Byblis (plant)
Encyclopedia
Byblis is a small genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of carnivorous plant
Carnivorous plant
Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic...

s, sometimes termed the rainbow plants for the attractive appearance of their mucilage
Mucilage
Mucilage is a thick, gluey substance produced by most plants and some microorganisms. It is a polar glycoprotein and an exopolysaccharide.It occurs in various parts of nearly all classes of plant, usually in relatively small percentages, and is frequently associated with other substances, such as...

-covered leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 in bright sunshine. Native to western Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, it is the only genus in the family Byblidaceae. The first species in the genus was described by the English botanist Richard Anthony Salisbury
Richard Anthony Salisbury
Richard Anthony Salisbury FRS was a British botanist. While he is remembered as a valuable worker in horticultural and botanical sciences, several bitter disputes caused him to be ostracised by his contemporaries.-Life:...

 in 1808. Seven species are now recognized (see below).

Byblis species look very similar to Drosera and Drosophyllum
Drosophyllum
Drosophyllum is a genus of carnivorous plants containing the single species Drosophyllum lusitanicum...

, but are distinguished by their zygomorphic flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

s, with five curved stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...

s off to one side of the pistil. These genera are in fact not closely related; modern classifications place Byblis in the Lamiales
Lamiales
Lamiales is an order in the asterid group of dicotyledonous flowering plants. It includes approximately 11,000 species divided into about 20 families...

, while the sundew
Sundew
Drosera, commonly known as the sundews, comprise one of the largest genera of carnivorous plants, with at least 194 species. These members of the family Droseraceae lure, capture, and digest insects using stalked mucilaginous glands covering their leaf surface. The insects are used to supplement...

s and Drosophyllum are now placed in the Caryophyllales
Caryophyllales
Caryophyllales is an order of flowering plants that includes the cacti, carnations, amaranths, ice plants, and many carnivorous plants. Many members are succulent, having fleshy stems or leaves.-Description:...

.

Plant characteristics

All species of the genus form upright growth supported by a weak, fibrous root system. The genus can be divided into two groups or "complexes": The B. liniflora complex and the B. gigantea complex (see below).

Leaves

The leaves of all species are round in cross section and highly elongated, tapering at the end. The surface of the leaves is densely studded with glandular hairs which secrete a mucilaginous
Mucilage
Mucilage is a thick, gluey substance produced by most plants and some microorganisms. It is a polar glycoprotein and an exopolysaccharide.It occurs in various parts of nearly all classes of plant, usually in relatively small percentages, and is frequently associated with other substances, such as...

 substance from their tip. These serve to attract small insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s, which upon touching the sticky secretions are ensnared. Unless they are strong enough to escape, the insect prey either die of exhaustion or asphyxiate as the mucilage envelops them and clogs their spiracles. Unlike the sundews, however, Byblis can move neither their tentacles nor the leaves themselves to aid trapping or digestion. As a result, they are grouped among the "passive flypaper traps" along with Pinguicula
Pinguicula
The butterworts are a group of carnivorous plants comprising the genus Pinguicula. Members of this genus use sticky, glandular leaves to lure, trap, and digest insects in order to supplement the poor mineral nutrition they obtain from the environments. Of the roughly 80 currently known species, 12...

, Drosophyllum
Drosophyllum
Drosophyllum is a genus of carnivorous plants containing the single species Drosophyllum lusitanicum...

, Roridula
Roridula
Roridula is a South African genus of plants that, whilst having many of the adaptations of a carnivorous plant, such as the possession of insect-trapping sticky hairs, does not directly digest the animals it traps. Instead, it has a mutualistic relationship with Pameridea roridulae, a species of...

, Stylidium and Triphyophyllum peltatum.

Along with the stalked mucilaginous glands, the leaves are also equipped with sessile
Sessility (botany)
In botany, sessility is a characteristic of plants whose flowers or leaves are borne directly from the stem or peduncle, and thus lack a petiole or pedicel...

 glands, which assumedly are responsible for the secretion of the digestive juices. Sessile glands are five to ten times as numerous as the stalked glands.

Flowers

Flowers in this genus are born singly at the end of unbranching, leaf-like inflorescence
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. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

s which emerge from the leaf axes. The five-petaled flowers are generally purple to pale violet, though B. gigantea und B. filifolia can sometimes produce white flowers. Except for the self-fertile B. liniflora, all species require pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

 from other individuals for fertilization. The pollen release of B. gigantea and B. lamellata is only triggered by the resonance frequency of a landing pollinator, helping ensure cross-pollination with other individuals.

Fruit and seeds

Fertilized flowers mature to form an egg shaped, two-parted seed capsule. As the seed capsule dries out it cracks open (dehisces), dropping the seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

 on the ground (see gravity dispersal). The black seeds are generally round and often bear webbed surface markings, although those of B. lamellata are strongly ridged (see http://seeds.carnivoren.org/images/156.jpg). The germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

 of many species is brought on by brush fires after the dry period; components of the smoke are responsible for triggering germination.

Distribution and habitat

All Byblis species are native to Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

. B. gigantea and B. lamellata are endemic to the Perth
Perth, Western Australia
Perth is the capital and largest city of the Australian state of Western Australia and the fourth most populous city in Australia. The Perth metropolitan area has an estimated population of almost 1,700,000....

 region of southwest Australia, while the species making up the B. liniflora complex are found only in north Australia. The exception here is B. liniflora itself, whose distribution extends into southern Indonesia
Indonesia
Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...

 and Papua New Guinea
Papua New Guinea
Papua New Guinea , officially the Independent State of Papua New Guinea, is a country in Oceania, occupying the eastern half of the island of New Guinea and numerous offshore islands...

.

Like many carnivorous plants, Byblis species usually grow in bog
Bog
A bog, quagmire or mire is a wetland that accumulates acidic peat, a deposit of dead plant material—often mosses or, in Arctic climates, lichens....

s and marsh
Marsh
In geography, a marsh, or morass, is a type of wetland that is subject to frequent or continuous flood. Typically the water is shallow and features grasses, rushes, reeds, typhas, sedges, other herbaceous plants, and moss....

es. They generally prefer seasonally wet sandy soil in partial or direct sunlight with temperatures between ~ 5-40 °C (40-105 °F).

Environmental status

As native plants of Australia, all Byblis species are protected. Until the year 2000, they were also given international protection under CITES appendix II, but were removed from the list when Australia entered the CITES agreement. Since then trade of the genus has been unregulated outside of Australia. However, due the sensitivity of the plant, interest in the genus has been restricted to a small portion of the carnivorous plant hobbyist community. The majority of plant material sold today is produced in cultivation, with the annual B. filifolia and B. liniflora being the most common. Most of the other species must be grown from seed, which is often collected from the wild for this purpose.

The West Australian species B. gigantea und B. lamellata are being threatened by habitat destruction for urban sprawl
Urban sprawl
Urban sprawl, also known as suburban sprawl, is a multifaceted concept, which includes the spreading outwards of a city and its suburbs to its outskirts to low-density and auto-dependent development on rural land, high segregation of uses Urban sprawl, also known as suburban sprawl, is a...

 from cities such as Perth
Perth, Western Australia
Perth is the capital and largest city of the Australian state of Western Australia and the fourth most populous city in Australia. The Perth metropolitan area has an estimated population of almost 1,700,000....

. Particularly damaging is the draining of wet habitats to produce arable land. B. gigantea is on the Internation Union for Conservation of Nature's Red List of Threatened Species and is considered critically endangered.

Carnivorous or protocarnivorous

The status of the genus as a truly carnivorous plant
Carnivorous plant
Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic...

 has been repeatedly put into question. In their natural habitat, all species have been observed playing host to live bug
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...

s of the genus Setocoris, which nourished themselves by eating prey caught by the plants. Following this discovery it was assumed that, as with the genus Roridula
Roridula
Roridula is a South African genus of plants that, whilst having many of the adaptations of a carnivorous plant, such as the possession of insect-trapping sticky hairs, does not directly digest the animals it traps. Instead, it has a mutualistic relationship with Pameridea roridulae, a species of...

, the plants don't actually digest their prey themselves, rather relying on the bugs to do that. The plants, it was reasoned, benefited by absorbing nutrients from the excrements of the bugs, either through their leaves or through the ground. An indirect digestion of these nutrients by a chitinase
Chitinase
Chitinases are hydrolytic enzymes that break down glycosidic bonds in chitin. As chitin is a component of the cell walls of fungi and exoskeletal elements of some animals , chitinases are generally found in organisms that either need to reshape their own chitin or dissolve and digest the chitin of...

 producing fungus
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...

 was even proposed. It wasn't until 2005 that direct digestion of insect prey by enzymes secreted by the sessile glands of B. filifolia was proven. Soon thereafter similar results were found with B. liniflora. These results clearly place this genus among the true carnivorous plants.

Systematics

Molecular genetics studies have placed the genus in the family Lamiales
Lamiales
Lamiales is an order in the asterid group of dicotyledonous flowering plants. It includes approximately 11,000 species divided into about 20 families...

. While its placement within the order is still unclear, it is closely related to Martyniaceae
Martyniaceae
Martyniaceae is a family of flowering plants in the Lamiales order that are restricted to the New World. The family was included in Pedaliaceae in the Cronquist system but is recognized as a separate family by the Angiosperm Phylogeny Group on the basis of phylogenetic studies that show that the...

, Lentibulariaceae
Lentibulariaceae
Lentibulariaceae is a family of carnivorous plants containing three genera, Genlisea, the corkscrew plants, Pinguicula, the butterworts, and Utricularia, the bladderworts....

 as well as Gesneriaceae
Gesneriaceae
Gesneriaceae is a family of flowering plants consisting of ca. 150 genera and ca. 3,200 species in the Old World and New World tropics and subtropics, with a very small number extending to temperate areas. Many species have colorful and showy flowers and are cultivated as ornamental plants.Most...

.

For a time, the genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 Roridula
Roridula
Roridula is a South African genus of plants that, whilst having many of the adaptations of a carnivorous plant, such as the possession of insect-trapping sticky hairs, does not directly digest the animals it traps. Instead, it has a mutualistic relationship with Pameridea roridulae, a species of...

was also assigned to the family Byblidaceae. Since that time, however, it have been recategorized into its own family, Roridulaceae.

Traditionally the genus was divided into only two species, namely B. gigantea and B. liniflora. Further species were described in the 1980s, particularly through the work of the Australian botanist Allen Lowrie
Allen Lowrie
Allen Lowrie is a West Australian botanist. He is living in Duncraig, a Perth suburb, is married and has two daughters.Lowrie, originally a businessman and inventor, got in contact with the carnivorous flora of western Australia in the late sixties and worked on it as an amateur...

. Seven species are currently recognized:
  • Byblis aquatica
    Byblis aquatica
    Byblis aquatica is an insectivorous plant belonging to the genus Byblis, commonly known as the rainbow plants. It was described by Allen Lowrie and John Godfrey Conran in 1998, assigned to a group of annual north Australian species known as the "Byblis liniflora complex"...

    (annual, scrambling stem up to 45 cm (17.7 in), semiaquatic habitats)
  • Byblis filifolia
    Byblis filifolia
    Byblis filifolia is a species of plant in the Byblidaceae family. It is endemic to Australia.-References:* Conran, J.G., Lowrie, A. & Leach, G. 2000. . Downloaded on 20 August 2007....

    (annual, up to 60 cm (23.6 in), anthers longer than filament
    Stamen
    The stamen is the pollen producing reproductive organ of a flower...

    s)
  • Byblis gigantea
    Byblis gigantea
    Byblis gigantea is a species of plant in the Byblidaceae family. It is endemic to Australia.-Source:* Conran, J.G., Lowrie, A. & Leach, G. 2000. . Downloaded on 20 August 2007....

    (perennial, up to 70 cm (27.6 in), seeds with honeycomb pattern)
  • Byblis guehoi
    Byblis guehoi
    Byblis guehoi is a species of carnivorous plant in the genus Byblis. It is a compact species and is tetraploid. It was described in 2008 by Allen Lowrie and John Godfrey Conran. It is endemic to the Kimberley region of Western Australia....

  • Byblis lamellata
    Byblis lamellata
    Byblis lamellata is a carnivorous plant in the Byblidaceae family. It is endemic to Australia.-References:* Conran, J.G., A. Lowrie & J. Moyle-Croft 2002. A revision of Byblis in south-western Australia. Nuytsia 15: 11–19....

    (perennial, up to 45 cm (17.7 in), deeply ridged seeds)
  • Byblis liniflora
    Byblis liniflora
    Byblis liniflora is a species of carnivorous plant in the Byblidaceae family. It is found in Australia, Indonesia, and Papua New Guinea.-References:* Conran, J.G., Lowrie, A. & Leach, G. 2000. . Downloaded on 20 August 2007....

    (annual, up to 15 cm (5.9 in), anthers shorter than filament
    Stamen
    The stamen is the pollen producing reproductive organ of a flower...

    s)
  • Byblis rorida
    Byblis rorida
    Byblis rorida is a species of plant in the Byblidaceae family. It is endemic to Australia.-References:* Conran, J.G., Lowrie, A. & Leach, G. 2000. . Downloaded on 20 August 2007....

    (annual, up to 30 cm (11.8 in), heavily set with glandular tentacles)

Subdivision of the genus

Byblis liniflora complex

The four species of this complex, B. liniflora, B. rorida, B. filifolia and B. aquatica, are annual
Annual plant
An annual plant is a plant that usually germinates, flowers, and dies in a year or season. True annuals will only live longer than a year if they are prevented from setting seed...

 herbaceous
Herbaceous
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...

 plants that reach a height of 15–50 cm (5.9–19.7 in) and a maximum leaf length of 4–15 cm (1.6–5.9 in). These species grow from seedlings to flowering plants in only a few months, setting seed and dying with the onset of the dry season. The original haploid chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...

 count of this complex is x=8. The diploid number is therefore 2n=16, whereas the tetraploid species B. liniflora is 2n=32.

Byblis gigantea complex

The remaining two species, B. lamellata und B. gigantea, make up what is known as the B. gigantea complex. These perennial species are both endemic to Southwest Australia
Southwest Australia
Southwest Australia is a biodiversity hotspot that includes the Mediterranean forests, woodlands, and scrub ecoregions of Western Australia. The region has a wet-winter, dry-summer Mediterranean climate, one of five such regions in the world...

, and reach heights of 45–70 cm (17.7–27.6 in). Unlike the annual members of the B. liniflora complex, these species survive the dry season by dying back to an underground rhizome
Rhizome
In botany and dendrology, a rhizome is a characteristically horizontal stem of a plant that is usually found underground, often sending out roots and shoots from its nodes...

, out of which they emerge come fall. The leaves of this complex can reach 20 cm (7.9 in) in length. The base chromosome count of the complex is x=9; since both species are diploid, their chromosome count is 2n=18.

Paleobotany

In the year 2004 a single fossil of a seed, resembling that of members of the modern day B. liniflora complex, was discovered in south Australia dating from the middle of the Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...

 epoch. The species was assigned to the Byblidaceae as a parataxon of the genus.

Etymology

The Latin generic name "Byblis" originates from a goddess from Greek mythology
Greek mythology
Greek mythology is the body of myths and legends belonging to the ancient Greeks, concerning their gods and heroes, the nature of the world, and the origins and significance of their own cult and ritual practices. They were a part of religion in ancient Greece...

, of whom Ovid
Ovid
Publius Ovidius Naso , known as Ovid in the English-speaking world, was a Roman poet who is best known as the author of the three major collections of erotic poetry: Heroides, Amores, and Ars Amatoria...

 wrote in his Metamorphoses (IX, l. 454-664). Byblis
Byblis
In Greek mythology, Byblis or Bublis was a daughter of Miletus. Her mother was either Tragasia, Cyanee, daughter of the river-god Meander, or Eidothea, daughter of King Eurytus of Caria. She fell in love with Caunus, her twin brother....

, niece of Apollo
Apollo
Apollo is one of the most important and complex of the Olympian deities in Greek and Roman mythology...

, fell deeply in love with her twin brother Caunus
Kaunos (mythology)
In Greek mythology, Caunus or Kaunos was a son of Miletus, grandson of Apollo and brother of Byblis.Caunus became the object of his own sister's passionate love. From some accounts it appears that Caunus was the first to develop the affection towards her; others describe Byblis' feelings as...

. At his rejection of her advances, she let forth an endless stream of glistening tears, eventually literally transforming into a spring. The droplets lining the leaves of the Byblis are said to resemble those tears.

The English vernacular name - "rainbow plants" - also denotes the mucilaginous droplets which, under the right lighting conditions and viewing angle, sparkle in a rainbow of colors.

Further reading

  • Conran, John G.: The embryology and relationships of the Byblidaceae, Australian Syst. Bot. 9, 243-254, 1996
  • Conran, John G.; Carolin, R.: Byblidaceae, in: Kadereit, J. (ed.): The Families and Genera of Vascular Plants, Vol. VII: Flowering Plants: Dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae), Springer, 2004, 45-49.
  • Fukushima, K., K. Imamura, K. Nagano & Y. Hoshi (2011). Journal of Plant Research 124(2): 231–244.

External links

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