THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore is a peer-reviewed journal publishing original papers and

reviews on a wide range of subjects: plant taxonomy (including revisions), phytogeography,

floristics, morphology, anatomy, as well as horticulture and related fields, such as ecology and

conservation, with emphasis on the plant fife of the Southeast Asian-Pacific region.

Dr David J. Middleton Dr Jana Leong-Skornickova

(Editor-in-Chief) (Managing Editor)

Felicia Tay Dr Daniel C. Thomas Christina Soh

(Graphics Editor) (Copy Editor) (Business Manager)

Editorial Advisory Board

Professor Sir Peter Crane

Yale University

U.S.A.

Dr Rogier P.J. de Kok

Royal Botanic Gardens, Kew

U.K.

Dr W. John Kress

National Museum of Natural History

Smithsonian Institution

U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh

U.K.

Dr Kiat W. Tan

Gardens By The Bay

Singapore

Dr Nigel P. Taylor

Singapore Botanic Gardens

National Parks Board, Singapore

Dr Ian M. Turner

Royal Botanic Gardens Kew

U.K.

Dr Jan-Frits Veldkamp

National Biodiversity Center

The Netherlands

Dr Jun Wen

National Museum of Natural History

Smithsonian Institution

U.S.A.

Professor Nianhe Xia

South China Institute of Botany

P.R. China

Published twice yearly by the National Parks Board, Singapore, the annual subscription for the journal

is Singapore $100.00 including postage. Overseas subscribers should make payment in the form of

bank draft or international money order in Singapore currency, payable to National Parks Board. Please

forward payment to “Accounts Receivable Section, National Parks Board, Headquarters, Singapore

Botanic Gardens, 1 Cluny Road, Singapore 259569”.

Instructions for contributing authors

(more details in http://www.sbg.org.sg/research/instructions-for-authors.pdf)

Manuscripts should be sent by e-mail to <gardbullsing@gmail.com> or posted to

The Editor, Gardens’ Bulletin Singapore, Singapore Botanic Gardens, 1 Cluny Road,

Singapore 259569.

The language used is English. Prior to submission, it may be advisable to have manuscripts checked by

someone fluent or proficient in that language. Contributions must not have been published or be under

consideration elsewhere. There are no page charges. Submitted material cannot be returned.

Authors will receive a pdf reproduction of their contribution, gratis.

Cover letter, submission and copyright. In your cover letter or message, state clearly you are submitting

your manuscript (state title) for publication in the Gardens’ Bulletin Singapore. By submitting the

manuscript(s), authors agree that upon acceptance, copyright of the entire work becomes the property

of the Singapore Botanic Gardens, National Parks Board, Singapore. Authors must ensure all required

permission has been obtained in writing to publish any material they present; proof may be required.

Manuscripts are best submitted in electronic form (Microsoft Word 97-2003 preferred). Use Times New

Roman, 10-point font, 1.5-line spacing. Do not send original artwork unless requested by the Editor.

Number all pages serially including the title and abstract on the first.

(continued on inside back cover)

The Gardens’ Bulletin

Singapore

VOL. 69(1) 2017 ISSN 0374-7859

CONTENTS

A.R. Rafidah

Taxonomy and conservation status of Microchirita

(Gesneriaceae) in Peninsular Malaysia.1

M. Rodda

Index of names and types of Hoy a

(Apocynaceae: Asclepiadoideae) of Borneo.33

Y.W. Low, A.K. Muhammad Ariffin, A.A. Joffre & D. Duratul Ain

Novitates Bruneienses, 8. Macrosolen brunsing (Loranthaceae),

a new hemiparasitic shrub from Brunei Darussalam.67

I Putu Gede P. Damayanto & E.A. Widjaja

A noteworthy Dendrocalamus (Poaceae: Bambusoideae)

from Sumatra, Indonesia.75

I. Haerida

Liverworts of Bali, Indonesia, with new records to the island.81

C.-W. Lin, D.C. Thomas, W.H. Ardi & C.-I Peng

Begonia ignita (sect. Petermannia, Begoniaceae), a new species

with orange flowers from Sulawesi, Indonesia.

N. Simonsson Juhonewe & M. Rodda

Contribution to a revision of Hoy a (Apocynaceae: Asclepiadoideae)

of Papuasia. Part I: ten new species, one new subspecies

and one new combination.97

S. Lindsay, P.H. Hovenkamp & D.J. Middleton

New combinations and typifications mAglaomorpha (Polypodiaceae).149

Date of publication: 24 May 2017

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 69(1): 1-31. 2017

doi: 10.3850/S201009811600002X

Taxonomy and conservation status of Microchirita

(Gesneriaceae) in Peninsular Malaysia

A.R. Rafidah

Forest Biodiversity Division, Forest Research Institute Malaysia (FRIM),

52109 Kepong, Selangor, Malaysia

rah dahar @fr im. go v. my

ABSTRACT. A revision of Microchirita (C.B.Clarke) Yin Z.Wang (Gesneriaceae) in Malaysia

is presented as a precursory paper for the Flora of Peninsular Malay sia account. The taxonomic

treatment includes a key for identification, detailed descriptions of the species, information on

seed morphology (illustrated by Scanning Electron Microscopy micrographs), and a discussion

on distributions, including maps, and the conservation status for each species.

Keywords. Chirita, limestone, seed morphology, SEM

Introduction

The genus Microchirita (C.B.Clarke) Yin Z.Wang was raised by Wang et al. (2011)

from Chirita section Microchirita C.B.Clarke based on molecular phylogenetic

studies by Moller et al. (2009) and Wang et al. (2011). This change of status was

also supported by Weber et al. (2011). Microchirita comprises approximately 28

species in India (Western Ghats and NE India), southern China, Indochina, Myanmar,

Thailand, Peninsular Malaysia, Borneo (Sarawak) and Indonesia (Sumatra, Java,

Bali and Sumba). The highest concentration of species is in Thailand (Weber et al.,

2011; Middleton & Triboun, 2013; Puglisi et al., 2016). Of the six taxa recognised in

Peninsular Malaysia two are endemic, Microchirita ruthiae Rafidah and M. sericea

(Ridl.) A. Weber & Rafidah (Rafidah, 2011). Microchirita species are herbs that usually

grow in forest or on exposed rocks in the open or in light shade. In Malaysia, the genus

is restricted to limestone areas (Henderson, 1939; Kiew, 2009).

In preparation for the Flora of Peninsular Malaysia account, complete

descriptions of all Malaysian Microchirita species were prepared. The aim of the study

was to examine morphological characters in detail, including ones that in previous

descriptions were incompletely described, through the observation of living plants and

herbarium specimens. These characters include colour, stigma form, hairs on corolla,

and seeds. Furthermore, the distribution and the conservation status of each species is

assessed. Recommendations have been made regarding their long-term protection.

Card. Bull. Singapore 69(1) 2017

A need for protection

Limestone karst hills and caves are an integral part of our natural heritage that have

taken millions of years to form and, therefore, cannot be sustainably exploited.

Limestone hills are especially rich in biodiversity which is often found in no other

habitats. Therefore, national conservation action plans must urgently be implemented

to protect these hills. National programmes should include long-term holistic strategies

that balance the use of limestone hills and the preservation of their unique biodiversity.

There are potential non-destructive uses of limestone hills for their caves, possible

sustainable bird’s nests harvesting, their value as scenic landscapes and monuments,

their archaeological and cultural heritage, and their recreational value.

Actions : Threatened species require conservation action plans that include more

intensive surveys, particularly for endemic species and threatened species. Species

categorised under guidelines from IUCN (2012) as Critically Endangered (CR) or

Data Deficient (DD) hold the greatest conservation concern due to a combination of

extreme rarity and loss of habitat.

Legal status : Chua et al. (2009) suggested limestone hills should be included in the

Permanent Forest Reserves network or that the protected status of Totally Protected

Areas should be properly enforced. Microchirita species are restricted to limestone

hills, so any disturbance to these limestone hills creates an impact on the included

flora.

Buffer zone : In Peninsular Malaysia, apart from the limestone hills within Taman

Negara, it is rare to find a hill with any surrounding forest left intact (Kiew, 1991).

Karst hills are a unique habitat for several rare and endemic plants. In Peninsular

Malaysia an estimated 1250 species are found on limestone (which is 22% of the

Peninsular Malaysia flora), of which at least 125 species are endemic to limestone.

Microchirita species mostly grow at the base of hills, especially the more widespread

species such as M. caliginosa , and it is therefore important to have a buffer zone

around these hills.

Awareness & education : To support the conservation of limestone hills, awareness

of the threats to this habitat in the general public and by state authorities needs to be

promoted.

Material and methods

Microchirita specimens were examined from the following herbaria: BK, BKF, BO,

E, K, KEP, KLU, L, SAN, SAR, SING and UKMB (Thiers, continuously updated).

Complete descriptions are based on herbarium specimens as well as observations

on living material from field studies, together with materials cultivated in the FRIM

Microchirita in Peninsular Malaysia

(Forest Research Institute Malaysia) nursery. Digital colour photographs were taken to

illustrate morphology and habitat.

To examine the testa surface, seeds of Microchirita species were removed

from fruits of herbarium specimens or from fresh material and were mounted using

double-sided tape on aluminium stubs. They were then sputter coated with gold at 20

in A for 90 seconds. The specimens were then examined under a Scanning Electron

Microscope, models FEI (Quantum 200) and JEOL (JSM-6400). Micrographs were

made to record details of the seed at various magnifications.

To prepare distribution maps of a particular species/taxon, data from herbarium

specimens were entered directly into BRAHMS (Botanical Research and Herbarium

Management System) database software. The conservation status assessment for all

species was made using the modified Malaysian Plant Red List Categories which is

used solely for species growing in Malaysia (Chua & Saw, 2006). The Malaysian

Rare (RA) category is an additional category to the IUCN Red List Categories (Chua,

2012) and refers to taxa that are rare but not necessarily in danger of extinction. A

taxon data information sheet was completed for each species and the information was

uploaded into the Malaysia Clearing House Mechanism website (http://www.chm.

frim.gov.my/). The Extent of Occurrence (EOO) and Area of Occupancy ( AOO) were

calculated for each species using Arc View GIS 3.2a software. Distribution maps of all

species are shown in Maps 1-7.

Conservation status and threats

In Malaysia, the limestone flora has repeatedly been recognised as critically endangered

(Davis et al., 1995a, 1995b; Chua et al., 2009; Saw et al., 2009; Kiew et al., 2011) and

has been identified as one of the Important Plant Areas (IPAs) (Kiew et al., 2011). In

Peninsular Malaysia, limestone karst hills are mostly concentrated in the central and

northern states and most hills lie on state land that has no legal protection (Chua et

al., 2009). According to Kiew (1991), the limestone flora is particularly vulnerable

because it occupies such a small area (0.4% of the land area of Peninsular Malaysia),

and is threatened by quarrying and mining, land clearance for agricultural practices,

burning, flooding by hydroelectric dams, temple building and damage associated with

recreational activities. Price (2001) stated that in 1975, a total of 62 limestone hills

were quarried (3 in Perlis, 8 in Kedah, 32 in Perak, 3 in Selangor, 7 in Pahang and 6

in Kelantan). However by 2013, there were 66 active quarries in Perak (Ramli et al.,

2014) alone. In Sarawak, the major threats to the vegetation of limestone hills include

farming activities, which can lead to the burning of limestone vegetation, quarrying

and the collection of plants for sale.

The current critical threats to Microchirita species are mostly from quarrying

and mining of the limestone, mostly for cement, thereby destroying the habitat and

the vegetation. Endemic species are the most vulnerable to extinction. Other threats

to the plant populations are land clearance around the foothills, mostly for agriculture

leading to changes in microclimate and increased risk of fire, particularly in Kelantan

Card. Bull. Singapore 69(1) 2017

and Pahang, and also from direct human disturbance.

Microchirita species are not economically valuable but rather have value as

flagship species of threatened limestone habitats because they are found nowhere

else. Table 1 summarises the conservation status of each species of Microchirita in

Malaysia. Two taxa are categorised as Teast Concern (LC), one as Near Threatened

(NT), one as Vulnerable (VU) and two as Endangered (EN). None of the Microchirita

species in Malaysia are categorised as Extinct (EX). The two taxa that are categorised

as LC occur within the network of Totally Protected Areas (in Taman Negara) and

have no known current threats. Of the three threatened taxa, the two Endangered

taxa, Microchirita ruthiae (found only in Kelantan) and M. sericea (found only in

Perak, Perlis and Kedah), are endemic to Peninsular Malaysia and the hills where

they grow are not legally protected and are threatened by quarrying. The Vulnerable

taxon, Microchirita rupestris (Ridl.) A.Weber & Rafidah, is categorised as Vulnerable

in Malaysia whereas globally it should be considered Least Concern (Middleton, pers.

comm.). Further details are under each species.

Table 1. Conservation status of Microchirita species in Peninsular Malaysia.

Species

No. of hills

IUCN Category

Microchirita caliginosa (C.B.Clarke) Yin Z.Wang

Least Concern

Microchirita involucrata (Craib) Yin Z.Wang

Least Concern

Microchirita rupestris (Ridl.) A.Weber & Rafidah

Vulnerable

Microchirita ruthiae Rafidah

Endangered

Microchirita sericea (Ridl.) A.Weber & Rafidah

Endangered

Microchirita viola (Ridl.) A.Weber & Rafidah

Near Threatened

Notes on selected habitat and morphological characters

In Peninsular Malaysia, as elsewhere, Microchirita species are morphologically very

similar, especially when sterile. Nevertheless, there are a few characters that separate

the species even though vegetative and reproductive characters can be variable.

Observations in the field, together with photos, are important in recording diagnostic

characters that cannot be seen or are difficult to see in dried herbarium specimens.

Microchirita caliginosa (C.B.Clarke) Yin Z.Wang and M. involucrata (Craib)

Yin Z.Wang are frequently found at the base of hills where there is a thin soil layer.

On this soil Microchirita caliginosa and M. involucrata always produce carpets

of seedlings that grow rapidly. Microchirita sericea, M. rupestris, M. viola (Ridl.)

A.Weber & Rafidah and M. ruthiae, however, inhabit rock crevices and cracks in

the limestone, sometimes in wet conditions. Microchirita species often have a short

lifespan or are annual with most species living for less than six months to one year

(pers. obs.). What would appear to be perennial plants have been observed, however,

Microchirita in Peninsular Malaysia

in Microchirita ruthiae and M. sericea. Observations of the developmental stages

(buds, flowering and fruiting stages) shows that Microchirita species produce flowers

at a very early stage when they have only one or two leaves, then flower continuously

throughout their short life (pers. obs.). Microchirita caliginosa and M. involucrata ,

and occasionally M. rupestris , are monopodial.

In general, the hair type of the lamina surface is different in each Microchirita

species. The type of hairs (whether pubescent, strigose or sericeous) and the density

(sparse or dense) vary across the species. It varies from sparsely pubescent (M.

involucrata ), to moderately densely pubescent (M. ruthiae ) to densely pubescent (M.

caliginosa ), while M. rupestris and M. viola have sparsely strigose hairs. Furthermore,

Microchirita sericea has densely sericeous hairs. The colour of the upper surface of the

fresh lamina varies from yellowish green, whitish green to dark green, while the lower

surfaces of Microchirita viola and M. caliginosa are sometimes reddish to reddish-

green. This character is useful in the field.

Some species have epiphyllous or crested inflorescences (. Microchirita ruthiae )

or they have cymose inflorescences (Af. involucrata and M. rupestris). The individual

inflorescences of Microchirita are always up to 6-flowered or rarely more.

Ridley (1905, 1908, 1923) relied on bract characters in his keys to Microchirita

species, thus treating this character as diagnostic. Two species of Microchirita have

conspicuous bracts: M. rupestris (saucer-shaped bract) and M. involucrata (foliose

bract) (Fig. 1).

There are differences in the size and indumentum of the calyx lobes but the

differences are small. In general, the indumentum of the calyx of Microchirita caliginosa

and M. ruthiae is similar in their glandular and eglandular hairs. Microchirita rupestris

and M. viola have ciliate hairs while M. sericea has densely sericeous hairs.

The shape of the corolla tube varies from slightly curved to more strongly curved

downwards. The length of the corolla tube ranges from 1.4-5.5 cm and the width from

2^1 mm. The colour of the corolla tube varies from cream, yellowish, pale purple to

dark purple, to occasionally whitish. The throat colour for all Microchirita species is

yellow or whitish, except for M. involucrata that usually possesses a dark red throat.

The glandular hairs in the throat of the corolla in some species are translucent

or glistening-translucent, bead-like hairs except for M. rupestris that has a cluster of

hairs (not bead-like) around the corolla mouth. Microchirita viola differs from the

other species in having conspicuously attractive golden yellow, glandular bead-like

hairs that can be seen from the corolla mouth.

In general, the corolla lobes are usually unequal in all Microchirita species, and

the lowest lobe is the largest. Microchirita viola is distinct from all the other species

by having the median lower lobe much longer than the other two lobes. The width

of the lobes was measured from lobe to lobe (across). The width of the Microchirita

sericea corolla reaches to 3 cm across while the rest are less than 2.5 cm wide and

most range from 1.2 to 2 cm. The colour of the lobes is always much darker than that

of the corolla tube and varies from whitish, pale blue, pale purple to deep purple or

violet. The lobes may be striped or almost plain-coloured (Fig. 2). Microchirita viola

and M. sericea have conspicuously striped lobes while the others possess faint stripes.

Card. Bull. Singapore 69(1) 2017

Fig. 1. Conspicuous bracts in Malaysian Microchirita. A, B. Microchirita rupestris (Ridl.)

A.Weber & Rafidah. C. Microchirita involucrata (Craib) Yin Z.Wang. (Photos: A.R. Rafidah)

In Peninsular Malaysia, flower colour is of great importance in separating the species

even though this character is lost on dried specimens. Flower colour is constant, only

a few have occasional white flowers in purple populations. Photographs of the flowers

are therefore crucial.

Microchirita species have two stamens and the filaments are usually curved

and geniculate with the exception of M. involucrata and M. ruthiae that have straight

filaments. The anthers are generally fused apically or joined. However, the anthers of

Microchirita ruthiae are free and not joined together.

Two types of hairs, glandular or eglandular, are found on the ovary and style in

Microchirita species. Microchirita caliginosa and M. ruthiae have both types of hairs,

while the others consistently only have eglandular hairs. In fresh flowers, stigma colour

varies from white, whitish green, pale purple to deep purple, independently of the

colour of the style or the corolla. The stigmas are shallowly bilobed or deeply bilobed

and covered with papillate trichomes. Unequal stigma lobes are seen in Microchirita

involucrata.

The seed shape in Microchirita species ranges from narrowly ellipsoid

through ellipsoid to broadly ellipsoid or broadly ovoid (Table 2). Species with a seed

length:width ratio of 3:1 (seen clearly in Microchirita viola ) are described as narrowly

ellipsoid, which distinguishes it from the other species. Those with a ratio 2:1, such as

Microchirita in Peninsular Malaysia

Fig. 2. Corolla shape, colour and pattern. A. Microchirita caliginosa (C.B.Clarke) Yin Z.Wang.

B. M. ruthiae Rafidah. C, D. M. involucrata (Craib) Yin Z.Wang. E, F. M. rupestris (Ridl.)

A.Weber & Rafidah. G. M. sericea (Ridl.) A.Weber & Rafidah. H. M. viola (Ridl.) A.Weber &

Rafidah. (Photos: A.R. Rafidah & K. Imin)

Microchirita caliginosa, M. involucrata , M. sericea and M. ruthiae , have a narrowly

ellipsoid to broadly ellipsoid or broadly ovoid shape. The cell edges of Microchirita

species also vary from weakly elevated ( Microchirita viola and M. sericea ) to elevated

(M. caliginosa and M. ruthiae ) to strongly elevated (M. involucrata and M. rupestris).

Table 2. Summary of seed characters in Microchirita species.

Card. Bull. Singapore 69(1) 2017

S-h

s__.

. .

P*h

>^>

D TP

03 D

D 3

| >

»*»

(00

£ 3

J-H

& 1

b.a

-4—>

h> D

£ £

‘3

.Pk

+->

too

S-h

CCS

.CD

-t—>

-C

C<3

§■

.Pk

+j>

Cod

C/2

-t—>

C/2

-(-<

»■*»

3 «

3c ^

o o

Z 3

-j—>

—H

v CJ

toD

+->

S-H

h->

C/2

+->

a* d

3 iP 1

'3 3

oo o:

Pk P

3 £

_&P

-t->

_ D

C/2

’3

P^h

ctf

PLh

/-H

O 3

Shape

COD

3 X

CCS o

k< S-H

> ^

> Pk

__ CS

S-H

C/2

Microchirita in Peninsular Malaysia

Fig. 3. SEM micrographs of seeds of Microchirita showing seed shape, size and surface pattern.

A, B. M. caliginosa (C.B.Clarke) Yin Z.Wang. C, D. M. involucrata (Craib) Yin Z.Wang. E,

F. M. rupestris (Ridl.) A.Weber & Rafidah. G, H. M. ruthiae Rafidah. I, J. M. sericea (Ridl.)

A.Weber & Rafidah. K, L. M. viola (Ridl.) A.Weber & Rafidah. (Photos: A.R. Rafidah)

Card. Bull. Singapore 69(1) 2017

The surface pattern of the seed is consistent in all species, that is, it is either

papillate or canaliculate (Fig. 3/Table 2). All Microchirita seeds have bumps or knobs

within a thick square (up to 10 bumps), except M. viola which does not form squares

and the bumps are arranged continuously in lines, while M. sericea forms weak

squares. This character is useful for distinguishing between species.

Taxonomic treatment

Microchirita (C.B.Clarke) Yin Z.Wang, J. Syst. Evol. 49: 59 (2011); Weber et al.,

Taxon 60: 778 (2011). - Chirita sect. Microchirita C.B.Clarke in A.DC. & C.DC.,

Monogr. Phan. 5(1): 127 (1883); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 134

(1974). - Roettlera sect. Microchirita (C.B.Clarke) Fritsch in Engl. & Prant, Nat.

Pflanzenfam. 4(3B): 148 (1895). - Didymocarpus sect. Microchirita (C.B.Clarke)

Chun, Sunyatsenia 6: 290 (1946). - TYPE: Microchirita hamosa (R.Br.) Yin Z.Wang,

lectotype designated by Burtt (1954).

Short-lived or rarely perennial, caulescent (erect or creeping) herbaceous plants; stem

branched or unbranched, sometimes slightly woody, often fleshy, indumentum of

white or silvery soft, glandular hairs. Leaves opposite, whorled or decussate; petiole

long or short, winged or not; lamina variable in shape, ranging from narrowly elliptic

to ovate, base sometimes unequal, margin entire or serrate. Inflorescences axillary,

epiphyllous, crested or cymose, one- to many-flowered; bracts free, cup-shaped, leaf¬

like or caducous. Flowers : calyx pentamerous, free to base, appressed in the fruit;

corolla bilaterally symmetrical, tubular to broadly funnel-shaped with 5 lobes, upper

lip 2-lobed, lower lip 3-lobed, lobes rounded; stamens 2, filaments adnate to the

corolla, anthers fused either apically or joined by a ligature, rarely free, anther-thecae

divergent, staminodes 2 or very rarely 3; nectary rim entire, cleft or lobed; carpels

two, rarely with one carpel sterile, style one, straight, stigma bilobed. Fruit a slender

capsule, plagiocarpic, dehiscence on both sides. Seeds numerous, tiny.

Distribution. Approximately 28 species in India (Western Ghats and NE India),

China, Indo-China, Myanmar, Thailand, Peninsular Malaysia, Borneo (Sarawak) and

Indonesia (Java, Sumatra, Bali, and Sumba) with centre of diversity in Thailand. In

Malaysia, six species are recorded.

Habitat and ecology. In Peninsular Malaysia restricted to limestone hills. In wet, light

to moderately shady places at cliff bases, on cliff walls in crevices and cracks, or at

cave entrances.

Microchirita in Peninsular Malaysia

la.

lb.

2a.

2b.

3a.

3b.

4a.

4b.

5a.

5b.

6a.

6b.

7a.

7b.

Key to Microchirita species in Malaysia

Lamina narrowly elliptic or elliptic with widest point of lamina at the middle ... 2

Lamina ovate, obovate, lanceolate or ovate with widest point of lamina below or

above the middle.4

Margin serrate.4. M. ruthiae

Margin serrulate.3

Stem erect; leaf lower surface pale green to yellowish green, sometimes reddish

green, base narrowly cuneate.1. M. caliginosa

Stem creeping; leaf lower surface very pale green or whitish green, base attenuate

or sometimes cordate.5. M. sericea

Inflorescences cymose, bracts present.5

Inflorescences epiphyllous and crested, bracts absent (or minute).6

Bracts connate-perfoliate into cup-like arrangement.3. M. rupestris

Bracts not fused at base, leaf-like.2. M. involucrata

Corolla tube pale violet, lobes with conspicuous dark purple stripes; glandular

hairs golden yellow, apically swollen in a cluster above the anther.6. M. viola

Corolla tube white or cream, pale yellow or pale purple, lobes conspicuously

striped or plain; glandular bead-like hairs translucent or glistening yellow or

brown. 7

Corolla tube white; anthers free. 4. M. ruthiae

Corolla tube cream, pale yellow or pale purple; anthers fused apically or joined

by apical ligature.5. M. sericea

1. Microchirita caliginosa (C.B.Clarke) Yin Z.Wang, J. Syst. Evol. 49: 60 (2011). -

Chirita caliginosa C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 122 (1883);

Clarke in Hooker, FI. Brit. India 4: 360 (1884); Ridley, J. Linn. Soc., Bot. 32: 516

(1896); Ridley, J. Straits Branch Roy. Asiat. Soc. 44: 58 (1905); Ridley, FI. Malay

Penins. 2: 524 (1923); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 186 (1974). -

Roettlera caliginosa (C.B.Clarke) Kuntze, Revis. Gen. PI. 2: 476 (1891). - TYPE:

Malaysia, Peninsular Malaysia, Kuala Lumpur [Klang], 1891, Kehding in Beccari 73

(holotype FI [FI013109], digital image seen). (Fig. 2A, 3A-B, 4, 5)

Chirita parvula Ridl., J. Fed. Malay States Mus. 10: 149 (1920); Ridley, FI. Malay

Penins. 2: 524 (1923). - TYPE: Malaysia, Peninsular Malaysia, Pahang, KotaTongkat,

10 th mile Benta-Kuantan road, June 1917, Evans s.n. (holotype K [K000450490]).

Card. Bull. Singapore 69(1) 2017

100W

loroc

102W

103W

1Q4W

Selected Localities

1. Bkt. Kaplu

2. Gua Badak

3. Ipoh

4. BktTakun

5. Bkt, Chintamani

6. Gn, Senyum

1. Kota Gelanggi

8, Ranching F.R.

9 Batu Biwa

10. Taman Negara

Extent of Occurrence (EDO)

46,616 sq km

Area of Occupancy (AOO):

112 sq km

Forest cover within EOO :

66 %

O-

104 "OC

1 : 3,Sou,00u

SO Kilometers

NFI ill Courtesy of Forestry Department Peninsular Malaysia

SINGAPORE

102.W

103°0Q‘

Location based on :

# Herbarium specimen(s)

□ Forest cover as in Nations!

Forest Inventory 111 (1991-1993)

100W

101 ^ 00 -

Fig. 4. Distribution of Microchirita caliginosa (C.B.Clarke) Yin Z.Wang in Peninsular

Malaysia.

Microchirita in Peninsular Malaysia

Fig. 5. Distribution of Microchirita caliginosa (C.B.Clarke) Yin Z.Wang in Sarawak.

Chirita fusca C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 129 (1883). -

Roettlerafusca (C.B.Clarke) Kuntze, Revis. Gen. PI. 2: 476 (1891). -TYPE. Malaysia,

Sarawak, June 1896, Lobb (holotype K [000858415]).

Branching, monopodial, herbaceous, short-lived plant. Stems brown to pale red or pale

maroon, succulent, erect, flowering at 1 cm tall, erect up to 60 cm, 2-7 mm diameter,

shortly decumbent at the base, hairs pubescent or strigose, rarely glabrous, internodes

to 10 cm long. Leaves opposite, decussate, lowermost solitary (resulting from one of

a pair dying); petioles green sometimes maroon, 2-6 cm long, hairs glandular and

eglandular; lamina narrowly elliptic or elliptic, 6-15.5 x 2.5-7 cm, yellowish green

to dark green above, pale green or greyish green or sometimes reddish beneath, soft

when fresh, papery when dry, densely pubescent, base narrowly cuneate, unequal,

margin serrulate, apex acuminate; midrib sunken above, prominent beneath, pubescent

beneath, lateral veins 5-10 pairs on each side of the midrib, intercostal veins reticulate.

Inflorescences epiphyllous, to 6-flowered; bracts absent; pedicels pale green.

Card. Bull. Singapore 69(1) 2017

sometimes greenish maroon, shortly fused to the petiole, to 1.5 cm long, sparsely

glandular hairy. Calyx green, clasping the corolla tube, lobes narrowly triangular or

very narrowly ovate, 10-12 mm long, 8-10 mm wide, margin ciliate, outer surface

glandular and eglandular hairy. Corolla very sparsely hairy outside and inside, tube

usually cream or yellowish, sometimes whitish or pale purple, 2.4-5.5 cm long,

slightly curved, 2-4 mm wide at the base, inside throat dark yellow to pale yellow,

glandular hairs in the mouth of the tube above the anthers translucent pale yellow;

upper lobes elliptic, not spreading, 4-5.5 x 5-6 mm, lower lateral lobes elliptic, 5-6

x 4-6 mm, lower central lobe elliptic, slightly elongated, recurved or not, 6-8 x 5-10

mm Stamens, filaments cream or white, inserted at 8-10 mm from the base of the

corolla, 3-6.5 mm long, slightly curved or geniculate, glabrous; anthers pale yellow,

pale brown or whitish, at base with long white deflexed hairs, anther thecae divergent,

2-3 mm long; staminodes 2 (rarely 3), cream or green, vestigial, inserted at 9 mm

from the base of the corolla tube, 2-7 mm long, glabrous. Nectary pale yellow or

cream, a shallowly lobed ring, to 1 mm high. Pistil pale green; ovary 1.8-2.5 cm long,

1 mm wide at the base, narrowing to 0.5 mm below the stigma, hairs glandular and

eglandular; style 6-28 mm long; stigma deeply 2-lobed, to 3 mm long, above with

very fine papillose hairs, beneath with very sparsely glandular hairs; ovules cream,

less than 1 mm long. Capsules green, cylindric, slender, slightly curved towards tip, to

8 cm long, 1.3-1.8 mm diameter, sparsely hairy; calyx persistent, green, hairy. Seeds

yellowish cream to brownish, 435-550 x 100-230 pm, ellipsoid to narrowly ellipsoid,

surface papillate or canaliculate.

Distribution. Endemic to Malaysia: Peninsular Malaysia (Kedah, Pahang, Perak,

Selangor and Terengganu) and Sarawak (Kuching Division).

Habitat and ecology. Damp places on limestone karst hills, limestone cliffs, crevices

and on open limestone rocks, on thin soil or on guano in caves or cave mouths, around

the base or at the foot of limestone hills and at the base of shaded rocks. Altitude to

240 m.

ProvisionalIUCNconservation status. Least Concern (LC). Populations are widespread

within the network of Totally Protected Areas in Taman Negara (the National Park),

both in Pahang and Terengganu.

Additional specimens examined. PENINSULAR MALAYSIA: Kedah: Kodiang, Bkt. Kaplu,

6 Nov 2009, Rafidah FRI 64417 (KEP); ibidem, 27 May 2010, Rafidah FRI 64545 (KEP).

Pahang: Bkt. Charas, 15 Oct 1931, Henderson 25233 (SING); ibidem, 26 Nov 1984, Kiew

RK 1557 (KEP); ibidem, 1 Apr 2008, Rafidah FRI 55717 (KEP); Bkt. Chintamani, 4 Oct 1931,

Henderson SFN 25033 (SING, BK); Gn. Jebak Puyuh, 10 Feb 1986, Kiew RK 2158 (KEP);

ibidem, 10 Feb 1986, Kiew RK s.n. (KEP); Gn. Senyum, 30 Jul 1929, Henderson s.n. (SING);

ibidem, 28 Nov 1984, Kiew RK 1587 (KEP); ibidem, 2 Apr 2008, Rafidah FRI 55721 (KEP);

Gua Bama, 3 Apr 2008, Rafidah FRI 55726 (KEP); Gua Cermin R.F., 31 Mar 2008, Rafidah

FRI 55713 (KEP); ibidem, 31 Mar 2008, Rafidah FRI 55714 (KEP); Gua Kechil, 30 Jul 2009,

Rafidah FRI 643 79 (KEP); Kota Glanggi, 4 Aug 1929, Henderson SFN22419 (SING); ibidem, 2

Microchirita in Peninsular Malaysia

Apr 2008, Rafidah FRI55724 (KEP); Panching, 26 Nov 1984, Kiew RK 1571 (KEP); Panching

F.R., 15 Oct 1931, Henderson SFN 25223 (SING); Taman Negara, Batu Subuh, 5 Oct 1984,

Dawn RK 1470 (KEP); Taman Negara, Kuala Keniyam, 29 Sep 1982, Kiew RK 1202 (KEP).

Perak: Batu Kurau, Dec 1884, Scortechini 1582 (SING); Ipoh, 17 Jul 1917, Burkill 2558

(SING); ibidem, s.d., Gordon GS 435 (KLU); ibidem, Feb 1904, Ridley s.n. (SING); Gopeng,

8 Mar 1993, Davison GD 4 (KEP); Gn. Lanno, 16 Apr 1925, Mills 15061 (SING); Gn. Mesah,

20 Apr 1962, Bnrtt B1665 (SING); ibidem, May 1902, Curtis s.n. (SING); Gn. Pipit, 23 Apr

1987, Kiew RK 2524 (KEP); Gn. Rapat, 21 Jul 2009, Rafidah FRI 64347 (KEP); ibidem, 9 Mar

1931, Samsuri SA 560 (KLU, SING); Kinta, Aug 1898, Curtis 3109 (SING); ibidem, Jan 1885,

King’s collector 7028 (SING); ibidem, 1885, King’s collector 937 (SING); Kuala Dipang F.R.,

Feb 1890, Curtis 2359 (SING); ibidem, 1898, Ridley s.n. (SING); Lenggong, Gua Badak, 28

Oct 2008, Imin FRI 63212 (KEP); ibidem, Jul 1909, Ridley s.n. (SING); Sg. Siput, 21 May

1985, Anthonysamy SA 842 (KEP); Sg. Siput Utara, 7 Jan 2015, Rafidah FRI 82007 (KEP).

Selangor: Batu Caves, 18 Oct 1983, Anthonysamy SA 379 (KEP); ibidem, 19 Nov 1916,

Burkill 2253 (SING); ibidem, s.d., Chung 331 (KLU), s.d., Chung 390 (KLU); ibidem, Feb

1890, Curtis 2359 (SING); ibidem, May 1902, Curtis s.n. (SING); ibidem, s.d., DingHou 715

(KEP); ibidem, 23 Jan 1966, Hardial 477 (SING); ibidem, 1889, Kelsall s.n. (SING); ibidem, 1

May 1981, Kiew RK 1023 (KEP); ibidem, Mohd. Kasim 391 (KLU); ibidem, 14 Oct 1966, Ng

FRI 1629 (KEP, SING); ibidem, 23 Jun 1889, Ridley s.n. (SING); ibidem, 4 Nov 1953, Sinclair

SFN 40066 (KEP, SING); ibidem, 29 Nov 1959, Smith KEP 85205 (KEP); ibidem, s.d., Yap

SK 26 (KLU); Bkt. Anak Takun, 27 Apr 2006, Phoon FRI 51570 (KEP, SING); ibidem, 3 May

2005, Sam FRI 50118 (KEP); Kanching F.R., 10 Jul 1995, Julia JS 26 (KEP); Bkt. Takun, 3

Nov 1937, Mohd. Nur 34389 (SING); ibidem, 9 Mar 1988, Saw FRI 36215 (SING); ibidem,

20 Nov 1962, Sinclair 10732 (SING); ibidem, Nov 1969, Stone 8934 (KLU); ibidem, 21 Sep

1969, Stone 8794 (SING); ibidem, 24 Jun 1933, Symington FMS 30796 (KEP); ibidem, 8

May 1935, Symington 39598 (KEP); Kanching, 16 Mar 1935, Symington FMS 37431 (KEP).

Terengganu: Taman Negara, Batu Biwa, 25 Oct 1986, Kiew RK 2339 (KEP, SING); ibidem,

22 Oct 1986, Kiew RK 2284 (SING).

Notes. This is overall the most common species in Malaysia but is less common

further north (Kedah mainland and Kelantan) where it is then replaced by Microchirita

involucrata. Where the distribution of these two species overlaps, they can occasionally

be found on the same hill, e.g. Gua Bama, Pahang. It is absent from Langkawi, Kedah.

It is the only species to occur in Borneo (Sarawak). Specimens for Sarawak are listed

in Kiew (2004). Microchirita caliginosa always produces carpets of seedlings that

grow very quicldy. One unusual Microchirita caliginosa population was found in

Bukit Kaplu, Kedah, where the stem and lower lamina surface was reddish and the

corolla was much smaller compared with the usual M. caliginosa.

2. Microchirita involucrata (Craib) Yin Z.Wang, J. Syst. Evol. 49: 60 (2011)

(“ involucrate ”). - Chirita involucrata Craib, Gard. Chron., Ser. 3, 83: 140 (1928);

Wood, Notes Roy. Bot. Gard. Edinburgh 33: 199 (1974). - TYPE: “Cult. Hort. Bot.

Abdn, 17/xi/27 from seed collected in Surat on Kaw Tao by A.F.G. Kerr, 30/xii/26”

(lectotype ABD, specimen with appended protologue, designated here by Carmen

Puglisi; isolectotype ABD). (Fig. 2C-D, 3C-D, 6)

Card. Bull. Singapore 69(1) 2017

Selected Localities

1. Langkawi 6.

2. Gunung Keriang 7.

3. Gua Labua 8,

4. Gn. Inas F.R. 9.

5. Gua Musang-K.Lipis Rd. 10,

Taman Negara

Gua Musang

Gua Ikan

Gua Setir

Kuala Betis

Extent of Occurrence (EOO):

21,004 sq km

Area of Occupancy (AOO):

72 sq km

Forest cover within EOO :

71%

□

■]

o'?

Location based on:

• Herbarium specimen(s)

□ Forest cover as in National

Forest Inventory III (1991-1993)

CF>

-o

O’

-CD

-o

-a

1C0“00’

100'W

loroo 1 2 3 4 5

ioroo'

102 * 00 '

102 J 00'

■W

*00'

104"GO’

104*00'

50 Kilometers

IMF! Ill Courtesy of Forestry Department Peninsular Malaysia

1 : 3,500.000

Fig. 6. Distribution of Microchirita involucrata (Craib) Yin Z.Wang in Peninsular Malaysia.

Microchirita in Peninsular Malaysia

Branching, monopodial, herbaceous, short-lived plant. Stems pale green, reddish green

or purplish, succulent, to 36 cm tall, 4-7 mm diameter, slightly pubescent, sometimes or

rarely glabrous, internodes to 11 cm long. Leaves decussate; petioles green, sometimes

maroon, 2-10 cm long, pubescent; lamina ovate to broadly ovate, 5.5-11 x 3.5-7 cm,

yellowish green to dark green above, pale green or whitish green beneath, soft when

fresh, chartaceous when dry, sparsely pubescent, base rounded or subcordate, unequal,

margin serrulate, apex acute; midrib sunken above, prominent beneath, pubescent

beneath, lateral veins 6-8 pairs on each side of the midrib, intercostal veins reticulate.

Inflorescences axillary, cymose, to 6-llowered; peduncles, bracts and pedicels

sparsely glandular hairy; peduncle 1.8-3 cm long, not fused to petiole; bracts dark

green, opposite, in pairs, leaf-like, 10-20 x 9-15 mm; pedicels pale green, sometimes

greenish maroon or downy, shortly fused to the petiole, 1-2 cm long, sparsely

glandular hairy. Calyx pale green, clasping the tube, lobes narrowly triangular or very

narrowly ovate, 5-6 mm long, 1-2 mm wide, margin ciliate, outer surface glandular

and eglandular hairy. Corolla very sparsely hairy outside, corolla tube usually lilac

or pale purple, sometimes dark purple, 1.4-2 cm long, slightly curved, 2-3 mm wide

at the base, throat dark red; lobes plain or very faintly striped, upper lobe sometimes

dark or violet blue, the lower lobes dark pink or dark purple, upper lobes triangular, not

spreading, 3-3.5 x 5-6 mm, lower lateral lobes elliptic, 5-6 x 4-5 mm, lower central

lobe elliptic, 5-6 x 8-9 mm Stamens, filaments white, inserted at 6-10 mm from the

base of the corolla, c. 6 mm long, straight, glabrous; anthers purplish white or dark

purple; anther-thecae divergent, 1-2 mm long; staminodes 2, white, vestigial, inserted

at 6-7 mm from the base of the corolla tube, 4-5 mm long, hairy, sometimes glabrous.

Nectary cream or pale yellow, entirely cylindric, to 1 mm long. Pistil whitish green;

ovary 5-9 mm long, 1.5 mm wide at the base, narrowing to 0.5 mm below the stigma,

glabrous; style to 6 mm long; stigma pale brown or whitish purple, shallowly bilobed,

1-2 mm long, slightly bent backwards when mature, with very fine papillose hairs

above; ovules cream, less than 1 mm long. Capsule pale to dark green, to 5.5 cm long,

1-2.5 mm wide, slightly curved, erect or horizontal, glabrous, slightly hairy towards

tip; calyx persistent, green, hairy. Seeds yellowish cream to brown, many, 315-380 x

120-250 pm, narrowly ellipsoid to broadly ellipsoid, surface canaliculate, each testa

cell or square with up to 6 knobs or bumps.

Distribution. Peninsular Thailand, Cambodia, southern Vietnam and Malaysia (Kedah,

Kelantan and Pahang).

Habitat and ecology. Growing in limestone crevices and shady places. On Langkawi,

Kedah, on limestone hills by the sea and, inland, at the base of limestone cliffs or on

rich damp soil around cave mouths.

Provisional IUCN conservation status. Least Concern (LC). Populations occur in

Taman Negara (The National Park) and it is the most common species found in the

northern and eastern part of Malay Peninsula.

Card. Bull. Singapore 69(1) 2017

Additional specimens examined. PENINSULAR MALAYSIA. Kedah: Gn. Inas L.R., 30 Oct

2007, Imin FRI58583 (KEP); Gn. Keriang, 6 Nov 2009, Rafidah FRI64418 (KEP); ibidem,

1898, Ridley s.n. (SING); Langkawi, Sep 1924, Burkill 1493 (SING); ibidem, 1978, Keng 79

(SING); Langkawi, P. Buyong, 3 Nov 2009, Rafidah FRI 64395 (KEP); Langkawi, P. Jerkom, 17

Nov 1941, Corner s.n. (SING); Langkawi, P. Langgun, Feb 1989, Tay 155 (SING); Langkawi,

P. Timun, 3 Nov 2009, Rafidah FRI 64394 (KEP); Ulu Muda F.R., Gua Labua, 4 Mar 2008,

Rafidah FRI 55687 (KEP). Kelantan: Elephant Cave, 15 May 1990, Kiew RK 2948 (KEP); Gn.

Reng, 14 Jan 2010, Rafidah FRI 64448 (KEP); Gua Batu Boh, 4 Mar 2008, Chew FRI 58699

(KEP); ibidem, 3 Aug 1962, UNESCO 268 (SING); Gua Ikan, 14 May 1990, Kiew RK 2940

(SING); Gua Setir, 19 May 1990, Kiew RK 3038 (SING); ibidem, 12 Jan 2010, Rafidah FRI

64426 (KEP); Kuala Betis, 15 Feb 2003, Kiew RK 5253A (SING); Batu Papan F.R., 9 May

1990, Kiew RK 2885 (SING); Gua Batu Boh, Gua Chawan, 14 Mar 2000, Kiew RK 4195

(SING); Sg. Jenera, 10 May 1990, Kiew RK 2896 (KEP); Sg. Keteh, Gua Ninik, 18 Feb 1924,

Nur 12130 (SING); Sg. Nenggiri, P. Raba, 12 May 1990, Kiew RK 2920 (SING). Pahang:

Kuala Lipis, 27 Apr 1996, Kiew RK 3913 (KEP); Gua Bama, 25 Aug 2008, Mohd Hairul FRI

60062 (KEP); ibidem, 3 Apr 2008, Rafidah FRI 55728 (KEP); Gua Tipus, Oct 1927, Henderson

SFN 19403 (SING); Taman Negara, Batu Kanok, 9 Oct 1984, Kiew RK 1505 (SING); Taman

Negara, Batu Subuh, 5 Oct 1984, Dawn RK 1472 (KEP); ibidem, 5 Oct 1984, Dawn RK 1474

(KEP); ibidem, 8 Oct 1984, Kiew RK 1491 (KEP); Taman Negara, Batu Kepayang, 3 Oct 1984,

Kiew RK 1424 (KEP); Taman Negara, Bkt. Batu Luas, 2 Oct 1984, Kiew RK 1404 (KEP);

ibidem, 11 Oct 1984, Kiew RK 1537 (KEP).

THAILAND. 1901, Curtis 3221 (SING); Kau Hoa Kwai, Tassateng, Seidenfaden 2134 (SING).

Notes. Microchirita involucrata is most common in the northern and eastern part of

the Malay Peninsula, especially in Kelantan, where it replaces M. caliginosa. Unusual

plants with whitish flowers and 4 stamens were recorded from Gunung Reng, Kelantan.

3. Microchirita rupestris (Ridl.) A.Weber & Rafidah, Taxon 60: 779 (2011). - Chirita

rupestris Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 60 (1905); Ridley, J. Asiat. Soc.

Bengal 74: 766 (1908); Craib, Curtis’s Bot. Mag. 136: t. 8333 (1910); Ridley, FI. Malay

Penins. 2: 525 (1923); Henderson, Malay. Wild FIs., Dicots. 349 (1959); Wood, Notes

Roy. Bot. Gard. Edinburgh 33: 201 (1974). - TYPE: Malaysia, Peninsular Malaysia,

Kedah, Langkawi, November 1889, Curtis 2120 (lectotype SING [SING0042989],

designated here by Carmen Puglisi; isolectoype SING [SING0042990]). (Fig. 2E-F,

3E-F, 7)

Chirita glasgovii Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 60 (1905); Ridley, J.

Asiat. Soc. Bengal 74: 766 (1908); Ridley, FI. Malay Penins. 2: 526 (1923). - TYPE:

Malaysia, Peninsular Malaysia, Perak, Waterloo Estate, December 1897, Robertson-

Glasgow s.n. (holotype SING).

Erect, monopodial, herbaceous, short-lived plant, axillary branches sometimes present.

Stems greenish purple or pale green, to 25 cm tall, 5 mm diameter, starting to flower

when 2-leaved, slightly curved at the root base, finely hairy; internodes not swollen,

5.5-8 cm long. Leaves decussate; petiole green sometimes reddish green, to 6 cm long,

Microchirita in Peninsular Malaysia

Selected Localities

1. P. Langkawi

2. Wang Kelian

3. Bkt. Lagi

4. Gn. Baling

Extent of Occurrence (EOO):

1,243 sq km

Area of Occupancy (AOO):

20 sq km

Forest cover within EOO :

□

Location based on :

# Herbarium specimen(s)

□ Forest cover as in National

Forest Inventory III (1991-1993)

50 Kilometers

NFI III Courtesy of Forestry Department Peninsular Malaysia

1 : 3,500,000

100W 10r00' 1Q2 D 0G' t03 n 00’

-o

SINGAPORE

1 oo;w

101 * 00 '

102 ° 00 '

IQ^CO 1

104*00'

Fig. 7. Distribution of Microchirita rupestris (Ridl.) A.Weber & Rafidah in Peninsular Malaysia.

Card. Bull. Singapore 69(1) 2017

shortly hairy; lamina ovate, sometimes obovate or lanceolate, 8-14 x 4.5-6.6 cm,

yellowish green to dark green above, pale green to whitish green beneath, coriaceous

when fresh and dried, sparsely strigose hairy on both surfaces, base oblique, unequal,

margin shallowly serrate, apex acute; midrib sunken above, prominent beneath, lateral

veins 6-8 pairs on each side of the midrib, intercostal veins reticulate. Inflorescence

axillary, branched cyme, with up to 6 flowers; peduncles, bracts and pedicels with

pubescent hairs; peduncles reddish green or pale green, to 5 cm long, not adnate to

petiole; bracts dark green above, pale green or whitish green beneath, 1.2-3.5 x 1.2-3

cm, connate-perfoliate, margin serrate; pedicels green, to 1 cm long. Calyx pale green,

clasping the tube, lobes narrowly triangular, 8-11 mm long, 7-11 mm wide, outer

surface ciliate, inner surface glabrous. Corolla sparsely hairy outside with a cluster

of hairs in the mouth of the tube above the anthers, hairs not bead-like, tube lilac,

pale purple or white outside, throat whitish, 1.5-2.8 cm long, curved downwards;

lobes faintly striped, dark purple, bluish purple, lilac or white, with 2 yellow patches

on the lower lobes, upper lobe sometimes dark or violet blue, the lower lobes dark

pink or dark purple, upper lobes elliptic, not spreading, 4-5 x 5-6 mm, lower lateral

lobes elliptic, 5-6 x 4-5 mm, lower central lobe elliptic, 5-6 x 8-10 mm. Stamens :

filaments slightly geniculate at the point of attachment, whitish with black-brown spots

at the knees, inserted at 6-10 mm from the base of the corolla, 4-5.5 mm long; anthers

cream with a purple mark, anther-thecae divergent, 2-3 mm long, with reflexed hairs

to 2 mm long on the lower surface; staminodes 2, brown, inserted at 6 mm above the

base of the corolla, less than 4 mm long, glabrous. Nectary pale green, a complete ring,

up to 1.1 mm deep. Pistil light green; ovary 9-10 mm long, 0.5 mm wide at the base,

narrowing to 0.2 mm below the stigma, hairs glandular; style to 4 mm long; stigma

deeply 2-lobed, 2 mm long, very fine glandular hairy (abaxial), slightly papillose

(adaxial); ovules cream, less than 1 mm long. Capsule green, to 8 cm long, slender,

erect; calyx persistent, pale to dark green, hairy. Seeds yellowish cream, 324-385 x

170-220 pm, ellipsoid, papillate or canaliculate.

Distribution. Southern, western and northern Thailand, Peninsular Malaysia (Kedah,

Perak and Perlis).

Habitat and ecology. On limestone cliffs or rock faces. On Langkawi on limestone by

the sea and, inland, at cave mouths or on rich damp or thinner soil layers at base of

limestone cliffs.

Provisional IUCN conservation status. Vulnerable VU Blab(iii) in Peninsular

Malaysia. In Peninsular Malaysia Microchirita rupestris has a patchy distribution in

Kedah (Gunung Baling and some parts of Langkawi), Perak (Waterloo Estate) and

Perlis, but none lie within the network of Totally Protected Areas. It is vulnerable

because it inhabits limestone cliffs and rock faces, cave mouths, rich damp soil at

the base of limestone hills, or, in Langkawi, co mm only on limestone by the sea that

is sensitive to disturbance. Although limestone is quarried on the main island of

Microchirita in Peninsular Malaysia

Langkawi, populations of this species are safe from quarrying on the smaller islands.

However, there also other threats, such as from environmental destruction caused by

tourism infrastructure and other development activities. The major threats on Gunung

Baling are also from quarrying. In neighbouring Thailand this species is common and

widespread (Middleton, pers. comm.).

Additional specimens examined. PENINSULAR MALAYSIA: Kedah: Gn. Baling, 25

Nov 1941, Comer s.n. (SING); ibidem, 1 Nov 2008, Imin LRI 63180 (KEP); Gn. Keriang,

6 Nov 2009, Rafidah LRI 64419 (KEP); Langkawi, Nov 1941, Corner s.n. (SING); ibidem,

1979, Stone 14294 (KLU); Langkawi, Gua Cerita, 4 Nov 2009, Rafidah LRI 64399 (KEP);

Langkawi, P. Langgun, 4 Nov 2009, Rafidah FR164397 (KEP); ibidem, 6 Nov 2009, Rafidah

FR164419 (KEP); ibidem, Feb 1989, Tay 149 (SING); Langkawi, Selat Panchor Air, Ahmad

Saktian Langgang Cl20 (KLU). Perlis: Bkt. Lagi, 16 Nov 1929, Henderson SFN 22816

(SING); ibidem, 16 Nov 1929, Henderson SFN 22848 (SING); Gua Cenderawasih, 25 Nov

1929, Haniff 7559 (SING); ibidem, 9 Jan 2007, Phoon FRI 53210 (KEP, SING); Wang Kelian,

29 Nov 2000, Kiew RK 5149 (KEP, SING).

Notes. Microchirita rupestris is not commonly found in Malaysia and has a narrow

distribution, being found only west of the Main Range. The population from Gunung

Baling, Kedah, was recorded with whitish flowers. Chirita glasgovii Ridl. is based

on material collected at Waterloo, Perak but the type material is not distinct from

Microchirita rupestris. An attempt was made to re-collect the species from a limestone

area close to the type locality to confirm this decision but this was unsuccessful.

4. Microchirita ruthiae Rafidah, Nordic J. Bot. 31: 612 (2013). - TYPE: Peninsular

Malaysia, Kelantan, Jeli, Gunung Reng, 24 October 2009, Rafidah FRI 64385 (holotype

KEP; isotypes E, SING). (Fig. 2B-H, 3G-H, 8)

Caulescent, herbaceous, branched plant. Stems pale green or whitish green, flowering

at c. 10 cm tall, erect or sub-erect to 30 cm, 6-8 mm diam., sparsely to densely hairy,

internodes 3.5-6.5 cm long, hairs eglandular, whitish. Leaves opposite, decussate;

petiole pale green, 5-15 mm long, hairs eglandular; lamina ovate or elliptic, 6.5-

14.5 x 6-6.5 cm, dark green, yellowish green above, pale green to whitish beneath,

thinly coriaceous, pubescent, base cuneate, sometimes unequal, margin serrate, apex

acute; midrib sunken above, prominent beneath, lateral veins 6-7 pairs on each side

of the midrib, hairy, intercostal veins reticulate. Inflorescence epiphyllous (axillary),

1^1-flowered, flowering from petiole base; bracts absent; pedicels very pale green,

shortly fused to the petiole, to 5-15 mm long, glandular hairy. Calyx pale green at

base, dark green towards tip, clasping the corolla tube, lobes 8-10 mm long, 1-2 mm

wide, apex acute, margin entire, hairy, outer surface glandular and eglandular hairy.

Corolla tube very sparsely hairy outside, whitish, pale green, or sometimes dull purple,

2-2.5 cm long, slightly curved, throat very pale green to whitish green, glandular

hairs above the anthers, glistening; lobes faintly striped, whitish, sometimes very pale

Card. Bull. Singapore 69(1) 2017

Selected Localities

1 Gn Reng

2. GuaSetir

3. Chiku 8

4. Gua Musang

5 Gua Maka

Extent of Occurrence (EOO)

3,186 sq km

Area of Occupancy (AOO):

24 sq km

Forest cover within EOO :

46 %

(£3

O-

■O

■t

□

tJJ

■o

1 : 3,500,000

50 Kilometers

NFl III Courtesy of Forestry Department Peninsular Malaysia

100 * 00 '

101 * 00 '

loroo'

102 * 00 '

102 * 00 ''

103 * 00 '

104 * 00 ’

104 * 00 '

Location based on :

Herbarium specimen(s)

Forest cover as in National

Forest Inventory III (1991-1993)

Fig. 8. Distribution of Microchirita ruthiae Rafidah in Peninsular Malaysia.

Microchirita in Peninsular Malaysia

purple, upper lobes rounded, spreading, slightly cleft at apex of each lobe, 4-5 x 5-7

mm, lower lateral lobes rounded, 4-5 x 5-7 mm, lower central lobe elliptic, elongated,

6-8 x 8-10 mm. Stamens, filaments pale green, inserted at 6-8 mm from the base of

the corolla, 3-4 mm long, straight, glabrous; anther-thecae divergent, 1-2 mm long,

yellow, not joined, with long reflexed hairs on the lower surface; staminodes 2 (rarely

3), pale green, inserted at 6 mm from the base of the corolla tube, 1.5-2 mm long.

Nectary pale yellow or cream, less than 1 mm deep. Pistil pale green; ovary 1-1.4 cm

long, c. 1.5 mm wide at the base, narrowing to 0.5 mm below the stigma, glandular and

eglandular hairy; style 6-8 mm long; stigma deeply 2-lobed, c. 2 mm long, 4 mm wide,

inserted between the anthers, very fine densely papillose hairy; ovules cream, less than

1 mm long. Capsule green, 3-5.5 cm long, 2-3 mm wide, slender, erect, densely hairy;

calyx persistent, pale green, hairy. Seeds yellowish cream, 325^140 x 190-220 pm,

broadly ovate or elliptic, slightly elongate with papillate surface.

Distribution. Endemic in Peninsular Malaysia: Kelantan.

Habitat and ecology. On limestone cliffs, crevices or soil pockets at base of limestone.

Some plants grow in places directly exposed to sunlight but most grow in shaded

habitats.

Provisional IUCN conservation status. Endangered EN Blab(iii). None of the six

Kelantan limestone hills where this species is found lie within the network of Totally

Protected Areas. The species is prone to disturbance because it grows on limestone

cliffs, in crevices or in soil pockets at the base of hills and the surrounding areas are

plantations with associated disturbance and microclimate changes. On Gunung Reng it

is additionally disturbed by recreational activities. None of these localities is protected

by a buffer zone.

Additional specimens examined. PENINSULAR MALAYSIA: Kelantan: Chiku 8, 24 May

1990, Kiew RK 3112 (KEP, SING); Gua Musang, Batu Serai, 17 May 1990, Kiew RK 2996

(SING); Gn. Reng, 18 May 1990, Kiew RK 2008 (KEP); ibidem, 14 Jan 2010, Rafidah FRI

64448 (KEP); ibidem, 7 Aug 1984, Weber UPM 3453 (KEP); Gua Maka, 19 May 1990, Kiew

RK 3022 (KEP); Gua Setir, 13 Jan 2010, Rafidah FRI 64441 (KEP).

Notes. Microchirita ruthiae occurs only east of the Main Range. Sterile material of

Microchirita ruthiae superficially looks similar to M. sericea.

5. Microchirita sericea (Ridl.) A.Weber & Rafidah, Taxon 60: 779 (2011). - Chirita

sericea Ridl., J. Straits Branch Roy. Asiat. Soc. 43: 59 (1905); Ridley, J. Asiat. Soc.

Bengal 74: 764 (1908); Ridley, FI. Malay Penins. 2: 525 (1923); Wood, Notes Roy.

Bot. Gard. Edinburgh 33: 187 (1974). - TYPE: Malaysia, Peninsular Malaysia, Perak,

Ipoh, August 1898, Curtis 3131 (lectotype SING [SING0042991], designated here;

isolectotype SING [SING0042992]). (Fig. 2H, 3I-J, 9)

Card. Bull. Singapore 69(1) 2017

Selected Localities

1. Bkt. Keteri

2 Gn. Baling

3. Tambun

4. ipoh

5. K. Dipang FR.

Extent of Occurrence (EOO) :

2,569 sq km

Area of Occupancy (AOO):

24 sq km

Forest cover within EOO :

51 %

50 Kilometers

NFS ill Courtesy of Forestry Department Peninsular Malaysia

1^.

O '

-o

C'i

■O

-o

Location based on :

SINGAPORE

IQCTQQ 1

10QW

loroc

iot v oo <

i o2;oo’

102W

103°G0‘

c 00‘

104°00'

104*00'

1 : 3 , 500,000

• Herbarium specimen(s)

p Forest cover as in National

Forest Inventory III (1991-1993)

Fig. 9. Distribution of Microchirita sericea (Ridl.) A.Weber & Rafidah in Peninsular Malaysia.

Microchirita in Peninsular Malaysia

Chirita sericea var. scortechinii Ridl., J. Straits Branch Roy. Asiat. Soc. 43: 59 (1905).

-TYPE: Peninsular Malaysia, Perak, 1884, Scortechini 112b (lectotype K, designated

by Wood (1974); isolectotype SING).

Herbaceous, apparently perennial plant. Stems, leaves and inflorescence densely hairy

with eglandular white or downy hairs. Stems woody, pale green, sometimes reddish

green, creeping to 30 cm long, internodes to 5.5 cm long. Leaves opposite; petiole

pale green, to 2.2 cm long; lamina narrowly elliptic or narrowly ovate, 4-11 x 1M cm,

pale green above, very pale green or whitish green beneath, coriaceous when fresh and

dry, densely sericeous hairs on both surfaces, base attenuate, or sometimes cordate or

unequal, margin serrulate, apex acuminate or acute; midrib sunken above, prominent

beneath, lateral veins 5-10 pairs on each side of the midrib, intercostal veins reticulate.

Inflorescences epiphyllous, 1-4-flowered; bracts absent; pedicels pale green, shortly

fused to the short petiole, 2-3 cm long. Calyx pale green, clasping the corolla tube,

lobes narrowly triangular, 16-20 mm long, 2-3 mm wide, outer surface densely

hairy. Corolla hairy outside, tube pale blue, lilac or white, to 2.6 cm long, narrowly

tubular, slightly curved, not pouched, throat yellow, glandular hairs translucent brown,

positioned above the anthers; lobes conspicuously striped, upper lobes rounded, not

spreading, 3.5-5 x 5-6 mm, lower lateral lobes elliptic, 5-6 x 5-7 mm, lower central

lobe elliptic, elongated, 6.5-8 x 8-10 mm. Stamens : filaments brown, inserted 1.1-

E4 cm from the base of the corolla, 4-8 mm long, slightly geniculate proximally;

anthers very pale green, sometimes brown, anther-thecae divergent, 2-3 mm long,

fused apically, with reflexed hairs on the lower surface; staminodes 2, cream, inserted

1.1-1.4 cm from the base of the corolla tube, to 2 mm long. Nectary pale green or

cream, 2-3 mm long, margin entire. Pistil pale green, densely eglandular hairy; ovary

15-20 x 1-2 mm; style 1-1.2 cm long; stigma slightly 2-lobed, heart-shaped, to 2 mm

long, abaxially velvety, adaxial with very fine papillose hairs. Capsule whitish green,

to 7 cm long, cylindric slender, slightly curved; calyx persistent, green, hairy. Seeds

yellowish cream, many in one row, 300-380 x 120-250 pm, broadly elliptic, rounded,

surface papillate or canaliculate.

Distribution. Endemic to Peninsular Malaysia (Perak, Perlis and Kedah).

Habitat & ecology. On limestone cliffs or rocks, in dry places at the foot of limestone

cliffs, often under overhangs or near caves.

Provisional IUCNconservation status. Endangered ENB lab(iii). This species does not

grow within the network of Totally Protected Areas and has a narrow distribution. The

populations are restricted to a few hills in Perak, i.e. Kuala Dipang, Ipoh limestone and

one record from Gunung Baling, Kedah. The last collection from Gunung Baling was

made in 1941 and recent field trips did not re-locate it from that area. Part of Gunung

Baling is currently disturbed by quarrying activity. In Perak, no karst hill is protected

and many hills are actively being quarried or the flora is being degraded by resort or

recreational development or through activities related to temples (Kiew et al., 2011).

Card. Bull. Singapore 69(1) 2017

Additional specimens examined. PENINSULAR MALAYSIA: Kedah: Gn. Baling, 25 Nov

1941, Corner s.n. (SING). Perak: Gn. Rapat, 21 Jul 2009, Rafidah FRI64347 (KEP); ibidem,

26 May 2010, Rafidah FRI 64544 (KEP); Gn. Tasek, Perak Tong Temple, 21 Jul 2009, Rafidah

FRI 64348 (KEP); ibidem, 23 Oct 1958, Sinclair 9844 (SING); Ipoh, 4 Jul 1917, Burkill 2552

(SING); ibidem, Aug 1898, Corner s.n. (KEP); ibidem, Feb 1904, Ridley 11952 (SING);

ibidem, Feb 1904, Ridley s.n. (KEP); ibidem, 18 Aug 1986, Weber UPM 4167 (KEP); ibidem,

18 Aug 1986, Weber s.n. (KEP); Kuala Dipang F.R., 1898, Ridley s.n. (SING); Tambun, 10 Sep

1920, Burkill 6284 (SING); Sg. Siput Utara, 29 Jan 2015, Rafidah FRI 82017 (KEP). Perlis:

Bkt. Keteri, 28 Nov 2013, Rafidah FRI 75880 (KEP).

Notes. Microchirita sericea occurs west of the Main Range, replacing M. rupestris in

the south.

6. Microchirita viola (Ridl.) A.Weber & Rafidah, Taxon 60: 779 (2011). - Chirita

viola Ridl., J. Linn. Soc., Bot. 32: 516 (1896); Ridley, J. Straits Branch Roy. Asiat. Soc.

43: 58 (1905); Ridley, FI. Malay Penins. 2: 525 (1923); Henderson, Malay. Wild FIs.,

Dicots. 350 (1959); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 190 (1974). - TYPE:

Malaysia, Peninsular Malaysia, Kedah, Langlcawi, Sep 1890, Curtis 2570 (lectotype

SING [SING0042993], designated here; isolectotypes SING [SING0042994,

SING0042995]). (Fig. 2H, 3K-L, 10)

Chirita hamosa auct. non R.Br.: Wood, Notes Roy. Bot. Gard. Edinburgh 33: 191

(1974), pro parte.

Short-lived, caulescent, branching herbs. Stems reddish or pale green, beginning to

flower at the one-leaf stage, to 30 cm tall, 5-7 mm diam., intemodes swollen, to 10 cm

long, finely hairy. Leaves opposite, decussate; petiole pale green, sometimes reddish,

to 6 mm long, with glandular hairs; lamina ovate, sometimes orbicular, 3-8.5 x 2-5.5

cm, yellowish green to pale green above, whitish green or reddish beneath, coriaceous

when fresh and dry, sparsely strigose hairy on both surfaces, base slightly cordate or

rounded, margin serrate, apex acute; midrib sunken above, prominent beneath, lateral

veins 6-16 pairs on each side of the midrib, intercostal veins reticulate. Inflorescence

crested epiphyllous, up to 3 2-paired flowers; bracts absent; pedicels fused to the

petioles, green or reddish, to 1 cm long, glandular and eglandular hairy. Calyx green or

reddish green, lobes narrowly ovate, 7-10 mm long, c. 1 mm wide, outer surface ciliate,

inner surface glabrous. Corolla tube pale violet, sparsely hairy outside, 2-2.2 cm long,

curved downwards, throat very pale purple, glandular bead-lilce golden yellow hairs, to

1.5 mm long, apically swollen in a cluster in the throat above the anthers; lobes violet,

each with conspicuous dark purple stripes, upper lobes rounded, slightly imbricate,

not spreading, 4-5 x 3.5-5 mm, lower lateral lobes elliptic, spreading, 5-6 x 4.5-6

mm, lower central lobe elliptic, elongated, curved downwards, 6.5-10 x 4.5-6 mm.

Stamens, filaments white, sometimes reddish at base, inserted at 5-8 mm from the

base of the corolla, 1.8-2 mm long, slightly geniculate at the point of attachment;

Microchirita in Peninsular Malaysia

Selected Local fifes

1* Tg. Rhu

2. Kuah

3. Selat Panchor F.R

4. 8kt. Malut F.R.

5. Gn, Keriang

Extent of Occurrence (EOO) :

119 sq km

Area of Occupancy (AOO):

32 sq km

Forest cover within EOO :

57 %

1 ; 3,500,000

□

Location based on

• Herbarium specimens)

|—] Forest cover as in National

^ Forest Inventory HI (1991-1993)

O-

□

100 00 '

ioroo’

102 ° 00 ’

WOO 1

-j

tji

50 Kilometers

NFI III Courtesy of Forestry Department Peninsular Malaysia

SINGAPORE

102W I03 n 00'

104 , 00‘

104 a 00'

O-

Fig. 10. Distribution of Microchirita viola (Ridl.) A.Weber & Rafidah in Peninsular Malaysia.

Card. Bull. Singapore 69(1) 2017

anthers pale yellow with purple markings, anther-thecae joined apically, 1.5-1.7 mm

long; staminodes 2 or 3, unequal in size, white, inserted 5-8 mm above the base of

the corolla, 1.3-2.2 mm long. Nectary cream, to 1.1 mm long. Pistil very pale green;

ovary 4-5 mm long, 0.5 mm wide at the base, narrowing to 2 mm below the stigma,

eglandular hairy; style to 4 mm long; stigma pale yellow, shallowly 2-lobed, c. 2 mm

long, very fine glandular hairy; ovules cream, less than 1 mm long. Capsule pale

green or reddish green, to 5 cm long, cylindric and slender, sparsely hairy, sometimes

glabrous; calyx persistent, reddish green, hairy. Seeds yellowish cream, many in one

row, 600-710 x 200-280 pm, very narrowly ellipsoid, surface rounded papillate.

Distribution. Peninsular Thailand and Peninsular Malaysia (Kedah: Langkawi and

Gunung Keriang).

Habitat and ecology. On limestone cliffs or crevices with a thin soil layer or, on

Langkawi, on limestone rocks near the sea.

Provisional IUCN conservation status. Near Threatened (NT). The species is common

on Langkawi Island, much of which is protected. Its Near Threatened status is based

on its localised and patchy distribution on Langkawi Island. The species flowers

abundantly from October to December in the wettest period. Previously it was recorded

in Gunung Keriang on the Kedah mainland but it has not been recollected in spite of

recent botanical collecting there.

Additional specimens examined. PENINSULAR MALAYSIA: Kedah: Gn. Keriang, Peb

1890, Allen s.n. (SING); Langkawi, 20 Nov 1941, Corner s.n. (SING); Langkawi, Ayer Hangat,

Chung 505 (KLU); Langkawi, Bkt. Malut P.R., 4 Nov 1968, Keng 80 (SING); Langkawi,

Batu Puteh, Aug 1941, Nauen 38120 (SING); Langkawi, Kuah, 8 Nov 1968, Chung RC 7

(KEP); ibidem, 5 Nov 2009, Rafidah FRI 64407 (KEP); ibidem, 1979, Stone 14349 (KLU);

Langkawi, P. Langgun, 4 Nov 2009, Rafidah FRI 64398 (KEP); Langkawi, P. Timun, 1926,

Holttum 17433 (SING); Langkawi, Selat Panchor L.R., 19 Nov 1941, Corner 37832 (SING);

ibidem, 21 Nov 1934, Henderson SFN 28931 (SING); ibidem, Nov 1934, Henderson SFN

29185 (SING); ibidem, 3 Nov 2009, Rafidah PRI 64388 (KEP); Langkawi, Tg. Rhu, 21 Nov

1993, Anthonysamy SA 1144 (KEP, SING); Langkawi, Tg. Sawah, 22 Nov 1941, Corner s.n.

(SING); Langkawi, Tg. Terai, 13 Nov 1941, Corner s.n. (SING).

ACKNOWLEDGEMENTS. This research was carried out as part of the flora of Peninsular

Malaysia Project (Project no. 01-04-01-0000 Khas 2), funded by the Ministry of Science,

Technology and Innovation (MOSTI), and ‘Documentation & Inventory of flora of Malaysia

Project’, both at the forest Research Institute Malaysia (PRIM), Kepong, the ‘Systematic

Studies of Selected Taxa of Plants Project’ ( Erycibe [Convolvulaceae], Chirita , Henckelia

section Boeopsis, Loxocarpus and Salicini [Gesneriaceae] and Utricularia [Lentibulariaceae]

in Peninsular Malaysia) (Research and Pre-commercialisation grant, GPP-TPBC-1208-001) at

PRIM, and a Postgraduate Research (IPPP) grant (Project no. PS171/2008B) at the University

of Malaya (UM), Kuala Lumpur. I am grateful to the curators and keepers of the herbaria at

BK, BKP, BO, E, K, KEP, KLU, SAN, SAR, SING and UKMB for permission to examine

Microchirita in Peninsular Malaysia

specimens in their care; to Dr Ruth Kiewand Dr Noorma Wati Haron (supervisor in Master of

Science) for their guidance; to the Flora Biodiversity Programme staff for their assistance in

the held; to Dr Saw Leng Guan and Dr Richard Chung for giving comments on this paper; and

Dr Lillian Chua for checking the conservation status assessment. I also thank Mr Roslee Halip

(University of Malaya) and Mr. Azizi Abdul Jalil (FRIM) for assisting in SEM techniques.

References

Burtt, B.L. (1954). Studies in the Gesneriaceae of the world: Types and lectotypes of certain

genera and groups of lower rank. Notes Roy. Bot. Gard. Edinburgh 21: 193-208.

Chua, L.S.L. (2012). Conservation. In: Kiew, R., Chung, R.C.K., Saw, L.G. & Soepadmo, E.

(eds) Flora of Peninsular Malaysia Series II, vol. 3, pp.3-10. Kepong: Forest Research

Institute Malaysia.

Chua, L.S.L. & Saw, L.G. (2006). Malaysia Plant Red List: Guide for contributors. Kepong:

Forest Research Institute Malaysia.

Chua, L.S.L., Kiew, R. & Chan, Y.M. (2009). Assessing conservation status of Peninsular

Malaysian Begonias. Blumea 54: 94-98.

Davis, S.D., Heywood, V.H. & Hamilton, A.C. (1995a). In: Davis, S.D., Heywood, V.H. &

Hamilton, A.C. (eds) Limestone Lora of Peninsular Malaysia. In: Centres of Plant

Diversity, a Guide and Strategy for Their Conservation, vol. 2. Asia. Australasia and the

Pacific, pp. 303-307. U.K., Cambridge: IUCN Publication Unit.

Davis, S.D., Heywood, V.H. & Hamilton, A.C. (1995b). In: Davis, S.D., Heywood, V.H. &

Hamilton, A.C. (eds) Limestone Lora of Borneo. In: Centres of Plant Diversity, a Guide

and Strategy for Their Conservation, vol. 2. Asia. Australasia and the Pacific, pp. 332-

336. U.K., Cambridge: IUCN Publication Unit.

Henderson, M.R. (1939). The Lora of limestone hills of the Malay Peninsula. /. Malayan

Branch Roy. Asiat. Soc. 17: 13—87.

IUCN (2012). The IUCN Red List Categories and Criteria. Version 3.1 (2nd edition). Gland:

IUCN Species Survival Commission, IUCN.

Kiew, R. (1991). The Limestone Flora. In: Kiew (ed) The State of Nature Conservation in

Malaysia, pp. 42-50. Kuala Lumpur: Malayan Nature Society.

Kiew, R. (2004). The limestone Lora of Sarawak. In: Yong, H.S., Ng, F.S.P. & Lee, E.Y.E.

(eds) Sarawak Bau limestone biodiversity, pp. 79-89. Sarawak Mus. J. 59(80), Special

issue 6. Kuching: Sarawak Museum Department.

Kiew, R. (2009). The natural history of Malaysian Gesneriaceae. Malayan Nat. J. 61(3):

257-265.

Kiew, R., Ummul-Nazrah, A.R. & Chua, L.S.L. (2011). Conservation status of Paraboea

species (Gesneriaceae) in Malaysia. Gard. Bull. Singapore 63(1&2): 433^150.

Middleton, D.J. & Triboun, P. (2013). New species of Microchirita (Gesneriaceae) from

Thailand. Thai Forest Bull, Bot. 41: 13-22.

Moller, M., Pfosser, M., Jandg, C.G., Mayer, V., Clark, A., Hollingsworth, M.L., Barfuss,

M.J.H., Wang, Y.Z., Kiehn, M. & Weber, A. (2009). A preliminary phylogeny of the

‘Didymocarpoid Gesneriaceae’ based on three molecular data sets: Incongruence with

available tribal classifications. Amer. J. Bot. 96(5): 989-1010.

Price, L. (2001). Caves and karst of Peninsular Malaysia. Kuala Lumpur: Gua Publications.

Puglisi, C., Middleton, D.J, & Suddee, S. (2016). Four new species of Microchirita

(Gesneriaceae) from Thailand. Kew Bull. 71: 2-7.

Card. Bull. Singapore 69(1) 2017

Rafidah, A.R. (2011). Revision ofChirita (Gesneriaceae) for Peninsular Malaysia. Unpublished

MSc thesis, University of Malaya, Kuala Lumpur, Malaysia.

Ramli, M.O & Meor Abdul Rahman, M.T. (2014). Estimation of Sub-Surface Limestone

Reserve Under Idle Ex-Mining Land in Perak. In: Taib, N.I. (ed) Proceedings of the

National Geoscience Conference 2014, pp. 183-192. Kuala Lumpur: Geological

Society of Malaysia.

Ridley, H.N. (1905). The Gesneraceae of the Malay Peninsula. J. Straits Branch Roy. Asiat.

Soc. 43: 1-92.

Ridley, H.N. (1908). Gesneriaceae. In: King, G. & Gamble, J.S. (eds) Materials for a Plora of

the Malayan Peninsula. J. Asiat. Soc. Bengal 74: 729-908.

Ridley, H.N. (1923). Gesneriaceae. In: The Flora of the Malay Peninsula, vol. 2: 495-505.

London: L. Reeve & Co.

Saw, L.G., Chua, L.S.L. & Abdul Rahim, N. (2009). Malaysia National Strategy for Plant

Conservation. Malaysia: Ministry of Natural Resources and the Environment & Lorest

Research Institute Malaysia.

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria

and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.

nybg.org/science/ih/. Accessed 22 Nov. 2016.

Wang, Y.Z., Mao, R.B., Liu, Y., Li, J.M., Dong, Y., Li, Z.Y. & Smith, J.P. (2011). Phylogenetic

reconstruction of Chirita and allies (Gesneriaceae) with taxonomic treatments. J. Syst.

Evol. 49, 1: 50-64.

Weber, A., Middleton, D.J., Porrest, A., Kiew, R., Lim, C.L., Rafidah, A.R., Yao, T.L. & Moller,

M. (2011). Molecular systematics and remodeling of Chirita and associated genera

(Gesneriaceae). Taxon 60: 767-790.

Wood, D. (1974) A revision of Chirita (Gesneriaceae). Notes Roy. Bot. Gard. Edinburgh 33:

123-205.

Appendix 1. Specimens Examined. Specimens are arranged by collector (with Erst initial,

when known) in alphabetical order, followed by collector’s number in numerical order (s.n. =

without number), followed by the taxon number in the text.

Ahmad Saktian Langgang 120 (3); Allen, M. s.n. (6); Anthonysamy, S. 379 (1), 842 (1), 1144

( 6 ).

Burkill, I.H. 1493 (2), 2253 (1), 2552 (5), 2558 (1), 6284 (5); Burtt, B.L. 1665 (1).

Chew, M.Y. 58699 (2); Chin, S.C. 331 (1), 390 (1), 505 (6), 2121 (6); Chung, R.C.K. 7 (6);

Corner, E.J.H. 37832 (6), s.n. (2), s.n. (3), s.n. (5), s.n. (6); Curtis, C. 2120 (3), 2359

(1) , 2570 (6), 3109 (1), 3131 (5), 3221 (2), s.n. (1).

Davison, G.W.H. 4 (1); Dawn, J. 1470 (1), 1472 (2), 1474 (2); Ding, H. 715 (1).

Evans, s.n. (1).

Gordon, H.S. 435 (1).

Hardial, S. 477 (1); Henderson, M.R. 19403 (2), 22419 (1), 22816 (3), 22848 (3), 25033 (1),

25223 (1), 25233 (1), 28931 (6), 29185 (6), s.n. (1); Holttum, R.E. 17433 (6).

Imin, K. 58583 (2), 63180 (3), 63212 (1), 71983 (2).

Julia, S. 26 (1).

Kamarul Hisham, M. 67229 (1); Kehding in Beccari 73 (1); Kelsall, H. s.n. (1); Keng, H. 79

(2) , 80 (6); Kiew, R. 1023 (1), 1202 (1), 1404 (2), 1424 (2), 1491 (2), 1505 (2), 1537

(2), 1557 (1), 1571 (\),1587 (1), 2008 (4), 2158 (1), 2284 (1), 2339 (1), 2524 (1), 2885

Microchirita in Peninsular Malaysia

(2), 2896 (2), 2920 (2), 2940 (2), 2948 (2), 2996 (4), 3022 (4), 3035 (2), 3112 (4), 3973

(2), 4915 (2), 5149 (3), 5253,4 (2), 65514 (1), 70496A (1), 70509 (1), s.n. (1); King’s

collector. 937 (1), 5028 (1), 7028 (1).

Lobb (1).

Maxwell, J.F. 78-131 (1); Mills, G.R. 15061 (1); Mohd Hairul, M.A. 54065 (3), 60062 (2),

72402 (3), 72449 (2); Mohd. Haniff. 7559 (3); Mohd. Kasim, R. 391 (1); Mohd. Nur

12130 (2), 34389 (1).

Nauen, J.C. 38120 (6); Ng, F.S.P. 1629 (1).

Phoon , S2V. 51570 (1), 53210 (3).

Rafidah, A.R. 55687 (2), 55713 (1), 55714 (1), 55717 (1), 55721 (1), 55724 (1), 55726 (1),

55728 (2), 64314 (2), 64319 (2), 64346 (5), 54347 (1), 64348 (5), 54349 (5), 54379 (1),

64385 (4), 54355 (6), 64394 (2), 64395 (2), 64397 (3), 64398 (6), 64399 (3), 64407 (6),

54477(1), 54475(2), 54479(3), 54425(2), 54447 (4), 54447(2), 54544(5), 54545(1),

54552 (1), 54505 (1), 54535 (1), 52005 (1), 52077 (4); Tftd/ey, 774V. 77952 (5), j./i. (1),

s.n. (2), s.rc. (5); Robertson-Glasgow s.n.( 3).

Sam, Y.Y. 50118 (1); Samsuri, A. 560 (1); Saw, L.G. 36215 (1); Scortechini, B. 112b (5), 1582

(1); Sinclair, J. 9844 (5), 10732 (1), 40055 (1); Smith, J.W. 85205 (1); Stone, B.C.

8934 (1), 8794 (1), 14294 (3), 14349 (6); Syahida-Emiza, S. 66733 (1); Symington, C.F.

30796(1), 37431 (1), 39598 ( 1).

Fay, E.P. 149 (3), 755 (2).

UNESCO. 268 (2).

Weber, A. 3453 (4), 4167 (5), 540507 (4).

Yap, S.K. 25(1).

Gardens’ Bulletin Singapore 69(1): 33-65. 2017

doi: 10.3850/S2010098116000039

Index of names and types of Hoya

(Apocynaceae: Asclepiadoideae) of Borneo

M. Rodda

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

rodda.michele@gmail.com

ABSTRACT. Types of all Hoya species occurring in Borneo and their synonyms are indicated

and clarified. Forty-six lectotypes, nine neotypes and five epitypes are designated.

Keywords . Brunei, epitype, holotype, Kalimantan, lectotype, neotype, Sabah, Sarawak

Introduction

The present paper is a precursor to a revision of Hoya R.Br. of Borneo. Its aim is to

list all Hoya taxa occurring on the island of Borneo and their synonyms, clarify and

standardize type citation, select lectotypes, epitypes and neotypes when necessary and

provide an extensive list of isotypes and syntypes.

The earliest description of a Hoya species that occurs in Borneo, H. multiflora

Blume, was described by the German born Dutch botanist Carl Ludwig Blume in 1823,

based, however, on Javanese materials. The first Hoya species based on a Bornean

collection, H. imperialis Lindl., was published by Lindley (1846). Later, in 1880,

Bentham published a peculiar leafless species with long photosynthetic peduncles

endemic to Borneo, Astrostemma spartioides Benth. (now Hoya spartioides (Benth.)

Kloppenb.). Until 1921 only nine Hoya species where known to occur in Borneo

(Merrill, 1921). Nutt (2001) listed 21 species in an unpublished checklist. Lamb et al.

(2014) estimated 60-70 species for Sabah alone. In the past 20 years almost 40 taxa

(species and subspecies) of Hoya from Borneo have been described, mostly by Ted

Green and Dale Kloppenburg (USA). The most recently published taxa have been

described predominantly based on collections from Sabah and Sarawak (Rodda &

Simonsson, 2011a, 2011b; Rodda & Simonsson Juhonewe, 2013a; Lamb et al., 2014;

Rodda et al., 2014, 2016; Rodda, 2015a; Lamb & Rodda, 2016) with only one from

Kalimantan (Rahayu et al., 2015).

A guidebook to Hoya R.Br. of Borneo has been recently published (Lamb &

Rodda, 2016) and includes a checklist of Bornean Hoya that lists 72 taxa (71 species and

one subspecies) but does not deal with typification of names. The current bibliographic

checklist of names is intended as a precursor to a formal taxonomic revision of the

genus for Borneo.

Card. Bull. Singapore 69(1) 2017

Materials and methods

The list published here includes accepted names, indicated in bold, of all species

occurring in Borneo following the guide book checklist by Lamb & Rodda (2016:

193-197). All taxa are listed regardless of where the type was collected. Under each

accepted name all known synonyms are listed, again regardless of where the type

was collected. The protologue of each name has been examined to establish which

original materials were listed and needed to be located. Literature was obtained on the

Biodiversity Heritage Library website [http://www.biodiversitylibrary.org], JSTOR

[http://www.jstor.org/] and in the libraries of the Singapore Botanic Gardens, Royal

Botanic Gardens Kew and Royal Botanic Garden Edinburgh.

Types have been located in person or via loans at A, BISH, BM, BO, BRUN,

CGE, E, FI, G, HITBC, HN, IBSC, K, KEP, KUN, L, OXF, P, SAN, SAR, SING, SNP,

TO, UC, US, VN, WRSL, Z and on JSTOR Global Plants (https://plants.jstor.org/

accessed on 28 July 2016).

If a suitable specimen is available, a lectotype is selected for names where the

protologue does not explicitly mention a single type specimen with a direct reference

to the institution in which it was deposited, strictly applying Art 9.1 & 9.2 of the

ICN (McNeill et al., 2012). A reference to a single specimen indicated as ‘type’ or

‘holotype’ in treatments published before 1 January 2001 is considered as effective

lectotypification under Art. 9.9 of the ICN (McNeill et al., 2012). Schlechter’s

specimens in B are usually considered to be holotypes (Nicholas, 1992). However,

for Hoya names this is incorrect under ICN Art. 9.1 (McNeill et al., 2012), as the

herbarium where the specimens are deposited is not clearly indicated by Schlechter

and it cannot be ascertained that only a single specimen of the type collection was

consulted. I have therefore selected lectotypes whenever original material could be

located.

Noltie (2005) clarified the type citations of taxa published by Wight. His

notation of types is followed here. When based on specimens from Wallich the sheets

usually bear a Wallich Asclepiadaceae number (Noltie, 2005), a number assigned to

sheets given to Wight from Wallich before Wallich numbers were assigned. These

sheets, listed in Linnean Society Manuscript SP1284 became part of Wight’s personal

working herbarium and bear an HRWP (Herbarium Robert Wight Proper) label and

may also bear a Wight number that is often a species number, an ‘Asclep’ number and

the later assigned Wallich number. Sheets from Wight’s personal working herbarium

are to be considered as Wight’s ‘top set’ (Noltie, 2005: 133), they are often annotated

by Wight himself and may have pencilled sketches of the flowers. These specimens

are here preferred over other duplicates in the selection of lectotypes. For taxa where

a single sheet of a taxon was mentioned in Linn Soc Mss SP1284, if the sheet is extant

and can be easily identified, it is considered a holotype.

Names and types of Bornean Hoya

Typification of recently published names

Twenty new Hoya names listed here have been published since 1995 in Fraterna,

the bulletin of the International Hoya Society, and two (Hoya ranauensis T.Green &

Kloppenb. and Hoya amoena Bakh.f. subsp. bogorensis T.Green & Kloppenb.) in the

online journal Hoya New (http://www.rare-hoyas.com/publication.htm). As already

mentioned by Forster (1991) in relation to taxa from the Philippines published by the

same authors, their style of type citation may be confusing as it often includes only a

number and the herbarium where the holotype is deposited. Frequently not explicitly

referred to a collector, this number may be a sheet number, a collector number or the

collection date of the specimen (see Rodda, 2015b). Without a full citation of the

holotype, simple citation of a number may refer to multiple specimens belonging to

different collectors. This is not strictly needed for valid publication but recommended

by ICN Art. 40A.3. (McNeill et al., 2012). I have therefore examined all holotypes or

specimens labelled as such in the herbaria where holotypes were deposited, matched

them against the protologue and provided a full citation that is corrected in case

mistakes were noted in the protologue. Herbarium acronyms followed by a number

are here interpreted as sheet number (e.g. UC102003 for Hoya kastbergii Kloppenb.)

but may potentially be dates as well (see below H. walliniana Kloppenb. & Nyhuus

and H. nyhuusiae Kloppenb.). Holotypes of six species have not been found. For three

names, Hoya clemensiorum T.Green, H. lambii T.Green and H. monetteae T.Green,

specimens made by the publishing author and labelled as neotypes by him have been

found, suggesting that the holotype is indeed missing or was never actually made.

The list of names

Hoya acicularis T.Green & Kloppenb., Fraterna 15(4): 7 (2002). - TYPE: Originally

from Malaysia, Sabah, Danum Valley, cultivated in USA, Hawaii, Oahu, Ka‘a‘awa,

Green Plant Research, vouchered on 24 January 1991 as Green, T. 93031 (holotype

BISH [BISH1000875]).

Hoya aeschynanthoides Schltr., Bot. Jahrb. Syst. 40 (Beibl. 92): 13 (1908). - TYPE:

Indonesia, Borneo, Koetei, auf Baumen am Long Dett, August 1901, Schlechter, R.

13550 (lectotype B [B100277178], designated here).

Notes. Schlechter (1908) mentioned only Schlechter 13550 for Hoya aeschynanthoides

Schltr. A single duplicate bearing buds and a drawing of dissected flower parts has

been found in B [B100277178] and it is therefore selected here as lectotype of H.

aeschynantho ides .

Hoya australis R.Br. ex J.Traill, Trans. Hort. Soc. London 7: 28 (1830); Hill, Telopea

3: 250 (1988); Smith, FI. Vit. Nova 4: 118 (1988); Forster & Liddle, Austrobaileya 3:

508 (1991); Liddle & Forster, Asklepios 102: 6 (2008). - TYPE: Australia, Queensland,

Cook District, Cape Grafton, 1770, Banks, J. & Solander, D. s.n. (lectotype BM

[BM001040528], designated by Hill (1988)).

Card. Bull. Singapore 69(1) 2017

Notes. Hoya australis R.Br. ex J.Traill is a widely variable species thought to occur

in Australia, Papuasia and Melanesia (Hill, 1988; Forster & Liddle, 1991; Liddle &

Forster, 2008). It is currently classified into seven subspecies based on vegetative

morphology and distribution (Liddle & Forster 2008; Kloppenburg, 2012). The

discovery of Hoya australis in Sabah (based on Jamirus, J. in Lamb , A. AL2500/2014,

SAN) considerably extends its distribution area. Preliminary molecular analysis

confirms that the Bornean collection belongs to Hoya australis but it is inconclusive

to its sub-specific ranking (Rodda, unpublished). This is the reason why synonymies

are not indicated here. For the most updated treatment of subspecies and synonymies

of Hoya australis see Liddle & Forster (2008).

The specimen here indicated as lectotype of Hoya australis has long been

considered a holotype (Hill, 1988; Smith, 1988; Forster & Liddle, 1991; Liddle &

Forster, 2008). The specimen is not a holotype as the protologue of Hoya australis is

not more specific than that the original material is from Australia in Brown’s herbarium.

It is therefore impossible to establish whether it was based on a single specimen only.

Hill (1988), in mentioning Banks , J. & Solander, D. s.n. (BM) as holotype of H.

australis , is here considered as an effective lectotypification following ICN Art. 9.9

(McNeill et al., 2012).

Hoya bakoensis Rodda, Phytokeys 53: 87 (2015) [21 July 2015]. - TYPE: Malaysia,

Sarawak, Bako National Park, 20 March 2015, Rodda, M. MR1042b (holotype SING;

isotypes SAR, KEP).

Hoya beccarii Rodda & Simonsson, Webbia 68: 13 (2013). - TYPE: Malaysia,

Sarawak, Matang, July 1866, Beccari, O. 1866 ( collezioni Beccari 6526a) (holotype

FI; isotype FI).

Hoya benchaii Gavrus, A.L.Lamb, Emoi & Gokusing, Sandakania 19: 18 (2014). -

TYPE: Originally from Malaysia, Sabah, east of Ranau, Telupid area, cultivated at

Kipandi Park, vouchered on 8 May 2011 as Chai, B. in Lamb, A. 2132/2011 (holotype

SAN; isotype SNP n.v.).

Hoya callistophylla T.Green, Fraterna 13(4): 2 (2000). - TYPE: Originally from

Malaysia, Sabah, Nabawan, cultivated in USA, Hawaii, Oahu, Ka‘a‘awa, vouchered

on unknown date as Green , T. 201 (holotype BISH [BISH1014775]).

Hoya caudata Hook.f., FI. Brit. India 4(10): 60 (1883); Rintz, Malayan Nat. J. 30:

484 (1978). - TYPE: Peninsular Malaysia, Malacca, Maingay, A.C. 1956 (Kew distr.

no. 1128) (lectotype K [K000895134], designated by Rintz (1978), first step, and

designated here, second step; isolectotypes K [K000895133], L [L0004315]).

Hoya crassifolia Ridl., J. Straits Branch Roy. Asiat. Soc. 61: 30 (1912), nom. illeg.,

non Hoya crassifolia (J.Jacq.) Haw. - Hoya caudata Hook.f. var. crassifolia Ridl., FI.

Malay Penins. 3: 395 (1923). - TYPE: Malaysia, Perak, Tapah, on the trees on the

Temoh Road, November 1908, Ridley, H.N. 14059 (lectotype SING [SING0059473],

designated here; isolectotype K [K001096330]).

Names and types of Bornean Hoya

Notes. Hooker indicated the type of Hoya caudata Hook.f. as Maingay 1128. This

is the K distribution number of the Maingay sheets that corresponds to Maingay

1956. The indication by Rintz (1978) of Maingay 1128 (K) as ‘type’ is an effective

lectotypification under Art. 9.9 of the ICN (McNeill et al., 2012). However in K there

are two sheets labelled Maingay (.Kew distr. no. 1128 ) and, therefore, a second-step

lectotypification is necessary. The duplicate with barcode K000895134 is a well-

preserved fertile specimen that also bears a pencilled sketch and is here selected as

lectotype of H. caudata.

Two duplicates have been found of the type of Hoya crassifolia Ridl. The SING

sheet [SING0059473] is a well-preserved fertile specimen and is here selected as

lectotype.

Hoya chewiorum A.L.Lamb, Gavras, Emoi & Gokusing, Sandakania 19: 25 (2014).

-TYPE: Originally from Malaysia, Sabah, Central Crocker Range, Sg. Kopongian on

the upper reaches of the Moyong River, cultivated at Kipandi Park, vouchered on 30

April 2009 as Gokusing, L. andLombika, J. in Lamb, A. AL1769/2009 (holotype SAN;

isotype SNP n.v.).

Hoya clemensiorum T.Green, Fraterna 14(3): 12 (2001). - TYPE: Originally from

Malaysia, Sabah, Poring, Rafflesia Centre, 16 September 1991, cultivated in USA,

Oahu, Ka‘a‘awa, Hawaii, reputedly vouchered on unknown date as Green, T. 91032

(holotype BISH, missing); Malaysia, Sabah, Mt. Kinabalu, Dallas, 8 Sepember 1931,

Clemens, J. & M.S. 26358 (neotype K, designated here; isoneotypes BM, BO, L).

Notes. The holotype of Hoya clemensiorum T.Green, Green 91032 dated 16

September 1991, cannot be found at BISH. In BISH there is instead a sterile specimen

with the same collector and number dated 09 June 2008 labelled as neotype of H.

clemensiorum , obtained in cultivation from a plant originally wild collected in Sabah

on 16 September 1991.1 have been unable to find the publication of a neotypification

and I therefore select Clemens 26358 (K) as the neotype of H. clemensiorum as it

matches the original description, is fertile, and is among one of the best preserved

specimens of H. clemensiorum collected by the Clemenses.

Hoya coriacea Blume, Bijdr. FI. Ned. Ind. 16: 1063 (1826); Rintz, Malayan Nat. J. 30:

495 (1978). -TYPE: Indonesia, Java, Salak [In fruticetis adpedemmontis Salak’], s.d.,

s.coll. (lectotype L [sheet number 898.168-117], designated here; possible isolectotype

P [P00639838]).

Hoya brunoniana Wight, Contr. Bot. India 37 (1834). - TYPE: Penang?, Wall Asclep.

no 37[= Wall. Cat. 8163] (holotype K; isotypes K [K001129108], E [E00179575]).

Hoya fraterna Blume, Rumphia 4: 32 (1849) [late October 1849]; Blume, Mus. Bot.

1(3): 44 (1849) [November 1849], - TYPE: Indonesia, Java, West Java, Kuripan [In

calcareis Kuripan Javae occidentalis] s.d., s.coll. (lectotype L [L0004318], designated

here) (Synonymy established by Backer & Bakhuizen van den Br ink , 1965).

Card. Bull. Singapore 69(1) 2017

Hoy a occlusaRidl., J. Straits Branch Roy. Asiat. Soc. 61: 31 (1912). - TYPE: Malaysia,

Selangor, top of the Batu Caves, December 1890, Ridley, H.N. s.n. (lectotype SING

[SING0059478], designated here).

Hoya angustisepala Elmer ex C.M.Burton, Hoyan 8(4) pt 2: 6 (1987). - Hoya

mindanaensis Elmer, Leafl. Philipp. Bot. 10: 3585 (1938), nom. inval. (missing Latin

description). - TYPE: Philippines, Davao, Mindanao, Todaya, Mt. Apo, Sibulan River,

3000 ft., June 1909, Elmer, A.D.E. 10829 (lectotype B [B100277215], designated here;

isolectotypes BISH [BISH1000876], BM [BM000945404], BO, E [E00288757], K

[K000911125], L [L0004313], L(U) [U1102604],NY [NY00318664], US [00170056]).

Notes. Blume did not directly indicate any specimen for Hoya coriacea Blume but

mentioned the provenance ‘ In fruticetis ad pedem montis Salak/. I have located one

specimen in L bearing an autograph label by Blume ‘ Hoya coriacea; Salak’ which is,

therefore, suitable for designation as the lectotype. A specimen sent by Blume to P in

1836 [P00639838] which is labelled ‘In fruticetis ad pedem montis Salak ’ is a perfect

match to Blume’s description. However there is no autograph label and therefore the L

specimen is preferred as lectotype for H. coriacea.

Duplicates of Wall. Asclep. 37[= Wall. Cat. 8163] have been found at E and K.

The K duplicate is the only one bearing a Herbarium Robert Wight Proper (HRWP)

label, it is annotated ‘37 H. brunoniana ’ has a sketch of the flower in Wight’s hand and

is therefore one of Wight’s specimens on which he based his descriptions. Linn Soc

Mss SP1284 indicates that only 1 sheet of Wall. Asclep. 37 was given to Robert Wight

for his personal working herbarium, therefore the K sheet can be considered a holotype.

The name Hoya fraterna was first mentioned in volume four of Rumphia (Blume,

1849: 32) [late October 1849] where Blume compared it to the similar species Hoya

coriacea (fraterna = brotherly, closely associated (Steam, 2008)). The description is

expanded in Blume (1849: 44) [November 1849] (Stafleu & Cowan, 1976). Blume

did not refer to any specimens but mentioned the provenance as ‘In calcareis Kuripan

Javae occidentals I have located one specimen in L bearing an autograph label in

Blume’s hand ‘Hoyafraterna Bl; in calcareis Kuripan’ that is to be considered original

material and is, therefore, selected as lectotype.

One duplicate has been found of the type of Hoya occlusa Ridl. at SING

[SING0059478], The specimen is well-preserved and fertile and is here selected as

lectotype.

Hoya coronaria Blume, Bijdr. FI. Ned. Ind. 16: 1063 (1826); Rintz, Malayan Nat.

J. 30: 505 (1978). - Eriostemma coronaria (Blume) Kloppenb. & Gilding, Fraterna

14(2): 1 (2001). - TYPE: Indonesia, Java, s.d., s.coll. ‘124/6’ (lectotype L, [sheet

no. 898.168-121], designated by Kleijn & Van Donkelaar (2001: 469); isolectotype L

[sheet no. 898.168-128].

Eriostemma obtusifolioides Gilding & T.Green, Fraterna 22(3): 5 (2009). - TYPE:

Originally from Malaysia, Sabah, Apin Apin, cultivated in USA, Hawaii, Oahu,

Ka‘a‘awa, Green Plant Research, vouchered in 2007 as Green, T. s.n. (holotype BISH

[BISH1072387]).

Names and types of Bornean Hoya

=Dischidia villosa nom. nud. (based on L sheet [L0004319, L0004320]).

Notes. Kleijn & Van Donkelaar (2001: 469) selected sheet no. 898.168-121 (L) as

lectotype of Hoya coronaria Blume. This is one of three sheets annotated ‘ Hoya

coronaria BT in Blume’s hand. It must be noted that both sheet no. 898.168-121 and

sheet no. 898.168-128, despite being specimens seen by Blume, cannot be directly

linked to the protologue of the taxon. The syntype sheet no. 898.168-124 instead has

a label in Blume’s handwriting where, among undecipherable words, is the common

name ‘Kilampahan’ and on a different label in Reinwardt’s handwriting the possible

variant ‘Tjilampahan’, in agreement with Blume’s protologue.

Another syntype of Hoya coronaria is the illustration of Corona Ariadna, published in

Rumph. Amb. 5. t. 172.

The synonymy of Hoya coronaria is complex and a taxonomic revision of

Hoya section Eriostemma Schltr. is required before a complete list of synonyms can

be given. The Bornean Eriostemma obtnsifolioides Gilding & T.Green, however, is

indistinguishable from H. coronaria so is included here. Eriostemma is nested within

Hoya but can be recognised at the sectional level ( Hoya sect. Eriostemma) (Wanntorp

et al., 2006a, 2006b, 2011, 2014; Rodda & Ercole, 2014; Rodda et al., 2014). The type

of Eriostemma obtnsifolioides was only mentioned as ‘Bishop 736375 and ‘Ex hort.

97048TG, Apinapin, Sabah, Malaysia’. The code ‘97048TG’ was not found on the

type sheet. Type citation has been here amended.

Hoya corymbosa Rodda & Simonsson, Syst. Bot. 38: 1126 (2014). - TYPE: Originally

from Malaysia, Sabah, Sipitang District, Mt. Rimau area, cultivated in Kipandi Butterfly

Farm, vouchered on 18 June 2009 as Gokusing, L. et al. All803/2009 (holotype SAN).

Hoya cumingiana Decne. in A.DC., Prodr. 8: 636 (1844). - TYPE: Philippines,

Luzon, Batangas, s.d., Cuming, H. 1480 (lectotype P [P00639839], designated here;

isolectotypes BM, C [C10006728], CGE (2 sheets), E [E00288755], G, G-Boiss (2

sheets), K [K001044838, K000911123], L [L0004322]).

Notes. In describing Hoya cumingiana , Decaisne (1844) mentioned a single gathering,

Cuming 1480, kept in Delessert’s herbarium ‘v.s. in herb. Delessert’. Delessert’s

herbarium was merged into G (Stafleu, 1970) where, however, I could not find any

Cuming 1480 specimen clearly labelled as originating from Delessert’s herbarium.

A suitable specimen from Delessert’s herbarium is instead present in P [P00639839],

being a well-preserved fertile specimen and is here selected as lectotype for Hoya

cumingiana.

Hoya curtisii King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74(2): 563

(1908); Rintz, Malayan Nat. J. 30: 505 (1978). - TYPE: Malaysia, Perak, Waterloo,

October 1892, Curtis, C. 2894 (lectotype SING [SING0059475], designated here;

isolectotypes SING [SING0059474], K [K000895130]).

Card. Bull. Singapore 69(1) 2017

Notes. King & Gamble (1908) described Hoy a curtisii based on Curtis 2894. However,

they did not mention where their types were deposited and, therefore, a lectotype needs

to be selected. Two suitable fertile specimens are present at SING and [SING0059475]

is here selected as lectotype.

Hoya danumensis Rodda & Nyhuus, Webbia 64(2): 164 (2009). - TYPE: Originally

from Malaysia, Sabah, Borneo, Danum Valley, January 1993, cultivated in Italy, Turin,

vouchered on 3 September 2008 as Rodda, M. 2008. lh (holotype FI; isotypes K, L,

TO).

Hoya devogelii Rodda & Simonsson, Webbia 66(1): 35 (2011) [6 June 2011], - TYPE:

Malaysia, Sarawak, Bahagian Kapit, N. part of Hose Mountains, below ridge leading

to Bt. Batu, 10 December 1991, Tech, L.S. and De Vogel, E.F. 9398 (holotype L;

isotypes K, SAR n.v.).

Hoya diversifolia Blume, Bijdr. FI. Ned. Ind. 16: 1064 (1826); Rintz, Malayan Nat.

J. 30: 517 (1978); Jagtap & Singh, Fasc. FI. India 24: 98 (1999). - TYPE: Rumphius

Herbarium Amboinense 5 Tab. 175 Fig. 2. (lectotype, designated here); ‘ Hoya

heterophylla , s.d., s.coll. (epitype L [sheet number 989168-147], designated here).

Hoya heterophylla nom. nud. (based on L sheet number 989168-147).

Hoya crassipes Turcz., Bull. Soc. Imp. Naturalistes Moscou 21: 261 (1848). - TYPE:

Indonesia, Java, Zollinger, H. 2581 (lectotype KW [KW001000511], designated here,

isolectotype A).

Hoya orbicidata Wall, ex Wight & Arn. in Wight, Contr. Bot. India 36 (1834). -TYPE:

Myanmar, Prome in Ava, 1826, Wallich, N., Wall. Asclep. 32 [= Wall. Cat. 8151]

(lectotype K (Herb. R. Wight Proper, with a sketch of the flower in Wight’s hand),

designated here; isolectotypes E [E00179565], K [K001129088], K 2 more sheets).

Hoya esculenta Tsiang, Sunyatsenia 3: 176 (1936), nom. illeg. - TYPE: Sussuela

esculenta Rumph. Herb. Amb. 5: 467, t. 175.28 (1747) (lectotype, designated here).

Hoya zollingeriana Miq., FI. Ned. Ind. 2: 518 (1857) [20 Aug. 1857]. - TYPE:

Indonesia, Java, bij Lalaei, Mei, Zollinger, H. s.n. (lectotypeP [P05029459], designated

in Rodda (2015b: 306)).

Hoya liangii Tsiang, Sunyatsenia 3(2-3): 177 (1936). - TYPE: China, Hainan,

Ngai Yuen, 3 September 1933, Liang, H.Y. 26867 (lectotype IBSC [IBSC0005685],

designated here; isolectotype NY [NY00318649]).

Hoya el-nidicus Kloppenb.,Ymterim 1991(3) supplement: I (1991). - Hoya diversifolia

Blume subsp. el-nidicus (Kloppenb.) Kloppenb., Fraterna 14(1): 13 (2001). - TYPE:

Philippines, Palawan, El Nido rest area, 15 August 1988, Pancho, J. s.n. (holotype

UPLB sheet no. 41931, n.v.)

Names and types of Bornean Hoya

Hoyapersicinicoronaria Shao Y.He & P.T.Ei, Novon 19: 475 (2009). - TYPE: China.

Hainan Prov., Bawangling, on a tree in montane tropical rainforest, 19°06'N 109°06'E,

214 m, 8 August. 2007, He, S.Y. & Lin, J.Y. 0708086 (holotype CANT, n.v., photo at

MO not found).

Notes. Blume (1826) indicated in the protologue of Hoya diversifolia a reference to

Rumphius’ Herbarium Amboinense 5 Tab. 175 Fig. 2 and no direct or indirect reference

to a specimen. Rumphius’s plate does not have a detailed illustration of the flowers and

does not entirely match Blume’s description as the leaves, described as ‘aveniis’ by

Blume have evident venation in the plate. However Blume’s description may be based

on live plants, as fresh leaves do not have distinct venation while veins are sometimes

visible in dry specimens. No specimen from Blume’s time annotated as collected in

‘Kuripan’ or ‘circa Bataviam’ has been found. Herbarium Amboinense 5 Tab. 175 Fig.

2 is therefore the only material available for lectotypification. In L there are numerous

specimens identified as Hoya diversifolia with annotations in Blume’s hand. Among

these there is one labelled Hoya heterophylla in Blume’s hand, a nomen nudum,

and with a very similar etymology as Hoya diversifolia. Sheet number 989168-147

is a well-preserved fertile specimen and is therefore designated as epitype for Hoya

diversifolia. Specimen [P00639841] in Pis labelled both Hoya heterophylla and Hoya

diversifolia Bl. Bijdr’ in Blume’s (or Decaisne?) hand, the locality is Java, ‘m.fisit]

Blume 1836’ and may be original material, however it is impossible to verily if it was

collected prior to 1826.

Hoya crassipes Turcz. was based on Zollinger 2581. The KW duplicate

[KW001000511] is a well-preserved fertile sheet and is designated here as lectotype.

Hoya orbiculata Wall, ex Wight & Arn. was based on Wall. Asclep. 32 [= Wall.

Cat. 8151]. A specimen belonging to what used to be Wight’s personal herbarium and

bearing a sketch of the flower in Wight’s hand is present at K and is the most suitable

lectotype for Hoya orbiculata (Noltie 2005: 134). The plate of Sussuela esculenta in

Rumphius (1747: t. 175.28) is a syntype of Hoya orbiculata.

Hoya esculenta Tsiang is an illegitimate name because it is clearly indicated as

a synonym of the earlier Hoya diversifolia Blume and Hoya orbiculata Wall, ex Wight

& Arn. and is therefore superfluous and illegitimate following ICN Art. 52.1 (McNeill

et al., 2012). It is lectotypified by Herb. Amb. 5: 467, t. 175.28.

The type of Hoya liangii Tsiang was indicated as Liang 26867. The IBSC

duplicate [IBSC0005685] is a well-preserved fertile sheet and is designated here as

lectotype. The syntype How, C.F. 73935 is also available at IBSC [IBSC0005687]).

Hoya elliptica Hoolc.f., FI. Brit. India 4: 58 (1883); Rintz, Malayan Nat. J. 30: 505

(1978). - TYPE: Malaysia, Malacca, 1 April 1868, Maingay, A.C. 3286, Kew Distrib.

1137 (lectotype K [K000895126], designated here; isolectotype K [K000895127]).

Notes. Hooker described Hoya elliptica citing specimen Kew Distrib. 1137. Two

sheets are preserved at K labelled Kew Distrib. 1137 as well as bearing the original

collection number Maingay 3286. The duplicate here selected as lectotype is well

preserved, fertile and bears a dissection and a sketch of the flowers by Gamble as an

aid to identification.

Card. Bull. Singapore 69(1) 2017

Hoya elmeri Merr., Univ. Calif. Publ. Bot. 15: 258 (1929); Rodda et al., Webbia 69: 44

(2014). - TYPE: Malaysia, Sabah, Tawao, October 1922 - March 1923, Elmer, A.D.E.

20652 (lectotype A [A00076433], designated by Rodda et al., (2014: 44); isolectotypes

BISH, BM, BO, IBSC, MICH, NY, P, SING, U, UC [UC312114], Z).

Hoya fauziana Rodda, Simonsson & A.L.Lamb, Webbia 70(2): 207 (2015) [8 June

2015]. - TYPE: Originally from Malaysia, Sabah, Ulu Tomani, 1200 m, cultivated at

Kipandi Park, vouchered on 30 October 2013 as Lamb, A. 2462/2013 (holotype SAN;

isotype SING).

Hoya finlaysonii Wight, Contr. Bot. India 38 (1834); Rintz, Malayan Nat. J. 30: 511

(1978); Noltie, Bot. Robert Wight 170 (2005). - TYPE: Penang?, herb. Finlayson,

Wall Asclep. no 42[B = Wall. Cat. 8166B] (lectotype K [K000895121], designated

here; isolectotypes K [K001129114], E [E00179577]).

Notes. The occurrence of Hoya finlaysonii Wight in Borneo is doubtful. However, as

numerous closely related taxa occur in Borneo and although their synonymies have

not been fully clarified, this name is also typified here. Wight (1834) mentioned the

materials for Hoya finlaysonii as ‘Wall Asclep. n. 42. Penang?; Wallich and Finlayson’.

The description did not include a description of the flowers and it is possible that the

specimens examined were all sterile, as also mentioned by King & Gamble (1908)

when referring to Kew’s materials of H. finlaysonii. A sterile specimen belonging to

what used to be Wight’s personal herbarium is present at K and is the most suitable

lectotype for Hoya finlaysonii (Noltie 2005: 134). Rintz (1978) mentioned a ‘holotype’

of Hoya finlaysonii as ‘Malaysia, Melaka, Wight (K)’ but it is impossible to ascertain

whether [K000895121] is the same specimen as no Wight specimen of the taxon was

collected in Melaka. Since the specimen is sterile a careful selection of a fertile epitype

will be needed. The syntype Wall Asclep. no 42[A = Wall. Cat. 8166A] is also present

at K [K001129113],

Hoya forbesii King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74(2): 574

(1908); Rintz, Malayan Nat. J. 30: 517 (1978). - TYPE: Indonesia, Sumatra, Kaba

Volcano, 1881, Forbes, H.O. 2896a (lectotype K [K000894730], designated here).

Notes. Three specimens were cited in the original description of Hoya forbesii ,

Scortechini 1679 & 1680 and Forbes 2896a. The only extant specimen I was able to

locate is Forbes 2896a (K), which also bears a dissection and a drawing of the flowers

in Gamble’s hand and is therefore selected as the lectotype.

Hoya gildingii Kloppenb., Fraterna 15(1): 10 (2002). - TYPE: Originally from

Malaysia, Sabah, Mt. Kinabalu, Bukit Tapai Trail, cultivated in Sweden, vouchered in

2001 as Wiberg, E.K., s.n. (holotype UC [UC1784972]).

Hoya glabra Schltr., Bot. Jahrb. Syst. 40 (Beibl. 92): 14 (1908). - TYPE: Borneo,

Long Wahau, 2 Aug 1901, Schlechter, R. 13458 (lectotype B [B100277199], designated

here).

Names and types of Bornean Hoya

Notes. Schlechter 13458 is extant in B and is therefore selected as lectotype for Hoya

glabra.

Hoya hamiltoniorum A.L.Lamb, Gavrus, Emoi & Gokusing, Sandalcania 19: 32

(2014). - TYPE: Originally from Malaysia, Sabah, Ulu Tomani, cultivated at Kipandi

Park, vouchered on 26 Aug 2009 as Gokusing, L. & Lombika, J. in Lamb, A. 1814/2009

(holotype SAN; isotype SNP n.v.).

Hoya ignorata T.B.Tran, Rodda, Simonsson & Joongku Lee, Novon 21(4): 509 (2011)

[29 December 2011], - TYPE: Vietnam. DakLak, Dak Mil, 11 December 1979, Bien,

L. K. 1072 (holotype HN; isotype MO).

Hiepia corymbosa V.T.Pham & Aver., Turczaninowia 14(3): 6 (2011). - TYPE:

Vietnam, Quang Tri prov., Da Krong distr., Ta Rut municipality, A Pul village,

around point 16°24'29"N 107°01 f 07"E, broad- leaved evergreen forest along rocky

river canyon composed with shale and sandstone at elev. about 300 m a.s.l., 10 May

2011, Averyanov, L., Loc, P.K., Hieu, N.Q., The, P.V., Vinh. N.T. CPC 3066 (holotype

Herbarium of the Center for Plant Conservation, Vietnam, n.v.; isotype LE, n.v.).

Notes. The monotypic genus Hiepia V.T.Pham & Aver, was published on 28 September

2011. Examination of the protologue, associated images and detailed photographs of

the taxon (Pham & Averyanov, pers. comm.) suggest that it is indistinguishable from

a taxon published as Hoya ignorata T.B.Tran et al. on 29 December 2011. Rodda

et al. (2014) showed that the taxon belongs to the genus Hoya and since the epithet

‘corymbosa ’ is unavailable in Hoya the correct name is Hoya ignorata under Art. 11.4

of the ICN (McNeill et al., 2012).

Hoya imperialis Lindl., Edwards’s Bot. Reg. 32: sub. t. 68 (1846); Rintz, Malayan

Nat. J. 30: 501 (1978); Forster & Liddle, Asklepios 56: 24 1992). - TYPE: Herb.

Lindley ‘Cultivated, from Sarawak via Lowe’s Nursery’, s.d., s.coll., s.n. (lectotype

CGE [CGE06041], designated here).

Hoya imperialisLmdl. var. rauschiiRegd, Gartenflora 282 (1855). -TYPE: Gartenflora

1855: 282. t.134 (erroneously numbered t.130) (lectotype designated here).

Notes. Forster & Liddle (1992a) mentioned that a lectotype for Hoya imperialis Lindl.

may have to be chosen from one of the illustrations published by Lindley (however

no illustration was provided together with the protologue) as no type specimen may

be extant in BM, K or L (Rintz, 1978). Lucas (2008) traces the Lindley herbarium to

CGE. There I have found a specimen originally belonging to Lindley’s herbarium that

is here selected as lectotype for Hoya imperialis.

Hoya imperialis Lindl. var. rauschii was described as having paler corolla

lobes than the type. The taxon was described from cultivated material growing in

Schaffhausen (Switzerland). Original material has not been found and it is possible

that a specimen was never made. The illustration published concurrently with the

description is selected as lectotype.

Card. Bull. Singapore 69(1) 2017

Hoya jiewhoeana Rodda, A.L.Lamb & Gokusing, Guide Hoyas Borneo 198 (2016)

[2 May 2016] (as ‘jiewhoeiana’). - TYPE: Originally from Malaysia, Sabah, Tawau,

Kalabakan area, local collector, cultivated in Sabah, vouchered on 17 January 2016 as

Bosuang, S. & Gokusing, L. in Lamb, A. AL2747/2016 (holotype SAN; isotype SING).

Hoya kastbergii Kloppenb., Fraterna 16(4): 1 (2003). - TYPE: Malaysia, Sarawak,

Bau, Kastberg, A. (holotype UC, missing). Malaysia, Sarawak, Near Mulu National

ParkH.Q., 23 March 1981, Collenette, I.S. 2357 (neotype L [L2726571], designated

here; isoneotype K).

Notes. The type specimen of Hoya kastbergii was indicated only as ‘UC 102003’.

The specimen in UC barcoded [UC 102003] belongs to Erigeron glacialis (Nutt.) A.

Nelson var. glacialis (Pia Nutt pers. comm.). It is possible that ‘102003’ may be a

collection date (October 2003) and not a barcode. See Hoya nyhuusiae below for a

similar case. No other specimen of Hoya kastbergii is present in UC and since no

collection information apart from the collector name and the UC barcode was provided

in the description of the taxon it is impossible to identify isotypes that may be present

in other herbaria. To fix the application of the name I have therefore selected another

specimen matching the original description, Collenette 2357 (L) as neotype of H.

kastbergii.

Hoya kipandiensis Gavrus, A.L.Lamb, Emoi & Gokusing, Sandakania 19: 42 (2014).

-TYPE: Originally from Malaysia, Sabah, central Crocker Range, Mongkusilad River,

nr. Kg. Kipandi, cultivated at Kipandi Park, vouchered on 25 Jan 2010 as Gokusing, L.

in Lamb, A. 1909/2010 (holotype SAN; isotype SNPn.v.).

Hoya kloppenburgii T.Green, Fraterna 14(2): 11 (2001). - TYPE: Originally from

Malaysia, Sabah, Nabawan, cultivated in USA, Hawaii, Oahu, Ka‘a‘awa, vouchered

on unknown date as Green, T. 97057 (holotype BISH [BISH1014779]).

Hoya lacunosa Blume, Bijdr. FI. Ned. Ind. 16: 1063 (1826); Rintz, Malayan Nat. J.

30: 490 (1978). - Otostemma lacunosum (Blume) Blume, Rumphia 4: 30 (1849) [late

October 1849]. - TYPE: [circa Buitenzorg at arbores] Hoya lacunosa, s.d., s.coll., s.n.

(lectotype L [sheet number 898168-188], designated here).

Notes. The only specimen belonging to Hoya lacunosa Blume that can be referred to

Blume is sheet number 898168-188 (L) that bears an autograph label ‘ Hoya lacunosa'.

Blume left Java in 1826 and returned to the Netherlands (Van Steenis-Kruseman,

1950). Therefore the specimen, if collected by Blume, is likely to have been collected

before the publication of the Asclepiadaceae account in the Bijdragen (Blume, 1826)

and is therefore a suitable lectotype for Hoya lacunosa.

Hoya lambii T.Green, Fraterna 13(2): 2 (2000). - TYPE: Originally from Malaysia,

Sabah, cultivated in USA, Hawaii, Oahu, Ka‘a‘awa, reputedly vouchered on unknown

date as Green, T. 9905 (holotype BISH, not found). Originally from Malaysia, Sabah,

Names and types of Bornean Hoya

Cultivated in USA, Hawaii, Oahu, Ka‘a‘awa, vouchered on unknown date as Green,

T. s.n. (neotype BISH [BISH1016619], designated here).

Notes : The holotype of Hoya lambii, Green 9905 is not present in BISH and no

duplicates have been located in other herbaria. A specimen labelled as neotype is

present in BISH ( Green s.n.) suggesting that the publishing author may be aware that

the holotype is missing. However, the neotypification has not been published and is

therefore formally published here.

Hoya lasiantha (Korth. exBlume) Miq., FI. Ned. Ind. 2: 526 (1857) [20 August 1857];

Rintz, Malayan Nat. J. 30: 498 (1978). - Plocostemma lasianthum Korth ex Blume,

Mus. Bot. 1(4): 60 (1849). - TYPE: Borneo, [ad montem Pamotton insulae Borneo]

s.d., Korthals, P. s.n. (lectotype U [U1102625] designated here); Mus. Bot. 1(4) Fig.

14 (1849) (epitype, designated here).

Notes. The name Hoya lasiantha is usually attributed to Blume (1849: 30) [late

October 1849] (Stafleu & Cowan, 1976) where it is, however, a nomen nudum as it

refers only to a specimen labelled Hoya lasiantha in Korthals herbarium. The taxon is

validated as Plocostemma lasianthum Korth. ex Blume (Blume 1849: 60) [November

1849] (Stafleu & Cowan, 1976) and later transferred to Hoya by Miquel (1857). The

authorship of the name is here corrected.

Blume mentioned a collection in Korthals herbarium from Borneo ‘ad montem

Pamotton insulae Borneo’. I have located a single specimen matching Blume’s

description and illustration (Blume, 1849: Fig. 14) with a manuscript annotation by

Korthals ‘ Hoya lasiantha Borneo’ in L (formerly in U). The label does not have further

details on the collection locality, but this is common with Korthals specimens (Van

Steenis-Kruseman, 1950). The specimen is only a fragment with a single mounted

flower. It is possible that an original complete specimen was in L but may have gone

missing, or that Korthal’s label may have been mixed during Miquel’s time (Van

Steenis-Kruseman, 1950), therefore the U specimen is the only available original

material for lectotypifying Hoya lasiantha. To aid in the application of the name the

detailed illustration in Blume (1849, Fig. 14) is selected as epitype.

Hoya latifolia G.Don, Gen. Hist. 4: 127 (1837); Rintz, Malayan Nat. J. 30: 508 (1978).

- Hoya macrophylla Wight, Contr. Bot. India 38 (1834), non Hoya macrophylla

Blume (1826). - TYPE: Malaysia, Penang, Wall. Asclep. no. 138[A = Wall. Cat

8161A (lectotype K [K000895124], designated here) ex parte; isolectotypes CGE (1

not mixed, 1 ex parte , E [E00179576], K [K000895125].

Notes. A suitable lectotype for Hoya latifolia G.Don can be selected from amongst

material of the two syntypes indicated in the protologue (Wall. Asclep. no. 138[A

= Wall. Cat 8161 A] and Wall. Asclep. no. 138[B = Wall. Cat 816IB]). Original

specimens have been found at CGE, E and K. The one here selected as lectotype is

the only one bearing a Herbarium Robert Wight Proper (HRWP) label indicating that

it was one of the specimens studied by Wight and on which he based his descriptions

Card. Bull. Singapore 69(1) 2017

(Noltie, 2005). The specimen is composed of two leaves and a follicle, excluding the

flowers in the envelope, which belong to Hoy a mitrata Kerr (Rodda, 2012). Often two

types of leaves are present on each specimen, some agreeing with Wight description

(palminerved) others easily separated because they are penninerved and with a thinner

texture. As suggested by Nutt (label on [K000895124]), the flowers do not agree with

H. latifolia as currently applied e.g. in Rintz (1978). A syntype Wall. Asclep. no. 138[B

= Wall. Cat. 8165B] is present at K [K000895123],

Hoya mappigera Rodda & Simonsson, Feddes Repert. 122(5-6): 338 (2012). - TYPE:

Malaysia, Dingdings, Lumut Dindings, 1894, Ridley H.N. s.n. (holotype SING

[SING0120871]).

Hoya medinillifolia Rodda & Simonsson, Webbia 66: 149 (2011). - TYPE: Malaysia,

Sarawak, Batu Laga Plateau, Batang Rejang, 7 th Division, 10 Sep 1984, A bang Mohtar

S.48205 (holotype K [K000894726]; isotypes L, SAR).

Hoya meredithii T.Green, Fraterna 20(3): 14 (2007). - Hoya meredithii T.Green,

Phytologia 64(4): 304 (1988), nom. inval. - TYPE: Originally from Malaysia,

Sarawak, Bau, Meredith, Y. 80-05, cultivated in USA. Hawaii, Oahu, Ka‘a‘awa,

vouchered in 1981 as Green, T. 81-002 (holotype BISH [BISH1014780]; isotypes

BISH [BISH1014781], NY).

Notes. The holotype of Hoya meredithii T.Green was cited by Green (2007) as ‘Bishop

Museum, 511439 hie designatus, exhort, T. Green of Oahu, Kaaawa, Hawaii, Lectotype

(NY)’. This is interpreted as holotype BISH [BISH1014780], with an isotype at NY. A

second isotype at BISH [BISH1014781] is present.

Hoya mindorensis Schltr., Philipp. J. Sci. 1 (Suppl. 4): 303 (1906). - TYPE:

Philippines, Mindoro, Baco River, April-May 1905, McGregor, R.C. 332 (lectotype B

[B100277216], designated by Rodda et al. (2014)).

Hoya minutiflora Rodda & Simonsson, Webbia 65(2): 174 (2010). - TYPE: Originally

from Borneo?, cultivated at Hortus Botanicus Leiden., vouchered in August 1999 as s.

coll. L30929, IPPS 7760 (holotype L, spirit material).

Hoya mitrata Kerr, Hooker’s Icon. PI. 35. t. 3406 (1940). - TYPE: Thailand, Surat,

Ban Tong Tao, Kerr, A.F.G. 13152 (lectotype K, designated by Rintz (1978)).

Hoya wallichiana Decne. inA.DC., Prodr. 8: 635 (1844) nom. rej. prop. - TYPE: -

[Malaysia, Penang] “Hoya Wallichiana Dne in DC. Prodr. Hoya coronaria Bl. (d’apres

Hooker FI. of British India IV 58)” (lectotype P designated in Rodda (2012)).

Notes. A lectotype was selected in Rodda (2012). However, Rintz (1978) mentioned

the same specimen and used the word ‘type’ which counts as an earlier lectotypification

under Art. 9.9 of the ICN (McNeill et al., 2012). Hoya mitrata has been proposed

Names and types of Bornean Hoya

as a nom. cons. prop, against the earlier Hoya wallichiana Decne. (Rodda, 2012). A

decision is still pending.

Hoya monetteae T.Green, Fratema 17(2): 10 (2004). - TYPE: Philippines, Palawan

Island, Tumarbong River delta, 4 November 2000, Green, T. 007 (holotype BISH, not

found). Malaysia, Sabah, Lahad Datu, Tabin, South Eastern Trail, 3-4 km from the

base camp, 100-200 m, 10 March 1998, Sugau, J.B., et al. JBS340 (neotype SAN,

designated here; isoneotypes K, L [L2718278]).

Notes. The specimen Green 007 , dated 4 November 2000, is not extant in BISH.

A sterile specimen, Green 007, dated 30 May 2008, labelled as neotype of Hoya

monetteae is instead present at BISH. I have been unable to find publication of the

neotypification. Specimen Sugau et al. JBS340 (SAN) is a fertile and well-preserved

specimen of H. monetteae and is here selected as a neotype.

Hoya multiflora Blume, Cat. Gew. Buitenzorg 49 (1823); Rintz, Malayan Nat. J.

30: 492 (1978); Jagtap & Singh, Fasc. FI. India 24: 108 (1999). - Centrostemma

multiflorum (Blume) Decne., Ann. Sci. Nat., Bot. ser. 2, 9: 272 (1838). - Cyrtoceras

multiflorum (Blume) Heynh., Nom. Bot. Hort. 1(2): 183 (1840). - TYPE: Indonesia,

Java [prope flumine Tjapaes? Kietpil?, Salleh], 16 Sep 1820, s.coll., s. num. (lectotype

L [L2727033], designated here).

Asclepias carnosa Blanco, FI. Filip. 208 (1837), nonL.f. (1782). -TYPE: Philippines,

Rizal Province, Luzon, Bosoboso, March 1915, Merrill Species Blancoanae 852

(neotype L [L2726976], designated here; isoneotype P [P00700474]).

Cyrtoceras reflexum Benn., PI. Jav. Rar. 90 (1838). - Centrostemma cyrtoceras (Benn.)

Meisn., PI. Vase. Gen. Ill (1840), nom. inv. -TYPE: Indonesia, Java, Bennett, PI. Jav.

Rar. (1838) Fig. 21 (lectotype, designated here). Indonesia, Java, ‘Ascl. 5’ 1802-1818,

Horsfield, T. s.n. (epitype K [K000894745], designated here; isoepitype K).

Cyrtoceras lindleyanum Decne. Hort. Universel 1: 81 (1839). - Hoya coriacea auct.

non Blume: Lindley, Edwards’s Bot. Reg. 25:1.18 (1839). - Centrostemma coriaceum

Meisn., PI. Vase. Gen. Ill (1840), nom. illeg. - Centrostemma lindleyanum (Decne.)

Decne. in A.DC., Prodr. 8: 634 (1844). - Cyrtoceras coriaceum Heynh., Nom. Bot.

Hort. I (2): 183 (1840), nom. illeg. - TYPE: Philippines, Manila, cultivated in UK,

Loddiges Nurseries, August 1830? (lectotype CGE [CGE06040], designated here).

Cyrtocerasfloribundum Maund, Botanist 4:1.178 (1840), nom. illeg. - TYPE: Maund,

Botanist 4:1.178 (1840) (lectotype, designated here).

Centro stemmamicranthum Blume, Mus. Bot. 1(3): 46 (1849 ).- Cyrtoceras micranthum

(Blume) Miq., FI. Ned. Ind. 2: 515 (1857). - Hoya celebica Boerl., Handl. FI. Ned.

Ind. 2: 440 (1899). - TYPE: ‘in sylvis insulae Celebes’ Fig. 12 in Blume, Mus. Bot.

1(3) (1849) (lectotype, designated here).

Card. Bull. Singapore 69(1) 2017

Centrostemma elegans Blume, Mus. Bot. 1(3): 46 (1849). - Cyrtoceras elegans

(Blume) Miq., FI. Ned. Ind. 2: 515 (1857). - Hoya elegans (Blume) Boerl., Handl.

FI. Ned. Ind. 2: 440 (1899), non Kostel. (1834). - TYPE: Borneo, s.d, s.coll, s.n.

(lectotype L [L2726969], designated here).

Centrostemma laurifolium Blume, Mus. Bot. 1(3): 46 (1849). - Cyrtoceras laurifolium

(Blume) Miq., FI. Ned. Ind. 2: 514 (1857). -Hoyajavanica Boerl., Handl. FI. Ned. Ind.

2: 440 (1899). -TYPE: Indonesia, Java, W Java [Tjunlcankan. In sylvismontanis Javae

occidentalis], Salalc, August 1825, s.coll., s.n. (lectotype L [L0004337], designated

here).

Cyrtoceras uncinatum Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indie 25: 408 (1863).

-TYPE: Indonesia, Sumatra, Palembang, ad arbores, s.d., Teijsmann, J.E. (not found).

Indonesia, Java [prope flumine Tjapaes? Kietpil ?, Salleh], 16 Sep 1820, s.coll., s. n.

(neotype L [L2727033], designated here).

Hoya reticulata Costantin in Lecomte, FI. Indo-Chine 4: 138 (1912), non Hoya

reticulata Moon (1824), nee Hoya reticulata Schltr. (1913), nee Hoya reticulata Merr.

(1921). - TYPE: Cambodia, Mt. Krewanh, Sep 1870?, Pierre, L., s.n. (lectotype P

[P00700502], designated here).

Centrostemma platypetalum Merr., Sunyatsenia 2: 16-17 (1934). - TYPE: China.

Guangdong. Sup Man Ta Shan, 19 July 1933, Tso, C.L. 23346 (lectotype NY

[NY00318540], designated here).

Hoya refracta nom. nud. on numerous sheets in L and P.

Notes. The publication of Hoya multiflora Blume only consists of a diagnosis ‘caule

fruticoso scandente; foliis oppositis oblong-lanceolatis utrinque acutis glabris subtus

pallidioribus; floribus fasciculatis terminal ibus’ without any information that could

link it to a specimen such as a collector name, a locality or a common name. Numerous

specimens of Hoya multiflora are present in L with Blume’s autograph labels. Among

these, only one [L2727033] has a date definitively setting its collection before

publication of the protologue, ‘16 September 1820’. It is a fertile and well-preserved

specimen and is therefore suitable lectotype material.

There are no known extant collections from Blanco, including for Asclepias

carnosa Blanco, non L.f. (Merrill, 1918). Merrill’s illustrative specimen 852 is

therefore selected as neotype for Asclepias carnosa Blanco.

No specimens are listed for Cyrtoceras reflexum Benn. The plate included

together with the species description (Bennett, 1838 Fig. 21) is therefore selected

as lectotype. A Horsfield collection from Java [K000894745] that may be original

material is selected as an epitype.

Cyrtoceras lindleyanum Decne. was a new name based on a plant misidentified

by Lindley as Hoya coriacea Blume (Lindley, 1839). Lindley’s treatment was based

on a plant ‘Sent by Mr Cuming to Messrs. Loddiges, from Manila; it flowered for

Names and types of Bornean Hoya

the first time in August 1838’. In Lindley’s herbarium at CGE there is a sheet labeled

‘Hoya coriacea Loddiges Aug. 1830’. Excluding the date that does not match, this

sheet matches the protologue and being the only original material available is here

selected as lectotype for Cyrtoceras lindleyanum. Miquel (1857: 515) cited Cyrtoceras

lindleyanum Miq. with himself as the author. This, however, is likely a typographical

error as he clearly also included Cyrtoceras lindleyanum Decne. in the citation.

Cyrtoceras floribundum is clearly indicated as a synonym of the earlier

Cyrtoceras reflexum Benn. and is therefore superfluous and illegitimate following ICN

Art. 52.1 (McNeill et al., 2012). The name was based, as per Hoya coriacea Lindl.,

on a live plant originally collected by Cuming in the Philippines and cultivated at

Loddiges Nursery (UK). No specimens have been found with such provenance. The

only material available for the selection of a lectotype is therefore the plate included

together with the species description.

No specimens are mentioned for Centrostemma micranthum Blume. The

illustration published concurrently with the description is therefore selected as the

lectotype.

No specimens are mentioned for Centrostemma elegans Blume. There is,

however, a specimen in L identified as Centrostemma elegans in Blume’s handwriting,

also labelled Cyrtoceras elegans (Blume) Miq in Miquel’s hand and Hoya elegans

(Blume) Boerl. in Boerlage’s hand, that is a suitable lectotype.

No specimen was mentioned in the description of Centrostemma laurifolium,

only ‘In sylvis montanis Javae occidentalis.’ A specimen bearing a label in Blume’s

hand from Salak is here selected as lectotype.

Cyrtoceras uncinatum Teijsm. & Binn. was described from plants cultivated in

Bogor Botanic Garden originally collected from Sumatra by Teijsmann. No specimens

have been found and it is possible that the description was based on a live plant. The

name is neotypified with the lectotype of Hoya multiflora.

In P herbarium there is only one specimen labeled in Costantin’s hand as Hoya

reticulata Costantin, collected by Pierre on Mt. Krewanh. This specimen [P00700502]

is therefore selected as lectotype.

Centrostemma platypetalum Merr. was described based on Tso 23346. The NY

duplicate is a fertile well-preserved specimen and is here selected as lectotype.

HoyanabawanensisKloypQnb. &Wiberg,Fratema 15(2): 2 (2002).-TYPE: Originally

from Malaysia, Sabah, Nabawan, cultivated in Sweden, Borlange, vouchered on

unknown date (before March 2002), as Wiberg, E.K. s.n. (holotype UC [UC1776376]).

Hoya nuttiana Rodda & Simonsson, Phytotaxa 140: 56 (2013). - TYPE: Malaysia,

Sarawak, near Kuching, 24 October 1894, Haviland, G.D. & Hose , G.F. 8530 (holotype

K [K000014447]).

Hoya nyhuusiae Kloppenb., Fraterna 16(3): 1 (2003). - TYPE: Originally from

Malaysia, Sabah, Mt. Kinabalu, cultivated in Sweden, Stockholm, vouchered in 2003

as Nyhuus, T. s.n. (holotype UC [UC 1784974]).

Card. Bull. Singapore 69(1) 2017

Notes. The protologue of Hoya nyhuusiae Kloppenb. states ‘(UC) Torill Nyhuus #2003

ex hort. Torill Nyhuus, Sweden. From Mt. Kinabalu, Sabah, Malaysia’. There is no

specimen Nyhuus 2003 in UC but the specimen labelled as holotype is Nyhuus s.n. and

‘2003’ is the year of collection as also noted for Hoya walliniana. Type citation has

been amended accordingly.

Hoya obscura Elmer ex C.M.Burton, Hoyan 8(1): 15 (1986). - Hoya obscura Elmer,

Leafl. Philipp. Bot. 10: 3586 (1938), nom. inval. (missing Latin description). - TYPE:

Philippines, Luzon, Sorsogon, Irosin, Mt. Bulusan, July 1906, Elmer, A.D.E. 16719

(lectotype US [US00026729], designated here; isolectotypes BISH [BISH1000882],

BM [BM000945396], BO, G, GH, K [K000911128], L [L0004338], L(U)

[U1102639], NY [NY00318665], P [P00639850], S [S12-12728], UC [UC272655],

US [US00026729]). Philippines, Luzon, Nueva Vizcaya, vicinity of Dupax, March-

April 1912, McGregor, R.C., 11377 (epitype BM [BM001135048], designated here).

Notes. No single holotype of Hoya obscura Elmer ex C.M.Burton was stated by Burton

and a lectotype needs to be designated among the duplicates of the type collection

Elmer 16719. All duplicates located and examined are sterile and a selection of an

epitype is necessary for a correct application of the name. Specimen McGregor 11377,

a fertile and well-preserved specimen is therefore selected as epitype of H. obscura.

Hoya obtusifolia Wight, Contr. Bot. India 38 (1834); Rintz, Malayan Nat. J. 30: 520

(1978); Jagtap & Singh, Fasc. FI. India 24: 113 (1999); Noltie, Bot. Robert Wight 170

(2005). - TYPE: sin. loc. (Malaysia, Penang?), Wallich, Waif Asclep. 38 [= Wall. Cat.

8167] (holotype K [K000895115]; isotypes E [E00179578], K [K001129115]).

Hoya teysmanniana Miq., FI. Ned. Ind. 2: 522 (1857). - TYPE: Indonesia, Sumatra,

Soengi Pagoe, s.d., Teijsmann, J.E., 1170HB (lectotype U [U0000686], designated

here; possible isolectotype K [K000894744]).

Notes. Duplicates of Wall. Asclep. 38[= Wall. Cat. 8167] have been found at E and K.

The K duplicate is the only one bearing a Herbarium Robert Wight Proper (HRWP)

label, it is annotated ‘38 Hoya obtusifolia RW’, and is therefore one of Wight’s

specimens on which he based his descriptions. Linn Soc Mss SP1284 indicates that

only 1 sheet of Wall. Asclep. 38 was given to Robert Wight for his personal working

herbarium, therefore the K sheet can be considered a holotype.

Hoya teysmanniana Miq. was described from Teijsmann material indicated as

being from ‘Sumatra, bij Soengi Pagoe’. The U specimen [U0000686] is the only one

found which matches this information and is therefore selected as lectotype. The K

specimen [K000894744] was also collected by Teijsmann in Sumatra and bears a label

in Miquel’s hand. However, it does not have a collection number or locality (Sumatra

only) and may be considered as a possible isolectotype.

Hoya omlorii (Livsh. & Meve) L.Wanntorp & Meve, Willdenowia 41(1): 99 (2011)

[Jun 2011], - Clemensiella omlorii Livsh. & Meve, Edinburgh J. Bot. 66: 454 (2009)

Names and types of Bornean Hoya

(‘omlori- TYPE: Indonesia, Sumatra, Aceh, Gunung Leuser Reserve, Camp

Simpang and vicinity, 19 August 1972, de Wilde, W.J.J.O & de Wilde-Duyfjes, B.E.E.

14377 (holotype L; isotype K).

Hoyapubera Blume, Bijdr. FI. Ned. Ind. 16: 1065 (1826). - Acanthostemmapuberum

Blume, Rumphia 4: 29 (1849). - TYPE: [in m ari d m is insulae Nusae Kambangae ad

arbores] (lectotype L [L0275692], designated here).

Notes. A few specimens can be found in L labelled Hoya pubera Blume. Among these,

[L0275692] bears a label, likely in Blume’s hand, with ‘1684 Dischidia elliptica Bl.

Hoya pubera Bl.’ another specimen [L0275693] bears ‘ Acanthostemma pubera Bl,

Hoya pubera Bl (illegible) Bl. 1684’, possibly in Blume’s hand. Both are sterile and

cannot without doubt be linked to the protologue of Hoya pubera. Another specimen,

in P [P00639851], was sent by Blume as Hoya pubera in 1836 and mentioned by

Decaisne (1844: 639). Once again this is sterile and it is impossible to ascertain

whether it was collected before the publication of H. pubera. Another specimen in

U [U1102645] is labelled as collected by Blume, but is undated and just a fragment.

Following Art. 9.12 of the ICN (McNeill et al., 2012), [L0275692] is to be considered

as ‘remaining original material’ and is therefore here selected as the lectotype for Hoya

pubera.

Hoya phyllura O.Schwartz, Mitt. Inst. Bot. Hamburg 7: 26 (1931). - TYPE:

Indonesia, Borneo, Lebang Hara, 24. November 1924, Winkler , H. 339 (lectotype

HGB [HBG506353], designated here; isolectotype HBG [HBG506354]).

Hoya linusii A.L.Lamb, Gavrus & Emoi, Sandakania 19: 50 (2014) [2 May 2014] . -

TYPE: Originally from Malaysia, Sabah, Crocker Range, upper reaches of the Moyog

River, cultivated at Kipandi Park, vouchered on 13 November 2007 as Gokusing, L in

Lamb, A. 1223/2007 (holotype SAN; isotype SNP, n.v.).

Notes. Schwartz cited only a single collection for Hoya phyllura , Winkler 339. Two

duplicates are found in HBG. The first, bearing a sketch of the flowers is selected as

the lectotype.

Hoya ranauensis T.Green & Kloppenb., Hoya New 2(3): 21 (2014) [before 22 May

2014]. -TYPE: Originally from Malaysia, Sabah, Ranau, Poring Hot Springs, cultivated

in USA, Hawaii, Oahu, Ka‘a‘awa, garden of Ted Green, vouchered on unknown date

as Green, T. s.n. (holotype BISH, sheet no. BISH757786 [BISH1049135]).

Hoya retrorsa Gavrus, A.L.Lamb, Emoi & Gokusing, Sandakania 19: 59 (2014). -

TYPE: Originally from Malaysia, Sabah, Pensiangan, Mount Lumatan, cultivated

at Kipandi Park, vouchered on 4 May 2010 as Gokusing, L. in Lamb, A. 1950/2010

(holotype SAN; isotype SNP n.v.).

Card. Bull. Singapore 69(1) 2017

Hoya revoluta Wight ex Hoolc.f., FI. Brit. India 4: 55 (1883). - TYPE: Malaysia, s.l.,

s.d., Maingay, A.C. 1127 (lectotype K [K000279860], designated by Forster & Liddle

(1992b)). Malaysia, Pahang, Tasek Bera, low altitude, 14 October 1930, Henderson,

M.R. 24439 (epitype SING, designated by Rodda & Simonsson Juhonewe (2013b)).

Hoya rintzii Rodda, Simonsson & S.Rahayu, Webbia 69: 44 (2014). -TYPE: Malaysia,

Selangor, Sungai Langat, 3 Jun 1976, Rintz, RE. RER61 (holotype KEP; isotype K).

Hoya rundumensis (T.Green) Rodda & Simonsson, Webbia 68: 13 (2013). - Hoya

plicata King & Gamble subsp. rundumensis T.Green, Asklepios 108: 19 (2010). -

TYPE: Originally from Malaysia, Sabah, Rundum, 3000 ft., cultivated in USA, Oahu,

Ka‘a‘awa, Hawaii, garden of Ted Green, vouchered on 12 August 2009 as Green, T.

2010.001 (holotype BISH [BISH1016412]).

Hoya ruthiae Rodda, Phytokeys 53: 84 (2015) [21 July 2015]. - TYPE: Malaysia,

Sabah, Lahad Datu, Bukit Baturong, on limestone, 7 July 2000, Kiew, R. RK5029

(holotype SING [SING0077484]).

Hoya scortechinii King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74: 567

(1908) [20 February 1908] - TYPE: Malaysia, Perak, s.d., Scortechini, B. 464b

(lectotype K [K000895122], designated here).

Notes. Three specimens are listed under the description of Hoya scortechinii : J.

Scott s.n. from Penang, B. Scortechinii 464 & 1557 from Perak. I could only locate

B. Scortechinii 464b in K. It is a complete and well-preserved specimen bearing a

dissection and drawings by Gamble made in November 1906 and is therefore a suitable

lectotype.

Hoya sammannaniana A.L.Lamb, Gavras, Emoi & Gokusing, Sandakania 19: 77

(2014). - TYPE: Originally from Borneo. Malaysia, Sabah, Beluran, Paitan River,

cultivated at Kipandi Park, vouchered on 27 December 2008 as Gokusing, L. in Lamb,

A. 1649/2008 (holotype SAN; isotype SNPn.v.).

Hoya sipitangensis Kloppenb. & Wiberg, Fraterna 15(3): 4 (2002). - TYPE: Originally

from Malaysia, Sabah, Sipitang, cultivated in Sweden, Borlange, reputedly vouchered

on unknown date as Wiberg, E.K. 52002 (holotype UC, missing). Originally from

Malaysia, Sabah (Sarawak sic), Sipitang, 1997, Wiberg, E.K. etal., s.n., cultivated and

vouchered in Sweden, Borlange, 2001, Wiberg, E.K. s.n. (neotype UC [UC 1784973]).

Hoyayapiana Kloppenb., Gard. Bull. Singapore 61: 327 (2010). - TYPE: Originally

from Brunei, Belait, Seria, Tutong District, 21 January 2008, Wang, L.K 090122,

cultivated in Singapore, vouchered on 20 April 2009 as Yap, KF. s.n. (holotype SING

[SING0124341]; isotypes BRUNn.v., K [K000898014]).

Names and types of Bornean Hoya

Notes. The type of Hoya sipitangensis was cited as ‘52002 (UC), collected by Eva-

Karin Wiberg et al. at Sipitang, Sarawak, Malaysia ex hort. Eva-Karin Wiberg at

Borlange, Sweden’. The specimen labelled as holotype of Hoya sipitangensis in

UC agrees in collecting information but does not bear the number 52002 which is,

in any event, probably a collection date rather than a number as observed for Hoya

walliniana and Hoya nyhuusiae. It is barcoded UC 1784973. As it is impossible to

verily if [UC 1784973] is the intended holotype or another specimen obtained from the

same plant in cultivation it is designated as neotype of Hoya sipitangensis.

The collection date of the type of Hoya yapiana Kloppenb. is indicated in

the protologue as ‘21 January 2008’ ; however, this is not the collection date of the

specimen but the collection date of a living plant in Brunei that later bloomed in

Singapore where it was cultivated by K.F. Yap, who made the type specimen from this

plant on 20 April 2009. This collection date is mentioned on the holotype in SING but

not on the duplicate in K.

Hoya sigillatis subsp. sigillatis T.Green, Fraterna 17(3): 2 (2004). - TYPE: Originally

from Malaysia, Sabah, Tenom Agricultural Park, cultivated in USA, Hawaii,

Oahu, Ka‘a‘awa, vouchered on unknown date as Green, T. 91024 (holotype BISH

[BISH1014783]).

Hoya sigillatis T.Green subsp. paitanensis Gavras, A.L.Lamb, Emoi & Golcusing,

Sandalcania 19: 68 (2014). - TYPE: Originally from Malaysia, Sabah, Beluran,

Valley of the Paitan River, cultivated at Kipandi Park, vouchered on 10 April 2011 as

Gokusing, L. & Lombika, J. in Lamb, A. 1757/2011 (holotype SAN; isotype SNPn.v.).

Hoya spartioides (Benth.) Kloppenb., Fraterna 14(2): 8 (2001). - Astrostemma

spartioides Benth., Hooker’s Icon. PI. 14: 7, t. 1311 (1880). - Absolmsia spartioides

(Benth.) Kuntze, Revis. Gen. PI. 2: 418 (1891). - TYPE: Malaysia, North Borneo,

1877-1878, Burbidge, F.W. s.n. (lectotype K [K000894739], designated by Wanntorp &

Forster (2007), first step, designated here, second step; isolectotype K [K000894738]).

Notes. Wanntorp & Forster (2007) selected Burbidge s.n. (K), a specimen not personally

examined by the authors and mounted on two sheets, as lectotype of Hoya spartioides.

However the two sheets are not clearly labelled as being part of the same specimen

and must therefore be considered as two duplicates following ICN Art. 8.3 (McNeill

et al., 2012). The specimen with barcode [K000894739] is therefore designated as the

lectotype of Hoya spartioides. Syntype Motley , J. 978 is extant at K [K000894737].

Hoya telosmoides Omlor, Novon 6: 290 (1996). - TYPE: Malaysia, Sabah, Mt

Kinabalu, Tenompok, 7 Jun 1932, Clemens, J. & M.S. 29828 (holotype BM; isotype

K).

Hoya undulata S.Rahayu & Rodda, Gard. Bull. Singapore 67: 86 (2015). - TYPE:

Indonesia, West Kalimantan, Putussibau, 300-380 m, on a slope above a stream,

February 2014, Sulaiman Hasim s.n. (holotype BO).

Card. Bull. Singapore 69(1) 2017

Hoya vacciniiflora O.Schwartz, Mitt. Inst. Bot. Hamburg 7: 261 (1931). - TYPE:

Indonesia, Bukit Mulu, 2 December 1924, Winkler, H. 518 (lectotype HBG

[HBG506352], designated here).

Notes. Schwartz cited only a single collection for Hoya vacciniiflora O.Schwartz,

Winkler 339. One sheet of this collection is present at HBG and is here selected as

lectotype.

Hoya verticillata (Vahl) G.Don, Gen. Hist. 4: 128 (1837); Rintz, Malayan Nat. J.

30: 514 (1978); Veldkamp et al., Blumea 40: 427 (1995). - Sperlingia verticillata

Vahl, Skr. Naturhist.-Selsk. 6: 113 (1810). - TYPE: Flohr inHb. Vahl ‘exlnd. Orient.

CynanchumV (lectotype C [C10006735], designated by Veldkamp et al., (1995);

possible isolectotype C [C10006736]).

Hoya acuta Haw., Revis. PI. Succ. 4 (1821). - TYPE: ‘Vidi crescentem sine floribus

in region horto Kewense A.D. 1819’; ‘Hort Liv 1825’ (neotype K, designated here).

Hoya pallida Lindl., Bot. Reg. 11: t. 951 (1826), non Hoya pallida Dalzell & Gibson

(1861). - TYPE: Bot. Reg. 11: t. 951 (1826) (lectotype, designated here). ‘Hort Liv

1825’ (epitype K, designated here).

Hoya nicobarica R.Br. ex J.Traill, Trans. Hort. Soc. London 7: 28 (1830). - TYPE:

India, Nicobar Islands, 1785, Hb. Banks, J.P. Rottler/J.G. Koenig s.n. (lectotype BM

[BM001014257], designated by Veldkamp et al. (1995).

Hoya angustifolia J.Traill, Trans. Hort. Soc. London 7: 29 (1830). - Hoya pottsii

J.Traill var. angustifolia (J.Traill) Tsiang & P.T.Li, Acta Phytotax. Sin. 12: 124 (1974).

- TYPE: China, Potts (not found, possibly just a living plant).

Hoya pottsii J.Traill, Trans. Hort. Soc. London 7: 25 (1830). - TYPE: China, Macao.

Cultivated by Mr. Sabine (lectotype Trans Hort. Soc. London 7, plate 1, designated by

Forster & Liddle (1992c)).

Hoya hookeriana Wight, Contr. Bot. India 37 (1834). - TYPE: Bangladesh, Sylhet,

Wall. Asclep 28B [=Wall. Cat. 8158B] HRWP (lectotype K [K000873071], designated

here; isolectotypes BM, E [E00179571], K [K001129101], K-W).

Hoya parasitica Wall, ex Wight, Contr. Bot. India 37 (1834); Veldkamp et al.,

Blumea 40: 427 (1995); Jagtap & Singh, Fasc. FI. India 24: 114 (1999). - Asclepias

parasitica Roxb., Hort. Bengal. 20 (1814), nom. inval. - Asclepias parasitica Roxb.,

FI. Ind. 2: 42 (1832), non Asclepias parasitica Wall, ex Hornem. (1819). - TYPE:

Roxburgh illustration number 1382 (lectotype K, designated here [http://apps.kew.org/

floraindica/img/illustration/large/57419.jpg]).

Names and types of Bornean Hoya

Hoya opposita (Vahl) G.Don, Gen. Hist. 4: 128 (1838); Veldkamp et al., Blumea 40:

427 (1995). -Sperlingia opposita Vahl, Skr. Naturhist.-Selsk. 6: 114 (1810). - TYPE:

Flohr in Hb. Vahl ‘ex Ind. Orient.’ (lectotype C [Cl0006734], designated by Veldkamp

et al. (1995)).

Hoya ridleyi King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74(2): 575 (1908);

Rintz, Malayan Nat. J. 30: 514 (1978). - TYPE: Singapore, Grange Road, Ridley, H.N.

10358 (lectotype SING [SING0047800], designated here; isolectotype K, a dissection

by Gamble).

Hoya parasitica Wight var. geoffrayi Costantin in Lecomte, FI. Indo-Chine 4: 136

(1912); Veldkamp et al., Blumea 40: 427 (1995). - TYPE: Cambodia, Campot, 7

April 1904, Geoffray, M 382 (lectotype P [P00639824], designated by Veldkamp et

al. (1995)).

Hoya parasitica Wight var. spirei Costantin in Lecomte, FI. Indo-Chine 4: 136 (1912);

Veldkamp et al., Blumea 40: 427 (1995). - TYPE: Laos, Luang Prabang, Dok Than,

s.d., Spire 1529 (lectotype P [P00639823], designated by Veldkamp et al. (1995)).

Hoya balansae Costantin in Lecomte, FI. Indo-Chine 4: 136 (1912). - TYPE:

Vietnam, Ha Long Bay, on limestone rocks, 3 July 1885, Balansa, B. 598 (lectotype P

[P00645997], designated here).

Hoya globiflora Ridl., J. Fed. Malay States Mus. 5: 164 (1915); Rintz, Malayan Nat.

J. 30: 514 (1978) (as ‘ globifera’): ; Veldkamp et al., Blumea 40: 427 (1995). - TYPE:

Thailand, Surat Thani, Robinson, H.C. 5756 (lectotype SING, designated by Veldkamp

et al. (1995) not found; lectotype K, designated here).

Hoya obscurinervia Merr., Philipp. J. Sci. 23: 263 (1923). - TYPE: China, Hainan,

San Tsuen, near Nodoa and Loh Hoe, 15 April 1922, McClure, F.A. 9819 (lectotype A

[A00076421], designated here; isolectotype IBSC).

Hoya amoena Bakh.f., Blumea 6: 380 (1950) - TYPE: Indonesia, Java, East Java,

Beoeki, Poeger, 19 October 1895, Koorders, S.H. 20341 (holotype L [L004312]).

Hoya recurvula Kloppenb., Fraterna 13(4): 9 (2000). - TYPE: Originally from

Malaysia? [Southern Malaysia], cultivated in USA, California, Fresno, vouchered in

2000 as Kloppenburg, RD. 81009 (holotype UC [UC1746636]).

Hoya wibergiae Kloppenb., Fraterna 14(1): 5 (2001). - TYPE: Originally from

Philippines, Palawan, Estrella waterfall area, cultivated in USA, California, Fresno,

vouchered in 2000 as Kloppenburg, RD. 951074 (holotype UC [UC 1746635]).

Hoya ubudensis Kloppenb. & Yap, Asklepios 108: 7 (2010). - TYPE: Originally from

Indonesia, Bali, Ubud, Tjampuhan, cultivated in Singapore, vouchered on 26 January

2008 as Toy, A. & Yap, K.F., BAL050208 (holotype SING [SING0143934]).

Card. Bull. Singapore 69(1) 2017

Hoy a recurvula Kloppenb. subsp. bokorensis Kloppenb. & Yap, Asklepios 108:

6 (2010). - TYPE: Originally from Cambodia, Kampot, Bolcor, cultivated in

Singapore, vouchered on 24 August 2008 as Yap, K.F., CBD070101 (holotype SING

[SING0143935]).

Hoya bawanglingensis Shao Y.He & P.T.Li, Novon 19: 357 (2009). - TYPE: China,

Hainan Province, Bawangling, 19°07'N 109°06'E, on tree in montane rainforest, 423

m, 8 August 2007, He, S.H. & Lin, J.Y. 0708081 (holotype CANT, n.v.; photo MO

n. v.).

Hoya amoena Bakh.f. subsp. bogorensis T.Green & Kloppenb., Hoya New 2(4): 14

(2014) - TYPE: Originally from Indonesia, Java, Bogor Botanic Garden, cultivated in

USA, Hawaii, Oahu, Oahu, Ka‘a‘awa, vouchered in 2014 as Green, T. s.n. (holotype

BISH [sheet number 759664]).

Sperlingia tetraphylla nom. mid. annotation on [Cl0006736].

Hoya albens nom. nud. in Miller, Bristol Nursery Cat. 55 (1826).

Hoya lanceolata Donn, Hort. Cantabrig., ed. 2: 92 (1826), nom. nud. non Hoya

lanceolata Wall, ex D.Don (1825).

Notes. The specimen here selected as lectotype of Hoya verticillata was indicated by

Veldkamp et al. (1995) as the holotype. This requires a correction to lectotype under

ICN Art. 9.9 (McNeill et al., 2012). A possible isotype is present at C [C10006736].

It bears the same annotation on the back of the sheet as the lectotype. However,

‘CynanchumV has been crossed out and substituted with Sperlingia tetraphylla , a

nomen nudum and I therefore doubtfully refer to it as an isotype.

Hoya acuta Haw. was described by Haworth based on a sterile live plant

cultivated in Kew ‘ Vidi crescentem sine floribus in region horto Kewense A.D. 1819’.

Following Traill (1830) and Veldkamp et al. (1995), Hoya pallida Lindl., Hoya acuta

Haw., and the nomina nuda Hoya lanceolata and Hoya albens, may all represent

the same plant, sent as a live specimen to K by Wallich and then spread to various

gardens in the UK. Since no original material is available for Hoya acuta the name

is neotypified by a specimen deposited in K that bloomed in ‘Hort Liv’ (Liverpool

botanic garden?) in 1825, likely originating from the same Wallich stock.

Hoya pallida Lindl. was likely based on the same live plant as for Hoya acuta

(Traill, 1830; Veldkamp et al., 1995) but cultivated in the garden of the Duke of

Northumberland. The only original material available is the illustration published in

Lindley (1826: 11, t. 951) that is selected as the lectotype. The neotype of Hoya acuta

is also selected as an epitype of H. pallida. Another specimen, also labelled Hoya

pallida that was obtained from cultivation in Chatsworth in 1850, is present in CGE.

The specimen [BM001014257] was indicated as the holotype of Hoya

nicobarica R.Br. ex J.Traill in Veldkamp et al. (1995) and is here considered as an

effective lectotypification following ICN Art. 9.9 (McNeill et al., 2012).

Names and types of Bornean Hoya

Hoya angustifolia J.Traill was possibly based on a live plant and no extant

specimens have been found.

Wight indicated the materials for Hoya hookeriana as Wall. Asclep. 28A&B and

Wall. Asclep. n. 39 (ex parte). Following Noltie (2005), the Wallich Catalogue numbers

associated with Wall. Asclep. 28A,B are Wall. Cat. 8158A&B, while Noltie (2005)

suggests excluding Wall. Asclep. n. 39 because it is unnamed in the Wallich Cataloge

and it does not bear a locality. Wall. Cat. 8153A (K) was incorrectly indicated as the

lectotype of Hoya hookeriana Wight by Veldkamp et al. (1995). The specimen here

selected as the lectotype belonged to Wight’s personal working herbarium, is labelled

‘Herb. R. Wight. Prop’ and ‘28B’ and bears a description and sketch in Wight’s hand.

Syntypes (Wall. Cat. 8158A) are to be found at E [E00179570], K [K001129100],

[K000873070] andK-W.

Veldkamp et al. (1995) cited a Roxburgh (BM) specimen as the holotype of

Hoya parasitica Wall, ex Wight. This counts as an effective lectotypification under

ICN Art 9.9 (McNeill et al., 2012). However no Roxburgh specimen identified as Hoya

parasitica is present in BM. The only original material is therefore the illustration in

K that is here selected as the lectotype.

Numerous specimens are cited in the protologue of Hoya ridleyi King & Gamble,

collected by Curtis and Ridley. Rintz indicates the type of the taxon as ‘Malaysia,

Ridley (K)’ The only original materials collected by Ridley in K available for Hoya

ridleyi are three dissections by Gamble mounted on a single card. Rintz (1978)

cannot therefore be considered a first step lectotypification. Ridley 10358 (SING) is

instead here selected as lectotype. Other syntypes found are Curtis s.n. [SING 121195,

SING0121079], Ridley 9476. ([SING0121189], K, dissection by Gamble), Ridley

5519 ([SING0121191], K, dissection by Gamble).

Hoya balansae Costantin was described from material collected by Balansa in

Ha Long Bay in Vietnam. The only Balansa specimen from Ha Long Bay labelled

Hoya balansae in Costantin’s handwriting has been found in P [P00645997] and is a

suitable lectotype.

The only collection for Hoya globiflora Ridl. was indicated as ‘Koh Pennan

No. 5756.’ Veldkamp et al. (1995) indicated a SING duplicate as the holotype that

would count as an effective lectotypification under ICN Art 9.9 (McNeill et al., 2012).

However, the specimen is not extant at SING and it is also not listed in the SING

card index suggesting it may never have been present. The K duplicate is instead here

selected as the lectotype for Hoya globiflora.

Hoya obscurinervia Merr. was based on McClure 9819. The duplicate at A

[A00076421] is a well-preserved fertile specimen and it is selected as lectotype.

Hoya vitellina Blume, Mus. Bot. 1(3): 45 (1849) - TYPE: Java, West Java, Blume,

CLP ., s.n. (lectotype L [L0004346], designated here; possible isolectotypes BO

[B01869758, B01869758], U [U1102651]).

Hoya fuscomarginata N.E.Br., Bull. Misc. Inform. Kew 278 (1910). - TYPE: Origin

unknown, cultivated at Glasnevin Botanic Gardens, Dublin, 13 April 1910, s. coll. (K

holotype [K000449169]).

Card. Bull. Singapore 69(1) 2017

Notes. Hoya vitellina Blume was indicated as originating from West Java, without

providing any additional specimen information. Miquel (1857: 522) indicated that

specimens of the taxon were collected by Blume and Van Hasselt. The only possible

material likely collected by Blume is in L [L0004346], Following Art. 9.12 of the

ICN (McNeill et al., 2012) [L0004346] this is to be considered as ‘remaining original

material’ and therefore here selected as lectotype. Three possible duplicates are present

in U and BO. However, due to the lack of information on these sheets it is impossible

to verily if they are actual duplicates.

Hoya fuscomarginata N.E.Br.was described from a specimen deposited in K

originating from cultivation at Glasnevin Botanic Gardens, Dublin and originally

purchased from Mr Pauwels, who stated that it had been imported with orchids from

British Guiana. This is impossible, as no Hoya species is known to occur in the New

World. There is only one specimen in K clearly labelled as type in Brown’s hand that

is therefore a holotype.

Hoya wallichii (Wight) C.M.Burton, Hoyan 18(1:2): 5 (1996); Noltie, Bot. Robert

Wight 111 (2005). - Physostelma wallichii Wight, Contr. Bot. India 40 (1834). -

TYPE: [Herb. R. Wight Prop], ‘Wall Asclep. n. 130’ (lectotype K [K000449753],

designated in Rodda et al., (2016).

Hoya walliniana Kloppenb. & Nyhuus, Fraterna 16(4): 9 (2003). - TYPE: Originally

from Malaysia, Sabah, Sipitang, cultivated in Sweden, Stockholm, vouchered in

November 2003 as Nyhuus, T. s.n. (holotype UC [UC1784975]).

Notes. The type of Hoya walliniana was indicated as ‘112003 (UC) ex hort. Torill

Nyhuus Sweden, originally collected in 1997 in Sipitang, Sartawalc [sic.], Sabah,

Malaysia’. The specimen labelled as holotype in UC agrees in collecting author and

provenance but does not bear collection number 112003. The number is here interpreted

as a collection date (November 2003) as seen in Hoya kastbergii and Hoya nyhuusiae.

The provenance of the plant is Sipitang, a division of the Malaysian state of Sabah,

and not Sarawak as alternatively indicated in the type citation. Provenance and date

are here updated in the holotype citation.

Hoya waymaniae Kloppenb., Fraterna 1995(2): 8 (1995). - TYPE: Originally

from Malaysia, Sabah, cultivated, USA, California, Fresno (holotype UC missing);

Indonesia, Kalimantan Timur, surroundings of Tong Sungai Barang, 15 October 1992,

van Valkenburg J.L.C.H JVV1190 (neotype K, designated here; isoneotypes BO n.v.,

L [L0200212]).

Notes. The type of Hoya waymaniae Kloppenb. was mentioned in the original

publication as ‘cultivated at Fresno, CA., USA from clone obtained from the Sydney

Botanical Garden by Ted Green in 1988 or there about. Holotype: UC#11’. In UC the

label of the specimen indicated as holotype of H. waymaniae reads: ‘Pressed specimen

prepared 23 April 1997 from material cultivated by Ted Green, cloned from plant in

Sydney Botanical Garden which was obtained in 1985 by Ben Wallace from the Tenom

Orchid Centre in Sabah, Borneo. Submitted by Dale Kloppenburg, 1999’. No collector

Names and types of Bornean Hoya

or collection number is present but the specimen is clearly derived from the original

living plant. This specimen cannot be the type as it was collected two years after the

description of H. waymaniae. Despite originating from the same living plant from

which the type was collected, UC1776371 is not a suitable neotype as it is sterile. We

select instead a well-preserved fertile specimen of the taxon, van ValkenburgJVV1190

(K) as the neotype.

Hoya wongii Rodda, Simonsson & L.Wanntorp, Blumea 56: 205 (2011). - TYPE:

Brunei, Tutong district, PasirPuteh, 5 October 1989. Wong, K.M. KMW1566 (holotype

BRUN; isotypes K, L, SAN, SING).

Index to the names (accepted names in bold)

Absolmsia spartioides (Benth.) Kuntze sub: Hoya spartioides (Benth.) Kloppenb.

Acanthostemmapuberum Blume sub: Hoya pubera Blume

Asclepias carnosa Blanco sub: Hoya multiflora Blume

Asclepiasparasitica Roxb. sub: Hoya verticillata (Vahl) G.Don

Astrostemma spartioides Benth. sub: Hoya spartioides (Benth.) Kloppenb.

Centrostemma coriaceum Meisn. sub: Hoya multiflora Blume

Centrostemma cyrtoceras Meisn. sub: Hoya multiflora Blume

Centrostemma elegans Blume sub: Hoya multiflora Blume

Centrostemma laurifolium Blume sub: Hoya multiflora Blume

Centrostemma lindleyanum (Decne.) Decne. sub: Hoya multiflora Blume

Centrostemma micranthum Blume sub: Hoya multiflora Blume

Centrostemma multiflorum (Blume) Decne. sub: Hoya multiflora Blume

Centrostemma multiflorum sensu Chun sub: Hoya multiflora Blume

Centrostemmaplatypetalum Merr. sub: Hoya multiflora Blume

Clemensiella omlorii Livsh. & Meve sub: Hoya omlorii (Livsh. & Meve) L.Wanntorp

& Meve

Cyrtoceras coriaceum Heynh. sub: Hoya multiflora Blume

Cyrtoceras elegans (Blume) Miq. sub: Hoya multiflora Blume

Cyrtocerasfloribundum Maund sub: Hoya multiflora Blume

Cyrtoceras laurifolium (Blume) Miq. sub: Hoya multiflora Blume

Cyrtoceras lindleyanum Decne. sub: Hoya multiflora Blume

Cyrtoceras micranthum (Blume) Miq. sub: Hoya multiflora Blume

Cyrtoceras multiflorum (Blume) Heynh. sub: Hoya multiflora Blume

Cyrtoceras reflexum Benn. sub: Hoya multiflora Blume

Cyrtoceras uncinatum Teijsm. & Binn. sub: Hoya multiflora Blume

Eriostemma obtusifolioides Gilding & T.Green sub: Hoya coronaria Blume

Hiepia corymbosa V.T. Pham & Aver, sub: Hoya ignorata T.B.Tran, Rodda, Simonsson

& Joongku Lee

Hoya acicularis T.Green & Kloppenb.

Hoya acuta Haw. sub: Hoya verticillata (Vahl) G.Don

Hoya aeschynanthoides Schltr.

Hoya albens nom. nud. sub: Hoya verticillata (Vahl) G.Don

Card. Bull. Singapore 69(1) 2017

Hoya arnoena Bakh.f. sub: Hoy a verticillata (Vahl) G.Don

Hoya arnoena Bakh.f. subsp. bogorensis T.Green & Kloppenb. sub: Hoya verticillata

(Vahl) G.Don

Hoya angustifolia J.Traill sub: Hoya verticillata (Vahl) G.Don

Hoya angustisepala Elmer ex C.M.Burton sub: Hoya coriacea Blume

Hoya australis R.Br. ex J.Traill

Hoya bakoensis Rodda

Hoya balansae Costantin sub: Hoya verticillata (Vahl) G.Don

Hoya bawanglingensis Shao Y.He & P.T.Li sub: Hoya verticillata (Vahl) G.Don

Hoya beccarii Rodda & Simonsson

Hoya benchaii Gavrus, A.L.Lamb, Emoi & Gokusing

Hoya brunoniana Wight sub: Hoya coriacea Blume

Hoya callistophylla T. Green

Hoya caudata Hook.f.

Hoya caudata Hook.f. var. crassifolia Ridl. sub: Hoya caudata Hook.f.

Hoya celebica Boerl. sub: Hoya multiflora Blume

Hoya chewiorum A.L.Lamb, Gavrus, Emoi & Gokusing

Hoya brunoniana Wight sub: Hoya coriacea Blume

Hoya clemensiorum T.Green

Hoya coriacea Blume

Hoya coriacea sensu Lindley sub: Hoya multiflora Blume

Hoya coronaria Blume

Hoya corymbosa Rodda & Simonsson

Hoya crassifolia Ridl. sub: Hoya caudata Hook.f.

Hoya erassipes Turcz. sub: Hoya diversifolia Blume

Hoya cumingiana Decne.

Hoya curtisii King & Gamble

Hoya danumensis Rodda & Nyhuus

Hoya devogelii Rodda & Simonsson

Hoya diversifolia Blume

Hoya diversifolia Blume subsp. el-nidicus (Kloppenb.) Kloppenb. sub: Hoya

diversifolia Blume

Hoya el-nidicus Kloppenb. sub: Hoya diversifolia Blume

Hoya elegans sensu (Blume) Boerl. sub: Hoya multiflora Blume

Hoya elliptica Hook.f.

Hoya elmeri Merr.

Hoya esculenta Tsiang sub: Hoya diversifolia Blume

Hoya fauziana Rodda, Simonsson & A.L.Lamb

Hoya finlaysonii Wight

Hoya forbesii King & Gamble

Hoya fraterna Blume sub: Hoya coriacea Blume

Hoya fuscomarginata N.E.Br. sub: Hoya vitellina Blume

Hoya gildingii Kloppenb.

Hoya glabra Schltr.

Hoya globiflora Ridl. sub: Hoya verticillata (Vahl) G.Don

Hoya hamiltoniorum A.L.Lamb, Gavrus, Emoi & Gokusing

Names and types of Bornean Hoya

Hoya heterophylla nom. nud. sub: Hoya diversifolia Blume

Hoya hookeriana Wight sub: Hoya verticillata (Vahl) G.Don

Hoya ignorata T.B.Tran, Rodda, Simonsson & Joongku Lee

Hoya imperialis Lindl.

Hoya imperialis Lindl. var. rauschii Regel sub: Hoya imperialis Lindl.

Hoya javanica Boerl. sub: Hoya multiflora Blume

Hoya jiewhoeana Rodda, A.L.Lamb & Gokusing

Hoya kastbergii Kloppenb.

Hoya kipandiensis Gavrus, A.L.Lamb, Emoi & Gokusing

Hoya kloppenburgii T.Green

Hoya lacunosa Blume

Hoya lambii T.Green

Hoya lanceolata nom. nud. sub: Hoya verticillata (Vahl) G.Don

Hoya lasiantha (Korth. ex Blume) Miq.

Hoya latifolia (Wight) G.Don

Hoya liangii Tsiang sub: Hoya diversifolia Blume

Hoya linusii A.L.Lamb, Gavrus & Emoi sub: Hoya phyllura O.Schwartz

Hoya macrophylla Wight sub: Hoya latifolia (Wight) G.Don

Hoya mappigera Rodda & Simonsson

Hoya medinillifolia Rodda & Simonsson

Hoya meredithii T. Green

Hoya mindanaensis Elmer sub: Hoya coriacea Blume

Hoya mindorensis Schltr.

Hoya mindorensis subsp. superb a Kloppenb. sub: Hoya elmeri Merr.

Hoya minutiflora Rodda & Simonsson

Hoya mitrata Kerr

Hoya monetteae T.Green

Hoya multiflora Blume

Hoya nabawanensis Kloppenb. & Wiberg

Hoya nicobarica R.Br. ex J.Traill sub: Hoya verticillata (Vahl) G.Don

Hoya nuttiana Rodda & Simonsson

Hoya nyhuusiae Kloppenb.

Hoya obscura Elmer ex C.M.Burton

Hoya obscura Elmer sub: Hoya obscura Elmer ex C.M.Burton

Hoya obscurinervia Merr. sub: Hoya verticillata (Vahl) G.Don

Hoya obtusifolia Wight

Hoya occlusa Ridl. sub: Hoya coriacea Blume

Hoya omlorii (Livsh. & Meve) L.Wanntorp & Meve

Hoya opposita (Vahl.) G.Don sub: Hoya verticillata (Vahl) G.Don

Hoya orbiculata Wall, ex Wight & Arn. sub: Hoya diversifolia Blume

Hoya pallida Lindl. sub: Hoya verticillata (Vahl) G.Don

Hoya parasitica Wall, ex Wight sub: Hoya verticillata (Vahl) G.Don

Hoya parasitica Wight var. geoffrayi Costantin sub: Hoya verticillata (Vahl) G.Don

Hoya parasitica Wight var. spirei Costantin sub: Hoya verticillata (Vahl) G.Don

Hoyapersicinicoronaria Shao Y.He & P.T.Li sub: Hoya diversifolia Blume

Hoya phyllura O.Schwartz

Card. Bull. Singapore 69(1) 2017

Hoya plicata King & Gamble subsp. rundumensis T. Green sub: Hoy a rundumensis

(T.Green) Rodda & Simonsson

Hoyapottsii J.Traill sub: Hoya verticillata (Vahl) G.Don

Hoya pottsii J.Traill var. angustifolia Tsiang & P.T.Li sub: Hoya verticillata (Vahl)

G.Don

Hoya pubera Blume

Hoya ranauensis T. Green & Kloppenb.

Hoya recurvula Kloppenb. sub: Hoya verticillata (Vahl) G.Don

Hoya recurvula Kloppenb. subsp. bokorensis Kloppenb. & Yap sub: Hoya verticillata

(Vahl) G.Don

Hoya refracta nom. nud. sub: Hoya multiflora Blume

Hoya reticulata Costantin sub: Hoya multiflora Blume

Hoya retrorsa Gavras, A.L.Lamb, Emoi, Gokusing

Hoya revoluta Wight ex Hook.f.

Hoya ridleyi King & Gamble sub: Hoya verticillata (Vahl) G.Don

Hoya rintzii Rodda, Simonsson & S.Rahayu

Hoya rundumensis (T.Green) Rodda & Simonsson

Hoya ruthiae Rodda

Hoya sammannaniana A.L.Lamb, Gavras, Emoi, Gokusing

Hoya scortechinii King & Gamble

Hoya sigillatis T.Green subsp. sigillatis

Hoya sigillatis T.Green subsp. paitanensis Gavras, A.L.Lamb, Emoi, Gokusing

Hoya sipitangensis Kloppenb. & Wiberg

Hoya spartioides (Benth.) Kloppenb.

Hoya telosmoides Omlor

Hoya teysmanniana Miq. sub: Hoya obtusifolia Wight

Hoya ubudensis Kloppenb. & Yap sub: Hoya verticillata (Vahl) G.Don

Hoya undulata S.Rahayu & Rodda

Hoya vacciniiflora O.Schwartz

Hoya verticillata (Vahl) G.Don

Hoya vitellina Blume

Hoya wallichii (Wight) C.M.Burton

Hoya walliniana Kloppenb. & Nyhuus

Hoya waymaniae Kloppenb.

Hoya wibergiae Kloppenb. sub: Hoya verticillata (Vahl) G.Don

Hoya wongii Rodda, Simonsson & L.Wanntorp

Hoya yapiana Kloppenb. sub: Hoya sipitangensis Kloppenb. & Wiberg

Hoya zollingeriana Miq. sub: Hoya diversifolia Blume

Otostemma lacunosum (Blume) Blume sub: Hoya lacunosa Blume

Physostelma wallichii Wight sub: Hoya wallichii (Wight) C.M.Burton

Plocostemma lasianthum Korth ex Blume sub: Hoya lasiantha (Korth. ex Blume)

Miq.

Sperlingia opposita Vahl sub: Hoya verticillata (Vahl) G.Don

Sperlingia tetraphylla nom. nud. sub: Hoya verticillata (Vahl) G.Don

Sperlingia verticillata Vahl sub: Hoya verticillata (Vahl) G.Don

Names and types of Bornean Hoya

ACKNOWLEDGEMENTS. This study is part of an on-going research project on the

systematics of Asian Apocynaceae. Financial support was received from the National Parks

Board (Singapore) which sponsored numerous herbarium study trips to Asian and European

institutions. The curators of A, BISH, BM, BO, BRUN, CGE, E, FI, G, HITBC, HN, IBSC,

K, KEP, KUN, L, OXF, P, SAN, SAR, SING, SNP, TO, UC, US, VN, WRSL and Z herbaria

are thanked for allowing access and/or for allowing loans or high quality images of herbarium

specimens. The librarians of the Singapore Botanic Gardens, Royal Botanic Gardens Kew

and Royal Botanic Garden Edinburgh have helped to find often obscure references. Elisabeth

Liljebjorn spent many hours scanning all the issues of the journals Fraterna and Hoyan as well

as creating PDFs of numerous papers. Lastly Anthony Lamb and Anthea Phillips are thanked

for their continuous support for my work on Bornean Hoya.

References

Bennett, J.J. (1838) Plantae Javanicae Rariores Part 1, page [1]—104. London: H. Allen et

socios.

Blume, C.L. (1823). Catalogus. Batavia: Lands Drukkerji

Blume, C.L. (1826). Bijdragen tot de Flora van Nederlandsch Indie , vol. 16. Batavia: Lands

Drukkerji.

Blume, C.L. (1849). Rumphia , vol. 4. Leiden, Amsterdam: C.G. Sulpke.

Backer, C.A. & Bakhuizen van den Brink, R.C. (1965). Hoya In: Flora of Java, vol. 2, pp.

266-271. Noordhoff, Groningen: Noordhoff.

Bentham, G. (1880). Astrostemma spartioides Benth. In: Hooker, J.D. (ed) Hooker’s leones

Plantarum, vol. 14, pp. 7-8, t. 1311. London: Williand and Norgate.

Decaisne, J. (1844) Hoya In: Candolle, A.L.P.P. de (ed) Prodromus Systematis Naturalis Regni

Vegetabilis, vol. 8. Paris: Fortin, Masson et sociorum.

Forster, PI. (1991). Clarification of the author citations, publication places and types for recently

described Hoya species (Asclepiadaceae) from the Philippines. Asklepios 5: 100-101.

Forster, PI. & Liddle, D.J. (1991). Variation in Hoya australis R.Br ex Traill (Asclepiadaceae).

Austrobaileya 3: 503-521.

Forster, PI. & Liddle, D.J. (1992a). Asclepiad Profile no. 2. Asklepios 56: 24-26.

Forster, PI. & Liddle, D. J. (1992b). Taxonomic studies on the genus Hoya R.Br. (Asclepiadaceae)

in Papuasia 4. Typification and synonymy of Hoya revoluta Wight ex J.D. Hook.

Austrobaileya 3: 638-639.

Forster, PI. & Liddle, D.J. (1992c). Synonymy of Hoya pottsii Traill. Austrobaileya 3: 635.

Hill, K.D. (1988). A revision of Hoya (Asclepiadaceae) in Australia. Telopea 3(2): 241-255.

King, G. & Gamble, J. (1908). Hoya. In: Flora of the Malayan Peninsula, J. Asiat. Soc. Bengal,

Pt. 2. Natural History. 74: 559-580.

Kleijn, D. & Van Donkelaar, R. (2001). Notes on the taxonomy and ecology of the genus Hoya

(Asclepiadaceae) in Central Sulawesi. Blumea 46: 457-483.

Kloppenburg, R.D. (2012). Two new hoyas by Dale Kloppenburg: Hoya samarensis & Hoya

australis ssp. nathalieae. Asklepios 113: 27-29.

Lamb, A. & Rodda, M. (2016). A Guide to Hoyas of Borneo. Kota Kinabalu: Natural History

Publications Borneo.

Lamb, A., Gavrus, A., Emoi, B. & Gokusing, L. (2014). The hoyas of Sabah, a commentary

with seven new species and a new subspecies. Sandakania 19: 1-89.

Liddle, D.J. & Forster, PI. (2008). Notes on some commonly cultivated Hoya species from

Australia, Papuasia and Melanesia: Hoya australis R.Br. ex Traill. Asklepios 102: 3-15.

Lindley, J. (1826). Hoya pallida. Bot. Reg. 11: t. 951.

Card. Bull. Singapore 69(1) 2017

Lindley, J. (1839). Hoya coriacea. Edwards’s Bot. Reg. 25: t.18.

Lindley, J. (1846). New garden plant. Hoya imperialis. Edwards’s Bot. Reg. 32: t. 68.

Lucas, A.M. (2008). Disposing of John Lindley’s library and herbarium: the offer to Australia.

Arch. Nat. Hist. 35(1): 15-70.

McNeill, J., Buck, W.R., Demoulin, V., Greuter, W., Hawkworth, D.L., Herendeen, P.S., Knapp,

S., Marhold, K., Prado, J., PrudTiomme van Reine, W.F., Smith, G.R, Wiersema, J.H. &

Turland, N.J. (2012). International Code of Nomenclature for algae, fungi, and plants

(Melbourne Code). Regnum Vegetabile 154. Konigstein: Koeltz Scientific Books.

Merrill, E.D. (1918). Species Blancoanae. Manila: Manila Bureau of Printing.

Merrill, E.D. (1921). A bibliographic enumeration of Bornean plants. J. Straits Branch Roy.

Asiat. Soc., Special Number. Singapore: Fraser & Neave Ltd.

Miquel, F.E.W. (1857). Flora van Nederlandsch Indie, vol. 2. Amsterdam: C. G. van der Post.

Nicholas, A. (1992). The Asclepiadaceous works of Rudolf F. Schlechter (1872-1925).

Willdenowia 22: 215-264.

Noltie, H.J. (2005). The Botany of Robert Wight. Regnum Vegetabile, vol. 145. Ruggell: A.R.G.

Gantner Verlag.

Nutt, P. (2001). Checkliste der Gattung Hoya auf Borneo (Apocynaceae-Asclepiadoideae).

Diploma thesis, Westfalische Wilhelms-Universitat Munster.

Rahayu, S., Meve, U. & Rodda, M. (2015). Hoya undulata (Apocynaceae, Asclepiadoideae), a

new myrmecophytic species from Borneo, and typification of H. darwinii. Gard. Bull.

Singapore 67(1): 85-94.

Rintz, R.E. (1978). The Peninsular Malaysian species of Hoya (Asclepiadaceae). Malayan Nat.

J. 30: 467-522.

Rodda, M. (2012). (2074) Proposal to conserve the name Hoya mitrata against H. wallichiana

{Apocynaceae, Asclepiadoideae). Taxon 61: 685-686.

Rodda, M. (2015a). Two new species of Hoya R.Br. (Apocynaceae, Asclepiadoideae) from

Borneo. PhytoKeys 53: 83-93.

Rodda, M. (2015b). A new typification of Hoya zollingeriana (Apocynaceae, Asclepiadoideae).

Gard. Bull. Singapore 67(2): 305-307.

Rodda, M. & Ercole, E. (2014). Hoya papaschonii (Apocynaceae: Asclepiadoideae), a new

species from southern Thailand with a peculiar corona. Phytotaxa 175(2): 97-106.

Rodda, M. & Simonsson, N. (2011a). Hoya devogelii (Apocynaceae: Asclepiadoideae), a new

species from Sarawak, Borneo. Webbia 66(1): 33-38.

Rodda, M. & Simonsson, N. (2011b). Hoya medinillifolia (Apocynaceae: Asclepiadoideae), a

new species from lowland forest of Sarawak, Borneo. Webbia 66(2): 149-154.

Rodda, M. & Simonsson Juhonewe,N. (2013a). Hoyanuttiana (Apocynaceae: Asclepiadoideae),

a new species from Sarawak, Malaysian Borneo. Phytotaxa 140(1): 56-60.

Rodda, M. & Simonsson Juhonewe, N. (2013b). The taxonomy of Hoya micrantha and Hoya

revoluta (Apocynaceae, Asclepiadoideae). Webbia 68: 7-16.

Rodda, M., Simonsson Juhonewe, N. & Ercole, E. (2014). Hoya corymbosa (Apocynaceae:

Asclepiadoideae), a new unusual species from Sabah, Borneo and its systematic position

based on phylogenetic analyses. Syst. Bot. 38: 1125-1131.

Rodda, M., Simonsson Juhonewe, N. & Middleton, D.J. (2016). The taxonomic status of the

presumed extinct Singaporean Hoya wallichii (Apocynaceae: Asclepiadoideae). Gard.

Bull. Singapore 68: 175-187.

Rumphius, G.E. (1747). Herbarium amboinense, vol. 4. Amsterdam: Apud Fransicum

Changuion, Joannem Catuffe, Hermannum Uytwerf.

Smith, A.C. (1988). Hoya. In: Flora Vitiensis Nova, A New Flora of Fiji, vol. 4, pp. 116-123.

Honolulu: SB Printers Inc.

Names and types of Bornean Hoya

Schlechter, R. (1908). Beitrage zur Kenntnis der Asclepiadaceen des Monsum-Gebietes. Bot.

Jahrb. Syst. 40 (Beibl. 92): 1-45.

Stafleu, F.A. (1970). Benjamin Delessert and Antoine Lasegue. Taxon 19: 920-936.

Stafleu, F.A. & Cowan, R.S. (1976). Taxonomic Literature 1, 2nd ed. Utrecht: Bohn, Scheltema

& Holkema.

Steam, W.T. (2008). Botanical Latin. Singapore: KHL Printing Co.

Traill, J. (1830). Accounts and descriptions of the several plants belonging to the genus Hoya

which are cultivated in the garden of the Horticultural Society at Chiswick. Trans. Hort.

Soc. London 7: 16-30.

Van Steenis-Kruseman, M.J. (1950). Malaysian plant collectors and collections: Being a

cyclopaedia of botanical exploration in Malaysia and a guide to the concerned literature

up to the year 1950. In: Flora Malesiana, Ser. 1, Spermat. 1, pp. 5-606. Jakarta:

P.Noordhoff-Kolff N.V.

Veldkamp, J.F., van Donkelaar, R. & Kloppenburg, R.D. (1995). The identity of Sperlingia

Vahl (Asclepiadaceae). Blumea 40: 425-428.

Wanntorp, L. & Forster, PI. (2007). Phylogenetic relationships between Hoya and the monotypic

Genera Madangia , Absolmsia, and Micholitzia (Apocynaceae, Marsdenieae): Insights

from flower morphology. Ann. Missouri Bot. Gard. 94: 36-55.

Wanntorp, L., Kocyan, A. & Renner, S.S. (2006a). Wax plants disentangled: a phylogeny

of Hoya (Marsdenieae, Apocynaceae) inferred from nuclear and chloroplast DNA

sequences. Molec. Phylogenet. Evol. 39: 722-733.

Wanntorp, L., Kocyan, A., van Donkelaar, R. & Renner, S.S. (2006b). Towards a monophyletic

Hoya (Marsdenieae, Apocynaceae): inferences from the chloroplast trnL region and the

rbcL-atpB spacer. Syst. Bot. 31: 586-596.

Wanntorp, L., Gotthardt, K. & Muellner, A.N. (2011). Revisiting the wax plants {Hoya,

Marsdenieae, Apocynaceae): Phylogenetic tree using the matK gene and psbA-trnH

intergenic spacer. Taxon 60: 4-14.

Wanntorp, L., Grudinski, M., Forster, P.I., Muellner-Riehl, A.N. & Grimm, G.W. (2014). Wax

plants {Hoya, Apocynaceae) evolution: Epiphytism drives successful radiation. Taxon

63: 89-102.

Wight, R. (1834). Contributions to the botany of India. London: Parbury, Allen & Co.

Gardens’ Bulletin Singapore 69(1): 67-73. 2017

doi: 10.3850/S2010098116000046

Novitates Bruneienses, 8. Macrosolen brunsing

(Loranthaceae), a new hemiparasitic shrub

from Brunei Darussalam

Y.W. Low 1 , A.K. Muhammad Ariffin 2 , A.A. Joffre 2 & D. Duratul Ain 2

herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

lo w_y ee_wen@nparks. gov. sg

2 Brunei National Herbarium, Forestry Department,

Ministry of Industry and Primary Resources, Jalan Menteri Besar,

Berakas, BB3910 Brunei Darussalam

ABSTRACT. Macrosolen brunsing Y.W.Low & Ariffin is described and illustrated here as a new

species of aerial hemiparasite based on two collections from the Ladan Hills Forest Reserve,

Tutong, Brunei Darussalam. The new species differs from all Macrosolen taxa enumerated in

Borneo by its distinct linear leaves ((4—)8—14.5 cm long, 0.1—0.2(—0.25) cm wide).

Keywords. Borneo, endemic, linear-leaved, Malesia, new species

Introduction

The presence of aerial hemiparasitic shrubs of the Loranthaceae in tropical rainforests

is often overlooked, even when they are flowering, as they are generally concealed

high-up in the canopy of their host. Although many species produce very spectacular

inflorescences, often their presence is only revealed by their withered flowers littered

on forest floor, usually around the base of the trunk of their host. On a recent botanical

excursion to the Nyamokning Dam construction site in the Ladan Hills Forest Reserve,

a sterile but striking hemiparasitic shrub with linear leaves (Y.W. Low et al. LYW1059 )

was collected from a species of Hydnocarpus Gaertn. (Achariaceae) host growing on

a ridge at about 150 m asl. Later, the same hemiparasite was spotted in flower and fruit

(Y.W. Low et al. LYW 1081) on a species of Mallotus Lour. (Euphorbiaceae) host, just

above the campsite on the same trip. These two collections were closely examined in

BRUN and SING (herbaria acronyms follow Thiers, (continuously updated)).

The Ladan Hills plant readily keys to Macrosolen Blume in the Flora Malesiana

account for Loranthaceae (Barlow, 1997). Based on fieldwork carried out for the

Brunei Checklist Project, Kirkup (1996) enumerated nine taxa, of which two were

unidentified. These are Macrosolen beccarii Tiegh. ex Becc., M. borneanus Danser,

M. cochinchinensis (Lour.) Tiegh., M. crassus Danser, M. curvinervis Danser, M. aff.

formosus (Blume) Miq., M. retusus (Jack) Miq., M. sp. A and M. sp. B. Of the nine taxa

listed by Kirkup (1996), only six were recognised by Barlow (1997) in his revision of

the genus for Malesia, as M. borneanus is considered a synonym of M. beccarii , and

Card. Bull. Singapore 69(1) 2017

M. curvinervis is a synonym of M. macrophyllus (Korth.) Miq. However, we were

unable to match the Ladan Hills taxon to any of the Macrosolen species preserved

in BO, BRUN and SING. Barlow (1997) enumerated 24 species of Macrosolen in

Malesia, and a survey of all the species listed in his revision reveals that only one

taxon, Macrosolen brevitubus Barlow was reported to have linear leaves, but only at a

juvenile stage. The leaf width measurement recorded for Macrosolen brevitubus at the

juvenile stage was 0.5-1 cm wide (Barlow, 1995, 1997), while the Ladan Hills taxon

differs in having much narrower leaf width at 0.1-0.2(-0.25) cm wide. Hence, the

Ladan Hills population which maintains this extremely narrow leaf even when mature

is distinct and described here as a new species.

Materials and Methods

A review of all the Bornean Macrosolen species was conducted based on herbarium

specimens preserved in BO, BRUN and SING. Conventional methods employed in

herbarium taxonomy were applied in this study. All measurements for vegetative

characters were taken from dried herbarium specimens, w hil e measurements of

reproductive structures were taken from materials preserved in spirit collection.

Photographic documentation was taken in the field. Type materials present at SING

were examined, as well as type images of all Malesian Macrosolen species available

at JSTOR® Global Plants website (http://plants.jstor.org). Botanical terms used in this

study largely follow Beentje (2012). Provisional conservation assessments are made

using the methodology proposed by IUCN (2012).

New species

Macrosolen brunsing Y.W.Low & Ariffin, sp. nov.

Similar to Macrosolen brevitubus Barlow but differs in having narrow linear leaves

((4-)8-14.5 cm long, 0.1-0.2(-0.25) cm wide), and inflorescence a raceme of two

opposite pairs of flowers. - TYPE: Brunei, Tutong District, Rambai, Ladan Hills

Forest Reserve, Nyamokning Dam, BRUN-SING botanical exploration campsite on

the edge of forest near water body, lowland mixed dipterocarp forest on yellow sandy

clay soils, 04°23'35.6"N 114°48'58.6"E, 75 m asl, 22 August 2016, Y.W. Low, M.I. Siti

Nur Bazilah, A.K. Muhd. Ariffin, A. Watu, E. Jangarun, P. Azlan, K. Muhd. Khairul

Nizam & Z.A. Muhd. Wafiuddin LYW 1081 (holotype BRUN (including spirit material

as part of a single specimen); isotypes E, K, L, SAN, SAR, SING [[SINGO166300] &

spirit material [SING0202921]). (Fig. 1, 2)

Aerial stem hemiparasitic shrub with epicortical runners bearing secondary haustoria,

glabrous. Leaves opposite, rarely sub-opposite; petioles 1-2 mm long; lamina

narrowly linear, (4—)8—14.5 cm long, 0.1—0.2(—0.25) cm wide, base truncate, apex

narrowly acute, coriaceous; midrib inconspicuous and sunken above, prominent and

A new Macrosolen from Brunei Darussalam

Fig. 1 . Macrosolen brunsing Y.W.Low & Ariffin displaying aggressive epicortical runners

bearing secondary haustoria along the branch of a Mallotus sp. (Euphorbiaceae). From type

material Y.W. Low et al. LYW1081. (Photo: Y.W. Low)

Card. Bull. Singapore 69(1) 2017

Fig. 2. Macrosolen brunsing Y.W.Low & Ariffin. A. Pendulous flowering branch showing

distichous almost needle-like linear leaves and a tenninal inflorescence. B. Close-up of mature

flower buds that somewhat resemble bowling pins. C. Close-up of open flowers. D. Close-up

of fruits. All from type Y.W. Low et al. LYW1081. (Photos: Y.W. Low)

raised below; secondary and tertiary veins inconspicuous on both sides. Inflorescences

axillary or rarely terminal, a raceme of 2 opposite decussate pairs of flowers; peduncle

1-3 mm long; pedicels c. 1.5 mm long. Bracts 3 under each flower, connate and

forming a cup. Ovary cylindrical, c. 0.3 cm long, mostly pale green except for the

darker green apex, smooth; calyx limb cylindrical, c. 1 mm long, truncate. Corolla

6-merous, gamopetalous, in mature bud with 6 prominent keels or wings below the

A new Macrosolen from Brunei Darussalam

Fig. 3. Habitat of Macrosolen brunsing Y.W.Low & Ariffin in the lowland mixed dipterocarp

forest of the Nyamoking Dam, Ladan Hills Forest Reserve. Foreground is the blue roof campsite

at the edge of lake. (Photo: A.K. Muhammad Ariffin)

point of reflexion along the sutures of the lobes extending above the neck to the apex

of the bud, somewhat zygomorphic, 17-21 mm long, inflated in the lower part, winged

below the middle, clavate above a long neck and acute at apex, mostly red with a

thick black band at neck and a black spot at apex; tube in open flowers 9-11 mm long

with the narrowly spathulate lobes reflexed c. 3 mm above the margin of the throat.

Stamens 6, inserted slightly below the throat of the corolla tube, at the middle of the

corolla lobes, greenish yellow, basifixed, immobile; filaments c. 5 mm long, c. 3 mm

protruding out from the corolla-tube mouth; anthers 3—4 mm long, base obtuse, apex

acute. Style c. 1.8 cm long, greenish yellow, glabrous; stigma club-shaped, c. 1 mm

long, red. Fruit broadly prolate to globose, c. 0.8-0.9 cm long, 0.6-0.7 cm wide,

mostly bright yellow except for the green apex, smooth.

Additional specimen examined. BORNEO: Brunei: Tutong District, Rambai, Ladan Hills

Forest Reserve, Nyamokning Dam, Hulu Sungai Tutong, ridge of lowland mixed dipterocarp

forest on yellow sandy clay soils, 04°20T4.2"N 114°49'076"E, 152 m asl, 20 Aug 2016, Y.W.

Low, M.I. Siti Nur Bazilah, A.K. Muhd. Ariffin, A. Watu, P. Azlan, K. Muhd. Khaind Nizam &

Z.A. Muhd. Wafiuddin LYW1059, sterile (BRUN, SING).

Distribution and Habitat. Macrosolen brunsing Y.W.Low & Ariffin is so far known

only from the Ladan Hills Lorest Reserve, in the lowland mixed dipterocarp forest on

yellow sandy clay soils. (Lig. 3, 4)

Card. Bull. Singapore 69(1) 2017

Etymology. The epithet brunsing is composed by merging two herbaria acronyms

together, namely BRUN (the Brunei National Herbarium) and SING (Herbarium of

the Singapore Botanic Gardens). This new species is named for the two herbaria to

celebrate the close working relationship between the two herbaria that can be traced

back to the early 80’s. The on-going MoU programme “The Botanical Survey of Brunei

Darussalam ’ continues this cooperation.

Proposed IUCN Conservation Assessment. Macrosolen brunsing Y.W.Tow & Ariffin

is so far known only from two locations in the almost complete Nyamokning Dam,

Tadan Hills Forest Reserve. The IUCN status proposed here for Macrosolen brunsing

is Data Deficient (DD) as the species is only know from two collections, both within

the same locality. Generally, the Tadan Hills Forest Reserve is considered to be

poorly explored botanically. In this area at least Macrosolen brunsing is not under any

imminent threats due to human activity as the species is protected in a forest reserve.

The Brunei Government officially announced that logging will be phased out in all

forest reserves in 2014 (Hamit, 2014).

A new Macrosolen from Brunei Darussalam

ACKNOWLEDGEMENTS. We are grateful to Hajah Noralinda binti Haji Ibrahim, the Acting

Director of Forestry Department, Brunei Darussalam for providing us with field support and

assistance for this study. Special thanks to Roslinah binti Haji Mohsin for her encouragement

and continuous support and assistance to this project. Various staff members of the Brunei

National Herbarium (BRUN) have kindly helped: Mediana Kampong and Muhd. Wafiuddin

Zainol Ariffin assisted with herbarium materials and the BRAHMS database; Watu Awok, Azlan

Pandai, Jangarun Eri, Muhd. Khairul Nizam Kampong and Rauzaidi Mureh provided excellent

assistance in the field, as well as maintaining the ex-situ research collection. We sincerely

thank Dr Joeni Setijo Rahajoe (Keeper of BO) for providing us with permission and access to

the Loranthaceae collection at BO; Dr Atik Retnowati and Ridha Mahyuni provided excellent

support and assistance at BO. Siti Nur Bazilah Mohamed Ibrahim (SING) provided valuable

field assistance on the botanical excursion at the Nyamokning Dam, as well as herbarium and

curatorial support in Singapore. Research funding for YWL is provided by the National Parks

Board, Singapore through the Singapore Botanic Gardens. Dr Robert Faden (US) and Dr Jan-

Frits Veldkamp (L) provided nomenclatural comments on the proposed specific epithet; and

Dr Max M.J. van Balgooy (L) confirmed that the new species is the first linear-leaved taxon

when mature of the genus Macrosolen that he was aware of. This research collaboration is

conducted under the MoU programme “ The Botanical Survey of Brunei Darussalam ’ between

the Brunei Forestry Department, Ministry of Industry and Primary Resources and the National

Parks Board, Singapore.

References

Barlow, B.A. (1995). New and noteworthy Malesian species of Loranthaceae. Blumea 40:

15-31.

Barlow, B.A. (1997). Loranthaceae. In: Kalkman, C., Kirkup, D.W., Nooteboom, H.P, Stevens,

P.F. & de Wilde, W.J.J.O. (eds) Flora Malesiana, ser. 1, Seed Plants, vol. 13, pp. 209-

401. Leiden: Rijksherbarium/Hortus Botanicus.

Beentje, H. (2012). The Kew Plant Glossary: An Illustrated Dictionary of Plant Terms. Revised

edition. London: Royal Botanic Gardens, Kew.

Hamit, R. (2014). Logging no longer allowed in forest reserves: MIPR. Brunei Times , 23

March 2014, p. A3.

IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. 2nd ed. Gland, Switzerland,

and Cambridge, UK: IUCN.

Kirkup, D.W. (1996). Loranthaceae. In: Coode, M.J.E., Dransfield, J., Forman, L.L., Kirkup,

D.W. & Said, I.M. (eds) A Checklist of the Flowering Plants and Gymnosperms of

Brunei Darussalam , pp. 176-182. Brunei Darussalam: Ministry of Industry and Primary

Resources.

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria

and associated staff. New York Botanical Garden’s Virtual Herbarium, http://sweetgum.

nybg.org/ih/. Accessed 1 Sep. 2016.

Gardens’ Bulletin Singapore 69(1): 75-80. 2017

doi: 10.3850/S2010098116000058

A noteworthy Dendrocalamus (Poaceae: Bambusoideae)

from Sumatra, Indonesia

I Putu Gede P. Damayanto 1 & E.A. Widjaja 2

herbarium Bogoriense, Research Center for Biology, Indonesian Institute of Sciences,

Jl. Raya Jakarta-Bogor Km. 46, Cibinong 16911, Indonesia

2 Present address RT/RW 03/01, Kp. Cimoboran, Ds. Sukawening,

Dramaga Sub District, Bogor District, Indonesia

parlida.damayanto.tab @ gmail.com

ABSTRACT. There are eight species of Dendrocalamus Nees ( Dendrocalamus asper (Schult.)

Backer, D. bengkalisensis Widjaja, D. brandisii (Munro) Kurz, D. buar Widjaja, D. giganteus

Munro, D. bait Widjaja, D. latiflorus Munro, D. membranaceus Munro), wild or cultivated,

already recorded for Sumatra. A new species is added to this list, Dendrocalamus luteus

Damayanto & Widjaja, which grows wild in the Indonesia Ecosystem Restoration area in

Sumatra’s Jambi and South Sumatra Provinces. The new species is described, illustrated, and

included in an identification key to all Sumatran Dendrocalamus species.

Keywords. Bambusoideae, Dendrocalamus luteus , new species, Sumatra

Introduction

According to Widjaja et al. (2014) Sumatra has the highest bamboo diversity in

Indonesia, with at least 80 species, of which 32 species are endemic to the island.

Three of these Sumatran endemic species are in the genus Dendrocalamus Nees

(Dendrocalamus bengkalisensis Widjaja, D. buar Widjaja, D. bait Widjaja), another

species of Dendrocalamus is newly described here, and five other species (D. asper

(Schult.) Backer, D. brandisii (Munro) Kurz, D. giganteus Munro, D. latiflorus Munro,

D. membranaceus Munro) are found only in cultivation.

An identification key to these nine Dendrocalamus spp. in Sumatra is presented.

Identification Key to Dendrocalamus species in Sumatra

la Dominant primary branch axes at middle to distal parts of culm growing long and

overarching onto neighbouring trees.2

lb. Dominant primary branch axes not especially elongated, never overarching onto

neighbouring trees.5

2a. Leaf blade pubescent on lower surface.3

2b. Leaf blade glabrous on lower surface.4

Card. Bull. Singapore 69(1) 2017

3a Culm sheath apex with white wax and white to brown hairs at the base, culm

sheath margin with white to brownish hairs; culm below nodes covered with

brownish and whitish hairs and white waxiness; leaf midrib prominent and leaf

base yellow. D. luteus

3b Culm sheath completely covered by white hairs and without wax, culm sheath

margin glabrous; culm below nodes glabrous; leaf midrib not prominent and leaf

base not yellow. D. bengkalisensis

4a Culm sheath auricles rim-like and crisped to blade base; ligules with 4-15 mm

long bristles. D. buar

4b. Culm sheath auricles rounded lobes; ligules glabrous. D. bait

5a. Culm sheath auricles rim-like. D. latiflorus

5b. Culm sheath auricles small or rounded lobes, with wavy folds or crisped to blade

base.6

6a. Lower culm of young culm with dense velvety brown hairs. D. asper

6b. Lower culm of young culm without velvety hairs.7

7a Culm sheath auricles crisped to the blade base; culm sheath ligules more than 1

mm high. D. giganteus

7b Culm sheath auricles small lobes with wavy folds; culm sheath ligules up to 0.5

mm high.8

8a Culm sheath auricle glabrous; blade usually shorter than half of the sheath length

. D. brandisii

8b Culm sheath auricle with long bristles; blade usually exceeding half of the sheath

length. D. membranaceus

Dendrocalamus luteus Damayanto & Widjaja, sp. nov.

Dendrocalamus luteus resembles D. bengkalisensis Widjaja (Widjaja, 1997) but differs

in having white to brown culm sheath hairs, lobe-like leaf sheath auricles, and brownish

and whitish hairs below the culm node (in contrast, D. bengkalisensis has consistently

white culm sheath hairs, rounded 0.5-1 mm high leaf-sheath auricles, and is glabrous

below the culm node). - TYPE: Sumatra, Jambi Province, Batanghari Subdistrict, Sei

Bahar (now Muaro Jambi District, Sei Bahar Subdistrict), 8 July 2006 Widjaja 8128

(holotype BO [one specimen over three sheets], accession No. BO-1934223 (shoot),

BO-1934220 (leafy branch), BO-1934224 (inflorescence); isotype K). (Fig. 1)

Culm yellowish green to yellow; brownish and whitish hairs and white waxiness

below the node when young, which disappears at maturity. Branch complement with

a dominant primary branch which becomes very long (up to 5 m) and overarches onto

neighbouring trees or even droops to the ground; culm and branch tips that droop to

New Dendroccilamus from Sumatra

2 cm

4 cm

Fig. 1. Dendroccilamus luteus Damayanto & Widjaja. A. Culm shoot. B. Detail of upper part

of culm sheath, showing rounded auricle with bristles. C. Leafy branch. D. Leaf sheath-blade

junction, showing auricle with bristles. E. Inflorescence. F. Glume. G. Lemma. H. Palea. I.

Pistil surrounded by six stamens with free filaments, without lodicules. J. Detail of pistil.

Drawn by Wahyudi Santoso from Widjaja 8128 (BO).

Card. Bull. Singapore 69(1) 2017

the ground occasionally rooting and forming a new clump. Culm sheaths 15.9-16.9

x 14.5-15.4 cm, orange in young shoots, covered by white wax on the upper part and

with dense appressed white to brownish hairs at the base, sheath margin with white

hairs along the apical part which are gradually replaced by brownish hairs (up to 0.3

cm long) toward the basal portion, eventually deciduous; auricles rounded, 0.4-0.5 cm

high, bristles many and long, 2-2.8 cm long; ligules dentate, 0.2-0.4 cm high, bristles

few, 0.4-0.6 cm long; blade spreading then reflexed, narrowly lanceolate, 10.7-12.4 x

2-2.7 cm. Leaf 23.2-253 x 2.3-4.5 cm, midrib base orange to yellow, pubescent on

lower surface; auricles lobe-like, less than 0.5 mm, bristles 0.9-1.7 cm long; ligules

entire, 1-2 mm high, glabrous. Inflorescences with 1-2 florets in each pseudospikelet,

without a terminal vestigial flower; empty glumes 2; lemma 6-7 mm high; palea

keeled at the apex, 8-8.5 mm long; stamens six, up to 4 mm long, anthers yellow with

free filaments; ovary obovoid with thickened and hairy apex, produced into long hairy

style 6-7 mm long and ending in a white plumose stigma.

Distribution. Sumatra, Jambi Province and South Sumatra Province, in the Indonesian

Ecosystem Restoration area.

Ecology. This species forms the dominant growth in secondary forest along the road at

around 60 m asl. Following logging of the forest in this area in 1991, this species has

become abundant, with a suggestion from the local people that it is being dispersed by

elephants.

Notes. The epithet luteus (meaning yellow) refers to the colour of the shoot, culm, base

of the leaf blade and midrib, and anthers. Table 1 compares D. luteus to its closely

related species.

Vernacular names and uses. Unknown.

Additional specimens examined. INDONESIA: Jambi Province: Batanghari District,

Bajubang Subdistrict, Hutan Harapan, Jembatan Jengkol, 3 Apr 2013, Wardi et al. BOHK 402

(BO); Batanghari District, Bajubang Subdistrict, Harapan Rainforest, 500 m before Jembatan

Jengkol (Jengkol bridge) from office, 02°10'23"S 103°19'54"E, 2 Mar 2016, Damayanto &

Widjaja 02 (BO), Damayanto & Widjaja 03 (BO), Damayanto & Widjaja 04 (BO); Batanghari

District, Bajubang Subdistrict, Harapan Rainforest, 1 km after Jembatan Jengkol from office,

02°1F17.5"S 103°20'22.7"E, 2 Mar 2016, Damayanto & Widjaja 05 (BO). South Sumatra

Province: Musi Banyuasin District, Batanghari Leko Subdistrict, Hutan Harapan, around

Bato Nursery, 7 Apr 2013, Wardi et al. BOHK 478 (BO); Musi Banyuasin District, Lubuk

Bintialo Subdistrict, Harapan Rainforest, Merajang, 02°18'08.1"S 103°17'48.0"E, 3 Mar

2016, Damayanto & Widjaja 12 (BO), Damayanto & Widjaja 13 (BO), Musi Banyuasin

District, Lubuk Bintialo Subdistrict, Harapan Rainforest, Near Danau Fokus 3, 02°18'08.1"S

103°17'48.6"E, 4 Mar 2016, Damayanto & Widjaja 14 (BO).

New Dendroccilamus from Sumatra

Table 1. Dendrocalamus luteus Damayanto & Widjaja compared to closely related species.

D. luteus

D. bengkalisensis

D. buar

D. bait

Shoot

Orange with white

hairs and white

wax on the upper

sheath

Orange with white

hairs or glabrous

on the upper

sheath, white wax

not seen

Orange with

white hairs and

white wax on the

upper sheath

Orange with white

hairs on the upper

sheath, white wax

not seen

Culm colour

Yellowish green to

yellow

Green

Culm sheath

auricles

Rounded, 0.4-0.5

cm high, bristles

2-2.8 cm long

Rounded, 0.7-1

cm high, bristles

0.7-2.5 cm long

Rim like and

crisped to blade

base, bristles

0.9-1.8 cm long

Rounded, bristles

1.0-1.5 cm long

Culm sheath

ligule

Dentate, 0.2-0.4

cm high, bristles

few 0.4-0.6 cm

long

Denticulate, up

to 0.2 cm high,

bristles up to 0.2

(-0.5) cm long

Denticulate,

irregular, 0.7-1.9

cm high, bristles

0.4 -1.5 cm

Irregular, up to 1

cm, glabrous

Culm sheath

blade

Spreading when

young, reflexed

when mature

Reflexed

Erect to reflexed

Culm sheath

margin

White hairs

and replaced by

brownish hairs

Absent

Lower leaf

surface

Pubescent

Glabrous

Leaf sheath

auricles

Lobe-like, bristles

0.9-1.7 cm long

Rounded lobes,

bristles 0.9-1.7

cm long

Small to incon¬

spicuous, bristles

0.1-0.2 cm long

Small to incon¬

spicuous, bristles

0.4-0.6 (-1.2) cm

long

Palea

Keeled

No data

Not keeled

ACKNOWLEDGEMENTS. We would like to thank the Director of the Herbarium Bogoriense

(BO), Botany Division, Research Center for Biology, LIPI for allowing the consultation of

research material and the use of research facilities. We would like also to thank the Director

and staff of Harapan Rainforest (Indonesian Ecosystem Restoration) for permission and

funding to collect specimens. The Asian Logging Company, who were the previous owner

of the Indonesian Ecosystem Restoration, kindly issued a permit to collect bamboo in their

area. Thanks are due to Messrs. Zelvin, Fery and Edi for helping to collect the specimens and

Mr Wahyudi Santoso (BO) for preparing the illustration. We sincerely thank Dr Wong Khoon

Meng (SING) for reviewing this manuscript before it was submitted.

Card. Bull. Singapore 69(1) 2017

References

Widjaja, E.A. (1997). New taxa in Indonesian bamboos. Reinwardtia 11(2): 57-152.

Widjaja, E.A., Rahayuningsih, Y., Ubaidillah, R., Maryanto, I. & Rahajoe, J.S. (eds) (2014).

Kekinian Keanekaragaman Hayati Indonesia 2014. Jakarta: LIPI Press.

Gardens’ Bulletin Singapore 69(1): 81-87. 2017

doi: 10.3850/S201009811600006X

Liverworts of Bali, Indonesia,

with new records to the island

I. Haerida

Herbarium Bogoriense, Botany Division, Research Center for Biology,

Indonesian Institute of Sciences, Cibinong Science Center,

Jl. Raya Jakarta Bogor Km. 46, Cibinong, Bogor

ihaerida@ gmail.com

ABSTRACT. A total of 72 species of liverworts have been found in Bali. Based on recent

collections and former records of the species including 31 genera and 17 families, among them,

43 species are new to Bali.

Keywords. Bali, liverworts, new records

Introduction

Bali is a major tourist destination in Indonesia. According to the Central Bureau of

Statistics of the Republic of Indonesia, in 2015, 4,002,000 people visited Bali and in

July 2016 alone 484,231 people visited. There is, therefore, great pressure to expand the

infrastructure and facilities for tourists in Bali. This inevitably leads to the degradation

or destruction of forest and other natural habitats and, consequently, the biological

diversity found there becomes threatened.

Information on the bryophyte flora of Bali is rather rare. Hegewald & van Zanten

(1986) reported 78 species of mosses and 17 species of liverworts from Bali. Touw

(1992) noted 167 species of mosses for Bali. Additional literature including information

on the liverworts of Bali include Eggers et al. (1998), Schafer-Verwimp (2006, 2009),

Haerida et al. (2010), Alam (2012), Heinrichs et al. (2012) and Girmansyah et al.

(2013). Based on the results of their studies, only 26 species of liverworts have been

recorded, therefore indicating that an inventory on liverwort diversity of Bali is

needed. The study by Girmansyah et al. (2013) was a more general survey of Bali’s

plant and fungal diversity and they recorded about 1700 species of Spermatophytes,

Pteridophytes, fungi (Basidiomycota and Glomeromycota), and bryophytes (mosses

and liverworts). All of the species reported were based on specimens housed in Bogor

Herbarium (BO), Naturalis in Leiden (L) and from the literature.

In 2013, 2015 and 2016 the author visited several localities in Bali to collect

material of bryophytes and other cryptogams. The localities were in Candikuning (Bali

Botanical Garden, Bedugul, Baturiti at c. 1250-1450 m asl, Mt Tapak at c. 1900 m

and Mt Pohen at c. 2060 m), Bangli (Mt Abang at c. 1300-2250 m asl, Mt Batur and

surrounding areas, including the lake, at c. 1085-1610 m asl, and Mt Penulisan west

and east side at c. 1490-1610 m asl), and Karangasem (Mt Seraya, Bukit Tabuan area

at c. 550-790 m asl, Pura Bhur, Bwah and Swah areas at c. 630-1050 m asl, and Pura

Card. Bull. Singapore 69(1) 2017

Lempuyang area at c. 800-1150 m asl). This resulted in c. 180 collections, among

them c. 92 liverworts.

Results and discussion

Based on the previous literature, such as Hegewald & van Zanten (1986), Eggers et al.

(1998), Schafer-Verwimp (2006, 2009), Haerida et al. (2010), Alam (2012), Heinrichs

et al. (2012) and Girmansyah et al. (2013), and on the additional species found in this

study, the total number of identified liverwort species recorded from Bali is now 72

species, of which 43 species are new records for Bali. Of these, 15 species are thalloid

liverworts and 57 species are leafy liverworts. All new collections have been deposited

in BO.

These 72 species are from 31 genera and 17 families (Aneuraceae, Aytoniaceae,

Frullaniaceae, Herbertaceae, Jackiellaceae, Lejeuneaceae, Lophocoleaceae,

Marchantiaceae, Metzgeriaceae, Pallaviciniaceae, Plagiochilaceae, Porellaceae,

Radulaceae, Ricciaceae, Scapaniaceae, Targioniaceae and Wiesnerellaceae).

The specimens were identified using the Guide to the Liverworts and Hornworts of

Java (Gradstein, 2011) and other literature such as Benedix (1953), Hattori (1973),

Zhu & So (2001), Zhu & Gradstein (2005) and Siregar et al. (2013).

Candikuning has a higher number of liverworts species than Bangli and

Karangasem. This is because the forests in this area are wetter, thereby suiting a

greater bryophyte diversity (c. 77% of all species). Although Bangli has similar

conditions to Candikuning in the undamaged forested areas, there is actually very

little forest left undamaged, particularly around Mt Batur where the vegetation has

been greatly impacted by volcanic eruptions. Most other areas around Mt Abang have

been converted to agricultural land. Other species of liverwort were found in Bangli

and in Karangasem. These areas are relatively dry and, therefore, fewer liverworts are

present.

The higher number of species found in the Candikuning areas is due to the

optimal climate for bryophytes, specifically liverworts. Bali Botanical Garden and

surrounding areas have an annual precipitation of 2,950 mm/y, an average temperature

of c. 20.06°C, and an average of humidity of c. 80.82%. Mt Abang and surrounding

areas in Bangli have an annual precipitation of c. 2,200 mm/y, an average temperature

of c. 24.9°C, and an average humidity of c. 88%. The driest area, Mt Seraya in

Karangasem, has an annual precipitation of c. 1,945 mm/y, an average temperature of

c. 23°C, and an average humidity of c. 79%.

Species list

All specimens cited are in BO unless otherwise stated. An asterisk (*) indicates a new

record for Bali. Nomenclature follows Soderstrom et al. (2010, 2016) and Gradstein

(2011). IH + number refers to a collection by the author. Records which are based only

on information from the literature are referenced.

Liverworts of Bali

ANEURACEAE

*Aneura maxima (Schiffn.) Steph. - by little pond near spring, shaded at Mt Seraya c.

700 m, IH2132.

*Aneura pinguis (L.) Dumort. - on soil, semi-shaded site on Mt Abang, c. 1200 m, IH

2133.

AYTONIACEAE

*Asterella vulcanica (Schiffn.) Pand et al. ex Kachroo & Bapna - on rock and on

soil, semi-shaded sites on Mt Penulisan Barat and Mt Seraya, trail to Pura

Lampuyang, c. 850-c. 1400 m, IH 2116, 2117, 2120.

*Reboulia hemisphaerica (L.) Raddi - on soil, semi-shaded to shaded sites in Bali

Botanical Garden, on Mt Penulisan Barat and on Mt Seraya, trail to Pura

Lempuyang, c. 900-1400 m, IH 2028, 2114, 2118.

FRULLANIACEAE

Frullania ericoides (Nees) Mont. - Record from Hegewald & van Zanten (1986).

^Frullania gaudichaudii (Nees & Mont.) Nees & Mont. - on fallen trunk, open site in

Bali Botanical Garden, IH 2037.

^Frullania gracilis (Reinw., Blume & Nees) Dumort. - on fallen trunk, open site in

Bali Botanical Garden, IH 2038.

* Frullania junghuhniana Gottsche - on fallen trunk, open site in Bali Botanical

Garden, IH 2052.

* Frullania meyeniana Lindenb. - on tree trunk and on fallen trunk, open sites in Bali

Botanical Garden, IH 2051, 2053.

* Frullania moniliata (Reinw., Blume & Nees) Mont. - on tree trunk and on fallen

trunk, shaded to open sites in Bali Botanical Garden, on Bukit Tapak and on

Bukit Pohen, IH 2013, 2015, 2032, 2043, 2058, 2086, 2098.

* Frullania ornithocephala (Reinw., Blume & Nees) Nees - on fallen branch, open site

on Bukit Pohen, IH 2062.

Frullania riojaneirensis (Raddi) Spruce - on rotten log, open site on Bukit Tapak, IH

2096.

Frullania ternatensis Gottsche - Record from Hegewald & van Zanten (1986).

Frullania tricarinata Sande Lac. - Record from Hegewald & van Zanten (1986).

HERBERTACEAE

Herbertus ramosus (Steph.) H.A.Mill. - Record from Hegewald & van Zanten (1986).

J A CKIELL A CE AE

*Jackiella javanica Schiffn. - on fallen branch, shaded site on Mt Seraya, IH 2130.

LEJEUNEACEAE

Acanthocoleus javanicus (Steph.) Kruijt - Record from Heinrichs et al. (2012).

Acrolejeunea aulacophora (Mont.) Steph. - Record from Hegewald & van Zanten

(1986).

Card. Bull. Singapore 69(1) 2017

Acrolejeunea fertilis (Reinw., Blume & Nees) Schiffn. - Record from Heinrichs et al.

( 2012 ).

*Cheilolejeunea trapezia (Nees) R.M.Schust. & Kachroo - on leaves, shaded site in

Bali Botanical Garden, IH 2080.

*Cololeieunea angustiflora (Steph.) Mizut. - on leaves, semi-shaded site on Bukit

Tapak, IH 2040.

^Cololejeunea appressa (A.Evans) Benedix - on leaves, shaded site on Bukit Tapak,

IH 2024.

^Cololejeunea cf. lanciloba - on leaves, shaded site on Bukit Tapak, IH 2092.

* Cololejeunea macounii (Spruce ex Underw.) A.Evans - on leaves, semi-shaded to

shaded sites in Bali Botanical Garden and on Bukit Tapak, IH 2041, 2074.

* Cololejeunea cf. serrulata - on leaves, semi-shaded site on Bukit Tapak, IH 2018.

* Cololejeunea goebelii (Gottsche ex K.I.Goebel) Schiffn. - on leaves, shaded site on

Bukit Tapak, IH 2082, 2083, 2084.

* Cololejeunea gottschei (Steph.) Mizut. - on leaves, semi-shaded site on Bukit Tapak,

IH 2090.

* Cololejeunea subfloccosa Mizut. - on leaves, shaded site on Bukit Tapak, IH 2023.

* Cololejeunea triapiculata (Herzog) Tixier - on leaves, semi-shaded site on Bukit

Tapak, IH 2019.

Colura leratii (Steph.) Steph. - Record from Eggers et al. (1998).

Diplasiolejeunea cavifolia Steph. - Record from Schafer-Verwimp (2006).

*Drepanolejeunea pentadactyla (Mont.) Steph. - on leaves, shaded sites on Bukit

Tapak and in Bali Botanical Garden, IH 2078, IH s.n.

*Lejeunea apiculata Sande Lac. - on leaves, shaded site in Bali Botanical Garden, IH

2079.

Lejeunea mimula Hurl. - Record from Heinrichs et al. (2012).

^Leptolejeunea epiphylla (Mitt.) Steph. - on leaves, semi-shaded to shaded sites on Mt

Seraya and on Bukit Tapak 825 m, IH s.n., IH 2128.

Leptolejeunea foliicola Steph. - Records from Eggers et al. (1998) and Bechteler et

al. (2017).

Leptolejeunea massartiana Schiffn. ex Herzog - Record from Eggers et al. (1998).

* Leptolejeunea subacuta Steph. - on leaves, semi-shaded to shade sites on Bukit

Tapak and on Mt Seraya, IH 2093, 2129.

Lopholejeunea eulopha (Taylor) Schiffn. - Record from Haerida et al. (2010).

Lopholejeunea horticola Schiffn. - Record from Haerida et al. (2010).

* Lopholejeunea nigricans (Lindenb.) Steph. - on soil, semi-shaded on Mt Seraya

1500 m, IH 2120.

Lopholejeunea recurvata Mizut. - Record from Haerida et al. (2010).

Lopholejeunea subfusca (Nees) Schiffn. - Records from Haerida et al. (2010) and

recent collections from Bali on fallen twigs, open site on Bukit Pohen, IH 2044.

* Lopholejeunea zollingeri (Steph.) Schiffn. - on rotten log, open site on Bukit Pohen,

IH 2031.

Ptychanthus striatus (Lehm. & Lindenb.) Nees - on tree trunk, semi-shaded site on

Bukit Pohen, IH 2085.

Liverworts of Bali

Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenb. - Records from

Heinrich et al. (2012) and recent collections from Bali, on tree trunk and on

fallen branch, semi-shaded site on Bukit Pohen, IH 2049, 2073.

LOPHOCOLEACEAE

*Heteroscyphus argutus (Nees) Schiffn. - on root, shaded site on Bukit Tapak, IH

2095.

*Heteroscyphus aselliformis (Reinw., Blume & Nees) Schiffn. - on rotten log, semi-

shaded site on Bukit Pohen, IH 2029.

Heteroscyphus coalitus (Hook.) Schiffn. - on soil, on root and on rock, semi-shaded

to shaded sites in Bali Botanical Garden, on Bukit Tapak and on Mt Seraya, IH

2045, 2094, 2126.

MARCHANTIACEAE

Dumortiera hirsuta (Sw.) Nees - on soil, semi-shaded to shaded sites in Bali Botanical

Garden, on Mt Abang and on Mt Seraya, c. 850-c. 1700 m, IH 2046, 2107, 2113,

2121.

*Marchantia acaulis Steph. - on soil, semi-shaded on Mt Abang, c. 1700, IH 2131.

Marchantia emarginata Reinw., Blume & Nees - on soil and on ‘pura’ wall, semi-

shaded to open sites in Bali Botanical Garden, on Mt Penulisan and on Mt

Seraya, IH 2030, 2108, 2122.

*Marchantia geminata Reinw., Blume & Nees - on soil, open site on Mt Penulisan,

IH 2100.

*Marchantia treubii Schiffn. - on soil, semi-shaded to open sites on Mt Penulisan, c.

1800 m, IH 2101, 2111.

METZGERIA CE AE

*Metzgeria ciliata Raddi - on soil, shaded site in Bali Botanical Garden, growing

together with Wiesnerella denudata, IH 2047.

*Metzgeria lindbergii Schiffn. - on tree trunk and on twigs, shaded sites in Bali

Botanical Garden and on Bukit Tapak, IH 2072, 2077.

PALLAVICINIACEAE

*Pallavicinia lyellii (Hook.) Gray - on soil, open site on Mt Seraya, IH 2125.

PLAGIOCHILACEAE

Chiastocaulon opposition (Reinw., Blume & Nees) S.D.F.Patzak et al. - Record from

Hegewald & van Zanten (1986).

Plagiochila javanica (Sw.) Nees & Mont. - Record from Hegewald & van Zanten

(1986).

Plagiochila salacensis Gottsche - Record from Hegewald & van Zanten (1986).

Plagiochila teysmannii Sande Lac. - Record from Hegewald & van Zanten (1986).

Card. Bull. Singapore 69(1) 2017

PORELLACEAE

Porella acutifolia (Lehm. & Lindenb.) Trevis. - Record from Hegewald & van Zanten

(1986).

RADULACEAE

*Radula acuminata Steph. - on leaves, shaded site on Bukit Tapak, IH 2088.

*Radula campanigera Mont. - on tree trunk, semi-shaded site on Bukit Tapak, IH

2026.

*Radula complanata (L.) Dumort. - on tree trunk, shaded site on Bukit Pohen, IH

2033.

*Radula javanica Gottsche - on climber, on leaves and on tree trunk, shaded to open

sites on Bukit Tapak, IH 2016, 2089, 2099.

Radula multiflora Gottsche ex Schiffn. - Record from Hegewald & van Zanten (1986).

*Radula ventricosa Steph. - on tree trunk, semi-shaded on Bukit Tapak, IH 2025.

RICCIACEAE

* Riccia junghulmiana Nees & Lindenb. - on soil, open site in Bali Botanical Garden,

IH 2017.

SCAPANIACEAE

Scapania javanica Gottsche - Record from Schafer-Verwimp (2009).

TARGIONIACEAE

* cf. Targionia hypophylla - on soil, semi-shaded on Mt Seraya, IH 2119.

WIESNERELLACEAE

*Wiesnerella denudata (Mitt.) Steph. - on rock, shaded site near Batur Lake, IH 2104.

ACKNOWLEDGEMENTS. The author is grateful to “Proyek DIPA Puslit Biologi - LIPI,

tahun 2013-2016” for financial support for the study. She is also grateful to Prof. Tanias Poes

for checking the Cololejeuna spp. and to Prof. Rob Gradstein for verifying the identification of

some thalloid species. The author also sincerely thanks the two anonymous reviewers for their

comments and suggestions on this manuscript.

References

Alam, A. (2012). Liverwort Flora of Parson’s valley, Nilgiri hills (Western Ghats), South India.

Arch. Bryol. 119: 1-20.

Bechteler, J., Schafer-Verwimp, A., Lee, G.E., Feldberg, K., Perez-Escobar, O.A., Poes, T.,

Peralta, D.F., Renner, M.A.M. & Heinrichs, J. (2017). Geographical structure, narrow

species ranges, and Cenozoic diversification in a pantropical clade of epiphyllous leafy

liverworts. Ecol. Evol. 7: 638-653.

Benedix, E.H. (1953). Indomalayische Cololejeuneen. Feddes Repert. 134: 1-88.

Liverworts of Bali

Eggers, J., Frahm, J.-P. & Pursell, R.A. (1998). New bryophyte taxon records for tropical

countries II. Trop. Bryol. 14: 81-84.

Girmansyah, D., Santika, Y., Retnowati, A., Wardani, W., Haerida, I., Widjaja, E.A. & van

Balgooy, M.M.J. (eds) (2013). Flora of Bali: an Annotated Checklist. Bogor: Herbarium

Bogoriense, Botany Division, Research Center for Biology-LIPI & Yayasan Pustaka

Obor Indonesia.

Gradstein, S.R. (2011). Guide to the liverworts and hornworts of Java. Bogor: SEAMEO

BIOTROP.

Haerida, I., Gradstein, S.R. & Tjitrosoedirdjo, S.S. (2010). Lejeuneaceae subfamily

Ptychanthoideae (Hepaticae) in West Java. Gard. Bull. Singapore 62(1): 53-103.

Hattori, S. (1973). Notes on Asiatic species of the genus Frullania, Hepaticae, II. J. Hattori

Bot. Lab. 37: 55-84.

Hegewald, E. & van Zanten, B.O. (1986). A list of bryophytes from Bali (Indonesia) collected

by E. and P. Hegewald in 1981. J. Hattori Bot. Lab. 60: 263-269.

Heinrichs, J., Dong, S., Feldberg, K. Schafer-Verwimp, A. & Schmidt, A. (2012).

Sphaerolejeunea (Lejeuneaceae, Porellales) is a synonym of Lejeunea. Phytotaxa 69:

7-15.

Schafer-Verwimp, A. (2006). A new species of Diplasiolejeunea (Lejeuneaceae,

Jungermanniopsida) from Sumatra, and a key for the genus in Asia. Herzogia 19: 239-

244.

Schafer-Verwimp, A. (2009). New national and regional bryophyte records, 22. J. Bryol. 31:

201 - 210 .

Siregar, E.S., Ariyanti, N.S. & Tjitrosoedirdjo, T.T. (2013). The liverwort genus Marchantia

(Marchantiaceae) of Mount Sibayak North Sumatra, Indonesia. Biotropia 20(2): 73-80.

Soderstrom, L., Gradstein, S.R. & Hagborg, A. (2010). Checklist of the hornworts and

liverworts of Java. Phytotaxa 9: 53-149.

Soderstrom, L., Hagborg, A., von Konrat, M., Bartholomew-Began, S., Bell, D., Briscoe, L.,

Brown, E., Cargill, D.C., Costa, D.P., Crandall-Stotler, B.J., Cooper, E.D., Dauphin, G,

Engel, J.J., Feldberg, K., Glenny, D., Gradstein, S.R., He, X., Heinrichs, J., Hentschel,

J., Ilkiu-Borges, A.L., Katagiri, T., Konstantinova, N.A., Larrafn, J., Long, D.G., Nebel,

M., Poes, T., Felisa Puche, F., Reiner-Drehwald, E., Renner, M.A.M., Sass-Gyarmati,

A., Schafer-Verwimp, A., Moragues, J.G.S., Stotler, R.E., Sukkharak, P., Thiers, B.M.,

Uribe, J., Vana, J., Villarreal, J.C., Wigginton, M., Zhang, L., Zhu, R.-L. (2016). World

checklist of hornworts and liverworts. PhytoKeys 59: 1-828.

Touw, A. (1992). A survey of the mosses of the Lesser Sunda Islands (Nusa Tenggara),

Indonesia. J. Hattori Bot. Lab. 71: 289-366.

Zhu, R.L. & So, M.L. (2001). Epiphyllous Liverworts of China. Nova Hedwigia Beih. 121:

1-148.

Zhu, R.L. & Gradstein, S.R. (2005). Monograph of Lopholejeunea (Lejeuneaceae, Hepaticae)

in Asia. Syst. Bot. Monogr. 74: 1-198.

Gardens’ Bulletin Singapore 69(1): 89-95. 2017

doi: 10.3850/S2010098116000081

Begonia ignita (sect. Petermannia, Begoniaceae),

a new species with orange flowers

from Sulawesi, Indonesia

C.-W. Lin' D.C. Thomas 2 , W.H. Ardi 3 & C.-I Peng 4

herbarium of Taiwan Forestry Research Institute,

No. 53, Nanhai Road, Taipei 100, Taiwan

2 Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

3 Bogor Botanic Gardens, Jl. Ir. H. Juanda No. 13, Bogor, Indonesia

4 Herbarium, Biodiversity Research Center, Academia Sinica,

Nangang, Taipei 115, Taiwan

bopeng @ sinica.edu.tw

ABSTRACT. Begonia ignita C.W.Lin & C.I Peng, a new species of Begonia sect. Petermannia

from Sulawesi, Indonesia, is here described and illustrated. It is distinct from other species

in Begonia section Petermannia by a character combination including a procumbent stem

ascending only at the apex, symmetric or subsymmetric leaves, the presence of a pale band or

maculation running parallel to the leaf margin, and orange tepals. A detailed comparison with

a morphologically similar species, the Sulawesi endemic Begonia mendumiae M.Hughes, is

provided.

Keywords. Begonia ignita , B. mendumiae, Indonesia, new species, Sulawesi

Introduction

Forty-six species of Begonia are presently known from the Indonesian island of

Sulawesi (Thomas et al., 2013), 22 of which have been described since 2006 (Hughes,

2006; Thomas & Hughes, 2008; Girmansyah et al., 2009; Thomas et al., 2009a,

2009b, 2011; Wiriadinata, 2013; Ardi et al., 2014). In addition, the examination

of major herbarium collections (BO, E, L, K, SING) and specimen images from

numerous additional herbaria provided by the Begonia Resource Centre (Hughes

et al., 2015) indicates that there are many more species awaiting description. This

high discovery rate is not surprising considering (i) that biological inventory rates

and specimen collection density on Sulawesi are among the lowest in Indonesia

(Cannon et al., 2007), (ii) the vast area (174,600 km 2 ) of the island, (iii) the markedly

lower number of currently accepted species in comparison to the neighbouring

Borneo: Sarawak (124,450 km 2 with c. 100 species of Begonia : Kiew et al., 2015)

and Sabah (72,500 km 2 with 82 species: Kiew et al., 2015); Brunei (5,765 km 2 with

22 species: Joffre et al., 2015; Low et al., 2015); and (iv) the lack of a revision of

central Malesian Begonia.

Card. Bull. Singapore 69(1) 2017

A very attractive orange-flowered Begonia , claimed to be of likely Sulawesi

origin by some growers, has recently entered into cultivation inside and outside of

Indonesia. It is immediately distinct from all other known Sulawesi species in its

symmetric or subsymmetric leaves and orange or pale tangerine to vermilion tepals. In

this study, we describe this species using measurements of herbarium specimens made

from living plants in cultivation. In addition to the full taxonomic account, colour

plates, line drawings, and a comparison to a phenetically similar species are provided

to aid in identification.

Begonia ignita C.W.Lin & C.I Peng, sp. nov. § Petermannia

This species is distinct from other species in Begonia section Petermannia by a character

combination including a procumbent stem ascending only at the apex, symmetric or

subsymmetric leaves, the presence of a pale band or maculation running parallel to

the leaf margin, and orange tepals. - TYPE: Collected in Indonesia, Sulawesi, precise

locality unknown, grown in cultivation in Bogor Botanic Gardens and vouchered on 9

October 2016 as Wisnu H. Ardi WI 117 (holotype BO; isotype SING). (Fig. 1, 2)

Plant perennial, monoecious. Stem procumbent and rooting at nodes, ascending at the

apex, olive to magenta, 8-20 cm long, 4-6 mm across, glabrous, internodes 0.5-5 cm

long; stipules widely ovate-triangular, 8-12 mm long, c. 6 mm wide, hyaline, reddish,

strongly keeled, margin entire, apex aristate, arista c. 2.5-4 mm long. Leaves 5-10,

held horizontally; petiole terete, 3.5-8 cm long, 3.5-6 mm across, olive-red to crimson,

sub-appressed sparsely velutinous, slightly grooved adaxially; lamina cordiform to

widely cordiform, basifixed, symmetric or subsymmetric, cordate at base, margin

slightly undulate and distantly denticulate, glabrous, apex acuminate, 7-12 cm long,

6.5-11 cm wide, basal lobes equal or subequal, to 2.5 cm long, thickly chartaceous,

adaxially dark olive-green to dark maroon, embellished with a silvery green band

(sometimes with irregular spots of variable sizes between veins) running parallel to the

margin, veins red towards base; slightly rugose between primary veins; abaxially pale

red to magenta; venation palmate-pinnate, midrib conspicuous, 6-9.5 cm long with c.

3 pairs of major lateral veins, each branching dichotomously. Inflorescence terminal,

bisexual, protogynous, cymose-paniculate, 13-22 cm long; bracts magenta to pale

rosy-brown, hyaline, persistent, widely ovate, (4-)8-12 mm long, (2-)4-7 mm wide,

decreasing in size towards the more distal inflorescence nodes, aristate at apex, arista

3^1 mm long, margin entire; pistillate flowers produced on up to 3 subsequent basal

nodes of the inflorescence, usually solitary or sometimes together with 1 staminate

flower; staminate flowers usually only produced in distal inflorescence part, up to 15,

in several (1—)2—3-flowered compressed monochasial partial inflorescences borne on

short lateral branches (0.1-2 cm), at the most distal nodes sometimes reduced to a single

flower, erect or ascending, reddish to crimson, glabrous. Staminate flower: pedicel

11-18 mm long, glabrous; tepals 2, orange or pale tangerine to vermilion, glabrous,

widely ovate to suborbicular, margin entire, base rounded, 10-15 mm long, 9-11 mm

wide; androecium subzygomorphic, stamens 35^15, yellow, filaments shortly fused

Fig. 1. Begonia ignita C.W.Lin & C.I Peng. A. Habit. B. Stipule. C. Bracts at lowermost and

uppermost parts of inflorescence. D. Male inflorescence parts showing basal few-flowered

cymes and distal nodes bearing single flowers. E. Staminate flower, face and side views. F.

Stamen, dorsal and ventral views. G. Pistillate flower, face and side views. H. Style, dorsal,

side and ventral views. I. Cross section of an ovary. Scale: A, D: 5 cm; E: 2 cm; B, C, G: 1 cm;

H: 5 mm; F: 2 mm. Drawn by C.W. Lin.

Card. Bull. Singapore 69(1) 2017

Fig. 2. Begonia ignita C.W.Lin & C.I Peng. A, B. Habit, showing variation in leaf colours.

C, D. 5-tepaled pistillate flower (occasional), face and side views. E. Inflorescence, showing

2-tepaled staminate flowers. F. Staminate flower, face view. G. 4-tepaled pistillate flower in

inflorescence. H. 2-tepaled pistillate flower (occasional). I. Cross section of ovary showing

axile, bilamellate placentae. (Photos: Y.-Z. Siaw)

Begonia ignita, a new species from Sulawesi

at base; anthers c. 0.8 mm long, oblongoid, subequal in length to filaments. Pistillate

flower: pedicel 6-13 mm long, glabrous; tepals 4 (rarely 2, 3 or 5), pale orange to

vermilion, ovate or obovate to oblanceolate, 9-15 mm long, 4-10 mm wide, glabrous;

ovary reddish, body trigonous-ellipsoid, 9-11 mm long, c. 7 mm across, 3-winged;

wings cuneate or rounded at base, truncate or rounded at the apex, 12-15 mm long,

2-4 mm wide, locules 3, placentae axile, bilamellate; styles 3, yellow, bifid, c. 4 mm

long; stigmas in a spiral band and papillose all around. Fruits not seen.

Distribution. Only known from cultivation; likely endemic to Sulawesi (see Notes).

Etymology. The specific epithet refers to the flame-coloured tepals (Latin: igneus -

flame-coloured).

Notes. Begonia ignita has entered into cultivation and is grown by various amateur

enthusiasts and Begonia collectors inside and outside of Indonesia. The origin of

these cultivated plants remains obscure. Enquiries about the species’ origin with

several growers indicated that most acquired the plants from material cultivated

by other growers. This is also the case for the plants cultivated at Bogor Botanic

Gardens on which the description is based. Some growers indicated that a market in

Sulawesi was the likely original source of the plants, but they could not provide any

additional information on the location of this market (province, city etc.). Phylogenetic

reconstructions using chloroplast DNA sequence data of Begonia ignita (ndhA intron,

ndhF-rpl32 spacer, rpl32-trnh spacer, unpublished data, D. Thomas) integrated with

previously published data of other Asian taxa (Thomas et al., 2012; Moonlight et

al., 2015), including accessions of about 74% of the currently accepted species from

Sulawesi, corroborate a putative Sulawesi origin. Begonia ignita is retrieved in a well-

supported clade of Sulawesi endemics in Begonia section Petermannia. It is sister

to a well-supported clade including two subclades: a subclade of endemic species

distributed on the northern arm of Sulawesi, and a subclade of species endemic to

eastern Central Sulawesi.

Begonia ignita is one of the most distinct and most beautiful Indonesian species

in cultivation. Its colourful cordiform leaves adorned with a silvery green band or

maculation ru nnin g inside the margin, and the showy flammate flowers immediately

make it a very attractive species. It is distinct from all other Sulawesi begonias by the

symmetric or subsymmetric cordiform leaves and in being the only known orange-

flowered Begonia on the island. The new species resembles the Sulawesi endemic

Begonia mendumiae M.Hughes (sect. Petermannia ) in the creeping habit, and the

band or maculation of the leaves but can be easily differentiated from B. mendumiae

by the characters shown in Table 1.

Card. Bull. Singapore 69(1) 2017

Table 1. Comparison of Begonia ignita C.W.Lin & C.I Peng and B. mendumiae M.Hughes.

B. ignita (Figures 1, 2)

B. mendumiae (Hughes,

2006: Figures 1,5)

Stem vestiture

Glabrous

Hairy

Leaf

Shape

Cordiform, symmetric or

subsymmetric

Ovate, strongly asymmetric

Size (cm)

7-12x6.5-11

to 9 x 6

Vestiture

Glabrous

Pinkish-red hairy

Bract length (mm)

4-12

c. 4

Inflorescence

Staminate part

Partial inflorescences with

(1 —)2—3 flowers

6-8 flowers

Staminate flower

Tepal colour

Orange or pale tangerine to

vermilion

Pale dusky pink

Anther colour

Yellow

Orange

Pistillate flower

Number of tepals

(2-)4(-5)

Tepal colour

Pale orange to vermilion

Dusky pink

Tepal margin

Glabrous

Sparsely fimbriate

Style colour

Yellow

Orange

Ovary

Glabrous

Hairy

ACKNOWLEDGEMENTS. We are grateful to Yu-Zhang Siaw and Yu-Pin Ang who kindly

provided samples and photos.

Begonia ignita, a new species from Sulawesi

References

Ardi, W.H., Ardaka, I.M., Hartutiningsih, Lugrayasa, I.N. & Thomas, D.C. (2014). Two new

species of Begonia (Begoniaceae) from Sulawesi, Indonesia. Edinburgh J. Bot. 71:

259-268.

Cannon, C.H., Summers, M., Harting, J.R. & Kessler, P.J.A. (2007). Developing conservation

priorities based on forest type, condition, and threats in a poorly known ecoregion:

Sulawesi, Indonesia. Biotropica 39: 747-759.

Girmansyah, D., Wiriadinata, H., Thomas, D.C. & Hoover, W.S. (2009). Two new species

and one new subspecies of Begonia (Begoniaceae) from Southeast Sulawesi, Indonesia.

Reinwardtia 13: 69-74.

Hughes, M. (2006). Four new species of Begonia (Begoniaceae) from Sulawesi. Edinburgh J.

Bot. 63: 191-199.

Hughes, M., Moonlight, P., Jara, A., Tebbitt, M. & Pullan, M. (2015). Begonia Resource Centre.

http://elmer.rbge.org.uk/begonia/. Accessed 6 Dec. 2016.

Joffre, A.A., Kiew, R., Julia, S. & Rimi, R. (2015). The Begoniaceae of Brunei Darussalam,

Borneo, including two new species. Sandakania 20: 7-50.

Kiew, R., Julia, S., Rimi, R. & Joffre, A.A. (2015). A Guide to Begonias of Borneo. Kota

Kinabalu: Natural History Publications (Borneo).

Low, Y.W., Joffre, A.A. & Muhammad Ariffin, A.K. (2015). Novitates Bruneienses, 2. A

remarkable new species of Begonia sect. Petermannia (Begoniaceae) from Brunei

Darussalam. Gard. Bull. Singapore 67: 61-68.

Moonlight, P.W., Richardson, J.E., Tebbitt, M.C., Thomas, D.C., Hollands, R. & Hughes,

M. (2015). Continental-scale diversification patterns in a mega-diverse genus: the

biogeography of Neotropical Begonia L. J. Biogeogr. 42: 1137-1149.

Thomas, D.C. & Hughes, M. (2008). Begonia varipeltata (Begoniaceae): Anew peltate species

from Sulawesi, Indonesia. Edinburgh J. Bot. 65: 369-374.

Thomas, D.C., Ardi, W.H. & Hughes, M. (2009a). Two new species of Begonia (Begoniaceae)

from Central Sulawesi, Indonesia. Edinburgh J. Bot. 66: 103-114.

Thomas, D.C., Ardi, W.H., Hartutiningsih & Hughes, M. (2009b). Two new species of Begonia

(Begoniaceae) from South Sulawesi, Indonesia. Edinburgh J. Bot. 66: 229-238.

Thomas, D.C., Ardi, W.H. & Hughes, M. (2011). Nine new species of Begonia (Begoniaceae)

from South and West Sulawesi, Indonesia. Edinburgh J. Bot. 68: 225-255.

Thomas, D.C., Hughes, M., Phutthai, T., Ardi, W.H., Rajbhandary, S., Rubite, R., Twyford, A.D.

& Richardson, J.E. (2012). West to east dispersal and subsequent rapid diversification

of the mega-diverse genus Begonia (Begoniaceae) in the Malesian archipelago. J.

Biogeogr. 39: 98-113.

Thomas, D.C., Ardi, W.H., Girmansyah, D. & Hughes, M. (2013). Sulawesi Begonia Data

Portal. http://portal.cybertaxonomy.Org/flora-malesiana-prospective/.Accessed 13 Dec.

2016.

Wiriadinata, H. (2013). A new species of Begonia (Begoniaceae) from South Sulawesi,

Indonesia. Reinwardtia 13: 445-448.

Gardens’ Bulletin Singapore 69(1): 97-147. 2017

doi: 10.3850/S2010098116000093

Contribution to a revision of Hoya

(Apocynaceae: Asclepiadoideae) of Papuasia.

Part I: ten new species, one new subspecies

and one new combination

N. Simonsson Juhonewe 1 & M. Rodda 2

‘Research affiliate at National Research Institute of Papua New Guinea

Home address: PO Box 1 - 524, Ukarumpa, EHP 444, Papua New Guinea

2 Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

rodda. michele@gmail. com

ABSTRACT. The large majority of Hoya R.Br. species known from Papuasia were described

in the first half of the 20th century and most of these are endemic. Along with Borneo and the

Philippines the region is a centre of diversity of the genus. Since 2006 herbarium materials

have been studied for a revision of Hoya of New Guinea along with field investigations on

Papuasian Hoya since 2010. In the present paper we publish ten new species, Hoya brassii

PI.Forst. & Liddle ex Simonsson & Rodda, H. carrii PI.Forst. & Fiddle ex Simonsson &

Rodda, H. edholmiana Simonsson & Rodda, H. evelinae Simonsson & Rodda, H. juhoneweana

Simonsson & Rodda, H. krusenstierniana Simonsson & Rodda, H. koteka Simonsson &

Rodda, H. stenakei Simonsson & Rodda, H. versteegii Simonsson & Rodda and H. yvesrocheri

Simonsson & Rodda, one subspecies, H. juhoneweana ssp. lindforsiana Simonsson & Rodda,

and make one new combination, H urniflora (PI.Forst.) Simonsson & Rodda, which is also

lectotypified.

Keywords. Indonesia, Marsdenieae, Papua New Guinea

Introduction

Hoya R.Br., with an estimated 350—450 species (Rodda, 2015), is the largest genus

in the Apocynaceae. The most species-rich regions for Hoya are in tropical and

subtropical South Asia, Southeast Asia and Papuasia. The genus also extends to the

Himalayan foothills to the northwest, Okinawa (Japan) to the northeast, Australia to

the south and the Fiji Islands to the southeast. The plants are generally epiphytic or

hemi-epiphytic climbers or have a pendulous habit, with a few shrubby species (Rodda

& Ercole, 2014; Rodda et al., 2014a).

Recent regional revisions are only available for Peninsular Malaysia (Rintz,

1978), China (Fi et al., 1995) and Australia (Forster & Fiddle, 1996). A guidebook

to Hoya of Borneo has just been published (Famb & Rodda, 2016) but a revision for

the island is still in preparation (Rodda, in prep.). Species diversity is highest in the

Malesian islands, in particular in the Philippines, Borneo and New Guinea. In both the

Philippines and Borneo the number of species known has dramatically increased in

Card. Bull. Singapore 69(1) 2017

recent years. Kloppenburg (1991) suggested that the Philippines might have in excess of

30 species, a number that has since climbed to 104 species (Aurigue et al., 2013) with a

further 80 new taxa published since 2013 (IPNI, 2016). However, the actual number of

species in the Philippines is pending a critical revision. Forster et al. (1998) noted the

species diversity of Hoy a in Borneo was ‘unknown’ but recent research has since led to

the description of numerous new species in preparation for a comprehensive revision

(Rodda & Nyhuus, 2009; Rodda et al., 2011, 2014a, 2014b, 2015, 2016; Rodda &

Simonsson, 2011a, 2011b; Rodda & Simonsson Juhonewe 2013a, 2013b; Lamb et al.,

2014; Rahayu et al., 2015; Rodda, 2015; Lamb & Rodda, 2016), and a checklist that

includes 71 species and one subspecies (Lamb & Rodda, 2016).

The island of New Guinea has 74 species according to Forster (1996), mostly

described in the first half of last century. Papuasia lies towards the eastern edge

of the distribution area of the genus Hoya and includes the Papua and West Papua

provinces of Indonesia, Papua New Guinea (PNG), and the Solomon Islands. The

largest land mass is the island of New Guinea, the largest tropical island in the world.

Major contributions to Hoya of New Guinea were published by Schlechter, almost

entirely based on his own collections obtained during expeditions between 1901 and

1909 (Schlechter, 1905, 1908, 1913). Schlechter’s collections are limited to German

New Guinea, now part of PNG. Moore (1916) described numerous Hoya species

from Indonesian New Guinea, based on specimens collected by Boden Kloss. For the

Solomon Islands Hoya species were first published by Hemsley (1891, 1892, 1894).

The most recent contributions to the Hoya of New Guinea were published by

Forster & Liddle (1991, 1992, 1993) and Forster et al. (1995). Forster et al. (1997)

described the monotypic genus Madangia PI.Forst., Liddle & I.M.Liddle which has

since been synonymised into Hoya (Wanntorp & Forster, 2007).

Despite the large number of species already recorded, the Hoya diversity of

PNG is still poorly known. Early exploration was focused mostly in Madang and

Morobe Provinces of PNG (Forster, 1996), leaving much of the island unexplored and,

consequently, the likelihood of large numbers of new taxa yet to be discovered (Forster

1996; Prance et al., 2000). Indonesian New Guinea is overall much less explored

than PNG (Forster, 2006) and only 16 species have so far been recorded from both

Indonesian New Guinea and PNG (Forster, 1996, 2006).

The island of New Guinea is dominated by the mountain ranges that run the

length of the island. These mountains can exceed 3000 m and divide the island in a

cratonic southern half and accreted terranes to the north (Brown et al., 2006; Heads,

2006). The varied topography, in combination with ever-wet versus monsoonal

climate zones (Whitmore, 1984; Van Welzen, 1998), offers a large number and variety

of biogeographic zones that support high species richness, as observed for example in

orchids (Vollering et al., 2016) and Rhododendron (Brown et al., 2006). Despite the

lack of biogeographic data on Hoya , based on our observations the lowland rainforests

and lower montane forests appear to harbour the majority of Hoya species, while the

montane forests have a high percentage of local endemics. Only a very few species

go beyond 2000 m, most notably Hoya krusenstierniana Simonsson & Rodda (here

Hoya of Papuasia I.

published) that inhabits the cold upper montane forests from 1800 m up to 2600 m.

That is surprisingly high, since Hoya has its upper limit at 2200 m on both Borneo

and Sumatra (Lamb & Rodda, 2016). Above 2600 m, in the sub-alpine vegetation, no

Hoya species have so far been discovered. Along the shoreline, with rough conditions

such as saline winds, strong sun exposure and little shade, there are only a few Hoya

species capable of growing, although often quite abundantly. These coastal species are

widespread all along New Guinea’s coastline and its outlying islands, e.g. Hoya anulata

Schltr., H. australis R.Br. ex J.Traill, H. globulifera Blume, H. nicholsoniae F.Muell.,

H. inconspicua Hemsl. and H. ischnopus (Warb.) Schltr. The coastal species, being

common and easily accessible, are thus the most abundantly represented New Guinean

species in herbaria and were also the most frequently encountered species during the

first author’s expeditions in lowland PNG. As these species are hardy, widespread,

and tolerant of weather extremes, they have also been sighted on big trees in cities, in

gardens, in secondary forests and even growing up to 1400 m (except H. inconspicua

that is limited to the lowlands). In contrast, almost all other Hoya species are confined

to primary forests or their margins and are often absent from secondary forests. They

are thus vulnerable due to forest clearance for agriculture and timber harvesting. Except

for the widespread coastal species, there is almost no overlap between species in the

highlands and lowlands. There are two common montane species that are capable of

growing outside of primary forests, recolonising secondary forests and even growing

on trees in villages and gardens; these are H. solaniflora Schltr. and H. subglabra

Schltr. Both species are frequently seen in bright locations between 1500 and 1800 m

all over the Highlands in PNG.

New Guinea has several species complexes of similar taxa that may turn out to

be fewer variable species once more collections are gathered from the more remote

areas of the island. These complexes include the Hoya ischnopus complex (which

includes H. dimorpha F.M.Bailey, H. dischorensis Schltr., H. flavescens Schltr., H.

kenejiana Schltr. and H. montana Schltr.) and the H. globulifera complex (including

H. lamingtoniae F.M.Bailey and H. rubida Schltr.). A third more challenging group

includes all members of Hoya section Eriostemma Schltr. which are co mm on and

widespread throughout the Malesian region and Papuasia. Species delimitation within

this section is particularly problematic.

New Guinea’s Hoya species harbour several extremes of morphology for

the genus, including the smallest leaved species such as Hoya microphylla Schltr.,

H. pulchella Schltr., H. oxycoccoides S.Moore and H. krusenstierniana. The largest

flowers are recorded from New Guinea, Hoya gigas Schltr. and H. lauterbachii

K.Schum., and the almost as large, newly discovered, H. stenakei Simonsson & Rodda.

The majority of New Guinea’s species of Hoya have large (occasionally > 3

cm), campanulate or urceolate corollas. Even species forming small plants such as

Hoya pulchella , H. patella Schltr. and H. juhoneweana ssp. lindforsiana Simonsson &

Rodda have large flowers. This is not observed in Borneo or Philippine Hoya , where

the majority of species have reflexed (Borneo) or revolute (Philippines) corollas and

only a few taxa have campanulate corollas.

100

Card. Bull. Singapore 69(1) 2017

The current project

In the current project we set out to study all herbarium specimens from Papuasia,

several of which were already known to represent new species (Forster & Liddle,

1992), and to conduct extensive field surveys in PNG to obtain new collections.

Herbarium materials were studied at B, BM, BO, CGE, G, K, L, LAE, OXF, P, SAN,

SAR, SING and Z. Loans from A and US were studied at SING. Forster & Liddle

(1992) stated that a comprehensive revision of the genus in Papuasia would require

extensive fieldwork and cultivation of field-collected material that would take many

years. Cultivation of field-collected materials for the description of new species and

photo documentation has been successfully adopted by Rintz (1978) and Lamb et

al. (2014) for Peninsular Malaysia and Sabah Hoy a respectively. Our observations

in the field confirmed the need to cultivate sterile field-collected specimens as only

approximately one out of every 300 Hoy a individuals observed were in flower during

the fieldwork. A revision based only on specimens collected in flower during fieldwork

would result in the omission of over 99 % of the total plants sighted in the field.

Therefore, the first author moved to PNG in 2010 to conduct extensive field work in

all provinces and to cultivate selected collections at two different altitudes to obtain

flowering material. The plants were grown in Lae, Morobe Province, for low altitude

species, and in Kainantu, Eastern Highlands Province, at 1500 m altitude, for high

altitude species.

By December 2015, 55 field expeditions in 11 provinces (Fig. 1) had been

carried out and data gathered on locality, habitat, population count and size, presence

of ants, size of host tree and forest type for more than 4000 Hoy a individuals. A

selected 1500 specimens have been brought into cultivation (Fig. 2). Once they flower

they are photographed and made into herbarium specimens. The materials gathered

will be used for conservation, taxonomic and molecular studies.

The purpose of the present paper is to describe 10 species and one subspecies

that have been identified as new among the herbarium specimens, new field collections

and cultivated collections that have already flowered. We also move Marsdenia

urniflora PI.Forst into Hoy a. All specimens that could be georeferenced have been

plotted in Fig. 1.

Taxonomy

Hoy a brassii PI.Forst. & Liddle ex Simonsson & Rodda, sp. nov.

Similar to Hoya globulifera Blume in leaf shape (linear-lanceolate) but distinguished

by its much shorter petioles (1-2 mm in H. brassi , > 4 mm in H. globulifera ), much

shorter peduncles (c. 1 mm vs. > 1 cm) and inflorescences with fewer flowers (1-10

vs. 10-30 flowers). - TYPE: Papua New Guinea, Western Province, Palmer River, 2

miles below junction with Black River, 100 m., June 1936. Brass, L.J. 6883 (holotype

A [A00353955], BM [BM001014248], BRI n.v.). (Fig. 3, 4)

Hoyci of Papuasia I.

101

Fig. 1 . Map of Papua New Guinea (shaded in grey) with A. Localities visited by the first author

between 2011 and 2015. B. Collection localities of the new species published in the present

paper, excluding Hoya versteegii and Hoy a koteka. Map constructed using SimpleMappr

(www. s implemappr. net).

102

Card. Bull. Singapore 69(1) 2017

Fig. 2. A. Recently wild-collected plants in cultivation at Ukarumpa, PNG. B. Hoy a edholmiana

in bloom in cultivation (live collection no. NS 11-232). C. Two of the many clones of Hoy a

calycina complex. (Photos: N. Simonsson Juhonewe)

Hoya of Papuasia I.

103

Epiphytic climber with white latex in all vegetative parts. Stems cylindrical, stiff,

2-3 m long, 1.5-3 mm in diameter, green to greyish brown, glabrous or sparsely

pubescent; older stems leafless, up to 5 mm in diameter, greyish brown, glabrous with

peeling bark; internodes (1—)3—15 cm long. Adventitious roots sparsely produced

along the stem. Leaves petiolate; petiole terete, 1-2 x c. 1.5 mm in diameter, greyish

green, glabrous; lamina linear-lanceolate, 1-2 mm thick, fleshy, 5-11 x 1-2.5 cm,

bright to dark green on adaxial surface, light green on abaxial surface, glabrous, apex

acute to acuminate, base round to attenuate, basal colleters 2, ovoid, 4-5 x 1.5-2 mm;

venation pinnate, midrib depressed on adaxial surface, secondary veins 2-5 each side,

barely evident in living material, more visible in dry material, diverging at 20-45°.

Inflorescences one per node, extra-axillary, positively geotropic, convex, consisting of

1-10 flowers; peduncle terete, c. 1 x 2 mm, older peduncles developing a rachis from

previous flowerings, new peduncles glabrous or sparsely pubescent, older peduncles

glabrous; pedicels terete, 10-15 x c. 1 mm in diameter, light green-yellow, often with

red spots, papillate to glabrous, warty. Flower buds apically flattened, cream white to

pale yellow with reddish spots. Calyx c. 3 mm in diameter, lobes deltoid, c. 1 x 1 mm,

apex broadly acute to obtuse, margin sparsely ciliate; basal colleters one in each calyx

lobe sinus, deltoid. Corolla rotate, concave, 1-1.5 cm in diameter (c. 1.5 cm when

flattened), cream white to pale yellow, often with red spots on abaxial surface, also

visible on the adaxial surface as pink spots; tube 3-3.5 mm long; lobes ovate, 4-5 x

5-6 mm, adaxially thinly pubescent, denser at base of the corona, abaxially glabrous,

apex acute, revolute. Corona staminal 1.5-1.7 mm high, c. 3.5 mm in diameter, fleshy,

cream white to pale yellow with maroon centre; lobes obovate, c. 1.5 x 1-1.2 mm,

inner processes slightly erect, apices acute, outer processes usually slightly erect to

same level as inner processes, apex obtuse when observed from above, deeply grooved

between inner and outer processes, not basally fused with the filament tube, with basal

revolute margins. Style-head convex to same level as inner corona processes, hidden

by anther appendages. Pollinia oblong slightly tapering at the base, with sterile edge,

c. 450 x 200 pm, corpusculum squarish, c. 170 x 150 pm, caudicles attached at the

sides of corpusculum, 70-100 pm long. Ovary broadly conical, glabrous. Fruits and

seeds not observed.

Distribution. Recorded only from one single locality at the confluence of Palmer River

and Black River in the northwestern corner of PNG Western Province at 100-150 m.

Etymology. Named after Leonard John Brass (1900-1971), an Australian botanist who

collected the type specimen in 1936 on the Fly River Expedition of the Am erican

Museum of National History (the second Archbold New Guinea Expedition).

Habitat and ecology. At the type locality Hoya brassii is commonly found at the top of

mature primary forest trees, in bright light conditions. It was absent in the understorey

where the new taxon Hoya evelinae Simonsson & Rodda was instead abundant.

104

Card. Bull. Singapore 69(1) 2017

500 {jm

Fig. 3. Hoya brassii Simonsson & Rodda. A. Corolla, top view. B. Corona, side view. C.

Corona, from underneath. D, E. Calyx. F. Pollinarium. All from Simonsson Juhonewe, N. &

Juhonewe, F. NS0034L. (Drawing: M. Rodda)

Hoyci of Papuasia I.

105

Fig. 4. Hoya brassii Simonsson & Rodda. A. Flowering branch. B. Inflorescence from above.

C. Corolla and calyx from underneath. All from Simonsson Juhonewe, N. & Juhonewe, F.

NS0042L. (Photos: N. Simonsson Juhonewe)

Provisional IUCN conservation assessment. Critically Endangered CR Bla,b(iii)

(IUCN, 2016). The species appears to be common at the type locality but it may be

locally endemic as it has not been recorded from elsewhere, either from herbarium

specimens or during the first author’s fieldwork. Much of PNG Western Province’s

lowland rainforest has been, or is being, logged (Shearman et al., 2009) and the type

locality is not inside a protected area and may be lost in the near future.

106

Card. Bull. Singapore 69(1) 2017

Additional specimens examined. PAPUA NEW GUINEA: Western Province: Palmer River,

2 miles below junction Black River, slender climber common in substage forest of ridges, 100

m, pale yellow flowers mottled with brown on outer side, Jun 1936, Brass, L.J. 6976 (A n.v.,

BM, BRI n.v., LAE [Sheet 72124]); Palmer River, 2 miles below junction Black River, 100 m,

small climber in ridge forest, Jun 1936, Brass, L.J. 7021 (BM, BRI n.v.); Palmer River, 2 miles

below junction with Black River, 100 m, slender climber in ridge forests, Jun 1936, Brass, L.J.

7072 (A); cultivated plant grown on from Black River, near Dahamo, living accession NS 11-

100, vouchered at Ukarumpa on 27 May 2012 as Simonsson Juhonewe, N. & Juhonewe, F.

NS0034L (LAE); cultivated plant grown on from Black River, near Dahamo, living accession

NS 11-129, vouchered in Ukarumpa on 22 Jun 2012 as Simonsson Juhonewe, N. & Juhonewe,

F. NS0042L (LAE).

Notes. A small-flowered species from the lowlands of PNG Western Province that

may be confused with the widespread and common Hoya globulifera Blume because

both species have linear-lanceolate leaves with pinnate venation and secondary veins

held at an acute angle. It is easily distinguished from Hoya globulifera by its very

short petioles that are only 1-2 mm long, by the peduncles that are c. 1 mm long and

the inflorescences that have 1-10 cream white to pale yellow flowers, often spotted in

(brownish) red with short hairs on the inside of the corolla. Hoya globulifera instead

has petioles > 4 mm long, peduncles > 1 cm long, and inflorescences bearing 10-30

flowers with a colour range from dull yellow-purple to rich maroon with corollas

covered by long hairs on the inside. The taxon was long recognised as a new species

by Paul Forster and David Liddle and the type specimen Brass, L.J. 6883 was already

labelled as such by them. P.I.Forst. & Liddle are therefore recognised as “ex” authors

for Hoya brassii.

Hoya carrii P.I.Forst. & Liddle ex Simonsson & Rodda sp. nov.

Similar to Hoya solaniflora Schltr. in the rotate corolla, but distinguished from it by the

plant being a vigorous climber with thick leaves (up to 3 mm thick) and with a corolla

3 cm in diameter when flattened, whereas H. solaniflora is a more slender vine, with

thinner laminae (0.5-1.5 mm thick), and flowers c. 2 cm in diameter when flattened. -

TYPE: Papua New Guinea, Oro Province, Isuarava, epiphytic climber, forest c. 4500

ft. [c. 1370 m] , 3 March 1936, Carr, C.E. 15904 (holotype SING [SING0227204];

isotype B [B100277188], BM [BM001014247], BRI n.v., K, NY [NY00074132],

SING [SING0227204], LAE). (Fig. 5-7)

Epiphytic climber , vigorous with white latex in all vegetative parts. Stems cylindrical,

2-4 m long, 1.5-3 mm in diam., greyish green, glabrous or rarely sparsely pubescent on

younger stems; older stems leafless, up to 5 mm in diameter, greyish brown, glabrous with

peeling bark; intemodes 1.5—15(—20) cm long. Adventitious roots sparsely produced

along the stems. Leaves petiolate; petiole terete, 6-12 x 1-1.5 mm in diam., green

when young, greyish brown when old, glabrous; lamina narrowly elliptic-acuminate,

very thick and fleshy, often up to 3 mm thick, (4—)7—13 x (1.5—)2—4 cm, dark green on

adaxial surface, light green on abaxial surface, glabrous, apex acuminate to acute, base

Hoya of Papuasia I.

107

rounded, sometimes slightly revolute, more evident on dry material, immature leaves

often reddish-maroon, with 1 basal colleter, globose, 3-4 x 3M mm; venation pinnate,

midrib depressed on adaxial surface, secondary veins 2-5 each side, light green and

sometimes slightly elevated, anastomosing near the leaf margin, branching from the

midrib at 40-90°. Inflorescences one per node, extra-axillary, positively geotropic,

convex, consisting of (2-)6-12 flowers; peduncle terete, 1-6 cm x 1-2 mm, older

peduncles forming a rachis from previous flowerings, glabrous or sparsely pubescent;

pedicels terete, 2-A cm x c. 0.6 mm in diameter, pedicel reddish green. Flower buds

apically flattened, cream-white, often red near the calyx. Calyx c. 6 mm in diameter,

lobes elliptic-deltoid, c. 1.5 x 1.5-2 mm, apex rounded or acute, glabrous to sparsely

pubescent; basal colleters one in each sepal sinus, round to deltoid. Corolla rotate,

concave, c. 2.5 cm in diameter (c. 3 cm when flattened), pure white to cream-white with

varying degree of crimson under the corona, often also visible on the outer surface;

tube 5-6 mm long, lobes ovate, 0.7-1 x 0.7-1 cm, thickly pubescent inside, glabrous

outside, margin recurved, apex acuminate, recurved. Corona staminal, c. 3 mm high,

8-10 mm in diameter, fleshy, creamy yellow(-greenish); corona lobes 5, rhomboid

when seen from above with ovate top and a central deep groove, 4-5 x c. 3 mm, inner

processes flat to slightly erect, apex acuminate, outer processes flat to slightly erect,

usually at same level as inner processes, not basally fused with the filament tube,

with basal revolute margins. Style-head convex, hidden by anther appendages at same

level or barely exceeding the apices of inner coronal processes. Pollinia oblong with

sterile edge, c. 920 x 370 pm, corpusculum rhomboid, c. 500 x 300 pm, caudicles

attached at the sides of corpusculum, c. 120-150 pm long. Ovary conical, often with a

mammillate tip, c. 2 mm long; each carpel c. 1 mm wide at base, glabrous. Fruits and

seeds not observed.

Distribution. Carr collected Hoya carrii (Carr 13084, Carr 13500, Carr 14768 ,

Carr 15366 ) at two sites c. 15 km apart on the Kokoda Track along the border of

Central and Oro Provinces, at c. 1370 m altitude (4500 ft.). Later, during the first

author’s fieldwork, two populations were found 25 km apart, c. 340 km northwest of

the type locality ( Simonsson Juhonewe & Juhonewe NS0066H , Simonsson Juhonewe

& Juhonewe NS0078H, Simonsson Juhonewe & Juhonewe NS0079H, Simonsson

Juhonewe & Juhonewe NS0097L ) in Obura-Wonenara, Eastern Highlands Province,

at c. 1800 m altitude in mossy forests on ridges, flowering in January and November.

The species seems to be uncommon, but where it is found it is locally abundant.

Etymology. Named after C. E. Carr (1892-1936), who first collected the taxon in

1935, and again in 1936, on a collecting expedition supported by the British Museum,

London, which started in 1934. He died of malaria in June 1936, just a few months

after discovering the second locality of Hoya carrii.

Habitat and ecology. Found on ridges at 1400-1800 m, on mature trees or mossy

undergrowth in primary forests. Not observed in nearby re-growth. In the wild and in

cultivation in PNG it usually flowers during November - January.

108

Card. Bull. Singapore 69(1) 2017

500 |jm

Fig. 5. Hoya carrii Simonsson & Rodda. A. Corolla, top view. B. Corona, side view. C. Corona,

from underneath. D, E. Calyx. F. Pollinarium. All from Simonsson Juhonewe & Juhonewe

NS0066H. (Drawing: M. Rodda)

Hoyci of Papuasia I.

109

Fig. 6. Hoya carrii Simonsson & Rodda. A. Proudly displayed as an adornment by a local

collector B. In cultivation at Ukarumpa, inflorescence from beneath (living plant NS 13-064).

C. In cultivation at Ukarumpa, inflorescence from above (living plant NS 13-064). (Photos: N.

Simonsson Juhonewe)

110

Card. Bull. Singapore 69(1) 2017

Fig. 7. Hoya carrii Simonsson & Rodda. A, B. Corolla, top view. C. Corona, side view. D, E.

lamina, from above. (A, D from live plant NS 13-064, C from NS 13-092, B, E from NS 13-

065). (Photos: N. Simonsson Juhonewe)

Provisional IUCN conservation assessment. Critically Endangered CR B2a,b(iii)

(IUCN, 2016). The type locality is along the famous Kokoda Track that is still covered

by thick forest, but further development along the track with expanding population

may reduce the forest cover greatly. The forest in the two recently recorded localities is

under high pressure from expanding population because of slash-and-burn agriculture

that later turns the landscape into eroded abandoned grassy fields. Hoya carrii has not

been recorded elsewhere, neither in herbaria nor during the first author’s fieldwork.

Hoya of Papuasia I.

Ill

Additional specimens examined. PAPUA NEW GUINEA: Central Province: Boridi, 10

Oct 1935, Carr, C.E. 13084 (BM, SING [SING0121812]); Boridi, 2 Nov 1935, Carr, C.E.

14768 (BM, SING [SING0121815]); Boridi, 25 Nov 1935, Carr, C.E. 13500 (BM, SING

[SING0121814]); Oro Province: Isuarava, 3 Feb 1936, Carr, C.E. 15366 (BM, SING

[SING0121813, SING0121816, SING0121817]); Eastern Highlands Province: Obura-

Wonenara, Baira, 25 Jan 2013, Simonsson Juhonewe, N & Juhonewe, F. NS0066H (LAE,

SING) (living accession NS 13-065); Obura-Wonenara, Kamanangkera, 27 Nov 2013,

Simonsson Juhonewe, N., Juhonewe, F. & ApaApako NS0079H (LAE) (living accession NS 13-

092); Obura-Wonenara, Kamanangkera, 27 Nov 2013, Simonsson Juhonewe, N., Juhonewe, F.

& Apa Apako NS0078H (LAE) (living accession NS 13-093); cultivated plant grown on from

Obura-Wonenara, Baira, living accession NS 13-064, vouchered at Ukarumpa on 14 Jan 2016

as Simonsson Juhonewe, N & Juhonewe, F. NS0097L (LAE).

Notes. A montane species not easily confused vegetatively with any other Hoya spp.

due to its unusual thick leaves. Amongst other known New Guinean species it most

resembles Hoya solaniflora but the stems are more vigorous and robust, the lamina

is larger and very thick (often 1.5-3 mm) and usually breaks easily if bent, and the

flowers are c. 3 cm in diameter when flattened, whereas H. solaniflora is a more slender

vine, having laminae 0.5-1.5 mm thick, and the flowers are c. 2 cm in diameter when

flattened.

The taxon was long recognised as a new species by Paul Forster and David

Liddle and the type specimen Carr, C.E. 15904 was already labelled as such by them.

Hoya edholmiana Simonsson & Rodda, sp. nov.

Similar to the species in the Hoya ischnopus (Warb.) Schltr. species complex (which

includes H. dimorpha F.M.Bailey, H. dischorensis Schltr., H. flavescens Schltr., H.

kenejiana Schltr. and H. montana Schltr.) in having numerous flowers bearing yellow

densely pubescent rotate corollas on slender peduncles, but easily distinguished by the

conspicuous bright, raised venation which is lacking in the Hoya ischnopus complex

and the corollas being slightly campanulate with grooves under each corona lobe and

1.5 to 2 times the size of the corolla in the Hoya ischnopus complex (measured flat).

- TYPE: Papua New Guinea, Morobe Province, Skindiwai, Kuper Range, 21 km E of

Wau, 7°19'S 146°55'E, 1600 m, Nothofagus forest, 24 November 1981, Kairo, A. 405

(holotype LAE). (Fig. 8, 9)

Slender liana with white latex in all vegetative parts. Stems cylindrical, 1.5-3

mm in diameter, greyish green to dark purple, glabrous; older stems often leafless,

lignified, up to 5 mm in diameter, grey-brown, with peeling bark; internodes 3-20 cm

long. Adventitious roots numerous along the stem. Leaves petiolate; petiole terete,

1.5-2 x c. 1.2 mm in diameter, greyish green, glabrous; lamina ovate-elliptic, thin,

coriaceous, 5-12 x 2-5 cm, dark green to purplish on adaxial surface, light green on

abaxial surface, glabrous, apex cuspidate to caudate, base round to acuminate, margin

entire, often undulate, slightly revolute; venation pinnate, midrib depressed on adaxial

surface, secondary veins 4-6 each side, reticulate or inconspicuous, raised adaxially,

much lighter in colour when young, anastomosing near the leaf margin, branching

112

Card. Bull. Singapore 69(1) 2017

from the midrib at 50-80°; basal colleters two, light brown. Inflorescence one per

node, extra-axillary, positively geotropic, pseudo-umbelliform, convex, consisting of

7-20 flowers; peduncle persistent, terete, slender, 4-15 cm x 1-1.5 mm, glabrous,

older peduncles forming a rachis from previous flowerings; pedicels terete, 2-A cm

x c. 1 mm in diameter, light green to purplish red, glabrous. Flower buds cream

coloured, flushed pink on parts exposed to the sun. Flowers lasting about one week in

cultivation, with a weak scent during the day, more intense at night. Calyx 4-10 mm

in diameter, calyx lobes triangular to lanceolate, 1.5-5 x c. 1.5 mm, glabrous, apex

rounded to acute, margin glabrous or ciliate. Corolla spreading, 16-22 mm in diameter

(19-25 mm when flattened), cream to pale yellow, flushed pink on the outer surface

in specimens growing in bright light; tube 5-7 mm, lobes triangular, slightly fleshy,

6- 9 x 5-7 mm, inside thickly and densely covered with 0.5-1 mm long hairs, outside

glabrous, margins recurved, apex acute, recurved. Corona staminal, 2.5-3.5 mm high,

7- 9 mm in diameter, fleshy, yellow, somewhat translucent; lobes ovate-elliptic, 3^.5

x 2-2.5 mm, inner processes cuspidate, slightly erect, not meeting (or just barely)

in the centre, outer processes apex rounded to obtuse (acute), slightly erect, c. 1 mm

higher than inner processes, carinate above, with basal revolute margins. Style-head

convex, hidden by anther appendages. Pollinia oblong, c. 500 x 200 pm, with sterile

edge, corpusculum ovate 200-250 x 100-120 pm, caudicles attached in the middle of

corpusculum, c. 70 x 30 pm. Ovary broadly conical c. 2 mm long; each carpel c. 1 mm

wide at the base, light green-yellow, glabrous. Fruits cylindric and tapering, c. 20 x 1

cm (unripe), dark green becoming red-pigmented in sunlight, seeds not seen mature.

Distribution. The species is recorded from the Kuper Range (holotype Kairo 405) and

from three additional collections by the first author in Morobe Province (Fig. 1).

Etymology. Named after the siblings Charlize, Diesel and Evelize Edholm of Sweden,

supporters of the first author’s work in PNG.

Habitat and ecology. Hoya edholmiana was rather abundant at the three localities

where it was observed by the first author. It was recorded only at altitudes between

c. 1450 and 1850 m. It was absent both further down and further up the mountain

during the extensive expeditions, even on the same slope. It is usually an epiphyte

but on shaded or partially exposed ridges was also found growing as a lithophyte on

mossy rocks, entangled amongst other plants. Where it occurs, it is common in the

primary forests and can occasionally be found in adjacent mature secondary forests.

It thrives on younger trees along disturbed paths but is absent in the surrounding sun-

exposed and drier subsistence gardens, where instead H. magnifica P.I.Forst. & Liddle,

H. subglabra Schltr. and Hoya sp. sect. Eriostemma were recorded. The specimens

brought into cultivation flower abundantly throughout the year.

Provisional IUCN conservation assessment. Least Concern (IUCN, 2016). It appears

to be rather common and somewhat capable of re-colonising moist secondary forests.

It has only been recorded in Morobe Province.

Hoyci of Papuasia I.

113

Fig. 8. Hoya edholmiana Simonsson & Rodda. A. Fruiting plant in situ in Finisterre Range,

Yawan. B. Whole inflorescence. A from live plant NS 11 -210; B from Simonsson Juhonewe, N.,

& Juhonewe, F. NS0009H. (Photos: N. Simonsson Juhonewe)

114

Card. Bull. Singapore 69(1) 2017

Fig. 9. Hoya edholmiana Simonsson & Rodda. A, F. Corolla, top view. B, G. Corolla, from

underneath. C, H. Corona, side view. D, I. Corona, from underneath. E, J. Calyx. K. Pollinarium.

A-E from Simonsson Juhonewe, N., & Juhonewe, F. NS()()()8H ; F-J from NS 11-232; K from

Simonsson Juhonewe N. & Juhonewe, F. NS0039L. (Photos A-J: F. Juhonewe; K: M. Rodda).

Hoya of Papuasia I.

115

Additional specimens examined. PAPUANEW GUINEA: Morobe Province: Finisterre Range,

Yawan, 1713 m, 1 Apr 2011, Simons son Juhonewe, N., & Juhonewe, F. NS0008H (LAE), (living

accession NS11-212); Finisterre Range, Yawan, 1740 m, 1 Apr 2011, Simonsson Juhonewe, N.,

& Juhonewe, F. NS0009H (LAE) (living accession NS 11-218); Finisterre Range, Yawan, 1800

m, 1 Apr 2011, Simonsson Juhonewe, N., & Juhonewe, F. NS0010H (LAE) (living accession

NS11-226); cultivated plant grown on from Finisterre Range, Yawan, 1713 m, living accession

NS 11-212, vouchered at Ukarumpa on 21 Jun 2012 as Simonsson Juhonewe, N. & Juhonewe F.

NS0038L (LAE, SING); cultivated plant grown on from Finisterre Range, Yawan, 1740 m, living

accession NS11-218, vouchered at Ukarumpa on 21 Jun 2012 as Simonsson Juhonewe N. &

Juhonewe, F. NS0039L (LAE, SING); cultivated plant grown on from Finisterre Range, Yawan,

1800 m, living accession NS 11-224, vouchered at Ukarumpa on 21 Jun 2012 as Simonsson

Juhonewe, N. & Juhonewe, F. NS0040L (LAE); Huon Peninsula, Hoboc, 1830 m, 24 Dec

2011, Simonsson Juhonewe, N. & Juhonewe, F. NS0015H (LAE) (living accession NS 11-333);

cultivated plant grown on from Huon Peninsula, Hoboc, 1450 m, living accession NS11-245,

vouchered at Ukarumpa on 8 Jun 2012 as Simonsson Juhonewe, N. & Juhonewe, F. NS0035L

(LAE); Waria Valley, Saureli, 1700 m, 1 Mar 2012, Simonsson Juhonewe, N., Juhonewe, F.,

Allison, A. & James, S. NS0022H (LAE) (living accession NS 12-096); cultivated grown on

from Waria Valley, Saureli, 1700 m, living accession NS 12-103, vouchered at Ukarumpa on 21

Jun 2012 as Simonsson Juhonewe, N. & Juhonewe, F. NS0041L (LAE, SING).

Notes. This new species is morphologically similar to the as yet unresolved Hoya

ischnopus complex, but is easily distinguished by the raised, conspicuous reticulate

secondary veins on the leaves, especially when young, which makes it easy to identify

even when not in flower. The venation of the leaves is most raised and reticulate in the

populations from the middle of Finisterre and Saruwaged ranges, and less raised in the

populations from the area where the Central Range meets the Owen Stanley Range.

Also its pedicels are more slender, bearing flowers 16-22 mm in diameter, and the

corona lobes are ovate-elliptic with rounded to obtuse (rarefy acute) outer processes.

Species in the Hoya ischnopus- complex have plain leaves lacking the raised venation,

stiffer peduncles, flowers < 1.5 cm in diameter, and the corona lobes are rhomboid with

acute outer processes. Hoya edholmiana is only found above 1400 m while species in

the Hoya ischnopus complex can be found from sea level to 1400 m.

Hoya evelinae Simonsson & Rodda, sp. nov.

Similar to Hoya epedunculata Schltr. in having much reduced peduncles, but differs in

having peduncles along climbing stems, leaves < 8 cm and (narrowly) elliptic corona

lobes, while H. epedunculata has terminal peduncles, leaves > 8 cm in length and

obovate corona lobes. - TYPE: Papua New Guinea, Chimbu-Gulf Province, Crater

Mt. Wildlife Management Area, near junction between Wara O and Pio River. 440-

460 m, 30 March 1997, Takeuchi, W. 11941 (holotype A; isotypes K [K000224390],

L, LAE, US). (Fig. 10, 11)

Epiphytic climber with white latex in all vegetative parts, glabrous. Stems up to 5

m long, cylindrical, 1-3 mm in diameter, green; older stems lignified, up to 5 mm in

diameter, greyish brown, internodes 2—10(—25) cm long. Adventitious roots sparsely

116

Card. Bull. Singapore 69(1) 2017

scattered along the stems. Leaves often held horizontally, petiolate; petiole terete, 1-3

x 0.5 mm; lamina elliptic-ovate, coriaceous, 5-10 x 2-3(-4) cm, medium green on

both surfaces, apex acute to caudate, base round to attenuate; venation pinnate, midrib

depressed on adaxial surface, secondary veins obscure. Inflorescence one per node,

extra-axillary, pseudo-umbellate, consisting of 2-5 flowers; peduncle terete, much

reduced, c. 1 x 1 mm, older peduncles forming a rachis from previous flowerings,

pedicels terete, 2-3 cm x 0.5-1 mm in diameter, light green, glabrous. Calyx c. 5 mm

in diameter, calyx lobes ovate, c. 2 x 0.7 mm, apex acute-rounded. Corolla broadly

campanulate, almost rotate, c. 3 cm in diameter (2 cm when dry), pale pink, translucent;

tube c. 1 cm (c. 0.5 when dry), lobes broadly triangular, c. 1 x 1.2 cm, thinly pubescent

inside, almost glabrous near centre, outside glabrous, margins recurved, apex obtuse.

Corona staminal, c. 2 mm high, c. 8 mm in diameter, fleshy, creamy yellow with

maroon centre; lobes from above (narrowly) elliptic, c. 4 x 1.8 mm (c. 3 x 1 when

dry), inner processes acuminate, outer processes rounded (acute), with basal revolute

margins along the entire length of the corona lobe, not basally fused with filament

tube. Pollinia oblong c. 330-380 x 120-200 pm, with sterile edge, corpusculum

oblong with constriction in the middle, 150-200 x 100-150 pm, caudicles attached at

the lower half of the corpusculum, 70-80 x c. 50 pm. Ovary conical, 2-3 mm long,

glabrous. Fruit and seed not observed.

Distribution. The species is only known from the type locality on Crater Mountain and

from Black River in Western Province. It may be locally endemic as it has not been

recorded elsewhere, neither in herbaria nor during the first author’s fieldwork.

Etymology. Named after the late Mrs Evelina Eriksson (1932-2009) of Sweden who

supported the first author’s work in PNG.

Habitat and ecology. A climber in riverine forest, often growing on trees or shrubs, or

even on mossy rocks, near or above streams. Not observed on taller trees or in sunnier

and drier areas, only in shade or filtered sunlight. Locally common.

Provisional IUCN conservation assessment. Critically Endangered CR Bla,b(iii)

(IUCN, 2016). The species co-occurs with Hoy a brassii at the type locality where

it is locally co mm on However much of PNG Western Province’s lowland rainforest

has been or is being logged (Shearman et al., 2009). The type locality is not inside a

protected area and may be lost in the near future.

Additional specimens examined. PAPUA NEW GUINEA: Western Province: Black River,

near Dahamo, 1 Mar 2011, Simonsson Juhonewe, N. & Juhonewe, F. NS0005H (LAE, SING)

(living accession NS 11 -115).

Notes. In Papuasia the only other species with much reduced peduncles are Hoya

epedunculata and Hoya brassii. Hoya evelinae can be separated from H. epedunculata

because it produces peduncles almost at every node when mature, and often has

numerous inflorescences with simultaneously open flowers, leaves < 8 cm long and

Hoya of Papuasia I.

117

Fig. 10. A. Hoya evelinae Simonsson & Rodda inflorescences with multiple flowers open. B.

C. Hoya epedunculata Schltr. inflorescences with a single flower open. (Photos: N. Simonsson

Juhonewe)

118

Card. Bull. Singapore 69(1) 2017

Fig. 11. Hoya evelinae Simonsson & Rodda. A. flower, top view. B. Corolla from underneath.

C. Corona, from underneath. D. Corona, top view. E. Pollinarium. All from Simonsson

Juhonewe, N. & Juhonewe, F. NS0005H. (Photos A-D: N. Simonsson Juhonewe; E: M. Rodda)

corona lobes (narrowly) elliptic, while H. epedunculata has fewer peduncles, often

terminal, leaves > 8 cm in length and its corona lobes are obovate. Hoya evelinae can be

separated from Hoya brassii because its flowers are > 2 cm in diameter (vs. 1-1.5 cm).

Hoya juhoneweana Simonsson & Rodda, sp. nov.

Similar to Hoya urniflora in the sunken leaf venation and bullate leaf surface, but

Hoya urniflora has corollas c. 1.5 cm wide whereas H. juhoneweana has corollas

> 2.5 cm wide. - TYPE: Papua New Guinea, Morobe Province, Huon Peninsula,

Hoboc, primary riverine forest, 1500 m, 31 December 2011, Simonsson Juhonewe,

N. & Juhonewe, F. NS0016H (holotype LAE; isotype SING) (living accession NS 11-

363). (Fig. 12-14).

Hoya of Papuasia I.

119

subsp. juhoneweana

Pendulous epiphyte with white latex in all vegetative parts. Stems up to 5 m long,

flexible, (horizontal) pendulous, cylindrical, 1—1.5 mm in diameter, green, pubescent

with very stiff, spreading 0.5 mm or longer stiff hairs; older stems lignified, up to

3 mm in diameter, greyish brown, often glabrous; internodes 3-6(-25) cm long.

Adventitious roots scattered along the stems. Leaves pendulous, petiolate; petiole

terete, 2-10 x 1-1.5 mm, green, covered with very stiff, erect, 1 mm or longer hairs;

lamina elliptic-ovate, chartaceous, stiff and brittle when mature, 3-8 x 2-4 cm, bright

green on adaxial surface, light green on abaxial surface, both surfaces covered by

very stiff, erect, 1 mm or longer hairs, especially near base and abaxial midrib, apex

cuspidate-caudate, base round to attenuate, margin entire and revolute; venation

pinnate, midrib much depressed on adaxial surface, secondary veins 2-5 each side,

evident and often depressed, giving the lamina a bullate appearance, anastomosing

near the leaf margin, branching from the midrib at 45-80°; leaf colleters one pair at

the lamina base, greyish white. Inflorescence one per node, extra-axillary, positively

geotropic, pseudo-umbellate, consisting of 1-5 flowers; peduncle terete, 1-3 cm x

c. 1 mm, older peduncles forming a rachis from previous flowerings, new peduncles

hirsute, often almost glabrous in old peduncles; pedicels terete, c. 2 cm x c. 1 mm

in diameter, light green, often spotted in purple near calyx, glabrous (sparsely

pubescent). Flower buds globular with flattened apex, creamy white with pink and

purple dots. Calyx c. 5 mm in diameter, calyx lobes ovate-lanceolate, c. 1 x 0.7 mm,

apex acute, purple spotted. Corolla deeply campanulate, fleshy, 2.5-3 cm in diameter,

cream coloured with a slightly greenish hue in centre fading to a pinkish hue on lobes

penetrating from the outer surface; tube 1—1.5 cm long x 1.5-2 cm in diameter, lobes

broadly triangular, c. 1 x 1.5 cm, thickly pubescent inside, usually almost glabrous

near centre, outside glabrous, margins recurved, apex acute. Corona staminal, c. 5

mm high, c. 9 mm in diameter, fleshy, cream coloured with a light green protruding

style-head; lobes round from above, c. 3.5 x 3 mm, inner processes much reduced

and linear, 0.5 x 0.2 mm, each outer process with a sharp central ridge and a basal

longitudinal protrusion on each side running from the base of the guide rails up to

the apex, with basal revolute margins right up to the apex. Style-head mamillate,

higher than anther appendages. Pollinia oval, 400-500 x 280-320 pm, no sterile

edge, retinaculum ovoid, 350^100 x c. 300 pm, caudicles attached at the base of

retinaculum, broadly triangular, 100-150 x c. 100 pm. Ovary conical, c. 3 mm long,

narrowing down at 2 mm from the base; each carpel c. 1.3 mm wide at base, light

green, glabrous. Fruit and seed not observed.

Distribution. Only found near the remote village of Hoboc at 1500-1800 m. Four

individuals of Hoya juhoneweana ssp. juhoneweana were originally seen, of which

two had disappeared when their locality was revisited for the last time in 2012 due

to expanding subsistence gardens and degradation of the riverine forest by the local

villagers. Hoya urniflora was collected at the same locality and is abundant there.

120

Card. Bull. Singapore 69(1) 2017

Fig. 12. Comparison between Hoya juhoneweana ssp. lindforsiana Simonsson & Rodda

(A, D, from Simonsson Juhonewe, N. & Juhonewe, F. NS0068L ), Hoya juhoneweana ssp.

juhoneweana Simonsson & Rodda (B, E, from Simonsson Juhonewe, N. & Juhonewe, F.

NS0016H ) and Hoya urniflora (RI.Forst.) Simonsson & Rodda (from Simonsson Juhonewe,

N. & Juhonewe, F. NS0102L ). A-C. Corolla from above. D-F. Corolla, side view. (Photos: F.

Juhonewe)

Etymology. Named after the first author’s colleague, Foreting Juhonewe, who is also

from Hoboc village. He has been of great importance for this research. He has eagerly

tried to find more of this rare and diminishing Hoya and promoted its conservation in

situ amongst his clan’s people.

Habitat and ecology. This taxon co-occurs with two other new species described here,

H. edholmiana and H. urniflora. At the type locality this taxon and Hoya urniflora

inhabit only mossy riverine forest on mature trees in primary forest and were not

found in the adjacent mature secondary forest and in the subsistence gardens. The co-

occuring taxon Hoya edholmiana , however, is more abundant and also occasionally

found in mature secondary forest.

Provisional IUCN conservation assessment. Critically Endangered CR Blab(ii,iii,v),

D (IUCN, 2016). This subspecies has only been recorded at the type locality, despite

extensive field expeditions in the Finisterre-Sarawaged Range by the first author and

Hoyci of Papuasia I.

121

Fig. 13. Hoya juhoneweana ssp. juhoneweana Simonsson & Rodda in habitat. A. The pendent

epiphyte at Hoboc, the type locality. B. Inflorescence with two open flowers. All based on

Simonsson Juhonewe, N. & Juhonewe, F. NS0016H. (Photos: A: S. Pimm Lyon; B: N.

Simonsson Juhonewe).

122

Card. Bull. Singapore 69(1) 2017

Fig. 14. Hoya juhoneweana subsp. juhoneweana Simonsson & Rodda. A. Flower from above.

B. Flower, side view. C. Corolla and calyx, from underneath. D. Corona, from underneath. E.

Corona, side view. F. Corona, top view. G. Pollinarium. All based on Simonsson Juhonewe, N.

& Juhonewe, F. NS0016H. (Photos A-F: F. Juhonewe; G: M. Rodda)

Hoya of Papuasia I.

123

many other botanists (Vollering et al., 2016). The population size at the type locality

was reduced from 2010 to 2012 due to expanding subsistence agriculture.

Additional specimens examined. PAPUA NEW GUINEA: Morobe Province: Huon Peninsula,

Hoboc, 1830 m, 24 December 2011, living accession NS 11-332, vouchered at Ukarumpa on 2

June 2013 as Simonsson Juhonewe, N. & Juhonewe, F. NS0074L (LAE).

Notes. Hoy a juhoneweana is easily distinguished from the large majority of Hoy a

species due to the conspicuous sunken venation causing a characteristic bullate leaf

surface, and by its dense cover of >1 mm long hairs on all vegetative parts, especially

when young. Its leaves are, however, indistinguishable from Hoya urniflora. The two

species can be separated when sterile because Hoya juhoneweana is pendulous while

H. urniflora is a climber. When in flower, the two taxa are easily distinguished, as Hoya

juhoneweana has >2.5 cm wide corollas whereas the corollas of H. urniflora are c. 1.5

cm wide. Interestingly, the two species co-exist and intermingle at the same locality

leading us to initially assume they were a single species during early expeditions.

Upon flowering in cultivation, however, we realised there were two distinct species

growing together in the same area.

Hoya juhoneweana subsp. lindforsiana Simonsson & Rodda, subsp. nov.

Similar to Hoya juhoneweana ssp. juhoneweana and Hoya urniflora in having sunken

leaf venation and bullate leaf surface, but easily distinguished by the diameter of its

corolla (4-5 cm), whereas it is 2.5-3 cm in H. juhoneweana subsp. juhoneweana and

c. 1.5 cm in Hoya urniflora. - TYPE: Papua New Guinea, Madang Province, Mebu,

disturbed primary riverine forest, 1300 m, living accession NS 12-279, vouchered

at Ukarumpa on 9 March 2013, Simonsson Juhonewe, N. & Juhonewe, F. NS0068L

(holotype LAE; isotype SING). (Fig. 12, 15)

Slender pendulous epiphyte with occasional twining branches, with white latex in all

vegetative parts. Stems up to 5 m long, flexible, often pendulous, cylindrical, 1-1.5

mm in diameter, green, pubescent with very stiff, spreading, 0.5 mm or longer hairs;

older stems lignified, up to 3 mm in diameter, greyish brown, pubescent or glabrous;

intemodes 3-6(-25) cm long. Adventitious roots scattered along the stems. Leaves

pendulous, petiolate; petiole terete, 2-10 x 1-1.5 mm, green, covered with very

stiff, erect 1 mm or longer hairs; lamina elliptic-ovate, chartaceous, stiff and brittle

when mature, 3-8 x 2-4 cm, bright green on adaxial surface, light green on abaxial

surface, both surfaces covered by very stiff, erect, 1 mm or longer hairs, especially

near base and abaxial midrib, apex cuspidate to acute, base round to attenuate, margin

recurved; venation pinnate, midrib depressed on adaxial surface, secondary veins

2-5 each side, evident and often depressed, giving the lamina a bullate appearance,

anastomosing near the leaf margin, branching from the midrib at 45-80°; leaf colleters

one pair at each lamina base, greyish white. Inflorescence one per node, extra-axillary,

124

Card. Bull. Singapore 69(1) 2017

positively geotropic, pseudo-umbellate, consisting of 1-5 flowers; peduncle terete,

1-3 cm x c. 1 mm, older peduncles forming a rachis from previous flowerings, new

peduncles hirsute, old peduncles often almost glabrous; pedicels terete, 2-3 cm x c.

1 mm in diameter, light green, often spotted in purple near calyx, glabrous (sparsely

pubescent). Flower buds apically flattened, creamy white with pink and purple dots.

Calyx c. 5 mm in diameter, calyx lobes ovate-lanceolate, c. 2 x 1.5 mm, apex acute

and ciliate. Corolla broadly campanulate, almost rotate, fleshy, 4-5 cm in diameter,

cream-white coloured with a slightly greenish-yellow translucent hue, outside often

pink-maroon if exposed to bright light; tube 1.2-1.5 cm long, glabrous to puberulous

within, glabrous near centre, outside glabrous; lobes broadly triangular with upturned

auricles at the sinuses, c. 1.5 x 1.5 cm, margins entire and slightly revolute, apex acute,

inside sparsely pubescent, outside glabrous. Corona staminal, c. 6 mm high, c. 14 mm

in diameter, fleshy, cream-white; lobes broadly elliptic, c. 6 x 4.5 mm, inner processes

much reduced and linear, 0.8 x 0.3 mm, each outer process with a sharp central ridge

and a basal longitudinal protrusion on each side running from the base of the guide

rails up to the apex, with basal revolute margins right up to the apex, not basally fused

with filament tube. Style-head basal convex process narrowed into a raised convex

tip, much exposed and not hidden by anther appendages, light green. Pollinia elliptic,

900-950 x 400-500 pm, without sterile edge, corpusculum ovate, 500-600 x 400-

450 pm, caudicles attached at lower half of corpusculum, 300-350 x 130-150 pm.

Ovary conical, c. 3 mm long, narrowing at 2 mm from the base; each carpel c. 1.5 mm

wide at base, light green, glabrous. Fruit and seed not observed.

Distribution. Only found in the Mebu area, Madang Province at 1300 m altitude,

where 12 specimens were sighted at only one site near a deep, moist montane valley. It

has never been recorded outside that locality, even in other parts of the same mountain

range.

Etymology. Named after Constantinus Lindfors in Sweden, a supporter of the first

author’s work in PNG.

Habitat and ecology. Hoya juhoneweana subsp. lindforsiana inhabits mossy riverine

primary forests where it grows on large mature trees but is absent in the adjacent

mature but drier secondary forests and in the subsistence gardens. The co-occuring

taxa Hoya oreostemma Schltr. and Hoya solaniflora are more abundant and grow well

on younger trees along paths and in mature secondary forest with more sunlight. The

inability to recolonise secondary habitats has also been observed for the two similar

taxa H. juhoneweana subsp. juhoneweana and H. urnijiora.

Provisional IUCN conservation assessment. Critically Endangered CR B2a,b(iii), D

(IUCN, 2016). Despite extensive field expeditions in the Finisterre-Sarawaged Range

by the first author and many other botanists (Vollering et al., 2016) this subspecies has

only been recorded at the type locality, where a single population including only 12

individuals was seen in an area of < 1 km 2 .

Hoyci of Papuasia I.

125

Fig. 15. Hoya juhoneweana subsp. lindforsiana Simonsson & Rodda. A. Flower from above.

B. Corolla and calyx, from underneath. D. Corona, side view E. Corona, top view. F. Calyx

and ovaries. G. Pollinarium. All based on Simonsson Juhonewe, N. & Juhonewe, F. NS0068L.

(Photos A-F: N. Simonsson Juhonewe; G: M. Rodda)

126

Card. Bull. Singapore 69(1) 2017

Notes. In vegetative appearance Hoya juhoneweana subsp. lindforsiana is not easily

distinguished from the typical subspecies. When in flower, however, they are easily

distinguished, even in the bud stage. Hoya juhoneweana subsp. juhoneweana has

globular buds with flattened apices, a more narrowly campanulate corolla, 2.5-3 cm

wide, and a corona c. 0.9 cm wide, whereas H. juhoneweana subsp. lindforsiana has

wide flattened buds, a broadly campanulate, almost rotate corolla, 4-5 cm wide, and

corona c. 1.3 cm wide. The two subspecies are so far only known from their respective

separate deep montane valleys on each end of the Finisterre-Sarawaged Range, about

170 km apart.

Hoya krusenstierniana Simonsson & Rodda, sp. nov.

Similar to Hoya solaniflora Schltr. being thin wiry climbers with small leaves, 1-7 x

0.5-4.5 cm, and even flowering herbarium material is difficult to distinguish. In the

living state, however, it is easily distinguished by its purple to rich crimson flowers

and by its scent, which is reminiscent of peeled unripe green bananas, whereas Hoya

solaniflora bears pale flowers in shades of white-yellow-pink, and with a scent more

similar to Gardenia jasminoides J.Ellis. - TYPE: Papua New Guinea, Morobe Province,

Mount Kaindi, 7°20'S 146°45'E, edge of mossy forest, climber straggling in regrowth,

7650 ft [c. 2330 m], 2 November 1966, Ridsdale, C.E. NGF 30193 (holotype LAE;

isotypes L n.v., BRI n.v., CANB n.v.). (Fig. 16, 17)

Epiphytic climber with white exudate in all vegetative parts. Stems cylindrical, 1-5

m long, 1-2 mm in diameter, dark green, glabrous or sparsely pubescent; older stems

leafless, up to 3 mm in diameter, greyish brown, glabrous; internodes 1—10(—20) cm

long. Leaves petiolate; petiole terete, 3-10 x c. 1 mm in diameter, green, pubescent

(glabrescent with age); lamina ovate-lanceolate, thick, fleshy, 1.5-7.5 x 1-3.6 cm, bright

to dark green on adaxial surface, light green on abaxial surface, glabrous or sparsely

pubescent on both surfaces when young, glabrous when old, apex obtuse to narrowly

acuminate to cuspidate, base round to attenuate with basal colleters, margin sometimes

recurved; venation pinnate, midrib depressed on adaxial surface, secondary veins 2-4

each side, often slightly depressed on adaxial surface and anastomosing near the leaf

margin, branching at 30-80°. Inflorescences one per node, extra-axillary, positively

geotropic, flat to convex, consisting of 5-15 flowers; peduncle terete, peduncle (0.5-

) 1.5—7 cm x 0.7-1 mm, older peduncles developing arachis from previous flowerings,

new peduncles sparsely pubescent, older peduncles often glabrous; pedicels terete,

1-3 cm x 0.5-0.7 mm in diameter, green, pubescent. Flower buds apically flattened,

green to maroon. Flower scent like peeled green, unripe bananas. Calyx 4-6 mm in

diameter, lobes ovate-elliptic, 1.5-2 x 1-1.5 mm, apex rounded to acute, glabrous on

both sides, margin ciliate; basal colleters absent. Corolla rotate, slightly concave, c.

1.8 cm diameter (c. 2 cm when flattened), pink to rich crimson; tube c. 6 mm long,

lobes triangular to broadly acuminate, c. 5 x 7 mm, adaxially thickly pubescent,

abaxially glabrous, margins entire, apex acute. Corona staminal 2-3 mm high, c.

7 mm in diameter, fleshy, intensely crimson; lobes variable in shape from broadly

ovate to lanceolate, even within the same population, 4 x 1.5-2 mm, inner processes

Hoyci of Papuasia I.

127

5 mm

SOOpm

5 mm

Fig. 16. Hoya krusenstierniana Simonsson & Rodda. A. Flower, top view. B. Corona, top view.

C, D. Corona, from underneath. E, F. Corona, side view. G. Calyx and ovaries, side view. H.

Calyx and ovaries, top view. I. Pollinarium. A, C, E, G, H from Simonsson Juhonewe, N. &

Juhonewe, F. NS0019H; B, D, F, I from Simonsson Juhonewe, N. & Juhonewe, F. NS0020H.

(Drawing: M. Rodda)

128

Card. Bull. Singapore 69(1) 2017

Fig. 17. Hoya krusenstierniana Simonsson & Rodda in habitat on Mount Kaindi. A. Mature

plant growing on tree trunks in mossy forest. B. Flowers from two different plants showing

variation in the shape of the corona (above from Simonsson Juhonewe, N. & Juhonewe,

F. NS0019H , below from from Simonsson Juhonewe, N. & Juhonewe, F. NS0020H ). C.

Inflorescence. (Photos: F. Juhonewe)

Hoya of Papuasia I.

129

elongated, flat to erect, outer processes apex rounded to acute when observed from

above, slightly erect both higher and lower than inner corona apex, grooved between

inner and outer processes, not fused with filament tube, with basal revolute margins.

Style-head convex, hidden by both anther appendages and inner corona processes.

Pollinia oblong, 650 x 270 pm, with sterile edge, corpusculum oblong, c. 310 x 170

pm, caudicles attached at the sides of corpusculum, c. 100 pm long. Ovary conical

with a rounded to distinct mammillate tip, c. 2 mm long; each carpel c. 1 mm wide at

base, glabrous. Fruits c. 18 cm long, 0.5 cm wide, purplish. Seeds not observed.

Distribution. Recorded pr im arily at 1800-2600 m on Mount Kaindi in Morobe

Province, Mount Pimbilyo in Enga Province and along the Kokoda Track in Central

and Oro Provinces.

Etymology. Named after the Krusenstiern family, supporters of the first author’s work

in PNG.

Habitat and ecology. Hoya krusenstierniana has the highest altitudinal record for

Hoy a in Malesia, as it grows at 1800-2600 m. The second highest record for the genus

is Hoya nyhuusiae Kloppenb. which is found up to 2200 m on Mt Kinabalu in Sabah

(Borneo). On the summit of Mount Kaindi, Hoya krusenstierniana is found creeping

and twining on dwarf mossy trees or on a thick layer of moss on the ground, both under

shade and in fully exposed locations. It is also found twining among taller growth

where it produces bigger leaves on the slopes near the summit or towards Edie Creek

mine. It flowers throughout the year.

Provisional IUCN conservation assessment. Endangered EN Blab(iii) (IUCN,

2016). The type locality was already disturbed when the species was first collected

in the 1960s; however it was still abundant during the most recent collection in 2012.

However, now the area faces a rapidly expanding population so when the first author

was there in 2012, people had begun settling down on the summit of Mount Kaindi and

were in process of clearing both the primary and secondary forest to create subsistence

gardens. This, in combination with people practising slash-and-bum agriculture

downhill which can lead to rapidly spreading out-of-control fires, may pose threats to

the future of the type locality. However, the species has also been recorded on other

mountain tops (Tari & Reeve 4352 , Carr 13714), suggesting it may be thriving on

several of Papua New Guinea’s uninhabited cold mountain tops too inaccessible for

expeditions and most subsistence gardeners. The type locality has seen a decline in the

population of Hoya krusenstierniana and only rather old collections are known from

the other localities.

Additional specimens examined. PAPUA NEW GUINEA: Morobe Province: Mount Kaindi,

7°25'S 146°45'E, Nothofagus forest understorey, 7500 ft [c. 2285 m], 9 Oct 1964, Womersley,

J.S. NGF 19432 (LAE, L n.v., BRI n.v., CANB n.v.); Mount Kaindi, S 7 20, E 146 45, 6500

ft [c. 1980 m], 9 April 1965, Taylor, G. J. NGF 20965 (LAE, L n.v.); Mount Kaindi, 7°25'S

146°45'E, regrowth of mossy forest, 7500 ft [c. 2285 m], 24 April 1965, Womersley, J.S. NGF

24514 (LAE); Edie Creek, 7°20'S 146°45E, vine on undergrowth, 6500 ft [c. 1980 m], 15 Jan

1966, Henty, E.E. NGF 27170 (LAE, L n.v.); Mount Kaindi, understorey of montane forest,

130

Card. Bull. Singapore 69(1) 2017

7500 ft, 2 June 1967, Womersley, J.S. NGF 24966 (LAE); Mount Kaindi, steep slope in moss

forest, twining in undergrowth, c. 8000 ft [c. 2440 m], 7 Jun 1968, Coode, M., Woods, P.

& Katik, P. NGF 32858 (LAE, L n.v., BRI n.v.); Mount Kaindi, near helicopter pad, 7°20'S

146°45'E, regrowth, 7500 ft [c. 2285 m], 18 Dec 1968, Vandenberg, J. NGF 42161 (A, LAE, L

n.v., CANB n.v., K); Mount Kaindi, 7°23'S 146°42'E, on side of road in disturbed area, climber

over secondary species, 7200 ft [c. 2195 m], 26 Feb 1970, Streimann, H, & Kairo, A. NGF

47516 (LAE, L n.v., BRI n.v.); Mount Kaindi, near summit, montane forest edge, 7800 ft [c.

2380 m], 1 Jul 1972, Beamish, K. 33 (LAE); Mount Kaindi, 7°29'S 146°40'E, regrowth forest,

2438 m, 28 May 1974, Katik, P. & Larivita, G. LAE 62075 (LAE,L n.v., BRI n.v.); Mount

Kaindi, summit region, S 7° E 146°, mossy forest, c. 2360 m, 28 Oct 1974, Allison, A. 273

(LAE); Mount Kaindi, near repeater station, 7°10'S 146°40'E, Nothofagus dominated forest,

2400 m, 30 May 1977, Conn, B. & Kairo, A. 168 (LAE, BFC n.v.); Mount Kaindi, mossy

forest, 2400 m, 15 Nov 1979, Tagawa, H. 0472 (LAE); Mount Kaindi, summit, 2350 m, 8 May

1981, Reeve, T.M. 3668 (LAE, K, NSW n.v.); Mount Kaindi, upper montane forest, 2350 m,

11 Nov 1984, Cruttwell, N. 2718 (LAE); Mount Kaindi, on summit, 10 Jan 2012, Simonsson

Juhonewe, N. & Juhonewe, F. NS0019H (LAE, SING) (living accession NS 12-042); Mount

Kaindi, on summit, 10 Jan 2012, Simonsson Juhonewe, N & Juhonewe, F. NS0020H (LAE,

SING) (living accession NS 12-045); Enga Province: Mount Pimbilyo, near Laiagam, primary

Nothofagus forest, 2600 m, 20 Nov 1981, Tari, J. & Reeve, T.M. 4352 (LAE, K, NSW n.v.);

Oro Province: Kokoda Track, Alola, c. 6000 ft [c. 1830 m], Dec 1935, Carr, C.E. s.n. (SING

[SING0121847]); Central Province: Kokoda Track, The Gap, c. 7000 ft [c. 2135 m], 9 Dec

1935, Carr, C.E. 13714 (SING [SING0121849, SING0121850, SING0121851, SING121852]),

Kokoda Track, Above The Gap, c. 8000 ft [c. 2440 m], 2 Jan 1936, Carr, C.E. s.n. (SING

[SING0121848]).

Notes. Hoya krusenstierniana may be easily confused with the more common and

widespread Hoya solaniflora which thrives at lower elevations of 1200-1800 m. Both

sterile specimens and dry flowers of the two species can be easily confused as both

species have variable leaf shape and thickness. In the living state Hoya krusenstierniana

is easily distinguished by its purple to rich crimson flowers and by its scent, which is

reminiscent of peeled unripe green bananas, whereas H. solaniflora has pale flowers in

shades of white- yellow-pink and with a scent more similar to Gardenia jasminoides

J.Ellis. Usually the corona is a key character for species delimitation in Hoya as

it is very stable but obviously not for this taxon. The corona morphology of Hoya

krusenstierniana is quite variable within the population on Mt. Kaindi (Fig. 16, 17)

and is therefore not a good morphological character to distinguish the species. Instead,

it is the intense crimson colouring (rarely pale pink) and the unusual flower scent that

suggests a different pollinator than that of the similar Hoya solaniflora.

Hoya koteka Simonsson & Rodda, sp. nov.

Similar to Hoya multiflora Blume in its conical buds with five basal humps in line with

the corolla lobe sinuses. However, Hoya koteka has single-flowered inflorescences

while H. multiflora has inflorescences with numerous flowers. - TYPE: Indonesia,

Papua Province, Geitenkamp, 12 July 1907, Versteeg, G. 1431 (holotypeL [L0794363];

isotypes A, BO, L [L0794364]). (Fig. 18)

Hoya of Papuasia I.

131

Climber or spreading lax shrub or climber; presence of latex unknown. Stems

cylindrical, 1.5—4 mm in diameter, hirsute, older stems leafless, glabrous; internodes

2-5 cm long. Leaves petiolate; petiole terete, channelled above, 3-5 x c. 1 mm in

diameter, sparsely hirsute below, lamina narrowly elliptic, coriaceous when dry,

(5-)7-10 x 1.7-3.5 cm, puberulous on the abaxial surface, adaxial surface glabrous,

older leaves glabrous, apex caudate, base acute; venation pinnate, midrib depressed on

adaxial surface, raised on adaxial surface, secondary veins 4-7 each side, anastomosing

near the leaf margin, branching from the midrib at c. 60°; leaf colleters not present

on dry specimens. Inflorescences single-flowered, peduncle extra axillary, perennial,

terete, 1.5-3 mm x c. 1 mm, older peduncles bearing overlapping scars from previous

flowerings forming a short rachis, glabrous or sparsely pubescent; pedicels terete,

1.5-2 cm x c. 0.5 mm in diameter, glabrous. Flower buds conical, with five basal

humps in line with the corolla lobe sinuses. Calyx c. 2 mm in diameter, calyx lobes

triangular, c. 0.5 x 1 mm, apex acute, glabrous with basal lanceolate colleters, c. 50 pm

long. Corolla probably spreading (not possible to verity on herbarium material alone),

1.5-2 cm in diameter, tube 2-2.5 mm, lobes narrowly lanceolate, 0.7—10 x 2.5-3 mm,

pubescent inside, glabrous outside, apex acuminate. Corona staminal, stipitate, stipe

elongated conical-truncate, c. 2 mm high, c. 1 mm wide at base, tapering to c. 0.6 mm

at apex; corona c. 2.5 mm high, c. 2.7 mm in diameter, fleshy; corona lobes carinate,

the inner half attached at the back of the anthers, the outer half free, c. 2 x 0.5 mm,

inner processes acute, slightly bilobed at the tip, outer processes truncate, lacking

basal revolute margins. Style-head linear with round apex extending above the corona

lobes and anther appendages, 2-3 mm long. Pollinia ovoid, c. 250 x 180 pm, without

sterile edge, corpusculum ovoid, c. 120 x 70 pm, caudicles attached at the base of

corpusculum, thin and narrow, c. 100 x 15 pm. Ovary linear, 1.5-2 mm long, glabrous.

Fruits linear, 15-20 cm long.

Distribution. Recorded from a single locality in Papua Province, Indonesia.

Etymology. Named after its elongated style-head that resembles the traditional koteka,

or penis-gourd traditionally worn by male natives of New Guinea.

Habitat and ecology. Unknown.

Provisional IUCN conservation assessment. Data deficient (IUCN, 2016) as it is

known only from the type specimen collected 110 years ago.

Notes. Hoy a koteka is similar to Hoya multiflora Blume in the vegetative morphology,

as it appears to be a non-climbing epiphyte with thin leaves and lacks basal lamina

colleters. The general flower morphology is also rather similar as the corolla has

lanceolate lobes and the corona is stalked with elongated corona lobes. The flowers,

however, are much smaller in Hoya koteka (corona c. 2.5 mm high, c. 2.7 mm in

diameter in Hoya koteka vs. > 5 mm high and > 7 mm in diameter in H. multiflora ) and

132

Card. Bull. Singapore 69(1) 2017

Fig. 18. Hoya koteka Simonsson & Rodda. A. Whole plant. B. Flower, top view. C. Corona,

side view. D. Bud, E. Calyx and ovaries. F. Pollinarium. All based on Versteeg, G. 1431.

(Drawing: M. Rodda)

Hoya of Papuasia I.

133

the pollinia do not have a sterile edge, as has also been observed in Hoy a juhoneweana

and Hoya urniflora but otherwise present in most other Hoya species. Notable

exceptions are also the species in Hoya section Eriostemma and the species formerly

included in the genus Clemensiella.

Hoya stenakei Simonsson & Rodda, sp. nov.

Similar to Hoya onychoides P.I.Forst., Liddle & I.M.Liddle in flower size and clawed

corolla but differs from it by the almost pentagonal corona and densely pubescent

corolla whereas H. onychoides has a narrowly star-shaped corona and glabrous

corolla. - TYPE: Papua New Guinea, Sandaun Province, Baiberi, 1 March 2011,

lowland rainforest. Simonsson Juhonewe, N., & Juhonewe, F. NS0026H (holotype

LAE; isotype SING). (Fig. 19, 20)

Vigorous epiphytic climber with white latex in all vegetative parts. Stems cylindrical,

3-10 mm in diameter, dark green, new growth often slightly pubescent, later glabrous;

older stems leafless and lignified, up to 15 mm in diameter, greyish-brown, glabrous

with peeling bark; internodes 3-30 cm long. Leaves pendulous, petiolate; petiole

terete, 20-50 x c. 4-8 mm in diameter, green to brown, glabrous (sparsely pubescent

when young); lamina lanceolate, very succulent and fleshy, 10-30 x 5-12 cm, dark

green on adaxial surface, light green on abaxial surface, glabrous, apex acute, base

cordate; pinnate, midrib depressed on adaxial surface, secondary veins 4-7 each side,

secondary veins sometimes protruding, anastomosing in the middle between midrib

and leaf margin, branching from the midrib at 45-80°. Inflorescences one per node,

extra-axillary, positively geotropic, pseudo-umbelliform, convex, consisting of 2-10

flowers; peduncle terete, 2-7 cm x 2-5 mm, older peduncles forming a rachis from

previous flowerings. Flowers scented like Gardenia jasminoides , especially at night.

Corolla spreading, c. 7 cm when flattened, pastel pink with 2-3 cm wide pale cream

yellow centre inside, outside cream; tube c. 1.5 cm long, inside pubescent, outside

glabrous, lobes fleshy, triangular-acuminate, c. 3 x 2 cm, inside thickly pubescent,

denser along margin and on apices, outside glabrous, margin c. 5 mm reflexed to

revolute, apex acute. Corona staminal c. 1 cm high, 2 cm in diameter, fleshy, cream

coloured; lobes rhomboid, centrally ridged and laterally grooved, c. 1 x 0.7 cm, inner

processes cuspidate, meeting in centre, outer processes laterally compressed, c. 0.8 cm

lower than inner processes, basal margins spreading, touching only towards the apex

of the outer corona lobe. Style-head convex, hidden by anther appendages, which in

turn are partially covered by the overlapping inner coronal processes. Pollinia linear,

E9-2.1 mm x 450-550 pm, lacking sterile edge, corpusculum 550-650 x 400^150

pm, caudicles attached at the middle of corpusculum, 250-300 x c. 150 pm. Ovary

conical, c. 8 mm long, glabrous. Fruits and seeds not observed.

134

Card. Bull. Singapore 69(1) 2017

Fig. 19. Hoya stenakei Simonsson & Rodda. A. Flower, top view. B. Flower, side view. C.

Corona, top view. D. Corona, from underneath. E, F. Corona, side view. G. Pollinarium.

All from Simonsson Juhonewe, N., & Juhonewe, F. NS0026H. (Photos A-F: N. Simonsson

Juhonewe; G: M. Rodda)

Hoyci of Papuasia I.

135

Fig. 20. Hoya stenakei Simonsson & Rodda. A. The plant was found growing at the top of a

large tree. A climber managed to reach the top of the tree and collect some material. B. The

plant in cultivation. C. The large, shiny leaves. (Photos: N. Simonsson Juhonewe)

136

Card. Bull. Singapore 69(1) 2017

Distribution. Known only from the type locality in Sandaun Province, Baiberi.

Etymology. Named after the late Mr Sten-Ake Svensson (1947 - 2011) of Sweden,

whose children supported the first author’s work in PNG.

Habitat and ecology. The single known specimen was growing as an epiphyte 20

metres up in a very tall tree, nestling inside a fern, Asplenium sp., in a species-rich

primary forest with limestone karsts.

Provisional IUCN conservation assessment. Critically Endangered CR B2ab(ii,iii),

Cl, C2(ai), D (IUCN, 2016). Several botanical expeditions have been conducted in the

vicinity of the type locality, both by the first author and several other earlier botanists,

but Hoya stenakei was found only once. The two similar species Hoya archboldiana

C.Norman and Hoya onychoides P.I.Forst., Tiddle & I.M.Tiddle, on the other hand,

were encountered several times during the first author’s expeditions in PNG. The type

locality was to be logged soon after our expedition and it must now be assumed that

this locality is destroyed.

Notes. A new species with one of the largest flowers in the entire genus. It belongs

to the Hoya macgillivrayi group, up until now consisting of only three species, H.

macgillivrayi F.M.Bailey from Cape York in Queensland, Australia, and two from

Papua New Guinea, H. archboldiana and H. onychoides . The new taxon is the most

distinct of these species as their coronas are narrowly star-shaped, when viewed from

above, and the inside of their corolla lobes are glabrous, whereas Hoya stenakei has

an almost pentagonal corona and the inside of the corolla lobes is densely pubescent.

Key to species in the Hoya macgillivrayi group

(modified from Forster et al., 1995):

1 a. Inside of corolla lobes pubescent; corona almost pentagonal. H. stenakei

lb. Inside of corolla lobes glabrous; corona narrowly star-shaped.2

2a. Corolla campanulate, lobes shorter than the tube and <15 mm long, reflexed.

. H. archboldiana

2b. Corolla campanulate-rotate or rotate, lobes as long as or longer than the tube

and >15 mm long, semi-erect to incurved.3

3a. Corolla lobes strongly incurved giving a ‘claw-like’ appearance to the lobe, with

margins strongly reflexed creating a protrusion at the base of sinus between the

corolla lobes. H. onychoides

3b. Corolla lobes weakly incurved, with margins slightly reflexed with no protrusion

formed at the base of the sinus between the corolla lobes. Hoya macgillivrayi

Hoya of Papuasia I.

137

Hoy a urniflora (P.I.Forst) Simonsson & Rodda, comb. nov. - Marsdenia urniflora

RI.Forst., Austral. Syst. Bot. 8: 752 (1995). - TYPE: Papua New Guinea, Oro Province,

Lala River, c. 5500 ft. [c. 1675 m], 28 December 1935, Carr, C.E. 14065 (lectotype

SING [SING0122003], designated here; isolectotypes BM [BM001014154], K, L,

SING [SING122001, SING122002]). (Fig. 12, 21, 22).

Similar to Hoya juhoneweana because both have sunken leaf venation and a bullate

leaf surface. Hoya juhoneweana has corollas >2.5 cm wide whereas H. urniflora have

corollas c. 1.5 cm wide.

Epiphytic climber with white exudate latex in all vegetative parts. Stems cylindrical,

1.5-3 mm in diameter, sparsely to densely hispid; internodes 3-25 cm long. Leaves

petiolate; petiole terete, 4-8 x 1-2 mm in diameter, hirsute; lamina lanceolate, stiff and

chartaceous, (1.5—)3—10 x 1—2.5 cm, hirsute on the abaxial surface, adaxial surface very

hispid when young, almost glabrous on old leaves, apex long acuminate, base attenuate

to acute with single, ovoid basal colleters; venation pinnate, midrib and secondary

veins depressed on adaxial surface, raised on abaxial surface, secondary veins slightly

visible, 2-5 each side, branching from the midrib at 45-80°, anastomosing along the

lamina edge. Inflorescences one per node, extra-axillary, convex, consisting of (l-)5-

10 widely spaced flowers, peduncle perennial, terete, 0.4-2(-3) cm x 1.5-2 mm, older

peduncles forming a rachis from previous flowerings, hirsute; pedicels terete, 1.5-2

cm x c. 1 mm in diameter, hirsute. Calyx 3^4 mm in diameter, calyx lobes oblong,

1-2 x 1-1.5 mm, apex round, thinly pubescent outside; colleters one in each calyx

lobe sinus, c. 50 pm long. Corolla base narrowly campanulate, terminating in free

spreading lobes, 14-16 mm wide; tube 5-6 x 5-6 mm, sparsely pubescent inside, more

thickly pubescent at the throat, outside glabrous, lobes triangular, 5-9 x 3.5-4.5 mm,

velvety, outside glabrous, apex acute. Corona staminal 4.5-5.5 mm high, 4.5-5.5 mm

wide, fleshy; corona lobes erect, almost completely fused to the back of the anthers,

c. 4 x 1.5-2 mm, inner processes acute with a round tip, outer processes round, with

basal revolute margins. Style-head conical, hidden by anther appendages. Pollinia

oblong 400-500 x 150-200 pm, without sterile edge, corpusculum ovoid, 350—400 x

230-300 pm, caudicles attached at the base of the corpusculum, 130-170 x c. 50 pm.

Ovary conical, c. 2 mm long. Fruits and seeds not observed.

Distribution. First collected by Carr in 1935 ( Carr 14065 ) along the Kokoda Track

over the Owen Stanley Range along the border of Central and Oro Provinces, where he

also collected Hoya carrii. In 1970 a specimen ( Streimann & Kairo NGF42473) was

collected 260 km NW in southern Morobe province on the Central Range. The species

was collected again in 2011 at the easternmost end of the Finisterre Range on Huon

Peninsula, 290 km N of the first collection and 190 km NE of the second collection

(N. S. Juhonewe & F. Juhonewe NS0069L, NS0096L, NS0098L, NS0099L, NS0100L,

NS0101L, NS0102L). This suggests a widespread distribution, although the taxon may

still be rare and very scattered since it has only been encountered at one locality during

the first author’s expeditions.

138

Card. Bull. Singapore 69(1) 2017

Fig. 21. Hoya urniflora (P.I.Forst.) Simonsson & Rodda inflorescence A. Side view. B. From

underneath. All from Simonsson Juhonewe, N. & Juhonewe, F. NS0098L. (Photos: F. Juhonewe)

Hoyci of Papuasia I.

139

Fig. 22. Hoya urniflora (P.I.Forst.) Simonsson & Rodda. A, B. Corolla from above. C. Corolla

and calyx, from underneath. D. Corona from above. E, F. Corona, side view. G. Corona

from underneath. H. Calyx and ovaries. I. Pollinarium. A from Simonsson Juhonewe, N. &

Juhonewe, F. NS0098L, C-I from Simonsson Juhonewe, N. & Juhonewe, F. NS0102L. (Photos

A-H: F. Juhonewe; I: M. Rodda)

140

Card. Bull. Singapore 69(1) 2017

Habitat and ecology. It inhabits primary moist cloud forest at 1800-2000 m. It climbs

until it finds support on branches and then sends out stiff spreading branches which

flower several times a year. In cultivation it proved to be quite sensitive to both warm

and dry conditions and was rather difficult to propagate and transplant, with several

failed attempts. This is, therefore, a very sensitive species and is likely to suffer under

human pressure and global warming.

Provisional IUCN conservation assessment. Endangered B2a,b(iii) (IUCN, 2016).

Hoya urniflora is a locally rare but widespread species that is very susceptible to forest

degradation.

Additional specimens examined. PAPUA NEW GUINEA: Morobe Province: Menyamya

district, Spreader Divide, between Aseki and Menyamya, 6600 ft [c. 2010 m], epiphyte

climbing over small vegetation, 7°18'S 146°07'E, 21 Nov 1970, Streimann & Kairo NGF 42473

(LAE); Huon Peninsula, Hoboc, 1850 m, 24 Dec 2011, living accession NS 11-321, vouchered

at Ukarumpa on 31 May 2013, Simonsson Juhonewe, N. & Juhonewe, F. NS0069L (LAE,

SING); Huon Peninsula, Hoboc, 1900 m, 24 Dec 2011, living accession NS 11-327, vouchered

at Ukarumpa on 9 Mar 2014, Simonsson Juhonewe, N. & Juhonewe, F. NS0096L (LAE); Huon

Peninsula, Hoboc, 1850 m, 24 Dec 2011, living accession NS11-322, vouchered at Ukarumpa

on 28 Aug 2014, Simonsson Juhonewe, N. & Juhonewe, F. NS0098L (LAE); Huon Peninsula,

Hoboc, 1850 m, 24 Dec 2011, living accession NS11-323, vouchered at Ukarumpa on 9 Jun

2014, Simonsson Juhonewe, N. & Juhonewe, F. NS0099L (LAE); Huon Peninsula, Hoboc,

1900 m, 24 Dec 2011, living accession NS11-326, vouchered at Ukarumpa on 31 Dec 2014,

Simonsson Juhonewe, N. & Juhonewe, F. NS0100L (LAE); Huon Peninsula, Hoboc, 2013 m,

29 Dec 2011, living accession NS11-349, vouchered at Ukarumpa on 31 Jan 2016, Simonsson

Juhonewe, N. & Juhonewe, F. NS0101L (LAE); Huon Peninsula, Hoboc, 1885 m, 29 Dec 2011,

living accession NS 11-354, vouchered at Ukarumpa on 29 Sep 2014, Simonsson Juhonewe, N.

& Juhonewe, F. NS0102L (LAE).

Notes. Hoya urniflora was described as Marsdenia urniflora P.I.Forst. (Forster 1995:

752) with a holotype in SING. However, there are three duplicates of Carr 14065

in SING with none of them labelled as the holotype. Following ICN Art. 8.1 & 8.2

(McNeill et al., 2012) a holotype may only be a single specimen. Therefore, we select

a lectotype here, specimen SING0122003.

The taxon should be placed in Hoya because, despite lacking the sterile edge of the

pollinia typical of Hoya , it does have valvate bud aestivation, a character difficult or

impossible to observe on dry specimens with only open flowers but easily seen in fresh

material. Bud aestivation in Marsdenia R.Br. is instead imbricate. Pollinia without

a sterile edge in Hoya are not limited to H. urniflora but typical of species in Hoya

section Eriostemma and, in Papuasia, is also lacking in Hoya juhonewe ana.

Hoya urniflora is easily distinguished from the majority of Hoya species due to the

conspicuous sunken venation causing a characteristic bullate leaf surface, and by

its dense cover of >1 mm long hairs on all vegetative parts, especially when young.

Vegetatively it can only be confused with the new taxon Hoya juhoneweana Simonsson

& Rodda. However, Hoya juhoneweana has pendulous stems while H. urniflora

Hoya of Papuasia I.

141

develops climbing stems. When in flower, the two taxa are easily distinguished, as

Hoy a juhoneweana has corollas >2.5 cm wide whereas the corollas of H. urniflora are

c. 1.5 cm wide (Fig. 12).

Hoya versteegii Simonsson & Rodda, sp. nov.

Similar to numerous other Hoya species with elliptic and succulent leaves but differing

from all other Hoya species in the infundibuliform corolla with a continuously narrow

tube. - TYPE: Indonesia, Papua Province, Voer Gelukshauvel, s. dat., Versteeg, G.

1236 (holotype BO). (Fig. 23)

Climber , presence of latex unknown. Stems cylindrical, 1.5-4 mm in diameter,

sparsely pubescent; intemodes 4-15 cm long. Leaves petiolate; petiole terete,

channelled above, 10-15 x 3-5 mm in diameter, pubescent; lamina elliptic, fleshy, stiff

and coriaceous when dry, 8-15 x 3-6 cm, pubescent on the abaxial surface, adaxial

surface glabrous to sparsely pubescent, apex acuminate-apiculate, base round with no

colleters on old leaves; venation pinnate, midrib depressed on adaxial surface, raised

on adaxial surface, secondary veins almost invisible, 3-5 each side, branching from

the midrib at c. 45°. Inflorescences one per node, extra-axillary, convex, consisting of

3 or more flowers, peduncle perennial, terete, 5-10 cm x 1.5-2 mm, older peduncles

forming a rachis, glabrous or sparsely pubescent; pedicels terete, 2.5-3 cm x c. 0.8

mm in diameter, glabrous. Calyx 3-3.5 mm in diameter, calyx lobes oblong, 1-2 x

0.7-1 mm, apex round, thinly pubescent outside, basal colleters one in each calyx

lobe sinus, lanceolate, c. 50 pm long. Corolla infundibuliform, with free spreading

lobes, c. 2.5 cm in diameter when flattened; tube 12-15 mm x c. 3 mm . sparsely

pubescent inside, more densely pubescent at throat, lobes broadly ovate, 7-10 x 9-12,

apex acuminate, glabrous. Corona staminal, stipitate, stipe 1-1.5 mm high, c. 1.5

mm wide; corona c. 3 mm high, c. 3 mm in diameter, fleshy; corona lobes carinate,

almost completely fused to the back of the anthers, c. 3 x 1 mm, inner processes

acute with a round tip, outer processes round, with basal revolute margins. Style-head

hidden by anther appendages. Pollinia oblong, 500-600 x 280-320 pm with pellucid

margin, corpusculum rhomboid, c. 400 x 250 pm, caudicles attached at the base of

corpusculum, 100-150 x 30^40 pm. Ovary linear, c. 2.5 mm long, glabrous. Fruits

and seeds not observed.

Distribution. Recorded from a single locality in Papua Province, Indonesia, Voer

Gelukshauvel.

Etymology. Named after its collector, G. Versteeg (1876-1943).

Provisional IUCN conservation assessment. Data Deficient (IUCN, 2016). This

species is known only from the type specimen collected in the early 20th century.

142

Card. Bull. Singapore 69(1) 2017

Fig. 23. Hoya versteegii Simonsson & Rodda. A. Corolla sliced longitudinally showing the

position of the corona. B. Corona, side view. C. Corona, from underneath. D. Calyx and

ovaries. E. Pollinarium. All based on Versteeg, G. 1236. (Drawing: M. Rodda)

Notes. The flowers of Hoya versteegii , with their infundibuliform corolla and a long

narrow tube, may at first be confused with those of other genera, e.g. Marsdenia s.l.

The gynostegium is instead typical of Hoya as the corona lobes have revolute margins,

albeit much reduced, and the pollinia have a narrow sterile edge. Among other Hoya

species, only H. telosmoides Omlor from Borneo and H. papaschonii Rodda from

Thailand have a tubular corolla, but in both cases they are not continuously narrow as

in H. versteegii. Additionally the corolla tube of Hoya versteegii is 12-15 mm long

white that of H. telosmoides and H. papaschonii is less than 10 mm long. Both species

have much thinner leaves, chartaceous when dry, while Hoya versteegii has stiff and

coriaceous leaves.

Hoya yvesrocheri Simonsson & Rodda, sp. nov.

Similar to Hoya stenophylla Schltr. by having linear leaves but differs in the wider

leaves and flowers not developing simultaneously, as usually observed in most Hoya

species, but rather with only one or rarely two flowers open at a time. - TYPE: Papua

Hoya of Papuasia I.

143

New Guinea, Morobe Province, Wagau, 4200 ft [c. 1280 m], found in large tree in old

garden near village, 6°50'S 146°50'E, 11 June 1964, Millar, A.N. NGF 23466 (holotype

LAE; isotypes L [L2726759], CANB n.v.). (Fig. 24)

Liana slender, glabrous with white latex in vegetative parts. Leafy stems cylindrical,

1-2 mm in diam; older stems lignified, max diameter (up to 3-4 mm in diameter), colour

greenish brown, bark peeling; internodes 1-5(10) cm long. Leaves usually hanging,

petiolate; petiole terete, c. 2-5 x 1-1.5 mm in diameter, green without markings;

lamina linear or slightly lanceolate, coriaceous, carinate, c. 5-10 x 0.5-1 cm, green on

both surfaces, apex acute base rounded or attenuate, margin entire; venation pinnate

but barely evident in living material, midrib depressed on adaxial surface, secondary

veins 4-6 each side, secondary veins anastomosing near leaf margin, branching from

the midrib at c. 60-90°; leaf colleters 1 or 2, conical 0.3-0.4 mm long. Inflorescences

one per node, extra-axillary, positively geotropic, pseudo-umbellate, usually 1-8 buds

at different stages with only one or rarefy two flower open at a time; peduncle terete, c.

0.5-3 x 0.1-0.15 cm, older peduncles bearing scars from previous flowerings, pedicels

terete, 2 cm x 0.5-1 mm, green. Calyx c. 3 mm in diameter, lobes triangular-ovate, c. 0.5

x 0.5 mm, glabrous. Corolla campanulate, c. 1.5 cm diameter (c. 2 cm when flattened),

white with a slight pink or purple hue, intensifying with more sunlight; tube c. 7 mm,

lobes triangular, acuminate, c. 6 x 7 mm, margins entire, apex slightly revolute, densely

pubescent with unicellular hairs, abaxially glabrous. Corona staminal c. 2 mm high, 5

mm in diameter, fleshy, creamy white-yellow with dark reddish centre; corona lobes

elliptic-ovate, c. 2 x 1 mm, inner processes slightly erect, outer processes rounded and

erect, with basal revolute margins. Style-head hidden by anther appendages. Pollinia

oblong, c. 350 x 130-150 pm, with sterile edge, corpusculum 150-200 x 100-110 pm,

caudicles attached to middle of corpusculum, c. 70 x 30 pm. Ovary conical, c. 0.8 mm

long, glabrous. Fruits and seeds unknown.

Distribution. Recorded from Morobe Province only, at altitudes between 720 and

1260 m.

Etymology. Named after the Yves Rocher Foundation which supported the first

author’s Hoya Project for five years during 2011-2015.

Habitat and ecology. It seems to be confined to altitudes between 700 and 1300 metres

within a small area in Morobe province. The first author has only seen it once, on a

fallen tree on a very sharp ridge in primary rainforest in Bui a, Morobe Province. Upon

re-visiting the area in 2013 the ridge had been burned for expanding slash-and-burn

subsistence gardens.

Provisional IUCN conservation assessment. Endangered EN Blab(i,iii) (IUCN,

2016). Hoya yvesrocheri has only been found in Morobe Province between 700 and

1300 m, an altitude where the few remnants of forest are rapidly being converted into

subsistence gardens or abandoned grasslands due to the expanding population. Most

144

Card. Bull. Singapore 69(1) 2017

Fig. 24. Hoya yvesrocheri Simonsson & Rodda. A. Inflorescence with a single open flower

and leaves. B. Flower, top view. C. Corona, top view. D. Corona, from underneath. E, F.

Corona, side view. G. Calyx and ovaries. H. Pollinarium. All from Simonsson Juhonewe, N. &

Juhonewe, F. NS0047L. (Photo A: T. Sulemana. B-H: M. Rodda)

Hoya of Papuasia I.

145

remaining forest in that area is above 1500 m and thus in a different climate zone,

making it unlikely that this taxon thrives in those forests.

Additional specimens examined. PAPUA NEW GUINEA: Morobe Province: Bupu Village

above Gurakor, rainforest, 2400 ft [c. 730 m], 7°00'S 146°40'E, 27 Jul 1968, Millar, A.N. NGF

12172 (L [L2726757], LAE); Mumeng subdistrict, Bupu village, 914 m, mountain ridges,

Castanopsis forest, 7°00'S 146°35'E, 15 May 1974, Katik, P. & Larivita, G. LAE 62031 (BRI

n.v., CANB n.v., L [L2726758], LAE); Bula, 744 m, 07°00'59"S 146°56'44"E, 16 Feb 2008,

living accession NS08-030, vouchered at Ukarumpa on 29 Jun 2012 as Simonsson Juhonewe,

N. & Juhonewe, F. NS0047L (LAE Spirit in SING).

Notes. Similar to Hoya stenophylla Schltr. in the linear leaves, a character unique

to these two species in PNG. They can be easily distinguished by Hoya yvesrocheri

having an evenly green colour in all vegetative parts except for older stems, leaves

twice the width, and in its unique floral development where the flowers of a single

inflorescence are in different stages with only one or rarely two flowers open at a time.

In Hoya stenophylla all flowers develop simultaneously, the usual pattern for most

Hoya species.

ACKNOWLEDGEMENTS: This study is part of an on-going research project on the

systematics of Hoya. Nadhanielle Simonsson Juhonewe received support from the Yves

Rocher Foundation, Helge Ax:son Johnsons Stiftelse and many individual contributors.

Michele Rodda received support from the National Parks Board (Singapore), which sponsored

numerous herbarium trips to Asian and European herbaria, and from the SYNTHESYS Project

http://www.synthesys.info/ which is financed by the European Union under the FP7 Integrating

Activities Programme, grants GB-TAF-5657, NL-TAF-676 and DE-TAF-675. We would like to

thank the curators of the herbaria mentioned in the text for allowing access and/or for providing

high quality images of herbarium specimens. We thank Paul Forster and the late David Liddle

for their encouragement to work on Papuasian Hoya , Theresa Sulemana for the permission to

use her photograph of Hoya yvesrocheri, Stephanie Pimm Lyon for the permission to use her

photographs of Hoya juhoneweana ssp. juhoneweana in habitat, Foreting Juhonewe for the

permission to use numerous photographs, and two anonymous reviewers for their valuable

comments on the manuscript.

References

Aurigue, F.B., Sahagun, J.R. & Suarez, W.M. (2013). Hoya cutis-porcelana (Apocynaceae): A

New Species from Samar and Biliran Islands, Philippines. J. Nat. Stud. 12(1): 12-17.

Brown, G.K., Nelson, G. & Ladiges, PY. (2006). Historical biogeography of Rhododendron

section Vireya and the Malesian Archipelago. J. Biogeogr. 33: 1929-1944.

Forster, PI. (1995). Circumscription of Marsdenia (Asclepiadaceae: Marsdenieae), with a

revision of the genus in Australia and Papuasia. Austral. Syst. Bot. 8: 703-933.

Forster, PI. (1996). A checklist of the Asclepiadaceae of Papuasia. Sci. New Guinea 22(1):

15-22.

146

Card. Bull. Singapore 69(1) 2017

Forster, P.I. (2006). The Asclepiad Flora of New Guinea. In: Marshall, A.J. & Beehler, B.M.

(eds) The Ecology of Papua, pt. 1, pp. 371-378. Hong Kong: Periplus Editions

Forster, P.I. & Liddle, D.J. (1991). Variation in Hoy a australis Br. ex Traill (Asclepiadaceae).

Austrobaileya 3: 503—521.

Forster, P.I. & Liddle, D.J. (1992). Taxonomic studies on the genus Hoya R.Br. (Asclepiadaceae)

in Papuasia, 1-5. Austrobaileya 3: 627-641.

Forster, PI. & Liddle, D.J. (1993). Taxonomic studies on the genus Hoya R.Br. (Asclepiadaceae:

Marsdenieae) in Papuasia, 6. Austrobaileya 4: 51-55.

Forster, PI. & Liddle, D.J. (1996). Hoya. In: Orchard, A.E. & Wilson, A. (eds) Flora of

Australia , vol. 28. Canberra: Australian Biological Resources Study.

Forster, PI., Liddle, D.J. & Liddle, I.M. (1995). Taxonomic studies on the genus Hoya R.Br.

(Asclepiadaceae: Marsdenieae) in Papuasia, 7. Austrobaileya 4: 401—406.

Forster, PI., Liddle, D.J. & Liddle, I.M. (1997). Madangia inflata (Asclepiadaceae:

Marsdenieae), a new genus and species from Papua New Guinea. Austrobaileya 5:

53-57.

Forster, PI., Liddle, D.J. & Liddle, I.M. (1998). Diversity in the genus Hoya (Asclepiadaceae-

Marsdenieae). Aloe 35: 44^18.

Heads, M. (2006). Biogeography, ecology and tectonics in New Guinea. J. Biogeogr. 33: 957-

958.

Hemsley, W.A. (1891). New Solomon Islands Plants. Ann. Bot. (Oxford) 5: 501-508.

Hemsley, W.A. (1892). Decades Kewenses. Bull. Misc. Inform. Kew 65/66: 125-128.

Hemsley, W.A. (1894). Flora of the Solomon Islands. Bull. Misc. Inform. Kew 91: 211-215.

IPNI, The International Plant Names Index (2016). http://www.ipni.org. Accessed 20 Apr.

2016.

IUCN Standards and Petitions Subcommittee (2016). Guidelines for Using the IUCN Red

List Categories and Criteria. Version 12. http://www.iucnredlist.org/documents/

RedListGuidelines.pdf. Accessed 16 Feb. 2017.

Kloppenburg, R.D. (1991). Philippine Hoya Species. Fresno: ORCA Publishing

Lamb, A, & Rodda, M. (2016). A Guide to Hoy as of Borneo. Kota Kinabalu: Natural History

Publications (Borneo).

Lamb, A., Gavrus, A., Emoi, B. & Gokusing, L. (2014). The hoyas of Sabah, a commentary

with seven new species and a new subspecies. Sandakania 19: 1-89.

Li, P.T., Gilbert, M.G. & Stevens, W.D. (1995). Asclepiadaceae. In: Wu, Z.Y. & Raven, PH.

(eds) Flora of China, vol. 16, pp. 189-270. Beijing: Science Press; and St. Louis:

Missouri Botanical Garden Press.

McNeill, J., Buck, W.R., Demoulin, V., Greuter, W., Hawkworth, D.L., Herendeen, PS., Knapp,

S., Marhold, K., Prado, J., Prud’homme van Reine, W.F., Smith, G.F., Wiersema, J.H. &

Turland, N.J. (2012). International Code of Nomenclature for algae, fungi, and plants

(Melbourne Code). Regnum Vegetabile, vol. 154. Konigstein: Koeltz Scientific Books.

Moore, S. (1916). Asclepiadaceae. In: Ridley, H.N. Report on the Botany of the Wollaston

Expedition to Dutch New Guinea, 1912-1913. Trans. Linn. Soc. London, Bot. 9: 112-

116.

Prance, G.T., Beentje, H., Dransfield, J. & Johns, R. (2000). The Tropical Flora Remains

Undercollected. Ann. Missouri Bot. Gard. 87: 67-71.

Rahayu, S., Meve, U. & Rodda, M. (2015). Hoya undulata (Apocynaceae, Asclepiadoideae), a

new myrmecophytic species from Borneo, and typification of H. darwinii. Gard. Bull.

Singapore 67: 85-94.

Rintz, RE. (1978). The Peninsular Malaysian species of Hoya (Asclepiadaceae). Malayan Nat.

J. 30: 467-522.

Hoya of Papuasia I.

147

Rodda, M. (2015). Two new species of Hoya R.Br. (Apocynaceae, Asclepiadoideae) from

Borneo. PhytoKeys 53: 83-93.

Rodda, M. & Ercole, E. (2014). Hoya papaschonii (Apocynaceae: Asclepiadoideae), a new

species from southern Thailand with a peculiar corona. Phytotaxa 175(2): 97-106.

Rodda, M. & Nyhuus, T. (2009). Hoya danumensis, a new species of Hoya (Apocynaceae,

Asclepiadoideae) from Borneo. Webbia 64(2): 163-167.

Rodda, M. & Simonsson, N. (2011a). Hoya devogelii (Apocynaceae: Asclepiadoideae), a new

species from Sarawak, Borneo. Webbia 66(1): 33-38.

Rodda, M. & Simonsson, N. (2011b). Hoya medinillifolia (Apocynaceae: Asclepiadoideae), a

new species from lowland forest of Sarawak, Borneo. Webbia 66(2): 149-154.

Rodda, M. & Simonsson Juhonewe,N. (2013a). Hoya nuttiana (Apocynaceae: Asclepiadoideae),

a new species from Sarawak, Malaysian Borneo. Phytotaxa 140(1): 56-60.

Rodda, M. & Simonsson Juhonewe, N. (2013b). The taxonomy of Hoya micrantha and Hoya

revoluta (Apocynaceae, Asclepiadoideae). Webbia 68: 7-16

Rodda, M., Simonsson, N. & Wanntorp, L. (2011). Hoya wongii (Apocynaceae,

Asclepiadoideae): a new campanulate flowered species from Brunei (Borneo). Blumea

56: 205-208.

Rodda, M., Simonsson Juhonewe, N. & Ercole, E. (2014a). Hoya corymbosa (Apocynaceae:

Asclepiadoideae), a new unusual species from Sabah, Borneo and its systematic position

based on phylogenetic analyses. Syst. Bot. 38 (4): 1125-1131.

Rodda, M., Simonsson Juhonewe, N. & Rahayu, S. (2014b). Taxonomic revision of the Hoya

mindorensis complex (Apocynaceae:-Asclepiadoideae). Webbia 69(1): 39—47.

Rodda, M. Simonsson Juhonewe, N. & Lamb, A. (2015). Hoya fauziana (Apocynaceae:

Asclepiadoideae), a new species from Sabah, Malaysian Borneo. Webbia 70: 207-210.

Rodda, M., Lamb, A., Gokusing, L. & Bosuang, S. (2016). Hoya jiewhoeiana (Apocynaceae,

Asclepiadoideae), a new species from Sabah, Malaysian Borneo. In: Lamb, A. &

Rodda, M. A Guide to Hoy as of Borneo, pp. 198-200. Kota Kinabalu: Natural History

Publications (Borneo).

Schlechter, R. (1905). Hoya. In: Schumann, K.M. & Lauterbach, C.A.G. Nachtrage zur Flora

der deutschen Schutzgebiete in der Siidsee mit Ausschluss Samoa ‘s und der Karolinen,

pp. 362-367. Leipzig :Gebriider Borntraeger

Schlechter, R. (1908). Beitrage zur Kenntnis der Asclepiadaceen des Monsun-Gebietes. Bot.

Jahrb. Syst. 40, Beibl. 92: 1-19.

Schlechter, R. (1913). Die Asclepiadaceen von Deutsch-Neu-Guinea. Bot. Jahrb. Syst. 50:

81-164.

Shearman, P.L., Ash, J., Mackey, B., Bryan, J.E. & Lokes, B. (2009). Forest Conversion

and Degradation in Papua New Guinea 1972-2002. Biotropica 41: 379-390.

Van Welzen, PC. (1998). Increased speciation in New Guinea: tectonic causes? In: Dransfield,

J., Coode, M. J.E. & Simpson, D.A. (eds) Plant Diversity in Malesia III: Proceedings of

the Third International Flora Malesiana Symposium 1995, pp. 363-387. Kew: Royal

Botanic Gardens.

Voilering, J., Schuiteman, A., de Vogel, E., van Vugt, R. & Raes, N. (2016). Phytogeography

of New Guinean orchids: patterns of species richness and turnover. J. Biogeogr. 43:

204-214.

Wanntorp, L. & Forster, PI. (2007). Phylogenetic Relationships between Hoya and the

Monotypic Genera Madangia,Absolmsia, mdMicholitzia (Apocynaceae, Marsdenieae):

Insights from Flower Morphology. Ann. Missouri Bot. Gard. 94: 36-55.

Whitmore, T.C. (1984). A vegetation map of Malesia at scale 1:5 million. J. Biogeogr. 11:

461^471.

Gardens’ Bulletin Singapore 69(1): 149-155. 2017

doi: 10.3850/S2010098116000104

149

New combinations and typifications

in Aglaomorpha (Polypodiaceae)

S. Lindsay 12 , P.H. Hovenkamp 3 & D.J. Middleton 4

‘Gardens by the Bay, 18 Marina Gardens Drive, Singapore 018953

2 Present Address: Native Plant Centre, Horticulture and Community Gardening

Division, National Parks Board, 1 Cluny Road, Singapore 259569

stuart_lindsay @ nparks. go v. sg

3 Naturalis Biodiversity Center, PO Box 2317, 2300 RA Leiden, The Netherlands

4 Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

ABSTRACT. In preparation for a new checklist of Singaporean plants, the Flora of Singapore

project, and the Ferns of Thailand, Laos and Cambodia website, new combinations in

Aglaomorpha Schott are made to accommodate species formerly placed in Drynaria (Bory)

J.Sm. and Christiopteris Copel. Ten names are lectotypified, seven of which are second step

lectotypifications, and one name is neotypified.

Keywords. Christiopteris, Drynaria

Introduction

A close relationship between Aglaomorpha Schott and Drynaria (Bory) J.Sm. has long

been suspected and led to the establishment of the tribe Drynarieae Subh.Chandra

to accommodate these two genera (see Hennipman et al., 1990). On the other hand,

the relationships of Christiopteris Copel. were presumed to be with other genera of

Polypodiaceae (Hennipman et al., 1990) or were said to be uncertain (Hennipman et

al., 1998).

Recent phylogenetic research on these genera has shown that both Aglaomorpha

and Christiopteris are nested within a paraphyletic Drynaria (Janssen & Schneider,

2005; Schneider et al., 2008) which led Christenhusz & Schneider (2012) to propose

the conservation of Drynaria against Aglaomorpha. After two rounds of voting

no consensus could be reached as to whether to conserve or reject this proposal

(Applequist, 2014). This means that the normal rules of priority should apply and

that the name Aglaomorpha should be adopted for the genus sensu Into. Despite this,

Christenhusz (2015) made combinations in Drynaria for all species in the clade which

did not already have names in the genus.

The Angiosperm Phytogeny Group has been deliberating on relationships and

the classification of flowering plants since 1998 (APG, 1998; APG II, 2003; APG III,

2009; APG IV, 2016) but only recently has there been an equivalent for lycophytes

and ferns, the Pteridophyte Phylogeny Group (PPG I, 2016). This has set the standard

for family delimitation in lycophytes and ferns based on a large synthesis of relevant

150

Card. Bull. Singapore 69(1) 2017

literature and community consensus, the latter particularly when better data is pending.

In this work Drynaria and Christiopteris are included in synonymy of Aglaomorpha.

As names are needed for a number of existing and upcoming projects, such as a new

checklist of Singaporean plants, the Flora of Singapore project, and the Ferns of

Thailand, Laos and Cambodia website (Lindsay & Middleton, 2012 onwards), we

hereby make the necessary new combinations, including for those species not strictly

necessary for these projects.

The expanded Aglaomorpha has about 50 species (PPG I, 2016) in tropical

Africa, the islands of the Indian Ocean, tropical Asia and tropical Australia to Fiji.

New Combinations

Aglaomorpha baronii (Diels) Hovenkamp & S.Linds., comb. nov. - Drynaria baronii

Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 330, f. 171C (1899). - Polypodium

baronii Christ, Nuovo Giom. Bot. Ital., n.s., 4(1): 100, pi. 2 (1897), nom. illeg., non

Baker (1887). - TYPE: China, Shensi Septentrionali [= Northern Shaanxi], Dalle

falde fino circa la meta del Monte Zu-lu, 10-12 Aug. 1894, Giraldi s.n. (lectotype

FI [FI017966], first step designated by Roos (1986), second step designated here;

isolectotype K [K000959789]).

Distribution. China (Gansu, Qinghai, Shaanxi, Shanxi, Sichuan, E Xizang [Tibet],

NW Yunnan).

Aglaomorpha bonii (Christ) Hovenkamp & S.Linds., comb. nov. - Drynaria bonii

Christ, Notul. Syst. (Paris) 1(6): 186 (1910). - TYPE: Vietnam, Tonkin [Ha Nam

Province], 8 Jul. 1886, in montibus Ban Phet, Bon 3204 (lectotype P [P00266163],

first step designated by Roos (1986), second step designated here; isolectotype P

[P00274620]).

Distribution. India, China, Vietnam, Laos, Cambodia, Thailand, Peninsular Malaysia.

Aglaomorpha delavayi (Christ) Hovenkamp & S.Linds., comb. nov. - Drynaria

delavayi Christ, Bull. Soc. Bot. France: Mem. 1: 22 (1905). - TYPE: China, Yunnan,

Mt. de Kouang-yun-chan, Delavay 1166 (lectotype P [P00266136], first step designated

by Roos (1986), second step designated here; isolectotypes K [K000959792], P

[P00274616, P00274615]).

Distribution. Bhutan, Myanmar, China.

Aglaomorpha descensa (Copel.) Hovenkamp & S.Linds., comb. nov. - Drynaria

descensa Copel., Philipp. J. Sci., C. 3: 36 (1908). - TYPE: Philippines, Luzon,

New combinations in Aglaomorpha

151

Nueva Ecija, Copeland 2061 [also labelled Pteridophyta Philippinensia Exsiccatae

42] (lectotype MICH [MICH 1190384], first step designated by Roos (1986), second

step designated here; isolectotypes B (n.v.), BM (n.v.), G (n.v.), K [K000572169],

L [L.0537125], MU [MU000020234], NSW (n.v.), S [S09-41634; S-P-4751], UC

[UC350958]).

Distribution. Philippines (Luzon).

Aglaomorpha fortunei (Kunze ex Mett.) Hovenkamp & S.Linds., comb. nov. -

Polypodium fortunei Kunze ex Mett., Abh. Senckenberg. Naturf. Ges. 2(1): 121, t. 3,

f. 42-45 (1856). - Drynaria fortunei (Kunze ex Mett.) J.Sm. in Seemann, Bot. Voy.

Herald: 425 (1857), nom. illeg., non T.Moore (1855). - Drynaria roosii Nakaike, New

LI. Jap. Pterid.: 841 (1992). -TYPE: China, Fortune A 78 (lectotype K [K000959791],

first step designated by Roos (1986), second step designated here; isolectotype GOET

[GOET007408]).

Distribution. China, Taiwan, Laos, Vietnam (according to Roos, 1986).

Aglaomorpha involuta (Alderw.) Hovenkamp & S.Linds., comb. nov. - Drynaria

involuta Alderw., Bull. Dept. Agric. Indes Neerl. 21: 8, t. 4 (1908). - TYPE: Borneo,

Teijsmann 8577 (lectotype L [L.3532943], designated by Roos (1986)).

Distribution. Borneo.

Aglaomorpha laurentii (Christ ex De Wild. & Durand) Hovenkamp & S.Linds., comb,

nov. - Polypodium propinquum var. laurentii Christ ex De Wild. & Durand, Ann. Mus.

Congo Beige. Bot., ser. 2, 1: 70 (1899). - Drynaria laurentii (Christ ex De Wild. &

Durand) Hieron., Veg. Erde 2: 57, f. 54. (1908). - TYPE: D.R.Congo (Kinshasa),

lower Congo, Inkisi, Laurent s.n. (lectotype BR [BR0000008053800], designated by

Roos (1986); isolectotype P [P00266236]).

Distribution. Widespread in tropical Africa (Gabon, Uganda, Kenya, Tanzania,

Principe Isl., Sao Tome, Bioko Isl. (Fernando Po), Guinea, Sierra Leone, Liberia,

Ivory Coast, Ghana, Togo, Nigeria, Cameroon, Central African Republic, Equatorial

Guinea, D.R.Congo, Ethiopia, Rwanda, Congo, Angola).

Aglaomorpha mollis (Bedd.) Hovenkamp & S.Linds., comb. nov. - Drynaria mollis

Bedd., Ferns Brit. India 1: pi. 216 (1866). - TYPE: India, Nynee Tal, Dyas 6 (lectotype

K [K000959785], designated by Roos (1986)).

152

Card. Bull. Singapore 69(1) 2017

Distribution. Nepal, Bhutan, N India, China (Xizang [Tibet], NW Yunnan).

Aglaomorpha parishii (Bedd.) Hovenkamp & S.Linds., comb. nov. - Pleopeltis

parishii Bedd., Ferns Brit. India 1: pi. 125 (1866). - Drynaria parishii (Bedd.) Bedd.,

Suppl. Ferns S. Ind.: 24 (1876). -TYPE: Myanmar, ‘Moulmein’, Parishs.n. (lectotype

K [K000959778], designated here).

Distribution. Myanmar, Vietnam, Laos, Thailand.

Aglaomorphapleuridioides (Mett.) Hovenkamp & S.Linds., comb, no \.~Polypodium

pleuridioides Mett. in Miquel, Ann. Mus. Bot. Lugduno-Batavi 2: 229 (1866). -

Drynaria pleuridioides (Mett.) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 330

(1899). - TYPE: Java, Blume s.n. (lectotype L [L.3532882], designated here).

Distribution. Indonesia (Sumatra, Java, Lesser Sunda Islands, Sulawesi, Moluccas).

Aglaomorpha propinqua (Wall, ex Mett.) Hovenkamp & S.Linds., comb. nov. -

Polypodium propinquum Wall, ex Mett., Abh. Senckenberg. Naturf. Ges. 2(1): 120,

t. 3, f. 50 (1856). - Drynaria propinqua (Wall, ex Mett.) J.Sm. ex Bedd., Lerns

Brit. India 1: 160, pi. 160 (1866). - TYPE: Nepal, 1821, Wallich 293 (lectotype B

[B200060689], first step designated by Roos (1986), second step designated here;

isolectotypes B [B200060690, B200060691, B200060692], BR [BR0000005794416],

K-W [K001109730 (left specimen only), K001109731, K000959780], L [L.3532834,

L.3532835], NY [00127764], US [US00065584], W [W0061364]).

Note. There are several other specimens of Wallich 293 listed in the literature and

specimen databases but as this is a mixed collection we have only listed those we have

verified as isolectotypes.

Distribution. India, S China, Myanmar, Vietnam, Laos, Thailand.

Aglaomorpha quercifolia (L.) Hovenkamp & S.Linds., comb. nov. - Poly podium

quercifolium L., Sp. PI. 2: 1087 (1753). - Drynaria quercifolia (L.) J.Sm., J. Bot.

(Hooker) 3: 398 (1841). - TYPE: “Habitat in India”, Herb. Hermann 1: 39, No. 382

(lectotype BM [BM000621366], designated by Sledge, Bull. Brit. Mus. (Nat. Hist.),

Bot. 2: 144 (1960); not Herb. Linn (LINN, Microfche no. 1251.12 ) as given by Roos

(1986) and Hovenkamp (2013)).

Distribution. Sri Lanka and India to S China and throughout SE Asia and Malesia to

Liji and tropical Australia.

New combinations in Aglaomorpha

153

Aglaomorpha rigidula (Sw.) Hovenkamp & S.Linds., comb. nov. - Polypodium

rigidulum Sw., J. Bot. (Schrader) 1800(2): 26 (1801). - Goniophlebium rigidulum

(Sw.) T.Moore, Index Fil.: 396 (1862). - Drynaria rigidula (Sw.) Bedd., Ferns Brit.

India 3: pi. 314 (1869). - TYPE: Java, Thunberg s.n. (lectotype UPS [Cat. No. 24565],

designated here).

Note. Roos (1986) chose a Thunberg specimen from Java at S to be the lectotype (and

the same specimen was cited in Flora Malesiana (Hovenkamp & Roos, 1998)) but we

have been unable to trace this specimen.

Distribution. SE Asia to the Pacific islands and tropical Australia.

Aglaomorpha sagitta (Christ) Hovenkamp & S.Linds., comb. nov. - Polypodium

sagittum Christ, Bull. Herb. Boissier 6: 199 (1898) [as ‘sagitta’]. - Christiopteris

sagitta (Christ) Copel. in Perkins, Fragm. FI. Philipp.: 188 (1905). - Selliguea sagitta

(Christ) Fraser-Jenk., Taxon. Revis. Indian Subcontinental Pteridophytes: 45 (2008).

- Drynaria sagitta (Christ) Christenh., Phytotaxa 230(3): 299 (2015) [as ‘ sagittata’].

- TYPE: Philippines, Luzon, La Trinidad. Loher s.n. (lectotype P [P01545178],

designated by Hennipman & Hetterscheid (1984)).

Distribution. Philippines (Luzon, Mindanao).

Aglaomorpha sparsisora (Desv.) Hovenkamp & S.Linds., comb. nov. - Polypodium

sparsisorum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten

Naturk. 5: 315 (1811). -Drynaria sparsisora (Desv.) T.Moore, Index Fil.: 348 (1862).

- TYPE: Papua New Guinea, Central Province, Cloudy Bay, 16 Feb. 2007, Johns

11056 (neotype SING [SING0195508] designated here; isoneotypes BRI (n.v.), BRIT

(n.v.), K (n.v.), L (n.v.), LAE (n.v.), UPNG (n.v.)).

Note. In the protologue Desvaux (1811) gave the distribution of the species as ‘Habitat

in America?’, clearly expressing doubt as to where the species was from. The species

is only known in tropical Asia and Australasia, not in the Americas. We have been

unable to trace any original material.

Distribution. Thailand, Cambodia, Malesia and tropical Australia.

Aglaomorpha tricuspis (Hook.) Hovenkamp & S.Linds., comb. nov. - Acrostichum

tricuspe Hook., Sp. Fil. 5: 272, t. 304 (1864). - Gymnopteris tricuspis (Hook.) Bedd.,

Ferns Brit. India 1: pi. 53 (1865). - Cheiropleuria tricuspe (Hook.) J.Sm., Hist. Fil.:

139 (1875) [as “bicuspe”]. - Leptochilus tricuspis (Hook.) C.Chr., Index Filic., fasc.

7: 388 (1906). - Christiopteris tricuspis (Hook.) Christ, J. Bot. (Morot), ser. 2, 1: 273

154

Card. Bull. Singapore 69(1) 2017

(1908) [= J. Bot. (Morot) 21: 273 (1908)]. - Selliguea tricuspis (Hook.) Fraser-Jenk.,

Taxon. Revis. Indian Subcontinental Pteridophytes: 45 (2008). - Drynaria tricuspis

(Hook.) Christenh., Phytotaxa 230(3): 299 (2015). - TYPE: India, Sikkim, Atkinson

s.n. (lectotype K [K000959645], first step designated by Hennipman & Hetterscheid

(1984), second step designated here; isolectotype K [K000959644]).

Distribution. India, Thailand, Peninsular Malaysia, Vietnam, China (Hainan).

Aglaomorpha volkensii (Hieron.) Hovenkamp & S.Linds., comb. nov. - Drynaria

volkensii Hieron., [Veg. Erde 2: 57 (1908), nom. nud.] Bot. Jahrb. Syst. 46: 393 (1911).

- TYPE: Tanzania, Moshi District, Kilimanjaro, Marangu, Volkens 735 (lectotype B

[B200061011], designated by Tardieu-Blot (1964); isolectotypes E [E00429072], K

[K000435913]).

Distribution. Widespread in tropical Africa from Cameroon, Bioko, Congo (Kinshasa),

Rwanda, Burundi, Sudan, Ethiopia, Kenya, Somalia, Uganda, Zambia, Malawi,

Tanzania, Mozambique.

Aglaomorpha willdenowii (Bory) Hovenkamp & S.Linds., comb. nov. - Polypodium

willdenowii Bory, Ann. Sci. Nat. (Paris) 5: 468, pi. 13 (1825). - Drynaria willdenowii

(Bory) T.Moore, Index Fil.: lxxix (1857). - TYPE: Mauritius, Mt du Pouce, Bory St.

Vincent s.n. (lectotype P [P00266223], designated by Roos (1986)).

Distribution. Madagascar, Mascarenes (Mauritius, ?Reunion), Comores.

ACKNOWLEDGEMENTS. We thank Arne Anderberg (S), Mats Hjertson (UPS), Chiara Nepi

(FI) and Carmen Puglisi (SING) for their help to locate original material of some names, John

McNeill (E) for his advice on the typification of Drynaria baronii and the reviewers for their

helpful comments. We also thank the Biodiversity Heritage Library and JSTOR Global Plants

for making much of the literature and many of the type specimens available online.

References

APG (1998). An ordinal classification for the families of flowering plants. Ann. Missouri Bot.

Gard. 85(4): 531-553.

APG II (2003). An update of the Angiosperm Phylogeny Group classification for the orders and

families of flowering plants: APG II. Bot. J. Linn. Soc. 141(4): 399-436.

APG III (2009). An update of the Angiosperm Phylogeny Group classification for the orders

and families of flowering plants. APG III. Bot. J. Linn. Soc. 161(2): 105-121.

APG IV (2016). An update of the Angiosperm Phylogeny Group classification for the orders

and families of flowering plants: APG IV. Bot. J. Linn. Soc. 181(1): 1-20.

New combinations in Aglaomorpha

155

Applequist, W.L. (2014). Report of the Nomenclature Committee for Vascular Plants 66. Taxon

63(6): 1358-1371.

Christenhusz, M.J.M. (2015). New combinations in Drynaria (Polypodiaceae subfam.

Polypodioideae). Phytotaxa 230(3): 299-300.

Christenhusz, M.J.M. & Schneider, H. (2012). Proposal to conserve the name Drynaria against

Aglaomorpha (Polypodiaceae). Taxon 61(2): 465-466.

Desvaux, A.N. (1811). Observations sur quelques nouveaux genre de fougeres. Ges. Naturf.

Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 297-330.

Hennipman, E. & Hetterscheid, W.L. A. (1984). The emendation of the fem genus Christiopteris,

including the transference of two taxa to the microsorioid Polypodiaceae. Bot. Jahrb.

Syst. 105: 1-10.

Hennipman, E., Hovenkamp, PH. & Hetterscheid, W.L.A. (1998). Christiopteris. In: Kalkman,

C. & Nooteboom, H.P. (eds) Flora Malesiana, ser. 2, Ferns and Fern Allies, vol. 3, pp.

34-36.

Hennipman, E., Veldhoen, P. & Kramer, K.U. (1990). Polypodiaceae. In: Kramer, K.U. &

Green, P.S. (eds) The Families and Genera of Vascular Plants, Volume 1: Pteridophytes

and Gynmosperms. pp. 203-230. Springer-Verlag.

Hovenkamp, P.H. (2013). Drynaria. In: Parris, B.S., Kiew, R., Chung, R.C.K. & Saw, L.G.

(eds) Flora of Peninsular Malaysia, ser. 1, Ferns and Lycophytes, vol. 2, pp. 112-118.

Kepong: Forest Research Institute Malaysia.

Hovenkamp, P.H. & Roos, M.C. (1998). Drynaria. In: Kalkman, C. & Nooteboom, H.P. (eds)

Flora Malesiana, ser. 2, Ferns and Fern Allies, vol. 3, pp. 36-44.

Janssen, T. & Schneider, H. (2005). Exploring the evolution of humus collecting leaves in

drynarioid ferns (Polypodiacae, Polypodiidae). PI. Syst. Evol. 252(3-4): 175-197.

Lindsay, S. & Middleton, D.J. (2012 onwards). Ferns of Thailand, Laos and Cambodia, http://

rbg-web2.rbge.org.uk/thaiferns/Accessed 1 Mar. 2017.

PPG I (2016). A community-derived classification for extant lycophytes and ferns. J. Syst.

Evol. 54(6): 563-603.

Roos, M.C. (1986). Phylogenetic systematics of the Drynarioideae (Polypodiaceae). Verb.

Kon. Ned. Akad. Wetensch., Afd. Natuurk., Sect. 2, vol. 85. Amsterdam: Noord-Holland

Uitg. Mij.

Schneider, H., Kreier, H.P, Hovenkamp, P.H. & Janssen, T. (2008). Phylogenetic relationships

of the fern genus Christiopteris shed new light onto the classification and biogeography

of drynarioid ferns. Bot. J. Linn. Soc. 157: 645-656.

Tardieu-Blot, M.L. (1964). Pteridophytes. In: Aubreville, A. (ed) Flore du Cameroun 3. Paris:

Museum national d’histoire naturelle.

Instructions for contributing authors (continued)

Title and authorship. The title should concisely describe the contents. If a scientific name is used, its

authority is normally excluded, but the family name would be provided. Authors’ names, affiliations

and postal/ e-mail addresses are stated below the title. If more than one author, indicate “corresponding

author”. Avoid footnotes. A short running title (up to six words) should also be provided.

Abstract. The abstract should be less than 200 words. It should concisely indicate the article’s contents

without summarising it; mentioning novelties and name changes. Keywords. Suggest at most eight

keywords, in alphabetical order.

Scientific names and author abbreviations. Genus and species names of organisms must be italicised and

followed by the authority (with family name in parentheses) when first mentioned in the text or diagnoses.

Standards for author abbreviations should follow:

IPNI <http ://www. ipn i. org/ipni/authorsearchpage. do>

Authors of Fungal Names <http://www.indexfungorum.org/authorsoffungalnames.htm>

Herbarium abbreviations follow

Thiers, B. (2013). [continously updated] Index Herbariorum: A global directory of public herbaria and

associated staff. New York Botanical Garden's Virtual Herbarium, <http://sweetgum.nybg.org/ih/>

Journal and book title abbreviations. For journals:

Bridson, G.D.R., Townsend, S.T., Polen, E.A. & Smith, E.R. (eds) (2004). BPH-2: periodicals with

botanical content: comprising a second edition ofBotanico-Periodicum-Hunlianum. Pittsburg: Hunt

Institute for Botanical Documentation.

For books:

Stafleu, F.A. & Cowan, R.S. (eds) (1976-88). Taxonomic Literature. 2nd edition. 7 vols [Regnum Veg.

vols 94, 98, 105, 110, 112, 115, 116],

Stafleu, F.A. & Mennega, E.A. (eds) (1992-). Taxonomic Literature. Supplements [Regnum Veg. vols

125, 130, 132],

A useful source of verifying names of publications is

<http://asaweb.huh.harvard.edu:8080/databases/publication_index.html>. If in doubt, list full titles.

Other abbreviations and units of measurement. If using standard abbreviations and acronyms, give the

full term on first mention. Dates are cited as: 1 Jan 2000. SI (metric) units of measurement are used and

spelled out except when preceded by a numeral; they are abbreviated in standard form: g, ml, km, etc.

Coordinates should be given in the form 11°46 , 44.8"N 109 o 10'44.5"E.

Tables. Tables are numbered in arabic numerals in the order they are first mentioned in the text and carry

an indicative legend at the head. Tables are given at the end of the manuscript.

Illustrations. All drawings, maps, graphs and photographic images (individually or collected in a plate)

are to be numbered in arabic numerals in the order they are first mentioned in the text, as Fig. 1, Fig. 2,

etc. (plate components would be referred to in the text as Fig. 1A, IB, Fig. 1A-D, etc.). If relevant, scale

bars should be used to indicate magnification.

When grouping photographs, the maximum page area 19.5 x 13 cm must be heeded. High resolution digital

images may be submitted as separate files (line drawings in black and white at 600 dpi, photographs at

300 dpi) sent electronically or in a CD. Captions on figures should be added as a layer so that they can be

further edited or send both captioned and uncaptioned versions. Do not embed images into the main text

file and provide the legends for all figures at the end of the manuscript.

References in the text. Citation in the text should take the form: King & Gamble (1886) or (King &

Gamble, 1886), or King et al. (1886) if more than two authors to a work. Use 2000a, 2000b, etc. if several

papers by the same author(s) in one year are cited.

References listed at the end. There, works mentioned in the text are listed alphabetically as follows:

Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1999). User’s Guide to the DELTA Editor, http://

biodiversity.uno.edu/delta/ (accessed on 2 Aug. 2010).

Persson, C. (2000). Phylogeny of Gardenieae (Rubiaceae) based on chloroplast DNA sequences from the

rps 16 intron and trn L(UA A)-F(GAA) intergenic spacer. Nordic J. Bot. 20: 257-269.

Ridley, H.N. (1930). The Dispersal of Plants Throughout the World. Ashford, U.K.: E. Reeve.

Smith, A.C. & Darwin, S.P (1988). Rubiaceae. In: Smith, A.C. (ed) Flora Viliensis Nova, A New Flora

of Fiji A. 143-193.

References to web-based resources should include either a doi (digital object identifier) specification

or full URL mentioning also the date it was accessed. Use of DNA sequences from GenBank should be

acknowledged and the studies for which the sequences were generated should be cited.

Style of nomenclatural summaries. The following style is required:

Ornithoboea arachnoidea (Diels) Craib, Notes Roy. Bot. Gard. Edinburgh 11: 251 (1920); Burtt, Notes

Roy. Bot. Gard. Edinburgh 22: 294 (1958).

Ornithoboea parishii C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 148 (1883).

If authors include full bibliographic data for these works in the list of references at the end of the paper,

they should also be mentioned in the text briefly, e.g., “Nomenclatural references researched include

Blume (1849) and Roxburgh (1824).”

Homotypic synonyms should be provided in a block, stating the type at the end.

Front cover picture: Hoy a urnifiora (Photo by F. Juhonewe)