Ranjeet Singh 1 & Harpreet Kaur 2 * Introduction

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1 Biologia 70/1: 85 93, 2015 Section Zoology DOI: /biolog Two new and one already known species of the genus Thelohanellus (Myxozoa: Myxosporea: Bivalvulida) parasitizing fresh water fishes in wetlands of Punjab, India Ranjeet Singh 1 & Harpreet Kaur 2 * 1 Department of Zoology and Environmental Sciences, Punjabi University, Patiala, Punjab , India 2 Department of Zoology and Environmental Sciences, Punjabi University, Patiala, Punjab , India; harpreetbimbra@gmail.com Abstract: A study of parasites of freshwater fishes in Harike and Ropar wetlands of Punjab of India revealed the presence of two new and one already known myxozoan species belonging to the genus Thelohanellus Kudo, 1933 infecting gills and caudal fin of two carps and gall bladder of a catfish in the wetlands of Punjab, India. Plasmodia of the first species, T. lamelliformis sp. n. are found infecting primary gill lamellae of Catla catla (Hamilton, vern. thail, spores measure µm, are elongate pyriform, tapering anteriorly in valvular view, having truncated anterior end and broad rounded posterior end. Shell valves are stained dark blue (with iron-haematoxylin) in the middle as well as in the posterior part of the spore body. Anteriorly, a prominent bent is present on one side of the shell valve. Polar capsule is broadly pyriform, measures µm and is placed in the middle of spore body cavity with a distinct neck leading to the outside. Plasmodia of the second species, T. rohi sp. n. are found infecting caudal fin of Labeo rohita (Hamilton, vern. rohu, spores measure µm, are elongately oval in valvular view having narrow, blunt anterior end with a prominent pore and broad rounded posterior end. A knob-like structure is present at the anterior end of the spore. Inner valve stains dark-blue (with iron-haematoxylin) in the middle and posterior part of the spore body. Polar capsule is oval to spherical in shape, measures µm, is placed anteriorly and occupies nearly one third of the spore body cavity. Plasmodia of the third species, T. wallagoi Sarkar, 1985 infect primary gill lamellae of Wallago attu (Bloch & Schneider, 1807), spores measure µm, are pyriform in valvular view, having bluntly pointed anterior end and rounded posterior end. Parietal folds are absent. Polar capsule is pyriform, measures µm, is eccentrically placed with narrow pointed anterior end. Earlier, this species was reported from West Bengal infecting gall bladder of Wallago attu. Key words: myxosporeans; Harike wetlands; Ropar wetland; India; freshwater fishes Abbreviations: LS Length of spore; WS Width of spore; LPC Length of polar capsule; WPC Width of polar capsule; NC Number of coils of polar filaments; SD Standard deviation. Introduction In Punjab, there are 12 natural, 10 man-made wetlands covering a 15,500 ha area and only three main wetlands are included in the Ramsar list of International importance, i.e., Harike, Kanjali and Ropar wetlands. These wetlands have extremely rich biodiversity as they support a variety of plant and animal life. Harike wetland is the largest freshwater wetlands in northern India occupying an area of 4,100 ha, harbouring 16 species of freshwater fishes (Punjab State Council for Science and Technology Chandigarh 2002). Ropar wetland is an important habitat of many species and has tremendous ecological value, spreads over 1,365 ha, and support as many as 35 species of fishes. These fishes are vulnerable to various parasitic infections, out of which Myxozoa is emerging as the major group. Phylum Myxozoa includes four malacosporean and 2,180 myxosporean species of a total of 62 genera (Lom & Dykova 2006). Three more genera (Soricimyxum, Gadimyxa, Thelohanelloid) with the type species S. fegati Prunescu et al., 2007 from liver of Sorex araneus L., 1758, G. atlantica Koie et al., 2007 from urinary system of Gadus morhua L., 1758 and T. bengalensis Sarkar, 2009 from gall bladder of Arius sagor (Hamilton, have been described (Prunescu et al. 2007; Koie et al. 2007; Sarkar 2009). The Phylum Myxozoa has been studied by only limited number of workers in the Indian subcontinent. Most of the work on this phylum has been done on fresh water as well as marine fishes of two states, West Bengal and Andhra Pradesh. In addition, some work has been carried out in north India by Gupta & Khera (1987, 1988a d, 1989a, b, 1990, 1991). * Corresponding author c 2015 Institute of Zoology, Slovak Academy of Sciences

2 86 R. Singh &H.Kaur Since 2007, there has been an addition to the existing number of myxosporean species in India, particularly those belonging to Myxobolus. Recently, Kaur & Singh (2008, 2009, 2010a, b, 2010/2011, 2011a f, 2012a) have described 17 new species of the genus Myxobolus and one new species of the genus Triangula from freshwater fishes in wetlands of Punjab. Kaur & Singh (2012b) also compiled and published a synopsis of 131 nominal species of Myxobolus Buetschli, 1882 (Myxozoa: Myxosporea: Myxobolidae) reported from India and a revised dichotomous key of 59 genera of the Phylum Myxozoa (Class Myxosporea). There are very few numbers of species of the genus Thelohanellus reported all over the world. In a monograph, Lom & Dykova (1992) listed 39 species of this genus. Basu & Haldar (1999) gave a checklist of its different species described from Indian fishes. Basu et al. (2006) provided a synopsis of 32 Indian species belonging to the genus Thelohanellus, including one new species T. disporomorphus infecting Indian major carp, Cirrhina mrigala (F. Hamilton,. Kalavati & Nandi (2007) gave a compilation of 27 species of the genus Thelohanellus infecting Indian fishes. Singh & Kaur (2014) have recorded two new species, i.e., T. kalavate and T. kalbensi from caudal fin of Cirrhina reba (Day, 1878) and gills of Labeo calbasu Hamilton,1822, respectively. Recently, Zhang et al (2013) gave a synopsis of 108 nominal species of the genus Thelohanellus reported globally. During the present study on fishes of Harike and Ropar wetlands of Punjab, two new species, T. lamelliformis sp. n. and T. rohi sp. n., and one already known species T. wallagoi Sarkar, 1985 were collected from fins and gills of Catla catla, caudal fins of L. rohita and gills of Wallago attu, respectively. The description has been prepared in accordance with the guidelines of Lom & Arthur (1989). Material and methods Fishes collected from Harike and Ropar wetlands were frozen in an icebox and brought to the laboratory and examined for myxozoan infections. Plasmodia when found were removed and teased on slide and were removed, placed on microscopic slides and examined in the light microscope under 100 oil objective (Magnus inclined Trinocular microscope MLX-Tr) for the presence of myxospores. Fresh spores were treated with 8% KOH for the extrusion of polar filaments. For permanent preparation, air-dried smears were stained with Ziehl-Neelsen and iron-haematoxylin. Drawings were made from stained material with the aid of camera lucida. Measurements of spores were done with the aid of a calibrated ocular micrometer. All measurements are presented in µm as range values followed by mean ± SD in parentheses. Fig. 1. Thelohanellus lamelliformis sp. n.: A, D Spore stained in Ziel-Neelsen (valvular view); B, E Spore stained in ironhaematoxylin (extruded polar filament); C Spore in side view.

3 Two new species of Thelohanellus 87 Table 1. Measurements (µm) and ratio of Thelohanellus lamelliformis sp. n. Characters Range Mean values SD LS WS LPC WPC Ratio: LS/WS 2.0 NC 6-7 Parietal folds Absent Results and discussion Thelohanellus lamelliformis sp. n. (Fig. 1) Plasmodia. Minute, present around gill lamellae. Each plasmodium contains spores. Spore description (Table 1). Measurements are based on 8 10 spores in frontal view. The spores are histozoic, measuring µm, elongately pyriform, tapering anteriorly in valvular view having truncated anterior end and broad rounded posterior end. Shell valves are thick, smooth, symmetrical and measure 0.5 µm in thickness. Shell valves are stained dark blue (with ironhaematoxylin) in the middle as well as in the posterior part of the spore body. Anteriorly, a prominent bent is present on one side of the shell valve. Parietal folds are absent. Polar capsule is broadly pyriform, measures µm and is placed in the middle of spore body cavity with a distinct neck leading to the outside. Polar filament form 6 7 coils arranged perpendicular to the polar capsule axis. Polar filament is thick, ribbonlike, measuring 50.8 µm in length and extrude through the anterior tip of the spore. Two capsulogenic nuclei measuring 1.5 µm and 0.9 µm in diameter are located just beneath the polar capsule. Sporoplasm is agranular, homogenous and occupies the whole extracapsular space behind the polar capsule and contains one sporoplasmic nucleus measuring µm indiameter. Iodinophilous vacuole is absent. Table 2. Thelohanellus lamelliformis sp. n. in comparison to morphologically similar species (measurements are in micrometers). Species Host Site of infection Locality Spore Polar capsule T. lamelliformis sp. n. (present study) T. catlae Chakravarty & Basu, 1948 T. seni (Southwell & Prashad, 1918) Chakravarty & Basu, 1948 T. gangeticus Tripathi, 1952 T. andhrae Qadri, 1962 T. chrysopomati Lalitha Kumari, 1969 T. potaili Lalitha Kumari, 1969 T. coeli Sarkar & Mazumdar, 1983 T. opthalmicus Haldar et al., 1983 T. wallagoi Sarkar, 1985 T. valeti Fomena & Bouix, 1987 T. assambai Fomena et al., 1994 T. chandannagarensis Basu & Haldar, 2003 T. endodermitus Mukhopadhyay & Haldar, 2004 T. zahrahae Szekely et al., 2009 Catla catla (F. Hamilton, gill lamellae Harike wetland, Punjab Catla catla branchiae West Bengal Catla catla branchiae West Bengal Chela bacaila (Hamilton, Labeo fimbriatus Bloch, 1795 Barbus chrysopoma Valenciennes, 1842 Labeo potail Sykes, 1839 Tachysurus tenuispinis (Day, 1877) Catla catla Wallago attu Bloch & Schneider, 1801 Barbus jae Boulenger, 1903 Barbus aspilus Boulenger, 1907 muscles West Bengal gills Andhra Pradesh gill contents Andhra Pradesh fin Andhra Pradesh gall bladder West Bengal internal musculature and sclera of eye West Bengal (12.5) (7.5) (6.5) (3.2) gall bladder West Bengal intestine, muscles Africa ; ; Labeo sp. Africa Catla catla gill lamellae West Bengal Labeo rohita F. Hamilton, 1822 Barbonymus gonionotus (Bleeker, 1850) undersurface of scales West Bengal gills Malaysia

4 88 R. Singh &H.Kaur Fig. 2. Thelohanellus rohi sp. n.: A, D Spore stained in Ziel-Neelsen (valvular view); B, E Spore stained in iron-haematoxylin (extruded polar filament); C Spore in side view. Taxonomic summary of T. lamelliformis sp. n. Family: Myxobolidae, Thelohan, 1982 Genus: Thelohanellus Kudo, 1933 Plasmodia: Minute and present in gill lamellae. Spores: Spores are elongatly pyriform, tapering anteriorly in valvular view having truncated anterior end and broad rounded posterior end. Shell valves are stained dark blue (with Iron-haematoxylin) in the middle as well as in the posterior part of the spore body. Anteriorly, a prominent bent is present on one side of the shell valve. Type host: Catla catla (Hamilton, vern. thail. Type locality: Harike wetland, Punjab, India. Type specimens: Paratypes are spores stained in Ziehl-Neelsen and Ironhaematoxylin, deposited in the museum of Department of Zoology, Punjabi University, Patiala, India- Slide no. C/P/ZN and C/P/IH Site of infection: Gill lamellae. Prevalence of infection: 1% (1/10). Clinical symptomatology: None. Etymology. The specific epithet lamelliformis highlights the location of plasmodia in the primary gill lamellae. Differential diagnosis. The studied species was compared to 14 representatives of the genus Thelohanellus infecting fish (Table 2). It differs from all of them by morphological and morphometric characters. The novel species possess elongately pyriform in shape with tapering, truncated anterior end and broad rounded posterior end, like T. chrysopomati, T. gangeticus, T. coeli, T. andhrae, T. potaili, and T. valeti but can be differentiated from them in having a prominent bent in the shell valves on one side at the anterior. However, in T. coeli, anterior end is narrow with slightly angled neck but presence of median longitudinal ridge and much wider (LS/WS: 1.7) spores in T. coeli demarcates them from the present species under consideration (LS/WS: 2.0). The parietal folds are absent in the present species, this differentiate it from T. chrysopomati, T. andhrae and T. potaili in which 4 6 parietal folds are present at the posterior region of spores. Furthermore, the polar capsule in the present species is broadly pyriform with a distinct neck leading to outside and is placed in the middle of spore body cavity in contrast to anteriorly situated larger polar capsule in T. gangaeticus, T. andhrae and T. valeti. Moreover, in the present species shell valves are stained dark blue (with iron-haematoxylin) in the middle as well as in the posterior part of the spore body. Polar filament is thick and ribbon-like in the present species. In view of the above differences, we assign a myxozoan parasite of Catla catla from Harike wetland to a new species of the genus Thelohanellus. Thelohanellus rohi sp. n. (Fig. 2) Plasmodia. Small, white, spherical to round, mm in diameter, located on the caudal fin in numbers 5 6 plasmodia per fin. Each plasmodium contains 9 11 spores. Spore description (Table 3). Measurements based on 7 9 spores in frontal view. The spores are histozoic,

5 Two new species of Thelohanellus 89 Table 3. Measurements (µm) and ratio of Thelohanellus rohi sp. n. Characters Range Mean values SD LS WS LPC WPC Ratio: LS/WS 1.8 NC 4 5 Parietal folds Absent measure µm, elongately oval in valvular view having narrow, blunt anterior end with a prominent pore and broad rounded posterior end. A knoblike structure is present at the anterior end of the spore. Shell valves are thick, smooth, symmetrical and measure 0.8 µm in thickness. Inner valve stains darkblue (with iron-haematoxylin) in the middle and posterior part of the spore body. Parietal folds are absent. Polar capsule is oval to spherical in shape, measure µm, is placed anteriorly and occupies nearly one third of the spore body cavity. Polar filament forms 4 5 coils arranged perpendicular to the polar capsule axis. Polar filament is thick, thread-like, measures 62 µm in length and extrudes through a prominent pore on a knob-like structure at the anterior end. One capsulogenic nucleus is present beneath the polar capsule and measures µm in diameter. Sporoplasm occupies Table 4. Thelohanellus rohi sp. n. in comparison to morphologically similar species (measurements are in micrometers). Species Host Site of infection Locality Spore Polar capsule T. rohi sp. n. (present study) T. niloticus Gurley, 1893 T. rohitae (Southwell & Prashad, 1918) Chakravarty, 1943 T. nikolski Akhmerov, 1955 Labeo rohita F. Hamilton, 1822 Labeo niloticus (L., 1758) Labeo rohita, Labeo bata F. Hamilton, 1822 Cyprinus carpio haematopterus Martens, 1876 T. shortii Qadri, 1967 Labeo fimbriatus Bloch, 1795 T. opthalmicus Haldar Catla catla et al., 1983 T. bengalensis Sarkar & Raychaudhury, 1986 T. sanjibi Sarkar & Ghosh, 1990 T. sudevi Sarkar & Ghosh, 1990 caudal fin Ropar wetland, Punjab skin of head Nile (Egypt) gills West Bengal fin Amur basin (Russia) fin internal musculature and sclera of eye Andhra Pradesh (12.53) (6.91) West Bengal (12.5) (7.5) Catla catla gall bladder West Bengal (10.95) (6.59) Mystus gulio (Hamilton, Amblypharyngodon mola (F. Hamilton, T. chilkensis Kalavati & Vaidehi, 1991 T. costeae Sakiti, 1997 Labeo senegalensis Valenciennes, 1842 T. ndjamenaensis Kostoıngue et al., 1999 T. bicornei Kabre et al., 2002 T. avijiti Basu & Haldar, 2003 T. endodermitus Mukhopadhyay & Haldar, 2004 T. habibpuri Acharya & Dutta, 2007 T. imphlaensis Hemananda et al., 2010/2011 T. kalbensi Singh & Kaur, 2012 kidney West Bengal (12.52) (8.27) kidney West Bengal (14.0) (5.87) (7.07) (4.2) (6.5) (3.2) (5.42) (3.47) (4.52) (4.0) (5.17) (2.65) Labeo rohita gall bladder Orissa Labeo parvus Boulenger, 1902 Labeo coubie Rüppell, 1832 gill Benin (Africa) (9.4) (5.6) gill Chad (Central Africa) (10.0) (7.3) intestine Burkina Faso (Africa) (13.5) (8.4) (4.8) (2.6) (4.2) (3.2) (7.2) (3.7) Labeo rohita dorsal fin West Bengal Labeo rohita undersurface of scales West Bengal Labeo rohita pectoral fin West Bengal (13.9) (8.5) Labeo rohita gills Imphal, Manipur Labeo calbasu (Hamilton, gills Harike wetland, Punjab (21.33) (9.43) (6.0) (4.9) (10.79) (3.78)

6 90 R. Singh &H.Kaur Fig. 3. Thelohanellus wallagoi: A, D Spore stained in Ziel-Neelsen (valvular view); B, F Spore stained in iron-haematoxylin (extruded polar filament); C Spore showing sporoplasmic nuclei and coiled polar filament; E Spore stained in iron-haematoxylin. the whole extracapsular space behind the polar capsule. Sporoplasm contains two nuclei measuring µm in diameter. Iodinophilous vacuole is absent. Taxonomic summary of T. rohi sp. n. Family: Myxobolidae, Thelohan, 1982 Genus: Thelohanellus Kudo, 1933 Plasmodia: Small, white, spherical to round, mm in diameter and are located on the caudal fin. Spores: Spores are elongately oval in valvular view having narrow, blunt anterior end with a prominent pore and broad rounded posterior end. A knob-like structure is present at the anterior end of the spore. Type host: Labeo rohita (Hamilton, vern. Rohu. Type locality: Ropar wetland, Punjab, India. Type specimens: Paratypes are spores stained in Ziehl-Neelsen and Iron-haematoxylin, deposited in the museum of Department of Zoology, Punjabi University,Patiala, India- Slide no. C/A/ZN and C/I/IH Site of infection: Caudal fin. Prevalence of infection: 25% (3/12). Clinical symptomatology: None. Etymology. The specific epithet rohi highlights the vernacular name of the host fish. Differential diagnosis. The studied species was compared to 17 representatives of the genus Thelohanellus infecting fish (Table 4). It differs from all of them by morphological and morphometric characters. The novel species possesses elongately oval in shape with Table 5. Measurements (µm) and ratio of Thelohanellus wallagoi Sarkar, Characters Range Mean values SD LS WS LPC WPC Ratio: LS/WS 2.0 NC 5 6 Parietal folds Absent blunt narrow anterior end and broad rounded posterior end like T. sudevi and T. chilkensis, but the spores are elongately ellipsoidal with anterior end acuminated in T. sudevi and elongate pyriform with flat tip in T. chilkensis. Shell valves are smooth and symmetrical unlike in spores of T. chilkensis in which inner wall of the spore with a distinct constriction at posterior 1/3 rd formed by 2 indentations on either side. In addition, the polar capsule in the present species is oval to spherical in shape situated at the anterior end of the spore body cavity. It occupies one third of the spore body cavity in contrast to flask-shaped polar capsule occupying more than half in T. chilkensis and is pyriform, occupying less than half in T. sudevi. Furthermore, a knob-like structure with a prominent pore is present at the anterior end of the spore

7 Two new species of Thelohanellus 91 Table 6. Comparison of Thelohanellus wallagoi Sarkar, 1985 in original description and in present species (measurements are in micrometers). Species Host Site of infection Locality Spore Polar capsule T. wallagoi (present study) Wallago attu Bloch & Schneider, 1801 gill lamellae Harike wetland, Punjab T. wallagoi Wallago attu gall bladder West Bengal (9.25) (4.85) (5.47) (2.71) and inner shell valve stained dark blue (with ironhaematoxylin) in the middle and posterior part of the spore body in species under consideration which differentiates it from the above mentioned species. In view of the above differences, we assign a myxozoan parasite of L. rohita from Ropar wetland to a new species of the genus Thelohanellus. Thelohanellus wallagoi Sarkar, 1985 (Fig. 3) Plasmodia. Minute, present around gill lamellae. Each plasmodium contains spores. Spore description (Table 6). Measurements based on 8 10 spores in frontal view. The spores are histozoic, measure µm, pyriform in valvular view having bluntly pointed anterior end and rounded posterior end. Shell valves are thin, smooth, symmetrical and measure 0.16 µm in thickness. Parietal folds are absent. Polar capsule is pyriform, measures µm, eccentrically placed with narrow pointed anterior end. Polar filament forms 5 6 coils arranged perpendicular to the polar capsule axis. Polar filament is thin, thread-like measure 25 µm in length and extrudes from the anterior end of the spore. Sporoplasm is agranular, homogenous occupying the whole extracapsular space behind the polar capsule and contains two nuclei measuring (0.8 ± 0.14) µm in diameter. An iodinophilous vacuole is present measuring 2.0µm in diameter (Table 5). Remarks. The present observations (LS/WS: 2.0) on T. wallagoi Sarkar, 1985 are in conformity with the original description (LS/WS: 1.9) except some minor variations in the size of the polar capsule. Earlier, this parasite was recorded from gall bladder of Wallago attu Bloch & Schneider, A new location (gill lamellae) and a new locality (Harike wetland) are recorded for this parasite. Acknowledgements The authors acknowledge the financial support by University Grants commission (UGC), Govt. of India (MRP). References Acharya S. & Dutta T Thelohanellus habibpuri sp. n. (Myxozoa: Bivalvulida) from the tropical freshwater fish rohu, Labeo rohita (Hamilton-Buchann, 1882) in West Bengal, India: Light and electron microscope observations. Anim. Biol. 57 (3): DOI: / Akhmerov A.K Ways of the origin of myxosporidian species of the genus Thelohanellus Kudo from Amur wild carp. Dokl. Akad. Nauk SSSR 105: (in Russian) Basu S. & Haldar D.P Thelohanellus bifurcata n. sp. a new species of the genus Thelohanellus from hybrid carps and checklist of the species of the genus described from Indian fishes. Proc. Zool. Soc. 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Protistology 6(4): Kabre G.B., Sakiti N.G., Marques A. & Sawadogo I Thelohanellus bicornei sp. n., (Myxosporea, Bivalvulida) a gill parasite of Labeo coubie Ruppel, 1832 (Osteichthyes, Cyprinidae) from Burkina Faso, West Africa. Parasite 9(3): Kalavati C. & Nandi N.C Handbook of Myxosporidean parasites of Indian fishes. ZSI, India, Kolkata. PMID: Kalavati C. & Vaidehi J A new myxosporidian, Thelohanellus chilkensis n. sp. from the gills of the common carp, Labeo rohita in Chilka lake, Orissa, India. Uttar Pradesh J. Zool. 11 (1): Kaur H. & Singh R Observations on one new species of genus Myxobolus M. naini and rediscription of M. magauddi recorded from freshwater fishes of Kanjali Wetland of Punjab, India, pp In: Proc. 20 th Natl. Congr. Parasitol., Department of Zoology North Eastern Hill University, Shillong (Meghalaya). 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Two new species of Myxobolus (Myxozoa: Myxosporea: Bivalvulida) infecting an Indian major carp in Ropar and Kanjali wetlands (Punjab). J. Parasit. Dis. 35 (1): DOI: /s Kaur H. & Singh R. 2011c. Myxobolus harikensis sp. nov. (Myxozoa: Myxobolidae) infecting fins of Cirrhina mrigala (Ham.) an Indian major carp in Harike Wetland, Punjab. Parasitol. Res. 109 (6): DOI: /s x Kaur H. & Singh R. 2011d. Two new and one already known species of Myxobolus (Myxozoa: Myxosporea: Bivalvulida) infecting gill lamellae of Indian major carp fishes in Ropar and Harike wetlands (Punjab), pp In: Bamdyopadhyay P.K. & Mandal D.R. (eds), Proc. 22 nd Natl. Congr. Parasitol., Oct. 30 Nov. 1, 2010 Advances in Parasitology: A Novel Approach Towards a Disease Free World, Department of Zoology, University of Kalyani, Kalyani, West Bengal, India, 343 pp. Kaur H. & Singh R. 2011e. Two new species of Myxobolus (Myxozoa: Myxosporea: Bivalvulida) infecting Indian freshwater fishes in Punjab Wetlands. Parasitol. 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9 Two new species of Thelohanellus 93 Sarkar N.K. & Mazumdar S Studies on myxosporidian parasites (Myxozoa: Myxosomatidae) from marine fishes in West Bengal, India. I. Description of three new species from Tachysurus sp. Arch. Protistenk. 127 (1): Sarkar N.K. & Raychaudhuri S Thelohanellus bengalensis sp. n. and Myxidium mystuium sp. n. (Myxozoa): Two new Myxosporidia from Indian freshwater teleost. Acta Protozool. 25 (3): Singh R. & Kaur H Two new and two already known species of genus Thelohanellus Kudo, 1933 (Myxozoa: Myxosporea: Bivalvulida) infecting Indian major carp fishes in Punjab wetlands. J. Parasit. Dis. 38 (1): DOI /s Southwell T. & Prashad B Notes from the Bengal Fisheries Laboratory. No. 5 Parasites of Indian fishes with a note on the carcinoma in the climbing perch. II. On some Indian Myxosporidia Plate XI, p In: Records of the Indian Museum 15: Szekely C., Shaharom-Harrison F., Cech G., Mohamed K. & Molnar K Myxozoan pathogens of Malaysian fishes cultured in ponds and net-cages. Dis. Aquat. Organ. 83 (1): DOI: /dao01990 Tripathi Y.R Studies on the parasites of Indian fishes. ň. Protozoa. Myxosporidia together with a Checklist of parasitic protozoa described from Indian fishes. Records of the Indian Museum 50: Zhang J.Y.,Gu Z.M., Kalavati C., Costa Eiras J., Liu Y., Guo Q.Y. & Molnar K Synopsis of the species of Thelohanellus Kudo, 1933 (Myxozoa: Myxosporea: Bivalvulida). Syst. Parasit. 86 (3): DOI: /s Received May 29, 2014 Accepted November 30, 2014