Junker_265-280.indd


INTRODUCTION

Helmeted guineafowls, Numida meleagris (Linnaeus, 
1758), are distributed throughout most of South Afri-
ca and almost the entire African continent (Del Hoyo, 
Elliot & Sargatal 1994). Studies to elucidate the hel-
minth fauna of these hosts in South Africa have been 
undertaken by Saayman (1966), Crowe (1977) and 
Verster & Ptasinska-Kloryga (1987), but were re-
stricted to the Eastern Cape, the Northern Cape and 
Gauteng Provinces.

Although relatively wide-spread in Africa, Crested 
guineafowls, Guttera edouardi (Hartlaub, 1867), are 
scarce and have a limited distribution within South 
Africa. They occur in the Limpopo, North West, Mpu-
malanga and KwaZulu-Natal Provinces and are listed 
as rare or accidental in Gauteng Province (Hockey, 
Dean & Ryan 2005; Lepage 2007). To date our 
knowledge concerning their helminth fauna is virtu-
ally non-existent.

Ortlepp (1937, 1938a,b, 1963) reported on the ces-
tode and nematode parasites of guineafowls of 
southern Africa present in the National Collection of 
Animal Helminths, formerly known as the Onderste-
poort Helminthological Collection, or material made 

265

Onderstepoort Journal of Veterinary Research, 74:265–280 (2007)

Helminths of guineafowls in Limpopo Province, 
South Africa

K. JUNKER and J. BOOMKER*

Department of Veterinary Tropical Diseases, Faculty of Veterinary Science, University of Pretoria, Private 
Bag X04, Onderstepoort, O110 South Africa

ABSTRACT

JUNKER, K. & BOOMKER, J. 2007. Helminths of guineafowls in Limpopo Province, South Africa. 
Onder   stepoort Journal of Veterinary Research, 74:265–280

Between July 2005 and November 2006 the gastro-intestinal helminths of 15 Helmeted guineafowls 
and a single Crested guineafowl from Musina, Limpopo Province were examined, and in July and 
August 2005 helminths were collected from five Helmeted guineafowls from Mokopane in the same 
province. The acanthocephalan Mediorhynchus gallinarum, the cestodes Abuladzugnia gutterae, Da-
vainea nana, Hymenolepis cantaniana, Numidella numida, Octopetalum numida, Ortleppolepis multi-
uncinata, Porogynia paronai, Raillietina angusta, Raillietina pintneri, Raillietina steinhardti and Rail-
lieti na sp. and the nematodes Ascaridia numidae, Cyrnea parroti, Gongylonema congolense, Hadjelia 
truncata, Sicarius caudatus, Subulura dentigera, Subulura suctoria, Subulura sp., Tetrameres numida 
and an unidentified subulurid were recovered. A single trematode species, Dicrocoelium macrosto-
mum, was present in the liver. Mediorhynchus gallinarum, A. gutterae, O. multiuncinata, H. truncata 
and S. caudatus are recorded for the first time from Helmeted guineafowls, as well as from South Afri-
ca. South Africa is a new geographic record for D. macrostomum, G. congolense and D. nana. 
Subulura suctoria, G. congolense and H. truncata from the Crested guineafowl constitute new host-
parasite associations.

Keywords:  Acanthocephalans, cestodes, guineafowls, Guttera edouardi, nematodes, Numida mele-
agris, trematodes

* Author to whom correspondence is to be directed. E-mail: 
joop.boomker@up.ac.za

Accepted for publication 13 April 2007—Editor



266

Helminths of guineafowls in Limpopo Province, South Africa

available to him by various collectors. He described 
several new species of cestodes and nematodes 
and added numerous parasites to the host-parasite 
list of guineafowls in South Africa. His reseach, how-
ever, was of a taxonomic nature and the material at 
his disposal represented incidental findings rather 
than complete collections.

In this paper we present data obtained from 16 birds, 
including a single Crested guineafowl, at Musina, 
Lim popo Province, and from five Helmeted guinea-
fowls at Mokopane, Limpopo Province, South Africa.

MATERIAL AND METHODS

In July and August 2005 a total of five Helmeted 
guineafowls were sampled in the vicinity of Moko-
pane (Potgietersrus), Limpopo Province. A complete 
helminth recovery was not possible, but some of the 
worms present in the small intestine of three of the 
birds, the complete caeca of one of them and part of 
the intestinal and caecal contents of another were 
collected and fixed separately in 70 % ethanol.

In July 2005 and in May, July and November 2006, 
three, five, three and four Helmeted guineafowls 
(eight males and seven females) were collected on 
a farm approximately 60 km west of Musina (Mes-
sina), Limpopo Province (22°22’ S, 29°30’ E, Altitude 
700–800 m). The vegetation type in the study area 
is classified as Mopani veld (Acocks 1988).

The birds were aged according to the criteria estab-
lished by Siegfried (1966) and in total ten adults and 
five juveniles were collected. The juveniles were be-
tween six and ten months old (Siegfried 1966). In 
November 2006 a single adult female Crested guin-
eafowl, found moribund in a wire snare, was made 
available to us for examination.

The carcasses of the birds were opened according 
to standard techniques for necropsies of chickens, 
and the viscera removed. The trachea was opened 
and macroscopically examined for helminths.

The crop, proventriculus, gizzard, small intestine 
and caecum/colon were separated and individually 
washed over a 150 μm sieve. The livers of nine Hel-
meted guineafowls and the single Crested guinea-
fowl were sliced into 5 mm wide sections and incu-
bated in phosphate-buffered saline at 40° C for 30 
min. Subsequently, the slices together with the sa-
line were washed over a 150 μm sieve. The gastro-
intestinal and liver residues left on the sieves, as 
well as the organs themselves were fixed separate-
ly in 70 % ethanol and transported to the laboratory 

at Onderstepoort. Each sample was examined un-
der a stereoscopic microscope and the helminths 
removed.

Cestodes were stained in haematoxylin and mount-
ed in Canada balsam or mounted and cleared in 
Hoyer’s medium. Acanthocephalans were cleared 
in Hoyer’s medium and studied as temporary mounts 
in the same medium. All nematodes were cleared in 
lactophenol for identification.

The ecological terms are used in accordance with 
the definitions of Margolis, Esch, Holmes, Kuris & 
Schad (1982).

RESULTS

All the guineafowls were infected and all were con-
currently parasitized by acanthocephalans, cestodes 
and nematodes.

Data on the prevalence, intensity and habitat prefer-
ence of the parasites from the Helmeted guineafowls 
in Musina are presented in Tables 1 and 2. Five of 
the nine hosts (55.6 %), whose livers were examined, 
harboured Dicrocoelium macrostomum, the intensity 
of infection ranging from 8 to 182 flukes. In addition, 
the livers of three of the nine birds yielded five, 11 
and five young specimens of Porogynia paronai. 
These had the typical three circles of large hammer-
shaped rostellar hooks and small, unarmed suckers. 
No differential development could be seen in any of 
the proglottids of the short strobilae which ranged 
from 2.3 to 3.8 mm (n = 5) in length. The scolices 
were 689–746 μm wide and the rostella were 261–
329 μm wide.

Birds from Mokopane yielded the nematodes Subu-
lura suctoria, Subulura dentigera and Ascaridia nu-
midae and seven cestodes, namely Ortleppolepis 
multiuncinata, Porogynia paronai, Raillietina stein-
hardti, Raillietina pintneri, Raillietina sp., Numidella 
numida and Octopetalum numida.

Subulura dentigera and S. suctoria were co-specific 
in the two hosts from Mokopane. One of these har-
boured a total of 579 nematodes consisting of 142 
male and 159 female S. suctoria, 134 male and 126 
female S. dentigera and 18 immature Suctoria spp. 
These nematodes were suspended freely in the 
contents of the posterior saccate part of the caeca, 
virtually occupying the entire lumen (Fig. 2D).

Eight of the 15 helmeted guineafowls from Musina 
harboured S. dentigera and S. suctoria concurrently, 
and in all these hosts S. suctoria by far outnumbered 



267

K. JUNKER & J. BOOMKER

T
A

B
L
E

 1
 

T
h
e
 s

ite
 p

re
fe

re
n
ce

, 
p
re

va
le

n
ce

 a
n
d
 i

n
te

n
si

ty
 o

f 
in

fe
ct

io
n
 o

f 
a
ca

n
th

o
ce

p
h
a
la

n
s 

a
n
d
 c

e
st

o
d
e
s 

co
lle

ct
e
d
 f

ro
m

 1
5
 H

e
lm

e
te

d
 g

u
in

e
a
fo

w
ls

 i
n
 L

im
p
o
p
o
 P

ro
vi

n
ce

, 
S

o
u
th

 A
fr

ic
a
. 

A
d
d
iti

o
n
a
l d

a
ta

 o
n
 g

u
in

e
a
fo

w
l h

e
lm

in
th

s 
in

 s
o
u
th

e
rn

 A
fr

ic
a
 f
ro

m
 v

a
ri
o
u
s 

a
u
th

o
rs

 a
re

 in
cl

u
d
e
d
 f
o
r 

co
m

p
a
ri
so

n

P
a
ra

s
it

e

T
h

is
 p

a
p

e
r

V
e
rs

te
r 

&
 P

ta
s
in

s
k
a
-K

lo
ry

g
a
 

(1
9
8
7
)

S
a
a
y
m

a
n

 (
1
9
6
6
)

C
ro

w
e
 

(1
9
7
7
)

O
rt

le
p

p
 

(1
9
6
3
)

S
it

e
P

re
v
a
le

n
c
e
 

(%
)

In
te

n
s
it

y
P

re
v
a
le

n
c
e
 

(%
)

In
te

n
s
it

y
P

re
v
a
le

n
c
e
 

(%
)

In
te

n
s
it

y
P

re
s
e
n

c
e

P
re

s
e
n

c
e

M
e
a
n

 (
±

 S
D

)
R

a
n

g
e

M
e
a
n

R
a
n

g
e

M
e
a
n

R
a
n

g
e

A
c
a
n

th
o

c
e
p

h
a
la

n
s

M
e
d
io

rh
yn

ch
u
s 

g
a
lli

n
a
ru

m

M
e
d
io

rh
yn

ch
u
s 

n
u
m

id
a
e

M
e
d
io

rh
yn

ch
u
s 

ta
e
n
ia

tu
s

S
I

S
I

S
I

1
0
0

– –

5
5
.7

 (
±

 7
8
.3

 )

– –

2
–
2
3
1

– –

– – 2
7

– – 1
.7

– – 0
–
2
2

– 3
9

–

– 1
1
.5

–

– ?
–
2
7

–

– – +

– – –

C
e
s
to

d
e
s

A
b
u
la

d
zu

g
n
ia

 g
u
tt
e
ra

e

A
b
u
la

d
zu

g
n
ia

 t
ra

n
sv

a
a
le

n
si

s

D
a
va

in
e
a
 n

a
n
a

H
ym

e
n
o
le

p
is

 c
a
n
ta

n
ia

n
a

N
u
m

id
e
lla

 n
u
m

id
a

O
ct

o
p
e
ta

lu
m

 n
u
m

id
a

O
rt

le
p
p
o
le

p
is

 m
u
lti

u
n
ci

n
a
ta

P
a
ro

n
ie

lla
 s

p
.a

P
o
ro

g
yn

ia
 p

a
ro

n
a
i

R
a
ill

ie
tin

a
 a

n
g
u
st

a

R
a
ill

ie
tin

a
 p

in
tn

e
ri

R
a
ill

ie
tin

a
 s

te
in

h
a
rd

ti

R
a
ill

ie
tin

a
 s

p
.

R
a
ill

ie
tin

a
 s

p
.a

 

S
kr

ja
b
in

ia
 d

e
w

e
ti

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

S
I

8
0

– 3
3

4
0

6
7

6
7

8
7

– 4
7

5
3

8
0

5
3

7
3

– –

1
1
.7

 (
±

 8
.2

)

– 5
.8

 (
±

 4
.4

)

4
2
.7

 (
±

 7
0
.4

)

5
5
.9

 (
±

 7
2
.7

)

9
1
.9

 (
±

 1
1
0
.7

)

9
.3

 (
±

 5
.2

)

– 1
2
.3

 (
±

 1
3
.3

)

1
0
.3

 (
±

 7
.9

)

5
.3

 (
±

 3
.9

)

4
9
.0

 (
±

 6
0
.2

)

1
5
.8

 (
±

 8
.8

)

– –

1
–
2
8

– 1
–
1
0

1
–
1
2
4

1
–
1
4
4

1
–
3
6
0

2
–
1
4

– 5
–
3
9

1
–
2
5

2
–
1
2

4
–
1
3
7

6
–
2
8

– –

– – – – 2
9

4
8

– 2
5

– 8 4
4

3
1

– 3
5

–

– – – – 1
.8

8 – 1
.5

– <
 1

.0

3
.9

1
.9

– 2
.7

–

– – – – 0
–
4
2

0
–
7
2

– 0
–
1
7

– 0
–
2
1

0
–
4
5

0
–
2
0

– 0
–
1
7

–

– – – – 4
7

7
5

– – 7
5

– 3
6

– – – –

– – – – 8
.7

1
6
.0

– – ? – 6
.3

– – – –

– – – – ?
–
1
4

? – – ?
–
5

– 3
–
2
7

– – – –

– – – – – + – – + – + – – – –

+ + + – + + + – + + + + – – +

a
 

L
is

te
d
 b

y 
V

e
rs

te
r 

&
 P

ta
si

n
sk

a
-K

lo
ry

g
a
  
(1

9
8
7
) 

a
s 

a
 n

e
w

 s
p
e
ci

e
s,

 b
u
t 
w

e
re

 n
o
t 
su

b
se

q
u
e
n
tly

 d
e
sc

ri
b
e
d

S
I 

=
 s

m
a
ll 

in
te

st
in

e



268

Helminths of guineafowls in Limpopo Province, South Africa

T
A

B
L
E

 2
 

T
h
e
 s

ite
 p

re
fe

re
n
ce

, 
p
re

va
le

n
ce

 a
n
d
 in

te
n
si

ty
 o

f 
in

fe
ct

io
n
 o

f 
n
e
m

a
to

d
e
s 

co
lle

ct
e
d
 f
ro

m
 1

5
 H

e
lm

e
te

d
 g

u
in

e
a
fo

w
ls

  
in

 L
im

p
o
p
o
 P

ro
vi

n
ce

, 
S

o
u
th

 A
fr

ic
a
. 
A

d
d
iti

o
n
a
l d

a
ta

 o
n
 g

u
in

e
a
-

fo
w

l n
e
m

a
to

d
e
s 

in
 s

o
u
th

e
rn

 A
fr

ic
a
  
fr

o
m

 v
a
ri
o
u
s 

a
u
th

o
rs

 a
re

 in
cl

u
d
e
d
 f
o
r 

co
m

p
a
ri
so

n

 N
e
m

a
to

d
e
s

T
h

is
 p

a
p

e
r

V
e
rs

te
r 

&
 P

ta
s
in

s
k
a
-K

lo
ry

g
a
 

(1
9
8
7
)

S
a
a
y
m

a
n

 (
1
9
6
6
)

C
ro

w
e
 

(1
9
7
7
)

O
rt

le
p

p
 

(1
9
3
7
, 

1
9
3
8
b

, 
1
9
6
4
)b

S
it

e
P

re
v
a
le

n
c
e
 

(%
)

In
te

n
s
it

y
P

re
v
a
le

n
c
e
 

(%
)

In
te

n
s
it

y
P

re
v
a
le

n
c
e
 

(%
)

In
te

n
s
it

y
P

re
s
e
n

c
e

P
re

s
e
n

c
e

M
e
a
n

 (
±

 S
D

)
R

a
n

g
e

M
e
a
n

R
a
n

g
e

M
e
a
n

R
a
n

g
e

A
sc

a
ri
d
ia

 g
a
lli

A
sc

a
ri
d
ia

 n
u
m

id
a
e

C
yr

n
e
a
 p

a
rr

o
ti

D
is

p
h
a
ry

n
x 

n
a
su

ta

G
o
n
g
yl

o
n
e
m

a
 c

o
n
g
o
le

n
se

G
o
n
g
yl

o
n
e
m

a
 in

g
lu

vi
co

la

H
a
d
je

lia
 in

e
rm

is

H
a
d
je

lia
 t
ru

n
ca

ta

H
e
te

ra
ki

s 
g
a
lli

n
a
ru

m

S
ic

a
ri
u
s 

ca
u
d
a
tu

s 

S
u
b
u
lu

ra
 d

e
n
tig

e
ra

S
u
b
u
lu

ra
 s

u
ct

o
ri
a

S
u
b
u
lu

ra
 s

p
.

U
n
id

e
n
tif

ie
d
 s

u
b
u
lu

ri
d

T
e
tr

a
m

e
re

s 
n
u
m

id
a

S
I

S
I

G
iz

P
ro

v

C
ro

p

C
ro

p

G
iz

G
iz

C
a
e
ca

G
iz

, 
S

I

C
a
e
ca

C
a
e
ca

C
a
e
ca

S
I

P
ro

v

– 6 1
0
0

– 4
0

– – 5
3

– 5
3

5
3

1
0
0

4
0

1
3

3
3

– 4
.0

a

1
3
.8

 (
±

 1
8
.2

)

– 2
3
.0

 (
±

 2
2
.0

)

– – 1
.6

 (
±

 0
.5

)

– 2
.1

 (
±

 1
.7

)

1
5
.9

 (
±

 1
3
.4

)

5
3
6
.3

 (
±

 5
8
9
.2

)

4
4
.0

 (
±

 6
5
.4

)

2
.5

 (
±

 0
.7

)

2
.4

 (
±

 1
.7

)

– 4
a

2
–
7
5

– 2
–
6
1

– – 1
–
2

– 1
–
6

1
–
3
1

9
–
2
 2

1
4

1
–
1
7
0

2
–
3

1
–
5

2 1
3

1
3

1
0

– – – – 4 – 6 2
3

1
0

– –

<
 1

<
 1

<
 1

1
.8

– – – – <
 1

– 1
.3

<
 1

<
 1

– –

0
–
2

0
–
1
9

0
–
1
6

0
–
5
9

– – – – 0
–
2

– 0
–
5
4

0
–
4
0

0
–
4

– –

6
4

– – – – – – – ? – – – – – –

5
.4

– – – – – – – 1
4
8

– – – – – –

?
–
9

– – – – – – – ?
–
2
5
7

– – – – – –

– – – – – – – – – – + + – – –

– + + – – + + – + – + + – – –

a
 

O
n
ly

 a
 s

in
g
le

 h
o
st

 h
a
rb

o
u
re

d
 t
h
is

 p
a
ra

si
te

b
 

In
cl

u
d
in

g
 u

n
p
u
b
lis

h
e
d
 r

e
co

rd
s 

o
f 
O

rt
le

p
p
 c

ite
d
 in

 V
e
rs

te
r 

&
 P

ta
si

n
sk

a
-K

lo
ry

g
a
 (

1
9
8
7
)

S
I 

=
 s

m
a
ll 

in
te

st
in

e
G

iz
 

=
 g

iz
za

rd
P

ro
v 

=
 p

ro
ve

n
tr

ic
u
lu

s



269

K. JUNKER & J. BOOMKER

S. dentigera, the ratio ranging from 4.5:1 to 53:1. In 
the remaining hosts only S. suctoria was present 
(Fig. 2E, F).

The Crested guineafowl harboured a single acan-
thocephalan species, Mediorhynchus gallinarum (n 
= 48), five species of cestodes, namely Abuladzugnia 

gutterae (n = 1), N. numida (n = 114), O. numida (n 
= 57), O. multiuncinata (n = 1) and P. paronai (n = 
52), as well as three species of nematodes, S. suc-
toria (n = 260), Gongylonema congolense (n = 56) 
and Hadjelia truncata (n = 2), representing a total of 
591 helminths.

FIG. 1 A, B. Cyrnea parroti male. A. Anterior end. B. Posterior end. C, D. Gongylonema congolense. C. Anterior extremity of female, 
ventral view. The arrow points to the excretory pore. D. Posterior extremity of male. The inset illustrates the barbed tip of the 
long spicule. E, F. Hadjelia truncata male. E. Ventral view of anterior extremity. F. Lateral view of anterior extremity

FE

DC

BA



270

Helminths of guineafowls in Limpopo Province, South Africa

Our finding of M. gallinarum, A. gutterae, H. multiunci-
nata, H. truncata and Sicarius caudatus in Hel meted 
guineafowls in South Africa constitutes new host 
associations, as well as new geographic records for 
these parasites. Dicrocoelium macrostomum, G. 
congolense and Davainea nana are recorded in 

South Africa for the first time, and the Crested guin-
eafowl is a new host for the nematodes S. suctoria, 
G. congolense and H. truncata.

Despite the generally high helminth burdens, the 
Helmeted guineafowls were in good physical condi-

FIG. 2 A, B, C. Sicarius caudatus. A. Anterior extremity of male. The deirids are marked by arrows. B. Posterior extremity of female. 
Note the finger-like protruberances (arrow) at the tip of the tail. C. Posterior extremity of male. D. Distal part of guineafowl 
caecum filled with Subulura spp. E. Subulura dentigera female, anterior part. The arrow indicates the cuticular denticles as 
described by Ortlepp (1937); x 400. F. Subulura suctoria female, anterior part; x 400

FE

DC

BA



271

K. JUNKER & J. BOOMKER

tion and no obvious lesions were associated with 
the presence of helminths. The crop mucosa of a 
single bird from Musina had an inflamed appear-
ance. This, however, did not seem to be related to 
G. congolense living in shallow tunnels under the 
crop lining, but rather to the presence of numerous 
thorny seeds of Tribulus terrestris.

TAXONOMIC REMARKS

Cyrnea parroti Seurat, 1917 (Table 3; Fig. 1A, B)

Ortlepp (1938b) described Habronema numidae 
from Helmeted guineafowls in Malawi, South Africa 
and Swaziland. This nematode has subsequently 
been included in the genus Cyrnea Seurat, 1914, 
but it is still listed under its original name in Yamaguti 
(1961) as well as in Verster & Ptasinska-Kloryga 
(1987).

In his work on the Habronematinae, Chabaud (1958) 
divided the genus Cyrnea into two subgenera, Pro-
cyrnea Chabaud, 1958 and Cyrnea Chabaud, 1958, 
which he later raised to genus level (Chabaud 1975). 
Following an in-depth study of the cephalic struc-
tures, he synonymized Cyrnea (Cyrnea) numidae 
(Ortlepp, 1938) with Cyrnea (Cyrnea) parroti Seurat, 
1917 (Chabaud 1958).

Specimens from our hosts mounted en face show 
the same arrangement of submedian lobes and sim-
ple lips as illustrated for C. parroti by Chabaud (1958) 
and otherwise conform well with the description and 
measurements supplied by Ortlepp (1938b) for C. 
numidae. The range of measurements in our speci-
mens was, however, generally wider than that pro-
vided by the latter author (Table 3). Ortlepp (1938b) 
himself stated that his new species most closely re-
sembled Cyrnea eurycerca and C. parroti and that 
the arrangement of the caudal papillae in the males 
as well as the spicules were very similar.

Gongylonema congolense Fain, 1955 (Table 4; 
Fig. 1C, D)

This parasite was first described by Fain (1955a) 
from domestic chickens, a single duck, Cairina mo-
schata domestica and from N. meleagris from the 
Democratic Republic of the Congo and Rwanda. 
Sub sequently it has been recorded from N. melea-
gris in Burundi, Nigeria, Ethiopia, Ghana and Burkina 
Faso (Fain & Thienpont 1958; Fabiyi 1972; Graber 
1976; Hodasi 1976; Vercruysse, Harris, Bray, Na-
galo, Pangui & Gibson 1985).

One of the main morphological characteristics of this 
species is the hook situated at a distance of about 

50 μm from the distal tip of the left spicule (Fain 
1955a) (Fig. 1D). The hook itself carries three fine 
barbs. In our specimens the barbed hook of the tip 
of the left spicule was often difficult to see, but in 
specimens where the distance could be determined 
it varied from 31 to 46 μm.

It is not always easy to judge whether the left spi-
cule is intact or damaged, which could lead to meas-
uring errors. There are, however, sufficient other 
characteristics, such as the gubernaculum, the ex-
tent and arrangement of the cuticular plaques (Fig. 
1C), as well as the length of the right spicule to dif-
ferentiate G. congolense from other species utiliz-
ing avian hosts (Fain 1955a).

While our specimens fit in well with Fain’s (1955a, 
b) description of G. congolense, we have not been 
able to confirm that the excretory pore opens on a 
transversally elongated plaque as was described by 
him. In our specimens it would seem that the two 
median ventral longitudinal rows of plaques are in-
terrupted, leaving a plaque-free zone immediately 
anterior and posterior to the excretory pore (Fig. 
1C).

Measurements of our specimens and those of Fain 
(1955a) taken from guineafowl hosts are presented 
in Table 4. These indicate that there is little geo-
graphic variation in the morphology of G. congo-
lense from the same host species.

Hadjelia truncata (Creplin, 1825) (Table 5; Fig. 
1E, F)

The most obvious differences between H. truncata 
and sympatric specimens of C. parroti are the posi-
tion of the vulva and the winged appearance of the 
lips of H. truncata in ventral view (Fig. 1E, F). In H. 
truncata the vulva is distinctly anterior and posi-
tioned in front of the posterior end of the oesopha-
gus. These characteristics are in accordance with 
the generic diagnosis of Hadjelia provided by Yama-
guti (1961).

Measurements of the specimens from the guinea-
fowls fall well within the range of measurements pro-
vided by Ortlepp (1964) for Hadjelia inermis (Ge-
doelst, 1919) (Table 5). Hadjelia inermis had been 
synonymized with H. truncata by Chabaud & Cam-
pana (1950), and Ortlepp (1964) commented on 
this, but chose to retain the former species. He lists 
his own measurements for H. inermis collected from 
Red- and Yellow-billed hornbills from South Africa, 
together with measurements for H. inermis taken 
from Gedoelst (1919) and for H. inermis and H. trun-
cata as provided by Cram (1927, cited by Ortlepp 



272

Helminths of guineafowls in Limpopo Province, South Africa

T
A

B
L
E

 3
 

T
h
e
 m

a
in

 m
o
rp

h
o
lo

g
ic

a
l c

ri
te

ri
a
 o

f 
C

yr
n
e
a
 p

a
rr

o
ti 

S
e
u
ra

t,
 1

9
1
7
 f

ro
m

 H
e
lm

e
te

d
 g

u
in

e
a
fo

w
ls

. 
T

h
e
 r

a
n
g
e
 o

f 
m

e
a
su

re
m

e
n
ts

 is
 p

ro
vi

d
e
d
. 

A
ll 

m
e
a
su

re
m

e
n
ts

 in
 m

ic
ro

m
e
tr

e
s 

u
n
le

ss
 

o
th

e
rw

is
e
 s

ta
te

d

S
o

u
rc

e
P

re
s
e
n

t 
s
tu

d
y

O
rt

le
p

p
 (

1
9
3
8
b

)

M
o

rp
h

o
lo

g
ic

a
l 
c
ri

te
ri

a
M

a
le

s
 (

n
 =

 6
)

F
e
m

a
le

s
 (

n
 =

 4
)

M
a
le

s
 

F
e
m

a
le

s

B
o
d
y 

le
n
g
th

 (
m

m
)

M
a
xi

m
u
m

 w
id

th

D
is

ta
n
ce

 a
p
e
x 

to
 n

e
rv

e
 r

in
g

D
is

ta
n
ce

 a
p
e
x 

to
 d

e
ir
id

s

D
is

ta
n
ce

 a
p
e
x 

to
 e

xc
re

to
ry

 p
o
re

D
e
p
th

 o
f 
b
u
cc

a
l c

a
p
su

le

W
id

th
 o

f 
b
u
cc

a
l c

a
p
su

le
 (

in
n
e
r)

M
u
sc

u
la

r 
o
e
so

p
h
a
g
u
s

G
la

n
d
u
la

r 
o
e
so

p
h
a
g
u
s

O
e
so

p
h
a
g
u
s 

to
ta

l l
e
n
g
th

L
e
n
g
th

 o
f 
ta

il

D
is

ta
n
ce

 v
u
lv

a
 t
o
 p

o
st

e
ri
o
r 

e
n
d

E
g
g
 le

n
g
th

 x
 e

g
g
 w

id
th

L
e
n
g
th

 o
f 
ri
g
h
t 
sp

ic
u
le

L
e
n
g
th

 o
f 
le

ft
 s

p
ic

u
le

L
e
n
g
th

 o
f 
g
u
b
e
rn

a
cu

lu
m

L
e
n
g
th

 o
f 
ca

u
d
a
l a

la
e

9
–
1
1

2
2
9
–
2
7
4

1
8
7
–
2
6
2

2
2
0
–
3
7
0

2
2
0
–
3
6
2

2
9
–
3
6

1
0
–
1
5

3
0
4
–
3
9
3

2
 2

3
4
–
2
 5

2
6

2
 2

8
4
–
2
 8

3
0

1
2
0
–
1
9
3

– – 4
1
0
–
5
1
0

a

8
3
4
–
1
 3

5
4

a

6
3
–
8
4

a

4
3
7
–
6
1
8

1
1
–
1
6

2
3
2
–
3
8
0

2
3
7
–
2
6
3

3
0
9
–
3
6
4

3
1
1
–
3
5
7

3
4
–
4
5

1
2
–
1
6

– – 2
 0

3
9
–
3
 0

5
6

1
2
8
–
1
5
0

6
6
1
–
8
9
7

4
5
–
4
6
 x

 2
5
–
2
7

– – – –

1
1
–
1
3

1
8
0
–
2
1
0

2
1
0
–
2
4
0

– 2
5
0
–
2
9
0

3
0

1
0

2
7
0
–
3
0
0

1
 7

0
0
–
2
 0

0
0

– – – – 4
2
0
–
4
3
8

1
 0

8
0
–
1
 1

1
0

7
0

4
2
0
–
5
2
0

1
8
–
1
9

3
0
0
–
3
6
0

2
1
0
–
2
4
0

– 2
5
0
–
2
9
0

3
6

1
2

3
3
0

2
 4

0
0
–
2
 6

0
0

– 1
2
6
–
1
3
0

~
 7

5
0

4
2
–
4
5
 x

 2
4

– – – –

a
  
  
 M

e
a
su

re
m

e
n
ts

 o
f 
th

e
 s

p
ic

u
le

s 
a
n
d
 t
h
e
 g

u
b
e
rn

a
cu

lu
m

 a
re

 d
e
ri
ve

d
 f
ro

m
 t
e
n
 m

a
le

s



273

K. JUNKER & J. BOOMKER

T
A

B
L
E

 4
 

T
h
e
 m

a
in

 m
o
rp

h
o
lo

g
ic

a
l c

ri
te

ri
a
 o

f 
G

o
n
g
yl

o
n
e
m

a
 c

o
n
g
o
le

n
se

 F
a
in

, 
1
9
5
5
 m

a
le

s 
fr

o
m

 H
e
lm

e
te

d
 g

u
in

e
a
fo

w
ls

  
fr

o
m

 S
o
u
th

 A
fr

ic
a
 (

p
re

se
n
t 
st

u
d
y,

 G
F

M
/N

 r
e
p
re

se
n
ts

 o
u
r 

sp
e
ci

m
e
n
 

n
u
m

b
e
r)

, 
a
n
d
 f
ro

m
 t
h
e
 D

e
m

o
cr

a
tic

 R
e
p
u
b
lic

 o
f 

th
e
 C

o
n
g
o
 a

n
d
 R

w
a
n
d
a
 (

F
a
in

 1
9
5
5
a
).

 A
ll 

m
e
a
su

re
m

e
n
ts

 in
 m

ic
ro

m
e
tr

e
s 

u
n
le

ss
 o

th
e
rw

is
e
 s

ta
te

d

S
o

u
rc

e
P

re
s
e
n

t 
s
tu

d
y

F
a
in

 (
1
9
5
5
b

)

M
o

rp
h

o
lo

g
ic

a
l 
c
ri

te
ri

a
G

F
M

3
/N

1
G

F
M

1
/N

1
6

G
F

M
1
/N

1
7

G
F

M
1
/N

2
1

G
F

M
1
/N

2
2

G
F

M
1
/N

2
3

G
F

M
1
/N

2
4

G
F

M
1
/N

2
5

(n
 =

 5
)

B
o
d
y 

le
n
g
th

 (
m

m
)

M
a
xi

m
u
m

 w
id

th

D
is

ta
n
ce

 a
p
e
x 

to
 d

e
ir
id

s

D
is

ta
n
ce

 a
p
e
x 

to
 n

e
rv

e
 r

in
g

D
is

ta
n
ce

 a
p
e
x 

to
 e

xc
re

to
ry

 p
o
re

D
is

ta
n
ce

 a
p
e
x 

to
 e

n
d
 o

f 
p
la

q
u
e
s

D
is

ta
n
ce

 a
p
e
x 

to
 c

e
rv

ic
a
l a

ile
s

D
e
p
th

 o
f 
b
u
cc

a
l c

a
p
su

le

M
u
sc

u
la

r 
o
e
so

p
h
a
g
u
s

G
la

n
d
u
la

r 
o
e
so

p
h
a
g
u
s

O
e
so

p
h
a
g
u
s 

to
ta

l l
e
n
g
th

L
e
n
g
th

 o
f 
ta

il

C
a
u
d
a
l a

la
e
 (

le
ft
; 
ri
g
h
t)

L
e
n
g
th

 o
f 
g
u
b
e
rn

a
cu

lu
m

L
e
n
g
th

 o
f 
ri
g
h
t 
sp

ic
u
le

L
e
n
g
th

 o
f 
le

ft
 s

p
ic

u
le

 (
m

m
)

1
7

2
6
6

8
4

2
1
0

3
0
5

4
4
0

1
0
2
; 
1
1
0

3
1

– – 4
 1

2
5

2
0
7

6
0
0
 (

le
ft
)

8
7

9
8

8
.7

1
5

2
2
8

7
0
; 

8
6

1
9
0

3
0
3

3
8
5

1
1
8

– 3
6
2

2
 9

6
7

3
 4

0
7

1
7
3

– 7
3

1
0
1

–

1
4

2
4
4

1
0
7

2
3
2

3
5
5

4
8
4

1
5
3

3
0

2
1
6

3
 9

3
4

4
 1

5
0

1
8
3

– 8
5

7
9

4
.8

1
3

2
1
5

8
8

1
8
8

3
2
2

4
7
0

1
0
7
; 
1
2
3

3
0

3
0
4

3
 0

7
5

3
 3

7
9

1
7
0

– 8
7

8
6

5
.5

– – 9
8

2
2
4

3
4
0

– – – 4
3
1

3
 8

7
9

4
 3

1
0

– – 8
2

9
9

7
.4

1
4

2
3
0

1
0
9
; 
1
0
4

2
2
3

3
6
2

4
8
6

1
5
0
; 
1
4
1

3
4

3
8
7

3
 4

1
7

3
 8

0
4

1
9
7

– – 9
8

5
.5

1
7

2
4
7

9
6
; 
8
3

1
9
1

3
3
7

5
0
0

1
2
5
; 
1
2
3

3
1

3
3
2

3
 5

2
3

3
 8

5
3

2
0
2

6
3
0
; 
5
8
3

8
0

1
0
0

8
.1

– 2
3
0

– – – – – – – – – 1
6
5

– 8
0

9
4

8
.2

1
2
–
2
4

1
7
0
–
2
0
0

8
5
–
1
2
5

1
9
6
–
2
3
5

3
1
0
–
3
5
0

4
5
0
–
4
7
5

1
2
5
–
1
7
5

3
0
–
4
5

2
9
0
–
4
0
0

2
 5

2
0
–
3
 9

2
0

– 1
8
5
–
2
0
0

5
7
5
–
7
0
0
; 
4
5
0
–
5
0
0

6
8
–
8
5

1
0
4
–
1
4
0

7
–
1
1



274

Helminths of guineafowls in Limpopo Province, South Africa

T
A

B
L
E

 5
 

T
h
e
 m

a
in

 m
o
rp

h
o
lo

g
ic

a
l c

ri
te

ri
a
 o

f 
H

a
d
je

lia
 t
ru

n
ca

ta
 (

C
re

p
lin

, 
1
8
2
5
) 

fr
o
m

 H
e
lm

e
te

d
 g

u
in

e
a
fo

w
ls

. 
T

h
e
 r

a
n
g
e
 o

f 
m

e
a
su

re
m

e
n
ts

 is
 p

ro
vi

d
e
d
. 
A

ll 
m

e
a
su

re
m

e
n
ts

 in
 m

ic
ro

m
e
tr

e
s 

u
n
-

le
ss

 o
th

e
rw

is
e
 s

ta
te

d

S
o

u
rc

e
P

re
s
e
n

t 
s
tu

d
y

G
e
d

o
e
ls

t 
(1

9
1
9
)

O
rt

le
p

p
 (

1
9
6
4
)

M
o

rp
h

o
lo

g
ic

a
l 
c
ri

te
ri

a
M

a
le

s
F

e
m

a
le

s
M

a
le

s
 

F
e
m

a
le

s
 

M
a
le

s
 

F
e
m

a
le

s
 

G
F

M
9
/1

G
F

M
/1

1
G

F
M

1
/1

0
G

F
M

1
/1

4
G

F
M

6
/1

B
o
d
y 

le
n
g
th

 (
m

m
)

M
a
xi

m
u
m

 w
id

th

D
is

ta
n
ce

 a
p
e
x 

to
 n

e
rv

e
 r

in
g

D
is

ta
n
ce

 a
p
e
x 

to
 d

e
ir
id

s

D
is

ta
n
ce

 a
p
e
x 

to
 e

xc
re

t.
 p

o
re

D
e
p
th

 o
f 
b
u
cc

a
l c

a
p
su

le

W
id

th
 o

f 
b
u
cc

a
l c

a
p
su

le
 (

in
n
e
r)

M
u
sc

u
la

r 
o
e
so

p
h
a
g
u
s

G
la

n
d
u
la

r 
o
e
so

p
h
a
g
u
s

O
e
so

p
h
a
g
u
s 

to
ta

l l
e
n
g
th

D
is

ta
n
ce

 a
p
e
x 

to
 v

u
lv

a

L
e
n
g
th

 o
f 
ta

il

E
g
g
 le

n
g
th

 x
 e

g
g
 w

id
th

L
e
n
g
th

 o
f 
le

ft
 s

p
ic

u
le

L
e
n
g
th

 o
f 
ri
g
h
t 
sp

ic
u
le

7 1
6
0

2
0
8

2
3
7
; 
2
3
9

2
7
5

4
4

5 3
6
9

1
 7

5
0

2
 1

1
9

– – – 1
 3

4
6

2
7
1

8 1
4
5

2
1
2

2
3
8
; 
2
3
1

2
5
9

4
2

7 3
9
7

1
 9

2
7

2
 3

2
4

– – – 1
 4

3
4

2
5
4

1
0

2
0
9

1
8
5

2
0
6
; 
2
0
4

2
3
4

3
9

5 3
5
8

1
 9

4
8

2
 3

0
6

1
 6

9
8

1
3
8

5
0
 x

 3
2

– –

– 2
1
7

1
5
9

1
6
0
; 
1
6
1

1
7
9

4
0

7 3
4
6

2
 0

7
6

2
 4

2
2

1
 6

9
1

– 5
3
 x

 3
5

– –

1
1

1
4
0

– 2
5
7
; 
2
5
9

2
9
0

4
1

6 4
9
5

1
 9

8
8

2
 4

8
3

2
 2

3
8

1
2
1

– – –

6
.1

–
6
.4

5

1
4
0
–
1
4
4

1
8
0
–
2
1
5

2
1
0
–
2
6
0

2
2
0
–
2
7
5

– – – – 2
 0

0
0

– 1
2
0

– 1
 6

0
0
–
1
 9

0
0

2
0
0

1
8
–
2
1
.8

2
4
0
–
2
6
0

2
6
0
–
2
7
5

3
3
0

3
6
0

– – – – 2
 4

0
0
–
3
 6

0
0

1
 8

6
0
–
2
 9

7
0

9
0
–
1
2
0

5
4
–
5
7
 x

 3
0
–
3
2

– –

6
–
7

– – – – 4
0
–
5
0

1
0

2
3
0
–
2
8
0

1
 9

0
0
–
2
 2

0
0

2
 1

3
0
–
2
 4

8
0

– 1
2
0
–
1
4
0

– 1
 2

0
0
–
1
 5

0
0

2
1
5
–
2
8
0

1
7
–
1
9

– – – – 4
7
–
5
2

1
2

4
0
0
–
4
5
0

2
 0

0
0
–
2
 3

0
0

2
 4

0
0
–
2
 7

5
0

2
 2

0
0
–
2
 5

0
0

1
1
0
–
1
2
0

3
2
–
3
7
 x

 2
5
–
2
7

– –



275

K. JUNKER & J. BOOMKER

T
A

B
L
E

 6
 

M
o
rp

h
o
lo

g
ic

a
l c

ri
te

ri
a
 o

f 
S

ic
a
ri
u
s 

d
ip

te
ru

m
 (

P
o
p
o
va

, 
1
9
2
7
),

 S
ic

a
ri
u
s 

h
o
o
p
o
e

  S
h
a
rm

a
, 
1
9
7
1
, 
S

ic
a
ri
u
s 

ca
u
d
a
tu

s 
Q

u
e
n
tin

 &
 W

e
rt

h
e
im

, 
1
9
7
5
 a

n
d
 S

ic
a
ri
u
s 

re
n
a
ta

e
 C

a
n
cr

in
i, 

B
a
lb

o
 

&
 I
o
ri
, 
1
9
9
1
 d

e
sc

ri
b
e
d
 f
ro

m
 a

vi
a
n
 h

o
st

s.
 A

ll 
m

e
a
su

re
m

e
n
ts

 in
 m

ic
ro

m
e
tr

e
s 

u
n
le

ss
 o

th
e
rw

is
e
 s

ta
te

d

S
o

u
rc

e
A

li
 (

1
9
6
1
)

 
S

h
a
rm

a
 (

1
9
7
1
)

Q
u

e
n

ti
n

 &
 W

e
rt

h
e
im

 
(1

9
7
5
)

P
re

s
e

n
t 

s
tu

d
y

C
a
n

c
ri

n
i,
 B

a
lb

o
 &

 I
o

ri
 

(1
9
9
1
)

M
o

rp
h

o
lo

g
ic

a
l 
c
ri

te
ri

a
S

ic
a
ri

u
s
 d

ip
te

ru
m

 
S

ic
a
ri

u
s
 h

o
o

p
o

e
S

ic
a
ri

u
s
 c

a
u

d
a
tu

s
S

ic
a
ri

u
s
 c

a
u

d
a
tu

s
S

ic
a
ri

u
s
 r

e
n

a
ta

e

M
a
le

F
e
m

a
le

M
a
le

F
e
m

a
le

M
a
le

F
e
m

a
le

M
a
le

F
e
m

a
le

M
a
le

F
e
m

a
le

B
o
d
y 

le
n
g
th

 (
m

m
)

1
0
.2

–
1
1
.9

1
2
.9

–
1
6
.1

6
.7

–
9
.4

1
1
.2

–
1
8
.2

7
.3

1
3
.3

5
.9

8
.3

4
.9

7
.3

D
is

ta
n
ce

 a
p
e
x 

to
 d

e
ir
id

s
7
0
–
7
4

a
8
0
–
9
0

a
6
0
–
7
0

5
0
–
6
0

8
5

8
5

6
3
; 
6
8

7
8
; 
8
7

6
8
; 
7
0

6
5
; 
7
5

D
is

ta
n
ce

 a
p
e
x 

to
 n

e
rv

e
 

ri
n
g

1
8
0
–
2
0
0

2
4
0
–
2
6
0

1
1
0
–
1
4
0

1
6
5
–
2
1
0

2
1
5

2
5
0

2
0
8

2
2
6
–
2
2
7

1
5
0

1
6
5

D
is

ta
n
ce

 a
p
e
x 

to
 e

xc
re

t.
 

p
o
re

2
4
0
–
2
6
0

2
8
0
–
3
2
0

1
3
0
–
1
8
0

1
8
0
–
2
2
0

2
7
0

3
1
0

–
2
7
8
–
2
9
8

–
–

D
e
p
th

 o
f 
b
u
cc

a
l c

a
p
su

le
4
3
–
4
5

5
2
–
5
8

1
4
–
1
7

2
5

2
8

3
8

2
8

3
1
–
3
3

2
5

2
5

M
u
sc

u
la

r 
o
e
so

p
h
a
g
u
s

5
1
0
–
5
3
0

5
6
0
–
6
1
0

2
4
0
–
3
2
0

3
2
0
–
3
8
0

3
8
0

4
1
0

–
–

2
5
0

2
3
6

G
la

n
d
u
la

r 
o
e
so

p
h
a
g
u
s

2
 8

8
0
–
2
 9

2
0

3
 1

6
0
–
3
 9

1
0

2
 8

0
0
–
3
 0

4
0

3
 2

0
0
–
3
 6

0
0

2
 1

7
0

2
 9

0
0

–
–

1
 9

5
0

2
 3

6
5

O
e
so

p
h
a
g
u
s 

to
ta

l 
le

n
g
th

3
 4

0
0
–
3
 5

0
0

3
 7

0
0
–
4
 5

0
0

–
–

2
 5

5
0

3
 3

1
0

2
 5

3
5

2
 7

2
2

2
 2

0
0

2
 6

0
1

L
e
n
g
th

 o
f 
ta

il
2
1
0

1
8
5
–
2
1
0

1
7
6
–
2
0
8

1
6
7
–
2
5
6

1
9
0

2
5
0

1
6
1

1
6
8

–
1
1
0

L
e
n
g
th

 o
f 
ri
g
h
t 
sp

ic
u
le

9
3
–
1
6
0

–
4
4
0
–
5
6
0

–
1
7
0
–
1
9
0

–
1
7
1

–
1
7
5

–

L
e
n
g
th

 o
f 
le

ft
 s

p
ic

u
le

6
2
0
–
6
9
0

–
4
7
0
–
6
0
0

–
4
0
0
–
4
5
0

–
4
1
3

–
3
6
0

–

D
is

ta
n
ce

 v
u
lv

a
 t
o
 t
ip

 o
f 

ta
il

–
5
 0

0
0

–
–

–
4
 9

5
0

–
2
 9

6
0

–
2
 8

0
0

E
g
g
 le

n
g
th

–
3
8
–
4
0

–
3
3
–
4
6

–
4
3

–
3
8

–
3
7
–
4
0

E
g
g
 w

id
th

–
3
0
–
3
7

–
2
9
–
3
9

–
3
0

–
2
9

–
2
5
–
2
7

M
a
x.

 w
id

th
 o

f 
a
la

e
–

–
–

–
4
5

4
5

3
6

4
0
–
5
0

–
–

E
xt

e
n
si

o
n
 a

la
e

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

W
h
o
le

 b
o
d
y

a
  
  
 C

e
rv

ic
a
l p

a
p
ill

a
e
 in

 A
li 

(1
9
6
1
) 

in
te

rp
re

te
d
 h

e
re

 a
s 

d
e
ir
id

s



276

Helminths of guineafowls in Limpopo Province, South Africa

1964) (Table 5). As we have not been able to exam-
ine the type specimens of either species, we have 
chosen to adopt the conclusion of the in-depth mor-
phological study of Chabaud & Campana (1950).

The single complete specimen in our collection ap-
pears slightly shorter than previously described 
ones. The depth of the buccal cavity of our speci-
mens corresponds with the lower range of the pha-
rynx sizes provided by Ortlepp (1964) and all three 
authors he quoted.

The oesophagus length is relatively uniform in all the 
sources quoted by Ortlepp (1964). In our specimens 
the total length of the oesophagus only reached 2.1 
and 1.9 mm which, considering that these speci-
mens are short, does not seem extraordinary. The 
egg size is very similar to that of H. inermis as re-
corded by Gedoelst (1919) and Cram (1927, cited 
by Ortlepp 1964), but larger than given by Ortlepp 
(1964). Ortlepp (1964) pointed out that this was the 
only noteworthy difference between his specimens 
and those described by Gedoelst (1919).

According to Chabaud (1958) the genus Hadjelia 
has been described from numerous birds, especial-
ly Coraciiformes, from Europe, Asia and Africa. Data 
pertaining to African hosts mainly list Bucerotiformes 
(Ortlepp 1964) and we are aware of only one refer-
ence to Hadjelia from galliform birds, namely Had-
jelia lhuillieri Seurat, 1916 from Alectoris barbara (= 
Caccabis petrosa from Algeria in Seurat 1916). Inci-
dentally, Chabaud (1958) speculated that the latter 
species might be identical with H. truncata, but 
thought that the characteristics of the single known 
female specimen of H. lhuillieri were not sufficient to 
draw a final conclusion.

Sicarius caudatus Quentin & Wertheim, 1975 
(Table 6; Fig. 2A, B, C)

Four species of the genus Sicarius are known from 
avian hosts, namely Sicarius dipterum (Popova, 
1927), Sicarius hoopoe Sharma, 1971, Sicarius cau-
datus Quentin & Wertheim, 1975 and Sicarius rena-
tae Cancrini, Balbo & Iori, 1991. The left spicule of 
S. dipterum is distinctly longer (660–670 μm) than 
that of our specimens, whereas the subequal spi-
cules of S. hoopoe are 440–560 and 470–600 μm in 
length (Cancrini, Balbo & Iori 1991). Our specimens 
best fit the description of S. caudatus, as they have 
six pairs of caudal papillae as opposed to the eight 
pairs of S. renatae (Cancrini et al. 1991).

According to Quentin & Wertheim (1975) the deirids 
in S. caudatus are situated at the origin of the lateral 

alae. In some of our specimens, we have observed 
the same arrangement, but in one male and one fe-
male the right and left deirids emerge 11 and 27 μm, 
and 17 and 37 μm anterior to the origin of the alae 
(Fig. 2A). We have too little material to comment on 
the significance of this observation.

Quentin & Wertheim (1975) describe the cuticular 
processes in the tail of S. caudatus as atrophied, 
the tail consisting merely of a smooth stump, which 
at best has rugged edges. Our specimens possess 
about seven distinct, albeit short, cuticular exten-
sions similar to those illustrated by Cancrini et al. 
(1991) for S. renatae (Fig. 2B). Despite these differ-
ences we have allocated our specimens to S. cau-
datus. Apart from the original description and their 
inclusion in some taxonomic reviews (Chabaud 
1958; Ali 1961), we have not found any other refer-
ences to S. caudatus in the literature. The measure-
ments of the specimens collected during this study 
are included in Table 6.

DISCUSSION

Despite the fact that various studies on the helminths 
of guineafowls in South Africa have been conducted, 
direct comparisons between the results of these 
studies are not always possible, as they had differ-
ent objectives. Ortlepp (1937, 1938a, b, 1963) stud-
ied the helminths of all the organs and the entire 
alimentary canal, but his work was of a taxonomic 
nature, based on incidental findings, and presented 
no epidemiological data. Crowe (1977) listed the 
helminth species recovered from the small intestine, 
caeca and rectum of guineafowls, but in his subse-
quent analysis grouped them as acanthocephalans, 
cestodes and nematodes respectively. The two 
studies providing data on the prevalence and inten-
sity of the helminths are those of Saayman (1966) 
and Verster & Ptasinska-Kloryga (1987). However, 
Saayman (1966) only examined the intestinal tract 
and Verster & Ptasinska-Kloryga (1987) collected 
helminths from the gizzard, intestine and caeca. 
Thus, their data on species richness would not re-
flect worms located in e.g. the crop or proventricu-
lus.

The study conducted on guineafowls in Burkina 
Faso by Vercruysse et al. (1985) lends itself best to 
comparison with ours, as they examined the com-
plete alimentary tract, including the crop and pro-
ventriculus. Of the total of 13 helminth species col-
lected by these authors, eight species coincide with 
species recovered from our hosts. If the single acan-
thocephalan present in the birds from Burkina Faso 



277

K. JUNKER & J. BOOMKER

is taken into account, this number will increase by 
one. Vercruysse et al. (1985) record the acantho-
cephalan Mediorhynchus selengensis, which has 
been synonymized with M. gallinarum by Schmidt & 
Kuntz (1977), and the nematodes Cyrnea parroti, S. 
suctoria, G. congolense and A. numidae, which are 
also recorded in this study. In addition to these spe-
cies, Vercruysse et al. (1985) recorded the cestode 
Cotugnia digonopora and the nematodes Eucoleus 
annulatus, T. fissispina and Dispharynx spiralis.

Nematodes

Cyrnea

With the exception of C. parroti, helminths were re-
covered from their usual predilection sites. According 
to Anderson (1992) members of the genus Cyrnea 
occur in the proventriculus of birds and he records 
Cyrnea colini in the wall of the proventriculus near 
the gizzard of Bobwhite quails. We did not recover 
C. parroti from the proventriculus, but in all infected 
guineafowls the parasites were situated under the 
gizzard lining and could only be seen after the horny 
layer had been removed. There seemed, however, 
to be a preference for the proventricular-gizzard 
isthmus as described for Cyrnea neeli from wild tur-
keys in the south-eastern United States (Davidson, 
Hon & Forrester 1977). Similarly, C. parroti recov-
ered from Helmeted guineafowls in Burkina Fasso 
were also present in the gizzard (Vercuysse et al. 
1985).

Subulura

The genus Subulura has a wide distribution in galli-
naceaous birds on the African continent and records 
exist from Zimbabwe, Tanzania, Ghana, Nigeria and 
Somalia (Nicholls, Bailey, Gibbons, Jones & Samour 
1995; Nfor, Ajanusi, Agbede & Esievo 1999; Poul-
sen, Permin, Hindsbo, Yelifari, Nansen & Bloch 
2000; Permin, Esmann, Hoj, Hove & Mukaratirwa 
2002; Magwisha, Kassuku, Kyvsgaard & Permin 
2002). However, the genus is not restricted to the 
African continent and, according to Yamaguti (1961) 
is a cosmopolitan species.

Ortlepp (1937) recovered S. suctoria in association 
with S. dentigera from guineafowls from various re-
gions in South Africa and Swaziland and concluded 
that the two species had a wide distribution. Contrary 
to our findings, he found S. dentigera to be far more 
abundant than S. suctoria.

Verster & Ptasinska-Kloryga (1987) collected hel-
minths from 48 guineafowls in the vicinity of Pretoria. 

Subulura suctoria was present in 11 and S. denti g-
era in three of the hosts examined. From these and 
our own results it is apparent that the two species, 
S. suctoria and S. dentigera often share the same 
habitat. It is difficult to judge from our data whether 
these two species are interactive and compete for 
the same resources. If so, S. suctoria would seem 
the stronger competitor as it consistently occurred 
in higher numbers than S. dentigera. However, the 
numbers of S. dentigera were not greater in hosts 
with relatively low burdens of S. suctoria, but rather 
the numbers of S. dentigera were low in these hosts 
as well. It is possible, that this association is similar 
to the major-minor species concept, as seen with 
Theladorsagia circumcincta and Theladorsagia dav-
tiani in sheep and goats.

A literature study confirms the dominance of S. suc-
toria in guineafowls and Vercruysse et al. (1985) 
recorded a 100 % prevalence of S. suctoria from 
103 Helmeted guineafowls in Burkina Faso. In addi-
tion to being the most prevalent nematode, these 
authors also found S. suctoria to be one of the most 
numerous parasites (26–1 071 worms per host). 
Subulura dentigera was not reported from these 
hosts.

Ascaridia numidae

Ascaridia numidae is another nematode commonly 
encountered in Helmeted guineafowls and has been 
recorded from various geographic localities in Africa. 
The prevalence and intensity of this parasite varies 
greatly from 98.1 % with a range of intensity from 1 
to 1 452 in hosts in Burkina Faso (Vercruysse et al. 
1985) and 86.7 % with intensities ranging from 1 to 
504 in birds in Ghana (Hodasi 1976) to a low preva-
lence of 13 % with a maximum of 19 worms per host 
in South Africa (Verster & Ptasinska-Kloryga 1987). 
In the present study A. numidae was present in a 
single host only.

Gongylonema

Both Hodasi (1976) and Vercruysse et al. (1985), 
record G. congolense from hosts they examined, 
with a prevalence of 48.9 and 73.8 %, respectively. 
This indicates that G. congolense not only forms a 
regular part of the helminth community of guinea-
fowls in South Africa, but throughout the African con-
tinent. With the exception of Gongylonema ingluvi-
cola allegedly recorded by Ortlepp (“1937, 1938, 
unpublished records” cited by Verster & Ptasinska-
Kloryga 1987), the absence of this genus in previ-
ous reports on helminths of guineafowls in South 



278

Helminths of guineafowls in Limpopo Province, South Africa

Africa, is most likely due to the fact that earlier au-
thors did not examine the crop of the hosts in their 
studies.

Tetrameres

While Tetrameres numida was recovered in low num-
bers from the Musina guineafowls, none of the more 
com monly reported species of this genus was pres-
ent in our material. A second species, which has 
been recorded from guineafowls and is also a com-
mon para site of domestic chickens, is Tetrameres 
fissispina Diesing, 1861. Vercruysse et al. (1985) 
re port a 48.5 % prevalence and an intensity of infec-
tion ranging from 1 to 146 worms per host from Hel-
meted guineafowls in Burkina Faso, and 23.3 % of 
126 Helmeted guineafowls in Nigeria were infected 
with T. fissispina (Fabiyi 1972). In Ghana the preva-
lence of infection in the same host was 8.9 % with a 
mean worm burden of 2.8, ranging from one to eight. 
Young scavenging chickens in Ghana had a preva-
lence of T. fissispina of 58 % (Poulsen et al. 2000).

We are aware of a single record of three females of 
T. fissispina from a single Helmeted guineafowl in 
South Africa (Le Roux 1926), and the same author 
reports a high percentage of infection (78 %) in 60 
domestic chickens in the same country. The prov-
entriculus of a single, heavily infected host contained 
a minimum of 150 females (Le Roux 1926).

A third species commonly infecting domestic chick-
ens, namely Tetrameres americana, which had a 60 
and 62 % prevalence in adult chickens in Tanzania 
and Zimbabwe, respectively (Per min, Magwisha, 
Kassuku, Nansen, Bisgaard, Frand sen & Gibbons 
1997; Permin et al. 2002), has not yet been recorded 
from guineafowls.

From the literature cited above it would appear that 
the prevalence of the genus Tetrameres is slightly 
higher in domestic chickens than in Helmeted guin-
eafowls. Since the data above concerning the do-
mestic chickens above pertain to free-ranging or 
scavenging chickens, guineafowls and domestic 
fowls probably had an equal chance of exposure to 
the parasite. Whether the higher infection rates in 
chickens are a result of higher host densities or 
whether guineafowls are generally more resistant 
towards helminth infections remains speculation.

Trematodes

The literature contains few reports of trematodes 
from guineafowl hosts, but a number of trematodes 
have been listed as parasitizing not only the diges-

tive tract and urogenital system, but also the respi-
ratory system of domestic fowls (Soulsby 1968). To 
our knowledge the only published records of trema-
todes from the liver of guineafowls are those of D. 
macrostomum, that occurs in the gall bladder and 
bile ducts of N. meleagris (= N. ptilorhyncha) in 
Egypt (Lesbouyries 1941) and Lutztrema sp. from 
the gallbladder of Helmeted guineafowls in Ghana 
(Hodasi 1976). The former parasite has also been 
found in the liver of Helmeted guineafowls in the 
Kruger National Park (Horak 2007, personal com-
munication) and was present in the Musina hosts. 
The results of this study and unpublished data of 
Horak (2007) suggest that D. macrostomum is not 
uncommon in South African guineafowls and can 
reach high intensities in individual hosts.

Postharmostomum gallinum has been reported from 
the gastro-intestinal tract from Crested guineafowls 
in Pakistan (Khan, Khan & Rayaz 1984). Hodasi 
(1969, 1976) collected Postharmostomum ntowi 
and Episthmium ghanense and Episthmium africa-
nus from the gastro-intestinal tracts of domestic 
chickens. Intensities and prevalences were low and 
the latter author concluded that trematodes were 
rare parasites in gallinaceous birds.

The fact that the intermediate hosts of trematodes 
are mainly molluscs or rarely annelids (Gibbons, 
Jones & Khalil 1996), both of which are typically as-
sociated with moist environmental conditions, might 
well explain why trematodes played a minor role as 
parasites of the guineafowls in our dry study area.

Cestodes

Porogynia

The presence of young stages of Porogynia paronai 
in the liver of infected hosts is unusual. Hodasi 
(1976), however, recovered adult Cotugnia melea-
gridis from the small intestine of Helmeted guinea-
fowls in Ghana, and recorded numerous young 
forms of this parasite from the host’s gallbladder. 
Since the life cycle of Porogynia is not known, one 
can only speculate on the presence of immatures in 
the liver.

During the normal course of cestode development 
in avian hosts, the cysticercoid is freed from the ar-
thropod intermediate host in the intestine as a result 
of mechanical and chemical actions. Subsequently, 
the scolex evaginates and the cysticercoid attaches 
itself to the gut wall (Reid 1962). The fact that young 
P. paronai were recovered from the liver of three 
birds and in relatively high numbers, in addition to 



279

K. JUNKER & J. BOOMKER

their uniform stage of development, suggests that 
their presence is not a result of post-mortem migra-
tion. Whether the newly freed cysticercoid, assum-
ing that an arthropod is the intermediate host, mi-
grates up the common bile duct to mature to a 
certain stage, before leaving the liver to resume its 
final maturation in the small intestine, or whether we 
have observed aberrant migration of juvenile stages 
will remain speculation until the development of P. 
paronai can be studied in more detail.

Abuladzugnia

Interestingly, the cestode A. gutterae, which was 
com mon in the guineafowls examined by us was not 
found in any of the previous surveys. Ortlepp (1963) 
originally described this species as Cotugnia gut-
terae from three specimens that had been collected 
from Crested guineafowls in Mozambique. Since 
then there seem to have been no further records of 
this parasite. Spasskii (1973) created the genus Abu-
ladzugnia to accommodate A. gutterae and another 
of Ortlepp’s (1963) species formerly described as 
Cotugnia transvaalensis.

Conclusion

The above findings suggest, that despite geograph-
ical variation in the prevalence and intensity of indi-
vidual helminth species, probably caused by envi-
ronmental conditions, such as temperature, rainfall 
and soil conditions, the helminth community of guin-
eafowls in Africa is composed of a relatively stable 
body of core and secondary species enriched by 
satellite species. The latter probably depend on lo-
cal conditions and can be influenced by abiotic con-
ditions, but also the presence or absence of certain 
intermediate hosts and other terrestrial birds which 
may serve as reservoir hosts for certain parasites. 
We interpret the relative uniformity in the helminth 
community of Helmeted guineafowls in Africa as 
flowing from a long host/parasite association during 
which parasites have spread in conjunction with 
their hosts.

ACKNOWLEDGEMENTS

We thank Dr W.J. Luus-Powell, University of Lim po-
po, who enabled the collection of Helmeted guinea-
fowls in Musina and Mr H.E. Hattingh, University of 
Limpopo, for collecting and placing the Mokopane 
material at our disposal as well as for collecting the 
birds at Musina. Mr K. Meyer and Mr M. Storm kind-
ly made the guineafowls available to us. The techni-

cal assistance of Mr. Ryno Watermeyer, University 
of Pretoria, especially with the processing of the ces-
todes, is greatly appreciated. This study was made 
possible by a Claude Leon Foundation Postdoctoral 
Fellowship grant to the first author.

REFERENCES

ACOCKS, J.P.H. 1988. Veld types of South Africa, with accom-
panying veld type map, 3rd ed. (Memoirs of the Botanical 
Survey of South Africa, no. 57).

ALI, S.M. 1961. On some new nematodes (Habronematinae) 
from birds in Hyderabad, India, and the relationships of the 
genus Habronema. Journal of Helminthology, 35:1–48.

ANDERSON, R.C. 1992. Nematode parasites of vertebrates, 
their development and transmission, 1st ed. Wallingford and 
New York: CAB International.

CANCRINI, G., BALBO, T. & IORI, A. 1991. Su un nuovo Habro-
nematidae parassita di Acryllium vulturinum: Sicarius renatae 
sp. n. Parassitologia (Roma), 33:133–136.

CHABAUD, A.G. 1958. Essai de classification des nematodes 
Habronematinae. Annales de Parasitologie, 33:445–508.

CHABAUD, A.G. 1975. Keys to genera of the order Spirurida, 
Part 2. Spiruroidea, Habronematoidea and Acuarioidea, in 
CIH keys to the nematode parasites of vertebrates, edited by 
R.C. Anderson, A.G. Chabaud & S. Willmott. Farnham Royal: 
Commonwealth Agricultural Bureaux.

CHABAUD, A.G. & CAMPANA, Y. 1950. Notes sur le genre Had-
jelia Seurat, 1916 (Nématodes-Spiruridae). Annales de Para-
sitologie Humaine et Comparée, 25:435–440.

CROWE, T.M. 1977. Variation in intestinal helminth infestation of 
the Helmeted guinea-fowl. South African Journal of Wildlife 
Research, 7:1–3.

DAVIDSON, W.R., HON, L.T. & FORRESTER, D.J. 1977. Status 
of the genus Cyrnea (Nematoda: Spiruroidea) in wild turkeys 
from the Southeastern United States. Journal of Parasitology, 
63:332–336.

DEL HOYO, J., ELLIOT, A. & SARGATAL, J. (Eds). 1994. Hand-
book of the birds of the world. Vol. 2, New World vultures to 
guineafowl. Barcelona: Lynx Edicions.

FABIYI, J.P. 1972. Studies on parasites of the grey-breasted hel-
met guineafowl (Numida meleagris galeata Pallas) of the 
Vom area of the Benue Plateau State, Nigeria. I. Helminth 
parasites. Bulletin of Epizootic Diseases of Africa, 20:235–
238.

FAIN, A. 1955a. Sur un nouveau Gongyloneme, G. congolense 
n. sp., parasite de la Poule, du Canard et des Gallinacés 
sauvages. Revue de Zoologie et de Botanique Africaines, 
51:1–10.

FAIN, A. 1955b. Le genre Gongylonema Molin 1857, au Congo 
Belge et au Ruanda-Urundi. Annales de Parasitologie Hu-
maine et Comparée, 30:202–218.

FAIN, A. & THIENPONT, D. 1958. Notes sur deux Gongylonèmes 
congolais parasites de l’Oryctérope et de la Pintade. Revue 
de Zoologie et de Botanique Africaines, 57:113–116.

GEDOELST, L. 1919. Un genre nouveau de Spiruridae. Comptes 
Rendus des Séances de la Société de Biologie, 82:1145–
1148.

GIBBONS, L.M., JONES, A. & KHALIL, L.F. 1996. Eighth inter-
national training course on identification of helminth parasites 
of economic importance. St. Albans: International Institute of 
Parasitology, Commonwealth Agricultural Bureaux.



280

Helminths of guineafowls in Limpopo Province, South Africa

GRABER, M. 1976. Helminthes et helminthiases des animaux 
domestiques et sauvages d’Ethiopie. Tome 2. Deuxieme Mis-
sion des Vétérinaires Français en Ethiopie, Janvier-Mars ’75.

HOCKEY, P.A.R., DEAN, W.R.J. & RYAN, P.G. (Eds). 2005. 
Roberts—Birds of Southern Africa, VIIth ed. Cape Town: The 
Trustees of the John Voelcker Bird Book Fund.

HODASI, J.K.M. 1969. Comparative studies on the helminth 
fauna of native and introduced domestic fowls in Ghana. 
Jour nal of Helminthology, 43:35–52.

HODASI, J.K.M. 1976. The helminth parasites of the helmet 
guinea fowl (Numida meleagris galeata Pallas) in Ghana. 
Bulletin of Animal Health and Production in Africa, 24:81–
87.

KHAN, S.W., KHAN, A.J. & RAYAZ, S. 1984. Helminth parasites 
found in the gastro-intestinal tract of guinea fowl, Guttera 
pucherani of Peshawar, N.W.F.P., Pakistan. Bulletin of Zool-
ogy, University of Peshawar, 2:57–59.

LEPAGE, D. 2007. Avibase, hosted by Bird Studies Canada, the 
Canadian copartner of Birdlife International.

 www.bsc-eoc.org/avibase/avibase.jsp?pg

LE ROUX, P.L. 1926. Helminths collected from the domestic fowl 
(Gallus domesticus) and the domestic pigeon (Columba livia) 
in Natal. 11th and 12th Reports of the Director of Veterinary 
Edu cation and Research, Part 1: 207–217.

LESBOUYRIES, G. 1941. La pathologie des oiseaux. Paris: Vigot 
Frères.

MAGWISHA, H.B., KASSUKU, A.A., KYVSGAARD, N.C. & PER-
MIN, A. 2002. A comparison of the prevalence and burdens 
of helminth infections in growers and adult free-ranging chick-
ens. Tropical Animal Health and Production, 43:205–214.

MARGOLIS, L., ESCH, G.W., HOLMES, J.C., KURIS, A.M. & 
SCHAD, G.A. 1982. The use of ecological terms in parasitol-
ogy (report of an ad hoc committee of the American Society 
of Parasitologists). Journal of Parasitology, 56:436–439.

NFOR, M.B., AJANUSI, O.J., AGBEDE, R.I.S. & ESIEVO, K.A.N. 
1999. Prevalence of parasites of guinea fowl (Numida me-
leagris galeata) in Zaria, Nigeria. Bulletin of Animal Health 
and Production in Africa, 47:103–106.

NICHOLLS, P.K., BAILEY, T.A., GIBBONS, L.M., JONES, A. & 
SAMOUR, J.H. 1995. Parasitic infection in a flock of rufous-
crested bustards (Eupodotis ruficrista) in the United Arab 
Emirates. Journal of Zoo and Wildlife Medicine, 26:590–596.

ORTLEPP, R.J. 1937. South African helminths, Part I. Onderste-
poort Journal of Veterinary Science and Animal Industry, 9: 
311–336.

ORTLEPP, R.J. 1938a. South African helminths, Part III. Some 
mammalian and avian cestodes. Onderstepoort Journal of 
Veterinary Science and Animal Industry, 11:23–50.

ORTLEPP, R.J. 1938b. South African helminths, Part V. Some 
avian and mammalian helminths. Onderstepoort Journal of 
Veterinary Science and Animal Industry, 11:63–104.

ORTLEPP, R.J. 1963. Observations on cestode parasites of 
guinea-fowl from Southern Africa. Onderstepoort Journal of 
Veterinary Research, 30:95–118.

ORTLEPP, R.J. 1964. Some helminths recovered from Red- and 
Yellow-billed Hornbills from the Kruger National Park. Onder-
stepoort Journal of Veterinary Research, 31:39–52.

PERMIN, A., ESMANN, J.B., HOJ, C.H., HOVE, T. & MUKARA-
TIRWA, S. 2002. Ecto-, endo- and haemoparasites in free-
ranging chickens in the Goromonzi District in Zimbabwe. Pre-
ventive Veterinary Medicine, 54:213–224.

PERMIN, A., MAGWISHA, H., KASSUKU, A.A., NANSEN, P., 
BISGAARD, M., FRANDSEN, F. & GIBBONS, L.M. 1997. A 
cross-sectional study of helminths in rural scavenging poultry 
in Tanzania in relation to season and climate. Journal of Hel-
minthology, 71:233–240.

POULSEN, J., PERMIN, A., HINDSBO, O., YELIFARI, L., NAN-
SEN, P. & BLOCH, P. 2000. Prevalence and distribution of 
gastro-intestinal helminths and haemoparasites in young 
scavenging chickens in upper eastern region of Ghana, Afri-
ca. Preventive Veterinary Medicine, 45:237–245.

QUENTIN, J.C. & WERTHEIM, G. 1975. Helminthes d’oiseaux et 
de mammifères d’Israël V. – Spirurides nouveaux ou peu con-
nus. Annales de Parasitologie (Paris), 50:63–85.

REID, W.M. 1962. Chicken and turkey tapeworms. Handbook to 
aid in the identification and control of tapeworms found in the 
United States of America. Poultry Department and Poultry 
Disease Research Center, College Experiment Station, 
Athens, Georgia.

SAAYMAN, J. 1966. A study of the diet and parasites of Ardeola 
(Bubulcus) ibis, Numida meleagris and Gallus domesticus 
from the Eastern Cape Province, South Africa. Ph.D. thesis, 
University of South Africa.

SCHMIDT, G.D. & KUNTZ, R.E. 1977. Revision of Mediorhynchus 
Van Cleave 1916 (Acanthocephala) with a key to species. 
Journal of Parasitology, 63:500–507.

SEURAT, L.G. 1916. Sur un nouveau type de Spiruridae. Comp-
tes Rendus des Séances de la Societé de Biologie, 79:517–
519.

SHARMA, R.K. 1971. A new nematode Sicarius hoopoe n. sp. 
(Habronematinae) from a bird. Zoologischer Anzeiger 
Leipzig, 187:303–306.

SIEGFRIED, W.R. 1966. Growth, plumage development and 
moult in the Crowned Guineafowl Numida meleagris coro-
nata Gurney. Department of Nature Conservation Investiga-
tional Report No. 8.

SOULSBY, E.J.L. 1968. Helminths, arthropods and protozoa of 
domesticated animals (Sixth edition of Mönnig’s veterinary 
helminthology and entomology). London: Baillière and Cas-
sell.

SPASSKII, A.A. 1973. [New genera of cyclophyllidean cestodes.] 
Parazity Zhivotnykh i Rastenii, 9:38–48 [in Russian].

VERCRUYSSE, J., HARRIS, E.A., BRAY, R.A., NAGALO, M., 
PANGUI, M. & GIBSON, D.I. 1985. A survey of gastrointesti-
nal helminths of the common helmet guinea fowl (Numida 
meleagris galeata) in Burkina Faso. Avian Diseases, 29:742–
745.

VERSTER, A. & PTASINSKA-KLORYGA, Y. 1987. Helminths of 
Helmeted guineafowl in southern Africa. South African Jour-
nal of Wildlife Research, Supplement I: 36–38.

YAMAGUTI, S. 1961. The nematodes of vertebrates, Vol. III, Part 
I & II, in Systema Helminthum. New York: Interscience Pub-
lishers.