Enzyklopädie der Holzgewächse – 58. Erg.Lfg. 06/11 1
Cinchona pubescens VAHL, 1790
syn.: Cinchona chomeliana WEDD., 1848,
Cinchona lutea PAV . in HOWARD, 1859,
Cinchona ovata (RUIZ & PAVO N , 1799),
Cinchona succirubra (PAV . ex KLOTZSCH, 1857)1
Red cinchona, Red quinine tree, Peruvian bark,
Jesuits’ bark, Countess bark Family: Rubiaceae
French: arbre à quinine, quinquina rouge
German: Chinarinde, Chinarindenbaum
Spanish: Cascarilla, Hoja ahumada, Hoja de zambo, Quina2
1For a list of synonyms see ANDERSSON (1998)
2Acosta Solís (1945b) lists 18 different common names for Ecuador alone
3Author’s note: Since C. succirubra and C. ledgeriana are synonyms for C. pubescens and C. calisaya, respectively, these names are substituted throughout
the text when synonyms were used in older publications
Fig. 1: Cinchona pubescens trees in the introduced range: highlands of Santa Cruz Island Galápagos (600 masl)
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Cinchona is the most commercially important genus of
the family Rubiaceae (coffee family) after the genus Cof-
fea, which produces the coffee of commerce. The genus
Cinchona comprises 23 species of tropical evergreen trees
and shrubs, which are distributed from Costa Rica to Bo-
livia. C. pubescens grows at altitudes between 300 and
3300 m. It is a tree with broad leaves and white or pink
fragrant flowers arranged in clusters. C. pubescens has
been cultivated in several tropical regions of the world for
its quinine-containing bark and roots. Quinine was used
as a remedy to treat malaria and therefore had significant
economic importance from the 17th to the beginning of
the 20th century. In 1944, quinine was synthesized and
therefore C. pubescens lost much of its importance, but
natural quinine is still used both where the synthetic is not
available and for other medicinal purposes.
The genus Cinchona was named after the COUNTESS OF
CHINCHÓN, wife of the Viceroy of Peru, by the Swedish
botanist LINNAEUS in 1742. According to the well cited le-
gend, the countess was cured of malaria by having been
administered the bark of Cinchona in 1638 after all other
remedies failed. Although this story is not true, Cinchona
ever since was frequently used as a malaria remedy, especi-
ally distributed by the Jesuits in their world travels. Cin-
chona is the national tree of Ecuador and is on the coat of
arms of Peru.
C. pubescens has the widest natural distribution of all Cin-
chona species, from the mountainous area of central Costa
Rica and Panama and the coastal zone of NE Venezuela,
over the Andean ridge through Colombia, Ecuador and
Peru, and south to central Bolivia at altitudes between 300
and 3300, where it was also traditionally cultivated [2, 7].
Due to the massive exploitation of quinine in past centu-
ries, this formerly common tree of the western Andean slo-
pes was exploited to the point of extinction in its native
range . The C. pubescens forests had almost disappeared
as early as 1859 in Ecuador  and elsewhere .
The genus Cinchona was introduced to other tropical re-
gions mainly for the production of quinine. C. pubescens
is known to have been cultivated (and is still being cultiva-
ted) in e.g. India, Indonesia, some African countries, Sri
Lanka, and Mexico . It was also introduced to some
Pacific islands and archipelagos, like Galápagos, Hawaii,
and Tahiti, where quinine was never exploited and where
the species has become highly invasive [37, 50, 66]. Here,
its distribution ranges from 600 to 1400 m in Tahiti ,
from 180 to 860 m in Galápagos , and from 792 to
1158 m on Maui .
In its native range, C. pubescens is an evergreen tree of
10–25 m height and a diameter at breast height (DBH) of
20–80 cm [3, 7] in Ecuador, and 30 m height and a DBH
of 90 cm in northern Peru . In Ecuador, the trees
usually have a main trunk that ramifies in the upper third
and has a semicircular crown .
In its introduced range, e.g. in Galápagos, trees can reach
a height of 15 m and a DBH of 25 cm (, H. JÄGER, un-
publ. data). They have a main trunk but also often deve-
lop several equally strong trunks a short distance away
from the original trunk, which emerge by suckering of
specialized underground stems (cf. ). In this way, C.
pubescens trees take on a multi-stemmed growth form,
with the individual stems still connected . In addition,
it resprouts from fallen and cut stems to produce vertical
shoots. Contrary to its habit in its natural range in Ecua-
dor, in Galápagos the tree produces branches already in
the lowest third of the trunk, which are slightly bent up-
wards, resulting in a semicircular to cylindrical crown (H.
JÄGER, pers. observ.). This observation contradicts that of
SHIMIZU  who reported that young trees produce hori-
zontal branches in Galápagos. In Tahiti, C. pubescens rea-
ches a height of 14 m and a DBH of 30 cm, but most trees
have a DBH below 10 cm .
Fig. 2: Natural distribution of Cinchona pubescens (drawing
by W. Roloff, modified after Andersson 1998)
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Leaves, leaf coloring
C. pubescens is characterized by its membranous, large,
broad leaf blades with a thin cuticle. Leaves are opposite,
8.3–23 cm long and 5.3–21 cm wide (those of saplings and
other rapidly growing shoots often much larger than
those of flowering shoots), elliptic or ovate to suborbicu-
lar with pubescent petioles (1.2–5 cm long) and stipules
(1.2–2.6 × 0.5–1.5 cm) . Stipules are often caducous
and leave a well-marked scar on the branch . Leaf bla-
des are matt above with 7–11 pairs of conspicuous secon-
dary veins, sometimes puberulent along veins, and usually
more or less densely hairy beneath, especially on the
young leaves. Leaf domatia are usually absent but, if pre-
sent, they are distinctly pouch-shaped . The leaf mor-
phology is variable though, especially with respect to size,
shape, and indumentum of the leaf blades. Reasons for
this variation could include local differentiation and clinal
variation in response to altitude [3, 34]. This already con-
fusing variation is worsened by hybridization, mainly
with C. calisaya and C. macrocalyx . Leaves turn red
with age and are shed continuously and in small quanti-
ties throughout the year in Galápagos [10, 56].
Flowers, fruits, and seeds
In Ecuador, around Loja, Cinchona flowers and bears
fruits at the same time all year round . The flowers of
C. pubescens are clustered in large, broadly pyramidal pa-
nicles, usually up to 20 cm but sometimes longer in size.
The corollas are pinkish or purplish, paler at base
(corollas outside may be white to light pink or red in Ha-
waii and Galápagos) and are fragrant [7, 63]. It was thus
assumed that C. pubescens is insect-pollinated but this
has not been confirmed anywhere [56, 63]. The corolla
tube is 9–14 mm long, pubescent outside and glabrous in-
side. The capsules are ellipsoid to subcylindrical and 13–
41 × 5–7 mm long, opening from the base to tip when ma-
ture. Seeds are 7–12 × 2.1–2.8 mm, including the irregu-
larly dentate wings . As in the native range, C. pubes-
cens trees in Galápagos develop flowers more or less all
year round but with a peak between August and Novem-
ber. Development from the opening of the flower to the
production of mature fruits takes about 19 weeks .
Highest fruit production is between 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 .
Each capsule contains about 60–70 seeds (J. L. RENTERÍA,
pers. comm. 2004). Similar observations are reported
from India, where flowers and fruits are found almost all
year round . In India, one gram of Cinchona seeds (in-
cluding C. pubescens) contains about 300 to 400 seeds.
Fig. 3: Leaves in Galápagos
Fig. 5: Flowers in Galápagos
Fig. 4: Flower panicles in Galápagos
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Wood and bark
Since the focus of research on Cinchona has always been on
the quinine content in the bark, there is only little informa-
tion available on its wood characteristics. In its natural
range, the wood of C. pubescens is brown-yellowish to
brown-orange, compact and of a fine texture , and has a
density of 0.54 g cm–3 [11, 69]. The outer bark is 3–20 mm
thick and light coffee-brown, turning cinnamon-brown
when dry. It may have white streaks caused by lichens gro-
wing on it [3, 42]. When the bark is being peeled off the
trunk, a milky sap leaks from the cuts and the cortex below
takes on a reddish color (hence the name “red bark”), which
quickly darkens. The fresh and dry bark has the very bitter
taste characteristic for all Cinchona species .
In several drawings, ACOSTA SOLÍS  describes adventi-
tious roots of Cinchona species in Ecuador as “stilt
roots”, which are either covered by the soil or exposed.
However, this observation does not concur with observati-
ons from Galápagos, where these roots are rather simple
and weak .
Taxonomy, genetic differentia-
tion, and hybridization
The genus Cinchona L. was first described by LACON-
DAMINE in 1738 . Since this genus was medicinally
important, it has historically attracted extraordinary
attention from taxonomists. Many different names
were established in the 19th century on the grounds of
very minor morphological differences [29, 54]. In his
revision of the genus Cinchona, ANDERSSON  consi-
dered more than 330 names (including many at variety
level) and recognized 23 species. The taxonomy of C.
pubescens is especially difficult since it frequently hy-
bridizes with other Cinchona species where they occur
together in nature [3, 7, 9, 29]. This confusion was al-
ready noticed by VON HUMBOLDT in 1808, who stated
that one might consider Cinchona leaves from the
same branch as being specimens of different species if
one did not have the opportunity to see the branch in
the wild . In 1946, four species were known to form
hybrids with C. pubescens , whereas ANDERSSON 
lists 7 species hybridizing with it (C. barbacoensis, C.
calisaya, C. lancifolia, C. lucumifolia, C. macrocalyx,
C. micrantha, C. officinalis). Hybrids between C. pu-
bescens and C. calisaya are the most commonly found
in nature and seem also to have been produced in culti-
The chromosome number of C. pubescens is 2n = 34 .
Fig. 6: Fruits in Galápagos with Ecuadorian 10-cent coin,
equal to the size of a US 10-cent coin (ca. 1.8 cm)
Fig. 8: Seeds (scale is in cm), in Galápagos
Fig. 7: Opened fruits in Galápagos
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Growth, development, and yield
In its natural range in Bolivia, a Cinchona tree reaches a
height of about 4 m and a diameter of about 15 cm when
6 years old, and over 30 cm in diameter when 10 or 12
years old . Cinchona trees in India flower after the
fourth year , whereas C. pubescens trees in Galápagos
flower when only 2 years old, with a DBH of 1.5 cm (H.
JÄGER, pers. observ.). A C. pubescens tree in Bolivia pro-
duces about 7–9 kg of seed per season, which are harve-
sted in November and December. In Jamaica, young seed-
lings of C. pubescens grow about 1 m during the first year
and reach about 1.8 m at the end of the second year, with
a DBH of 3.6 cm . The maximum DBH of C. pubes-
cens trees found in Tahiti was 30 cm  and 10-year-old
trees had a DBH of 5 cm .
In Galápagos, C. pubescens demography was studied over
7 years in the Fern-Sedge highland vegetation zone of
Santa Cruz Island, which is largely invaded by C. pubes-
cens . Results showed that the density of C. pubescens
trees larger than 1.5 m increased from 123 per hectare in
1998 to 439 in 2005. Concurrently, the number of stems
per hectare increased from 355 to 1652, which represents
an average number of stems per tree of 2.9 and 3.8, res-
pectively. These results parallel those of SHIMIZU , who
counted 3.4 stems per tree in an area nearby.
Fig. 11: Growth forms and habitus of Cinchona pubescens in
Galápagos (drawing by W. ROLOFF, modified after
Fig. 9: Tree trunk (overgrown with mosses and liches), in
Fig. 10: Part of trunk cut open to show the wood, in
Galápagos. Note the red coloration after cutting
the trunk, hence the name „red guinine”
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At the same time, basal area increased from 1.0 to 4.2 m2
ha1, while mean C. pubescens cover increased from 6.6 to
16.4 % .
In Tahiti, C. pubescens reached a maximum density of
9200 stem ha–1 and 14,500 seedlings ha–1 . However,
these data are from small study plots in the most invaded
areas of Tahiti (J.-Y. MEYER, pers. comm. 2010).
In Bolivia, the bark of C. pubescens is thicker and heavier
when older, but does not contain as much quinine. It is
stripped between October and January, producing about
2 kg on average . The bark is harvested after 6 to 10
years in Ecuador  and after 8 to 12 years in Jamaica
. The bark of Cinchona in India is harvested after the
third or fourth year from trees that are uprooted to thin
the plantation. A second harvest can be carried out on the
remaining trees after the seventh or eighth year and a final
harvest after about 12 years . In commercial Cinchona
plantations in Peru, 2860 kg of C. pubescens bark was
harvested between 1943 and 1945 and sold at a price of
US $ 109,500 .
In Ecuador, naturally occurring C. pubescens in the
province of Bolívar and on the western slopes of the
Andes produces the highest alkaloid content [2, 49].
However, the alkaloid content varies considerably with
locality, from 0.1 to 7.8 % [3, 60], and with the age of
the tree and the part of the tree the samples were taken
from. Younger trees and thicker barks contain more al-
kaloids . This especially applies to the quinine con-
tent, which is up to 5 % in the wild form collected
from the slopes of Chimborazo in Ecuador, whereas
the wild form from Costa Rica contains hardly any
. The bark of trunks, twigs, and roots of cultivated
C. pubescens in India, Java and Sri Lanka yields bet-
ween 4–8 % of total alkaloids, of which 0.8–1.4 % is
quinine [12, 60]. The alkaloid content of C. pubescens
in India is highest from the fourth to seventh year and
then declines again . The same is true for the alka-
loid content for C. pubescens cultivated in Bolivia .
A 16-year study in Java showed that the quinine con-
tent of Cinchona clones increased in the early years
and then declined again in the fifth or seventh year.
Manuring maintained the quinine content or reduced
the rate of decline in Java . The quinine content of
the bark from the same seedling variety or clone varies
with the soil and environmental conditions and can be
increased by stunting the vertical growth .
In 1891, C. pubescens was the species of choice for culti-
vation and provided the largest amount of bark, especially
in Sri Lanka and Java . Nowadays, the main Cinchona
alkaloid producing countries are Indonesia, the Democra-
tic Republic of Congo, Tanzania, Kenya, Rwanda, Sri
Lanka, Bolivia, Colombia, Costa Rica, and India. These
countries produce between 400–700 tons of alkaloids ob-
tained from 8000–10,000 tons of bark produced annually
[20, 35]. The bark yield in India alone is about 9000–
16,000 kg ha-1 .
Fig. 12: Young seedlings in Galápagos
Fig. 13: Dense C. pubescens stand with native vegetation in
the understorey in Galápagos
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Due to the overexploitation of Cinchona before the synthesis
of quinine in 1944, Cinchona species have drastically been re-
duced in their abundance in South America, e.g. in Bolivia,
Colombia and Ecuador [9, 32]. As a consequence, C. pubes-
cens is now extremely rare in Ecuador and has a low regene-
ration rate (, C. I. ESPINOSA, pers. comm. 2010). It used to
grow best at an altitude of 850–1350 m, in areas with steep
slopes and in disturbed habitats . C. pubescens was also
cultivated here and the plantations were most successful in
the province of Bolívar, in an area where other trees had been
felled and burnt with the rest of the vegetation . A type of
C. pubescens with high quinine yield was cultivated in small
plantings in the Andean foothills of central Ecuador . In
1860, seeds and seedlings of this type were taken to India, Sri
Lanka, and Guatemala [49, 54].
In its native and introduced range, Cinchona can be propaga-
ted through seeds and vegetative means but most commercial
plantations are raised by seeds, and methods used were simi-
lar [12, 23, 39]. Typically, Cinchona seedlings are produced in
raised seedbeds in soil mixed with leaf compost and manure.
Seeds are thickly spread over the bed, for example, 2 g m-2.
The seeds are covered with a thin layer of soil under partial
shade provided by roofing. The sown beds are kept moist and
seeds germinate in 10–40 days, with a germination rate vary-
ing between 50 and 85 %. This way, 400 g of seeds can pro-
duce about 100,000 seedlings. After three months the seed-
lings have grown a pair of leaves and are planted out in rows
with sufficient spacing (e.g. 5 × 5 cm). When seedlings are
10 cm tall, bearing three to four pairs of leaves, they are sepa-
rated at 10–12 cm spacing. The partial shade can be removed
after 3 months to expose the plants to the sun for the next
four months. This way, it takes about 16 months to obtain
seedlings of 20–25 cm in height, which are ready to be plan-
ted in the field at 1–2 m spacing. Trees are thinned after the
third year for extracting the bark, leaving 50 % of the original
number of trees at the end of the fifth year. In India, about
3000 C. pubescens individuals are planted per hectare, leaving
space in between for the planting of shade trees, like Crotala-
ria anagyroides and Tephrosia candida which also serve as
nitrogen fixers or for Alnus nepalensis and Aleurites spp.,
also as shade trees. In addition, about 100 kg each of nitrogen
and phosphorus are applied per hectare as well as other
Greenhouse experiments in Puerto Rico revealed that C.
pubescens grows best at temperatures ranging from 21 to
27 °C and that the total alkaloids and quinine content in
roots and stems are also higher compared to cultivation at
lower temperatures . However, seedling survival is lo-
wer since leaves grow larger at these temperatures which
apparently resulted in death of the weaker plants .
Seeds from India begin to lose their viability after six to
eight weeks and lose it completely after one year .
Seeds in Galápagos also lost their viability after approxi-
mately one year .
Abb. 14: C. pubescens trees in the natural range: Botanical
Garden of the Universidad National de Loia, Ecua-
dor, 2160 masl (approx. height: 7 m)
Fig. 15: New shoots emerging from a cut Cinchona pubes-
cens tree in Galápagos
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In some countries like India, Sri Lanka, and Guatemala,
C. pubescens is propagated using vegetative techniques,
such as grafting, budding, and softwood cuttings, which
are safer than propagation by seedlings [12, 39, 41]. The
best wood for cuttings is the thin wood of the current
year’s growth . The cuttings are taken just below the
joint or point where a pair of leaves originated. They are
then put out in thatched beds as already described for
seedlings. They can also be grown in boxes about 5 cm
deep, filled with a layer of vegetable mould mixed with
sand. A layer of pure sand above the mould promotes
drainage, since cuttings are apt to rot off at the level of
the soil if not thoroughly drained. These cuttings form
new roots in two to four months, according to season and
temperature . Even though C. calisaya yields more
quinine, C. pubescens is often the predominant species in
plantations because it is hardier, easier to propagate, and
has a much wider range of growth conditions . There-
fore, it is used as a root stock for the grafting of C. cali-
saya (e.g. in Indonesia) because of its hardy and vigo-
rously growing nature at lower elevations . Hormonal
treatments help to induce better rooting . Nowadays,
the propagation of C. pubescens is much easier. Twenty
thousand shoots of C. pubescens can be obtained from
the apical meristem of a plant by micropropagation with-
in one year. However, this way the root development of
the shoots is often inadequate so that the transfer of the
shoots to the soil can be complicated .
Habitat and climate requirements
Generally, in its natural range, Cinchona trees need warm
climates with high precipitation and humidity almost all
year round for optimal growth [2, 22]. In Ecuador, tempe-
ratures vary between 10 and 23 °C and C. pubescens often
grows in steep gorges that are difficult to access and in di-
sturbed habitats . Soils are volcanic and rich in organic
matter, nitrogen and phosphorus, but poor in potassium
, are warm (16.3 °C on average), and slightly acid (pH
5.1 ). Trees also grow well in clay soils and in very
rocky areas, where the roots are exposed to the air .
Soils around Loja in southern Ecuador, where several Cin-
chona species grow, mainly comprise decomposed mica-
ceous schist and gneiss, covered by a layer of vegetable
mould . In Bolivia, cultivated C. pubescens trees grow
best on the sides of valleys or ridges of the Andes at alti-
tudes of about 900–1200 m. They will also grow up to
2400 m in altitude but then acquire a stunted growth form
and will not yield much quinine .
Experimental investigations in Guatemala, India and Java
showed that Cinchona grows best in well-drained soils with
high organic matter, high nitrogen content, and low car-
bon/nitrogen ratio. These soils are also usually rich in citric-
soluble phosphate, lime, and hydrolytic acidity and trees de-
veloped well in gentle as well as steep slopes with protection
against erosion [12, 13, 41, 54]. In India, C. pubescens grows
best at elevations between 460 and 2000 m asl, where it with-
stands both high humidity and dry conditions [12, 16]. As in
its native range, C. pubescens grows better in recently cleared
forests than in grasslands . In Tahiti, C. pubescens has its
growth optimum between 600 and 950 m elevation in secon-
dary and primary rainforests  but was also found at up to
1400 m elevation in undisturbed cloud forest . In São
Tome and Príncipe and in Java it grows best at elevations of
650–1350 m and 1250–1950 m, respectively . Optimal gro-
wing temperatures in India are 21–24 °C and should not go
below 8 °C in winter and 30 °C in summer .
In Ecuador, C. pubescens does not usually occur as isola-
ted individual trees nor as a forest but in small groups of
trees, in “patches” [4, 22]. This was already the case in the
area of Loja in 1737 and in other parts of the country, and
was attributed to the cutting of large trees [25, 62]. C. pu-
bescens is not a dominant tree species and has almost
been brought to extinction through overexploitation du-
ring the 200 years up to World War II . It is now consi-
dered endangered in its natural range in the south of
Ecuador . In contrast, in northern Peru, C. pubescens
occurs as a dominant forest tree where it can reach a
height of up to 30 m and a DBH of 90 cm .
In its introduced range, C. pubescens is much more com-
petitive in its growth and has become invasive in some
areas, meaning that it successfully reproduces and spreads
without human intervention. This is especially the case in
Galápagos, Hawaii, and Tahiti, where it outshades and re-
duces the cover of native plant species [19, 36, 37, 65].
Cultivation of Cinchona was initiated in Hawaii in 1868
and, by 1978, C. pubescens had naturalized . In
Galápagos, C. pubescens was introduced in the 1940s [28,
45] and had naturalized by 1972 . In Tahiti, C. pubes-
cens was introduced in 1938  and has naturalized since
the 1970s . C. pubescens has also been reported to oc-
cur outside of plantations in Java and New Guinea , in
Jamaica , Guatemala , and on St Helena .
Biological characteristics of C. pubescens in its introduced
range that make it a good invader include its abundant seed
production, e.g. in Hawaii and Galápagos. However, its light,
wind-borne seeds do not seem to be dispersed very far from
the parent tree. The maximum dispersal distance in Galápa-
gos was 15 m  and in Hawaii it was reported that the spe-
cies does not disperse further than 100 m from the parent tree
. The successful invasion of C. pubescens on both archi-
pelagoes was a slow but continuous one. This is reflected in
the long time that passed between the introduction of the
species and the recognition that it had naturalized (over 100
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years in Hawaii, and about 30 years in Galápagos [28, 63]).
The youngest mature trees were observed in Galápagos,
where 2-year-old seeding trees of 1.8 m height with a DBH of
1.5 cm produced fruit (H. JÄGER, pers. observ.). Another inva-
sive characteristic of C. pubescens is the ability of the seeds to
germinate in dense understorey vegetation as well as under a
dense C. pubescens canopy, as in Hawaii and Galápagos [52,
63]. Last but not least, C. pubescens has a rapid vegetative re-
production by vigorous suckering from roots and stems [37,
46]. In addition, it demonstrates fast growth in its introduced
range: about 1 m per year in Galápagos and 1–2 m elsewhere
(, H. JÄGER, pers. observ.).
C. pubescens also may benefit from increased nutrient uptake
due to its association with arbuscular mycorrhizal fungi,
which are mainly lacking in native species in Galápagos .
In Ecuador, Cinchona species tend to be associated with
certain other tree species, like Palicourea spp., Joosia pul-
cherrima, Hoffmania ecuatoriana, Hieronyma macro-
carpa, Ocotea spp., and Weinmannia fagarioides . Old
trunks of C. pubescens are generally covered with moss,
liverwort, and lichen species , and this is also the case
in Galápagos (mainly Frullania and Omphalanthus spe-
cies, F. ZIEMMECK, pers. comm. 2007). In Peru at an eleva-
tion of about 2445 m, C. pubescens is mainly associated
with various Miconia spp., Clusia spp., Weinmannia pin-
nata, Hedyosmum scabrum, Chusquea scandens, Ilex sp.,
and Hesperomeles sp. Below this elevation in the dwarf
forest with short vegetation (1.5 m), C. pubescens is asso-
ciated with other trees of the genera Clusia, Befaria,
Wemmannia, and Brunellia .
The databases of the Systematic Mycology and Microbio-
logy Laboratory lists 27 fungi species as associated with
C. pubescens but 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 JENKINS was also recorded from C. pubescens
from western Ecuador (H. C. EVANS, pers. comm. 2009).
However, there is no evidence that these species are im-
portant economic pathogens .
In plantations in Guatemala, India, and Java, Cinchona is
susceptible to many pests and diseases. Cinchona seed beds
are frequently affected by the fungi Rhizoctonia solani J. G.
KÜHN, Phytophthora spp. (root rot fungus), and Rosellinia ar-
cuata PETCH (black-root fungus), causing a disease known as
“damping off” [12, 13, 54]. These fungi penetrate the seed-
lings through their roots and cause sudden wilting and rot-
ting of seedlings. In Java and India, plants were also attacked
by the insects Helopeltis (tea mosquito) and Pachypeltis (leaf
scorch) [12, 13]. Cinchona stem bark disease due to Phytoph-
thora cinnamomi RANDS is a major disease in central African
countries (Rwanda and the Democratic Republic of Congo)
. In Galápagos, secondary pathogens were isolated from
C. pubescens, mainly Fusarium spp. and Botryodiplodia the-
obromae PAT. (H. C. EVANS, pers. comm. 2009).
Because of its alkaloids, Cinchona is one of the most wi-
dely produced and traded medicinal plants , and the
alkaloids extracted from C. pubescens, “red bark”, and C.
calisaya, “yellow bark”, are commercially most important
[7, 20]. Chemical components of interest extracted from
the bark of the trunk (but also of twigs and roots) are
quinine, quinidine, cinchonidine, and cinchonine [12, 20].
The most important is quinine, which, in the form of qui-
nine salts, like sulfate, bisulfate, hydrochloride, and dihy-
drochloride, was used for the prevention and treatment
of malaria  before the chemical synthesis of quinine
was achieved by the American chemists WOODWARD and
DOERING in 1944 .
There are three common ways of harvesting Cinchona bark:
uprooting, coppicing, and mossing. In the first case, the com-
plete tree is uprooted and the bark stripped off the trunk .
When coppicing a tree, the trunk is cut close to the ground,
thereby inducing the production of basal sprouts to produce
a second crop of bark . However, the trees recover slowly
from this treatment and the bark does not contain as much
alkaloid as first-stem bark . Since it was observed that the
best Cinchona was always covered with moss, in “mossing”
the bark was stripped off the trunk and the bare trunk subse-
quently covered with moss. This system was practiced in Java
and Sri Lanka, and induced the cambium to produce a se-
cond bark which contained up to three times higher alkaloid
content than the first [23, 60]. A fourth approach for the bark
harvest was used in Loja (Ecuador) where people stripped the
bark from standing tall trees as far as they could reach. Then
they made steps by tying sticks to the tree with ropes so they
could harvest the bark further up until they reached the top
of the tree . Apparently, this practice was limited and fel-
ling the Cinchona trees was much more common, leading to
the near-extinction of Cinchona species (especially C. pubes-
cens) in that area [4, 32].
For commercial distribution, the stems and roots are cut
into pieces, the bark is then separated, dried, graded,
packed, and traded . In Bolivia, the bark is placed in
paved yards and generally dries in four days to three
weeks, depending on the weather conditions . As early
as 1737 it was noted that the Cinchona with the “red
bark” from Loja (probably C. pubescens) has a much bet-
ter curative effect when used fresh and green, and turns
worthless when dry and long kept .
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The wood of Cinchona also contains alkaloids and there-
fore was the preferred building material for houses in Sou-
thern Ecuador, since it was one of the more termite-resi-
stant woods . Even though the wood is of inferior qua-
lity in Galápagos, it is now increasingly used for construc-
tion, since alternative wood is getting scarce due to over-
exploitation or governmental regulations (H. JÄGER, pers.
Since World War II, the main part of the quinine yield has
gone towards the production of pharmaceutical com-
pounds, like antipyretic drugs against colds, flu, and rheu-
matism. The remaining part in the form of quinine hydro-
chloride and quinine sulfate is used in the beverage indu-
stry, mainly as a bitter flavor in tonic water and bitter le-
mon [12, 20] and might still have some prophylactic value
against malaria. Quinidine is used as a cardiac depressant
and its sulfate form is used for the treatment of cardiac
arrhythmia . Today, between 400–700 tons of alka-
loids are obtained from 8000–10,000 tons of bark produ-
ced annually [20, 35]. In 2000, 1 kg of quinine cost ca. US
$ 500 and quinidine ca. US $ 1500 because of its naturally
lower occurrence .
C. pubescens has long been recognized as a potential risk
to native vegetation in its introduced range, especially in
Hawaii, Galápagos, and Tahiti. In the Galápagos National
Park, several control methods to combat C. pubescens
have been implemented. Manual methods include felling
and uprooting of trees, as well as hand-pulling of smaller
plants, but trees resprout from cut stumps [8, 46]. Chemi-
cal control methods consist of applying herbicides by a
range of means: hack and squirt, basal bark, cut stump,
girdle and squirt, branch filling, tree injections, and foliar
spray . However, most of these methods were ineffec-
tive in the long run. Eventually, a hack and squirt techni-
que was developed in which a mixture of picloram and
metsulfuron was applied to connecting machete cuts
around the circumference of tree trunks, which killed 73–
100 % of the trees. Smaller shoots and saplings were pul-
led out by hand . This method is now successfully ap-
plied for C. pubescens control by the Galápagos National
Park Service on a small scale, as well as the uprooting of
trees in conservation priority areas . Control of C. pu-
bescens would be difficult in Hawaii (mainly Maui), since
it primarily grows in steep gulches and thick understorey
vegetation. However, control of small populations in
spots of native vegetation could prevent further spread
and help to slow down the degradation of the remaining
native forest in the area .
Pathogens attacking C. pubescens in its native range in
Ecuador have been isolated but it remains to be seen
whether these might be potential biological control agents
(H. C. Evans, pers. comm. 2009).
History of discovery and cultivation of
The early history of the use of quinine is obscure and
the question of whether the indigenous people of South
America knew about its curative properties remains
controversial. Some authors argue that quinine was used
to treat different fevers (not malaria) in Peru and Ecua-
dor before the arrival of the Spaniards [32, 47] and some
argue that the people did not know about its fever-redu-
cing characteristics [33, 62]. However, GRAY et al. 
reports that it was the current opinion in Loja (Ecua-
dor) that the qualities and uses of Cinchona bark were
known by the indigenous people before the arrival of
the Spaniards and that it was applied by them to cure
“intermittent fevers”, which were “… frequent over all
that wet unhealthy country” .
The way quinine made its way to Europe is another my-
stery. The most frequently cited version is that the
COUNTESS OF CHINCHÓN returned to Spain in 1642 and
brought with her a stock of Cinchona bark to cure mala-
ria there . This was reason enough for LINNAEUS to
name the genus Cinchona in her honor in 1742, erro-
neously leaving out an “h” when doing so . However,
this version is not true since the first wife of the Viceroy
died before she left Spain for Peru and his second wife
died in Colombia before she could return to Spain . In
addition, the official diary of the Viceroy, kept by his
secretary, was found in 1930 and revealed that his second
wife had never fallen ill with a fever .
Even the point of time when Cinchona was brought to
Europe is not clear, since the bark of the Peruvian bal-
sam tree (Myroxylon peruiferum) had been shipped to
Rome at the beginning of the 17th century under the
name “quina-quina” (Quechua word meaning “bark of
barks”)  and the actual Cinchona bark was only
exported to Europe from 1631  or the mid-1600s
 onwards. Cinchona was superior to Peruvian bal-
sam in its feverreducing qualities [32, 40]. The Augusti-
nian and Jesuits were apparently really the first to
bring Cinchona bark to Europe and distribute it as a
remedy against malaria (hence the name “Jesuit’s
bark” ). In 1677, Cinchona bark entered the Lon-
don Pharmakopoeia as “Cortex Peruvianus” (Peruvian
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Cinchona was the first effective treatment for malaria, and
in 1820 PELLETIER and CAV E N TO U isolated quinine and cin-
chonine from the Cinchona bark, of which quinine was
the main antimalarial ingredient . After this successful
isolation, quinine extract quickly replaced the untreated
Cinchona preparations in the treatment of malaria. Sup-
plies of quinine were limited and the need was great. Ma-
laria was affecting millions of people in the Asiatic tropics
and in Africa, and the imperialistic activities of Europe-
ans in these areas required high amounts of quinine,
which prompted the cultivation of Cinchona [21, 54].
It is notable that 200 years passed between the discovery
of the genus Cinchona and its cultivation. The reasons are
likely the rather difficult cultivation of the tree and the in-
itially abundant supplies of wild bark, sufficient to meet
demands in Europe and America . Several investiga-
tors visiting the natural sources of Cinchona in the Andes
had pointed out that the trees were in danger of extermi-
nation as a consequence of overexploitation [23, 25, 54].
VON HUMBOLDT reported that 25,000 trees were cut an-
nually in the Andes in 1795 . The Dutch were the first
in reacting to the increase in the demand for the Cinchona
bark by sending the German botanist HASSKARL to the
Andean region in 1852 to obtain Cinchona seeds . The
British followed shortly after by sending SPRUCE into the
forest of the western slopes of Mount Chimborazo in
Ecuador to collect Cinchona seeds and plants (also of C.
pubescens) to start a plantation in India in 1860 .
MARKHAM also investigated the Cinchona-producing
areas in South America and collected plants and seeds for
these plantations . In 1865, LEDGER obtained seeds
from a superior Cinchona strain (C. calisaya), enabling
the Dutch to assume world dominance in quinine produc-
tion from the 1890s onwards through their plantations in
Java [54, 55]. Before World War II, about 90 % of the
world supply of Cinchona bark was produced in Java
. When Java was occupied by the Japanese in 1942,
natural quinine became very scarce, which led to the deve-
lopment of a synthetic production of quinine alkaloid in
Significance of Cinchona today
The need for effective antimalarial drugs has persisted
over the nearly two centuries since quinine was first isola-
ted . The evolution of strains of the malaria parasites
resistant to the synthetic drugs (mainly chloroquine), and
the consequent rise in the malaria incidence in Asia and
Africa, have revitalized interest in quinine and in the anti-
malarial properties of the other alkaloids of Cinchona
[21, 35]. Other natural antimalarials in the bark’s extract,
such as quinidine, appear to potentiate the effects of qui-
nine . This, coupled with the increasing use of qui-
nidine as an antiarrhythmic compound, has increased the
demand for Cinchona bark .
 ANONYMOUS, 1889: Bulletin of Miscellaneous Information
(Royal Gardens, Kew) 34, 244–247.
 ACOSTA SOLÍS, M., 1945a: Habitat y distribución de las Cin-
chonas en el Ecuador. Flora. Revista al servicio de las Cien-
cias Naturales y Biológicas, Instituto Ecuadoriano de Cien-
cias Naturales 6, 15/16, 9–18.
 ACOSTA SOLÍS, M., 1945b: Botánica de las Cinchonas. Flora.
Revista al servicio de las Ciencias Naturales y Biológicas, In-
stituto Ecuadoriano de Ciencias Naturales 6, 15/16, 29–55.
 ACOSTA SOLÍS, M., 1946: Cinchonas del Ecuador. Editorial
del Ecuador, Quito.
 ACOSTA SOLÍS, M., 1961: Los bosques del Ecuador y sus
productos. Editorial del Ecuador, Quito.
 ALONSO, L. E.; ALONSO, A.; SCHULENBERG, T. S.; DALLMEIER,
F. (eds.), 2001: Biological and social assessments of the
Cordillera de Vilcabamba, Peru. RAP Working Papers
12 and SI/MAB Series 6, Conservation International,
 ANDERSSON, L., 1998: A revision of the Genus Cinchona
(Rubiaceae-Cinchoneae). Memoirs of the New York Bota-
nic Garden, Volume 80.
 BUDDENHAGEN, C. E.; RENTERÍA, J. L.; GARDENER, M.; WIL-
KINSON, S. R.; SORIA, M.; YÁNEZ, P.; TYE, A.; VALLE, R.,
2004: The control of a highly invasive tree Cinchona pubes-
cens in Galapagos. Weed Technology 18, 1194–1202.
 CAMP, W. H., 1949: Cinchona at high altitudes in Ecuador.
Brittonia 6, 394–430.
 CEDEÑO, W. E., 1990: Estudio fenológico de especies vegeta-
les introducidas y nativas en la Isla Santa Cruz, Galápagos.
Tesis de Ingeniería Forestal, Universidad Téchnica de Esme-
 CHAVE, J.; COOMES, D.; JANSEN, S.; LEWIS, S. L.; SWENSON,
N. G.; ZANNE, A. E., 2009: Towards a worldwide wood eco-
nomics spectrum. Ecology Letters 12, 351–366.
 CHOPRA, V. L.; PETER, K. V., 2005: Handbook of industrial
crops. Haworth Press, London.
 COSTER, C., 1942: The work of the West Java Research In-
stitute in Buitenzorg. Empire J. Exper. Agriculture 10,
 CRONK, Q. C. B.; FULLER, J. L., 1995: Plant invaders. The
threat to natural ecosystems. Chapman and Hall, London.
 DEL TREDICI, P., 2001: Sprouting in temperate trees: a mor-
phological and ecological review. Botl. Rev. 67, 121–140.
 ENGLER, A., 1896: Empfehlung der Anlage von Cinchona-Plan-
tagen im Kamerungebirge. Notizblatt des Königlichen Botani-
schen Gartens und Museums Berlin, 6, Band 1, 186–189.
 FARR, D. F.; ROSSMAN, A. Y., 2010: Fungal Databases, Syste-
matic Mycology and Microbiology Laboratory, ARS,
USDA. http://nt.ars-grin.gov/fungaldatabases/ (last acc.: 18
 FERREIRA, J. F. S., 2004: Artemisia annua L.: the hope against
malaria and cancer. Medicinal and aromatic plants: produc-
tion, business and applications. Proc. Meeting Jan 15–
17/2004. Mountain State Univ. Beckley, WV.
ArtemisiavsMalariaandCancer.pdf (last acc.: 18 Oct. 2010).
15_III-4_Cinchona_pubescens_58EL:00_Musterseite_NEU 13.05.2011 14:46 Uhr Seite 11
12 Enzyklopädie der Holzgewächse – 58. Erg.Lfg. 06/11
 FISCHER, L. K.; VON D ER LIPPE, M; KOWARIK, I., 2009: Tree
invasion in managed tropical forests facilitates endemic spe-
cies. J. of Biogeography 36, 2251–2263.
 FRANCKENPOHL, J., 2000: Entwicklung von Methoden zur
diastereoselektiven Funktionalisierung der Cinchona Alka-
loide und Synthese neuartiger funktionalisierter Chinucli-
dine. Doctoral thesis, Hannover Univ., Germany.
 GAL, J., 2006: Chiral drugs from a historical point of view,
1–26. In: FRANCOTTE, E.; LINDNER, W. (eds): Chirality in
Drug Research, Wiley-VCH Verlag, Weinheim.
 GARMENDIA SALVADOR, A., 2005: El árbol de la Quína (Cin-
chona spp.). Distribución, caracterización de su habitat y
arquitectura. Editoral Universidad Técnica Particular de
Loja, San Cayetano Alto, Ecuador.
 GIBBS. M., 1885: Cultivation of Cinchona in Bolivia. Am. J.
Pharmacy (1835–1907), 57, Botanical Medicine Monogra-
phs and Sundry. http://www.swsbm.com/AJP/AJP.html (last
acc. 18 Oct. 2010).
 GISP, 2009: Global Invasive Species Programme, database.
=1&sts=sss&lang=EN (last acc. 18 Oct. 2010).
 GRAY, J.; ARROT, W.; MILLER, P., 1737: An account of the Pe-
ruvian or Jesuit’s Bark. Philosophical Transactions (1683–
1775) 40, 81–86.
 GÜNTER, S.; STIMM, B.; WEBER, M., 2004: Silvicultural contribu-
tions towards sustainable management and conservation of fo-
rest genetic resources in Southern Ecuador. Lyonia 6, 1, 75–91.
pdf (last acc. 18 Oct. 2010).
 HAGGIS, A. W., 1941: Fundamental errors in the early hi-
story of Cinchona. Bull. History of Medicine 10, 417–459.
 HAMANN, O., 1974: Contribution to the flora and vegeta-
tion of the Galápagos Islands. III. Five new floristic records.
Botaniska Notiser 127, 309–316.
 HARLING, G.; ANDERSSON, L. (eds.), 1994: Rubiaceae – Cin-
choneae – Coptosapelteae. Flora of Ecuador 50, 1/4, 162.
 HOBHOUSE, H., 1986: Seeds of Change: Five plants that
transformed mankind. Harper & Row, New York.
 HODGE, W. H., 1948: Wartime Cinchona procurement in
Latin America. Economic Botany 2, 229–257.
 HONIGSBAUM, M., 2003: The fever trail. In search of the
cure for malaria. Macmillan, New York.
 HUMBOLDT, VO N F. W. H. A., 1808a: Über die Chinawälder
in Südamerika, 1. Die Gesellschaft Naturforschender
Freunde zu Berlin, Magazin für die neuesten Entdeckungen
in der gesamten Naturkunde 1, 57–68.
 HUMBOLDT, VO N F. W. H. A. 1808b: Über die Chinawälder
in Südamerika, 2. Der Gesellschaft Naturforschender
Freunde zu Berlin Magazin für die neuesten Entdeckungen
in der gesamten Naturkunde 1, 104–120.
 HUSAIN, A., 1991: Economic aspects of exploitations of me-
dicinal plants. In: AKERELE, O.; HEYWOOD, V.; SYNGE, H.
(eds): Conservation of medicinal plants. Cambridge Uni-
versity Press, Cambridge.
 JÄGER, H.; TYE, A.; KOWARIK, I., 2007: Tree invasion in na-
turally treeless environments: impacts of quinine (Cinchona
pubescens) trees on native vegetation in Galápagos. Biologi-
cal Conservation 140, 297–307.
 JÄGER, H.; KOWARIK, I.; TYE, A., 2009: Destruction without
extinction: long-term impacts of an invasive tree species on
Galápagos highland vegetation. J. Ecology 97, 1252–1263.
 JÄGER, H.; KOWARIK, I., 2010: Resilience of native plant com-
munity following manual control of invasive Cinchona pube-
scens in Galápagos. Restoration and Ecology 18, 103–112.
 JOY, P. P.; THOMAS, J.; MATHEW, S.; SKARIA, B. P., 2001: Me-
dicinal plants, 449–632. In: BOSE, T. K.; KABIR, J.; DAS,
P.; JOY, P. P. (eds.): Tropical Horticulture, Volume 2, Naya
 KEEBLE, T. W., 1997: A cure for the ague: the contribution
of ROBERT TALBOR (1642–81). J. Royal Society of Medicine
 KING, G., 1880: A manual of Cinchona cultivation in India.
Superintendent of Government Printing, Calcutta, India. Ac-
cess through Open Library. http://openlibrary.org/
(last acc. 18 Oct. 2010).
 KLOTZSCH, J. F., 1857: Über die Abstammung der im Han-
del vorkommenden rothen Chinarinde. Abhandlungen der
Königlichen Akademie der Wissenschaften zu Berlin, 51–75.
 KOBLITZ, H.; KOBLITZ, H.; SCHMAUDER, H. P.; GROGER, D.,
1983: Studies on tissue cultures of the genus Cinchona L.
Plant Cell Reports 2, 95–7.
 KUNTZE, O., 1878: Cinchona. Arten, Hybriden und Cultur
der Chininbäume. Monographische Studie nach eigenen
Beobachtungen in den Anpflanzungen auf Java und im Hi-
malaya. H. Hässel, Leipzig.
 LUNDH, J. P., 2006: The farm area and cultivated plants on
Santa Cruz, 1932–1965, with remarks on other parts of Ga-
lapagos. Galapagos Research 64, 12–25.
 MACDONALD, I. A. W.; ORT I Z , L.; LAWESSON, J. E.; NOWAK,
J. B., 1988: The invasion of highlands in Galápagos by the
red quinine-tree Cinchona succirubra. Environmental Con-
servation 15, 215–220.
 MARKHAM, C. R., 1862: Travels in Peru and India. John
 MARKHAM, C. R., 1874: A Memoir of the Lady Ana de
Osorio, Countess of Chinchon and Vice-Queen of Peru
(A.D. 1629–39), with a Plea for the Correct Spelling of the
Chinchona Genus. Trübner & Co, London.
 MARTIN, W. E.; GANDARA, J. A., 1945: Alkaloid content of
Ecuadorian and other American Cinchona barks. Botanical
Gazette 2, 184–199.
 MEYER, J.-Y., 2004: Threat of invasive alien plants to native
flora and forest vegetation of Eastern Polynesia. Pacific
Science 58, 357–375.
 MEYER, J.-Y.; MALET, J.-P., 2000: Forestry and agroforestry
alien trees as invasive plants in the Pacific Islands. Paper,
FAO Workshop on Forestry Data Collection for the Pacific
Region, 4–8 September 2000, Apia, Samoa.
918_112565 (last acc. 18 Oct. 2010).
 PALACIOS, J. H., 1993: Efecto de Cinchona succirubra
KLOTZSCH sobre la comunidad de Miconia robinsoniana
COGN. en la Isla Santa Cruz, Galápagos. Tesis de Licencia-
tura, Universidad Central del Ecuador, Quito, Ecuador.
 PIER, Pacific Island Ecosystems at Risk, 2009: Weed risk as-
sessment for Cinchona pubescens. http://www.hear.org/pier
/wra/pacific/cinchona_pubescens_htmlwra.htm (last acc. 18
 POPENOE, W., 1949: Cinchona cultivation in Guatemala: a brief
historical review up to 1943. Economic Bot. 3, 150–157.
 PRENDERGAST, H. D. V.; DOLLEY, D., 2001: Jesuit’s bark
(Cinchona [Rubiaceae]) and other medicines. Economic
Bot. 55, 3–6.
15_III-4_Cinchona_pubescens_58EL:00_Musterseite_NEU 13.05.2011 14:46 Uhr Seite 12
Enzyklopädie der Holzgewächse – 58. Erg.Lfg. 06/11 13
 RENTERÍA, J. L., 2002: Ecología y manejo de la cascarilla
(Cinchona pubescens VAHL) en Santa Cruz, Galápagos. Te-
sis de Ingeniería Forestal, Universidad Nacional de Loja,
 ROCCO, F., 2003: The miraculous fever-tree. Malaria and
the quest for a cure that changed the world. Harper Collins,
 SAUER, J. D., 1988: Plant migration: the dynamics of geo-
graphic patterning in seed plant species. Univ. of California
 SCHMIDT, S. K.; SCOW, K. M., 1986: Mycorrhizal fungi on
the Galápagos Islands. Biotropica 18, 236–240.
 SCHUMANN, K., 1891: Rubiaceae. Cinchona, 44–45. In:
ENGLER, A.; PRANTL, K. (eds.): Die natürlichen Pflanzenfa-
milien nebst ihren Gattungen und wichtigeren Arten insbe-
sondere den Nutzpflanzen, Engelmann, Leipzig.
 SHIMIZU, Y., 1997: Competitive relationships between tree
species of Scalesia (S. pedunculata, S. cordata, S. microce-
phala) and introduced plants (Cinchona succirubra, Psi-
dium guajava, Lantana camara) with reference to regenera-
tion mechanism of Scalesia forests in the Galápagos Is-
lands. Repr. from Regional Views 11, 23–172. Institute for
Applied Geography, Komazawa Univ., Tokyo.
 SPRUCE, R., 1861: Report on the expedition to procure seeds
and plants of the Cinchona succirubra or red bark tree. G.
E. Eyre and W. Spottiswoode, London.
 STARR, F.; STARR, K.; LOOPE, L., 2003: Cinchona pubescens. Re-
port for the Hawaiian Ecosystems at Risk project (HEAR).
escens.htm (last acc. 18 Oct. 2010).
 TAYL O R , N., 1943: Quinine: the story of Cinchona. Reprin-
ted from Scientific Monthly (July), publ. by AAAS, Smith-
sonian Institute, Washington, D.C.
 VANQUIN, V., 2006: Ecologie et dynamique de l’invasion de
l’arbre à quinine Cinchona pubescens (Rubiacées) dans les
forêts humides de l’île de Tahiti (Polynésie française). Rap-
port de stage de licence. Universite Paul Verlaine, Metz &
Délégation à la Recherche, Papeete.
 WEBER, E., 2003: Invasive Plant Species of the World. A reference
guide to environmental weeds. CABI Publishing, Cambridge.
 WINTERS, H. F.; LOUSTALOT, A. J.; CHILDERS, N. F., 1946: In-
fluence of temperature on growth and alkaloid content of
Cinchona seedlings. Fed. Exper. Station, U.S. Dep. of Agri-
culture, Mayaguez, Puerto Rico.
 WOODWARD, R. B.; DOERING, W. E., 1945: The total synthe-
sis of quinine. J. American Chemical Society 67, 860–874.
 ZANNE, A. E.; LOPEZ-GONZALEZ, G.; COOMES, D. A.; ILIC,
J.; JANSEN, S.; LEWIS, S. L.; MILLER, R. B.; SWENSON, N. G.;
WIEMANN, M. C.; CHAVE, J., 2009: Global wood density da-
tabase. Dryad. Identifier: doi:10.5061/dryad.234/1 (last acc.
18 October 2010).
Dr. HEINKE JÄGER
Department of Ecology
Technische Universität Berlin
Fig. 16: C. pubescens trees in the introduced range: highlands of Santa Cruz Island, Galápagos (670 masl)
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