Pentapetalae

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Pentapetalae
Schattenmorelle Bluete 01.jpg
Cerasus (Rosaceae)
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Class: Eudicotyledoneae
(unranked): Gunneridae
Clade: Pentapetalae
DE Soltis, PS Soltis & WS Judd 2007
Clades and orders [1]

In phylogenetic nomenclature, the Pentapetalae are a large group of eudicots that were informally referred to as the "core eudicots" in some papers on angiosperm phylogenetics. [2] They comprise an extremely large and diverse group accounting for about 65% of the species richness of the angiosperms, with wide variability in habit, morphology, chemistry, geographic distribution, and other attributes. Classical systematics, based solely on morphological information, was not able to recognize this group. In fact, the circumscription of the Pentapetalae as a clade is based on strong evidence obtained from DNA molecular analysis data. [3] [4] [5]

Contents

The Pentapetalae clade is composed of the orders Berberidopsidales—including the family Aextoxicaceae [6] [7] —Caryophyllales, Santalales and Saxifragales, the families Dilleniaceae and Vitaceae and all members of the clades Asteridae and Rosidae. [8]

Phylogenetic analyses of complete chloroplast genome sequences have provided a reliable outline of the relationships among the major Pentapetalae lineages and also provide a framework for investigating the evolutionary processes that generated a large proportion of the diversity of extant angiosperms. [9] In light of these phylogenetic results, the current challenge for scientists in this area of botany is to identify the characters that are unique to the superasterid and superrosid clades and those that arose in parallel in both, and then to explore their evolutionary implications. [10]

Description

As the name of the clade suggests, the Pentapetalae have a characteristic type of flower made up of whorls of five pieces each. The perianth is formed by a sepal and a corona perfectly differentiated. The sepals are innervated by three or more vascular bundles—called leaf scars [11] —originating in the vascular system of the petiole, while the petals have only one trace. The stamen usually has twice as many pieces as the calyx and corona, which are arranged in two whorls. When the number of stamens is greater than twice the number of pieces of the perianth, they are arranged in fascicles or in a centrifugal spiral. [2] [8]

Scanning electron microscope image of a tricolpate pollen grain in Arabis Arabis voch1-4.jpg
Scanning electron microscope image of a tricolpate pollen grain in Arabis

Pollen grains in the Pentapetalae are characteristically tricolpate. This type of pollen grain has three or more pores within grooves called "colpos". In contrast, most other spermatophytes—that is, gymnosperms, monocots and paleodicots—have monoculcate pollen, with a single pore located in a groove called a "sulcus". [2] [8]

The gynoecium of Pentapetalae plants is usually composed of five carpels joined together, although gynoeciums formed by three carpels are also quite common. In cases where the gynoecium is composed of only two carpels, they overlap. In general, they present "compitum", a region of the style where the stylar canals of the different carpels are united in a single cavity and in which the pollen tubes can change direction of growth from one carpel to another. The ovules are usually of axillary placentation. The pistil, finally, commonly terminates in a style and a stigma that is not decurrent. The fruit is dry and dehiscent, when it is a capsule it shows loculicidal dehiscence. Regarding the interaction between pollen and pistil, pentapetalous plants have a gametophytic incompatibility system based on the RNAase system. [2] [8] Another anatomical characteristic of Pentapetalae is the presence of a closed root apical meristem. From the phytochemical point of view, this group of plants present cyanogenesis—that is, they biosynthesize cyanogenetic glycosides that by hydrolysis originate cyanide—through the metabolic pathway of branched amino acids, such as leucine, isoleucine and valine. [2] [8]

Diversity

Dilleniales

Flower of Dillenia indica. Dillenia indica.jpg
Flower of Dillenia indica .

Dilleniales are recognized by their leaves with usually strong and parallel secondary veins that go straight to the teeth; being common the tertiary scalariform venation. The leaf lamina is usually rough. Also, the leaves tend to elongate when still rolled. The wood is usually vivid brown. The peduncles are jointed near the apex and persist after the flower falls off; the flowers are usually conspicuous, with ruffled petals and numerous stamens that are reflexed in the bud, usually having porous anthers. The fruits are small follicles containing seeds with aril, the calyx is persistent, sometimes acrescent, and the filaments are also persistent. [12]

Berberidopsidales

Flowers of Berberidopsis corallina Berberidopsis corallina 1.jpg
Flowers of Berberidopsis corallina

Berberidopsidales is an order formally accepted only in the most recent phylogenetic classifications of angiosperms, [13] [14] comprising two small families, Aextoxicaceae and Berberidopsidaceae, which together include only three genera and four species distributed in Chile and eastern Australia. The distinctive characters of the order are largely related to its anatomy, such as the presence of crystals—especially drusen—in the leaves and petioles, the vascular bundles of the petiole form a ring and the stoma of the leaf epidermis are of a particular type called "cyclocytic". The androecium has stamens with rigid filament and the seeds present endotesta.

The flowers of Berberidopsis corallina do not differ in sepals and petals, but exhibit a gradual transition from small outer tepals to larger, brightly colored inner tepals. The androcecium consists of a ring of stamens and there are three carpels with parietal placentation. In contrast, Aextoxicon has unisexual and pentamerous flowers. Male flowers have a distinct calyx and corona and a haplostomous androcecium. Female flowers have the same type of perianth, but the number of pieces is more variable. [15] [16]

Given these differences between the two genera belonging to the same family, it has been suggested that the floral development of Berberidopsidales is a "link" in the evolution of the Gunneridae [17] [18] flower, and that the floral morphology of Aextoxicon , with features such as the highly variable number of sepals and spirally arranged petals, is also compatible with this hypothesis. [19] However, the position of the Berberidopsidales in the phylogenetic tree is not congruent with it. [20]

Santalales

Flowers of Santalum album Santalum album (Chandan) in Hyderabad, AP W2 IMG 0023.jpg
Flowers of Santalum album

The order Santalales, with its approximately 160 genera and 2,200 species distributed worldwide, is a monophyletic group that has long occupied an unresolved position at the base of the Gunneridae. [21] Recently, phylogenetic studies based on molecular data from the complete chloroplast genome indicate a position of Santalales at the base of the Asterids. [22] Santalales is ecologically diverse and includes free-living plants, such as Erythropalum , as well as (hemi)parasites. Among the latter are species that parasitize stems or shoots, such as the mistletoes of the genus Misodendrum , and root parasites, including the well-known and economically important sandalwood tree ( Santalum album ), whose aromatic wood is a component of many perfumes. [23] [24] [25] The APG III classification system recognized seven families in Santalales: Balanophoraceae, Misodendraceae, Opiliaceae, Schoepfiaceae, Loranthaceae, Santalaceae (including Viscaceae), and Olacaceae. [1] In 2010, however, new molecular data have made it possible to revise that conclusion, so that Aptandraceae, Balanophoraceae, Coulaceae, Erythropalaceae, Loranthaceae, Misodendraceae, Octonemaceae, Olacaceae, Opiliaceae, Santalaceae, Schoepfiaceae, Strombosiaceae, and Ximeniaceae are now recognized. [26]

Rosids

Rose uchiyamane flower. Rosa uchiyamane.jpg
Rose uchiyamane flower.

The rosids are a large group of eudicotyledons containing approximately 70 000 species, [27] more than a quarter of the total number of angiosperm species. [28] It has been subdivided into some 16 to 20 orders, depending on the circumscription and classification adopted. These orders, in turn, comprise about 140 families. [29] Together with the asterids, they constitute the two largest groups of eudicotyledons.

The rosids share a few morphological characteristics that distinguish them from other groups, such as the presence of nectaries in the floral receptacle, the long embryo and the distinctive mucilaginous cells. At the molecular level, the rossids are characterized by the loss of function of the chloroplast infA gene and the absence of the coxII.i3 intron in the mitochondrion. [2]

Asterids

Inflorescence of an asteraceae. IMG 7411-Asteraceae sp..jpg
Inflorescence of an asteraceae.

The asterids are a large group of eudicotyledons that includes approximately 80 000 species, grouped in 13 orders and more than a hundred families, [27] and between a third and a quarter of the total number of angiosperm species. [28] [29] Together with the rosids, they constitute the two largest groups of eudicotyledons. They represent the most apotypic clade of the angiosperms, or, as it is inappropriately called, "most evolved". The group most likely originated in the Cenozoic, about 50 million years ago, and its success is related to its adaptation to insect pollinators. Four of the largest families of angiosperms belong to this clade: the Asteraceae, the Rubiaceae, the Lamiaceae and the Apocynaceae. [2]

The plants belonging to this clade are characterized by being herbaceous, with hermaphrodite, zygomorphic flowers—that is, they admit only one plane of symmetry—that are pollinated by insects. In addition, the stamens are arranged in a circle and the petals of the corona are joined together forming a tube. The gynoecium is formed by two welded carpels. The flowers are often arranged in tight inflorescences, such as the ears of the labiatae and plantaginaceae or the capitula (heads) of the compositae. [2]

Evolution and phylogeny

The age of this clade has been estimated at 113 to 116 million years. [30] [31] [20] The oldest macrofossils of eudicots, which unfortunately cannot be attributed to any extant group, belong to the Cretaceous-Cenomanian, just 96–94 million years old. [20]

Analyses of complete chloroplast genome sequences allowed us to resolve the relationships among the major Pentapetalae clades. Immediately after diverging from the Gunnerales, the Pentapetalae diverged into three major subclades: (i) the Dileniaceae, (ii) the superrosid clade including the Saxifragales, Vitaceae and Rosids, and (iii) the superasterid clade composed of the Berberidopsidales, Santalales, Caryophyllales and Asterids. The close relationship demonstrated at the molecular scale between Saxifragales, Vitaceae and rosids is congruent with their morphological affinities. In fact, these clades form the so-called "core of the rosids" in the Cronquist and Takhtajan classification systems. [32] [33] They all have an androcecium with jointed anthers, leaves with stipules, the endosperm with nuclear formation, and a micropyle that forms from the outer integument or both integuments. [34] [35] [20]

Several putative synapomorphies may also characterize the superrosid clade. Thus, the "psyllulate" pollen or with a granular structure of the exine, the presence of sclereids in the leaves, the isomerism of the androcecium and the fused carpels unite the santalales, the caryophyllales and the asterids. Likewise, leaves without stipules may be another synapomorphic character, although Berberidopsidaceae have stipules and Aextoxicaceae lack them; thus, the ancestral status of Berberidopsidales and asterids is still unclear. [2] [20]

The initial divergence between the Dilleniales, superrosids and superasterids must have occurred very quickly, within a period of one million years after the initial separation of the Pentapetalae from the Gunnerales. Likewise, the superrosids and superasterids show an early and very rapid divergence since the lineages that led to the Vitaceae, Saxifragales, and rosids arose within a period of only five million years, as did those that led to the appearance of the Berberidopsidales, Caryophyllales, and asterids. [20]

The following cladogram summarizes the phylogenetic relationships within the Pentapetalae clade and this clade. [36] [37]

eudicots
basal eudicots

(paraphyletic group: Ranunculales, Proteales, Trochodendrales, Buxales)

core eudicots

Gunnerales

Pentapetalae

Dilleniales

superrosids

Saxifragales

rosids

Vitales

eurosids

fabids (8 orders)

malvids (8 orders)

superasterids

Santalales

Berberidopsidales

Caryophyllales

asterids

Cornales

Ericales

euasterids

campanulids (7 orders)

lamiids (8 orders)

The names lamiids (for euasterids I) and campanulids (for euasterids II) were suggested by Bremer et al. (2002) and later suggested fabiids (for eurosids I) and malvids (for eurosids II).

Related Research Articles

<span class="mw-page-title-main">Apiales</span> Order of eudicot flowering plants in the asterid group

The Apiales are an order of flowering plants. The families are those recognized in the APG III system. This is typical of the newer classifications, though there is some slight variation and in particular, the Torriceliaceae may also be divided.

<span class="mw-page-title-main">Malpighiales</span> Eudicot order of flowering plants

The Malpighiales comprise one of the largest orders of flowering plants, containing about 36 families and more than 16,000 species, about 7.8% of the eudicots. The order is very diverse, containing plants as different as the willow, violet, poinsettia, manchineel, rafflesia and coca plant, and are hard to recognize except with molecular phylogenetic evidence. It is not part of any of the classification systems based only on plant morphology. Molecular clock calculations estimate the origin of stem group Malpighiales at around 100 million years ago (Mya) and the origin of crown group Malpighiales at about 90 Mya.

<span class="mw-page-title-main">Saxifragales</span> Order of Eudicot flowering plants in the Superrosid clade

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 vegetative and floral features makes it difficult to define common features that unify the order.

Urticales is an order of flowering plants. Before molecular phylogenetics became an important part of plant taxonomy, Urticales was recognized in many, perhaps even most, systems of plant classification, with some variations in circumscription. Among these is the Cronquist system (1981), which placed the order in the subclass Hamamelidae [sic], as comprising:

<span class="mw-page-title-main">Geraniales</span> Order of flowering plants in the rosid subclade of eudicots

Geraniales is a small order of flowering plants, included within the rosid subclade of eudicots. The largest family in the order is Geraniaceae with over 800 species. In addition, the order includes the smaller Francoaceae with about 40 species. Most Geraniales are herbaceous, but there are also shrubs and small trees.

<span class="mw-page-title-main">Celastrales</span> Order of flowering plants, mostly from tropics and subtropics

The Celastrales are an order of flowering plants found throughout the tropics and subtropics, with only a few species extending far into the temperate regions. The 1200 to 1350 species are in about 100 genera. All but seven of these genera are in the large family Celastraceae. Until recently, the composition of the order and its division into families varied greatly from one author to another.

<span class="mw-page-title-main">Primulaceae</span> Family of flowering plants that includes the primroses

The Primulaceae, commonly known as the primrose family, are a family of herbaceous and woody flowering plants including some favourite garden plants and wildflowers. Most are perennial though some species, such as scarlet pimpernel, are annuals.

<span class="mw-page-title-main">Angiosperm Phylogeny Group</span> Collaborative research group for the classification of flowering plants

The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships discovered through phylogenetic studies.

<span class="mw-page-title-main">Saxifragaceae</span> Family of flowering plants in the Eudicot order Saxifragales

Saxifragaceae is a family of herbaceous perennial flowering plants, within the core eudicot order Saxifragales. The taxonomy of the family has been greatly revised and the scope much reduced in the era of molecular phylogenetic analysis. The family is divided into ten clades, with about 640 known species in about 35 accepted genera. About half of these consist of a single species, but about 400 of the species are in the type genus Saxifraga. The family is predominantly distributed in the northern hemisphere, but also in the Andes in South America.

<span class="mw-page-title-main">Eudicots</span> Clade of flowering plants

The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves (cotyledons) upon germination. The term derives from Dicotyledons.

<span class="mw-page-title-main">Rosids</span> Large clade of flowering plants

The rosids are members of a large clade of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms.

<span class="mw-page-title-main">Peridiscaceae</span> Family of flowering plants in the order Saxifragales

Peridiscaceae is a family of flowering plants in the order Saxifragales. Four genera comprise this family: Medusandra, Soyauxia, Peridiscus, and Whittonia., with a total of 12 known species. It has a disjunct distribution, with Peridiscus occurring in Venezuela and northern Brazil, Whittonia in Guyana, Medusandra in Cameroon, and Soyauxia in tropical West Africa. Whittonia is possibly extinct, being known from only one specimen collected below Kaieteur Falls in Guyana. In 2006, archeologists attempted to rediscover it, however, it proved unsuccessful.

Dipentodon is a genus of flowering plants in the family Dipentodontaceae. Its only species, Dipentodon sinicus, is a small, deciduous tree native to southern China, northern Myanmar, and northern India. It has been little studied and until recently its affinities remained obscure.

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

Huerteales is the botanical name for an order of flowering plants. It is one of the 17 orders that make up the large eudicot group known as the rosids in the APG III system of plant classification. Within the rosids, it is one of the orders in Malvidae, a group formerly known as eurosids II and now known informally as the malvids. This is true whether Malvidae is circumscribed broadly to include eight orders as in APG III, or more narrowly to include only four orders. Huerteales consists of four small families, Petenaeaceae, Gerrardinaceae, Tapisciaceae, and Dipentodontaceae.

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

Lepidobotryaceae is a family of plants in the order Celastrales. It contains only two species: Lepidobotrys staudtii and Ruptiliocarpon caracolito.

<span class="mw-page-title-main">Mesangiospermae</span> One of two clades of flowering plants

Mesangiospermae is a clade of flowering plants (angiosperms), informally called "mesangiosperms". They are one of two main groups of angiosperms. It is a name created under the rules of the PhyloCode system of phylogenetic nomenclature. There are about 350,000 species of mesangiosperms. The mesangiosperms contain about 99.95% of the flowering plants, assuming that there are about 175 species not in this group and about 350,000 that are. While such a clade with a similar circumscription exists in the APG III system, it was not given a name.

When the APG II system of plant classification was published in April 2003, fifteen genera and three families were placed incertae sedis in the angiosperms, and were listed in a section of the appendix entitled "Taxa of uncertain position".

<span class="mw-page-title-main">Superrosids</span> Clade of flowering plants

The superrosids are members of a large clade of flowering plants, containing more than 88,000 species, and thus more than a quarter of all angiosperms.

<i>Berberidopsis beckleri</i> Species of flowering plant

Berberidopsis beckleri is a species of climbing plant found in cool rainforests in eastern Australia. Its common name is the montane tape vine. Ferdinand von Mueller described the plant as Streptothamnus beckleri from collections at the Clarence River.

<span class="mw-page-title-main">Superasterids</span> Clade of flowering plants

The superasterids are members of a large clade of flowering plants, containing more than 122,000 species.

References

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  3. ↑ Hoot, S. B; Magallón, S.; Crane, P. R. (1999). "Phylogeny of basal eudicots based on three molecular data sets: atpB, rbcL, and 18S nuclear ribosomal DNA sequences". Annals of the Missouri Botanical Garden. 86 (86): 1–32. doi:10.2307/2666215. JSTOR   2666215. Archived from the original on 2023-10-29. Retrieved 2022-11-14.
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  8. 1 2 3 4 5 Cantino, P.D.; Doyle, J.A.; Graham, S.W.; Judd, W. S.; Olmstead, R.G.; Soltis, D. E.; Soltis, P.S.; Donoghue, M.J. (2007). "Towards a phylogenetic nomenclature of Tracheophyta". Taxon. 56 (3): 822–84. doi:10.2307/25065864. JSTOR   25065864.
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  14. ↑ Soltis, D.E.; Soltis, P.S.; Chase, M.W.; Mort, M.E.; Albach, D.C.; Zanis, M.; Savolainen, V.; Hahn, W.H.; Hoot, S.B.; Fay, M.F. (2000). "Angiosperm phylogeny inferred from 18 S rDNA, rbcL, and atpB sequences". Botanical Journal of the Linnean Society. 133 (4): 381–461. doi:10.1006/bojl.2000.0380.
  15. ↑ Baillon, H. (1870). "Histoire des Plantes II". Hachette (in French). Paris: Elaeagnacées: 487–495.
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  18. ↑ Ronse De Craene, Louis P. (2007). "Are Petals Sterile Stamens or Bracts? The Origin and Evolution of Petals in the Core Eudicots". Ann Bot. 100 (3): 621–630. doi:10.1093/aob/mcm076. PMC   2533615 . PMID   17513305. Archived from the original on 2022-11-07. Retrieved 2022-11-07.
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  20. 1 2 3 4 5 6 Moore, M.J.; Soltis, Pamela S.; Bell, Charles D.; Burleigh, J. Gordon; Soltis, Douglas E. (2010). "Phylogenetic analysis of 83 plastid genes further resolves the early diversification of eudicots". PNAS. 107 (10): 4623–4628. Bibcode:2010PNAS..107.4623M. doi: 10.1073/pnas.0907801107 . PMC   2842043 . PMID   20176954.
  21. ↑ Malécot, V.; Nickrent, D. L. (2008). "Molecular phylogenetic relationships of Olacaceae and related Santalales". Systematic Botany. 33 (33): 97–106. doi:10.1600/036364408783887384. S2CID   62879334.
  22. ↑ Moore M., Bell C., Soltis P. S., Soltis D. E.. 2008. Analysis of an 83-gene, 86-taxon plastid genome data set resolves relationships among several deep-level eudicot lineages Archived 2022-11-10 at the Wayback Machine . From Botany 2008: Botany without Borders, annual meeting of the Botanical Society of America, Vancouver, British Columbia, Canada, 97.
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  25. ↑ Vidal-Russell, R.; Nickrent, D. L. (2008). "The first mistletoes: Origins of aerial parasitism in Santalales". Molecular Phylogenetics and Evolution. 47 (2): 523–537. doi:10.1016/j.ympev.2008.01.016. PMID   18378469. Archived from the original on 2023-11-25. Retrieved 2022-11-07.
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  27. 1 2 Wang, Hengchang; Moore, Michael J.; Soltis, Pamela S.; Bell, Charles D.; Brockington, Samuel F.; Roolse, Alexandre; Davis, Charles C.; Latvis, Maribeth; Manchester, Steven R.; Soltis, Douglas E. (March 10, 2009). "Rosid radiation and the rapid rise of angiosperm-dominated forests". Proceedings of the National Academy of Sciences. 106 (10): 3853–3858. Bibcode:2009PNAS..106.3853W. doi: 10.1073/pnas.0813376106 . PMC   2644257 . PMID   19223592.
  28. 1 2 Scotland, Robert W.; Wortley, Alexandra H. (2003). "How many species of seed plants are there?". Taxon. 52 (1): 101–104. doi:10.2307/3647306. JSTOR   3647306.
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  30. ↑ Leebens-Mack, Jim; Raubeson, Linda A.; Cui, Liying; Kuehl, Jennifer V.; Fourcade, Matthew H.; Chumley, Timothy W.; Boore, Jeffrey L.; Jansen, Robert K.; de Pamphilis, Claude W. (2005). "Identifying the Basal Angiosperm Node in Chloroplast Genome Phylogenies: Sampling One's Way Out of the Felsenstein Zone". Mol Biol Evol. 22 (10): 1948–1963. doi: 10.1093/molbev/msi191 . PMID   15944438.
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  33. ↑ Takhtajan, A.L. (1997). Diversity and Classification of Flowering Plants. New York: Columbia Univ Press.
  34. ↑ Nandi, O.I.; Chase, M.W.; Endress, P.K. (1998). "A combined cladistic analysis of angiosperms using rbcL and nonmolecular data sets". Ann Mo Bot Gard. 85 (1): 137–212. doi:10.2307/2992003. JSTOR   2992003. Archived from the original on 2022-06-23. Retrieved 2022-11-14.
  35. ↑ Stevens PF. 2011 Phylogeny, version 9. Accessed September 1, 2011.
  36. ↑ Cole, Theodor C. H.; Hilger, Hartmut H. (2013). "Angiosperm Phylogeny" (PDF). Flowering Plant Systematics. Archived from the original (PDF) on 2017-05-17. Retrieved 2022-11-07.
  37. ↑ Moore, Michael J.; Hassan, Nasr; Gitzendanner, Matthew A.; Bruenn, Riva A; Croley, Matthew; Vandeventer, Alexia; Horn, James W.; Dhingra, Amit; Brockington, Samuel F.; Latvis, Maribeth; Ramdial, Jeremy; Alexandre, Roolse; Piedrahita, Ana; Soltis, Pamela S.; Soltis, Douglas E. (May 2011). "Analysis of the Plastid Inverted Repeat for 244 Species: Insights into Deeper-Level Angiosperm Relationships from a Long, Slowly Evolving Sequence Region". International Journal of Plant Sciences. 172 (4): 541–558. doi:10.1086/658923. S2CID   39521750.
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