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In mathematics and
phylogenetics In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
, Newick tree format (or Newick notation or New Hampshire tree format) is a way of representing graph-theoretical trees with edge lengths using parentheses and commas. It was adopted by James Archie, William H. E. Day, Joseph Felsenstein,
Wayne Maddison Wayne Paul Maddison (born 1958) is a Canadian evolutionary biologist, arachnologist, and biological illustrator. He is Canada Research Chair in Biodiversity and a professor at the departments of zoology and botany at the University of British ...
, Christopher Meacham, F. James Rohlf, and David Swofford, at two meetings in 1986, the second of which was at Newick's restaurant in
Dover Dover ( ) is a town and major ferry port in Kent, southeast England. It faces France across the Strait of Dover, the narrowest part of the English Channel at from Cap Gris Nez in France. It lies southeast of Canterbury and east of Maidstone. ...
, New Hampshire, US. The adopted format is a generalization of the format developed by Meacham in 1984 for the first tree-drawing programs in Felsenstein's PHYLIP package.


Examples

The following tree: could be represented in Newick format in several ways ((,)); ''no nodes are named'' (A,B,(C,D)); ''leaf nodes are named'' (A,B,(C,D)E)F; ''all nodes are named'' (:0.1,:0.2,(:0.3,:0.4):0.5); ''all but root node have a distance to parent'' (:0.1,:0.2,(:0.3,:0.4):0.5):0.0; ''all have a distance to parent'' (A:0.1,B:0.2,(C:0.3,D:0.4):0.5); ''distances and leaf names'' (popular) (A:0.1,B:0.2,(C:0.3,D:0.4)E:0.5)F; ''distances and all names'' ((B:0.2,(C:0.3,D:0.4)E:0.5)F:0.1)A; ''a tree rooted on a leaf node'' (rare) Newick format is typically used for tools like PHYLIP and is a minimal definition for a
phylogenetic tree A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...
.


Rooted, unrooted, and binary trees

When an ''unrooted'' tree is represented in Newick notation, an arbitrary node is chosen as its root. Whether rooted or unrooted, typically a tree's representation is rooted on an internal node and it is rare (but legal) to root a tree on a leaf node. A '' rooted binary tree'' that is rooted on an internal node has exactly two immediate descendant nodes for each internal node. An ''unrooted binary'' tree that is rooted on an arbitrary internal node has exactly three immediate descendant nodes for the root node, and each other internal node has exactly two immediate descendant nodes. A ''binary tree rooted from a leaf'' has at most one immediate descendant node for the root node, and each internal node has exactly two immediate descendant nodes.


Grammar

A grammar for parsing the Newick format (roughly based on ):


The grammar nodes

Tree: The full input Newick Format for a single tree Subtree: an internal node (and its descendants) or a leaf node Leaf: a node with no descendants Internal: a node and its one or more descendants BranchSet: a set of one or more Branches Branch: a tree edge and its descendant subtree. Name: the name of a node Length: the length of a tree edge.


The grammar rules

Note, ", " separates alternatives. Tree → Subtree ";" Subtree → Leaf , Internal Leaf → Name Internal → "(" BranchSet ")" Name BranchSet → Branch , Branch "," BranchSet Branch → Subtree Length Name → ''empty'' , ''string'' Length → ''empty'' , ":" ''number'' Whitespace (spaces, tabs, carriage returns, and linefeeds) within ''number'' is prohibited. Whitespace within ''string'' is often prohibited. Whitespace elsewhere is ignored. Sometimes the Name ''string'' must be of a specified fixed length; otherwise the punctuation characters from the grammar (semicolon, parentheses, comma, and colon) are prohibited. The Tree → Subtree ";" production is instead the Tree → Branch ";" production in those cases where having the entire tree descended from nowhere is permitted; this captures the replaced production as well because Length can be ''empty''. Note that when a tree having more than one leaf is rooted from one of its leaves, a representation that is rarely seen in practice, the root leaf is characterized as an Internal node by the above grammar. Generally, a ''root node'' labeled as Internal should be construed as actually internal if and only if it has at least two Branches in its BranchSet. One can make a grammar that formalizes this distinction by replacing the above Tree production rule with Tree → RootLeaf ";" , RootInternal ";" RootLeaf → Name , "(" Branch ")" Name RootInternal → "(" Branch "," BranchSet ")" Name The first RootLeaf production is for a tree with exactly one leaf. The second RootLeaf production is for rooting a tree from one of its two or more leaves.


Notes

* An unquoted ' may not contain blanks, parentheses, square brackets, single_quotes, colons, semicolons, or commas. Underscore characters in unquoted ''s'' are converted to blanks. * A ' may also be quoted by enclosing it in single quotes. Single quotes in the original string are represented as two consecutive single quote characters. * Whitespace may appear anywhere except within an unquoted ' or a * Newlines may appear anywhere except within a ' or a . * Comments are enclosed in square brackets. They can appear anywhere newlines are allowed. Comments starting with are generally computer-generated for additional data. Some dialects allow nested comments.


Dialects


New Hampshire X format

The New Hampshire X (NHX) format is an extension to Newick that adds key-value data (gene duplication, etc.) to Newick nodes. This is done by putting the additional data in square brackets &NHX:''key''=''value'':.../code> in the node labels. The brackets are used because they represent comments in the
Nexus file The extensible NEXUS file format is widely used in phylogenetics, evolutionary biology, and bioinformatics. It stores information about taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more population ...
format, so any parser not understanding these additional information will ignore them.


Extended Newick

While the standard Newick notation is limited to phylogenetic trees, ''Extended Newick'' (Perl Bio::PhyloNetwork) can be used to encode explicit phylogenetic networks. In a
phylogenetic network A phylogenetic network is any graph used to visualize evolutionary relationships (either abstractly or explicitly) between nucleotide sequences, genes, chromosomes, genomes, or species. They are employed when reticulation events such as hybridi ...
, which is a generalization of a
phylogenetic tree A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...
, a node either represents a divergence event (
cladogenesis Cladogenesis is an evolutionary splitting of a parent species into two distinct species, forming a clade. This event usually occurs when a few organisms end up in new, often distant areas or when environmental changes cause several extinctions, ...
) or a reticulation event such as hybridization,
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introg ...
, horizontal (lateral) gene transfer or recombination. Nodes that represent a reticulation event are duplicated, annotated by introducing the # symbol into the Newick format, and numbered consecutively (using
integer An integer is the number zero (0), a positive natural number (1, 2, 3, ...), or the negation of a positive natural number (−1, −2, −3, ...). The negations or additive inverses of the positive natural numbers are referred to as negative in ...
values starting with 1). For example, if leaf Y is the product of hybridisation (x) between lineages leading to C and D in the tree above, : one can express this situation by defining two trees in standard Newick notation (A,B,((C,Y)c,D)e)f; ''and'' (A,B,(C,(Y,D)d)e)f; ''standard Newick'', ''all nodes are named (internal nodes lowercase, leaves upper case)'' or in extended Newick notation (A,B,((C,(Y)x#H1)c,(x#H1,D)d)e)f; ''extended Newick, all nodes are named; 1 is the integer identifying the hybrid node x'' The here is a hybrid node. It will be joined by the program into a single node when drawn. This is the picture drawn by Dendroscope for this example: : The production rules above is modified by the following for labelling hybrid nodes (in general, nodes representing reticulation events): Leaf → Name Hybrid Hybrid → ''empty'' , "#" Type ''integer'' -- The #i part is an obligatory identifier for a hybrid node Type → ''empty'' , ''string'' -- type of reticulation, e.g., H = hybridisation, LGT = lateral gene transfer, R = recombination. In the visualization of LGT events, for a given reticulate node, one incoming edge is usually drawn as an "acceptor" edge and all other incoming edges are drawn as "transfer" edges. Some programs (e.g. Dendroscope and
SplitsTree SplitsTree is a freeware program for inferring phylogenetic trees, phylogenetic networks, or, more generally, split graphs, from various types of data such as a sequence alignment, a distance matrix or a set of trees. Software SplitsTree impleme ...
) allow exactly one copy of the reticulate node to be labeled with to indicate that it corresponds to the acceptor edge. Extended Newick is backward-compatible: a hybrid node would simply be interpreted as a few strangely-named nodes for legacy parsers.


Rich Newick format

The Rich Newick format, also known as the
Rice Rice is a cereal grain and in its Domestication, domesticated form is the staple food of over half of the world's population, particularly in Asia and Africa. Rice is the seed of the grass species ''Oryza sativa'' (Asian rice)—or, much l ...
Newick format, is a further extension of Extended Newick. It adds support for: * Unrooted phylogenies. This is simply done by writing an unrooted tree as usual (i.e., pick an arbitrary root at a binary branch point) and prefixing to the string. , on the other hand, can be used to force a rooted tree. * Bootstrap values and probabilities. This is done by adding additional fields after the length; fields can be left empty as long as the colons are present. This may be backward-incompatible.


Ad hoc extensions

Some other programs, like NWX, uses comments starting with to encode additional information in an ad hoc manner: * MrBayes and BEAST add additional information like probability, length in years, standard deviation for values to the nodes. They also use .


Visualization

Many tools have been published to visualize Newick tree data. Specific examples include the ETE toolkit ("Environment for Tree Exploration") and T-REX. Phylogenetic software packages such as
SplitsTree SplitsTree is a freeware program for inferring phylogenetic trees, phylogenetic networks, or, more generally, split graphs, from various types of data such as a sequence alignment, a distance matrix or a set of trees. Software SplitsTree impleme ...
and the tree-viewer Dendroscope as well as the online tree viewing tool IcyTree can handle standard and extended Newick notation, while the phylogenetic network software ''PhyloNet'' makes use of both the Extended Newick and Rich Newick format.


See also

* phyloXML * T-REX (Webserver) allows handling phylogenetic trees and networks in the Newick format. * Smart Game Format is an application of Newick format and is widely used for recording board games.


References


External links


Miyamoto and Goodman's Phylogram of Eutherian Mammals
An example of a large phylogram with its Newick format representation.
Phylogenetic tree (newick) viewer
(By Huerta-Cepas et al. 2016) {{Use dmy dates, date=December 2024 Trees (data structures) Graph description languages Phylogenetics