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'''''Ceodes umbellifera''''', [[Synonym (taxonomy)|synonym]] '''''Pisonia umbellifera''''', commonly known as the '''birdlime tree'''<ref name="APNI">{{APNI| name = Pisonia umbellifera| id = 44631| accessdate = April 28, 2016}}</ref> or '''bird catcher tree,''' is a species of [[plant]] in the [[Nyctaginaceae]] family. It grows throughout the tropical Indo-Pacific. It is native to the [[Andaman Islands]], [[Indonesia]], [[Malaysia]], the [[Philippines]], [[Thailand]], [[Vietnam]], [[China]], [[Taiwan]], [[Hawaii]], [[Africa]] and [[Madagascar]] and the states of [[New South Wales]] and [[Queensland]] in [[Australia]].<ref name=FOC>{{cite web|title=''Pisonia umbellifera''|work=Flora of China|url=http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=242338473|accessdate=2009-04-14}}</ref><ref name=NSW>{{cite web |url=http://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=sp&name=Pisonia~umbellifera |title=''Pisonia umbellifera''|accessdate=2009-04-14 |author= Jacobs, S.W.L.|author2= G. J. Harden|work= PlantNET - New South Wales Flora Online |publisher=Royal Botanic Gardens & Domain Trust, Sydney Australia}}</ref> A variegated form is widely cultivated in frost-free climates.
'''''Ceodes umbellifera''''', [[Synonym (taxonomy)|synonym]] '''''Pisonia umbellifera''''', commonly known as the '''birdlime tree'''<ref name="APNI">{{APNI| name = Pisonia umbellifera| id = 44631| accessdate = April 28, 2016}}</ref> or '''bird catcher tree,''' is a species of [[plant]] in the [[Nyctaginaceae]] family. It grows throughout the tropical Indo-Pacific. It is native to the [[Andaman Islands]], [[Indonesia]], [[Malaysia]], the [[Philippines]], [[Thailand]], [[Vietnam]], [[China]], [[Taiwan]], [[Hawaii]], [[Africa]] and [[Madagascar]] and the states of [[New South Wales]] and [[Queensland]] in [[Australia]].<ref name=FOC>{{cite web|title=''Pisonia umbellifera''|work=Flora of China|url=http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=242338473|accessdate=2009-04-14}}</ref><ref name=NSW>{{cite web |url=http://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=sp&name=Pisonia~umbellifera |title=''Pisonia umbellifera''|accessdate=2009-04-14 |author= Jacobs, S.W.L.|author2= G. J. Harden|work= PlantNET - New South Wales Flora Online |publisher=Royal Botanic Gardens & Domain Trust, Sydney Australia}}</ref> A variegated form is widely cultivated in frost-free climates.
[[Image:Pisonia umbellifera variegata.jpg|thumb| ''P. umbellifera'' 'Variegata']]
[[Image:Pisonia umbellifera variegata.jpg|thumb| ''P. umbellifera'' 'Variegata']]
Its former genus, ''Pisonia'', is named after a Dutch scientist, [[Willem Piso]], and ''umbellifera'' is derived from Latin ''umbelliferum'', denoting the species' big, 'shade-carrying' foliage.<ref name=":0">{{Cite web|title=Native Plants Hawaii|url=http://nativeplants.hawaii.edu/|access-date=2021-04-13|website=nativeplants.hawaii.edu}}</ref>{{clear left}}
Its former genus, ''[[Pisonia]]'', is named after a Dutch scientist, [[Willem Piso]], and ''umbellifera'' is derived from Latin ''umbelliferum'', denoting the species' big, 'shade-carrying' foliage.<ref name=":0">{{Cite web|title=Native Plants Hawaii|url=http://nativeplants.hawaii.edu/|access-date=2021-04-13|website=nativeplants.hawaii.edu}}</ref>{{clear left}}


== Description ==
== Description ==
''Ceodes umbellifera'' is a shrub with large, medium green leaves. Other variegated varieties exist (''Ceodes umbellifera'' 'Variegata') with marbling of white, light and dark green on the shrub's leaves. The tree's elliptic leaves may be between 6 to 20cm long, and 4 to 10cm wide.<ref name=":1">{{Cite web|last=National Herbarium of New South Wales|date=2004|title=Pisonia umbellifera (J.R.Forst. & G.Forst.) Seem|url=https://plantnet.rbgsyd.nsw.gov.au/cgibin/NSWfl.plpage=nswfl&lvl=sp&name=Pisonia~umbellifera|url-status=live|access-date=15 April 2021|website=PlantNET FloraOnline}}</ref>
''Ceodes umbellifera'' is a shrub with large, medium green leaves. Other variegated varieties exist (''Ceodes umbellifera'' 'Variegata') with marbling of white, light and dark green on the shrub's leaves. The tree's elliptic leaves may be between 6 to 20cm long, and 4 to 10cm wide.<ref name=":1">{{Cite web|last=National Herbarium of New South Wales|date=2004|title=Pisonia umbellifera (J.R.Forst. & G.Forst.) Seem|url=https://plantnet.rbgsyd.nsw.gov.au/cgibin/NSWfl.plpage=nswfl&lvl=sp&name=Pisonia~umbellifera|url-status=live|access-date=15 April 2021|website=PlantNET FloraOnline}}</ref>


There are discrepancies between sources regarding the height of the shrub. Most state that ''Ceodes umbellifera'' will grow to between 4 and 12 metres.<ref name=":0" /><ref name=":2">{{Cite web|last=Fern|first=K|date=2019|title=''Pisonia umbellifera''|url=http://tropical.theferns.info/viewtropical.php?id=Pisonia+umbellifera|url-status=live|access-date=15 April 2021|website=Useful Tropical Plants}}</ref> Some sources suggest the tallest known specimen is 20 metres tall,<ref name=":3">{{Cite web|last=Pucci, P. & Beltramini, M.|date=2003|title=''Pisonia umbellifera''|url=21 April 2021|url-status=live|website=Monaco Nature Encyclopedia}}</ref> whilst another reports a height of 28 metres.<ref name=":2" /> Once mature, the tree has a spreading, rounded crown5 which may cover over 20 feet.<ref name=":0" />
There are discrepancies between sources regarding the height of the shrub. Most state that ''Ceodes umbellifera'' will grow to between 4 and 12 metres.<ref name=":0" /><ref name=":2">{{Cite web|last=Fern|first=K|date=2019|title=''Pisonia umbellifera''|url=http://tropical.theferns.info/viewtropical.php?id=Pisonia+umbellifera|url-status=live|access-date=15 April 2021|website=Useful Tropical Plants}}</ref> Some sources suggest the tallest known specimen is 20 metres tall,<ref name=":3">{{Cite web|last=Pucci, P. & Beltramini, M.|date=2003|title=''Pisonia umbellifera''|url=21 April 2021|url-status=live|website=Monaco Nature Encyclopedia}}</ref> whilst another reports a height of 28 metres.<ref name=":2" /> Once mature, the tree has a spreading, rounded crown which may cover over 20 feet.<ref name=":0" />


In summer, the tree will start to bud and produce flowers. Before blooming, the buds of the flower stem from a complex [[inflorescence]].<ref name=":1" /> Buds are around 6cm in height, and have a pale brown, chickpea-like appearance.<ref name=":1" /> The flowers consist of small pink or yellow petals in a rounded funnel formation, with short white stems protruding from the centre.<ref name=":3" /> The flowers are very fragrant.
In summer, the tree will start to bud and produce flowers. Before blooming, the buds of the flower stem from a complex [[inflorescence]].<ref name=":1" /> Buds are around 6cm in height, and have a pale brown, chickpea-like appearance.<ref name=":1" /> The flowers consist of small pink or yellow petals in a rounded funnel formation, with short white stems protruding from the centre.<ref name=":3" />


After flowering, the shrub produces small fruit.<ref name="NSW" /> These are the colloquial namesake of the tree, which is also known as '[[birdlime]]';<ref name=":0" /> due to the fruit's secretion of a sticky substance, small birds may often become ensnared in this substance and are unable to fly away.<ref name=":4">{{Cite journal|last=Priddel, D. & Carlile, N.|date=1995|title=Mortality of Adult Gould's Petrels ''Pterodroma leucoptera leucoptera'' at the Nesting Site on Cabbage Tree Island, New South Wales|journal=Emu - Austral Ornithology|volume=95|issue=4|pages=259–264|doi=10.1071/MU9950259}}</ref> The viscid fruit are dark brown and purple, cylindrical in shape and about 2-4 cm long.<ref name=":1" /> They are vertically ribbed. Both the leaves and fruit are hairless.<ref name=":1" />
After flowering, the shrub produces small fruit.<ref name="NSW" /> These are the colloquial namesake of the tree, which is also known as '[[birdlime]]';<ref name=":0" /> due to the fruit's secretion of a sticky substance, small birds may often become ensnared in this substance and are unable to fly away.<ref name=":4">{{Cite journal|last=Priddel, D. & Carlile, N.|date=1995|title=Mortality of Adult Gould's Petrels ''Pterodroma leucoptera leucoptera'' at the Nesting Site on Cabbage Tree Island, New South Wales|journal=Emu - Austral Ornithology|volume=95|issue=4|pages=259–264|doi=10.1071/MU9950259}}</ref> The viscid fruit are dark brown and purple, cylindrical in shape and about 2-4 cm long.<ref name=":1" /> They are vertically ribbed. Both the leaves and fruit are hairless.<ref name=":1" />
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== Taxonomy ==
== Taxonomy ==
The species was first described by [[Johann Reinhold Forster]] and [[Georg Forster]] in 1776 as ''Ceodes umbellifera''.<ref name=":11">{{Cite journal|last=Rossetto, E. F. & Caraballo, M.|date=2019|title=Splitting the Pisonia birdcatcher trees: re-establishment of Ceodes and Rockia (Nyctaginaceae, Pisonieae)|url=https://www.researchgate.net/publication/342721410_Splitting_the_Pisonia_birdcatcher_trees_re-establishment_of_Ceodes_and_Rockia_Nyctaginaceae_Pisonieae|journal=PhytoKeys|volume=152(3)|pages=121-136}}</ref><ref name=":13">{{Cite book|last=Forster, J. R. & Forster, G.|title=Characteres Generum Plantarum|year=1776|location=B. White, T. Cadell, & P. Elmsly, London.}}</ref> The genus ''Pisonia'' of the Pisonieae tribe has a "convoluted taxonomic history"<ref name=":11" /> that has involved ''Ceodes umbellifera'' being moved at various times to the genus ''Pisonia''.
The species was first described by [[Johann Reinhold Forster]] and [[Georg Forster]] in 1776 as ''Ceodes umbellifera''.<ref name=":11">{{Cite journal|last=Rossetto, E. F. & Caraballo, M.|date=2019|title=Splitting the Pisonia birdcatcher trees: re-establishment of Ceodes and Rockia (Nyctaginaceae, Pisonieae)|url=https://www.researchgate.net/publication/342721410_Splitting_the_Pisonia_birdcatcher_trees_re-establishment_of_Ceodes_and_Rockia_Nyctaginaceae_Pisonieae|journal=PhytoKeys|volume=152(3)|pages=121-136}}</ref><ref name=":13">{{Cite book|last=Forster, J. R. & Forster, G.|title=Characteres Generum Plantarum|year=1776|location=B. White, T. Cadell, & P. Elmsly, London.}}</ref> The genus ''Pisonia'' of the Pisonieae tribe has a "convoluted taxonomic history"<ref name=":11" /> that has involved ''Ceodes umbellifera'' being moved at various times to the genus ''Pisonia''.
''Ceodes umbellifera'' was the first species placed in the genus ''Ceodes''.<ref name=":11" /> It was first discovered in 1776 by J.R. Forster and G. Forster in Vanuatu.<ref name=":11" /> It was distinguished from other like species by the "absence of stalked glands along the ribs of anthocarps".<ref name=":11" />. Forster and Forster placed the species in ''Ceodes'' rather than ''Pisonia''.<ref name=":14">{{Cite journal|last=Skottsberg, C.|date=1926|title=Vascular plants from the Hawaiian Islands|journal=Acta Horti Gothoburgensis|volume=2|pages=185–284}}</ref><ref name=":11" /><ref name=":13" />
''Ceodes umbellifera'' was the first species placed in the genus ''[[Ceodes]]''.<ref name=":11" /> It was first discovered in 1776 by J.R. Forster and G. Forster in Vanuatu.<ref name=":11" /> It was distinguished from other like ,species by the "absence of stalked glands along the ribs of [[Anthocarp|anthocarps]]".<ref name=":11" />. Forster and Forster placed the species in ''Ceodes'' rather than ''Pisonia''.<ref name=":14">{{Cite journal|last=Skottsberg, C.|date=1926|title=Vascular plants from the Hawaiian Islands|journal=Acta Horti Gothoburgensis|volume=2|pages=185–284}}</ref><ref name=":11" /><ref name=":13" />


In 1804, Du Petit-Thouars found similar species in Mauritius which also lacked glands on their anthocarps.<ref name=":11" /> He classified them under a new genera ''Calpidia'', which modern botanists suggest essentially had the same characteristics as ''Ceodes''.<ref name=":11" /> It is likely that, as Elson Felipe Rossetto and Marcos Caraballo suggest, Du Petit-Thouars was not aware of the description and categorisation of ''Ceodes'' "28 years earlier".<ref name=":11" /> An 1880 classification by Bentham and Hooker retained both ''Ceodes'' and ''Calipidia'' as genera interchangable with ''Pisonia''.<ref>{{Cite book|last=Bentham, G. & Hooker, J. D.|title=Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita|publisher=Reeves & Co., London.|year=1880}}</ref><ref name=":12">{{Cite journal|last=Rossetto, E. F., Faria, A. D., Ruas, P. M., Ruas, C., Douglas, N. A.,& Ribeiro, J.|date=2019|title=Clarifying generic delimitation in Nyctaginaceae tribe Pisonieae after more than a century of taxonomic confusion.|url=|journal=Botanical Journal of the Linnean Society|volume=189(4)|pages=378-396}}</ref> They introduced ''Timeroyea'' as a new genus apart from these, characterised by "inconspicuous anthrocarpal glands"<ref name=":11" /> but with more than 25 stamens.<ref name=":11" />
In 1804, [[Louis-Marie Aubert du Petit-Thouars|Du Petit-Thouars]] found similar species in [[Mauritius]] which also lacked glands on their anthocarps.<ref name=":11" /> He classified them under a new genera ''Calpidia'', which modern botanists suggest essentially had the same characteristics as ''Ceodes''.<ref name=":11" /> It is likely that, as Elson Felipe Rossetto and Marcos Caraballo suggest, Du Petit-Thouars was not aware of the description and categorisation of ''Ceodes'' "28 years earlier".<ref name=":11" /> An 1880 classification by [[George Bentham|Bentham]] and [[Joseph Dalton Hooker|Hooker]] retained both ''Ceodes'' and ''Calipidia'' as genera interchangable with ''Pisonia''.<ref>{{Cite book|last=Bentham, G. & Hooker, J. D.|title=Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita|publisher=Reeves & Co., London.|year=1880}}</ref><ref name=":12">{{Cite journal|last=Rossetto, E. F., Faria, A. D., Ruas, P. M., Ruas, C., Douglas, N. A.,& Ribeiro, J.|date=2019|title=Clarifying generic delimitation in Nyctaginaceae tribe Pisonieae after more than a century of taxonomic confusion.|url=|journal=Botanical Journal of the Linnean Society|volume=189(4)|pages=378-396}}</ref> They introduced ''Timeroyea'' as a new genus apart from these, characterised by "inconspicuous anthrocarpal glands"<ref name=":11" /> but with more than 25 stamens.<ref name=":11" />


In 1866, [[Berthold Carl Seemann]] transferred it to the genus ''[[Pisonia]]'' as ''Pisonia umbellifera'', along with the other species placed in ''Ceodes''.<ref>{{Cite journal|last=Seemann, B.|date=1863|title=On the genus Ceodes of Forster|journal=Journal of Botany, British and Foreign|volume=1|pages=244–246}}</ref> Subsequent work by Heirmel in 1889 specified ''Timeroyae'' as a subcategory of ''Pisonia'', with another new subcategory ''Prismatocarpae'' introduced, for specimens with fewer than 15 stamens.<ref name=":11" /> Rossetto and Caraballo note that, "in this treatment, ''Calpidia'' was listed as a synonym of ''Pisonia'', while ''Ceodes'' was not mentioned at all".<ref name=":11" />
In 1866, [[Berthold Carl Seemann]] transferred it to the genus ''[[Pisonia]]'' as ''Pisonia umbellifera'', along with the other species placed in ''Ceodes''.<ref>{{Cite journal|last=Seemann, B.|date=1863|title=On the genus Ceodes of Forster|journal=Journal of Botany, British and Foreign|volume=1|pages=244–246}}</ref> Subsequent work by Heirmel in 1889 specified ''Timeroyae'' as a subcategory of ''Pisonia'', with another new subcategory ''Prismatocarpae'' introduced, for specimens with fewer than 15 stamens.<ref name=":11" /> Rossetto and Caraballo note that, "in this treatment, ''Calpidia'' was listed as a synonym of ''Pisonia'', while ''Ceodes'' was not mentioned at all".<ref name=":11" />


Heirmel revised this in 1913, splitting ''Calpidia'' and ''Pisonia'', with the former to include ''Prismatocarpae'' and ''Timeroyae'', both of which did not have anthrocarpal glands.<ref>{{Cite journal|last=Heimerl, A.|date=1913|title=Die Nyctaginaceen-Gattungen Calpidia und Rockia|journal=Österreichische Botanische Zeitschrift|volume=63(7)|pages=279–290}}</ref> ''Calpidia'' and ''Pisonia'' were distinguished by "the absence of bracteoles, having a reduced perisperm that forms gelatinous traces and starch accumulation in the embryo".<ref name=":11" />
[[Anton Heimerl|Heirmel]] revised this in 1913, splitting ''Calpidia'' and ''Pisonia'', with the former to include ''Prismatocarpae'' and ''Timeroyae'', both of which did not have anthrocarpal glands.<ref>{{Cite journal|last=Heimerl, A.|date=1913|title=Die Nyctaginaceen-Gattungen Calpidia und Rockia|journal=Österreichische Botanische Zeitschrift|volume=63(7)|pages=279–290}}</ref> ''Calpidia'' and ''Pisonia'' were distinguished by "the absence of [[Bract|bracteoles]], having a reduced [[Ovule|perisperm]] that forms gelatinous traces and starch accumulation in the embryo".<ref name=":11" />


In a 1926 paper, Skottsberg suggested ''Ceodes'' as a genus instead of ''Calpidia'',<ref name=":14" /> using an argument, described by Rossetto and Caraballo, "that there were extant original specimens and that the scant description of the genus was similar to other names published during the late 18th century".<ref name=":11" /> However this work<ref name=":14" /> ignored the specimen ''umbellifera''. Skottsberg's later research, published in 1936, acknowledged only one specific ''Ceodes'' type, ''Ceodes umbellifera'',<ref>{{Cite journal|last=Skottsberg, C.|date=1936|title=The arboreous Nyctaginaceae of Hawaii|journal=Svensk Botanisk Tidskrift|volume=30|pages=722–743}}</ref> but also including "''Ce. excelsa'' and ''Ce. forsteriana'' as synonyms".<ref name=":11" /> Heimerl’s 1937 work also included this change but featured ''Ceodes umbellifera'' as the only ''Ceodes'' taxon.<ref name=":11" />
In a 1926 paper, [[Carl Skottsberg|Skottsberg]] suggested ''Ceodes'' as a genus instead of ''Calpidia'',<ref name=":14" /> using an argument, described by Rossetto and Caraballo, "that there were extant original specimens and that the scant description of the genus was similar to other names published during the late 18th century".<ref name=":11" /> However this work<ref name=":14" /> ignored the specimen ''umbellifera''. Skottsberg's later research, published in 1936, acknowledged only one specific ''Ceodes'' type, ''Ceodes umbellifera'',<ref>{{Cite journal|last=Skottsberg, C.|date=1936|title=The arboreous Nyctaginaceae of Hawaii|journal=Svensk Botanisk Tidskrift|volume=30|pages=722–743}}</ref> but also including "''Ce. excelsa'' and ''Ce. forsteriana'' as synonyms".<ref name=":11" /> Heimerl’s 1937 work also included this change but featured ''Ceodes umbellifera'' as the only ''Ceodes'' taxon.<ref name=":11" />


In 1964, Stemmerik proposed a change which reincorporated ''Ceodes'' into ''Pisonia''.<ref name=":12" /> Several morphological traits were omitted from this classification, such as "the absence of bracts and bracteoles at the upper portion of the pedicels, presence of starch in the embryo and number of apertures in pollen grains".<ref name=":11" /> Stemmerik suggested that since not all of these were constant traits then they should not be regarded in the classification.<ref name=":11" />
In 1964, Stemmerik proposed a change which reincorporated ''Ceodes'' into ''Pisonia''.<ref name=":12" /> Several morphological traits were omitted from this classification, such as "the absence of bracts and bracteoles at the upper portion of the [[Pedicel (botany)|pedicels]], presence of starch in the embryo and number of apertures in [[Pollen|pollen grains]]".<ref name=":11" /> Stemmerik suggested that since not all of these were constant traits then they should not be regarded in the classification.<ref name=":11" />


In 2019, a study by Rossetto and colleagues used the molecular data of the species' DNA to reveal that the ''Pisonia'' genus was non-monophyletic.<ref name=":12" /> It was suggested to resurrect the genera ''Rockia'' and ''Ceodes'' in replacement of ''Pisonia''.<ref name=":11" /> This rendered the genera monophyletic. In 2020, it was generally accepted that ''Ceodes'' should be restored, including ''Ceodes umbellifera.''<ref name=":11" /> This placement is accepted by [[Plants of the World Online]], as of May 2021.<ref>{{Cite web|title=Ceodes umbellifera J.R.Forst. & G.Forst. {{!}} Plants of the World Online {{!}} Kew Science|url=http://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:604437-1|access-date=2021-05-29|website=Plants of the World Online|language=en}}</ref>
In 2019, a study by Rossetto and colleagues used the molecular data of the species' DNA to reveal that the ''Pisonia'' genus was [[Monophyly|non-monophyletic]].<ref name=":12" /> It was suggested to resurrect the genera ''Rockia'' and ''Ceodes'' in replacement of ''Pisonia''.<ref name=":11" /> This rendered the genera monophyletic. In 2020, it was generally accepted that ''Ceodes'' should be restored, including ''Ceodes umbellifera.''<ref name=":11" /> This placement is accepted by [[Plants of the World Online]], as of May 2021.<ref>{{Cite web|title=Ceodes umbellifera J.R.Forst. & G.Forst. {{!}} Plants of the World Online {{!}} Kew Science|url=http://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:604437-1|access-date=2021-05-29|website=Plants of the World Online|language=en}}</ref>


== Distribution and habitat ==
== Distribution and habitat ==

Revision as of 23:13, 29 May 2021

Ceodes umbellifera
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Nyctaginaceae
Genus: Ceodes
Species:
C. umbellifera
Binomial name
Ceodes umbellifera
(J. R. Forster & G. Forster) Seemann
Synonyms[1]
  • Cedrota guianensis Blanco
  • Labordia fauriei H.Lév.
  • Bougainvillea racemosa Blanco
  • Calpidia excelsa (Blume) Heimerl
  • Calpidia nishimurae (Koidz.) Rehder & E.H.Wilson
  • Calpidia pancheriana Heimerl
  • Ceodes excelsa (Blume) Skottsb.
  • Pisonia aruensis Barg.-Petr.
  • Pisonia excelsa Blume
  • Pisonia gammillii Merr.
  • Pisonia macrocarpa C.Presl
  • Pisonia mooriana F.Muell.
  • Pisonia nishimurae Koidz.
  • Pisonia umbellata Seem.
  • Pisonia umbellifera (J.R.Forst. & G.Forst.) Seem.
  • Pisonia viscosa Seem.

Ceodes umbellifera, synonym Pisonia umbellifera, commonly known as the birdlime tree[2] or bird catcher tree, is a species of plant in the Nyctaginaceae family. It grows throughout the tropical Indo-Pacific. It is native to the Andaman Islands, Indonesia, Malaysia, the Philippines, Thailand, Vietnam, China, Taiwan, Hawaii, Africa and Madagascar and the states of New South Wales and Queensland in Australia.[3][4] A variegated form is widely cultivated in frost-free climates.

P. umbellifera 'Variegata'

Its former genus, Pisonia, is named after a Dutch scientist, Willem Piso, and umbellifera is derived from Latin umbelliferum, denoting the species' big, 'shade-carrying' foliage.[5]

Description

Ceodes umbellifera is a shrub with large, medium green leaves. Other variegated varieties exist (Ceodes umbellifera 'Variegata') with marbling of white, light and dark green on the shrub's leaves. The tree's elliptic leaves may be between 6 to 20cm long, and 4 to 10cm wide.[6]

There are discrepancies between sources regarding the height of the shrub. Most state that Ceodes umbellifera will grow to between 4 and 12 metres.[5][7] Some sources suggest the tallest known specimen is 20 metres tall,[8] whilst another reports a height of 28 metres.[7] Once mature, the tree has a spreading, rounded crown which may cover over 20 feet.[5]

In summer, the tree will start to bud and produce flowers. Before blooming, the buds of the flower stem from a complex inflorescence.[6] Buds are around 6cm in height, and have a pale brown, chickpea-like appearance.[6] The flowers consist of small pink or yellow petals in a rounded funnel formation, with short white stems protruding from the centre.[8]

After flowering, the shrub produces small fruit.[4] These are the colloquial namesake of the tree, which is also known as 'birdlime';[5] due to the fruit's secretion of a sticky substance, small birds may often become ensnared in this substance and are unable to fly away.[9] The viscid fruit are dark brown and purple, cylindrical in shape and about 2-4 cm long.[6] They are vertically ribbed. Both the leaves and fruit are hairless.[6]

Taxonomy

The species was first described by Johann Reinhold Forster and Georg Forster in 1776 as Ceodes umbellifera.[10][11] The genus Pisonia of the Pisonieae tribe has a "convoluted taxonomic history"[10] that has involved Ceodes umbellifera being moved at various times to the genus Pisonia. Ceodes umbellifera was the first species placed in the genus Ceodes.[10] It was first discovered in 1776 by J.R. Forster and G. Forster in Vanuatu.[10] It was distinguished from other like ,species by the "absence of stalked glands along the ribs of anthocarps".[10]. Forster and Forster placed the species in Ceodes rather than Pisonia.[12][10][11]

In 1804, Du Petit-Thouars found similar species in Mauritius which also lacked glands on their anthocarps.[10] He classified them under a new genera Calpidia, which modern botanists suggest essentially had the same characteristics as Ceodes.[10] It is likely that, as Elson Felipe Rossetto and Marcos Caraballo suggest, Du Petit-Thouars was not aware of the description and categorisation of Ceodes "28 years earlier".[10] An 1880 classification by Bentham and Hooker retained both Ceodes and Calipidia as genera interchangable with Pisonia.[13][14] They introduced Timeroyea as a new genus apart from these, characterised by "inconspicuous anthrocarpal glands"[10] but with more than 25 stamens.[10]

In 1866, Berthold Carl Seemann transferred it to the genus Pisonia as Pisonia umbellifera, along with the other species placed in Ceodes.[15] Subsequent work by Heirmel in 1889 specified Timeroyae as a subcategory of Pisonia, with another new subcategory Prismatocarpae introduced, for specimens with fewer than 15 stamens.[10] Rossetto and Caraballo note that, "in this treatment, Calpidia was listed as a synonym of Pisonia, while Ceodes was not mentioned at all".[10]

Heirmel revised this in 1913, splitting Calpidia and Pisonia, with the former to include Prismatocarpae and Timeroyae, both of which did not have anthrocarpal glands.[16] Calpidia and Pisonia were distinguished by "the absence of bracteoles, having a reduced perisperm that forms gelatinous traces and starch accumulation in the embryo".[10]

In a 1926 paper, Skottsberg suggested Ceodes as a genus instead of Calpidia,[12] using an argument, described by Rossetto and Caraballo, "that there were extant original specimens and that the scant description of the genus was similar to other names published during the late 18th century".[10] However this work[12] ignored the specimen umbellifera. Skottsberg's later research, published in 1936, acknowledged only one specific Ceodes type, Ceodes umbellifera,[17] but also including "Ce. excelsa and Ce. forsteriana as synonyms".[10] Heimerl’s 1937 work also included this change but featured Ceodes umbellifera as the only Ceodes taxon.[10]

In 1964, Stemmerik proposed a change which reincorporated Ceodes into Pisonia.[14] Several morphological traits were omitted from this classification, such as "the absence of bracts and bracteoles at the upper portion of the pedicels, presence of starch in the embryo and number of apertures in pollen grains".[10] Stemmerik suggested that since not all of these were constant traits then they should not be regarded in the classification.[10]

In 2019, a study by Rossetto and colleagues used the molecular data of the species' DNA to reveal that the Pisonia genus was non-monophyletic.[14] It was suggested to resurrect the genera Rockia and Ceodes in replacement of Pisonia.[10] This rendered the genera monophyletic. In 2020, it was generally accepted that Ceodes should be restored, including Ceodes umbellifera.[10] This placement is accepted by Plants of the World Online, as of May 2021.[18]

Distribution and habitat

Distribution

Ceodes umbellifera may be found in a diverse range of climates worldwide. The plant is native to Australia, Southern China, Thailand, Vietnam, Taiwan, Malaysia, Philippines, Indonesia, Hawaii and many Polynesian islands.[7][19] It is also naturally occurring in some parts of Africa and Madagascar.[7]

In Australia, the species is found along the east coast in the states of New South Wales and Queensland, where it is 'widespread, but not common'.[6] In NSW, it is found along the North Coast and Central Coast regions. It mostly grows in rainforests north of the Shoalhaven region.[6]

In Hawaii, Ceodes umbellifera is known as 'pāpala' or 'pāpala kēpau'.[5] It is particularly endemic in the areas of Kauaʻi, Oʻahu, Lānaʻi and Maui, but has also be found on mainland Hawaii and Molokaʻi.[5] In Hawaii, the species will be found in areas receiving "50-100 inches" of rainfall annually and in regions between "150-1999" feet above sea level.[5]

Habitat

Generally, its ideal growing conditions are between about 16 and 20 degrees Celsius.[20] It may only be grown in frost-free environments and gardens, and similarly will not tolerate sustained heat.[20] However, some rare accounts describe the tree surviving to as cold as 26 degrees Fahrenheit.[19]

The plant requires high humidity and moderate moisture, although "established plants are drought tolerant".[6] Most often, reliably moist soil with high potassium is needed, but the species will survive dry soil with occasional watering.[7]

Moderate amounts of sunlight are needed, with most growing instructions designating part shade or full sun. When grown as an indoor plant, gardeners suggest "four hours of direct sunlight ... bright light the rest of the time".[20]

In the wild, Ceodes umbellifera is often found in rainforests, on islands and along coastlines and in shady, moist gullies.[21] Therefore, it is largely found in countries with tropical regions.[21]

Ecology

The fruit of Ceodes umbellifera exude a sticky sap, which often sticks to wildlife passing by the tree. Small animals, such as birds, reptiles and insects are often trapped in the substance and starve to death.[7][8][19]

Largely it is seabirds, dwelling on off-shore islands, that die in the fruits of the shrub.[9] However, there are some accounts of mainland birds becoming stuck. Scientists in the Goldsborough Valley, south of Cairns in Queensland, Australia, discovered a female Buff-breasted Paradise Kingfisher who was "completely immobilised, unable even to flap her wings".[22]

It is generally agreed that the tree's birdlime is an adaptation, enabling the tree to pollinate; pollen will adhere to a bird, and be deposited somewhere else, where a new plant may grow.[5] However, some botanists suggest that the extreme stickiness may have evolutionary advantages outside of pollination.[22] Since Ceodes umbellifera is widespread on tropical island rainforests, it is plausible that when the plant "traps and kills animals" it is "creating pockets of nutrients in the immediate soil environment for seedlings".[22] This is especially important on islands because often their soil is not as fertile and nutrient rich.

Gould's Petrels

The tree presents a threat to the Gould's Petrel population on Cabbage Tree Island, since the birds are classified as a threatened species. Petrels' entanglement in the viscous fruit sap has been identified as a major cause of mortality by Australian ornithologists.[9] In some cases, a single fruit was enough to trap a bird and cause its death. During 1992 and 1993, some of the Ceodes umbellifera plants on Cabbage Tree Island were subject to herbicide poisoning, in an attempt to reduce the threat to Gould's Petrels and improve their endangerment status.[9] This was largely successful, although other natural threats to the petrel remain.

Uses 

Traditional medicine

In Hawaii, the tree was used for a variety of purposes by early indigenous people. Leaves of Ceodes umbellifera is used as a general purpose medicine for digestive issues and childhood disease.[5]

In their paper on saponins found in the shrub's leaves, Lavaud and Beauvir state that "In Vanuatu, the leaves of P. umbellifera are also used as a treatment for ciguatera poisoning and as a 'green manure'.[23]

Culture

In Hawaii, the tree's leaves and bark are also used as a stable source of food for pigs.[23]

The adhesive sap from the fruit is also commonly used by indigenous Hawaiians. The sap may be made into a paste with okra and used to mend broken bowls.[24] Additionally, it is used by indigenous men to catch birds, from which the feathers are plucked and used for traditional garments.[24]

See also

References

  1. ^ "Ceodes umbellifera J.R.Forst. & G.Forst". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 26 May 2021.
  2. ^ "Pisonia umbellifera". Australian Plant Name Index (APNI), IBIS database. Centre for Plant Biodiversity Research, Australian Government. Retrieved 28 April 2016.
  3. ^ "Pisonia umbellifera". Flora of China. Retrieved 14 April 2009.
  4. ^ a b Jacobs, S.W.L.; G. J. Harden. "Pisonia umbellifera". PlantNET - New South Wales Flora Online. Royal Botanic Gardens & Domain Trust, Sydney Australia. Retrieved 14 April 2009.
  5. ^ a b c d e f g h i "Native Plants Hawaii". nativeplants.hawaii.edu. Retrieved 13 April 2021.
  6. ^ a b c d e f g h National Herbarium of New South Wales (2004). "Pisonia umbellifera (J.R.Forst. & G.Forst.) Seem". PlantNET FloraOnline. Retrieved 15 April 2021.{{cite web}}: CS1 maint: url-status (link)
  7. ^ a b c d e f Fern, K (2019). "Pisonia umbellifera". Useful Tropical Plants. Retrieved 15 April 2021.{{cite web}}: CS1 maint: url-status (link)
  8. ^ a b c Pucci, P. & Beltramini, M. (2003). [21 April 2021 "Pisonia umbellifera"]. Monaco Nature Encyclopedia. {{cite web}}: Check |url= value (help)CS1 maint: multiple names: authors list (link) CS1 maint: url-status (link)
  9. ^ a b c d Priddel, D. & Carlile, N. (1995). "Mortality of Adult Gould's Petrels Pterodroma leucoptera leucoptera at the Nesting Site on Cabbage Tree Island, New South Wales". Emu - Austral Ornithology. 95 (4): 259–264. doi:10.1071/MU9950259.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  10. ^ a b c d e f g h i j k l m n o p q r s t u Rossetto, E. F. & Caraballo, M. (2019). "Splitting the Pisonia birdcatcher trees: re-establishment of Ceodes and Rockia (Nyctaginaceae, Pisonieae)". PhytoKeys. 152(3): 121–136.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ a b Forster, J. R. & Forster, G. (1776). Characteres Generum Plantarum. B. White, T. Cadell, & P. Elmsly, London.{{cite book}}: CS1 maint: multiple names: authors list (link)
  12. ^ a b c Skottsberg, C. (1926). "Vascular plants from the Hawaiian Islands". Acta Horti Gothoburgensis. 2: 185–284.
  13. ^ Bentham, G. & Hooker, J. D. (1880). Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita. Reeves & Co., London.{{cite book}}: CS1 maint: multiple names: authors list (link)
  14. ^ a b c Rossetto, E. F., Faria, A. D., Ruas, P. M., Ruas, C., Douglas, N. A.,& Ribeiro, J. (2019). "Clarifying generic delimitation in Nyctaginaceae tribe Pisonieae after more than a century of taxonomic confusion". Botanical Journal of the Linnean Society. 189(4): 378–396.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  15. ^ Seemann, B. (1863). "On the genus Ceodes of Forster". Journal of Botany, British and Foreign. 1: 244–246.
  16. ^ Heimerl, A. (1913). "Die Nyctaginaceen-Gattungen Calpidia und Rockia". Österreichische Botanische Zeitschrift. 63(7): 279–290.
  17. ^ Skottsberg, C. (1936). "The arboreous Nyctaginaceae of Hawaii". Svensk Botanisk Tidskrift. 30: 722–743.
  18. ^ "Ceodes umbellifera J.R.Forst. & G.Forst. | Plants of the World Online | Kew Science". Plants of the World Online. Retrieved 29 May 2021.
  19. ^ a b c "Pisonia umbellifera 'Variegata', Map Plant, Shrub, [Heimerliriodendron]". www.smgrowers.com. Retrieved 17 May 2021.
  20. ^ a b c "Bird Catcher Tree/Pisonia umbellifera 'Variegata'". 7 August 2013. Retrieved 17 May 2021.
  21. ^ a b Australian Botanic Garden (2021). "Pisonia umbellifera". Retrieved 21 April 2021.{{cite web}}: CS1 maint: url-status (link)
  22. ^ a b c Murphy, S. & Legge, S. (2003). "A first record of a land-bird being trapped by Pisonia umbellifera". Sunbird. 33 (3): 111–112.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  23. ^ a b Lavaud, D., Beauvir, S., Massiot, G., Le Men-Olivier, L, & Bourdy, G. (1996). "Saponins from Pisonia umbellifera". Saponins from Pisonia Umbellifera. 43 (1): 189–194.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  24. ^ a b Buck, P.H. (1964). Arts and crafts of Hawaii. Bishop Museum Press. pp. 45, 217–218.