Transitional fossil
A transitional fossil is any fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

ized remains of a lifeform that exhibits characteristics of two distinct taxonomic groups. A transitional fossil is the fossil of an organism near the branching point where major individual lineages (clade
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

s) diverge
Genetic divergence
Genetic divergence is the process in which two or more populations of an ancestral species accumulate independent genetic changes through time, often after the populations have become reproductively isolated for some period of time...

. It will have characteristics typical of organisms on both sides of the split, but because of the incompleteness of the fossil record, there is usually no way to know exactly how close it is to the actual point of divergence.

Transitional fossils serve as a reminder that taxonomic divisions are human constructs that have been imposed in hindsight on a continuum of variation. Numerous examples exist, including those of humans and other primates, tetrapods and fish, and birds and dinosaurs. The phrase "missing link" has been used extensively in popular writings on human evolution to refer to a perceived gap in the hominid evolutionary record. It is most commonly used to refer to any new transitional fossil finds. Scientists, however, do not use the term as it is misleading and inaccurate.


These diagrams plot the set of Hominine
Homininae is a subfamily of Hominidae, which includes humans, gorillas and chimpanzees, and some extinct relatives; it comprises all those hominids, such as Australopithecus, that arose after the split from orangutans . Our family tree, which has 3 main branches leading to chimpanzees, humans and...

 species known to science as of a given year. Each species is plotted as a box showing the range of cranial capacities for specimens of that species, and the range of dates at which specimens appear in the fossil record. The sequence of diagrams shows how an apparent "missing link" or gap between species in the fossil record may become filled as more fossil discoveries are made.

In 1859, when Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

's On the Origin of Species was first published, the fossil record was poorly known, and Darwin described the lack of transitional fossils as "the most obvious and gravest objection which can be urged against my theory", but explained it by relating it to the extreme imperfection of the geological record. He noted the limited collections available at that time, but described the available information as showing patterns which followed from his theory of descent with modification
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 through natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....

. Indeed, Archaeopteryx
Archaeopteryx , sometimes referred to by its German name Urvogel , is a genus of theropod dinosaur that is closely related to birds. The name derives from the Ancient Greek meaning "ancient", and , meaning "feather" or "wing"...

was discovered just two years later, in 1861, and represents a classic transitional form between dinosaurs and birds. Many more transitional fossils have been discovered since then and it is now considered that there is abundant evidence of how all the classes
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...

 of vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

s are related, much of it in the form of transitional fossils.


The reconstruction of the evolution of the horse and its relatives assembled by Othniel Charles Marsh
Othniel Charles Marsh
Othniel Charles Marsh was an American paleontologist. Marsh was one of the preeminent scientists in the field; the discovery or description of dozens of news species and theories on the origins of birds are among his legacies.Born into a modest family, Marsh was able to afford higher education...

 from surviving fossils that form a single, consistently developing lineage with many "transitional" types, is often cited as a family tree. However, modern cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 gives a different, multi-stemmed shrublike picture, with multiple innovations and many dead ends. Other specimens cited as transitional forms include the "walking whale" Ambulocetus
Ambulocetus was an early cetacean that could walk as well as swim. It lived during early Eocene some 50-49 million years ago. It is a transitional fossil that shows how whales evolved from land-living mammals. The Ambulocetus fossils were found in Pakistan by anthropologist Johannes Thewissen...

, the recently-discovered lobe-finned fish Tiktaalik
Tiktaalik is a genus of extinct sarcopterygian from the late Devonian period, with many features akin to those of tetrapods . It is an example from several lines of ancient sarcopterygian "fish" developing adaptations to the oxygen-poor shallow-water habitats of its time, which led to the...

and various hominid
The Hominidae or include them .), as the term is used here, form a taxonomic family, including four extant genera: chimpanzees , gorillas , humans , and orangutans ....

s considered to be proto-human
Homo (genus)
Homo is the genus that includes modern humans and species closely related to them. The genus is estimated to be about 2.3 to 2.4 million years old, evolving from australopithecine ancestors with the appearance of Homo habilis....


A middle Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...

 precursor to seed plants from Belgium has been identified predating the earliest seed plants by about 20 million years. Runcaria, small and radially symmetrical, is an integumented megasporangium surrounded by a cupule. The megasporangium bears an unopened distal extension protruding above the mutlilobed integument. It is suspected that the extension was involved in anemophilous pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

. Runcaria sheds new light on the sequence of character acquisition leading to the seed. Runcaria has all of the qualities of seed plants except for a solid seed coat and a system to guide the pollen to the seed.

Limitations of the fossil record

Not every transitional form appears in the fossil record because the fossil record is nowhere near complete. Organisms are only rarely preserved as fossils in the best of circumstances and only a fraction of such fossils have ever been discovered. The paleontologist Donald Prothero noted that this is illustrated by the fact that the total number of species of all kinds known through the fossil record was less than 5% of the number of known living species, which suggests that the number of species known through fossils must be less than 1% of all the species that have ever lived.

The fossil record is very uneven, and with few exceptions are heavily slanted toward organisms with hard parts, leaving most groups of soft-bodied organisms
Soft-bodied organisms
The term "soft-bodied organism" is used to describe animals without skeleton, roughly corresponding to the group Vermes as proposed by Carl von Linné. All animals have muscles, but since muscles can only pull, never push, a number of animals have developed hard parts that the muscles can pull on,...

 with little to no fossil record. The groups considered to have a good fossil record, including a number of transitional fossils between traditional groups, are the vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

s, the echinoderm
Echinoderms are a phylum of marine animals. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone....

s, brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...

s and some groups of arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...


Evolutionary taxonomy and cladistics

In evolutionary taxonomy
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 and overall similarity. This type of taxonomy considers taxa rather than single species, so that...

, the prevailing form of taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 during much of the 20th century and still used in basal textbooks, taxa based on morphological similarity are often drawn as "bubbles" branching off from each other, forming evolutionary trees. Transitional forms, are seen as falling between the various groups in term of anatomy, and are placed at the borders of these.

With the establishment of cladistic methods
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

, relationships are now strictly expressed in so-called cladograms, illustrating the branching of the evolutionary lineages. The different so-called 'natural' or 'monophyletic' groups form nested units that do not overlap. Within cladistics there is thus no longer a transition between established groups, but a differentiation that occurs within groups, represented as a branching in the cladogram
A cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...

. In this context, transitional organisms can be conceptualized as representing early examples on the different branches of a cladogram, lying between a particular branching point and the "crown-group", i.e. the most-derived group, which is placed at the end of a lineage.

Transitional vs ancestral

A source of confusion is the concept that a transitional form between two different taxonomic group must be directly ancestral to one or both groups. This was exacerbated by the fact that one of the goals of evolutionary taxonomy was the attempt to identify taxa that were ancestral to other taxa. However, it is almost impossible to be sure that any form represented in the record is actually a direct ancestor of any other. In fact because evolution is a branching process that produces a complex bush pattern of related species rather than a linear process that produces a ladder like progression, and because of the incompleteness the fossil record, it is unlikely that any particular form represented in the fossil record is a direct ancestor of any other. Cladistics deemphasized the concept of one taxonmic group being an ancestor of another, and instead emphasizes the concept of identifying sister taxa that share a common ancestor with one another more recently than they do with other groups. There are a few exceptional cases, such as some marine plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...

 micro-fossils, where the fossil record is complete enough to suggest with confidence that certain fossils represent a population that was actually ancestral to a later population of a different species, but in general transitional fossils are considered to have features that illustrate the transitional anatomical features of actual common ancestors of different taxa rather than to be actual ancestors.

Comparison to 'intermediate' forms

The terms 'transitional' and 'intermediate' are for the most part used as synonyms; however, a distinction between the two can be made:
  • "Transitional" can be used for those forms that do not have a significant number of unique derived traits that the derived relative does not possess as well. In other words, a transitional organism is morphologically close to the actual common ancestor it shares with its more derived relative.
  • "Intermediate" can be used for those forms that do have a large number of uniquely derived traits not connected to its derived relative.

According to this definition, Archaeopteryx
Archaeopteryx , sometimes referred to by its German name Urvogel , is a genus of theropod dinosaur that is closely related to birds. The name derives from the Ancient Greek meaning "ancient", and , meaning "feather" or "wing"...

, which does not show any derived traits that more derived birds do not possess as well, is transitional. In contrast, the platypus
The platypus is a semi-aquatic mammal endemic to eastern Australia, including Tasmania. Together with the four species of echidna, it is one of the five extant species of monotremes, the only mammals that lay eggs instead of giving birth to live young...

 is intermediate because it retains certain reptilian traits no longer found in modern mammals and also possesses derived traits of a highly specialized aquatic animal.

Following this definition, all living organisms are in fact to be regarded as intermediate forms when they are compared to some other related life-form. Indeed there are many species alive today that can be considered to be intermediate between two or more groups.

Missing links

A popular term used to designate transitional forms is "missing links". The term tends to be used in the popular media, but is avoided in the scientific press as it relates to the links in the great chain of being
Great chain of being
The great chain of being , is a Christian concept detailing a strict, religious hierarchical structure of all matter and life, believed to have been decreed by the Christian God.-Divisions:...

, a static pre-evolutionary concept now abandoned. In reality, the discovery of more and more transitional fossils continues to add to knowledge of evolutionary transitions, making many of the "missing links" missing no more (see List of transitional fossils).

The term "missing links" was used by Charles Lyell
Charles Lyell
Sir Charles Lyell, 1st Baronet, Kt FRS was a British lawyer and the foremost geologist of his day. He is best known as the author of Principles of Geology, which popularised James Hutton's concepts of uniformitarianism – the idea that the earth was shaped by slow-moving forces still in operation...

 in a somewhat different way in his Elements of Geology of 1851, but was popularized in its present meaning by its appearance in Lyell's Geological Evidences of the Antiquity of Man of 1863, p. xi. By that time geologists had abandoned a literal Biblical account and it was generally thought that the end of the last glacial period marked the first appearance of humanity, a view Lyell's Elements presented. His Antiquity of Man drew on new findings to put the origin of human beings much further back in the deep geological past. Lyell's vivid writing fired the public imagination, inspiring Jules Verne
Jules Verne
Jules Gabriel Verne was a French author who pioneered the science fiction genre. He is best known for his novels Twenty Thousand Leagues Under the Sea , A Journey to the Center of the Earth , and Around the World in Eighty Days...

's Journey to the Center of the Earth
Journey to the Center of the Earth
A Journey to the Center of the Earth is a classic 1864 science fiction novel by Jules Verne. The story involves a German professor who believes there are volcanic tubes going toward the center of the Earth...

, and Louis Figuier
Louis Figuier
Louis Figuier was a French scientist and writer. He was the nephew of Pierre-Oscar Figuier and became Professor of chemistry at L'Ecole de...

's 1867 second edition of La Terre avant le déluge which included dramatic illustrations of savage men and women wearing animal skins and wielding stone axes, in place of the Garden of Eden
Garden of Eden
The Garden of Eden is in the Bible's Book of Genesis as being the place where the first man, Adam, and his wife, Eve, lived after they were created by God. Literally, the Bible speaks about a garden in Eden...

 shown in the 1863 edition.

The idea of a "missing link" between humans and so-called "lower" animals remains lodged in the public imagination. The concept has been fueled by the successive discoveries of Australopithecus africanus
Australopithecus africanus
Australopithecus africanus was an early hominid, an australopithecine, who lived between 2–3 million years ago in the Pliocene. In common with the older Australopithecus afarensis, A. africanus was slenderly built, or gracile, and was thought to have been a direct ancestor of modern humans. Fossil...

(Taung Child
Taung Child
The Taung Child — or Taung Baby — is the fossilized skull of a young Australopithecus africanus individual. It was discovered in 1924 by quarrymen working for the Northern Lime Company in Taung, South Africa...

), Australopithecus sediba
Australopithecus sediba
Australopithecus sediba is a species of Australopithecus of the early Pleistocene, identified based on fossil remains dated to about 2 million years ago....

, Homo erectus
Homo erectus
Homo erectus is an extinct species of hominid that lived from the end of the Pliocene epoch to the later Pleistocene, about . The species originated in Africa and spread as far as India, China and Java. There is still disagreement on the subject of the classification, ancestry, and progeny of H...

(Peking Man
Peking Man
Peking Man , Homo erectus pekinensis, is an example of Homo erectus. A group of fossil specimens was discovered in 1923-27 during excavations at Zhoukoudian near Beijing , China...

, Java Man
Java Man
Java Man is the name given to fossils discovered in 1891 at Trinil - Ngawi Regency on the banks of the Solo River in East Java, Indonesia, one of the first known specimens of Homo erectus...

, Turkana boy
Turkana Boy
Turkana Boy, also occasionally, Nariokotome Boy is the common name of fossil KNM-WT 15000, a nearly complete skeleton of a hominid who died in the early Pleistocene. This specimen is the most complete early human skeleton ever found. It is 1.5 million years old...

), and other Hominina
The more anthropomorphic primates of the Hominini tribe are placed in the Hominina subtribe. Referred to as hominans, they are characterized by the evolution of an increasingly erect bipedal locomotion. The only extant species is Homo sapiens...


Creationist arguments

Proponents of creationism
Creationism is the religious beliefthat humanity, life, the Earth, and the universe are the creation of a supernatural being, most often referring to the Abrahamic god. As science developed from the 18th century onwards, various views developed which aimed to reconcile science with the Genesis...

 have frequently made claims about the existence or implications of transitional fossils that paleontologists consider to be false, and in some cases deliberately misleading. Some of these claims include:
  • 'There are no transitional fossils.' This is a claim made by groups such as Answers in Genesis
    Answers in Genesis
    Answers in Genesis is a non-profit Christian apologetics ministry with a particular focus on supporting Young Earth creationism and a literal interpretation of the Book of Genesis. The organization has offices in the United Kingdom and the United States...

     and the Institute for Creation Research
    Institute for Creation Research
    The Institute for Creation Research is a Christian institution in Dallas, Texas that specializes in education, research, and media promotion of Creation Science and Biblical creationism. The ICR adopts the Bible as an inerrant and literal documentary of scientific and historical fact as well as...

    . Such claims may be based on a misunderstanding of the nature of what represents a transitional feature but are also explained as a tactic employed by creationists seeking to distort or discredit evolutionary theory and has been called a "favorite lie" of creationists. Some creationists dispute the lack of transitional forms.

  • 'No fossils are found with partially functional features.' Vestigial organs are common in whales (legs), flightless birds (wings), snakes (pelvis and lung), and numerous structures in humans (the coccyx
    The coccyx , commonly referred to as the tailbone, is the final segment of the vertebral column. Comprising three to five separate or fused vertebrae below the sacrum, it is attached to the sacrum by a fibrocartilaginous joint, the sacrococcygeal symphysis, which permits limited movement between...

    , plica semilunaris, and appendix
    Vermiform appendix
    The appendix is a blind-ended tube connected to the cecum , from which it develops embryologically. The cecum is a pouchlike structure of the colon...


  • Henry M. Morris
    Henry M. Morris
    Henry Madison Morris was an American young earth creationist and Christian apologist. He was one of the founders of the Creation Research Society and the Institute for Creation Research...

     and other creationists have claimed that evolution predicts a continuous gradation in the fossil record, and have misrepresented the expected partial record as having "systematic gaps". Due to the specialized and rare circumstances required for a biological structure to fossilize, only a very small percentage of all life-forms that ever have existed can be expected to be represented in discoveries and each represents only a snapshot of the process of evolution. The transition itself can only be illustrated and corroborated by transitional fossils, but it will never demonstrate an exact half-way point between clearly divergent forms.

  • The theory of punctuated equilibrium
    Punctuated equilibrium
    Punctuated equilibrium is a theory in evolutionary biology which proposes that most species will exhibit little net evolutionary change for most of their geological history, remaining in an extended state called stasis...

     developed by Stephen Jay Gould
    Stephen Jay Gould
    Stephen Jay Gould was an American paleontologist, evolutionary biologist, and historian of science. He was also one of the most influential and widely read writers of popular science of his generation....

     and Niles Eldredge
    Niles Eldredge
    Niles Eldredge is an American paleontologist, who, along with Stephen Jay Gould, proposed the theory of punctuated equilibrium in 1972.-Education:...

     and first presented in 1972 is often mistakenly drawn into the discussion of transitional fossils. This theory, however, pertains only to well-documented transitions within taxa or between closely related taxa over a geologically short period of time. These transitions, usually traceable in the same geological outcrop, often show small jumps in morphology between extended periods of morphological stability. To explain these jumps, Gould and Eldredge envisaged comparatively long periods of genetic stability separated by periods of rapid evolution. Gould made the following observation of creationist misuse of his work to deny the existence of transitional fossils:

External links

  • Transitional vertebrate fossils FAQ, at the TalkOrigins Archive
    TalkOrigins Archive
    The TalkOrigins Archive is a website that presents mainstream science perspectives on the antievolution claims of young-earth, old-earth, and "intelligent design" creationists...

  • A Smooth Fossil Transition: single celled Radiolarian
  • Transitional species in insect evolution (discusses the evolution of termite
    Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...

    s from cockroach
    Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...

    es, with particular focus on extant intermediate species).
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