History, Biology, and Conservation of Pacic Endemics.

1. The Royal Creeper, Oxera pulchella (Lamiaceae),

a New Caledonian Ornamental Plant1
Gildas Gtebl 2
Abstract: A thorough review of Oxera pulchella Labill., also called the royal
creeper, was carried out to better understand the amazing story of this almost
forgotten New Caledonian ornamental species. The history of this plant is interestingly related to the history of New Caledonia. The literature records analysis
shows that taxonomic placement of the genus Oxera Labill. has been widely discussed from its discovery until very recently. This review also offers the opportunity to understand and hypothetize how Oxera pulchella was exported and disseminated worldwide through botanical gardens and private nurseries at the end
of the nineteenth century. Along with a review of Oxera pulchellas etymology,
vernacular names, iconography, and horticultural fame, this review considers its
current taxonomy, horticultural importance, conservation, biology, and various
economic issues with a perspective from our ongoing research on this genus of
special ornamental potential.
The creeper Oxera pulchella Labill. is native
to New Caledonia. This species was once well
known in Europe as an ornamental plant, and
its history is interestingly linked to the history
of New Caledonia. It was discovered to the
wider world by the rst explorers landing in
New Caledonia, and it appears in the rst accounts of the ora of New Caledonia (Figure
1). Later, its history has been linked to the
rst settlements in Nouma and with the rst
New Caledonian horticulturists, botanists,
and gardeners. Oxera pulchella was one of the
rst native New Caledonian ornamental
plants to be exported to the Old World, and it
is associated with some of the earliest and
most famous plant hunters and botanic gardens who were on the lookout for attractive
owering plants. Oxera pulchella acquired a
1
This work was partly supported by a grant (11026051-D) from the Ministre charg de lOutre-Mer.
Manuscript accepted 7 December 2011.
2
Institut Agronomique no-Caldonien, Station de
Recherche Agronomique de Saint Louis, BP 711, 98810
Mont Dore, New Caledonia (e-mail: gateble@iac.nc).

degree of horticultural fame in Europe at the

end of the nineteenth century. Since it was
rst described, O. pulchella has been at the
center of a long debate regarding its taxonomic afliations. Meanwhile, in its homeland, it was nearly forgotten, it was rarely used
ornamentally, and it was rapidly losing its
limited natural habitats on the Noumean peninsula to urban developments. More recently,
one subspecies, O. pulchella subsp. grandiora,
has been protected in two New Caledonian
provinces. It now has International Union for
Conservation of Nature status and belongs to
a genus of considerable ornamental interest.
In this paper I review the taxonomic history
of O. pulchella from the family level to the
infraspecic level, discussing its etymology,
detailing the major historical events from its
discovery to its international dissemination,
summarizing its horticultural history, and
briey discussing its conservation, biology, and
a number of issues of economic importance.
taxonomic history
Above Species Level

Pacic Science (2012), vol. 66, no. 4:413 433

doi: 10.2984/66.4.1
2012 by University of Hawaii Press
All rights reserved

The taxonomic placement of the genus Oxera

has provoked numerous articles from the time
of its original description to the present day.

413

414

PACIFIC SCIENCE October 2012

Figure 1. Habit of a owering branch with inorescence and cluster of two fruiting calyces of Oxera pulchella subsp.
pulchella, drawn by Laurence Ramon.

De Labillardire (1824) did not have fruits

available so was unable to place the genus
properly, writing that Oxera probably lay
somewhere between Vitex and the Bignoniaceae. De Kok and Mabberley (1999a) provided a good taxonomic history of the turbulent
generic and intrageneric delimitation of the
genus Oxera. In 1999, Oxera lost its New
Caledonian endemic status when an Oxera
species was described in Vanuatu (de Kok and
Mabberley 1999b). Now, Oxera is generally
recognized as being in the Lamiaceae family
(Steane et al. 2004), but this position could
evolve still further because relationships
among many families in Lamiales, and to a
certain extent also their limits, are still unclear (APG III 2009:117). The infrageneric
taxonomy was revised recently by de Kok
(1997) based on a comprehensive cladistic
analysis. De Kok and Mabberley (1999a, b)

and Mabberley and de Kok (2004) then divided the genus into ve informal groups.
Oxera pulchella was inserted in the pulchella
group along with four other species. The
pulchella group also nds support from a
chemotaxonomic study carried out on the
amounts of avonoids and verbascosides
(Grayer and de Kok 1998).
Below Species Level
Some taxonomic changes for Oxera pulchella
have also occurred at the infraspecic level
(Figure 2). Based on ower morphology differences (corolla and calyx), Dubard (1906)
created three new varieties (O. pulchella var.
grandiora Dubard, O. pulchella var. deplancheana Dubard, and O. pulchella var. microcalyx
Dubard) and described a new species closely
related to the latter variety, O. balansae

Pacic Endemic: Royal Creeper, Oxera pulchella Gtebl

Dubard. Guillaumin (1933) added yet another variety to Dubards three, O. pulchella var.
sinuata Guillaumin. According to Guillaumin
(1933), this variety is intermediate between
the O. pulchella type and O. pulchella var. grandiora. Moldenke (1975a) also described O.
pulchella var. brevicalyx Moldenke as a new variety having a shorter calyx. The same year,
Moldenke (1975b) pointed out a new New
Caledonian taxon, O. pulchella var. brevifolia
Moldenke, but was certainly mistaken in citing his own description. Later, Moldenke
and Moldenke (1983) pointed out that O.
pulchella Dubard ex Moldenke is a synonym
of O. pulchella Labill. because Moldenke
(1940) put Oxera pulchella Labill. as the correct name instead of Oxera pulchella Dubard.
Finally, de Kok and Mabberley (1999b) simplied matters by distinguishing two subspecies within the O. pulchella species: O. pulchella
Labill. subsp. pulchella and O. pulchella Labill.
subsp. grandiora (Dubard) de Kok. In their
synopsis, de Kok and Mabberley (1999b)
combined the two varieties of O. pulchella, O.
pulchella var. brevicalyx and O. pulchella var.
microcalyx, to the level of species under Oxera
brevicalyx (Moldenke) de Kok. The three
other varieties, O. pulchella var. grandiora, O.
pulchella var. sinuata, and O. pulchella var. deplancheana, were combined under the new
combination O. pulchella subsp. grandiora
(Figure 2).
Taxonomic Discussion
The new genus organization and the one
within the pulchella informal group claried
the relationships between the species (de Kok
and Mabberley 1999b, Mabberley and de
Kok 2004). Although the key they provided
for the pulchella group is usually helpful,
there are still some remaining issues to be
resolved to help eld botanists and horticulturists to properly identify the different taxa.
Indeed, although O. morierei Vieill. and O.
crassifolia Virot are easily distinguished within
the pulchella group, it is more difcult to
distinguish between some other taxa, for example:
(1) Oxera balansae and O. brevicalyx if seen
outside their natural environments, because

415

their natural distribution patterns are clearly

disjunct although closely related [O. brevicalyx
(Moldenke) de Kok is the former O. pulchella
var. microcalyx Dubard] according to Dubard
(1906).
(2) Oxera balansae and O. pulchella subsp.
grandiora are sympatric taxa in the Nouma
peninsula. Leprdours islet specimens are
sometimes classied under O. balansae (Mabberley and de Kok 2004) and sometimes under
O. pulchella (Guillaumin 1911, Jaffr et al.
2008).
(3) The two O. pulchella subspecies, especially for plants growing in the Npoui and
Pindai peninsulas (MacMilan 5041, 1952, P)
where they are sympatric with O. brevicalyx.
Indeed, except in this case, O. pulchella subsp.
pulchella is a taxon with a thick or eshy calyx
and growing at medium to high altitude in
humid forests, and O. pulchella subsp. grandiora is a taxon having a thinner calyx and restricted to dry forests at low altitude in the
Nouma peninsula. According to Mabberley
and de Kok (2004), Moldenke (1970) even
wrongly illustrated O. pulchella with a gure
(photograph 283) of O. brevicalyx.
Therefore, more work is required to clearly
understand the infra- and inter-relationships
of the aforementioned four taxa. In a previous
hybridization study, Gtebl et al. (2009) have
shown that there was good homogeneity in
this pulchella group, at least for those four
taxa. From our results with some other crosses, and in hindsight, we have observed that
our F1 putative hybrids are fertile and can
give rise to F2 progenies. We have also observed a wide range of variation, especially in
ower and calyx sizes within the F1 putative
hybrids (Figure 3). At one point, we even
wondered whether O. pulchella subsp. grandiora might be a naturally occurring polyploid
living in sympatry with O. balansae and /or O.
brevicalyx. After some preliminary cytological
work on O. brevicalyx and O. pulchella pollen
mother cells in which we were not able to precisely determine their chromosome numbers
because they were quite numerous (2n = 56
to 64) and because they were relatively small
in size (1 to 2 m), we decided that hypothesis was not sustainable (Klein and Gtebl
2010).

Figure 2. Diagrammatic representation of the taxonomic relationships of Oxera pulchella and its close relatives.

Figure 3. Flower and calyx variations of both Oxera pulchella subspecies compared with closely related species and with
putative hybrid progenies (scale bar = 2 cm).

418

etymology and vernacular names

In the literature, neither native nor local
French vernacular names have been recorded
for Oxera pulchella, but there are several native
names for the closely related species Oxera balansae Dubard (Mabberley and de Kok 2004).
Indeed, as a creeper vine, stems of O. balansae
are traditionally used to tie crab claws in Lifou
( J. Ounmoa, pers. comm.) and to tie many
other things in Mar ( N. Robert, pers.
comm.). To repair this oblivion, in 2001,
the dry forest program (Programme de Conservation des forts sches) organized a school
contest in which children invented vernacular
names for several species. The name selected
for O. pulchella was liane perruche, given by
M. Song (Papineau 2005), which can be translated into English as parakeet creeper. In
his account of A Voyage in Search of La Prouse,
and before publishing the rst account of the
New Caledonian ora, de Labillardires
translator (1802) wrote, in the second volume
of the English translation, of some new Bignoniae, of which one is called trumpet ower in the index of the rst volume. Trumpet
ower could therefore be said to be the rst
vernacular name given to O. pulchella even if
it was a bit over-translated. Indeed, in the
original French version, de Labillardire
(1799 1800) wrote about some bignones,
but there is no index and no reference to a
French translation of trumpet ower. In his
Sertum Austro-Caledonicum, de Labillardire
(1824) did not mention any vernacular names
but explained the signicance of Oxera as
coming from a Greek word related to propter
corolla fauce dilatatam in Latin, which can be
translated (Gafot 1934) as peculiar to the
swollen throat of the corolla. According to
Lemaire (1867), de Labillardires translation
of the greek is incorrect as it should
be written Oncera instead of Oxera. This
Greek and de Labillardires original thought
and translation were then followed by Don
(1838) and by de Kok and Mabberley (1999a).
However, for most authors, Oxera was a
Greek name derived (depending on the author) from acetosus, oxeros, oxus, or oxys, meaning sour, dubious, tasting or smelling like
vinegar (Pfeiffer 1874, Nicholson 1889, Rodi-

PACIFIC SCIENCE October 2012

gas 1889, Bennett 1901, Bailey and Miller

1906, Fraser and Hemsley 1916, Burras and
Grifths 1994, Suprin 1996). De Kok and
Mabberley (1999a) also explained that the
term Oncoma as used by Sprengel (1827) was a
Greek word synonym of Oxera. Oncoma is
also a medical term meaning swelling or tumor, a genus of hemipterous insects (Oncoma
Fieb.), and a molluscan genus (Oncoma Mayer)
synonym of Strombus Linn. The epithet pulchella, from the Latin meaning beautiful or
pretty, used by de Labillardire to describe his
new species anticipated the later ornamental
value of this plant. After its original description, several vernacular names were given to
O. pulchella in different languages: Oxera lgant in French (Poiret 1825) translated into
Osera elegante in Italian by Correzioni
(Poiret 1846), Oxra in French (Boiste 1843),
Oxre lgante in French (Dominguez 1846),
Oxre trs belle in French (Drapiez 1853),
Pretty Oxera in English (Guilfoyle 1883),
Oxre jolie in French (Duchartre 1887), Oxera mignon in French (Rodigas 1889), Oxera
gentil in French (Bellair and Saint-Lger
1900), and Royal climber and Showy Oxera in
English (Moldenke 1970), which was nally
translated into French as Liane royale (Mabberley and de Kok 2004).
major historical events
Discovery
Oxera pulchella was rst discovered in 1793 by
a French naturalist, J.-J. H. de Labillardire,
assisted by the naturalist L. A. Deschamps
and the gardener F. Delahaie, during the
dEntrecasteaux expedition in search for La
Prouses ships. Some 20 yr later, de Labillardire (1824) published his Sertum AustroCaledonicum and briey described the new
genus Oxera Labill. along with a single new
species, Oxera pulchella Labill. Although de
Labillardire did not mention where he collected his type specimen, he almost certainly
collected it during one of his inland explorations around Balade (northeast of Grande
Terre) and probably in the back mountains
during a 3-day expedition from 3 May to 5
May 1793 (rst year of the Republic, Floral
14 to 16). De Labillardire (1802, 2:244 245)

Pacic Endemic: Royal Creeper, Oxera pulchella Gtebl

wrote that I here observed a great number of

plants which I had not yet found in any of the
locations I had made on this island. They belonged for the most part to the family of the
Proteae and to that of the Bignoniae. Although
Proteaceae are quite well represented in the
New Caledonian ora, native Bignoniaceae
are not. Apart from O. pulchella, only Deplanchea speciosa Vieill. and Pandorea pandorana
(Andrews) Steenis might match this family at
that elevation and place. The rst specimen of
O. pulchella was then probably collected somewhere between what is now called Col dAmos
and Mount Mandjlia, where the species still
occurs (see specimens of McPherson 3774,
1981, P and McKee 33086, 1977, P). These
specimens (de Labillardires and the two
others) are now placed in O. pulchella Labill.
subsp. pulchella. The discussion in the following paragraphs focuses on a taxon now placed
in O. pulchella subsp. grandiora (Dubard) de
Kok.
Export
In the Labiatae treatment in the Flora of New
Caledonia, Mabberley and de Kok (2004)
studied 31 specimens of Oxera pulchella subsp.
grandiora collected by 18 botanists. Nearly
half of those specimens (14) were collected,
mostly in the Nouma (Port de France) peninsula, by 10 botanists (Balansa, Baudouin,
Brousmische, Deplanche, Germain, Mueller,
Pancher, Veitels, and Vieillard) at the end of
the nineteenth century. The rst literature
record found of a cultivated O. pulchella in
New Caledonia is in the Extracts from the
journal of Mr. John Gould Veitch, during a
trip to the Australian colonies and the South
Sea Islands published by his father ( Veitch
1866) and briey translated by Naudin (1866).
J. G. Veitch, as a collector of ne plants for
Veitchian nurseries, immediately spotted a
plant called O. pulchella in the gardens of Government House directed by J. A. I. Pancher.
He thought that this species loaded with
bunches of white owers would prove a valuable conservatory plant in England and that
the three other species of Oxera were less
striking ( Veitch 1866:291). J. G. Veitch arrived in Nouma on 30 September 1865 and

419

left a week later with the HMS Curacoa expedition. During his stay in New Caledonia,
Veitch also visited the Model Farm at Yahou
along with the naturalist Brenchley (1873)
and the ofcer Foljambe (1868) but did not
mention any species of Oxera there. From the
literature survey, although Veitch collected
many plants in the South Sea islands during
that trip I could not nd any record that he
took samples (cuttings and /or seeds) of O. pulchella. At that time, the living plants collected
were packed in Wardian cases and sent to
Veitch Nurserys customers in Australia and
some were sent directly to the Veitch nurseries in Chelsea, England, with the HMS Falcon, which was in Nouma at the same time
(Heriz-Smith 1990). Back in Sydney, Veitch
also exhibited his fresh and healthy plants,
collected during the trip, at a Sydney Botanical Garden grand ower show on 18 October
1865 (Foljambe 1868) but still without any
reference to O. pulchella. Later on, in the famous Hortus Veitchii, J. H. Veitch (1906) described several plants collected in New Caledonia by J. G. Veitch, but still there was no
reference to O. pulchella. In fact, those plants
were collected on previous visits to other
South Sea islands, which is why Hortus Veitchii
gave the wrong origin as New Caledonia for
several plants described in the catalogue
(Heriz-Smith 1990). From Heriz-Smiths
point of view (1990), J. G. Veitch was unable
to collect at all in New Caledonia due to social commitments such as cricket games between the French and English warship teams!
However, this was not Panchers opinion
(1871 1873), who gave living plants of Aralia
tenuifolia Pancher [Schefera veitchii (Carrire)
Frodin & Lowry] to M. Weight (i.e.,
Veitch), who managed to bring them back in
good shape. J. G. Veitch is apparently not the
rst exporter of a living plant of O. pulchella
to Australia or England. W. R. Guilfoyle, an
Australian landscape gardener, nurseryman,
and botanist, could have been the rst exporter of O. pulchella even if he had already possessed this species before his trip to New
Caledonia (Guilfoyle 1866). This was because
on his way to the Yahou Model Farm, he
spotted the yellow owers of Oxera pulchella
among the conspicuous plants he saw (Guil-

420

foyle 1869:134). Indeed, during his botanical

expedition among the South Sea islands on
board HMS Challenger, Guilfoyle collected
living plants and also made a quick stop in
Nouma in 1868 (Morat 2010).
Among other early horticulturists and /or
naturalists who sent or could have sent living
plants from New Caledonia to Australia,
France, and other parts of the world at the
end of the nineteenth century (deduced from
Morat [2010]), it is worth mentioning J. Paddon (settler who sent living plants and seeds
to Sydney before 1861), E. Deplanche (plant
collector between 1855 and 1867), E. Vieillard ( botanist and horticulturist in New Caledonia between 1855 and 1867), J. A. Boutan
(agronomist and Yahou Model Farm founder), D. Cooper (collector of seeds for Veitchian nurseries around 1860), Landman (a
policeman who sent ferns for the Great London Exhibition in 1862), E. de Greslan (settler from 1863 who sent seeds), Veitels (O.
pulchella collector in 1866), A. Baudouin (plant
collector between 1867 and 1868), E. M.
Heckel (scientist who sent seeds between
1867 and 1869), B. Balansa (Director of
Nouma acclimatation garden between 1868
and 1872), F. A. Campbell (collector during
an expedition in February March 1873), J.
Caldwell (collector of native sugarcane plants
in 1868), E. L. Layard (honorary British Consul and naturalist in New Caledonia during
the 1870s who sent living plants to Melbourne), F. Jensen (plant collector in 1876),
R. Germain (plant collector beween 1875 and
1879), A. Hughan (Australian photographer
and collector around 1880), A. Brousmiche
(plant collector between 1881 and 1884), F.
Gervais (sent seeds around 1885 1886), V.
Perret (sent seeds in 1885 1886), E. Hodgson
(collector in New Caledonia in 1886 and
1887), E. F.-A. Raoul (sent living plants to
Sydney in 1889), J. Dybowski (colonial agricultural inspector), Moriceau (sent seeds), L.
Gauharou (sent seeds), C. de Touzalin (sent
seeds), and J. Bernier and L. Cribs (collectors
for the Paris Universal Exposition in 1900).
According to Morat (2010), the following
botanists and /or horticulturists were doubtful
collectors of plants in New Caledonia and are
thus unlikely to have been responsible for ex-

PACIFIC SCIENCE October 2012

ports of Oxera pulchella from New Caledonia:

F. J. von Mueller (Director of Melbourne
Botanic Garden, cited as a O. pulchella subsp.
grandiora collector in Mabberley and de Kok
[2004]), C. Moore (Director of Sydney Botanical Garden), R. Schomburgk (Adelaide
Botanical Garden), T. and T. W. Sheperd
(horticulturists in Sydney), and J. F. Roberts
(horticulturist in Melbourne). Some of these
might however have helped to disseminate O.
pulchella, at least within Australia.
The most probable rst and major exporter of living Oxera pulchella plants or plant
parts (seeds, rooted cuttings) is Jean Armand
Isidore Pancher, who was a botanist and gardener in New Caledonia for the French Museum National dHistoire Naturelle between
1857 and 1869. He went back to France after
retirement but returned to New Caledonia in
1874 with de Maerschalk on duty for the renewed Belgian Linden horticulture and acclimatation company. From 1875 to his death in
1877, Pancher sent several cases with living
plants from New Caledonia to Belgium (Sagot
1879). Among the plants sent to Belgium
by Pancher, Sagot (1879) made mention of
Oxera robusta Vieill. but not of O. pulchella.
Later on, Rodigas (1889) wrote that O. pulchella was rst cultivated and introduced in
Linden houses at Ghent in 1875 directly from
New Caledonia through Pancher and de
Maerschalk. Sagot (1879) also stated that
Pancher sent several living plants from New
Caledonia to Australia and from New Caledonia to France and de Rothschild houses in
1859 and 1860 (Figure 4).
World Dissemination of O. pulchella
Having reviewed how and by whom O. pulchella was probably exported from New Caledonia, I now discuss its ethelochory (deliberate dispersal as a garden plant). It is interesting
that O. pulchellas dissemination was inuenced by either small or powerful private
nurseries, by plant hunters, and by British and
French colonial empires within their tropical
botanic gardens distributed around the world.
Figure 4 is an attempt to illustrate the dissemination of O. pulchella during the second
half of the nineteenth century.

Pacic Endemic: Royal Creeper, Oxera pulchella Gtebl

421

Figure 4. Ethelochory of Oxera pulchella during the second half of the nineteenth century. Solid arrows show documented routes of introduction, dashed arrows show probable routes of introduction. 1, Sagot (1879); 2, Baptist (1864),
Guilfoyle (1866), Graham (1874), Moore (1895); 3, Abbott (1869); 4, Andr (1874), Dufresne (1874); 5, Rodigas
(1889); 6, dEprmesnil (1878); 7, Schomburgk (1878); 8, Cantley (1880); 9, Anonymous (1888a); 10, Ross (1888); 11,
Carrire (1890), Guillaumin (1921a, b, 1952); 12, Guilfoyle (1883); 13, Bailey (1885); 14, Saul (1891), Bailey (1892),
Bailey and Miller (1906); 15, Hart (1897); 16, Willis (1898); 17, Stewart (1900); 18, Mabberley and de Kok (2004).

oceania: Vanuatu: Mabberley and de

Kok (2004) and de Kok and Mabberley
(1999b) hypothesized that Oxera pulchella
subsp. pulchella was introduced to Vanuatu
long ago. This hypothesis was based on a
specimen ( Veillon 2940, nou) collected at
Dillon Bay, Erromango, and a personal communication of J.-M. Veillon. According to my
interpretation of Veillon 2940, the specimen is
more related to Oxera balansae (see discussion
in the section on Taxonomic History), a species found in the Loyalty Islands of New
Caledonia, which are about 300 km from
Erromango. This plant may have arrived on
Erromango through natural long-distance
dispersal.
Australia: It is surprising that O. pulchella
appeared in the literature as a cultivated plant
for the rst time in Sydney, Australia, in Baptist and Sons nursery catalogue in 1864 (Baptist 1864). Soon after it also appeared in Guilfoyles (1866) Sydney nursery catalogue 2 yr
before Guilfoyles trip (1869) to New Caledonia. Oxera pulchella then appeared in another

private nursery in Sydney along with Dammara moorii [Agathis moorei (Lindl.) Mast.],
another New Caledonian species (Graham
1874). Within Australia, the other records of
living O. pulchella were in botanical gardens in
Tasmania in 1868 (Abbott 1869), in Adelaide
in 1878 (Schomburgk 1878), in Melbourne in
1883 (Guilfoyle 1883), in Brisbane in 1885
(Bailey 1885), and nally in Sydney in 1895
(Moore 1895). A more thorough unpublished
literature review of Sydney Botanical Garden
directed by C. Moore, who had several collectors in New Caledonia, might prove that O.
pulchella was rst cultivated in Sydney Botanical Garden before being distributed (Figure
4) to private nurseries in Sydney and to the
other botanical gardens (Tasmania, Adelaide,
Melbourne, and Brisbane).
africa: Algeria: Dufresne (1874) and
Andr (1874) wrote about O. pulchella as a
handsome plant that owered every year in
the gardens of Hamma, Algiers, Algeria (a
French colony at that time). No literature record was found about that early introduction

422

to Algeria, but the gardens of Hamma were, at

that time, a famous acclimatation and testing
garden. I hypothesize that O. pulchella was introduced to Algiers in 1859 or 1860 during a
port call by Admiral J. Saisset, who ordered
Pancher to prepare some living plants before
going back to France (Sagot 1879). However,
O. pulchella was still not listed as a plant to be
sold by the Hamma gardens in 1865 1866
(Anonymous 1865). Anyway, Jarry-Desloges
(1940) was wrong in saying that O. pulchella
was introduced for the rst time in the
Hamma gardens in 1883. N. Mauri (1937),
the Hamma gardens head of propagators, was
surprisingly also wrong in saying that O. pulchella was introduced in his garden in 1886.
Mauritius: In 1880, Oxera pulchella was
under cultivation in the Royal Botanical Gardens of Mauritius, Jardin des Pamplemousses
(Cantley 1880). I suppose that it was introduced through British botanic gardens exchanges and probably through the Sydney
Botanical Garden (Figure 4).
Other African countries: According to
Verdcourt (1992), O. pulchella was found in
nurseries at Amani in Tanzania in 1948 and
might also have been in Kenya.
europe: France and Belgium: Sagot
(1879) stated that living plants were sent to
France and de Rothschild houses in 1859 and
1860 by Pancher, but there is no proof that O.
pulchella was among them (Figure 4). The rst
reliable literature record found in Europe for
the successful introduction of O. pulchella is
Rodigas (1889), who said that it was rst introduced in Ghent by Linden through Pancher and de Maerschalk in 1875. A short time
later, between 1876 and 1878, O. pulchella
was introduced to the Cannes acclimatation
gardens to evaluate its ornamental potential
(dEprmesnil 1878). Carrire (1890), Guillaumin (1921a, b, 1952), and Jarry-Desloges
(1940) agreed that O. pulchella was cultivated
and spread in France by V. Lemoine, horticulturist in Nancy, from an introduction from
the Hamma gardens. However they disagreed
about the date of introduction: 1885 (1888 in
Lemoines horticultural catalogue) for Carrire (1890), 1883 for Guillaumin (1921a, b,
1952), and Bellair and Saint-Lger (1900)
stated 1886 for Europe (Figure 4).

PACIFIC SCIENCE October 2012

United Kingdom of Great Britain and Ireland: There was a great deal written about the
rst introduction of O. pulchella to England at
the end of the 1880s. According to Watson
(1888) and Anonymous (1890), Sir George
Macleay introduced it directly from New
Caledonia by seeds, but F. Ross (1888), who
was Sir George Macleays gardener, said that
it was introduced from the Hamma gardens in
Algiers by Macleay, who had gone on a visit
there 7 yr earlier (1881). This latter story is
the most reliable (Figure 4). I could not nd
precisely when O. pulchella was introduced
under cultivation in the Royal Botanic Gardens, Kew, but it was there before 1896 (Hart
1897). Anonymous (1888a) also wrote about
unfortunate attempts to acclimatize O. pulchella at Fota Island, Cork, Ireland, after a trip
to Algiers in 1880 (Figure 4).
Other European countries: Oxera pulchella
was probably rst introduced to other European countries through the Lemoine and /or
Clibran nurseries. For example, Berger (1912)
referred to the presence of this plant in Italy,
and Moldenke (1976) referred to it in Germany.
america: In North America, the rst introduction of O. pulchella was probably at
John Sauls nurseries (Maple Grove Farms) in
Washington, D.C., in 1891 (Figure 4), perhaps via England (Saul 1891, Bailey 1892,
Bailey and Miller 1906). A few years later, it
was recorded in California (Stewart 1900).
Wilson (1938) and Moldenke (1959) conrmed that it was still under cultivation in
California.
In South America, Hart (1897) reported
that O. pulchella was introduced into the Royal
Botanic Gardens of Trinidad in 1896 from
the Royal Botanic Gardens, Kew (Figure 4).
asia: Oxera pulchella was probably introduced in Asia through the Royal Botanic Gardens of Ceylon (Sri Lanka) at the end of the
nineteenth century (Figure 4). Indeed, Willis,
the director of the gardens, put O. pulchella on
his Desiderata list in 1898 ( Willis 1898). Oxera pulchella was then seen under cultivation in
Ceylon by Macmillan (1914) and Moldenke
(1974).
Later on, Moldenke and Moldenke (1983)
said Oxera pulchella is now widely cultivated

Pacic Endemic: Royal Creeper, Oxera pulchella Gtebl

for ornament in both hemispheres. Even

though O. pulchella was distributed widely
around the world before and during the twentieth century, its current distribution remains
quite restricted, usually being found only in
specialized nurseries such as Lancasters
(Lancaster 1999) and known only to keen amateur gardeners and collectors.
horticultural history
General Issues
The rst recorded mention of the culture
of Oxera pulchella dates back to 1838. Don
(1838) explained that the culture of Oxera is
to be related to that of Jacaranda Juss. species. This reference to the culture and propagation of O. pulchella is doubtful because
there is no recorded herbarium specimen between that of de Labillardire in 1793 and
that of Vieillard between 1855 and 1860
and no other references during that period
(1793 1855). During the 1870s and based
on the experience of the Hamma gardens in
Algiers, several references advised that O.
pulchella should be cultivated outdoors in the
south of France or under protection in the
Paris climate (Andr 1874, 1878, 1879, Dufresne 1874). Anonymous (1886) commented
on the culture and recent owering of O.
pulchella at Pendell Court, Bletchingley, Surrey, in December 1886. This rst owering
date was later the center of a debate between
Anonymous (1905a, b) and Kennedy (1905).
Hooker (1887) then published a brief description and a concise plate under tab. 6938
in Curtiss Botanical Magazine, from a Pendell Court specimen. Hookers paper was
followed and cited by many others (Anonymous 1887, 1888b, Duchartre 1887, Bois
1891, for example). Popular and dramatic descriptions were used at that time, such as
gem, will be one of the nest new plants of
the year, the most graceful and beautiful
ower we have seen, and one of the handsomest stove plants recently gured (Anonymous 1887, 1888c, d ). F. Ross, Sir George
Macleays gardener, even received a FirstClass Certicate from the Royal Horticultural oral committee for his success in culti-

423

vating and owering O. pulchella (Anonymous

1888e, 1892, Morris and Wilks 1889). Oldeld Nurseries, Manchester, then sold plants
of O. pulchella by placing at least 26 advertisements in the Gardeners Chronicle during the years 1889 and 1890 under titles
such as Stove plants, greenhouses plants,
Gems, gems, gems, of choice stove plants,
and Clibrans gems of greenhouse and stove
(Clibran 1889a, b, 1890). This Oxera pulchella fever born in England soon spread,
at least in the literature, such as in France
(Carrire 1890, van den Heede 1890, Le Beau
1891), in Germany (Beck and Abel 1889,
1891), in Denmark (Helweg 1889 cited in Bay
1890), in Switzerland (Haasis 1890), in the
United States (Goldring 1888, Bailey 1892),
and in Canada (Saul 1891). Many other
papers were written to describe the best propagation and cultivation techniques for O.
pulchella, mainly from F. Ross primary experience (Anonymous 1888f, g, 1891a, b, Daveau 1893 1899, Watson 1896, Wright 1902).
Hudson (1891) and Plantsman (1891) suggested an interesting use for O. pulchella as a
stove climber, well suited to training upon the
roofs of stoves to give some shade to other
plants.
After this bullish period from 1887 to the
end of the nineteenth century, Oxera pulchella
was gradually almost forgotten and the literature becomes comparatively scarce during the twentieth century ( Weathers and
Worthington-Williams 1910, Watson 1915,
Battandier et al. 1914, Bailey 1928, Jex-Blake
1950, Hoyt 1958, Brilmayer 1962, Van der
Spuy 1976, Macmillan 1989, Graf 1992, Lancaster 1999, Reich 1999, 2010). Cathey (1977)
even suggested that O. pulchella was suitable as
a bonsai. The main cultivation issues discussed regarding O. pulchella were its frost resistance and its owering time.
Frost Resistance
Since the rst attempts to cultivate O. pulchella in Europe and North Africa, its frost resistance has always been a major topic of conversation. Rivire (1882) stated that leaves and
young outdoor plants were sometimes frost
damaged in the winter in the Algiers climate.

424

For Anonymous (1888b), O. pulchella required

the warmth of a stove or intermediate house
in Great Britains climate, and Anonymous
(1888f ) said that a minimum temperature
of 12C 15C is needed for owering. Beck
and Abel (1893) suggested that O. pulchella
can withstand temperatures of 5C 8C.
Later, O. pulchella was cultivated outdoors
with more or less success (for example, in a
cliff garden of St Lawrence, Isle of Wight,
in southern England [Fitzherbert 1905]; in
Vintimiglia, Italy [Berger 1912]; in southern
California [Miller and Bailey 1919]; and in
southern France [ Jarry-Desloges 1940]). For
Mathias (1982), O. pulchella was less sensitive
to cold than Petrea L. or Mandevilla Lindl. but
is still touchy to frost, and Graf (1992) and
Llamas (2003) said that it is a tropical climber
that can withstand hardiness zones 10 or
10 11.
Flowering Time
In New Caledonia, O. pulchella subsp. grandiora owers naturally during the cool season
from June July to August September. Most
herbarium specimens deposited at nou and
having owers were collected during that
period. In the Northern Hemisphere, most
of the literature also indicates that owering
is during the European (Anonymous 1886,
Hooker 1887, Carrire 1890), North African
(Dufresne 1874, Rivire 1912), or Asian
(Moldenke and Moldenke 1983) winter periods, though Rivire (1889) talked about spring
owering in Algeria. Anonymous (1891a)
advised that for good owering O. pulchella
subsp. grandiora should be exposed to full
light and to a minimum of 15C, which are
conditions similar to Noumas climate in
July August. Very few Northern Hemisphere
authors mention owering times outside their
winters. For Mathias (1982) owering time
was extremely variable, from early spring
to summer to autumn, and Lady Plowden
entered some owers in a competition in
England in mid-August (Anonymous 1905c).
From earlier trials under controlled-light
conditions, it seems that O. pulchella subsp.
grandiora could be a short-day plant (Lemay
and Gtebl 2008).

PACIFIC SCIENCE October 2012

Illustrations
De Labillardires (1824) rst botanical iconography of the species, tab. 28, somewhat
redrawn with other New Caledonian plants,
page 415, in Charton (1863) included O. pulchella subsp. pulchella (sensu de Kok and Mabberley 1999b). The rst published plate of O.
pulchella subsp. grandiora is from Curtiss
Botanical Magazine, tab. 6938 (Hooker 1887).
Following this rst plate, others were soon
published in other horticultural magazines,
such as plate 651 drawn by H. G. Moon in
The Garden ( Watson 1888) somewhat redrawn in Mo llers Deutsche Ga rtner-Zeitung
(Credner 1888), g. 34 in The Gardeners
Chronicle (Anonymous 1888h), g. 15 in the
Journal of Horticulture and Cottage Gardener (Anonymous 1888f ) later republished by
Anonymous (1895) and English (1907), plate
LXXVI in LIllustration Horticole (Rodigas
1889), g. 80 in the Revue Horticole (Carrire 1890), g. 474 in Les plantes de serre
(Bellair and Saint-Lger 1900), and plate 148
in Wild owers of the world (Everard and
Morley 1974). The rst account of a published photographic engraving seems to be
the one of Miss Wilmott from Warley Place
in The Garden (Anonymous 1894, 1895). I
publish here a new iconography account of
the less-known O. pulchella subsp. pulchella,
drawn by Laurence Ramon (Figure 1).
some issues of biology, conservation,
and economy
Biology
Both subspecies of Oxera pulchella grow exclusively on nonserpentine soils (de Kok 2002),
though the species has previously been
wrongly listed on serpentine soils (Heckel
1892, Jeanneney 1894, Morat et al. 1986).
According to de Kok (2002), the pulchella
subspecies is restricted to soils derived from
schists, and the grandiora subspecies can be
found either on calcareous or schist-derived
soils. Apart from one specimen (McMilan
5041, see discussion in the Taxonomic History section), O. pulchella subsp. grandiora is
restricted to dry forests at low altitude, and O.
pulchella subsp. pulchella is restricted to humid

Pacic Endemic: Royal Creeper, Oxera pulchella Gtebl

forests generally between 160 and 600 m (de

Kok and Mabberley 1999b, Jaffr et al. 2001,
Mabberley and de Kok 2004).
In preliminary work on pollination biology, de Kok (1997, 2001) concluded that Oxera
pulchella and the other taxa belonging to the
informal pulchella group were likely to be
pollinated by birds such as honeyeaters (Phylidonyris undulata) even if some introduced
honey bees (Apis mellifera) are sometimes seen
in the owers. In gardens in New Caledonia,
gray-eared honeyeaters (Lichmera incana ssp.
incana) are often seen collecting nectar from
the owers of both O. pulchella subspecies by
perforating the base of the corolla. This behavior is unlikely to pollinate the owers.
Along with local buttery and moth enthusiasts ( J.-J. Gervolino and T. Salesnes), we suspect that moths play a role in the pollination
of O. pulchella. De Kok (1997) hypothesized
that there was a shift from buttery pollination to bird pollination and even a reverse
shift for species of the neriifolia group. The
pollination biology of O. pulchella requires
further eld observation, especially to determine if the pollination vectors differ between
the two subspecies.
The moderately eshy fruits of O. pulchella
actively attract birds so putative hybrid fruits
under study must be protected with nets. The
dissemination of O. pulchella seeds is likely
then to be by an endozoochory mechanism
if large birds such as Ptilinopus greyii are
involved (Tassin et al. 2010) or by a shortdistance zoochory mechanism (consumption
of only the eshy part of the fruit either on
the plant or a few meters away) if smaller birds
such as Zosterops spp. are involved as was reported for Oxera sulfurea (Boissenin et al.
2006; F. Desmoulins, pers. comm.). Further
studies on seed dissemination by birds in O.
pulchella would be of great interest to increase
our understanding of the distribution patterns
of the two subspecies.
Further work is ongoing on seed desiccation tolerance of O. pulchella and its relatives.
According to our preliminary observations,
the seeds of O. pulchella are recalcitrant, and it
would be useful to characterize its seed desiccation tolerance more precisely to learn if
desiccation tolerance correlates with ecosys-

425

tem (rain forest for pulchella subspecies and

dry forest for grandiora subspecies) and /or
with species. The observed sensitivity to desiccation of O. pulchella seeds supports the idea
that worldwide dissemination during the second half of the nineteenth century was carried
out using rooted cuttings or young plants.
Conservation
Conservation awareness appeared only during
the 1990s for O. pulchella subsp. grandiora,
when Bouchet et al. (1995) argued in favor of
an urgent need for protecting sclerophyll forests. From that time until very recently, the
town of Nouma was expanding, and more
and more patches of dry-forest relicts (Kamr, Montravel, Tina, and Ouen Toro, for
example) were cleared for building. Despite
the formal protection of relictual patches
of dry forest on the Nouma peninsula, the
development of spontaneous settlements or
shacks with associated agroforestery systems continues in some dry forests ( Nouville, Montravel, Tina). Last-chance transplantation of natural mature plants was
sometimes performed before clearing (Anonymous 1996, Suprin 1996, Lemay and Gtebl 2008). De Kok (1998) suggested that O.
pulchella subsp. grandiora should be classied
EN (Endangered) according to IUCN Red
List criteria because of the urban expansion
threat. Morat et al. (2001) then suggested
CR (Critically Endangered) status, and Hequet (2007) recommended VU ( Vulnerable)
status. According to the online IUCN Red
List (Hequet 2010), O. pulchella subsp. grandiora is now assessed as VU [A2c; B1ab(iii) +
2ab(iii)], and O. pulchella subsp. pulchella is
under LC (Least Concern) status. According to local legislation, O. pulchella is also a
protected species in the Northern Province
(Anonymous 2008) and in the Southern Province (Anonymous 2009). Though O. pulchella subsp. grandiora is formally restricted
to the Nouma peninsula (Mabberley and de
Kok 2004), some very closely related plants
are found outside Nouma, such as on Leprdour islet, on Npoui-Pinda peninsula (250
km from Nouma), and in a Pata dry forest.
Further research on pollination biology,

426

PACIFIC SCIENCE October 2012

dissemination mechanisms, and genetic diversity is a necessary prerequisite for an optimal

conservation plan for the O. pulchella subsp.
grandiora subspecies.
Economy
Suprin (1996) and de Kok (1998) suggested
bringing O. pulchella subsp. grandiora under
cultivation in New Caledonia for its ornamental features. However, like most native
plants, O. pulchella subsp. grandiora remained
almost unknown to the New Caledonian gardener until the setting up of a research and
development program aimed at promoting
and domesticating the ornamental native ora
(Gtebl 2009a, b). In partnership with the
Dry Forest Project, a technical sheet with
propagation, cultivation tips, and uses was
written on O. pulchella subsp. grandiora
(Gtebl et al. 2006). This technical sheet and
mother plants were given to growers so that
private nurseries could produce and sell their
own plants. With the help of the Dry Forest
Project, extension services, the landscaping
association, and growers, the home gardener
can now buy pot plants of this species. This
type of product is still marginal in the New
Caledonian ornamental horticulture industry,
but its popularity is expected to rise. From an
amelioration research program point of view,
the Oxera genus is of great interest and a main
concern, because its theoretical potential as
an ornamental is high (Gtebl 2010). Other
ways of promoting O. pulchella economically
are also conceivable. A team from the University of New Caledonia recently found an
interesting brown color as a natural dye extracted from O. pulchella leaves (Hnawia et al.
2010). Biological activities of avonoids and
verbascosides found in the species (Grayer
and de Kok 1998) may also offer some opportunities.

New Caledonia was the original supplier of

this biological resource. Academic and practical work is still needed to improve the
knowledge and attractiveness of the species.
A combination of morphological analyses,
cross-pollination studies, isozyme analyses,
cytometric approaches, and molecular insights should greatly clarify the specic and
infraspecic taxa delimitations of O. pulchella
and determine its closest relatives. These
sorts of studies may also help to bring characters of interest known in other Oxera species
(e.g., fragrance, color range, plant architecture, multiple owering) into modern O. pulchella progenies.
acknowledgments
For a South Sea islander, this comprehensive bibliography review would have not been
possible without the use of the Biodiversity
Heritage Library Web site. For bibliographic
research, I am also very grateful to Rogier
de Kok from the Royal Botanic Gardens,
Kew; Muriel Beros from Astredhor; Lu Anne
Schwarz from the Atlanta Botanical Garden;
Marsha Winter from the University of the
West Indies; Pascale Perret from Socit Nationale de Protection de la Nature; Mary Fruneau from Socit Nantaise dHorticulture;
Helen Carey from Country Life magazine;
Danielle Saintpierre from Programme Fort
Sche; Fiona Patrick from Cornell University; Vanessa Hequet and Jacqueline FambartTinel from NOU; and Peter Butcher. For
designing Figure 4, I thank Armelle Tardivel.
I am also indebted to Nigel Heriz-Smith,
Jean-Marie Veillon, and Philippe Morat for
some related discussions; to Caradoc Doy for
arguing about Veitchs history; and to two
anonymous reviewers for constructive comments.
Literature Cited

closing comments
Through this comprehensive review article
telling the story of Oxera pulchella and summarizing the major aspects of the species,
I hope to revive interest in this almostforgotten gem and to remind ourselves that

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