteristic thick-shelled spirurid egg. X 240.
25.6. Draschia megastoma in the stomach of a
horse. Note that this spirurid also produces larvae
(I) and not the small , thick-shelled egg. X 150.
26.3 26.4

26.5
Figures 26.1-26.6. Spirurids. 26. 1. Trichospirura intestine (i), and the larvae (I) within this gravid adult
sp. in the pancreatic duct of a monkey. Note the female. X 50. 26.5. Tetrameres sp. in the
uteri filled with the characteristic spirurid eggs {e). proventriculus of a woodcock. Note that the worm is
X 100. 26.2. Crassicauda sp. in the subcutaneous hypertrophied and filled with myriads of eggs. Note
fat of a porpoise. Note the eggs within the two uteri. the eosinophilic fluid (x) within the pseudocoelom.
X 60. 26.3 . Gnathostoma sp. in the stomach of a X 100. 26.6. Higher magnification of Tetrameres sp.
raccoon. Note the lateral alae (a) and the in the proventriculus of a woodcock. Note the egg has
musculature. X 100. 26.4. Dracunculus insignis in polar filaments (f) and abundant eosinophilic fluid (x)
the subcutaneous tissues of an opossum. Note the in the pseudocoelom. X 300.
musculature (m), inapparent lateral chords (c), small
FILAR IDS

The filarids, or filarial nematodes, are usually small

parasites that infect many groups of animals. Most
members of the group produce distinctive larvae called
microfilariae (Figs. 27.5, 28.2, 28.6, 29.1 ). The presence
of these within the adult female or within the tissues of the
host is important in their identification. Some primitive
filarids produce thick egg shells to protect their larvae
(Figs. 30.2, 30.5, 30.6).
Filarids have coelomyarian musculature (Figs. 27.1-
30.6). In Onchocerca these muscles often atrophy and
are replaced by hypodermal tissue (Figs. 29.1 ). The
lateral chords are usually small (Figs. 27.3, 28.4, 28.6,
29.3) and often there is a lateral internal ridge of the
cuticle at the chords (Figs. 27.4, 29.3, 29.4). Th e most
important diagnostic feature of the filarids is the presence
of a very small intestine (Figs. 27.1-27.3, 28.1, 28.3, 28.4,
28.6, 29.1, 29.5). Genera of "primitive," filarids , however,
often have large, thin-walled intestines (Figs. 30.2, 30.3).
The cuticle in some groups, e.g. , the Dirofilaria , can be
helpful in identification. Some members of the genus
have evenly spaced , external longitudinal cuticular ridges
(Figs. 29.5, 29.6). Adult females of Onchocerca, on the
other hand, have cuticular rings (annulations) that encircle
the worm and thus appear as cuticular bumps in longitudi-
nal section (Fig. 29.2).
Figures 27.1-27.6. Fi larids. 27.1. Elaephora
schneideri in the carotid artery of an elk. Note the
coelomyarian musculature, the small intestine (i), and
that the worm is a male since there is one genital
tube (g). X 60. 27.2 Elaephora schneideri in the
carotid artery of an elk. Note the small intestine (i)
and the sperm (s) within the vas deferens. X 120.
27.3. Cordophilus sagittus in the lung of a kudu.
Note the "low" lateral chords (c), the coelomyarian
musculature, the small intestine (i) and the numerous
sections of ovary (o) within this adult female worm.
X 120. 27.4 Higher magnification of Cordophilus sagittus
illustrating the lateral chords (c), the lateral internal
cuticular ridge (r), and the musculature. X 300. 27.5.
Stephanofilaria stilesi in the skin of a cow. Note that the
intestine (i) is rather large and that the uteri contain
microfilariae (m) and eosinophilic discs (e). X 250.
27.6. Stephanofilaria stilesi in the skin of a cow. Note
the lateral alae (a) , intestine (i) and testis (t) of the adult
male worm . X 750.
28.3
28.5
Figures 28.1-28.6. Filarids. 28.1. Dipetalonema
sp. in the subcutaneous tissues of a monkey. Note
the very small size of the worms. The intestine (i) is
small and the uteri (u) are filled with microfilariae.
X 400. 28.2. Brugia sp. in the lymph node of a cat.
Note the coelomyarian muscles and the microfilariae
(m) within the adult female. X 1000. 28.3.
Elaephora poeli in the aorta of a water buffalo. Note
the musculature and the small intestine (i). X 240.
28.4. Elaephora poe/i in the aorta of a water buffalo.
•
P'
28.6
Note the lateral chords (c) and the intestine (i)
contain black pigment. X 300. 28.5. Filarial
nematode in a brain of a starling. Note the
developing microfilariae (m) and the large intestine
(i). X 480. 28.6. Cardianema sp. in the heart of a
turtle. Note the small intestine (i), the lateral chords
(c), and microfilariae (m) within the sections of uteri.
X 470.

29.5
Figures 29.1-29.6. Filarids. 29.1. Onchocerca gibsoni
in the skin of a cow. Note the extremely small intestine (i)
and the uteri (u) filled with microfilariae. The musculature
has become hypotrophied and replaced by hypodermis
(h). X 120. 29.2. Onchocerca sp. in perioccular tissue of
a dog. Note the evenly spaced cuticular annulations (a)
that can be seen in this longitudinal section of worm.
X 300. 29.3. Dirofilaria immitis in the heart of a dog.
Note the small intestine (i), the coelomyarian muscles,
lateral chords (c), lateral internal cuticular ridges (r) and
29.4
29.6
the muscular vagina (v). X 50. 29.4. Dirofilaria immitis in the
lung of a dog. The worm is degenerating but the lateral
internal cuticular ridges (r) can still be identified between the
bands of coelomyarian musculature (m). X 140. 29.5.
Dirofilaria ursi from the diaphragm of a bear. Note the
coelomyarian musculature and small intestine (i). The
external cuticle has evenly spaced external, longitudinal
cuticular ridges (r). X 70. 29.6. Higher magnification of
Dirofilaria ursi. Note the coelomyarian muscles (m) and the
evenly spaced, external cuticular ridges (r). X 300.
 ......_e
Figures 30.1-30.6. Filarids. 30.1. Hastospiculum
varani in the lung of a monitor lizard. Note the large, tri-
radiate esophagus (e) and the uteri (u) filled with eggs.
X 70. 30.2. Hastospiculum varani in the lung of a
monitor lizard. Note the intestine (i) is rather large and
not typical of a filarial nematode. Also, the uteri (u) are
filled with thick-shelled eggs. The musculature (m) is
coelomyarian. X 100. 30.3. Edesonfilaria malayensis
in the peritoneal tissue of a monkey. Note the glandular
esophagus (e) lacks a lumen. The intestine (i) is larger
than that normally seen in a filarial nematode. X 75. 30.4.
Edesonfilaria malayensis in the peritoneal tissue of monkey.
Note the large glandular esophagus (e) and the section of
uterus (u) filled with microfilariae. X 75. 30.5. Filaria
taxideae in the subcutaneous tissues of a lesser panda. Note
the glandular intestine (i) and, even though the uteri (u) are
empty, thick-shelled eggs (e) containing microfilariae can be
found in the vicinity of this female. X 520. 30.6.
Hastospiculum varani in the lung of a monitor lizard. Note the
thick-shelled eggs (e) containing microfilariae. X 750.
APHASMIDS

The aphasmid nematodes differ from the phasmid

nematodes (all the preceeding groups) in that they lack a
tiny pair of sensory papillae (i.e., the phasmids) on the
caudal end . These structures are not readily identifiable
on histologic section.
Aphasmid nematodes, however, do have several
important morphological features that distinguish them
from the phasmid nematodes. The distinct lateral chords
of the phasmids are not present in aphasmids. There are,
howe"Ver, hypodermal bands with associated nuclei called
bacillary bands, but these structures do not resemble the
chords of the phasmids (Fig . 31.1, 31.3-31.5, 32.3). The
germ cell formation is hologonic. Thus all stages of
development of the germ cells can be found in any section
of ovary or testis (Figs. 31.2, 33.4). The adult female has
only one genital tract (Figs. 31.3, 32.3). In contrast, most
phasmids have two tracts (although most rhabditoids have
only one).

One of the most characteristic feature of the group is

that many species contain a row of esophageal gland
cells (stichocytes) that form a stichosome (Figs. 31.1 ,
31 .3- 31.6 , 32.1 , 32.5, 32.6). This structure surrounds
the esophagus and usually is quite basophilic.
Eggs in many genera are bipolar plugged (e.g.,
Trichuris) but may either be unembryonated or contain a
larva (Figs. 31.3, 31.6, 32.2, 32.3). Some members , such
as Trichinella produce larvae (Fig. 32.6).
Eustrongylides sp. and Dioctophyma sp. have opercu-
lated eggs (Fig. 33.4). In histologic section the esopha-
gus is very glandular (Figs. 33.3, 33.5). The ventral
chord is enlarged (Figs. 33.5, 33.6). The enteron is
attached to the body wall by pseudomembranes (Figs.
33.3, 33.5, 33.6) in the anterior part of the worm .

31.5
Figures 31.1-31.6. Aphasmids. 31.1. Trichuris vu/pis in
the intestine of a dog. There are two sections of the anterior
end of a worm embedded in the intestinal epithelium. The
sections contain a stichosome (s) surrounding the
esophagus (e). The stichosome nucleus (n) can be seen in
one section. Large lateral hypodermal chords are absent
but a large bacillary band (b) is present. X 400. 31 .2.
Trichuris vu/pis in the intestine of a dog. Note the testis (t)
undergoing hologonic cleavage, sections of spicule (s), and
ciromyarian muscles associated with the spicule (m). X 150.
...."'•....
31.2
31.6
31.3. Capillaria sp. in the esophagus of a chicken. Note the
stichosome (s) surrounding the esophagus (e), the hypodermal
bacillary bands (h), intestine (i) and one uterus (u) within the
adult female. X 300. 31.4. Higher magnification of Capillaria
sp in esophagus of chicken. Note the stichosome (s),
esophagus (e), and hypodermal bacillary bands (h). X 750.
31.5. Capillarid in esophagus of shrew. Note the stichosome
(s), esophagus (e), and hypodermal bacillary bands (h).
X 750. 31 .6. Capillarid in shrew esophagus. Note the
embryonated eggs (e). X 480.
Figures 32.1-32.6. Aphasmids. 32.1. Capillaria
aerophilia in lung of opossum. Note the stichosome
(s) in both longitudinal (left section) and cross
section. X 300. 32.2. Capillaria sp. in the
esophagus of a chicken. Note the characteristic
bipolar plugs (p) of the egg. X 900. 32.3.
Anatrichosoma sp. in the skin of a monkey. Note
the hypodermal bacillary bands (h) and the
developing eggs (e) within the uterus. X 100. 32.4.
Egg of Anatrichosoma in the skin of a monkey. Note
the bipolar plugs (p) of the egg and that it is
embryonated. X 750. 32.5. Trich inella spiralis adult
female in the small intestinal epithelium of a mouse.
Note the stichosome (s) and sperm (sp) within the
uterus (u). X 750. 32.6. Trichinella spiralis larva in
the muscle of a pig. Note the hypodermal bacillary
bands (h) and stichosome (s). X 480.
33.3
Figures 33.1-33.6. Aphasmids . 33.1.
Eustrongylides ignotus in the proventriculus of an
egret. Note the characte ristic muscles (m). X 75.
33.2. Higher magnification of Eustrongylides
ignotus. Note the muscles (m), intestine (i), and
spicule (s). X 300. 33.3. Eustrongylides ignotus in
the proventriculus of an egret. Note the glandular
esophagus (e) , musculature, ventral nerve chord
(v), and pseudomembranes (p). X 100. 33.4.
Eustrongylides ignotus in the proventricu lus of an
egret. Note the thick shelled, mammilated eggs (e)
and the ovary (o) undergoing hologonic cleavage.
X 300. 33.5. Dioctophyma renale in the liver of an
otter. Note the esophagus (e) with associated
pseudomembranes (p), the musculature, and the
ventral nerve chord (v). X 40. 33.6. Higher
magnification of Dioctophyma renale in an otte r.
Note the pseudomembranes (p) and ventral nerve
chords (v). X 100.
MORPHOLOGICAL
CHARACTERISTIC OF
ACANTHOCEPHALAN$
IN TISSUE SECTION

Acanthocephalans, as adults, are parasitic in the intestine

of their hosts. The most characteristic feature of these
parasites is the presence of a spined (hooked) proboscis
(Figs. 34.1-34.3), and thus these parasites are often
referred to as thorny-headed worms. ~he worms have a
cuticle and lack a digestive tract. The hypodermal layer is
thicker than the muscular layer (Figs. 34.4, 34.6) and
contains a felted layer under the cuticle (Figs. 34.5 and
34.6) and a thicker layer of cross fibers containing lucunar
channels (Figs. 34.4-34.6). Two layers of muscles, one
circular and one longitudinal, line the pseudocoelom (Fig.
34.5).
Acanthoc::ephalans possess a structure that no other
parasite has. This structure is called a lemniscus and is
associated with the eversion or retraction of the spined
proboscis. Lemnisci are found in the anterior portion of

the worm in close association with the proboscis. The

lemnisci have lemnisci compressor muscles at the
periphery (Fig. 34 .5) .
Sexes are separate in the acanthocephalans. Ova are
developed in "egg balls" within the pseudocoelom of the
female and embryonated eggs are laid when mature
(Figs. 34.5, 34.6).
Encysted juvenile acanthocephala are regularly found
in vertebrates other than the definitive host, in such
animals as fish, frogs, lizards, snakes, etc. These
juveniles have the features of an adult acanthocephalan
with the exception of reproductive organs (Fig. 34.3).
34.3
34.5
Figures 34.1-34.6. Acanthocephalans. 34.1. Anterior
end of an acanthocephalan attached to the intestinal wall
of a swan. Note the proboscis (p). X 150. 34.2. Anterior
end of an Prosthenorchis sp. attached to the intestinal wall
of a monkey. Note the spines (s) (hooks) on the proboscis
(p). X 100. 34.3. Larval acanthocephalan encysted in the
intestinal wall of an armadillo. Note the proboscis (p) with
associated spines (s) and the hypodermis (h) containing
lacanar channels (1). X 75. 34.4. Cross sections of
Neoechinorhynchus sp. in the intestine of a turtle. Note
the thin cuticle (c) , thick hypodermis (h), lucanar channels (1),
developing eggs (e), and lack of intestinal tract. X 75. 34.5.
Macracanthorhynchus sp. in the intestine of a raccoon. Note
the thick hypodermis (h) with both felted (f) and cross fibers
(c), circu(ar (em) and longitudinal (lm) muscle fibers, lemnisci
(1), and lemnisci compressor muscles (m), egg balls (eb), and
maturing eggs (e). X 75. 34.6. Prosthenorchis sp. in the
intestine of a monkey. Note the thin cuticle (c), the
hypodermis with both felted (f) and cross fibers (cf), lucanar
channel (1), egg balls (eb), and maturing eggs (e). X 150.
MORPHOLOGICAL
CHARACTERISTICS OF
TREMATODES IN TISSUE SECTION

Trematodes are commonly found in all species of verte-

brates. Adults are parasitic in all organ systems, but are
most common to the digestive system. The schisto-
somes, which live in the vascular system, can be found
thoughout the body of its host. Trematodes are flat and
twisted upon themselves in histological section, so much
so that the relationship among their internal organs (so
important in their identification) is often hard to discern.
The trematode body is limited by a tegument which
may or may not contain spines. Spines are often on the
anterior portion of the organism but can be found in all the
tegument (Figs. 35.3, 35.6) . Additionally, some trema-
todes, e.g., the echinostomes, have large spines around
the anterior oral sucker (Fig. 35.4).
An oral sucker can be found surrounding the mouth at
the anterior end (Figs. 35.2, 35.4, 35.5, 37.3). Many
trematodes have ventral suckers (Fig. 35.2, 35.5, 35.6,
37.4); others have terminal suckers on the posterior end .
The mouth gives rise to a muscular pharynx (Fig. 35.2)
and in turn to paired intestinal caeca (very rarely is there
only one caecum) (Figs. 35.1, 35.5, 35.6, 37.3, 37.4 ,
37.6). These caeca are blind and end in the posterior
portion of the body. The caeca often contain brown to
black pigment associated with the breakdown of blood
cells (Fig. 37.1 ). This "blood pigment" when extruded
from the body of the parasite can be found in the tissues
as well (Fig. 37.2).
Trematodes, both as adults and larvae , have no body
cavity and are filled with parenchyma (Figs. 35.1-37.6).
 Trematodes are hermaphroditic, i.e. , have both male
and female reproductive organs in one organism (Figs.
35.1, 35.5, 36.3). The exception to this rule is found in
the schistosomes. Reproductive organs are found within
the parenchyma and consist of peripherally located
vitellaria (yolk forming glands) (Figs. 35.1 , 35.3, 35.5,
36.2), one ovary, and paired testes (Figs. 35.1 , 35.5,
36.3). Vitellaria, when stained with H&E, contain eosino-
philic globules (i.e. , yolk material) (Figs. 35.1, 35.3, 35.5,
36.2). Mature sperm, when stained with H&E, are baso-
philic, elongate strands (Fig. 36.3). Eggs can be found
within the organism and the shell, when mature, is often
yellow to brown (Figs. 35.5, 36.1-36.4, 36.6). Most
trematode eggs are operculated, i.e., have a small cap
that pops off the eggs to allow the escape of the enclosed
larva (Fig. 36.3). Schistosomes, however, do not have an
operculum in their eggs and since the adults are found in
the vascular system , eggs are commonly found through-
out the body. Schistosome eggs, when viable, contain a
miracidium (Figs. 36.4, 36.5).
Larval trematodes can be found in tissue. Metacer-
cariae, resistant larval stages of some trematodes, can be
identified by the resistent surrounding cyst wall , suckers,
and paired caeca (Figs. 37.5, 37.6). Other larvae, such as
the mesocercariae, can be found migrating in the tissues
of the host (Figs. 37.3, 37.4).
35.5
Figures 35.1-35.6. Trematodes. 35.1. Trematode
in the liver of an opossum. Note the organism lacks a
body cavity and is filled with parenchyma (p). This
parenchyma contains paired intestinal caeca (c), a
testis (t) and vitellaria (v). X 180. 35.2. Trematode in
pancreas of racoon. Note muscular anterior oral
sucker (s), ventral sucker (vs) and pharynx (p).
X 150. 35.3. Paragonimus sp. in lung of dog. Note
the tegumental spines (s) and vitellaria (v) found
within the parenchyma (p). Yellow eggshells (e) are
35.6
present in the tissue surrounding the worm. X 150.
35.4. Echinostome trematode in the intestine of a goose.
Note the large spines (s) surrounding the oral sucker (o).
X 180. 35.5. Trematode in the pancreas of a raccoon.
Note the relationship among the oral sucker (s), ventral
sucker (vs), intestinal caecum (c), vitellaria (v), testis (t),
and uterus (u) containing eggs. X 75. 35.6.
Echinostome in the intestine of a goose. Note the spines
(s) in the tegument, large ventral sucker (vs), testis (t),
and paired intestinal caeca (c). X 160.

Figures 36.1-36.6. Eggs of trematodes. 36.1.
Trematode in pancreas of raccoon. Note the
parenchyma surrounds numerous coils of the uterus
containing yellow-shelled eggs (e). X 75. 36.2.
Wed/ia sp. in muscle of swordfish. Note the
vitellaria (v) and yellow-shelled eggs (e). X 50.
36.3. Trematode in the bile duct of a cockatoo.
Note the testis (t) with developing and mature sperm
(sp), and that the eggshells are brown and contain
an operculum (o). X 150. 36.4. Schistome eggs
36.6
in liver of turtle. Note the lack of an opercu lum in
eggshell and that a miracidium (m) is contained
within. X 400. 36.5. Schistosome egg in liver of
duck. Note the eggshell (e) and miricidium (m).
X 300. 36.6. Brain of dolphin. Note that this
trematode egg is yellowish in color and that on cross
section is triangular in shape (arrows). X 150.
 .......
·~
··~· -

..
•,
..
..- .
~
~ ,~

•
37.3
·" -. .. "' '

37.5
Figures 37.1-37.6. Trematodes. 37.1.
Paragonimus sp. in lung of dog. Note the brown to
black pigment (p) within the intestinal caecum (c). X
100. 37.2. Lymph node of cow with pigment from
migrating Fasciola magna. Note that the black
pigment (p) can be found in tissue long after the
passage of the parasite. X 45. 37.3. Mesocercaria
of A/aria sp. in connective tissue of water snake.
Note that this migrating larva has an oral sucker (s)
and paired intestinal caeca (c) surrounded by
37.6
parenchyma (p) . X 300. 37.4. Mesocercaria of
Hematoloechus sp. in liver of bullfrog. Note that the
ventral sucker (vs) and paired intestinal caeca (c)
surround by parenchyma (p) are easily identified.
X 450. 37.5. Metacercaria in peritoneum of fish . Note
the resistent walls (w) of the larvae and that they are
filled with parenchyma (p). X 100. 37.6. Metacercaria
in peritoneum of fish. Note the resistent cyst wall (w)
and the the parenchyma (p) of the larva surrounds two
intestinal caeca (c). X 200.
MORPHOLOGICAL
CHARACTERISTICS OF
CESTODE$ IN TISSUE SECTION
Cestodes resemble trematodes in histological section but
lack a digestive tract, i.e. , paired caeca. Adult cestodes
are commonly found in the intestine of their final host and
as larvae in the tissues or cavities of the intermediate host
in their life cycle.
Adult cestodes are segmented into proglottids and this
feature is ofte n apparent in tissue sections (Fig. 38.3,
38.4). Each proglottid contains both male and female
reproductive organs (Figs. 38.3-38.5). The uterus appears
as a cavity in the parenchyma of the worm and is usually
filled with myriads of eggs (Figs. 38.5, 38.6). A "skirt" of
parenchyma (called a velum) may extend from the proglot-
tids of some genera of adult cestodes (Figs. 38.4, 38.5).
When present the proglottids are described as craspedote;
when absent, acraspedote.
The anterior end of both adult and larval cestodes often
contain suckers or other types of holdfast organs (i.e. ,
bothria, bothridia) (Figs. 38.1 and 38.2). In histological
section suckers appear as muscular circles due to radial
striations of the muscle fibers (Figs. 38.1 , 38.2). Some
cestode anterior ends may also contain hooks and thus are
referred to as armed (Figs. 38.1 , 39.2, 39.3 and 40.4) .
Cestod es have calcareous corpuscles embedded within
their parenchyma. These exact function of these struc-
tures is not known but are very helpful for identification
purposes in histological section. Calcareous corpuscles
appear as basophilic to clear ovals that sometimes have a
concentric ringed appearance. Sometimes these cor-
puscles have been "dissolved" in fixation or histological
processing. In t hese cases, however, the parenchyma
continues to conform to the shape of the corpuscles.
Calcareous corpuscles are more numerous in the head
and neck region of both the adult and larval cestode (Figs.
39.3, 39.4, 40.1, 40.6, 42.5). Sometimes corpuscles are
inapparent if other portions of the parasite are studied.
The main groups of cestodes encountered in histo-
logical section are the cyclophyllidians and th e pseudo-
phyllidians. Histologically it is almost impossible to
determine th e exact species of the parasite in question.
However, two differences between members can be used
to narrow the differential. Cyclophyllidians have four
suckers on the anterior end (Figs. 38.1, 39.2, 39.3). Both
the adults and larvae have these structures.
Pseudophyllidians lack these suckers. Cyclophyllidians
also have a muscles (within the parenchyma) that separate
the medullary and cortical regions (Fig. 38.6).
Pseudophyllidians lack these muscles (Figs. 41.5, 41.6).
Larval cestodes are often found in histologic section.
Larval cestodes may be cystic or solid in nature. There
are several types of cystic larval cestodes, i.e., cysticer-
coid, cysticercus, coenurus, and hydatid cyst. The most
common type of solid larva is called a plerocercoid (some
of which are referred to as a sparganum).
Cysticercoids are very small larvae that have a tiny
bladder and the scolex is surro unded by parenchymous
arms (Figs. 39.1, 39.2). These larva can be found within
the lamina propria of the small intestines.
The cysticercus is very common in tissues and is t he
larval stage of many different species. The cysticercus is
a bladder that has an inverted neck with scolex. The
scolex may be armed (Figs. 39.3, 40.1 , 40.3, 40.4), and
always has four suckers ( Fig. 40.4), even though all of
these structures do not show due to histological section-
ing. The tegument, or covering of the parenchyma, is
external to the bladder but when the neck is studied it is
found to be in the interior (Fig. 40.1 ). This is due to the
invagination of the neck. On rare occasions the neck may
be evaginated (Fig. 40.2). This is true of the
strobilocercus (Figs. 39.5 , 39.6). Calcareous corpuscles
are numerous in the neck and scolex (Figs. 39.3, 39.4)
and very rare in the bladder.
The coenurus resemb les the cysticercus but has more
than one scolex (Fig. 40.3). In histological section it has
all the morphologic characterist ics of a cysticercus.
The hydatid cysts are a bladder with myriads of
associated scolices (Figs. 40.5, 40.6, 41.1 -41.4). These
scolices are very small and referred to a protoscolices.
Often the protoscolices are grouped into small clusters
termed brood capsules (Figs. 40.6, 41.2, 41.4). Each
protoscolex has suckers and a scolex. Calcareous
corpuscles are usually present. The hydatid cyst may be
uniloccular (Figs. 40.5, 40.6), multiloccular (Figs. 41.1,
41.2), or a hybrid of these two (Figs. 41.3 , 41.4).
The plerocercoid is a solid-bodied cestode larva that
lacks suckers (Figs. 41.5, 41.6). When a plerocercoid is
suspected, multiple histological sections should be made
to ensure that neither scolex nor suckers are present.
The tetrathyridium is a unique type of larval cestode.
This parasite is solid-bodied, like the plerocercoid, but has
suckers (Fig. 42.2). The tetrathyridium is the larval stage
of Mesocestoides sp., a cyclophyllidium , and must be
differentiated from th e plerocercoid.
Some larval cestodes undergo asexual reproduction
within the tissues of its host. Tetrathyridia of
Mesocestoides is a prime example. Other cestodes ,
presently of undetermined taxon omic status , undergo this
reproduction and are said to be "aberranf' in nature (Fig.
42).

38.3
38.5
Figures 38.1-38.6. Adult cestodes. 38.1. Scolex
of Taenia pisiformis in the small intestine of a fox.
Note the rostellum with hooks (h) and the large,
muscular suckers (s). X 200. 38.2. Scolex of an
adult cestode attached to the intestinal mucosa of a
pygmy sperm whale. Note the suckers (s) and
associated bothrid ia (b). x 200. 38.3. Adult cestode
in the intestine of a canvasback duck. Note the
segmentation of the proglottids (p) and the paired
(i.e., male and female) reproductive organs (r) in
38.2
~-·. ..· .
..~:~. _
rJ
38.4
each proglottid. X 100. 38.4. Adult cestode in the
intestine of a duck. The extensions, vellum (v) of the
proglottids make this cestode craspedote. X 120.
38.5. Higher magnification of adult cestode in
intestine of duck. Note the male cirrus (c) and
female uterus (u) filled with eggs (e). X 150. 38.6.
Dipylidium caninum in the intestine of a dog. Note
that the eggs (e) within the uterus are in "ball" like
masses. Muscles (m) divide the parenchyma into
cortical (c) and medullary (md) regions. X 150.

Figures 39.1-39.6. Larval cestodes. 39.1.
Cysticercoid of Hymenolepsis nana in the abdomen
of a beetle. Note the small bladder (b), scolex with
one sucker (s) visible, and the parenchymous tissue
(p) surrounding the scolex. X 300. 39.2.
Cysticercoid of Hymenolepsis sp. in the lamina
propria of the intestine of a mouse. Note the hooks
(h) on the scolex and the suckers (s). X 300. 39.3.
Cysticercus in the lung of a rabbit. Note the hooks
(h) on the scolex, and two of the four suckers (s).
39.6
The parenchyma of the scolex is filled with calcareous
corpuscles (c). X 75. 39.4. Higher magnification of
calcareous corpuscles (c) in the neck of a cysticercus.
Note the uniformity in shape and size of these
structures . X 300. 39.5. Strobilocercus (an
evaginated cysticercus) from the peritoneal cavity of a
mouse. Note the well developed body, i.e., strobilum
(st). X 7. 39.6. Strobilocercus from the peritoneal
cavity of a mouse. Note the scolex containing suckers
(s) and the body or strobilum (st). X 10.
40.1
Figures 40.1 - 40.6. Larval cestodes. 40.1. Invaginated
cysticercus of Taenia crassiceps in the peritoneal cavity of
a rat. Note the armed (i.e., hooked) rostellum (r) , hook
(h), suckers (s) , calcareous corpuscles (c), and that the
tegument (t) seems to be in the interior of the parasite.
X 75. 40.2. Evaginated cysticercus of Taenia crassiceps
in the peritoneal cavity of a rat. Note the rostellum (r),
suckers (s) , and that the tegument (t) is now at the
exterior of the parasite. X 75. 40.3. Coenurus in the
subcutaneous tissue of a rabbit. Note the three
40.2
40.4
40.6
invaginated scolices (s) and the bladder wall (b). X 30. 40.4.
Coenurus from the subcutaneous tissues of a monkey. The
scolex has four suckers (s) and the rostellum contains hooks (h).
X 200. 40.5. Uniloccular hydatid cyst of Echinococcus
granulosus from the lung of a came l. Note the thick, laminated
cyst wall (w) and numerous protoscolices (p). X 30. 40.6.
Uniloccular hydatid cyst of Echinococcus granulosus from the
lung of a camel. Note the thick, laminated wall (w), calcareous
corpuscles (c), and the numerous protoscolices (p). Groupings of
protoscolices are termed brood capsules (be). X 60.

41.5
Figures 41.1-41-6. Larval cestodes . 41 .1.
Multiloccular hydatid cyst of Echinococcus
multilocularis in the liver of a vole . Note the thin
bladder wall (w) and the numerous protoscolices (p).
X 30. 41.2. Higher magn ification of Echinococcus
multilocularis hydatid cyst. Note the th in bladder wall
(w) and numerous protoscolices (p). Some
protoscolices are grouped into brood capsules (be).
X 75. 41 .3. Hydatid cyst of Echinococcus vogeli from
the liver of a paca. Note the bladder wall (w) and the
41.2
.. , ,
. ' ,1,
41 .6
compartmentalization of the cyst. X 30. 41.4.
Echinococcus vogeli hydatid cyst from the liver of a
paca. Note the bladder wall (w) and the numerous
protoscolices (p) arranged into brood capsules (be).
X 75. 41.5. Sparganum in the muscles of an
opossum. Note the lack of a scolex and the sol id
nature of this larva fi ll ed with parenchyma (p). X 20.
41 .6. Sparganum in the muscles of an opossum. Note
the solid, parenchymus body containing muscles (m)
and excretory canals (c). X 100.
42.1
Figures 42.1-42.6. Aberrant larval cestodes.
42 .1. Proliferating larval cestode in the wall of the
intestine of a dog. Note the solid (s) and bladder-
like (b) morphology of the larvae. X 15. 42.2.
Larval cestodes that resemble tetrathyridia in the
liver of a lizard. Note the invaginated scolices (s) of
these solid-bodied larvae. X 75. 42.3. Proliferating
larval cestode in the intestine of a dog . Note that
this larva has an invaginated scolex with a sucker
(s). X 300. 42.4. Proliferating larval cestode in the
stomach wall of a cat. Note that one larva appears
to be a solid-bodied parasite (s) and there appears
to be asexual reproduction in the sections on the
right (r). X 15. 42.5. Proliferating larval cestode
from the stomach wall of a cat. Note the
calcareous corpuscles (c) embedded within the
parenchyma. X 75. 42.6. Proliferating larval
cestode in the stomach wall of a cat. Note the
infolding of the tegument and the lack of scolex
and calcareous corpuscles. X 150.
MORPHOLOGICAL
CHARACTERISTICS OF
ARTHROPODS IN TISSUE SECTION

In histological section, this group of parasites are repre-

sented as fly larvae (i.e., maggots or bats), mites, and
occasionally as ticks. They can have any combination of
the following features: a highly chitinized body wall (i.e.,
exoskeleton) (Fig.43.4), striated muscles, a trachael ring
system for respiration, and jointed appendages (in the
mites and ticks). The intestine in these organisms is a
simple tube lined by columnar to cuboidal cells (Figs. 43.2,
43.3).
Sots and maggots may have spines on their surface
(Fig. 43.1 ). The muscle is striated and usually prominent
(Figs. 43.1 , 43.2, 43.4). A chitinized tracheal ring system
is easily identified (Figs. 43.2-43.5) and terminates in a
highly chitined spiracular plate (Fig. 43.6).
There are numerous genera of mites that occur both in
the skin and in deeper organs of their host. Mites have
jointed appendages (Figs. 44.2, 44.6) that may have
setae or chitinized ''hairs." Mouth parts are usually
readily identifiable (Figs. 44.1 , 44.2, 44.6). The exoskel-
eton may be highly chitinized (usually in the head area,
the legs, and areas of muscle attachment) (Figs. 44.2-
44.4) but often lacks appreciable chitin. Some mites have
spines (Fig. 44.4) on the exoskeleton; other have ridging
that resembles spines. Egg yolk in the female mite is
brilliantly red with Hematoxylin and Eosin and is present
both in egg yolk glands (Figs. 44.1, 44.3) and within eggs
(Fig. 44.1).
Ticks are sometimes found attached to the skin of any
number of mammals (Fig. 43.5). Their size and thick
"leathery" exoskeleton allow differentiation from mites.

Figures 43.1-43.6. Arthropods. Sots. 43.1. Bot
of Chrysomya sp. in skin of dog. Note the yellow
spines (s) in the thick cuticle (c) and the striated
muscle (m). X 150. 43.2. Bot of Chrysomya sp. in
the skin of dog. Note the spines (s), striated
muscle (m), chitinized trachael rings (t), intestine
(i), and excretory malpighian tubules (mt). X 30.
43.3. Bot of Hypoderma bovis in the skin of a cow.
Note the intestine (i) with engorged blood cells and
chitinized trachael rings (t). X 40. 43.4. Bot of
43.6
Chrysomya sp. in skin of dog. Note the thick
cuticle (c), striated muscle (m) , and tracheal ring
(t). X 30. 43.5. Dermacentor sp. embedded in the
skin of dog. Note the tracheal ring (t). X 40. 43.6.
Bot of Dermatobia sp. in skin of dog. Note the
highly cuticularized spiracular plate (sp). X 50.
Figures 44.1 - 44.6. Arthropods. Mites. 44.1.
Pneumonyssus sp. in the lung of a monkey. Note
the mouthparts (c), chitinized appendages (a),
striated muscle (m), and the yolk material (y) found
both in the yolk glands (yg) and developing egg (e).
X 150. 44.2. Pneumonyssus sp. in lung of monkey.
Note the mouthparts (c), appendages (a) that are
highly cuticularized at points of muscle attachment,
and striated muscle (m). X 300. 44.3. Pneuno-
nyssus sp. in lung of monkey. Note cuticle with
chitinized muscle attachments (ma), striated muscle
(m), and eosinophilic yolk in yolk glands (yg).
X 480. 44.4. Sarcoptes sp. in the skin of a fox.
Note chitinized appendage bases (a) and cuticular
spines (s) on the dorsal aspect of the mite. X 300.
44.5. Demodex canis in hair follicle of dog. Note
the large number of mites and elongated bodies (b).
X 10. 44.6. Demodex canis in hair follicle of pig.
Note the mouth parts (c) and attachments of four
pairs of appendages (b). X 400.
MORPHOLOGICAL
CHARACTERISTICS OF
PENTASTOMES
IN TISSUE SECTION

The pentastomes are a primitive phylum of parasites. As

adults, most pentastomes parasitize reptiles but a few
genera occur in birds and mammals. As larval parasites,
pentastomes parasitize a large range of animals.
Pentastomes have two pairs of hooks surrounding the
mouth (Fig. 45.1 ). The appearance of these hooks led
early researchers to believe the organism had five
mouths, therefore , "penta stoma.'' The worms often
appear segmented (Figs. 45.2, 45.4) but since they are
not, they are referred to as pseudosegmented . Muscula-
ture is striated (Fig . 45.2). The intestine is multicellular
and is bordered by two acidophilic glands for most of its
length (Figs. 45.3, 45.4). These glands are brilliantly
eosinophilic when stained with H&E.
The most characteristic feature of a pentastome is the
presence of sclerotized openings in the body wall. These
openings are associated with the production of new
cuticle during molts. The sclerotized openings have two
cells associated with them. The openings appear as
eosinophilic rings when stained with H&E (Fig. 45.5).
When stained with Movat pentachrome stain, however,
the rings are black in color (Fig. 45.6). This is an impor-
tant fact to remember since it is the cuticle that is last to
be reabsorbed by the body after the parasite dies. Many
times the cuticle of a nymph (a larval stage) can be
identified by the presence of these structures. Penta-
starnes are the only group of animals with this structure.

Figures 45.1-45.6. Pentastomes. 45.1. Anterior
end of a pentastome nymph in the liver of a
monkey. Note the cross sections of hooks (h).
X 150. 45.2. Body wall of a pentastome nymph in
the liver of a monkey. Note the pseudo-
segmentation , acidophilic glands (a) , and striated
muscle (m). X 150. 45.3. Pentastome nymph in
the mesentery of a monkey. Note the acidophilic
glands (a) that surround the intestine (i). X 40.
45.4. Pentastome nymph in the mesentery of a
45.2
45.4
monkey. Note the acidophilic glands (a), the
pseudosegmentatin of the body wall (p), and the
shed cuticle (c) from a previous molt. X 60. 45.5.
Adult Armillifer sp. in the lung of a boa constrictor.
Note the sclerotized openings (s) in the cuticle.
Two cells (c) are associated with the opening. X
750. 45.6. Adult Armillifer sp. in the lung of a boa
constrictor. This section is stained with Movat
pentachrome stain. Note the sclerotized opening
(s) is black. X 750.
RECOMMENDED REFERENCES

Ash LR, Orihel TC. Atlas of Human Parasitology. 4th ed .

Chicago, IL: American Society of Clinical Pathologists;
1997, 41 0 pp.

Beaver PC, Jung RC , Cupp EW. Clinical Parasitology. 9th

ed. Philadelphia, PA: Lea & Febiger; 1984, 825 pp.

Binford CH, Connor DH. Pathology of Tropical and

Extraordinary Diseases: An Atlas. Washington, DC:
Armed Forces Institute of Pathology; 1976, 696 pp.

Chitwood M, Lichtenfels LR. Identification of parasitic

metazoa in tissue sections. Exp Parasitol. 1972:32:407-
519.

Levine HD. Nematode Parasites of Domestic Animals and

of Man. 2nd ed. Minneapolis, MN: Burgess Publishing
Co. ; 1980, 477 pp.

Orihel TC, Ash LR. Parasites in Human Tissues. Chi-

cago, IL: American Society of Clinical Pathologists; 1995,
386 pp.
 INDEX collar, cuticular, 2, 4, 30, 32
Contracaecum, 4, 9, 13, 19, 21
A Cooperioides, 5, 25
cordons, 2, 4, 30, 32
acanthocephalan, 44-45 Cordophilus, 7, 10, 36
acraspedote, 50 corpus, esophageal, 9, 14, 15, 17, 18
Aelurostrongylus, 29 corpuscles, calcareous, 50, 52, 53, 55
alae craspedote, 50, 51
cephalic, 2, 32 Crassicauda, 34
caudal, 2, 3,12 cross fibers, 45
lateral, 2, 4, 5, 15-21 , 33, 34, 36 cuticle, 2, 22, 23, 31, 45, 59, 60
A/aria, 49 cyclophyllidian, 50
Anasakis, 20 cysticercoid, 50, 52
Anatrichosoma, 12, 42 cysticercus, 50, 52, 53
Ancylostoma, 6 , 23 Cystocaulus, 29
Angiostrongylus, 10, 12, 13, 28
annulations, 6, 35, 38 D
aphasmids, 2, 3, 40-43
appendages, 56, 58 Demodex, 58
Armilliter, 60 Dermacentor, 57
arthropods, 56-58 Dermatobia, 57
Ascaridia, 20 Dictyocaulus, 29
ascarids, 19-21 Dioctophyma, 40, 43
Ascaris, 11 , 21 Dipetalonema, 37
atractids, 14, 16 Dipylidium, 51
Dirofilaria, 5, 6, 8, 35, 38
B discs, eosinophilic, 36
diverticulum , intestinal, 9
bacillary bands, 2, 7, 40, 41 Drachia, 6, 9, 12, 30, 32, 33
Baylisascaris, 20, 21 Dracunculus, 34
bladder, 50, 52-55
bosses, cuticular, 2, 4, 33 E
bothria, 50
bothridia, 50, 51 Echinococcus, 53, 54
bots, 56, 57 Echiura, 4 , 32
Brevimulticaecum, 21 Edesonfilaria, 39
brood capsule, 50, 53, 54 egg
brush border, intestinal, 3, 9, 22-24, 30, 33 acanthocephalan, 44, 45
Brugia, 37 arthropod, 56, 58
buccal cavity, 2, 3, 6-9, 22, 23, 30, 32 balls, 44, 45, 51
bulb cestode, 50, 51
cuticular, 2, 4, 32 nematode, 3, 11, 12, 14, 15, 17-20, 22, 24, 27, 28, 30 ,
esophageal, 9, 15, 17, 18 31 ' 33-35, 39-45
bursa, 3 trematode , 46-48
Elaephora, 8, 10, 11 , 36, 37
c Enterobius, 5, 9, 18
esophagus, 3, 9, 19, 23, 32, 37, 41 , 43
caecum, intestinal, 19 intestinal junction, 9
Capillaria, 41, 42 isthmus, 9, 15, 17, 18
capillarid, 41 lumen, 9, 32
Cardianema, 37 rhabditoid , 9, 14, 15, 17
cestodes, 50-55 Eustrongylides, 40, 43
chords excretory cell, 2, 19, 21
accessory hypodermal, 2, 7, 22, 27-29 excretory canals, 54
lateral, 2, 5, 7, 14, 16, 19-24, 28-3 1, 33-38 exoskeleton, 56
Chrysomya, 57
cleavage, F
hologonic, 3, 13, 40, 41
telegonic, 3, 13 Fasciola, 49
coenurus, 50, 53 felted layer, 44
filaments, polar, 12, 34

62
Filaria, 39 metastrongyles, 3, 4, 7, 22, 27-29
filarids, 35-39 microfi laria, 3, 13, 35, 37, 39
Filariopsis, 27 Micronema, 14
Filaroides, 27 miracidium, 46, 48
fluid, eosinophilic, 30, 32-34 mites, 56, 58
Molineus, 25
G muscles
acanthocephalan, 44, 45
glands cestode, 50, 51 , 54
acidophilic, 59-60 ciromyarian, 3, 41
eosinophilic, 21 coelomyarian , 2, 8, 19-22, 27, 28,30, 31,34-39,43
esophageal, 40 lemnisci comp ressor, 44, 45
yolk, 56, 58 longitudinal, 45
Gnathostoma, 4, 32, 34 meromyarian, 2
Gonglyonema, 4 , 33 nematode , 2, 8
Graphidium, 10, 12, 25 platymyarian, 2, 8, 14-18, 22-25
polymyarian, 2
H striated
arthropod , 56-58
Habronema, 8 pentastomid, 59 , 60
Haemonchus, 26
Halicephalobus, 14, 15 N
Hastospiculum, 9, 39
Hematoloechus, 49 nematode, 2-43
Heterakis, 20 Neoechinorhynchus, 45
hooks nerve chord , 2, 40, 43
cephalic, 59 , 60 Nippostrongylus, 6, 26
proboscis, 44, 45 Nochtia, 59, 60
scolex, 52 , 53 nymph
hydatid cyst
uniloccular, 50 , 53 0
multiloccular, 50, 54
hybrid , 50 Oesophagostomum, 8, 24
Hymenolepsis, 52 01/ulanus, 26
Hypoderma, 57 Onchocerca, 6, 35, 38
hypodermal bands, 2, 40 openings, sclerotized , 59, 60
hypodermis, 2, 7, 35, 44, 45 operculum, 46, 48
Ostertagia, 26
ovary
nematode, 3, 11, 13, 15, 26, 28, 36, 40, 43
intestines trematode, 46
arthropod, 56, 57 ovum (ova), 3, 11 , 13, 14, 27, 28,44
nematode, 3, 10, 11, 14-18, 22-31,34-39, 41, 43 oxyurid, 11 , 17, 18
pentastome , 59, 60 Oxyuris, 18
trematode, 46, 47, 49
p
L
Parafilaroides, 11 , 27
lacunar channels, 44. 45 Paragonimus, 47, 49
larva, 3, 11 Parelaphostrongylus, 29
lemniscus, 44, 45 parenchyma, 46-52, 54, 55
Longistrongylus, 26 Passalurus, 18
Pelodera, 14, 15
M pentastoma, 59, 60
pharynx, 46, 47
Macracanthorhynchus, 45 phasmids, 2, 3
maggots, 56 Physaloptera, 4, 7, 10, 11, 30-32
mesocercaria, 46, 49 pinworm, 17, 18
Mesocestoides, 50 plerocercoid , 50
metacercaria, 46, 49 pneumonyssus, 58
Pneumostrongylus, 29
Polydelphis, 21 T
Porrocaecum, 20
proboscis, 44, 45 Taenia, 51 , 53
proglottid, 50, 51 tegument, 46, 50 , 53
Prosthenorchis, 45 testis
protoscolices, 50, 53, 54 nematode, 3, 36, 40 , 41
pseudocoelom, 2, 19, 21 , 30, 31 , 33, 34, 44 trematode, 46-48
pseudomembrane, 40, 43 Tetrameres, 12, 34
Pseudophyllidian, 50 tetrathyridium, 50, 54
Pterygodermatites, 5, 32 Thelazia, 30, 33
ticks, 56
R Toxocara , 20
tracheal rings, 56, 57
rachis, ovary, 13 trematodes, 46-49
reproductive tracts Trichinella , 3, 13, 40, 42
cestode , 51 Trichospirura, 34
nematode, 3, 11 , 14-16 trichostrongyles, 22 , 25, 26
Rhabdias, 14, 16 Trichuris, 7, 13, 40, 41
rhabditoids, 14-16 tubules, malpighian, 57
ridges, cuticu lar
external, 2, 5, 6, 22, 25, 26, 35 , 38 u
internal, 2, 5 , 7, 22 , 23, 35, 36, 38
rostellum , 51 , 53 Uncinaria, 23
uterus,
s cestode, 50, 51
nematode, 3, 11 , 16, 20, 24, 28, 29, 32-34, 36-39, 41
Sarcoptes, 58 trematode , 47, 48
scolex, 50-52, 55
seminal receptical, 3 v
setae , 56
sheath, cuticular, 22, 27 vagina, 3 , 38
sparganum, 50, 54 vas deferens, 3, 11 , 36
sperm vellum, 50, 51
cestode, 51 vitellaria, 46-48
nematode, 3, 11 , 12, 18, 27, 28, 36, 42
trematode, 46, 48 w
spicule, 3, 12, 25, 41 , 43
spines Wedlia, 48
cuticular, 4, 30, 32, 45, 56-58
tegumental, 46, 47 y
spiracular plate, 56, 57
Spirocerca, 33 yolk, egg, 46, 56, 58
spirurids, 7, 30-34
Stephanofilaria, 13, 36
Stephanurus, 8, 13, 24
stichocyte, 40
sti chosome, 3 , 40-42
strobilocercus, 50, 52
stro billum, 52
strongyles, 22-29
Strongyloides, 3, 14, 15
Strongulus, 6, 8, 23 , 24
sucker
cestode, 50, 51 , 53, 55
oral, 46, 47, 49
ventral, 46, 47, 49
Syngamus, 22
An Atlas
of Metazoan Parasites
in Animal Tissues

C.H. Gardiner
Registry of Veterinary Pathology
Armed Forces Institute of Pathology
Washington, DC

S.L. Poynton
Registry of Veterinary Pathology
Armed Forces Institute of Pathology
Washington, DC
 In Memory of

Paul Chester Beaver

1905-1993
Abstract

Gardiner, C.H. and Poynton, S. L. 1999.

An Atlas of Metazoan Parasites in Animal Tissues.
Registry of Veterinary Pathology
Armed Forces Institute of Pathology
American Registry of Pathology
Washington , DC , 63 pp.

This atlas illustrates metazoan parasites in animal tissues.

To facilitate identification , it provides a brief description of
parasites, hosts, transmission, and pathogenesis of the
most important metazoans. Also included are 270 color
photographs of metazoans and associated lesions,
reco rded using optimal con ditions for identification.

Copies of this publication can be purchased from the

American Registry of Pathology Bookstore, Armed Forces
Institute of Pathology, 14th St. & Alaska Ave. , NW, Room
1077, Washington, DC 20306-6000.
Preface Acknowledgments

For years people have been identifying parasites found in This atlas was made possible by the generosity of hun-
histological sections. Often this identification centers dreds of contributors who shared their material with the
around the organ and host in which the parasite is found, Registry of Veterinary Pathology. We thank them . In
with relatively little emphasis on the morphological addition , the following professionals were invaluable: Ms.
characteristics of the parasite intself. How embarrassed Robyn-Anne Ferris, Mr. Ken Stringfellow, Ms . Michele
would you be if you diagnosed a worm in the lung of a cat Richman , and Ms. Fran Card . Thanks to Dr. Lawrence Ash
as Aelurostrongylus when in fact it was Paragonimus! for editorial suggestions. And to all of the histologic
This atlas focuses on the morphological characteristics of technicians at the Armed Forces Institute of Pathology,
all groups of metazoan parasites found in histological Wash ington , DC, who for years cut and stained thousands
section. I suggest you initially study the atlas from of sections of worms, we say thanks. There are none better
beginning to end- it tel ls a story. When you have become than you!
proficient at recognizing the characteristics that differenti-
ate one group from the others, then delve into more
specific identification. A word of caution . Too often
scientists try to identify the parasite too specifically, i.e.,
they identify the parasite to species. For example , the
relationship among internal organs must be taken into
account to identify trematodes . This is often impossible in
histological section since these worms often coil and turn
on themselves. Therefore, be as specific as you can, but
don 't feel bad if you can only go to family or group of
parasite in your opinion .
The large majority of histological sections in the atlas
were stained with hematoxylin and eosin. When other
stains were used they are listed. Examples of over 1 00
metazoan parasites are present in this book. As ever,
experience is the best teacher. But, if you are not initially
proficient, do as we have always done-match your
parasite to the picture. It works!

C. H. Gardiner
Contents
Morphological Characteristics of Metazoan Parasites ..... 1
Morphological Characteristics of Nematodes
in Tissue Section ......................................................... 3
Cuticular manifestations in nematodes ...................... 4
Hypodermis in nematodes .......................................... 7
Musculature of nematodes .......................................... 8
Esophagi of nematodes ........ .... ........... ..... .. .. .. ............ 9
Digestive tract......................... .......... ......... ..... ........... 10
Reproductive tract .. .. ... ...... .... .. .. .. .. .. .. ........... ... .. .. ...... 11
Nematode groups
Rhabditoids ..... ........................................................... 14
Oxyurids .................. .. ....... .. .. .. ...... .. ............................ 17
Ascarids ..................................... .. ............................... 19
Strongyles ..................................................... .. .. ... ...... 22
True strongyles .............................. ... .. ............... 23
Trichostrongyles ..... .. ...... .. ................ .. ............... 25
Metastrongyles ..................................... .. ........... 27
Spirurids ....................... ........ ... ....................... ............ 30
Filarids ....................................................................... . 35
Aphasmids .. ..... .. ... .. .. ................ ...... ....................... .... 41
Morphological Characteristics of
Acanthocephalans in Tissue Sections .. ................... 44
Morphological Characteristics of Trematodes in Tissue ....
Sections ..... ............................. .... .... ........................... . 46
Morphological Characteristics of Cestodes in Tissue ...... ..
Section s .. ......... ...................... ... ... ..... ...... ................. ... 50
Morphological Characteristics of Arthropods in Tissue ..... .
Sections ...... ... ..... ... ................ ........................ ............. 56
Morphological Characteristics of Pentastomes
in Tissue Sections .................... .... ..... ...... ...... ... .. ...... . 59
Recommended References .. ........ .. ..... .. .............. .. ..... ..... 61
Index ............................................ ........................... ........... 62
Figure s
Figures 1. 1-1.6. Cuticular manifestations in nematodes ......... 4
Figures 2. 1-2.6. Cuticular manifestations in nematodes ......... 5
Figures 3.1 - 3.6. Cuticular manifestations in nematodes ......... 6
Figures 4. 1-4. 6. Hypodermis m nematodes .............................. 7
Figures 5.1 -5.6. Musculature of nematodes ............................. 8
Figures 6.1 -6. 6. Esophagi of nematodes .................................. 9
Figures 7.1-7. 6. Intestines of nematodes .. ..................... .... .... 10
Figures 8.1 -8. 6. Reproduction in nematodes .......................... 11
Figures 9.1-9. 6. Reproduction m nematodes .......................... 12
Figures 10.1-10.6. Repro duction in nematodes .......... ........... 13
Figures 11.1-11.6. Rhabditoids ................................................ 15
Figures 12. 1- 12.6. Rhabditoids ........... .. ........ .. .......... .............. . 16
Figures 13. 1- 13.6. Oxyurids ..................................................... 18
Figures 14.1-14.6. Adult ascarids ... .. .......... .... ........ ... ......... ..... 20
Figures 15.1-15.6. Larval ascarids .......................................... 21
Figures 16.1-16.6. True strongyles ........................ ......... ......... 23
Figures 17. 1- 17.6. Tru e strongyles .......................................... 24
Figures 18. 1-18.6. Trichostrongyles ................................ ........ 25
Figures 19. 1- 19.6. Trichostrongyles ................. ............. .. ........ 2 6
Figures 20.1-20.6. Metastrongyles ....... ................................... 27
Figures 21 . 1- 21.6. Metastrongyles ...... .. ............ ...................... 28
Figures 22. 1- 22.6. Metastrongyles .......................................... 29
Figures 23.1-23.6. Spirurids .. ..................................... .. ............ 31
Figures 24. 1- 24.6. Spirurids ... .................................... ............ .. 32
Figures 25. 1- 25.6. Spirurids ... .................................................. 33
Figures 26. 1- 26.6. Spirurids ..................................................... 34
Figures 27. 1-27.6. Filarids .............................. ......................... 36
Figures 28. 1-28.6. Filarids .. .............. .. ........... ..................... .... . 37
Figures 29.1-29.6. Filarids ....................................................... 38
Figures 30.1- 30.6. Filarids ................ .......... ............ .............. ... 39
Figures 31.1 -31. 6. Aphasmids ............... ......................... ......... 41
Figures 32. 1- 32.6. Aphasmids .. .. .. ................................... ........ 42
Figures 33. 1- 33.6. Aphasmids .... ......... .. .................................. 43
Figures 34.1-34.6. Acanthocephalans ............................. ........ 45
Figures 35. 1-35.6. Trematodes ... ..................... ............. .. ......... 47
Figures 36. 1- 36.6. Eggs of trematodes ................................... 48
Figures 37. 1-37.6. Trematodes .. ......... .... .................... .. ........... 49
Figures 38. 1-38.6. Adult cestodes ...... ..................................... 51
Figures 39. 1-39.6. Larval cestodes ......................... ................ 52
Figures 40.1-40.6. Larval cestodes ........... ...................... .. ...... 53
Figures 41.1-41-6. Larval cestodes ............. ..................... .. ..... 54
Figures 42. 1- 42.6. Aberrant larval cestodes ..................... .. .... 55
Figures 43. 1-43.6. Arthropods. Bots ........... .......... .................. 57
Figures 44. 1-44.6. Arthropods. Mites ..... ........... ........... .. .. ....... 58
Figures 45. 1- 45.6. Pentastomes ................ .............................. 60