Review of Fiji Hibiscus (Malvaceae-Malvoideae) Species in Section Lilibiscus

Abstract

The taxonomy of Hibiscus species in section Lilibiscus in Fiji is
reappraised in the light of recent field and morphological studies. Six pre-
European contact species are recognized. These comprise four Fiji endemic
species, three of which are new and described: H. bennettii L.Thoms. and Braglia
from Mt. Delaikoro, Vanua Levu; H. bragliae L.Thoms. from Ovalau; and H.
macverryi L.Thoms. and Braglia from Macuata Province, Vanua Levu and
Taveuni. The fourth Fiji endemic, Hibiscus storckii Seem., was first collected on
Taveuni in 1860 and is here reinstated. Hibiscus cooperi Hort. Ex Lemaire, a
species generally associated with the South Pacific region, has been confused in
recent times with H. storckii. Hibiscus cooperi is lectotypified and reinstated.
Hibiscus cooperi and H. rosa-sinensis L. are both considered ancient, pre-European
introductions into the Pacific Islands. The Fiji archipelago is recognized as a
secondary center of diversity and speciation in sect. Lilibiscus in the South Pacific.
A botanical key is provided for the six Lilibiscus species that were present in the
South Pacific Islands prior to European contact. All four Fiji endemic Lilibiscus
species fall into the IUCN Critically Endangered (CR) Red List category and are
highly threatened in the wild from climate change and associated extreme events
including tropical cyclones, flooding, and landslides, as well as displacement by
environmentally invasive species and through hybridization with introduced
Hibiscus species. They are in urgent need of more intensive field survey followed
by complementary in situ and ex situ conservation actions.

Full text

Review of Fiji Hibiscus (Malvaceae-Malvoideae)
Species in Section Lilibiscus
1
Lex A. J. Thomson,
2,4
and Luca Braglia
3
Abstract: The taxonomy of Hibiscus species in section Lilibiscus in Fiji is
reappraised in the light of recent field and morphological studies. Six pre-
European contact species are recognized. These comprise four Fiji endemic
species, three of which are new and described: H. bennettii L.Thoms. and Braglia
from Mt. Delaikoro, Vanua Levu; H. bragliae L.Thoms. from Ovalau; and H.
macverryi L.Thoms. and Braglia from Macuata Province, Vanua Levu and
Taveuni. The fourth Fiji endemic, Hibiscus storckii Seem., was first collected on
Taveuni in 1860 and is here reinstated. Hibiscus cooperi Hort. Ex Lemaire, a
species generally associated with the South Pacific region, has been confused in
recent times with H. storckii. Hibiscus cooperi is lectotypified and reinstated.
Hibiscus cooperi and H. rosa-sinensis L. are both considered ancient, pre-European
introductions into the Pacific Islands. The Fiji archipelago is recognized as a
secondary center of diversity and speciation in sect. Lilibiscus in the South Pacific.
A botanical key is provided for the six Lilibiscus species that were present in the
South Pacific Islands prior to European contact. All four Fiji endemic Lilibiscus
species fall into the IUCN Critically Endangered (CR) Red List category and are
highly threatened in the wild from climate change and associated extreme events
including tropical cyclones, flooding, and landslides, as well as displacement by
environmentally invasive species and through hybridization with introduced
Hibiscus species. They are in urgent need of more intensive field survey followed
by complementary in situ and ex situ conservation actions.
LILIBISCUS (Hochreutiner 1900; lectotype H.
rosa-sinensis L., Van Borssum Waalkes 1966)is
a relatively small section in Hibiscus, nested
within section Bombicella (Pfeil and Crisp
2005). The progenitor of this group is
considered to have arisen within the past
one million years in Madagascar (Koopman
and Baum 2008). Section Lilibiscus comprises
about 23 species found mainly on midoceanic
volcanic islands in the Pacific and Indian
Oceans. Defining the features that set apart
species in sect. Lilibiscus is problematic: As
with other sections of Hibiscus, the currently
recognized characteristic features of these
section groups do not appear to represent
synapomorphies (Pfeil and Crisp 2005).
Updating the original description of the
section by Hochreutiner (1900), with infor-
mation from molecular and phylogenetic
studies of the Tribe Hibisceae (Malvaceae)
on Madagascar (Koopman and Baum 2008),
the recent review of the Hawaiian Lilibiscus
species (Huppman 2013), Van Borssum
Waalkes (1966),Cafferty and Cheek (1996),
Giraud (2013) and our work in Fiji, the sect.
Lilibiscus is described as follows:
Habit evergreen, woody shrubs to small,
multistemmed trees, usually 2–8 m tall.
Branches typically erect to spreading, semi-
pendulous in H. schizopetalus (Dyer) Hook.f.
and semi prostrate-ramified in H. fragilis DC.
Leaves±discolorous, glossy on upper surface,
glabrous or glabrescent, penninerved, most
frequently entire/unlobed, ±ovate/lanceolate,
1
Manuscript accepted 12 March 2018.
2
Faculty of Science, Health, Education and Engineer-
ing, University of the Sunshine Coast, Maroochydore,
Queensland, 4558, Australia.
3
Institute of Agricultural Biology and Biotechnology,
National Research Council (CNR), Milano, 20133, Italy.
4
Corresponding author
(e-mail: Lex.Thomson@gmail.com).
Pacific Science (2019), vol. 73, no. 1:79–121
doi:10.2984/73.1.5
©2019 by University of Hawai‘i Press.
All rights reserved.
79

less commonly elliptic or orbicular; hetero-
phyllous in several species, including deeply
divided,3(–5)-lobed,juvenile andintermediate
leavesin threeMascarene Islandspecies(viz. H.
boryanus DC., H. genevii Bojer ex Hook., H.
liliiflorus Cav.). Divided leaves often persisting
on mature plants of H. boryanus and H.
macverryi sp. nov, and some cultivars of H.
bennettii sp. nov.andH. bragliae sp. nov. Flowers
solitary, axillary, frequently showy with var-
iously coloured petals –white, cream, yellow,
orange, red or pink petals often with darker red
eye zone (and often multi-coloured in modern
hybrids); devoid of fragrance except for five
Hawaiian species, H. arnottianus A.Gray, H.
hannerae (O. Degener and I. Degener) Hupp-
man, H. immaculatus M. Roe, H. punaluuensis
(Skottsb.)O.Deg.and I.Deg.and H. waimeae A.
Heller. Peduncle articulated above middle.
Epicalyx 5–9 linear to narrowly triangular free
lobes, glabrous or glabrescent, tiny and scale-
like in H. schizopetalus and its hybrids; epicalyx
lobes shorter than calyx except in H. boryanus.
Calyx glabrous or glabrescent, united at base
into a tube, 5-lobed, ±non-inflated although
bulbous in H. kahilii C.N. Forbes. Corolla
medium to large, usually 8 to 20cm across,
somewhat asymmetrical, often lily-like or vase-
shaped. Petals 5 in single flower forms, or >35
petals in the double-flowered forms of H. rosa-
sinensis, (3.5–)5–10cm long, flat or recurved,
brightly coloured (red, pink, orange, yellow) or
white. Staminal column exserted and longer
than the petals, monadelphous, and antherifer-
ous in the upper/distal half. Fruits±obovate, 5-
sectioned papery capsule, glabrous or scabrous
but never woolly. Seeds irregular/blocky to
kidney-shaped, brown to black, hairy to sub-
glabrous. Ploidy variable chromosome number
with a probable base number of n=7or9;
diploid number of 42 (36 to 54), triploids (63),
tetraploids (72 to84), andnumerousaneuploids
and higher level polyploids (e.g. 144 and 168)
recordedespeciallyin modernhybrids.Ecology
and distribution found naturally on mid-
oceanic volcanic islands in the Pacific and
Indian Oceans, with one species in east Africa.
Species in sect. Lilibiscus have been the
source of taxonomic confusion as they readily
hybridize to produce fertile offspring. There
is also a relatively high frequency of genetic
mutation or sporting within Lilibiscus species
and their hybrids. There has also been
uncertainty concerning the native origins
and identity of H. rosa-sinensis, coupled with
the frequent misuse of this name for inter-
specific hybrids. Furthermore, other Lilibiscus
species/varieties, including from Fiji, were
brought into cultivation in Europe during the
1800s with incomplete or incorrect/specula-
tive information on their origins.
The sect. Lilibiscus is here considered to
include 23 species originating from six
geographic regions:
Kenya and Tanzania, east Africa: H. schizo-
petalus: the most morphologically distinctive
species with pendulous flowers, a very long
floral tube and laciniate petals; widely used in
early Hibiscus hybridization programs.
Madagascar: H. bernieri Baill., H. liliastrum
Hochr., and H. perrieri Hochr.
Mauritius, Reunion and Rodriguez Islands
(Mascarene Islands): H. boryanus,H. fragilis,
H. genevii and H. liliiflorus.
Hawaiian Island group in the northeastern
Pacific Ocean, comprising a clade of white-
flowered species (H. arnottianus,H. hannerae,
H. immaculatus,H. punaluuensis and H.
waimeae) and a clade of red-flowered species
(H. clayi O. Deg. and I. Deg., H. kahilii,H.
kokio Hillebr. ex Wawra and H. saintjohnianus
Roe).
Species known only in cultivation but
considered likely to have originated in
Indonesia/SE Asia: H. rosa-sinensis and H.
cooperi Hort. ex Lemaire.
Fiji: H. storckii Seem., H. bennettii sp. nov., H.
bragliae sp. nov. and H. macverryi sp. nov.
These four species are the main subject of
this paper.
Note: Hibiscus denisonii Burb. is also only
known in cultivation, and while sometimes
suggested as originating in the Pacific Islands,
this entity is morphologically quite different
to Fiji Lilibiscus species. Further genetic
research is needed to identify the relationship
ofH. denisonii to otherLilibiscusandwhetherit is
a species, or more likely a hybrid from the mid-
nineteenth century. An increase in leaf thick-
nessis often associated withpolyploidy (e.g. Ta l
1980), and the unusual leathery leaves of H.
denisonii are likely associated with its high
80 PACIFIC SCIENCE •January 2019

ploidy level, viz. 2n=84 (tetraploid) or 168
(octoploid) (Nakamura and Nomoto 1985).
About half the species in sect. Lilibiscus are
rare, including. H. clayi,H. hannerae,H.
immaculatus,H. saintjohnianus and other
Hawaiian Lilibiscus species (Huppman 2013);
H. fragilis (Cafferty and Cheek 1996), H.
liliiflorus (Stearns and Stearns 2000), H. genevii
–rediscovered in Le Mondrain, Mauritius in
1968 by Dr. L. Bernardi and Mr. J. Guého; H.
storckii (Seemann 1865), or else unknown in
the wild (H. rosa-sinensis and H. cooperi).
Lilibiscus species are usually better known in
cultivation due to their ornamental appeal,
cultural significance and medicinal qualities.
Although H. rosa-sinensis has yet to be
definitively located in nature, its name is often
misused for Lilibiscus interspecific hybrids. The
native origin of H. rosa-sinensis has been
variously suggested as China, South-east Asia,
South Asia, East Africa and Indian Ocean
Islands (Loureiro 1788,Hochreutiner 1900,
Bates 1965,Gast 1980,Florence 2004,Lawton
2014,Giraud 2013,Sampath 2014). The type
specimen of H. rosa-sinensis is a semidouble (or
double) flowered hibiscus from Sri Lanka
(Linnaeus 1753). Van Borssum Waalkes
(1966) notes that the protologue of H. rosa-
sinensis L. in Species Plantarum begins with a
citation of Flora Zeylanica and with a corre-
sponding specimen in the Hermann Herbar-
ium (Vol. III, fol. 4, Linn. n. 260) that he then
designated as the lectotype. Van Borssum
Waalkes (1966) also noted that there were
authentic specimens of H. rosa-sinensis in the
Linnean Herbarium: Linn 875.16 and as
recorded in the letter from C. Linnaeus of 18
August 1763 to G. H. Bradd, a senior official in
the Swedish East India Company (Jarvis 2007).
Hibiscus rosa-sinensis is considered an
ancient cultivated plant throughout the Poly-
nesian countries in the South Pacific Ocean
including Cook Islands (Figure 1), French
Polynesia (Figure 2), Samoa, Tonga, and
Polynesian outlier islands in Fiji, Vanuatu,
Solomon Islands and Papua New Guinea; it
has been present in the eastern Pacific Island
region for at least 700 years. This assertion is
based on the recorded presence of the species
in Tahiti in 1769 by Daniel Solander, botanist
on Captain Cook’s HMS Endeavour voyage,
along with an illustration of the single-flower
form of the species illustrated by Cook’s
botanical artist, Sydney Parkinson (Carr 1983)
and double-flower (flore pleno) form, and
coupled with the knowledge that Tahitians
are widely thought to have ceased long-
distance oceanic voyages around the four-
teenth or fifteenth century (e.g. Nunn 2007).
Hibiscus rosa-sinensis is a culturally impor-
tant and widely used medicinal plant in the
South Pacific region, including the Melanesian
countries and territories of Fiji (Weiner 1984,
Cambie and Ash 1984), New Caledonia
(Rageau 1973), Papua New Guinea (WHO
2009,Coiffier 2012), Solomon Islands, and
Vanuatu (Layard 1942), and in the Polynesian
countries and territories of Cook Islands
(Whistler 1990), Niue (Sykes 1970), French
Polynesia (Florence 2004), Samoa (Whistler
1992, 2000), and Tonga (Whistler 1991). In Fiji
H. rosa-sinensis and related species in sect.
Lilibiscusareknownlocallyby about 20different
names (Ta b l e 1 ), and indicative of their
importance for medicine, personal adornment,
house decoration, and as ornamentals and
hedge plants in village gardens and formarking
walking trails between villages.
With the exception of H. storckii Seem.,
other sect. Lilibiscus specimens collected in
Fiji, have generally been referred to as H. rosa-
sinensis by their collectors but have hitherto
never been subjected to rigorous botanical
investigation.
The objectives of this paper are to:
1. Review Hibiscus in sect. Lilibiscus in Fiji,
which hitherto have been neglected and
considered to be forms of H. rosa-sinensis.
2. Clarify the identity of H. storckii, which has
beenlosttosciencesinceitsdiscoveryin1860
on Taveuni, and which has become confused
with other species in the collections of
botanical gardens in the UK and in Europe.
3. Recognize and provide botanical descrip-
tions for the four Fiji endemic species in
sect. Lilibiscus, including diagnostic mor-
phological traits to be used in their
identification.
4. Draw attention to the critically endangered
status of Fiji’s endemic Hibiscus species, and
their further research and conservation
needs.
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 81

FIGURE 1. Hibiscus rosa-sinensis (Polynesian introduction; flore pleno), Pokata Marae (sacred place), Takitumu District,
Rarotonga, Cook Islands. (Photo: Randolph Thaman.).
82 PACIFIC SCIENCE •January 2019

METHODS
This study began through the chance dis-
covery of a wild species of Hibiscus on Mt
Delaikoro, Vanua Levu, Fiji in November
2015, which was originally thought to be H.
storckii, but which subsequent studies showed
to be a closely allied undescribed species. The
methods deployed in this study are described
in the following subsections.
Observations and Measurements on Fiji
Hibiscus
Field studies in the South Pacific Islands —
Between 2015 and 2017, fieldwork was
undertaken in the Fiji archipelago including
the five major islands of Viti Levu, Vanua
Levu, Kadavu, Taveuni, and Ovalau: this
involved collection, measurement, and photo-
graphy of Hibiscus specimens, both growing in
the wild and cultivated. Field studies on
Hibiscus were also undertaken in Solomon
Islands (Guadalcanal and Western Province),
Tonga (Tongatapu, ‘Eua, Vava‘u) and Vanuatu
(Efate) during the same period.
Botanical Specimens —Examination and
measurements of high-resolution images of
botanical specimens in sect. Lilibiscus from
major herbaria with Pacific collections, espe-
cially early botanical collections from Fiji.
The herbaria from which Hibiscus sect.
Lilibiscus specimens were examined, including
high-resolution images, were University of
South Pacific (USP, Fiji), Solomon Islands
National Herbarium (BSIP, Solomon Islands),
Papua New Guinea National Herbarium (LAE,
Papua New Guinea), Bishop Museum (BISH,
Hawai‘i, USA), National Tropical Botanical
FIGURE 2. Hibiscus rosa-sinensis (Polynesian introduction; single petals), Tahuata, Marquesas, French Polynesia
(cultivated). (Photo: Jean-François Butaud.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 83

Garden (PTBG, Hawai‘i, USA), Australian
National Herbarium (CANB, Australia), Auck-
land War Memorial Museum (AK, New
Zealand), Herbarium Bogoriense (BO, Indo-
nesia), Royal Botanic Gardens Kew (KEW,
UK), British Museum of Natural History (BM,
UK), Muséum National d’Histoire Naturelle
(P, France), Naturalis Biodiversity Center
(L, Netherlands), Conservatoire et Jardin
botaniques de la Ville de Genève
(G, Switzerland), New York Botanical Garden
(NY, USA), and Harvard University Herbaria
(GH, USA).
Determination and measurement of the
traits of key taxonomic significance in Fiji
species in sect. Lilibiscus included floral traits
(flower shape, size, and pigmentation, ped-
uncle/pedicel, petal, calyx, epicalyx, staminal
column, filaments, anthers, style, stigma) and
foliar traits (leaf shape, dimensions, petiole,
indumentum and leaf dentition pattern). In
Fiji Lilibiscus the shape of plant tissues and
associated ratios, such as the proportion of
pedicel above and below point of articulation
and leaf length:width, are generally more
informative than absolute measurements due
to the impact of environmental factors and
plant age/development on size of tissues—for
example, leaf size is extremely variable and
impacted by growing environment, including
developmental factors.
Observations on Hibiscus sect Lilibiscus
Under Cultivation —Propagation and culti-
vation of 12 species and >60 named hybrids
of Hibiscus sect. Lilibiscus including all those of
known and putative Fijian and Pacific origins
was undertaken in Suva, Fiji, and Melbourne,
TABLE 1
Local Names for Hibiscus (sect. Lilibiscus) in Fiji (Source: Paul Geraghty, USP, and local informants.)
Local Name Meaning Fiji Language Group Location
drigidrigi possibly “growing densely”West Fijian Ba (Viti levu), Vatulele
kauta borrowed from kaute, probably
(East) Futunan or Uvean Rotuman Rotuma
kaute, ’aute as Rotuman East Fijian Naselesele (Taveuni)
loloru unknown West Fijian Nadroga (Viti levu)
sakelu East Fijian Rakiraki (Viti levu)
seqelu East Fijian Somosomo (Taveuni): recorded by
Seemann but no longer in use
sekelu unknown East Fijian Macuata, Gone, and Dau Islands
(Vanua levu)
senikaute, seni’aute se “flower”ni “of”+kaute East Fijian Central and eastern Vanua levu
senilato perhaps “tusk flower”East Fijian Bua (Vanua levu)
senitoa “chicken flower”East Fijian Dawasamu, Wainibuka, Waidina,
Verata, Bau, Rewa, Suva, Namosi
(Viti levu), Beqa, Kadavu, Gau, Lau
senitoatoa variant of senitoa East Fijian Moce (Lau)
senitoo “chicken flower”West Fijian Waya, Vuda, Rakiraki, Wainibuka,
Waimaroiwai, Nadrau, Keiasi (Viti
levu), Mamanuca, Vatulele
sikelu, si’elu unknown East Fijian Seqaqa, Vanua levu interior
to “chicken”West Fijian Namosi
tokalau tradewind, “southeast wind”or
nineteenth-century name of
islands comprising Tokelau,
Tuvalu, and Kiribati
West Fijian Savatu, Nadrau, Navosa, Magodro
(Viti levu)
torilau perhaps related to “tokalau”West Fijian Serua (Viti levu)
84 PACIFIC SCIENCE •January 2019

Australia in order to observe and record
morphological differences when grown under
the same environmental conditions. Hibiscus
plants were propagated from seed, cuttings,
and grafts and grown for a period of 6–24
months in a freely draining potting media (a
mixture of compost, sand, perlite, coir fiber,
and dust) in both the open and in protected
environments. Plants were regularly fertilized
using a soluble fertilizer formulated to provide
an NPK ratio of 15:3:20, with added Mg, trace
elements, and chelated iron; plants were
treated as necessary for insect pests, mites,
and fungal pathogens using sprays containing
neem oil, petroleum oil, detergent, pyre-
thrum, piperonyl butoxide, tau-fluvalinate
and myclobutanil.
Assessment of Foliar and Floral Traits —In
total more than 3,000 individual measure-
ments of foliar and floral traits were under-
taken on H. bragliae (four wild plants, four
herbarium specimens, and seven cultivated
plants), H. bennettii (three wild plants, four
herbarium specimens, and 15 cultivated
plants), H. cooperi (nine herbarium specimens
and 18 cultivated plants) H. macverryi (one
wild plant, two herbarium specimens, and 14
cultivated plants) and H. storckii (six herbar-
ium specimens and 19 cultivated plants). The
main petal color was assessed using MS Paint
on images of a minimum of six flowers per
species/variant. Petal eye color was also
assessed but was similar between all of the
Fijian species: dark crimson red.
Consultations and Communication with
Hibiscus Experts —Wide consultation,
including discussions and correspondence, on
the subject of Fiji Lilibiscus species was under-
takenwithmore than 25Pacific,Australian,and
international botanists, ethnobotanists, lin-
guists, and Hibiscus researchers, growers, and
breeders (see acknowledgments). The main
aims of these consultations were to seek out
gray literature and undocumented field obser-
vations on sect. Lilibiscus in Fiji and other
PacificIslandnations,discuss researchprogress
and review findings, and develop a better
understanding of Fiji traditional knowledge,
local names, and meanings for Hibiscus in sect.
Lilibiscus.
Interviews were undertaken with key Fijian
informants living in proximity to native
Lilibiscus populations (on Taveuni and Ova-
lau), including seeking information on the
distribution, names, uses, and traditional
folklore associated with the Fiji species.
RESULTS
This study has identified three new endemic
Fijian species, viz. H. bennettii L.Thoms. and
Braglia, H. bragliae L.Thoms. and H. macver-
ryi L.Thoms. and Braglia, with the collection
locations for these species as shown in
Figure 3. This study supports the reinstate-
ment of the Fiji endemic H. storckii Seem. and
H. cooperi Hort. ex Lemaire, the latter coming
to international horticultural attention from
collections originating from the South Pacific
Islands in the mid-1800s. These five species
are morphologically distinct, and well-differ-
entiated from one another and the original
(unhybridized) forms of H. rosa-sinensis as
indicated in the botanical descriptions and key
provided in this paper. These five species are
now described and discussed in turn.
Hibiscus storckii Seem.
Hibiscus storckii was first collected near Somo-
somo on the island of Taveuni, Fiji in 1860
(Figures 4 and 5). The species was formally
describedbyGermanbotanist,Dr.Berthold
Seemann(Seemann1865), and named in honor
of his botanical assistant, Mr. Jacob Storck. The
specimens were collected during Seemann’s
ascent of DesVoeux Peak, on the30 May, 1860
(Seemann 1862). Hibiscus storckii has never been
recollected and therefore has continued to
elude botanists and hibiscus enthusiasts (e.g.
Gast 1980,Phillips 1990). Smith (1981),who
didnotobserve H. storckiiin thewild,treatedthe
species asconspecific with H.rosa-sinensis.Inthe
absence of any subsequent field collections,
Smith’s treatment has been widely accepted
until now.
Taveuni is a large shield volcano complex
(437km
2
) in which volcanism has occurred
over the past 780,000 years, with the most
recent activity about 350–550 years ago
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 85

(Whelan et al. 1985,Cronin 1999). The
topography of the main ridge is rugged, with
steep, dissected ranges and peaks, and over
150 distinctive scoria cones, craters, and
fissure zones (Cronin 1999). During 2016
and 2017, three visits were made to Taveuni by
the first author in search of H. storckii.No
surviving individuals of H. storckii were
located in the wild and, as indicated in its
original description, H. storckii appears to be
exceedingly rare, at least in more accessible
mountainous areas. Interviews with Fijian
landowners from the two villages in the area of
the original Seemann collections of H. storckii
revealed three reported sightings of wild
Hibiscus plants having large pink flowers with
a dark center:
1. One plant was observed by Mr. Pio Lele
(Somosomo Village) in 1997/1998 growing
about 60 m inside the Department of
Forestry’s“blue line”on the walking track
from Somosomo to Lake Tagimoucia (16°
480S, 179°560W). This Hibiscus was
destroyed during the establishment of a
dalo garden and Mr. Lele stated that Hibiscus
is no longer present in this location, at least
adjacent to the track.
2. One plant was observed by Mr. Solomoni
Tikoisuva (Tavuki Village) in 2014 near to
the “blue line”on the road from Tavuki to
Des Voeux Peak (16°490400 S, 179°5804500
W elevation 730 m). A thorough search in
April 2017 failed to relocate this individual.
3. Several plants were near the summit of Des
Voeux Peak (16°500300 S, 179°5705400 W
elevation 1,160 m). Mrs. Maria Botea
(Tavuki Village) observed these plants prior
to Tropical Cyclone Winston, but exten-
sive searching in March and June 2016
failed to relocate these plants, which are
presumed to have been destroyed by
Tropical Cyclone (TC) Winston.
FIGURE 3. Map of Fiji showing collection locations for the three new Fijian sect. Lilibiscus species.
86 PACIFIC SCIENCE •January 2019

FIGURE 4. Holotype specimen of Hibiscus storckii (Seemann 23). Kew Herbarium.
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 87

During the fieldwork on Taveuni a Hibiscus
that morphologically precisely matches with
Seemann’s botanical specimens and his type
description of H. storckii was observed grow-
ing in gardens in several villages (Figures 6
and 7). Taveuni informants indicated that this
Hibiscus matched the wild pink-flowered
Hibiscus they had observed at high elevation
on Taveuni, and accordingly this hibiscus is
hereby identified as H. storckii Seem. Hibiscus
storckii Seem. is here reinstated given that the
species differs considerably in its morphology
and DNA (Braglia, unpublished data) from H.
rosa-sinensis and other known South Pacific
species of sect. Lilibiscus. Following, we
provide a botanical description of H. storckii
based on our examination of the holotype,
isotypes and living cultivated specimens in Fiji
and Tonga.
Hibiscus storckii Seem., Flora Vitiensis [See-
mann] Vol. 1, p. 17 (March, 1865).
Holotype: Fiji, Taveuni, Somosomo (wild),
May 1860. B. Seemann 23 (K; K000659901).
Isotypes: BM (BM000645554), G (G00388550
and G00388551), GH (00052856), P
(P00646014).
FIGURE 5. Illustration of Hibiscus storckii by Walter Fitch in
Seemann (1873).
FIGURE 6. Hibiscus storckii. Matei, Taveuni, Fiji (cultivated) (Photo: Lex Thomson.).
88 PACIFIC SCIENCE •January 2019

Etymology: The specific epithet honors
Seemann’s botanical assistant, Mr. Jacob
Storck, who assisted with the collection of
the specimens on Taveuni.
Vernacular names: Segelu was recorded as its
local name by Seemann in 1860 but this name
appears to be no longer currently used on
Taveuni. There are similar Fijian names for
Hibiscus on the neighboring island of Vanua
Levu, viz. sekelu and sikelu in Macuata
Province (Paul Geraghty, University of South
Pacific, pers. comm.).
References: Hibiscus storckii Seem., Bon-
plandia 9: 254 (Seemann 1861), nom. nud.
Illustration by William Fitch, Plate 4 in
Flora Vitiensis: A description of the plants of
the Viti or Fiji Islands, with an account of
their history, uses and properties. Vol. 2.
Seemann (1873).
Description: Multistemmed woody shrubs
or small trees, 2–4 m tall, the upper stems light
gray. Branchlets terete, green, smooth. Stipules
filiform, 3–6(–9) mm, straight or gently
recurved, light green. Petioles green to red-
dish-green, ±swollen and lighter or reddish
for 6–7mm at each end, puberulent, (0.7–)
2–4(–5.5) cm long; petiole:leaf lamina length
ratio averaging between 0.26 and 0.32. Leaves
semi-glossy, simple, alternate, glabrous, dis-
colorous mid- to dark green above and light
green below, ±flat-undulate/inwardly cur-
ving, elliptic to lanceolate-elliptic, highly
variable in size 4–12cm long 1.7–6.5cm
wide, leaf base obtuse-rounded, length:width
ratio between (2–) 2.4 and 2.8 (–3). Leaf
margin entire or sparsely and irregularly
dentate with an average of two (up to 6–7)
teeth on the distal 40%–60% of each leaf
margin; leaf dentition pattern diagnostic with
the long edge of the tooth following the leaf
margin and the short base of each tooth ±
perpendicular to the mid-rib. Intermediate and
regrowth leaves on strongly growing new
shoots, up to 14.5cm long 6.5cm wide,
broadly lanceolate or infrequently tri-lobed,
acuminate, length:width ratio c. 2.5 and ±
uniformly serrated on distal portion of leaf
blade, (4–)7–8(–9) teeth on each side; petiole:
intermediate leaf lamina length ratio aver-
aging between 0.35 and 0.43. Venation
palmate, typically with three main veins;
midrib raised on lower surface. Flowers
solitary in the upper leaf axils, c. 14.5–15cm
diameter, borne on slender, puberulent,
FIGURE 7. Hibiscus storckii close-up of flower showing characteristic dark-red eye zone and red stigma pads –Matei,
Taveuni, Fiji (cultivated) (Photo: Lex Thomson.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 89

articulated pedicels, (3–)4–6(–7) cm long,
with articulation point 7–13mm from base of
flower (or av. 20% of total length above joint).
Pedicels grading into whitish cupulate recep-
tacle. Epicalyx segments 7–10, green, glabrous,
linear ±1–1.5mm wide, straight to spreading,
of variable length (3–)10–20 (–22) mm with
an acuminate tip. Calyx light green, pub-
erulent, forming a slightly swollen or campa-
nulate tube c. 20–28mm long, with five acute-
isosceles triangular lobes (7–13mm long and
5mm wide at base). Petals 5, united with floral
tube for lower 10–12mm; obovate, (6.5–)7–9
(–10) cm long4.5–7.5 cm wide; ±recurved,
ruffled and slightly overlapping, light lavender
pink (av. RGB 245, 150, 207, Hex Code
F596CF) with a distinctive large, dark
crimson serrated eye zone with radius of
2.8–3.5cm; the crimson points of inner eye
running into fine whitish veins. Outer surface
of petals light lavender pink with whitish
flecks. Staminal column dark crimson near
base, then pink grading into white in the
apical portion, (8–)10–13 (–14) cm long.
Stamens numerous (65–)70–85 (–95), thin,
pink filaments c. 5–8mm long with light
yellow kidney-shaped pollen sacs on the upper
one-third of the staminal column. Style white
±4mm, style branches 5, red, ±9 mm, spreading
to semi-erect. Stigma pads ±globular-capitate,
red, villose. Fruits and seeds not seen.
Natural distribution, habitat and associates:
Hibiscus storckii is a rare component of upland
rainforest (above 700m) and cloud forest on
Taveuni, Cakaudrove Province, Fiji. The
extent of cloud forest in Fiji is extremely
limited, occupying only the ridges on the
highest peaks. At Des Voeux Peak the
associated cloud forest species of H. storckii
include Astronidium sp., Blechnum milnei,
Cyathea alata,Cyathea medularis,Cyrtandra
tempestii,Ficus vitiensis,Freycinetia sp., Macar-
anga seemannii,Medinilla waterhousei,Pleran-
dra sp., Scaevola floribunda,Smilax vitiensis,
Spiraeanthemum serratum and Weinmannia
richii. The Des Voeux Peak cloud forest is
frequently subject to low light levels,
typically 20%–40% daylight under the
almost permanent cloudy conditions. The
annual rainfall is recorded as 997cm, with
high humidity (>90%) and mean daily
maximum temperatures around 17–20°C
(Ash 1987).
Genetic relationships and breeding system:
Tubulin-based polymorphism (TBP) analysis
suggests that H. storckii’s closest relative is the
new Fijian species H. bennettii (L. Braglia,
unpub. data).
Early horticultural references: No early
horticultural references have been located
for H. storckii, and a review of Kew Gardens
(UK) records indicates that living material of
the species was not sent to them by either
Berthold Seemann or Jacob Storck.
Horticultural varieties and importance: Exam-
ination of about 40 specimens of H. storckii,
growing in gardens on Taveuni (Caukadrove
Province), Suva, Tailevu, and Serua Provinces
VitiLevu andTongatapu(Tonga)did notreveal
any horticultural varieties.
Conservation status:H. storckii is here
assessed as critically endangered (CR B2 ab;
IUCN 2016) in the wild. This assessment is
based on its highly restricted natural geo-
graphic range, fragmented on mountain peaks
(above 700 m) on Taveuni, with no surviving
individuals known in the wild. Fijian infor-
mants have reported that H. storckii occurs at
an extremely low frequency in low, open
rainforest above 700 m and in high elevation
sites at 1,000–1,150 m on Taveuni—on
precipitous rocky cliffs. These habitats are
threatened by climate change, as small
increases in temperature (1–2°C) will lead
to shrinkage of the more open upland rain-
forest and cloud forest associations. The H.
storckii population on Des Voeux Peak is
highly threatened and plants have not been
observed since Category 5 TC Winston
struck in February 2016. The planting of a
native reed known as gasau (Miscanthus
floridulus (Labill.) Warb. ex K. Schum. and
Lauterb., Poaceae) to control soil erosion
around the Telecom Tower on Des Voeux
Peak is another threatening factor. This
densely clustering reed is likely to choke
out and prevent natural seedling regeneration
of H. storckii.
There is an urgent need for more com-
prehensive field surveys to ascertain the
species’complete native distribution and
abundance in open forest habitats above 700
90 PACIFIC SCIENCE •January 2019

m on Taveuni: These surveys ought to focus
on cloud forest communities on mountain
peaks above 800–850 m that occur along
approximately 10km of the central ridge
(Ash 1987,Cronin 1999), and including Des
Voeux Peak (1,194 m), Mt. Koroturaga (864
m), Mt. Manuka (842m), and Mt. Uluiqalau
(1,241 m).
Translocation of H. storckii to other secure
conservation sites (ex situ conservation in
botanical gardens and long-term seed storage)
is an essential complementary conservation
measure. Hibiscus storckii is occasionally culti-
vated in home gardens in lowland Fiji, Tonga
and Samoa. However, its long-term survival is
far from secure in such settings given the
fickle nature of the ornamental plant sector, its
reduced ability to set seed at low elevation and
possible hybridization with introduced, exotic
Hibiscus species for any seed set.
Hibiscus bennettii L. Thoms. and Braglia sp.
nov.
This new Hibiscus species was discovered
growingnear thepeak of Mt.Delaikoro, Vanua
Levu on 5 November 2015, during which
collections were made from two morphologi-
cally similar Hibiscus plants (Figure 8). These
Hibiscus plants, which appeared to be extremely
old, were growing on a steep rocky slope at an
elevationof930 mnear the Telecomtower.The
site is remote from villages and there was no
evidence of gardening or of them having been
planted. Subsequent examination revealed that
these specimens matched no other currently
circumscribedspecies ofHibiscus.H. bennettii sp.
nov. closelyresemblesand is relatedto H.storckii
(based on its TBP profile, Braglia unpub.) but
exhibits consistent differences in floral and
foliar traits as indicated in the botanical
FIGURE 8. Hibiscus bennettii—Mt. Delaikoro, Vanua Levu, Fiji (Photo: Sonu Dutt, Fiji Department of Forestry.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 91

description and key. It differs from H. rosa-
sinensis in its broadlanceolate leaves witheither
entire margins and/or with irregular dentitions
in the apical portion of the leaf blade, and by
their lighter-colored petals with white veins
and trim and a large lobate dark red eye-zone,
and light yellow stigma pads (Figure 9) and
epicalyx segments (Figure 10).
Hibiscus bennettii L. Thoms. and Bragliasp. nov.
Holotype: Fiji, Vanua Levu, Mt. Delaikoro
(wild). Alivereti Naikatini ANX 121, ANX 122,
27 Jul 2016 (SUVA).
Etymology: The specific epithet honors
John Edwin Bennett (born 21 January 1949)
in recognition of his assistance with field work
and research contributions on Fiji’s native
Hibiscus species during 2016 and 2017.
Vernacular name: Mt. Delaikoro hibiscus;
sikelu, sekelu—Fijian names for Hibiscus (sect.
Lilibiscus) in Macuata Province.
References: See later under genetic relation-
ships.
Description: Multistemmed woody shrubs or
small trees, 2–4 m tall, the upper stems light
gray. Branchlets terete, green, smooth, or
sparsely minutely hairy. Stipules filiform, 3–6
(–9) mm, straight or gently recurved, light
green. Petioles green, soon turning reddish-
green,(0.4–)0.6–1.5(–2.4)cm long,puberulent
in central channel on upper side of petiole (very
short stellate hairs mostly < 0.02mm long),
petiole:leaf lamina length ratio averaging
between0.14 and 0.19. Leaves simple,alternate,
glabrous, discolorous mid- to dark green, dull
to semiglossy above and light green below, ±
FIGURE 9. Hibiscus bennettii flower (typical pink petal form)—Rukuruku, Ovalau, Fiji (cultivated) (Photo: Nicolas
Perrault.).
92 PACIFIC SCIENCE •January 2019

FIGURE 10. Hibiscus bennettii flower—showing epicalyx and calyx region. Kuala Lumpur, Malaysia (cultivated) (Photo:
Alex Quek.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 93

flat-undulate/inwardly curving, highly variable
in size mostly 3.5–10cm long 2.0–4.5cm
wide, broad lanceolate, leaf base obtuse-
rounded, length:width ratio between (1.7–)
2.1 and 2.4 (–3.0). Leaf margin irregularly
serrate on the upper portion with 2–3(–7) teeth
per margin or infrequently entire; the inner
edge of the leaf teeth is frequently at ±45°
angle to the midrib (or sometimes up to 90°).
Regrowth leaves on strongly growing new shoots
larger, up to 15cm long7.5 cm wide, broad
lanceolate or tri-lobed, irregularly serrated-
dentate, typically 4–7 teeth per margin (on
distal 0.6–0.75 of leaf). Juvenile and intermediate
leaves 2, 3–5-lobed, mostly 7–12cm long 4–8
cm wide, length:width ratio between 1.4
and 1.6, teeth per margin (1–)2–3(–7).
Venation palmate, typically with three main
veins; midrib raised on lower surface. Flowers
solitary in the upper leaf axils borne on slender,
puberulent, articulated pedicels, (4–)5–7(–9)
cm long, with articulation point 5–11 mm from
base of flower (or av. 13% of total length above
joint for type; av. 21% for juvenile-intermedi-
ate leaf variant). Pedicels grade into whitish
cupulate receptacles. Epicalyx segments 7–10,
green, sparsely hairy, linear ±1.5–2mm wide,
straight to spreading, of variable length (6–)
10–14 (–15) mm with an acuminate tip. Calyx
light green, forming a slightly swollen or
campanulate tube c. 17–30mm long, with five
acute-narrow triangular lobes (12–18 mm long
and c. 5mm wide at base). Petals 5, united with
floral tube for lower 10–1 2mm, obovate,
(5.5–)6–7(–8.5) cm long4–6.5cm wide; ±
recurved, ruffled, and slightly overlapping;
rose pink (Thulian pink, av. RGB 245, 150,
207, Hex Code F596CF) with white veins and
trim, and lobate, dark red basal zone (±15–25
mm long). Outer surface of petals light pink
with whitish raised veins and flecks. Staminal
column dark crimson near base, then pink
grading into white in the apical portion, (7–)
8–10 (–12) cm long. Stamens numerous (60–)
70–80 (–90), thin, white filaments c. 5–7mm
long with yellow kidney-shaped pollen sacs and
grains positioned along the upper one-third of
staminal column. Style cream, branches 5,
spreading to semi-erect. Stigma pads ±glob-
ular-capitate, light yellow, villose. Fruits and
seeds not seen.
The new species may be confused with H.
storckii, especially when only sterile specimens
are available, but is morphologically differ-
entiated as follows:
Petals –H. bennettii: Thulian pink with
white veins and trim, and lobate, dark crimson
basal zone (1.9–2.8cm long). White and pale
pink petaled forms are known in cultivation.
H. storckii: lighter lavender pink with a large,
dark crimson red serrated eye zone of radius
2.8–3.6cm; the crimson points of the inner
eye running into fine whitish veins. Staminal
column –H. bennettii: dark crimson near base,
then pink grading into white in the apical
portion (pink-petalled form). H. storckii: dark
crimson to rose pink near base, then pink
throughout. Stamens –H. bennettii: white. H.
storckii: light lavender pink. Stigma pads –H.
bennettii: light yellow. H. storckii: red. Pedicel:
flower stalk ratio—i.e. the proportion of the
flower stalk above the joint or articulation—is
considerably less in H. bennettii (type), av.
0.13, than in H. storckii, av. 0.21.The key
differences in the foliage of pressed dried
specimens of the two species are as follows:
Adult leaf shape—mainly broad lanceolate in
H. bennettii with a leaf length-width ratio of ±
2.3 cf. mainly elliptic in H. storckii with leaf
length-width ratio of ±2.6. Leaf dentition –
the inner edge of the leaf teeth is frequently at
±45°(to 90°) angle to the midrib in H.
bennettii cf. H. storckii in which the base of the
leaf teeth is frequently ±perpendicular (to
45°) to the midrib. Petiole: leaf lamina length
ratio –less in H.bennettii (averaging between
0.14 and 0.19) than H. storckii (averaging
between 0.30 and 0.32).
Hibiscus bennettii and H. storckii are readily
distinguished from H. macverryi by their adult
leaves that are unlobed (except in juvenile
foliage variant of H. bennettii), whereas in H.
macverryi the adult leaves are heterophyllous
including some deeply three-lobed (divided
almost to the leaf stalk and resembling those
of red maple Acer rubrum). The petals of H.
bennettii and H. storckii are often larger and
pale pinkish purple/lavender pink in color,
whereas in H. macverryi they are a rich deep
rose. Hibiscus storckii and H. bennettii differ
from H. cooperi in the absence of a white halo
around the darker eye of its petals and in their
94 PACIFIC SCIENCE •January 2019

more elliptic leaf shape and dentition, includ-
ing some larger teeth or lobes, most evident in
intermediate foliage. Hibiscus bennettii and H.
storckii differ from H. rosa-sinensis in their
elliptic to broad lanceolate leaves with either
entire margins and/or with irregular denti-
tions in the apical portion of the leaf blade,
and by their lighter-colored petals with a
distinctive, large serrate or lobate dark
crimson red eye-zone.
Other specimens examined: Fiji, Vanua Levu,
Mt. Delaikoro. L. A. J. Thomson LAJT 1,5
November 2015, University of South Pacific
—South Pacific Regional Herbarium, Fiji
(SUVA); Fiji, Kadavu, Western end of island
near Cape Washington (cult.), AC Smith 323,
28 Oct 1933 (BISH,P).
Early horticultural references: No early (i.e.
pre-1900) horticultural references were
located that may be referrable to H. bennettii.
Natural distribution, habitat and associates:
Known only from four plants, mainly on steep
slopes (west to northwest aspect) near the
summit of Mt. Delaikoro, Vanua Levu. The
habitat is low open cloud forest with Syzygium
spp. and Cyathea spp being the major
associated species. Other cloud forest species
on Mt Delaikoro include Agathis macrophylla,
Clinostigma exorrhizum, Elaeocarpus cheloni-
morphus, Litsea spp., Saurauia rubicunda,
Metrosideros collina,Neuburgia collina, Pandanus
vitiensis, Physokentia thurstonii, Polyscias corti-
cata,P. joskei and Trimmenia weinmanniifolia.
Genetic relationships and breeding system:
Morphological and genetic studies (L. Brag-
lia, unpublished) have shown H. bennettii to be
most closely related to H. storckii, and indeed
sterile specimens of the species may be readily
confused. In 1935, R. R. Anson of the
Colonial Cotton Growers Corporation (Fiji)
was reported to have provided seeds of H.
storckii to the Danish cytologist Mr.. A.
Skovsted in Trinidad (Phillips 1990), but it
is almost certain that Anson provided
Skovsted with seeds of H. bennettii (rather
than H. storckii). Skovsted (1941) reported
2n=84 with a base number of 7. In a letter to
Mr. Keith Woolliams at Waimea Arboretum,
Hawai‘i, Ross Gast wrote “In 1965, I located
Skovsted in Copenhagen where, in retire-
ment, he was curator of the Botanical
Museum at the University of Copenhagen.
We exchanged a number of letters, and I
learned that when he was working for the
Colonial Cotton Company in Trinidad, he
received seed of H. storckii from another
geneticist working for the company in Fiji.
From this seed he grew plants and made his
chromosome studies. As the species made an
excellent garden plant, it became quite
popular in the West Indies, and later in
Florida under the name of David May. He
described the plants as very free blooming,
large flat white, yellow style, small with good
heavy foliage—very showy—with a pink flush
which fades to white as soon as it opens.”The
International Hibiscus Society database
(http://www.internationalhibiscussociety.org)
description of ‘David May’is: “bloom type:
single, regular; size: 18–20cm; bloom color:
yellow white, slight pastel pinkish, fades later
in day to white, straw style, lemon yellow
stigmas; bloom characteristics: medium tex-
ture, slightly cupped, overlapped; leaf char-
acteristics: medium green, and bush
characteristics: good growth habits.”Both
descriptions of ‘David May’closely corre-
spond with the lighter-colored petal variant of
H. bennettii.
Horticultural varieties and importance: During
fieldwork for this study, several variants of H.
bennettii were found in cultivation. These
included white and pale pink-petaled variants.
Thefoliage ofboth variantscloselymatches the
type form while the petals are either white
(flushed very pale pink during the cool season),
or pale-pastel pink. The staminal column is
dark crimson near its base, then white through-
out. These two variants are in cultivation in
Fiji (Vuna, Taveuni; Lautoka and Waila, Viti
Levu) and Vanuatu (Port Vila, Efate). There is
another variant that completely retains its
juvenile–intermediate foliage on mature speci-
mens; it was observed in cultivation in both Fiji
—Namoka, Naitasiri Province, Viti Levu, and
To n g a —Nuku‘alofa, Tongatapu, and Neiafu,
Vava’u. The leaves on flowering bushes are
typically deeply and narrowly 3, 5-lobed, with
3–15 smaller teeth per leaf, and a low leaf
length:width ratio between 1.4 and 1.6. There
appears to be potential forbreeding this unique
and ornamental foliar trait into modern
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 95

hibiscus hybrids. Note: Further genetic and
field research may indicate introgression of
genes from other Lilibiscus species into these
presumed cultivars, and/or that these entities
warrant recognition as separate subspecies or
species.
Conservation status:Hibiscus bennettii is here
assessed as critically endangered (CR B2 ab,
IUCN 2016) in the wild. This threat category
is based on its highly restricted natural
geographic range (< 10km
2
) on a single
mountain peak (Lat 16°35013.235400S Long
179°18052.812”E, elevation 930 m) on Vanua
Levu, with only four surviving individuals
known in the wild, and highly threatened due
to climate warming and increase in more
intense cyclones. There is an urgent need for
more comprehensive field surveys to ascertain
the species’complete native distribution and
abundance including on mountain peaks
above 700 m on the island of Vanua Levu,
including Mt. Batini (1,111 m) and Mt.
Sorolevu (1,032 m), Mt. Delaikoro (941m),
Mt. Delanacau (744m), Mt. Dikeva (957 m)
Mt. Mariko (881m), Mt. Setura (842m), Mt.
Valili (903m), with a focus on cloud forest on
mountain tops and ridges above 850 m within
the proposed Greater Delaikoro Protected
Area (Tuiwawa and Pene 2014).
Related species: Morphological and genetic
studies (L. Braglia, unpubl.) have shown H.
storckii and H. bennettii to be closely related.
Both species are also related to H. macverryi sp.
nov., H. bragliae sp. nov., and H. cooperi.Hibiscus
bennettii and H. storckii are readily distin-
guished from H. macverryi by their adult
leaves that are unlobed (except in the juvenile
foliage variant of H. bennettii), whereas in H.
macverryi the adult leaves are heterophyllous
including some deeply three-lobed (divided
almost to the leaf stalk and resembling those
of red maple Acer rubrum). The fully opened
petals of H. bennettii and H. storckii are larger
and pale pinkish purple in color, whereas in H.
macverryi they are a rich deep rose. Hibiscus
storckii and H. bennettii differ from H. cooperi in
the absence of a white halo around the darker
eye of its petals and in their more elliptic leaf
shape and dentition, including some larger
teeth or lobes, most evident in intermediate
foliage. Hibiscus bennettii and H. storckii differ
from H. rosa-sinensis in their elliptic to broad
lanceolate leaves with either entire margins
and/or with irregular dentitions in the apical
portion of the leaf blade, and by their lighter-
colored petals with a distinctive, large serrate
or lobate dark crimson red eye zone.
Hibiscus bragliae L. Thoms. sp. nov.
The origins of H. bragliae date from the
nineteenth century and the historical port
town of Levuka on Ovalau in Lomaiviti
Province. Levuka was founded around 1820
by European settlers and traders and soon
became an important South Pacific port with
periodic visits being undertaken by scientific
expeditions. In October 1838, Honore Jac-
quinot, the French zoologist on Dumont
d’Urville’s Astrolabe expedition (1837–1840),
collected the first specimen of Hibiscus in Fiji
from near Levuka. This specimen of H. rosa-
sinensis L. is held in the Herbier Muséum Paris
(P06705102) and is typical of pre-European
Pacific introductions of H. rosa-sinensis with a
truncate leaf base, ±10 uniform serrations per
leaf side and a leaf length:width ratio of 2.1.
Four subsequent botanical collections of
Hibiscus from Ovalau and the neighboring
island Moturiki, undertaken between 1854
and 1928, have been erroneously referred to
H. rosa-sinensis. In October 1854, William
Milne, the Scottish botanist on board the H.
M. S. Herald (a US exploratory expedition
vessel) collected a Hibiscus specimen (Milne
241) on “high grounds”of South Ovalau. This
scant specimen comprised a twig with three
leaves and a flower. A few years later, in 1860,
the German Botanist Berthold Seemann,
collected a Hibiscus specimen (Seemann 22),
presumably from a garden plant with double-
petaled flowers on Moturiki. The next Hibiscus
specimen collection from Ovalau was by U.S.
naturalist Edwin Bryan Jr. (606) on 15
October 1924: a double-petaled red-flowered
plant growing near a walking trail (possibly
planted) at about 200m elevation. On 26 June
1928 U.S. botanist John Gillespie collected a
single red-petaled Hibiscus (4453) on a stream
near the summit of main ridge west of Levuka
(500m elevation). These four Hibiscus speci-
mens differ from H. rosa-sinensis in their
96 PACIFIC SCIENCE •January 2019

narrower leaves, rounded cf. truncate leaf-
base, and their leaf margin dentition.
Following on fieldwork on other Fiji
Islands, which resulted in the discovery of
two new Lilibiscus species, three field visits
were undertaken by the author to Ovalau in
2017. The first visit was undertaken from 3 to
4 April 2017 with local ethnobotanist John
Bennett. During this visit, villagers from
Arovudi reported a location near the base of
Mt Tomuna in the northern part of Ovalau
known as Vei Senitoa—or “place of Hibiscus”in
Fijian—in which wild Hibiscus were to be
found growing among rocks.
During the second visit (8–10 June 2017) an
attempt was made to reach Vei Senitoa from
Rukuruku village. On this trek, an unusual
crimson red-flowered hibiscus was collected
south of Mt. Tomuna (LAJT3, Lat 17°
38.877’S,Long178°47.055’E,350melevation;
9 June 2017, Figures 11,12). This Hibiscus was
growingonanarrow ridge,adjacenttoa disused
walking track between Lovoni and Taviya
Villages, and it was morphologically similar
to the four earlier botanical collections of
Hibiscus in sect. Lilibiscus on Ovalau.Itis
described in the next part of this paper as a
new Fiji endemic species, H. bragliae sp. nov.
Subsequent attempts to reach Vei Senitoa
by a three-person Fijian team led by Bobo
Ahtack, including during the third visit (23–26
July 2017) were hampered due to damage to
tracks and the forested landscape wrought by
TC Winston. Mr. Ahtack reported that Vei
Senitoa had been obliterated by landslides
during and subsequent to TC Winston, and
that the Hibiscus is no longer present at this
location. Mr. Ahtack’s team located a further
seven plants of the crimson red-flowered
hibiscus in the same locality as the earlier find
FIGURE 11. Flower of H. bragliae—south of Mt. Tomuna, Ovalau, Fiji (Photo: Lex Thomson.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 97

in June. Interviews during the third visit with
Fijian informants, including the former Chief
of Taviya Village and senior students from St
John’s College, Cawaci, indicated that the
crimson red-flowered Hibiscus occurs spor-
adically in proximity to and away from tracks
at higher elevations in the interior of north-
eastern Ovalau. The double-petaled variant
that was collected by Edwin Bryan (E. H.
Bryan Jr. 606) in 1924 has been re-collected by
Amato Pauliasi on a ridge west of Ovalau on
25 July 2017 as part of the current study.
Hibiscus bragliae L.Thoms. sp. nov.
Holotype: Fiji, Ovalau, Mt Tomuna. Lex A.
J. Thomson, LAJT3. University of South
Pacific—South Pacific Regional Herbarium,
Suva, Fiji (SUVA).
Etymology: The specific epithet honors Dr.
Luca Braglia (born 10 December 1980) in
recognition of his pioneering DNA studies on
Hibiscus in sect. Lilibiscus, including on the new
species.
Vernacular names: Ovalau hibiscus, senitoa
(vernacular).
References:Wilcox and Holt (1913) p. 37.
Accessions 460 and 461.
Description: Multistemmed large woody
shrubs 3–4 m tall3–5 m across. Lower trunk
somewhat roughened, mottled whitish-light
gray and light brownish-gray often with
patches of moss and lichen; upper stems light
brownish-gray, finely furrowed and with
small, circular lenticels. Branchlets terete,
green with irregularly punctate, vertically
oriented, brown lenticels. Stipules filiform,
3–4mm, light green; soon deciduous. Petioles
midgreen, lighter green and swollen at base
and for apical section with small reddish
marks±above and below swollen apical sec-
tion, terete, slightly twisted, and semi-erect;
1.5–2.5cm long on smaller leaves and 4–5cm
FIGURE 12. Mature leaves of H. bragliae—south of Mt. Tomuna, Ovalau, Fiji (Photo: Lex Thomson.).
98 PACIFIC SCIENCE •January 2019

long on larger leaves. Leaves dull, alternate,
discolorous mid- to dark green above and
mid- to light green below, lanceolate with
rounded leaf base, irregularly shallowly to
deeply crenate-dentate leaf margin (below the
broadest part of the leaf), with typically (3–)
5–10 (–12) teeth on each leaf side. Mostly
4–10cm long 2–5cm wide, with an average
leaf length:width ratio between 2.1 and 2.3,
and average petiole: leaf lamina length ratio
between 0.3 and 0.6. Intermediate and regrowth
leaves on strongly growing new shoots may be
larger, up to 9–12cm long4.5–5.5 cm wide,
and sometimes deeply three-lobed. Venation
palmate typically with three main veins;
midrib raised on lower surface. Flowers c.
9–10cm across, solitary in the upper leaf axils.
Pedicel slender, light green, minutely hairy,
(3–)4–6(–7) cm long, articulated; with
articulation point 8–12mm from base of
flower (or av. 24% of total length above joint).
Both ends of pedicels swollen with the apical
portion grading into a whitish cupulate recep-
tacle. Epicalyx segments 8–10, light green,
whitish near base, linear±1 mm wide, typically
horizontally spreading (or recurved to semi-
erect), highly variable in length with longest
segments typically 6–9mm long. Calyx light
green, whitish near base, fused into straight-
sided tube c. 20–24mm long, calyx lobes 5
acute-isosceles triangular 6–9(–10) mm long.
Petals 5, regular/single,±oblong, 7.5–9cm
long2.5–3.3cm wide and fused for lower
10–12mm; typically recurved, spirally
arranged, burgundy/crimson (crimson to
Indian red, av. RGB 217, 77, 105, Hex Code
D94D69) with a darker eye zone of radius of
25–42mm. Outer surface of petals dark red
with lighter pinkish red veins, and minutely
hairy on petal veins and margins. Staminal
column dark pink near base, becoming progres-
sively lighter along its length, 9–10cm long.
Stamens numerous c. 108–120, thin light
mauvish pink filaments c. 7–9mm long, often
arranged in pairs on the terminal±30% of
staminal column; cream-pale yellow kidney-
shaped pollen sacs, pollen grains pale-yellow.
Style pink branches 5, erect, villose. Stigma pads
±globular, dark pink, villose. Flowers typically
remain open for one day. Fruits immature
capsular, domed cylindric, minutely hairy.
Other specimens examined: Fiji, Ovalau,
Lomaiviti Province. High grounds, South
Ovalau, U.S. exploratory expedition, Milne
241. MacGillivray, October 1854 (K). On
stream, near summit of main ridge west of
Levuka. Alt 500 m. J. W. Gillespie 4453, 26
June 1928 (BISH). Double-petaled variant near
trail (likely planted). Alt±200 m. E. H. Bryan
Jr. 606, 15 October 1924 (BISH).
Natural Distribution, habitat, and associates:
Knownonly fromrocky sitesand drainagelines
inmountainouslocations (±300–500m)on the
volcanic island of Ovalau. It may have also been
planted along traditional walking tracks. Asso-
ciated rainforest tree and plant species include
Acacia richii,Agathis macrophylla,Aleurites
moluccana,Alphitonia zizyphoides,Bischofia java-
nica,Calophyllum spp., Cananga odorata,Dios-
corea nummularia, Dysoxylum spp.,
Endospermum macrophyllum,Ellatostachys fal-
cata,Ficus spp., Gironniera celtidifolia,Glochidion
spp., Gymnostoma vitiense,Gnetum gnemon,
Intsia bijuga,Macaranga spp.,Meremmia peltata,
Pleiogynium timoriense,Syzygium spp., Retro-
phyllum vitiense, and Veitchia spp. The leaves of
the new species appear to be especially
attractive to leaf-chewing insects, at least in
its native habitats.
Genetic relationships and breeding system: The
phylogenetic interpretation of the Tubulin-
based Polymorphism (TBP) data for many
sect. Lilibiscus species and cultivars suggest a
close association between H. bragliae and the
other known Fijian endemic Lilibiscus species
(L. Braglia, unpubl. data). Koopman and
Baum (2008) have suggested that one dispersal
event from Madagascar, likely around one
million Ma, gave rise to the predominantly
Indo-Pacific Hibiscus sect. Lilibiscus group.
The new Fijian species shares floral and foliar
traits with Mascarene Island species, espe-
cially H. genevii, such as small flowers with
spirally arranged narrow petals and narrowly
trilobed juvenile and intermediate leaves and
likely represents an early or ancestral species
of Lilibiscus in the South Pacific Islands.
Hibiscus bragliae is distinguished from other
Fijian species in sect. Lilibiscus by its ±funnel-
shaped flowers with conspicuously spirally
arranged, overlapping (near base) narrow
petals and also as follows: H. storckii has
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 99

larger±flat flowers c. 16–18 cm across, laven-
der pink petals with a distinctly serrated, dark-
red basal petal zone cf. H. bragliae (open
flowers c. 9–10cm across). H. bennettii has
considerably larger±flat flowers c. 16–18 cm
across, white or pink petals with a white trim.
H. macverryi differs in its broader, deep rose-
colored flowers with a large serrated dark red
eye zone; its bright yellow anthers, its
heterophyllous mature foliage, which invari-
ably includes some trifoliate leaves. H. rosa-
sinensis differs in its broader leaves (leaf length:
width ratio between 1.2 and 1.7 cf. ≥2.1) that
have a ±regularly serrate margin cf. an
irregularly crenate-dentate margin in H.
bragliae.
Early Horticultural References —The earliest
horticultural reference to the single and
double crimson-petaled forms are in a report
on Hibiscus breeding in Hawai‘i in the early
1900s (Wilcox and Holt 1913), in which they
are recorded as being imported from Fiji by
Mr J. D. McInerny. The descriptions of these
two forms, slightly adapted from Wilcox and
Holt (1913) and with measurements con-
verted to SI units, are as follows:
Fijian crimson single.Bush: slow growing,
sparsely branching, with smooth gray bark
throughout. Leaves: elliptic or narrowly ovate,
coarsely dentate, acuminate, smooth, light
green, prominent veins, 2.5–3.8cm wide,
5–7.5cm long, petiole 3.8cm long. Flower:
dark crimson in throat, gradually paling to
light crimson pink at border, 7.6cm wide,
funnel shaped, with recurved petals, column
7.5cm long, pale crimson, stigmas far pro-
truding, peduncle, 2.5cm long, involucre
bracts 6 or 7, green, poor seeder.
Fijian double crimson.Bush: low growth,
upright, sparsely branched, dense foliage, bark
light gray, green twigs. Leaves: lanceolate or
ovate, smooth, crenate, 2.4 to 6.4 cm wide, 6.4
to 8.9cm long, petiole 2.5 cm. Flower: 6.4cm
wide, double, petals crimson pink, with white
veins and dark petal claws. Peduncle 5cm
long, bracts 8, short, green, recurved, not self-
seeder.
It is likely that U.S. Hibiscus breeder Ross
Gast was responsible for the introduction of
‘Fiji White’into Hawai‘i. In his letters to J. W.
Staniford, Gast describes a Hibiscus (Fiji #1) he
collected in Fiji in 1963 as follows “I did get
one outstanding variety, I think. This is a
creamy white single with heavy raspberry
colored rays extending from the base of the
petals almost to the tip. It appears to be a
strong grower with good foliage, and I am
sure that it will become a welcome addition to
our commercial list when we introduce it.”In
another letter Gast described it as “the most
unusual hibiscus I’ve ever seen.”Gast’s Fiji #1
closely matches the description of the juvenile
foliage variant of H. bragliae.
Horticultural varieties and importance: Three
Hibiscus entities that appear to be selections of
H. bragliae were found in cultivation as
follows:
1. Double-petaled variant (Figures 13 and
14): This variant has likely been planted on
a walking trail, west of Levuka, Ovalau, and
was an early introduction into Hawai‘i
around 1900. The double-petaled variant
has functionally male flowers with±16–20
petals, which decrease in size toward the
center of the flower from 4014 mm in
outer petals to 10mm wide in inner petals.
The petal color is crimson in the center,
with whitish veins, becoming pinkish
toward the petal edge; inner petals are
lighter, whereas the back of the petal varies
in color from crimson to pink. The leaves
are the same as the type form.
2. Multicolored petal variant (Figure 15):
This lighter multicolored petal variant is
widespread in cultivation, including Fiji
(Viti Levu and Taveuni), Tonga (Tonga-
tapu) and New Caledonia (Nepou, Grand
Terre, and Lifou, Loyalty Islands). The
inner petal pigmentation comprises a well-
defined dark crimson eye zone, then a rose-
pink zone fading paler to white on the outer
zone of the petal. The stigma pads are light
pink. The leaves are typically lanceolate,
but sometimes approaching elliptic.
3. Juvenile foliage, multicolored petal variant
(Figure 16): This variant has been grown
under the name ‘Fiji White’in both the
United States (Hawai‘i) and Japan. It
retains its juvenile to intermediate foliage,
typically deeply and narrowly trilobed, on
flowering plants with only a small percen-
100 PACIFIC SCIENCE •January 2019

FIGURE 13. Flower of double-petal variant of H. bragliae—west of Levuka, Ovalau, Fiji (Photo: Lex Thomson.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 101

tage of entire mature leaves. The petal
colors are similar to the multicolored petal
variant (2), but with a broader white outer
zone and with bright yellow pollen grains
and creamy-lemon stigma pads.
Note: Further genetic and field research
may indicate introgression of genes from
other Lilibiscus species into cultivars 2 and 3,
and/or that these entities warrant recognition
as separate subspecies or species.
Conservation Status:The speciesisassessed as
criticallyendangered(CE C1,IUCN2016) due
to its small population size (likely < 250 mature
plants in the wild), highly restricted distribu-
tionona singleisland, andthreatsfrom extreme
climatic events, especially severe tropical
cyclones (category 3 and above), environmen-
tally invasive weeds coupled with a reduced
interest amongst Fijians in growing and
maintaining Hibiscus in villages and as markers
for paths between villages, and loss of species
purity due to hybridization with introduced,
exotic Hibiscus species. The H. bragliae popula-
tion that had been growing at Vei Senitoa (near
the base of Mt Tomuna, northern Ovalau)
appears to have been submerged by landslides
associated with category 5 TC Winston in
February 2016. Given its critically endangered
status, it is suggested that ex situ populations of
H. bragliae, including all four known variants,
be established in Savurua Botanical Gardens
and Thurston Gardens in Fiji as well as in other
Botanical Gardens internationally.
There is an urgent need for more com-
prehensive field surveys to ascertain the
species’complete native distribution and
abundance in open forest habitats above 350
m elevation on Ovalau, especially focusing on
rocky slopes and drainage line habitats above
450 m elevation on the north, east, and
southeastern portions of Ovalau.
Hibiscus macverryi L. Thoms. and Braglia sp.
nov.
In March 2016, on a visit to Taveuni in search
of H. storckii, the first author was informed by
Mr. Suliano Manabua, a Fijian farmer, that a
Hibiscus morphologically near-identical with
Hibiscus ‘Ruby Rose’was growing wild in the
Waibula River Catchment on Taveuni. On
8 June 2016, the first author walked with
FIGURE 14. Habit of H. bragliae (double-petal variant)—west of Levuka, Ovalau, Fiji (Photo: Amato Pauliasi.).
102 PACIFIC SCIENCE •January 2019

FIGURE 15. Tri-color petal variant of H. bragliae—Nepou, New Caledonia (cultivated) (Photo: Celine Babouline.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 103

Mr. Manabua from the Villa Maria Settlement
into the area where the wild Hibiscus was
growing. A single surviving specimen was
located in lowland tropical moist forest
growing adjacent to Waibula River in gravelly
sand with boulders (16°480S and 179°520W,
elevation 50 m, Figures 17,18). Three other
Hibiscus plants growing within and on the
margin of the bouldery riverbed had been
destroyed during the flooding by category 5
TC Winston earlier in the year. Subsequently
a number of Fijian villagers from northeastern
Taveuni, including from Qeleni Village,
informed the senior author that the new
species is rare in more open forest areas in the
mid-upper catchment Waibula River, in
rugged dissected terrain. It also has great
local cultural significance: Di Lutu, the
ancestor goddess of a region stretching from
Naselesele to Lavena on the northeastern
coast of Taveuni, wears a flower of this
hibiscus behind her right ear. Di Lutu is
reputed to both “protect”the region and also
to “haunt”a pool adjacent to Qeleni village
(Paul Geraghty, University of South Pacific,
pers. comm.). The new species has both
highly distinctive flowers and heterophyllous
mature foliage and is here described as H.
macverryi.
Hibiscus macverryi L.Thoms. and Braglia, sp.
nov.
Holotype: Fiji, Taveuni, Waibula River. Lex
A. J. Thomson. LAJT2. 8 June 2016. Uni-
versity of South Pacific—South Pacific Regio-
nal Herbarium, Suva, Fiji (SUVA).
Etymology: The specific epithet honors
Father Michael McVerry (born 4 September
1943) in recognition of his assistance with
locating the new species in the wild and for his
contributions to Taveuni, including the work
of Marist Tutu Rural Training College to train
young farmers and guide married couples, and
the Taveuni Flower Society.
Vernacular names: Di Lutu’s‘aute (pre-
ferred); jungle hibiscus, ruby rose hibiscus,
Mrs. Hassinger’s hibiscus
Description: Multistemmed large woody
shrubs 3–4 m tall3–5 m across. Lower trunk
somewhat roughed, mottled light gray and
brownish-gray often with patches of moss and
lichen; upper stems smooth, light gray.
FIGURE 16. Flower of juvenile leaf variant of H. bragliae “Fiji white”Big Isle, Hawai‘i, USA (cultivated) (Photo: Kenton
Sanchez.).
104 PACIFIC SCIENCE •January 2019

Branchlets terete, green to brown. Stipules
filiform with recurved tip, 5–6mm, light
green; soon turning light brown and decid-
uous. Petioles reddish green or sometimes light
green tinged red (0.8–) 2.1–4.8 (–6.2) cm long;
upper margin puberulent. Leaves semiglossy
to glossy, alternate, discolourous, mid- to dark
green above and light green below, hetero-
phyllous, extreme variation in leaf shape from
broad lanceolate with irregular, sometimes
deep serrations or lobes, through to deeply
palmately lobed into three segments, highly
variable in size (4–)6–10 (–12) cm long (2–)
4–9(–10) cm wide, leaf margin irregularly
serrated, base obtuse/rounded. Intermediate
and regrowth leaves on strongly growing new
shoots larger up to 13cm long 11cm wide
and new growth more frequently palmately
trilobed. Leaf length:width ratio extremely
variable ranging from (0.9–) 1.0 to 2.0 (–2.1).
Venation palmate typically with three main
veins; midrib raised on lower surface. Flowers
10–14cm across, solitary in the upper leaf
axils, borne on (3.5–)5–8(–9) cm long
slender, light green articulated pedicels with
articulation point 4–11mm from base of
flower (or av. 10% of total length of pedicel).
Epicalyx segments (5–)6–7(–8), light green,
whitish near base, linear, straight/ascending,
7–12mm long 1–2 mm wide, sometimes
with an acuminate tip. Calyx light green,
whitish near base, fused into slightly swollen
or campanulate tube c. 22–28mm long, calyx
lobes 5 acute-isosceles triangular 10–14mm
long. Petals 5, regular/single, obovate, (5–)
6.4–6.6 cm long(4.2–)4.6–5.0cm wide and
fused for lower 10–12mm; flat or recurved
before closing, slightly overlapping, deep
rose (French rose, av. RGB 246, 76, 133, Hex
Code F64C85), with a large, dark crimson
FIGURE 17. Hibiscus macverryi in lowland tropical moist forest—Waibula River, Taveuni, Fiji (Photo: Lex Thomson.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 105

serrated eye zone of radius of (2.1–)2.5–3.2
cm. The outer surface of petals have
prominent raised veins, rose-colored with
whitish flecks on one-third of petal (eye zone
not visible on outer petal surface except with
strong back light). Staminal column dark
crimson near base and then midcrimson, (6–)
7–8(–9) cm long. Stamens numerous c. 70–90,
thin light pink-colored filaments c. 6–8mm
long with yellow kidney-shaped pollen sacs
(and pollen grains)±tightly grouped along the
upper third of staminal column. Style crimson,
branches 5, spreading/semi erect, villose.
Stigma pads±globular, dark crimson, villose.
Flowers usually remain open for one day. Fruits
and seeds not observed.
Other specimens examined: Fiji: Vanua
Levu, Macuata Province, mountains near
coast. W Greenwood 637, 2 September
1923, K; Wakaya Island, Lomaiviti Province,
low grounds. Milne 36, July 1855, K.
Natural distribution, habitat, and associates:
Known only from six locations—all lowland
(50–300m) tropical moist forest, typically in
gravelly sand amongst volcanic rocks and
boulders in open to shaded forest sites
adjacent to creeks and rivers—in the Waibula
River catchment, Taveuni, Caukadrove Pro-
vince and adjacent to creeks in near-coastal
mountain ranges in Macuata Province, Vanua
Levu, and Wakaya Island, Lomaiviti Province.
Associated tree species on Taveuni include
Geissois ternata A. Gray, Calophyllum sp.,
Pometia pinnata J. R. Forst. & G. Forst., and
Premna serratifolia L. The Waibula River
specimen was infected with the Hibiscus
erineum mite (Aceria hibisci (Nalepa) Keifer),
a parasitic mite that was first recorded in Fiji
FIGURE 18. Hibiscus macverryi flower—Waibula River, Taveuni, Fiji (Photo: Lex Thomson.).
106 PACIFIC SCIENCE •January 2019

(Nalepa 1906). The flowers are visited by the
orange-breasted myzomela (Myzomela jugu-
laris Peale), which may assist pollination.
Genetic relationships and breeding system:
According to previous published DNA-based
studies (Braglia et al. 2010), H. macverryi (as
‘Ruby Rose’) is closely related to Hibiscus ‘Fiji
Island’and H. cooperi (as H. storckii). Hibiscus
macverryi is readily distinguished from other
South Pacific Island species in sect. Lilibiscus
by its mature heterophyllous foliage, invari-
ably including some trilobed leaves. The
flowers of H. macverryi differ from the four
other Lilibiscus species as follows:
H. bennettii: Has rose pink or pale pink
petals with a white edge or white petals, 7–9
cm long, petal veins white, distinctive lobate
dark-red basal zone (c. 1.9–2.8cm radius),
filaments white, stigma pads light-yellow
whereas in H. macverryi the petals are an
intense rose, < 7cm long, petal veins rose to
deep rose, with a distinctive serrated dark red
basal zone (c. 2.5–3 cm radius) due to dark-red
pigmentation extending along each petal vein,
filaments pink, stigma pads crimson.
H. cooperi: Main petal color variable
(Turkey red, salmon, or pale rose pink or
white), but with a small (<1.7cm) basal dark
red zone surrounded by a large whitish-pale
halo and with white petal veins, whereas in H.
macverryi the main petal color is invariably
intense deep rose with a distinctive very large
(radius of 2.5–3cm) serrated, dark-red basal
zone due to the dark eye zone extending along
each main petal vein.
H. rosa-sinensis: Petals may be in single or
multiple whorls, typically deep rose red to
bright red; each petal with a small dark red
basal zone (radius <1.0cm), whereas H.
macverryi flowers have a highly distinctive
very large (radius of 2.5–3cm) serrated, dark-
red basal zone.
H. storckii: Has larger lavender pink petals
(flowers c. 14–18cm across) cf. H. macverryi
flowers (c. 10–14cm across) and with a larger
serrated, dark-red basal zone (with the
dark eye zone also extending along each main
petal vein).
Early horticultural references: The earliest
horticultural reference appears to be Bull
(1872) under the name Hibiscus (rosa-sinensis)
carminata perfecta:“A charming stove plant, in
habit resembling H. rosa-sinensis, recently
imported from the South Sea Islands. It has
stalked ovate obscurely three-lobed leaves
which are deeply toothed, and very large
flowers, nearly 5 inches across, with broad and
slightly undulated petals, forming a full round
flower, of perfect shape, and of a rich soft
carmine rose with a deep crimson eye. The
staminal column is very prominent, and adds
much to the beauty of the flowers, the column
being of a rosy hue, the numerous stamens
bright yellow, and the five stigmas a rich
velvety crimson.”Hibiscus macverryi is the
same entity as Hibiscus ‘Mrs Hassinger’that
was introduced and used extensively in the
early Hawaiian hybrid breeding programs
(Wilcox and Holt 1913). The description of
‘Mrs Hassinger’(adapted from Hoyt 1950 and
Wilcox and Holt 1913) is: Moderately
vigorous to vigorous shrub. Typical flower
13–15cm across, extremely flat–faced with no
recurve, petals overlap 0.3cm from base, out
halfway to periphery, slightly ruffled on entire
margin except in eye. Basal 2.5cm of petal
deep reddish-black merging abruptly with
rose red for next 1.3cm, fading gradually to
lighter shade of rose red. Back of petals from
base up to 2.5cm is cardinal red fading to
spiraea red; column 8–10cm long, slender,
basal half same color as base of petals fading to
spiraea red; stamens start at 5.7cm from base
and continue for 2.5 cm, filaments 6.5mm, not
close packed; stigmas just above, 15mm
diameter, spread 0.3–0.6cm crimson. Calyx
1.9–3.2cm long, medium green. Epicalyx
segments 5–6, 1cm 0.15cm, medium green.
Peduncle 5–7cm long, stiff, slender. Leaves
4.5–9cm long 2.5–6cm, lanceolate-ovate or
lobed-deeply cleft on each side of midrib
forming a three-parted leaf, serrate, flexible,
leathery; dark semiglossy green on top,
medium semiglossy green on bottom, glab-
rous both sides; petiole 1.5–3cm long, green-
ish-red, glabrous except for hirsute line on
dorsal surface. Bark light grayish-brown,
comparatively smooth; branchlets light green,
glabrous. Flower held full-face, opening early
morning, closing late afternoon, one-day
bloom, blooms year round, no perfume;
successful on own root or grafted; origin
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 107

unknown; crosses readily but not self-seeding;
good consistent blooming form.
Horticultural varieties and importance:Hibiscus
macverryi has been available in horticulture
over a long period, including in the UK,
Europe, Asia, Hawai‘i, and mainland United
States, in more recent times under the name
“Ruby Rose.”It has been an important parental
species contributing to modern tropical Hibis-
cus hybrids, e.g. 34 hybrid crosses involving
‘Mrs Hassinger’are described in Wilcox and
Holt (1913), including 30 as the female parent.
During the current study, only rather minor
morphological variations were observed in
cultivated plants of H. macverryi in Fiji. These
included one individual with slightly larger
flowers and leaves, and brown branchlets
(Rukuruku, Ovalau) andanother with a slightly
lighter, less “star-shaped”inner petal pigmen-
tation (Raiwaqa, Viti Levu).
Conservation status: The species is assessed
as critically endangered (CE C1, IUCN 2016)
due to its small population size (likely < 250
mature plants in the wild), restricted distribu-
tion (known with certainty from only two
locations) and threats from environmentally
invasive weeds, hybridization with introduced
Hibiscus species, and extreme climatic events,
especially severe tropical cyclones (category 3
and higher) and flash flooding given its
exclusively riparian distribution. Of the six
mature individuals the senior author was
taken to visit during 2016–2017, five had
been destroyed by flooding associated with
TC Winston. These included three on the
Waibula River, the famous Di Lutu-associated
Hibiscus plant at the Qeleni pool and another
plant on Wai ni Aute (Nakaute Creek) near
Bainikarasa. A small population of H. mac-
verryi around Bainikarasa was reported to
have been destroyed by much earlier farming
activities. There is an urgent need for more
comprehensive field surveys to ascertain the
species’complete native distribution and
abundance including along drainage lines on
the eastern side of Taveuni from Naselesele to
Lavena, and especially in the mid-upper
catchment of the Waibula River, as well as
along drainage lines in near-coastal mountain
ranges in Macuata Province, Vanua Levu.
Given its critically endangered status, it is
recommended that ex situ populations from
surviving populations and individuals of H.
macverryi be established in Fiji (Savurua
Botanical Gardens and Thurston Gardens)
as well as in other botanic gardens inter-
nationally.
Hibiscus cooperi Hort. ex Lemaire
Hibiscus cooperi is a highly ornamental plant
with red, pink or white flowers. It is best
known in horticulture through its original
form with tricolored variegated leaves. H.
cooperi quickly became popular in heated
glasshouses in the northern hemisphere after
its introduction into Britain around 1858 from
the South Pacific via Sydney, Australia.
Shortly after its introduction into Europe,
several exquisite illustrations of the species
were published (Lemaire 1864, van Houtte
1864). H. cooperi Hort. (in Journal Général
d’Horticulture, p. 109, t. 1556; 1864) was
synonymized under H. rosa-sinensis by
Hochreutiner (1900). Since that time, the name
H. cooperi has been used loosely in horticultural
circles—more recently on social media—to
encompass almost any variegated hibiscus in
sect. Lilibiscus. The Plant List reports Hibiscus
cooperi auct.as an unresolvednamewithoriginal
publication details as J. Gén. Hort. 5: 109 1863
(http://www.theplantlist.org/tpl1.1/record/
kew-2849722 accessed 16/2/2017).
It is proposed here to reinstate H. cooperi
given that the species is well-marked morpho-
logically and genetically distinctive from H.
rosa-sinensis and other Lilibiscus species (Braglia
et al. 2010 Amplified Fragment Length Poly-
morphism[AFLP]study inwhich H. cooperi was
labeled as H. storckii)), freely seeding and
breeding true-to-type for flower and leaf
characters.
Hibiscus cooperi Hort. ex Lemaire (1864).Ibiscus
cooperii, ketmie deCooper. L’Illustration horti-
cole (illustration by Stroobant) 11: Pl. 412.
Designation of lectotype: Because there was
no type specimen designated for H. cooperi in
Lemaire’s description in 1864, the accompa-
nying illustration by P. Stroobant is
hereby designated as the lectotype (Fig. 19).
Although Meehan’s (1863) rudimentary
108 PACIFIC SCIENCE •January 2019

FIGURE 19. Early illustration of Hibiscus cooperi. Stroobant (1864) (lectotype).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 109

description of H. cooperi predates that of
Lemaire (1864), the latter’s description is
complete, including an accurate illustration as
lectotype, and accordingly has been selected
for the protologue.
Etymology: The species name honors Sir
Daniel Cooper (1821–1902), a merchant with
trading connections to South Pacific Islands,
as well as being a philanthropist, a member of
the New South Wales (NSW) Legislative
Council and Speaker in the NSW Legislative
Assembly. Cooper has been credited with
bringing the species into cultivation in
Australia from the South Pacific Islands.
Vernacular name: Cooper’s hibiscus
References:Anon (1863).Hibiscus cooperi (as
nom nud.) by “G.A.”on 15 September 1863.
Journal of Horticulture and Cottage Gardens.
Vol. V (new series): 213.
Description: Multistemmed large woody
shrubs or small trees, 3–5 m tall, sometimes
reduced to a low sprawling shrub to 1–1.5 m.
Bark grayish, somewhat rough. Branchlets
terete, reddish-brown to green, ageing to
light gray with leaf scars prominent. Aerial
roots may form on lower parts of the trunk
and main branches. Stipules filiform, 5–12
mm, light green, reddish near base. Petioles
dark reddish-brown to brownish-green, (12–)
17–27mm long, with sparse to medium
density of stellate and single hairs (mostly
0.2–0.3mm long). Leaves semiglossy, simple
(never lobed), alternate, thin, discolorous
mid- to dark green to reddish-green above
and light green below and/or variegated
(mosaic of green, light gray-green, white,
often with rose pink patches especially near
leaf margin), lanceolate to broad lanceolate,
5–11.5 (–13) cm long(2.5–)3–5(–6.5) cm
wide, broadest above midpoint, leaf length:
width ratio 1.9 to 3, leaf base obtuse-rounded.
Leaf margin entire to sparsely irregularly
serrate. Cotyledons green±orbicular 15 mm
diameter. Intermediate and coppice regrowth
leaves±regularly serrated along the upper
half to two-thirds of leaf margin. Venation
palmate typically with three main veins±
dark-reddish; midrib raised on lower surface.
Leaf margins and veins sparsely pubescent
(short stellate and single hairs). Flowers
solitary in the upper leaf axils, borne on
3.5–4.6cm long slender, articulated flower
stalks, with articulation point 9–20mm from
base of flower (or av. 32% of total length above
joint). Epicalyx segments 7–9, green (or whitish-
light green in variegated form), linear, acumi-
nate, spreading to erect, c. 15–20mm long.
Calyx light green, fused into slightly swollen
tubec.30–35mm long,calyx lobes(2–) 5 acute-
elongated isosceles triangular with a raised
longitudinal vein (L:W –at base of lobe–ratio
4.2–6.8, lobes frequently remaining partly
fused at anthesis). Petals 5, regular, single,
obovate 5.5–6.5cm long 3.5–5 cm wide,
fused for lower 9–12mm; separate to slightly
overlapping, margin sometimes recurved, red
or pink (Type form: salmon red, av. RGB 243,
75, 86, Hex Code F34B56; Naselai pink: rose
pink, av. RGB 244,126,169, Hex Code
F47E49), cream, with a small red eye zone
(10–16mm radius) enclosed by a substantial
white halo and with white veins radiating into
upper portion of the petal. Outer surface of
petals light to rose pink with whitish flecks.
Staminal column dark crimson near base and
then light crimson to red, 6–7cm long. Stamens
numerous (65–)70–80 (–85), thin pink-colored
filaments, 3–4.5mm long, with light yellow
kidney-shaped pollen sacs, along the terminal
quarter of the staminal column. Style white,
then five-branched, red, spreading/semi-erect,
villose, 1–1.5cm long. Stigma pads ±globular,
red (cream in the light-petalled form), villose.
Fruit five-valved capsule, oblong-cylindric, c.
2.5–3cm long, green turning light to dark
brown at maturity; typically with c. 30–40seeds
per capsule.Seedblack/dark gray or brown,
short sparse hairs, irregular-blocky kidney to
triangular c. 42.5–3mm.
Other specimens examined: In addition to the
protologue and the lectotype, the sources used
for the preceding description included culti-
vated plants and high-resolution images of
cultivated specimens (both living plants and
herbarium specimens). The living specimens
and their high-resolution images examined
were those being cultivated in the South Pacific
Islands including Lautoka, Viti Levu and
Tokou, Ovalau, Fiji; Erromango (Figure 20)
and Port Vila, Efate, Vanuatu; and Medana,
Guadalcanal, Solomon Islands as well as
the Auckland Domain Wintergardens and
110 PACIFIC SCIENCE •January 2019

Wellington Botanic Garden, New Zealand;
Royal Botanical Gardens, Melbourne (Fig. 21);
Royal Botanic Gardens, Kew London, UK
(Figures 22 and 23); the National Tropical
Botanical Gardens, Kaua‘i, Hawai‘i, USA; and
private gardens in O‘ahu, Hawai‘i, USA and
Oak Park, Melbourne, Australia.
The herbarium specimens and their high-
resolution images examined were as follows:
Variegated leaved specimens: Venezuela,
Puerto Caballo (cultivated). 1893–1894. (ex
Herb. E. Drake), P06612454, P; Venezuela,
Puerto Caballo (cultivated). (ex Herb. E.
Drake), P06612357, P; Caribbean, Lesser
Antilles, Guadeloupe (cultivated). 1893. (ex
Herb. De La Guadeloupe et Dépendances,
Pere Duss 122) P06612453, P; Brazil, Rio de
Janerio (cultivated). 27 December 1873. (ex
Herb. Glaziou. Pl. de Rio Janeiro)
P06612459, P; Caribbean, French Antilles,
Martinique, Forte de France (cultivated) M.
Mouret 595 May 1904, P06612452, P; Brazil,
Sao Paulo, Rue Prof. Luis Rosa, Campinas
(cultivated). 16 November 1953. Herbarium
of P.A. Fryxell. 00386906, NY. Nonvariegated
leafed specimens: United States, Hawai‘i,
Ka’ua‘i (cultivated). Kava Vale 001, 1 Oct
2016. National Tropical Botanical Garden
Herbarium, Hawai‘i, 1000054778, NTBG.
Natural distribution, habitat and associates:
Hibiscus cooperi is unknown in the wild, but has
been presumed native to Melanesia. It is likely
that H. cooperi was first introduced to Australia
from either Vanuatu or New Caledonia. H.
cooperi is widely cultivated in Vanuatu, e.g.
Efate, Erromango, Tanna, but was unknown
on New Caledonia in the nineteenth century
(Gildas Gâteblé, pers. comm.). Daniel Coop-
er’s Uncle’s (also Daniel Cooper) firm Cooper
and Levy procured the first cargoes of
FIGURE 20. Hibiscus cooperi salmon pink petals, green-foliaged form—Happylands, Erromango Island, Vanuatu
(cultivated). (Photo: Gildas Gâteblé, IAC, Noumea.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 111

sandalwood from Vanuatu, presumably from
Erromango in the 1840s, and may well have
acquired the hibiscus during these voyages. In
the Naturalis Biodiversity Centre there is a
Hibiscus specimen (http://data.biodiversity
data.nl/naturalis/specimen/L.2358217) col-
lected in 1840 from Tondano, North Sula-
wesi, Indonesia that closely matches H. cooperi.
Further research will be required to clarify the
identity of Tondano hibiscus: If confirmed as a
non-variegated form of H. cooperi, then the
early date of Tondano collection may suggest
that H. cooperi is an aboriginal introduction
from Sulawesi into the South Pacific.
Genetic relationships and breeding system:H.
cooperi is distinguished from other species in
sect. Lilibiscus by the combination of its unique
petal pigmentation pattern, long linear epi-
calyx segments, long narrow triangular calyx
lobes (often remaining partly fused at
anthesis), dark reddish-brown petioles, and
lanceolate–broad lanceolate leaves with entire
FIGURE 21. Hibiscus cooperi variegated-foliaged type form with red petals (RHS color chart 46 C, Turkey Red)—Royal
Botanic Gardens, Melbourne, Australia (cultivated) (Photo: David Robbins.).
112 PACIFIC SCIENCE •January 2019

to irregularly serrated margins. A DNA-based
study performed by Braglia et al. (2010)
applying the widely recognized AFLP method
indicated that H. cooperi (labeled in the study
as H. storckii) is genetically distinctive from H.
rosa-sinensis cultivars with its closest relatives
being the Fijian entities H. macverryi sp nov.
(labeled in the study as ‘Ruby Rose’) and ‘Fiji
Island’, together with a several hybrid culti-
vars including ‘Psyche’and ‘Snow Queen’that
are likely to have H. cooperi and H. schizopetalus
in their ancestry. Hibiscus cooperi differs from
the four Fijian Lilibiscus species in its entire
seedling and intermediate leaves, which are
never divided or deeply lobed (Figure 24).
Early horticultural references: The earliest
complete Latin description of H. cooperi has
been provided by Lemaire (1864). As indi-
cated previously, the earliest known descrip-
tion of H. cooperi is that of Thomas Meehan
(1863). Meehan (1826–1901) was a British-
born nurseryman and botanist who worked at
Kew Gardens from 1846 to 1848 before
moving to Philadelphia, United States It is
likely that Meehan’s brief description is a
second-hand account provided through a
connection in his British horticultural circles:
“Hibiscus cooperi –a beautiful plant, intro-
duced from New South Wake, by Sir Daniel
Cooper, Bart., after whom it is named. It
forms a shrub of dense habit, producing large
scarlet flowers and tri-colored foliage of great
beauty. The leaves are of a bright green color,
distinctly striped and flaked with crimson and
white; and the various shadow of color
produced by the foliage, as it assumes different
stages of growth, gives it a very pleasing and
attractive appearance. Its free growth and
FIGURE 22. Hibiscus cooperi “Naselai pink”rose pink petals, green-foliaged form—Kew Gardens, UK (cultivated).
(Photo: Kew Gardens.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 113

beautiful foliage render it a plant that can be
strongly recommended for decorative pur-
poses generally.”
A variegated-leaved hibiscus, undoubtedly
referrable to H. cooperi, was introduced by
James Veitch into the European horticulture
and nursery trade around 1858 (Lindley
1858). However, Veitch is unlikely to have
coined the name H. cooperi because in Hortus
Veitchianus (1906, p. 264) it is cited as “H.
Cooperi, Hort.,”i.e. “hortulanorum,”under
the name H. rosa-sinensis var. cooperi. The first
publication of the name H. cooperi appears to
have been by “G.A.”on 15 September 1863 in
the Journal of Horticulture and Cottage Gardens
(p 213, nom nud.). Another early reference to
H. cooperi,asH. cooperi Hortul., which
included an illustration of a sterile twig, was
provided by Van Houtte (1864). Based on the
nursery catalogue of James Veitch, it was
reported that H. cooperi was brought from
Australia by Sir Daniel Cooper: It was further
noted that the hibiscus bore large red flowers,
struck relatively easily from cuttings and
needed to be grown in a glasshouse during
winter months in Europe.
A further early reference to H. cooperi is
provided by Mettler and Otto (1865), and
these authors listed H. tricolor Hort. Belg as a
synonym. Hibiscus tricolor Dehnh. (Cat. Horti
Camald. ed. 2: 20 1832) is here considered to
be a form of H. rosa-sinensis. Dehnhardt’s
description of H. tricolor indicates several
differences with H. cooperi including diagnos-
tically that the petals are veined white to the
base in H. tricolor, whereas in H. cooperi the
petals have a dark red eye surrounded by a
distinct white halo. There are several other
minor differences between these two entities
such as 6–8 epicalyx segments (cf. 7–9in
FIGURE 23. Hibiscus cooperi flower bud showing long thin epicalyx segments and narrow calyx lobes; variegated form—
Kew Gardens, UK (cultivated). (Photo: Kew Gardens.).
114 PACIFIC SCIENCE •January 2019

H. cooperi) and the stigma pads are recorded as
golden (cf. red in H. cooperi). Furthermore,
Dehnhardt’s description of H. tricolor makes
no mention of variegated leaves and the
specific epithet likely refers to the three colors
found on the outer surface of the petals.
Horticulturalvarietiesandimportance:Hibiscus
cooperi is widely cultivated as an ornamental
both in the tropics/subtropics and in heated-
indoor environments in cooler climates. The
type form and the most commonly cultivated
varieties follow.
Type form—this is the variety with varie-
gatedfoliagethatwas originallyintroducedinto
cultivation from the South Pacific Islands
under the name H. cooperi and accordingly
selected as lectotype. The flower petal color is
typically an intense salmon pink-red but pink
petaled variegated plants also exist in cultiva-
tion. We have observed that selfed seedlings of
the variegated form may include seedlings with
different foliage pigmentation types, whereas
selfed seedlings of green-leaf forms may
include variegated seedlings at low frequency.
‘Copper leaf’—a dwarf shrub form with
bronzed foliage and scarlet-crimson petals
(Wilcox and Holt 1913)
‘Thumbelina’—A sprawling shrub form
found in cultivation in Fiji. It was reported
to have been produced from a crossing of H.
storckii x‘Honey Do’—Hans and Sonja Drost
and registered in 1982—but more likely was
derived from a selfed seed of H. cooperi or is a
natural variant. It has a large white zone
FIGURE 24. Regrowth leaves collected from grafted scions of four endemic Fiji Lilibiscus species (H. bennettii, H. bragliae,
H. macverryi, and H. storckii), plus H. cooperi. (Photo: Tong Wang.).
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 115

surrounding the eye on small flowers about 9
cm across (with individual petals measuring
about 5018 mm). The leaves are smaller
than typical H. cooperi, mostly 5.5–7.5
2.5–3.7cm wide with a length:width ratio of
between 1.9 and 2.5.
‘Naselai pink’—Naselai pink typically has
pink flowers and green foliage. For several
decades, a Hibiscus entity referred to as H.
storckii has been grown in botanic gardens of
Europe and has been circulating under that
name in the global horticultural trade (e.g.
Rivers-Smith 2004,Giraud 2013). The sources
of most of these plants in Europe and the
United States appears to have come either
directly or indirectly from the plants grown at
Kew Royal Botanic Gardens over the past 40
years. Information provided by Kew indicates
that their material was provided by a private
collector, K. Beckett, who obtained it from the
National Tropical Botanical Garden (NTBG)in
Kaua‘i, Hawai‘i in 1972 (Pers. comm. from
NoeliaAlvares, KEW and Kava Vale, NTBG).The
source of the Hibiscus plant at NTBG was from
seed provided by noted United States Hibiscus
breeder, Ross Gast. In 1965 Gast had collected
seed of this Hibiscus in Naselai Village, near
Nakelo, Fiji (Gast 1980). Gast considered it
might be H. storckii, later referring to it as
‘Naselai pink’and indeed the accession at NTBG
is under that name. Examination of images and
specimens of ‘Naselai pink’from NTBG
(Hawai‘i) and KEW (UK), show that the plant
is synonymous with H. cooperi.
‘Byron Watkins’–a form with cream-white
petals, which may occur at low frequency in
the progeny from selfed ‘Naselai pink’plants
(Brian Kerr, pers. comm.). K. Beckett also
supplied the cream-white petaled form
to Kew Gardens under the name ‘Storckii
cream’(www.svenlandrein.com/virtherb
pages/19726091.html, accessed on 11/1/
2018). Further genetic research may indicate
introgression of genes from other Lilibiscus
species into one or more of these cultivars.
H. cooperi was one of the most widely used
species in the early Hawaiian Hibiscus breed-
ing programs, both as a male and female
parent. The species was reported to set self-
seeds and to cross readily with several species
in sect. Lilibiscus and with numerous cultivars
(Wilcox and Holt 1913). It is estimated here
that at least 47 of the cultivars documented by
Wilcox and Holt (1913) have H. cooperi in
their recent ancestry. Furthermore, our
morphological and DNA studies (unpublished
and Braglia et al. 2010) indicate that H. cooperi
is likely involved as a parent in many of today’s
widely grown Hibiscus varieties including
‘Lillian Wilder’/‘Apple Blossom,’‘Woodrow
Wilson’/‘Dawn,’‘Snow Queen,’‘Snow
Flake,’and ‘Rose Flake.’In cultivation, H.
cooperi readily produces seed without the need
for hand pollination and further research is
needed to understand its breeding system
more fully. Given its major ornamental
potential and its contribution to many widely
grown Hibiscus varieties, H. cooperi is highly
recommended for cultivation by botanical
gardens, hibiscus breeders, and enthusiasts.
Conservation status: The conservation status
is categorized as data deficient (IUCN 2016)
because H. cooperi is unknown in the wild,
although presumed to be extremely rare or
extinct. Its ex situ conservation status is well
assured through botanical gardens, including
through the efforts of the Botanic Gardens
Conservation International network.
KEY TO FIJI HIBISCUS SPECIES IN SECT.
LILIBISCUS (INCLUDING PRE-EUROPEAN
INTRODUCTIONS)
1. Flowers –single petaled (invariably five
petals) 2
Flowers –double-petaled, i.e. many petals
and petaloids in a tight formation, giving a
full ball-shaped appearance 7
2. Flowers±funnel shaped, spirally arranged
narrow petals < 3.3cm wide (at widest
point), style branches±erect. (Epicalyx
segments < 9 mm long; seedling, inter-
mediate and coppice regrowth including
trilobed leaves) H. bragliae sp. nov.
Flowers opening±flat, petals >3.3 cm
wide (at widest point), and later becoming
recurved during the course of the day, style
branches spreading to semierect 3
3. Leaves heterophyllous, invariably includ-
ing deeply trilobed leaves on mature
plants H. macverryi sp. nov.
116 PACIFIC SCIENCE •January 2019

Leaves entire or with a few large lobed
leaves on coppice/regrowth shoots 4
4. Petals >6.5cm long with dark red basal
zone >1.5cm radius. Seedling leaves
divided and intermediate leaves (including
coppice/regrowth) often with one to three
lobes 5
Petals < 6.5cm long with dark red basal
zone < 1.5cm radius. Seedlings and inter-
mediate leaves entire, never lobed 6
5. Stigma pads light yellow. Petals rose or
light pastel pink with white veins and trim,
or white petals (±flushed pale pink); lobate,
dark red basal zone±15–22 mm radius.
Leaves mainly lanceolate; petiole: leaf
length ratio 0.14–0.19. Leaf dentition:
Inner edge of the leaf teeth frequently at
±45°(to 90°) angle to the mid-rib H.
bennettii sp. nov.
Stigma pads red. Petals light lavender pink;
distinctive large, dark red serrated basal
zone±28–35 mm radius. Leaves lanceo-
late-elliptic; petiole: leaf length ratio 0.30-
–0.42. Leaf dentition: base of the leaf teeth
frequently±perpendicular (to 45°) to the
mid-rib H. storckii
6. Leaves lanceolate (L:W ratio 1.9–3),
margin entire or with irregular serrations,
typically 4–5 (range 0–12) teeth per
margin. Calyx lobes acute-elongated iso-
sceles triangular (L:W at base ratio 4.2–6.8,
lobes frequently remaining partly fused at
anthesis); calyx lobes extending beyond
mature fruit. Petals red, pink, or rarely
cream, with a small, lobate, dark red basal
zone, enclosed by a substantial white halo
and with white veins radiating into upper
portion of the petal H. cooperi
Leaves deltoid to broad lanceolate (L:W
ratio 1.2–1.7). Leaf dentitions±uniform,
typically 7–11 (range 5–12) teeth per
margin. Calyx lobes triangular (L:W at
base ratio 1.6–2); calyx lobes not extending
beyond mature fruit. Petals red or rose pink
with a small, ±serrated dark red basal zone,
±white petal veins (in “Waimea Beauty”)
H. rosa-sinensis
7. Crimson petals have whitish veins turning
pink near edge (±16–20petals/petaloids),
which decrease in size from the outer petals
(c. 4014 mm) toward the center of the
flower; functionally male H. bragliae sp.
nov.(flore pleno)
Deep pink or red petals and petaloids (±20
in number), with dark red basal zone,
extending along the floral tube, petaloids
decreasing in size toward the center of the
flower (Outer petals±50 13mm) H.
rosa-sinensis (flore pleno)
Striped dark pink and white/light yellow
petals and petaloids, (±20–25 in number)
H. rosa-sinensis (‘Baptistii’)
DISCUSSION
This study has resulted in the recognition of
four endemic Hibiscus species in sect. Lilibiscus
in Fiji including H. storckii, first collected in
1860, and the reinstatement of H. cooperi.
Hibiscus storckii was described in 1865 by
Seemann but subsumed under H. rosa-sinensis
by Smith (1981). The three new taxa, H.
bennettii,H. bragliae and H. macverryi, had each
been collected previously, but early collectors
had failed to appreciate that they represented
distinctive Fiji endemic taxa. The hitherto lack
of recognition of Fiji Lilibiscus species is mainly
attributable to their limited distributions in
difficult and inaccessible terrain and associated
limited collections, lack of rigorous morpho-
logical assessments, coupled with confusion
overtheidentityand circumscriptionof H. rosa-
sinensis.Thisstudy hasalso reinstatedH. cooperi,
a distinctive species whose description was
basedon plantedmaterialsoriginatingfrom the
South Pacific Islands. Hibiscus cooperi is here
considered more likely to have originated in
north Sulawesi, Indonesia and to have been an
ancient pre-European introduction into Mel-
anesia.
The existence of undescribed Fijian Lili-
biscus species had been hypothesized by
Hibiscus breeders such as the late Australian
Hibiscus breeder Geoff Harvey, who consid-
ered ‘Ruby Rose’(i.e. H. macverryi) and ‘Fijian
White’(i.e. H. bragliae form) to be unde-
scribed species from Fiji (pers. comm. to Gil
Bujanda, 2010). Likewise an AFLP study of
109 Hibiscus accessions found ‘Ruby Rose’and
‘Fiji Island’to cluster (Braglia et al. 2010),
being well-differentiated from other Lilibiscus
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 117

species and cultivars, and most similar to H.
cooperi, which was labeled in this study as H.
storckii. Nevertheless, the discovery of three
new species in Fiji was totally unexpected,
especially given the assertion of Smith (1981)
that Fiji had no native species in sect. Lilibiscus.
The Fiji archipelago, with four endemic
Lilibiscus species, can now be regarded a
secondary center of diversity in sect. Lilibiscus,
and with analogies to the Hawaiian archipelago
in the North Pacific that has nine endemic
species in sect. Lilibiscus (Huppman 2013).
Further DNA studies are needed to
elucidate more clearly the genetic and evolu-
tionary relationships among the Fijian species
and with other members of Lilibiscus, espe-
cially H. cooperi, H. rosa-sinensis and H. genevii;
the latter appearing to be the morphologically
closest ally of the Indian Ocean Island species
to H. bragliae. DNA studies are also needed to
ascertain the extent of genetic variation within
and between individuals and current levels of
inbreeding, in order to inform planning for
their conservation.
There is an urgent need for in situ
conservation measures for the four endemic
and critically endangered Fijian Hibiscus
species. Each species is facing increasing
threats from extreme climatic events, envir-
onmentally invasive weed species, hybridiza-
tion with introduced exotic Hibiscus species,
and their interactions. Prior to implementing
appropriate in situ conservation measures for
Fiji Hibiscus species, there is a need for more
comprehensive field surveys to better ascer-
tain their distribution and abundance. The
feasibility of using unmanned aerial vehicles,
or drones, for new field surveys ought to be
investigated given the often inaccessible and
treacherous nature of the habitats in which all
four Hibiscus species occur. It is also empha-
sized that Fiji Hibiscus field survey work, and
associated collections, and subsequent con-
servation actions will need to be both
informed by and involve the local Fijian
communities and traditional landowners.
The Polynesian kaute or ‘aute, which was
moved by Polynesians throughout the South
Pacific Islands, arguably represents a pure or
unhybridized form of H. rosa-sinensis. Alter-
natively there is a linguistic contention that H.
rosa-sinensis was given the name kaute in East
Polynesia—which originally meant Broussone-
tia papyrifera, and still does in many places—
and that this name was borrowed as this
hibiscus was spread westward West Polynesia,
then to Fiji and Rotuma, and to Polynesian
outliers in the Solomon Islands and Papua
New Guinea (Paul Geraghty, pers. comm.).
The possibility that “H. rosa-sinensis”didn’t
have its own original name in Eastern
Polynesia may suggest—contrary to the long
held-prevailing view—that H. rosa-sinensis, or
at least kaute/’aute, was not part of the original
plants brought into the Pacific by Polynesians.
Further genetic and field research in Eastern
Polynesia is needed to clarify the relationship
between kaute/’aute and H. rosa-sinensis from
Asia.
There is also a need to search for wild
populations of H. rosa-sinenis in South-east
Asia and Western Melanesia, especially
along the northern coastline of the island
of New Guinea. In particular the reports on
herbarium labels from botanical collections
of H. rosa-sinensis in Indonesia that suggest
H. rosa-sinensis is a wild species in West
Papua need to be followed up: these include
a report of it being common on the edge of
swamp forest, north of Simboro Strait, Lake
Sentani (R. S. Sigafoos 118, BO, 1945). The
earliest known specimen of H. cooperi appears
to be have been collected from Tondano,
North Sulawesi, Indonesia in 1840, and
accordingly Sulawesi ought to be the focus
for any future searches for wild populations
of H. cooperi.
ACKNOWLEDGMENTS
The authors are indebted to Professor David
Mabberley (Wadham College, University of
Oxford, UK and National Herbarium New
South Wales, Sydney, Australia) for his assis-
tanceandearlyguidanceinthiswork.Wewould
liketo sincerelyacknowledgethegeneroushelp
of the following people and organizations in
providing information, assistance, images,
access to botanical specimens and seeds: Noelia
Alvarez (Royal Botanic Gardens, Kew, UK),
Celine Babouline (New Caledonia), John
118 PACIFIC SCIENCE •January 2019

Bennett (Fiji), David Bush (CSIRO, Canberra,
Australia),Jean-FrançoisButaud(FrenchPoly-
nesia), Sam Chanel (Department of Forests,
Port Vila, Vanuatu), Gildas Gâteblé (Institut
Agronomique Néo-Calédonien, Mont-Dore,
New Caledonia), Paul Geraghty (University of
South Pacific, Fiji), Margaret Hanes (Eastern
Michigan University, United States), Elizabeth
Huppman (Lyon Arboretum, Hawai‘i, United
States), Angela Kay Kepler (Hawai‘i, United
States), Bruce Maslin (Western Australian
Herbarium, Perth/Singapore Botanic Gar-
dens), Gerald McCormack (Cook Islands
Natural Heritage Trust), Alivereti Naikatini
(IAS/USP South Pacific Regional Herbarium),
Chigira Osamu (Forest Tree Breeding Center,
Okinawa, Japan), Amato Pauliasi (Fiji), Nicolas
Perrault (McGill University, Department of
Biology, Canada), Paul Quek (Malaysia), David
Robbins (Royal Botanic Gardens, Melbourne,
Australia), Kenton Sanchez (United States),
Myknee Sirikolo (National Herbarium and
Botanical Garden, Ministry of Forestry and
Research, Honiara, Solomon Islands), Paul
Smith (Botanic Gardens Conservation Inter-
national, UK), Randolph Thaman (University
of South Pacific, Fiji), Kava Vale (National
Tropical Botanical Garden, Kaua‘i, Hawai‘i,
United States), Tong Wang (map illustrations;
Melbourne, Australia), herbaria (Australian
National Herbarium, Canberra, Museum
National d’Histoire Naturelle, Paris, France;
Herbarium Bogoriense, Indonesia; Naturalis
Biodiversity Center, Leiden, Netherlands;
Auckland War Memorial Museum, New Zeal-
and; Solomon Islands National Herbarium,
Honiara; Conservatoire et Jardin Botaniques,
Geneva, Switzerland; Kew Gardens and Nat-
ural History Museum London, UK; Bishop
Museum, New York Botanical Garden and
Harvard University Herbaria, United States),
and hibiscus experts and enthusiasts from
around the globe for their valued inputs and
considered thoughts including Gil Bujanda
(United States), Jill Coryell (Hawai‘i, United
States), Ortwin Heckl (Canary Islands, Spain),
Colleen and Geoff Keena (Australia), Oliver
Joshua (Vanuatu), Brian Kerr (Australia),
Ursula Lengdobler (Germany), Wally Morgan
(Australia),DavidOrr(Hawai‘i, UnitedStates),
Richard Thomson (Fiji), Robin Yarrow (Fiji)
and Father Michael McVerry (Fiji).
The Mohammed bin Zayed Species Conserva-
tion Fund through a grant to Botanic Gardens
Conservation International has been support-
ing the “Survey, inventory and conservation of
Hibiscus storckii, a highly threatened Fijian
species.”The assistance and support of this
fund and BGCI/ Paul Smith and Fiji colla-
borators Nunia Moko and Siteri Tikoca
(Nature Fiji Mareqeti Viti) and Marika Tui-
wawa (IAS/USP South Pacific Regional Her-
barium) is most gratefully acknowledged in the
support of the research undertaken in prepara-
tion of this manuscript.
Literature Cited
Anon. 1863. Hibiscus cooperi (as nom nud.) by
“G.A.”on 15 September 1863. J. Hort.
Cottage Gdnr. Vol. V (new series): 213.
Ash, J. 1987. Stunted cloud-forest in Taveuni,
Fiji. Pac. Sci. 41:191–199.
Bates, D. M. 1965. Notes on the
cultivated Malvaceae. 1. Hibiscus.Baileya
13 97–131.
Braglia, L., S. Bruna, S. Lanteri, A. Mercuri,
and E. Portis. 2010. An AFLP-based
assessment of the genetic diversity within
Hibiscus rosa-sinensis and its place within the
Hibiscus genus complex. Sci. Hortic.
123:372–378.
Bull, W. 1872. Special list of magnificent new
and choice varieties of Gladiolus and new
plants. William Bull. New Plant Merchant,
Chelsea, London.
Cafferty S., and M. Cheek. 1996. Hibiscus fragilis
Malvaceae. Curtis’sBot.Mag.13:210–214.
Cambie, R. C., and J. Ash. 1984. Fijian
medicinal plants. CSIRO, Australia.
Carr,D.J.ed.1983.SydneyParkinson,artist
Cook’sEndeavour voyagepl.73p.79,col.pl.;
Bacstrom.S.Ms.:102.Solander,D.Pl.
Otaheit.: 2 Hibiscus Rosa Sinensis;Solander,D.
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 119

Slip Catalogue XIV: 693–695; Banks, J. Cat.
Pl.: 27. Natural History Museum, London.
Coiffier, C. 2012. L’hibiscus rosa–sinensis,
Représentation de la division dualiste du
cosmos Iatmul en Papouasie Nouvelle-
Guinée (Hibiscus rosa-sinensis, a representa-
tion of the dualistic division of the Iatmul
cosmos in Papua New Guinea). J. des
Anthropol. 128–129:115–138.
Cronin, S. 1999. Volcanic hazard and risk
assessment for Taveuni, Fiji Islands. SOPAC
TechnicalReport298.South PacificApplied
Geoscience Commission, Suva.
Florence, J. 2004. Hibiscus rosa-sinensis. Pages
210–211 in Flore de la Polynésie française
Vol. 2. IRD Éditions, Publications Scien-
tifiques du Muséum, Paris.
Gast, R. H. 1980. Hibiscus around the world.
Letters to J.W. Staniford from Ross H.
Gast. American Hibiscus Society, Florida.
Giraud, J. F. 2013. Merveilleux Hibiscus
(Wonderful Hibiscus). Éditions Orphie,
Saint Denis, Reunion Island.
Hochreutiner, B. P. G. 1900. Révision du
genre Hibiscus (Revision of the genus
Hibiscus). Conserv. Jard. Bot. Genéve
Annuaire 4:23–191.
Hoyt, C. 1950. A study of synonymy in
Chinese Hibiscus in Florida. Masters thesis.
University of Florida, Gainesville.
Huppman, E. R. H. 2013. Analysis of
relationships among endemic Hawaiian
Hibiscus. Dissertation submitted to the
graduate division of the University of
Hawai‘iatMānoa in partial fulfillment of
the requirements for the degree of Doctor
of Philosopy in Horticulture, Hawai‘i.
International Union for Conservation of
Nature, IUCN. 2016. Guidelines for using
the IUCN red list categories and criteria.
Version 12. Prepared by the Standards and
Petitions Subcommittee, IUCN Gland,
Switzerland.
Jarvis, C. E. 2007. Chapter 6. Sources of
information: Collectors, suppliers of pro-
pagules, intermediaries and recipients. Page
194 in C. E. Jarvis, Order out of chaos:
Linnaean plant names and their types. The
Linnean Society of London in association
withthe NaturalHistory Museum,London.
Koopman, M. M., and D. Baum. 2008.
Phylogeny and biogeography of tribe
Hibisceae (Malvaceae) on Madagascar. Syst.
Bot. 33: 364–374.
Lawton, B. P. 2014. Hibiscus: Hardy and
tropical plants for the garden. Timber
Press, Portland, Oregon.
Layard, J. 1942. Stone men of Malekula.
Chatto and Windus, London.
Lemaire, C. A. 1864. Ibiscus cooperii, ketmie de
Cooper.L’Illustration horticole (illustration
by P. Stroobant) 11: Pl. 412. Imprimerie et
lithographie de F. et E. Gyselnyck, Gand,
Belgium.
Linnaeus, C. 1753. Species Plantarum. Vol. 1,
p. 694. Laurentius Salvius, Stockholm.
Loureiro, J. D. 1788. Fl. Cochinch I: 419.
Lindley, J. 1858. Report on Crystal Palace
Horticultural Exhibition. Page 439 in The
Garden Chronicle and Agricultural Gaz-
ette, London.
Meehan, T. ed. 1863. Hibiscus cooperi in Gard.
Monthly & Hort. Advertiser 5:313. Brinck-
loe, Philadelphia.
Mettler, R., and E. Otto. 1865. Neue
allgemeine deutsche Garten- und Blumen-
zeitung (New general German garden and
flower newspaper) 21:74–75.
Nakamura, T., and N. Nomoto. 1985. The
cytological studies in family Malvaceae, II.
Chromosome numbers and karyotypes in
genus Hibiscus. Chromosome Inf. Serv.
39:36–37.
Nalepa, A. 1906. Ueber zwei neve Eriophyi-
den von den Fidschiinseln (About two new
Eriophyidae from the Fiji Islands) J. Econ.
Biol. 1:147–151.
Nunn, P. D. 2007. Climate, environment and
society in the Pacific during the last
millennium. Elsevier, Amsterdam.
Pfeil, B. E., and M. D. Crisp. 2005. What
to do with Hibiscus? A proposed nomen-
clatural resolution for a large and well-
known genus in Malvaceae and com-
ments on paraphyly. Aust. Syst. Bot.
18:49–60.
Phillips, R. H. 1990. Hibiscus storckii—fact or
fiction—A non-botanist’s view. Paper pre-
sented to the Second International Aus-
tralian Hibiscus Society Inc. Convention
120 PACIFIC SCIENCE •January 2019

28 April to 3 May, 1990. Brisbane,
Queensland.
Rageau, J. 1973. Plantes medicinales de La
Nouvelle-Calédonie, Office de la
recherche scientifique et technique outre-
mer (ORSTOM), Paris.
Rivers-Smith, B. 2004. A Fijian Hibiscus
odyssey—Some facts and history relating
to Hibiscus in Fiji. Hibiscus International: 4
(I).(Jan–March 2004).
Sampath, P. 2014. The origins of Hibiscus rosa-
sinensis (also known as the shoe flower).
Hibiscus International: 14 (1), Issue 58:
16–19.
Seemann, B. 1861. Plantae Vitienses in
Bonplandia 9:254.
—. 1862. Viti: An account of a government
mission to the Vitian or Fijian Islands in
the years 1860–61. Cambridge, UK.(Rep-
rinted in 1973, with an introduction by P.
A. Snow, Colonial History Series No. 85).
—. 1865. Flora Vitiensis: A description of the
plants of the Viti or Fiji Islands, with an
account of their history, uses and properties.
Vol. 1 p. 17. L. Reeve and Co, London.
—. 1873. Flora Vitiensis: A description of the
plants of the Viti or Fiji Islands, with an
account of their history, uses and properties.
Vol. 2 Plate 4. L. Reeve and Co, London.
Smith, A. C. 1981. Hibiscus rosa-sinensis. Pages
419–420 in Flora Vitiensis Nova—A new
flora of Fiji. Vol. 2. National Trop. Bot.
Garden, Kaua‘i, Hawai‘i.
Skovsted, A. 1941. Chromosome numbers in
the Malvaceae II. Comptes Rendus Des-
travaux Laboratoire Carlsberg. Ser. Phy-
siol. 23:195–240.
Stearns, B. P., and S. C. Stearns. 2000.
Watching, from the edge of extinction.
Yale University Press, New Haven, CT.
Sykes, W. R. 1970. Contributions to the flora
of Niue. New Zealand Department of
Scientific and Industrial Research Bulletin
200, Wellington, NZ.
Tal, M. 1980. Physiology of polyploids. Pages
61–76 in W.H. Lewis (ed.). Proceedings of
the International Conference on Poly-
ploidy: Biological Relevance held at
Washington University, St Louis, Missouri,
May 24–27, 1979. Plenum Press, New York.
Tuiwawa, M., and S. Pene. 2014. A biodiversity
assessment, socioeconomic study and
archaeological survey of the Greater Delai-
koro Area, Vanua Levu. Report compiled by
the Institute of Applied Science, University
of the South Pacific for the GEF-PAS
project “Proposed protection of the Greater
Delaikoro Area.”Suva.
Van Borssum Waalkes, J. 1966. Malesian
Malvaceae revised. Blumea 14:1–251.
Van Houtte, L. 1864. Hibiscus cooperi in “Flore
des serres et des jardin de l’Europe: annales
générales d’horticulture”(Plants of green-
houses and gardens of Europe). 15 July
1864. 15: 109.
Weiner, M. A., 1984. Secrets of Fijian
medicine. Publisher unstated.
Whelan, P. M., J. B. Gill, E. Kollman, R. A.
Duncan,and R.E.Drake.1985.Radiometric
dating of magmatic stages in Fiji. Pages
415–440 in D. W. Scholl, T. L. Vallier (eds.)
GeologyandoffshoreresourcesofthePacific
Island arcs—Tonga region. Circum-Pacific
Council for Energy and Mineral Resources,
Earth Science Series, 2. Houston.
Whistler, W. A.1990.EthnobotanyoftheCook
Islands: The plants, their Maori names, and
their uses. Allertonia, 5:347–424.
———. 1991. The ethnobotany of Tonga:
The plants, their Tongan names and their
uses. Bishop Museum Bulletin in Botany
2:1–155.
———. 1992. Polynesian herbal medicine.
National Trop. Bot. Garden, Kaua‘i,
Hawai‘i.
———. 2000. Plants in Samoan culture. The
ethnobotany of Samoa. Isle Botanica,
Hawai‘i.
WHO. 2009. Medicinal plants in Papua New
Guinea.WorldHealthOrganizationRegio-
nal Office for the Western Pacific, Manila.
Wilcox, E. V., and V. S. Holt. 1913.
Ornamental Hibiscus in Hawaii. Bulletin
No. 29. Hawai‘i Agricultural Experiment
Station. Paradise of the Pacific Press,
Honolulu, Hawai‘i.
Fiji Hibiscus Species in Section Lilibiscus •Thomson and Braglia 121