Winteraceae

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Winteraceae
Temporal range: Albian - recent [1] [2]
Drimys winteri.jpg
Drimys winteri
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Magnoliids
Order: Canellales
Family: Winteraceae
R.Br. ex Lindl.
Genera [3]

Winteraceae is a primitive family of tropical trees and shrubs including 93 [4] species in five genera. [3] It is of particular interest because it is such a primitive angiosperm family, distantly related to Magnoliaceae, though it has a much more southern distribution. [5] Plants in this family grow mostly in the southern hemisphere, and have been found in tropical to temperate climate regions of Malesia, Oceania, eastern Australia, New Zealand, Madagascar and the Neotropics, [5] with most of the genera concentrated in Australasia and Malesia. The five genera, Takhtajania, Tasmannia, Drimys, Pseudowintera, and Zygogynum s.l. all have distinct geographic extant populations. Takhtajania includes a single species, T. perrieri, endemic only to Madagascar, Tasmannia has the largest distribution of genera in Winteraceae with species across the Philippines, Borneo, New Guinea, Eastern Australia, and Tasmannia, Drimys is found in the Neotropical realm, from southern Mexico to the subarctic forests of southern South America, Pseudowintera is found only in New Zealand, and Zygogynum has species in New Guinea and New Caledonia. [6]

Contents

This family has been estimated to be anywhere from 105 to at least 35 million years ago. [3] [7] Being one of few angiosperms forming persistent tetrads with prominent sculpturing, pollen of Winteraceae is rare but easy to identify in the fossil record. [7] Pollen samples found in Gabon may indicate that the family is at least 120 million years old, [8] but the association of these fossils with Winteraceae is uncertain. [7] Oldest unambiguous Winteraceae fossils are from the middle to late Albian of Israel (~110 million years old; described as Qatanipollis ). [2] Pollen fossils indicate that the range has been much wider than it is now, [3] reaching north as far as Greenland during the Paleocene (Danian), [7] and disappearing from continental Africa (Cape Peninsula, South Africa) in the Miocene. [9] Equally characteristic is Winteraceae wood, which lacks xylem vessels in contrast to most other flowering plants. [10] Fossil Winteraceae wood has been found in the Late Cretaceous to Paleogene (c. 85–35 million years ago) of Antarctica (Santonian-Campanian), [11] western North America (Central Valley, California; Maastrichian) [12] and Europe (Helmstedt, Germany; Eocene). [13]

According to the 1998 APG I system, it did not belong to any order, [14] but it has since been placed in Canellales by the APG II, APG III and APG IV systems. [15] [16] [17]

Winteraceae

Zygogynum

Pseudowintera

Drimys

Tasmannia

Takhtajania

Description

Members of the family Winteraceae are trees or shrubs. The leaves are alternate, with light green dots and a fragrant aroma. Some are used to produce essential oils. Stipules are absent. Flowers are small, mostly appearing in cymes or fascicles. They have two to six free, valvate sepals, though they are united in Drimys. [5]

The Winteraceae have no vessels in their xylem. [10] This makes them relatively immune to xylem embolisms caused by freezing temperatures. In addition, vascular occlusion can occur near the openings of the stomata, preventing excess water from entering. [3]

Among all species, the distinctive characters of released pollen tetrads are easily recognized using light and electron microscopy. [7] [18]

Evolution of vesselless wood

Winteraceae was initially placed as a basal group within the Angiosperms due to its vesselless wood. [19] Xylem vessels were seen as an important evolved character for the diversification and success of Angiosperms, so vesselless wood was seen as an archaic trait, resulting in basal placement of the Winteraceae. However, molecular phylogenetic work placed Winteraceae within the Magnoliids, well within the angiosperms. [19] This placement suggests that the vesselless wood of the Winteraceae was a derived character rather than ancestral. Through the fossil pollen record, it is hypothesized that Winteraceae moved from Northern Gondwana through Southern Gondwana in the Cretaceous. [20] This meant movement from hot humid environments to temperate humid environments where freeze-thaw events occurred. Vesselless wood has 20% of the water conductivity of vessel-bearing wood, however, under freeze events, wood with vessels loses up to 85% of water conductivity while vesselless wood loses at most 6% of water conductivity. [21] The ability to avoid serious water limitation and therefore the shedding of leaves is hypothesized to be a major evolutionary pressure behind the reversion to vesselless wood. This is further supported by the heteroxylly hypothesis in which “primitive” vessels conferred little difference in stem hydraulic efficiency under normal conditions as compared to vesselless angiosperms. This would indicate that the pressure of freeze-thaw events and the subsequent risk of embolism would be a stronger evolutionary factor compared to the weaker hydraulic constraints of vesselless wood compared to “primitive” vessels. [22] This movement from hot humid environments to temperature humid environments where freeze-thaw events occurred is seen as the evolutionary pressure behind the unique reversion to vesselless wood in Winteraceae.

Another character of Winteraceae that was seen to indicate a basal position in the phylogeny was the presence of waxy stomatal plugs, seen as limiting water loss in respiration and therefore an archaic trait to limit water loss. [23] However, further research showed that in these humid environments, water cover on the surface of leaves decreased photosynthetic rates and waxy stomatal plugs reduce this water cover and therefore reduce the negative impacts on photosynthetic ability. [24] Winteraceae species with stomatal plugs removed saw decreases in photosynethic rates of up to 40%. [20] This further shows that characters once thought to be archaic could rather be derived adaptations to temperate humid environments.

Rediscovery of Takhtajania

Takhtajania perrieri was first collected 1909 on the Manongarivo Massif of central Madagascar at an elevation of 1700 meters. In 1963, the French botanist René Paul Raymond Capuron examined the unidentified plant sample, which he identified as a new species, which he named Bubbia perrieri, after the French botanist Henri Perrier de la Bâthie, classifying it in the Australasian genus Bubbia. In 1978, the botanists Baranova and J. F. Leroy reclassified the plant into its own genus, Takhtajania, after the Russian botanist Armen Takhtajan. Many subsequent expeditions to find the species were futile, but in 1994 Malagasy plant collector Fanja Rasoavimbahoaka collected a specimen in Anjahanaribe-Sud Special Reserve 150 km from the location at which the 1909 specimen was collected, which George E. Schatz identified in May 1997 as Takhtajania. A subsequent expedition discovered a large grove of the species at the spot where the second sample was collected. [25]

Notable species

Drimys winteri (Winter's bark) is a slender tree native to the Magellanic and Valdivian temperate forests of Chile and Argentina. It is a common garden plant grown for its fragrant mahogany-red bark, bright-green leaves, and its clusters of creamy white, jasmine-scented flowers. The bark has historically been used to prevent scurvy. [26]

Tasmannia piperita is notable for the great range of numbers for petal, stamen and pistil counts.[ citation needed ] Tasmannia lanceolata , known as Tasmanian pepper, is grown as an ornamental shrub, and is increasingly being used as a condiment.[ citation needed ]

Related Research Articles

<span class="mw-page-title-main">Flowering plant</span> Clade of seed plants that produce flowers

Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae, commonly called angiosperms. They include all forbs, grasses and grass-like plants, a vast majority of broad-leaved trees, shrubs and vines, and most aquatic plants. The term "angiosperm" is derived from the Greek words ἀγγεῖον / angeion and σπέρμα / sperma ('seed'), meaning that the seeds are enclosed within a fruit. They are by far the most diverse group of land plants with 64 orders, 416 families, approximately 13,000 known genera and 300,000 known species. Angiosperms were formerly called Magnoliophyta.

<span class="mw-page-title-main">Canellales</span> Order of flowering plants

Canellales is the botanical name for an order of flowering plants, one of the four orders of the magnoliids. It is recognized by the most recent classification of flowering plants, the APG IV system. It is defined to contain two families: Canellaceae and Winteraceae, which comprise 136 species of fragrant trees and shrubs. The Canellaceae are found in tropical America and Africa, and the Winteraceae are part of the Antarctic flora. Although the order was defined based on phylogenetic studies, a number of possible synapomorphies have been suggested, relating to the pollen tube, the seeds, the thickness of the integument, and other aspects of the morphology.

<span class="mw-page-title-main">Nymphaeales</span> Order of flowering plants

The Nymphaeales are an order of flowering plants, consisting of three families of aquatic plants, the Hydatellaceae, the Cabombaceae, and the Nymphaeaceae. It is one of the three orders of basal angiosperms, an early-diverging grade of flowering plants. At least 10 morphological characters unite the Nymphaeales. One of the traits is the absence of a vascular cambium, which is required to produce both xylem (wood) and phloem, which therefore are missing. Molecular synapomorphies are also known.

<span class="mw-page-title-main">Nymphaeaceae</span> Family of plants

Nymphaeaceae is a family of flowering plants, commonly called water lilies. They live as rhizomatous aquatic herbs in temperate and tropical climates around the world. The family contains five genera with about 70 known species. Water lilies are rooted in soil in bodies of water, with leaves and flowers floating on or emergent from the surface. Leaves are round, with a radial notch in Nymphaea and Nuphar, but fully circular in Victoria and Euryale.

<span class="mw-page-title-main">Aristolochiaceae</span> Family of flowering plants

The Aristolochiaceae are a family, the birthwort family, of flowering plants with seven genera and about 400 known species belonging to the order Piperales. The type genus is Aristolochia L.

<span class="mw-page-title-main">Hamamelidaceae</span> Witch-hazel, a shrub or small tree

Hamamelidaceae, commonly referred to as the witch-hazel family, is a family of flowering plants in the order Saxifragales. The clade consists of shrubs and small trees positioned within the woody clade of the core Saxifragales. An earlier system, the Cronquist system, recognized Hamamelidaceae in the Hamamelidales order.

<span class="mw-page-title-main">Canellaceae</span> Family of flowering plants

The Canellaceae are a family of flowering plants in the order Canellales. The order includes only one other family, the Winteraceae. Canellaceae is native to the Afrotropical and Neotropical realms. They are small to medium trees, rarely shrubs, evergreen and aromatic. The flowers and fruit are often red.

<span class="mw-page-title-main">Nelumbonaceae</span> Family of flowering plants

Nelumbonaceae is a family of aquatic flowering plants. Nelumbo is the sole extant genus, containing Nelumbo lutea, native to North America, and Nelumbo nucifera, widespread in Asia. At least five other genera, Nelumbites, Exnelumbites, Paleonelumbo, Nelumbago, and Notocyamus are known from fossils.

<span class="mw-page-title-main">Saururaceae</span> Family of flowering plants

Saururaceae is a plant family comprising four genera and seven species of herbaceous flowering plants native to eastern and southern Asia and North America. The family has been recognised by most taxonomists, and is sometimes known as the "lizard's-tail family". The APG IV system assigned it to the order Piperales in the clade magnoliids.

<span class="mw-page-title-main">Vessel element</span> Component of Xylem

A vessel element or vessel member is one of the cell types found in xylem, the water conducting tissue of plants. Vessel elements are found in angiosperms but absent from gymnosperms such as conifers. Vessel elements are the main feature distinguishing the "hardwood" of angiosperms from the "softwood" of conifers.

<span class="mw-page-title-main">Trochodendraceae</span> Family of flowering plants

Trochodendraceae is the only family of flowering plants in the order Trochodendrales. It comprises two extant genera, each with a single species along with up to five additional extinct genera and a number of extinct species. The living species are native to south east Asia. The two living species both have secondary xylem without vessel elements, which is quite rare in angiosperms. As the vessel-free wood suggests primitiveness, these two species have attracted much taxonomic attention.

<span class="mw-page-title-main">Chloranthaceae</span> Family of flowering plants

Chloranthaceae is a family of flowering plants (angiosperms), the only family in the order Chloranthales. It is not closely related to any other family of flowering plants, and is among the early-diverging lineages in the angiosperms. They are woody or weakly woody plants occurring in Southeast Asia, the Pacific, Madagascar, Central and South America, and the West Indies. The family consists of four extant genera, totalling about 77 known species according to Christenhusz and Byng in 2016. Some species are used in traditional medicine. The type genus is Chloranthus. The fossil record of the family, mostly represented by pollen such as Clavatipollenites, extends back to the dawn of the history of flowering plants in the Early Cretaceous, and has been found on all continents.

Ctenolophon is the only genus in the flowering plant family Ctenolophonaceae. It has two recognized species:

<span class="mw-page-title-main">Cabombaceae</span> Family of flowering plants

The Cabombaceae are a family of aquatic, herbaceous flowering plants. A common name for its species is water shield. The family is recognised as distinct in the Angiosperm Phylogeny Group IV system (2016). The family consists of two genera of aquatic plants, Brasenia and Cabomba, totalling six species.

<i>Degeneria</i> Genus of flowering plants

Degeneria is a genus of flowering plants endemic to Fiji. It is the only genus in the family Degeneriaceae. The APG IV system of 2016, recognizes this family, and assigns it to the order Magnoliales in the clade magnoliids.

<i>Didymeles</i> Genus of trees

Didymeles is a genus of flowering plants. It is variously treated as the only genus of the family Didymelaceae — or in the family Buxaceae, as in the APG IV system.

<span class="mw-page-title-main">Triuridaceae</span> Family of flowering plants

Triuridaceae are a family of tropical and subtropical flowering plants, including nine genera with a total of approximately 55 known species. All members lack chlorophyll and are mycoheterotrophic. The heterotrophic lifestyle of these plants has resulted in a loss of xylem vessels and stomata, and a reduction of leaves to scales.

<span class="mw-page-title-main">Asteliaceae</span> Family of flowering plants

Asteliaceae is a family of flowering plants, placed in the order Asparagales of the monocots.

<span class="mw-page-title-main">Icacinales</span> Order of flowering plants

Icacinales is an order of Angiosperms.

Frost resistance is the ability of plants to survive cold temperatures. Generally, land plants of the northern hemisphere have higher frost resistance than those of the southern hemisphere. An example of a frost resistant plant is Drimys winteri which is more frost-tolerant than naturally occurring conifers and vessel-bearing angiosperms such as the Nothofagus that can be found in its range in southern South America.

References

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