Journal of Helminthology (1979) 53, 73-78
Dunnifilaria dilli sp. n. (Nematoda: Filarioidea) from Rattus koratensis in Thailand J. W. MAK Institute for Medical Research, Kuala Lumpur, Malaysia and R. E. WEAVER Department of Entomology, SEATO, Medical Research Laboratory, Bangkok, Thailand ABSTRACT Dunnifilaria dilli, a new species of filariid from Rattus koratensis from Thailand is described. Adult worms were recovered from the lymphatic system. They differ from D. ramachandrani in their location in the host, the tail length, the shape of the spicules, and in the number of caudal papillae. Microfilariae are much smaller and unsheathed. Additional hosts include R. rattus and R. sabanus.
During a survey for filarial infections in small mammals trapped in Sakaret and Chanthaburi, Thailand, a number of filarial parasites were recovered. Of these, Breinlia booliati, Breinlia sergenti and Dunnifilaria ramachandrani were recovered for the first time in Thailand. This will be the subject of another paper. In addition to the above, microfilariae and adult worms were recovered from Rattus koratensis in Chanthaburi and from Rattus sabanus and Rattus rattus in Sakaret, which belonged to the genus Dunnifilaria Mullin and Balasingam, 1973. However, these filarial worms were found to be different from Dunnifilaria ramachandrani Mullin and Balasingam, 1973, the sole representative of the genus, recovered from Rattus sabanus in Malaysia. This new species is named after Dr. G. S. Dill, the original collector of these worms in recognition of his work in filariasis in Thailand. MATERIALS AND METHODS Animals trapped were identified and screened for microfilaraemia by the method of Dennis and Kean (1971). Thick blood smears prepared from positive animals were allowed to dry overnight and then stained with dilute Giemsa (2-2.5 ml to 100 ml buffered distilled water at pH 7.2 for 1 hour) or hot haematoxylin after methanol fixing. Animals were then killed and subjected to an exhaustive post-mortem. Lymph nodes were examined by the method of Buckley and Edeson (1956). Adult worms recovered were killed in hot 70% ethanol and transferred to a solution of 70 % ethanol and 5 % glycerine. The adult worms were then slowly brought into pure glycerine by evaporation and studied. All measurements and drawings were made with the aid of a camera lucida. RESULTS Adult worms belonging to this species were recovered from 6 Rattus koratensis trapped in Canthaburi and from a R. sabanus and R. rattus trapped in Sakaret, Thailand. In all a total of 8 complete males and females, 2 portions of male and 4 portions of female worms were recovered from the lymphatic system, in particular the lymph nodes of the posterior aspect. The following description of the parasite is based on the study of 8 males and females each. All measurements are in millimetres unless stated otherwise, the mean being in parenthesis. 73
J. W. MAK and R. E. WEAVER
Dunnifilaria dilli sp. n. Description Filarioidea: Onchocercidae (Leiper, 1911) Chabaud and Anderson, 1959; Splendidofilariinae Chabaud and Choquet, 1953; Dunnifilaria Mullin and Balasingam, 1973. Adult worms small and slender, tapering at both extremities. Mouth opening small, simple, and surrounded by four pairs of minute submedian papillae. Cuticle generally smooth but with fine transverse striations at both extremities, starting a short distance from both ends. Oesophagus fairly long with a short muscular anterior portion poorly demarcated from a longer, glandular posterior portion. Vulval opening just posterior to midoesophageal region. Anus in female inconspicuous and subterminal. Tail in male short, without caudal alae. Two pairs of inconspicuous postanal papillae. Spicules subequal and similar. Microfilariae without sheath, circulating in peripheral blood. Male: (Figs. 3-5; Table 1). Male shorter than female worm. Body length 11.8-19.7 (14.9). Maximum width 81-110 (96.2) urn. Posterior end coiled into 2-3 turns. Tail short, without caudal alae. Spicules subequal and similar, the left being 70-95 (81.2) um and the right 60-85 (71.5) um long. Both slightly curved and tapers sharply at the tip, the right being more pointed. Gubernaculum absent. Two pairs of inconspicuous postanal papillae. TABLE 1 Comparison between male Dunnifilaria sp. n. and Dunnifilaria ramachandrani Mullin and Balasingam, 1973. All measurements in microns unless stated otherwise. (Mean measurements in parenthesis) Dunnifilaria sp. n.
Range
Mean
Total body length (mm) Distance from anterior end to: Nerve ring testis
11.8-19.7
14.9
110-175 430-675
146.8 537
Length of oesophagus Length of muscular portion Length of tail Left spicule Right spicule
635-740 185-220 60-90 70-95 60-85
687.5 202.5 74.2 81.2 71.5
Spicular ratio Left: right
1.0-1.3:1
1.1:1
Max. width
81-110
96.2
44-58
51.1
60-90 52-58
74.3 48.1
Width at Nerve ring Oesophageal-intestinal junction Anus Midbody Papillae
2 pairs postanals
Dunnifilaria ramachandrani
Range 9.6-14.8
Mean 13.6 60
500-950 75-100 110-160 70-90 65-80
650 130 77 70 1.1:1 —
90-130 4 pairs postanals
Female: (Figs. 1 and 6; Table 2). Longer than male. Body length 22.1-30.4 (25.9). Maximum width 150-160 (153.3) urn. Vulval opening anterior to oesophageal intestinal junction, 328.8 um from the anterior end. Ovijector slightly globular and muscular. Vagina long and muscular. Didelphic, uterine coils extend nearly to tip of tail. Anus subterminal, about 10 um from tip. 74
Dunnifilaria dilli sp. n. from rat in Thailand TABLE 2 Comparison between female Dunnifilaria sp. n. and Dunnifilaria ramachandrani Mullin and Balasingam, 1973. All measurements in microns unless stated otherwise (Mean measurements in parenthesis) Dunnifilariasp. n. Mean Range
Dunnifilaria ramachandrani Range Mean
22.1-30.4
25.9
18.5-30.3
25.2
125-160 185^160
142.5 328.8
320-450
100 410
Length of: Oesophagus Glandular portion
670-840 330
737.5 330
Maximum width
150-160
153.3
Width at: Nerve ring O-I junction Vulval opening Mid-body
80 110-125 90-115
80 117.5 101.3
Total body length (mm) Distance of anterior end to: Nerve ring Vulval opening
— 150
120-200
Tail length
180
10
25
Microfilaria: (Fig. 2; Table 3). Very short; in Giemsa stained specimens 70-93 (08.5) pirn long by 3-4 (3.7) urn at its widest. A large proportion of the microfilariae with their posterior fourth characteristically bent back on itself and sometimes doubling back again. No sheath. Innenkorper poorly visible. Nuclei overlapping and blob-like. Cephalic space very short. Landmarks other than nerve ring poorly visible. TABLE 3 Comparison of microfilarial measurements between Dunnifilaria ramachandrani and Dunnifilaria sp. n. Giemsa stain D. ramachandrani Dunnifilaria sp. n. Total length Distance from ant. end to: 1st nucleus nerve ring ex. pore anal pore Width at 1st nucleus nerve ring ex. pore anal pore
Haematoxylin stain D. ramachandrani Dunnifilaria sp. n.
122-144(133)
70-93 (80.5)
112-136(123)
70-102 (87.0)
(2.2) 38^8 (42)
1-2(1.6) 15-32(22.3) 26-30 (28.7) 58-72 (65)
(2) 32^3 (38) 68-77 (74)
1-2(1.6) 17-25 (20.3) 23-38 (30) 62-80 (72)
2-3.5 (2.6) 3-4 (3.4) 3-3.5 (3.2) 2-3 (2.5)
(4)
2-3 (2.6) 25-4 (3.2) 2-3.5 (2.9) 2(2)
107-122 (114) (6.5)
Max. width
—
3-^t (3.7)
—
3-4 (3.3)
No. measured
10
20
10
20
75
J. W. MAK and R. E. WEAVER
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: / •
w
FIG. 1. Anterior end female worm; FIG. 2. Microfilaria; FIG. 3. Posterior end of male worm; FIG. 4. Left spicule; FIG. 5. Right spicule; FIG. 6. Posterior end of female worm. 76
Dunnifilaria dilli sp. n. from rat in Thailand
Type Host: Rattus koratensis Additional Hosts: Rattus sabanus and Rattus rattus Location: Lymphatic system Type locality: Chanthaburi and Sakaret, Thailand Specimens: Holotype (male). I.M.R. Fil. Coll. No. 1-76. Deposited in the Filariasis Research Division, Institute for Medical Research, Jalan Pahang, Kuala Lumpur, Malaysia. Paratypes: from the same host and locality, deposited in the World Health Organization Collaborating Centre for the Filarioidea, London School of Hygiene and Tropical Medicine, and I.M.R. Fil. Coll. No. 2-76. DISCUSSION This worm is placed in the genus Dunnifilaria Mullin and Balasingam, 1973. It is a small worm with uniform diameter and tapering at both extremities. It has a round, smooth head, without any peribuccal chitinous formations and with small cephalic papillae. The oesophagus is fairly long, with an anterior muscular portion indistinctly demarcated from the posterior glandular portion. The vulva is anterior in position. The anus is subterminal and difficult to visualize. In the male, the tail is digitiform, without any caudal alae, with only postanal papillae, without gubemaculum, and with subequal, similar spicules. Anderson and Bain (1976) in a recent review of the classification of the filarioids and their allies listed 5 of the 16 genera in the subfamily Splendido filariinae that are found in mammals, these being Micipsella Seurat, 1921, Meningonema Orihel & Esslinger, 1973, Dunnifilaria Mullin and Balasingam, 1973, Onchocercella Yorke and Maplestone, 1931, and Protofilaria Chandler, 1929. This worm is not placed in Micipsella because of its divided oesophagus and absence of cuticular bosses. It does not have the numerous preanal papillae in the male nor the quadridelphic female as in Meningonema. Unlike Onchocercella, it does not have fusiform cuticular thickenings nor adanal papillae. The female worm unlike Protofilaria does not have a truncated tail with a dorsoventral cleft dividing it into two short lobes. While Dunnifilaria ramachandrani were recovered from the right ventricle and pulmonary arteries, Dunnifilaria dilli worms were from the lymphatic system, being recovered mostly from the lymph nodes of the posterior part of the animal. Unlike D. ramachandrani, it has fine transverse striations at both extremities. It is of the same length but slightly narrower than the former. The tail is very much shorter in this worm being about 74.2 um long compared to 130 um in D. ramachandrani (Table 1). The oesophagus is of the same length but the muscular portion is longer in the present male worm. The left spicule is slightly longer in the present worm being 81.2 um compared to 77 um respectively. However, the spicular ratios are similar. The spicules in this worm appear to be narrower and tapers more sharply than in D. ramachandrani. Unlike D. ramachandrani it has only 2 pairs of post-anal papillae. In both male and female of the present worm the nerve ring is further back from the anterior end (Tables 1 and 2). The microfilariae of D. dilli are without sheath and very much smaller than those of D. ramachandrani, being 80.5 Mm and 87.0 um in Giemsa and haematoxylin stained specimens compared to 133 um and 123 um respectively. It is very much narrower and the excretory pore is situated at a distance of 34.5% of the total length from the anterior end compared to 60.2 % in D. ramachandrani.
ACKNOWLEDGEMENTS
The authors are grateful to Dr. Douglas J. Gould, Chief, Department of Medical Entomology, SEATO, Medical Research Laboratory, Bangkok, Thailand who initiated the project and to Dr. George F. deWitt, Director, Institute for Medical Research, Kuala Lumpur, Malaysia, for permission to publish this paper. 77
J. W. MAK and R. E. WEAVER
REFERENCES ANDERSON, R. C. and BAIN, O. (1976) Keys to genera of the order Spirurida. Part 3. Diplotriaenoidea, Aproctoidea and Filarioidea. In CIH Keys to the Nematode Parasites of Vertebrates (eds R. C. Anderson, A. G. Chabaud and S. Willmott) Commonwealth Agricultural Bureaux: Farnham Royal, Bucks, pp. 59-116. BUCKLEY, J. J. C. and EDESON, J. F. B. (1956) On the adult morphology of Wuchereria sp. {malayi?) from a monkey (Macaca irus) and from cats in Malaya, and on Wuchereria pahangi n. sp. from a dog and a cat. Journal of Helminthology, 30, 1-20. CHABAUD, A. G. and ANDERSON, R. C. (1959) Nouvel essai de classification des filaires (superfamille des Filarioidea) II. 1959. Annales de Parasitologie, 34, 64-87. DENNIS, D. T. and KEAN, B. H. (1971) Isolation of microfilariae: report of a new method. Journal of Parasitology, 57, 1145-1146. MULLIN, S. and BALASINGAM, S. (1973) Dunnifilaria ramachandrani gen. n., sp. n. (Nematoda: Filarioidea) from the long-tailed giant rat (Rattus sabanus) in Malaysia. Proceedings of the Helminthological Society of Washington, 40, 47-49. Received 10 November, 1977.
78
Dunnifilaria dilli sp. n. (Nematoda: Filarioidea) from Rattus koratensis in Thailand J. W. MAK Institute for Medical Research, Kuala Lumpur, Malaysia and R. E. WEAVER Department of Entomology, SEATO, Medical Research Laboratory, Bangkok, Thailand ABSTRACT Dunnifilaria dilli, a new species of filariid from Rattus koratensis from Thailand is described. Adult worms were recovered from the lymphatic system. They differ from D. ramachandrani in their location in the host, the tail length, the shape of the spicules, and in the number of caudal papillae. Microfilariae are much smaller and unsheathed. Additional hosts include R. rattus and R. sabanus.
During a survey for filarial infections in small mammals trapped in Sakaret and Chanthaburi, Thailand, a number of filarial parasites were recovered. Of these, Breinlia booliati, Breinlia sergenti and Dunnifilaria ramachandrani were recovered for the first time in Thailand. This will be the subject of another paper. In addition to the above, microfilariae and adult worms were recovered from Rattus koratensis in Chanthaburi and from Rattus sabanus and Rattus rattus in Sakaret, which belonged to the genus Dunnifilaria Mullin and Balasingam, 1973. However, these filarial worms were found to be different from Dunnifilaria ramachandrani Mullin and Balasingam, 1973, the sole representative of the genus, recovered from Rattus sabanus in Malaysia. This new species is named after Dr. G. S. Dill, the original collector of these worms in recognition of his work in filariasis in Thailand. MATERIALS AND METHODS Animals trapped were identified and screened for microfilaraemia by the method of Dennis and Kean (1971). Thick blood smears prepared from positive animals were allowed to dry overnight and then stained with dilute Giemsa (2-2.5 ml to 100 ml buffered distilled water at pH 7.2 for 1 hour) or hot haematoxylin after methanol fixing. Animals were then killed and subjected to an exhaustive post-mortem. Lymph nodes were examined by the method of Buckley and Edeson (1956). Adult worms recovered were killed in hot 70% ethanol and transferred to a solution of 70 % ethanol and 5 % glycerine. The adult worms were then slowly brought into pure glycerine by evaporation and studied. All measurements and drawings were made with the aid of a camera lucida. RESULTS Adult worms belonging to this species were recovered from 6 Rattus koratensis trapped in Canthaburi and from a R. sabanus and R. rattus trapped in Sakaret, Thailand. In all a total of 8 complete males and females, 2 portions of male and 4 portions of female worms were recovered from the lymphatic system, in particular the lymph nodes of the posterior aspect. The following description of the parasite is based on the study of 8 males and females each. All measurements are in millimetres unless stated otherwise, the mean being in parenthesis. 73
J. W. MAK and R. E. WEAVER
Dunnifilaria dilli sp. n. Description Filarioidea: Onchocercidae (Leiper, 1911) Chabaud and Anderson, 1959; Splendidofilariinae Chabaud and Choquet, 1953; Dunnifilaria Mullin and Balasingam, 1973. Adult worms small and slender, tapering at both extremities. Mouth opening small, simple, and surrounded by four pairs of minute submedian papillae. Cuticle generally smooth but with fine transverse striations at both extremities, starting a short distance from both ends. Oesophagus fairly long with a short muscular anterior portion poorly demarcated from a longer, glandular posterior portion. Vulval opening just posterior to midoesophageal region. Anus in female inconspicuous and subterminal. Tail in male short, without caudal alae. Two pairs of inconspicuous postanal papillae. Spicules subequal and similar. Microfilariae without sheath, circulating in peripheral blood. Male: (Figs. 3-5; Table 1). Male shorter than female worm. Body length 11.8-19.7 (14.9). Maximum width 81-110 (96.2) urn. Posterior end coiled into 2-3 turns. Tail short, without caudal alae. Spicules subequal and similar, the left being 70-95 (81.2) um and the right 60-85 (71.5) um long. Both slightly curved and tapers sharply at the tip, the right being more pointed. Gubernaculum absent. Two pairs of inconspicuous postanal papillae. TABLE 1 Comparison between male Dunnifilaria sp. n. and Dunnifilaria ramachandrani Mullin and Balasingam, 1973. All measurements in microns unless stated otherwise. (Mean measurements in parenthesis) Dunnifilaria sp. n.
Range
Mean
Total body length (mm) Distance from anterior end to: Nerve ring testis
11.8-19.7
14.9
110-175 430-675
146.8 537
Length of oesophagus Length of muscular portion Length of tail Left spicule Right spicule
635-740 185-220 60-90 70-95 60-85
687.5 202.5 74.2 81.2 71.5
Spicular ratio Left: right
1.0-1.3:1
1.1:1
Max. width
81-110
96.2
44-58
51.1
60-90 52-58
74.3 48.1
Width at Nerve ring Oesophageal-intestinal junction Anus Midbody Papillae
2 pairs postanals
Dunnifilaria ramachandrani
Range 9.6-14.8
Mean 13.6 60
500-950 75-100 110-160 70-90 65-80
650 130 77 70 1.1:1 —
90-130 4 pairs postanals
Female: (Figs. 1 and 6; Table 2). Longer than male. Body length 22.1-30.4 (25.9). Maximum width 150-160 (153.3) urn. Vulval opening anterior to oesophageal intestinal junction, 328.8 um from the anterior end. Ovijector slightly globular and muscular. Vagina long and muscular. Didelphic, uterine coils extend nearly to tip of tail. Anus subterminal, about 10 um from tip. 74
Dunnifilaria dilli sp. n. from rat in Thailand TABLE 2 Comparison between female Dunnifilaria sp. n. and Dunnifilaria ramachandrani Mullin and Balasingam, 1973. All measurements in microns unless stated otherwise (Mean measurements in parenthesis) Dunnifilariasp. n. Mean Range
Dunnifilaria ramachandrani Range Mean
22.1-30.4
25.9
18.5-30.3
25.2
125-160 185^160
142.5 328.8
320-450
100 410
Length of: Oesophagus Glandular portion
670-840 330
737.5 330
Maximum width
150-160
153.3
Width at: Nerve ring O-I junction Vulval opening Mid-body
80 110-125 90-115
80 117.5 101.3
Total body length (mm) Distance of anterior end to: Nerve ring Vulval opening
— 150
120-200
Tail length
180
10
25
Microfilaria: (Fig. 2; Table 3). Very short; in Giemsa stained specimens 70-93 (08.5) pirn long by 3-4 (3.7) urn at its widest. A large proportion of the microfilariae with their posterior fourth characteristically bent back on itself and sometimes doubling back again. No sheath. Innenkorper poorly visible. Nuclei overlapping and blob-like. Cephalic space very short. Landmarks other than nerve ring poorly visible. TABLE 3 Comparison of microfilarial measurements between Dunnifilaria ramachandrani and Dunnifilaria sp. n. Giemsa stain D. ramachandrani Dunnifilaria sp. n. Total length Distance from ant. end to: 1st nucleus nerve ring ex. pore anal pore Width at 1st nucleus nerve ring ex. pore anal pore
Haematoxylin stain D. ramachandrani Dunnifilaria sp. n.
122-144(133)
70-93 (80.5)
112-136(123)
70-102 (87.0)
(2.2) 38^8 (42)
1-2(1.6) 15-32(22.3) 26-30 (28.7) 58-72 (65)
(2) 32^3 (38) 68-77 (74)
1-2(1.6) 17-25 (20.3) 23-38 (30) 62-80 (72)
2-3.5 (2.6) 3-4 (3.4) 3-3.5 (3.2) 2-3 (2.5)
(4)
2-3 (2.6) 25-4 (3.2) 2-3.5 (2.9) 2(2)
107-122 (114) (6.5)
Max. width
—
3-^t (3.7)
—
3-4 (3.3)
No. measured
10
20
10
20
75
J. W. MAK and R. E. WEAVER
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: / •
w
FIG. 1. Anterior end female worm; FIG. 2. Microfilaria; FIG. 3. Posterior end of male worm; FIG. 4. Left spicule; FIG. 5. Right spicule; FIG. 6. Posterior end of female worm. 76
Dunnifilaria dilli sp. n. from rat in Thailand
Type Host: Rattus koratensis Additional Hosts: Rattus sabanus and Rattus rattus Location: Lymphatic system Type locality: Chanthaburi and Sakaret, Thailand Specimens: Holotype (male). I.M.R. Fil. Coll. No. 1-76. Deposited in the Filariasis Research Division, Institute for Medical Research, Jalan Pahang, Kuala Lumpur, Malaysia. Paratypes: from the same host and locality, deposited in the World Health Organization Collaborating Centre for the Filarioidea, London School of Hygiene and Tropical Medicine, and I.M.R. Fil. Coll. No. 2-76. DISCUSSION This worm is placed in the genus Dunnifilaria Mullin and Balasingam, 1973. It is a small worm with uniform diameter and tapering at both extremities. It has a round, smooth head, without any peribuccal chitinous formations and with small cephalic papillae. The oesophagus is fairly long, with an anterior muscular portion indistinctly demarcated from the posterior glandular portion. The vulva is anterior in position. The anus is subterminal and difficult to visualize. In the male, the tail is digitiform, without any caudal alae, with only postanal papillae, without gubemaculum, and with subequal, similar spicules. Anderson and Bain (1976) in a recent review of the classification of the filarioids and their allies listed 5 of the 16 genera in the subfamily Splendido filariinae that are found in mammals, these being Micipsella Seurat, 1921, Meningonema Orihel & Esslinger, 1973, Dunnifilaria Mullin and Balasingam, 1973, Onchocercella Yorke and Maplestone, 1931, and Protofilaria Chandler, 1929. This worm is not placed in Micipsella because of its divided oesophagus and absence of cuticular bosses. It does not have the numerous preanal papillae in the male nor the quadridelphic female as in Meningonema. Unlike Onchocercella, it does not have fusiform cuticular thickenings nor adanal papillae. The female worm unlike Protofilaria does not have a truncated tail with a dorsoventral cleft dividing it into two short lobes. While Dunnifilaria ramachandrani were recovered from the right ventricle and pulmonary arteries, Dunnifilaria dilli worms were from the lymphatic system, being recovered mostly from the lymph nodes of the posterior part of the animal. Unlike D. ramachandrani, it has fine transverse striations at both extremities. It is of the same length but slightly narrower than the former. The tail is very much shorter in this worm being about 74.2 um long compared to 130 um in D. ramachandrani (Table 1). The oesophagus is of the same length but the muscular portion is longer in the present male worm. The left spicule is slightly longer in the present worm being 81.2 um compared to 77 um respectively. However, the spicular ratios are similar. The spicules in this worm appear to be narrower and tapers more sharply than in D. ramachandrani. Unlike D. ramachandrani it has only 2 pairs of post-anal papillae. In both male and female of the present worm the nerve ring is further back from the anterior end (Tables 1 and 2). The microfilariae of D. dilli are without sheath and very much smaller than those of D. ramachandrani, being 80.5 Mm and 87.0 um in Giemsa and haematoxylin stained specimens compared to 133 um and 123 um respectively. It is very much narrower and the excretory pore is situated at a distance of 34.5% of the total length from the anterior end compared to 60.2 % in D. ramachandrani.
ACKNOWLEDGEMENTS
The authors are grateful to Dr. Douglas J. Gould, Chief, Department of Medical Entomology, SEATO, Medical Research Laboratory, Bangkok, Thailand who initiated the project and to Dr. George F. deWitt, Director, Institute for Medical Research, Kuala Lumpur, Malaysia, for permission to publish this paper. 77
J. W. MAK and R. E. WEAVER
REFERENCES ANDERSON, R. C. and BAIN, O. (1976) Keys to genera of the order Spirurida. Part 3. Diplotriaenoidea, Aproctoidea and Filarioidea. In CIH Keys to the Nematode Parasites of Vertebrates (eds R. C. Anderson, A. G. Chabaud and S. Willmott) Commonwealth Agricultural Bureaux: Farnham Royal, Bucks, pp. 59-116. BUCKLEY, J. J. C. and EDESON, J. F. B. (1956) On the adult morphology of Wuchereria sp. {malayi?) from a monkey (Macaca irus) and from cats in Malaya, and on Wuchereria pahangi n. sp. from a dog and a cat. Journal of Helminthology, 30, 1-20. CHABAUD, A. G. and ANDERSON, R. C. (1959) Nouvel essai de classification des filaires (superfamille des Filarioidea) II. 1959. Annales de Parasitologie, 34, 64-87. DENNIS, D. T. and KEAN, B. H. (1971) Isolation of microfilariae: report of a new method. Journal of Parasitology, 57, 1145-1146. MULLIN, S. and BALASINGAM, S. (1973) Dunnifilaria ramachandrani gen. n., sp. n. (Nematoda: Filarioidea) from the long-tailed giant rat (Rattus sabanus) in Malaysia. Proceedings of the Helminthological Society of Washington, 40, 47-49. Received 10 November, 1977.
78
