Kalanchoe pinnata (cathedral bells)

Pictures

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Picture

Title

Caption

Copyright

Seedlings

Kalanchoe pinnata (cathedral bells); seedlings amongst litter layer.

©Sheldon Navie

Young plant

Kalanchoe pinnata (cathedral bells); young plant.

©Sheldon Navie

Leaves

Kalanchoe pinnata (cathedral bells); leaves.

©Sheldon Navie

Leaves

Kalanchoe pinnata (cathedral bells); close-up showing the pinnate structure of the upper leaves.

©Arne Witt

Leaves

Kalanchoe pinnata (cathedral bells); upper leaves, note reddish tinge.

©Sheldon Navie

Flowers

Kalanchoe pinnata (cathedral bells); open flowers.

©Sheldon Navie

Habit

Kalanchoe pinnata (cathedral bells); growth habit, showing a dense stand.

©Sheldon Navie

Habit

Kalanchoe pinnata (cathedral bells); potted specimen, showing the cylindrical stem and crenate leaves.

©Arne Witt

Habit

Kalanchoe pinnata (cathedral bells); habit. Guanica Dry forest, Puerto Rico. May; 2011

©Smithsonian Institution/Julissa Rojas-Sandoval

Habit

Kalanchoe pinnata (cathedral bells); habit. Guanica Dry forest, Puerto Rico. May; 2011

©Smithsonian Institution/Julissa Rojas-Sandoval

Identity

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Preferred Scientific Name

Kalanchoe pinnata (Lam.) Pers. 1805

Preferred Common Name

cathedral bells

Other Scientific Names

Bryophyllum calycinum Salisb. 1805
Bryophyllum pinnatum (Lam.) Oken 1841
Bryophyllum pinnatum var. calcicolum H. Perrier
Calanchoe pinnata Pers.
Cotyledon calyculata Solander
Cotyledon pinnata Lam.
Crassuvia floripenula Comm.
Kalanchoe brevicalyx (Raym.-Hamet & H. Perrier) Boiteau
Kalanchoe calcicola (H. Perrier) Boiteau
Kalanchoe calycinum Salisb.
Kalanchoe pinnata var. brevicalyx Raym.-Hamet & H. Perrier
Kalanchoe pinnata var. calcicola H. Perrier
Kalanchoe pinnata var. floripendula Pers.
Kalanchoe pinnata var. genuina Raym.-Hamet
Sedum madagascariense Clus.
Verea pinnata (Lam.) Spreng. 1825

International Common Names

English: air plant; airplant; floppers; lifeplant
Spanish: hoja del aire; hoja santa
French: bryophylle

Local Common Names

Australia: green mother of millions; leaf of life; live leaf plant; miracle leaf; resurrection plant; sprouting leaf
Bahamas: life plant
Brazil: coirama; folha da fortuna; saiao
China: lao di sheng gen
Colombia: colombiano
Cook Islands: nganga'ere toka
Cuba: belladona; bruja; flor de aire; hoja de aire; prodigiosa; siempreviva; vibora; yerba coronel
Dominica: herbe mal tete
Dominican Republic: bruja; hegüey; higüey; oken; topetope; yerba bruja
Fiji: bulatawamudu
Germany: Brutblatt
Haiti: fébrivier; feuilles loup-garou; feuilles socier; feuilles sorcier; l’âne blanc; loup garou; sanglou; tope; tope-tope; z’herbe mal tête
India: panphooti; rankalli
Jamaica: leaf-of-life
Kiribati: teang
Lesser Antilles: baby bush; colic bush; fey chofi; gewitout; kalabana; kawakte lezom; lamowi; love bush; lucky bush; malotet; plok; salt fish; sweetheart bush; temetic; travel life; wonder-of-the-world
Madagascar: sodifafana; soutou-fafan; tsilafafa
Marshall Islands: kibilia
Myanmar: ywet-kya-pin-pauk
Netherlands: wonderblad
Niue: tupu he lau; tupu noa
Puerto Rico: yerba bruja
Samoa: pagi
South Africa: chandelier plant; finger plant
Taiwan: luo di sheng ken
Tonga: pipi vao
Trinidad and Tobago: never dead; parvu; wonder-of-the-world
USA: Canterbury bells; Mexican love plant; mother of thousands
USA/Hawaii: 'oliwa ku kahakai

EPPO code

KANPI (Kalanchoe pinnata)

Summary of Invasiveness

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In Queensland, Australia, K. pinnata was ranked 47 of 200 invasive naturalized plants (Batianoff and Butler, 2002). In New South Wales, Australia, it is a declared noxious weed under that state's Noxious Weed Act (1993) in the shire of Maclean. The species is listed as W2 requiring that the weed must be fully and continuously suppressed and destroyed by land managers. K. pinnata is described as a moderate invader of the Pacific islands of Hawaii, French Polynesia and Palau (Meyer, 2000). In the Galapagos Islands, K. pinnata invades disturbed sites and native vegetation (Soria et al., 2002). K. pinnata is recognized as a threat to island ecosystems by the Pacific Island Ecosystems at Risk project (PIER, 2004). It continues to be made available as a garden ornamental in many countries and is therefore likely to spread further.

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Plantae
Phylum: Spermatophyta
Subphylum: Angiospermae
Class: Dicotyledonae
Order: Rosales
Family: Crassulaceae
Genus: Kalanchoe
Species: Kalanchoe pinnata

Notes on Taxonomy and Nomenclature

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The family Crassulaceae includes about 34 genera and 1400 species distributed mostly in arid and dry regions in Mexico, Africa (Cape region), South America and Australia (Stevens, 2012). Plants within this family can be recognized by their succulent herbaceous or soft-stemmed habit, and by their flowers, which have the same number of sepals, petals and carpels (Stevens, 2012). Recent studies circumscribe the family Crassulaceae into three sub-families: Crassuloideae, Kalanchoideae and Sempervivoideae (Ham and 't Hart, 1998; Mort et al., 2001; Stevens, 2012).

The sub-family Kalanchoideae includes the genera Adromischus, Bryophyllum, Kalanchoe and Tylecodon (Mort et al., 2001; Stevens, 2012). Classification within this family is difficult, mainly because many of the species hybridize in both the wild and in cultivation. Some authors consider all Bryophyllum and Kitchingia species within a broadly circumscribed Kalanchoe (Boiteau and Allorge-Boiteau, 1995; Rauh, 1995) while others consider all three as distinct genera when considering a larger set of morphological characters and evidence of polyphyly within Kalanchoe sensu lato (Lauzac-Marchal, 1974; Toelken, 1985; Wickens, 1987; Forster, 1997; Staples et al., 2002). Genetic studies (Gehrig et al., 1997, 2001) and examination of the relative extent of utilization of CAM (Kluge et al., 1991; Kluge et al., 1993) support the differentiation of Bryophyllum and Kalanchoe. The Plant List (2010) has Bryophyllum pinnatum as the preferred name, while USDA-ARS (2013) uses K. pinnata, and only lists Bryophyllum as a non-preferred name for all species.

The genus Kalanchoe includes about 145 species native to the Old World, especially southern Africa, Arabia and South East Asia. Some species of Kalanchoe are characterized by the capability to produce plantlets in notches at the margin of the leaf blade. Two varieties of K. pinnata are recognized in a revision of the Kalanchoe of Madagascar (Boiteau and Allorge-Boiteau, 1995), namely K. pinnata var. pinnata and var. genuina.

While many botanists have adopted the APG III system of classification for the orders and families of flowering plants which places the Crassulaceae family in the order Saxifragales, the CAB Thesaurus continues to use the Cronquist system which places it under Rosales.

Description

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K. pinnata is a succulent glabrous glaucous perennial shrub 0.3-2 m high, with cylindrical erect, little-branched stems suckering at the base. The leaves are opposite and decussate and, in adult plants, are only present on the upper parts. The upper leaves are often 3- or 5-foliate while the lower leaves are simple. The leaf-blade is elliptic, 5-25 cm long and 2-7 cm wide with the apex obtuse to truncate, sometimes obscurely so. The leaf edge is crenate and young plants are often produced in the marginal notches especially after leaf detachment. The leaves of the inflorescence are similar to the foliage leaves but smaller, lower, 3- to 5-foliate, simple above and on the ultimate branch. The inflorescence is paniculate and the pedicels are slender (1-2.5 cm long) with young plants being produced at the points of insertion of the floral pedicel. The flowers are pendulous; calyx papery and much inflated, 25-40 mm long. The corolla is up to 7 cm long, yellowish-green tinged with pink or reddish, to twice as long as the calyx. Fruits have four slender papery tubes enclosed in the base of the corolla tube and the seeds are ellipsoid-oblong approximately 0.5 mm long, obscurely longitudinally striate and brown (Stanley and Ross, 1986; Wickens, 1987; Evett and Norris, 1990; Wagner et al., 1999).

Plant Type

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Herbaceous
Perennial
Seed propagated
Succulent
Vegetatively propagated

Distribution

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According to Boiteau and Allorge-Boiteau (1995), the native range of K. pinnata is in the northern half of Madagascar. However, it has also been found on the eastern seaboard, in the south, just north of Fort Dauphin (ABR Witt, ARC-PPRI, South Africa, personal communication, 2004). It has naturalized widely in the tropics where it has become invasive in many countries. Interestingly, K. pinnata has not become invasive in South Africa although it is sometimes found near habitation in large numbers (Toelken, 1985).

Distribution Table

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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.

Last updated: 17 Feb 2021

Continent/Country/Region	Distribution	Origin	Invasive	Reference	Notes
Africa
Cabo Verde	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
Comoros	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
Kenya	Present	Native	Naturalized	Witt and Luke (2017) Wickens (1987)	Naturalized
Madagascar	Present, Localized	Native		Boiteau and Allorge-Boiteau (1995)
Malawi	Present	Introduced	Naturalized	Witt and Luke (2017) Fernandes (1983)	Naturalized
Mauritius	Present	Introduced		Boiteau and Allorge-Boiteau (1995) CABI (Undated)
Mozambique	Present	Introduced		Fernandes (1983)
Réunion	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
Seychelles	Present	Introduced		CABI (Undated b)
South Africa	Present	Introduced		Toelken (1985)
Tanzania	Present	Introduced	Naturalized	Witt and Luke (2017) Wickens (1987)	Naturalized
Uganda	Present	Introduced	Naturalized	Witt and Luke (2017) Wickens (1987)	Naturalized
Zambia	Present	Introduced	Invasive	Witt and Luke (2017) Wickens (1987)
Zimbabwe	Present	Introduced		Fernandes (1983)
Asia
Bhutan	Present	Introduced		Dobremez et al. (2002)
China	Present	Introduced		Boiteau and Allorge-Boiteau (1995) Kunjun and Gilbert (2001) Tang et al. (2016)
-Fujian	Present	Introduced		Kunjun and Gilbert (2001)
-Guangdong	Present	Introduced		Kunjun and Gilbert (2001)
-Guangxi	Present	Introduced		Kunjun and Gilbert (2001)
-Yunnan	Present	Introduced		Kunjun and Gilbert (2001)
India	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
Indonesia	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
-Java	Present	Introduced		Boiteau and Allorge-Boiteau (1995) CABI (Undated)
-Maluku Islands	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
Japan	Present	Introduced		Enomoto (2001)
Myanmar	Present	Introduced		Kress et al. (2003)
Nepal	Present	Introduced		Dobremez et al. (2002)
Philippines	Present	Introduced		CABI (Undated)	Original citation: Missouri Botanical Garden (2003)
Sri Lanka	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
Taiwan	Present	Introduced		Li et al. (1977) Kunjun and Gilbert (2001)
Europe
Portugal	Present			CABI (Undated a)	Present based on regional distribution.
-Azores	Present	Introduced		USDA-ARS (2004) DAISIE (2013)
-Madeira	Present	Introduced		CABI (Undated)	Original citation: Silva Vieria RMda (2002)
Spain	Present			CABI (Undated a)	Present based on regional distribution.
-Canary Islands	Present	Introduced		USDA-ARS (2004) Acebes et al. (2001)
North America
Anguilla	Present, Widespread	Introduced		Broome et al. (2007)
Antigua and Barbuda	Present, Widespread	Introduced		Broome et al. (2007)
Bahamas	Present	Introduced		Boiteau and Allorge-Boiteau (1995) Correll and Correll (1982)
Barbados	Present, Widespread	Introduced		Broome et al. (2007)
Belize	Present	Introduced		CABI (Undated)
Bermuda	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
British Virgin Islands	Present	Introduced		Acevedo-Rodríguez and Strong (2012)	Anegada, Guana, Tortola, Virgin Gorda
Cayman Islands	Present	Introduced		Proctor (1984)
Costa Rica	Present	Introduced		CABI (Undated)	Original citation: Missouri Botanical Garden (2003)
Cuba	Present	Introduced	Invasive	Oviedo Prieto et al. (2012) Acuña G. (1974) González-Torres et al. (2012)
Dominica	Present, Widespread	Introduced		Broome et al. (2007)
Dominican Republic	Present	Introduced		Holm et al. (1991) Acevedo-Rodríguez and Strong (2012)
Grenada	Present, Widespread	Introduced		Broome et al. (2007)
Guadeloupe	Present, Widespread	Introduced		Broome et al. (2007)
Guatemala	Present	Introduced		CABI (Undated)	Original citation: Missouri Botanical Garden (2003)
Haiti	Present	Introduced		Acevedo-Rodríguez and Strong (2012)
Honduras	Present	Introduced		CABI (Undated)	Original citation: Missouri Botanical Garden (2003)
Jamaica	Present	Introduced		Adams (1972) CABI (Undated)
Martinique	Present, Widespread	Introduced		Broome et al. (2007)
Mexico	Present	Introduced		Boiteau and Allorge-Boiteau (1995) CABI (Undated)
Montserrat	Present, Widespread	Introduced		Broome et al. (2007)
Netherlands Antilles	Present, Widespread	Introduced		Broome et al. (2007)	Saba, St. Eustatius, St. Martin
Nicaragua	Present	Introduced		CABI (Undated)	Original citation: Missouri Botanical Garden (2003)
Panama	Present	Introduced		CABI (Undated)	Original citation: Missouri Botanical Garden (2003)
Puerto Rico	Present	Introduced		USDA-NRCS (2004) Acevedo-Rodríguez and Strong (2012) CABI (Undated)
Saint Lucia	Present, Widespread	Introduced		Broome et al. (2007)
Saint Vincent and the Grenadines	Present, Widespread	Introduced		Broome et al. (2007)
Trinidad and Tobago	Present	Introduced		Acevedo-Rodríguez and Strong (2012)
U.S. Virgin Islands	Present	Introduced		Ting (1989) USDA-NRCS (2004) Acevedo-Rodríguez and Strong (2012)	St. Croix, St. John, St. Thomas
United States	Present			CABI (Undated a)	Present based on regional distribution.
-Florida	Present	Introduced		USDA-NRCS (2004)
-Hawaii	Present	Introduced		USDA-NRCS (2004) CABI (Undated)
Oceania
American Samoa	Present	Introduced		Space and Flynn (2002)
Australia	Present			CABI (Undated a)	Present based on regional distribution.
-New South Wales	Present, Localized	Introduced	Invasive	Evett and Norris (1990)
-Queensland	Present, Localized	Introduced	Invasive	Stanley and Ross (1986)
Cook Islands	Present	Introduced		Space and Flynn (2002a) CABI (Undated)
Federated States of Micronesia	Present	Introduced		Fosberg et al. (1979)
Fiji	Present	Introduced		CABI (Undated b)
French Polynesia	Present	Introduced		Allan Herbarium (2000) Meyer (2000)
Guam	Present	Introduced		Fosberg et al. (1979)
Kiribati	Present	Introduced		Fosberg et al. (1979) Orton Williams (1980)
Marshall Islands	Present	Introduced		Fosberg et al. (1979)
Nauru	Present	Introduced		Fosberg et al. (1979)
New Caledonia	Present	Introduced		Boiteau and Allorge-Boiteau (1995) Swarbrick (1997)
New Zealand	Present	Introduced		Allan Herbarium (2000)
Palau	Present	Introduced		Fosberg et al. (1979) Space et al. (2003)
Tonga	Present	Introduced		Space and Flynn (2001)
Tuvalu	Present	Introduced		Swarbrick (1997)
South America
Bolivia	Present	Introduced		CABI (Undated)	Original citation: Missouri Botanical Garden (2003)
Brazil	Present	Introduced		Boiteau and Allorge-Boiteau (1995)
-Paraiba	Present	Introduced		CABI (Undated)	Original citation: New York Botanical Garden (2003)
-Parana	Present	Introduced		CABI (Undated)	Original citation: New York Botanical Garden (2003)
-Rio de Janeiro	Present	Introduced		CABI (Undated)	Original citation: New York Botanical Garden (2003)
-Santa Catarina	Present	Introduced		CABI (Undated)	Original citation: New York Botanical Garden (2003)
Colombia	Present	Introduced		Boiteau and Allorge-Boiteau (1995) CABI (Undated)
Ecuador	Present	Introduced	Invasive	Soria et al. (2002) CABI (Undated)
-Galapagos Islands	Present	Introduced		USDA-ARS (2004)
French Guiana	Present	Introduced		Boggan et al. (1997) BDG Program (2003)
Guyana	Present	Introduced		Boggan et al. (1997) BDG Program (2003) CABI (Undated)
Paraguay	Present	Introduced		Boiteau and Allorge-Boiteau (1995) CABI (Undated)
Peru	Present	Introduced		Boiteau and Allorge-Boiteau (1995) CABI (Undated)
Suriname	Present	Introduced		Boggan et al. (1997) BDG Program (2003)
Venezuela	Present	Introduced		Boiteau and Allorge-Boiteau (1995) BDG Program (2003)

History of Introduction and Spread

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The extensive distribution of K. pinnata throughout the tropics relative to related species is speculated to be due, in part, to its medicinal usage by early navigators (Boiteau and Allorge-Boiteau, 1995). These include the Malayo-Polynesian speaking navigators from the Indo-Malay archipelago who visited and colonized Madagascar some time after the first century AD. It is possible that the introduction of K. pinnata to other islands and the lands adjoining the Indian Ocean occurred some time after this. Movement of K. pinnata to the Caribbean and South America may have occurred after the arrival of early European traders in the Indian Ocean or possibly during the slave trade from West Africa. Movement of K. pinnata into Oceania may have occurred with the Polynesian navigators or much later during European colonial times and to the present day. Bailey (1885) mentions it in his census of the Brisbane Botanic Garden and Bowen Park (the garden of the Queensland Acclimatisation Society), Australia, which is an indication that it was utilized more than 100 years ago as a garden ornamental in Australia.

Risk of Introduction

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Further spread is probable because of the popularity of K. pinnata as a garden ornamental and medicinal plant. Seeds are available from commercial nurseries via mail-order catalogues and web-sites.

Habitat

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K. pinnata is native to Madagascar where it is fairly abundant along the coast growing in sandy soils and along granite outcrops. It is also found in the temperate humid and subhumid climates of the central highlands of Madagascar, with annual rainfall between 1000 and 2000 mm (Paulian, 1984).

In Hawaii, USA, it has naturalized and is sometimes very abundant at low elevations, in dry to mesic, disturbed areas (Wagner et al., 1999). On Guam it is often found naturalized near beaches (Stone, 1970). In Fiji it often forms dense stands, occurring from near sea level to an elevation of 550 m as a weed of waste places, cultivated fields and roadsides, naturalized on rocky coasts and slopes and sometimes in dry forest (Smith, 1985). In French Polynesia, K. pinnata occurs on the mesic uplands of the island of Rurutu (Meyer, 2000) and, on the US Virgin Island of St John, it is becoming a serious weedy pest in the dry scrub (Ting, 1989). In tropical East Africa, K. pinnata is found in riverine areas, amongst rocks and also in the shade of trees in areas ranging in altitude from 10 to 1600 m (Wickens, 1987). In Queensland, Australia, K. pinnata occurs along sandy and rocky shores and estuaries, densities increasing from the coast to inland areas (Batianoff and Franks, 1997).

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
Terrestrial	Managed	Managed forests, plantations and orchards	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Managed grasslands (grazing systems)	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Disturbed areas	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Rail / roadsides	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Urban / peri-urban areas	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Natural forests	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Wetlands	Present, no further details	Harmful (pest or invasive)
Littoral		Coastal areas	Present, no further details	Harmful (pest or invasive)

Hosts/Species Affected

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K. pinnata is not normally known as a weed of crops, although it has been reported to displace crops in the Galapagos Islands (Soria et al., 2002).

Biology and Ecology

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Genetics

Kalanchoe species in their Madagascan native range have shown a haploid chromosome number of n=17 except for K. pinnata which has 2n=40 (Taylor, 1926; Baldwin, 1938; Uhl, 1948; Friedmann, 1971). A few of the chromosomes are markedly smaller than the rest so the haploid number could be interpreted as n=17+3 or n=18+2. K. pinnata can be considered as an aneuploid with 6 (or 4) supplementary chromosomes and may be a recent derivation from an ancestor with n=17 or n=18 (Uhl, 1948).

Physiology and Phenology

All Kalanchoe species utilize crassulacean acid metabolism (CAM). This is an ecophysiological adaptation to arid or other water-stressed environments. CAM was first described from K. pinnata (for a review, see Kluge and Ting, 1978).

Many Kalanchoe species show a high phenotypic variability in CAM expression with water relations, light intensity, photoperiod and the diel temperature regime being major modulating factors (Kluge and Brulfert, 1996). Carbon isotope ratios of the biomass of a plant are a convenient method for obtaining information about the frequency and variability of photosynthetic carbon assimilation pathways (Kluge et al., 1993). The range in ratio values for K. pinnata indicates acquisition of external carbon by both the PEP and C<(sub)>3 pathways when external conditions such as light and water are altered. Winter and Holtum (2002) showed that decreasing night temperature also decreased night fixation of carbon dioxide. This occurs both in the laboratory and field situations (Luettge et al., 1991) and is likely to facilitate the occupation of ecological niches where such changes frequently occur (Kluge et al., 1993).

Reproductive Biology

K. pinnata reproduces both sexually and vegetatively. In vegetative reproduction, suckers are produced from the base of the plant and daughter plantlets are formed along the edges of detached leaves (epiphyllous buds). These plantlets can also be formed on the inflorescences.

The epiphyllous buds differentiate by producing two roots from the lower side, followed by a shoot from the upper side from which the first pair of leaves emerges. Root and shoot initiation occurs 9 and 12 days after leaf detachment, respectively (Sawhney et al., 1994).

Batygina et al. (1996) found that epiphyllous bud development in K. pinnata proceeds via embryoidogenesis (i.e. somatic embryos). Vegetative propagule organs arise from dormant meristem derivatives. Their development proceeds through globular, heart-shaped and torpedo-shaped stages. Comparative morphological and embryological analysis of vegetative propagules and sexual embryos revealed a strong similarity in their developmental patterns. According to Naylor (1932), the primordia for the epiphyllous buds already exist in the leaf notches. These remain dormant until conditions are favourable for plantlet formation (Yarbrough, 1932). An increased cytokinin/auxin ratio appears to be responsible for bringing about the effect. This is supported by studies on endogenous levels of these hormones (Henson and Wareing, 1977), as well as cytokinin promotion and auxin inhibition of budding when applied exogenously to K. pinnata (Mohan Ram, 1963; Houck and Rieseberg, 1983).

In southeast Queensland, Australia, K. pinnata flowers from June to September (Forster, 1985). After flowering, K. pinnata can produce vegetative buds on the inflorescences at the points of insertion of the floral pedicel (Friedmann, 1975).

Environmental Requirements

In Madagascar, K. pinnata is found in temperate and subhumid climates which receive between 1000 and 2000 mm of rain per annum (Paulian, 1984). It is relatively drought intolerant and may become almost totally defoliated during periods of extended drought (Forster, 1985). It is fairly abundant along the coast, growing in sandy soils and along granite outcrops.

In its introduced range it often occurs in similar habitats along sandy and rocky shores and estuaries (Batianoff and Franks, 1997). In Queensland, Australia, it is most commonly associated with loamy sand, alluvial soils, clay loams and skeletal soils (Queensland Herbarium database, Brisbane Botanic Gardens, Australia, 1999) and where soil moisture persists with surface organic matter.

K. pinnata cannot withstand trampling by animals and therefore only tends to thrive on rock piles and along fence lines in pastures.

Associations

A typical association for K. pinnata is at Nundah Creek near Brisbane, Australia, in a coastal heath verging on mangrove swamps. Dominant native species include Eucalyptus tereticornis, E. racemosa and Casuarina equistifolia, with Melaleuca quinquenervia and M. nodosa. Other invasive plants such as Schinus terebinthifolia, Asparagus spp., Lantana camara and Passiflora sp. also occur in this area (Sparkes et al., 2002).

Allelopathic chemicals released from roots and fallen leaves may facilitate displacement of native flora. Leaf extracts of K. pinnata inhibited the root length of Vigna radiata cuttings by 38%; inhibited the rooting rate, root number and root length of pea cuttings by 14%, 23% and 27%, respectively; and inhibited the germination rates of Brassica juncea and Brassica oleracea var. caulorapa seeds by 42% and 54%, respectively (Huang et al., 1997). The allelochemicals have been identified as p-hydroxybenzoic-, protocatechuic-, gallic-, p-coumaric and coffeic acid (Bär et al., 1997).

Latitude/Altitude Ranges

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Latitude North (°N)	Latitude South (°S)	Altitude Lower (m)	Altitude Upper (m)
		1000	2000

Rainfall Regime

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Summer

Soil Tolerances

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Soil drainage

free
seasonally waterlogged

Soil texture

heavy
light
medium

Natural enemies

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Natural enemy	Type	Life stages
Alcidodes sedi	Herbivore	Stems
Aphthona	Herbivore	Leaves
Leptomyrina phidias	Herbivore	Leaves

Notes on Natural Enemies

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K. pinnata was observed with a stem blight disease in Pune, India, caused by two fungi Colletotrichum dematium and Glomerella cingulata. The disease symptoms were visible over the entire stem as irregular, discoloured areas that later exhibited growth of punctate or slightly elongated, dark, acervular fruit bodies (Rao et al., 1989).

Kalanchoe isometric virus (KIV) was identified in crude sap of K. pinnata plants showing a mild mosaic on the leaves (Izaguirre-Mayoral et al., 1990). The virus could be transmitted mechanically to several test plants but not to healthy Bryophyllum. Ultrastructural studies of infected leaves indicated that KIV caused an increase in chloroplast volume, distortion of the grana and reduced the number of thylakoids per granum. KIV infection also impaired the diurnal pattern of CAM synthesis and hydrolysis of starch. Young Kalanchoe plantlets acquire KIV through their physical connections with infected mature leaves.

Athelia rolfsii, a leaf rot, has been isolated from leaves of K. pinnata from Calicut, India. Leaves of infected plants exhibited slightly sunken, large circular patches with abundant sclerotia during periods of heavy rainfall and high humidity (Leelavathy, 1973).

In Kiribati, K. pinnata is a recorded host of the root-knot nematode Meloidogyne sp. and the dagger nematode Xiphinema brevicolle (Orton Williams, 1980).

Three insect species, Spilosoma lutescens, Leptomyrina phidias (Lepidoptera: Lycaenidae) and an unknown Aphthona sp. (Coleoptera: Chrysomelidae) were collected on K. pinnata in Madagascar. Aphthona sp. is a leaf feeder and particularly damaging to plants in the field. It was not collected on other Kalanchoe species surveyed in the field and is considered a potential biological control agent.

Other species collected on K. pinnata in its introduced range in South Africa include Aphis sedi (Hemiptera: Aphididae), Alcidodes sedi (Coleoptera: Curculionidae), Scirtothrips aurantii (Thysanoptera: Thripidae) and an unidentified weevil (Coleoptera: Curculionidae: Conoderinae). In host range trials, Alcidodes sedi could complete its development on a number of Bryophyllum and Kalanchoe species but failed to develop on other related species in the Crassulaceae (ABR Witt, ARC-PPRI, South Africa, unpublished data, 2004).

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

Propagation is either by seeds, suckers from the base of the plant or by the formation of daughter plantlets along the edges of detached leaves (epiphyllous buds) and inflorescences. As plantlets fall to the ground, they rapidly colonize the surrounding area.

Accidental Introduction

This species has appeared along the sand dunes of Peregian Beach, Sunshine Coast, southeast Queensland, Australia, as a consequence of dumping of garden trash into bush adjoining the beach. K. pinnata was not present in transects at this site in 1982 but was present in 1997 (Batianoff and Franks, 1998). This corresponds to a period of rapid urban development in this holiday/tourist area. Plants also invade bushland from abandoned homesteads.

Intentional Introduction

Deliberate introductions of this plant are quite likely because it is such a popular garden ornamental and has medicinal properties.

Pathway Vectors

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Vector	Notes	Long Distance	Local	References
Clothing, footwear and possessions		Yes
Mail		Yes
Soil, sand and gravel		Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	Borne externally	Visibility of pest or symptoms
Flowers/Inflorescences/Cones/Calyx	seeds
Fruits (inc. pods)	seeds
Leaves
Seedlings/Micropropagated plants
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)	seeds

Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Growing medium accompanying plants
Roots
Wood

Impact Summary

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Category	Impact
Animal/plant collections	None
Animal/plant products	None
Biodiversity (generally)	Negative
Crop production	None
Environment (generally)	Negative
Fisheries / aquaculture	None
Forestry production	None
Human health	None
Livestock production	Negative
Native fauna	None
Native flora	Negative
Rare/protected species	None
Tourism	None
Trade/international relations	None
Transport/travel	None

Economic Impact

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Four species of Kalanchoe, including K. pinnata were responsible for 41 recorded poisoning incidents affecting 379 cattle in Queensland, Australia, between 1960 and 1984 (McKenzie and Armstrong, 1986). In New South Wales, Australia, two cattle reportedly died from ingestion of K. pinnata (Reppas, 1995).

In the Galapagos Islands, K. pinnata has been reported to displace crops (Soria et al., 2002). It has also been reported as a host for crop pests, including Scirtothrips aurantii, Athelia rolfsii and Xiphinema brevicolle (see Notes on Natural Enemies for more details). Other economic impacts are associated with the costs of control.

Environmental Impact

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The major impact of K. pinnata is the formation of thick stands that displace existing vegetation, reducing local biodiversity. On the island of St John, US Virgin Islands, K. pinnata forms dense stands crowding out native herbaceous vegetation (Ting, 1989), and in the Galapagos Islands, K. pinnata not only replaces the herb layer with a monospecific stand, but forms a dense carpet that inhibits the regeneration of the shrub and tree layers (Tye, 2001). Allelopathic chemicals released from roots and fallen leaves may facilitate this invasion and displacement of native flora.

Threatened Species

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Threatened Species	Conservation Status	Where Threatened	Mechanism	References
Phyllostegia knudsenii (Waimea phyllostegia)	NatureServe; USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2009a)
Cyanea recta (Kealia cyanea)	National list(s); USA ESA listing as threatened species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (1995)
Cyrtandra limahuliensis (Limahuli cyrtandra)	NatureServe; USA ESA listing as threatened species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (1995)
Isodendrion longifolium (longleaf isodendrion)	EN (IUCN red list: Endangered); USA ESA listing as threatened species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2011b)
Lobelia monostachya (Waianae Range lobelia)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (1998)
Myrsine linearifolia (narrowleaf colicwood)	EN (IUCN red list: Endangered); NatureServe; USA ESA listing as threatened species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2003)
Nototrichium humile (kaala rockwort)	EN (IUCN red list: Endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2008b)
Peucedanum sandwicense (makou)	NatureServe; USA ESA listing as threatened species	Hawaii	Competition - smothering	US Fish and Wildlife Service (2011a)
Phyllostegia mollis (Waianae Range phyllostegia)	USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2009d)
Phyllostegia renovans (red-leaf phyllostegia)	NatureServe; USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources; Competition - shading	US Fish and Wildlife Service (2010i)
Pittosporum napaulense	No Details	Hawaii	Competition - monopolizing resources; Competition - shading	US Fish and Wildlife Service (2010i)
Platydesma rostrata	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources; Competition - smothering	US Fish and Wildlife Service (2010i)
Poa mannii (Mann's bluegrass)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2010b)
Pritchardia napaliensis	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010c)
Psychotria grandiflora (large-flowered balsamo)	EN (IUCN red list: Endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010i)
Psychotria hobdyi (Hobdy's wild-coffee)	USA ESA listing as endangered species; USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010i)
Pteralyxia kauaiensis (Kauai pteralyxia)	EN (IUCN red list: Endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (1995); US Fish and Wildlife Service (2010d)
Remya kauaiensis (Kauai remya)	EN (IUCN red list: Endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010e)
Remya montgomeryi (Kalalau Valley remya)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010a)
Scaevola coriacea (dwarf naupaka)	NatureServe; USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010k)
Schiedea apokremnos (Kauai schiedea)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010l)
Schiedea hookeri (sprawling schiedea)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources; Ecosystem change / habitat alteration	US Fish and Wildlife Service (2011c)
Schiedea kauaiensis (Kauai schiedea)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2008a)
Schiedea membranacea	CR (IUCN red list: Critically endangered); NatureServe; USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2010f)
Schiedea spergulina var. leiopoda	National list(s); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2010m)
Schiedea stellarioides	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources; Ecosystem change / habitat alteration	US Fish and Wildlife Service (2010g)
Solanum sandwicense	National list(s); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2009e)
Spermolepis hawaiiensis (Hawaii scaleseed)	USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources; Ecosystem change / habitat alteration	US Fish and Wildlife Service (2010n)
Stenogyne campanulata (Kalalau Valley stenogyne)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition - monopolizing resources	US Fish and Wildlife Service (2009b)
Stenogyne kanehoana (Oahu stenogyne)	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (1998)
Tetramolopium filiforme (ridgetop tetramolopium)	USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2011a)
Urera kaalae	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Ecosystem change / habitat alteration; Pest and disease transmission	US Fish and Wildlife Service (2011d)
Wilkesia hobdyi (dwarf iliau)	CR (IUCN red list: Critically endangered); National list(s); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2010h)
Xylosma crenata	CR (IUCN red list: Critically endangered); USA ESA listing as endangered species	Hawaii	Competition (unspecified)	US Fish and Wildlife Service (2009c)

Social Impact

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The plant has no direct impact on health or social activities although dense infestations can interfere with access to amenity areas and cause a reduction in biodiversity, which affects activities such as bird watching.

Risk and Impact Factors

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Invasiveness

Proved invasive outside its native range
Highly adaptable to different environments
Highly mobile locally
Has high reproductive potential

Impact outcomes

Negatively impacts animal health
Reduced native biodiversity

Impact mechanisms

Competition - monopolizing resources
Competition - shading
Competition - smothering
Competition - strangling
Competition (unspecified)
Pest and disease transmission

Likelihood of entry/control

Highly likely to be transported internationally deliberately
Difficult to identify/detect as a commodity contaminant

Uses

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Morton (1990) has reviewed the medicinal uses of K. pinnata. In Africa, India and South America it is used for the treatment of insect bites, wounds, ulcers, abscesses and burns, inflammations, swellings and discolorations (Safford, 1905; Kirtikar and Basu, 1935; Dalziel, 1937; Dastur, 1951; Chopra et al., 1956; Boakye-Yiadom, 1977); rheumatic afflictions, erysipelas and boils (Quisumbing, 1951); smallpox, diarrhoea, dysentery, lithiasis and phthisis (Nadkarni, 1954); whitlow (Gaind and Gupta, 1971) and cholera (Siddiqui et al., 1989); mange, and as a cold remedy, especially for small children (Raponda-Walker and Sillans, 1961); bronchitis (Quisumbing, 1951); as a diuretic (Oliver-Bayer, 1986); conjunctivitis, earache and sore throats (Oliver-Bayer, 1986); and in the induction of labour and the removal of ovarian cysts (Onwuliri and Anekwe, 1992). The Caiçaras Indians of Brazil use K. pinnata for the treatment of general aches, cough, bronchitis, influenza, pneumonia and dermatitis (Begossi et al., 2002). The juice of the fresh leaves is used very effectively for the treatment of jaundice in folk medicines of Bundelkhand region of India. In west tropical Africa, the leaf juice is fed into a newborn infant's mouth, while an infusion is drunk by the child and the mother (Dalziel, 1937). In Singapore, the juice is given as a febrifuge (Burkill, 1935). Apart from making concoctions, decoctions and infusions of plant parts, some leaves are regularly eaten in soups as a tonic for the stomach and kidneys.

The many ethnobotanical uses of K. pinnata have led to investigations of its chemical composition and properties. The aqueous extract of K. pinnata yielded bryophyllin B, with activity against the in vitro growth of tissue culture cells (Yamagishi et al., 1989). The methanolic extract of K. pinnata has shown potent cytotoxicity in vitro against tumour cell cultures (Yamagishi et al., 1988), significant anti-ulcer activity in nine experimental animal models (Pal and Chaudhuri, 1991), and potent central nervous system depressant action (Pal et al., 1999). The methanolic extract of K. pinnata leaves showed significant anti-inflammatory activity on different experimental models (Pal and Chaudhuri, 1990, 1992). The juice from the leaves of K. pinnata exhibited histamine-blocking activity (Nassis et al., 1992). A methanolic extract of K. pinnata leaves has been studied for anti-inflammatory, analgesic and antipyretic activities in animal models. The extract produced a significant inhibition of carrageenan-induced paw oedema, a significant reduction in cotton pellet granuloma in rats, an inhibition of acetic acid induced writhing in mice, and a significant dose-dependent reduction of yeast-induced pyrexia (Olajide et al., 1998).

The juice of leaves of K. pinnata has antibiotic activity against Staphylococcus aureus, Bacillus subtilis, Escherichia coli and Pseudomonas aeruginosa (Boakye-Yiadom, 1977). The juice was also bactericidal to Streptococcus pyrogenes, Streptococcus faecalis, Proteus spp., Klebsiella spp., Shigella spp., Salmonella spp. and Serratia marcescens, including the clinical isolates of these organisms possessing multiple antibiotic resistances (Obaseiki-Ebor, 1985).

The methanolic extract of leaves had activity against Staphylococcus aureus, Bacillus subtilis, Escherichia coli, Shigella dysenteriae and Proteus vulgaris, but not against Pseudomonas aeruginosa or Klebsiella pneumoniae (Akinpelu, 2000). Ethanolic extract had activity against seven bacteria (Staphylococcus aureus, Salmonella typhimurium, S. paratyphi,S. typhi, E. coli, Shigella dysenteriae and Pseudomonas aeruginosa) and two filamentous fungi (Alternaria alternata and Rhizoctonia bataticola [Macrophomina phaseolina]), and a yeast Candida albicans. The ethyl acetate extract also showed appreciable antimutagenic activity (Obaseiki-Ebor et al., 1993).

It has been shown that oral K. pinnata leaf extract is strongly effective against murine leishmaniasis, a tropical protozoal disease (Torres-Santos et al., 2003).

K. pinnata has shown insecticidal properties related to a bufadienolide constituent (Supratman et al., 2000).

Onwuliri and Anekwe (1992) found that K. pinnata is a rich source of some elements that are important in the maintenance of human health. This, plus the physiological activities mentioned above, may explain the traditional medicinal roles attributed to the plant.

Uses List

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General

Ornamental

Medicinal, pharmaceutical

Traditional/folklore

Similarities to Other Species/Conditions

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Boiteau and Allorge-Boiteau (1995) place K. pinnata in Group IX Proliferae with eight other closely related species, two of which have pinnate leaves and may be confused with K. pinnata. These species are Bryophyllum rubellum and B. proliferum.

Prevention and Control

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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.

Mechanical Control

K. pinnata can be controlled manually provided that the whole plant, especially the leaves and roots, are removed completely (Soria et al., 2002). A hand weeding exercise to remove K. pinnata was costed at Au$2095/ha (Sparkes et al., 2002). It was noted that the difficulty in removing plants with their root systems intact and the avoidance of leaf fall made this method less effective and more expensive than treatment with 2,4-D (approximately Aus$160/ha).

Chemical Control

In efficacy tests (Sparkes et al., 2002), 2,4-D and fluroxypyr gave best control of K. pinnata with >90% kill rates. Metsulfuron and triclopyr/picloram did not give good control. Sparkes et al. (2002) conjectured that these poor results may have been due to insufficient wetting agent and poor penetration of these herbicides through the plant's waxy cuticle. Sparkes et al. (2002) recommended 2,4-D for K. pinnata control because this has the lowest cost, non-target damage and environmental impact of the herbicides trialled.

Foliar application of glyphosate is used for the control of K. pinnata in the Galapagos Islands (Soria et al., 2002).

References

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US Fish and Wildlife Service, 2008. Kulu`i (Nototrichium humile). 5-Year Review. In: Kulu`i (Nototrichium humile). 5-Year Review : US Fish and Wildlife Service.11 pp.

US Fish and Wildlife Service, 2009. 5-Year Review, Short Form Summary: Species Reviewed: Phyllostegia knudsenii (no common name). In: 5-Year Review, Short Form Summary: Species Reviewed: Phyllostegia knudsenii (no common name) : US Fish and Wildlife Service.6 pp.

US Fish and Wildlife Service, 2009. 5-Year Review, Short Form Summary: Species Reviewed: Stenogyne campanulata (no common name). In: 5-Year Review, Short Form Summary: Species Reviewed: Stenogyne campanulata (no common name) : US Fish and Wildlife Service.6 pp.

US Fish and Wildlife Service, 2009. 5-Year Review, Short Form Summary: Xylosma crenatum (no common name). In: 5-Year Review, Short Form Summary: Xylosma crenatum (no common name) : US Fish and Wildlife Service.7 pp.

US Fish and Wildlife Service, 2009. Phyllostegia mollis (no common name). 5-Year Review: Summary and Evaluation. In: Phyllostegia mollis (no common name). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.13 pp.

US Fish and Wildlife Service, 2009. Solanum sandwicense (Popolo 'aiakeakua). 5-Year Review: Summary and Evaluation. In: Solanum sandwicense (Popolo 'aiakeakua). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.13 pp.

US Fish and Wildlife Service, 2010. 5-Year Review Short Form Summary: Remya montgomeryi (no common name). In: 5-Year Review Short Form Summary: Remya montgomeryi (no common name) : US Fish and Wildlife Service.7 pp. https://ecos.fws.gov/docs/five_year_review/doc3343.pdf

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Poa mannii (Mann's bluegrass). In: 5-Year Review, Short Form Summary: Species Reviewed: Poa mannii (Mann's bluegrass) : US Fish and Wildlife Service.10 pp.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Pritchardia napaliensis (loulu palm). In: 5-Year Review, Short Form Summary: Species Reviewed: Pritchardia napaliensis (loulu palm) : US Fish and Wildlife Service.10 pp.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Pteralyxia kauaiensis (kaulu). In: 5-Year Review, Short Form Summary: Species Reviewed: Pteralyxia kauaiensis (kaulu) : US Fish and Wildlife Service.9 pp.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Remya kauaiensis (no common name). In: 5-Year Review, Short Form Summary: Species Reviewed: Remya kauaiensis (no common name) : US Fish and Wildlife Service.12 pp.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Schiedea membranacea (no common name). In: 5-Year Review, Short Form Summary: Species Reviewed: Schiedea membranacea (no common name) : US Fish and Wildlife Service.9 pp.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Species Reviewed: Schiedea stellarioides (no common name). In: 5-Year Review, Short Form Summary: Species Reviewed: Schiedea stellarioides (no common name) : US Fish and Wildlife Service.7 pp.

US Fish and Wildlife Service, 2010. 5-Year Review, Short Form Summary: Wilkesia hobdyi (dwarf iliau). In: 5-Year Review, Short Form Summary: Wilkesia hobdyi (dwarf iliau) : US Fish and Wildlife Service.8 pp.

US Fish and Wildlife Service, 2010. Determination of Endangered Status for 48 Species on Kauai and designation of Critical Habitat: Final Rule. In: Determination of Endangered Status for 48 Species on Kauai and designation of Critical Habitat: Final Rule : US Fish and Wildlife Service.i + 205 pp.

US Fish and Wildlife Service, 2010. Recovery Outline for the Kauai Ecosystem. In: Recovery Outline for the Kauai Ecosystem : US Fish and Wildlife Service.38 pp. + 3 maps.

US Fish and Wildlife Service, 2010. Scaevola coriacea (dwarf naupaka). 5-Year Review: Summary and Evaluation. In: Scaevola coriacea (dwarf naupaka). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.19 pp.

US Fish and Wildlife Service, 2010. Schiedea apokremnos (maolioli). 5-Year Review: Summary and Evaluation. In: Schiedea apokremnos (maolioli). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.16 pp.

US Fish and Wildlife Service, 2010. Schiedea spergulina var. leiopoda (no common name). 5-Year Review: Summary and Evaluation. In: Schiedea spergulina var. leiopoda (no common name). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.11 pp.

US Fish and Wildlife Service, 2010. Spermolepis hawaiiensis (no common name). 5-Year Review: Summary and Evaluation. In: Spermolepis hawaiiensis (no common name). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.19 pp.

US Fish and Wildlife Service, 2010. Tetramolopium filiforme (no common name). 5-Year Review: Summary and Evaluation. In: Tetramolopium filiforme (no common name). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.11 pp.

US Fish and Wildlife Service, 2011. Endangered and Threatened Wildlife and Plants; Listing 23 Species on Oahu as Endangered and Designating Critical Habitat for 124 Species. In: Federal Register , 76(148) : US Fish and Wildlife Service.46362-46593. http://www.gpo.gov/fdsys/pkg/FR-2011-08-02/pdf/2011-17162.pdf

US Fish and Wildlife Service, 2011. Isodendrion longifolium (aupaka). 5-Year Review: Summary and Evaluation. In: Isodendrion longifolium (aupaka). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.28 pp. http://ecos.fws.gov/docs/five_year_review/doc3809.pdf

US Fish and Wildlife Service, 2011. Schiedea hookeri (no common name). 5-Year Review: Summary and Evaluation. In: Schiedea hookeri (no common name). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.20 pp.

US Fish and Wildlife Service, 2011. Urera kaalae (opuhe). 5-Year Review: Summary and Evaluation. In: Urera kaalae (opuhe). 5-Year Review: Summary and Evaluation : US Fish and Wildlife Service.19 pp.

USDA-ARS, 2004. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

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Links to Websites

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Website	URL	Comment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway	https://doi.org/10.5061/dryad.m93f6	Data source for updated system data added to species habitat list.

Contributors

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22/05/13 Updated by:

Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

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Kalanchoe pinnata (cathedral bells)

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