NOTE ON A NEW KEY OF THE GENUS PROCAMALLANUS WITH A HISTORICAL REVIEW

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Japan, J. Med. Sci. Biol., 19, 165-170, 1966 ON A NEW KEY OF THE GENUS PROCAMALLANUS WITH A HISTORICAL REVIEW The genus Procamallanus was created by Baylis, 1923 with P. laeviconchus (Wedl, 1862) as its type. While creating this genus he also described in 1923 Procamallanus spiralis from a single male specimen from a siluroid fish Hetrobranchus anguillaris from the Nile river at Cairo. Later in the same year Baylis (1923 b) expanded the description by adding detailed measurements of the female but certain details relating to the caudal papillae of the male were lacking. In 1927, Fujita from Japan described P. parasilum from a siluroid fish. Travassos (1928) added three more species to the genus Procajnallanus namely P. inopinatus, P. iherengi and P. rarus from siluroid fishes in Brazil. Baylis (1929) discovered P, xenopoides from a betrachian in Egypt. Agrawal (1930) described P. mehrii from an Indian siluroid fish which was redescribed by Khera in 1955. Tornquist (1931) redescribed the species (P. spiralis), cited its new host (Host being Crenidens forskalli, Sargus noct and Chilinus trilobatus) and added details of the morphological and cytological features of both male and female worms. Tornquist (1931) also published a monograph on Cucullanidae and Carnallanidae in which he described P. sphaerochonchus from a siluroid fish in Europe. Vaz and Pereira (1934) described two species P. hilari and P. amaradi from Brazil fishes. In 1935, Pearse adde P. kerri from Siamese fish basing his descriptions on a single female. In the same year, i. e., 1935, six more species were added to the genus Procamallanus, three of these, P. wrighti, P, fariasi and P. barroslomai by Pereira from Brazil, P. sigani by Yamaguti from Japan, P. fulvidraconis by Li from China and P. planoratus by Kulkarni from India, all from siluroid fishes. In the subsequent year Pereira added one more species, P. cearnsis from Brazil. In 1937 P. solmei was described by Southwell and Krishner from a betrachian (toad) in South Africa. Johnston and Mawson (1940) reported another species, P. murreyensis from an Australian fish. In 1941, Yamaguti described P. lonis from a Japanese fish. In 1946, P. pereirari another species was described by Annereaux who developed a key, for the 22 existing species of the genus Procamallanus. His paper included the first report of the genus in North America. In 1948, P. brevis was added by Kung from a betrachian host in Africa. In the year 1952, Olsen published a paper which was concerned with P. pereirai and also divided the genus into Procamallanus and Spirocamallanus. He also described a new species which he placed in the new genera Spirocamallanus. The genus Procamallanus was restricted to those species having a smooth buccal capsule lining. The new genus was characterised by spiral thickening on the inner wall of the buccal capsule. In 1953, Winter reviewed the literature on these nematodes and redescribed Spirocamallanus spiralis. He worked on parasites from Mexico and based his account on females only. In 1955, Sogandares-Bernal established a new host-record for S. pereirai from Lousiana. In 1955 S. Khera described two new species of Procamallanus, P. aspiculus from B. bagarius and P. gubernaculus from Rita rita. Ali (1956) in India modified the key developed by Annereaux. He developed 165

166 Vol. 19 a key for the identification of the species of the original genus Procamallanus, but the key is based on certain measurements which do not correspond to some of those previously reported. The worms described by Ali were, P. heteropneustes, P. clarius, P. singhi, P. hydrabadensis and P. viviparus all recorded from siluroid fishes of fresh water of India. In 1960, he again reported two more new species of Procamallanus namely P. globochonchus and P. ophiocephalus from Rita hastata and Ophiocephalus giving a fresh key to the genus Procamallanus. In the same year i. e., 1960 Yeh Liang Sheng wrote on a reconstruction of the genus Camallanus in which he divided Carnallanidae into two subfamilies, Procamallaninae and Spirocamallanus while the latter includes Camallanus (Railliet and Henry, 1915), Camallanides (Baylis and Daubney, 1922), Procamallanus (Yorke and Maplestone, 1926), Piscilanid gen. nov, Serpenema, gen. nov, and Zeylanema Yeh, 1960). In 1965, Sinha and Sahy wrote on the validity of the genus Indocamallanus Chakravarty et all and discovered P. (monospiculus) devendri n. sp.; in the same year Sinha and Sahay reported a new Spirocamallanus from E. Vacha. Sahay reported in late 1965 Spiro Camallanus Chauhani n. sp.; from Mystus cavasius. Key to the Genus Procamallanus The key to the genus Procamallanus has been given by Annereaux (1946), Khera (1955), Aui (1956; 1960). If the key is amended in view of the recent works of Olsen (1952) and Yeh Liang Sheng (1960), it would facilitate the study of known species of the genus. The author does not agree with Khera (1955( who included P. gubernaculus in Procamallanus and not in Spirocamallanus. The author also fails to understand why Aui (1960) included P. gubernaculus Khera, 1955 in Procamallanus when P. gubernaculus is provided only with one spicule. Ali (1960) seems to be mis-informed about it as is evident from his key in which he mentions P. gubernaculus as having two spicules, the right one measuring 0.29 mm and the left one. 0.06 mm. It has been clearly mentioned in the account given by Khera (1955) published in Anales de la Escuela Nacional de Ciencias Biologicas, VIII (3-4) : 243-252 Mexico that the spicules reffered to above is without the left spicule. An accessory piece cannot be regarded as a spicule. Besides the inclusion of all the Spirocamallanids in the subgenus Procamallanus does not seem to be sound. Ali gave his key in 1956 modifying the key developed by Annereaux and he described five new species of Procamallanus three of which should have been placed in Spirocamallanus. Most probably Ali did not note the paper by Olsen. The following key is a modification on Annereaux's, Khera's and Ali's key to the genus Procamallanus. P. kerri Pearse, 1933 known from female specimens only has not been included in the key. 1. Subgenus-Aspiculus Au, 1960. Key to the Species of the Genus Procamallanus Buccal capsule with semi-lunar cuticular thickenings and a pair of golf-club-shaped-projections, spicules absent...p. aspiculus, Khera 1955. 2. Sub-genus-Monospiculus Ali, 1956. 1. Buccal capsule without spiral thickenings...2 2. With only one spicule...p slomei Southwell and Krishner 1937 With one spicule+y shaped gubernaculum+tail of female bifur-

1966 167 cated at tip...p. (monospiculus) devendri n. sp. Sinha and Sahay 1965 Gubernaculum otherwise+female tail without bifurcation...p. (monospiculus) heteropnuesti Chakravarty et al. 1961. 3. Subgenus-Isospiculus Ali, 1956. 1. Buccal capsule without spiral thickening, Spicule 0.45 mm long, 8 caudal papillae,...p. wrighti Pereira, Buccal capsule without spiral thickenings & spicule shorter...2 2. 7 pairs of caudal papillae, spicule 0.05 mm long...p. cearnsis Pereira, 8 pairs of caudal papillae, spicules 0.12 mm long...p. inopinatus Travassos, 1928. 13 pairs of caudal papillae, spicules 0.11 mm long...p. xenopoidis Baylis,1929. 4. Subgenus \Procamallanus Baylis, 1923 and Ali, 1956. 1. Buccal capsule without spiral thickenings...2 2. Gubernaculum present...p. annulatus Yamaguti, 1954. Gubernaculum absent...3 3. Spicules short, more than 12 pairs of caudal papillae...4 Spicules longer, 12 pairs or fewer Caudal papillae...5 4. Vulva equatorial or subequatorial, female tail with caudal spines. 6 Vulva near the posterior end, female tail without caudal spines...7 5. Caudal papillae 12 pairs, female tail with caudal spines...p. sphaeroconchus, Tornquist, 1931. Caudal papillae 9 pairs, female tail without caudal spines...8 6. Right spicule 0.15 mm long, left spicule 0.05 mm long. 14 pairs of caudal papillae...p, laeviconchus (Wedl) Baylis, 1923 Right spicule 0.089 mm long, left spicule 0.039 mm long, 16 pairs of caudal papillae...p. brevis Kung, 1948 7. Right spicule less than twice as long as left spicule (R. S-0.07; L. S-0.04 mm) 13 pairs of caudal papillae...p. heteropnuestes Au, 1956. Right spicule three times as long as left spicule (R. S-0.085 mm ; L. S-0.029 mm), 14 pairs of caudal papillae...p. clarius Ali, 1956 8. Right spicule 0.04 mm long ; L, spicule 0.12 mm ; circum anal papillae three pairs...p. sigami Yamaguti, Right spicule 0.05 mm ; L. Spicule-0.22 mm long circum anal papillae absent...p. lonis Yamaguti, 5. Subgenus - Spirocamallanus Subgenus Nov. 1. Buccal capsule with spiral thickenings...2 2. Single spicule...3 Isospiculus...4 Spicules unequal...5 3. Buccal capsule with lateral finger shaped thickenings parallel to it, gubernaculum present...3a 3a. Tail of female bluntly rounded with vulvular appendage...p. gubernaculus Khera, 1955. 3b. Tail of female bifurcated without vulvular appendage...s. chauhani n. sp. Buccal capsule otherwise...p. parasiluri Fujita, 1927. 4. Spicules 0.25 mm long, 11 pairs of caudal papillae...p. mehrii Agarwal, 1930. Spicules 0.12 mm long, 8 pairs of caudal papillae...p. planoratus Kulkarni,

168 Vol. 19 5. Spicules similar in shape...6 Spicules dissimilar in shape...7 6. Gubernaculum present...p. bagarii Karve and Naik, 1951. Gubernaculum absent...8 7. Right spicule three times longer than the left (R. S-0.18 ; L. S- 0.06 mm), caudal alae meet ventrally...p. viviparus, Ali, 1956. Right spicules more than three times longer than the left, caudal alae lateral...9 8. Inner thickenings of the buccal capsule with few thick spirals...10 Inner thickenings of the buccal capsule with 8 or more thick spirals...11 9. Circum anal papillae present...12 Circum anal papillae absent...13 10. Right spicule 0.53 mm left spicule 0.25 mm, spirals in males 4-5, female 14-16...P. Right spicule 0.43 mm, left spicule 0.17, spirals in males 3-4, female unknown...p. iheringii Travassos et al. 1928. rarus Travassos et al. 1928. 11. Spicules long, right spicule at least 0.2 mm...14 Spicules much smaller...15 12. 2 pairs of circumanal papillae...p. ophicephalus A1i, 1956. 3 pairs of circumanal papillae...16 13. 10 pairs of caudal papillae, right spicule four times longer than left (R. S-0.2 mm ; L. S-0.05 mm)...p. hydrabadensis Ali, 1956. 11 pairs of caudal papillae right spicule five times longer than the left (R. S-0.2 mm, L. S-0.04 mm) female tail with caudal spine P. singhi Ali, 1956 14. 7 pairs of caudal papillae...17 More than 7 pairs of caudal papillae...18 and P. murrayensis Johnston and Mawson, 1940. 15. Spicules 0.1 mm or more, 9 or more caudal papillae...19 Spicules less than 0.1 mm, fewer caudal papillae...20 16. Right spicule five times longer than left (R. S-0.15 mm L. S-0.03 mm), 13 pairs of caudal papillae...p. globoconchus Ali, 1960. Right spicule eight times longer than left (R. S-0.24 mm ; L. S- 0.03 mm) 14 pairs of caudal papillae...p. fulvidraconis Li, 17. Caudal alae meet ventrally, 2 pairs of sessile adanals and one pair of adanal...p. monotaxis Olsen, 1952. 18. 9 pairs of caudal papillae, female tail with caudal spines...p. pereirai Annereaux, 1946. 14 pairs of caudal papillae, fcmale tail without caudal spines...p. amarali Vaz and Pereira, 1934. 19. Right spicule 1/3 longer than left (R. S-0.15 mm L. S-0.10 mm), 7 caudal papillae...p, spiralis Baylis, 1923. Right spicule less than 1/3 longer than left (R. S-0.11 mm ; L. S- 0.10 mm) 9 caudal papillae...p fariasi Pereira, 20. 8 pairs of caudal papillae, right spicule 0.08 mm, left spicule 0.06 mm...p. hilari Vaz and Pereira, 1934. 6 pairs of caudal papillae, right spicule 0.06 mm left spicule 0.05 mm...p. barroslimai Pereira,

1966 169 Summary: Various workers have given the key to the genus Procamallanus. Such keys have been modified after proper amendment in view of the work of Olsen (1952) and Yeh Liang Sheng (1960). A new subgenus Spirocamallanus has been added to the existing list of four subgenera created by Ali in 1960. The forms having spiral thickenings in the buccal capsule have been referred to this new subgenus. P. parasiluri Fujita, 1927 formerly kept in the subgenus Monospiculus by Ali, 1960; P. mehrii Agrawal, 1930 and P, planoratus Kulkarni, 1935 both referred to in the sugenus Iso spiculus by Au, 1956 and those forms formerly placed in the subgenus Procammallanus with the spiral thickenings in their buccal capsule have been transferred in the new subgenus. S. chauhani sp. nov. has been referred to this new subgenus. P. (monospiculus) devendri sp. nov. Sinha and Sahay, 1965 and P. (monospiculus) heteropneustei Chakravarty et al., 1961 have been kept in the subgenus Monospiculus created by Ali in the year 1956. The author wishes to express his thanks to Dr. Devendra Prasad, Professor of Zoology, S cience College Patna-5 Patna University (Bihar) under whose guidance this study was made. The author is also indebted to Dr. S. M. Ali, Professor of Zoology, Marathwada University, Aurangabad, Dr. S. Khera, Reader, University of Jodhpur for their constant encouragement valuable suggestions. The author records his sincere thanks to the authorities of the University Grants Commission, New Delhi for kindly bestowing on him a research grant-in-aid and to Dr. K. C. Bose Prof, of Zoology, Ranchi University, for providing the necessary facilities. The author is also indebted to Dr. Y. Komiya, the National Institute of Health, Tokyo Japan, f or very kindly going through the manuscript. REFERENCES AGARWAL, M. P. (1930) : A new nematode, Procamallanus mehrii nov. sp., from a local siluroid fish Wallago attu. Alld. Univ. Stud., VI, 59-64. ALI, S. M. (1956) : Studies on the nematode parasites of fishes found in Hydrabad state. Indian. J. Helminthol, 8, 1-83. (WL. 9939g). ANNEREAUX, R. F. (1946) : A new nematode Procamallanus pereirai with a key to the genus. Trans. Am. Micr. Soc., LXV, 299-303. BAYLIS, H. A. (1939) : The fauna of British India, including Ceylon and Burma. Nematoda, Vol. I and II. Taylor and Francis Ltd. Red Lion Court, Fleet Street, London. BAYLIS, H. A. (1924) : Note on Procamallanus spiralis Baylis, 1923 (Ndmatoda), Parasitology, 15, 137-138. JOHNSTON, T. H. AND MAWSON, P. M. (1951) : Some nematode parasitic in Australian fresh water fish. Trans. Roy. Soc. Austr., 64, 95-100. KARVE, J. N. AND NAIK, G. G. (1951) : Some paratic nematodes of fishes. J. Univ. Bombay, 19, 1-37. (W. L. 11594c) KHERA, S. (1956) : On some species of Procamallanus Baylis, 1923 from INDIA. Annals de la Escuela National de Cencias Biologicas., 8, 243-252 (W. L. 7196). KULKARNI, R. B. (1935) : A second species of Proeamallanus Baylis, 1923 from India. Proc. Ind. Acad. Sci., 11, 29-32. LI, H. C. (1935) : The taxonomy and Early development of Procamallanus fulviraconis nov. sp. J. Parasitol., XXI, 103-113. OLSEN, L. S. (1952) : Some nematodes parasitic in Marine fishes. Pub. Inst. Mar. Sci. Univ. Text., 11, 173-215 (W. L. 1736f). PEARSE, A. S. (1933) : Parasites of Siamese fishes and Crustaccans. J. Siam. Soc. Nat. Hist.,

170 Vol. 19 Suppl. IX, 179-191. PEREIRA, G. (1935) : Ascaridata e Spirurata parasitos de peires nordeste Brasileiro, Arch. Inst. Biol., 6, 53-62. SINHA, D. P. AND SAHAY, U. (1965) : On a new species of Procamallanus (Camallanidae, Nematoda) with a discussion on the validity of the genus Indocamallanus Chakravarty et al., Zoologischer Anzeiger (sent). SINHA, D. P. AND SAHAY, U. (1965) : On a new species of Spirocamallanus Olsen, 1952 (Camallanidae, Procamallaninae, Nematoda) from Indian. J. Helminthol, 17, 49-53. SAHAY, U. (1965) : On a new nematode of the genus Spirocamallanus with a review of the genus. Indian. J. Helminthol. (sent for publication). TRAVASSOS, L, ARTIGAS, P. E. AND PEREIRA, R (1928) : Fauna Helminthologiea dos Peixas de Agna Doce do Brazil. Arch Inst. Biol., 1, 5-63. YAMAGUTI, S. (1954) : Parasitic worms mainly from Celebes Pt. 9 Nematodes from fishes. Acta. Med. Okayama, 9, 122-133. YAMAGUTI, S. (1961) : Systema Helminthum Vol III part I and II Inter Science Publishers, INC New York, London. YEH-LIANG-SHENG (1960) : J. Helminthol., 34, 117-124. YORKE, W. AND MAPLESTONE, P. A. (1926) : The nematode parasites of the vertebrates J. A. Churchill, 7 Great Marlborough Street. Department of Zoology, University of Ranchi, Bihar, India UMAPATI SAHAY Received : February 9th, 1966