Cephalotaceae
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''Cephalotus'' ( or ; Greek: ''κεφαλή'' "head", and ''οὔς''/''ὠτός'' "ear", to describe the head of the anthers) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
which contains one
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
, ''Cephalotus follicularis'' the Albany pitcher plant, a small
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other ...
pitcher plant. The pit-fall traps of the modified leaves have inspired the
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
s 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.


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 A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
are small and have the appearance of
moccasins A moccasin is a shoe, made of deerskin or other soft leather, consisting of a sole (made with leather that has not been "worked") and sides made of one piece of leather, stitched together at the top, and sometimes with a vamp (additional panel o ...
, 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 Peristome (from the Greek ''peri'', meaning 'around' or 'about', and ''stoma'', 'mouth') is an anatomical feature that surrounds an opening to an organ or structure. Some plants, fungi, and shelled gastropods have peristomes. In mosses In mosse ...
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
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s inside. Insects trapped in this digestive fluid are consumed by the plant. The operculum has
translucent In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable scattering of light. On a macroscopic scale (one in which the dimensions ...
cells which confuse its
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
prey as they appear to be patches of sky. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are forme ...
is groupings of small, hermaphroditic, six-parted, regular
flowers A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
, 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 to
King George Sound King George Sound ( nys , Menang Koort) is a sound on the south coast of Western Australia. Named King George the Third's Sound in 1791, it was referred to as King George's Sound from 1805. The name "King George Sound" gradually came into use ...
in December 1801 and January 1802. On 2 January 1802 the expedition's botanist, Robert Brown, wrote in his diary: 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 Ferdinand Lucas Bauer (20 January 1760 – 17 March 1826) was an Austrian botanical illustrator who travelled on Matthew Flinders' expedition to Australia. Biography Early life and career Bauer was born in Feldsberg in 1760, the youngest son ...
and
William Westall William Westall (12 October 1781 – 22 January 1850) was a British landscape artist best known as one of the first artists to work in Australia. Early life Westall was born in Hertford and grew up in London, mostly Sydenham and Hampstead. ...
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", 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 Jean-Baptiste Louis Claude Théodore Leschenault de La Tour (13 November 1773 – 14 March 1826) was a French botanist and ornithologist. Born at the family seat (since 1718), Le Villard, near Chalon-sur-Saône, Leschenault de la Tour arrived in ...
, botanist to
Nicolas Baudin Nicolas Thomas Baudin (; 17 February 1754 – 16 September 1803) was a French explorer, cartographer, naturalist and hydrographer, most notable for his explorations in Australia and the southern Pacific. Biography Early career Born a comm ...
's expedition. In 1806,
Jacques Labillardière Jacques-Julien Houtou de Labillardière (28 October 1755 – 8 January 1834) was a French biologist noted for his descriptions of the flora of Australia. Labillardière was a member of a voyage in search of the La Pérouse expedition. He pub ...
used these specimens as the basis of his publication of the species in ''
Novae Hollandiae plantarum specimen ''Novae Hollandiae Plantarum Specimen'' is a two-volume work describing the flora of Australia. Facsimiles of the originals can be found in the onlinBiodiversity Heritage Library (Vol.1)anVol 2) The author was the French botanist Jacques Labillar ...
''. 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 Rosaceae (), the rose family, is a medium-sized family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus ''Rosa''. Among the most species-rich genera are '' Alchemilla'' (270), ''Sorb ...
. 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.


Current placement

The Australian pitcher plant is an advanced
rosid The rosids are members of a large clade ( monophyletic group) of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms. The clade is divided into 16 to 20 orders, depending upon circumscription and classifica ...
, and thus closer related to
apple An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple trees are cultivated worldwide and are the most widely grown species in the genus '' Malus''. The tree originated in Central Asia, where its wild ancest ...
s and oaks than to other pitcher plants like Nepenthaceae ( basal
core eudicots The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dicots ...
) and
Sarraceniaceae Sarraceniaceae are a family of pitcher plants, belonging to order Ericales (previously Nepenthales). The family comprises three extant genera: ''Sarracenia'' (North American pitcher plants), '' Darlingtonia'' (the cobra lily or California pitc ...
(basal
asterid In the APG IV system (2016) for the classification of flowering plants, the name asterids denotes a clade (a monophyletic group). Asterids is the largest group of flowering plants, with more than 80,000 species, about a third of the total flo ...
s). The placement of its
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Cephalotaceae in the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Saxifragales The Saxifragales (saxifrages) are an order of flowering plants (Angiosperms). They are an extremely diverse group of plants which include trees, shrubs, perennial herbs, succulent and aquatic plants. The degree of diversity in terms of vegeta ...
has been abandoned. It is now placed within the order of
Oxalidales Oxalidales is an order of flowering plants, included within the rosid subgroup of eudicots. Compound leaves are common in Oxalidales and the majority of the species in this order have five or six sepals and petals. The following families are typic ...
where it is most closely related to Brunelliaceae,
Cunoniaceae Cunoniaceae is a family of 27 genera and about 335 species of woody plants in the order Oxalidales, mostly found in the tropical and wet temperate regions of the Southern Hemisphere. The greatest diversity of genera are in Australia and Tasmania ...
, and
Elaeocarpaceae Elaeaocarpaceae is a family of flowering plants. The family contains approximately 615 species of trees and shrubs in 12 genera."Elaeocarpaceae" In: Klaus Kubitzki (ed.). ''The Families and Genera of Vascular Plants'' vol. VI. Springer-Verlag: Be ...
. The monotypic arrangement of the family and genus is indicative of a high degree of
endemism Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...
, one of four such species of the region.


Ecology

The plant occurs in southern coastal districts of the
Southwest botanical province Southwest Australia is a biogeographic region in Western Australia. It includes the Mediterranean-climate area of southwestern Australia, which is home to a diverse and distinctive flora and fauna. The region is also known as the Southwest Aus ...
in Australia; recorded in the Warren, southern
Jarrah Forest Jarrah forest is tall open forest in which the dominant overstory tree is ''Eucalyptus marginata'' (jarrah). The ecosystem occurs only in the Southwest Botanical Province of Western Australia. It is most common in the biogeographic region named in ...
, and the
Esperance Plains Esperance Plains, also known as Eyre Botanical District, is a biogeographic region in southern Western Australia on the south coast between the Avon Wheatbelt and Hampton bioregions, and bordered to the north by the Mallee region. It is a ...
. 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 Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
and overcollecting; it is therefore classified as
Vulnerable species A vulnerable species is a species which has been categorized by the International Union for Conservation of Nature as being threatened with extinction unless the circumstances that are threatening its survival and reproduction improve. Vulnera ...
(VU A2ac; C2a(i)) by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. However, this classification is not in unison with Australia's national EPBC Act List of Threatened Species or Western Australia's
Wildlife Conservation Act 1950 The ''Wildlife Conservation Act 1950'' is an act of the Western Australian Parliament that provides the statute relating to conservation and legal protection of flora and fauna. Text was copied from this source, which is available under Attri ...
, which both list the species as Not Threatened. The larvae of ''
Badisis ambulans ''Badisis'' is a stilt-legged fly genus with only one known species, ''Badisis ambulans''. This is a wingless, haltere-less fly with an ant-like appearance. It is only found in the Southwest Australian bioregion of Western Australia. Dependent ...
'', an ant-like wingless micropezid
fly Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
, develop inside the pitchers. They have never been found anywhere else.


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 Perlite is an amorphous volcanic glass that has a relatively high water content, typically formed by the hydration of obsidian. It occurs naturally and has the unusual property of greatly expanding when heated sufficiently. It is an industrial ...
, and
sand Sand is a granular material composed of finely divided mineral particles. Sand has various compositions but is defined by its grain size. Sand grains are smaller than gravel and coarser than silt. Sand can also refer to a textural class ...
, a reasonable
humidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity dep ...
(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 Rear Admiral Phillip Parker King, FRS, RN (13 December 1791 – 26 February 1856) was an early explorer of the Australian and Patagonian coasts. Early life and education King was born on Norfolk Island, to Philip Gidley King and Anna ...
in 1823. A specimen flowered in 1827 and provided one source for an illustration in ''
Curtis's Botanical Magazine ''The Botanical Magazine; or Flower-Garden Displayed'', is an illustrated publication which began in 1787. The longest running botanical magazine, it is widely referred to by the subsequent name ''Curtis's Botanical Magazine''. Each of the issue ...
''. This plant is a recipient of the
Royal Horticultural Society The Royal Horticultural Society (RHS), founded in 1804 as the Horticultural Society of London, is the UK's leading gardening charity. The RHS promotes horticulture through its five gardens at Wisley (Surrey), Hyde Hall (Essex), Harlow Carr (Nor ...
's
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established annual award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. History The Award of Garden Merit ...
. There are several dozen ''Cephalotus'' clones that exist in cultivation; nine have been officially registered as
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s. One of the most well-known is 'Eden Black', a cultivar with unusually dark-coloured pitchers.


Genomics

The
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
of the pitcher plant ''Cephalotus follicularis'' has been sequenced. 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 Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
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.


See also

*
Carnivorous plants of Australia ''Carnivorous Plants of Australia'' is a three-volume work on carnivorous plants by Allen Lowrie. The three tomes were published in 1987, 1989, and 1998, by University of Western Australia Press. An entirely updated three-volume work by Lowrie w ...


References


External links

* *
Inner World of ''Cephalotus follicularis'' from the John Innes Centre
* {{Taxonbar, from=Q2181577 Monotypic Oxalidales genera Oxalidales Oxalidales of Australia Carnivorous plants of Australia Rosids of Western Australia Vulnerable flora of Australia Endemic flora of Southwest Australia