Systematic Implications for the Genus Diplazium

285 presumed to be closely related are in the same tereminal clade. The same depiction was also showed in the clade that containing D. malaccense,D. sorzogonense and D. tricholepis, and ‘D. porphyrachis group’. It is indicate that morphological data of Diplazium contain phylogenetic signal although many characters are polymorphic and homoplastic. Backer et al 1998 stated that morphological datasets contain more phylogenetic signal per characters and can swamp much larger molecular datasets. Therefore, study that include more morphological characters from living materials as well as anatomical characters, such as stipe or rachis, would give an inferred phylogenetic tree with strong statically support. As showed at Chapter 4, anatomical characters of stipe give an indication that the closely related species are showing the similar anatomical characters. Knowing the status of species, whether originally hybrid or not, that will be included in the phylogenetic analysis of Diplazium is important as many species of Diplazium are apogamous and they are presumed to be originally hybrids. Because if hybridization has occurred among the species of a taxon under cladistic analysis the results are varied. Hybridization results in incongruent intersecting data that obscure the underlying hiearachy Funk 1985. All hyphotheses regarding hybrids identification of course must be corroborated by chromosomal, distributional, and ecological data. Therefore some species of Diplazium that presumed to be originally hybrids based on result of this, such as D. procumbens, D. subvirescens , and D. asymmetricum, should be confirmed in the future study.

10.3. Systematic Implications for the Genus Diplazium

It is too premature to propose a systematic frame within the genus Diplazium because: 1 The most parsimonious tree generated from morphological data is not supported by the objective statistical arguments; 2 Phylogenetic tree generated from gene rbcL sequences data has not given robust phylogentic tree as most of the key species inferred from morphological tree are not included in analysing; 3 Phylogenetic analysis on data sets combining both molecular and non molecular for inferring the phylogeny of Diplazium has not been conducted 286 yet due to the incomplete data on many taxa the gene rbcL sequences data are only provided on 29 species of the 69 West Malesian species. However, this results of this study is interesting because it shows that some clades generated from morphological data are congruence with those generated from gene rbcL sequence data, such as the separation of D. porphyrorachis from the other species of Diplazium and the consistency of some species that included in the terminal clades of both morphological tree and gene rbcL tree. 292 LITERATURES Abraham A, Ninan CA, Mathew PM. 1962. Studies on the cytology and phylogeny of the pteridophytes: VII. Observation on one hundred species of south Indian ferns. J Indian Bot Soc 41: 339-421. Adams KL, Cronn R, Percifield R, Wendel JF. 2003. Genes duplicated by polyploidy show unequal contributions to the transcription and organ-specific reciprocal silencing. Proc Natl Acad Sci 100: 4649-4654. Alston AHG. 1956. The Subdivision of the Polypodiaceae. Taxon 5: 23-25. Andrews SB. 1990. Ferns of Queensland. Brisbane: Queensland Department of Primary Industries. Backer CA en O Posthumus. 1939. Varenflora voor Java. Uitgave van’s Lands Plantetuin, Buitenzorg. Backer RH, Yu X, DeSalle R. 1998. Assessing the relationship contribution of molecular and morphological characters in simultaneous analysis trees. Mol Phylog Evol 9: 427 – 436. Barrington DS, Paris CA, Ranker TA. 1986. Systematic inferences from spores and stomata size in the ferns. Amer Fern J 76 3: 149-159. Baack EJ. 2004. Cytotype segregation on regional and microbiographic scales in snow buttercups Rannunculus adoneus: Ranunculaceae. Amer J Bot 91: 1783-1788. Bhavanandan KV, Ammal LS. 1991. Cytological investigations on family Aspidiaceae sensu Copeland from south India. Indian Fern J 8: 78-86. Bir SS. 1962. Taxonomy of the Indian members of family “Aspleniaceae”. Bull Bot Surv India 4: 1-16. Bir SS. 1969. The stelar anatomy of Diplazium latifolium Moore. Amer Fern J 59: 23 – 26. Blackmore S. 2000. The palynological compass: the contribution of palynology to systematics. In: Nordenstan B, El-Ghozaly G, Kassas M, editors. Plant Systematics for the 21 st Century. London: Portland Press. Pp 161-177. Bower FO. 1912. Studies in the phylogeny of the Filicales. II. Lophosoria and its relation to the Cyatheoideae and other ferns. Annals Bot 26: 269-325. 293 Bower FO. 1913. Studies in the phylogeny of the Filicales. III. On Metaxya and certain other relative primitive ferns. Annals Bot 27: 443-477. Bower FO. 1923-1928. The Ferns Filicales. 3 Vols. England: Cambridge. Bremer K, Wanntorp HE. 1978. Phylogenetic systematic in botany. Taxon 27: 317-329. Brummit RK 1992. Vascular Plant Families and Genera. Kew: Royal Botanic Garden. Bull JJ, Huelsenbeck JP, Cunningham CW, Swofford DL, Waddel PJ. 1993. Partaining and combining data in phylogenetic analysis. Syst Biol 42: 384- 397. Chase MW, Soltis DE, Olmstead RG, Morgan D, Les DH, Mishler BD, Duvall MR, Price RA, Hill HG, Qiu Y-L, Kron KA, Rettig JH, Conti E, Palmer JD, Manhart JR, Sytsma KJ, Michaels HJ, Kress WJ, Karol KG, Clark WD, Hedrén M, Gaut BS, Jansen RK, Kim K-J, Wimpee CF, Smith JF, Furnier GR, Strauss SH, Xiang Q-Y, Plunkett GM, Soltis PS, Swensen SM, Williams SE, Gadek PA, Quinn CJ, Eguiarte LE, Golenberg E, Learn GH Jr, Graham SW, Barrett SCH, Dayanandan S, Albert VA. 1993. Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. Ann Missouri Bot Gard 80: 528-580. Chin WY. 1997. Ferns of the Tropics. Singapore: Times Editions. Ching R-C. 1940. On natural classification of the family ‘Polypodiaceae”. Sunyatsenia 5: 201-286. Ching R-C. 1954. A systematic arrangement of the Chinese fern families and genera with corresponding names in Chinese. Acta Phytotax Sin 3: 93-98. Ching R-C. 1964a. On some confused genera of the family Athyriaceae. Acta Phytotax Sin 9 1: 41-84. Ching R-C. 1964b. On the genus Diplaziopsis C. Chr. Acta Phytotax Sin 91: 31- 36. Christensen C. 1911. On the Genus Dryopteris. Amer Fern J 1: 33-37. Christensen C, Holttum RE. 1934. The ferns of Mount Kinabalu. Gard Bull Sing 7: 191-279. 294 Colwell RK, Hurtt GC. 1994. Non biological gradients in species richness and a spurious Rapoport effect. American Naturalist 144: 570-595. Copeland EB. 1908. A revision of the Philippine species of Athyrium. Philip J Sci IIIc: 285-230. Copeland EB. 1929. The oriental genera of Polypodiaceae. Univ Cal Publ Bot 16: 45-128. Copeland EB. 1947. Genera Filicum. MASS, USA: Waltham. Crane EH. 1997. A revised circumscription of the genera of the fern family Vittariaceae . Syst Bot 22 3: 509-517. Crane EH, Farrar DR, Wendel JH. 1995. Convergent simplification leads to a polyphyletic Vittaria. Amer Fern J 85: 283-305. Darnaedi D. 1991. Kromosom dalam Taksonomi. Makalah dalam Kursus Singkat Metodologi Penelitian Taksonomi Tumbuhan, PAU Hayat IPB, 12 Desember 1990-12 Februari 1991. Bogor. Darnaedi D. 1992. A preliminary cytologicalstudy of fern flora of Gede- Pangrango National Park West Java. Proc Sem Asian Pterid II: 73-78. Darnaedi D, Kato M, Iwastsuki K. 1989. A cytotaxonomic study of Dryopteris sparsa and closely related species Dryoptridaceae. J Jap Bot 64: 330- 340. Davis PH, Heywood VH. 1963. Principles of Angiosperm Taxonomy. Edinburg and London: Oliver and Boyd. de Vogel EF. 1987. Guidelines for the preparations of revisions. In de Vogel EF, editor. Manual of Herbarium Taxonomy Theory and Practice. Jakarta: UNESCO. Dickason FG. 1946. A phylogenetic study of the ferns of Burma. Ohio J Sci 42 2: 73 – 108. Doyle JJ, Doyle JL. 1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem Bull 19: 11-15. Dubuisson J-Y, Hebant-Mauri R, Galtier J. 1998. Molecules and morphology: Conflicts and congruence within the fern genus Trichomanes Hymenophyllaceae. Mol Phylog Evol 9 3: 390-397. 295 Duvall MR, Clegg MT, Chase MW, Clark WD, Kress WJ, Hills HG, Eguiarte LE, Smith JF, Gaut BS, Zimmer EA, Learn GH Jr. 2003. Phylogenetics hyphothesis for the Monocotyledons constructed from rbcL sequence data. Ann Missouri Bot Gard 80: 607-619. Eastwood W, Cronk QCB, Vogel JC, Hemp A, Gibby M. 2004. Comparison of molecular and morphological data on St. Helena: Elaphoglossum. Plant Syst Evol 245: 93-106. Edie, H. 1978. Ferns of Hongkong. Hong Kong: Hong Kong University Press. Colwell RK, Hurtt GC. 1994. Nonbiologcal gradients in species richness and a spurious Rapoport effect. Amer Naturalist 144: 570-595. Endress PK, Bass P, Gregory M. 2000. Systematic plant morphology and anatomy –50 years of progress. Taxon 49 3: 401-434. Estabrook GF. 1978. Some concept for estimation of evolutionary relationships in Systematic Botany. Syst Bot 3: 146 – 158. Felsenstein J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: 783 -791. Ferrarini E, Ciampolini F, Pichi Sermolli REG, Marchetti D. 1986. Iconographia palynologica Pteridophytorum Italiae. Webbia 401: 1-202. Funk VA. 1985. Phylogenetic pattern and hybridization. 1985. Ann Missouri Bot Gard. 72: 681-715. Gastony GJ. Spore morphology in the Dicksoniaceae. II. The genus Cibotium. Cannad J Bot 60: 955-972. Gastony GJ, Ungerer MC. 1997. Molecular systematics and a revised taxonomy of the onocleoid ferns Dryopteridaceae: Onocleeae. Amer J Bot 84: 840- 849. Gastony GJ, Johnson WP. 2001. Phylogenetic placements of Loxoschape thecifera Aspleniaceae and Actiniopteris radiate Pteridaceae based on analysis of rbcL nucleotide sequences. Amer Fern Jour 91: 197-213. Geiger JMO, Ranker TA. 2005. Molecular phylogenetic and historical biogeography of Hawaiian Dryopteris Dryopteridaceae. Mol Phylog Evol 34: 392-407. 296 Gentry AH, Dodson CH. 1987. Diversity and biogeography of neotropical vascular epiphytes. Annals Missouri Bot Gard 74: 205-233. Givnish TJ, Sytsma KJ. 1997. Consistency, characters, and the likelihood of correct phylogenetic inference. Mol Phylog Evol 7: 320-330. Golenberg EM, Giannasi DE, Clegg MT, Smiley CJ, Durbin M, Henderson D, Zurawski G. 1990. Chloroplast DNA sequence from a Miocene Magnolia species. Nature 344: 656-658. Godfray HC, Lawton JH. 2001. Scale and species numbers. TRENDS in Ecol Evol. 16 7: 400-404. Grant V. 1981. Plant Speciation. 2 nd Edition. New York: Columbia University Press. Harris WF. 1955. A Manual of the Spores of New Zealand Pteridophyta. Department of Scientific and Industrial Research Bulletin 116. Government Printer, Wellington. Hasebe M et al. 1994. rbc L gene sequences provide evidence for the evolutionary lineages of leptosporangiatae ferns. Proc Natl Acad Sci USA 91: 5730-5734. Hasebe M et al. 1995. Fern phylogeny based on rbcL nucleotida sequences. Amer Fern J 85: 134-181. Hassler M, Brian S. 2002. Family Athyriaceae, genus Diplazium; world species list. Published on the internet. http:homepages.caverock.net.nz~bjferndiplazium.htm accessed 6 November 2005. Haufler CH, Ranker TA. 1995. rbcL sequences provide phylogenetic insights among sister species of the ferns genus Polypodium. Amer Fern J 85: 361- 374. Haufler CH, Soltis DE. 1986. Genetic evidence suggests that homosporous fern with high chromosome numbers are diploid. Proc Natl Acad Sci 83: 4389- 4393. 297 Haufler CH, Hooper EA, Therrien JP. 2000. Modes and mechanism of speciation in pteridophytes: Implications of contrasting patterns in ferns representing temperate and tropical habitats. Plant Spec Biol 15: 223-236. Hauk WD, Parks CR, Chase MW. 2003. Phylogenetic studies of Ophioglossaceae : evidence from rbcL and trnL-F plastid DNA sequences and morphology. Mol Phylog Evol 28: 131-151. Hennequin S, Ebihara A, Ito M, Iwatsuki K, Dubuisson J-Y. 2006. New insights into the phylogeny of the genus Hymenophyllum s.l. Hymenophyllaceae: Revealing the polyphyly of Mecodium. Syst Bot 312: 271-284. Herter WG. 1949. Flora de Uruguay. I. Pteridophyta. Rev Sudamer Bot 9: 1- 28. Hillis DM. 1987. Molecular versus morphological approaches to systematics. Ann Rev Ecol Syst 18: 23-42. Holbrook-Walker S, Lloyd RM. 1973. Reproductive biology and gametophyte morphology of the Hawaiian fern genus Sadleria Blechnaceae relative to habitat diversity and propensity for colonization. Bot J Linn Soc 67: 157-174. Holttum, R.E. 1938. The Ecology of Tropical Pteridophytes. In: Verdoorn F, editor. Manual of Pteridology. The Netherlands: Martinus Nijhoff. Pp. 420- 450. Holttum RE. 1940. The fern genus Diplazium in the malay Peninsula. Gard Bull Sing 9 2: 74 – 107. Holttum RE. 1947. A revised classification of Leptosporangiate ferns. J Linn Soc Bot 53: 123-158. Holttum RE. 1954. A Revised Flora of Malaya. II. Ferns of Malaya. Singapore: Gov Print Office. Holttum RE. 1966. A Revised Flora of Malaya. II. Ferns of Malaya. Singapore: Gov Print Office. 658p. + Appendix. Holttum RE, Sen U. 1961. Morphology and classification of the tree ferns. Phytomorphology 11: 406-420. Holttum RE, Roy SK. 1965. Cytological observation on some ferns from New Guinea with descriptions of new species. Blumea 13:129-139. 298 Huang Y-M, Chou H-M, Wang J-C, Chiou W-L. 2007. The distribution and habitats of the Pteris fauriei complex in Taiwan. Tawania 521: 49-58. Hyrabayashi H. 1974. Cytogeographic Studies on Dryopteris of Japan. Tokyo: Harashobo. Imaichi R, Kato M. 1995. Speciation and morphological evolution in rheophytes. In: Iwatsuki K, Raven PH, editors. Evolution and Diversification of Land Plants. Tokyo: Springer-Verlag. Pp. 309- 318. IPNI The International Plant names Index. 2006. Published on the internet http:www.ipni.org accessed 17 July 2006. Iwatsuki K, Kato M. 1984. Additions to the enumeration of East Kalimantan Pteridophytes 3. Acta Phytotax Geobot 35 1-3: 59 -67. Jian-xiu LZ, Feng-Qin, Yu-long Z. 1988. Studies on the spores morphology of Hypodematium in China. Pp. 269-272. In KH. Shing K.U. Kramer Eds.. Proceeding of the International Symposium on Systematic Pteridology. September 5-10. Beijing. John RJ. 1995. Malesia-An Introduction. Curtis’s Bot Mag 12 2: 52-62. Kato M. 1972. The vascular structure and its taxonomic significant in the the Athyriaceae . Acta Phyototax Geobot 25 2-3: 79 – 91. Kato M. 1973. Taxonomical evaluation of the articulate hairs found in the Athyriaceae . Acta Phytotax Geobot 25 4-6:119-125. Kato M. 1977. Classification of Athyrium and allied genera of Japan. Bot Mag Tokyo 90: 23-40. Kato M. 1979. Taxonomic study of the genus Cornopteris Athyriaceae. Acta Phytotax Geobot 334-6: 101-118. Kato M. 1984. A taxonomic study of the Athyrioid fern genus Deparia with main reference to the Pacific species. J Fac Sci Univ Tokyo IV 13: 375- 430. Kato M. 1987. A phylogenetic classification Ophioglossaceae. Gard Bull Sing 40 1: 1 – 14. Kato M. 1992. Taxonomic study of Pteridophytes of ambon and Seram Molluccas collected by Indonesian – Japanese botanical expedition IX. 299 Woodsiaceae , Lindsaeaceae, and Adiantaceae. J Fac Sci Univ Tokyo Sect III 15: 135-152. Kato M. 1994. Taxonomic Studies of Pteridophytes of Ambon and Seram Mollucas Collected by Indonesian-Japanese Botanical Expedition IX. Woodsiaceae , Lindsaeaceae and Adiantaceae. J Fac Sci Univ Tokyo 154: 315-347. Kato M. 1995. Woodsiaceae. In: Iwatsuki K, Yamasaki Y, Boufford DE, Ohba F, editors. Flora of Japan. Volume I. Pteridophyta Gymnospermae. Tokyo: Kodhansha. Pp. 195-231. Kato M, Setoguchi H. 1999. An rbcL-based phylogeny and heteroblastic leaf morphology of Matoniaceae. Syst Bot 234: 391-400. Kato M, Darnaedi D, Iwatsuki K. 1991. Fern rheophytes of Borneo. J Fac Sci Univ Tokyo III 15: 91-110. Kato M, Yatabe Y, Sahashi N, Murakami N. 2001. Taxonomic studies of Cheiropleuria Dipteridaceae. Blumea : Kikkawa J, Williams EE. 1971. Vegetational structure and avian diversity in several New World areas. American Naturalist 105: 425-435. Kitching IJ, Forey PL, Humphries CJ, Williams DM. 1998. Cladistics: The Theory and Practice of Parsimony Analysis . Ed. 2 . Oxford: Oxford University Press. Kilpper K. 1964. Über eine RätLias-Flora aus dem Nördlichen Abfall des Albursgebirges in Nordiran. 1. Bryophyta und Pteridophyta. Palaeontographica, Abt B, Paläophytol 114: 1-78. Kluge AG. 1989. A concern for evidence and a phylogenetic hyphothesis of relationships among Epicrates Boidae, Serpentes. Syst Zool 38: 7 – 25. Kramer KU, Holttum RE, Moran RC, Smith AR. 1990. Dryopteridaceae. In: Kramer KU and Green PS eds. Pteridophytes and Gymnosperms. Berlin: Springer-Verlag. Pp 101-144. Large MF, Braggins JE. 1991. Spore Atlas of New Zealand Ferns and Ferns Allies . Wellington: SIR Publishing. 300 Lee MSY. 2004. Molecular and morphological datasets have similar numbers of relevant phylogenetic characters. Taxon 53 4: 1019-2022. Levin DA. 2002. The Role of Chromosomal Change in Plant Evolution. Oxford: Oxford University Press. 240pp. Lewis WH. 1980. Polyploidy in species population. In: Lewis WH, editor. Polyploidy: Biological Relevance . NY – USA: Plenum Press. Pp. 103-144. Lincoln RJ, Boxshall GA, Clark PF. 1982. A Dictionary of Ecology, Evolution and Systematics . New York: Cambridge University Press. Linder HP. 2001. On areas of endemism, with an example from the Restionaceae . Syst Biol 50 6: 892-912. Lloyd RM. 1974. Mating system and genetic load in pioneer and non-pioneer Hawaiian Pteridophyta. Bot J Linn Soc 69: 23-35. Lomolino MV. 2001. Elevational gradients of species-density: historical and prospective views. Global Ecology Biogeography 10: 3-13. Löve A, Löve D, Pichi Sermoli REG. 1977. Cytotaxonomical Atlas of the Pteridophyta. Vaduz: J Cramer. 398p. Lovis JD. 1977. Evolutionary patterns and processes in ferns. Adv Bot Res 4: 229-415. Lu J-M, Barrington DS, Li D-Z. 2007. Molecular phylogeny of the Polystichoid ferns in Asia based on rbcL sequences. Syst Bot 32 1: 26-33. Maddison WP. 1993. Missing data versus missing characters in phylogenetic analysis. Syst Biol 42: 576-581. Manton I. 1950. Problems of Cytology and Evolution in the Pteridophyta. Cambridge: Cambridge Univ. Press. Manton I. 1953. The cytological evolution of the fern flora of Ceylon. Symp Soc Expl Biol 7: 174-185. Manton I. 1954. Cytological notes on one hundred species of Malayan ferns. In: Holttum RE. A Revised Flora of Malaya. II. Appendix: 623 – 627. Manton I, Sledge WA. 1954. Observations on the cytology and taxonomy of the Pteridophyte flora of Ceylon. Phil Trans B. 238. Maxted N. 1992. Towards defining a taxonomic revision methodology. Taxon 414: 653-660. 301 McCoy ED. 1990. The distribution of insects along elevational gradients. Oikos 58: 313-322. Mickel JT. 1974. Phyletic lines in the modern fern. Annals Missouri Bot Gard 61 2: 474 – 482. Manen, J.-F., V. Savolainen, S. De Marchi, and B. Rion. 1995. Chloroplast DNA sekuense from Mioece diatomite deposit in Ardeche France Comptes Rendu. Academy of Science Paris 318: 971-975. Matsumoto S. 2003. Species ecological study on reproductive systems and speciation of Cyrtomium falcatum complex Dryopteridaceae in Japanese Archipelago. Ann Tsukuba Bot Gard 22: 1-141. Mitsui K. 1986. The development of the perispore in the fern family Thelypteridaceae. In Blackmore S, Ferguson IK eds. Pollen and spores: form and function. Linnean Society Symposium Series No 12 : 401-403. London: Academic Press. Mitsuta S. 1985. List of West Sumatran ferns and fern allies collected by dr. Hotta and others during 1980-83 3. Acta Phytotax Geobot 36 1-3: 73-86. Moore S-J, Hsieh T-H, Huang YM, Chiou W-L. 2002. Diplazium maoenense Ching, a poorly known species of the Athyriaceae Pteridophyta in Taiwan. Taiwan J For Sci 171: 113-118. Murakami N, Yokoyama J, Cheng X, Iwasaki H, Imaichi R, Iwatsuki K. 1998. Molecular -taxonomy of Hymenasplenium obliquissimum complex Aspleniaceae based on rbcL sequences. Plant Spec Biol 13: 51-56. Murakami N, Nagomi S, Watanabe M, Iwatsuki K. 1999a. Phylogeny Aspleniaceae inferred from rbcL nucleotide sequences. Amer Fern J 89: 232- 243. Murakami N, Watanabe M, Yokoyama J, Yatabe Y, Iwasaki H, Serizawa S. 1999b. Molecular taxonomic study and revision of the three Japanese species of Asplenium sect. Thamnopteris. J Plant Res 112: 15-25. Myamoto MM, Fitch WM. 1995. Testing species phylogenies and phylogenetic methods with congruence. Syst Biol 44: 64 – 76. 302 Nagalingum NS, Schneider H, Pryer KM. 2007. Molecular phylogenetic relationship and morphological evolution in the heterosporous fern genus Marsilea. Sys Bot 32 1: 16 – 25. Nakato I. 1981. Notes chromosomes of Japanese pteridophytes 1. J Jap Bot 56: 200-205. Nayar B. 1970. A phylogenetic classification of the homosporous ferns. Taxon. 19: 229-236. Nelson G, Platnick N. 1981. Systematics and Biogeography . New York: Columbia Univ Press. Nishida M, Nishida H. 1982. Histology of the rhizome of Loxsomopsis and affinity of Solenostelopsis loxsomoides Ogura. Acta Phytotax Geobot 33: 302 – 307. Ohta N, Takamiya M. 1999. Taxonomic studies of the Diplazium mettenianum complex Woodsiaceae; Pteridophyta in Japan: Morphology, cylotogy and taxonomy of plants with normal-shaped spores. J Plant Res 112: 67-86. Pacheco L. 2004. A new species of Diplazium Woodsiaceae from Ecuador. Brittonia 562: 121-123. Palmer JD, Jansen RK, Michael HJ, Chase DW, Manhart JH. 1988. Chloroplast DNA variation and plant phylogeny. Ann Missouri Bot Gard 75: 1180-1206. Park CH, Kato M. 2003. Apomixis in the interspecific triploid hybrid fern Cornopteris christenseniana Woodsiaceae. J Plant Res 116: 93-103. Parris BS, Beaman RS, Beaman, JH. 1992. The Plants of Mount Kinabalu. I. Ferns and Fern Allies. Kew: Royal Botanic Gardens. Parris BS. 2003. The distribution of Grammitidaceae Filicales inside and outside Malesia. Telopea 101: 451-466. Pettit JM. 1966. Exine stucture in some fossil and recent spores and pollen as revealed by light and electron microscopy. Bull Brit Mus Nat Hist Geol 13: 221-257. Pichi-Sermolli REG. 1977. Tentamen Pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 31 2: 313-512. Praptosuwiryo TNg. 1999. Biosystematic Study of Diplazium in Java. Program Pasca Sarjana, Institut Pertanian Bogor. [Thesis] 303 Praptosuwiryo TNg. 2003. Studi Kromosom Somatik Beberapa Jenis Tumbuhan Paku di Pulau Lombok dan Sumbawa. Widyariset 5 : 87-96. Praptosuwiryo TNg, Darnaedi D. 1994. Cytological Study of the Ferns Genus Diplazium in Gede-Pangrango National Park, Java. Floribunda 115: 57- 60. Praptosuwiryo TNg, Darnaedi D. 2004. Cytological Study of Some Species of Ferns Genus Diplazium in Java I. Floribunda 25: 128-137. Price MG. 1983. Several unusual Malesian diplazia. Gard Bull Sing 36 : 25-29. Qiu Y-J, White RA, Turner MD. 1995. The developmental anatomy of Metaxya rostrata Filicales: Metaxyaceae. Amer J Bot 82 8: 969-981. Queiroz A de, Donoghue MJ, Kim J. 1995. Separate versus combined analysis of phylogenetic evidence. Annu Rev Ecol Syst 26: 657-681. Rahbek C. 1995. The elevational gradient of species richness: a uniform pattern? Ecography 18: 200-205. Rangker TA. 1989. Spore morphology and generic delimitation of new world Hemionitis , Gymnopteris, and Bommeria Adiantaceae. Amer J Bot 762: 297-306. Ranker TA, Floyd SK, Trapp PG. 1994. Multiple colonization of Asplenium adiantum-nigrum onto the Hawaiian Archipelago. Evolution 48: 1364-1370. Ramsey J, Schemske WD. 1998. Pathways, mechanism, and rates of polyploidy formation in flowering plants. Ann Rev Ecol Syst 29: 467-501. Rifai MA. 1976. Sendi-sendi Botani Sistematika. Bogor: Lembaga Biologi Nasional – LIPI. Roos MC. 1993. State of affairs regarding Flora Malesiana: progress in revision work and publication schedule. Flora Malesiana Bulletin 11: 133-142. Roos M. 1995. Flora Malesiana: progress, needs and prospects. In: J. Dransfield et. al. Pp. 231-246. Plant Diversity in Malesia III. Proceedings of the Third International Flora Malesiana Symposium. Roubik, D.W. 2003. Pollen and spores of Barro Colorado Islands. 2003. http:striweb.si.eduroubik [24 Feb. 2007]. Rouhan G, Dubuisson J-Y, Rakotondrainibe F, Motley TJ, Mickel JT, Labat J-N, Moran RC. 2004. Molecular phylogeny of the fern genus Elaphoglossum 304 Elaphoglossaceae based on chloroplast non-coding DNA sequences: contributions of species from the Indian Ocean area. Mol Phylog Evol 33: 745 – 763. Sano R, Takamiya M, Kurita S, Ito M, Hasebe M. 2000a. Diplazium subsinuatum and Di. Tomitaroanum should be moved to Deparia according to molecular, morphological, and cytological characters. J Plant Res 113: 157- 163. Sano R, Takamiya M, Ito M, Kurita S, Hasebe M. 2000b. Phylogeny of the lady fern group, tribe Physematieae Dryopteridaceae, based on chloroplast rbcL gene sequences. Mol Phylog Evol 15: 403-413. Schölch A. 2000. Relations between sumbarginal and marginal sori in ferns. I. The sori of selected Hypolepidaceae and Dennstaedtiaceae. Plant Syst Evol 220: 161 – 183. Shmida A, Wilson MV. 1985. Biological determinants of species diversity. Journal of Biogeography 12: 1-20. Shivas MG. 1961. Contributions to the cytology and taxonomy of species of Polypodium in Europe and America: I. Cytology. J Linn Soc Bot 58: 13- 25. Skog JE, Mickel JT, Moran RC, Volovsek ME, Zimmer EA. 2004. Molecular studies of representative species in the fern genus Elaphoglossum Dryopteridaceae based on chloroplast DNA sequences rbcL, trnL-F, and rps4-trnS. Int J Plant Sci 165 6: …. Sledge WA. 1962. The athyrioid ferns of Ceylon. Bull Brit Mus Nat Hist Bot 2: 277-323. Smith AR. 1972. Comparison of fern and flowering plant distribution with some evolutionary interpretation for ferns. Biotropica 4: 4-9. Smith AR, Kreier H-P, Haufler CH, Ranker TA, Schneider H. 2006. Serpocaulon Polypodiaceae, a new genus segregated from Polypodium. Taxon 554: 919-930. Soltis DE, Soltis PS. 1998. Choosing an approach and an appropriate gene for phylogenetic analysis. In DE Soltis, PS Soltis, JJ Doyle. Pp 1-42. Mollecular 305 Systematics of Plant II. DNA Sequencing. Boston: Kluwer Academic Publishers. Soltis PS, Soltis DE, Smiley CJ. 1992. An rbcL sequences from a Miocene Taxodium Bald Cypress. Proceeding of the National Academy of Sciences U.S.A. 89:449-451. Soltis PS and Soltis DE. 2000. The role of genetic and genomic attributes in the success of polyploids. Proc Natl Acad Sci 97: 7051-7057. Stebbins GL. 1971. Chromosomal Evolution in Higher Plants. MA – USA: Addison – Wesley. Steenis, C.G.G.J. van. 1957. Specific and Infraspecific Delimitation. In: Flora Malesiana. Ser. I, Vol. 5. Steenis CGGJ van. 1981. Rheophytes of the World. Sijthoff Noordhoff, Alpena an den Rjn: Sijthoff Noordhoff. Strong EE, Lipscomb D. 1999. Character coding and inapplicable data. Cladistics 15: 363-371. Swartz, O. 1801. Achrad. J. Bot. 18002: 61. Swofford DL. 1998. PAUP. Phylogenetic Analysis Using Parsimony and Other Methods. Version 4. Sinauer Associates, Sunderland, Massachusetts. Tagawa M. 1972. Ferns of Borneo, collected by M Hirano and M. Hotta 6. Acta Phytotax Geobot. 25 2-3: 61-68. Tagawa M, Iwatsuki K. 1988. Flora of Thailand. Vol. 3. Part 3: 436-466. Takamiya M ed. 1996. Index to Chromosomes Number of Japanese Pteridophyta 1910-1996. Tokyo: Japan Pteridological Society. Takamiya M. 1997. Current status and problems of cytotaxonomical studies of Japanese pteridophytes. Bull Jap Pterid Soc 70: 1-23. Takamiya M, Takaoka C, Ohta N. 1999. Cytological and reproductive studies on Japanese Diplazium Woodsiaceae; Pteridophyta: Apomictic reproduction in Diplazium with evergreen bi- to tripinnate leaves. J Plant Res 112: 419-436. Takamiya M, Ohta N, Fujimaru-Takaoka C, Uki K. 2000. J Plant Res 113: 203- 215. 306 Takamiya M, Ohta N, Yatabe Y, Murakami N. 2001. Cytological, morphological, genetic, and molecular phylogenetic studies on intraspecific differentiation within Diplazium doederleinii Woodsiaceae: Pteridophyta. Int J Plant Sci 162 3: 625-636. Tardieu-Blot. 1932. Les Apleniees Du Tonkin. Theses. Prentees A la Faculte des Science de Paris. Toulouse Imprimerie henri Basuyau C’. 190p. Thiele K. 1993. The holy grail of the perfect characters: the cladistic treatment of morphometric data. Cladistics 9: 275-304. Tiegham P van, Douliot H. 1886. Sur la polystelie. Ann Sci Nat Bot Ser 7 3: 275 – 322. Tindale MD SK Roy. 2002. A Cytotaxonomic Survey of the Pteridophyta of Australia. Australian Syst Bot15: 839-937. Triono T. 2006. A Phylogenetic Study of Pouteria Sapotaceae in Malesia and Australia using Morphology and Molecular Data. The Australian National University. Canberra, Australia. [Doctoral Thesis] Tryon A. 1986. Stasis, diversity andfunction in spores based on an electron microscope survey of the Pterudophyta. In: Blackmore S, Ferguson IK, editors. Pollen and spores: form and function. Linnean Society Symposium Series No 12 : 233-249. London: Academic Press. Tryon AF, Lugardon B. 1991. Spore of the Ptridophyta. New York: Springer Verlag. Tseng-Chieng, H. 1981. Spore Flora of Taiwan. 1981. Taiwan: National Taiwan University. Tryon RM, Stolze RG. 1991. Pteridophyta of Peru. Part IV 17. Dryopteridaceae . Fieldiana No 17: 1-176. Tsutsumi,C. and Kato,M. 2006. Evolution of Epiphytes in Davalliaceae and Related Ferns. Bot J Linn Soc 151, 495-510. Van Alderwerelt van Rosenburgh CRWK. 1908. Malayan Ferns. Batavia: Landsdrukkerij. Van Konijnenburg-van Cittert JHA. 1975. Some notes on Marattia anglica from the Jurassic of Yorkshire. Rev Palaeobot Palynol 20: 205-214. 307 Van Konijnenburg-van Cittert JHA. 1999. On the evolution of fern spore anchitecture. In: Kurmann MH, Hemsley AR Eds. The Evolution of Plant Architecture. Pp 179-287. Kew: Royal Botanic Garden. Vida G. 1976. The role of polyploidy in evolution. In: Nivac VJA Pacltova, editors. Evolutionary Biology . Rechoslovak: Prague-CCzechoslovak Acad Sci Prague. Pp.267-294. Vogel JC, Rumsey FJ, Schneller JJ, Barret J, Gibby M. 1999. Where are the glacial refugia in Europe? Evidence from pteridophytes. Bot J Linn Soc 66: 23-37. Wang M-L, Chen Z-D, Zhang X-C, Lu S-G, Zhao G-F. 2003. Phylogeny of Athyriaceae : evidence from chloroplast trnL-F region sequences. Acta Phytotax Sin 41 5: 416-426. Wagner WH. 1961. Problems in the classification of ferns. Recent Advances in Botany. Pp. 841-844. University of Toronto Press. Wagner WH. 1972. Disjunctions in homosporous vascular plants. Ann Missouri Bot Gard 59: 203-217. Wagner WH. 1974. Structure of spores in relation to fern phylogeny. Ann Missouri Bot Gard

61: 332-353.