Archaeamphora longicervia
Encyclopedia
Archaeamphora longicervia is an extinct
species of pitcher plant
bearing close affinities to extant members of the family
Sarraceniaceae
. Fossil material assigned to this taxon
originates from the Yixian Formation
of northeastern China
, dated to the Early Cretaceous
(around ).
Archaeamphora longicervia is the earliest known carnivorous plant
and represents the only fossil record of pitcher plants (with the possible exception of some palynomorph
s of uncertain nepenthacean
affinity). Furthermore, the monotypic Archaeamphora is one of the three oldest known genera
of angiosperms
(flowering plants). Li (2005) notes that "the existence of a so highly derived Angiosperm in the Early Cretaceous suggests that Angiosperms should have originated much earlier, maybe back to 280 mya as the molecular clock
studies suggested".
: αρχαίος, archaios, combining form in Latin
archae- and the Greek
amphoreus (pitcher). The specific epithet longicervia is derived from the Latin
longus (long) and cervicarius (with a neck). The latter refers to the characteristic constriction in the pitchers of this species.
, western Liaoning
, China
. These Early Cretaceous
beds constitute the lower part of the Yixian Formation
, which is dated at 124.6 million years old. Nine specimens of A. longicervia have been found, including holotype
CBO0220 and paratype
CBO0754.
plant growing to around 50 mm (2 in) in height. The stem, at least 21 mm (.8 in) long and 1.2 mm (.04 in) wide, bore distinctive vertical ridges and grooves. Pitchers were ascidiate in form and 30 to 40 mm (1.2 to 1.6 in) long. Mature pitchers and underdeveloped pitchers or phyllodia-like leaves were arranged spirally around the stem. Pitchers consisted of a tubular base, expanded middle section, constriction around the mouth, and a vertical, spoon-shaped lid. A single wing ran down the adaxial side of each pitcher. Three to five parallel major veins were present on the pitchers, along with a few intercostal veins and numerous small veinlets.
Two unusual bag-like structures were present on each pitcher, one on either side of the central wing. Similar but semi-circular structures were found on the margin of the lid. These structures exhibited strong yellow-green intrinsic fluorescence
when exposed to 500 nm
wavelength
light.
Tiny gland
s, approximately 4 µm
in diameter, were found on the inner surface of the pitchers and partially embedded in the grooves along the veins. These also showed very strong golden-yellow fluorescence.
A single seed
was found intimately associated with the fossil material of A. longicervia and is presumed to belong to the same species. It is winged and reticulate-tuberculate in morphology
, closely resembling the seeds of Sarraceniaceae
taxa
. The seed is oval-shaped, covered with black-brown warts, and measures 0.9 mm by 1.25 mm (.03 in by .05 in).
, considered a key marker differentiating angiosperms from gymnosperm
s. Oleanane was detected in the pitcher plant specimens, suggesting that they belong to the angiosperms.
Several morphological features of A. longicervia suggest a close relationship to Sarraceniaceae
; both taxa exhibit one or two pitcher wings, a smooth peristome
, and pitchers that extend vertically from the top of a short petiole
.
Li (2005) suggests that A. longicervia is morphologically similar to modern Sarracenia purpurea
. It shares with this species the spiral arrangement of its pitchers and phyllodia-like tubular leaves with parallel major veins. Archaeamphora longicervia also shows close affinity to species of the genus Heliamphora
in having pitchers with a long neck and upright lid. Of particular note is the similarity between the thick semi-circular structures on the lid of A. longicervia and the large nectar-secreting "spoon" present on the upper posterior portion of Heliamphora exappendiculata
pitchers.
Li (2005) mentions the discovery of another type of pitcher plant from the same formation. This variety differs from the type material of A. longicervia in having pitchers that lack any constriction before the mouth, instead gradually expanding from the petiole
into a hollow trumpet-like shape. He suggests that it "should be a different species" from A. longicervia. An intermediate form with a wider neck is also reported, suggesting that pitcher plants were already a diversified group in the Early Cretaceous.
in the region was mostly composed of lacustrine
sediments and volcanic rock
s.
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
species of pitcher plant
Pitcher plant
Pitcher plants are carnivorous plants whose prey-trapping mechanism features a deep cavity filled with liquid known as a pitfall trap. It has been widely assumed that the various sorts of pitfall trap evolved from rolled leaves, with selection pressure favouring more deeply cupped leaves over...
bearing close affinities to extant members of the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Sarraceniaceae
Sarraceniaceae
Sarraceniaceae is a family of pitcher plants , belonging to order Ericales .The family comprises three extant genera: Sarracenia , Darlingtonia , and Heliamphora . The extinct Archaeamphora longicervia may also belong to this family...
. Fossil material assigned to this taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
originates from the Yixian Formation
Yixian Formation
The Yixian Formation is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans 11 million years during the early Cretaceous period...
of northeastern China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
, dated to the Early Cretaceous
Early Cretaceous
The Early Cretaceous or the Lower Cretaceous , is the earlier or lower of the two major divisions of the Cretaceous...
(around ).
Archaeamphora longicervia is the earliest known 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...
and represents the only fossil record of pitcher plants (with the possible exception of some palynomorph
Palynomorph
Palynomorph is the geological term used to describe a particle of a size between five and 500 micrometres, found in rock deposits and composed of organic material such as chitin, pseudochitin and sporopollenin...
s of uncertain nepenthacean
Nepenthes
The Nepenthes , popularly known as tropical pitcher plants or monkey cups, are a genus of carnivorous plants in the monotypic family Nepenthaceae. The genus comprises roughly 130 species, numerous natural and many cultivated hybrids...
affinity). Furthermore, the monotypic Archaeamphora is one of the three oldest known genera
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 angiosperms
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...
(flowering plants). Li (2005) notes that "the existence of a so highly derived Angiosperm in the Early Cretaceous suggests that Angiosperms should have originated much earlier, maybe back to 280 mya as the molecular clock
Molecular clock
The molecular clock is a technique in molecular evolution that uses fossil constraints and rates of molecular change to deduce the time in geologic history when two species or other taxa diverged. It is used to estimate the time of occurrence of events called speciation or radiation...
studies suggested".
Etymology
The generic name Archaeamphora is derived from the GreekGreek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
: αρχαίος, archaios, combining form in Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
archae- and the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
amphoreus (pitcher). The specific epithet longicervia is derived from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
longus (long) and cervicarius (with a neck). The latter refers to the characteristic constriction in the pitchers of this species.
Fossil material
All known fossil material of A. longicervia originates from the Jianshangou Formation in BeipiaoBeipiao
Beipiao is a city in Chaoyang prefecture, Liaoning province, in Northeast China. It has a population of 202,807. The main industry in the area is coal mining. With vertical shafts of almost 1000m, these are some of the deepest coal mines in China. The coal produced is used for coking...
, western Liaoning
Liaoning
' is a province of the People's Republic of China, located in the northeast of the country. Its one-character abbreviation is "辽" , a name taken from the Liao River that flows through the province. "Níng" means "peace"...
, China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
. These Early Cretaceous
Early Cretaceous
The Early Cretaceous or the Lower Cretaceous , is the earlier or lower of the two major divisions of the Cretaceous...
beds constitute the lower part of the Yixian Formation
Yixian Formation
The Yixian Formation is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans 11 million years during the early Cretaceous period...
, which is dated at 124.6 million years old. Nine specimens of A. longicervia have been found, including holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...
CBO0220 and paratype
Paratype
Paratype is a technical term used in the scientific naming of species and other taxa of organisms. The exact meaning of the term paratype when it is used in zoology is not the same as the meaning when it is used in botany...
CBO0754.
Description
Archaeamphora longicervia was a herbaceousHerbaceous
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...
plant growing to around 50 mm (2 in) in height. The stem, at least 21 mm (.8 in) long and 1.2 mm (.04 in) wide, bore distinctive vertical ridges and grooves. Pitchers were ascidiate in form and 30 to 40 mm (1.2 to 1.6 in) long. Mature pitchers and underdeveloped pitchers or phyllodia-like leaves were arranged spirally around the stem. Pitchers consisted of a tubular base, expanded middle section, constriction around the mouth, and a vertical, spoon-shaped lid. A single wing ran down the adaxial side of each pitcher. Three to five parallel major veins were present on the pitchers, along with a few intercostal veins and numerous small veinlets.
Two unusual bag-like structures were present on each pitcher, one on either side of the central wing. Similar but semi-circular structures were found on the margin of the lid. These structures exhibited strong yellow-green intrinsic fluorescence
Fluorescence
Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation of a different wavelength. It is a form of luminescence. In most cases, emitted light has a longer wavelength, and therefore lower energy, than the absorbed radiation...
when exposed to 500 nm
Nanometre
A nanometre is a unit of length in the metric system, equal to one billionth of a metre. The name combines the SI prefix nano- with the parent unit name metre .The nanometre is often used to express dimensions on the atomic scale: the diameter...
wavelength
Wavelength
In physics, the wavelength of a sinusoidal wave is the spatial period of the wave—the distance over which the wave's shape repeats.It is usually determined by considering the distance between consecutive corresponding points of the same phase, such as crests, troughs, or zero crossings, and is a...
light.
Tiny gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...
s, approximately 4 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...
in diameter, were found on the inner surface of the pitchers and partially embedded in the grooves along the veins. These also showed very strong golden-yellow fluorescence.
A single 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...
was found intimately associated with the fossil material of A. longicervia and is presumed to belong to the same species. It is winged and reticulate-tuberculate in morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
, closely resembling the seeds of Sarraceniaceae
Sarraceniaceae
Sarraceniaceae is a family of pitcher plants , belonging to order Ericales .The family comprises three extant genera: Sarracenia , Darlingtonia , and Heliamphora . The extinct Archaeamphora longicervia may also belong to this family...
taxa
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
. The seed is oval-shaped, covered with black-brown warts, and measures 0.9 mm by 1.25 mm (.03 in by .05 in).
Taxonomy
The fossil material of A. longicervia was subjected to chemical analysis for oleananeOleanane
Oleanane is a natural triterpene. It forms the central core for a wide variety of chemical compounds found in flowering plants which are referred to collectively as oleanane triterpenes.Some oleanane triterpenes have a suppressing effect on insect pests...
, considered a key marker differentiating angiosperms from gymnosperm
Gymnosperm
The gymnosperms are a group of seed-bearing plants that includes conifers, cycads, Ginkgo, and Gnetales. The term "gymnosperm" comes from the Greek word gymnospermos , meaning "naked seeds", after the unenclosed condition of their seeds...
s. Oleanane was detected in the pitcher plant specimens, suggesting that they belong to the angiosperms.
Several morphological features of A. longicervia suggest a close relationship to Sarraceniaceae
Sarraceniaceae
Sarraceniaceae is a family of pitcher plants , belonging to order Ericales .The family comprises three extant genera: Sarracenia , Darlingtonia , and Heliamphora . The extinct Archaeamphora longicervia may also belong to this family...
; both taxa exhibit one or two pitcher wings, a smooth peristome
Peristome
The word peristome is derived from the Greek peri, meaning 'around' or 'about', and stoma, 'mouth'. It is a term used to describe various anatomical features that surround an opening to an organ or structure. The term is used in plants and invertebrate animals, such as in describing the shells of...
, and pitchers that extend vertically from the top of a short petiole
Petiole (botany)
In botany, the petiole is the stalk attaching the leaf blade to the stem. The petiole usually has the same internal structure as the stem. Outgrowths appearing on each side of the petiole are called stipules. Leaves lacking a petiole are called sessile, or clasping when they partly surround the...
.
Sarracenia purpurea
Sarracenia purpurea, commonly known as the purple pitcher plant, northern pitcher plant, or side-saddle flower, is a carnivorous plant in the family Sarraceniaceae...
. It shares with this species the spiral arrangement of its pitchers and phyllodia-like tubular leaves with parallel major veins. Archaeamphora longicervia also shows close affinity to species of the genus Heliamphora
Heliamphora
The genus Heliamphora contains 23 species of pitcher plants endemic to South America. The species are collectively known as sun pitchers, based on the mistaken notion that the heli of Heliamphora is from the Greek helios, meaning "sun"...
in having pitchers with a long neck and upright lid. Of particular note is the similarity between the thick semi-circular structures on the lid of A. longicervia and the large nectar-secreting "spoon" present on the upper posterior portion of Heliamphora exappendiculata
Heliamphora exappendiculata
Heliamphora exappendiculata is a species of Marsh Pitcher Plant endemic to the Chimanta massif and Aprada Tepui in Venezuela. It was for a long time considered a variety of H. heterodoxa, but has recently been raised to species rank.-References:* McPherson, S. 2007. Pitcher Plants of the Americas...
pitchers.
Li (2005) mentions the discovery of another type of pitcher plant from the same formation. This variety differs from the type material of A. longicervia in having pitchers that lack any constriction before the mouth, instead gradually expanding from the petiole
Petiole (botany)
In botany, the petiole is the stalk attaching the leaf blade to the stem. The petiole usually has the same internal structure as the stem. Outgrowths appearing on each side of the petiole are called stipules. Leaves lacking a petiole are called sessile, or clasping when they partly surround the...
into a hollow trumpet-like shape. He suggests that it "should be a different species" from A. longicervia. An intermediate form with a wider neck is also reported, suggesting that pitcher plants were already a diversified group in the Early Cretaceous.
Habitat
The area inhabited by A. longicervia is thought to have experienced significant climatic fluctuations during the Early Cretaceous, ranging from arid or semi-arid to more humid conditions. The substrateSubstrate (biology)
In biology a substrate is the surface a plant or animal lives upon and grows on. A substrate can include biotic or abiotic materials and animals. For example, encrusting algae that lives on a rock can be substrate for another animal that lives on top of the algae. See also substrate .-External...
in the region was mostly composed of lacustrine
Lake
A lake is a body of relatively still fresh or salt water of considerable size, localized in a basin, that is surrounded by land. Lakes are inland and not part of the ocean and therefore are distinct from lagoons, and are larger and deeper than ponds. Lakes can be contrasted with rivers or streams,...
sediments and volcanic rock
Volcanic rock
Volcanic rock is a rock formed from magma erupted from a volcano. In other words, it is an igneous rock of volcanic origin...
s.