Cinchona pubescens (quinine tree)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Biology and Ecology
- Latitude/Altitude Ranges
- Air Temperature
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Plant Trade
- Wood Packaging
- Impact Summary
- Economic Impact
- Environmental Impact
- Impact: Biodiversity
- Threatened Species
- Social Impact
- Risk and Impact Factors
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Cinchona pubescens Vahl
Preferred Common Name
- quinine tree
Other Scientific Names
- Cinchona asperifolia Wedd.
- Cinchona caloptera Miq.
- Cinchona chomeliana Wedd.
- Cinchona hirsuta Ruiz & Pavon
- Cinchona lechleriana Schlecht.
- Cinchona succirubra Pav. ex Klotzsch
- Quinquina pubescens (Vahl) Kuntze
International Common Names
- English: Jesuits' Peruvian-bark; red Cinchona; red Peruvian-bark; sacred Peruvian-bark
- Spanish: cascarilla amarga; cascarilla gallinazo; varona
- French: quinquina rouge
Local Common Names
- Brazil: quina-do-Amazonas
- Ecuador: cascarilla; roja-roja
- CIHPU (Cinchona pubescens)
Summary of InvasivenessTop of page
C. pubescens has been widely introduced throughout the tropics over the past 150 years because of its medicinal properties. However, it is only in small island systems, notably the Galapagos and Hawaii, that it has become invasive, after a relatively long period of adaptation. In the moist upland regions of these islands, it is beginning to have a significant impact on biodiversity affecting endemic species in both the flora and fauna.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Dicotyledonae
- Order: Gentianales
- Family: Rubiaceae
- Genus: Cinchona
- Species: Cinchona pubescens
Notes on Taxonomy and NomenclatureTop of page
Andersson (1998) finally resolved and simplified the taxonomy of the genus Cinchona and, in particular, the confusion surrounding the identification of the quinine species of commerce. His was the first revision since 1878 (Prendergast and Dolley, 2001). One of these, collected by the botanist and explorer Richard Spruce from the western slopes of the Ecuadorian Andes (Drew, 1996; Naranjo, 1996) was, until recently, more commonly known as C. succirubra, particularly in its invasive range (Cronk and Fuller, 1995; Macdonald et al., 1988). Andersson (1998) determined this as C. pubescens and, thus far, 104 synonyms have been cited (Missouri Botanical Garden, 2003). Obviously, only a selection of the more commonly quoted synonyms can be included here under the non-preferred scientific names. Earlier, Purseglove (1968) had suggested that C. succirubra probably represented a form or variety of the highly variable species C. pubescens. The genus was named after the Countess of Chinchón, wife of the Viceroy of Peru, but was mis-spelled by Linnaeus in his description of the type species, Cinchona officinalis (Naranjo, 1996; Prendergast and Dolley, 2001).
DescriptionTop of page
C. pubescens is a large, erect, rapidly growing evergreen tree; up to 30 m tall, but most frequently reported as 4-10 m in height, attaining heights of 15 m on Santa Cruz, Galapagos (Shimizu, 1997); sparsely branched as lower branches shed, with young branches glabrous or pubescent. Bark brown, smooth; inner bark reddening when cut. Leaves deciduous, opposite, large, broadly elliptic-ovate to broadly oblong, 20-40 (-50) x 10-30 (-40) cm; upper surface light green, puberulent or glabrate, rather thin, membranous to papery, conspicuously veined (9-11 lateral pairs), margins entire, apex rounded, base broadly to narrowly cuneate, petioles 1.5-4.5 cm long, stipules ovate, caducous; lower surface with tufts of hairs in axils of lateral veins; old leaves reddish-orange. Flower numerous, in large panicles, rose-pink, fragrant; calyx densely appressed, pubescent, teeth deltate: heterostylous; in microstyled plants 5 exerted anthers alternate with corolla lobes and bifid stigma reaches half length of corolla tube; in macrostyled plants the situation is reversed with the stigma being exserted and the stamens hidden in pale pink corolla tube (1.0-1.2 cm long), which is villous within and comprising 5 spreading lobes, 4-5 mm long. Capsules ovoid-fusiform to oblong, 2-3 cm long; dehiscent from base to apex. Seeds 7-10 x 2-3 mm, with a broad ciliate wing. Note that this species description is a composite from several sources which vary considerably, especially in dimensions of the tree, leaves and seeds, emphasizing the variability of this species.
DistributionTop of page
C. pubescens has a natural range extending from Bolivia, through Peru, Ecuador and Colombia to Costa Rica in Central America. It was widely distributed throughout the tropics as a source of quinine, particularly by the British during the latter half of the 1800s (Moureau, 1945). It is still held in many tropical botanical gardens worldwide for its medicinal interest.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Planted||Reference||Notes|
|India||Present||Present based on regional distribution.|
|-Assam||Present||Introduced||Not invasive||Planted||Naranjo, 1996|
|-Sikkim||Present||Introduced||Not invasive||Planted||Purseglove, 1968|
|-Tamil Nadu||Present||Introduced||ca. 1865||Not invasive||Purseglove, 1968; Naranjo, 1996|
|Indonesia||Present||Present based on regional distribution.|
|-Java||Present||Introduced||ca. 1865||Not invasive||Planted||Purseglove, 1968|
|Philippines||Present||Introduced||Not invasive||Planted||Alejandro and Liede, 2000|
|Sri Lanka||Present||Introduced||Not invasive||Planted||Purseglove, 1968|
|Burundi||Present||Introduced||Not invasive||Starr et al., 2003|
|Cameroon||Present||Introduced||Not invasive||Planted||Prendergast and, 2001|
|Congo||Present||Introduced||Not invasive||Planted||Purseglove, 1968|
|Congo Democratic Republic||Present||Introduced||Starr et al., 2003|
|Kenya||Present||Introduced||Not invasive||Planted||Moureau, 1945|
|Nigeria||Present||Introduced||Not invasive||Planted||Okonkwo and Msonthi, 1995|
|Réunion||Present||Introduced||Not invasive||Planted||Sejourne et al., 1986|
|Rwanda||Present||Introduced||Not invasive||Husain, 1991|
|Saint Helena||Restricted distribution||Introduced||Invasive||Planted||Cronk and Fuller, 1995|
|Tanzania||Present||Introduced||1900||Not invasive||Purseglove, 1968; Missouri Botanical Garden, 2003|
|Uganda||Present||Introduced||1918||Not invasive||Moureau, 1945; Purseglove, 1968|
|USA||Present||Present based on regional distribution.|
|-Hawaii||Localised||Introduced||1868||Invasive||Wagner et al., 1999; Starr et al., 2003||Hawaii (Big Island), Maui, Oahu|
Central America and Caribbean
|Costa Rica||Present||Native||Not invasive||Standley and Williams, 1975|
|Guatemala||Present||Introduced||Not invasive||Planted||Standley and Williams, 1975|
|Jamaica||Present||Introduced||Not invasive||Planted||USDA-ARS, 2009|
|Panama||Present||Native||Not invasive||Natural||USDA-ARS, 2009|
|Puerto Rico||Present||Introduced||Not invasive||Planted||Harper and Winters, 1946|
|Bolivia||Present||Native||Not invasive||Natural||Missouri Botanical Garden, 2003|
|Colombia||Present||Native||Not invasive||Natural||USDA-ARS, 2009|
|Ecuador||Restricted distribution||Native||Not invasive||Harmann, 1984; Tye, 2001|
|-Galapagos Islands||Localised||Introduced||1925||Invasive||Planted||Tye, 2001||Santa Cruz|
|Peru||Present||Native||Not invasive||Natural||Missouri Botanical Garden, 2003|
|Venezuela||Present||Native||1925||Not invasive||Natural||Missouri Botanical Garden, 2003|
|French Polynesia||Localised||Introduced||1940||Invasive||Planted||Meyer, 2004||Tahiti|
|Palau||Absent, formerly present||Introduced||Not invasive||Fosberg et al., 1993|
History of Introduction and SpreadTop of page
Richard Spruce's collections from western Ecuador were sent to Kew in the late 1850s and thence to the British colonies, particularly India and Sri Lanka to establish plantations for quinine production (Drew, 1996; Naranjo, 1996). By 1880, Indian plantations were producing approximately 500,000 kg of cinchona bark (Purseglove, 1968). The Dutch also introduced C. pubescens into Java during this period, but this was soon replaced by the yellow-barked Cinchona (C. calisaya = C. ledgeriana) because of its much higher quinine content. However, C. pubescens was still used as a rootstock (Drew, 1996). According to Tye (2001), C. pubescens was first introduced into the Galapagos sometime before 1925, specifically for quinine production, and not in 1946 as cited by Macdonald et al. (1988). In Hawaii, plantations were established as early as 1868 on Maui Island (Starr et al., 2003) and on Oahu and the island of Hawaii in the early twentieth century (Starr et al., 2006) Invasive spread from such plantations seems to have occurred only in small island systems, such as the Galapagos where spread was first recorded in 1965-66 (Tye, 2001) and Hawaii where it was reported as becoming naturalized from 1978-87 (Starr et al., 2003) and now occupying about 6000 ha on Maui (Buddenhagen et al., 2004). There are no corroborated reports of C. pubescens spreading from the much larger cultivated areas in Africa and Asia and becoming invasive. To reduce the reliance on quinine from the East Indies, plantations were established at or near the sites of the plant’s origin in the New World beginning in the 1930s. C. pubescens was introduced to Tahiti in the 1940s and observed to be naturalized by the 1970s (Meyer, 2004). It has naturalized on St. Helena island in the south Atlantic, although currently occupying a limited distribution there (Cronk, 1989; Cronk and Fuller, 1995). Quinine was first artificially synthesized in 1944, resulting in a reduced demand for bark from C. pubescens (Starr et al., 2003). Large quinine tree plantations were established in Africa during World War II, and by 1998 Zaire was the world’s top supplier, along with contributions from Burundi, Cameroon and Kenya (Bruce-Chwatt, 1988; Dagani, 2005). Limited production continues in South America, Indonesia and India.
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|French Polynesia||South America||1940||Crop production (pathway cause)||Yes||No||Meyer (2004)|
|Galapagos Islands||South America||1925||Crop production (pathway cause)||Yes||No||Tye (2001)|
|Hawaii||South America||1868||Crop production (pathway cause)||Yes||No||Starr et al. (2003); Wagner et al. (1999)|
|Saint Helena||South America||1870s||Crop production (pathway cause)||Yes||No||Cronk and Fuller (1995); Cronk and Fuller (1995)|
Risk of IntroductionTop of page There remains a possibility that it could still be transported between countries and continents for cultivation as a herbal or ornamental plant (PIER, 2002), but the phytosanitary risk is perceived as low.
HabitatTop of page
The native range extends from Andean South America in Bolivia north to Costa Rica. Unusually, it occurs on both sides of the Cordillera in sub-montane rainforest, predominately at altitudes of 800-2800 m but can be found up to 3500 m. There are also reports of its occurrence in the lowland Amazonian rainforest of Peru at altitudes of 120-160 m (Missouri Botanical Garden, 2003). This wide geographic and altitudinal native range explains the high variability of this species and the plethora of synonyms. It was first cultivated in similar moist montane habitats in the Indian states of Tamil Nadu (Nilgiri Hills) and Sikkim (Himalayan foothills) at 700-1800 m altitude (Purseglove, 1968), but is now cultivated through a broad range in the tropics, although it fails to produce an economic harvest in some of its introduced range (Tye et al., 2003). Similarly moist, fertile habitats favour growth and spread in its introduced range (Itow, 2003; Starr et al., 2003).
Habitat ListTop of page
|Terrestrial – Managed||Cultivated / agricultural land||Present, no further details|
|Managed forests, plantations and orchards||Principal habitat||Harmful (pest or invasive)|
|Managed forests, plantations and orchards||Principal habitat||Productive/non-natural|
|Disturbed areas||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Rail / roadsides||Present, no further details|
|Terrestrial ‑ Natural / Semi-natural||Natural forests||Present, no further details||Harmful (pest or invasive)|
|Natural forests||Present, no further details||Natural|
|Scrub / shrublands||Principal habitat||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page
C. pubescens is not a weed of agricultural crops, although there are reports of it invading Eucalyptus plantations in Hawaii (Wagner et al., 1999; Starr et al., 2003). In the Galapagos, it has invaded two of the upper vegetation zones (500-700 m), posing a threat to the endemic flora (Macdonald et al., 1988; Cronk and Fuller, 1995).
Biology and EcologyTop of page
The chromosome number is 2n=34, which appears to be shared by most, if not all, Cinchona species (Purseglove, 1968). C. pubescens is a highly variable taxa which is compounded by its ability to readily hybridize, examples being C. pubescens x C. calisaya = C. hybrida; C. pubescens x C. officinalis = C. robusta (Purseglove, 1968). Molecular analysis by Andersson and Antonelli (2005) confirmed close relationships among Cinchona species, with the genus representing a distinct clade.
Physiology and Phenology
Seeds require light for germination, which can take 2-3 weeks (Moureau, 1945), although seedlings can also grow under shade which suggests that it is a pioneer species in its montane forest habitat. Seedling shade tolerance also enables C. pubescens to produce high density stands (Eliasson, 1982; Jager et al., 2009). In its exotic range, C. pubescens flowers and sets seeds throughout the year (Purseglove, 1968) and can seed copiously from about 5 years of age (Tye, 2001).
C. pubescens is a hermaphrodite and insect-pollinated, with high heterozygosity. Pollinating insects are purportedly non-specialized, such as bees, butterflies and various dipterans attracted to the fragrant flowers, but this is speculative rather than confirmed (Starr et al., 2003). Attempts at self pollination in both flower types have not been successful. Plants mature early, and can flower after 1-2 years, with prolific seed production numbering in the thousands per tree (Schofield, 1989). Seeds are surrounded by a paper-like wing which aids wind dispersal. However, observations in Hawaii suggest that the seeds do not travel long distances since most seedlings are found within 100 m of the parent tree (Starr et al., 2003). Seeds are short-lived and quickly lose viability (Purseglove, 1968). C. pubescens can re-grow from root fragments (Macdonald et al., 1988); the species also produces root suckers and readily re-sprouts from cut or damaged stems (Starr et al., 2003). Thus C. pubescens may spread vigorously by vegetative means (Hamann, 1974).
In its natural habitat, predominately montane rainforest, C. pubescens favours precipitous, well-drained slopes with a well-distributed annual rainfall of over 2000 mm with an average minimum temperature of 14ºC. It can, however, tolerate more extremes of temperature since as herbarium records show it has been collected occasionally from both cloud and lowland rainforest (Missouri Botanical Garden, 2003). Although associated primarily with moist environments, C. pubescens does persist under drier conditions (Itow, 2003). It grows best on light, virgin forest soil, rich in organic matter and preferably of volcanic origin. It cannot tolerate waterlogging and grows poorly on soils exposed to fire. Where invasive, it appears to favour mesic wet forest and moist tropical montane scrub always based on volcanic soil, as shown in the Galapagos, at 600-800 m altitude and with an even rainfall of 2000-3000 mm per annum. It inhabits high elevation cloud forests in Tahiti, 1000 to 2200 m in altitude (Vanquin, 2006).
C. pubescens was shown to have high levels of vesicular-arbuscular mycorrhizal fungi in its roots in the Galapagos ecosystem (Schmidt and Scow, 1986), unlike many natives in the same system (Henderson et al., 2006).
ClimateTop of page
|A - Tropical/Megathermal climate||Tolerated||Average temp. of coolest month > 18°C, > 1500mm precipitation annually|
|Af - Tropical rainforest climate||Preferred||> 60mm precipitation per month|
|Am - Tropical monsoon climate||Tolerated||Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Air TemperatureTop of page
|Parameter||Lower limit||Upper limit|
|Absolute minimum temperature (ºC)||0||5|
|Mean annual temperature (ºC)||15||21|
|Mean maximum temperature of hottest month (ºC)||25||30|
|Mean minimum temperature of coldest month (ºC)||12||16|
RainfallTop of page
|Parameter||Lower limit||Upper limit||Description|
|Dry season duration||0||4||number of consecutive months with <40 mm rainfall|
|Mean annual rainfall||1200||3500||mm; lower/upper limits|
Notes on Natural EnemiesTop of page
There have few studies on the natural enemies of Cinchona spp. in their native ranges. A specific and damaging disease on C. pubescens is caused by Elsinoe cinchonae Jenkins (Elsinoaceae, Ascomycota) resulting in leaf spots, stem cankers and fruit scab (Jenkins, 1945). Cinchona species are infected by Phytophthora quininea in Central America (Guatemala), the Caribbean Islands (Puerto Rico), and South America (Bolivia, Peru) (Crandall, 1947; Cline et al., 2008). In contrast, there are a number of reports from its exotic, cultivated range of natural enemies, which are invariably generalist opportunistic pests and diseases of plantation tree crops in the tropics. For example, both Purseglove (1968) and Frohlich and Rodewald (1970) list damping-off of nursery seedlings due to Rhizoctonia solani Kuhn (Corticiaceae, Basidiomycota), as well as various root rots in the plantation trees caused by the honey fungus, Armillaria mellea (Fr.) Quel. (Tricholomataceae, Basidiomycota), and grey root rot, Rosellinia arcuata Petch (Xylariaceae, Ascomycota). However, Frohlich and Rodewald (1970) considered that Phytophthora species, especially P. cinnamomi Rands (root rot and stripe canker) and P. parasitica Dastur (tip blight and girdle canker) were the most destructive pathogens. The most serious pest was listed as Helopeltis (Miridae) by Purseglove (1968) but Frohlich and Rodewald (1970) also included mites of the genera Brevipalpus, Taponemus and Tetranychus; with mealybugs (Planococcus citri Risso) and scale insects, such as Coccus hesperidum L., Coccus viridus Green and Saissetia coffeae Walk, especially important since they affected bark quality. Earlier, Butler (1918) reported the occurrence of pink disease, Corticium salmonicolor Berk. & Broome in Cinchona plantations in India and noted that the dense bushy trees of C. ledgeriana (= C. calisaya) suffered severely from the disease whilst the more open, sparsely branched C. pubescens was rarely affected.
Means of Movement and DispersalTop of page
Major spread is by wind-dispersed seed which are produced in large numbers; anecdotal evidence from both Hawaii and the Galapagos suggests that dispersal is localized (GISP, 2002) and in Hawaii most seedlings are found within 100 m of the parent tree (Starr et al., 2003). Meyer (2004) observed isolated trees naturalized in upland areas of Tahiti. On Santa Cruz, Galapagos, the species was able to spread rapidly in treeless area since the 1940s and by the beginning of the twenty-first century it occupied approximately 10% of the island (Buddenhagen et al., 2004). There are no documented examples of biotic transmission, spread via agricultural practices and it is unlikely to be introduced accidentally or as a contaminant of crop seed or produce. There are recommendations in Hawaii that C. pubescens should not be sold or planted. There remains a possibility that it could still be transported between countries and continents for cultivation as a herbal or ornamental plant (PIER, 2008).
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Fruits (inc. pods)||seeds|
|Plant parts not known to carry the pest in trade/transport|
|Growing medium accompanying plants|
|Stems (above ground)/Shoots/Trunks/Branches|
|True seeds (inc. grain)|
Wood PackagingTop of page
|Wood Packaging not known to carry the pest in trade/transport|
|Loose wood packing material|
|Processed or treated wood|
|Solid wood packing material with bark|
|Solid wood packing material without bark|
Impact SummaryTop of page
|Cultural/amenity||Positive and negative|
|Economic/livelihood||Positive and negative|
|Fisheries / aquaculture||None|
Economic ImpactTop of page
There are no records of direct impact on crops. The species is said to be invading Eucalyptus plantations in Hawaii (Starr et al., 2003) but there are no economic data. Tye (2001) gave estimates for eradication costs in the Galapagos at US$2 million over 15 years. Buddenhagen and Yanez (2005) estimated that eradicating the 11000 ha infestation on the Galapagos Island of Santa Cruz would cost US $1.65 million. Costs ranged from $14 to $2225 per ha depending on stem density. The initial knockdown alone would require more than 275 000 man-hours of effort, equivalent to employing 75 people for two years; further search and control effort would be required in later years.
Environmental ImpactTop of page
Impact on Habitats
C. pubescens has now become the most common invasive alien plant in the Miconia vegetation zone on the Galapagos island of Santa Cruz, and forests of the red quinine tree cover the upper volcanic slopes (Mauchamp, 1997). It was listed among 32 land plants within a listing of “100 of the world’s worst invasive alien species” (Lowe et al., 2000). It would seem that after a prolonged lag phase, C. pubescens has moved into an explosive invasive phase within the last two decades (Macdonald et al., 1988; Schofield, 1989; Moll, 1998; Mack et al., 2000; Tye, 2001), making it the major threat to biodiversity and habitat stability in the highland region of Santa Cruz Island. In Hawaii, C. pubescens has shown an ability to invade both disturbed forests and primary moist or mesic forests. The dense canopy and rapid growth rate (1-2 m per year) out-competes and shades out native vegetation, impacting forest structure severely, with a reduction in photosynthetically active radiation of 87% recorded on Santa Cruz (Jager et al., 2009). More litter was produced under Cinchona trees in Santa Cruz and the litter decomposed quickly, impacting the soil environment and in turn the vegetation cover (Capa, 2006). Water cycle modifications via enhanced fog interception were also observed due to the thick C. pubescens canopy on Santa Cruz (Jager et al., 2009). Despite these pronounced impacts on some locations where it occurs, to date C. pubescens is naturalized on relatively few islands (Meyer, 2004).
Impact: BiodiversityTop of page
On St. Helena, C. pubescens invaded tree-fern thickets (Cronk, 1989). C. pubescens apparently has impacted the biodiversity of upland Hawaiian forest reserves, out-competing indigenous plants, from understorey trees such as Acacia koa, to herbs such as the Uluhe fern, Dichanopteris linearis (Wagner et al., 1999; Starr et al., 2003). However, more in-depth study by Fischer et al. (2009) indicated that C. pubescens actually facilitated higher diversity of endemic species in the three forest types studied: Pinus plantations, Eucalyptus plantations and “near-natural” Acacia koa forests. Fischer et al. (2009) attributed the 20% increase in species richness and 50% increase in the proportion of endemic species to shading by C. pubescens in these simplified ecosystems.
In the Galapagos, areas invaded by C. pubescens on Santa Cruz went from open shrubland to forest within a few decades (Buddenhagen et al., 2004; Jager et al., 2007). See Itow (2003) for photographs showing the successive invasion from 1970 to 1991 to 2001. Itow (1990; 2003) identified 11 community types in the Santa Cruz highlands, all of which were altered by the invasion of C. pubescens. Species composition of the native community type dominated Miconia robinsoniana shrubs has been less impacted than the fern-sedge zone, likely because Miconia communities were already adapted to somewhat shaded conditions (Jager et al., 2007). The invasion of the Santa Cruz highlands has impacted endemic shrubs such as M. robinsoniana, and endemic herbs notably Justicia galapagana and Pilea baurii reducing percentage cover of most plant species by 50%, as well as on several endemic arthropods and birds such as the Galapagos rail, Laterallus spilonotus or the Galapagos petrel, Pterodroma galapagensis (Macdonald et al., 1988; Cruz and Cruz, 1996; Caraval, 2002; Gibbs et al., 2003; Jager et al., 2003; Buddenhagen et al., 2004; Jager et al., 2009). Gibbs et al. (2003) observed that the Galapagos rail avoided C. pubescens, even though the tree was found at 75% of points surveyed in 2000. Many species impacted by C. pubescens on Santa Cruz are red-listed by IUCN (Jager et al., 2007). Even if C. pubescens is controlled in these areas, other non-native species thriving in the shade of the quinine tree might continue to spread (Jager et al., 2007). The demonstrated transformation of the habitat itself by C. pubescens likely leads to drastic and unpredictable biodiversity changes over the long-term (Jager et al., 2009).
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Cyathea weatherbyana||EN (IUCN red list: Endangered) EN (IUCN red list: Endangered)||Galapagos Islands||Jäger et al., 2009|
|Miconia robinsoniana||EN (IUCN red list: Endangered) EN (IUCN red list: Endangered)||Galapagos Islands||Jäger et al., 2009|
|Scalesia pedunculata||EN (IUCN red list: Endangered) EN (IUCN red list: Endangered)||Galapagos Islands||Jäger et al., 2009; Shimizu, 1997|
Social ImpactTop of page
Further invasion poses a threat of long-term impact on tourism in affected islands. Tourism is the leading source of income in both the Hawaiian Islands and the Galapagos. The promotion of the Galapagos as a world “showcase for evolution” is jeopardized (Trillmich, 1992). Yet there is not enough funding available to adequately manage the populations present on Santa Cruz, Galapagos, or the infestations on the Hawaiian islands.
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Abundant in its native range
- Highly adaptable to different environments
- Tolerant of shade
- Benefits from human association (i.e. it is a human commensal)
- Long lived
- Fast growing
- Has high reproductive potential
- Reproduces asexually
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Increases vulnerability to invasions
- Modification of hydrology
- Modification of nutrient regime
- Modification of successional patterns
- Monoculture formation
- Negatively impacts forestry
- Negatively impacts tourism
- Reduced amenity values
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Competition - monopolizing resources
- Competition - shading
- Competition - smothering
- Rapid growth
- Highly likely to be transported internationally deliberately
- Difficult/costly to control
UsesTop of page
Cinchona bark was being used in Europe as a treatment for malaria as early as the 1650s before the plant was authenticated botanically (Raintree, 2003). The Jesuits had introduced it from the New World, hence the name Jesuit's bark or powder (Prendergast and Dolley, 2001). The bark was collected in its native Andean range and probably comprised a mixture of species including C. pubescens. One area of supply came from the western slopes of the Ecuadorian Andes in Cotapaxi Province, and that was where Richard Spruce was commissioned by the British Government to collect germplasm of the red quinine tree in the 1850s (Drew, 1996; Naranjo, 1996). It was this material which formed the basis of the plantations established by the British in the Indian subcontinent in the 1860s, and in Indonesia by the Dutch. However, it was soon discovered by the Dutch that the yellow-barked Cinchona from Peru gave much higher yields of the active alkaloid quinine (Purseglove, 1968), which was used both as a prophylactic and for treatment of malaria. Since the 1940s, new synthetic drugs replaced quinine but production still continues, principally in Africa, and an estimated 300-500 t are produced annually from 5,000 – 10,000 t of raw bark (Starr et al., 2003; Dagani, 2005), mainly for use in the food industry for tonic water and as an FDA-approved bitter food additive. Quinine producing countries in other parts of the world include Indonesia, Sri Lanka, Bolivia, Colombia, Costa Rica and India (Husain, 1991). It also has minor uses in the pharmaceutical industry for sunburn lotions, antiseptics and insecticides. Extracts from C. pubescens were found to have some activity against the fungus Candida albicans (Nino et al., 2003). It is still much favoured in herbal medicine worldwide. Recently, natural quinine is making a comeback for the management of new strains of malaria which have developed resistance to the synthesized quinine drugs. An impressive list of current uses for natural quinine was produced recently by Raintree (2007), from treatment of anaemia to muscle spasms to cancer. Quinine is also used as an additive in some carbonated beverages.
Similarities to Other Species/ConditionsTop of page
C. pubescens has been separated from the three other species of economic importance: C. calisaya Wedd., C. ledgeriana Moens ex Tremen. and C. officinalis L., on bark colour, hence the vernacular epithets red quinine, roja-roja and quinquina rouge. C. calisaya, with which C. ledgeriana has now been synonymized (Andersson, 1998), has thick whitish bark which yields the yellow Peruvian bark of commerce; whilst C. officinalis has rough brown bark and is a much less robust, slender tree.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
In the Galapagos, recommendations have been made to plant other non-invasive tree species in place of invaders like C. pubescens (Richardson, 1998). On Hawaii recommendation has been to discourage further planting of C. pubescens as a crop (Starr et al., 2003). Although the plant is highly susceptible to fire, this method is untenable in the delicate island ecosystems typically invaded by C. pubescens. C. pubescens appears to spread less rapidly in forested areas than in open areas (Buddenhagen et al., 2004).
There remains a possibility that it could still be transported between countries and continents for cultivation as a herbal or ornamental plant (PIER, 2008), but the phytosanitary risk is perceived as low.
Manual removal has been practiced in the Galapagos, but this is labour intensive and hence expensive (Macdonald et al., 1988) since it includes felling, grubbing out the stumps and hand-pulling seedlings. Tens of thousands of trees were removed on the Galapagos in the 1980s by this method within a 1000 ha area of Santa Cruz (Macdonald et al., 1988). Nevertheless, the plant can regrow from root fragments and also resprouts vigorously from cut stumps. Ring-barking appears ineffective because of rapid bark regeneration.
No targeted natural enemy surveys have been undertaken in the native range of C. pubescens and thus classical biological control has not been pursued. Hoffmann and Moran (1998) have advocated that biocontrol agents that reduce seed set should be used routinely as a first line of attack to prevent further invasions. Limited evaluation of indigenous wood-rotting fungi for stump-treatment application to prevent resprouting has been undertaken recently in the Galapagos, an approach successfully exploited in The Netherlands for the management of invasive alien tree species (de Jong, 2000).
C. pubescens is resistant to many herbicides but initial attempts to develop chemical control methods were largely ineffective (Macdonald et al., 1988; Starr et al., 2003; Buddenhagen et al., 2004). More recent research on Santa Cruz in the Galapagos concluded that the most effective herbicide combination was metsulfuron-methyl and picloram applied to a trunk cut with a machete (Buddenhagen et al., 2004). This combination was 80% effective, and was less labour intensive than five other methods tested. Some combinations with triclopyr also produced satisfactory control (Buddenhagen et al., 2004). Although a previous report claimed glyphosate was effective as a foliar application to small trees, Buddenhagen et al. (2004) observed no more than 2% control when applying glyphosate via the machete method.
There is an on-going campaign in the Galapagos to eradicate C. pubescens on Santa Cruz Island but thus far less than 2% of the 11,000 ha area has been controlledusing a combination of manual and chemical measures (Macdonald et al., 1988; Tye, 2001; Caraval, 2002; Buddenhagen and Yanez, 2005). An estimated 110 ha was controlled within the Galapagos National Park between 1998 and 2003 (Buddenhagen et al., 2004). For eradication purposes, Buddenhagen and Yanez (2005) recommended repeating control measures every two years because of the approximately 2 years taken by quinine trees to mature, and continuing an eradication program for 10-20 years. Despite these challenges, the development of techniques for its control, its isolation on a single island, ease of identification and consensus on its control make C. pubescens a good target for eradication in the Galapagos (Tye et al., 2003). In Hawaii, eradication is considered to be a difficult option because of the dense infestations and rough terrain (Starr et al., 2003). Spot control of invasive pockets in selected areas with a high biodiversity is being proposed to slow the invasion and preserve rare native species.
Monitoring and Surveillance
Some surveying has been conducted in the Hawaiian Islands on a roadside basis (Starr et al., 2006) but there is a need to collect more data on local distribution there. The distribution is well characterized in the Galapagos Islands (Buddenhagen et al., 2004) where the trees are more visible.
Recommendations have been made to plant other species in the areas affected by C. pubescens on Santa Cruz in the Galapagos (Richardson, 1998). Although cover of native plant species on Santa Cruz has been severely reduced in many cases, there is no evidence that any plant species have been extirpated from the area (Jager et al., 2009). Because the species pool for the affected habitat is still complete, if cover of C. pubescens can be reduced, the prognosis for eventual restoration of the ecosystem is good.
Gaps in Knowledge/Research NeedsTop of page
More research is needed on distribution, abundance and impacts in the Hawaiian Islands, St. Helena and Tahiti. Now that control methods have been better developed for Santa Cruz, Galapagos, research is needed to verify that proposed steps towards eradication can be effective.
ReferencesTop of page
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OrganizationsTop of page
USA: Haleakala Field Station, HFS P.O. Box 246, Makawao, (Maui) HI 96768, http://www.hear.org/usgs-brd-pierc-hfs/index.html
Galapagos Islands: Charles Darwin Research Station (CDRS), run by the Charles Darwin Foundaton, Puerto Ayora, Isla Santa Cruz, Galapagos, http://www.darwinfoundation.org/english/pages/index.php
ContributorsTop of page
15/12/2009 Updated by:
David Clements, Biology and Environmental Studies, Trinity Western University, 7600 Glover Road, Langley, British Columbia, V2Y 1Y1, Canada
Distribution MapsTop of page
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