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Protistology 7 (4), 209-217 (2012)
Protistology
Myxosporean species of the genus Thelohanellus Kudo, 1933 (Myxozoa: Myxosporea: Bivalvulida) from freshwater fishes of Punjab wetlands, India
Ranjeet Singh and Harpreet Kaur
Department of Zoology and Environmental Sciences, Punjabi University, Patiala, India
Summary
A study on the myxosporean infections in freshwater fishes in Harike and Kanjali wetlands of Punjab (India) has revealed the presence of three myxozoan species belonging to the genus Thelohanellus. Plasmodia were teased out with the help of a fine needle to liberate the spores on a clean slide. Spores were studied fresh as well as stained after fixing in Bouins fixative. T. batae Lalitha Kumari, 1969 was located within the wall lining of the duodenum and on the pectoral fin of the cat fish, Wallago attu. Earlier, this parasite was recorded from gill filaments of Labeo bata in Andhra Pradesh (India). Spores of T. mrigalae Tripathi, 1952 were found parasitizing the caudal fin of Catla catla. Spores of the new species, T. thaili sp. nov. were found infecting gill lamellae of Catla catla. Spores were pyriform in valvular view, tapering towards the anterior end with bluntly pointed tip. Spores measured 11.67*7.22 ^m in size. Polar capsule was flask-shaped, long-necked, measuring 7.3*4.4 ^m in size. It was placed eccentrically in the spore body cavity (aligned to one side of the inner wall of the spore) occupying nearly three-quarters of the spore body cavity. In the present study, new host, new site of infection and new locality for T. batae and T. mrigalae have been reported.
Key words: caudal fin, gill lamellae, Harike, Kanjali, plasmodium, thelohanelloid
Introduction
Punjab (India) has 3 main wetlands, i.e. Harike, Kanjali and Ropar wetlands which are included in Ramsar list of International importance. Harike wetland is the largest freshwater wetland (in northern India) of 4100 ha. area and is a suitable habitat for as many as 26 species of fishes. Kanjali wetland is a man made wetland covering an area of 185 ha., which nurtures up to 17 fish species. These wetlands
form major natural fisheries resource in whole of the Punjab state. A large variety of these fishes in these wetlands are vulnerable to myxozoan infections, beside other parasitic infections.
Myxozoans are economically important group of microscopic metazoan parasites of marine and freshwater fishes from natural and aquaculture resource causing production losses and render them unfit for human consumption. Up to now, phylum Myxozoa include more than 2,180 species attributed
© 2012 The Author(s)
Protistology © 2012 Protozoological Society Affiliated with RAS
to 65 genera (Singh and Kaur, 2012a). Thelohanellus Kudo, 1933 is the second most prevalent genus after Myxobolus B^schli, 1882 having 60 valid species (Lom and Dykova, 2006). Lom and Dykova (1992) in a monograph enlisted 39 species of the genus Thelohanellus. Kaur and Singh (2008, 2008-2009, 2009, 2010a, 2010b, 2010/2011, 2011a, 2011b, 2011c, 2011d, 2011e, 2011f, 2012a, 2012b) have recorded many myxosporeans infecting freshwater fishes of Punjab wetlands and described as many as 18 new species of the genera Myxobolus and Triangula. Kaur and Singh (2012a) gave a synopsis of Indian myxobolids and revised key to the phylum Myxozoa. Basu and Haldar (1999) and Basu and coauthors (2006) gave a checklist and a synopsis of the genus Thelohanellus enlisting 32 Indian species, respectively. Kalavati and Nandi (2007) compiled a handbook on myxosporean parasites ofIndian fishes. Valuable work has been contributed by Gupta and Khera (1987, 1988a, 1988b, 1988c, 1988d, 1989a, 1989b, 1990, 1991) on these parasites in north India. More recently, Singh and Kaur (2012a, 2012b) have recorded four new species, i.e. Thelohanellus kalavate, T. globulosa from caudal fin of Cirrhina reba and T. kalbensi, T. kanjalensis from gills of Labeo calbasu and skin of snout of Catla catla, respectively. Singh and Kaur (2012c) also studied the biodiversity of the myxozoan parasites in these wetlands. They recorded 36% infection in carp fishes with various genera: Myxobolus, Thelohanellus, Triangula and Neothelohanellus.
Spores ofthe genus Thelohanellus are histozoic, characterized in having smooth shell. Spores are tear to pyriform in shape, broadly ellipsoidal in valvular view and slim in sutural view. Spores contain a pyriform polar capsule with a 1—2 coils of polar filament. Sporoplasm contain two sporoplasmic nuclei.
During the present study, two already known species, i.e. T. batae Lalitha Kumari, 1969, T. mrigalae Tripathi, 1952 and one new species, T. thalli sp. nov. has been reported of which complete description is given. Spores up to generic level were identified with the help of the key given by Kaur and Singh (2012a). The descriptions have been prepared in accordance to the guidelines by Lom and Arthur (1989).
Material and methods
Fishes collected from Harike and Kanjali wetlands were brought to the laboratory and examined for myxozoan infections. Plasmodia were removed,
placed on clean slides and examined with 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 solution for the extrusion of polar filaments. For permanent preparation, air-dried smears were stained with Ziehl-Neelsen and Iron-haematoxylin. Drawings of stained material were made with the aid of camera lucida. Spores were measured with calibrated ocular micrometer. All measurements are presented in ^m as range values followed by mean ± SD (standard deviation) in parentheses. The abbreviations used in the paper are as follows: LS: Length of spore, WS: Width of spore, LPC: Length ofpolar capsule, WPC: Width ofpolar capsule, NC: Number of coils of polar filaments, SD: Standard deviation.
Results and discussion
Thelohanellus batae Lalitha Kumari, 1969
Plasmodia. Small, located within the wall lining of the duodenum and on the pectoral, 2-3 in number and measure 0.2-0.3 mm and 0.7-0.9 mm in diameter, respectively. Each plasmodium contains 12-13 spores.
Spores were histozoic, measure 8.9*3.2 ^m (measurements based on 8-10 spores in frontal view), pyriform to egg shaped in valvular view having pointed anterior end and rounded posterior end (Figs 1a, 1b, 3a, 3b) (Table 1). Shell valves were thin, smooth, symmetrical and measuring 0.2 ^m in thickness. Parietal folds absent. Polar capsule was elongately pyriform, measuring 4.5* 1.7 ^m, occupying nearly half of the spore body cavity. Polar filament form 4-6 coils arranged perpendicular to the polar capsule axis which extrude at the anterior tip of the spore (Figs 2a, 2b). Polar filament thin, thread-like measuring 30 ^m in length. Sporoplasm occupies the whole extracapsular space behind the polar capsule and contain a sporoplasmic nucleus measuring 1.5 ^m in diameter. An iodinophillous vacuole was absent.
Remarks. The present observations (LS/WS: 2.7) on T. batae Lalitha Kumari, 1969 are in conformity with the original description (LS/WS: 1.9) except some variations in the size of the spore and the polar capsule. Spore and polar capsule are smaller in size in the present species. Earlier, this parasite was recorded from the gill filaments of Labeo bata. In a new host Wallago attu, two new sites of infection (wall of duodenum and pectoral fin), and a new locality (Harike wetland) are recorded for this parasite (Table 2).
b
Fig. 1. Line drawing (Camera Lucida) of T. batae Lalitha Kumari, 1969 spores. a — Spore stained with Ziehl-Neelsen (valvular view); b — spore stained with Iron-haematoxylin; the polar filament is extruded. Scale bar: 0.005 mm.
Thelohanellus mrigalae T ripathi, 1952
Plasmodia. Small, white to pale yellow, rounded, located on the caudal fin, 2—5 in number and measuring 0.7—0.9 mm in diameter. Each plasmodium contains 8—10 spores.
Spores were histozoic, measuring 11.6* 8.3 ^m (measurements based on 12-13 spores in frontal view), elongately oval in valvular view having rounded blunt with a knob-like projection at the tip and broad rounded posterior end (Figs 5a, 5b, 6) (Table 3). Shell valves were thick, smooth, symmetrical and measuring 0.62 ^m in thickness. They were stained dark blue with Iron-haematoxylin throughout the length of spore (Figs 4a, 4b, 5a, 5b). Parietal folds absent. Polar capsule was broadly
Table 1. Measurements (|-im) of Thelohanellus batae Lalitha Kumari, 1969.
Character Range Mean value SD
LS 8.5-9.3 8.9 0.56
WS 3.0-3.4 3.2 0.28
LPC 4.2-4.8 4.5 0.42
WPC .8 i—i <JD i—i 1.7 0.14
LS/WS 2.7
NC 4-6
Parietal folds absent
Fig. 2. Micrographs of T. batae Lalitha Kumari, 1969 spores. a — Spore stained with Ziehl-Neelsen; b — spores stained with Iron-haematoxylin; the polar filament is extruded. Scale bars: 10 ^m.
pyriform to ovoid in shape, measuring 5.0*3.0 ^m, having a distinct neck and broad rounded posterior end. It was placed anteriorly occupying nearly half of the spore body cavity. Polar filament form 6—7 coils arranged obliquely to the polar capsule axis. It
(I
a
b
Fig. 3. Micrographs of T. batae Lalitha Kumari, 1969 spores. a , b — fresh spores. *1000.
Fig. 4. Line drawing (Camera Lucida) of T. mrigalae Tripathi, 1952 spores. a — Spore stained with Ziehl-Neelsen (valvular view); b — spore stained with Iron-haematoxylin; the polar filament is extruded. Scale bar: 0.005 mm.
was thick, thread-like and measuring 28 ^m in length after eversion (Fig. 5a, b and Fig. 6). Capsulogenic nucleus was absent. Sporoplasm occupies whole of the extracapsular space behind the polar capsule and contain a nucleus measuring 0.16 ^m in diameter. An iodinophillous vacuole was present measuring 2.9—3.1 (3.0+0.14) ^m in diameter.
Remarks. The present observations (LS/WS: 1.3) on T. mrigalaeTripathi, 1952 are in conformity with the original description (LS/WS: 1.7) except some variations in the size of the spore. Earlier, this parasite was recorded from skin on the head of Cirrhina mrigala. A new host — Catla catla, a new site of infection — caudal fin and a new locality — Kanjali Wetland are recorded for this parasite (Table 4).
• #
i----1
a
a
i----1
Fig. 5. Micrographs of T. mrigalae Tripathi, 1952 spores. a — Spore stained with Ziehl-Neelsen; b — spores stained with Iron-haematoxylin; the polar filament is extruded. Scale bars: 10 ^m.
Fig. 6. Micrographs of T. mrigalae Tripathi, 1952 fresh spore. *1000.
&
b
Table 2. Comparison of Thelohanellus batae in original description (Lalitha Kumari, 1969) and in present study (measurements are in jm).
Species Host Site of infection Locality Spore Polar capsule
T. batae (present study) Wallago attu wall of duodenum, pectoral fin Harike wetland, Punjab (India) 8.9x3.2 4.5x1.7
T. batae Lalitha Kumari, 1969 Labeo bata gill filaments Andhra Pradesh (India) 12.3x6.2 7.7x3.0
Table 3. Measurements (jm) of Thelohanellus Table 5. Measurements (jm) of Thelohanellus
mrigalae Tripathi, 1952. thaili sp. nov.
Character Range Mean value SD Character Range Mean value SD
LS 11.2-12.0 11.6 0.56 LS 11.27-12.07 11.67 0.56
WS 7.9-8.7 8.3 0.56 WS 7.02-7.42 7.22 0.28
LPC 4.0-6.0 5.0 1.4 LPC 6. 9 7. 7 7.3 0.56
WPC 2.9-3.0 3.0 0.14 WPC 4.1-4.7 4.4 0.42
LS/WS 1.3 LS/WS 1.6
NC 6-7 NC 4-5
Parietal Folds absent Parietal Folds absent
Thelohanellus thaili sp. nov.
Plasmodia. Minute and present around gill lamellae. Each plasmodium contains 10—11 spores.
Spores were histozoic, measuring 11.67* 7.22 ^m (measurements based on 4—7 spores in frontal view), pyriform in valvular view, tapering towards the anterior end with bluntly pointed tip and broad rounded posterior end (Table 5). Shell valves were thick, smooth, symmetrical and measuring 0.51 ^m in thickness. Parietal folds absent (Figs 7a, 7b). Polar capsule was flask-shaped, long-necked, measuring 7.3*4.4 ^m in size. It was placed eccentrically in the spore body cavity (aligned to one side of the inner wall of the spore) occupying nearly three-quarters of the spore body cavity (Fig. 9). Polar filament contain 4—5 coils arranged obliquely to the polar capsule axis, thread-like, measuring 58.3 ^m and extrude through a distinct pore at the anterior end ofthe spore (Figs 8a, 8b). One capsulogenic nucleus measuring 1.03—1.63 (1.33+0.42) ^m in diameter was present. Sporoplasm occupies the whole of the extracapsular space behind the polar capsule. Two sporoplasmic nuclei are equal in size, measuring 1.35 ^m in diameter. An iodinophillous vacuole was absent.
Differential diagnosis. The studied species was compared to 17 representatives of the genus Thelohanellus infecting fish (Table 6). It differs from all of them by morphometric characters.
Morphologically, spores of the present species resemble T. batae, T. chelae, T. chrysopomati, T. coeli, T. potaili, T. wallagoi, T. chandannagarensis, T. anilae, T. imphalensis and T. niloticus. It can be differentiated from all of the above mentioned species in having a flask-shaped polar capsule
Fig. 7. Line drawing (Camera Lucida) of T. thaili sp. nov. a — Spore stained with Ziehl-Neelsen (valvular view); b — spore stained with Iron-haematoxylin; the polar filament is extruded. Scale bar: 0.005 mm.
with a long neck opening to the exterior through a distinct pore. Furthermore, the polar capsule is eccentric and occupying nearly three-quarters of the spore body cavity in contrast to pyriform, terminal and oblique polar capsule in T. batae. The
Table 4. Comparison of Thelohanellus mrigalae in original description (Tripathi, 1952) and in present study (measurements are in jm).
Species Host Site of infection Locality Spore Polar capsule
T. mrigalae (present study) T. mrigalae Tripathi,1952 Catla catla Cirrhina mrigala caudal fin skin on the head Kanjali wetland, Punjab (India) West Bengal (India) 11.6x8.3 10.8-12.0x6.3 7.2 5.0x3.0 5.4-7.2x3.6-5.0
a
b
Fig. 8. Micrographs of T. thaili sp. nov. spores.
a — Spore stained with Ziehl-Neelsen; b — spores
stained with Iron-haematoxylin. Scale bars: 10
^m.
polar capsule is anteriorly placed and oval in T. chandannagarensis, terminal, oval or elongated in T. chelae. It is bifurcated in T. imphalensis and presence of pointed and anteriorly placed polar capsule in T. niloticus demarcates them from the spores of the present species in this study.
Spores of the species under consideration are pyriform, bluntly pointed at the anterior end with a distinct pore unlike anteriorly pointed spore of T. wallagoi, elongated pyriform spore in T. anilae and tear shaped spore of T. niloticus.
Furthermore, spores in the present species lack parietal folds and have shell valves equal and smooth unlike spores of T. chrysopomati, T. potaili, T. chelae and T. coeli.
In view of the above differences, it is concluded that the species under present study is really a new species. The name Thelohanellus thaili sp. nov. is proposed after the vernacular name, i.e. thaili, of the host fish, Catla catla.
Taxonomic summary.
Plasmodia. Minute and present around gill lamellae.
Spores. Spores are pyriform in valvular view. A single polar capsule is flask-shaped.
Type host. Catla catla (Ham.) vern. thail.
Type locality. Kanjali wetland, Punjab, India.
Fig. 9. Micrographs of T. thaili sp. nov. fresh spore. *1000.
Type specimen. Paratypes are spores stained in Ziehl-Neelsen and Iron-haematoxylin, deposited in the museum of department of Zoology, Punjabi University, Patiala, India. Slide no. CC/Zn/22.05. 2008 and CC/IH/22.05.2008.
Site of infection. Gill lamellae.
Prevalence of infection. 50% (10/20).
Clinical symptomatology. None.
Etymology. The specific epithet thaili has been given after the vernacular name of the host fish.
Acknowledgements
The authors express thanks to the University Grants Commission (UGC) for the financial support.
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Address for correspondence: Ranjeet Singh, Harpreet Kaur. Department of Zoology and Environmental Sciences, Punjabi University, Patiala-147002, India; e-mail: ranjitsrana@gmail.com; harpreet_bimbra@yahoo.com
