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A phylogenetic tree , phylogeny or evolutionary tree is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time. In other words, it is a branching diagram or a tree showing the evolutionary relationships among various biological species or other entities based upon similarities and differences in their physical or genetic characteristics. In evolutionary biology, all life on Earth is theoretically part of a single phylogenetic tree, indicating common ancestry . Phylogenetics is the study of phylogenetic trees. The main challenge is to find a phylogenetic tree representing optimal evolutionary ancestry between a set of species or taxa. Computational phylogenetics (also phylogeny inference) focuses on the algorithms involved in finding optimal phylogenetic tree in the phylogenetic landscape.

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32-912: The Filozoa are a monophyletic grouping within the Opisthokonta . They include animals and their nearest unicellular relatives (those organisms which are more closely related to animals than to fungi or Mesomycetozoa ). Three groups are currently assigned to the clade Filozoa: From Latin filum meaning "thread" and Greek zōion meaning "animal". A phylogenetic tree of Filozoa and its most closely related clades: Fonticulida Nucleariida [REDACTED] BCG2 True Fungi [REDACTED] Aphelida BCG1 Rozella [REDACTED] Namako-37 Microsporidia [REDACTED] Ichthyosporea [REDACTED] Syssomonas Corallochytrium [REDACTED] Filasterea [REDACTED] Choanoflagellatea [REDACTED] Animalia [REDACTED] The ancestral opisthokont cell

64-506: A binary tree ), and an unrooted bifurcating tree takes the form of an unrooted binary tree , a free tree with exactly three neighbors at each internal node. In contrast, a rooted multifurcating tree may have more than two children at some nodes and an unrooted multifurcating tree may have more than three neighbors at some nodes. Both rooted and unrooted trees can be either labeled or unlabeled. A labeled tree has specific values assigned to its leaves, while an unlabeled tree, sometimes called

96-593: A clade – that is, a grouping of taxa which meets these criteria: Monophyly is contrasted with paraphyly and polyphyly as shown in the second diagram. A paraphyletic grouping meets 1. but not 2., thus consisting of the descendants of a common ancestor, excepting one or more monophyletic subgroups. A polyphyletic grouping meets neither criterion, and instead serves to characterize convergent relationships of biological features rather than genetic relationships – for example, night-active primates, fruit trees, or aquatic insects. As such, these characteristic features of

128-480: A rooted phylogenetic tree, each node with descendants represents the inferred most recent common ancestor of those descendants, and the edge lengths in some trees may be interpreted as time estimates. Each node is called a taxonomic unit. Internal nodes are generally called hypothetical taxonomic units, as they cannot be directly observed. Trees are useful in fields of biology such as bioinformatics , systematics , and phylogenetics . Unrooted trees illustrate only

160-409: A clear outgroup. Another method is midpoint rooting, or a tree can also be rooted by using a non-stationary substitution model . Unrooted trees illustrate the relatedness of the leaf nodes without making assumptions about ancestry. They do not require the ancestral root to be known or inferred. Unrooted trees can always be generated from rooted ones by simply omitting the root. By contrast, inferring

192-814: A combination of genes that come from different genomic sources (e.g., from mitochondrial or plastid vs. nuclear genomes), or genes that would be expected to evolve under different selective regimes, so that homoplasy (false homology ) would be unlikely to result from natural selection. When extinct species are included as terminal nodes in an analysis (rather than, for example, to constrain internal nodes), they are considered not to represent direct ancestors of any extant species. Extinct species do not typically contain high-quality DNA . The range of useful DNA materials has expanded with advances in extraction and sequencing technologies. Development of technologies able to infer sequences from smaller fragments, or from spatial patterns of DNA degradation products, would further expand

224-427: A function of the number of tips. For 10 tips, there are more than 34 × 10 6 {\displaystyle 34\times 10^{6}} possible bifurcating trees, and the number of multifurcating trees rises faster, with ca. 7 times as many of the latter as of the former. A dendrogram is a general name for a tree, whether phylogenetic or not, and hence also for the diagrammatic representation of

256-609: A more suitable metaphor than the tree . Indeed, phylogenetic corals are useful for portraying past and present life, and they have some advantages over trees ( anastomoses allowed, etc.). Phylogenetic trees composed with a nontrivial number of input sequences are constructed using computational phylogenetics methods. Distance-matrix methods such as neighbor-joining or UPGMA , which calculate genetic distance from multiple sequence alignments , are simplest to implement, but do not invoke an evolutionary model. Many sequence alignment methods such as ClustalW also create trees by using

288-536: A number of different formats, all of which must represent the nested structure of a tree. They may or may not encode branch lengths and other features. Standardized formats are critical for distributing and sharing trees without relying on graphics output that is hard to import into existing software. Commonly used formats are Although phylogenetic trees produced on the basis of sequenced genes or genomic data in different species can provide evolutionary insight, these analyses have important limitations. Most importantly,

320-441: A phylogenetic tree. A cladogram only represents a branching pattern; i.e., its branch lengths do not represent time or relative amount of character change, and its internal nodes do not represent ancestors. A phylogram is a phylogenetic tree that has branch lengths proportional to the amount of character change. A chronogram is a phylogenetic tree that explicitly represents time through its branch lengths. A Dahlgrenogram

352-457: A polyphyletic grouping are not inherited from a common ancestor, but evolved independently. Monophyletic groups are typically characterised by shared derived characteristics ( synapomorphies ), which distinguish organisms in the clade from other organisms. An equivalent term is holophyly . The word "mono-phyly" means "one-tribe" in Greek. These definitions have taken some time to be accepted. When

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384-445: A tree shape, defines a topology only. Some sequence-based trees built from a small genomic locus, such as Phylotree, feature internal nodes labeled with inferred ancestral haplotypes. The number of possible trees for a given number of leaf nodes depends on the specific type of tree, but there are always more labeled than unlabeled trees, more multifurcating than bifurcating trees, and more rooted than unrooted trees. The last distinction

416-604: Is a diagram representing a cross section of a phylogenetic tree. A phylogenetic network is not strictly speaking a tree, but rather a more general graph , or a directed acyclic graph in the case of rooted networks. They are used to overcome some of the limitations inherent to trees. A spindle diagram, or bubble diagram, is often called a romerogram, after its popularisation by the American palaeontologist Alfred Romer . It represents taxonomic diversity (horizontal width) against geological time (vertical axis) in order to reflect

448-481: Is assumed to have possessed slender filose (thread-like) projections or 'tentacles'. In some opisthokonts (Mesomycetozoa and Corallochytrium ) these were lost. They are retained in Filozoa, where they are simple and non-tapering, with a rigid core of actin bundles (contrasting with the flexible, tapering and branched filopodia of nucleariids and the branched rhizoids and hyphae of fungi). In choanoflagellates and in

480-506: Is most true of genetic material that is subject to lateral gene transfer and recombination , where different haplotype blocks can have different histories. In these types of analysis, the output tree of a phylogenetic analysis of a single gene is an estimate of the gene's phylogeny (i.e. a gene tree) and not the phylogeny of the taxa (i.e. species tree) from which these characters were sampled, though ideally, both should be very close. For this reason, serious phylogenetic studies generally use

512-539: Is the most biologically relevant; it arises because there are many places on an unrooted tree to put the root. For bifurcating labeled trees, the total number of rooted trees is: For bifurcating labeled trees, the total number of unrooted trees is: Among labeled bifurcating trees, the number of unrooted trees with n {\displaystyle n} leaves is equal to the number of rooted trees with n − 1 {\displaystyle n-1} leaves. The number of rooted trees grows quickly as

544-529: The book Elementary Geology , by Edward Hitchcock (first edition: 1840). Charles Darwin featured a diagrammatic evolutionary "tree" in his 1859 book On the Origin of Species . Over a century later, evolutionary biologists still use tree diagrams to depict evolution because such diagrams effectively convey the concept that speciation occurs through the adaptive and semirandom splitting of lineages. The term phylogenetic , or phylogeny , derives from

576-404: The cladistics school of thought became mainstream in the 1960s, several alternative definitions were in use. Indeed, taxonomists sometimes used terms without defining them, leading to confusion in the early literature, a confusion which persists. The first diagram shows a phylogenetic tree with two monophyletic groups. The several groups and subgroups are particularly situated as branches of

608-413: The fact that a monophyletic group includes organisms (e.g., genera, species) consisting of all the descendants of a unique common ancestor. Conversely, the term polyphyly , or polyphyletic , builds on the ancient Greek prefix πολύς ( polús ), meaning "many, a lot of", and refers to the fact that a polyphyletic group includes organisms arising from multiple ancestral sources. By comparison,

640-419: The most primitive animals, namely sponges , they aggregate into a filter-feeding collar (made from microvilli , that are also made from actin) around the cilium or flagellum ; this is thought to be an inheritance from their most recent common filozoan ancestor. Monophyletic In biological cladistics for the classification of organisms , monophyly is the condition of a taxonomic grouping being

672-439: The optimal tree using many of these techniques is NP-hard , so heuristic search and optimization methods are used in combination with tree-scoring functions to identify a reasonably good tree that fits the data. Tree-building methods can be assessed on the basis of several criteria: Tree-building techniques have also gained the attention of mathematicians. Trees can also be built using T-theory . Trees can be encoded in

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704-403: The parent of all other nodes in the tree. The root is therefore a node of degree 2, while other internal nodes have a minimum degree of 3 (where "degree" here refers to the total number of incoming and outgoing edges). The most common method for rooting trees is the use of an uncontroversial outgroup —close enough to allow inference from trait data or molecular sequencing, but far enough to be

736-405: The range of DNA considered useful. Phylogenetic trees can also be inferred from a range of other data types, including morphology, the presence or absence of particular types of genes, insertion and deletion events – and any other observation thought to contain an evolutionary signal. Phylogenetic networks are used when bifurcating trees are not suitable, due to these complications which suggest

768-473: The relatedness of the leaf nodes and do not require the ancestral root to be known or inferred. The idea of a tree of life arose from ancient notions of a ladder-like progression from lower into higher forms of life (such as in the Great Chain of Being ). Early representations of "branching" phylogenetic trees include a "paleontological chart" showing the geological relationships among plants and animals in

800-574: The root of an unrooted tree requires some means of identifying ancestry. This is normally done by including an outgroup in the input data so that the root is necessarily between the outgroup and the rest of the taxa in the tree, or by introducing additional assumptions about the relative rates of evolution on each branch, such as an application of the molecular clock hypothesis . Both rooted and unrooted trees can be either bifurcating or multifurcating. A rooted bifurcating tree has exactly two descendants arising from each interior node (that is, it forms

832-417: The simpler algorithms (i.e. those based on distance) of tree construction. Maximum parsimony is another simple method of estimating phylogenetic trees, but implies an implicit model of evolution (i.e. parsimony). More advanced methods use the optimality criterion of maximum likelihood , often within a Bayesian framework , and apply an explicit model of evolution to phylogenetic tree estimation. Identifying

864-630: The term paraphyly , or paraphyletic , uses the ancient Greek prefix παρά ( pará ), meaning "beside, near", and refers to the situation in which one or several monophyletic subgroups are left apart from all other descendants of a unique common ancestor. That is, a paraphyletic group is nearly monophyletic, hence the prefix pará . On the broadest scale, definitions fall into two groups. The concepts of monophyly, paraphyly , and polyphyly have been used in deducing key genes for barcoding of diverse group of species. Phylogenetic tree Phylogenetic trees may be rooted or unrooted. In

896-411: The tree before hybridisation takes place, and conserved sequences . Also, there are problems in basing an analysis on a single type of character, such as a single gene or protein or only on morphological analysis, because such trees constructed from another unrelated data source often differ from the first, and therefore great care is needed in inferring phylogenetic relationships among species. This

928-493: The tree to indicate ordered lineal relationships between all the organisms shown. Further, any group may (or may not) be considered a taxon by modern systematics , depending upon the selection of its members in relation to their common ancestor(s); see second and third diagrams. The term monophyly , or monophyletic , derives from the two Ancient Greek words μόνος ( mónos ), meaning "alone, only, unique", and φῦλον ( phûlon ), meaning "genus, species", and refers to

960-529: The trees that they generate are not necessarily correct – they do not necessarily accurately represent the evolutionary history of the included taxa. As with any scientific result, they are subject to falsification by further study (e.g., gathering of additional data, analyzing the existing data with improved methods). The data on which they are based may be noisy ; the analysis can be confounded by genetic recombination , horizontal gene transfer , hybridisation between species that were not nearest neighbors on

992-422: The two ancient greek words φῦλον ( phûlon ), meaning "race, lineage", and γένεσις ( génesis ), meaning "origin, source". A rooted phylogenetic tree (see two graphics at top) is a directed tree with a unique node — the root — corresponding to the (usually imputed ) most recent common ancestor of all the entities at the leaves of the tree. The root node does not have a parent node, but serves as

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1024-404: The variation of abundance of various taxa through time. A spindle diagram is not an evolutionary tree: the taxonomic spindles obscure the actual relationships of the parent taxon to the daughter taxon and have the disadvantage of involving the paraphyly of the parental group. This type of diagram is no longer used in the form originally proposed. Darwin also mentioned that the coral may be

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