On the morphology and taxonomy of species of Crenosoma Molin., 1861 (Nematoda: Crenosomatidae) in Bulgaria


Jancev, J.; Genov, T.

Khelmintologiya (25): 45-62

1988


Nematodes of the genus Crenosoma M o 1 in., 1861 parasitize in the bronchioles of Carnivora and Insectivora mammals in Holarctic region. In the opinion of numerous specialists they manifest a specificity in respect to the definitive host. A great number of publications about the species of the genus Crenosoma have appeared in the last decades. Some of them describe new species, others only report data about their distribution. Very few give any information about their morphometric characterization and this information in most cases is incomplete. The lack of sufficient studies of Crenosoma morphometry and particularly of the copulatory organ of the male individuals and the area around the vulva in the females is a serious obstacle in elucidating the taxonomic status of some doubtful species and in improving the taxonomy of nematodes from the genus Crenosoma. The present investigation, including abundant helminthological material, represents a study on the morphology and taxonomy of species from the genus Crenosoma found in Bulgaria.

fi'1,11FAPCKA
AKAAEMH51
HA
HAYKYITE.
BULGARIAN
ACADEMY
OF
SCIENCES
XEJIMHHT0J1Of
HSI.
25.
HELMINTHOLOGY.
25
Co0un.
1988.
Sofia
On
the
morphology
and
taxonomy
of
species
from
the
genus
Crenosoma
Mo
1
i
n.,
1861
(Nematoda:
Crenosomatidae)
in
Bulgaria*
Janao
Janaev,
Todor
Genov
Central
Laboratory
of
Helminthology
Bulgarian
Academy
of
Sciences
Nematodes
of
the
genus
Crenosoma
M
o
1
i
n.,
1861
parasitize
in
the
bronchioles
of
Carnivora
and
Insectivora
mammals
in
Holaretic
region.
In
the
opinion
of
nu-
merous
specialists
they
manifest
a
specificity
in
respect
to
the
definitive
host.
A
great
number
of
publications
about
the
species
of
the
genus
Crenosoma
have
appeared
in
the
last
decades.
Some
of
them
describe
new
species,
others
only
report
data
about
their
distribution.
Very
few
give
any
information
about
their
morphometric
characterization
and
this
information
in
most
cases
is
incomplete.
The
lack
of
sufficient
studies
of
Crenosoma
morphometry
and
particularly
of
the
copulatory
organ
of
the
male
individuals
and
the
area
around
the
vulva
in
the
females
is
a
serious
obstacle
in
elucidating
the
taxonomic
status
of
some
doubt-
ful
species
and
in
improving
the
taxonomy
of
nematodes
from
the
genus
Crenosoma.
The
present
investigation,
including
abundant
helminthological
material,
represents
a
study
on
the
morphology
and
taxonomy
of
species
from
the
genus
Crenosoma
found
in
Bulgaria.
Material
and
Methods
Four
species
from
various
regions
in
Bulgaria
were
collected
and
we
had
at
our
disposal
the
following
materials:
1.
Crenosoma
vulpis
(Rudolphi,
1819;
from
Vulpes
vulpes
Linn
a-
e
u
s,
in
170
of
516
(32,94%);
from
1
to
82
specimens,
total
1759
specimens.
Canis
lupus
Linnaeus
in
2
of
39
(5,12%);
from
1
to
7
specimens,
to-
tal
9
specimens.
*
This
project
has
been
completed
with
the
financial
support
of
the
Committee
for
Science
at
the
Council
of
Ministers
under
contract
No
471.
45
Canus
familiaris
Linnaeus
in
2
of
319
(0,62%);
from
2
to
9
speci-
mens,
total
11
specimens.
2.
Crenosoma
petrowi
M
o
r
o
s
o
w,
1939
of
Martes
foina
Erx-
leben
in
7
of
70
(10%);
from
1
to
19
specimens,
total
57
specimens.
3.
Crenosoma
lophocara
G
e
r
i
c
h
t
e
r,
1951
of
Erinaceus
roumanicus
B
a
r-
r
et-Hamilt
o
n;
in
57
of
76
specimens
(75%);
from
5
to
735,
total
6,517
specimens.
4.
Crenosoma
melesi
sp.
nov.
of
Meles
meles
Li
n
n
a
e
u
s;
in
2
of
59
(3,39%);
73
and
63,
total
80
specimens.
The
nematodes
were
fixed
in
solutio
barbagali.
They
were
cleared
with
lac-
tophenol
and
lactic
acid.
Shape
and
size
of
the
spicule
and
the
gubernaculum
were
studied
after
dissection
from
the
body
of
the
nematodes.
Results
and
Discussion
Genus
Crenosoma
Moli
n,
1861
is
represented
in
Bulgaria
by
four
species
1
parasite
of
insectivorous
mammals
and
3
parasites
of
carnivorous
mammals:
Crenosoma
lophocara
G
e
r
i
c
h
t
e
r,
1951
(Fig.
1)
Host:
Erinaceus
roumanicus
Barr-Ham.
Localization:
bronchi.
Distribution:
Northern
Syria,
Georgia,
Turkey,
Bulgaria
(Strandja
the
vil-
lages
Fakia
and
Gramatikovo,
the
towns
Groudovo
and
Malko
Tarnovo,
Bo-
urgas,
district
and
Sofia
district).
The
species
was
found
in
72,82%
of
the
E.
roumanicus
investigated.
The
invasion
of
hedgehogs
of
various
populations
varied
within
wide
ranges
(40%
in
the
Sofia
district
and
90%
in
Strandja).
Description:
The
nematodes
have
a
cylindrical
body
covered
with
cuticule
on
which
well-shaped,
cuticular
rings
are
observed.
They
are
formed
by
longitu-
dinal
folds
and
by
the
striae
between
them.
At
the
anterior
portion
of
the
nemato-
des
the
longitudinal
cuticular
folds
are
shorter,
do
not
curve
considerably
at
the
posterior
portion,
so
that
the
curves
are
not
so
pronounced
as
in
other
species
of
the
same
genus.
The
first
rings
are
shaped
at
different
distance
from
the
cepha-
lic
extremity,
but
are
situated
always
a
short
distance
in
front
of
the
posterior
portion
of
the
oesophagus
so
that
the
last
ones
are
at
various
distances
near
the
caudal
bursa
of
the
males
and
the
anus
of
the
females.
The
curves
of
the
folds
shap-
ed
at
the
end
of
the
rings
are
better
expressed
in
the
anterior
portions
of
the
nematode's
body.
The
mouth
is
surrounded
by
six
papillae
and
two
amphids
and
opens
into
the
oesophagus
which
is
wider
at
the
posterior
portion.
The
excretory
pore
is
situated
approximately
at
the
level
of
the
middle
of
the
oesophagus,
while
the
cervical
papillae
are
at
the
level
of
the
first
third
of
the
oesophagus.
Males
(30
specimens).
Length
of
the
body
5,95-7,88
mm
(6,78
mm
on
the
average).
Maximal
width
268-400
pm
(ay.
367).
Width
of
the
anterior
extremity
41-52
gm
(ay.
47),
at
the
junction
of
the
oesophagus
with
the
intestine
111-145
gm
(ay.
126),
in
front
of
the
bursa
173-192
pm
(ay.
187).
The
excretory
pore
is
situated
at
123-192
;in
(ay.
144)
and
the
cervical
papillae
at
76--112
p.m
(ay.
78,5)
from
the
anterior
extremity
of
the
body.
The
oesophagus
is
238-277
urn
long
(ay.
256),
its
anterior
portion
is
26--37
pm
wide
(ay.
31)
and
the
posterior
one
55-76
pm
(ay.
61).
The
spicules
are
greatly
hitinized,
almost
equal
in
length
317-376
gm
(ay.
360)
and
their
width
at
the
proximal
portion
is
19-34
nm
(ay.
26),
a
t
the
distal
portion
11-13
gm
(ay.
12,9).
The
proximal
portion
of
the
spicules
46
\
I
11/1111111/111
1
#
4
'
1
0,20
min
•••
Fig.
I.
Crenosoma
lophocura
Gorichte
r,
1951
(orig.)
Male
specimen:
a
anterior
extremity,
lateral;
b
anterior
extremity,
ventral;
c
caudal
bursa,
lateral;
d
caudal
bursa,
ventral;
e
spicules;
gubcrnaculum.
Female
specimen:
g
area
of
the
vulva
47
up
to
the
division
into
two
branches
is
138-166
pm
long
(ay.
151),
the
length
of
the
thinner,
dorsal
branch
is
173-194
gm
(ay.
113)
and
the
width
22-34
pm
(ay.
30).
The
bursa
is
with
three
lobes,
well
developed,
326-455
gm
(ay.
389)x
416-472
pm
(ay.
453)
in
size.
The
ventral
rays
are
longest.
They
have
a
common
stem
and
divide
almost
at
their
middle.
Anteroventral
rays
are
131-150
gm
long
(ay.
140),
the
posteroventral
ones
124-142
gm
(ay.
136).
The
lateral
rays
have
also
a
common
stem.
The
shorter
anterolateral
rays,
long
111-131
pm
(ay.
121)
branch
first.
The
longer
medio-lateral
rays,
with
a
length
of
131-152gm
(ay.
141)
continue
in
the
common
stem,
branch
in
the
second
half
and
reach
the
edge
of
bursa
copulatrix.
The
externo-dorsal
rays
with
a
length
of
124-145
pm
(ay.
135)
are
separate
but
closer
to
the
lateral
rays.
The
relatively
long
dorsal
rays
117-
138
gm
(ay.
127)
are
widest
and
possess
three
small
papillae,
in
contrast
to
the
other
rays
which
end
with
large
globe-shaped
papillae.
Females
(30
specimens).
Body
length
13,82-16,37
mm
(ay.
15,32),
ma-
ximal
width
493-603
pm
(ay.
561).
Width
of
the
anterior
extremity
49-87
pm
(ay.
69),
at
the
junction
of
the
oesophagus
with
the
intestine
166-192
gm
(ay.
178),
at
the
area
of
the
vulva
—510-542
gm
(ay.
533).
Length
of
oesophagus
263-
315
pm
(ay.
286),
width
at
the
anterior
extremity
27-55
pm
(ay.
39),
at
the
pos-
terior
69-102
(ay.
81).
The
excretory
pore
is
at
a
distance
of
131-187
gm
(ay.
152),
and
the
cervical
papillae
at
89-119
gm
(ay.
95)
from
the
anterior
extremity
of
the
body.
The
vulva
is
5,43-7,11
mm
(ay.
6,71)
from
the
anterior
extremity
of
the
body,
the
anus
157-227
pm
(ay.
180)
from
the
posterior
ex-
terior
extremity.
The
uterus
is
filled
with
eggs
in
all
stages
of
development.
Eggs
without
larvae
are
69-76
gm
(ay.
78)x41-43
gm,
in
size
but
eggs
with
larvae
in
them
are
62-83
1.un
(ay.
78)x
41-55
gin
(ay.
50).
Embryos
larvae
are
252-
284
pm
(ay.
272,2)
long
and
14-15
gm
wide.
Notes:
Studies
of
specimens
from
the
genus
Crenosonta
collected
in
Erina-
ceus
routnanicus
in
various
parts
of
Bulgaria
show
that
one
species
(Crenosotna
lophocara)
parasitizes
in
this
host.
Comparing
the
results
from
our
investigations
about
the
morphology
of
C.
lop-
hocara
with
data
from
the
descriptions
of
other
authors
(G
e
r
i
c
h
t
e
r,
1951;
P
o
,a
0
H
a
A,
1955;
F
e
H
o
B,
1984)
we
notice
that
the
position
of
the
first
and
last
tegumental
ring
is
not
absolutely
constant.
The
tegumental
rings
can
be
si-
tuated
both
before
the
level
of
the
posterior
extremity
of
the
oesophagus
and
at
various
distances
near
the
caudal
bursa
in
the
male
and
the
anal
pore
of
the
fe-
male
individuals.
Besides
that,
in
the
specimens
investigated
in
Bulgaria
the
an-
tero-lateral
rays
are
not
separated,
the
three
lateral
rays
come
from
a
common
stem.
At
the
end
of
the
dorsal
ray
are
found
three
ventral
papillae,
not
one,
as
described
by
G
e
r
i
c
h
t
er
(1951).
Crenosotna
melesi
sp.
n.
(Fig.
2)
Host:
Metes
aides
Linnaeus.
Localization:
bronchi
and
bronchioles.
Habitat:
Teteven
(Lovech
district)
on
July
3rd
and
5th,
1984.
Materials:
80
specimens
(64(3`
and
74
29).
In
the
description
of
the
species,
dimensions
of
the
holotype
and
allotype
are
presented
in
parenthesis
after
these
of
the
paratype
specimens.
Length
of
body
is
in
millimeters,
the
remaining
dimen-
sions
in
microns.
All
specimens
have
been
deposited
in
the
museum's
collection
of
the
Central
Helminthological
Laboratory
at
the
Bulgarian
Academy
of
Scien-
ces,
Sofia,
Bulgaria.
Etymology
of
the
species
name:
the
name
of
the
species
Crenosoma
tnelesi
sp.
n.
comes
from
the
latin
name
of
the
definitive
host.
48
14
****Amamia404
111
E
C
E
C
C
}!
O
_
C
'8
J
Fig.
2
Crennsoma
melesi
sp.
n.
(oig.)
Male
specimen:
a
anterior
extremity,
ventral;
b
structure
of
the
anterior
rings
with
cuticular
folds;
b
1
middle
rings;
c
caudal
bursa,
ventral;
d
caudal
bursa,
lateral;
e
spicules;
f—
gubernaculum
Female
specimen:
g
anterior
extremity,
lateral;
h
area
of
the
vulva;
i
area
of
the
anus;
j
eggs
with
larvae
4
XerIMHHT0.11011{H,
KH.
25
Description:
Nematodes
with
relatively
small
size
of
the
body,
completely
covered
by
a
cuticular
sheath
on
which
are
observed
longitudinal
folds
and
fur-
rows
components
of
a
series
of
rings
along
almost
the
entire
length
of
the
body,
usually
from
the
space
around
the
middle
of
the
oesophagus
to
that
before
the
bursa
or
the
anal
pore.
Some
of
the
folds
are
wider
and
between
them,
depending
on
the
localization
of
the
rings,
are
seen
one,
two,
three
or
more
narrower
folds.
The
posterior
ends
of
the
folds
are
seen
particularly
well.
They
form
an
acute,
right
or
obtuse
angle
depending
on
the
extent
of
cuticule
contraction.
In
the
rear
the
cuticular
folds
are
stretched
with
sharp
ends,
therefore
in
the
lateral
portion
of
the
body
they
do
not
protrude.
The
mouth
opens
in
the
oesophagus
with
a
not
very
large
funnel-like
slit.
Males
(6
specimens).
Body
length
4,84-6,76
mm,
ay.
6,27
(6,44).
At
the
anterior
extremity
the
width
of
the
body
is
31-43
gm,
ay.
36
(38),
at
the
end
of
the
oesophagus
103-167
gm,
ay.124
(102),
in
front
of
the
bursa
170-200
gm,
ay.
188
(190).
The
maximal
width
is
300-396
gm,
ay.
255
(300).
Cervical
papil-
lae
are
located
asymmetrically,
laterally,
at
a
distance
of
90-103
gm,
ay.
93,2
(91-101)
from
the
anterior
extremity
of
the
body.
The
excretory
pore
is
found
120-140
gm,
ay.
137
(146)
from
the
head.
The
curves
at
the
back
part
of
the
folds
of
the
cuticular
rings
are
clearly
seen
until
the
25-26th
ring,
then
the
folds
stretch
and
can
be
observed
only
with
greater
magnification.
The
anterior,
well
outlined
cuticular
folds
of
the
rings
are
37-50
gm
long,
the
following
ones
--
81-86
gm.
At
about
the
middle
of
the
body
their
length
reaches
100
gm
and
after
that
they
diminish
once
more. The
last
ring
is
found
at
a
distance
of
76-200
gm
from
the
bursa.
The
oesophagus
is
259-280
pm
long,
ay.
265
(269).
In
front
it
is
much
nar-
rower
13-20
gm,
ay.15.7
(13)
but
at
the
back
it
becomes
wider,
reaching
38-45
gm,
ay.
41,4
(38).
The
two
spicules
are
approximately
equal
in
length,
curving
ventrally
more
more
or
less.
Their
length
varies
within
a
range
of
264-292
pm,
ay.
288,4
(269
and
272).
The
wider
proximal
portion
has
a
size
of
19-25
gm,
ay.
23
(24
and
25)
and
is
followed
by
a
great
constriction
of
13-17
pm,
ay.
15
(16
and
17),
but
their
widest
portion
is
behind
it
28-31
gm,
ay.
30
(30
and
31).
The
proximal
portion
of
the
spicules
before
they
separate
in
two
branches
is
110-126
gm
long,
ay.
117
(110
and
111).
The
ventral
branch,
which
in
its
posterior
part
makes
a
considerable
ventral
curve
is
longer
148-171
pm,
ay.
160
(159-161).
The
shorter
dorsal
branch
is
sharpened,
with
a
length
of
109-132
gni,
ay.
126
(116-118).
Viewed
laterally
the
gubernaculum
makes
two
curves
and
is
86-104
pm
long,
ay.
92
(92).
From
the
ventral
side
it
has
a
more
constricted
middle
part
11-12
gm
and
wider
anterior
and
posterior
portions
with
size
13
to
15
gm.
The
bursa
is
three-lobed,
when
spread
out
296-382
pm
long,
ay.
333
(331)
and
439-596
gm
wide,
ay.
551
(439).
The
bursa
is
supported
by
rays
differing
in
length
and
interposition.
Best
developed
and
longest
are
the
two
ventral
rays
which
are
directed
forward
and
accreted
in
the
first
third
of
their
stem.
They
are
138-152
gm
long,
ay.
147-143.
The
anterolateral
rays
are
the
earliest
to
be
separated
from
the
common
stem
of
all
three
lateral
rays,
their
length
is
86-90
pm
and
they
do
not
reach
the
edge
of
the
bursa.
Mediolateral
and
postlateral
rays
continue
in
a
common
stem
until
they
come
to
their
mid-length
and
reach
the
bursa's
edge.
The
mediolateral
rays
are
102-108
pm
long,
the
rather
slanting
posterolateral
rays
are
equal
in
length
with
the
anterolateral
rays.
Externodorsal
rays
are
separated
from
the
lateral
ones
and
are
76-80
pm
long.
Their
end
is
at
a
certain
distance
from
the
bursal
edge.
The
dorsal
ray
is
wider
and
longer
than
the
externodorsal
rays
and
has
two
lateral
protrusions
in
its
distal
part,
on
which
two
small
papillae
are
seen,
the
po-
sterior
part
is
narrower
and
elongated
reaching
the
bursa's
edge.
Its
length
is
50
93-103
Am,
ay.
99
(98).
The
remaining
rays
have
large
globe-like
papillae
at
their
ends.
Female
s
(30
specimens).
The
body
length
of
sexually
ripe
individuals
with
eggs
is
8,39-16,66
mm,
ay.
11,80
(12,89)
and
their
maximal
width
is
290-
610
gm,
ay.
503
(592).
The
width
of
the
cephalic
extremity
(measured
without
the
cuticular
sheath)
is
25-66
pm,
ay.
42
(56),
at
the
level
just
behind
the
oesophagus
117-191
gm,
ay.
139;
191.
Curves
of
the
cuticular
folds
are
clearly
observed
from
mid-oesophagus
level
to
the
39th
ring
and
after
that
they
gradually
stretch
and
reach
back
near
to
the
anal
pore.
The
oesophagus
is
241-326
pm
long,
ay.
278
(262),
its
width
at
the
anterior
portion
is
20-32
gm,
ay.
25
(25)
and
at
the
posterior
portion
(which
is
widest)
44-69
pm,
ay.
55
(66).
Cervical
papillae
are
situated
la-
terally
at
a
distance
of
43-91
gm,
ay.
77
(89)
from
the
cephalic
extremity
and
the
excretory
pore
is
a
little
behind
at
a
distance
121-237
gm,
ay.
149
(126).
Viewed
ventrally
the
vulva
represents
a
transverse
slit,
situated
quite
in
front
of
the
body's
middle,
at
a
distance
of
3,32-5,49
mm,
ay.
4,39
(5,29)
from
the
head.
The
ratio
between
the
distances
from
the
anterior
extremity
of
the
body
to
the
vulva
and
the
total
body
length
varies
within
the
range
1:3,36-1:3,46,
ay.
1:2,89.
Viewed
laterally
the
vulva
protrudes
considerably
above
the
cuticule's
surface
in
the
shape
of
oval
formations
in
front
of
and
behind
the
slit.
The
vagina
is
relatively
short
and
passes
into
two
uteruses,
which
are
often
amphidelphically
situated,
but
there
are
cases
when
just
after
the
vagina
they
are
at
first
parallel
one
to
the
other.
The
uteruses
are
filled
with
a
very
great
number
of
eggs
at
various
stages
of
develop-
ment,
including
the
stage
of
fully
shapes
larvae.
The
size
of
eggs
without
larvae
varies
from
66
to
78
gm,
ay.
74
x
36-56
pm,
ay.
42
and
of
eggs
with
larvae
from
70-81
pm,
ay.
74,7
x38-51
gm,
ay.
44.
Measured
after
coming
out
of
the
eggs
larvae
are
252-306
gm
long,
ay.
281
and
13-17
pm
wide,
ay.
15,4.
The
anus
is
situated
at
a
distance
of
142-216
nm,
ay.
170
(148).
The
tail
is
cone-shaped
with
a
little
wider
oval
part
at
the
end
and
with
two
papillae,
placed
slightly
in
front
of
the
tail's
end.
Differential
Diagnosis
Up
to
the
present
in
Meles
me/es
of
the
Palaearctic
region
have
been
found
the
Crenosomes:
Crenosoma
taiga,
C.
schulzi
and
C.
vulpis.
Cr
a
i
g
and
An
de
r-
s
o
n
(1972)
point
out
that
Crenosoma
petrowi
parasitizes
in
North-American
bad-
ger
Taxidea
taxus.
KoriTpxmaslitiyc
(1969),
KoH
-
rprimaaillryc
et
al.
(1976),taking
into
account
the
specificity
of
Crenosomes
to
definitive
hosts
are
of
the
opinion
that
the
reports
about
finding
C.
vulpis
in
Metes
metes
are
not
accompanied
with
morphological
data
and
probably
regard
C.
schulzi,
which
they
consider
as
an
independent
species.
111
a
x
m
a
T
o
B
a
(1966)
doubts
that
C.
schulzi
is
an
independent
species,
while
Cr
a
i
g
and
An
d
e
r
s
on
(1972)
suppose
it
is
a
synonym
of
C.
petrowi.
'
The
species
Crenosom
melesi,
described
by
us
is
morphologically
and
metri-
cally
slightly
similar
to
Crenosoma
taiga,
C.
schulzi,
C.
petrowi
and
C.
schachma-
tovae.
Despite
that
it
can
be
clearly
differentiated
from
them
by
the
following
cha-
racters:
from
Crenosoma
taiga
S
k
r
j
a
b
in
et
Petro
w,
1928,
parasite
of
Mus-
telidae
in
Palaearctic
region
by
the
shape
and
the
much
smaller
size
of
the
gubernaculum
(see
Table
1),
by
the
shape
and
width
of
the
anterolateral
ray
(in
C.
taiga
it
is
several
times
wider
than
the
remaining
lateral
rays),
51
Table
1
Comparative
table
with
the
dimensions
of
Crenosoma
melesi
sp.
n.
and
of
some
telidae
species
in
the
Palaearctic
(total
length
and
distance
to
the
vulva
in
mm,
ing
measurements
in
p.m)
other
Mus-
the
remain-
Parameters
C.
melesi
sp.
n.
C.
petrowi
authors'data
C.
schulzi
after
r
a
r
a-
p
H
H,
1956
C.
taiga
Skrjabin,
Petrow,
1928
C.
schach-
matovae
K
OHTpH-
MaBHtlyC,
1969
Males
No.
of
rings
25-26
12-14
17-18
18-20
Length
4,84-6,27
2,26-4,94
4,5-4,7
6,16-7,0
2,78
Width
(max.)
300-396
170-236
220
317
270
Oesophagus
Length
259-280
124
—236
240
Width
38-45
26-30
40
Spicules
Length
264-292
239-268
300-340
286-294
294-265
of
dorsal
protrusion
109-132
48-52
50
131
109-125
Gubernaculum
Length
86-104
89-116
110-120
129
Width
13-15
20-26
35
106
Rays
Ventral
138-156
79-86
150
Anterolateral
86-90
66-74
109
Mediolateral
192-108
79-86
125
Posterolateral
86-90
72-76
93
Externodorsal
76-80
59-68
78
Dorsal
93-103
62-86
-
-
Females
No
of
rings
in
front
30
16-21
20-23
Length
8,56-16,6
2,88-6,09
4,7-8,5
13,7
4,2
Width
(max.)
290-610
212-306
250-350
475
607
Oesophagus
Length
241-326
206-254
Tail
142-216
114-196
105
Eggs
78-81
X
52-67X
60-65
64-68X
38-51
29-38
42
34-36
Larvae
I
stage
252-216
196-249
269-289
-
by
the
smaller
size
and
shape
of
the
dorsal
ray
which
has
at
the
distal
end
two
branches
with
ventral
papillae;
from
Crenosorna
schulzi,
Gaga
r
i
n,
1959
parasite
of
Metes
metes
in
USSR
(Kirghizia)
by
the
smaller
size
of
the
spicules
and
the
longer
(more
than
two
times)
dorsal
protrusion
of
the
spicules,
by
the
relatively
smaller
size
of
the
gubernaculum
(see
Table
1)
and
the
larger
number
of
clearly
seen
cuticular
rings;
from
Crenosoma
petrowi,
M
o
r
o
s
o
v,
1939,
parasite
of
Mustelidae
in
the
Holarctic,
by
the
more
than
two
times
longer
dorsal
protrusion
on
the
spicules,
by
the
type
of
accretion
of
the
ventral
bursal
rays
(in
C.
petrowi
they
ac-
crete
for
2/3
of
the
length,
while
in
the
new
species
for
only
1/3
of
the
length),
by
the
longer
dorsal
ray,
which
is
with
branches
in
the
distal
end
and
with
papillae
at
the
ventral
side;
52
from
Crenosoma
schachmatovae
K
o
n
t
r
i
m
a
v
i
c
h
u
s,
1969,
parasite
of
Mustela
erminea
R.
in
Karelia
(USSR)
by
the
1:18
ratio
between
length
of
spicules
269
mm
and
body
length
4,89
mm.
In
C.
schachmatovae
this
ratio
is
1:11
(spicules
249
mm
long
and
body
2,78
mm),
by
the
shape
and
structure
of
the
dorsal
bursa
ray,
which
is
a
little
wider
than
the
externodorsal
and
with
two
very
short
lateral
protrusions
at
the
distal
end
(in
C.
schachmatovae
it
is
with
a
wide
base,
several
times
wider
than
the
ex-
ternodorsal
rays
and
with
greatly
developed
protrusions
at
the
distal
end),
by
the
significantly
shorter
extenodorsal
rays
(for
C.
schachmatovae.
K
o
H-
Tplimasatryc
et
al.,
1976,
writes
"with
equal
dorsal
and
externodorsal
rays"),
by
the
shape
of
the
distal
end
of
the
spicules
and
the
gubernaculum.
On
the
basis
of
all
these
facts
we
consider
the
species
Crenosoma
melesi
sp.
n.
found
in
Metes
metes
in
Bulgaria
a
new
species.
Crenosoma
petrowi
M
o
r
o
s
o
v,
1939
(Fig.
3)
Host:
Martes
foina
E
r
x
l
e
b
e
n.
Localization:
lung.
Distribution:
URSS
(Litovska
SSR,
Karelska,
Mariiska,
Tadjikska
and
Bash-
kirska
ASSR,
Gorkovska
region,
Caucasus,
Upper
Altai,
the
Krasnojar,
Irkutsk,
Habarovski
regions,
Kamchatka),
Czechoslovakia,
USA,
Canada
(B
a
s
H
o
and
Tomes,
1961;
Benorypos,
1965;
KouTpnivrastimyc,
1963,
1969;
KOHTPRMA.1311
nye,
Cxpst
6
uria,
1963;Me.abitwynaccree,
1959;Mopo3
Os,
1939;
Pomaitos,
1960,
1964;
Pyxrisi,aep
andPyx-
rist,aeBa,
1959;Capi
bixoB,
1955;Tponuxam,
1960,
1962,
1965;
Max-
maTosa,
1966;
Craig, Anderson,
1972;
Craig,
Borecky,
1976;
Addison,
1978).
Description:
Small
nematodes,
covered
by
transparent
cuticule
with
longi-
tudinal
folds,
elements
of
a
series
of
rings,
with
well
protruding
posterior
por-
tion
because
of
the
curves
of
the
cuticular
folds.
These
folds
are
clearly
visible
up
to
the
12th-14th
ring
in
the
males
and
to
the
16th-21st
ring
in
the
females.
After
that
the
folds
are
stretched
and
are
seen
only
with
great
magnification.
The
body
is
cylindrical,
slightly
narrower
at
the
two
extremities.
The
mouth
is
round
and
opens
straight
into
the
cylindrical
oesophagus,
which
is
wider
in
its
posterior
portion.
The
excretory
pore
is
situated
at
a
distance
of
about
1/3-1/4
of
the
oeso-
phagal
length
and
the
deirides
a
little
forward.
Males
(16
specimens).
Body
length
2,26-4,94
min
(ay.
2,02),
maximal
width
170-236
gm
(a.
192),
width
at
the
junction
of
oesophagus
with
the
intesti-
ne
86-98
gm
(ay.
88),
at
the
cephalic
extremity
without
cuticule
20-30
gm
(ay.
88),
in
front
of
the
bursa
146-152
gm
(ay.
148).
The
excretory
pore
is
at
a
distance
of
62-77
gm,
the
cervical
papillae
at
59-68
pm
from
the
anterior
ex-
tremity
of
the
body.
The
oesophagus
is
124-236
gm
long
(ay.
203),
12-18
gm
wide
in
the
anterior
portion
and
26-30
gm
in
the
posterior
portion.
The
caudal
bursa,
which
has
an
oval
shape
is
maintained
by
well-developed
rays.
The
ventral
rays
are
longest
and
originate
from
a
common
stem,
dividing
only
at
the
distal
third
or
fourth,
reaching
almost
to
the
bursal
end.
Anteroventral
rays
are
72-86
gm
long,
the
postero-ventral
—99-119
pm.
Antero-lateral
rays
are
shorter
(70-74
gm)
and
wider.
Mediolateral
and
posterolateral
rays
are
accreted
in
the
basic
two-
thirds,
the
longer
ray
is
the
mediolateral
one.
It
reaches
a
length
of
80-84
1.tm.
The
posterolateral
is
74-76
pm
long.
Externodorsal
rays
are
independent
and
shortest
52-60
gm,
The
dorsal
ray,
which
has
lateral
papillae
at
its
end,
is
long
64-76
gm.
Anterolateral
and
externodorsal
rays
do
not
reach
the
end
of
the
bursa.
The
two
almost
equally
long
spicules
are
slightly
bent
and
reach
a
length
53
E
E
a
b
ill
a
a
C
I
E
011111111
!WO..
r,s
E
Fig.
3.
Crenosoma
petrowi
M
o
r
o
s
o
v,
1939
(orig.)
Male
specimen:
a
caudal
bursa,
lateral;
b
caudal
bursa,
ventral;
c
gubernaculum;
d
spi-
cule,
distal
end
Female
specimen:
e
anterior
extremity;
f
area
of
the
vulva;
g
area
of
the
anus
5
5
O
E
O
54
of
239-268
pm
(ay.
257).
Their
dorsal
branch
is
relatively
short
48-52
pm
(ay.
50)
and
thin.Viewed
laterally
the
gubernaculum
has
the
shape
of
a
little
bark.
It
is
89-116
pm
long
(ay.
106).
Viewed
ventrally
it
has
an
enlargement
in
the
front
and
is
sligtly
sharp
distally.
Its
widest
part
reaches
23-26
pm.
Females
(26
specimens).
Body
length
2,88-8,09
mm
(ay.
4,59)
and
maxi-
mal
width
212-306
pm
(ay.
269,6).
The
width
of
the
cephalic
extremity
is
22-25
pm,
of
the
junction
between
oesophagus
and
intestine
101-112
pm.
The
length
of
the
oesophagus
is
206-254
pm
(ay.
229),
its
maximal
width is
31-38
p.m.
The
vulva,
which
represents
a
transverse
slit
is
situated
at
a
distance
of
1,26-3,71
mm
(ay.
1,79)
from
the
cephalic
extremity.
The
vagina
passes
through
two
opposite
situated
uteruses.
The
anus
is
at
114-196
pm
(ay.
147)
from
the
caudal
extremity
of
the
body.
The
size
of
eggs
is
52-67
pm
(ay.
59)
x
29-38
pm
(ay.
34).
The
length
of
larvae
is
196-249
pm
(ay.
226).
Notes:
The
first
description
of
Crenosoma
petrowi
(M
or
o
z
o
v,
1939)
made
on
one
male
specimen
is
far
from
perfect.
The
presence
of
a
dorsal
protrusion
on
the
spicules
is
not
reported,
but
Kontrimavichus
has
found
dorsal
protrusions
on
dissected
spicules
in
Crenosomas
similar
to
C.
petro
ri
from
Habarovsk
and
Altai
regions.
Pomanoa
(1960)
and
III
axma'rona
(1966)
report
the
pre-
sence
of
such
-
a
protrusion
on
Crenosomas
of
Martes
martes
and
M.
zibellina
in
Krasnoiarsk
region.
In
all
investigated
male
specimens
in
Bulgaria
we
found
that
the
dorsal
spicules
have
a
relatively
thin
and
short
dorsal
protrusion.
Comparison
of
the
morphology
and
the
metric
characteristics
of
the
Crenosoma
petrowi
specimens
we
studied
with
the
reports
of
other
authors
concerning
various
portions
in
the
area
and
various
hosts
(Table
2)
shows
the
indisputable
similarity
between
them
but
also
the
existence
of
obvious
differences.
Thus
the
length
of
spicules
in
Martes
foina
specimens
in
Czechoslovakia
(P
r
o
k
o
p
i
Z,
1965)
is
175-215
pm.
The
same
host
in
Bulgaria
(data
from
the
present
investigations)
has
spicules
239-264
pm
long.
In
Martes
Martes
from
the
Gorki
region
(M
o-
p
o
3
o
n,
1939)
they
are
225
pm,
in
the
same
host
in
Altai
(K
o
n
T
p
to
a
e
y
c,
1969)
301
pm
and
in
Mantes
pennanti
from
USA
(Craig
and
Anderson,
1972)
300-346
tim.
Differences
exist
also
in
the
size
of
eggs,
the
studies
of
KonTparvtaanityc
(1969)
show
that
the
females
are
43-47
x
23-29
pm,
in
our
investigations
they
are
52-67
x
2
9
3
8
pm
and
in
Czechoslovakia
70-
36
pm.
It
is
also
interesting
that
although
the
males
from
Czechoslovakia
are
very
long
(5-6
mm)
their
spicules
are
relatively
very
short.
We
found
in
Martes
foina
from
the
Sofia
region
only
one
female
specimen,
8,94
mm
long
and
with
maximal
width
255
pm.
The
length
of
the
oesophagus
is
269
pm,
its
maximal
width
43
pm,
the
anal
pore
is
at
124
pm
from
the
posterior
extremity
and
the
eggs
are
62
x
38
pm.
The
Crenosoma
found
in
Ursus
americanus
from
Ontario,
Canada
(A
d
d
i-
s
o
n,
1978)
which
are
considered
to
be
of
the
species
Crenosoma
petrowi
have
larger
male
and
female
body
size,
longer
spicules
and
gubernaculums
(Table
2).
Also
longer
bursa!
rays.
The
lack
of
sufficiently
complete
data
about
Crenosoma
from
various
parts
of
the
C.
petro
ri
area
does
not
provide
possibilities
for
an
exact
evaluation
of
the
type
of
the
indicated
differences.
Evidently,
a
well
expressed
intraspecific
variation
exists
in
C.
petrowi
and
it
seems
very
probable
that
the
fur-
ther
thorough
studies
of
the
differences
existing
between
specimens
from
Ame-
rica
will
prove
the
existence
of
new
species
in
the
genus
Crenosoma.
Crenosoma
vulpis
(R
u
d
o
1
p
h
i,
1819)
(Fig.
4)
Hosts:
Vulpes
vulpes
B
a
c
h
s
t
e
i
n,
Canis
lapis
Linnaeus,
Canis
fa-
rniliaris
L.
Localization:
Bronchi,
trachea,
nasal
cavity.
55
I
(
E
,
$
b
0
C
d
E
Y
I
FYY
FT,
uUIP
Fig.
4.
Crenosoma
vulpis
(R
udolph
i,
1819)
(orig
)
Male
specimen:
a
anterior
extremity;
b
caudal
bursa,
lateral;
c
caudal
bursa,
ventral;
d
gubernaculum;
e
spicules,
proximal
and
distal
Female
specimen:
f—
anterior
extremity;
g
area
of
the
vulva;
h
area
of
the
anus
1
11
E
56
Table
2
Dimensions
of
Crenosoma
petrowi
Moroso
v,
1939,
after
different
authors
(total
length
and
distance
from
Cephalus
to
vulva
in
mm,
the
remaining
dimensions
in
pm)
Parameters
M0p
0
3013,
1
939,
fr
om
M.
mar
te
s
Pr
o
kop
i
(:,
1965,
fr
o
m
M.
fo
ina
KouTp
uN
ta
B
1
4-
y
c,
1
96
9,
fro
m
M.
z
ibe
l
lin
a
Cra
ig,
An
der
-
son,
197
2,
fr
om
M.
p
en
n
a
ta,
T.
taxe
s
A
ddison,
19
78,
from
U.
a
mer
ic
an
a
Au
t
hor
s
'
da
ta
fr
om
M.
fo
in
a
Males
Length
1,5
5,0-6,0
3,70
3,9-5,6
5,3-7,9
2,26-4,94
Width.
(max.)
178
300-320
234
260-380
170-236
Oesophagus
Length
221
201
236-285
230-320
124-236
Width
(max.)
36
36-37
26-30
Spicules
Length
(total)
225
175-215
301
300-347
320-440
239-264
of
dorsal
protrusion
43
63-80
48-52
Gubernaculum
Length
113
120
90-114
91-140
88-116
Width
14
22
17-30
23-26
Rays-length
Anteroventral
95-160
79-86
Posteroventral
94-168
99-119
Anterolateral
75-110
66-74
Medolateral
85-117
79-86
Posterolateral
82-117
72-76
Externodorsal
65-100
59-68
Dorsal
100-150
62-86
Females
Length
6,8-8,0
3,75-5,55
6,9-10,3
10,2-17,8
2,88-8,06
Width
(max.)
300-360
224-258
270-320
370-500
212-306
Oesophagus
Length
259-319
206-254
Distance
from
vulva
to
cephalic
extremity
1,44-1,93
4,6-8,0
1,25-3,71
Tail
12,0
158-323
125-165
100-255
114-196
Eggs
43-47
52-67
Larva
I
stage
20,29
29-38
Distribution:
Europe
,
the
Asian
part
of
USSR,
USA
(B
en
nen
a,
1959-
Tarapnx,
1958;
rEeTpon
and
rarapun,
1938;
PyxrisineB,
1940;
Cr
a
i
g,
A
n
d
e
r
s
o
n,
1972;
A
d
d
i
s
o
n,
1978),
Bulgaria
(all
over
the
coun;
try,
from
20%
in
Eastern
Bulgaria
to
45,91%
in
Western
Bulgaria)
Description:
Nematodes
with
cylindrical
body,
slightly
narrower
at
both
ends.
The
cuticular
scheath
has
visible
lorigitudinal
folds
on
the
first
rings
in
male
specimens
up
to
the
14th-22nd
and
in
the
females
24th-31st.
The
cuticular
folds
form
a
curve
at
their
posterior
portion
at
different
angles
obtuse,
right
and
even
acute,
depending
on
the
contraction
and
maceration
of
the
cuticule.
At
the
back
the
folds
are
stretched
and
can
be
seen
with
greater
magnification.
The
mouth
is
oval
and
opens
into
a
cylindrical
oesophagus
which
is
different
in
its
pos-
terior
portion.
The
excretory
pole
is
at
the
level
of
the
first
third
of
the
oesopha-
gus
and
the
cervical
papillae
are
a
little
in
front
of
it.
57
Males
(50
specimens
from
V.
vulpes).
Body
length
3,39-8,36
mm
(ay.
6,06)
and
maximal
width
246-361
gm
(ay.
310).
Width
of
the
cephalic
extremity
31-46
pm
(ay.
125)
in
front
of
the
bursa
146-220
vim
(ay.
179).
The
excre-
tory
pore
is
situated
86-108
pm
and
the
cervical
papillae
78-84
gm
from
the
cephalic
extremity.
The
length
of
the
oesophagus
is
252-309
pm
(ay.
278)
and
the
width
at
the
anterior
portion
18-28
gm
and
at
the
posterior
45-55
gm.
Spread
out
caudal
bursa
has
a
size
269-427
Rm
(ay.
354)
x
356-483
pm
(ay.
402).
Ventral
rays
are
the
longest
146-188
pm
(ay.
158),
accreted
at
the
base,
branch-
ing
slightly
after
their
middle
and
reaching
the
edge
of
the
bursa.
The
three
lateral
rays
originate
in
a
common
stem
and
the
first
to
be
separated
is
the
antero-late-
ral
one
which
is
63-99
pm
long
(ay.
83).
Medio-
and
postero-lateral
rays
continue
in
a
common
stem
until
the
last
third
and
then
branch.
The
length
of
the
medio-
lateral
rays
is
73-111
pm
(ay.
90)
and
of
the
shorter
posterolateral
it
is
69-96
pm
(ay.
84).
The
externodorsal
rays
are
separated,
independent,
59-81
pm
long
(ay.
70).
Of
the
lateral
rays
only
the
mediolateral
one
reaches
near
the
edge
of
the
bursa,
the
remaining
lateral
and
dorsal
rays
do
not
reach
so
far.
The
dorsal
ray
is
rela-
tively
long
101.-151
gm
(ay.
116)
and
wider
than
the
remainig
rays
coming
near
the
posterior
extremity
of
the
bursa,
where
it
becomes
slightly
wider
and
then
forms
a
hardly
visible
protrusion.
The
two
almost
equal
and
slightly
bent
spi-
cules
are
336-401
pm
long
(ay.
371).
At
the
distal
part
of
the
spicules
a
thin
dorsal
branch
is
separated,
68-96
pm
long
(ay.
79).
The
length
of
the
gubernaculum
is
106-139
gm
(ay.
128)
and
its
width
in
the
proximal
part
is
17-25
gm,
in
the
middle
15-17
gm
and
in
the
distal
part
17-22
gm.
Females
(50
specimens
of
V.
vulpes).
Body
length
9,11-16,16
mm
(ay.
13,69),
maximal
width
246-510
pm
(ay.
401).
Width
at
the
anterior
extremity
34-56
pm
(ay.
45),
at
the
junction
between
oesophagus
and
intestine
127-
163
gm
(ay.
141).
The
excretory
pore
is
at
a
distance
of
92-139
pm,
and
the
cer-
vical
papillae
of
78-90
pm
from
the
cephalic
extremity.
The
length
of
the
oeso-
phagus
is
252-424
pm
(ay.
306),
the
width
at
the
anterior
portion
26-30
gm,
at
the
posterior
one
45-68
pm.
The
vulva
is
a
little
forward
from
the
middle
of
the
body
at
a
distance
of
4,96-7,33
mm
from
the
cephalic
extremity.
The
anus
is
142-188
pm
(ay.
167)
from
the
caudal
extremity.
Two
lateral
papilae
are
seen
on
the
tail.
Eggs
with
larvae
have
a
size
of
62-81
x
36-45
pm.
Larvae
are
269-
286
gm
long
and
15-16
gm
wide.
Notes:
C.
vulpis
is
a
parasite
of
foxes,
dogs
and
other
Canidae.
The
results
of
our
investigations
show
that
in
Bulgaria
this
parasite
affects
most
foxes,
rarely
wolves
and
only
per
chance
domestic
dogs.
The
morphological
and
metric
charac-
teristics
of
this
species
differentiate
it
clearly
from
the
remaining
species
of
the
genus
Crenosorna
in
Bulgaria.
It
is
particularly
well
distinguished
by
the
size
of
its
spicules,
the
length
of
their
dorsal
protrusion,
the
type
of
ventral
and
la-
teral
ray
accretion
of
the
bursa,
etc.
Comparison
between
the
data
we
obtained
for
Crenosoma
vulpis
and
the
data
of
other
authors
from
different
hosts
(Table
3)
shows
the
presence
of
certain
dif-
ferences.
In
the
descriptions
of
II
erpoa
(1941)
and
Kofurplimannilyc
(1969)
the
length
of
the
gubernaculum
is
550
pm.'
Most
probably
there
is
a
techni-
cal
mistake
in
this
case.
According
to
Ko3J10
El,
(1977)
it
is
96
pm,
to
A
d d
i-
s
o
n,
(1978)
106-140
pm
and
to
our
data
106-139
pm.
The
guiding
Tables
differentiating
C.
vulpis
from
C.
schulzi
indicate
that
while
in
the
first
species
the
elements
building
up
cuticular
rings
form
an
obtuse
angle,
the
second
one
they
form
an
acute
angle.
Results
obtained
in
our
investigations
about
the
structure
and
distribution
of
cuticular
rings
in
Crenosorna
vulpis
and
other
species
encountered
in
Bulgaria
58
Table
3
Dimensions
of
Crenosoma
vulpis
Rudolphi
after
various
authors
(total
length
and
distance
from
cephalus
to
vulva
in
mm,
the
remaining
dimensions
in
gm)
Parameters
1
-
1
eTpon,
1941
K
3
11
0
13,
1977
Graig
and
Anderson
1972
Addison,
1978
U.
americana
Authors'
data
Males
Length
3,5-5,0
3,36
—5,92
5,4-5,8
4,4-6,4
3,3-8,4
Width
(max.)
280-300
274
278
—390
246-362
Oesophagus
Length
300
160-330
255-308
252-309
Width
(max.)
55
40
32-59
45-55
Spicules
Length
370
232-300
354-404
373-428
336-401
sp.
protrusion
100
49
79-90
68-96
Gubernaculum
Length
550
96
121-130
106
—140
106-139
Width
18-33
17-25
Rays-length
A
nteroventral
133
—180
146-188
Posteroventral
133-180
Anterolateral
65-110
63-99
Medolateral
75-100
73-111
Posterolateral
60-98
69-96
Externodorsal
60-69
59-81
Dortal
112-135
101-151
Females
Length
12-15
5-12,9
6,0
—10.1
6,0-8,8
9,11-16,16
Width
(max.)
300-450
230-500
385-575
246-510
Oesophagus
Length
230-490
282-286
247-395
252-424
Distance
from
vulva
to
cephalic
extremity
8,2-6,1
Tail
870
150
132-193
142-188
Eggs
70
—76
X
7
0
X
37
62-84X
50-52
36-45
Larva
I
stage
264-340
280
246-308
269-289
show
that
the
distance
to
the
first
and
to
the
last
ring,
as
well
as
the
angle
formed
by
curving
the
folds
at
the
posterior
portion
of
the
rings
are
subject
to
variations
and
depend
to
a
great
extent
on
the
degree
of
contraction
and
maceration
of
the
cuticule.
For
that
reason
we
accept
that
the
taxonomical
importance
of
this
character
is
relative.
It
is
of
interest
that
the
C.
vulpis
specimens
from
foxes
we
investigated
are
similar
to
those
of
the
American
bear
from
Ontario,
but
with
much
larger
size.
Besides
that,
the
distal
portion
of
the
spicules
differs
considerably
in
shape.
In
the
American
specimens
there
is
a
sheath
and
a
rounded
shape
of
the
distal
end,
while
in
our
specimens
they
are
sharp.
It
should
be
noted
that
due
to
the
abundant
serial
material
of
the
four
species
from
the
genus
Crenosoma
,
which
have
been
found
in
Bulgaria,
the
possibly
most
complete
morphometric
characteristics
were
made
which
could
be
used
for
com-
parison
with
species
of
this
not
well-known
group
of
nematodes
found
in
other
59
regions
of
the
world.
Shape
and
size
of
the
spicules,
gubernaculum
and
caudal
bursa
rays
can
be
accepted
as
the
most
stable
and
promising
characters.
Moreover,
results
obtained
in
our
investigations
show
that
every
species
of
the
genus
Crenosoma
was
found
in
definite
hosts
not
only
from
one
family
but
even
from
one
and
the
same
genus
or
species.
For
example,
Crenosoma
melesi
was
found
only
in
M.
meles,
C.
petrowi
only
in
Mantes
fbina,
C.
vulpis
mainly
in
V.
vulpes
and
more
rarely
in
C.
lupus
and
very
rarely
in
C.
fanhiliaris
and
C.
lo-
phocara
in
E.
roumanicus.
These
results
are
in
support
of
the
opinion
of
some
authors
that
in
species
of
the
genus
Crenosoma
exists
a
high
host
specificity,
therefore
the
data
about
finding
species
of
the
genus
Crenosoma
in
nonspecific
hosts
are
to
be
doubted.
References
A
d d
i
so
n,
E.
M.
1978.
Crenosoma
spp.
(Nematoda:
Metastrongyloidea)
from
black
bears,
Ur-
sus
americanus,
of
central
Ontario.
Can.
J.
Zool.,
56,
1993-1997.
Baru
1,
V.,
J.
P
r
o
k
o
p
i
6.
1972.
The
systematic
position
and
the
distribution
of
nematodes
of
the
genus
Crenosoma
M
o
1
i
n,
1861,
parasitic
in
the
Redgehog
(Erinaceus
europaeus).
Vestnik
Ceskoslovenske
spolecnosti
Zoologicke,
36,
No
1,
6-11.
Br
g
1
e
z,
J.,
S.
Valentin6i
c.
1968.
Nametnice
medvjeda
(Ursus
urctos)
L.
Parasites
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MaTepaanbi
no
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xyHbfix
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B:
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xotaormAx
H
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T.
2.
KFLIHHH011,
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H
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n,
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e
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0.
14.
1965.
riapawiTipleciate
‘repBx
Ha3emmax
1703BOHO'lliblX
maTepmcoBoro
110-
6epexhg
Oxorcxoro
mopH
(cpayna,
3K051041451,
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KaHn.
AEtce.,
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BengeB
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M.
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K
inrremno
re.abminfrocimpibi
mneiconaTaionuax
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IIyruiL
Tpygm
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B.
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MaTepHanm
no
renbmturrocbayHe
MUM
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BCHBOTHMX
Ha
lore
KHprH3HH.
B:
Pa6orm
no
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60-nemo
P.
C.
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Anma-ATa,
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FeHo
B,
T.
1984.
Ken:minx
Ha
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14
rpH3aHrre
B
RsnrapHH.
C.,
BAH.
348
c.
T
p
o
B
a,
E.
T.
r
ell()
B.
1961.
XenmHwrotiayHa
Ha
Erineaceus
rounumicus
B
a
r
—H
a
m.
OT
Crpafirma.
B:
Koivinn.
143C.R.
na
npFrpo/nia
orttnula
ffa
3apa3a
B
Orpaiuma.
C.,
BAH,
123-133.
3
y
e
B
a,
H.
C.,
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H.
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y
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o
B.
1965.
K.
(Davie
rensmaaToB
xingusix
mnexortHratotusx
(Mustelidae,
Canidae,
Felidae)
6accertH
p.
Y1:11,1.
—B:
MaTepHanbt
Harm.
xoncl).
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II.
2.
M.,
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60
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172
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1963.
rOJMMIIHTErI
ICYHEPHX
ABIIIHCF0
BOCTOKB.
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Tpyom
TEJIAH,
13,
26-47.
K
0
H
T
p
H
M
a
B
H
Y
y
c,
B.
JI.
1969.
Fenbmtutrocitaytta
xyuutx
H
nyrn
ce
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432
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Konrp
H
M
a
B
H
u
y
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C.
C
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Si
6
it
n
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1963.
Fensmsurrotbayna
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it
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K
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tur
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B
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e
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Crenesomatidae,
Schul
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Meracrpolirsnomten
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)10413031114X
-
B:
OCHOBIA
lie-
maTonormi.
T.
26.
M.,
Hayxa,
44-68.
M
a
H
ynbc
Kt{
it
C.
14.
1953.
reTHWHIITLI
xylltax
Bypscr-Monronbcxoll
ACCP.
-
B.
Pa-
6orm
no
rensmintronormn
(K
75-nermo
man.
K.
H.
Cxpn6ttna).
M.
All
CCCP,
398-405.
M
en
lox
4
y
H
a
N
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r
e
c,
C.
1959.
Hoxoropme
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o
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xynnueo6pa3irmx
SIHTOBCKOit
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Acta
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2,
No
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M
o
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o
30
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1939.
F1apa3nrmtcoote
nepan
nyoutmx
3uepeft
cemelicrna
Mustelidac
(tcynmsx)
rOpItX0BCKOn
o6nacrn.
-
Tpyrtm
rOpbKOBCK.
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ati-ra,
4,
1-44.
My
M
H
H
0
8,
H.
1963.
MaTCP1111.161
HO
feHL.M11111.04ritylie
MOM%
WIOTOSTHHIAX
ACHBOTHIAX
1/36eKVI-
cirma.
-
Y36.
6non.
wyptt.,
6,
44-49.
H
e
T
p
o
B,
A.
M.
1940.
llapa3urnmectcrte
gepan
KYBIAIX
MOCKOBCKOPO
3oonapxa.
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Tpyom
Mom.
3oonapxa,
1,
202-231.
11
e
T
p
0
B,
A.
M.
1941.
FHPICTELIAC
60HC3HH
HYLUILTAX
313epen,.
M.,
Mewcoynapoonau
xmtra,
227
C.
II
T
p
o
B,
A.
M.,
B.
F.
Farapn
H.
1938.
Flpmmrutennan
(onclxbepenimanbnan)ottarttocrinca
ne-
TO.IHMX
T0JIbMIIHT030B
11)3111161X
3Bepeil.
-
Tpyou
Bcec.
140.-ra
TeRBMIIIFITOHOTHJI,
3,
33-47.
P000na
H,
T.
E.
1955.
Fensmunroltaytta
AltICKX
mnexonnTatontax
JIarortexcitoro
rocyoapc
-
raen-
HOTO
3anoneomixa.
Tp.
14H-Ta
300.1.
AH
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CCP,
14,
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Pomano
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H.
B.
1960.
Fenbmartrocpayna
co6oJur
nopyrnx
xyH6Hx
Kpactionpcxoro
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xypnan.,
39,
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Pomanoa,
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B.
1964.
Feabmturrocbayna
KyH6I4X
Cpermero
110BOYINMSI.
-
Ygeti.
3an.
Fopbx•
neg.
att-ra,
48,
120-133.
Pyxn
ri
o
e
B,
H.
1940.
Jlertymbie
renhmturrm
6apcyxon
(3mopoc
-
3pourane3
H
xpeno30ma-
To3).
Harmo-meronmi.
3art.
rnaB.
ynp.
no
3anonernmxam,
6,
87-99.
Pyxnnoe
I1.,M.
Pyxange
B
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1959.
Fenblvinttrorbayna
NYHI611X
n
opyrnx
xmunsix
Cenepo-
Kamsa3cxoro
3anonerunixa.
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Tpyrtm
Kanxa3cx.
3anoaeruntica,
5,
125-135.
Caouxo
B,
H.
A.
1955.
Fern.mmrrootayna
nyumbtx
3Bepelt
A3cp6a11,o3cancxon
CCP.
Kano.
Alice.
Laxy.
T
p
o
H
it
Ica
51,
A.
A.
1960.
Fern,mnirrodeptaXIKKX
nyuntsix
3acpck
TaTapcxofl
ACCP.
-
Tpygm
06-13a
CCTCCTBOHCHIAT.
Hplf
Kaaancxom
yn-Te.
63,
335-358.
Tponnxa
st,
A.
A.
1962.
K
axonoriptecxoll
xaparrepttertuce
rem,mintiotbayttm
necnoll
xynnum
Bocromno-Kamexoro
Roan.
-
Te3ncm
poxnagou
nay
,
moll
xoncitep.
BOF.
LI
.
2.
M.,
202-204.
Tponn
Ica
A,
A.
A.
1965.
Onur
axonormectcoro
anann3a
renbmtnnorbaynm
necnoll
K)1I11104.
-
Tpyribt
06utecrna
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nptt
Ka3ancx.
ytt-re,
67,
124-151.
Maxmar
o
a
a,
B.
14.
1966.
FOMPIAllrbl
xyltuttx
Kapenwn.
-
Tpyow
FEJ1AH,
17,
277-289.
AH‘ICB,
51.
19o1.
Bspxy
xernstottrodtaynara
Ha
ancattaxa
(
Vulpes
ropes
crucideru
B
e
c
11
s
t
e
i
n)
a
Certepo3ana4na
Bwiraplin.
-
XCHIVIHISTOJI01
-
HR,
4,
73-96.
e
B,
51.
H.
Part
xca
xo
B.
1977.
Xenmairrocbayna
Ha
nnewitara
(Vulpes
'wipes
cricidera
Bechstei
n)
B
Ceaepo3anaona
B.b.orapast.
-
Xenmmtronortm,
4,
73-
96.
61
O
mopt.onornn
H
Taxconomnn
BI4g0B
Nita
Crenosoma
Moli
n,
1861
(Nematoda:
Crenosomatidae)
B
BoarapHn
RHEIO
Auaea,
T000p
Teuoe
IfetunpalbuaR
fraeopumoptur
ze.mmuunroaoruu
EAR
(P
e
3
10
NI
e)
il3rIeHa
MOPC130.301
-
1451
14
THKCOHOMIIA
3Hat1Hnesibtroro
HHcria
HemaTog,
npHHagne-
watuttx
lc
4
Him
am
poHa.
Crenosoma
Moli
n,
1861,
o6HappgeHtibtx
y
XHWHbIX
H
HaceKomommax
maexonwratow,nx
B
BonrapHH.
CilertaHa
HOJIHHH
MOIXDOMeTpiPleCKBA
xapaxreplicTinca
B14410B
11
Ha
OCHOBBHH11
Hali6anee
Hagexcrunza
11pH311BKOB
(Dopmbi
14
pa3mepoH
CIIHKyJI
H
pyJmKOB,
pa3-
mepoH,
ttopmbi
14
pacnonoxcerntm
pe6ep
IcayganbHort
6ypcbl
onpetkeneHbr
cne-
,aytoulHe
B14)4bI
xpetto3om:
Crenosoma
lophocara
G
e
r
i
c
h
t
e
r,
1951
y
Fri-
naceus
roumanicus;
Crenosoma
melesi
n.
sp.
—y
Meles
meles;
Crenosoma
petrowi
M
o
r
o
s
o
w,
1932
y
Modes
foina;
Crenosoma
vulpis
(R
u
d
o
1
p
h
i,
1819)—
y
Vulpes
vulpes,
Canis
lupus
14
Canis
familiaris.
YCTHHOBBeHHE4e
angm
OTJ114LIBIOTCH
y3KOii
CIle11144MILIHOCTb}0
B
OTHOHNHHH
X035IeB.
62