Na t. His t. Res., Vo l. 3 No. 2: 115-122. March 1995

Chromosome Base Numbers for Tectα， riαand Allied Genera in Peninsular Malaysia

A. Aziz Bidin and R usea Go

Botany Department , Faculty of Life Sciences , Universiti Kebangasaan Malaysia 43600 43600 UKM Bangi, Selangor , Malaysia

Abstract Abstract Cytological studies , especially with regard to th 巴 determination of chromosome base numbers numbers for the and allied genera have rev 巴 aled the existence of a series of three base numbers, 39, 40 and 41. or strict multiple of these numbers. Four genera, Ctenitis, Cyclopeltis, Pleocnemia, and Pt eridrys have been determined to have a base number of 4 1. Heterogonium has π= 40 , while Tectaria has been confirmed in the present study to harbour two chromosome base numbers i.e. i.e. 39 and 40.

Key words: Chromosome base number, Ctenitis, Cyclopeltis, Heterogonium , Pleocnemia, PteridηIS ， Tectaria , Peninsular Malaysia.

The development of cytology is more Bearing this in mind, an initiative was underｭ recent recent than that of ftowering cytology, taken to carry out an investigation based on which extends back to the last century. The Tectariαand allied genera found in Peninsular reason reason is mainly because accurate techniques Malaysia involving the determination of chroｭ were not available to counter the problems of mosome numbers in order to establish the high chromosome numbers and material that exact chromosome base numbers for the group. was extremely to difficult handle, such as root Live specimens were collected from various tips. tips. localities (mainly limestone) in Peninsular Maｭ Before Before the application of Mantons technique laysia and planted at the University Fern Garｭ (Manton, 1950) most of the thousands of publiｭ dens (Universiti Kebangsaan Malaysia Fern cations cations on the pteridophytes were based on Gardens). During the course of the work , it was morphological morphological and anatomical studies. Only found that the genus is fairly variable in its very few reliable chromosome counts were behavior in cultivation, some taxa reaching made before that time. The high success rates maturity very rapidly whilst others take years of of Manton's squash techniques has enabled reｭ to do so. They are also particularly susceptible searches searches to explore the evolutionary studies of to damage from snails, slugs and insects. many fern ftoras from geographical areas such Materials Materials and Methods as as the Americas, Britain, West Africa, India, Japan, Malaysia and Australia. Source of materials Cytological Cytological work on Malaysian pteridoｭ The materials used for cytological studies phytes was initiated by Manton, with a total of were derived from two sources: 101 101 distributed among 43 genera anaｭ i. Fixed sporangial materials from lyzed lyzed for their chromosome numbers (Manton, grown in cultivation at Universiti Kebangsaan 1954). 1954). This was a major initial study but unｭ Malaysia Fern Gardens. fortunately fortunately it was not followed by other workｭ ii. Material fixed in situ at various localities ers, apart from a few sporadic counts made by in Peninsular Malaysia. Jermy on the allied fern genus Selaginella Methods (J ermy et a l., 1967), Bidin on a number of Maｭ In all the cases sporangial materials were laysian laysian Adiantum, Psilotum and Osmunda fixed in one part glacial acetic acid to three (Bidin, 1983, 1984, 1988) and Walker (1 979) on parts of absolute alcohol stored in the freezer at Christensenia. Christensenia. -25 0 C.

hu戸 A. A. A. A. Bidin and R. Go

The squashing, staining in aceto-carmine a diploid species commonly inhabiting humid and recording techniques are standard undisturbed forest , especially in shaded alluviｭ (Manton, 1950). al places near rivers or streams. Pleocnemia irregularis irregularis is larger and dimorphous, and comｭ Results Results and Di scussion monly found on the limestones of the west The taxonomic position of the fern genus coast of Peninsular Malaysia. It is also diploid. Tectaria Tectaria and allied genera in Malaysia had The emphasis of the present study has been been investigated by Holttum and published in almost exclusively upon the determination of numerous publications (Holttum, 1950, 1954, chromosome numbers (particularly on level of 1968, 1974, 1984, 1991). Holttum, in his book ploidy) in order to establish the chromosome of of 1966, subdivided the subfamily Tectarioidea base numbers for the Tectariαgroup. The specｭ of of the family Dennstaedtiaceae into two major imens examined for this purpose are listed in groups centred on Tectaria and Pleocnemia. Table 2. However, Holttum (1991), based on various If the results (gametic chromosome numbers) aspects aspects of morphology, anatomy and some cyｭ of all individuals examined are added to the tology tology and cell ultrastructure, dissolved the records already available in the literature Pleocnemia group and incorporated it into the (Manton 1954), the chromosome numbers of single, enlarged Tectaria group. Thus Tectaria some 34 taxa are now known (Table 3). This is sensu Holttum (1 991) comprises 11 genera of distributed among the di 仔erent ploidy levels which six are recorded for Peninsular Malayｭ as follows: sia: sia: Ctenitis, Cyclo.ρ eltis ， Heterogonium, Pleoｭ 2x 4x Total cnemia, PteridηIS ， and Tecta巾 (Table 1). N um ber of taxa: 24 10 34 In In terms of the number of species as well as % of each class: 70.6 29 .4 100 the the number of individual plants for each The diploids comprise 70.6 of the total with species, the Tectα， ria group forms a conspicuｭ the polyploids being at the tetraploid level ous part of the fern ftora of Peninsular Malayｭ only i.e. polyploidy is at a generally low level sia. sia. The majority of the species are found in with none of the taxa investigated above that undisturbed undisturbed habitats of the lowlands forests on leve l. These findings agree with the chromoｭ granite granite as well as limestone. some numbers for the Tectaria group reported The two most distinctive and widely disｭ from other geographical areas (Lovis, 1977). tributed tributed species are the simple-fronded Tectaｭ These figures favour the hypothesis that Tectaｭ ria ria singaporeana and the pinnately-divided ria and allied genera had their origin in the Pleocnemia irregularis. Tectariα singaporeana is Malesian region. The present study has also revealed the exｭ istence istence of two base numbers for the genus and and species numbers in Table Table 1. List of genera Tectaria, i. e. 39 and 40 and verifies the formerly the the Tectarioid for the Malesian region and in reported reported basic chromosome numbers in other Peninsular Peninsular Malaysia (After Holttum , 1991). genera allied to Tectaria (40, 41: Table 4). Reｭ Number of Species presentatives of these species are shown in Genus Malesia Malesia Peninsular Malaysia Figures 1 and 2. It is suggested that the number 40 is the more primitive one from eridrys Pt eridrys 6 3 Pleocnemia Pleocnemia 20 4 which the base numbers 39 and 41 may have Ctenitis Ctenitis 28 2 been derived as a result of aneuploidy, i. e. 士1. Tectaria Tectaria 105 28 Heterogonium Heterogonium 22 4 Acknowledgements Cyclopeltis Cyclopeltis 5 We are most grateful to the sta百 of the Uniｭ Lastreopsis Lastreopsis 4 。 Fern Gardens for Psomiocarpa Psomiocarpa l 。 versiti Kebangsaan Malaysia Aenigmopteris Aenigmopteris 5 。 the upkeep of live specimens and assistance. Chlamydogramme 2 。 We are also indebted to the Keepers of the Tectaridium Tectaridium 。 herbaria at Kew and Singapore for the loan of

-116- Chromosome base numbers for Tec俗ria and allied genera in Peninsular Malaysia

Table 2. Li st of specimens examin 巴 d. (Notes: K=The Herbarium, Kew, England; UKMB=The Herbarium, Universiti Universiti Kebangsaan Malaysia, Bangi; SING=The Herbarium, Singapore). Co l1 etion Herbarｭ Species Species Locality Co l1 ector Dated Number lum

1. 1. Ctenitis subobsαm Assam, Dighoi Jan 1891 K 2. 2. C. vilお Malaysia, Pahang, Rusea Go RA593, RA599 25/5/91 UKMB Taman Negara 3. 3. Cyclopeltis crenata Malaysia, Perak, Felda Lasah Rusea Go RA276, RA278 28/10/90 UKMB Malaysia, Pahang, Taman Negara B. S. Parris & 10362 28/2185 UKMB P. P. J. Edwards 4. 4. Heterogonium alderwereltii Malaysia, Selangor, Bkt. Takun Rusea Go RA700, RA701 , 16/3/92 UKMB Malaysia, Selangor, Batu Caves C. G. Mathew R.N. RA707 4/1107 K 5. 5. H. pinnatum Malaysia, Perak, Larut Dr. King 2236 8/18/81 K 6. 6. H. pinnatum Malaysia, Selangor, Bkl Takun Rusea Go RA162, RA165 26/9/90 UKMB Kelantan, Gua Cha Rusea Go RA540 5/1/91 UKMB Perak, Sg. Siput Rusea Go RA250 24/10/90 UKMB 7. 7. H. sagaenoides Malaysia, Pahang, Taman Rusea Go RA598, RA628 18/5/91 UKMB Negara

8. 8. Pleocn酔nia conjugata Malaysia, Pahang, Pulau Razali Jaman RJ2827 3/8/90 UKMB Pulau Pulau Tioman, G. Kajang Zulkifti Mohamad 9. 9. P. irregularis Malaysia, Perak, Lata Kekabu Rusea Go RA267 27/10/90 UKMB 10. 10. P. olivacea Malaysia, Pahang, Taman B. S. Parris & 10458 7/3/85 UKMB Negara; Negara; Kuala Kenyam P. J. Edwards

l. 1l. Pterid:ηIS australis Malaysia, Pahang, Taman B. S. Parris & 10419 4/3/85 UKMB Negara, Sg. Renuis Croxa l1 12. 12. P. syrmatica Malaysia, Pahang, Pulau Razali Jaman & RJ2826 3/8/90 UKMB Tioman, G. Kajang Zulki 日 i Mohamad 13. 13. Tecta7匂 angulata Malaysia, Selangor, Isma i1 Sahid 231 17/8/74 UKMB Bukit Bukit Takun

14. 14. T. baγberi Malaysia, Perak, KM 18, Razali Jaman & RJ3185 15/7/91 UKMB Jln. Jln. Tapah-Cameron Highlands M. Di near 15. 15. T. brachiata Nalaysia, Kedah, Pulau Razali Jaman & PL34 16/1/86 UKMB Langkawi, Perak, Pulau A. Zainuddin Pangkor, Telok Nipah Razali Jaman, RJ3078 30/12/89 UKMB Lagani Lagani Sahid & Sani Sani Miran 16. 16. T. cherasica Malaysia, Pahang, Gua Charas Rusea Go RA585, RA588 26/7/91 UKMB RA596

17. 17. T. c7酔回 ta Malaysia, Pahang, Batu 23 Rusea Go RA517, RA518 15/3/91 UKMB Jln. Jln. Tapah-Cameron Highlands

8. 18. T. CI臼uiunata Malaysia, Pahang, Gua Charas Rusea Go RA583, RA586 26/7/91 UKMB 19. 19. T. curtisii Malaysia, Kelantan, Jeli, Rusea Go RA634, RA637 3/9/91 UKMB G.Reng

20. 20. T. deω庁側S Malaysia, Perak, Gopeng しB. M. Allen 4486 13/2/60 K 1. 21. T. de，ω庁四S Malaysia, Selangor, Bk t. Takun Rusea Go RA202, RA207, 4110/90 UKMB RA212 Batu Batu Caves Rusea Go RA180, RA183 27/9/90 UKMB Kelantan, Gua Cha Rusea Go RA541 1/5/91 UKMB 22. 22. T. fauriei Thai1 and, Ching Rai, K. Iwatsuki, Tl 0888 23/9/67 K Nam Mae Kok N. Fukuoka, M. M. Hutoh & D. D. Chaiglom 23. 23. T. βssa Malaysia, Selangor, Rusea Go RA391 , RA397, 12/2/91 UKMB Klang Gates R A4 00

24. 24. T. grand:泌'ßntata Malaysia, Selangor, Kerling, Rusea Go RA533, RA536 17/10/91 UKMB Bk t. Tarek 25. 25. T. griffithii Malaysia, Selangor, Bk t. Takun Rusea Go RA151, RA152, 26/9/90 UKMB Pahang, Cameron Highlands, Rusea Go RA526 16/3/91 UKMB Kuala Boh A. A. A. Bidin and R. Go

Table 2. (continued)

Collection Collection Dated Herbar- Species Species Locality Collector Number lU m

26. 26. T. heゅ etocaulos Thailands, Koa Kanching Dr. Eryl Smith 8647 11/3/24 K 27. 27. T_ impressa Malaysia, Perlis, Wang Klian Rusea Go RA506, RA51 0 28/2/91 UKMB Kedah, Pulau Langkawi A. A. Bidin PL222 18/1/86 UKMB 28. 28. T. keckii Malaysia, Selangor, Bkt Rusea Go RA142, RA153 26/9/90 UKMB Takun Templer Park Rusea Go RA338 UKMB 29. 29. T. manilensis var. Malaysia, Perlis, Bk t. C. G. Mathew R.N. 703 3/2/14 K chup開gensお Chupeng 30. 30. T. melanocaula Sulawesi, M t. Wawonseru E. Hennipman 6088 1/7/79 K Malaysia, Cameron Highlands し B. M. Allen 3958 K 1. 31. T. semibiPinnata Malaysia, Johor, Kota Tinggi, Rusea Go RA650, RA653 8/9/91 UKMB Sg. Sg. Lalang Johor, Sg. Tersap R. E. Holttum 24910 6/7/31 SING Johor, Kota Tinggi, Kpg. Sg. L. B. M. Allen 29/6/59 SING Lalang

32. 32. T. s田町þinnata Malaysia, Kedah, Pulau Singa, Rusea Go R A4 74 25/2/91 UKMB Pulau Pulau Langkawi, Lubuk Rusea Go R A4 85 28/2/91 UKMB Sembilang Pulau Pulau Langkawi, Hutan Rusea Go RA481 26/2/91 UKMB Simpan Machinchang

33. 33. T. siifol抱 Ma1aysia, Selangor, Bk t. Rusea Go RA349 22/11/90 UKMB Anak Takun Perak, Sg. Siput Rusea Go RA263 26/10/90 UKMB 34. 34. T. simonsii Malaysia, Pahang, Batu 23, Rusea Go RA515, RA520, 15/3/91 UKMB Jalan Jalan Tapah-Cameron RA521 , RA525 Highlands Highlands

35. 35. T. 幻 ngaporeana Malaysia, Pahang, Taman B. S. Parris & 10400 6/3/85 UKMB Negara P. J. Edwards 36. 36. T. translucens Malaysia, Selangor, Bk t. Rusea Go RAI55, RA156 26/9/90 UKMB Takun Bk t. Takun Rusea Go RA704, RA706 16/3/92 UKMB Pahang, Taman Negara, Bkt B. S. Parris & 10450 7/3/85 UKMB Batu Luas P. J. Edwards 37. 37. T. tricuspis Malaysia, Pahang D r. king 1975 1/18/80 K 38. 38. T. vasta Malaysia, Pahang, Batu 23, Rusea Go RA515, RA516 15/3/91 UKMB Jalan Jalan Tapah-Cameron RA519 Highlands Highlands 39. 39. T. zeilanica Malaysia, Pulau Tioman, Rusea Go RAllO, RA 1l 5 26/8/90 UKMB Tekek, Tanjung Pak Seman 40. 40. Tectaria sp. 1 Malaysia, Perlis, Wang Klian Rusea Go RA506, RA507, 28/2/91 UKMB RA509, RA512 4 1. Tectaria sp. JI Malaysia, Kelantan, Gua Cha Rusea Go RA540, RA543, 1/5/91 UKMB RA546

-118- Chromosome base numbers for Tectaria and allied genera in Peninsular Malaysia

Table 3. Li st of species studied and chromosome numbers determined. The list includes chromosome numbers determined by Manton (1954) and Manton & sledge (1954). Chromosome Numbers Genus/Speci 巴s η Ploidy References

4t-- 公一勧勾一似おおお恥“おむ Ctenitis Ctenitis vilis Present Investigation C. C. subobscura 一 Fresh specimen not available 44， Cyclopeltis Cyclopeltis crenata AAU Present investigation Heterogonium Heterogonium alderwereltii Present Investigation H. H. giganteum 一 一 Fresh specimen not available 844444444747444nunu'i'i'i'I'i'inunonunonununUnLau H. ρinnatum Manton (1954) H. H. sagaenoides Manton (1954) Pleocnemia Pleocnemia conjugata Manton (1954) P. P. hemiteliiformis Manton (1954) P. P. irregularis Present investigation P. P. olivacea Present investigation Pteridrys australis ω Manton (1 954) 批恥叫ん P. P. syrmatica Pr巴 sent Investigation Tectaria Tectaria angulata Present Investigation nu T. T. barberi ub Manton (1954) T. T. barberi 叫ん叫批 Present investigation T. T. brachiata Present investigation cherasica cherasica Present investigation T. T. bu T. T. coadunata Pr巴 sent investigation T. T. curtisii Present investigation T. T. decurrens 一 一円 一勧 Fresh specimen not available 4 T. T. devexa U-AUnonU Monton (1 954) T. T. fauriei 一 一 Fresh specimen not available 474 hhuh T. T. fissa Present investigation T. T. grandidentata Present investigation griffithii T. griffithii Present investigation T. T. herpatocaulos 一 一口 一批お一 Fresh specimen not available 74 T. T. impressa onU Present investigation T. T. keckii Manton (1954, as T. amplifolia) T. T. manilensis va r. chupengensis Fresh specimen not available T. T. melanocaula 一批公 Fresh specimen not availabl 巴 T. T. semibipinnata 39 Present investigation T. T. semipinnata 40-1 Manton (1 954, as T. maingayi) μ T. T. siifolia 78 叫ん Present Investigation T. T. simonsii 78 uuhu Present Investigation T. T. singaporeana 39 Manton (1954) T. T. translucens 78 Present Investigation 一勧おお批 T. T. triωsPis Fresh specimen not available T. T. vasta 78 Present Investigation T. T. zeilanica 40 Manton and Sledge (1954) Tectaria Tectaria sp. 1 40 Present Investigation Tectaria Tectaria sp. II 78 Present Investigation

Table 4. Comparison of chromosome base numbers in Tectaria and its allied genera under investigation. Manton Walker Lovis Present Genus 1955 in Holttum, 1966 1966-73 1977 Investigation 'i'lnU Ctenitis Ctenitis 4ti 41 A-A 41 Cyclopeltis Cyclopeltis 一 一日 41 佳 41 4444 A- Heterogonium Heterogonium U'i'inU 40 Pleocnemia Pleocnemia 41 Pteridrys Pteridrys 41 Tectaria Tectaria 40 40 39 & 40

-119 A . A . Bidin and R. Go

B b

Fig. Fig. 1. Meiotic plates and silhouettes of representative species with base numbers n=39 and n=78 . A, a: Tectaria Tectaria semibipinnata (Wal l. ex Hook.) Cope l.: A, meiotic plate (base number n=39); a , silhouette (RA650) B , b: Tectaria impressa (F馥) Holt t.; B, moiotic plate (base number η=78); b , silhouette (RA506)

-120 一 Chromosome base numbers for Tectaria and allied genera in Peninsular Malaysia

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Fig. Fig. 2. Meiotic plates and silhouettes of representative species with base numbers n=40 and n=4 1. A , a: Tectaria Tectaria cherasica Holt t.; A, meiotic plate (base number n =40); a, silhouett巴 (RA585). B, b: Ctenitis vilis (Kunze) (Kunze) Ching.; B., meiotic (base plate number n=41); b , silhouette (RA593) .

-121 一 A. A. A. Bi din and R. Go specimens and visits to their herbaria for the in ferns. Ann. Bo t. Res. 4: 229-415. study. study. The work was funded by research grant Manton,1. 1950. Problems of cytology and evolution No. IRPA 4・ 07 ・ 03-007. in the Pteridophyta. 316 pp. Cambridge University Press, London. References Manton , 1. 1954. Cytological notes on one hundred Bidin; Bidin; A. 1983. Notes on the fern and fern-allies of species of Malayan ferns. In Holttum, R. E.: A rev ・ Carey Carey Island. Malayan Naturalist 37 (1): 13-19. ised tlora of Malaya II, Appendix: 623 -6 27. Bidin, A. 1984. A chromosome count for Osmunda Manton , 1. and W. A. Sledge. 1954. Observation on from Peninsular Malaysia. Fern Gazette 12(6): the cytology and of the pteridophyte 36 0- 362. tlora of Ceylon. Philos. Trans. Roy. Soc. London, B. Bidin, A. 1988. A further chromosome count for 238: 127-185. Osmunda (Osmundales) from Peninsula Malaysia. Walker, T. G. 1979. A further chromosome count for Gard. Gard. Bull. Sing. 41(2): 93-94. Christensenia. Fern Gazette 12 (1): 50-5 1. Holttum, R. E. 1950. Further notes on the fern-genus Heterogonium Heterogonium Pres l. Reinwardtia 1(1): 27-3 1. (Accepted on 8 November 1994) Holttum, R. E. 1954. A revised tlora of Malaya II. Ferns. Ferns. 643 pp. Government Printing Ofice, Singaｭ pore. pore. Holttum, R. E. 1968. A revised tlora of Malaya II. マレ一半島産のナナパケシダ属(シダ植物)と Ferns. Ferns. 2nd ed. 653 pp. Government Printing Office, その近縁属の染色体の基本数 Singapore. Singapore. A. A. Aziz Bidin and Rusea Go Holttum, R. E. 1974. The fern genus Pleocnemia. Kew Bull. 29(2): 341-357. Botany Department , Faculty of Li fe Sciences, Univesiti Univesiti Kebangsaan Malaysia. R. R. E. 1984. Studies in the fern-genera allied Holttum, 43600 UKM Bangi. Selangor. Malaysia to to Tectaria III. Blumea 30: 1-1 1. Holttum, R. E. 199 1. Tectaria Group. Flora Malesｭ マレ一半島産のナナパケシダ属とその近縁属の染色体 iana , ser. II , 2(1): 1-132. 数を調べた.その結果，カツモウイノデ属 ， Pleocnemia Jermy, A. C., K. Jones, and C. Colden, 1967. Cytomorｭ 属 ， f切 ridηs 属 ， Cyclopeltis 属のそれぞれの染色体の phological phological variation in Se旬以nella. J. Li nn. Soc. 基本数が x=41 ， Heterogonium 属の基本数が x=40 ， ナ (Bo t. ).60: 147-158. ナパケシダ属の基本数が x=39 ， 40 であることが明らか Lovis , J. D. 1977. Evolutionary patterns and process になった.

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