Cytogenetic Studies of Fishing Cat, Prionailurus Viverrinus

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Cytogenetic Studies of Fishing Cat, Prionailurus viverrinus (Bennett 1833) and Asiatic Golden Cat, Catopuma temminckii (Vigors and Horsfield 1827) by Conventional Staining, G-banding and High-resolution Techniques

Alongkoad Tanomtong1,*, Puntivar Kaewmad1, Sumpars Khunsook1 and Sarawut Kaewsri2

1 Department of Biology, Faculty of Science, Khon Kaen University, Muang, Khon Kaen 40002, Thailand 2 Program in Applied Biology, Department of Science, Faculty of Science, Buriram Rajabhat University, Muang, Buriram 31000, Thailand

Received October 10, 2008; accepted February 1, 2008

Summary The fishing cat, Prionailurus viverrinus (Bennett 1833) and the Asiatic golden cat, Catopuma temminckii (Vigos and Horsfield 1827) are two of many wild cat species currently listed as vulnerable or endangered in Thailand. 20 conventional staining, G-banding and high-resolution metaphase spreads obtained from lymphocyte cultures were analyzed for each animal. The female fishing cat has diploid chromosome number of 2n38 and the fundamental number (NF) was 74. There are 6 autosome types: A type had 6 large submetacentric, B type had 8 large acrocentric, C type had 4 large metacentric, D type had 4 medium and 4 small submetacentric, E type had 8 small metacentric and F type had 2 small telocentric chromosomes. The female Asiatic golden cat had 2n38, and the NF was 72. There are 6 autosome types: A type had 4 large and 2 medium submetacentric, B type had 6 large and 2 medium acrocentric, C type had 4 large metacentric, D type had 8 small submetacentric, E type had 6 small metacentric and F type had 4 small telocentric chromosomes. The female fishing cat and the Asiatic golden cat have a pair of the short arm of chromosome E1 (chromosome pairs 14) showed clearly observable satellite chromosomes. The X chromosome was submetacentric chromosome. From G-banding and high-resolution technique, number of bands were 184, 211 and 204, 245, respectively. We found that chromosomes A1, A2, A3, B1, B2, B4, X (fishing cat) and A1, A2, A3, C2, D1, D4, E1, E2, E3, F1, F2, X (Asiatic golden cat) patterns are same as domestic cat (Felis catus) chromosomes. Chromosomes B3, C1, C2, D1, D2, D3, D4, E1, E2, E3, F1 (fishing cat) and C1, D2, D4 (Asiatic golden cat) are similar to those of domestic cat chromosomes. These results show the evolutionary relationship between the wild cat (fishing cat, Asiatic golden cat) and the domestic cat. The karyotype formula of the female fishing cat and the Asiatic golden cat are as follows: m sm a sm m sm t Fishing cat: 2n(38) L4 L6 L8 M4 S8 S4 S2 sex chromosomes. m sm a sm a m sm t Asiatic golden cat: 2n(38) L4 L4 L6 M2 M2 S6 S8 S4 sex chromosomes.

Key words Fishing cat (Prionailurus viverrinus), Asiatic golden cat (Catopuma temminckii), Chromosome, Karyotype.

The fishing cat and the Asiatic golden cat are member of the order Carnivora (meat-eating mammals) in family Felidae. The family Felidae has only 9 species of 6 genera in Thailand, namely the jungle cat, Felis chaus (Gueldenstaedt 1776), the leopard cat Prionailurus bengalensis (Kerr 1792), The fishing cat, [Prionailurus viverrinus (Bennett 1833), the flat-headed cat, Prionailurus

* Corresponding author, e-mail: [email protected] 4 A. Tanomtong et al. Cytologia 74(1) planiceps (Vigors and Horsfield 1827), the Asian golden cat, Catopuma temminckii (Vigors and Horsfield 1827), the marbled cat Pardofelis marmorata (Martin 1837), the clouded leopard Pardofe- lis nebulosa (Griffith 1821), the leopard or the panther, Panthera pardus (Linnaeus 1758) and the tiger, Panthera tigris (Linnaeus 1758) (Lekagul and McNeely 1977, 1988, Par et al. 2003). Common characteristic of the fishing cat are most closely related to the leopard cat among Thai species. Larger and more stocky, gray or olive brown in basic pelage color, with smaller spots on the flanks, shorter legs, shorter ears, more elongated head, a plain breast, and a tail less than 50% of the head and body length. The fur is short and coarse, with the spots of the back tending of form more or less continuous stripes. The cheeks and muzzle are grayish-white, with two stripes angling from the eye to below the ear. The tail is gray with black spots at the base and black rings toward the tip, which is also black. The common characteristics of the Asiatic golden cat are the pelage color varies from golden brown to grayish brown, with the under-parts lighter; a black, phase is known, but is extremely rare. There are no definite spots and stripes on the body, but in certain pos- tures the black bases of the hairs may show through and resemble faint stripes, at least in some indi- viduals. There is on the light spot on the back of the ear (Lekagul and McNeely 1977, 1988, Parr et al. 2003). There are several reports on cytogenetic studies of the family Felidae included (Makino and Tateishi 1952, Thuline and Norby 1961, Ohno et al. 1962, Hsu 1962, Hsu et al. 1963, Matano 1963, Chu et al. 1964, Hus and Rearden 1965, Benirschke and Low 1966, Leyhausen and Tonkin 1968, Sutton 1968, Wurster and Benirschke 1967, 1968a, b, Hard 1968, Wurster 1969, Hsu and Benirschke 1971, Milosevic et al. 1972, Wurster-Hill 1973, Wurster-Hill and Gray 1973, Wurster- Hill and Meritt 1974, Yang et al. 2000, Nie et al. 2002, Keawmad et al. 2007, Tanomtong et al. 2008a, b, c). The recent gained results, cytogenetic knowledge of the fishing cat and the Asiatic golden cat (Carnivora, Felidae), can be a useful information for further conservation of family Feli- dae in Thailand.

Materials and methods Blood samples of the ﬁshing cat (2 females) and the Asiatic golden cat (2 females) were col- lected from Songkla Zoo, Thailand and then applied for cytogenetic studies by lymphocyte culture of whole blood samples. The culture cells were treated with a colchicine-hypotonic-ﬁxation-air-dry- ing technique followed by conventional staining, G-banding and high-resolution techniques (Rooney 2001, Campiranon 2003). 20 cells of each individual chromosome checks, length mea- surements, karotyping and idiograming were accomplished by using a light microscope as previous- ly described (Chaiyasut 1989).

Results Cytogenetic study of the female fishing cat using lymphocyte culture demonstrated that the chromosome number was 2n38 and the fundamental number (NF, number of chromosome arms) was 74. The autosomes of the fishing cat composed of 6 types: A type had 6 large submetacentric, B type had 8 large acrocentric, C type had 4 large metacentric, D type had 4 medium and 4 small submetacentric, E type had 8 small metacentric and F type had 2 small telocentric chromosomes. The female Asiatic golden cat revealed that 2n38, and NF was 72. There are 6 autosome types: A type had 4 large and 2 medium submetacentric, B type had 6 large and 2 medium acrocentric, C type had 4 large metacentric, D type had 8 small submetacentric, E type had 6 small metacentric and F type had 4 small telocentric chromosomes. The fishing cat and the Asiatic golden cat have a pair of the short arm of chromosome E1 (chromosome pairs 14) showed clearly observable satellite chromosomes. The X chromosome was medium submetacentric chromosome (Figs. 1, 2, 3). 2009 Cytogenetic Studies of Fishing Cat and Asiatic Golden Cat 5

Fig. 1. Metaphase chromosome plates and karyotypes of the female ﬁshing cat (A) and the female Asiatic golden cat (B) 2n (diploid)38 by conventional staining technique. Arrow heads were indicated satellite chromosomes with nucleolar organizer regions, NORs and sex-chromosomes. Scale bar represents 10 mm for A and B.

The female ﬁshing cat and the Asiatic golden cat from G-banding revealed that the number of G-bands on 1 set of haploid chromosomes, which includes autosomes, and the X chromosome, were 184, 211 bands, respectively (Figs. 5, 7). The number of bands in 1 set of prometaphase haploid chromosomes from the high-resolution method were 204, 245 bands, respectively (Figs. 6, 8). The present study showed lower number of bands because of only clearly observable bands of the chromosomes were counted. Comparison of chromosome banding patterns between the female ﬁshing cat and the domestic cat (Felis catus) exhibited that 7 chromosome pairs appear as the same pattern (pairs A1, A2, A3, B1, B2, B4 and X chromosome) and 11 chromosome pairs share similarities (pairs B3, C1, C2, D1, D2, D3, D4, E1, E2, E3 and F1 chromosome) (Fig. 4). Comparison of chromosome banding pattern between female Asiatic golden cat and domestic cat revealed that 12 chromosome pairs have the 6 A. Tanomtong et al. Cytologia 74(1)

Fig. 2. Metaphase chromosome plates and karyotypes of the female fishing cat (A) and the female Asiatic golden cat (B) 2n (diploid)38 by G-banding technique. Arrow heads were indicated satellite chromosomes with nucleolar organizer regions, NORs and sex-chromosomes. Scale bar represents 10 mm for A and B. same pattern (pairs A1, A2, A3, C2, D1, D4, E1, E2, E3, F1, F2 and X chromosome) and 3 chromosome pairs share similarities (pairs C1, D2 and D4 chromosome) (Fig. 4). Our results indicated that there is an evolutionary relationship between the wild cat (fishing cat, Asiatic golden cat) and the domestic cat. The length in centimeters of 20 cells of female chromosomes in mitotic metaphase cells was measured. The mean length of the short arm chromosome (Ls), the length of long arm chromosome (Ll), total length of arm chromosome (LT), relative length (RL), centromeric index (CI), standard deviation (SD) of RL, CI, size and the type of presented in Table 1 (fishing cat) and Table 2 (Asiatic golden cat). The fishing cats and the Asiatic golden cat revealed that the chromosome marker is chromosome pair 1, which is the largest submetacentric chromosome. The important chromosome 2009 Cytogenetic Studies of Fishing Cat and Asiatic Golden Cat 7

Fig. 3. Prometaphase chromosome plates and karyotypes of the female fishing cat (A) and the female Asi- atic golden cat (B) 2n (diploid)38 by high-resolution technique. Arrow heads were indicated satellite chromosomes with nucleolar organizer regions, NORs and sex-chromosomes. Scale bar represents 10 mm for A and B. marker of the fishing cat and the Asiatic golden cat are asymmetrical karyotype, which is all four types of chromosomes were found (metacentric, submetacentric, acrocentric and telocentric chromosome). The largest and smallest chromosomes show different sizes (approximately 4 folds). Figs. 5, 7 shows idiograms of fishing cat and Asiatic golden cat from G-banding technique, while Figs. 6, 8 shows idiograms from the high-resolution staining technique with landmarks, bands and sub- bands. The karyotype formulas of the female fishing cat and the Asiatic golden cat were as follows: m sm a sm m sm t Fishing cat: 2n(38) L4 L6 L8 M4 S8 S4 S2 sex chromosomes. m sm a sm a m sm t Asiatic golden cat: 2n(38) L4 L4 L6 M2 M2 S6 S8 S4 sex chromosomes. 8 A. Tanomtong et al. Cytologia 74(1)

Fig. 4. A comparison of the animal species in the family Felidae in Thailand, between the domestic cat, DC and the ﬁshing cat, FC (I); the domestic cat, DC and the Asiatic golden cat, AGC (II) in this study, showing the same (A), similar (B) and different (C) by G-banding patterns. 2009 Cytogenetic Studies of Fishing Cat and Asiatic Golden Cat 9

Fig. 5. Idiogram of the ﬁshing cat 2n (diploid)38 by G-banding technique. The arrow indicates nucleolar organizer region, NOR (satellite chromosome).

Discussion Cytogenetic studies of the female ﬁshing cat and the Asiatic golden cat using lymphocyte culture demonstrated that chromosome number was 2n38. The result is in agreement with the previous studies by Wurster and Benirschke (1968), Hsu and Benirschke (1971), Wurster-Hill and Gray (1973). This corresponds to 2n for the member of the family Felidae according to reports in jungle cat (Felis chaus), clouded leopard (Neofelis nebulosa), puma (Felis concolor), Canadian lynx (F. lynx), snow leopard (Uncia uncia), jaguarondi (F. yagouaroundi), cheetah (Acinonyx jabatus jabatus), serval (F. serval), Asian leopard cat (Prionallurus bengalensis), bob cat (Lynx rufus), marbled cat (F. marmorata), European wild cat (F. silvestris), black footed cat (F. nigripes), leopard (Panthera pardus), tiger (P. tigris), domestic cat (F. catus) (Makino and Tateishi 1952, Thuline and Norby 1961, Hsu 1962, Hsu et al. 1963, Hsu and Rearden 1965, Ohno et al. 1962, Matano 1963, Chu et al. 1964, Benirschke and Low 1966, Leyhausen and Tonkin 1968, Sutton 1968, Hard 1968, Wurster and Benirschke 1967, 1968a, b, Wurster 1969, Hsu and Benirschke 1971, Milosevic et al. 1972 Wurster-Hill 1973, Wurster-Hill and Gray 1973, Wurster-Hill and Meritt 1974, Keawmad et 10 A. Tanomtong et al. Cytologia 74(1)

Fig. 6. Idiogram of the ﬁshing cat 2n (diploid)38 by high-resolution technique. The arrow indicates the nucleolar organizer region, NOR (satellite chromosome).

al. 2007, Tanomtong et al. 2008a, b, c). However, it is interestingly that it differs from the chromosome number of Geoffroy’s (F. g eoffroyi), Marguay (F. wiedi) and ocelot (F. pardalis) which is 2n36 (Hsu 1962, Hsu et al. 1963a, Dubost and Royere 1993). The autosomes of the female ﬁshing cat can be separated into 6 types: A type had 6 large submetacentric, B type had 8 large acrocentric, C type had 4 large metacentric, D type had 4 medium and 4 small submetacentric, E type had 8 small metacentric and F type had 2 small telocentric chromosomes. The autosomes of the female Asiatic golden cat can be also separated into 6 types: A type had 4 large and 2 medium submetacentrics, B type had 6 large and 2 medium acrocentrics, C type had 4 large metacentrics, D type had 8 small submetacentrics, E type had 6 small metacentrics and F type had 4 small telocentric chromosomes. Different chromosomal features were reported by Wurster and Benirschke (1968), Hsu and Benirschke (1971), Wurster-Hill and Gray (1973) which all reported that the ﬁshing cat has 34 metacentrics and submetacentrics, 2 acrocentric chromosomes and Wurster-Hill and Gray (1973) that the Asiatic golden cat has 32 metacentrics and submetacentrics, 4 acrocentric chromosomes. 2009 Cytogenetic Studies of Fishing Cat and Asiatic Golden Cat 11

Fig. 7. Idiogram of the Asiatic golden cat 2n (diploid)38 by G-banding technique. The arrow indicates nucleolar organizer region, NOR (satellite chromosome).

The NF of the fishing cat and the Asiatic golden cat were 74 and 72 in female, respectively. This number is same as NF of the fishing cat as reported by Wurster and Benirschke (1968), Hsu and Benirschke (1971). For the sex-chromosomes, the X chromosome was medium submetacentric chromosome. These feature is similar to the report of Wurster and Benirschke (1968), Hsu and Benirschke (1971), Wurster-Hill and Gray (1973). The comparison of the fishing cat and the Asiatic golden cat X chromosome to those of the domestic cat indicated that the X chromosome was the medium submetacentric chromosome (Thuline and Norby 1961, Ohno et al. 1962, Matano 1963, Chu et al. 1964, Hsu and Rearden 1965). The comparison of the fishing cat and the Asiatic golden cat X chromosome to animals in family Felidae in Thailand namely, leopard (P. pardus), clouded leopard (Pardofelis nebulosa), leopard cat (Prionailurus bengalensis) and jungle cat (Felis chaus) found that X chromosome was the submetacentric chromosome (Keawmad et al. 2007, Tanomtong et al. 2008a, b, c). 12 A. Tanomtong et al. Cytologia 74(1)

Fig. 8. Idiogram of the Asiatic golden cat 2n (diploid)38 by high-resolution technique. The arrow indicates nucleolar organizer region, NOR (satellite chromosome).

To recent study, the fishing cat and the Asiatic golden cat have chromosome marker at short arm of chromosomes E1 (chromosome pairs 14) that are satellite chromosomes with nucleolar organizer regions (NORs). The result is in agreement with previous reports about on chromosomes E1 of cats in the family Felidae having a satellite chromosomes (Makino and Tateishi 1952, Thuline and Norby 1961, Ohno et al. 1962, Hsu 1962, Hsu et al. 1963, Matano 1963, Chu et al. 1964, Hus and Rearden 1965, Benirschke and Low 1966, Leyhausen and Tonkin 1968, Sutton 1968, Wurster and Benirschke 1967, 1968a, b, Hard 1968, Wurster 1969, Hsu and Benirschke 1971, Milosevic et al. 1972, Wurster-Hill 1973, Wurster-Hill and Gray 1973, Wurster-Hill and Meritt 1974, Yang et al. 2000, Nie et al. 2002, Keawmad et al. 2007, Tanomtong et al. 2008a, b, c). G-banding and high-resolution staining techniques revealed that the number of G-bands on one set of haploid, which includes autosomes and X chromosome, were 184 (fishing cat), 211 (Asi- atic golden cat) and 204 (fishing cat), 245 (Asiatic golden cat), respectively. In domestic cat, there are 317 bands on one set of haploid chromosomes (Yang et al. 2000). Our study showed a lower number of bands compared with previous study because of only clearly observable bands of the 2009 Cytogenetic Studies of Fishing Cat and Asiatic Golden Cat 13

Table 1. The mean value of the short arm chromosome length (Ls), of the long arm chromosome length (Ll), of the total arm length of the chromosome (LT), of the relative length (RL) and of the centromeric index (CI). The standard deviation (SD) of RL and CI, the size and type of the chromosomes of 20 cells in female the ﬁshing cat (Prionailurus viverrinus), 2n (diploid)38

Chromosome Chromosome Chromosome Ls Ll LT CISD RLSD pairs size type

A1 0.666 1.202 1.868 0.6430.017 0.0850.006 L Submetacentric A2 0.549 0.942 1.491 0.6320.018 0.0680.003 L Submetacentric A3 0.504 0.780 1.284 0.6070.007 0.0590.001 L Submetacentric B1 0.396 1.310 1.706 0.7680.026 0.0780.004 L Acrocentric B2 0.310 0.993 1.303 0.7620.034 0.0590.002 L Acrocentric B3 0.355 0.934 1.289 0.7250.021 0.0590.001 L Acrocentric B4 0.342 0.906 1.248 0.7260.018 0.0570.002 L Acrocentric C1 0.890 0.929 1.819 0.5110.006 0.0830.004 L Metacentric C2 0.686 0.728 1.414 0.5150.003 0.0640.001 L Metacentric D1 0.397 0.713 1.110 0.6420.025 0.0510.002 M Submetacentric D2 0.339 0.606 0.945 0.6410.039 0.0430.003 M Submetacentric D3 0.335 0.588 0.923 0.6370.032 0.0420.003 S Submetacentric D4 0.310 0.558 0.868 0.6430.032 0.0400.002 S Submetacentric E1 0.417 0.454 0.871 0.5210.008 0.0400.002 S Metacentric E2 0.303 0.357 0.660 0.5410.017 0.0300.004 S Metacentric E3 0.301 0.331 0.632 0.5240.016 0.0290.004 S Metacentric E4 0.246 0.281 0.527 0.5330.017 0.0240.004 S Metacentric F1 0.000 0.792 0.792 1.0000.000 0.0360.001 S Telocentric X 0.473 0.714 1.187 0.6020.004 0.0540.002 M Submetacentric

Remarks: Llarge chromosome (LT1.198), Mmedium chromosome (LT0.934–1.198) and Ssmall chromosome (LT0.934).

Table 2. The mean value of the short arm chromosome length (Ls), of the long arm chromosome length (Ll), of the total arm length of the chromosome (LT), of the relative length (RL) and of the centromeric index (CI). The standard deviation (SD) of RL and CI, the size and type of the chromosomes of 20 cells in female the Asiatic golden cat (Catopuma temminckii), 2n (diploid)38

Chromosome Chromosome Chromosome Ls Ll LT CISD RLSD pairs size type

A1 0.665 1.132 1.797 0.6300.018 0.0930.006 L Submetacentric A2 0.488 0.827 1.315 0.6290.022 0.0680.004 L Submetacentric A3 0.392 0.663 1.055 0.6280.030 0.0550.005 M Submetacentric B1 0.348 1.259 1.607 0.7830.040 0.0830.005 L Acrocentric B2 0.277 0.936 1.213 0.7720.024 0.0630.003 L Acrocentric B3 0.279 0.864 1.143 0.7560.029 0.0590.005 L Acrocentric B4 0.274 0.806 1.080 0.7460.038 0.0560.003 M Acrocentric C1 0.806 0.866 1.672 0.5180.020 0.0870.007 L Metacentric C2 0.584 0.651 1.235 0.5270.020 0.0640.004 L Metacentric D1 0.280 0.612 0.892 0.6770.029 0.0500.003 S Submetacentric D2 0.272 0.531 0.803 0.6610.023 0.0420.003 S Submetacentric D3 0.247 0.499 0.746 0.6690.029 0.0390.003 S Submetacentric D4 0.230 0.453 0.683 0.6630.038 0.0350.002 S Submetacentric E1 0.331 0.380 0.711 0.5340.023 0.0370.006 S Metacentric E2 0.251 0.306 0.557 0.5490.018 0.0290.002 S Metacentric E3 0.182 0.227 0.409 0.5550.025 0.0210.003 S Metacentric F1 0.000 0.690 0.690 1.0000.000 0.0360.004 S Telocentric F2 0.000 0.603 0.603 1.0000.000 0.0310.004 S Telocentric X 0.369 0.622 0.991 0.6280.027 0.0510.003 M Submetacentric

Remarks: Llarge chromosome (LT1.103), Mmedium chromosome (LT0.899–1.103) and Ssmall chromosome (LT0.899). 14 A. Tanomtong et al. Cytologia 74(1) chromosomes were counted. We found that chromosomes A1, A2, A3, B1, B2, B4, X (fishing cat) and A1, A2, A3, C2, D1, D4, E1, E2, E3, F1, F2, X (Asiatic golden cat) patterns are same as those appear as domestic cat chromosomes. Chromosomes B3, C1, C2, D1, D2, D3, D4, E1, E2, E3, F1 (fishing cat) and C1, D2, D4 (Asiatic golden cat) are all similar to those found in domestic cat. It indicates that there is evolutionary relationship between the wild cat (fishing cat, Asiatic golden cat) and the domestic cat. For further studies, more novels genetic information accomplished by using molecular biology or molecular genetics is an essential requirement.

Acknowledgements The ﬁnancial support from The Zoological Park Organization Under the Royal Patronage of H.M. The King is gratefully acknowledged. We also thank Dr. Sopon Dumnui, Director of the organization, Dr. Sumat Kamolnaranath, Chief of the Educational Division and Director of Songkla Zoo, for valuable help. Thanks are also intended to the staffs of this zoo for good cooperation.

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