Living fossil

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The coelacanths were thought to have gone extinct 66 million years ago, until a living specimen belonging to the order was discovered in 1938. Coelacanth CAS 1.JPG
The coelacanths were thought to have gone extinct 66  million years ago, until a living specimen belonging to the order was discovered in 1938.

A living fossil is an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of origin of the extant clade. Living fossils commonly are of species-poor lineages, but they need not be. While the body plan of a living fossil remains superficially similar, it is never the same species as the remote relatives it resembles, because genetic drift would inevitably change its chromosomal structure.

Contents

Living fossils exhibit stasis (also called "bradytely") over geologically long time scales. Popular literature may wrongly claim that a "living fossil" has undergone no significant evolution since fossil times, with practically no molecular evolution or morphological changes. Scientific investigations have repeatedly discredited such claims. [1] [2] [3]

The minimal superficial changes to living fossils are mistakenly declared as an absence of evolution, but they are examples of stabilizing selection, which is an evolutionary process—and perhaps the dominant process of morphological evolution. [4]

Characteristics

Fossil and living ginkgos
Gingkoites huttoni 1.jpg
170 million-year-old fossil Ginkgo leaves
Ginkgo Biloba Leaves - Black Background.jpg
Living Ginkgo biloba plant

Living fossils have two main characteristics, although some have a third:

  1. Living organisms that are members of a taxon that has remained recognisable in the fossil record over an unusually long time span.
  2. They show little morphological divergence, whether from early members of the lineage, or among extant species.
  3. They tend to have little taxonomic diversity. [5]

The first two are required for recognition as a living fossil; some authors also require the third, others merely note it as a frequent trait.

Such criteria are neither well-defined nor clearly quantifiable, but modern methods for analyzing evolutionary dynamics can document the distinctive tempo of stasis. [6] [7] [8] Lineages that exhibit stasis over very short time scales are not considered living fossils; what is poorly-defined is the time scale over which the morphology must persist for that lineage to be recognized as a living fossil.

The term living fossil is much misunderstood in popular media in particular, in which it often is used meaninglessly. In professional literature the expression seldom appears and must be used with far more caution, although it has been used inconsistently. [9] [10]

One example of a concept that could be confused with "living fossil" is that of a "Lazarus taxon", but the two are not equivalent; a Lazarus taxon (whether a single species or a group of related species) is one that suddenly reappears, either in the fossil record or in nature, as if the fossil had "come to life again". [11] In contrast to "Lazarus taxa", a living fossil in most senses is a species or lineage that has undergone exceptionally little change throughout a long fossil record, giving the impression that the extant taxon had remained identical through the entire fossil and modern period. Because of the mathematical inevitability of genetic drift, though, the DNA of the modern species is necessarily different from that of its distant, similar-looking ancestor. They almost certainly would not be able to cross-reproduce, and are not the same species. [12]

The average species turnover time, meaning the time between when a species first is established and when it finally disappears, varies widely among phyla, but averages about 2–3 million years.[ citation needed ] A living taxon that had long been thought to be extinct could be called a Lazarus taxon once it was discovered to be still extant. A dramatic example was the order Coelacanthiformes, of which the genus Latimeria was found to be extant in 1938. About that there is little debate – however, whether Latimeria resembles early members of its lineage sufficiently closely to be considered a living fossil as well as a Lazarus taxon has been denied by some authors in recent years. [1]

Coelacanths disappeared from the fossil record some 80 million years ago (in the upper Cretaceous period) and, to the extent that they exhibit low rates of morphological evolution, extant species qualify as living fossils. It must be emphasised that this criterion reflects fossil evidence, and is totally independent of whether the taxa had been subject to selection at all, which all living populations continuously are, whether they remain genetically unchanged or not. [13]

This apparent stasis, in turn, gives rise to a great deal of confusion – for one thing, the fossil record seldom preserves much more than the general morphology of a specimen. To determine much about its physiology is seldom possible; not even the most dramatic examples of living fossils can be expected to be without changes, no matter how persistently constant their fossils and the extant specimens might seem. To determine much about noncoding DNA is hardly ever possible, but even if a species were hypothetically unchanged in its physiology, it is to be expected from the very nature of the reproductive processes, that its non-functional genomic changes would continue at more-or-less standard rates. Hence, a fossil lineage with apparently constant morphology need not imply equally constant physiology, and certainly neither implies any cessation of the basic evolutionary processes such as natural selection, nor reduction in the usual rate of change of the noncoding DNA. [13]

Some living fossils are taxa that were known from palaeontological fossils before living representatives were discovered. The most famous examples of this are:

All the above include taxa that originally were described as fossils but now are known to include still-extant species.

Other examples of living fossils are single living species that have no close living relatives, but are survivors of large and widespread groups in the fossil record. For example:

All of these were described from fossils before later being found alive. [14] [15] [16]

The fact that a living fossil is a surviving representative of an archaic lineage does not imply that it must retain all the "primitive" features (plesiomorphies) of its ancestral lineage. Although it is common to say that living fossils exhibit "morphological stasis", stasis, in the scientific literature, does not mean that any species is strictly identical to its ancestor, much less remote ancestors.

Some living fossils are relicts of formerly diverse and morphologically varied lineages, but not all survivors of ancient lineages necessarily are regarded as living fossils. See for example the uniquely and highly autapomorphic oxpeckers, which appear to be the only survivors of an ancient lineage related to starlings and mockingbirds. [17]

Evolution and living fossils

The term living fossil is usually reserved for species or larger clades that are exceptional for their lack of morphological diversity and their exceptional conservatism, and several hypotheses could explain morphological stasis on a geologically long time-scale. Early analyses of evolutionary rates emphasized the persistence of a taxon rather than rates of evolutionary change. [18] Contemporary studies instead analyze rates and modes of phenotypic evolution, but most have focused on clades that are thought to be adaptive radiations rather than on those thought to be living fossils. Thus, very little is presently known about the evolutionary mechanisms that produce living fossils or how common they might be. Some recent studies have documented exceptionally low rates of ecological and phenotypic evolution despite rapid speciation. [19] This has been termed a "non-adaptive radiation" referring to diversification not accompanied by adaptation into various significantly different niches. [20] Such radiations are explanation for groups that are morphologically conservative. Persistent adaptation within an adaptive zone is a common explanation for morphological stasis. [21] The subject of very low evolutionary rates, however, has received much less attention in the recent literature than that of high rates.

Living fossils are not expected to exhibit exceptionally low rates of molecular evolution, and some studies have shown that they do not. [22] For example, on tadpole shrimp ( Triops ), one article notes, "Our work shows that organisms with conservative body plans are constantly radiating, and presumably, adapting to novel conditions... I would favor retiring the term 'living fossil' altogether, as it is generally misleading." [23] Some scientists instead prefer a new term stabilomorph, being defined as "an effect of a specific formula of adaptative strategy among organisms whose taxonomic status does not exceed genus-level. A high effectiveness of adaptation significantly reduces the need for differentiated phenotypic variants in response to environmental changes and provides for long-term evolutionary success." [24]

The question posed by several recent studies pointed out that the morphological conservatism of coelacanths is not supported by paleontological data. [25] [26] In addition, it was shown recently that studies concluding that a slow rate of molecular evolution is linked to morphological conservatism in coelacanths are biased by the a priori hypothesis that these species are 'living fossils'. [1] Accordingly, the genome stasis hypothesis is challenged by the recent finding that the genome of the two extant coelacanth species L. chalumnae and L. menadoensis contain multiple species-specific insertions, indicating transposable element recent activity and contribution to post-speciation genome divergence. [27] Such studies, however, challenge only a genome stasis hypothesis, not the hypothesis of exceptionally low rates of phenotypic evolution.

History

The term was coined by Charles Darwin in his On the Origin of Species from 1859, when discussing Ornithorhynchus (the platypus) and Lepidosiren (the South American lungfish):

All fresh-water basins, taken together, make a small area compared with that of the sea or of the land; and, consequently, the competition between fresh-water productions will have been less severe than elsewhere; new forms will have been more slowly formed, and old forms more slowly exterminated. And it is in fresh water that we find seven genera of Ganoid fishes, remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world, as the Ornithorhynchus and Lepidosiren, which, like fossils, connect to a certain extent orders now widely separated in the natural scale. These anomalous forms may almost be called living fossils; they have endured to the present day, from having inhabited a confined area, and from having thus been exposed to less severe competition.

—  On the Origin of Species , 1859 [28]

Other definitions

Long-enduring

Elephant shrews resemble the extinct Leptictidium of Eocene Europe. Rhynchocyon petersi from side.jpg
Elephant shrews resemble the extinct Leptictidium of Eocene Europe.

A living taxon that lived through a large portion of geologic time.[ citation needed ]

The Australian lungfish (Neoceratodus fosteri), also known as the Queensland lungfish, is an example of an organism that meets this criterion. Fossils identical to modern specimens have been dated at over 100 million years old. Modern Queensland lungfish have existed as a species for almost 30 million years. The contemporary nurse shark has existed for more than 112 million years, making this species one of the oldest, if not actually the oldest extant vertebrate species.

Resembles ancient species

A living taxon morphologically and/or physiologically resembling a fossil taxon through a large portion of geologic time (morphological stasis). [29]

Retains many ancient traits

More primitive trapdoor spiders, such as this female Liphistius sp., have segmented plates on the dorsal surface of the abdomen and cephalothorax, a character shared with scorpions, making it probable that after the spiders diverged from the scorpions, the earliest unique ancestor of trapdoor species was the first to split off from the lineage that contains all other extant spiders. Liphistius.sp.small.jpg
More primitive trapdoor spiders, such as this female Liphistius sp., have segmented plates on the dorsal surface of the abdomen and cephalothorax, a character shared with scorpions, making it probable that after the spiders diverged from the scorpions, the earliest unique ancestor of trapdoor species was the first to split off from the lineage that contains all other extant spiders.

A living taxon with many characteristics believed to be primitive.[ citation needed ] This is a more neutral definition. However, it does not make it clear whether the taxon is truly old, or it simply has many plesiomorphies. Note that, as mentioned above, the converse may hold for true living fossil taxa; that is, they may possess a great many derived features (autapomorphies), and not be particularly "primitive" in appearance.

Relict population

Any one of the above three definitions, but also with a relict distribution in refuges.[ citation needed ]

Some paleontologists believe that living fossils with large distributions (such as Triops cancriformis ) are not real living fossils. In the case of Triops cancriformis (living from the Triassic until now), the Triassic specimens lost most of their appendages (mostly only carapaces remain), and they have not been thoroughly examined since 1938.

Low diversity

Any of the first three definitions, but the clade also has a low taxonomic diversity (low diversity lineages).[ citation needed ]

Oxpeckers are morphologically somewhat similar to starlings due to shared plesiomorphies, but are uniquely adapted to feed on parasites and blood of large land mammals, which has always obscured their relationships. This lineage forms part of a radiation that includes Sturnidae and Mimidae, but appears to be the most ancient of these groups. Biogeography strongly suggests that oxpeckers originated in eastern Asia and only later arrived in Africa, where they now have a relict distribution. [17]

The two living species thus seem to represent an entirely extinct and (as Passerida go) rather ancient lineage, as certainly as this can be said in the absence of actual fossils. The latter is probably due to the fact that the oxpecker lineage never occurred in areas where conditions were good for fossilization of small bird bones, but of course, fossils of ancestral oxpeckers may one day turn up enabling this theory to be tested.

Operational definition

An operational definition was proposed in 2017, where a 'living fossil' lineage has a slow rate of evolution and occurs close to the middle of morphological variation (the centroid of morphospace) among related taxa (i.e. a species is morphologically conservative among relatives). [30] The scientific accuracy of the morphometric analyses used to classify tuatara as a living fossil under this definition have been criticised however, [31] which prompted a rebuttal from the original authors. [32]

Examples

Some of these are informally known as "living fossils".

Ginkgos not only have existed for a long time, but also have a long life span, with some having an age of over 2,500 years. Six specimens survived the atomic bombing of Hiroshima, 1 to 2 kilometers from ground zero. They still live there today. Ginkgo Stamm.jpg
Ginkgos not only have existed for a long time, but also have a long life span, with some having an age of over 2,500 years. Six specimens survived the atomic bombing of Hiroshima, 1 to 2 kilometers from ground zero. They still live there today.
Ferns were the dominant plant group in the Jurassic period, with some species, such as Osmunda claytoniana, maintaining evolutionary stasis for at least 180 million years. OsmundaRegalis.jpg
Ferns were the dominant plant group in the Jurassic period, with some species, such as Osmunda claytoniana , maintaining evolutionary stasis for at least 180 million years.

Bacteria

Protists

Plants

Fungi

Animals

Echidnas are one of few mammals to lay eggs. Wild shortbeak echidna.jpg
Echidnas are one of few mammals to lay eggs.
Vertebrates
Hoatzin hatch with two visible claws on their wings, but the claws fall out once the birds reach maturity. Hoatzins in Ecuador.jpg
Hoatzin hatch with two visible claws on their wings, but the claws fall out once the birds reach maturity.
Crocodilians survived the K-Pg extinction event that killed off the non-avian dinosaurs. Crocodylus acutus jalisco mexico.jpg
Crocodilians survived the K–Pg extinction event that killed off the non-avian dinosaurs.
Tuataras are reptiles, yet retain more primitive characteristics than lizards and snakes. Tuatara.jpg
Tuataras are reptiles, yet retain more primitive characteristics than lizards and snakes.
The goblin shark is the only extant representative of the family Mitsukurinidae, a lineage some 125 million years old (early Cretaceous). Mitsukurina owstoni, Pengo.jpg
The goblin shark is the only extant representative of the family Mitsukurinidae, a lineage some 125 million years old (early Cretaceous).
Nautilus retain the external spiral shell that its other relatives have lost. Nautilus profile.jpg
Nautilus retain the external spiral shell that its other relatives have lost.
With little change over the last 450 million years, the horseshoe crabs appear as living fossils. Horseshoe Crab.jpg
With little change over the last 450 million years, the horseshoe crabs appear as living fossils.
Invertebrates

See also

Related Research Articles

<span class="mw-page-title-main">Coelacanth</span> Order of lobe-finned fishes

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 than to ray-finned fish.

<span class="mw-page-title-main">Osmundaceae</span> Family of ferns

Osmundaceae is a family of ferns containing four to six extant genera and 18–25 known species. It is the only living family of the order Osmundales in the class Polypodiopsida (ferns) or in some classifications the only order in the class Osmundopsida. This is an ancient and fairly isolated group that is often known as the "flowering ferns" because of the striking aspect of the ripe sporangia in Claytosmunda, Osmunda, Osmundastrum, and Plensium. In these genera the sporangia are borne naked on non-laminar pinnules, while Todea and Leptopteris bear sporangia naked on laminar pinnules. Ferns in this family are larger than most other ferns.

<span class="mw-page-title-main">Lungfish</span> A type of bony fish

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, including the presence of lobed fins with a well-developed internal skeleton. Lungfish represent the closest living relatives of the tetrapods. The mouths of lungfish typically bear tooth plates, which are used to crush hard shelled organisms.

<span class="mw-page-title-main">Sarcopterygii</span> Class of fishes

Sarcopterygii — sometimes considered synonymous with Crossopterygii — is a clade of bony fish commonly referred to as lobe-finned fish. They are characterised by prominent muscular limb buds (lobes) within their fins, which are supported by articulated appendicular skeletons. This is in contrast to the other clade of bony fish, the Actinopterygii, which have only skin-covered bony spines supporting the fins.

The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleotide sequences for DNA, RNA, or amino acid sequences for proteins.

<span class="mw-page-title-main">Squamata</span> Order of reptiles

Squamata is the largest order of reptiles, comprising lizards and snakes. With over 11,500 species, it is also the second-largest order of extant (living) vertebrates, after the perciform fish. Members of the order are distinguished by their skins, which bear horny scales or shields, and must periodically engage in molting. They also possess movable quadrate bones, making possible movement of the upper jaw relative to the neurocranium. This is particularly visible in snakes, which are able to open their mouths very wide to accommodate comparatively large prey. Squamates are the most variably sized living reptiles, ranging from the 16 mm (0.63 in) dwarf gecko to the 6.5 m (21 ft) reticulated python. The now-extinct mosasaurs reached lengths over 14 m (46 ft).

<span class="mw-page-title-main">Lazarus taxon</span> Taxon that disappears from the fossil record, only to reappear later

In paleontology, a Lazarus taxon is a taxon that disappears for one or more periods from the fossil record, only to appear again later. Likewise in conservation biology and ecology, it can refer to species or populations that were thought to be extinct, and are rediscovered. The term Lazarus taxon was coined by Karl W. Flessa and David Jablonski in 1983 and was then expanded by Jablonski in 1986. Paul Wignall and Michael Benton defined Lazarus taxa as, "At times of biotic crisis many taxa go extinct, but others only temporarily disappeared from the fossil record, often for intervals measured in millions of years, before reappearing unchanged". Earlier work also supports the concept though without using the name Lazarus taxon, like work by Christopher R. C. Paul.

<span class="mw-page-title-main">Pseudoextinction</span> Phenomenon where a species is perpetuated by a daughter species

Pseudoextinction of a species occurs when all members of the species are extinct, but members of a daughter species remain alive. The term pseudoextinction refers to the evolution of a species into a new form, with the resultant disappearance of the ancestral form. Pseudoextinction results in the relationship between ancestor and descendant still existing even though the ancestor species no longer exists.

<i>Crocodylus</i> Genus of reptiles

Crocodylus is a genus of true crocodiles in the family Crocodylidae.

Neontology is a part of biology that, in contrast to paleontology, deals with living organisms. It is the study of extant taxa : taxa with members still alive, as opposed to (all) being extinct. For example:

<span class="mw-page-title-main">West Indian Ocean coelacanth</span> Species of lobe-finned bony fish

The West Indian Ocean coelacanth is a crossopterygian, one of two extant species of coelacanth, a rare order of vertebrates more closely related to lungfish and tetrapods than to the common ray-finned fishes. The other extant species is the Indonesian coelacanth.

In phylogenetics, basal is the direction of the base of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to the root. Note that extant taxa that lie on branches connecting directly to the root are not more closely related to the root than any other extant taxa.

<span class="mw-page-title-main">Euphyllophyte</span> Clade of vascular plants

The euphyllophytes are a clade of plants within the tracheophytes. The group may be treated as an unranked clade, a division under the name Euphyllophyta or a subdivision under the name Euphyllophytina. The euphyllophytes are characterized by the possession of true leaves ("megaphylls"), and comprise one of two major lineages of extant vascular plants. As shown in the cladogram below, the euphyllophytes have a sister relationship to the lycopodiophytes or lycopsids. Unlike the lycopodiophytes, which consist of relatively few presently living or extant taxa, the euphyllophytes comprise the vast majority of vascular plant lineages that have evolved since both groups shared a common ancestor more than 400 million years ago. The euphyllophytes consist of two lineages, the spermatophytes or seed plants such as flowering plants (angiosperms) and gymnosperms, and the Polypodiophytes or ferns, as well as a number of extinct fossil groups.

<span class="mw-page-title-main">Ghost lineage</span> Phylogenetic lineage that is inferred to exist but has no fossil record

A ghost lineage is a hypothesized ancestor in a species lineage that has left no fossil 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. 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.

<span class="mw-page-title-main">Evolution of reptiles</span> Origin and diversification of reptiles through geologic time

Reptiles arose about 320 million years ago during the Carboniferous period. Reptiles, in the traditional sense of the term, are defined as animals that have scales or scutes, lay land-based hard-shelled eggs, and possess ectothermic metabolisms. So defined, the group is paraphyletic, excluding endothermic animals like birds that are descended from early traditionally-defined reptiles. A definition in accordance with phylogenetic nomenclature, which rejects paraphyletic groups, includes birds while excluding mammals and their synapsid ancestors. So defined, Reptilia is identical to Sauropsida.

<span class="mw-page-title-main">Fish fin</span> Bony skin-covered spines or rays protruding from the body of a fish

Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the spine and are supported only by muscles.

<span class="mw-page-title-main">Evolution of fish</span> Origin and diversification of fish through geologic time

The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include Haikouichthys. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans.

<i>Amia</i> (fish) Genus of ray-finned fishes

Amia, commonly called bowfin, is a genus of bony fish related to gars in the infraclass Holostei. They are regarded as taxonomic relicts, being the sole surviving species of the order Amiiformes, which dates from the Jurassic to the Eocene, persisting to the present. There are two living species in Amia, Amia calva and Amia ocellicauda, and a number of extinct species which have been described from the fossil record.

This list of fossil fish research presented in 2021 is a list of new taxa of jawless vertebrates, placoderms, acanthodians, fossil cartilaginous fishes, bony fishes, and other fishes that were described during the year, as well as other significant discoveries and events related to paleoichthyology that occurred in 2021.

This list of fossil fish research presented in 2022 is a list of new taxa of jawless vertebrates, placoderms, acanthodians, fossil cartilaginous fishes, bony fishes, and other fishes that were described during the year, as well as other significant discoveries and events related to paleoichthyology that occurred in 2022.

References

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