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Biology and Impacts of Pacific Island Invasive Species. 11. Cinchona pubescens (Red Quinine Tree) (Rubiaceae)

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Cinchona pubescens Vahl (red quinine) is an evergreen tree ranging in height from 10 to 25 m with broad leaves and white or pink fragrant flowers arranged in clusters. Growing at altitudes between 130 and 3,300 m, it is one of 23 species in the genus Cinchona and has a natural distribution from Costa Rica to Bolivia. Cinchona pubescens has been cultivated in tropical regions (e.g., in South America, Africa, China, India, and Indonesia) for its quinine-containing bark and has become invasive in some regions. This is especially the case in the Pacific region, where C. pubescens has invaded humid highland areas of Galápagos, Hawai‘i, and Tahiti. It shades out and reduces cover of native plant species and adversely affects endemic birds. In addition, it changes microclimate and nutrient cycling in the soil, especially phosphorus, in Galápagos. Characteristics that make it such a successful invader include production of numerous, windborne seeds and vigorous vegetative reproduction by resprouting from underground stems and fallen trees. In Galápagos, C. pubescens is currently being manually controlled by uprooting the trees and by applying herbicides to cuts in the bark. However, this method requires continuous hand pulling of seedlings to be successful. Disturbance by control actions appears to facilitate establishment and invasion by other nonnative plant species, especially blackberry (Rubus niveus). Quinine and other alkaloids extracted from Cinchona bark are still being used for medicinal purposes today and the wood is increasingly used as construction material in Galápagos. Ironically, C. pubescens is now considered rare and endangered in its native range in Ecuador.
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Pacific Science (2015), vol. 69, no. 2:133 153
doi:10.2984/69.2.1
© 2015 by University of Hawai‘i Press
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133
Cinchona L. is likely the most commercially
important genus of the family Rubiaceae
after the genus Coffea L. due to its quinine-
containing bark. Cinchona pubescens Vahl (syn.
C. succirubra Pav. ex Klotzsch, 1857) (red qui-
nine tree) has been introduced worldwide,
mainly from Bolivia, Colombia, Ecuador, and
Peru (Andersson 1998) (Figure 1), and is con-
sidered among the 100 worst in
vaders glob-
ally (
ISSG 2015). It is highly invasive on
several Pacific island groups, especially Galá-
pagos, Hawai‘i, and the Society Islands
(
Weber 2003), where it forms dense stands
in the humid highland vegetation, reducing
plant species diversity and abundance. The
invaded islands are Santa Cruz in Galápagos
( Ja¨ger et al. 2009); the islands of Hawai‘i,
Maui, and O‘ahu in Hawai‘i (Starr et al. 2003);
and Tahiti in the Society Islands (
Meyer
2000). The introduction of the novel “tree”
life-form to a formerly treeless environment
in Galápagos led to sig nificant changes in
vegetation stand structure and changes in
light, water, and nutrient regimes ( Ja¨ger et al.
2007, 2009, 2013) (Figures 2 and 3). Con-
sequently, C. pubescens is now considered an
ecosystem engineer in Galápagos ( Jones
et al. 1994, Ja¨ger et al. 2009). In Hawai‘i,
Biology and Impacts of Pacific Island Invasive Species. 11. Cinchona
pubescens ( Red Quinine Tree) ( Rubiaceae)1
Heinke Ja¨ger
2,3
Abstract: Cinchona pubescens Vahl (red quinine) is an evergreen tree ranging in
height from 10 to 25 m with broad leaves and white or pink fragrant flowers ar-
ranged in clusters. Growing at altitudes between 130 and 3,300 m, it is one of 23
species in the genus Cinchona and has a natural distribution from Costa Rica to
Bolivia. Cinchona pubescens has been cultivated in tropical regions (e.g., in South
America, Africa, China, India, and Indonesia) for its quinine-containing bark
and has become invasive in some regions. This is especially the case in the Pa-
cific region, where C. pubescens has invaded humid highland areas of Galápagos,
Hawai‘i, and Tahiti. It shades out and reduces cover of native plant species and
adversely affects endemic birds. In addition, it changes microclimate and nutri-
ent cycling in the soil, especially phosphorus, in Galápagos. Characteristics that
make it such a successful invader include production of numerous, windborne
seeds and vigorous vegetative reproduction by resprouting from underground
stems and fallen trees. In Galápagos, C. pubescens is currently being manually
controlled by uprooting the trees and by applying herbicides to cuts in the bark.
However, this method requires continuous hand pulling of seedlings to be suc-
cessful. Disturbance by control actions appears to facilitate establishment and
invasion by other nonnative plant species, especially blackberry (Rubus niveus).
Quinine and other alkaloids extracted from Cinchona bark are still being used
for medicinal purposes today and the wood is increasingly used as construction
material in Galápagos. Ironically, C. pubescens is now considered rare and en-
dangered in its native range in Ecuador.
1 This work was financially supported by a Marie
Curie International Outgoing Fellowship within the
7th European Community Framework Programme, the
Deutsche Forschungsgemeinschaft (DFG), the Charles
Darwin Foundation, and the A. F. W. Schimper Stiftung.
Manuscript accepted 19 June 2014.
2 Department of Ecology, Technische Universita¨t
Berlin, Rothenburgstr. 12, 12165 Berlin, Germany.
3 Current address: Charles Darwin Foundation,
Puerto Ayora, Santa Cruz, Galápagos, Ecuador.
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C. pubescens has invaded native understory
and forest canopy vegetation, as well as non-
native forests and disturbed areas, where it
reduces the cover of other plant species (Starr
et al. 2003, Fischer et al. 2009) (Figure 4). In
Tahiti, C. pubescens has invaded secondary and
primary rain forests and un disturbed cloud
forest (Meyer 2004, Vanquin 2006) (Figure 4).
This review synthesizes information on
the ecology, impact, and management of C.
pubescens worldwide, which is partly published
in gray literature and therefore not easily
accessible.
name
Cinchona pubescens Vahl
Phylum Angiospermae, class Magnoliop-
sida, order Gentianales, family Rubiaceae,
tribe Cinchoneae, genus Cinchona L.
Synonyms: C. asperifolia Wedd., C. calop-
tera Miq., C. chomeliana Wedd., C. colorata
Laubert ex B. D. Jacks., C. cordifolia Mutis, C.
cordifolia var. macrocarpa Wedd. ex Howard,
C. cordifolia var. microcarpa Howard, C. cordifo-
lia var. peruviana Howard, C. cordifolia var. ro-
Figure 1. Natural distribution of Cinchona pubescens
(modified after Andersson [1998], Ja¨ger [2011]: Cinchona
pubescens, Enzyklopa¨die der Holzgewa¨chse, Vol. 58,
14 pp. Copyright Wiley-VCH Verlag GmbH & Co.
KGaA. Reproduced with permission).
Figure 2. Growth form of Cinchona pubescens in the fern-sedge vegetation on Santa Cruz Island, Galápagos (note in-
vasive Psidium guajava trees in the surroundings) (
H. Ja¨ger).
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tundifolia (Pav. ex Lamb.) Wedd., C. cordifolia
var. vera Wedd., C. coronulata Miq., C. decur-
rentifolia Pav. in Howard, C. delondriana
Wedd., C. discolor Hayne, C. elliptica Wedd.,
C. erythroderma Wedd., C. goudotiana Klotzsch
ex Triana, C. govana Miq., C. grandifolia Ruiz
& Pav., C. hirsuta Ruiz & Pav., C. lechleriana
Schltdl., C. howardiana Kuntze, C. lutea Pav.
in Howard, C. microphylla Mutis ex Lamb., C.
morada Ruiz, C. obovata Pav. ex Howard, C.
ovata Ruiz & Pav., C. ovata var. erythroderma
Wedd., C. ovata var. rufinervis Wedd., C. ovata
var. vulgaris Wedd., C. palescens Vell., C.
pallescens Vitman, C. pallescens var. ovata (Ruiz
& Pav.) Howard, C. pelalba Pav. ex DC., C.
pelletieriana Wedd., C. peruviana Howard, C.
platyphylla Wedd., C. pubescens var. cordata
DC., C. pubescens var. ovata (Ruiz & Pav.)
DC., C. pubescens var. pelletieriana Wedd., C.
pubescens var. purpurea (Ruiz & Pav.) Wedd.,
C. purpurascens Wedd., C. purpurea Ruiz &
Pav., C. purpurea Vell., C. rosulenta Howard ex
Wedd., C. rotundifolia Pav. ex Lamb., C. rubi-
cunda Tafalla ex Wedd., C. rufinervis Wedd.,
C. rugosa Pav. ex DC., C. scrobiculata Humb. &
Bonpl., C. scrobiculata var. delondriana Wedd.,
C. scrobiculata var. genuina Wedd., C. subcor-
data Pav. ex Howard, C. subsessilis Miq., C. suc-
cirubra Pav. ex Klotzsch, C. succirubra var. con-
glomerata Howard, C. succirubra var. cuchicara
Howard, C. succirubra var. erythroderma How-
ard, C. succirubra var. spruceana Howard, C.
succirubra var. vera Howard, C. tenuis Ruiz ex
DC., C. tucujensis H. Karst., C. viridiflora Pav.
ex Howard, Quinquina obovata (Pav. ex How-
ard) Kuntze, Q. ovata (Ruiz & Pav.) Kuntze,
Q. pubescens (
Vahl) Kuntze, Q. succirubra
(Pav. ex Klotzsch) Kuntze (Andersson 1998,
Grandtner 2005, IPNI 2005).
Common names: English: red cinchona,
red quinine tree, Peruvian bark, Jesuits’ bark,
Countess bark; Dutch: kinaboom; French:
quinine, quinquina rouge; German: Chi-
narinde, Chinarindenbaum; Spanish: casca-
rilla, quina, quinina.
Cinchona pubescens can be easily confused
with C. antioquiae L. Andersson, C. barbacoen-
sis H. Karst, C. capuli L. Andersson, and
Figure 3. Invasion of Cinchona pubescens in fern-sedge vegetation on Santa Cruz Island, Galápagos ( H. Ja¨ger).
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C. micrantha Ruiz ex. Pav. (Andersson 1998).
However, none of these is found in the Pacific
islands.
The genus Cinchona was named after the
Countess of Chinchón, wife of the Viceroy of
Peru, by Linnaeus in 1742. A confusion over
Spanish, Latin, and Italian orthography re-
sulted in the loss of the first “h” in the genus
name, and “Cinchona” became established
(Keeble 1997). According to a well-cited leg-
end, the countess was cured of malaria by hav-
ing been administered the bark of Cinchona in
1638 after all other remedies failed (
Hill 1952,
Prendergast and Dolley 2001), and it was not
until 1930 that this story was proven to be
false (Keeble 1997).
Due to its medical importance, Cinchona
has historically attracted a lot of attention
from taxonomists. During the nineteenth
century, many different names were estab-
lished based on minor morphological differ-
ences (Popenoe 1949, Harling and Andersson
1994). The taxonomy of C. pubescens is espe-
cially complicated because it frequently hy-
bridizes with other Cinchona species (Acosta
Solís 1945b, Camp 1949, Harling and Anders-
son 1994, Andersson 1998), especially with C.
barbacoensis H. Karst., C. calisaya Wedd., C.
lancifolia Mutis, C. lucumifolia Pav. ex Lindl.,
C. macrocalyx Pav. ex DC., C. micrantha
Ruiz ex. Pav., and C. officinalis L. (Andersson
1998).
description and account of variation
Species Description
Cinchona pubescens is an evergreen tree, 10
25 m tall, with a diameter at breast height
(DBH
) of 20
80 cm in Ecuador (Acosta Solís
1945b, Andersson 1998) and 30 m tall with a
DBH of 90 cm in northern Peru (
Hodge
1948). In Ecuador, the trees usually have a
main trunk that branches in the upper third
Figure 4. Habitats invaded by Cinchona pubescens. Left, Acacia koa forests in the Makawao Forest Reserve on Maui,
Hawai‘i (L. Fischer); right, Atara Mountain on
Tahiti ( J.-Y. Meyer).
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Pacific Island Invasive Species: Cinchona pubescens, Red Quinine Tree · Ja¨ger 137
forming a semicircular-shaped crown (Acosta
Solís 1945b). In its introduced range in Galá-
pagos and Tahiti, trees can reach a height of
up to 15 m and a DBH of 10
30 cm (Shimizu
1997, Vanquin 2006, Ja¨ger 2011). The maxi-
mum height of trees measured in the Makawao
Forest Reserve on Maui, Hawai‘i, was 16 m
(Fischer 2007).
The leaves are opposite, broad and mem-
branous, 8.3
23 cm long and 5.3
21 cm wide,
ovate to suborbicular with pubescent petioles
(1.2
5 cm long) and stipules that can be cadu-
cous (Andersson 1998) (Figure 5). Leaf blades
have 7
11 pairs of prominent secondary veins
and are commonly hairy beneath, especially
the young ones. Leaf domatia are usually ab-
sent, but if present they are pouch-shaped
(Andersson 1998).
The inflorescence of C. pubescens is usually
large, with broad panicles up to 20 cm and
sometimes longer (Figure 5). Flower corollas
are pinkish or purplish, paler at base (corollas
outside may be white to light pink or red in
Hawai‘i, Tahiti, and Galápagos), and are
fragrant (Andersson 1998, Starr et al. 2003;
J.-Y. Meyer, pers. comm., 2013) (Figure 5).
The corolla tube is 9
14 mm long, glabrous
inside and pubescent outside. The seed cap-
sules are ellipsoid to subcylindrical and 13 41
by 5
7 mm long, opening from the base to tip
when mature (Figure 5). Seeds are 7
12 by
2.1
2.8 mm, light, and have dentate wings
(Andersson 1998) (Figure 5).
Distinguishing Features
A conspicuous feature of C. pubescens is that its
senescing leaves turn yellow or bright red.
Cinchona pubescens also possesses unique
growth characteristics in its Pacific island
distribution. In Galápagos, it produces a
main trunk but also often develops several
trunks a short distance away, which emerge
by suckering of underground stems (see Del
Tredici 2001). This way, C. pubescens trees
take on a multistemmed growth form, with
the individual stems connected below the
surface (Shimizu 1997) (Figure 6). In addi-
tion, it resprouts from fallen and cut stems to
produce vertical shoots (Macdonald et al.
1988). Contrary to its growth form in main-
land Ecuador, trees produce branches on the
lower third of the trunk, which are bent
upward, resulting in a semicircular to cylin-
drical crown in the Galápagos (Figure 6). The
same is true for trees in the Makawao Forest
Reserve on Maui (L. Fischer, pers. comm.,
2014).
Intraspecific Variation
The leaf morphology is variable, especially
with respect to size, shape, and hairs of the
leaf blades. Reasons for this variation could
include local differentiation and clinal varia-
tion in response to altitude (von Humboldt
1808, Acosta Solís 1945b), as well as hybrid-
ization with other Cinchona species (Anders-
son 1998). To my knowledge, no work on the
potential genetic variation within C. pubescens
has been carried out. However, because the
species is so widespread and hybridizes with
other species, it is an important topic for
investigation.
Figure 5. Cinchona pubescens morphology. Top left, inflo-
rescence and leaves on Santa Cruz Island, Galápagos (
H.
Ja¨ger); top right, fruits from the Taravao plateau, Tahiti
( J.-Y. Meyer); bottom left, flowers on Santa Cruz Island,
Galápagos (
H. Ja¨ger); bottom right, seeds on Santa Cruz
Island, Galápagos (cm scale, two long vertical bars is
1 cm) (F. Bungartz).
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Illustrations
In addition to the figures provided in this re-
view, a good cross section of images is avail-
able on the following Web sites:
Photos from Hawai’i by Kim and Forest
Starr: http://www.starrenvironmental.com
/
images/species/?q=cinchona+pubescens&o
=plants
Global Invasive Species Database: http://
www.issg.org/database/species/ecology.asp?si
=63&fr=1&sts=sss&lang=EN, Feb. 2015
CABI Bioscience Invasive Species Com-
pendium, Datasheet Cinchona pubescens http://
www.cabi.org/isc
/datasheet
/13484
economic importance and
environmental impact
Detrimental Aspects
Studies carried out on C. pubescens in different
Pacific island ecosystems generally indicate
detrimental impacts on the native vegetation.
Common negative effects included the reduc-
tion of species abundance and diversity. In
Hawai‘i, C. pubescens outcompetes other spe-
cies in dense Dicranopteris linearis vegetation
zones, as well as in mesic and wet forests, es-
pecially Acacia koa forests in the Makawao
Forest Reserve on Maui (Starr et al. 2003)
(Figure 4). The herb and shrub cover in this
reserve were also reduced in invaded areas
(Fischer et al. 2009). In Tahiti, C. pubescens
has invaded secondary and primary rain for-
ests and undisturbed cloud forest (Meyer
2004, Vanquin 2006). It now dominates na-
tive forests in West Java ( Junaedi and Mu-
taqien 2010). In the highlands of Santa
Cruz Island in Galápagos, C. pubescens domi-
nates the Fern-Sedge zone and codominates
the Miconia zone, formed by the endemic
shrub Miconia robinsoniana Cogn. In both veg-
etation zones, C. pubescens is reducing species
diversity and is adversely affecting species
cover, especially that of endemic IUCN Red
List species, such as Justicia galapagana Lin-
dau and the tree fern Cyathea weatherbyana
(C. V. Morton) C. V. Morton ( Ja¨ger et al.
2007, 2009), as well as Scalesia pedunculata
Hook. f. in the Scalesia zone (Shimizu 1997).
These effects were attributed to the introduc-
tion of a tree to formerly treeless vegetation
zones of the Galápagos Islands, where C.
pubescens is now considered an ecosystem en-
gineer ( Jones at al. 1994, Ja¨ger et al. 2009).
Cinchona pubescens affects the microclimate in
areas where it has invaded by reducing the
photosynthetic active radiation by 87% and
by increasing precipitation as it intercepts the
highland fog with its broad leaves, which also
increases relative humidity ( Ja¨ger et al. 2009).
The concentration of phosphorus in senesced
C. pubescens leaves is double that of the con-
centration found in green leaves, and conse-
quently the phosphorus concentration in the
leaf litter and soil of Cinchona-invaded areas
is significantly higher compared with non-
invaded areas ( Ja¨ger et al. 2013). This, along
with a high specific leaf area and a faster de-
composition rate, suggests that Cinchona en-
hances phosphorus cycling in the soil ( Ja¨ger
et al. 2013). This pattern was confirmed over
a 2-yr study (
H.J., unpubl. data), indicating an
Figure 6. Growth form of Cinchona pubescens in
Galápagos (modified after Shimizu [1997], [ Ja¨ger [2011]:
Cinchona pubescens, Enzyklopa¨die der Holzgewa¨chse, Vol.
58, 14 pp. Copyright Wiley-VCH Verlag GmbH & Co.
KGaA. Reproduced with permission).
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increase in the soil phosphorus concentration
over time. In other systems, enhanced phos-
phorus levels in soils may drive species losses
(
Wassen et al. 2005), but whether or not this
has a negative impact on the native Galápagos
flora, which is adapted to phosphorus-poor
soils, is currently unknown. It is likely though
that increased phosphorus concentrations in
the soils facilitate the establishment of intro-
duced species, which are often adapted to
nutrient-rich soils.
Overall, plant community changes associ-
ated with the C. pubescens invasion in the
vegetation zones of Santa Cruz Island in
Galápagos appear to represent a new stable
state of the highlands plant communities. The
same is probably true for Hawai‘i and Tahiti.
Due to its ability to vigorously regenerate
sexually and asexually, it seems unlikely that
C. pubescens will be replaced over time as a
part of succession following disturbance. The
relatively long persistence of C. pubescens on
the islands where it has been introduced does
not suggest the invasion to be an ephemeral
one. However, more research is needed on all
the invaded oceanic islands to address this
question.
Animals are also affected by the invasion
of C. pubescens in Galápagos. For instance,
Shriver et al. (2011) found a 31% reduction in
the abundance of the Galápagos rail, Lateral-
lus spilonotus Gould, between 2000 and 2007
and related this to the increasing C. pubescens
cover in the study area. These results confirm
those of an earlier study that showed reduced
rail populations in areas invaded by C. pubes-
cens (Gibbs et al. 2003). It is also suspected
that the Galápagos petrel, Pterodroma phaeopy-
gia Salvin, is adversely affected by habitat loss
as C. pubescens overgrows the petrel colonies
( Wiedenfeld and Jiménez-Uzcátegui 2008).
Beneficial Aspects
In Galápagos, C. pubescens may provide a
new habitat for epiphytes such as locally rare
epiphytic fern and orchid species that occur
on C. pubescens trees covered by old lichens
and liverworts ( Ja¨ger et al. 2007). In aban-
doned eucalyptus and pine forests in Hawai‘i,
both the total number (including nonnatives)
and the number of endemic species are higher
in invaded compared with noninvaded plots
(Fischer et al. 2009). This observation was as-
cribed to C. pubescens (and C. calisaya Wedd.)
adding an additional vegetation layer to the
simplified ecosystem (Fischer et al. 2009). In
both cases, it seems that the trees change the
microclimate toward a more humid environ-
ment and thus facilitate the establishment of
hygrophilous species, especially ferns. How-
ever, those were short-term studies, and the
results may change over time and with an in-
creasingly severe C. pubescens invasion.
From a medicinal and economic point of
view, alkaloids extracted from C. pubescens
(and other species, such as C. calisaya) are
commercially valuable (Andersson 1998,
Franckenpohl 2000). The most important al-
kaloid is quinine, which was used for the pre-
vention and treatment of malaria before the
chemical synthesis of quinine in 1944 ( Wood-
ward and Doering 1945, Chopra and Peter
2005). However, the evolution of strains of
malaria parasites resistant to the synthetic
drugs has revived interest in natural quinine
and in the antimalarial properties of the other
alkaloids of Cinchona (
Husain 1991, Gal
2006). This, coupled with the increasing use
of the alkaloid quinidine as an antiarrhythmic
compound, has increased the demand for
Cinchona bark ( Husain 1991).
The wood of C. pubescens is now increas-
ingly used for house and fence construc-
tion in Galápagos because alternative wood
(such as Piscidia carthagenensis Jacq. or Cedrela
odorata L.) is becoming scarce due to govern-
mental regulations and overexploitation (
H.J.,
pers. obs.).
Regulatory Aspects
Cinchona pubescens is included among the 100
worst invaders worldwide (
ISSG 2015) due to
its rapid spread and detrimental impacts on
plant and animal species in the Pacific region.
In Tahiti, C. pubescens is one of 35 invasive
plant species that have been declared “threat-
ening to biodiversity” by the Government of
French Polynesia (LEXPOL 2015). “These
plants are subject to a ban on new imports,
propagation and planting, and prohibition of
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transfer from one island to another of any
whole plant, fragment of plant, cutting, fruit
or seed. Their destruction is permitted”
(
ISSG 2015). In Galápagos, spread to other
islands, which would provide highly suitable
growing conditions for it (e.g., San Cristóbal,
Isabela, Floreana), is prevented by the quar-
antine protocol for travels between islands
(Galápagos National Park Directorate 2015).
Starr et al. (2003) stated that C. pubescens
should not be further sold or planted in the
Hawaiian Islands. In addition, it has received
a High Risk designation by the Hawai‘i
Weed Risk Assessment program, which is
used to help guide planting choices (http://
www.plantpono.org/ hpwra-plant.php?id=
319; F. Starr, pers. comm., 2013).
geographic distribution
Cinchona pubescens has the broadest natural
distribution of all Cinchona species, ranging
from Costa Rica to Bolivia (Andersson 1998).
It has been cultivated in many countries for
the production of quinine, mainly in Bolivia,
China, Colombia, Costa Rica, the Demo-
cratic Republic of Congo, Guatemala, India,
Indonesia ( Java and Sumatra), Jamaica,
Kenya, Mexico, Rwanda, Sri Lanka, Taiwan,
and Tanzania (Table 1); it is still grown
in some of these locations (Popenoe 1949,
Husain 1991, Andersson 1998, Franckenpohl
2000, Zhengyi et al. 2013). Some introduc-
tions were only short-lived, such as the ones
to Algeria and Honduras (Sinclair and Fyfe
1883, Schumann 1891, Popenoe 1949) and to
the southern United States (Fosberg 1947).
Surprisingly, C. pubescens is now considered
rare and endangered in its native range in
Ecuador (Günter et al. 2004).
habitat
Climatic Requirements and Limitations
The ability of C. pubescens to tolerate a wide
range of environmental conditions probably
makes it the most widespread of all Cinchona
species, as well as a successful invader. Its
natural distribution seems to be restricted
to humid areas with an annual rainfall of
>1,400 mm (Garmendia Salvador 2005). In
Galápagos, C. pubescens withstands standing
water accumulated through heavy rain fall
during the El Niño (
H.J., pers. obs., 1998) as
well as prolonged dry periods (
Itow 2003).
Seeds tolerate both shady and sunny condi-
tions for germination (
Itow 2003) (see Figure
8). However, it appears that C. pubescens does
not occur below 130 m and above 3,300 m
above sea level (Gibbs 1885, Andersson 1998,
Rentería 2002, Missouri Botanical Garden
2015). Neither does it seem to flourish below
8°C, whereas it can endure temperatures as
high as 33°C in India and Galápagos (Chopra
and Peter 2005; H.J., unpubl. data). In the
Pacific region, the distribution of C. pubescens
ranges from 600 to 1,400 m in Tahiti (
Van-
quin 2006), from 180 to 860 m in Galápagos
(Rentería 2002), and from 792 to 1,158 m on
Maui (Starr et al. 2003). In West Java, it oc-
curs at approximately 1,600 m ( Junaedi and
Mutaqien 2010). Precipitation for optimal
growth ranges from 1,500 to 4,000 mm yr1 in
Hawai‘i (Fischer 2007), 2,350
5,000 mm yr1
in Tahiti (
Vanquin 2006), and averages
1,400 mm yr1 in Galápagos ( Ja¨ger and
Kowarik 2010).
Habitat and Resource Requirements and
Limitations
Cinchona pubescens thrives in a variety of habi-
tat types, which helps to explain its success
as an invader. It grows in steep gorges that
are difficult to access and in disturbed habitats
in its native range in Ecuador (King 1880,
Acosta Solís 1945a), as well as in river bot-
toms, pasture fields, and abandoned clearings
(Steere 1945). In its introduced range in
Hawai‘i, Tahiti, and Galápagos, it grows
especially well in areas that are disturbed and
difficult to access (Starr et al. 2003, Vanquin
2006, Ja¨ger and Kowarik 2010). In Ecuador,
roots are often exposed to the air (Acosta Solís
1945a), which is also the case in Galápa-
gos, where it thrives in shallow soils less than
20 cm deep (
H.J., unpubl. data). Cinchona pu-
bescens grows well in a range of soil types, such
as volcanic soils, which are rich in organic
matter, nutrients, and metal oxides (Coster
1942, Acosta Solís 1945a, Chopra and Peter
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Pacific Island Invasive Species: Cinchona pubescens, Red Quinine Tree · Ja¨ger 141
2005, Garmendia Salvador 2005, Vanquin
2006, Fischer 2007). However, C. pubescens
flourishes in Galápagos despite the fact that
the soils are poor in nitrogen and phosphorus
( Ja¨ger et al. 2013). Here, C. pubescens may
benefit from increased nutrient uptake due to
its association with arbuscular my
corrhizal
fungi, which are not so abundant in associa-
tion with native species (Schmidt and Scow
1986, Ja¨ger et al. 2013).
Ecosystem and Community Types Invaded
Cinchona pubescens most commonly invades
moist highland vegetation zones, which match
the wet and warm weather conditions of its
native range (Acosta Solís 1945a). In Hawai‘i,
C. pubescens has mainly invaded mesic and
wet forests, growing in steep gulches and
thick understory vegetation (Starr et al. 2003).
In Galápagos, C. pubescens spread from the
agricultural zone, where it was originally
planted, into moist upland zones, including
the Miconia zone, the semitropical forest
(dominated by the endemic tree Scalesia pe-
dunculata Hook. f.), and the Fern-Sedge zone
(Shimizu 1997, Ja¨ger et al. 2007). In Tahiti, C.
pubescens spread into secondary and primary
rain forests and undisturbed cloud forest (up
to 1,400 m on the peninsula of Tahiti Iti
[ Vanquin 2006]), and in West Java it invaded
native forests ( Junaedi and Mutaqien 2010).
history
It is notable that 200 yr passed between the
discovery of the genus Cinchona and its culti-
vation for medicinal use. The reasons for the
long delay are likely related to difficulty in
cultivating the tree, particularly due to low
seed germination rates (Popenoe 1949). Also,
the initially abundant supply of wild bark was
sufficient to meet early demands in Europe
and America (Popenoe 1949).
Cultivated for the quinine of commerce in
many tropical countries and islands, C. pubes-
cens was planted on Maui in 1868 and later by
state foresters on O‘ahu, Maui, and the island
of Hawai‘i in the first half of the 1900s (Starr
et al. 2003). It had naturalized on the island of
Hawai‘i by 1978 and on Maui by 1987 (Starr
et al. 2003). On Maui, C. pubescens is locally
abundant in the Makawao Forest Reserve of
East Maui in areas where it was originally
planted and in plantations of nonnative forest
trees, as well as along old roads, and in thick
native forests (Starr et al. 2006). In Galá-
pagos, C. pubescens is so far present only on
Santa Cruz Island, where it was introduced in
the 1940s by two farmers as a cash crop (
Ha-
mann 1974, Lundh 2006) and had naturalized
by 1972 ( Hamann 1974). The crop was not
economically viable, and quinine production
never commenced there. It spread from initial
plantings, and its distribution increased from
4,000 ha in 1987 to 8,500 ha in 1990 and to
more than 11,000 ha in 2004 (Buddenhagen
et al. 2004).
“In Tahiti, C. pubescens was first introduced
by Pétard (a pharmacist and chemist) and
Boubée (then head of the agronomical station
of Tahiti) as a medicinal plant in 1938 in two
locations (Fautaua Valley at about 480 m and
Afaahiti-Taravao plateau between 280 and
620 m above sea level)” ( J.-Y. Meyer, pers.
comm., 2013). In 1945, about 2,000 trees
were cultivated in Fautaua and 375 in Tara-
vao, but the plantations were abandoned
around 1951. In 1965, Pétard wrote that the
plantations of Fautaua “were left to the bush,
and in Afaahiti they were bulldozed and
turned into pastureland, except a few hun-
dreds of trees.” ( J.-Y. Meyer, pers. comm.,
2013). In 1948, a private farmer had also
planted 10,000 plants of mainly C. pubescens
on the Taravao plateau at about 400 m eleva-
tion. That plantation was also neglected in
1959. Pétard went back to that plantation in
1969 and noted that “Although the place is
invaded by the bush . . . some trees reach the
enormous height of 10
13 m and many of
them bear flowers or fruits. . . .” He also
checked the Fautaua plantation in 1969 where
he found “about fifty Cinchona succirubra [syn.
for C. pubescens] . . .” (Pétard 1964, 1986, ac-
cording to J.-Y. Meyer, pers. comm., 2013).
The successful invasion of C. pubescens on
all of these islands was a slow but continuous
one, which is reflected in the long time that
passed between the introduction of the spe-
cies and the recognition that it had become
invasive: about 40 yr in Galápagos (Ortiz and
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TABLE 1
Geographic Distribution of Cinchona pubescens in the Pacific Region
Locality Present Absent No Infor-
mation Citation
American Samoa X Presumed absent. Not listed in U.S. Forest Service
(2015)
Australia X Presumed absent. Not listed in the Flora of Australia
Online (2015)
Bonin Islands X Presumed absent. Not listed in Kawakami and
Okochi (2010)
Cedros Island, Mexico X Presumed absent. Not listed in Oberbauer (1993)
Clarion Island, Mexico X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Clipperton Island X Presumed absent. Not listed in Sachet (1962)
Cocos Island, Costa Rica X Presumed absent. Not listed in Fosberg and Klawe
(1966)
Cook Islands X Presumed absent. Not listed in U.S. Forest Service
(2015)
Easter Island (
Isla de
Pascua or Rapa Nui) X Presumed absent. Not listed in Dubois et al. (2013)
Fiji XaKew Gardens (2015) ( but see U.S. Forest Service
2015)
French Polynesia
Austral Islands X Presumed absent. Not listed in DIREN (2007)
Marquesas X Presumed absent. Not listed in Wagner and Lorence
(2002)
Society Islands X On Tahiti (Meyer 2000)
Tuamotu Islands X Presumed absent. Not listed in DIREN (2007)
Galápagos Islands X Hamann (1974)
Gorgona Island X Presumed absent. Not listed in Vásquez-Vélez (2014)
Guadalupe Island, Mexico X Presumed absent. Not listed in Oberbauer (2005)
Guafo Island, Chile X Presence unlikely, given the low annual temperatures
Hawaiian Islands X Starr et al. (2003)
Henderson Island, Chile X Presumed absent. Not listed in Waldren et al. (1995)
Indonesia X On Java (Andersson 1998)
Isla de la Plata, Ecuador X Presence unlikely, given the semiarid climatic
conditions
Japan (main islands) XbOn Okinawa ( Nagumo et al. 2010)
Juan Fernandez X Presumed absent. Not listed in Cuevas et al. (2004)
or Greimler et al. (2013)
Kiribati
Gilbert Islands X Presumed absent. Not listed in Wagner et al. (2012)
Line Islands X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Phoenix Islands X Presumed absent. Not listed in PIPA (2015)
Lobos Islands, Peru X Presence unlikely, given the arid climatic conditions
Lord Howe Island X Presumed absent. Not listed in the Flora of Australia
Online (2015)
Malpelo Island, Colombia X Presence unlikely, given the arid climatic conditions
Micronesia
Caroline Islands XcWagner et al. (2012)
Mariana Islands
(including Guam) X Presumed absent. Not listed in U.S. Forest Service
(2015)
Marshall Islands
(including Wake
Island)
X Presumed absent. Not listed in U.S. Forest Service
(2015) or Wagner et al. (2012)
Nauru X Presumed absent. Not listed in U.S. Forest Service
(2015)
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Pacific Island Invasive Species: Cinchona pubescens, Red Quinine Tree · Ja¨ger 143
Lawesson 1987), about 100 yr in Hawai‘i
(Starr et al. 2003), and about 60 yr in Tahiti
(
Vanquin 2006). In Galápagos, it has been
assumed that C. pubescens requires disturbance
to be able to proliferate since it spread after
prolonged man-made fires in the highlands in
the late 1960s (Kastdalen 1982). However, it
is possible that the expansion of C. pubescens
simply went unnoticed for a long time before
it became obvious. Starr et al. (2003) reported
that C. pubescens spreads along bulldozed for-
est roads in Hawai‘i but it seems that “C. pu-
bescens does not need bare, open soil or sunny
locations [. . . to spread] and that some of the
densest stands are in the shade of forestry
planting of eucalyptus” ( H. Oppenheimer,
pers. comm., 2013). On Maui, it appears that
it spreads from the area it was planted, re-
gardless of the surrounding habitat (K. and
F. Starr, pers. comm., 2013). There are
TABLE 1 (continued)
Locality Present Absent No Infor-
mation Citation
Palau XdFosberg et al. (1993) ( but see U.S. Forest Service
2015)
New Caledonia X Presumed absent. Not listed in Beauvais et al. (2006)
New Zealand X Presumed absent. Not listed in Breitwieser et al.
(2010)
Niue X Presumed absent. Not listed in Space and Flynn
(2000)
Norfolk Island X Presumed absent. Not listed in the Flora of Australia
Online (2015)
Pacific Equatorial Sporades
(Baker, Howland, and
Starbuck Island)
X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Palmyra X Presence unlikely, given the climatic conditions
Papua New Guinea X Andersson (1998)
Philippines X FAO (2015)
Pitcairn Island X Presumed absent. Not listed in Kingston and
Waldren (2003)
Sala y Gómez X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Samoa X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Socorro Island X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Solomon Islands X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Taiwan X Zhengyi et al. (2013)
Tokelau Island X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Tonga X Presumed absent. Not listed in U.S. Forest Service
(2015)
Tuvalu X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Vanuatu X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
Wallis and Futuna X Presumed absent. Not listed in Mueller-Dombois
and Fosberg (1998)
a Only a single collection record from Kew Gardens, collected in 1868 by the Colonial and Indian Exhibition, Fiji Commission.
b Nagumo et al. (2010) mentioned the establishment of Cinchona plantations on Okinawa and Kagoshima in 1876 but did not men-
tion which species. Plants had died by 1884.
c Listed as Cinchona succirubra.
d Introduction reported but not confirmed.
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144 PACIFIC SCIENCE · April 2015
indications that disturbance facilitated the
spread in Tahiti (
Vanquin 2006). There, the
initial spread of C. pubescens is likely to have
been slowed down by competition with the
invasive Miconia calvescens DC., which pro-
duces intense shade, and is itself shade toler-
ant (
Vanquin 2006). Cinchona pubescens has
also been reported to occur outside planta-
tions in Java and Papua New Guinea (Anders-
son 1998) and outside the Pacific region in
Jamaica (Sauer 1988), Guatemala (Martin and
Gandara 1945) and on Saint Helena (Cronk
and Fuller 1995).
physiology
Cinchona pubescens sheds its leaves continu-
ously throughout the year (Cedeño 1990,
Rentería 2002). It does not seem to resorb
phosphorus from senesced leaves before de-
taching them, resulting in a two-fold higher
phosphorus concentration in the senesced
leaves compared with mature green leaves
(6.04 mg g1 versus 3.15 mg g1 [ Ja¨ger et al.
2013]). A similar pattern was found for Ole-
aria axillaris (DC.) F. Muell. ex Benth. in the
coastal area of southwestern Australia with
phosphorus-poor soils (
like the ones in Galá-
pagos) and where senesced leaves also con-
tained the double amount of phosphorus
compared with green leaves (5.65 mg g1 ver-
sus 2.94 mg g1 [Hayes et al. 2013]). In both
cases, a possible explanation for this unusual
finding might be arbuscular mycorrhizal fun-
gi associated with both C. pubescens and O. ax-
illaris, which might enhance the phosphorus
uptake of their roots ( Hayes et al. 2013, Ja¨ger
et al. 2013).
A study of root samples collected from
C. pubescens adult trees in Galápagos showed
that roots were highly colonized with arbus-
cular mycorrhizal fungi (20% 100%), and
PCR analysis revealed that 95.5% of these be-
longed to the family Glomeraceae and 4.5%
to the Diversisporaceae (Serrano Tamay
2013).
reproduction
Cinchona pubescens has hermaphroditic flowers
that are probably pollinated by insects. Studies
to confirm this are lacking, but the sweet fra-
grance of flowers suggests this mode of polli-
nation (Starr et al. 2003). Biological charac-
teristics of C. pubescens that make it a successful
invader in its introduced range include its
abundant seed production. The youngest
seed-producing trees observed were 1 2 yr
old in Tahiti (
Vanquin 2006) and 2 yr old in
Galápagos; the latter were 1.8 m in height
with a DBH of 1.5 cm (
H.J., unpubl. data). As
in the native range (Garmendia Salvador
2005), C. pubescens trees in Galápagos de-
velop flowers more or less all year round but
with a peak between August and November
(Rentería 2002). Fruit production peaks be-
tween November and April, but mature fruits
persist on the trees for a long time so that
they can be found during all months of the
year (Rentería 2002). Development from the
opening of the flower to the production of
mature fruits takes about 19 weeks (Rentería
2002). Similar observations have been re-
ported from India, where flowers and fruits
are found almost all year round (Kuntze
1878), but trees only flower after the fourth
year (Chopra and Peter 2005). In Galápa-
gos, each capsule contains about 60 70 seeds
( J. L. Rentería, pers. comm., 2013), and the
thousand-seed weight is 0.26 g (
H.J., unpubl.
data). A C. pubescens tree in Bolivia produces
about 7
9 kg of seed per year, which were
harvested in November and December dur-
ing the height of Cinchona exploitation (Gibbs
1885).
The light, winged, wind-borne seeds do
not seem to be dispersed far from the parent
tree in the introduced range. In Hawai‘i, the
vast majority of seeds do not reach farther
than 100 m from the parent tree (Starr et al.
2003), and the maximum dispersal distance
reported from Galápagos was about 15 m
(Rentería 2002). Another characteristic of C.
pubescens is the ability of the seeds to germi-
nate in dense understory vegetation as well as
under a dense C. pubescens canopy, as shown
for Hawai‘i, Tahiti, and Galápagos (Pala-
cios 1993, Meyer 2000, Starr et al. 2003). In
addition, C. pubescens spreads vegetatively by
rapid suckering from roots and stems (Mac-
donald et al. 1988, Fischer 2007, Ja¨ger et al.
2009).
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Pacific Island Invasive Species: Cinchona pubescens, Red Quinine Tree · Ja¨ger 145
Because there is limited information avail-
able on the germination and growth of C. pu-
bescens in the Pacific region, general informa-
tion is drawn from experimental plantations
in the native and introduced range. In India,
seeds sown in soil beds germinate in 10 20
days, with a germination rate from 50% to
85%. These seedlings usually grow a pair of
leaves within 3 months and grow to a height
of about 20
25 cm after 16 months ( Joy et al.
2001, Chopra and Peter 2005).
Apparently, introduced C. pubescens grows
better in the field than under greenhouse con-
ditions. The reasons for this include different
nutrient and climate regimes (
H.J., unpubl.
data). In a greenhouse experiment in Loja,
Ecuador, seeds of C. pubescens from the native
range in Loja germinated after 19 days and
reached a total germination rate of 95%,
whereas seeds from Galápagos germi
nated
after 17 days with 87% total germination
(
H.J. and M. E. Apolo Chamba, unpubl. data).
Seedlings grown in the greenhouse from
Galápagos seeds grew about 13 cm in 13
months, which contrasts with field observa-
tions indicating that seedlings grew about 1 m
during the first year, attaining about 1.8 m
height at the end of the second year (
H.J.,
unpubl. data). Seedlings in a Jamaican planta-
tion also grew about 1 m per year (Sinclair
and Fyfe 1883).
Seeds from India began to lose their viabil-
ity after 6 to 8 weeks and lost it completely
after 1 yr (Kuntze 1878). In Galápagos, seeds
also lost their viability after approximately
1 yr (Rentería 2002).
population dynamics
Cinchona pubescens is a fast-growing species in
its introduced range. In Tahiti, trees grow
about 1
2 m per year and a 10-yr-old tree
reaches a DBH of 5 cm (
Vanquin 2006).
Trees 15 m tall with a DBH of 25 cm were
observed in Galápagos (Shimizu 1997; H.J.,
unpubl. data). As early as 1974, the invasive
character of C. pubescens was visible in
the highlands of Santa Cruz Island in Ga-
lápagos (
Hamann 1974). Figure 7 shows the
increase of C. pubescens density from 1987 to
2005. In 1987, C. pubescens density was esti-
mated at 1 100 individuals ha1 (Macdonald
et al. 1988). In the area toward the top of
the island, where Macdonald et al. (1988) had
counted one individual ha1, about 60 100 in-
dividuals ha1 were recorded only 4 yr later
(
Valdebenito 1991). This rapid increase in
density of C. pubescens was confirmed by a 7-yr
study in the fern-sedge highland vegetation
( Ja¨ger et al. 2009). Results showed that the
density of trees larger than 1.5 m increased
from 123 in 1998 to 439 ha1 in 2005. Con-
currently, the number of stems ha1 increased
from 355 to 1,652, which represents an aver-
age number of stems of 2.9 and 3.8, per tree,
respectively. These results parallel those of
Shimizu (1997), who recorded an average
of 3.4 stems per tree in an area nearby. The
basal area increased from 1.0 to 4.2 m² ha1
and mean C. pubescens cover increased from
6.6% to 16.4% over 7 yr ( Ja¨ger et al. 2009).
The highest C. pubescens cover recorded in
individual 20 by 20 m plots was 42% ( Ja¨ger
et al. 2009).
Cinchona pubescens reaches similar abun-
dance levels on other Pacific islands. In ex-
perimental plots on the Taravao plateau and
on the Atar Mountain in Tahiti (600
955 m
above sea level), the maximum percentage
cover of C. pubescens was 55%, maximum
Figure 7. Increase of Cinchona pubescens individuals (in-
cluding seedlings) at Media Luna between 1987 and 2005
on Santa Cruz Island, Galápagos. Value for the year 1987
from Macdonald et al. (1988), for the year 1991 from Val-
debenito (1991), and for the years 1998 to 2005 from
Ja¨ger et al. (2009).
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146 PACIFIC SCIENCE · April 2015
density of stems was 9,200 ha1, and that of
seedlings was 14,500 ha1 ( Vanquin 2006).
However, these densities were recorded from
small study plots in the most invaded areas of
Tahiti and may not necessarily be represen-
tative of the current situation ( J.-Y. Meyer,
pers. comm., 2013). In abandoned eucalyptus
and pine plantations, as well as in natural
Acacia koa stands in the Makawao Forest Re-
serve on Maui, Hawai‘i, average percentage
cover of C. pubescens reached 27%, the average
density of stems (0.2 1 m) 4,000 ha1, and that
of seedlings 114 ha1 (Fischer 2007). Popula-
tion density in the Patengan Natural Reserve
in the tropical mountain rain forests of West
Java was 222 individuals ha1, which is the
highest density of tree population in all three
research locations around Mount Patuha
( Junaedi and Mutaqien 2010).
response to management
Cinchona pubescens has long been recognized as
a potential risk to native vegetation in its in-
troduced range, especially in Hawai‘i, Galá-
pagos, and Tahiti. On Santa Cruz Island, sev-
eral control methods to combat C. pubescens
have been implemented in the Galápagos Na-
tional Park area and in the agricultural area.
Manual methods include felling and uproot-
ing of trees as well as hand pulling of smaller
plants, but trees resprout from cut stumps
(Macdonald et al. 1988, Buddenhagen et al.
2004) (Figure 8). Chemical control methods
consist of applying herbicides by a range of
means such as hack and squirt, basal bark, cut
stump, girdle and squirt, branch filling, tree
injections, and foliar spraying (Buddenhagen
et al. 2004). Most of these methods were inef-
Figure 8. Cinchona pubescens regeneration and control. Top left, germination under dense Miconia robinsoniana vegeta-
tion; top right, resprouting after manual control; bottom left, chemical control (“hack and squirt”); bottom right, manual
control. (Photos by H. Ja¨ger on Santa Cruz Island, Galápagos.)
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Pacific Island Invasive Species: Cinchona pubescens, Red Quinine Tree · Ja¨ger 147
fective in the long run. Eventually, a hack and
squirt technique was developed where a mix-
ture of picloram and metsulfuron was applied
to connecting machete cuts around the cir-
cumference of tree trunks, killing 73%
100%
of the trees (Buddenhagen et al. 2004) (Figure
8). Smaller shoots and saplings were pulled
out by hand. This method is now being suc-
cessfully applied for C. pubescens control by
the Galápagos National Park Directorate on a
small scale (approximately 110 ha between
1998 and 2003 [Buddenhagen and Yánez
2005]) in addition to uprooting trees in con-
servation priority areas ( Ja¨ger and Kowarik
2010).
Studies on the impacts of these measures
revealed that despite an initial decline in spe-
cies cover, native plants recovered and total
species cover as well as species diversity in-
creased after the control action occurred
( Ja¨ger and Kowarik 2010; H.J., unpubl. data).
However, both control actions caused severe
disturbance to the surrounding vegetation
and soil (Figure 8). This probably also facili-
tated the establishment of other introduced
species. For example, there was a significant
increase in cover of the highly invasive black-
berry Rubus niveus Thunb. ( Ja¨ger and Kowarik
2010; H.J., unpubl. data). In addition, con-
tinuous hand pulling of emerging seedlings
over a long period would be necessary to
guarantee lasting control success ( Ja¨ger and
Kowarik 2010).
Buddenhagen and Yánez (2005) estimated
the costs for chemical control of C. pubescens
on Santa Cruz Island at between US$14
2,225 ha1, depending on stem density. Treat-
ing all existing plants once in the invaded area
of at least 11,000 ha would require about
276,500 hr of labor (equivalent to 150 people
working for a year) at an estimated cost of
US$1.65 million in 2005 (Buddenhagen and
Yánez 2005). Treated areas would still have
to be revisited subsequently for several years
to pull out germinating seedlings ( Ja¨ger and
Kowarik 2010).
Control of C. pubescens is difficult in Hawai‘i
(mainly Maui) because it primarily grows in
steep gulches and thick understory vegeta-
tion. On the other hand, controlling small
populations in patches of native vegetation
could prevent further spread and help slow
down the degradation of the remaining native
forest in the area (Starr et al. 2003). An inter-
esting method was recently developed to kill
invasive Miconia calvescens DC. in inaccessible
areas in Hawai‘i by shooting encapsulated
herbicide projectiles at Miconia plants from a
helicopter (Leary et al. 2013). Something
similar might be possible for C. pubescens in
Hawai‘i and elsewhere.
natural enemies
Although the databases of the Systematic
Mycology and Microbiology Laboratory
(Farr and Rossman 2010) list 27 fungi species
as associated with C. pubescens, only seven of
these occur in the native range of C. pubescens
(e.g., Elsinoe cinchonae Jenkins, Phytophthora
cinnamomi Rands, Prillieuxina cinchonae J. A.
Stev.). The scab-causing pathogen Elsinoe cin-
chonae was also recorded from C. pubescens
from western Ecuador (
H. C. Evans, pers.
comm., 2013). However, there is no evidence
that these species are important economic
pathogens (PIER 2015) and therefore might
not be likely candidates for biological control.
In plantations in Guatemala, India, and
Java, Cinchona is susceptible to many pests
and diseases. Cinchona seedbeds are frequently
affected by the fungi Rhizoctonia solani J. G.
Kühn, Phytophthora spp. (root-rot fungus),
and Rosellinia arcuata Petch (
black-root fun-
gus), causing a disease known as “damping
off
(Coster 1942, Popenoe 1949, Chopra
and Peter 2005). These fungi penetrate the
seedlings through their roots and cause
sudden wilting and rotting of seedlings. In
Java and India, plants are also attacked by the
insects Helopeltis (tea mosquito) and Pachy-
peltis (
leaf scorch) (Coster 1942, Chopra and
Peter 2005). Cinchona stem bark disease due
to Phytophthora cinnamomi is a major disease
in central African countries (Rwanda and the
Democratic Republic of Congo [Chopra and
Peter 2005]). In Galápagos, secondary patho-
gens were isolated from C. pubescens, mainly
Fusarium spp. and Botryodiplodia theobromae
Pat. (
H. C. Evans, pers. comm., 2013). Van-
quin (2006) reported that in 1939 Roger
Heim found a mushroom of the Agaricaceae
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148 PACIFIC SCIENCE · April 2015
family in Papua New Guinea that attacks the
roots of C. pubescens, which leads to a desicca-
tion of the seedlings and a partial destruction
of different plant parts.
For a detailed description of the natural
enemies of C. pubescens see the CABI Invasive
Species Compendium (CABI 2015).
prognosis
An interval ranging from 40 to 100 yr passed
before C. pubescens was recognized as an inva-
sive species after its initial naturalization in
Tahiti, Hawai‘i, and Galápagos. One pos-
sible reason for this could be that the area
lacked openings for invasion windows (“safe
sites”) in the form of disturbance or climate
change (Kowarik 1995). Alternatively, the
invasion may have simply been unnoticed for
several decades. Starr et al. (2003) stated that
other mesic and wet forests on Maui currently
free of C. pubescens are potentially threatened
by invasion in the future. A future expansion
is also possible on Tahiti (
Vanquin 2006). In
addition, C. pubescens has been reported to oc-
cur outside plantations in Papua New Guinea
and Jamaica (Sauer 1988, Andersson 1998).
This means that there is still a risk that C. pu-
bescens might become invasive in areas where
it has been introduced but its ability to spread
without human intervention has not yet been
recognized. For example, C. pubescens was
introduced to Java in 1856 (de Padua et al.
1999), but it was only recently considered
invasive ( Junaedi and Mutaqien 2010). In
Galápagos, C. pubescens could easily grow on
islands other than Santa Cruz that receive
enough rainfall to allow agriculture, such as
San Cristóbal, Isabela, and Floreana, if seeds
were brought there by humans. The same is
probably the case for the Hawaiian Islands.
Cinchona pubescens produced a score of 9 (high
risk) in the Weed Risk Assessment from Pa-
cific Island Ecosystems at Risk (Daehler et al.
2004, PIER 2015) and therefore could be-
come invasive in many islands if introduced
and found optimal growing conditions. Pope-
noe (1949) reported that Cinchona seedlings in
the first plantations in Guatemala died shortly
after planting, and it was assumed that the
soils were not sufficiently acidic. Thus it is
possible that C. pubescens would more likely
become invasive in volcanic (= acidic) soils. As
a result, known populations on all Pacific is-
lands as well as in Asia (e.g., the Philippines or
Taiwan) should be monitored carefully. In
the case of small C. pubescens populations, it
would be best to eliminate these by control
methods currently applied in the Galápagos
National Park (Galápagos National Park
Directorate 2015).
acknowledgments
I thank Jon Witman and Alan Tye for their
constructive comments on the manuscript.
My thanks also to David Clements and two
anonymous reviewers who substantially im-
proved the content of the article. Jean-Yves
Meyer, Kim and Forest Starr, Hank Oppen-
heimer, and Leonie Fischer shared their
knowledge and photos of Cinchona pubescens
from Tahiti and Hawai‘i, for which I am very
grateful, as well as for the information by
Harry Evans on Cinchona pathogens. This
publication is contribution number 2094 of
the Charles Darwin Foundation for the Galá-
pagos Islands.
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... Therefore, it had been introduced to many parts of the world, mainly from Bolivia, Colombia, Ecuador, and Peru (Andersson 1998). The main area of introduction was the Pacific region, were quinine had been introduced to 11 islands or archipelagos (Jäger 2015) and has become invasive at least in Galapagos (Macdonald et al. 1988), Hawaii (Starr et al. 2003), Tahiti (Meyer 2004), and West Java (Junaedi and Mutaqien 2010). Quinine is now considered among the 100 worst invasive species globally (ISSG 2014). ...
... Red quinine tree has a natural distribution from Costa Rica to Bolivia (Andersson 1998). In Galapagos, this evergreen tree with wide leaves and aromatic, lightly pink flowers reaches a maximum height of 15 m (Shimizu 1997) and a DBH of 25 cm (Jäger 2015). ...
... Over a 7-year study in the highlands of Santa Cruz, the basal area of quinine trees increased from 1.0 to 4.2 m 2 ha −1 in the study plots, while mean quinine cover increased from 6.6% to 16.4% over the same time period. The mean density of quinine stems in 2005, including seedlings, was 2193 individuals ha −1 (Jäger 2015). ...
Chapter
One of the most invasive species in Galapagos Islands is the red quinine tree, Cinchona pubescens Vahl (Rubiaceae). Though considered rare and endangered in its native range in Ecuador, quinine is being controlled as an invasive and used as a timber source in Galapagos. Introduced to Santa Cruz Island in the 1940s, it started spreading in the 1970s and now covers a vast area in the humid highlands of the island. Quinine is considered an ecosystem engineer, changing plant species diversity and abundance as well as impacting endemic birds in the invaded area. It also alters the microclimate and increases phosphorus concentrations in the soil. The production of abundant small and wind-borne seeds, paired with a vigorous vegetative reproduction, makes it a very successful invader. Quinine is manually and chemically controlled by Galapagos National Park Directorate, but this method requires constant follow-up control of seedlings germinating from the seed bank. In addition, disturbances caused by these control measures seem to facilitate the establishment of other introduced plant species, especially blackberry (Rubus niveus). The quinine invasion in Galapagos Islands provides an opportunity to help understand the ecology of plant invasions in Galapagos and in island ecosystems in general.
... Está introducida en Tahití, zona del Pacífico, Hawai, Asía y en Tanzania, África. Esta especie crece en altitudes entre 300 y 3 300 m (Jäger, 2015), muy robusta por lo que se utilizaba como patrón de injerto. El contenido en alcaloides totales es del 3,8 %, de ellos menos de 50 % de quinina. ...
... El contenido en alcaloides totales es del 3,8 %, de ellos menos de 50 % de quinina. Se considera rara y en peligro de extinción en su área de distribución nativa en Ecuador (Günter et al., 2004), mientras que (Jäger y Kowarik, 2010) y (Jäger, 2015), la consideran como invasora, en las condiciones insulares de Galápagos. ...
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The management of forest seeds in Ecuador, in relation to their storage, is still insufficient, in particular the Cinchona pubescens Vahl. The objective was to determine the quality of the seeds and the best methods of seed storage of C. pubescens, using different times, containers and conditions. Seed collection was in Intag, Imbabura. The factors under study were: storage time, types of packaging and storage conditions. The quality of the seeds was determined at the time of harvest and in each treatment. Purity reached 76.5 %, the weight of 1,000 seeds was 0.31 g and the humidity content was 14 %. In the first storage trial, no germination was observed in any treatment from the first month. In the second trial, the best storage method was transparent cover stored in refrigeration for one week, which showed 9.25 % germination power. Cinchona pubescens seeds lose their viability quickly, which becomes null after one month of storage.
... Many invasive plant species spread rapidly and are difficult to eradicate once they are established and cover more than 1 ha (Richardson et al. 2000), resulting in changes in ecosystem functions, including the alteration of physical conditions which can act as irreversible barriers to restoration (Jäger, Kowarik, & Tye, 2009;Gardener et al., 2013). A few examples of the environmental impacts caused by quinine (Cinchona pubescens) and blackberry (Rubus niveus) are (a) changes in species composition and community structure, (b) changes in microclimate regimes (light, humidity, precipitation, etc.), and (c) changes in nutrient cycling (increase in nitrogen and/or phosphorus), in combination with a faster decomposition of the leaves (Rentería et al., 2012;Jäger, 2015). ...
... The interviews were carried out in English and Spanish, recorded, and the Spanish subsequently translated to English. Other methods used to understand the different components encompassing land management practices and debates in Galapagos include a review and analysis of technical reports, published literature, and existing data (e.g., maps, socio-economic and species abundance/change data) as well as of current and past policies, specifically the LOREG, 1999, 2015(CGREG, 2015. ...
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Terrestrial invasive species have been identified as one of the largest threats to endemic plants and wildlife in Galapagos and their spread remains one of the biggest challenges for the region. The management of these species is a common link among all land use activities and their spread impacts all residents as economic activities in Galapagos are linked to its status as a unique landscape. The study aims, through the use of key informant interviews, policy documents and literature, to provide new insights into plant invasive species management by exploring two land use interventions ‐ and the associated challenges and opportunities – currently being proposed by policymakers, academics and other relevant actors. These are 1) local sustainable agricultural production and 2) policies and mechanisms, specifically the ‘Buen vivir’ paradigm with/and Payments for Ecosystem Services (PES). It explores how these can create bridges and be beneficial to both conservation and development. However, whilst the initiatives offer real opportunities to manage and control invasive species, challenges remain in the form of how these activities will be carried out and by whom. Findings show that probable success is dependent on community inclusion with coordinated and integrated approaches from robust institutions with connectivity among land use actors/managers. In addition, support is needed for organisations/stakeholders that are currently tackling the invasive species issue. Studies on land use remain crucial as relatively contained and pristine landscapes such as Galapagos are likely to be increasingly important as a means to detect human‐induced alterations at the frontiers of ecology.
... Cinchona pubescens es también una especie arbórea de amplia distribución y fuera de su rango es considerada entre las malezas más agresivas, en especial en varias islas del Pacifico incluyendo Hawái, Tahití y Galápagos (Jäger 2011(Jäger , 2015. Por ese carácter agresivo, la literatura sobre ecología e impacto ambiental es amplia. ...
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Full-text available
Este trabajo se centra en las recientes decisiones tomadas por el Ministerio de Agricultura del Perú sobre la recolección, propagación y siembra de especies de quina. Señalamos las deficiencias de estas decisiones a la luz de los avances en el conocimiento de la taxonomía y sistemática de Cinchona. Destacamos el papel de la ciencia analizando e informando sobre las decisiones en el uso sostenible de los recursos naturales.
... Introduced species such as Cinchona pubescens, Rubus niveus and Psidium guajava have transformed highland habitats on the larger islands with fertile soils (Jäger et al. 2013;Jäger 2015). Using a high resolution study of macro-remains extracted from bogs on Isla Santa Cruz in Galápagos, Coffey et al. (2010) confirmed that Ageratum conyzoides, Solanum americanum, Ranunculus flagelliformis, Brickellia diffusa, Galium canescens and Anthephora hermaphrodita were introduced by people to the islands. ...
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The initial relationships between the first human colonizers and the native vegetation of Isla San Cristóbal, Galápagos, were studied by the analyses of wood charcoal, plant macro-remains, phytoliths and historical records. Archaeological and modern botanical samples were collected from four archaeological sites within the former farmland of the 19th century Hacienda El Progreso, a sugar plantation located in the moist highlands of the island. The archaeobotanical remains show the use of native timber, the introduction of crops and weeds, some aspects of local diet, and evidence of vegetation clearance. Ecological impact is shown by the changes to the native vegetation caused by human colonization of the island and the expansion of agricultural land for the plantation enterprise. This paper provides a synthesis of the archaeobotanical study at El Progreso which forms a baseline for future research in the Galápagos islands.
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Las cortezas y raíces de las diferentes especies de los árboles del género Cinchona L. (entre ellas, C. calisaya Wedd., C. officinalis L. y C. pubescens Vahl), han sido ampliamente utilizadas por su gran virtud febrífuga, igualmente para las arritmias cardiacas, los calambres musculares, resfriados, indigestión, fibrilación auricular, tónico eupéptico, catarros, acelerante del parto y como tónico capilar (Pérez Arbelaez, 1947; García-Barriga, 1975; Loayza-O, et al., 2010).
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Abstrak. Mutaqien Z. 2017. Pendugaan pelepasan senyawa alelopati pada proses dekomposisi serasah daun tumbuhan invasif: Calliandra calothyrsus dan Cinchona pubescens. Pros Sem Nas Masy Biodiv Indon 3: 334-338. Jenis asing invasif adalah salah satu faktor utama penyebab perubahan dramatis pada berbagai sistem ekologi di seluruh dunia dan disinyalir sebagai salah satu penyebab utama kepunahan jenis asli. Namun masih diperlukan banyak pekerjaan untuk memahami bagaimana mekanisme, termasuk peran alelopati pada berbagai jenis tumbuhan invasif. Cinchona pubescens Vahl. dan Calliandra calothyrsus Meissn. diketahui sebagai jenis invasif di beberapa tempat, bahkan C. pubescens termasuk salah satu dari 100 organisme paling invasif di dunia. Penelitian ini merupakan studi pendahuluan dari dugaan adanya mekanisme alelopati yang berasal dari proses dekomposisi daun dari dua jenis tumbuhan invasif tersebut. Percobaan perkecambahan Vigna radiata (L.) R. Wilczek dengan Rancangan Acak Lengkap dengan dua perlakuan berupa pemberian cairan rendaman daun C. pubescens dan C. calothyrsus dengan masa rendaman 7-180 hari dengan masing-masing tiga kali ulangan dilakukan untuk merepresentasikan proses penyebaran alelopati melalui proses dekomposisi yang mungkin terjadi di alam. Panjang akar dan batang kecambah diukur dan diproses menggunakan Anova dan uji lanjutan Ducan. Air rendaman daun C. calothyrsus tidak berpengaruh nyata pada perkecambahan V. radiata. Air rendaman C. pubescens berpengaruh nyata terhadap perkecambahan. Diduga kuat terdapat senyawa alelopatik yang dilepaskan pada proses dekomposisi daun C. pubescens. Semakin lama proses dekomposisi, semakin berkurang dampaknya terhadap perkecambahan. Abstract. Mutaqien Z. 2017. Presumption of allelopathic compound(s) released in the leaf litter decomposition process of invasive plants: Calliandra calothyrsus and Cinchona pubescens. Pros Sem Nas Masy Biodiv Indon 3: 334-338. Invasive alien species is one of the main cause of ecological systems dramatically changes and also native species extinction worldwide. However, more studies are needed to discover its mechanism, including role of allelopathy compound in invasive plants competition. Cinchona pubescens Vahl. and Calliandra calothyrsus Meissn. are recorded as invasive species in some regions. Moreover, C. pubescens is stated as one of 100 of the world's worst invasive alien species by IUCN. This study aimed to confirm indication of the existence of allelopathic mechanism in invasion process of these two invasive alien species by releasing allelopathic compound(s) from its leaves fall over decomposition process. To clarify this hypothesis, a preliminary study had been conducted by testing the effect of the compounds released from decomposition process over the times (7-180 days) to germination of Vigna radiata (L.) R. Wilczek in the laboratory. Completly Randomize Design was used by applying liquid produced by decomposition process of these two invasive species and control (three repetitions) to test its effects. Radicles and hypocotyls length were measured, compared and Anova analyzed by using R-statistic 3.1.3. Germination of V. radiata only significantly inhibited by compound(s) released by decomposition processes of C. callisaya's leaves. Its inhibition effect was reduced over the times
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Loss of biodiversity on tropical and subtropical oceanic islands is one of the most pressing conservation issues. These oceanic islands are well known for their unique fauna and ? ora, which evolved over long periods in isolation from external perturbation. However, the maj- ity of these islands in the Paci? c were eventually settled by Polynesians and then by Europeans; by about 200 years ago, only a few island groups remained untouched. The Bonin Islands are one of these groups. The Bonin Island group is one of the most remote in the world. The islands are located 1,000 km south of Japan off the eastern fringe of Eurasia. They were ? rst discovered by the Japanese in 1670, settled by Westerners from Hawaii in 1830, and ? nally recognized as a Japanese territory in 1862 on condition that previous settlers would be protected and allowed to remain with full rights. Because of this complicated history, the Bonins have two names.
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Interactions between organisms are a major determinant of the distribution and abundance of species. Ecology textbooks (e.g., Ricklefs 1984, Krebs 1985, Begon et al. 1990) summarise these important interactions as intra- and interspecific competition for abiotic and biotic resources, predation, parasitism and mutualism. Conspicuously lacking from the list of key processes in most text books is the role that many organisms play in the creation, modification and maintenance of habitats. These activities do not involve direct trophic interactions between species, but they are nevertheless important and common. The ecological literature is rich in examples of habitat modification by organisms, some of which have been extensively studied (e.g. Thayer 1979, Naiman et al. 1988).