
A ghost lineage is a hypothesized ancestor in a species
lineage that has left no
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
evidence, but can still be inferred to exist or have existed because of gaps in the fossil record or genomic evidence.
The process of determining a ghost lineage relies on fossilized evidence before and after the hypothetical existence of the lineage and extrapolating relationships between organisms based on
phylogenetic analysis
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 data ...
.
Ghost lineages assume unseen diversity in the fossil record and serve as predictions for what the fossil record could eventually yield; these hypotheses can be tested by unearthing new fossils or running phylogenetic analyses.
Ghost lineages and
Lazarus taxa are related concepts, as both stem from gaps in the fossil record.
A ghost lineage is any gap in a
taxon
In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
's fossil record, with or without reappearance, while a Lazarus taxon is a type of ghost lineage wherein a species is believed to have gone extinct due to an absence of it in the fossil record, but then reappears after a period of time.
Examples of Lazarus taxa include the famous
coelacanths, as well as the
Philippine naked-backed fruit bat.
Name
In 1992, an article stated: "These additional entities are taxa
roupsthat are predicted to occur by the internal branching structure of
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 ...
s.... I refer to these as ghost lineages because they are invisible to the fossil record."
Phylogenetic trees constructed based on fossil records and
Darwin's theory of evolution often give an indication that species with similar phenotypes existed, although its fossil has not been discovered.
It is important to note that ghost lineages and ghost taxa are not the same. A ghost lineage is a one direct connection between the descendant and the ancestor, whereas a ghost taxon has many split descendants.
Examples

When looking back at extinct organisms, there are some groups of organisms (or lineages) that have gaps in their fossil records. These organisms or species may be closely related to one another, but there are no traces in the fossil records or sediment beds that might shed some light on their origins. Biologists may infer the existence of ghost lineages by examining sequential
stratigraphic
Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks.
Stratigraphy has three related subfields: lithost ...
units in the fossil record.
Fossils can then be mapped onto
cladogram
A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
s and range charts to assess which lineages are missing in the fossil record.
A classic example is the
coelacanth
Coelacanths ( ) are an ancient group of lobe-finned fish (Sarcopterygii) in the class Actinistia. As sarcopterygians, they are more closely related to lungfish and tetrapods (the terrestrial vertebrates including living amphibians, reptiles, bi ...
, a type of fish related to the
lungfish
Lungfish are freshwater vertebrates belonging to the class Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, inc ...
es and to primitive
tetrapod
A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s. Coelacanths have been around for the past 80 million years but have failed to leave any fossils. The reason for this is their environment, which is deep water near volcanic islands; therefore, these sediments are hard to get to, giving these coelacanths an 80 million year gap or ghost lineage.
Another ghost lineage was that of the
averostran
theropods, a ghost lineage now reduced considerably due to the discovery of ''
Tachiraptor''.
Duration and diversification
The duration between distinct fossils can be measured and used to derive information about the evolutionary history of a species. A study conducted in 1998 showed that a correlation exists between the diversification of a species and the duration of its ghost lineage; namely that a shorter ghost lineage implies that there will be greater species diversification.
Genetic evidence
Genetic evidence has revealed
ghost populations in many species, including modern bonobos and chimpanzees,
allopolyploid frogs, polyploid
parthenogenetic
Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
crayfish, a variety of plants, and humans.
A study comparing the genomes of 69 modern bonobos and chimpanzees found between 0.94.2% of gene flow from ancient bonobos and an archaic great ape lineage to modern bonobos, allowing researchers to reconstruct 4.8% of this ghost population's genome.
Furthermore, previous models for European ancestry suggested that European populations descended from two ancient populations, but genetic evidence now suggests that a third ghost population, the
Ancient North Eurasians, has also contributed to European ancestry.
See also
*
Lazarus taxon
In paleontology, a Lazarus taxon (plural ''taxa'') is a taxon that disappears for one or more periods from the fossil record, only to appear again either in later fossil records, or as actual living organisms, and often in isolated, obscure, ...
References
{{Phylogenetics
Phylogenetics