In
biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditar ...
, phylogenetics (; from
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
φυλή/
φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups of organisms. These relationships are determined by
Computational phylogenetics, phylogenetic inference methods that focus on observed
heritable
Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic inf ...
traits, such as
DNA sequences,
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respon ...
amino acid
Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
sequences, or
morphology. The result of such an analysis is a
phylogenetic tree—a diagram containing a hypothesis of relationships that reflects the evolutionary history of a group of organisms.
The tips of a phylogenetic tree can be living taxa or fossils, and represent the "end" or the present time in an evolutionary lineage. A phylogenetic diagram can be rooted or unrooted. A rooted tree diagram indicates the hypothetical common ancestor of the tree. An unrooted tree diagram (a network) makes no assumption about the ancestral line, and does not show the origin or "root" of the taxa in question or the direction of inferred evolutionary transformations.
In addition to their use for inferring phylogenetic patterns among taxa, phylogenetic analyses are often employed to represent relationships among genes or individual organisms. Such uses have become central to understanding biodiversity, evolution, ecology, and genomes.
Phylogenetics is part of
systematics
Biological systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: cladograms, phylogenetic t ...
.
Taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
is the identification, naming and
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood.
Classification is the grouping of related facts into classes.
It may also refer to:
Business, organizat ...
of organisms. Classifications are now usually based on phylogenetic data, and many systematists contend that only
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
taxa should be recognized as named groups. The degree to which classification depends on inferred evolutionary history differs depending on the school of taxonomy:
phenetics
In biology, phenetics ( el, phainein – to appear) , also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary r ...
ignores phylogenetic speculation altogether, trying to represent the similarity between organisms instead;
cladistics
Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived ch ...
(phylogenetic systematics) tries to reflect phylogeny in its classifications by only recognizing groups based on shared, derived characters (
synapomorphies
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
);
evolutionary taxonomy
Evolutionary taxonomy, evolutionary systematics or Darwinian classification is a branch of biological classification that seeks to classify organisms using a combination of phylogenetic relationship (shared descent), progenitor-descendant relat ...
tries to take into account both the branching pattern and "degree of difference" to find a compromise between them.
Inference of a phylogenetic tree
Usual methods of
phylogenetic inference
Computational phylogenetics is the application of computational algorithms, methods, and programs to phylogenetic "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary hist ...
involve computational approaches implementing the Optimality criterion, optimality criteria and methods of Maximum parsimony (phylogenetics), parsimony,
maximum likelihood
In statistics, maximum likelihood estimation (MLE) is a method of estimating the parameters of an assumed probability distribution, given some observed data. This is achieved by maximizing a likelihood function so that, under the assumed sta ...
(ML), and
MCMC-based
Bayesian inference. All these depend upon an implicit or explicit
mathematical model
A mathematical model is a description of a system using mathematical concepts and language. The process of developing a mathematical model is termed mathematical modeling. Mathematical models are used in the natural sciences (such as physics, ...
describing the evolution of characters observed.
Phenetics
In biology, phenetics ( el, phainein – to appear) , also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary r ...
, popular in the mid-20th century but now largely obsolete, used
distance matrix
In mathematics, computer science and especially graph theory, a distance matrix is a square matrix
In mathematics, a square matrix is a matrix with the same number of rows and columns. An ''n''-by-''n'' matrix is known as a square matrix of orde ...
-based methods to construct trees based on overall similarity in
morphology or similar observable traits (i.e. in the
phenotype
In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological prop ...
or the
overall similarity of DNA, not the
DNA sequence
DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
), which was often assumed to approximate phylogenetic relationships.
Prior to 1950, phylogenetic inferences were generally presented as
narrative
A narrative, story, or tale is any account of a series of related events or experiences, whether nonfictional ( memoir, biography, news report, documentary, travelogue, etc.) or fictional (fairy tale, fable, legend, thriller
Thriller may r ...
scenarios. Such methods are often ambiguous and lack explicit criteria for evaluating alternative hypotheses.
History
The term "phylogeny" derives from the German , introduced by Haeckel in 1866, and the
Darwinian
Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations that ...
approach to classification became known as the "phyletic" approach.
Ernst Haeckel's recapitulation theory
During the late 19th century,
Ernst Haeckel
Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...
's
recapitulation theory, or "biogenetic fundamental law", was widely accepted. It was often expressed as "
ontogeny
Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
recapitulates phylogeny", i.e. the development of a single organism during its lifetime, from germ to adult, successively mirrors the adult stages of successive ancestors of the species to which it belongs. But this theory has long been rejected. Instead,
ontogeny evolves – the phylogenetic history of a species cannot be read directly from its ontogeny, as Haeckel thought would be possible, but characters from ontogeny can be (and have been) used as data for phylogenetic analyses; the more closely related two species are, the more
apomorphies their embryos share.
Timeline of key points

*14th century, ''lex parsimoniae'' (parsimony principle),
William of Ockam, English philosopher, theologian, and Franciscan friar, but the idea actually goes back to
Aristotle
Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical Greece, Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatet ...
, precursor concept
*1763, Bayesian probability, Rev. Thomas Bayes, precursor concept
*18th century, Pierre Simon (Marquis de Laplace), perhaps first to use ML (maximum likelihood), precursor concept
*1809, evolutionary theory, ''
Philosophie Zoologique
''Philosophie zoologique'' ("Zoological Philosophy, or Exposition with Regard to the Natural History of Animals") is an 1809 book by the French naturalist Jean-Baptiste Lamarck, in which he outlines his pre-Darwinian theory of evolution, part of ...
,''
Jean-Baptiste de Lamarck
Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolog ...
, precursor concept, foreshadowed in the 17th century and 18th century by Voltaire, Descartes, and Leibniz, with Leibniz even proposing evolutionary changes to account for observed gaps suggesting that many species had become extinct, others transformed, and different species that share common traits may have at one time been a single race, also foreshadowed by some early Greek philosophers such as
Anaximander in the 6th century BC and the atomists of the 5th century BC, who proposed rudimentary theories of evolution
*1837, Darwin's notebooks show an evolutionary tree
*1843, distinction between
homology and
analogy (the latter now referred to as
homoplasy
Homoplasy, in biology and phylogenetics, is the term used to describe a feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from homology, which is the term used to characterize ...
), Richard Owen, precursor concept
*1858, Paleontologist Heinrich Georg Bronn (1800–1862) published a hypothetical tree to illustrating the paleontological "arrival" of new, similar species following the extinction of an older species. Bronn did not propose a mechanism responsible for such phenomena, precursor concept.
*1858, elaboration of evolutionary theory, Darwin and Wallace, also in Origin of Species by Darwin the following year, precursor concept
*1866,
Ernst Haeckel
Ernst Heinrich Philipp August Haeckel (; 16 February 1834