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Biodi Borneo's ~CDtatBlical Secret

Although Borneo's soils are poor in nutrients, they have, over millions of years, spawned a rich biodiversity. The island enjoys a humid tropical climate with temperatures ranging between 25°C and 35°C at sea level and a yearly rainfall of between 2,000 and 4,000 millimetres (MacKinnon et a/., 1996). More importantly, it has maintained its tropical position uninterruptedly over almost 100 million years. Whereas continental drift has pushed Borneo's neighbours like the Philippines, Sulawesi, and New Guinea to and from different climate zones, Borneo has stayed put at its balmy equatorial location and evolution has been able to develop an immensely varied flora and fauna (Hall & Holloway, 1998; Hall, 2006).

Sadly, the past decades have put an end to Borneo as a pristine tropical paradise. Large-scale logging of tropical hardwood and, later, clearing of forest for oil palm and other plantations have increased to such an extent that a large part of Borneo is expected to have lost all of its lowland forest (the richest forest type in the world) by 2010 (Jepson et a/., 2001). With destruction pushing higher and higher up the mountain slopes, immediate action is needed to save as large an area

of forest as possible. The Heart of Borneo is an initiative aimed at preserving a continuous 220,000-square-kilometre area in the

centre of the island, which combines parts of all three countries

(Brunei Darussalam, Indonesia, and Malaysia) that share

J ,

l~

Borneo. This report describes the plant biodiversity

in this area, its value and future generations' losses if it were to disappear.

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IfJ Forest cover on Borneo in 2005.

The heart of the island is the only area where contiguous forest survives.

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Image Credil : 2 WWF Germany

Biodiscoveries - Borneo's Botanical Secret 5

A thousand shades of green

For most of the past 30 million years, Borneo was part of a single land mass (,Sundaland') that also included Java, Sumatra, Peninsular Malaysia, and the seas in between. The last time this happened was during the last 'ice age' just 12,000 years ago, when sea levels were more than 100 metres

lower than today (Voris, 2000). In those days, animals could roam between Borneo and mainland Asia without having to cross any seas. No wonder, then, that the Bornean fauna is very similar to the ones of other parts of Sundaland. For example, only 9% of Borneo's bird species are unique to the island (MacKinnon et al., 1996).

In plants, the situation is different. Because the pollen or seeds of many plants do not travel far, and also because many have adapted to Bomeo's high mountains and unusual rock types, Borneo's flora has developed in a unique way. From early on, botanists have realised that the island offers rich pickings, and many focused their attentions on the fascinating Mount Kinabalu (Wong & Phillipps, 1996). One of the first was Hugh Low, the colonial secretary for Labuan and, coming from a horticultural family, keenly interested

in plants. He was the first European to climb the mountain in 1851 and then twice more in 1858, collecting many plant specimens as he went along.

Since then until the Second World War, important contributions were made by a long line of European botanists. In the second part of the 20th century, Bornean botany was more and more conducted during large expeditions by local and overseas' universities and research organisations. Yet although one may think that after so much work, not much new remains to be discovered,

the rate of finding new species does not appear to be letting up. Over the past 25 years, at least 422 new species were added to the list (see Appendix), and dozens more still await study and formal naming.

EI Hoya waymaniae, II H. nabawanensis, and B H. kloppenburgii, three stunningly beautiful species of milkweed that were recently discovered in Sabah.

Photo Credit: 3, 4, 5 Peter Koomen

Today, we know that the island has the richest plant life among the areas that once formed Sundaland. It boasts a staggering 15,000 species of flowering plants, a diversity as great as all of Africa, more than three times that of Java (MacKinnon et al., 1996), and more than ten times that of European countries like the Netherlands (van der Meijden et al., 1995). This includes some 1 ,700 species of orchid, and 3,000 species of tree. In addition to flowering plants, Borneo has more than 1 ,000 species of ferns, many of which live on Mount Kinabalu, which has more fern species than all of Africa (for comparison: The Netherlands only have 50 species) and 1 ,344 species of bryophytes, the mosses and liverworts (see Table 1 for some more figures on species richness in different plant groups). The diversity of algae must be enormous, too, but this group has been much neglected by scientists so far.

For these reasons, Borneo is listed among the six areas in the world with the highest plant diversity (Bartlott et al., 1996). It applies to a small scale as well: the island holds the world record for tree species richness, with a mindboggling 1 ,186 species in a 0.5 square kilometre-chunk of Sarawak forest (Condit et al., 2000; Wills et al., 2006). Forests in Peninsular Malaysia are at most only two-thirds as rich (Harrison & Ashton, 2003).

iE, m Two of Borneo's more than 1,000 fern species: the epiphytic staghorn fern, Platycerium coronarium and the bracken fern, Oicronopteris sp.

Borneo is home to 721 species of mosses and 623 species of liverworts.

Photo Credit: 10, Menno Schilthuizen 11,12 Peter Koomen

A selection of Borneo's orchids:

III Pha/aenopsis philippinense, D Ph. fuscata,

B Ph. pantherina, and II Paphiopedilum rothschildianum.

Photo Credit: 6, 7, 8, 9 Peter Koomen

Not only is Borneo rich in species, many of the species are unique to the island as well. Over 5,000 species or 34% of the flowering plants are endemic: they are found nowhere else on earth (Wilson, 1994). Compare this with, say, Sumatra (with only 12% endemic plant species) and the botanical value of Borneo becomes clear. Among some groups, the percentage of endemicity is even higher: 58% of the highly-valued timber tree family Dipterocarpaceae, for example, are unique to the island. One family of small trees, the Scyphostegiaceae, is even entirely confined to Borneo.

Besides the large numbers of species, the Bornean flora is one of other kinds of superlatives as well. It has two contenders (an aroid and a Macaranga tree) that vie for the honour of having the world's largest undivided leaf (yorath & Teo, 1994). It also hosts the world's largest moss (a Oawsonia species), the world's largest pitcher plant (of four litre capacity), and eight species of the world's largest flower, Raft/esia (up to one metre across), a parasite that does not have any chlorophyll of its own (Nais, 2001). Another oddity is the fig Ficus albipila, one of the world's rarest trees: besides 1 or 2 individuals growing along the banks of Kinabatangan river in Sabah and the upper Tabalong River in South Kalimantan, it only occurs as a handful of scattered

trees in Australia and Peninsular Malaysia (Azmi, 2003).

Truly special assemblages of plants occur on those patches of Bomeo that have unusual physical propeties, such as the extremely cold and wet summit of Mount Kinabalu, the nutrient-poor 'kerangas' soils, and ultramafic rocks (very acidic and rich in toxic heavy metals). Another example is limestone. Limestone outcrops are scattered in their hundreds over Borneo's lowlands. The flora of such 'karst' hills is unusual, because not all plants can tolerate the alkaline, dry, steep, and calcium-rich substrate. Some limestone plants are so specialised that they cannot live anywhere else, and many species of herbs, rattans, screw-pines, and other plants are only found here (Kiew, 2001). In fact, among all lowland forest types, limestone forest has the fewest tree species in common with the other types (Procter et al., 1982). Not only that, but many

Borneo (% endemic) Mt. Kinabalu Netherlands
Flowering plants
Total ca. 15,000 (34%) ca. 5,000 ca. 1,400
Orchids ca. 1,700 840 30
Aroids 160 40 10
Trees 3,000 59
Dipterocarps 265-270 (58%) 0
Palms 280 0
Rattans 146 0
Figs 135 0
Non-flowering plants
Ferns >1,000 608 50
Bryophytes 1,344 234 507 Table 1: Figures on species richness and endemism for selected plant groups in Borneo and Mt. Kinabalu. Based on Akiyama et a/. (2001); Andersen et a/. (2003); Beaman (2001); Beaman & Beaman (1998); Dransfield & Johnson (1989); Lamb (pers. comm.); MacKinnon et aI. (1996); Payne et a/. (2000); Phillipps & Liew (2000); Rautner & Alfred (2005); Said (2005); Suleiman (pers. comm.);

Suleiman & Akiyama (2005); Suleiman & Edwards (2002); van der Meijden et a/. (1995);

van der Meijden (2005); Wong & Phillipps (1996).

are to just a single limestone outcrop. Batu Punggul in Sabah, for instance, is home to three recently discovered species, the orchids Ania ponggolensis and T richoglottis tinekeae, and Begonia amphioxus all of which occur nowhere else on earth (Kiew, 2001).

1m Borneo's lowland rainforest can contain over 1,000 species of tree in half a square kilometre.

1m Limestone outcrops, like Batu Punggul in Sabah, are home to unique vegetation, often with species endemic to just a single outcrop.

Photo Credit: 15 Peter Keemen, 16 Menne Schilthuizen

Biodiscoveries - Borneo's Botanical Secret 9

Botan'ea treasure trove

10

Biodiscoveries - Borneo's Botanical Secret

11

Another historically important export product is the nut of various species of Shorea trees. These 'illipe nuts' or 'tengkawang' are harvested from the forest and exported as nuts or as illipe oil for use as cocoa butter substitute, as well as in soap, candles, medicines and 'rainforest cosmetics'. Like most dipterocarps, Shorea trees fruit irregularly, once every seven years or so (MacKinnon et al., 1996). In a fruiting year, a hectare of forest floor may yield over one tonne of dried nuts, and total export can be worth up to ten million US$ (Iqbal, 1993).

However, these figures are dwarfed by those for the spiny, climbing palms that we know as rattan (the palm subfamily Lepidocaryoidae). Borneo is home to almost 150 species, which snake through the vegetation in abundance. Many of them are coveted for their long, regular, and hard stems that, after de-spining and boiling, can be turned into light-weight but strong and durable furniture. Calamus species such as C. caesius and C. optimus produce thin (less than 17 mm in diameter) but very long (more than 50 m) stems, whereas C. scipionum, C. omatus, and C. subinermis yield the thicker, but equally long, variety (Pearce, 1989; Chan, 2000). Throughout the 1990s, Indonesia and Malaysia exported an average of 125,000 tonnes of rattan and rattan products (valued about 125 million US$) per year (Vantomme et al., 2002), and much of the raw material came from Bornean forests.

or ages, the forests of Borneo have been a source of valued plant-derived materials, although the importing countries, the popularity of the commodities, and their uses have varied much over the centuries. Already two millennia ago, the Chinese traded on Borneo's shores to obtain coveted goods like tree resins, rattan, nuts, bananas, and fragrant 'gaharu' wood (Dunn, 1975; Whitmore, 1998). Since then, Borneo has provided the outside world with anything from charcoal and latex to fibres and fruits. Today, timber and rattan are the only remaining multimillion-dollar export products derived from Borneo's natural vegetation. However, a very large range of forest products continue to be gathered by the local communities for food, medicine, construction, and many other uses. A lot of those could also be commercialised in the future--provided the forest that harbours them survives.

One of the most important export products in the early days was the tree resin 'damar', which was (and still is) tapped from Shorea trees and the tall conifers Agathis bomeensis, A. dammara, and A. lenticula. A. bomeensis grows in lowlands, the other two in highlands. Originally, damar was used as fuel for lighting fires and to repair boats, but later it mostly found use in the preparation of paints, varnish, and linoleum (MacKinnon et al., 1996; Payne et al., 2000; Whitmore, 1998). Even today, Indonesia and Malaysia still export yearly some 2,000 tonnes (worth about two million US$) of damar for the production of specialised paints and varnishes, and much of it comes from Borneo (FAO, 1995).

An Agathis tree oozing the valued 'damar', and the dried resin.

1m Rattan after boiling in oil Photo Credit: 19 WWF-Malaysia / LYS

Photo Credit: 17. 18 Aida Lapis

Biodiscoveries - Borneo's Botanical Secret

Although rattan exports are important, most revenue comes, of course, from timber. Among the 3,000 tree species that grow in Borneo's forests are many with ideal properties for use as wood in construction and - to a lesser extent - for furniture and floors. The most popular timber family is

the Dipterocarpaceae, but the most highly-valued species is actually a member of the laurel family: Borneo ironwood or 'belian' (Eusideroxylon zwagert). Untreated, its timber can last for up to a century. Hence, it was (and is) the wood of choice for

the main beams of Borneo longhouses. Other much sought-after trees are the Agathis and Shorea species, 'merbau' (/ntsia palembanica), and 'ramin' (Gonystylus bancanus). Borneo timber constitutes a multibillion-dollar market. Over the past two decades, more cubic metres of wood were exported from Borneo than from Africa and tropical America combined (Curran et a/., 2004). For Malaysian Borneo alone, the total value of wood (legally) exported in 2004 amounted to 2.5 billion US$ (MTC, 2006).

Wild Supermarket

Besides for international trade, many of Borneo's plants are vital for the daily life of local communities. Even city folk could not live without some of its forest produce, like 'durian' (Durio zibethinus) and 'tarap' (Artocarpus odoratissimus) fruits, the 'tuhau' ginger (Etlingera punicea; Phillipps & Liew, 2000), and the ornamental orchids, palms, and aroids that adorn many a city garden.

Communities in villages, especially those far away from major towns are even more dependent on what can be obtained from the forest, such as edible ferns and other vegetables (for example, the recently discovered Begonia lazat, which is eaten by Sabah's 'Orang Sungai' people [Azmi, 1999]), fruits like various local mango species,

12

Biodiscoveries - Borneo's Botanical Secret

Em A young 'belian' tree (Eusideroxy/on zwagen), which produces one of Borneo's most durable types of timber.

ED intsia paiembanica,

an important timber tree.

Em Phaiaenopsis amabilis, one of the most popular orchids in Borneo gardens.

Photo Credit:

20 Menno Schi~huizen 21 Menno Schilthuizen 22 Peter Koomen

rattans for turning into mats, baskets, fish traps, hats, and ropes (Pearce, 1989), timber for building houses, the versatile bamboo, the football-sized fruits of Pangium edule, which have antimicrobial properties and are used for preserving meats, the poisonous 'tuba' (Derris elliptica) for catching fish, and an entire pharmacopoeia of herbal cures and medicines.

The sheer diversity of plants used by rural communities highlights the degree of underutilisation of Borneo's flora. The legendary durian fruit Durio zibethinus, hugely popular throughout Southeast Asia (the trade is estimated to be worth 1 .5 billion US$ yearly; Ross, 1997), is originally a species from Borneo (Baja-Lapis et a/., 2004; Tate, 1999). Eighteen more species of durian are found on the island, each with a different taste, and many are enjoyed by local people, though none are commercially grown on any scale. The same applies to the 24 wild mango species, and the 25 wild breadfruit species (Payne et a/., 2000). Another possibility are the many Vanilla orchids that occur in Borneo, of which some might be as useful as a spice as the globally used South American V. planifolia. At Tenom Agricultural Park in Sabah, some experiments have been started to assess the potential of the five Vanilla species of Sabah (Lamb, personal

Em Selakau man using rattan to make a fish trap.

fII A Bomean Vanilla orchid, V. pi/it era (not flowering).

Photo Credit:

23 WWF-Malaysia I R.Rajanathan 24 Menno Schilthuizen

Em Wild durian fruits,

an underexploited Borneo forest product

Em A native durian,

Durio grandiflora, flowering

ED A red durian tree, with steps installed by local durian collectors

Photo Credit:

25 WWF-Malaysia I Junaidi Payne 26 Peter Koomen

27 WWF-Malaysia I Reza Azmi

Biodiscoveries - Borneo's Botanical Secret

13

Photo Credit:

28, 30 Menno Schilthuizen 29, 31 Peter Koomen

Em Em mil m:I Aroids, like Alocasia cuprea (28) and begonias from Borneo are popular as ornamental plants.

m Em The bark of the langsat (Lansium domesticum) contains a powerful drug against malaria.

Pharmaceuticals, a joint venture between the Sarawak government and the US-based company Advanced Life Sciences (Sarawak MediChem, 2006).

communication). Even more revenue may be obtained from commercialising the many ornamental plants, like orchids, hoyas, begonias, palms, gingers, rhododendrons, and aroids, which might join a huge international market.

Hidden Cures

But perhaps most promising is tapping into local people's traditional herbal medicine. Scientists have only just started exploring this vast, and rapidly vanishing knowledge base. Interviews with elders of local comunities such as the Dusun and the Orang Sungai of Sabah have yielded long lists of plants that are used for a wide variety of ailments (Ahmad & Ismail, 2000; Phillipps & Liew, 2000; Yusoff et a/., 2003). Pharmacological investigation is beginning to show that some of these plants can be sources for new, powerful drugs of global importance.

Already, a Sarawak shrub (the identity of which is kept secret) has yielded a promising substance (code-named CBL316) that kills human cancer cells (Rouhi, 2003). The chemical was found by an Australian company, Cerylid Biosciences, which claims it is effective against 57 of 60 kinds of human cancer cells kept in the laboratory, including those that cause brain and breast cancer and melanoma. It is now being tested on laboratory mice and according to the US National Cancer Institute, appears to be a completely novel kind of anticancer agent, at least as effective as a leading anticancer drug, paclitaxel (Cerylid Biosciences, 2006).

The latex of bintangor trees (Ca/ophyllum lanigerum and C. teysmannH, relatives of the mangosteen), also from Sarawak, was found to contain two compounds (calanolides A and B, respectively), effective against a wide range of strains of the Human Immunodeficiency Virus HIV. The chemicals appear to block the replication of the virus. Moreover, calanolide A does not have the nasty side-effects that many other anti-AIDS drugs have, and it also acts against the tuberculosis bacterium, which many AIDS patients also suffer from. Clinical trials and development of this possible medicine is done by the company Sarawak MediChem

Finally, the bark of the 'Iangsat' tree (Lansium domesticum), used by the Kenyah of Kalimantan to treat malaria, was indeed discovered to contain a powerful and previously unknown antimalarial substance (Omar et a/., 2003). Researchers found that the compound, a 'triterpenoid', killed the malaria parasite, Plasmodim falciparum, in the test tube. It also cured mice that were ill with the mouse version of the disease. More such drugs could be discovered in projects such as the one started by Universiti Malaysia Sarawak in collaboration with the Japanese Yamanouchi Pharmaceutical company. This project tries to do rapid chemical screenings of extracts from hundreds of Borneo plants used in traditional medicine, and come up with potential drugs (Ahmad & Ismail, 2000).

So, the forests of Borneo are an unexplored botanical treasure trove, which, if kept intact, could be sustainably used to generate potentially billions of dollars' worth of additional revenue for the island. But even more valuable than the possible commercialisation of individual species are the 'ecosystem services' that the vegetation can provide. Regenerating rainforest stores more than one kilogramme of carbon dioxide per square metre per year in its wood, thus mopping up the greenhouse gas from the atmosphere. Mature forest also stabilises the climate, controls soil erosion, maintains soil quality and fertility, and is essential for a steady supply of clean drinking water. These priceless services are literally a matter of life or death for Borneo's 20 million people.

14 Biodiscoveries - Borneo's Botanical Secret

Currently, logging itself is no longer the greatest threat to Borneo's flora. Instead, the trend to convert logged-over forest to plantations has a much more severe impact on biodiversity. Whereas even moderately damaged natural forest can still contain fairly complete representations of the natural flora and fauna, tree plantations (for timber, rubber, gaharu, or palm oil, for example) do not (Payne, 2001).

Oil palm plantations in particular are currently enjoying a boom due to an anticipated replacement of fossil fuels by vegetable oils. Throughout Borneo, they have grown by almost ten percent per year between 1998 and 2003, and now cover about three million hectares, an area almost the size of the Netherlands (Rautner & Alfred, 2005). This trend seems to be only increasing, now that European laws call for the introduction of 'bio-diesel', mixtures of petroleum and vegetable oil (Pearce, 2005).

Ell] m Going, going .•. In the last 20 years, Borneo has exported as much timber as Africa and tropical America together.

Photo Credit: 35 Annadel Caban ban. 36 Peter Koomen

Biodiscoveries - Borneo's Botanical Secret 17

In addition to wholesale disappearance of forest, even of supposedly 'protected' ones in Kalimantan (Curran et a/., 2004), individual plant species suffer as well. Borneo ironwood is now so rare in the wild that it has been placed on the IUCN Red List of Endangered Species (IUCN, 2004). Many rattans were overcollected in Sarawak and Kalimantan when demand rose in the late 1980s (Pearce, 1989) and almost half of them are now considered endangered. And in Maliau Basin, Sabah, 'gaharu' collectors often kill many of the Aquilaria ma/accensis trees (now also on the Red List) in their search for suitably fragrant wood (Ali, 2001).

Some species endemic to limestone hills seem to have disappeared altogether: the slipper orchid Paphiopedilum philippinense, which used to grow on the limestone outcrops of Balambangan island off Borneo's north coast, has been exterminated by commercial collectors (Kiew, 2001). And the fate of the many plant species that were only known from Kinabalu's Pinusok Plateau also looks bleak, now that the area has been converted for vegetable farming and a golf course (Wong & Phillipps, 1996).

18

Biodiscoveries - Borneo's Botanical Secret

HEART OF BORNEO

Provisional Scope and Main Protected Areas

1. Satang Ai National Park

2. Lanjak Entlmau Wildlife Sanctuary

3. Betung Kerfhun National Park

4. Apan Entulu (Proposed)

5. Ballah (Proposed)

6. Oanau Unau (Proposed)

7. Apad Runan (Proposed)

8. 'Pulong Tau National Park

9. Ufu BeJait Protection Forest Reserve

10. Gunung Mutu National Park

11. Gunung Buda National Park

12. Utu Tembur6ng National Park

13. Maligan Virgin JUIlgle'Raserve

14. Crocker Range NatiOnal Park

15. 1mbak Canyon Conservation Area

16. Klnabalu Park

17. Mallau Basin Protection Forest Reserve

lB. 19. 20. KaYi\n

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23. Danau $entarum

With SO many setbacks, Borneo's flora is likely to be suffering devastating blows. More than 5,000 species are endemic to the island, and many occur in very small pockets. Habitat destruction, overcollecting and sometimes more subtle threats, such as the disappearance of pollinators (Harrison, 2000) will drive many plant species to extinction. Researchers from the National University of Singapore have estimated that up to 44% of the plant species of Southeast Asia are going to go extinct within the 21st century (Sodhi et a/., 2004), and unless active steps are taken soon, Borneo's flora will not escape this onslaught.

The Heart of Borneo initiative involving Brunei Darussalam, Indonesia, and Malaysia, is a last -ditch attempt to conserve a large chunk of the island's beleaguered biodiversity. The 220,000 square kilometres of lowland and highland rainforest in the heartland of Borneo could

preserve much of its flora for posterity, and continue its function as a water catchment area and carbon sink. The area is large and pristine enough for the complex ecosystems to continue as before, providing opportunities for scientists to go on studying the myriad of subtle interactions between species. The Heart of Borneo will also serve as a refuge for the many threatened species of plants that are now on the brink of extinction.

With dozens of new plant species discovered in Borneo's forests each year, it is likely that the Heart of Borneo will continue to be an important source for new biodiscoveries for the next centuries. And naming new species will only be the beginning. If sustainably managed, the area could be a source for valuable plant species that can be cultivated and commercialised for new foods and medicines for Borneo, Southeast Asia, and the rest of the world. But if we lose the Heart of Borneo, it will take its secrets to its grave.

m rml Throughout Borneo, natural forest is giving way to oil palm plantations.

Photos Credit

37 Uew Thor Seng

38 Menno Schilthuizen

m Botanists studying plant samples in a previously unexplored area in Borneo.

Photo Credit:

39 Menno Schilthuizen

Biodiscoveries - Borneo's Botanical Secret

19

List of plant species,

newy dfscovered in Borneo between 1 980 and 2005

Begonia, 7 new species (Beaman et al., 2001 a) Begonia, 12 new species (Kiew, 2001 a) Begonia, 6 new species (Sands, 1990)

Bombacaceae--Durian family Coelostegia montana (Sidiyasa, 2001) Durio bukitrayaensis (Kostermans, 1990)

This list of 422 species was compiled from references in the Flora Malesiana Bulletin, the Thomson lSI Web of Knowledge, and random literature searches. While attempts have been made to aim for completeness, it is unavoidable that many species (in particular those hidden in large revisions, and those published in minor journals) remain unlisted. The literature reference usually refers to the original description, although in some cases it refers to a secondary source.

Adiantaceae--Maidenhair ferns

Adiantum, new species (Bidin et al., 1988)

Anacardiaceae--Cashew family Semecarpus calcicola (Kiew, 2001 b)

Annonaceae--Custard Apple family

Mitrephora c/emensiorum, M. vittata, M. woodii (Weerasooyia & Saunders, 2001)

Tetrapetalum 1 new species (Momose, 1998) Goniothalamus, 10 new species (Mat -Salleh, 1993) Goniothalamus 1 new species (Beaman et al., 2001 a) Polyalthia, 4 new species (Johnson & Murray, 1999) Monocapia bomeensis (Mols & Kessler, 2000)

Apocynaceae--Periwinkle family Secamone badia (Klackenberg, 2004) Wil/ughbeia lunduensis (Middleton, 1996)

Araceae--Aroids

Schismatoglottis bulbifera (Okada et al., 1999) Amorphophallus julaihii, A. angulatus, A. brachyphy/lus, A. ebumeus, A. hottae, A. infundibuliformis, A. pendulus, A. lambii, A. rugosus, A. tinekeae, A. venustus, A. costatus,

20

Biodiscoveries - Borneo's Botanical Secret

A. linguiformis

(Hetterscheid, 1994; P. Boyce, cited at: http://www.snowsexotics.com/firstdivexp.htm) Nephthythis,1 new species (Hay et al., 1994)

Aridarum, Bucephalandra, and Hottarum, 1 new species each (Okada & Mori, 2000)

Aiocasia reginula, A. pangerana, A. princeps, A. puteri (Kiew, 2001 b)

Colocasia oresbia and Hottarum kinabaluense (Beaman & Beaman, 1998)

Burmanniaceae

Thismia lauriana (Jarvie, 1996) Thismia goodii (Kiew, 1999)

Capparaceae--Caper family

Capparis buwaldae (Maschwitz et al., 1996)

Chrysobalanaceae

Kostermanthus robustus (Prance, 2001)

Araliaceae--Ivy family

Aralia, 1 new species (Wen, 1994)

Crypteroniaceae

Payena, 5 new species (Pereira, 1997)

Crypteronia, 3 new species (Pereira & Wong, 1995)

Araucariaceae--Monkey Puzzle family Agathis kinabaluensis and A. lenticula (Beaman & Beaman, 1998)

Dipteridaceae

Cheiropleuria parva (Kato et al., 2001)

Asclepiadaceae--Milkweed family Hoya telosmoides (Omlor, 1996) Hoya nabawanensis (Kloppenburg & Wiberg, 2002)

Hoya, at least 2 new species (Kloppenburg & Wiberg, 2002) Hoya callistophylla, H. clemensiorum, H. kloppenburgii, H. lambii, H. meredithii, H. spatiodes, H. waymaniae (Lamb, 2003)

Elaeocarpaceae--Jamaican Cherry family Elaeocarpus, 5 new species (Coode, 1998)

Ericaceae--Heather family

Rhododendron, 10 new species (Argent, 2003; Argent et al., 2006).

Euphorbiaceae--Spurge family

Macaranga, at least 2 new species (Davies, 1999)

Aytoniaceae

Asterella limbata (Long & Grolle, 1994)

Fagaceae--Beeches, Oaks, Chestnuts Lithocarpus palungensis (Cannon & Manos, 2000)

Begoniaceae--Begonias Begonia lazat (Azmi, 1999)

Begonia, 2 new species (Kiew, 1998a) Begonia keithii (Kiew, 1998b)

Gesneriaceae--African Violet family Agalmyla macrocalyx (Hilliard & Burtt, 1999)

Monophyllaea, 1 new species (Kiew, 2002) Epithema dolichopodum (Kiew, 2001 b)

Paraboea madaiensis, P. sabahensis (Kiew, 2001 b)

Hanguanaceae

Hanguana major (Beaman & Beaman, 1998)

Hookeriaceae

Chaetomitrium, 2 new species (Akiyama & Suleiman, 2001) Distichophy/lum scabrisetum (Akiyama & Yamaguchi, 1999)

Lauraceae--Laurel family

Beilschmiedia, 1 new species (Nishida, 2005) Endiandra, 3 new species (Arifiani, 2000)

Leguminosae--Legumes

Mil/ettia bomeensis (Adema, 2000)

Kunstleria sarawakensis (Ridder-Numan & Kornet, 1994) Bauhinia, 1 new species (Jarvie, 1998)

Bauhinia campanulata (Larsen, 1994)

Dalbergia, 4 new species (Sunarno & Ohashi, 1997)

Lejeuneaceae

Metalejeunea winkleri (Zhu & Grolle, 2002)

Loganiaceae--Buddleja and Strychnine family Fagraea, 21 new species (Wong & Sugau, 1996)

Loranthaceae--Mistletoe family

Macrosolen brevitubus and Trithecanthera sparsa (Barlow, 1995)

Lowiaceae

Orchidantha, 4 new species (Pedersen, 2001) Orchidantha inouei (Nagamasu & Sakai, 1999) Orchidantha holttumii (Larsen, 1993)

Marattiaceae--Giant Fern family Marattia pellucida (Maideen et al., 2002)

Biodiscoveries - Borneo's Botanical Secret

21

22

Urticaceae--Nettle family

Elatostema, 8 new species (Beaman & Cellinese, 2004). Poikilospermum, 1 new species (Church et a/., 1998)

Violaceae--Violets

Rinorea belalongii (Stevens, 2000)

Vitaceae

Tetrastigma steenisii and T. megacarpum (Latiff, 1991)

Zingiberaceae--Gingers

Zingiber, 6 new species (Theilade & Mood, 1999) Zingiber, 5 new species (Theilade & Mood, 1997a) Zingiber kelabitianum (Theilade & Christensen 1998) Zingiber, 2 new species (Theilade & Mood, 1997b) Zingiber pseudopungens (Beaman & Beaman, 1998) Etlingera kenya/ang (Poulsen & Christensen, 2004) Etlingera, 3 new species (Poulsen et a/. 1999)

Etlingera muluensis (Beaman & Beaman, 1998) Boesenbergia, 2 new species (Poulsen, 1993) Boesenberia armeniaca, B. hosensis (Cowley, 2000) Boesenbergia, 2 new species (Cowley, 1998) Boesenbergia grandis (Beaman & Beaman, 1998) Amomum roseisquamosum (Nagamasu & Sakai, 1996) Amomum botryoideum (Cowley, 2000)

Amomum, 2 new species (Smith, 1989a)

Amomum anomalum, A. longipedunculatum and A. flavoalbum (Beaman & Beaman, 1998)

Plagiostachys, 2 new species (Cowley, 1999) Plagiostachys bracteolata, P. glandulosa (Julius, 2006) Plagiostachys, 8 new species (Julius, 2006) Zingiberaceae, 4 new species (Smith, 1990) Zingiberaceae, 2 new species (Smith, 1987)

Alpinia beamani (Smith 1989b)

Hedychium muluense (Beaman & Beaman, 1998) Homstedtia incana (Beaman & Beaman, 1998)

Yet undescribed (not included in the total tally): 1 fern species (Parris in Kiew, 2001)

20 species of Hoya (Lamb, 2003)

1 species of Pinanga (Andersen, 2003) 3 species of Plagiostachys (Julius, 2006)

12 species of orchids (Lamb, personal communication)

5 species of Orchidantha (Lamb, personal communication)

Melastomataceae--Melastoma family Sonerila verticillata (McDonald, 1995) Sonerila, 1 new species (Cellinese, 1997)

Myrsinaceae--Myrsine family

Embelia, 1 new species, Maesa, 1 new species (Stone, 1989)

Musci (unknown family)--mosses

At least 2 new moss species (Tan et a/., 1997)

Myrtaceae--Myrtle family Syzygium velutinum (Davis, 1997)

Nepenthaceae--Pitcher Plants

Nepenthes vogelii (Schuiteman & de Vogel, 2002) Nepenthes faizaliana (Adam & Wilcock, 1991) Nepenthes, 1 new species (Turnbull & Middleton, 1988) Nepenthes platychila (Lee, 2002)

Nepenthes hurrelliana (Cheek et a/., 2003)

Orchidaceae--Orchids

Podochilus marsupia/is (Schuiteman, 1998)

Trichoglottis tinekeae (Schuiteman, 1998)

Trichoglottis, 1 new species (Wood et a/., 1998b) Crepidium mieczys/awii (Margonska & Szlachetko, 2001) Crepidium, 1 new species (Margonska, 2002) Bulbophyllum, 9 new species (Vermeulen & Lamb, 1994) Bulbophyllum gemma-reginae (Vermeulen, 1996) Bulbophyllum,41 new species (Vermeulen, pers. comm.; in addition to the 10 mentioned above)

Trias, 1 new species (Vermeulen & Lamb, 1994) Liparis anopheles (Wood, 1991)

Appendicula, 1 new species (Wood, 1997a) Dendrochilum, 1 new species (Wood, 1997b) Dendrochilum, 1 new species (Beaman et a/., 2001 b) Dendrochilum 5 new species (Wood, 2001)

Dendrobium lamrianum (Chan, 1994; cited in Chan 1999)

Biodiscoveries - Borneo's Botanical Secret

Dendrobium, 2 new species (yukawa, 1998) Coelogyne, 1 new species (Wood et a/., 1998a) Coelogyne marthae, C. verrucosa (Sierra et a/., 2000) Paphiopedilum platyphyllum (GruB, 2001) Paphiopedilum supardii (Braem & Loeb, 1985) Paphiopedilum kolopakingii (Fowl ie, 1984)

Ania ponggolensis (Kiew, 2001 b)

Various genera, 14 new species (Chan et a/., 1994)

Rafflesiaceae--Rafflesias

Rafflesia tengku-adlinii (Salleh & Latiff, 1989) Rafflesia keithii, R. pricei (Meijer, 1984)

Biodiscoveries - Borneo's Botanical Secret

23

Protaceae

Helicia and Heliciopsis, each at least one new species (Chung, 1998)

Rhizophoraceae--Mangroves Anisophyllea ismailii (McDonald, 1995)

Palmae--Palms, Rattans

Calamus fimbriatus, C. nigricans, Daemonorops pumilus (van Valkenburg, 1995)

Daemonorops banggiensis (Dransfield, cited in Kiew 2001) Daemonorops ingens (Beaman & Beaman, 1998)

28 palm species endemic to Sarawak described post -1980 by Dransfield and Kiew (Pearce, 1989)

Areca rheophytica, Calamus amplijugus, C. comptus, C. convallium, C. e/opurensis, C. mesi/auensis, C. praetermissus, Kortha/sia furtadoana, K. jala, P/ectocomiopsis mira, Sa/acca lophospatha (Beaman & Beaman, 1998)

Rubiaceae--Bestraw family

Schradera grandiflora and S. pseudonervulosa (Puff et a/., 1998a)

Lecananthus peduncularis (Puff et a/., 1998b) Aleisanthiopsis multiflora (T ange, 1996) Xanthophytum bullatum (Tange, 1995) Neomussaenda kostermansiana (Tange, 1994) Antirhea inaequalis (Kiew, 2001 b)

Sapindaceae--Soapberry family Zollingeria, 1 new species (Adema, 1992)

Pandanaceae--Screw Pines

Pandanus ca/cinatus and P. lepatophilus (Stone, 1982) Pandanus beccatus and P. leuconotus (Beaman & Beaman, 1998)

Schisandraceae

Kadsura acsmithii (Saunders, 1997)

Pittosporaceae--Pittosporum family Pittosporum, 2 new species (Sugau, 1994)

Scrophulariaceae--Figwort family Brookea, 1 new species (Yamazaki, 1994)

Poaceae--Grasses

Kinabaluchloa nebulosa, Racemobambos hepburnii, Sphaerobambos hirsuta (Beaman & Beaman, 1998)

Sematophyllaceae

Trismegistia ma/iauensis (Akiyama & Suleiman, 2003)

Staphyleaceae--Bladder-nut family Turpinia,1 new species (Pereira, 1994)

Podocarpaceae--Podocarp family Podocarpus laubenfelsii (Tiong, 1984)

Podocarpus confertus, P. globulus (Beaman & Beaman, 1998)

Dacrydium gracilis (Beaman & Beaman, 1998)

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