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The only species of Nervilia Comm. ex Gaudich. included in national checklists and redlists of the Singapore flora is Nervilia punctata (Blume) Makino. This species is treated as presumed nationally extinct. There are three historic collections from Singapore, all collected by H.N. Ridley during the late 19th and early 20th centuries. However, the identity of these specimens has recently been cast into doubt as the Nervilia adolphi/punctata species alliance has become the subject of taxonomic scrutiny. The lack of visible characters on the existing specimens has so far made it impossible to pinpoint the correct identity of the Singapore specimens. We recently discovered a small population of Nervilia in Bukit Timah Nature Reserve in Singapore, which in our opinion is the same taxon that was collected by Ridley. The plants do not agree with other species in the Nervilia adolphi/punctata species alliance and the taxon is here described as a new species based on the clinandrial tissue surrounding the anther cap as well as the narrow oblong and truncate labellum with curled sides. We currently consider the species to be endemic to Singapore.

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Dracaena breviflora Ridl. (Asparagaceae) is newly recorded here for Singapore. A description and colour plates of this unusual species are included. Provisional conservation assessments of Endangered globally and Critically Endangered in Singapore are proposed.

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Combretum perakense M.Gangop. & Chakrab., described from specimens collected in Perak, Peninsular Malaysia, is here reduced to a synonym of Lophopyxis maingay Hook.f. (Lophopyxidaceae). Lectotypes are designated for three other synonyms of Lophopyxis maingayi.

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The history of Haplochorema K.Schum. (Zingiberaceae) is reviewed, its morphology is compared to Boesenbergia Kuntze and the molecular phylogenetic position is shown in relation to other Zingiberaceae. Based on a comparative analysis of molecular and morphological data, Haplochorema is reduced to Boesenbergia with eight new combinations. A lectotype for Boesenbergia loerzingii (Valeton) K.Larsen ex M.F.Newman, Lhuillier & A.D.Poulsen is designated here.

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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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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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         Dr Alistair Hay Dr Mark Large
    (Supplement Editor) (Supplement Editor)

Dr David J. Middleton            Lily Chen      Felicia Tay
    (Editor-in-Chief)              (Managing Editor) (Graphics Editor)

  Ada Davis               Daniel C. Thomas                Christina Soh 
(Copy Editor)                (Copy Editor)               (Business Manager)

Preface

Introduction

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.

David Mabberley’s distinguished and productive academic and administrative positions, roles, honours, books and other publications are given.

Personal reflections are offered on the botanical life of David Mabberley during his time at Oxford University.

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.

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.

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.

Aglaia mabberleyi Pannell is described as new from Borneo. It is distinguished from Aglaia rufibarbis Ridl. under which name it has hitherto been treated.

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.

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.

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.

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.

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.

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. 

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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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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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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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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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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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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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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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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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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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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Book review

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Book review

     Dr Lena Chan Dr Geoffrey Davison
(Supplement Editor) (Supplement Editor)

Dr David J. Middleton              Lily Chen
    (Editor-in-Chief)                   (Managing Editor)

      Felicia Tay                Ada Davis                 Christina Soh 
(Graphics Editor)          (Copy Editor)          (Business Manager)

Content Page

Preface

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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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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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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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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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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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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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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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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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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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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.