Cephalotus

Life-forms

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Rosids
Cladus: Eurosids I
Ordo: Oxalidales

Familia: Cephalotaceae
Genus: Cephalotus
Name

Cephalotaceae Dumort., Anal. Fam. Pl. 59, 61. 1829.
References
Primary references

Dumortier, B.-C. 1829. Analyse des Familles de Plantes: avec l'indication des principaux genres qui s'y rattachent. 104 pp., Tournay, J. Casterman. BHL Reference page. : 59, 61.

Links

Govaerts, R. et al. 2019. Cephalotaceae in World Checklist of Selected Plant Families. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2019 October 07. Reference page.
International Plant Names Index. 2019. Cephalotaceae. Published online. Accessed: October 07 2019.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page.
FloraBase Cephalotaceae

Vernacular names
Deutsch: Zwergkrug
English: Australian Pitcher-plant Family
中文: 土瓶草科

Cephalotus (/ˌsɛfəˈloʊtəs/ or /ˌkɛfəˈloʊtəs/; 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]

Description

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
Taxonomic history

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

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

The plant occurs in southern coastal districts of the Southwest botanical province in Australia; recorded in the Warren, southern Jarrah Forest, and the Esperance Plains. 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
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

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

Carnivorous plants of Australia

References

Bourke, G.; Cross, A.; Nunn, R.; Kalfas, N. (17 January 2020). "Cephalotus follicularis". IUCN Red List of Threatened Species. doi:10.2305/iucn.uk.2020-1.rlts.t39635a19631881.en. Retrieved 14 March 2022.
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.
Hooker, William Jackson (1831). "Cephalotus follicularis. Follicled Cephalotus". Curtis's Botanical Magazine. Samuel Curtis. 58: Pl. 3118 & 3119.
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.
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.
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): 1–9. doi:10.1038/s41559-016-0059. ISSN 2397-334X. Retrieved 14 March 2022.
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. Retrieved 11 March 2022.