Karyological Studies of two Japanese Noctule Bats (Chiroptera)

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1 Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics ISSN: (Print) (Online) Journal homepage: Karyological Studies of two Japanese Noctule Bats (Chiroptera) M. Harada, T. A. Uchida, T. H. Yosida & S. Takada To cite this article: M. Harada, T. A. Uchida, T. H. Yosida & S. Takada (1982) Karyological Studies of two Japanese Noctule Bats (Chiroptera), Caryologia, 35:1, 1-9, DOI: / To link to this article: Published online: 30 Jan Submit your article to this journal Article views: 59 Citing articles: 5 View citing articles Full Terms & Conditions of access and use can be found at

2 CARYOLOGIA Vol. 35, n. 1: 1-9, 1982 KARYOLOGICAL STUDIES OF TWO JAPANESE NOCTULE BATS (CHIROPTERA) M. HARADA,* T. A. UCHIDA,** T. H. YOSIDA,*** S. TAKADA **** *Laboratory of Experimental Animals, Osaka City University Medical School, Osaka, Tapan: **Zoological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan; ***Department of Cytogenetics, National Institute of Genetics, Yata 1111, Misima, Japan; ****Department of Medical Zoology, Osaka City University Medical School, Osaka, Japan. SUMMARY - Karyological investigations of two Japanese Nyctalus species were made based on G- and Cbanding pattern analysis. The karyotype of N. furvus (2n = 44, FN =50) was similar to the standard karyotype of Myotis species. On the other hand, the karyotype of N. lasiopterus aviator (2n = 42, FN = 50) was characterized by having an additional large metacentric pair. By comparison of the G-banding pattern between the two Nyctalus species, it is clear that the metacentric pair of N. lasiopterus aviator has derived from the Robertsonian fusion of the two pairs (nos. 8 and 10) of N. furvus. There was also a significant difference in amount of constitutive heterochromatin between the two Nyctalus species. In N. furvus the additional heterochromatin was observed on the majority of autosomes, while in N. lasiopterus aviator a small Chand was restricted to the centromeric area of each autosome. Thus, the Chand pattern of N. lasiopterus aviator would be interpreted by assuming the decrease of heterochromatin in the course of its speciation from N. furvus. INTRODUCTION G- and/ or C-banding patterns of the chromosomes of about 48 bat species have been analysed by PATHAK et al. (1973), STOCK (1975), OBARA et al. (1976), ANTHONY and KITCHIN (1976), OBARA and SAITOH (1977), BICKHAM and HAFNER (1978), HARADA and YosmA (1978), PATTON and BAKER (1978), BICKHAM (1979a, 1979b) and ANno et al. (1980). These studies have revealed that the banding patterns of chromosome arms or segments are strongly conservative. Thus, the banding patterns can be used to detect karyotypic alterations between species of the same genus and between related genera. The karyotypes of four Nyctalus species have been studied with conventional staining technique; Nyctalus noctula 2n = 42 (Duuc et al. 1967; VoRONTsov et al. 1969), N. lasiopterus aviator 2n = 42 (TsuCHIYA et al. 1972; HARADA 1973; ANno et al. 1977), N. leisleri 2n = 46 (FEDYK and FEDYK 1970; ZIMA 1978), and N. furvus 2n = 44 (ANno et al. 1977)

3 2 HARADA, UCHIDA, YOSIDA and TAKADA whose C-banding pattern also has been analysed (ANno et al. 1980). The present paper deals with the karyotype analysis of two Japanese noctule bats by G- and C.banding procedures. MATERIALS AND METHODS Nyctalus lasiopterus aviator was collected in Minamitsuru, Y amanashi Prefecture (3 males) on 25 July 1975 and in Miyako, Iwate Pre. (1 female) on 14 September 1976, and Nyctalus furvus in Miyako, Iwate Pre. (3 males, 1 female) on 14 September 1976 and 29 August For the karyotype analysis, tissues of lung and tail were used in accordance with the procedure described in a previous paper (HARADA and YosmA 1978). G-band staining was identical with the tripsin treatment technique of SEABRIGHT (1971 ), and C-hand was stained by a slight modification of the method by SUMMER (1972). RESULTS 1. Conventional karyotype. «Nyctalus furvus» 2n = 44, FN = The karyotype of Nyctalus furvus resembles closely those of Myotis species examined so far, with 2n = 44 (Fig. 1A). The autosomes consist of three large metacentric pairs, one small submetacentric pair and 17 near acrocentric pairs ranging from medium to small. Although some of the near acrocentric elements have a short arm, none of them was considered as biarmed when calculating the fundamental number. The X chromosome is a medium-sized submetacentric, and the Y is a dot-like element. In addition, out of the uniarmed autosomes one pair of chromosomes (no. 11) possesses maker-secondary constriction away from the centromere. «Nyctalus lasiopterus aviator» 2n = 42, FN = The karyotype of this species consists of four large metacentric pairs, one small submetacentric pair, 15 acrocentric pairs ranging from medium to small, a mediumsized submetacentric X and a dot-like Y chromosome (Fig. 1B). The autosomal elements of this species are more by one large metacentric pair and are fewer Fig Conventional karyotypes of two Japanese Nyctalus species. A, N. furvus; B. N. lasiopterus avzator.

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5 4 HARADA, UCHIDA, YOSIDA and TAKADA by two medium-sized acrocentric pairs than those of N. furvus. One pair of maker chromosomes (no. 11) with secondary constriction adjacent to the centromere is recognized also in this species. 2. G-banding pattern. The G-banding patterns of the two Nyctalus species, in conformity with the proposed standard karyotype of M yo tis hosonoi whose genus seems to be most primitive in Vespertilionidae, are easily matched (Fig. 2). The genomes of the two Nyctalus species appear directly homologous in structural arrangement. Thus, it is revealed that N. l. aviator has more one large metacentric pair (no. 8) instead of lacking one medium-sized acrocentric pair (no. 10) as compared with the autosomes of N. furvus. 3. C-banding pattern. The C-banding patterns differ greatly in amount of constitutive heterochromatin between the two Nyctalus species (Fig. 3 ). In N. furvus the short arms of near acrocentric chromosomes (nos. 6, 7, 12, 13, 16, 18, 19, 20 and 21) possess a large amount of heterochromatin, being entirely heterochromaic. In addition, a large block of heterochromatin occurs near the centromere of every autosome. On the other hand, in N. l. aviator a small C-hand is mainly restricted to the centromeric region. The X chromosome of N. furvus also possesses larger amounts of heterochromatin than does that of N. l. aviator. The Y chromosome of both species is stained as intensely as the Y chromosome of many other mammals. DISCUSSION The karyotypes of four Nyctalus species examined so far, as stated above, have 2n = 42-46, and are strikingly similar to that of Myotis which morphologically seems to be the primitive stock of family V espertilionidae (DoBSON 1978; MILLER 1907; TATE 1942) and karyologically has extreme uniformity of 2n = 44 (standard karyotype). A similar karyotype with 2n = 44 is widespread in Pipistrellus (CAPANNA and CIVITELLI 1966, 1967), Nyctalus (ANno et al. 1977) and Murina (HARADA 1973; ANDo et al. 1977) of Vespertilionidae. Nyctalus is placed as a member of pipistrelloid genera from the standpoint of morphological characters and considered to be a direct offshoot of Pipistrellus (TATE 1942). As for the direction of karyotypic evolution in Nyctalus, it seems that the ancestral form of Pipistrellus had the karyotype of 2n = 44 with a FN = 50 at the time of divergence from Myotis-like form, and that Nyctalus subsequently differen-

6 KARYOTYPES OF JAPANESE NOCTULE BATS 5 Fig. 2. ~ Pair matching of G-banded chromosomes between Myotis and Nyctalus. M, M. hosonoi; F, N. furvus; L, N. lasiopterus aviator. tiated in a similar way from a stem of Pipistrellus without a great change of the karyotype (ANno et al. 1977). Taking their opinions into consideration, it is assumed that the karyotype of N. furvus ( 2n = 44) is the most primitive among those of Nyctalus examined so far. The karyotype of N. l. aviator (2n = 42) is strikingly similar to that of N. noctula (2n = 42), especially in having an additional metacentric pair (no. 8 ). In this connection, it has been believed that the large meta-

7 6 HARADA, UCHIDA, YOSIDA and TAKADA centric pair in question originates from Robertsonian fusion of certain two pairs of acrocentric autosomes found in N. furvus (ANno et al. 1977). This assumption was crearly verified by determination of their chromosome number (nos. 8 and 10) in the present comparative analysis of G-banding patterns (Fig. 2). On the other hand, N. leisleri has the karyotype of 2n = 46 lacking one pair of small submetacentric chromosomes but with additional two pairs of small acrocentric autosomes, compared with the standard karyotype with 2n = 44. With respect to the karyotypic origin of this species, considering that the karyotype of N. leisleri with a higher number of acrocentric chromosomes is more primitive than that of N. noctula (2n = 42), FEDYK and FEDYK (1970) suggested that the karyotype of N. noctula might have originated, as a result of two centric fusions, from that of N. leisleri. One pair of small submetacentric lacking in N. leisleri, however, is extensively found not only in N. noctula (Duuc et al. 1967; VoRONTSOV et al. 1969) and N. l. aviator (TsucHIYA et al. 1972; HARADA 1973; ANno et al. 1977), but also in Myotis (e. g., HARADA and YosmA 1978; ANno et al. 1977), Murina (HARADA 1973; ANno et al. 1977}, Pipistrellus (FEDYK and Ru PRECHT 1976; CAPANNA and CIVITELLI 1967}, Vespertilio (VORONTSOV et al. 1969; OBARA and SAITOH 1977; ANno et al. 1977; ZrMA 1978}, Plecotus (HARADA 1973; BAKER and PATTON 1967; FEDYK and FEDYK 1970; ANno et al. 1977; ZIMA 1978} and Barbastella (CAPANNA et al. 1968; UcHIDA and ANno 1972; ZrMA 1978) within Vespertilionidae. It is accordingly emphasized that the karyotype of N. leisleri has been derived, as a result of centric fission of the small submetacentric autosomes (no. 4 ), from such standard karyotype as that found in N. furvus (2n = 44 ). As stated above, it is worthy of note that there is a significant difference in amount of constitutive heterochromatin between the two Nyctalus species (Fig. 3 ). Namely, the autosomes of N. furvus contained a very high amount of C-hand material as compared with those of N. l. aviator. Such a C-band variation found in the two Nyctalus species would be explained as follows: It is well known that the chromosomes of Myotis species have a small but distinct C-band only on each centromeric part excepting the pair no. 5 (HARADA and YosmA 1978; BICKHAM 1979b; ANno et al. 1980). Thus, the high amounts of heterochromatin contained in the autosomal complement of N. furvus appear to have been brought about, during the.course of its karyotypic alteration from a certain pipistrelle species with a Myotis- Fig Comparison of C-hand karyotypes between two Nyctalus species. A, N. furvus; B, N. lasiopterus aviator.

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9 8 HARADA, UCHIDA, YOSIDA and TAKADA like karyotype, by the extensive addition of C-band material, as stated by ANno et al. ( 1980 ). On the other hand, the C-band pattern of N. l. aviator seems to have been accompanied by loss of heterochromatin in the process of its speciation from N. furvus. Although in N. furvus more than half of the near acrocentric pairs have an additional short arm in this study (Fig. 1A), ANno et al. (1977) reported the karyotype having only one pair of such chromosomes. If so, polymorphism concerning the short arm of the near acrocentric pairs will be interpreted by difference in amount of constitutive heterochromatin (Fig. 3A). From the above considerations, it is concluded that Robertsonian translocation (centric fusion or :fission) and/or the addition and deletion of heterochromatin play an important part in the karytypic evolution considered responsible for bat speciation within Vespertilioninae. Acknowledgement. - The authors are very grateful to Mr. K. ENDO for help in collection, and Professor E. W. ]AMESON, Jr. of the University of California for comments on the manuscript. REFERENCES ANno K., TAGAWA T. and UCHIDA T. A., Considerations of karyotypic evolution within Vespertilionidae. Experientia, 33: , The C-banding pattern of 6 Japanese species of vespertilionine bats (Mammalia: Chiroptera). Experientia, 36: ANTHONY M. C. and KITCHIN R. M., Sex chromosome and Giemsa banding pattern of Plecotus townsendii. ]. Mammal., 57: BA~R R. and PATTON J. L., Karyotypes and karyotypic variation of north american vespertilionid bats. ]. Mammal., 48: BICKHAM]. W., 1979a.- Chromosomal variation and evolutionary relationships of Vespertilionid bats. J. Mammal., 60: , 1979b. - Banded karyotypes of 11 species of American Bats (genus Myotis). Cytologia, 44: BICKHAM ]. W. and HAFNER J. C., A chromosomal banding study of three species of Vespertilionid bats from Yugoslavia. Genetica, 48: 1-3. CAPANNA E. and CIVITELLI M. V., I chromosomi del Pipistrello albolimbato. Caryologia, 19: , I cromosomi del Pipistrellus savii. Caryologia, 20: CAPANNA E., CoNTI L. and RENZI G. D., I cromosomi di Barbastella barbastellus (Mammalia-Chiroptera). Caryologia, 21: DoBSON G. E., Catalogue of the Chiroptera in the collection of the British Museum. British Mus., London, XLII pp. Duuc B., SOLDATOVIC B. and RIMSA D., La formule chromosomique de la Noctule, Nyctalus noctula Schreber (Mammalia, Chiroptera). Experientia, 23: 1-4. F'EDYK A. and FEDYK S., Karyotypes of species of vespertilionid bats from Poland. Acta Theriol., 25: FEDYK S. and RuPRECHT A. L., Karyotypes of Pipistrellus pipistrellus (Schreber 1774) and P. nathusii (Keyserling and Blasius 1839) (Chiroptera: Vespertilionidae). Caryologia, 29: HARADA M., Chromosomes of nine Chiropteran species in Japan (Chiroptera). La Kromosomo, 91:

10 KARYOTYPES OF JAPANESE NOCTULE BATS 9 HARADA M. and YosmA T. H., Karyological study of four ;apanese Myotis bats (Chiroptera, Mammalia). Chromosoma, 65: MILLER G. S., The families and genera of bats. U. S. Nat. Mus. Ball., 57: XVII pp. OBARA Y. and SAITOH K., Chromosome studies in the Japanese Vespertilionid Bats: IV. Karyotypes and C-banding pattern of Vespertilio orientalis. Japan. J. Genet., 52: OBARA Y., ToMIYASU T. and SArTOH K., Chromosome studies in the Japanese vespertilionid bats. 1. Karyotypical variation in Myotis macrodactylus Temminck. Japan. J. Genet., 51: PATHAK S., Hsu T. C. and UTAKOJI T., Relationship between patterns of chromosome banding and DNA synthetic sequences: a study on the chromosomes of the Sebas bat, Carollia perspicillata. Cytogenet. Cell Genet., 12: PATTON J. C. and BAKER R. J., Chromosomal homology and evolution of Phyllostomatoid bats. Syst. Zoo!., 27: SEABRIGHT M., A rapid banding technique for chromosomes. Lancet, 11: STOCK A. D., Chromosome banding pattern homology and its phylogenetic implications in the bat genera, Carollia and Choeroniscus. Cytogenet. Cell. Genet., 14: SuMNER A. T., A simple technique for demonstrating centromeric heterochromatin. Exp. Cell Res., 75: TATE G. H. H., Results of the Archbold expeditions. No. 47. Review of the vespertilionine bats, with special attention to genera and species of the Archbold collections. Bull. Amer. Mus. Nat. Hist., 80: TsucHIYA K., HARADA M. and YosiDA T. H., Karyotypes of four species of bats collected in Japan. Ann. Rep. Nat. Inst. Genet. Japan, 22: UcHIDA T. A. and AND6 K., Karyotype analysis in Chiroptera. 1. Karyotype of Eastern barbastelle, Barbastella leucomelas darielingensis and comments on its phylogenetic position. Sci. Bull. Fac. Agr., Kyushu Univ., 26: VoRONTsov N. N., RADJABLI S. I. and VoLOBUEV V. T., The comparative karyology of the vespertilionid bats, Vespertilionidae (Chiroptera). In: «The mammals» (ed. Vorontsov). 2nd. All Union Mamm. Con. Moscow., pp ZrMA J., Chromosome characteristics of Vespertilionidae from Czechoslovakia. Acta Sc. Nat. Brno, 12: Received 22 May 1981; accepted 14 October