Unplaced in APG II

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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". [1]

Contents

By the end of 2009, molecular phylogenetic analysis of DNA sequences had revealed the relationships of most of these taxa, and all but three of them had been placed in some group within the angiosperms. In October 2009, APG II was superseded by the APG III system. [2] In APG III, 11 of the genera listed above were placed in families, or else became families whose position within their orders was approximately or exactly known. The family Rafflesiaceae was placed in the order Malpighiales, close to Euphorbiaceae and possibly within it. Mitrastema became a monotypic family, Mitrastemonaceae. This family and Balanophoraceae were placed incertae sedis into orders, that is, their positions within these orders remained completely unknown. Metteniusa was found to belong to a supraordinal group known as the lamiids, which has not been satisfactorily divided into orders. Cynomorium was raised to familial status as Cynomoriaceae, and along with Apodanthaceae and Gumillea, remained unplaced in APG III. Five taxa were unplaced among the angiosperms in APG III because Nicobariodendron and Petenaea were added to the list.

Leptaulus

There is no apparent reason for the inclusion of Leptaulus in the list of unplaced taxa, other than the time lag between submission and publication. In 2001, in a phylogenetic study based on morphological and DNA data, Leptaulus was found to belong to a group of six genera that most authors now consider to be the family Cardiopteridaceae. [3] This was confirmed in a study of wood anatomy in 2008. [4] The genus is placed in the Cardiopteridaceae in the APG III system of 2009. [1] Before 2001, Leptaulus and the rest of Cardiopteridaceae had usually been placed in a broadly circumscribed Icacinaceae, which turned out to be polyphyletic.[ citation needed ]

Some botanists do not recognize Cardiopteridaceae as a family of six genera. Instead, they segregate Cardiopteris into a monogeneric Cardiopteridaceae sensu stricto and place the other five genera in the family Leptaulaceae. [5] The monophyly of Leptaulaceae has never been tested with molecular data.[ citation needed ]

Pottingeria

It had long been thought, at least by some, that the small Southeast Asian tree Pottingeria might belong in the order Celastrales. [6] In a phylogenetic study of that order in 2006, Pottingeria was found to be a member of the order, but not of any of its families. It was in an unresolved pentatomy consisting of Parnassiaceae, Pottingeria , Mortonia , the pair ( Quetzalia + Zinowiewia ), and the other genera of Celastraceae. [7] When the APG III system was published in October 2009, the Angiosperm Phylogeny Group expanded Celastraceae to include all members of the pentatomy mentioned above. [8]

Dipentodon

Dipentodon has one species Dipentodon sinicus. [9] It is native to southern China, Burma, and northern India. [10] In 2009, in a molecular phylogenetic study of the order Huerteales, it was shown that Dipentodon and Perrottetia belong together as the two genera of the family Dipentodontaceae. [11]

Medusandra and Soyauxia

In 2009, in a molecular phylogenetic study of Malpighiales, Kenneth Wurdack and Charles Davis sampled five genera and one family that had been unplaced in APG II. They placed some of these for the first time and confirmed the previous placement of others with strong statistical support. [12]

In their outgroup, they included four genera from Saxifragales. These were Daphniphyllum, Medusandra, Soyauxia , and Peridiscus . In their phylogeny, Medusandra and Soyauxia formed a strongly supported clade with Peridiscus, a member of the family Peridiscaceae, the most basal clade in Saxifragales. Wurdack and Davis recommended that Medusandra and Soyauxia both be transferred to Peridiscaceae. Thus the monogeneric family Medusandraceae is subsumed into Peridiscaceae. Soyauxia had been found to be close to Peridiscus in another study two years before. [13] Wurdack and Davis also found that the family Rafflesiaceae and the genera Aneulophus , Centroplacus , and Trichostephanus belong in the order Malpighiales. [12]

Aneulophus

Aneulophus consists of two species of woody plants from tropical West Africa. [14] Wurdack and Davis found the traditional placement of Aneulophus in Erythroxylaceae to be correct. [12] Its position within the family remains uncertain.[ citation needed ]

Erythroxylaceae is a family of four genera. Erythroxylum has about 230 species. Nectaropetalum has eight species and Pinacopodium has two. No one has yet produced a molecular phylogeny of the family.[ citation needed ]

Centroplacus

Centroplacus has a single species, Centroplacus glaucinus, a tree from West Africa. It was found to be close to Bhesa , a genus that had only recently been removed from Celastrales. [7] Bhesa was grouped with Centroplacus to become the second genus in Centroplacaceae. [12] Bhesa consists of five species of trees from India and Malesia.

Trichostephanus

Trichostephanus has two species, both in tropical West Africa. It had usually been assigned to Achariaceae, but it was found to be deeply embedded in Samydaceae. [12] [15] Many taxonomists do not recognize Samydaceae as a separate family from Salicaceae.[ citation needed ]

Rafflesiaceae

Several genera have been removed from Rafflesiaceae, so that it now consists of only three genera: Sapria , Rhizanthes , and Rafflesia . All of these are holoparasites and, as discussed below, finding their relationships by molecular phylogenetics has presented special challenges. Rafflesia and its relatives were the subject of several papers from 2004 to 2009, and as the world's largest flower, Rafflesia has attracted special interest. In 2009, Wurdack and Davis confirmed earlier work in which it was found that Rafflesiaceae is nested within Euphorbiaceae sensu stricto, a circumscription of Euphorbiaceae that excludes Phyllanthaceae, Picrodendraceae, Putranjivaceae, Pandaceae, and a few other very small groups that had been included in it until the 1990s. [16] In order to preserve Rafflesiaceae, Wurdack and Davis split Euphorbiaceae sensu stricto into Euphorbiaceae sensu strictissimo and Peraceae, a new family comprising Pera and four other genera. [12]

Parasites

Four of the unplaced genera, and all three of the unplaced families of APG II consist of achlorophyllous holoparasites. In these, the chloroplast genes that are usually used in phylogenetic studies of angiosperms have become nonfunctional pseudogenes. If these evolve rapidly, they may be saturated with repeated mutations at the same site and consequently not be useful for phylogenetic reconstruction.[ citation needed ]

The relationships of some parasitic taxa have been elucidated in studies of nuclear and mitochondrial DNA sequences. But these sequences sometimes produce artifactual topologies in the phylogenetic tree, because horizontal gene transfer often occurs between parasites and their hosts. [17]

Bdallophyton and Cytinus

The parasitic genera Bdallophyton and Cytinus have been found to be closely related and have been placed together as the family Cytinaceae. On the basis of mitochondrial DNA, Cytinaceae has been placed in Malvales, as sister to Muntingiaceae. [18]

Mitrastemon

The parasitic family Mitrastemonaceae has one genus, known either as Mitrastemon or Mitrastema. The genus name and the corresponding family name have been a source of much confusion. [19] A phylogeny based on mitochondrial genes places Mitrastemon in the order Ericales, but this result had only 76% maximum likelihood bootstrap support. [20]

Hoplestigma

Hoplestigma consists of two species of African trees, notable for their large leaves, up to 55 cm long and 25 cm wide. [21] It is usually placed by itself in the family Hoplestigmataceae which is thought to be related to Boraginaceae. [22] In 2014, a phylogeny of Boraginaceae was published in a scientific journal called Cladistics . [23] By comparing the DNA sequences of selected genes, the authors of that study showed that Hoplestigma is related to members of Boraginaceae subfamily Cordioideae, and they recommended that Hoplestigma be placed in that subfamily. Other authors have suggested that, while Hoplestigma is the closest relative of Cordioideae, it should perhaps not be placed within it. [24]

Metteniusa

Metteniusa consists of seven species of trees in Central America and northwestern South America. Ever since Hermann Karsten proposed the name Metteniusaceae in 1859, some authors have placed Metteniusa by itself, in that family. [25] Most authors, however, placed it in Icacinaceae until that family was shown to be polyphyletic in 2001. [3]

In 2007, in a comparison of DNA sequences for three genes, it was found that Metteniusa is one of the basal clades of the lamiids. The authors recommended that the family Metteniusaceae be recognized. [26] Nothing is yet known about relationships among the groups of basal lamiids. The groups in this polytomy include the order Garryales, the families Icacinaceae, Oncothecaceae, and Metteniusaceae, as well as some unplaced genera, including Apodytes , Emmotum , and Cassinopsis . [24]

No phylogenetic study has focused on the lamiids, but phylogenies have been inferred for the asterids, a group composed of Cornales, Ericales, the lamiids, and the campanulids. [27] [28]

Balanophoraceae

Balanophoraceae is a family of holoparasites with 44 species in 17 genera. [24] For a long time, Cynomorium was usually included in this family, but it is now known to be unrelated. [20] In 2005, Balanophoraceae was shown to be in the order Santalales, but its position within that order has not been determined. [29]

Two researchers in Taiwan announced on the internet in 2009 that they have results supporting the placement of Balanophoraceae in Santalales. [30] They have yet to publish anything in a scientific journal.[ citation needed ]

Cynomorium

Many names have been published in Cynomorium , [31] but there are probably only two species. [32] It is not closely related to anything else, so it is placed in the monogeneric family Cynomoriaceae. [20]

Attempts to find its closest relatives have demonstrated with special clarity that molecular phylogenetics is not a sure-fire, problem-free method of determining systematic relationships. One study placed it in Saxifragales, but not at any particular position within that order. [29] Doubts have been expressed about the results of this study. Another study placed Cynomorium in Rosales based on analysis of the two invert repeat regions of the chloroplast genome, which evolve at one fifth the rate of the two single copy regions. [33]

Gumillea

Gumillea has a single species, Gumillea auriculata, [34] and is known from only one specimen which was collected in the late 18th century in Peru. [35] It was named by Hipólito Ruiz López and José Antonio Pavón Jiménez. [36]

George Bentham and Joseph Hooker placed it in Cunoniaceae, [37] and this treatment was followed by Adolf Engler and most others. [34] The last comprehensive treatment of Cunoniaceae, however, excludes it from the family. [38] In 2009, Armen Takhtajan placed Gumillea in Simaroubaceae. [39] A 2007 article on Simaroubaceae contains a list of the genera in the family. Gumillea is not on that list, but the authors do not provide a list or section on excluded genera. [40]

Gumillea has also been called a synonym of Picramnia , [14] [41] but the ultimate source of this information is obscure and it is not mentioned in either of the recent treatments of Picramnia. [42] [43] It is worth noting that on their plate for Gumillea, Ruiz and Pavón showed 11 ovules or immature seeds that had been extracted from a 2-locular ovary. But the ovary in Picramnia has (sometimes 2), usually 3 to 4 locules and there are always two ovules in each locule. [43]

It might be possible to determine the affinities of Gumillea if DNA could be extracted from the existing specimen. DNA has been successfully amplified from specimens of similar age. [44] Any material used in such research, however, will never be replaced.

Apodanthaceae

The family Apodanthaceae comprises 22 to 30 species of endoparasitic herbs. These are distributed into three genera: Pilostyles , Apodanthes , and Berlinianche . [45] Attempts to determine the relationships of Apodanthaceae have produced only uncertain results and they have remained enigmatic, [20] [46] until the family was shown to be confidently placed in Cucurbitales [47]

Related Research Articles

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

The Cucurbitales are an order of flowering plants, included in the rosid group of dicotyledons. This order mostly belongs to tropical areas, with limited presence in subtropical and temperate regions. The order includes shrubs and trees, together with many herbs and climbers. One major characteristic of the Cucurbitales is the presence of unisexual flowers, mostly pentacyclic, with thick pointed petals. The pollination is usually performed by insects, but wind pollination is also present.

<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

Saxifragales is an order of angiosperms, or flowering plants, containing 15 botanical families and around 100 genera, with nearly 2,500 species. Of the 15 families, many are small, with eight of them being monotypic. The largest family is the Crassulaceae (stonecrops), a diverse group of mostly succulent plants, with about 35 genera. Saxifragales are found worldwide, primarily in temperate to subtropical zones, rarely being encountered growing wild in the tropics; however, many species are now cultivated throughout the world as knowledge of plant husbandry has improved. They can be found in a wide variety of environments, from deserts to fully aquatic habitats, with species adapted to alpine, forested or fully-aquatic habitats. Many are epiphytic or lithophytic, growing on exposed cliff faces, on trees or on rocks, and not requiring a highly organic or nutrient-dense substrate to thrive.

<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">Phyllanthaceae</span> Family of flowering plants

Phyllanthaceae is a family of flowering plants in the eudicot order Malpighiales. It is most closely related to the family Picrodendraceae.

<span class="mw-page-title-main">Asterids</span> Clade of eudicot angiosperms

In the APG IV system (2016) for the classification of flowering plants, the name asterids denotes a clade. Asterids is the largest group of flowering plants, with more than 80,000 species, about a third of the total flowering plant species. Well-known plants in this clade include the common daisy, forget-me-nots, nightshades, the common sunflower, petunias, yacon, morning glory, lettuce, sweet potato, coffee, lavender, lilac, olive, jasmine, honeysuckle, ash tree, teak, snapdragon, sesame, psyllium, garden sage, table herbs such as mint, basil, and rosemary, and rainforest trees such as Brazil nut.

<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.

<span class="mw-page-title-main">Boraginales</span> Order of flowering plants within the lammiid clade of eudicots

Boraginales is an order of flowering plants in the asterid clade, with a total of about 125 genera and 2,700 species. Different taxonomic treatments either include only a single family, the Boraginaceae, or divide it into up to eleven families. Its herbs, shrubs, trees and lianas (vines) have a worldwide distribution.

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

The Rafflesiaceae are a family of rare parasitic plants comprising 36 species in 3 genera found in the tropical forests of east and southeast Asia, including Rafflesia arnoldii, which has the largest flowers of all plants. The plants are endoparasites of vines in the genus Tetrastigma (Vitaceae) and lack stems, leaves, roots, and any photosynthetic tissue. They rely entirely on their host plants for both water and nutrients, and only then emerge as flowers from the roots or lower stems of the host plants.

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">Balanophoraceae</span> Family of flowering plants

The Balanophoraceae are a subtropical to tropical family of obligate parasitic flowering plants, notable for their unusual development and formerly obscure affinities. In the broadest circumscription, the family consists of 16 genera. Alternatively, three genera may be split off into the segregate family Mystropetalaceae.

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

Medusandra is a genus of flowering plants in the family Peridiscaceae. It has two species, Medusandra richardsiana and Medusandra mpomiana. M. richardsiana is the most common and well known. Both species are native to Cameroon and adjacent countries.

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

The Icacinaceae, also called the white pear family, are a family of flowering plants, consisting of trees, shrubs, and lianas, primarily of the tropics.

Metteniusa santanderensis is a species of flowering plants in the family Metteniusaceae. It was formerly placed in the family Cardiopteridaceae. It is endemic to Colombia.

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

Cardiopteridaceae is a eudicot family of flowering plants. It consists of about 43 species of trees, shrubs, and woody vines, mostly of the tropics, but with a few in temperate regions. It contains six genera, the largest of which is Citronella, with 21 species. The other genera are much smaller.

<i>Oncotheca</i> Genus of trees

Oncotheca is a genus of tree endemic to New Caledonia. There are two species, Oncotheca balansae and Oncotheca humboldtiana.

The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a further revision, the APG IV system.

<span class="mw-page-title-main">Metteniusaceae</span> Family of trees

Metteniusaceae are a family of flowering plants, the only family in the order Metteniusales. It consists of about 10 genera and 50 species of trees, shrubs, and lianas, primarily of the tropics. The family was formerly restricted to just Metteniusa, but it is now expanded with a number of genera that were formerly placed in the widely polyphyletic Icacinaceae.

<span class="mw-page-title-main">Pentapetalae</span> Group of eudicots known as core eudicots

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. 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.

References

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  40. ↑ Joshua W. Clayton, Edwino S. Fernando, Pamela S. Soltis, and Douglas E. Soltis. 2007. "Molecular phylogeny of the tree-of-heaven family (Simaroubaceae) based on chloroplast and nuclear markers". International Journal of Plant Sciences168(9):1325-1339.
  41. ↑ Lois Brako and James L. Zarucchi. 1993. "Catalogue of the flowering plants and gymnosperms of Peru". Monographs in Systematic Botany from the Missouri Botanical Garden. monograph number 45.
  42. ↑ Edwino S. Fernando and Christopher J. Quinn. 1995. "Picramniaceae, a new family, and a recircumscription of Simaroubaceae". Taxon44(2):177-181.
  43. 1 2 Klaus Kubitzki. 2007. "Picramniaceae" pages 301-303. In: Klaus Kubitski (editor). The Families and Genera of Vascular Plants volume IX. Springer-Verlag: Berlin;Heidelberg, Germany.
  44. ↑ Katarina Andreasen, Mariette Manktelow, and Sylvain G. Razafimandimbison. 2009. "Successful DNA amplification of a more than 200-year-old herbarium specimen: recovering genetic material from the Linnean era". Taxon58(3):959-962.
  45. ↑ Albert Blarer, Daniel L. Nickrent, and Peter K. Endress. 2004. "Comparative floral structure and systematics in Apodanthaceae (Rafflesiales)". Plant Systematics and Evolution245(1-2):119-142.
  46. ↑ Apodanthaceae at Parasitic Plant Connection website
  47. ↑ Filipowicz, N. & Renner, S.S. 2010. The worldwide holoparasitic Apodanthaceae confidently placed in the Cucurbitales by nuclear and mitochondrial gene trees. BMC Evolutionary Biology 10: 219
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