FERNS & FERN-ALLIES of EQUATORIAL EAST ASIA
Malaysia - Singapore - Brunei Darussalam - Indonesia - Philippines - Papua New Guinea

5th International Flora Malesiana Symposium

9-14 September 2001, SYDNEY, Australia

Pteridophyte Mini-symposium
a symposium and workshop on the ferns and fern allies of the Malesian region

Abstracts

V.B. Amoroso & L.V. Rufila, Central Mindanao University, Philippines
Updates on Philippine fens allies

This paper will present the current status of Philippine fern-allies. Present survey of fern allies showed 74 species, 6 genera an 5 families. The morphology, ecology, status relationship and distribution of the lower vascular plants in the Philippines will be discussed.

V.B. Amoroso, P.M. Zamora & L.V. Rufila, Central Mindanao University, Philippines
Species richness, assesment and distribution of Lycopodium in the Philippines

Examinations of herbarium vouchers and fresh specimens including its aerial stem anatomy have revealed 46 taxa of Lycopodium. Cladogram analysis of 31 species showed that the most primitive species of Lycopodium are isodichotomously branched, non-strobili forming and stem stele smaller. Of the total number of species, 1 is endangered, 9 are endemic, 23 rare, 18 economically important species, 11 new record in the Philippines and 2 possibly new species. With Luzon and Mindanao as important centers of Lycopodium distribution, the high mountain ecosystems are characterized by high species diversity starting from 900 m up to 2600 masl. Due to alarming forest destruction and over-collection of Lycopodium plants in the wild, the population is declining. To increase the population, an ex situ conservation was successfully conducted.

J.F. Barcelona & R.J. Hickey, Philippine National Herbarium, Philippines & Miami University, Oxford, Ohio, USA.
Systematics of the fern genus Odontosoria sensu lato (Lindsaeaceae)

In 1990, Kramer and Green expanded the genus Odontosoria to include essentially all lindseaoid species with continuous axial sulcation and laterally adnate indusia. As so defined, Odontosoria appears to be unnaturally broad and contains widely disparate elements. The goals of this study were to provide a set of data sufficiently extensive to confirm or reject this generic delimitation, as well as to define the generic and species boundaries among the taxa. Toward this end, a series of collecting trips in the Bahamas (1995), the Philippines (1996, 1997), Puerto Rico (1997), Hawaii (1997), and Jamaica (1997) was undertaken. These collections were studied in conjunction with 4,798 sheets of herbarium specimens representing 3,134 collection numbers and 35 photographs from 22 herbaria. One to 30 specimens per species were sampled for a total of 109 morphological and anatomical characters. Putative phylogenetic relationships of the odontosorioids as well as of the dennstaedtioid and the lindsaeoid genera are discussed in light of both traditional (morphological and anatomical) and/or molecular data. Odontosoria sensu Kramer and Green is considered to be a paraphyletic assemblage of taxa held together by pleisiomorphies. A generic circumscription for the Odontosoria complex is proposed which consists of five component genera, **** (four species and three varieties), **** (four species), **** (one species), Odontosoria sensu stricto (10 species), and Sphenomeris (five species), the first three genera being new. One new species, **** celebesiana, is to be described, one variety is elevated to species rank (**** divaricata), one variety of **** chinensis is renamed as var. capitata, and 14 new combinations are proposed.

**** = new unpublished genera

J.F. Barcelona & Tom Hollowel, Philippine National Herbarium, Manila, Philippines & U.S. National Herbarium, Washington, DC, USA.
The distribution of threatened endemic pteridophytes in the Philippines

The Philippines is home to 1,100+ species of pteridophytes in 142 genera and 39 families (my classification system). These data reveal that 10% of the total described species in the world today are concentrated in an area only approximately 0.06 % of the earth’s land. Of these, 290 species (48%) are found only in the Philippines and nowhere else in the world. A significant percentage of the country’s endemic pteridophytes (32.75% or 95 species) are known only from types, i.e. from single collections, 73% (69 species) of which were collected from Luzon (44 species) and Mindanao (25 species) alone. In a country where the population represents 1.25 % of that of the world’s, forest cover has been diminishing in an alarming rate. Current estimates place the remaining original forest cover in the Philippines to be only less than 10%. Large scale commercial logging, slash and burn agriculture, and indiscriminate urbanization have been the major threats to the integrity of the country’s biodiversity, particularly the pteridophytes. In addition to intensive field studies of this plant group, rallying for awareness to our environmental problems through outreach educational programs, and possibly, in situ conservation, seem to be the only remaining option to save these plants from extinction.

H. Beukema & M. van Noordwijk, University of Groningen, Groningen, Netherlands, ICRAF S.E.Asia, Bogor, Indonesia
Biodiversity conservation in rubber agroforests?

Most rubber on Sumatra, introduced as a plantation tree, is produced in what looks like native forest to an outsider. This project assessed the biodiversity associated with smallholder rubber agroforests, known as 'jungle rubber'. Planted and managed by local farmers, these agroforests provide refuge for plants and animals of the natural forest that is rapidly disappearing in Sumatra. Can these agroforests, planted to provide owners with income from tapping rubber trees, really contribute to the conservation of tropical rain forest species? Plants and animals from neighbouring forest or agroforest areas establish themselves spontaneously in the gardens. Often only half of the trees in these agroforests are rubber trees. But are the species found in jungle rubber really the ones that are under threat? Limiting the scope of research to one indicator group, we studied ferns to compare jungle rubber with natural forest and with intensively managed rubber plantations in Jambi province, Sumatra. We assessed the share of 'forest species' in the total species richness of jungle rubber, and the degree to which rubber gardens are repetitive in species composition over larger areas All ferns were documented in 51 plots distributed over an area of about 200 by 150 kilometres. The results show that the average number of fern species per 40 x 40 m plot is approximately the same for forests, agroforests and rubber plantations. However, the species composition and variation beyond plot scale differed substantially between the three land use types. Ferns found growing in rubber plantation plots were mostly species that prefer open habitat. Jungle rubber showed the highest variation, but only about half of the species were classified as 'forest ferns'. As undisturbed forest was found to be about twice as rich and varied in 'forest ferns' as jungle rubber, the researchers conclude that agroforests such as jungle rubber are certainly no substitute for forest conservation. Priority for conservation of more natural forest types remains the only chance for the more sensitive species of rain forest flora and fauna to survive. Yet, in areas where most of the natural forest is gone, the extensive agroforest management style can make a substantial contribution to biodiversity conservation when compared to the alternative of intensively managed plantations.

P.J. Brownsey, Te Papa Tongarewa, National Museum of New Zealand, Wellington, New Zealand
New Zealand's pteridophyte flora - plants of ancient lineage but recent arrival?

A hypothesis is presented that most pteridophytes have arrived in New Zealand relatively recently by long-distance dispersal. The flora comprises 194 native species, of which 89 (46%) are endemic and 105 (54%) are widespread. Of the latter, 90% are shared with temperate Australasia, 53% with tropical Pacific and Malesian regions, 14 % with temperate southern Africa and 13% with the circum-antarctic islands and South America. New Zealand has undergone such dramatic changes in location, land area and topography since initial separation from Gondwana 85 Ma that it seems improbable that the 95 species shared with temperate Australasia could have remained conspecific throughout that time. Modern fossil and molecular evidence strongly suggest that many families of ferns had not even evolved prior to separation, and palynological evidence from New Zealand indicates that 78% of pteridophyte genera first appeared there only after separation from Gondwana. Present-day distributions in New Zealand suggest that ferns have greater dispersal potential than flowering plants, and that pteridophyte distributions are more heavily influenced by temperature, rainfall and geothermal activity than by geological history. Most endemic pteridophyte species have a predominantly southern distribution pattern and are characteristic of cool, lowland to montane forest. Pteridophytes in the northern part of New Zealand show a lower level of endemism and tend to be widespread species that have arrived from temperate Australasian and tropical regions. There is also evidence that some pteridophytes have migrated from New Zealand to Australia. It is suggested that the hypothesis of long-distance dispersal of pteridophytes across the Tasman Sea could be tested by molecular techniques, and is as relevant to Malesia as it is to Australasia.

J.M. Camus, Department of Botany, The Natural History Museum, London, UK
Pteridophyte Flora of Sulawesi, Indonesia

Sulawesi first attracted the attention of biologists in the 1800's when Alfred Russell Wallace found its fauna a strange combination of Asian and Australasian animals. This island has been central to much biogeographic discussion ever since. The island of Sulawesi lies on the equator near the rim of the Pacific Basin between continental Asia and Australia. The curious distorted shape of the island is the result of a series of dramatic geological events involving the collision and fusion of land masses of Sundaic and Australia-New Guinea origin. Different theories have been proposed suggesting past land links with the Moluccas and New Guinea to the East, The Philippines to the North, and Borneo and the Sunda Islands to the West and South. The physical history of the island makes it one of the most biogeographically complex regions of the world. The distribution patterns of the components of its pteridophyte flora echoe this. The island is mountainous throughout with the highest peak at 3505m and has active volcanoes, thermal springs, alluvial and sedimentary deposits. The terrain and climate have produced a number of vegetation types including mangrove and peat swamp forests, limestone monsoon rainforest and tropical montane rainforest. These provide a great diversity of habitats for ferns and fern allies. The pteridophyte flora of Sulawesi is poorly documented. Christ's publications on the fern flora of Celebes in 1898 and 1904 gave 337 species of pteridophytes. The current total is just over 500 taxa and some interesting range extensions have been discovered. Thorough exploration of the island would no doubt yield more data. The four main collecting trips by pteridologists during the 20th Century concentrated on montane areas so more information is needed for the lowland areas. Although relatively limited, current data allows some assessment of the degree of protection afforded to the pteridophyte flora by the conservation areas of the island.

R. Cranfill, Y. Nakahira & M. Kato, University of California, Berkeley, USA & University of Tokyo, Japan
A molecular phylogeny of Blechnaceae inferred from three plastid gene sequences

The Blechnaceae are composed of 200-250 species assigned to about nine genera, which have been defined by a few characters such as venation, indusium, sorus, etc. To understand the intergeneric and interspecific relationships of the family, a molecular phylogenetic analysis was made based on the rbcL, rps4 coding gene sequences and the non-coding trnS intergenic spacer. The analysis demonstrates that the family is monophyletic with high bootstrap support. The family itself is divided into two major clades with high bootstrap support: Woodwardia and the rest of the family. Woodwardia is a monophyletic group that includes species referred to the segregate genera Anchistea, Chieniopteris and Lorinseria and may retain primitive characters such as reticulate venation, and discrete sori. Stenochlaena is basal in the other clade, along with Salpichlaena and Blechnum serrulatum. Brainea, with exindusiate sori, is the next to diverge and appears to be isolated. Other primitive groups include Sadleria, Struthiopteris and the Blechnum orientale group. The isolated position of Sadleria, which is today endemic to Hawaii, suggests that it may have evolved originally in eastern or southeastern Asia with subsequent dispersal to Hawaii followed by extinction on the mainland. These basal-most genera and species groups are all restricted to or predominantly Laurasian in distribution, while the crown group of the family is predominantly Gondwanan in distribution. Within the crown group, Pteridoblechnum, Steenisioblechnum and Blechnum diversifolium form a clade, which in turn is sister to Blechnum fraseri. Doodia appears to have a biphyletic origin, with Doodia maxima sister to Blechnum brasiliense and with the remaining species sister to the Blechnum gibbum group of New Caledonian and Australia. Brainea, Doodia, Pteridoblechum, Sadleria, and Steenisioblechnum, which have been treated as monotypic or oligospecific genera, are nested within the genus Blechnum as currently treated in monographs and floras. It indicates that Blechnum is a paraphyletic unit in need of division into a number of smaller monophyletic genera. The tree also suggests that the separate sori of Woodwardia are primitive, whereas similar sori of Doodia, Pteridoblechnum, and Steenisioblechnum are secondarily interrupted ones derived from the coenosori of Blechnum, associated with the evolution from free venation to reticulate venation.

J.R. Croft, Australian National Herbarium, Canberra, Australia
On-line Flora of Australia pteridophytes: an information model for an electronic Flora Malesiana series 2.

While filling a valuable niche in the the area of biodiversity knowledge, traditional floras have major limitations in the way they store, present and distribute information. A collaborative project between the Centre for Plant Biodiversity Research and the Australian Biological Resources Study is investigating ways to store information from the printed volumes of the Flora of Australia in a relational database and deliver this informtion via the Internet in a variety of formats for different purposes. Text and images (line drawings and photographs) are being included. The pteridophyte volumes of Volumes 48 (mainland Australia) and volumes 49 & 50 (Australian oceanic islands) are being used as one of the prototypes for this treatment. The text and image information from the published floras is converted from its word-processor format to a structured XML datafile and loaded into an Oracle relational database from where it can be exported entirely or in part in response to standard database queries. The internet gateway to the database allows fields of flora information to be delivered to standard browsers and other devices in XML and HTML formats in a vast range of definable styles. It is proposed that future volumes of the flora be compiled directly into the database, from which camera ready copy will be generated; This not only allows for greater comparability and consistency of the data provided, but provided a mechanism for on-going maiantence of the data as new information becomes available. Published floras have a lot in common and only minor adjustments would be needed to the Flora of Australia data structure to enable it to used for Flora Malesiana treatments; this approach is recommended to the editors.

Y.B. Farida & B.C. Tan, Cryptogamic Laboratory, National University of Singapore, Singaporre
A comparison of pteridophytic diversity of various forests in Malaysia using quadrat sampling method

A study on the diversity of ferns and fern allies was carried out in various rainforests at different elevations and places in Malaysia (Malaya Peninsula and Sarawak) using quadrats of different sizes. The objectives are: 1) to compare the biodiversity of fern and fern allies in various rainforests at different elevations, and 2) to assess the effectiveness of quadrat sizes in capturing the species richness of rain forests in Malaysia. The results show that the fern diversity is low per unit area when compared to the diversity pattern at high elevations where there are more number of taxa per unit area. As such, there is a need to conduct more quadrat samplings in the lowland rain forest than in montane forest if only to capture the total biodiversity in the lowland rain forests.

R.J. Johns, Royal Botanic Gardens, Kew, UK
Observations based on field studies of Cyathea (Cyatheaceae) from Mt Jaya, New Guinea

Most herbarium collections of tree fern consist of a single fertile pinna, sometimes with a small piece of attached rachis, and often lack even a piece of stipe or such basic measurements as the length of the stipe and rachis. Consequently the majority of herbarium collections of Cyathea are completely inadequate for the careful description of the species. A field proforma has been developed for the collection of field data for Cyathea and Dicksonia. The results of the field studies using these data sheets for Cyathea collections are presented with particular reference to the high altitude tree ferns from Mt Jaya (Papua, Indonesia). Using the field data it is possible to make a detailed key to the species - difficulties however arise when trying to apply the published species names to the 'field' species. Other characters used for the identification of the Cyathea species in New Guinea are discussed, in particular characters resulting from the detailed study of the morphology using the SEM - scales, hairs, perispore ornamentation etc. Where practical trunk material was collected for anatomical studies. The sections of the Cyathea trunks show a marked uniformity in anatomy within a single plant (species), but marked differences in anatomy between the different species. These anatomical differences are discussed.

R.J. Johns, Royal Botanic Gardens, Kew, UK
Asplenium section Thamnopteris (Aspleniaceae) - new information leading to a better taxonomy of the section

In 1976 Holttum published the first overview of the species which he included in Asplenium Section Thamnopteris. New field studies and detailed morphological work, particularly using SEM studies of the perispore, provide powerful tools for the identification of species within this section. In 1950 Baker and Posthumus identified A. nidus as the only member of Section Thamnopteris in Java. Murakame collected several specimens from Java for DNA studies. The perispore morphology of the Murakame specimens from Java has been studied and the differences in ornamentation supports the conclusions on species delimitation based on the DNA studies by Murakame. Independently Johns collected two species from West Java around the type locality of A. nidus. DNA analysis by Murakame shows that these two species differ from his collections in Java. The perispore ornamentation of both also differs from the Murakame specimens. These recent studies of perispore ornamentation show that section Thamnopteris is represented by at least seven species in Java. What is Asplenium nidus? As the type specimen of Section Thamnopteris (Linnean Herbarium) from West Java is infertile and lacks scales, the application of this name is difficult. Most collections in horticultural trade are not A. nidus. One of the two species collected from near the type locality in Java probably represents 'true A. nidus'. It is proposed that an epitype for Asplenium nidus L. be selected (Article 7). The taxonomy of the species in Java is discussed. New Guinea is also an important centre of diversity within the section and an overview is given of species in this area and also brief comments made on the section in East Africa/Madagascar. Further work on perispore ornamentation could provide a valuable tool for developing our understanding of the natural sections within Asplenium.

S. Lindsay, Harvard University Herbaria, Cambridge, MA, USA
A change in the generic framework of the fern family Vittariaceae in Malesia.

In 1998 a paper appeared in the American journal Systematic Botany (vol. 22, no. 3, pp. 509-517) that boldly challenged the traditional concepts of genera in the fern family Vittariaceae. The new generic circumscriptions proposed in that paper appear, as yet, to have had little impact on the curatorial practices in most herbaria and to have been over-looked in a number of recent publications. Nevertheless, the biggest change proposed in that paper will be adopted in the forthcoming revision of Vittariaceae for Flora Malesiana. This could result in the complete loss of the genus Vittaria from Malesia with most (or all?) species traditionally recognized as Vittaria (in that region) being moved to the genus Haplopteris (an old and abandoned genus resurrected in 1998); 11 Malesian "Vittaria" species have already been re-classified including those as well known as Vittaria elongata Sw. [now Haplopteris elongata (Sw.) E.H.Crane], Vittaria ensiformis Sw. [now Haplopteris ensiformis (Sw.) E.H. Crane], Vittaria scolopendrina (Bory) Schkuhr ex Thwaites & Hook. [now, Haplopteris scolopendrina (Bory) C.Presl] and Vittaria zosterifolia Willd. [now, Haplopteris zosterifolia (Willd.) E.H.Crane]. Decisions on whether other Malesian "Vittaria" species (approximately 30) should be moved to Haplopteris, moved elsewhere, or treated as synonyms, will be dependent on the outcome of a thorough comparison of micro-morphological characters.

N. Murakami & Y. Yatabe, Kyoto University, Japan
Recognition of biological species in Asplenium nidus complex using molecular data and crossing experiments

Bird-nest fern, Asplenium nidus is a well known epiphytic fern species with large simple leaves. Since A. nidus lacks the good morphological characters available for species recognition, multiple cryptic species may be contained within this species. Recently, it has become easier to collect DNA nucleotide sequence data from wild plants using PCR and direct sequencing techniques. This kind of molecular information has mostly been used for phylogenetic analyses, but can also be useful for a-taxonomy (recognition of naturally existing species). In Mt. Halimun National Park, West Java, Indonesia, we found five rbcL sequence types of A. nidus. All plants regardless of rbcL sequence were tetraploids of 2n = 144 based on our cytological observation. According to the molecular tree based on the comparison of rbcL sequences with the plants of other localities, the five rbcL types growing in Mt. Halimun National Park were shown to be not closely related to each other. Although these rbcL types were morphologically continuous, ecological differentiation, especially in growing altitude or in position where they grow on the tree trunks, was observed. Crossing experiments among these rbcL types revealed that they are also reproductively isolated because hybrids failed to form between at some pairs of the rbcL types. Moreover, we examined allozyme polymorphisms of the five rbcL types of Asplenium nidus in order to clarify whether these assemblages of rbcL are genetically distinct populations also in their nuclear genomes. It was shown that these rbcL types are differentiated in allozyme loci even in the locality where several rbcL types are sympatrically distributed. These results strongly suggest that these rbcL types are cryptic species because they are reproductively isolated, genetically and ecologically well differentiated. The three rbcL types in Mt. Halimun National Park should be treated as separate species even though they are difficult to be distinguish only by their gross morphology.

B.S.Parris, Fern Research Foundation, Kerikeri, Bay of Islands, New Zealand
Biogeography of Grammitidaceae (Filicales) inside and outside Malesia

More than 700 species of Grammitidaceae are known. At least 250 occur in the New World and at least 450 in the Old World. Of the Old World species, > 300 are in Malesia, > 140 are in the Pacific including Australia and New Zealand, > 60 are in Asia excluding Malesia and > 50 are in Africa including Macaronesia, Madagascar and the Mascarene Islands. Lomaphlebia (2 spp.) is endemic to the New World; Ceradenia (c. 50 spp.), Cochlidium (16 spp.), Enterosora (c. 9 spp.), Melpomene (c. 25 spp.) and Terpsichore (c. 56 spp.) are centred in the New World, each with 3 or fewer species in Africa; Micropolypodium (c. 29 spp.) is New World-centred with up to 3 Asian/Malesian species; Lellingeria (c. 63 spp.) is New World-centred with 9 African species and 2 Pacific species; Grammitis sensu stricto has 14 New World spp., 7 African spp. and 4 Pacific spp.; Zygophlebia has at least 15 African spp. and 7 New World spp. Acrosorus (4 spp.), Calymmodon (c. 47 spp.), Chrysogrammitis (2 spp.), Prosaptia (c. 50 spp.), Scleroglossum (c. 10 spp.) and Themelium (6 spp.) are centred in Malesia and extend to Asia and/or the Pacific, as are several species groups of Ctenopteris, Grammitis sensu lato and Xiphopteris. Adenophorus (6 spp.) and Oligadenus (4 spp.) are endemic to the Pacific. A few species groups of Grammitis sensu lato are endemic to Africa. The Ctenopteris blechnoides species group has 8 species in the Asia-Malesia-Pacific region and 2 in Africa. The Grammitis billardierei species group is largely in Old and New World south temperate regions (14 spp.), with 1 species in Malesia and 2 in tropical Africa.

L. Perrie, Botany Department, Massey University, New Zealand
Investigating species boundaries : morphology and molecules

Many aspects of taxonomy have benefited greatly from molecular methods. However, given their expense, it is unlikely that molecular methods will challenge the primacy of morphological analysis in alpha taxonomy. Instead, molecular methods are best viewed as tools to investigate taxonomically difficult groups whose boundaries, but not constituents, have been resolved in the alpha taxonomy. An example is our use of AFLP (Amplified Fragment Length Polymorphism) DNA fingerprinting in the characterisation of New Zealand's 'Polystichum richardii' as an allopolyploid complex.

T. N. Praptosuwiryo and D. Darnaedi, Bogor Botanical Gardens, Bogor, Indonesia
The fern genus Diplazium in Indonesia

The study on fern genus Diplazium in Java was conducted based on gross morphology by examining 498 collections of specimens housed at Herbarium Bogoriense (BO) and 32 collections of living plants representing 10 species collected from 6 sites of Java. For supporting taxonomic evidence, anatomical, palinological, and cytological investigations were conducted. It is concluded that Javanese Diplazium comprises of 22 species with 4 varieties. Seven species, namely D. kustleri, D. latisquamatum, D. procumbens, D. riparium, D. simplicivenium, D. subpolypodioides and D. xiphophyllum are newly recorded for this island. One species concept has been changed by treating D. montanum as variety of D. pallidum. Anatomical evidence of the stipe is found to be useful to support species delimitation, but the distribution of stomatal types cannot be used for taxonomix purpose. Perine ornamentation is expected to represent good character for supporting species delimitation, but the data obtained from light microscope observation still need to be associated with the data obtained from scanning electron analysis. Four ploidy levels with x=41, diploid (D. accedens, D. esculentum, D. polypodioides, D. umbrosum), triploid (D. dilatatum, D. procumbens), tetraploid (D. bantamense, D. cordifolium, D. lobbianum, D. pallidum) and pentaploid (D. cordifolium) are reported. Two cytotypes of different ploidy levels of one species are not different morphologically. It is pressumedthat that the polyploidy mechanism occurred in the species of Javanese Diplazium investigated are autopolyploid. Further studies in this respect are needed.

G. Rusea, Universiti Putra Malaysia, Serdang Selangor, Malaysia
Towards the revision of Malaysian Lycopodiaceae

The family Lycopodiaceae in Malaysia consists of 27 known species from 3 genera, namely the Huperzia, Lycopodium and Lycopodiella. Currently there is no complete revision on this group of fern-allies and all publications related scattered in Kiew et. al. (1987), Parris (1997a, 1997b, 1997c; Parris et al. (1992); Parris & Edwards (1988); Tagawa (1967, 1974). Parris & Latiff (1997) published a cornerstone accounts on provisional checklist of Malaysian pteridophyte taxa which its currently employed as baseline information for this research. Turner's (1995) Catalogue of the Vascular Plants of Malaya is also another resourceful guideline. Holttum's account on ferns of Malaya does not include Lycopodiaceae, therefore this is the first attempt to put together all the known species as well as a potential new findings as research on herbarium collections and fresh collections from unbotanized areas continues. This revision is partly to revise Holtum's Ferns of Malaya first printed in 1965 as well as venturing into producing a Fern Flora of Malaysia. This taxonomic revision will utilize morphological and molecular evidence to solve some taxonomic uncertainties within this family.

Siti Khadijah Rambe, Nanyang Technological University, Singapore
Distribution of ferns in Malesia: the Aspleniaceae as a case study

The Phytogeography of ferns in Malesia is poorly known, despite the fact that studies on ferns have been conducted in this region since the mid- 1700's. As a region with an extremely rich fern flora, the phytogeography of Malesian ferns is well worth investigating. The Malesian region contains 4,400 species of pterodiphyte of which is 5.55% of this total belongs to the Aspleniaceae. Out of a total of 700 species, Aspleniaceae in Malesia comprise 35% of the family total, which is higher than previous estimates. Even though wind can distribute spores widely, there are localized patterns of fern distribution in Malesia. It was found there are recognizable groups of Malesian, Australian and Indochinese Aspleniaceae. Moreover, within Malesia, Aspleniaceae exhibit distinct distribution patterns, such as the West Malesian, East Malesian and South Malesian elements. Factors contributing to these patterns of distribution are believed to include: (1) wind, (2) position ! of Islands groups in Malesia, (3) topography and (4) spores size and architecture.

Siti Khadijah Rambe, Nanyang Technological University, Singapore
Important biological characters used for the taxonomic study of Aspleniaceae

According to previous researchers, vascular bundle characters define the family Aspleniaceae. Within the family, only Hymenasplenium is believed to be a natural genus in the Aspleniaceae due to the dorsiventral vascular bundles in the rhizome, which in Asplenium are radial. Due to highly variable morphological and anatomical characters in Aspleniaceae, molecular studies were conducted in order to answer taxonomic problems in the family. How reliable is single character data to define genera in the Aspleniaceae? What combination of characters - morphological, anatomical, or molecular will be required to define the genera of Aspleniaceae?

Y. Yatabe, H. Nishida, N. Murakami & K. Iwatsuki, Kyoto University, Chuo University &The University of the Air, Japan
Phylogeny of Osmundaceae inferred from rbcL nucleotide sequences and comparison to the fossil evidences

We determined rbcL sequences of 11 of 15 extant species of Osmundaceae which represent all three genera, Osmunda, Todea and Leptopteris. Our phylogenetic analysis concluded: 1) Osmunda subg. Osmunda and subg. Plenasium are monophyletic groups, but subg. Osmundastrum is not. The genus Osmunda is not monophyletic because Todea and Leptopteris are positioned within Osmunda. 2) Osmunda cinnamomea is the most basally positioned species in Osmundaceae, and it can be called as "a living fossil" because a fossil species (O. claytoniites) with almost the same morphology as this species was recorded from the Triassic. 3) Greater nucleotide variation was found among conspecific samples of O. cinnamomea, O. claytoniana and O. regalis collected from various localities. Each of these species may comprise more than two biologically differentiated species.


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