Table Of ContentJ. Helminthol. Soc. Wash.
64(1), 1997 pp. 113-119
Triodontophorus burchetti sp. n. and Triodontophorus hartmannae sp.
n. (Nematoda: Strongylidae) from the Burchell's, Hartmann's, and
Cape Mountain Zebras in Southern Africa
R. C. KRECEK,' V. A. KHARCHENKO, G. M. DvojNOS, F. S. MALAN, AND T. E. KRECEK '
2 2 3
1 Department of Veterinary Tropical Diseases, Faculty of Veterinary Science, University of Pretoria, Private Bag
X04, Onderstepoort 0110, South Africa,
2 Department of Parasitology, Institute of Zoology, Khmel'nytsky Street 15, Kiev-30, 252601 Ukraine, and
3 Hoechst Research Farm, P.O. Box 124, Malelane 1320, South Africa
ABSTRACT: Triodontophorus burchelli sp. n. is described from the ventral colons of 44 Burchell's zebras, Equus
burchelli antiquorum, in Kruger National Park in South Africa and Etosha National Park, Namibia. Triodonto-
phorus hartmannae sp. n. is described from the ventral colons of 21 Hartmann's mountain zebras, Equus zebra
hartmannae, in Etosha National Park, Namib-Naukluft Park, and the farm "Kelpie," Namibia, as well as from
13 Cape mountain zebras, Equus zebra zebra in Cape Province, South Africa. Triodontophorus burchelli sp. n.
is distinguished from other members of the genus by generally greater total body length, the absence of serration
on upper edges of teeth, and the shape of the gubernaculum. Triodontophorus hartmannae sp. n. is differentiated
from other species by a denticulation with medium serration.
KEY WORDS: Triodontophorus burchelli sp. n., Triodontophorus hartmannae sp. n., Strongylidae, new species,
Nematoda, taxonomy, Burchell's zebra, Hartmann's mountain zebra, Cape mountain zebra, Namibia, South
Africa.
Equids harbor at least 107 known species of studies and the nematodes were killed in Lugol's io-
dine and fixed in 10% formaldehyde (Malan et al.,
helminths, of which 69 are strongyles (Lichten-
198la, b). The specimens were cleared in lactophenol
fels, 1975). Like the other equine strongyles,
and examined with a Nikon Optiphot light microscope
Triodontophorus attaches to the intestinal mu- fitted with disc interference contrast. En face sections
cosa, which causes irritation, loss of blood, and of some of the specimens were cut in the region of the
ulcers (Levine, 1980). Parasitological investiga- buccal capsule and mounted in lactophenol to study
the structures of the head region. For scanning electron
tions were carried out on 3 species and subspe-
microscopy (SEM), the formalinized nematodes were
cies of equids in southern Africa including Bur- dehydrated in ethanol and critical-point dried in liquid
chell's zebras (Equus burchelli antiquorum), CO2. The dried nematodes were oriented onto a stub
Hartmann's mountain zebras (Equus zebra hart- bearing adhesive and coated with gold/palladium.
They were examined by SEM at 5-20 kV. Type spec-
mannae), and Cape mountain zebras (Equus ze-
imens are deposited in the United States National Par-
bra zebra) (Scialdo et al., 1982; Scialdo-Krecek,
asite Collection at Beltsville, Maryland, U.S.A., and
1983a, b; Scialdo-Krecek et al., 1983; Krecek et Onderstepoort Helminthological Collection, Onderste-
al., 1994). Two new Habronema, 1 new Cyli- poort Veterinary Institute, Onderstepoort, South Afri-
codontophorus, and 1 new Cylicostephanus spe-
cies recovered in these studies were previously
Results
described (Scialdo-Krecek and Malan, 1984;
Krecek, 1989; Scialdo-Krecek, 1983b), and this GENERAL: Strongylida, Strongylina, Stron-
paper reports on 2 previously unknown Triodon- gylidae, Strongylinae, Triodontophorus. Medi-
tophorus spp. um-sized worms. Buccal capsule subglobular
with 3 large esophageal teeth protruding into
Materials and Methods buccal cavity to about Vi its depth. Anterior rim
of buccal capsule surrounded by 6 platelike
Adult worms were recovered from 44 Burchell's ze-
bras (Equus burchelli antiquorum H. Smith, 1841) in structures that give appearance of the capsule
Kruger National Park (KNP), South Africa, and Etosha being thickened anteriorly. Mouth collar well
National Park (ENP), Namibia; 21 Hartmann's moun- developed with peripheral edge rounded or de-
tain zebras (Equus zebra hartmannae Matschie, 1898)
pressed as a ridge. Externo-labial papillae small,
in ENP, Namib-Naukluft Park, and the farm "Kelpie"
conical or slender. Cephalic papillae not promi-
in Namibia; and 13 Cape mountain zebras (Equus ze-
bra zebra) in Mountain Zebra National Park, South nent. External leaf-crown (ELC) of numerous
Africa. The zebras were processed for parasitological slender elements protrude from buccal collar. In-
113
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114 JOURNAL OF THE HELMINTHOLOGICAL SOCIETY OF WASHINGTON, 64(1), JAN 1997
ternal leaf-crown (ILC) of oval plates of same arately. All 3 laterals are of equivalent length
number as ELC elements. Dorsal gutter extends and thickness. Ventral rays each arise indepen-
to anterior edge of buccal capsule. Each of 3 dently and the prebursal papillae are evident.
esophageal teeth composed of 2 plates joined at Gubernaculum is pistol-shaped and the genital
an angle medially. cone is slightly elongated with hair-like genital
MALE: Well-developed dermal collar on appendages at the cone's tip. Spicule tips are
genital cone. Bursa with finely denticulated mar- hooked and sharply pointed distally.
gin and closed ventrally. FEMALES (11 specimens): Length 14.2-19.2
FEMALE: Vulva close or up to 3.0 mm from (16.3 ± 1.6) mm. Width 499-939 (758 ± 12.5).
the anus. Uteri parallel. Length of buccal capsule 114-180 (140 ± 18.9).
Width of buccal capsule 129-210 (149 ± 7.3).
Triodontophorus burchelli sp. n.
Number of elements of ELC 61. Esophagus 0.9-
(Figs. 1-9, Table 1)
1.2 (1.0 ± 0.1) mm long and 177-210 (194 ±
This new species was recovered from 11 of 11.5) wide. Distance of excretory pore from
the ventral colons of 44 Burchell's zebras in base of buccal capsule 650-824 (715 ± 62.6)
KNP and ENP in moderate numbers (1-2,500) and from anterior end 0.79-1.0 (0.86 ± 0.074)
and most closely resembles Triodontophorus mm. Distance of nerve ring from anterior end
brevicauda. Hence, measurements of T. brevi- 464-661 (539 ± 69.6) and of cervical papillae
cauda are included in Table 1 with this new 742-962 (843 ± 69.8). Distance from vulva to
Triodontophorus species. tip of tail 592-812 (667 ± 71) and from anus to
GENERAL: Mouth collar flattened with a rath- tip of tail 157-202 (178 ± 14.8). Egg 66-102
er acute erect edge around outside perimeter. Ce- (85.4 ± 10.9) long and 33-57 (45.8 ± 7.4) wide.
phalic papillae are not prominent while externo- TYPE SPECIMENS: One female holotype and 1
labial papillae long. Buccal capsule subglobular, male allotype; 1 male paratype (T-2168), On-
wider posteriorly. Teeth have no serration of up- derstepoort Helminthological Collection, Onder-
per edges. Esophagus moderately long, the stepoort Veterinary Institute, Onderstepoort,
nerve ring distinct and situated just above the South Africa.
middle of the esophagus. Two male and 2 female paratypes (USNPC
DESCRIPTION: Dimensions given as range No. 84404), U.S. National Parasite Collection,
(mean in micrometers ±1 standard deviation) U.S. Department of Agriculture (USDA), Belts-
unless otherwise indicated. ville, Maryland, U.S.A.
MALES (13 specimens): Length 12.9-18.0 TYPE HOST AND TYPE LOCALITY: EqilUS blir-
(15.5 ± 1.4) mm. Width 556-858 (650 ± 81.7). chelli antiquorum H. Smith, 1841. KNP, South
Length of buccal capsule 120-165 (144 ± 15.5). Africa (25°12'-24°24'S, 31°36'-32°2'E).
Width of buccal capsule 120-153 (140 ± 10.4). HABITAT IN HOST: Ventral colon.
Number of elements of ELC 60. Esophagus 0.8- ETYMOLOGY: This species is named after the
1.2 (1.0 ± 0.1) mm long and 165-213 (183 ± host, Burchell's zebra.
14.7) wide. Distance of excretory pore from
Triodontophorus hartmannae sp. n.
base of buccal capsule 510-788 (664 ± 70.9)
(Figs. 10-17, Table 1)
and from anterior end 684-928 (810 ± 66.1).
Distance anterior end from nerve ring 464-754 This species was recovered from all of the
(554 ± 74.1) and from deirids 638-904 (791 ± ventral colons of 21 Hartmann's mountain ze-
72). Dorsal ray 464-754 (571 ± 106) long. Gen- bras in ENP, Namib-Naukluft Park, and "Kel-
ital cone 224-314 (282 ± 25.9) long and 305- pie" farm, Namibia, in numbers of 2-1, 865 and
399 (345 ± 48.3) wide. Spicule length 1.5-2.2 from 8 of 13 Cape mountain zebras, South Af-
(1.7 ± 0.1) mm and gubernaculum length 224- rica, in numbers up to 889.
285 (245 ± 21.6). GENERAL: Mouth collar flattened with a rath-
Dorsal lobe of the male bursa is long, narrow, er acute erect edge around outside perimeter. Ce-
and demarcated from the lateral lobes. Dorsal phalic papillae not prominent whereas externo-
ray divided along its length into 2 small branch- labial papillae are long. Buccal capsule subgl-
es. Externodorsal ray is thicker than the lateral obular, and slightly wider posteriorly. Teeth have
rays and posterolateral and mediolateral rays medium serration of upper edges and protrude
arise together while the externolateral arises sep- well into the buccal capsule. Esophagus mod-
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KRECEK ET AL.—TRIODONTOPHORUS BURCHELLI SP. N. AND T. HARTMANNAE SP. N. 115
Figures 1-9. Drawing tube illustrations of Triodontophorus burchelli sp. n. 1. Buccal capsule, dorsal
view of the head. Scale bar = 100 urn. 2. Buccal capsule, lateral view of the head. Scale bar = 100 am.
3. En face view of head. Scale bar = 50 am. 4. Anterior end dorsal view. Scale bar = 500 am. 5. Male
tail, lateral view. Scale bar = 500 am. 6. Male tail, dorsal view. Scale bar = 500 am. 7. Female tail, lateral
view. Scale bar = 500 am. 8. Genital cone of male with gubernaculum, lateral view. Scale bar = 200 am.
9. Fused spicule tips of male. Scale bar = 50 am.
erately long, the nerve ring distinct and situated MALES (10 specimens): Length 12.6-17.3
just above the middle of the esophagus. (15.1 ± 1.2) mm. Width 557-696 (610 ± 53.6).
DESCRIPTION: Dimensions given as range Length of buccal capsule 180-204 (190 ± 8.4).
(mean in micrometers ± 1 standard deviation) Width of buccal capsule 135-183 (157 ± 12.8).
unless otherwise indicated. Number of elements of ELC 55-73. Esophagus
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Table 1. Principal measurements of Triodontophorus burchelli sp. n., T. brevicauda, T. hartmannae sp. n., T. nipponicus, and T. minor (all measurements in
micrometers umess oinerwise siaicu;.
T. burchelli T. brevicauda T. hartmannae T. nipponicus T. minor
13 $ 11 9 16* 39 10 $ 9 9 3 cJ 3 9 \J 3 9
•fl
Total length (mm) 12.9-18.0 14.2-19.2 10.4 15.1-17.6 12.6-17.3 12.5-16.8 11.2-13.1 13.1-14.8 12.4 11.8-13.6 H
Width 556-858 499-939 660 820-900 557-696 510-777 600-700 700-720 600 700-720 Xrn
X
Buccal capsule rn
Length 120-165 114-180 192 224-232 180-204 165-204 112-120 128-136 128 144-168 2
Width 120-153 129-210 135-183 135-177 160-180 178-184 176 192-208 z
H
Number of elements in external leaf-crown 60 61 * * 55-73 67-72 * * * * X
O
r
Esophagus 8
Length (mm) 0.8-1.2 0.9-1.2 1.0 1.1-1.2 0.7-1.2 1.0-1.2 0.9-1.0 1.0-1.1 1.0 1.0-1.2 n
Width 165-213 177-210 200 220-340 150-219 1 80-240 200-208 216-256 136 200-220 r
Distance of excretory pore from 0On0
base of buccal capsule 510-788 650-824 640 576-608 626-731 336-731 472-640 536-704 576 416-600
5
Dorsal ray H
Length 464-754 — 672 — 441-580 — 416-544 — 544 — 0•n
CC Genital cone
oo ^
pypy Length 224-314 — * * 224-336 — 184-280 — 320 — 00
righrigh Width 305-399 — ** _ 299-598 — 208-248 — 256 — Z
t ©t © Spicule length (mm) 1.5-2.2 2.24 — 0.9-1.1 0.9-1.0 1.2 O
20 20 Gubernaculum length 224-285 — — — 224-280 — — — — — 0
11
11 Distance from vulva to tip of tail (mm) — 592-812 — 256-360 — 661-916 — 616-776 — 592-680
, Th, Th Distance from anus to tip of tail (mm) — 157-202 — 96-152 — 157-252 — 160-176 — 120-144 O4^S
ee
He He Egg C
lmlm Length — 66-102 — 56-64 — 75-93 — 64-80 — 64-76 Z
inthinth Width — 33-57 — 32 — 39-48 — 36-40 — 40-52 VO
oloolo Type of denticulation No serration No serration Medium serration Strong serration Fine serration -J
gg
icic Externo-labial papillae Short, conical Short, broad, conical Long Long, narrow, pointed Long, narrow, pointed
aa
l Sl S with short tips
oo
cc
ietyiety * Not possible to cut heads in borrowed specimens.
o o ** Could not roll borrowed specimens—old and fragile.
f Wf W
aa
ss
hh
inin
gg
toto
nn
O LANRUOJ
KRECEK ET AL.—TRIODONTOPHORUS BURCHELLI SP. N. AND T. HARTMANNAE SP. N. 117
11 12
18
Figures 10-18. Drawing tube illustrations of Triodontophorus hartmannae sp. n. 10. Buccal capsule,
dorsal view of the head. Scale bar = 100 u.m. 11. Buccal capsule, lateral view of the head. Scale bar =
100 (xm. 12. En face view of head. Scale bar = 50 ftni. 13. Anterior end dorsal view. Scale bar = 500 n-rn.
14. Male tail, lateral view. Scale bar = 500 |un. 15. Male tail, dorsal view. Scale bar = 500 p-m. 16. Female
tail, lateral view. Scale bar = 500 (im. 17. Genital cone of male with gubernaculum, lateral view. Scale
bar = 200 u.m. 18. Fused spicule tips of male. Scale bar = 50 u.m.
0.7-1.2 (1.0 ± 0.1) mm long and 150-219 (197 Distance anterior end from nerve ring 534—626
± 22.5) wide. Distance of excretory pore from (578 ± 25.5) and from deirids 777-870 (822 ±
base of buccal capsule 626-731 (660 ± 37.9) 36.4). Dorsal ray 441-580 (512 ± 41.6) long.
and from anterior end 812-940 (850 ± 42.8). Genital cone 224-336 (289 ± 33.9) long and
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118 JOURNAL OF THE HELMINTHOLOGICAL SOCIETY OF WASHINGTON, 64(1), JAN 1997
299-598 (308.3 ± 117.6) wide. Spicule length tenfels (1975) but could not be assigned to a
0.9-1.1 (1.1 ± 0.07) mm and gubernaculum known species are therefore designated Triodon-
length 224-280 (243 ± 15.2). tophorus burchelli sp. n. and Triodontophorus
Dorsal lobe of the male bursa, is short and hartmannae sp. n.
demarcated from the lateral lobes. Dorsal ray di- According to Lichtenfels (1975), the species
vided along its length into 2 branches. Exter- of Triodontophorus are T. brevicauda Boulen-
nodorsal ray is thicker than the lateral rays and ger, 1916; T. brochotribulatus Martinez Gomez,
posterolateral and mediolateral rays arise togeth- 1966; T. minor (Looss, 1900); T. nipponicus Ya-
er while the externolateral arises separately. maguti, 1943; T. popovi Ershov, 1931; T. ser-
Ventral rays arise together and prebursal papillae ratus (Looss, 1900); and T. tenuicollis Boulen-
are evident. Gubernaculum is slender, pistol- ger, 1916. Other authors, Dvojnos and Khar-
shaped and the genital cone slightly rounded chenko (1985), considered T. brochotribulatus
with finger-like genital appendages. Spicule tips and T. popovi synonyms as well as T. nipponicus
are hooked and splayed, and distally tapered to and T. tenuicollis.
a rounded point. Triodontophorus burchelli sp. n. bears the
FEMALES (9 specimens): Length 12.5-16.8 closest resemblance to T. brevicauda. The new
(14.6 ± 1.6) mm. Width 510-777 (625 ± 80.9). species can be distinguished from T. brevicauda
Length of buccal capsule 165-204 (186 ± 14.4). in the male by a greater total body length, great-
Width of buccal capsule 135-177 (160 ± 15.8). er distances of the vulva to the tip of tail (592-
Number of elements of ELC 67-72. Esophagus 812 |xm as compared to 256-360 u,m) and anus
1.0-1.2 (1.1 ± 0.069) mm long and 180-240 to tip of tail (with 157-202 jjim as compared to
(207 ± 16.9) wide. Distance of excretory pore 96—152 (Jim). Triodontophorus burchelli also
from base of buccal capsule 336-731 (546 ± differs with an absence of serration on upper
138) and from anterior end 568-940 (749 ± edges of teeth and a more slender gubernaculum.
13.8) mm. Distance of nerve ring from anterior All the remaining species have some degree
end 487-661 (563 ± 53.2) and of cervical pa- of serration of the denticulation that differs from
pillae 556-893 (710 ± 135). Distance from vul- the smooth teeth of T. burchelli sp. n. This char-
va to tip of tail 661-916 (803 ± 93) and anus acteristic is salient for each species even while
to tip of tail 157-252 (212 ± 30.8). Eggs 75-93 the degree of teeth serration varies within a spe-
(81 ± 6.2) long and 39-48 (42 ± 3.8) wide. cies (Dvojnos and Kharchenko, 1985). The de-
TYPE SPECIMENS: One female holotype and 1 scription of T. minor by Skrjabin and Erschov,
male allotype; 2 female and 3 male paratypes 1933, in Popova, 1964 (English translation), is
(T-2167), Onderstepoort Helminthological Col- similar in many characteristics to this new spe-
lection, Onderstepoort Veterinary Institute, On- cies. These are the absence of serration of teeth,
derstepoort, South Africa. the long median lobe of the dorsal ray, and the
Two male and 2 female paratypes (USNPC spicules length as well as the female vulva and
No. 84405), U.S. National Parasite Collection, anus to tip of tail distances. The most apparent
USDA, Beltsville, Maryland, U.S.A. difference is the number of elements of the ex-
TYPE HOST AND TYPE LOCALITY: Equus zebra ternal leaf-crowns which they report as 50.
hartmannae Matschie, 1898. Farm "Kelpie," Triodontophorus burchelli sp. n. is character-
Namibia (22°43'S, 16°43'E). ized by a greater body length, the absence of
HABITAT IN HOST: Ventral colon. serration on the upper edges of the teeth, greater
ETYMOLOGY: This species is named after the vulva to tip of tail (519-812 |jim) length and
host, Hartmann's mountain zebra. shape of gubernaculum when compared with T.
For the purpose of comparison, measurements brevicauda.
of Triodontophorus brevicauda, Triodontopho- Triodontophorus hartmannae sp. n. differs
rus nipponicus, and Triodontophorus minor are from T. nipponicus Yamaguti, 1943, both in the
included with those for the 2 new species in Ta- longer distance of the vulva to the tip of the tail
ble 1. (661-916 fjim as compared to 616-776 jxm) and
less pointed serration of the teeth. Triodonto-
Discussion phorus hartmannae sp. n. has shorter spicules
Nematodes that agreed with the generic de- (0.9-1.1 mm) than T. minor according to Theiler
scription of Triodontophorus according to Lich- (1923) and Diaz-Ungria (1965), who reported a
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KRECEK ET AL.—TRIODONTOPHORUS BURCHELL1 SP. N. AND T. HARTMANNAE SP. N. 119
spicule length of 1.7 mm. Additionally, only 1 Horak, and F. S. Malan. 1994. Helminth para-
sites of Cape mountain zebras from Cape Prov-
type specimen of a male T. minor from a horse
ince, South Africa. Journal of Wildlife Diseases
was available from the U.S. National Parasite 30:277-280.
Collection, and its spicules measured 1.2 mm. Levine, N. D. 1980. Nematode Parasites of Animals
Spicules of males of this species from the col- and Man. Burgess Publishing, Minneapolis, Min-
nesota. 477 pp.
lection of the Institute of Zoology, Ukrainian
Lichtenfels, J. R. 1975. Helminths of domestic
National Academy, were measured and were
equids. Illustrated keys to genera and species with
1.2-1.4 mm in length. According to Theiler emphasis on North American forms. Proceedings
(1923), the ELC elements for T. minor number of the Helminthological Society of Washington
50, whereas Diaz-Ungria (1965) reported 44-50, (Special Issue) 42:1-92.
Malan, F. S., R. K. Reinecke, and R. C. Scialdo.
far fewer than the 61-73 of the new species T.
198la. Recovery of helminths postmortem from
hartmannae sp. n.
equines. I. Parasites in arteries, subperitoneum,
Triodontophorus hartmannae sp. n. is most liver and lungs. Onderstepoort Journal of Veteri-
closely related to T. nipponicus Yamaguti, 1943, nary Research 48:141-143.
and T. minor (Looss, 1900) in all features except , , and . 1981b. Recovery of
helminths postmortem from equines. II. Helminths
denticulation. The medium serration of dentic-
and larvae of Gasterophilus in the gastro-intesti-
ulation of the new species ranges between the nal tract and oestrids from the sinuses. Onderste-
fine serration of T. minor and the coarse serra- poort Journal of Veterinary Research 48:145-147.
tion of T. nipponicus. Popova, T. I. 1955. Strongyloids of animals and man.
Osnovy Nematodologii, Vol. V. Akad. Nauk
Triodontophorus hartmannae sp. n. is char-
SSSR, Moscow. 241 pp. (In Russian; English
acterized by medium serration of the teeth.
translation, 1964, Israel Program for Scientific
Translations, Jerusalem. Also available from the
Acknowledgments National Technical Information Service, U.S. De-
We thank Dr. J. R. Lichtenfels for comments partment of Commerce, Springfield, Virginia,
U.S.A.).
regarding specimens of the 2 new species de-
Scialdo, R. C., R. K. Reinecke, and V. de Vos. 1982.
scribed in this paper and Prof. J. D. F. Boomker
Seasonal incidence of helminths in the Burchell's
for comments on the manuscript. Financial sup- zebra. Onderstepoort Journal of Veterinary Re-
port for these studies originated in South Africa search 49:127-130.
from the University of Pretoria, Foundation for Scialdo-Krecek, R. C. 1983a. Studies on the para-
sites of zebras. I. Nematodes of the Burchell's ze-
Research Development, Hoechst, and the Fritz
bra in the Kruger National Park. Onderstepoort
Visser Agricultural Bursary and in the Ukraine Journal of Veterinary Research 50:111-114.
from the International Science Foundation. This . 1983b. Studies on the parasites of zebra. II.
study forms part of the Wildlife Research Pro- Cylicostephanus longiconus n. sp. (Nematoda:
Strongylidae) from the mountain zebra, Equus ze-
gramme at the Faculty of Veterinary Science,
bra hartmannae (Matschie, 1898). Onderstepoort
University of Pretoria.
Journal of Veterinary Research 50:169-172.
, R. K. Reinecke, and H. C. Biggs. 1983.
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