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Cephalotus

Cephalotus (/ˌsɛfəˈltəs/ or /ˌkɛfəˈltəs/; Greek: κεφαλή "head", and οὔς/ὠτός "ear", to describe the head of the anthers)[3] is a genus which contains one species, Cephalotus follicularis the Albany pitcher plant,[4] a small carnivorous pitcher plant. The pit-fall traps of the modified leaves have inspired the common names for this plant, which include 'Albany pitcher plant", "Western Australian pitcher plant", "Australian pitcher plant", or "fly-catcher plant." It is an evergreen herb that is endemic to peaty swamps in the southwestern corner of Western Australia.[5]

Cephalotus
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Oxalidales
Family: Cephalotaceae
Dumort.[2]
Genus: Cephalotus
Labill.
Species:
C. follicularis
Binomial name
Cephalotus follicularis
Labill.
Global range

Description edit

Cephalotus follicularis is a small, low growing, herbaceous species. Evergreen leaves appear from underground rhizomes, are simple with an entire leaf blade, and lie close to the ground. The insectivorous leaves are small and have the appearance of moccasins, forming the 'pitcher' of the common name. The pitchers develop a dark red colour in high light levels but stay green in shadier conditions. The foliage is a basal arrangement that is closely arranged with outward facing adapted leaf blades. These leaves give the main form of the species a height around 20 cm.

The 'pitcher' trap of the species is similar to other pitcher plants. The peristome at the entrance of the trap has a spiked arrangement that allows the prey to enter, but hinders its escape. The lid over the entrance, the operculum, prevents rainwater entering the pitcher and thus diluting the digestive enzymes inside. Insects trapped in this digestive fluid are consumed by the plant. The operculum has translucent cells which confuse its insect prey as they appear to be patches of sky.

The inflorescence is groupings of small, hermaphroditic, six-parted, regular flowers, which are creamy, or whitish.

In the cooler months of winter (down to about 5 degrees Celsius), they have a natural dormancy period of about 3–4 months, triggered by the temperature drop and reduced light levels.

Taxonomy edit

Taxonomic history edit

Botanical specimens were first collected during the visit of HMS Investigator to King George Sound in December 1801 and January 1802. On 2 January 1802 the expedition's botanist, Robert Brown, wrote in his diary:

"Remaind on board. Described a few plants. Mr Good went in search of the pitcher plant wch Messrs Bauer & Westall had found yesterday in flower. He returned with it in the evening."[6]

 
Plate 3119, Curtis's Botanical Magazine, details by Ferdinand Bauer

This represents the earliest documentary reference to this species; and although not entirely unambiguous as to the first collection, it is usually taken as evidence that the plant was discovered by Ferdinand Bauer and William Westall on 1 January 1802. Whether or not there was an earlier collection is largely immaterial, however, as all collections were incorporated into Brown's collection without attribution, so Brown is treated as the collector in botanical contexts.

Brown initially gave this species the manuscript name "'Cantharifera paludosa' KG III Sound",[7] but this name was not published, and it would not be Brown who published the first description.

The following year, further specimens were collected by Jean Baptiste Leschenault de la Tour, botanist to Nicolas Baudin's expedition. In 1806, Jacques Labillardière used these specimens as the basis of his publication of the species in Novae Hollandiae plantarum specimen.[8] Labillardière did not attribute Leschenault as the collector, and it was long thought that Labillardière had collected the plant himself during his visit to the area in the 1790s; in particular, Brown wrongly acknowledged Labillardière as the discoverer of this plant.

Leschenault's specimen was a fruiting plant, but the fruit was in poor condition, and as a result Labillardière erroneously placed it in the family Rosaceae. This error was not corrected until better fruiting specimens were collected by William Baxter in the 1820s. These were examined by Brown, who concluded that the plant merited its own family, and accordingly erected Cephalotaceae. It has remained in this monogeneric family ever since.[9]

Current placement edit

The Australian pitcher plant is an advanced rosid, and thus closer related to apples and oaks than to other pitcher plants like Nepenthaceae (basal core eudicots) and Sarraceniaceae (basal asterids). The placement of its monotypic family Cephalotaceae in the order Saxifragales has been abandoned. It is now placed within the order of Oxalidales where it is most closely related to Brunelliaceae, Cunoniaceae, and Elaeocarpaceae.[10] The monotypic arrangement of the family and genus is indicative of a high degree of endemism, one of four such species of the region.

 
Cephalotus follicularis in typical habitat in coastal SW-Australia

Ecology edit

The plant occurs in southern coastal districts of the Southwest botanical province in Australia; it has been recorded in the Warren, southern Jarrah Forest, and Esperance Plains regions. Its habitat is on moist peaty sands found in swamps or along creeks and streams, but it is tolerant of less damp situations. Its population in the wild has been reduced by habitat destruction and overcollecting; it is therefore classified as vulnerable species (VU A2ac; C2a(i)) by the IUCN.[1] However, this classification is not in unison with Australia's national EPBC Act List of Threatened Species[11] or Western Australia's Wildlife Conservation Act 1950, which both list the species as Not Threatened.[4]

The larvae of Badisis ambulans, an ant-like wingless micropezid fly, develop inside the pitchers. They have never been found anywhere else.[12]

Cultivation edit

 
Cephalotus follicularis: a young plant of about 2–3 years, grown in cultivation

Cephalotus are cultivated worldwide. In the wild, they prefer warm day-time temperatures of up to 25 degrees Celsius during the growing season, coupled with cool night-time temperatures. It is commonly grown in a mixture of sphagnum peat moss, perlite, and sand, a reasonable humidity (60–80%) is also preferred. It is successfully propagated from root and leaf cuttings, usually non-carnivorous leaves although pitchers can also be used. A dormancy period is probably crucial to long-term health of the plant.

The plants become colourful and grow vigorously when kept in direct sunlight, while plants cultivated in bright shade remain green.

Living plants were delivered to Kew Gardens by Phillip Parker King in 1823. A specimen flowered in 1827 and provided one source for an illustration in Curtis's Botanical Magazine.[3]

This plant is a recipient of the Royal Horticultural Society's Award of Garden Merit.[13]

 
Inflorescence of Cephalotus follicularis in cultivation.

There are several dozen Cephalotus clones that exist in cultivation; nine have been officially registered as cultivars. One of the most well-known is 'Eden Black', a cultivar with unusually dark-coloured pitchers.[14]

Genomics edit

The genome of the pitcher plant Cephalotus follicularis has been sequenced.[15][16] Its carnivorous and non-carnivorous leaves have been compared to identify genetic differences associated with key features relating to the attraction of prey and their capture, digestion and nutrient absorption. Results support the independent convergent evolution of Cephalotus and other carnivorous plant lineages. but also suggest that different lineages co-opted similar genes in developing digestive functions. This implies that the ways in which carnivory can be developed are limited.[17][18][16]

See also edit

References edit

  1. ^ a b Bourke, G.; Cross, A.; Nunn, R.; Kalfas, N. (17 January 2020). "Cephalotus follicularis". doi:10.2305/iucn.uk.2020-1.rlts.t39635a19631881.en. S2CID 243022719. {{cite journal}}: Cite journal requires |journal= (help)
  2. ^ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x.
  3. ^ a b Hooker, William Jackson (1831). "Cephalotus follicularis. Follicled Cephalotus". Curtis's Botanical Magazine. Samuel Curtis. 58: Pl. 3118 & 3119.
  4. ^ a b Attractions, Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and. "FloraBase—the Western Australian Flora". florabase.dpaw.wa.gov.au. Retrieved 2018-08-29.{{cite web}}: CS1 maint: multiple names: authors list (link)
  5. ^ Heywood, V.H.; Brummitt, R.K.; Culham, A.; Seberg, O. (2007). Flowering Plant Families of the World. Ontario, Canada: Firefly Books. p. 94. ISBN 978-1-55407-206-4.
  6. ^ Vallance, T. G.; Moore, D. T.; Groves, E. W. (2001). Nature's Investigator: The Diary of Robert Brown in Australia, 1801–1805. Canberra: Australian Biological Resources Study. ISBN 0-642-56817-0.
  7. ^ "Cephalotus follicularis Labill". Robert Brown's Australian Botanical Specimens, 1801–1805 at the BM. Retrieved 2009-01-10.
  8. ^ Hopper, Stephen (2003). "South-western Australian, Cinderella of the world's temperate floristic regions 1". Curtis's Botanical Magazine. 20 (2): 101–126. doi:10.1111/1467-8748.00379.
  9. ^ Mabberley, D. J. (1985). "Chapter IX: The natural system". Jupiter Botanicus: Robert Brown of the British Museum. Braunschweig: J. Cramer. pp. 141–176. ISBN 3-7682-1408-7.
  10. ^ Angiosperm Phylogeny Website, retrieved 9 January 2016
  11. ^ "Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act)". Australian Government. Retrieved 14 March 2022.
  12. ^ McAlpine, David K. (1998). "Review of the Australian stilt flies (Diptera : Micropezidae) with a phylogenetic analysis of the family". Invertebrate Systematics. 12 (1): 55. doi:10.1071/IT96018.
  13. ^ "Cephalotus follicularis". www.rhs.org. Royal Horticultural Society. Retrieved 12 April 2020.
  14. ^ "The Carnivorous Plant FAQ: Cephalotus: various types". www.sarracenia.com. Retrieved 2020-06-02.
  15. ^ "Evolution of the Oxalidales Carnivore". International Carnivorous Plant Society (ICPS). Retrieved 14 March 2022.
  16. ^ a b Fukushima, Kenji; Fang, Xiaodong; Alvarez-Ponce, David; Cai, Huimin; Carretero-Paulet, Lorenzo; Chen, Cui; Chang, Tien-Hao; Farr, Kimberly M.; Fujita, Tomomichi; Hiwatashi, Yuji; Hoshi, Yoshikazu; Imai, Takamasa; Kasahara, Masahiro; Librado, Pablo; Mao, Likai; Mori, Hitoshi; Nishiyama, Tomoaki; Nozawa, Masafumi; Pálfalvi, Gergő; Pollard, Stephen T.; Rozas, Julio; Sánchez-Gracia, Alejandro; Sankoff, David; Shibata, Tomoko F.; Shigenobu, Shuji; Sumikawa, Naomi; Uzawa, Taketoshi; Xie, Meiying; Zheng, Chunfang; Pollock, David D.; Albert, Victor A.; Li, Shuaicheng; Hasebe, Mitsuyasu (6 February 2017). "Genome of the pitcher plant Cephalotus reveals genetic changes associated with carnivory". Nature Ecology & Evolution. 1 (3): 59. doi:10.1038/s41559-016-0059. ISSN 2397-334X. PMID 28812732. S2CID 6223440.
  17. ^ Pain, Stephanie (2 March 2022). "How plants turned predator". Knowable Magazine. doi:10.1146/knowable-030122-1. Retrieved 11 March 2022.
  18. ^ Hedrich, Rainer; Fukushima, Kenji (17 June 2021). "On the Origin of Carnivory: Molecular Physiology and Evolution of Plants on an Animal Diet". Annual Review of Plant Biology. 72 (1): 133–153. doi:10.1146/annurev-arplant-080620-010429. ISSN 1543-5008. PMID 33434053. S2CID 231595236. Retrieved 11 March 2022.

External links edit

cephalotus, been, suggested, that, australian, pitcher, plant, merged, into, this, article, discuss, proposed, since, november, 2023, confused, with, cephalotes, greek, κεφαλή, head, οὔς, ὠτός, describe, head, anthers, genus, which, contains, species, follicul. It has been suggested that Australian pitcher plant be merged into this article Discuss Proposed since November 2023 Not to be confused with Cephalotes Cephalotus ˌ s ɛ f e ˈ l oʊ t e s or ˌ k ɛ f e ˈ l oʊ t e s Greek kefalh head and oὔs ὠtos ear to describe the head of the anthers 3 is a genus which contains one species Cephalotus follicularis the Albany pitcher plant 4 a small carnivorous pitcher plant The pit fall traps of the modified leaves have inspired the common names for this plant which include Albany pitcher plant Western Australian pitcher plant Australian pitcher plant or fly catcher plant It is an evergreen herb that is endemic to peaty swamps in the southwestern corner of Western Australia 5 CephalotusConservation statusVulnerable IUCN 3 1 1 Scientific classificationKingdom PlantaeClade TracheophytesClade AngiospermsClade EudicotsClade RosidsOrder OxalidalesFamily CephalotaceaeDumort 2 Genus CephalotusLabill Species C follicularisBinomial nameCephalotus follicularisLabill Global range Contents 1 Description 2 Taxonomy 2 1 Taxonomic history 2 2 Current placement 3 Ecology 4 Cultivation 5 Genomics 6 See also 7 References 8 External linksDescription editCephalotus follicularis is a small low growing herbaceous species Evergreen leaves appear from underground rhizomes are simple with an entire leaf blade and lie close to the ground The insectivorous leaves are small and have the appearance of moccasins forming the pitcher of the common name The pitchers develop a dark red colour in high light levels but stay green in shadier conditions The foliage is a basal arrangement that is closely arranged with outward facing adapted leaf blades These leaves give the main form of the species a height around 20 cm The pitcher trap of the species is similar to other pitcher plants The peristome at the entrance of the trap has a spiked arrangement that allows the prey to enter but hinders its escape The lid over the entrance the operculum prevents rainwater entering the pitcher and thus diluting the digestive enzymes inside Insects trapped in this digestive fluid are consumed by the plant The operculum has translucent cells which confuse its insect prey as they appear to be patches of sky The inflorescence is groupings of small hermaphroditic six parted regular flowers which are creamy or whitish In the cooler months of winter down to about 5 degrees Celsius they have a natural dormancy period of about 3 4 months triggered by the temperature drop and reduced light levels Taxonomy editTaxonomic history edit Botanical specimens were first collected during the visit of HMS Investigator to King George Sound in December 1801 and January 1802 On 2 January 1802 the expedition s botanist Robert Brown wrote in his diary Remaind on board Described a few plants Mr Good went in search of the pitcher plant wch Messrs Bauer amp Westall had found yesterday in flower He returned with it in the evening 6 nbsp Plate 3119 Curtis s Botanical Magazine details by Ferdinand BauerThis represents the earliest documentary reference to this species and although not entirely unambiguous as to the first collection it is usually taken as evidence that the plant was discovered by Ferdinand Bauer and William Westall on 1 January 1802 Whether or not there was an earlier collection is largely immaterial however as all collections were incorporated into Brown s collection without attribution so Brown is treated as the collector in botanical contexts Brown initially gave this species the manuscript name Cantharifera paludosa KG III Sound 7 but this name was not published and it would not be Brown who published the first description The following year further specimens were collected by Jean Baptiste Leschenault de la Tour botanist to Nicolas Baudin s expedition In 1806 Jacques Labillardiere used these specimens as the basis of his publication of the species in Novae Hollandiae plantarum specimen 8 Labillardiere did not attribute Leschenault as the collector and it was long thought that Labillardiere had collected the plant himself during his visit to the area in the 1790s in particular Brown wrongly acknowledged Labillardiere as the discoverer of this plant Leschenault s specimen was a fruiting plant but the fruit was in poor condition and as a result Labillardiere erroneously placed it in the family Rosaceae This error was not corrected until better fruiting specimens were collected by William Baxter in the 1820s These were examined by Brown who concluded that the plant merited its own family and accordingly erected Cephalotaceae It has remained in this monogeneric family ever since 9 Current placement edit The Australian pitcher plant is an advanced rosid and thus closer related to apples and oaks than to other pitcher plants like Nepenthaceae basal core eudicots and Sarraceniaceae basal asterids The placement of its monotypic family Cephalotaceae in the order Saxifragales has been abandoned It is now placed within the order of Oxalidales where it is most closely related to Brunelliaceae Cunoniaceae and Elaeocarpaceae 10 The monotypic arrangement of the family and genus is indicative of a high degree of endemism one of four such species of the region nbsp Cephalotus follicularis in typical habitat in coastal SW AustraliaEcology editThe plant occurs in southern coastal districts of the Southwest botanical province in Australia it has been recorded in the Warren southern Jarrah Forest and Esperance Plains regions Its habitat is on moist peaty sands found in swamps or along creeks and streams but it is tolerant of less damp situations Its population in the wild has been reduced by habitat destruction and overcollecting it is therefore classified as vulnerable species VU A2ac C2a i by the IUCN 1 However this classification is not in unison with Australia s national EPBC Act List of Threatened Species 11 or Western Australia s Wildlife Conservation Act 1950 which both list the species as Not Threatened 4 The larvae of Badisis ambulans an ant like wingless micropezid fly develop inside the pitchers They have never been found anywhere else 12 Cultivation edit nbsp Cephalotus follicularis a young plant of about 2 3 years grown in cultivationCephalotus are cultivated worldwide In the wild they prefer warm day time temperatures of up to 25 degrees Celsius during the growing season coupled with cool night time temperatures It is commonly grown in a mixture of sphagnum peat moss perlite and sand a reasonable humidity 60 80 is also preferred It is successfully propagated from root and leaf cuttings usually non carnivorous leaves although pitchers can also be used A dormancy period is probably crucial to long term health of the plant The plants become colourful and grow vigorously when kept in direct sunlight while plants cultivated in bright shade remain green Living plants were delivered to Kew Gardens by Phillip Parker King in 1823 A specimen flowered in 1827 and provided one source for an illustration in Curtis s Botanical Magazine 3 This plant is a recipient of the Royal Horticultural Society s Award of Garden Merit 13 nbsp Inflorescence of Cephalotus follicularis in cultivation There are several dozen Cephalotus clones that exist in cultivation nine have been officially registered as cultivars One of the most well known is Eden Black a cultivar with unusually dark coloured pitchers 14 Genomics editThe genome of the pitcher plant Cephalotus follicularis has been sequenced 15 16 Its carnivorous and non carnivorous leaves have been compared to identify genetic differences associated with key features relating to the attraction of prey and their capture digestion and nutrient absorption Results support the independent convergent evolution of Cephalotus and other carnivorous plant lineages but also suggest that different lineages co opted similar genes in developing digestive functions This implies that the ways in which carnivory can be developed are limited 17 18 16 See also editCarnivorous plants of AustraliaReferences edit a b Bourke G Cross A Nunn R Kalfas N 17 January 2020 Cephalotus follicularis doi 10 2305 iucn uk 2020 1 rlts t39635a19631881 en S2CID 243022719 a href Template Cite journal html title Template Cite journal cite journal a Cite journal requires journal help Angiosperm Phylogeny Group 2009 An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants APG III Botanical Journal of the Linnean Society 161 2 105 121 doi 10 1111 j 1095 8339 2009 00996 x a b Hooker William Jackson 1831 Cephalotus follicularis Follicled Cephalotus Curtis s Botanical Magazine Samuel Curtis 58 Pl 3118 amp 3119 a b Attractions Western Australian Herbarium Biodiversity and Conservation Science Department of Biodiversity Conservation and FloraBase the Western Australian Flora florabase dpaw wa gov au Retrieved 2018 08 29 a href Template Cite web html title Template Cite web cite web a CS1 maint multiple names authors list link Heywood V H Brummitt R K Culham A Seberg O 2007 Flowering Plant Families of the World Ontario Canada Firefly Books p 94 ISBN 978 1 55407 206 4 Vallance T G Moore D T Groves E W 2001 Nature s Investigator The Diary of Robert Brown in Australia 1801 1805 Canberra Australian Biological Resources Study ISBN 0 642 56817 0 Cephalotus follicularis Labill Robert Brown s Australian Botanical Specimens 1801 1805 at the BM Retrieved 2009 01 10 Hopper Stephen 2003 South western Australian Cinderella of the world s temperate floristic regions 1 Curtis s Botanical Magazine 20 2 101 126 doi 10 1111 1467 8748 00379 Mabberley D J 1985 Chapter IX The natural system Jupiter Botanicus Robert Brown of the British Museum Braunschweig J Cramer pp 141 176 ISBN 3 7682 1408 7 Angiosperm Phylogeny Website retrieved 9 January 2016 Environment Protection and Biodiversity Conservation Act 1999 EPBC Act Australian Government Retrieved 14 March 2022 McAlpine David K 1998 Review of the Australian stilt flies Diptera Micropezidae with a phylogenetic analysis of the family Invertebrate Systematics 12 1 55 doi 10 1071 IT96018 Cephalotus follicularis www rhs org Royal Horticultural Society Retrieved 12 April 2020 The Carnivorous Plant FAQ Cephalotus various types www sarracenia com Retrieved 2020 06 02 Evolution of the Oxalidales Carnivore International Carnivorous Plant Society ICPS Retrieved 14 March 2022 a b Fukushima Kenji Fang Xiaodong Alvarez Ponce David Cai Huimin Carretero Paulet Lorenzo Chen Cui Chang Tien Hao Farr Kimberly M Fujita Tomomichi Hiwatashi Yuji Hoshi Yoshikazu Imai Takamasa Kasahara Masahiro Librado Pablo Mao Likai Mori Hitoshi Nishiyama Tomoaki Nozawa Masafumi Palfalvi Gergo Pollard Stephen T Rozas Julio Sanchez Gracia Alejandro Sankoff David Shibata Tomoko F Shigenobu Shuji Sumikawa Naomi Uzawa Taketoshi Xie Meiying Zheng Chunfang Pollock David D Albert Victor A Li Shuaicheng Hasebe Mitsuyasu 6 February 2017 Genome of the pitcher plant Cephalotus reveals genetic changes associated with carnivory Nature Ecology amp Evolution 1 3 59 doi 10 1038 s41559 016 0059 ISSN 2397 334X PMID 28812732 S2CID 6223440 Pain Stephanie 2 March 2022 How plants turned predator Knowable Magazine doi 10 1146 knowable 030122 1 Retrieved 11 March 2022 Hedrich Rainer Fukushima Kenji 17 June 2021 On the Origin of Carnivory Molecular Physiology and Evolution of Plants on an Animal Diet Annual Review of Plant Biology 72 1 133 153 doi 10 1146 annurev arplant 080620 010429 ISSN 1543 5008 PMID 33434053 S2CID 231595236 Retrieved 11 March 2022 External links edit nbsp Wikimedia Commons has media related to Cephalotus follicularis nbsp Wikispecies has information related to Cephalotus nbsp Wikisource has original text related to this article Curtis s Botanical Magazine 58 Pl 3118 amp 3119 Cephalotus follicularis nbsp Wikisource has original text related to this article Cephalotus follicularis Cephalotus FloraBase Western Australian Government Department of Biodiversity Conservation and Attractions Cephalotus follicularis FloraBase Western Australian Government Department of Biodiversity Conservation and Attractions Inner World of Cephalotus follicularis from the John Innes Centre Archived 2014 03 16 at the Wayback Machine Cephalotus Labill Atlas of Living Australia Retrieved from https en wikipedia org w index php title Cephalotus amp oldid 1189343889, wikipedia, wiki, book, books, library,

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