Year of Publication: 2019, Vol. 71 (2)

Date Published 16 December 2019
ISSN2382-5812
D.J. Middleton, K. Armstrong, Y. Baba, H. Balslev, K. Chayamarit, R.C.K. Chung, B.J. Conn, E.S. Fernando, K. Fujikawa, R. Kiew, H.T. Luu, Mu Mu Aung, M.F. Newman, S. Tagane, N. Tanaka, D.C. Thomas, T.B. Tran, T.M.A. Utteridge, P.C. van Welzen, D. Widyatmoko, T. Yahara & K.M. Wong

Progress on Southeast Asia’s Flora projects [Page 267 - Page 319]
Abstract:
Southeast Asia (Myanmar, Thailand, Cambodia, Laos, Vietnam, Malaysia, Singapore, Brunei Darussalam, Indonesia, Timor Leste, the Philippines and Papua New Guinea) is a region of high plant diversity with an estimated 50,000 flowering plant species. Estimates of plant diversity in the region continue to grow as large numbers of new species are described even though there have been suggestions that there are few new species to be found in some parts of Southeast Asia. It is likely that most estimates of species numbers in the countries of Southeast Asia are too low due to the lack of taxonomic work on groups which have many locally endemic species. Differing collecting densities across the region can profoundly affect our understanding of plant diversity and lead to large underestimates of species diversity in poorly collected countries and regions. Progress on each of the major Flora projects in Southeast Asia, Flora of Thailand, the Flora of Cambodia, Laos and Vietnam/Flore du Cambodge, du Laos et du Viêtnam, Flora Malesiana, Flora of Peninsular Malaysia, Tree Flora of Sabah and Sarawak, Flora of Singapore and Flora of Vietnam, along with floristic research in Myanmar, the only country not covered by at least one of these Flora projects, is discussed. In addition to the formal Flora projects, there is much floristic activity occurring in the countries otherwise covered by transnational Floras. 
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Z. Dennehy & R. Cámara-Leret

Quantitative ethnobotany of palms (Arecaceae) in New Guinea [Page 321 - Page 364]
Abstract:
We conducted a bibliographic review of palm use in New Guinea to quantify palm-utilisation patterns across the region’s habitats, countries, and indigenous groups, and to identify the most useful species. We reviewed 187 bibliographic references and 140 herbarium specimens, spanning the years 1885–2018. We found 1178 use-reports and 894 palm-uses for 119 palm species. Lowland tropical rainforest is the best-studied habitat, and Indonesian New Guinea and Papua New Guinea have each received similar research effort. Most palms are used for Utensils and tools, Construction and Human food, and the stem, leaf and fruit are the most utilised palm parts. Only 5% of New Guinea’s indigenous groups have been studied, and <10 use-reports are recorded for most of the indigenous groups studied. Important species included Actinorhytis calapparia H.Wendl. & Drude, Adonidia maturbongsii W.J.Baker & Heatubun, Areca catechu L., Areca macrocalyx Zipp. ex Blume, and Metroxylon sagu Rottb. Overall, our study highlights the importance of palms for fulfilling subsistence needs in New Guinea, indicates that palm ethnobotany is neglected in the world’s most bioculturally diverse island, and gives directions for future research.

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T.F. Stuessy

Information content for biological classifications [Page 365 - Page 372]

Abstract:

Classification is a fundamental activity of the human species. The aim of all forms of classification is to establish a hierarchical structure of information that serves as a reference system to answer specific questions. In biological classification the objective is to store data in a conveniently retrievable fashion, to infer evolutionary relationships, and to predict undocumented characteristics of the included organisms. Different kinds of data have been used to form a basic data matrix from which to construct biological classifications. Dendrograms have been traditionally used to illustrate relationships among taxa, although such two-dimensional diagrams do not capture all relationships from the original data matrix. Controversies have existed on which algorithms are best suited to construct dendrograms. Explicit phyletic (evolutionary), phenetic, and cladistic schools of quantitative classification have each offered methods for doing do, and each has made claims for capturing maximum information. Decisions on which type of data and algorithms to use depend upon the nature of the systematic and evolutionary questions being posed. Important is the need for detailed evolutionary investigations so that inferred relationships can be properly evaluated. Information theory, a separate discipline, is viewed as having high potential to enrich information content of biological classifications.
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P. Bernhardt, G.R. Camilo & P.H. Weston

Shaken vs scraped: floral presentation contributes to pollinator guild segregation in co-blooming Symphionema montanum and sopogon anemonifolius (Proteaceae) [Page 377 - Page 396]

Abstract:

Sympatric populations of Symphionema montanum R.Br. and Isopogon anemonifolius (Salisb.) Knight showed overlapping flowering periods during November 2009 in the Blue Mountains (New South Wales, Australia). Symphionema montanum has porose anthers encircling the protruding style, and lacks a pollen presenter. In contrast, sessile, longitudinally dehiscent anthers of Isopogon anemonifolius deposit pollen grains on the subterminus of the style (pollen presenter). Neither species secretes nectar. The majority of foragers on Symphionema montanum were polylectic, female bees (Halictidae). Their pollen foraging resembled sonication and shaking. Polylectic, female bees (Colletidae) were dominant foragers on Isopogon anemonifolius grasping styles with their mandibles while scraping the pollen presenter. Exoneura species (Apidae) visited both shrubs. Only two specimens of Callomelitta antipodes on Isopogon anemonifolius carried pollen of both shrub species. Most bees, collected on either shrub, carried the grains of their host mixed with one to six pollen morphotypes of co-blooming, nectariferous taxa. We report a positive correlation between an increase in bee size and the number of morphotypes carried but colletids of Isopogon anemonifolius carried fewer morphotypes than halictids on Symphionema montanum.
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C.E. Jarvis

Georg Rumphius’ Herbarium Amboinense (1741–1750) as a source of information on Indonesian plants for Carl Linnaeus (1707–1778) – an Addendum [Page 397 - Page 399]

Abstract:

An Addendum, discussing Linnaeus’ generic names with particular reference to Rumphia, is provided to a recently published article.
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L.M.J. Chen, B.E.E. Duyfjes, Ali Ibrahim & W.J.J.O. de Wilde

Flora of Singapore precursors, 16: New records and notes on the plant diversity of Singapore [Page 401 - Page 406]

Abstract:

Due to ongoing work for the Flora of Singapore, a new family record for Singapore, the Stemonaceae, with one species, Stemona curtisii Hook.f., is recorded. In addition, Ammannia crassicaulis Guill. & Perr. in the Lythraceae is newly recorded as naturalising in Singapore. Notes on two rare species, Hernandia nymphaeifolia (C.Presl) Kubitzki in the Hernandiaceae and Securidaca philippinensis Chodat in the Polygalaceae, currently being revised for the Flora of Singapore are presented.
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W.W. Seah, S.M.X. Hung & K.Y. Chong

Flora of Singapore precursors, 17: Clarification of some names in the genus Calophyllum as known in Singapore [Page 407 - Page 411]

Abstract:

The species, Calophyllum soulattri, is found to have been wrongly included in Singapore’s native flora. The name Calophyllum wallichianum var. wallichianum is also found to have been misapplied to a taxon in Singapore and should rather be called Calophyllum rufigemmatum. The nomenclatural history and problems of both taxa are discussed in this paper.
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I.M. Ardaka & W.H. Ardi

A new species of Begonia (Begoniaceae) from the Moluccas, Indonesia [Page 415 - Page 419]

Abstract:

A new species of Begonia, Begonia mufidahkallae Ardaka & Ardi, is described from Sawai, Seram Utara District, Seram Island, Indonesia. The species is endemic to Seram Island and belongs to Begonia section Petermannia.

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M.B. Atmaja & I.G. Tirta

Notes on the orchids of Bali, Indonesia: six new species records [Page 421 - Page 427]

Abstract:

Six species of orchids are reported as new records for Bali. Two of the species were found to be growing wild in Bali Botanic Garden, which was formerly a part of the Batukahu Nature Reserve, while the rest were collected from other forests in Bali. The six newly recorded species are Bulbophyllum apodum Hook.f., Ceratostylis longipedunculata J.J.Sm., Dendrobium arcuatum J.J.Sm., Dendrobium connatum (Blume) Lindl., Habenaria reflexa Blume and Taeniophyllum hirtum Blume. Brief descriptions and photographs are provided.
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M.J.C. Arshed, E.M. Agoo & M. Rodda

The identity of Marsdenia parasita (Apocynaceae, Asclepiadoideae) [Page 429 - Page 433]

Abstract:

A neotype is designated for Marsdenia parasita Blanco and lectotypes are designated for its synonyms Dischidiopsis philippinensis Schltr. and Conchophyllum merrillii Schltr. ex Merr. The new combination, Dischidia parasita (Blanco) Arshed, Agoo & Rodda is proposed. We explain why a specimen collected by Llanos and identified by him as Marsdenia parasita is not original material and thus cannot serve as a lectotype. Moreover, this gathering belongs to a different species, Dischidia vidalii Becc. Dischidia bulacanensis Kloppenb. et al.
is here treated as a synonym of D. parasita.
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M.A.K. Naive, J.A.G.P. Dalisay, E.P.T. Maglangit, G.C. Café & O.M. Nuňeza

Free radical scavenging effects of the Philippine endemic medicinal plant Alpinia elegans (Zingiberaceae) [Page 435 - Page 444]

Abstract:

Alpinia elegans (C.Presl) K.Schum. is an endemic Philippine medicinal plant used in the treatment of various conditions such as muscoloskeletal diseases, hemoptysis, headache, migraine, stomach ache, and as an anti-relapse for women. The major phytochemical constituents of the ethanolic extract from the leaves of Alpinia elegans were screened and their antioxidant activity was evaluated using an in vitro 1,1-Diphenyl-2-picrylhydrazyl (DPPH) free radical-scavenging (antioxidant) activity assay. Flavonoids, steroids, saponins, tannins, alkaloids, and cyanogenic glycosides were found to be present in the ethanolic leaf extract of Alpinia elegans, while anthraquinone was not detected. High DPPH radical scavenging (antioxidant) activity was observed in the ethanolic leaf extract of Alpinia elegans with a percentage DPPH inhibition of 95.11±1.00 at 500 μg/ml. The present study suggests that the leaf extract is a source of medicinal or pharmaceutical antioxidants. Information derived herein provides a preliminary scientific basis for the existing ethnobotanical knowledge of local
communities.
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R.V.A. Docot, N.P. Mendez & C.B.M. Domingo

A new species of Hornstedtia and a new species record of Globba (Zingiberaceae) from Palawan, Philippines [Page 445 - Page 457]

Abstract:

During recent botanical exploration in the province of Palawan, Philippines specimens were collected of a new species, Hornstedtia crispata Docot, and a new species record for the Philippines, Globba francisci Ridl., both from the ginger family Zingiberaceae. The new species is described and illustrated here along with an assessment of its conservation status.
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R.V.A. Docot, K.D. Gutierrez, R.E.E. Mamalias, N.B.R. Espino, A.A.B. Java, C.D. Dineros & E.M.L. Mijares

Two new Zingiber species (Zingiberaceae) from Sorsogon, Philippines [Page 459 - Page 475]

Abstract:

Two new species of gingers (Zingiberaceae), Zingiber aguingayae Docot and Z. subroseum Docot, are described and illustrated here based on recent collections from Mount Bulusan, Sorsogon, Philippines. Our rediscovery of the poorly known Zingiber bulusanense Elmer at the same locality not only allowed us to clarify its identity but also gave us stronger assurance that the two new species are indeed undescribed. Evidence from morphological and molecular data using the ITS region supported the placement of both new species and Zingiber bulusanense within Zingiber sect. Zingiber. The conservation status of the two new species were also assessed.
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J.D. Mood, †J.F. Veldkamp, T. Mandáková, L.M. Prince & H.J. de Boer

Three new species of Boesenbergia (Zingiberaceae) from Thailand and Lao P.D.R. [Page 477 - Page 498]

Abstract:

Boesenbergia bella Mood & L.M.Prince, B. phengklaii Mood & Suksathan, and B. putiana Mood & L.M.Prince are described with photographs and a comparative table. The description of Boesenbergia petiolata Sirirugsa is revised to include morphology not previously noted. Molecular phylogenetic analyses of the relevant taxa using plastid and nuclear DNA sequence data are provided.
 
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O. Theanphong, T. Jenjittikul & W. Mingvanish

Essential oils composition of nine Curcuma species from Thailand: a chemotaxonomic approach [Page 499 - Page 518]

Abstract:

The chemical composition of the essential oils from fresh rhizomes of nine Curcuma L. species was investigated using the GC-MS technique. A total of 136 compounds,representing 97.19–99.11% of the total content of the essential oils, were identified. A dendrogram obtained from the cluster analysis based on their chemical composition was divided into two main clusters. The first cluster, with a high content of sesquiterpene hydrocarbon (e.g. β-curcumene) and oxygenated sesquiterpene (e.g. xanthorrhizol) was composed of Curcuma alismatifolia Gagnep., C. larsenii Maknoi & Jenjitt., C. sparganiifolia Gagnep. and C. harmandii Gagnep. The second cluster was subdivided into two groups, IIA and IIB. Group IIA with a high content of monoterpene hydrocarbons (e.g. camphene), sesquiterpene hydrocarbons (e.g. a-copaene), caryophyllene, and oxygenated sesquiterpenes (e.g. caryophyllene oxide), comprised Curcuma parviflora Wall. and C. rhabdota Sirirugsa &
M.F.Newman. The other, IIB, with a high content of oxygenated monoterpenes (e.g. camphor) and oxygenated sesquiterpenes (e.g. germacrone), included Curcuma rubrobracteata Škorničk. et al., C. angustifolia Roxb. and C. singularis Gagnep.
 
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M. Murugesan, A.A. Mao, L.R. Meitei & S.S. Kambale

Ceropegia khasiana (Apocynaceae: Ceropegieae), a new species from Meghalaya, Northeast India [Page 519 - Page 525]

Abstract:

A new species of Ceropegia, Ceropegia khasiana Murug., A.A.Mao, Meitei & Kambale (Apocynaceae), is described and illustrated from Meghalaya, Northeast India. The new species is superficially similar to Ceropegia macrantha Wight but it differs in having fewer, shorter fascicled roots up to 8 cm long, linear-lanceolate leaves with long acuminate apices, smaller flowers up to 4.7 cm long, 12–18-flowered inflorescences with two flowers open at a time, two umbels per node, corolla tube pinkish outside with dark reddish stripes, reddish inside in mature flowers, greenish or pinkish to reddish at the apex of corolla lobes which are densely ciliate hairy, inflated base with reddish patch at middle, outer corona with very sparse small ciliate hairs, and each pair of follicular mericarps unequal in length.
 
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Year of Publication: 2019, Vol. 71 (Supplement 2)

Date Published 27 September 2019
ISSN2382-5812
B. G. Briggs & K. L. Wilson

David Mabberley and Australian botany [Page 7 - Page 24]
David Mabberley has worked on five continents but chose Australia as his home, moving there in 1996. By then, he already had an outstanding international reputation and his contributions to Australian botany and Australian botanical history had started with his biographies of botanist Robert Brown and botanical artist Ferdinand Bauer. Joseph Banks, Brown and Bauer have remained continuing interests for him with further publications and lectures. In Australia he has contributed to the treatments of Meliaceae and Rutaceae in the Flora of Australia, drawn attention to the work of John Bidwill and other botanical figures, established important collaborations on the phylogeny and diseases of Citrus, investigated Red Cedar (Toona ciliata), given master classes in economic botany, and much more. Moving to Australia did not deflect David from his global reach in tropical botany, the world’s flora in The Plant-book, and economically important plants. He has contributed greatly to Australian botany, but his career of outstanding achievement continues to be global, not limited to a single continent.
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M.F. Large

Mabberley’s scholarship [Page 25 - Page 41]
David Mabberley’s distinguished and productive academic and administrative positions, roles, honours, books and other publications are given.
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A.M. Sing

Recollections of the ‘MabLab’, Department of Plant Sciences, University of Oxford, 1977–1996 [Page 43 - Page 46]
Personal reflections are offered on the botanical life of David Mabberley during his time at Oxford University.
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N.P. Taylor

E.J.H. Corner — Mabberley’s mentor — and his contributions to the Singapore Botanic Gardens’ heritage [Page 47 - Page 52]
The life and work of E.J.H. Corner during his time at Singapore Botanic Gardens, and how this contributed to the Gardens’ eventual inscription as a UNESCO World Heritage Site, is discussed.
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I. Friis

G.C. Oeder’s conflict with Linnaeus and the implementation of taxonomic and nomenclatural ideas in the monumental Flora Danica project (1761–1883) [Page 53 - Page 56]
Hitherto unpublished parts of the history of the IconesFlorae Danicae (1761–1883), one of the largest illustrated botanical works published, are analysed; it covered the entire flora of the double monarchy of Denmark–Norway, Schleswig and Holstein and the North Atlantic dependencies. A study of the little noticed taxonomic and nomenclatural principles behind the Icones is presented. G.C. Oeder, founder of the project, approved the ideas of Buffon and Haller and rejected Linnaean binary nomenclature because of its lack of stability of genera. In the Icones …, Oeder cited all names used for each plant in chronological order, with the binary Linnaean name last, to which principle Linnaeus reacted. By the end of the 18th century, Linnaean nomenclature had become standard, apart from in Flora Danica and a very few other botanical works. Applying Linnaean nomenclature elsewhere, O.F. Müller, editor 1775–1782, and M. Vahl, editor 1787–1799, followed Oeder’s norm in the Icones. J.W. Hornemann, editor 1810–1840, followed Oeder in his first fascicles, but began experimenting with changes towards Linnaean nomenclature from 1810. After 1840, subsequent editors consistently applied Linnaean principles for accepted names and synonyms.
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C.E. Jarvis

Georg Rumphius’ Herbarium Amboinense (1741–1750) as a source of information on Indonesian plants for Carl Linnaeus (1707–1778) [Page 87 - Page 107]
The Herbarium Amboinense (1741–1750) and the supplementary Herbarii Amboinensis Auctuarium (1755) of Georg Eberhard Rumpf (Rumphius) (1627–1702) provided detailed descriptions and illustrations of the plants of the island of Ambon, then a Dutch colony in the Maluku Islands (Moluccas) of Indonesia. The initial work, published in six volumes, contained a great deal of new botanical, medicinal and ethnographical information from a part of the world then little-known. Published in the Netherlands long after Rumphius’ death by the Director of the Amsterdam Botanic Garden, Johannes Burman, the work appeared prior to Carl Linnaeus’ Species Plantarum (1753) in which the consistent use of binomial names was introduced. However, in that work, Linnaeus referred to only a handful of Rumphius’ species accounts. More detailed studies by Linnaeus of Rumphius’ work soon followed, notably in the dissertation Herbarium Amboinense (May 1754; November 1759). In all, only about 100 of the nearly 700 taxa illustrated by Rumphius were referred to by Linnaeus in his various publications, though many of those that were cited serve as nomenclatural types for their corresponding Linnaean binomials. The reasons for Linnaeus’ apparent neglect of such an important source of novel information are explored.
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H.W. Lack

Flora Graeca on the European continent [Page 109 - Page 122]
Flora Graeca is a work in ten folio volumes, published in London between 1806 and 1840. It is based on the botanical collections made by John Sibthorp during his travels in what is now Cyprus, Greece, Italy and Turkey and the pictorial documentation of the plants produced by Ferdinand Bauer. The text was written by James Edward Smith, Robert Brown and John Lindley. Flora Graeca was printed in a tiny edition with the result that many botanists considered it to be non-existent or as only a manuscript. This paper deals with the few copies of Flora Graeca kept today in libraries on the European continent and their historical backgrounds – three in Paris, one in Vienna and Darmstadt respectively, all apparently subscription copies, while the copies in St. Petersburg and Copenhagen were previously owned by Fredrick North, 5th Earl of Guildford, and John Platt. The botanical affiliations of the previous owners, among them Franz I, Emperor of Austria, and Ludwig I, Grand Duke of Hesse, are explained. The libraries on the European continent that have copies of the so-called Bohn reissue of Flora Graeca are listed. Particular attention is given to the Belgrade copy, a hybrid consisting of printed title matter with some of the landscapes copied by hand and the 966 engravings, whereas all text pages, indices and appendices have been written on a type writer.
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C. Riedl-Dorn & M. Riedl

Ferdinand Bauer or Johann and Joseph Knapp? A rectification [Page 123 - Page 142]
The first part of the following paper deals with the life, travels, professional career as a natural history painter, use of colour charts and heritage of Ferdinand Bauer. The use of Ferdinand Bauer’s colour chart(s) by other painters is also discussed. It is followed by a short history of the use of colour charts. The final part discusses the attribution of watercolours preserved at the Vienna Natural History Museum that have hitherto been ascribed to Ferdinand Bauer. They are compared to pictures at the Austrian National Library, at the Museum of Applied Arts in Vienna and at the Museum für Kunst und Gewerbe in Hamburg that were produced by Joseph Knapp and should be attributed to him using evidence from the use of specific paper and some technical details.
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P.J. de Lange, M.F. Large, L. Shepherd, J.R. Rolfe & R.O. Gardner

The endemic that never was — resolving the status of Coprosma solandri (Rubiaceae) [Page 143 - Page 153]
Coprosma solandri, an enigmatic species known only from the type collection, was described in 1897 by New Zealand botanist Thomas Kirk, based on material included in a duplicate set of Banks & Solander specimens, sent at Kirk’s request, by the Natural History Museum (BM) in London to the Colonial Museum in Wellington in 1895. Here we revisit Coprosma solandri concluding that the specimens on which that name was based were not collected in New Zealand. We conclude the type of Coprosma solandri was derived from Hawaiian material of C. ernodeoides A.Gray that was accidentally sent to New Zealand on the assumption that Banks & Solander had collected it from there. Although available evidence precludes definite linkage to a specific collector, we raise the possibility that the original material derives from an Archibald Menzies collection made in 1793 in Hawai’i. The recognition that Coprosma solandri and C. ernodeoides are conspecific finally removes a puzzling Coprosma from the New Zealand flora 122 years after it was described and accepted uncritically as an endemic to that country.
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M. Cheek & D. Bridson

Three new threatened Keetia species (Rubiaceae—Vanguerieae), from the forests of the Eastern Arc Mountains, Tanzania [Page 155 - Page 168]
Keetia sp. A, of the Flora of Tropical East Africa, based on Mabberley & Salehe 1496, is formally named as Keetia davidii following the discovery of a second specimen, also from the Ukaguru Mountains of the Eastern Arc of Tanzania. This small tree or shrub is assessed as Endangered using the IUCN 2012 standard. Keetia sp. B, a shrub from Kwiro Forest near Mahenge, is described as Keetia wasumbii. Based on Cribb et al. 11027, it is assessed as Critically Endangered. Keetia sp. C, formally named as Keetia semsei based on Semsei 2025, from the Shikurufumi Forest near the Uluguru Mountains, is also assessed as Critically Endangered. The three species each derive from separate threatened patches of forest that are among the least well-surveyed and least protected in the Eastern Arc Mountain archipelago. For this reason, naming them, so enabling inclusion in the IUCN Red List, is important if they are to be included in conservation prioritisation exercises.
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†D.G. Frodin

Should a flora account be taken for granted? A fresh look at Polyscias serratifolia (Araliaceae) [Page 171 - Page 188]
Polyscias serratifolia (Miq.) Lowry & G.M.Plunkett (Gastonia serratifolia (Miq.) Philipson) currently encompasses a range of small to large trees endemic to Malesia save for two localities in the Solomon Islands. Latterly it has been referred to Polyscias subgen. Tetraplasandra but, lacking genomic analyses, only by assumed association with Polyscias spectabilis (Harms) Lowry & G.M.Plunkett. Nothing should, however, be taken for granted: collections accounted for since 1979, notably from ‘Wallacean’ Malesia, suggest that Polyscias serratifolia s.l. is a cluster of species, some previously described. From a total of 100 records and utilising both specimens and digital images I examined several likely diagnostic morphological characters, some not before used, and show that these taxa represent varying, but distinguishable, combinations of their states. Hopefully this will lead to additional field studies and collections including material for genomic analyses. Many of these taxa have not been recollected for decades or even a century or more. As a first step towards a revision, evaluations of the existing published taxa are presented, and for all eight new combinations are made or revived (Tetraplasandra koordersii Harms, Gastonia papuana Miq., Tetraplasandra paucidens Miq., T. philippinensis Merr., and T. solomonensis Philipson along with the imperfectly known Gastonia eupteronoides Teijsm. & Binn., Polyscias serratifolia proper, and Gastonia winkleri Harms).
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C.M. Pannell

Aglaia mabberleyi Pannell (Meliaceae), a new species from Borneo [Page 189 - Page 195]
Aglaia mabberleyi Pannell is described as new from Borneo. It is distinguished from Aglaia rufibarbis Ridl. under which name it has hitherto been treated.
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Z. Ezedin and G.D. Weiblen

Additions and changes to Ficus (Moraceae) in New Guinea with comments on the world’s largest fig [Page 197 - Page 216]
Two new species of Ficus in New Guinea are described and a third species is resurrected to improve the classification of Ficus sect. Dammaropsis, sect. Sycidium, and sect. Papuasyce. Ficus sect. Dammaropsis is extraordinary in producing by far the world’s largest known figs. New morphological, ecological, and molecular observations support recognising as separate species the lowland and highland forms of Ficus dammaropsis (Warb.) Diels sensu lato. The lowland form, described here as Ficus brusii Weiblen, is distinguished from the highland form by its much smaller figs that bear recurved, acuminate lateral bracts. The description of Ficus dammaropsis is amended to refer to the highland form with the world’s largest fig and includes F. dammaropsis var. obtusa Corner. Evidence from DNA suggests pollination by distinct fig wasp species and reproductive isolation between Ficus brusii and F. dammaropsis. Ficus hystricicarpa Warb. (sect. Sycidium) is resurrected from synonymy under Ficus wassa Roxb. on account of its conspicuously warty and hispidulous syconia. Lastly, an ecological genetic study led to the discovery of a third species occupying an altitudinal contact zone between Ficus itoana Diels of hill forest and Ficus microdictya Diels of montane forest. These members of the New Guinea endemic Ficus sect. Papuasyce point to a rare evolutionary shift in the sexual system from functional dioecy to monoecy. Here we describe Ficus umbrae Ezedin & Weiblen on the basis of morphological intermediacy and molecular evidence to report the first functionally gynomonoecious Ficus species.
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D.C. Thomas & W.H. Ardi

Begonia mabberleyana (Begoniaceae), a new species from Central Sulawesi, Indonesia [Page 217 - Page 223]
Based on collections from the Banggai Regency, Sulawesi, Indonesia, the new species Begonia mabberleyana D.C.Thomas & Ardi is described and illustrated. This species is restricted to limestone habitats and endemic to Central Sulawesi. A provisional conservation assessment indicates a Critically Endangered (CR) status for the species.
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R.P.J. de Kok

Cinnamomum mabberleyi, a new species from Vietnam and Laos [Page 225 - Page 229]
A new species of Cinnamomum (Lauraceae), C. mabberleyi, is named from Vietnam and Laos. A formal description, notes on distribution, conservation status and ecology, and a map are given; it is differentiated from Cinnamomum tsoi C.K.Allen.
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P. Baas, B.-J. van Heuven, X.Y. Ng & N. Vander Velde

Biomechanical and hydraulic challenges for a tropical swamp forest and driftwood tree – Alstonia spatulata Blume (Apocynaceae) [Page 231 - Page 244]
Rootwood and basal stemwood of Alstonia spatulata is polystyrene-like in texture and softness when dry. It has been traditionally used for pith helmets, rafters, and as cork substitute, and can be dispersed over long distances as driftwood. This driftwood is so common on the beaches across the Central Pacific that in the Marshall Islands a special traditional use of the wood for floaters or cork substitutes named wũj has emerged. Here we describe this ultralight driftwood and the rootwood and basal stemwood of Alstonia spatulata, a tree from swamp forests of Southeast Asia. The  round-tissue is composed of very thinwalled modified fibres without tip growth and vestured pits without borders. Axial parenchyma is in narrow marginal bands, and scanty paratracheal. Vessels are narrow. Rays are extremely low and mostly uniseriate. We discuss the biomechanical and hydraulic conductivity paradox of small to medium-sized trees resting on an extremely weak and soft trunk base (at the root collar), and the parallel evolution of similar very soft woods in swamp forests of both the Old and the New World and in the Deccan fossil record of India.
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D.A. Baum

Plant parts: processes, structures, or functions? [Page 245 - Page 256]
It is usually taken for granted that plants are composed of a series of discrete parts that can readily be compared both within a single plant and between closely or even distantly related organisms. The biological meaning of ‘part’ needs to be made more rigorous such that the naming of a part or the homologising of parts among organisms constitutes a scientific hypothesis that is testable, at least in principle. I explore three alternative approaches to defining parts. The parts-as-structures approach holds that parts (or “phenes”) are aspects of an organism that would not have been formed during development in the absence of causal genetic factors. A structural phene hypothesis is refuted by the lack of a hypothetical genetic deletion that would ablate just this part of the organism. The parts-as-functions approach focuses on pieces of the organism that could have been functionally different if one or more gene in the genome were mutated. A functional phene hypothesis is refuted by showing that all mutations affecting the phene have pleiotropic consequences. The part-as-process approach equates parts with developmental modules, making it especially helpful for addressing the
concept of serial homology. I conclude that, depending on the context, parts are best understood sometimes as structures, sometimes as functions, and sometimes as processes, but in each case we need to develop rigorous concepts rather than falling back on human perception as the ultimate arbiter of part-ness.
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A. Hay

Durianology, discovery, and saltation — the evolution of aroids [Page 257 - Page 313]
It is argued that E.J.H. Corner’s ‘durianology’ is an integrative, holistic approach to the evolution of angiosperm form which complements reductive, atomistic phylogenetic methods involving the reification of individuated high-level abstractions in the concept of morphological ‘character evolution’. A case is made that the Durian Theory involved in part the advanced, holistic cognitive mode of insight, and, drawing on recent findings from cognitive science, it is proposed that insight problem-solving may overcome some of the limitations and distortions of dis-integrative character analysis, and lead to discovery of novel morphological relations and global pattern recognition. Evidence drawn from molecular phylogenetic analyses, developmental studies, and from gross morphology is presented that supports an insight-based hypothesis of direct, saltatory derivation of the Araceae from an ancestor with shoot apices not enclosed by sheathing leaf bases, acropetally developed, reticulate-veined compound leaves, and a terminal polymerous strobiloid flower. It is proposed that this saltation led to an array of morphologically hybrid and compound decanalised structures blurring conventional morphological categories such as rhachis, rhachilla, petiolule and venation; leaf base and stipule; leaf and leaflet; leaf and perianth; flower and inflorescence; flower and floral organ; fruit and infructescence; and fruit and seed. The associated perturbation of developmental routines led both to great diversification and to widespread parallel simplification series. It is argued that holistic evolutionary hypotheses cannot usefully be tested using current atomistic phylogenetic methodology applied to  orphological characters. It is suggested that holarchical 258 Gard. Bull. Singapore 71 (Suppl. 2) 2019 (nested-hierarchical) rather than matrix character sets may provide a more holistic framework for evolutionary hypothesis-testing involving the interplay between molecular phylogeny, evodevo data and hypotheses, and the quantitative and/or probabilistic analysis of contextualised character distribution.
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W.D. Hawthorne & C.A.M. Marshall

Rapid Botanic Survey, Bioquality and improving botanical inventory in the tropics by integrating across spatial scales [Page 315 - Page 333]
We review the Rapid Botanic Survey method (RBS), in the context of botanical recording to date. The concept of bioquality, a biodiversity value respecting global rarity, is summarised. Bioquality assessment involves the Star system for categorising species by global rarity; and a Genetic Heat Index (GHI) which aggregates Stars into community scores. All vascular plant species in tropical Africa have Stars, and >3.1 million botanical records have been databased across the continent (Marshall et al., 2016). Presented here are updated bioquality scores from continental tropical Africa, and especially coastal East Africa, and calculated for sample units of various shapes and sizes: East African Flora regions, one degree squares, forest reserves, to fine scale (sample-level) hotspots along the East African coast. GHI is globally standardised and has been calculated for survey data outside Africa, though seldom in tropical Asia. RBS data can also be used to distinguish vegetation types and can include ethnobotanical data. It is recommended as a way to standardise biodiversity or environmental Impact assessment nationally and globally, and for integrating such survey results in databases that will be increasingly useful as the tension between conservation and deforestation increases, and the climate changes.
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M. Heads

Biogeography and ecology in a pantropical family, the Meliaceae [Page 335 - Page 461]
This paper reviews the biogeography and ecology of the family Meliaceae and maps many of the clades. Recently published molecular phylogenies are used as a framework to interpret distributional and ecological data. The sections on distribution concentrate on allopatry, on areas of overlap among clades, and on centres of diversity. The sections on ecology focus on populations of the family that are not in typical, dry-ground, lowland rain forest, for example, in and around mangrove forest, in peat swamp and other kinds of freshwater swamp forest, on limestone, and in open vegetation such as savanna woodland. Information on the altitudinal range of the genera is presented, and brief notes on architecture are also given. The paper considers the relationship between the distribution and ecology of the taxa, and the interpretation of the fossil record of the family, along with its significance for biogeographic studies. Finally, the paper discusses whether the evolution of Meliaceae can be attributed to ‘radiations’ from restricted centres of origin into new morphological, geographical and ecological space, or whether it is better explained by phases of vicariance in widespread ancestors, alternating with phases of range expansion.
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K.M. Wong & L. Neo

Species richness, lineages, geography, and the forest matrix: Borneo’s ‘Middle Sarawak’ phenomenon [Page 463 - Page 496]
Contemporary studies into the spectrum of plant life assembled on the island of Borneo continue to demonstrate an astonishing richness for some groups. Not all lineages are equivalent in their richness, and both biogeographic and ecological factors are the principal correlates of species richness and lineage diversification. The ways in which population genetic factors may influence the generation and persistence of variation, and their interaction with environmental change, could have fundamental importance in how diversity is maintained. Central Sarawak in the northwest Borneo hotspot is a premier ecological theatre where the interplay of such factors operates: its plant species richness is astounding, floristic documentation continues perhaps too slowly, and research and conservation priorities continue to loom large. Unfortunately, this resource has been severely modified in the several decades spanning the turn of the 21st century. The importance of increasing public perception, especially with well-illustrated accounts of this biological richness and its significance through a natural history perspective, will be as critical as the slowly advancing frontiers of the scientific platform on which our understanding depends. 
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P.C. Boyce & S.Y. Wong

Borneo and its disproportionately large rheophytic aroid flora [Page 497 - Page 524]
On the basis of decades of field observations and multidisciplinary research, in particular directed at species of Araceae exhibiting rheophytism, we offer an overview of morphological diversity among the more than 130 aroid species so adapted on Borneo. Based on a combination of morphological and concomitant ecological occurrence, a preliminary scheme of subcategories of van Steenis’ “rheophytic landplants” is outlined with the purpose of encouraging study to better understand the impetus of obligate rheophytic aroids’ evolution. It is hoped that the proposed subcategories will encourage targeted research with abundant field-based observations. Criticism is directed at the current demand by scientific journals that computer algorithm-generated statistical ‘proof’ be provided for all observational life science writing and further insisting that all such observations be linked to a ‘big picture’ by comparison within the context of a global perspective, preferably incorporating whatever themes are currently fashionable in the field. Such requirements discourage researchers, and particularly students, from undertaking purely observational research, and effectively result in the suppression of publication of vital observational data while encouraging publication of statistically well-supported biological nonsense.
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R. Kiew & L.G. Saw

Corner’s Riau Pocket and other hytogeographical provinces in Peninsular Malaysia [Page 525 - Page 538]
Four phytogeographical provinces have been recognised in Peninsular Malaysia — the Northern Province, the Perak Province, the Continental Intrusion and the Riau Pocket. The Riau Pocket, originally spelt Riouw, was restricted to SE Johor, Singapore, Banka, Riau Islands, SE Sumatra and NW Borneo but later was expanded to cover the entire east coast of Peninsular Malaysia as far north as Kelantan on the premise that this area harboured the Borneo element of the flora. However, based on a combination of a characteristic assemblage of species, a significant number of endemic species, and the absence of species from adjacent areas, the Riau Pocket is reinstated in its original sense. The boundary between the Riau Pocket and the east coast flora appears to be the Sungai Anak Endau. The Northern Province is a distinct province that lies in the northwest of Peninsular Malaysia that experiences a monsoon climate. It harbours a characteristic assemblage of species and has close affinity with the flora of southern Thailand. Based on the distribution of a sample of 969 taxa, the continental element is shown to be a significant part of the Peninsular Malaysian flora (about a fifth of species) but it is not restricted to a particular area. The Continental Intrusion originally defined by Corner is therefore not supported. The Perak Province is also not recognised because the 969-taxon sample demonstrates that it does not harbour a characteristic assemblage of species nor does it have distinct boundaries. In addition, although the 969-taxon sample shows that for the flora in general there are more Sumatra species than Borneo species in Peninsular Malaysia, the Sumatra species are not better represented in Perak, nor are Borneo species better represented on the east coast.
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Year of Publication: 2019, Vol. 71 (1)

Date Published 28 June 2019
ISSN2382-5812
I.M. Turner, D.J. Middleton, H. Duistermaat & J.F. Veldkamp

Flora of Singapore precursors, 6: Typification of grass names relevant to the flora of Singapore [Page 1 - Page 45]
Abstract:

The typification of more than 200 names of grass taxa relevant to the flora of Singapore is reviewed. A total of 52 lectotype designations are made here, including 22 which are posthumously published by the late J.F. Veldkamp. In addition, 13 second-step lectotypes (one by JFV), and two neotypes are designated. An earlier place of publication than generally cited for the combination Zizania latifolia is highlighted.


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I.M. Turner

Flora of Singapore precursors, 12. Notes on various Rubiaceae genera [Page 45 - Page 59]
Abstract:
In preparation for the account of the Rubiaceae for the Flora of Singapore various notes are presented concerning the genera Discospermum Dalzell, Lasianthus Jack, Mussaenda L. and Psychotria L. Nomenclatural comments on the generic synonyms Landia Comm. ex Juss. (= Mussaenda) and Uragoga Baill. (= Psychotria) are made. Lectotypes are designated at the first or second step for 42 names, and one neotype is designated. Recent collections of Lasianthus griffithii Wight and Psychotria morindiflora Wall. ex Hook.f. from Singapore are highlighted.

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P. Hovenkamp

Flora of Singapore precursors, 13. New names and lectotypifications in Athyriaceae and Polypodiaceae [Page 61 - Page 67]
Abstract:
A review of names and types in Diplazium (Athyriaceae) and Polypodiaceae relevant to the Flora of Singapore has been undertaken. A new name and 11 lectotype designations (including one second step lectotypification), are given. 
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M. Rodda & D.J. Middleton

Flora of Singapore precursors, 14. Notes on Apocynaceae [Page 69 - Page 80]
Abstract:
Lectotypes are designated for 28 names of Apocynaceae, seven of which are second-step lectotypifications. Two names are neotypified, Leptostemma hirsutum Blume is designated as the type of the genus Leptostemma Blume, and a new combination in Secamone R.Br. is coined for Genianthus maingayi Hook.f.
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J. Sangrattanaprasert, B.C. Ho, S. Chantanaorrapint & R.-L. Zhu

The genus Colura (Lejeuneaceae, Marchantiophyta), new to Singapore [Page 81 - Page 85]
Abstract:
Colura brevistyla Herzog is reported here as a new genus and species record for Singapore. A complete description and illustrations of C. brevistyla are provided based on the newly identified specimen from Singapore.
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G.T. Prance

The correct name for Atuna excelsa (Chrysobalanaceae) [Page 87 - Page 88]

Abstract:
A correction is made for the use of the name Atuna excelsa (Jack) Kosterm. which has been erroneously placed as a subspecies of A. racemosa Raf. despite being described sixteen years earlier. Atuna excelsa is restored as the correct name for this species and A. racemosa is reduced to subspecific rank. Since the holotype of Atuna racemosa is an old illustration, an epitype is proposed for this name.

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R.P.J. de Kok

A revision of Cinnamomum Schaeff. (Lauraceae) for Peninsular Malaysia and Singapore [Page 89 - Page 139]

Abstract:
A revision of all species of the genus Cinnamomum Schaeff. (Lauraceae) occurring in Peninsular Malaysia and Singapore is presented with a summary of taxonomic history, notable features ofmorphology, a key to species, and a description, distribution map, Provisional IUCN Conservation Assessment for each species, together with notes on ecology and ethnobotany. In this revision, twenty-two distinct species are recognised, including one former variety that is elevated to species level as Cinnamomum selangorense (Ridl.) de Kok. Eleven names are lectotypified (Camphora inuncta Nees; Cinnamomum cinereum Gamble, C. graciliflorum Gamble, C. malaccense Meisn., C. ridleyi Gamble, C. rhynchophyllum Miq., C. sintoc Blume, C. subcuneatum Miq., C. velutinum Ridl., and C. vimineum Wall. ex Nees; Laurus malabathrum Wall. ex Nees), and five names are placed into synonymy for the first time. A small number of species are considered to have a Provisional IUCN Conservation Assessment of Least Concern, while most are either Endangered or Critically Endangered, with one species considered to be Data Deficient (Cinnamomum trintaense Kosterm.).

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R.P.J. de Kok

Three new taxa, two new combinations and thirty-one lectotypifications in several Lauraceae genera from Peninsular Malaysia
[Page 141 - Page 161]
Abstract:
Three new taxa of Lauraceae are described, two new species: Lindera kochummenii de Kok and Machilus kochummenii de Kok, and a new subspecies, Actinodaphne sesquipedalis Hook.f. & Thoms. ex Meisn. subsp. glabra Kochummen ex de Kok. In addition, two new combinations are made: Machilus declinata (Blume) de Kok and Phoebe scortechinii (Gamble) Kochummen ex de Kok, and thirty-one names are lectotypified, including one second-step lectotypification.
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W.W. Seah

A new combination in Gynochthodes (Rubiaceae) [Page 163 - Page 165]
Abstract:
Morinda scortechinii (King & Gamble) Ridl. was recently placed in the synonymy of Gynochthodes umbellata (L.) Razafim. & B.Bremer. However, the two taxa were found to differ in a number of leaf and flowering head characters and hence should be considered distinct species. A new combination Gynochthodes scortechinii (King & Gamble) W.W.Seah is necessary owing to recent changes in the circumscription of several genera in the Morindeae tribe.
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W.A. Mustaqim & I.P. Astuti

New and noteworthy orchid records from Buru Island, Maluku Archipelago [Page 167 - Page 174]
Abstract:
A short note on the orchids of Buru Island, Maluku Archipelago, is presented. Bulbophyllum cruciatum J.J.Sm. and Dendrobium bicaudatum Reinw. ex Lindl. are new records for Buru. Trichotosia buruensis (J.J.Sm.) S.Thomas et al. has been rediscovered after a lapse of 84 years. The distribution of a species with a previously uncertain distribution in Maluku, Dilochia wallichii Lindl., is now confirmed for Buru. We also discovered a population of the endemic Vanda saxatilis J.J.Sm. All species are briefly discussed and illustrated by photographs.
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F. Wen

Petrocosmea weiyigangii (Gesneriaceae), a new species from a limestone cave in southern China [Page 175 - Page 183]
Abstract:
A new species, Petrocosmea weiyigangii F.Wen (Gesneriaceae), is described from southern China. It differs from all other species of Petrocosmea in its deeply lobed leaf blades. It is only known from a limestone cave in northwestern Guangxi, China. Following the IUCN Red List categories and criteria, Petrocosmea weiyigangii is assessed as Critically Endangered.
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A. Kartonegoro, P. Hovenkamp, P.C. van Welzen

A taxonomic revision of Macrolenes (Melastomataceae) [Page 185 - Page 241]
Abstract:
Macrolenes (Melastomataceae: Dissochaeteae), a genus of woody climbers in Malesia, is taxonomically revised. Seventeen species are recognised, of which three are new to science. The genus is characterised by its scrambling habit, a pair of hair cushion domatia on the base of the leaves, axillary inflorescences, and fimbriate connective appendages on the alternipetalous stamens. An identification key, nomenclature, descriptions, typification, geographic distributions and taxonomic notes are provided. The affinities with Dissochaeta, also woody climbers, are discussed.
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Y.M. Chan and L.S.L. Chua

Flowering phenology and seed production of three threatened tropical palms, Johannesteijsmannia spp. (Arecaceae)
[Page 243 - Page 260]
Abstract:
The reproductive behaviour and fruit production of three endangered species of Johannesteijsmannia H.E.Moore were studied for two years. Flowering occurred annually in Johannesteijsmannia magnifica J.Dransf. and sub-annually in J. lanceolata J.Dransf. and J. perakensis J.Dransf. The flowering of all species peaked from March to May and occurred during the wet seasons. High seed loss of 84–98% was recorded among the species. On average, Johannesteijsmannia perakensis produced the highest number of mature fruits per year (229) and J. lanceolata the lowest (69). This preliminary study provides vital information on species fecundity and demonstrates that each species, although within the same genus, has different flowering and fruiting behaviour.
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Year of Publication 2019, Vol. 71 (Supplement 1)

Date Published 29 May 2019
ISSN2382-5812
L. Chan & G.W.H. Davison

Introduction to the Comprehensive Biodiversity Survey of Bukit Timah Nature Reserve, Singapore, 2014–2018 [Page 3 - Page 17]

Abstract:
We introduce the Comprehensive Biodiversity Survey of Bukit Timah Nature Reserve, Singapore, for which planning began in 2014, field work was conducted largely in 2015–2016, and results analysed and written up in 2017–2018. The comprehensive survey is placed in the context of previous surveys in 1991–1992 and in 1992–1997. Bukit Timah is historically important and continues to be nationally and internationally significant for nature conservation.

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G.W.H. Davison & P.T. Chew

Historical review of Bukit Timah Nature Reserve, Singapore [Page 19 - Page 40]

Abstract:
Biological interest in Bukit Timah, Singapore, long pre-dated its declaration as a forest reserve (1887) and nature reserve (1951). The administrative and land use changes affecting this fragment of coastal hill dipterocarp forest are described. Boundary changes have determined the areas now supporting primary, old secondary or maturing secondary forest, and account for many of the features described in an accompanying set of papers on the current biodiversity of Bukit Timah Nature Reserve (BTNR). Research interest in the nature reserve continues to be high, with roughly 25 new research projects initiated each year.

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B.C. Ho, H.K. Lua, Bazilah Ibrahim, R.S.W. Yeo, P. Athen, P.K.F. Leong, Ali Ibrahim, S.L. Koh, Hassan Ibrahim, S. Lindsay, L.L. Chin, W.W. Seah & D.J. Middleton

The plant diversity in Bukit Timah Nature Reserve, Singapore [Page 41 - Page 134]
Abstract:
The plant diversity of Bukit Timah Nature Reserve (BTNR) is relatively well studied due to concerted effort over several decades, particularly as part of the worldwide system of ecological plots set up by the Center for Tropical Forest Science (CTFS), now called the Forest Global Earth Observatory. Publications arising from previous works have set baseline data for the species diversity, suggested that the forest resilience was greater than would be expected in such a small forest fragment, but that there was low recruitment of primary forest tree species into the secondary forest. In order to assess the overall vascular plant diversity, and to compare the diversity of the various forest types within BTNR to each other, 52 plots were set up, each 20 × 5 m, along nine different transects that covered the full range of topography and forest types, primary, old secondary and maturing secondary forests, within the reserve. The vascular plant diversity within each plot was recorded. In total, 1250 species in 148 families were recorded, including an additional 167 species newly listed for BTNR. The primary forest had the highest number of species not found in the other forest types. It nevertheless had a very large overlap with species in the old secondary forest but not with the maturing secondary forest.
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S.M.L. Lee

Macro-fungal diversity of Bukit Timah Nature Reserve, Singapore [Page 135 - Page 144]
Abstract:
The Bukit Timah Nature Reserve is an extremely important type locality for many species of macrofungi, especially for those collected by Edred John Henry Corner between 1929 and 1945 and later described by him. There has been little work done on the fungi of Singapore in recent years and much remains to be done. Current efforts to curate the fungal collection at Singapore Botanic Gardens are discussed, a new collecting programme is highlighted, and the very long way to go before we have a decent understanding of macrofungal diversity of Bukit Timah and the rest of Singapore is emphasised.
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R.C.H. Teo & N.J. Thomas

Updated Inventory of Amphibians, Reptiles and Mammals of the Bukit Timah Nature Reserve, Singapore [Page 145 - Page 183]
Abstract:
A two-year survey at Bukit Timah Nature Reserve, Singapore, found a high diversity of amphibian, reptile and mammal species. A total of 81 indigenous species was recorded – 18 amphibians, 40 reptiles and 23 mammals. The updated inventory comprises 115 species – 21 amphibians, 60 reptiles and 34 mammals ever historically recorded within the reserve.
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K.S. Lim

Birds of Bukit Timah Nature Reserve, Singapore [Page 185 - Page 208]
Abstract:
A survey of birds in Bukit Timah Nature Reserve (BTNR), Singapore, in 2015– 2016 yielded a cumulative total of 1663 individual sightings, amounting to 68 species. The total ever recorded for BTNR is now 146 bird species. The 68 species recorded in the present survey included four globally threatened, six globally near-threatened species, 14 nationally threatened and six nationally near-threatened species. Few of the species are entirely and strictly forest dependent, though many make use of both primary and disturbed forest environments.
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J.K.H. Koh & D.J. Court

Spider diversity in the Bukit Timah Nature Reserve, Singapore [Page 209 - Page 243]
Abstract:
This paper discusses the preliminary results of the first comprehensive survey of the spiders of the Bukit Timah Nature Reserve (BTNR) in Singapore. Two plots were established in each of the three zones of vegetation, viz., primary forest, old secondary forest, and maturing secondary forest. They were repeatedly sampled over an 18-month period. Sorting of the collection so far suggests that the three vegetation zones harbour rather different spider assemblages. Only ~9% of the total spider fauna recovered was shared by all three zones. The results have also yielded a preliminary picture of dominance, abundance and rarity. Although first intended to obtain a baseline for future quantitative analyses, the survey became a testing ground to modify and refine methodology so as to conduct future quantitative surveys with greater scientific rigour. Taxonomic work on the samples so far shows that the spiders in the BTNR span over 43 families, of which six families are listed for the first time in Singapore. The tally is summarised in an interim checklist of BTNR spiders. The checklist, with a total of 317 entries, shows that there are 158 described species of spiders in BTNR, of which 25 species are new records for Singapore. Another 159 morphospecies are provisionally recognised as distinct species, some of which may be new to science. Our observations during the survey have allowed us to provide a narrative of BTNR spider diversity against a backdrop of their microhabitat specialisation.
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J. S. Ascher, Z. W. W. Soh, B. M. Ho, R. Y. Y. Lee, A. Q. E. Leon, S. X. Chui1, J. J. L. Lai, J. X. Q. Lee, M.S. Foo, E. J. Y. Soh

Bees of the Bukit Timah Nature Reserve and vicinity [Page 245 - Page 271]
Abstract:
As a unique coastal hill dipterocarp forest remnant in Singapore, Bukit Timah Nature Reserve is a key refuge for flowering plants, but little information has been available about its bee pollinators and their floral associations. Historical and recent surveys of bees (Hymenoptera: Apoidea: Anthophila) at Bukit Timah and vicinity were compiled, yielding a total known fauna of four families, 23 genera, and 75 species (including unnamed morphospecies). Of these, 55 bee species, several known only from historical collections, have been recorded from the Bukit Timah Nature Reserve (BTNR) itself, which is dominated by mature, shady forest with few apparent flowers along the trails. More bee species (61) have been recorded from nearby Dairy Farm Nature Park (DF), which has more open, sunnier secondary forest with more conspicuous floral resources. Sampling methods included net collecting and honey baiting along transects, malaise trapping within the forest, and observations at flowers. Accounts are provided for species of particular taxonomic or conservation interest, and two new provisional synonymies are indicated. Floral associations are summarised for 32 floral hosts from BTNR (only 10 plant species) and from DF (28 plant species, including 5 shared with BTNR). For all species known from BTNR and vicinity, earliest and most recent dates of capture for both this area and for Singapore as a whole are provided. Four eusocial stingless bee species formerly collected in Singapore but not recently recorded are considered to be nationally extinct. An additional few poorly known solitary bee species may also be nationally extinct. By contrast, solitary bee species new to Singapore continue to be discovered at BTNR, notably Megachile resin and leafcutter bees attracted to reintroduced Tiger Orchids in 2014 during a mass bloom. Despite high species richness of native angiosperms persisting at BTNR, especially in its core, few bee species and individuals were found in recent bee surveys, likely reflecting limited availability of floral resources in the shady forest understorey. However, additional bee species are likely to be found in BTNR if further sampling is done during infrequent mass bloom events and traps are deployed at canopy level.
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S.K. Khew & H. Tan

Butterflies of the Bukit Timah Nature Reserve, Singapore, and its vicinity [Page 273 - Page 292]
Abstract:
A survey of butterflies has been undertaken within the Bukit Timah Nature Reserve (BTNR), Singapore, and in the adjacent well-vegetated areas that form a buffer to BTNR, namely Hindhede Park, Singapore Quarry and the Dairy Farm Nature Park. Sampling was mainly though observations and photographic records along the transects in BTNR, and through baiting (mainly at Dairy Farm). More butterfly species (85) were recorded from the buffer areas, which have more open, sunnier secondary forest with more conspicuous food sources for adult butterflies, than there were from BTNR (63). Despite the high plant species richness at BTNR, the butterfly diversity observed within the forest transects was rather low. The greater butterfly diversity at Dairy Farm Nature Park is attributable in part to higher abundance of nectar sources (flowering trees and bushes, including many non-native plants growing along the forest edge and along open trails and footpaths). However, there are likely to be more species within BTNR, such as in the tree canopy, that were beyond the sampling scope of this survey.
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Y. Cai, Y.P.Q. Nga & R.W.J. Ngiam

Diversity and Distribution of Dragonflies in Bukit Timah Nature Reserve, Singapore [Page 293 - Page 316]
Abstract:
Biodiversity baselines were established for dragonflies (Insecta: Odonata) of Bukit Timah Nature Reserve, Singapore, based on quantitative and qualitative samplings. Surveys were conducted from December 2014 to February 2016. Hydrological, physicochemical parameters and habitats were analysed to understand the main drivers structuring the dragonfly community. A total of 829 odonate specimens were recorded during the quantitative sampling, comprising 36 species of 28 genera in 11 families. The species diversity in each of the six sampling sites was compared using the Shannon-Wiener Index (H’). Hierarchical clustering and Detrended Correspondence Analysis (DCA) indicated that three main groupings of sites existed, each with a distinct community of associated species. Further analysis by Non-metric multidimensional scaling (NMDS) on the relevant distance based on species composition at the six sampling sites, together with 21 environmental variables showed that these groups were significantly associated with respective environmental variables. An updated species list of Bukit Timah dragonflies is provided for future reference, with 55 species of 43 genera in 12 families. Disturbance and threats to the odonate fauna of the Bukit Timah Nature Reserve are identified and conservation management measures are discussed. 
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R. Karam & J.H. Chong

Moths of Bukit Timah Nature Reserve, Singapore [Page 317 - Page 330]
Abstract:
The moth fauna of Bukit Timah Nature Reserve, Singapore, was studied using light trapping. Specimens and photographs were sorted into morphospecies including macro and micro moths. A total of 399 species has been found, of which nearly 200 have been identified to species level. Several are notably rare or otherwise of interest. The figures do not reach an asymptote, suggesting that the total moth fauna may be considerably greater. The nature reserve may be too small to yield statistically significant differences in the moth fauna between forest vegetation zones.
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M.K. Tan

Orthoptera species checklist of Bukit Timah Nature Reserve in the Zoological Reference Collection, Singapore [Page 331 - Page 338]
Abstract:
While work on Orthoptera in Singapore is not lacking, there is no species checklist for the species found within the Bukit Timah Nature Reserve (BTNR). Here, a checklist of orthopterans in the reserve is given based on specimens deposited in the Zoological Reference Collection of the Lee Kong Chian Natural History Museum, Singapore. In total, 83 species, 30 from the suborder Caelifera (grasshoppers and relatives) and 53 from the suborder Ensifera (crickets, katydids, and relatives), are recorded from the reserve.
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L.F. Cheong

Estimating saproxylic beetle (Coleoptera: Cerambycidae and Buprestidae) diversity in Bukit Timah Nature Reserve, Singapore, with a methodological and biological review [Page 339 - Page 368]
Abstract:
Approximately one third of all forest insect species worldwide depend directly or indirectly on dying or dead wood (i.e., they are saproxylic). They are a highly threatened ecological group but the status of many species remains undocumented. There is an urgent need to develop a better appreciation for the diversity and ecology of saproxylic insects so as to inform management strategies for conserving these organisms in tropical forests. Two of the historically better studied beetle groups, Cerambycidae and Buprestidae, are highlighted with a brief discussion of the methods for studying them and their ecology, and a systematic attempt  to survey these two beetle groups in the Bukit Timah Nature Reserve, Singapore, is described. From a comparison with the historical data, it is inferred that the decline of the saproxylic insect fauna must be happening at a rate that would certainly be considered alarming if only it were more widely noticed. Finally, the implications for overall conservation of the insect fauna and of the reserve are considered.
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P.Grootaert & M.S. Foo

The springtail catchers of the genus Neurigona (Insecta, Diptera, Dolichopodidae) in the primary forest of Bukit Timah Nature Reserve, Singapore [Page 369 - Page 379]
Abstract:
Three species of Neurigona Rondani, 1856, are recorded from the primary  of Bukit Timah Nature Reserve in the centre of the city of Singapore. Neurigona squamifera Parent, 1935, originally described from Peninsular Malaysia, is a species common in Bukit Timah. Neurigona temasek sp. nov., the most common species, and Neurigona timahensis sp. nov., a very rare species, are described as new for science. Gross morphology images are provided as well as illustrations of the male terminalia. A key is given to the five species hitherto known from Singapore.
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F. Seow-Choen I. Seow-En, E.K. Chua & M.E. Choo

A Survey of Stick-Insects in Bukit Timah Nature Reserve, Singapore [Page 381 - Page 389]
Abstract:
A survey of stick insects in Bukit Timah Nature Reserve was conducted as part of a broader biodiversity survey which covered the period from 1 April 2014 to 30 April 2018. Bushes and trees by the sides of the trails in Bukit Timah Nature Reserve were searched with hand-torches after dusk. Of the 16 species known from historical records to occur in Bukit Timah Nature Reserve, 11 species were found during the current survey. Five species that were not encountered are known to be rare. However, this does not exclude the possibility of their continued occurrence in the nature reserve. Rare phasmids are rare due mainly to scarcity of food plants and their habitat must be maintained if a wide array of biodiversity is to be conserved.
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J. K. I. Ho, M.S. Foo, D. Yeo, R. Meier

The other 99%: exploring the arthropod species diversity of Bukit Timah Nature Reserve, Singapore [Page 391 - Page 417]
Abstract:
Bukit Timah Nature Reserve (BTNR) is one of Singapore’s most important conservation areas because it is likely to be the last refuge for many species that belong to Singapore’s original forest biodiversity. We report here the results obtained from a first broad-scale survey of arthropods in BTNR. The focus was on insects because Singapore’s insect fauna remains largely unknown despite the fact that insects constitute much of the animal biomass and perform many ecologically important tasks. The survey relied on specimens collected with passive traps (e.g., Malaise traps) that were set along several transects in primary and different types of secondary forests. Specimens representing several thousand species were obtained. In order to process the specimens rapidly, we sorted them based on DNA sequences of the COI gene. Sequences for more than 9,000 specimens were obtained and the DNA data were used to group the specimens into putative species. Here, we compare the species numbers, composition, and species overlap between secondary and primary forests for “true bugs” (Hemiptera). Overall, the sequences belonged to more than 1850 insect species of which ca. 450 belonged to Hemiptera. A very large proportion of the “true bug” species are only represented by 1 or 2 specimens each and we find that BTNR’s species diversity is much higher than the diversity in mangroves and on the National University of Singapore (NUS) campus. We also report and illustrate some notable insect species found during the survey. They range from ship-timber beetles to beetle-flies mimicking leaf beetles and mantis-flies resembling praying mantises.
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K. Chatterjea

Bukit Timah Nature Reserve: a forest in transition [Page 419 - Page 440]
Abstract:
Bukit Timah Nature Reserve (BTNR), being virtually the only primary rain forest in Singapore, is of very high environmental importance. The forest is located amidst a highly developed residential area and is visited by large numbers of people who seek respite from urban stresses and are invigorated by the natural environment offered by the forest interiors. Trails within the forest are heavily frequented and are, therefore, under constant stress. Results of heavy usage are seen in extreme surface compaction and slope deterioration, which in turn affect the forest bio-physical environment. This paper covers research spanning over 13 years from 2004 (‘before forest closure’) to 2017 (‘after forest reopening’), tracks the degeneration of the forest under such severe patronage by visitors, and reports on changes in the Nature Reserve’s physical conditions during and after a period of closure for renovation of public facilities. Several factors of importance to the forest bio-physical environment, such as soil compaction, soil bulk density, soil surface organic matter status, and infiltration rates of surface soils were monitored during these 13 years and visible and measurable longitudinal changes are recorded. Significant improvements are seen in soil compaction values, organic matter content, and bulk density of the trail surface soils subsequent to the forest closure, indicating that the closure and the resultant natural regeneration of the forest has significantly improved the forest interior bio-physical environment. The success of this management strategy can be taken as evidence of good management procedure and can, therefore, set an example to follow in future to sustain the bio-physical status of Bukit Timah Nature Reserve. 
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Y. Cai

Hydrogeomorphic characteristics of streams in Bukit Timah Nature Reserve, Singapore [Page 441 - Page 490]
Abstract:
Field data and information from the literature on hydrology, hydraulic and geomorphic characteristics of the ten streams in Bukit Timah Nature Reserve were collected to form a baseline dataset with the aim of evaluating various functions that these streams perform, as well as the stresses and disturbances that they experience. Preliminary results show that majority of the channel reaches studied are functioning well, except that moderate to intense erosion was observed at the middle reaches of Lasia and Dairy Farm, and the lower reaches of Fern Valley and Wallace streams, where instability, undercutting and failure of banks have caused significant channelisation (bank widening and channel down-cutting). In general, the streams in Bukit Timah have little sediment, with low and small-sized bars visible only at the lower reaches of Fern Valley, Lasia and Wallace streams, which should not significantly affect the stream function if culverts and lined drains downstream are well maintained. However, several forced step-pools (artificial ponds) near the outlets of Taban, Seraya and Catchment streams and the middle of Jungle Fall stream continuously receive upstream sediments loads and accumulation of sediment with distinct siltation have obviously degraded the instream habitats. Follow-up actions on stream rehabilitation, adaptive management, future research and future monitoring are discussed. 
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C.T.T. Nguyen & Y. Cai

Physicochemical characteristics of streams in Bukit Timah Nature Reserve, Singapore [Page 491 - Page 556]
Abstract:
Spatial and temporal surveys were conducted to better understand the physicochemical characteristics of the streams in Bukit Timah Nature Reserve (BTNR). A total of 87 survey stations were selected along ten streams for an in-situ water quality study conducted in June 2018. Temporal investigation, including in-situ and ex-situ samplings, was conducted from late November 2017 to July 2018 for Fern Valley and Jungle Fall streams. The in-situ physicochemical parameters included pH, Dissolved Oxygen (DO), Conductivity, Oxidation-Reduction Potential (ORP), Total Dissolved Solids (TDS), Turbidity, Salinity, and Temperature. Water samples were collected for further analysis of Total Suspended Solids (TSS), anions, cations, elements, Total Organic Carbon (TOC), Total Nitrogen (TN) and Total Phosphorus (TP). Water discharge was calculated as stream crosssectional area multiplied by water velocity. Groundwater samples (down to maximum 2 m) were collected at upstream and downstream locations in Fern Valley and Jungle Fall catchments and analysed for water chemistry. Seventy-five soil samples (surface and subsurface) were collected to investigate the hydrogeomorphic conditions of the catchments in an attempt to understand the influence of hillslopes on water quality within the stream channel. Physicochemical baselines of the streams in Bukit Timah Hill were established, the data suggesting that stream temperature, TDS, salinity, the amount of TOC, TP, anions and cations vary within their expected natural ranges. Some parameters including DO and conductivity are slightly lower than expected, which may not be favorable for large aquatic animals. Some issues needing further investigations include low stream pH in Jungle Fall and Seraya streams as well as the significantly high concentration of Cd, Sb and Se in all streams.Follow-up actions are recommended to further investigate the drivers and monitor the effect of stream acidification, the causes and effects of high concentration of Cd, Sb and Se, as well as possible stream rehabilitation measures leading to improvement of water quality.
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T. Li, Y. X. Loh, W. Lim, M. Nyanasengeran, B. W. Low, H. H. Tan, D. C. J. Yeo & Y. Cai

The fish fauna of Bukit Timah Nature Reserve, Singapore [Page 557 - Page 573]
Abstract:
Bukit Timah Nature Reserve (BTNR) holds the largest remnant patch of primary rainforest in Singapore and its hill streams supports many native and threatened freshwater fauna. As the last comprehensive survey of freshwater fishes in BTNR was published nearly two decades ago, a series of surveys was carried out at 12 streams permeating BTNR and the Singapore Quarry to update the status of the fishes. Of the 27 species documented, 16 are native to Singapore and 11 are introduced. Two native species known from other parts of the Central Catchment Nature Reserve are recorded for the first time in BTNR. Species distribution, richness and abundance have been qualitatively assessed in relation to habitat characteristics. Potential threats and mitigation measures are discussed, which are important in the formulation of conservation and management strategies to safeguard the current diversity of freshwater fishes, many of which are threatened in rapidly developing Singapore. 
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M. D. Y. Khoo, N. J. L. Tiong, T. Li1, W. Lim, D. J. J. Ng, M. Nyanasengeran, D. C. J. Yeo & Y. Cai

The freshwater decapod crustaceans of Bukit Timah Nature Reserve, Singapore [Page 575 - Page 581]

Abstract:
Freshwater decapod crustaceans serve important ecological functions in tropical stream ecosystems. Stream surveys for decapod crustacean fauna were conducted between February 2015 and April 2016 at nine sites in Bukit Timah Nature Reserve (BTNR). A total of eight species (four freshwater crab, three freshwater shrimp and one crayfish species) out of the nine species known from BTNR were recorded. In general, the species were recorded from sites matching environmental conditions previously known or reported for each species, with some crab species showing minor overlap in their spatial distributions. This highlights the importance of broad-based conservation efforts focusing on the entire stream network in BTNR rather than on specific/selected streams as the various species each appear to have largely unique habitat requirements. The updated information on freshwater decapod crustaceans in BTNR can be used to inform the formulation of conservation and management measures including the establishment of monitoring programs for the protection of our native and threatened aquatic fauna in BTNR.

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L. Chan & G.W.H. Davison

Synthesis of results from the Comprehensive Biodiversity Survey of Bukit Timah Nature Reserve, Singapore, with recommendations for management [Page 583 - Page 610]

Abstract:
Twenty-two papers arising from the Comprehensive Biodiversity Survey of Bukit Timah Nature Reserve (BTNR), Singapore (2014–2018) have provided a thorough update and expansion of information on many groups of flora and fauna. In spite of the small size and isolation of BTNR there is still remarkable diversity, and new records and new species continue to accumulate. A large body of information on the terrestrial and aquatic physical environment is now available, providing a context for many of the biological results. Various management recommendations emerge.

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