126-600: Sceloporus graciosus arenicolus Degenhardt & Jones, 1972 Sceloporus arenicolus , the dunes sagebrush lizard , formerly known as the sand dune lizard and the dunes-sagebrush lizard, is an insectivorous spiny lizard species which only occurs in the shinnery oak sand dune systems of southeast New Mexico and only four counties in adjacent Texas . Their habitat is restricted to semi-stabilized sand dunes which include large networks of shinnery oak ( Quercus havardii ), short (<2 m) shrubs, and sand sagebrush ( Artemisia filifolia ) on sloping, sandy topography, where
252-471: A jigsaw puzzle . Rocks normally form relatively horizontal layers, with each layer younger than the one underneath it. If a fossil is found between two layers whose ages are known, the fossil's age must lie between the two known ages. Because rock sequences are not continuous, but may be broken up by faults or periods of erosion , it is very difficult to match up rock beds that are not directly next to one another. However, fossils of species that survived for
378-423: A mutation–selection balance . It is predicted that a viral quasispecies at a low but evolutionarily neutral and highly connected (that is, flat) region in the fitness landscape will outcompete a quasispecies located at a higher but narrower fitness peak in which the surrounding mutants are unfit, "the quasispecies effect" or the "survival of the flattest". There is no suggestion that a viral quasispecies resembles
504-405: A ring species . Also, among organisms that reproduce only asexually , the concept of a reproductive species breaks down, and each clone is potentially a microspecies. Although none of these are entirely satisfactory definitions, and while the concept of species may not be a perfect model of life, it is still a useful tool to scientists and conservationists for studying life on Earth, regardless of
630-400: A "classical" method of determining species, such as with Linnaeus, early in evolutionary theory. However, different phenotypes are not necessarily different species (e.g. a four-winged Drosophila born to a two-winged mother is not a different species). Species named in this manner are called morphospecies . In the 1970s, Robert R. Sokal , Theodore J. Crovello and Peter Sneath proposed
756-424: A 'smallest clade' idea" (a phylogenetic species concept). Mishler and Wilkins and others concur with this approach, even though this would raise difficulties in biological nomenclature. Wilkins cited the ichthyologist Charles Tate Regan 's early 20th century remark that "a species is whatever a suitably qualified biologist chooses to call a species". Wilkins noted that the philosopher Philip Kitcher called this
882-552: A collision that formed the Moon about 40 million years later, may have cooled quickly enough to have oceans and an atmosphere about 4,440 million years ago . There is evidence on the Moon of a Late Heavy Bombardment by asteroids from 4,000 to 3,800 million years ago . If, as seems likely, such a bombardment struck Earth at the same time, the first atmosphere and oceans may have been stripped away. Paleontology traces
1008-602: A common ancestor. Ideally the "family tree" has only two branches leading from each node ("junction"), but sometimes there is too little information to achieve this, and paleontologists have to make do with junctions that have several branches. The cladistic technique is sometimes fallible, as some features, such as wings or camera eyes , evolved more than once, convergently – this must be taken into account in analyses. Evolutionary developmental biology , commonly abbreviated to "Evo Devo", also helps paleontologists to produce "family trees", and understand fossils. For example,
1134-428: A connected series of neighbouring populations, each of which can sexually interbreed with adjacent related populations, but for which there exist at least two "end" populations in the series, which are too distantly related to interbreed, though there is a potential gene flow between each "linked" population. Such non-breeding, though genetically connected, "end" populations may co-exist in the same region thus closing
1260-456: A constant rate. These " molecular clocks ", however, are fallible, and provide only a very approximate timing: for example, they are not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first evolved, and estimates produced by different techniques may vary by a factor of two. Earth formed about 4,570 million years ago and, after
1386-403: A data source that is not limited to animals with easily fossilised hard parts, and they reflect organisms' behaviours. Also many traces date from significantly earlier than the body fossils of animals that are thought to have been capable of making them. Whilst exact assignment of trace fossils to their makers is generally impossible, traces may for example provide the earliest physical evidence of
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#17327827813241512-432: A different species from its ancestors. Viruses have enormous populations, are doubtfully living since they consist of little more than a string of DNA or RNA in a protein coat, and mutate rapidly. All of these factors make conventional species concepts largely inapplicable. A viral quasispecies is a group of genotypes related by similar mutations, competing within a highly mutagenic environment, and hence governed by
1638-573: A fortunate accident during other research. For example, the 1980 discovery by Luis and Walter Alvarez of iridium , a mainly extraterrestrial metal, in the Cretaceous – Paleogene boundary layer made asteroid impact the most favored explanation for the Cretaceous–Paleogene extinction event – although debate continues about the contribution of volcanism. A complementary approach to developing scientific knowledge, experimental science ,
1764-508: A genetic boundary suitable for defining a species concept is present. DNA barcoding has been proposed as a way to distinguish species suitable even for non-specialists to use. One of the barcodes is a region of mitochondrial DNA within the gene for cytochrome c oxidase . A database, Barcode of Life Data System , contains DNA barcode sequences from over 190,000 species. However, scientists such as Rob DeSalle have expressed concern that classical taxonomy and DNA barcoding, which they consider
1890-821: A minor group until the first jawed fish appeared in the Late Ordovician . The spread of animals and plants from water to land required organisms to solve several problems, including protection against drying out and supporting themselves against gravity . The earliest evidence of land plants and land invertebrates date back to about 476 million years ago and 490 million years ago respectively. Those invertebrates, as indicated by their trace and body fossils, were shown to be arthropods known as euthycarcinoids . The lineage that produced land vertebrates evolved later but very rapidly between 370 million years ago and 360 million years ago ; recent discoveries have overturned earlier ideas about
2016-492: A misnomer, need to be reconciled, as they delimit species differently. Genetic introgression mediated by endosymbionts and other vectors can further make barcodes ineffective in the identification of species. A phylogenetic or cladistic species is "the smallest aggregation of populations (sexual) or lineages (asexual) diagnosable by a unique combination of character states in comparable individuals (semaphoronts)". The empirical basis – observed character states – provides
2142-449: A particular species, including which genus (and higher taxa) it is placed in, is a hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further information comes to hand, the hypothesis may be corroborated or refuted. Sometimes, especially in the past when communication was more difficult, taxonomists working in isolation have given two distinct names to individual organisms later identified as
2268-555: A rapid increase in knowledge about the history of life on Earth and to progress in the definition of the geologic time scale , largely based on fossil evidence. Although she was rarely recognised by the scientific community, Mary Anning was a significant contributor to the field of palaeontology during this period; she uncovered multiple novel Mesozoic reptile fossils and deducted that what were then known as bezoar stones are in fact fossilised faeces . In 1822 Henri Marie Ducrotay de Blainville , editor of Journal de Physique , coined
2394-543: A relatively short time can be used to link up isolated rocks: this technique is called biostratigraphy . For instance, the conodont Eoplacognathus pseudoplanus has a short range in the Middle Ordovician period. If rocks of unknown age are found to have traces of E. pseudoplanus , they must have a mid-Ordovician age. Such index fossils must be distinctive, be globally distributed and have a short time range to be useful. However, misleading results are produced if
2520-400: A short way of saying that something applies to many species within a genus, but not to all. If scientists mean that something applies to all species within a genus, they use the genus name without the specific name or epithet. The names of genera and species are usually printed in italics . However, abbreviations such as "sp." should not be italicised. When a species' identity is not clear,
2646-404: A specialist may use "cf." before the epithet to indicate that confirmation is required. The abbreviations "nr." (near) or "aff." (affine) may be used when the identity is unclear but when the species appears to be similar to the species mentioned after. With the rise of online databases, codes have been devised to provide identifiers for species that are already defined, including: The naming of
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#17327827813242772-523: A species as groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups. It has been argued that this definition is a natural consequence of the effect of sexual reproduction on the dynamics of natural selection. Mayr's use of the adjective "potentially" has been a point of debate; some interpretations exclude unusual or artificial matings that occur only in captivity, or that involve animals capable of mating but that do not normally do so in
2898-400: A species as determined by a taxonomist. A typological species is a group of organisms in which individuals conform to certain fixed properties (a type), so that even pre-literate people often recognise the same taxon as do modern taxonomists. The clusters of variations or phenotypes within specimens (such as longer or shorter tails) would differentiate the species. This method was used as
3024-491: A species. All species definitions assume that an organism acquires its genes from one or two parents very like the "daughter" organism, but that is not what happens in HGT. There is strong evidence of HGT between very dissimilar groups of prokaryotes , and at least occasionally between dissimilar groups of eukaryotes , including some crustaceans and echinoderms . The evolutionary biologist James Mallet concludes that there
3150-685: A species. Generally the term includes the unknown element of a distinct act of creation. Many authors have argued that a simple textbook definition, following Mayr's concept, works well for most multi-celled organisms , but breaks down in several situations: Species identification is made difficult by discordance between molecular and morphological investigations; these can be categorised as two types: (i) one morphology, multiple lineages (e.g. morphological convergence , cryptic species ) and (ii) one lineage, multiple morphologies (e.g. phenotypic plasticity , multiple life-cycle stages). In addition, horizontal gene transfer (HGT) makes it difficult to define
3276-608: A steady increase in brain size after about 3 million years ago . There is a long-running debate about whether modern humans are descendants of a single small population in Africa , which then migrated all over the world less than 200,000 years ago and replaced previous hominine species, or arose worldwide at the same time as a result of interbreeding . Life on earth has suffered occasional mass extinctions at least since 542 million years ago . Despite their disastrous effects, mass extinctions have sometimes accelerated
3402-528: A taxonomic decision at the discretion of cognizant specialists, is not governed by the Codes of Zoological or Botanical Nomenclature, in contrast to the PhyloCode , and contrary to what is done in several other fields, in which the definitions of technical terms, like geochronological units and geopolitical entities, are explicitly delimited. The nomenclatural codes that guide the naming of species, including
3528-517: A traditional biological species. The International Committee on Taxonomy of Viruses has since 1962 developed a universal taxonomic scheme for viruses; this has stabilised viral taxonomy. Most modern textbooks make use of Ernst Mayr 's 1942 definition, known as the Biological Species Concept as a basis for further discussion on the definition of species. It is also called a reproductive or isolation concept. This defines
3654-447: A variation on the morphological species concept, a phenetic species, defined as a set of organisms with a similar phenotype to each other, but a different phenotype from other sets of organisms. It differs from the morphological species concept in including a numerical measure of distance or similarity to cluster entities based on multivariate comparisons of a reasonably large number of phenotypic traits. A mate-recognition species
3780-515: A variety of reasons. Viruses are a special case, driven by a balance of mutation and selection , and can be treated as quasispecies . Biologists and taxonomists have made many attempts to define species, beginning from morphology and moving towards genetics . Early taxonomists such as Linnaeus had no option but to describe what they saw: this was later formalised as the typological or morphological species concept. Ernst Mayr emphasised reproductive isolation, but this, like other species concepts,
3906-438: Is "an entity composed of organisms which maintains its identity from other such entities through time and over space, and which has its own independent evolutionary fate and historical tendencies". This differs from the biological species concept in embodying persistence over time. Wiley and Mayden stated that they see the evolutionary species concept as "identical" to Willi Hennig 's species-as-lineages concept, and asserted that
Sceloporus arenicolus - Misplaced Pages Continue
4032-400: Is a group of sexually reproducing organisms that recognise one another as potential mates. Expanding on this to allow for post-mating isolation, a cohesion species is the most inclusive population of individuals having the potential for phenotypic cohesion through intrinsic cohesion mechanisms; no matter whether populations can hybridise successfully, they are still distinct cohesion species if
4158-458: Is a set of organisms adapted to a particular set of resources, called a niche, in the environment. According to this concept, populations form the discrete phenetic clusters that we recognise as species because the ecological and evolutionary processes controlling how resources are divided up tend to produce those clusters. A genetic species as defined by Robert Baker and Robert Bradley is a set of genetically isolated interbreeding populations. This
4284-414: Is between 8 and 8.7 million. About 14% of these had been described by 2011. All species (except viruses ) are given a two-part name , called a "binomial". The first part of a binomial is the genus to which the species belongs. The second part is called the specific name or the specific epithet (in botanical nomenclature , also sometimes in zoological nomenclature ). For example, Boa constrictor
4410-414: Is called speciation . Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book The Origin of Species . Speciation depends on a measure of reproductive isolation , a reduced gene flow. This occurs most easily in allopatric speciation, where populations are separated geographically and can diverge gradually as mutations accumulate. Reproductive isolation
4536-595: Is composed only of eukaryotic cells, and the earliest evidence for it is the Francevillian Group Fossils from 2,100 million years ago , although specialisation of cells for different functions first appears between 1,430 million years ago (a possible fungus) and 1,200 million years ago (a probable red alga ). Sexual reproduction may be a prerequisite for specialisation of cells, as an asexual multicellular organism might be at risk of being taken over by rogue cells that retain
4662-403: Is described formally, in a publication that assigns it a unique scientific name. The description typically provides means for identifying the new species, which may not be based solely on morphology (see cryptic species ), differentiating it from other previously described and related or confusable species and provides a validly published name (in botany) or an available name (in zoology) when
4788-671: Is further weakened by the existence of microspecies , groups of organisms, including many plants, with very little genetic variability, usually forming species aggregates . For example, the dandelion Taraxacum officinale and the blackberry Rubus fruticosus are aggregates with many microspecies—perhaps 400 in the case of the blackberry and over 200 in the dandelion, complicated by hybridisation , apomixis and polyploidy , making gene flow between populations difficult to determine, and their taxonomy debatable. Species complexes occur in insects such as Heliconius butterflies, vertebrates such as Hypsiboas treefrogs, and fungi such as
4914-726: Is hard or even impossible to test. Later biologists have tried to refine Mayr's definition with the recognition and cohesion concepts, among others. Many of the concepts are quite similar or overlap, so they are not easy to count: the biologist R. L. Mayden recorded about 24 concepts, and the philosopher of science John Wilkins counted 26. Wilkins further grouped the species concepts into seven basic kinds of concepts: (1) agamospecies for asexual organisms (2) biospecies for reproductively isolated sexual organisms (3) ecospecies based on ecological niches (4) evolutionary species based on lineage (5) genetic species based on gene pool (6) morphospecies based on form or phenotype and (7) taxonomic species,
5040-482: Is hard to decide at what level to place a new higher-level grouping, e.g. genus or family or order ; this is important since the Linnaean rules for naming groups are tied to their levels, and hence if a group is moved to a different level it must be renamed. Paleontologists generally use approaches based on cladistics , a technique for working out the evolutionary "family tree" of a set of organisms. It works by
5166-403: Is no easy way to tell whether related geographic or temporal forms belong to the same or different species. Species gaps can be verified only locally and at a point of time. One is forced to admit that Darwin's insight is correct: any local reality or integrity of species is greatly reduced over large geographic ranges and time periods. The botanist Brent Mishler argued that the species concept
Sceloporus arenicolus - Misplaced Pages Continue
5292-478: Is not valid, notably because gene flux decreases gradually rather than in discrete steps, which hampers objective delimitation of species. Indeed, complex and unstable patterns of gene flux have been observed in cichlid teleosts of the East African Great Lakes . Wilkins argued that "if we were being true to evolution and the consequent phylogenetic approach to taxa, we should replace it with
5418-443: Is often said to work by conducting experiments to disprove hypotheses about the workings and causes of natural phenomena. This approach cannot prove a hypothesis, since some later experiment may disprove it, but the accumulation of failures to disprove is often compelling evidence in favor. However, when confronted with totally unexpected phenomena, such as the first evidence for invisible radiation , experimental scientists often use
5544-400: Is one of the species of the genus Boa , with constrictor being the species' epithet. While the definitions given above may seem adequate at first glance, when looked at more closely they represent problematic species concepts. For example, the boundaries between closely related species become unclear with hybridisation , in a species complex of hundreds of similar microspecies , and in
5670-594: Is one that contained an extinct "crocodile-like" marine reptile, which eventually came to be known as the mosasaurid Mosasaurus of the Cretaceous period. The first half of the 19th century saw geological and paleontological activity become increasingly well organised with the growth of geologic societies and museums and an increasing number of professional geologists and fossil specialists. Interest increased for reasons that were not purely scientific, as geology and paleontology helped industrialists to find and exploit natural resources such as coal. This contributed to
5796-401: Is our only means of giving rocks greater than about 50 million years old an absolute age, and can be accurate to within 0.5% or better. Although radiometric dating requires very careful laboratory work, its basic principle is simple: the rates at which various radioactive elements decay are known, and so the ratio of the radioactive element to the element into which it decays shows how long ago
5922-586: Is similar to Mayr's Biological Species Concept, but stresses genetic rather than reproductive isolation. In the 21st century, a genetic species could be established by comparing DNA sequences. Earlier, other methods were available, such as comparing karyotypes (sets of chromosomes ) and allozymes ( enzyme variants). An evolutionarily significant unit (ESU) or "wildlife species" is a population of organisms considered distinct for purposes of conservation. In palaeontology , with only comparative anatomy (morphology) and histology from fossils as evidence,
6048-578: Is sometimes an important source of genetic variation. Viruses can transfer genes between species. Bacteria can exchange plasmids with bacteria of other species, including some apparently distantly related ones in different phylogenetic domains , making analysis of their relationships difficult, and weakening the concept of a bacterial species. Paleontologist Paleontology ( / ˌ p eɪ l i ɒ n ˈ t ɒ l ə dʒ i , ˌ p æ l i -, - ən -/ PAY -lee-on- TOL -ə-jee, PAL -ee-, -ən- ), also spelled palaeontology or palæontology ,
6174-434: Is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity . Other ways of defining species include their karyotype , DNA sequence, morphology , behaviour, or ecological niche . In addition, paleontologists use the concept of the chronospecies since fossil reproduction cannot be examined. The most recent rigorous estimate for the total number of species of eukaryotes
6300-456: Is the scientific study of life that existed prior to the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossils to classify organisms and study their interactions with each other and their environments (their paleoecology ). Paleontological observations have been documented as far back as the 5th century BC. The science became established in
6426-503: Is thought to have been propelled by coevolution with pollinating insects. Social insects appeared around the same time and, although they account for only small parts of the insect "family tree", now form over 50% of the total mass of all insects. Humans evolved from a lineage of upright-walking apes whose earliest fossils date from over 6 million years ago . Although early members of this lineage had chimp -sized brains, about 25% as big as modern humans', there are signs of
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#17327827813246552-594: Is threatened by hybridisation, but this can be selected against once a pair of populations have incompatible alleles of the same gene, as described in the Bateson–Dobzhansky–Muller model . A different mechanism, phyletic speciation, involves one lineage gradually changing over time into a new and distinct form (a chronospecies ), without increasing the number of resultant species. Horizontal gene transfer between organisms of different species, either through hybridisation , antigenic shift , or reassortment ,
6678-530: The ICZN for animals and the ICN for plants, do not make rules for defining the boundaries of the species. Research can change the boundaries, also known as circumscription, based on new evidence. Species may then need to be distinguished by the boundary definitions used, and in such cases the names may be qualified with sensu stricto ("in the narrow sense") to denote usage in the exact meaning given by an author such as
6804-582: The Middle Ages the Persian naturalist Ibn Sina , known as Avicenna in Europe, discussed fossils and proposed a theory of petrifying fluids on which Albert of Saxony elaborated in the 14th century. The Chinese naturalist Shen Kuo (1031–1095) proposed a theory of climate change based on the presence of petrified bamboo in regions that in his time were too dry for bamboo. In early modern Europe ,
6930-538: The Neogene - Quaternary . In deeper-level deposits in western Europe are early-aged mammals such as the palaeothere perissodactyl Palaeotherium and the anoplotheriid artiodactyl Anoplotherium , both of which were described earliest after the former two genera, which today are known to date to the Paleogene period. Cuvier figured out that even older than the two levels of deposits with extinct large mammals
7056-637: The Permian–Triassic extinction event . Amphibians Extinct Synapsids Mammals Extinct reptiles Lizards and snakes Extinct Archosaurs Crocodilians Extinct Dinosaurs Birds Naming groups of organisms in a way that is clear and widely agreed is important, as some disputes in paleontology have been based just on misunderstandings over names. Linnaean taxonomy is commonly used for classifying living organisms, but runs into difficulties when dealing with newly discovered organisms that are significantly different from known ones. For example: it
7182-516: The Permian–Triassic extinction event . A relatively recent discipline, molecular phylogenetics , compares the DNA and RNA of modern organisms to re-construct the "family trees" of their evolutionary ancestors. It has also been used to estimate the dates of important evolutionary developments, although this approach is controversial because of doubts about the reliability of the " molecular clock ". Techniques from engineering have been used to analyse how
7308-454: The embryological development of some modern brachiopods suggests that brachiopods may be descendants of the halkieriids , which became extinct in the Cambrian period. Paleontology seeks to map out how living things have changed through time. A substantial hurdle to this aim is the difficulty of working out how old fossils are. Beds that preserve fossils typically lack the radioactive elements needed for radiometric dating . This technique
7434-399: The fly agaric . Natural hybridisation presents a challenge to the concept of a reproductively isolated species, as fertile hybrids permit gene flow between two populations. For example, the carrion crow Corvus corone and the hooded crow Corvus cornix appear and are classified as separate species, yet they can hybridise where their geographical ranges overlap. A ring species is
7560-507: The jaguar ( Panthera onca ) of Latin America or the leopard ( Panthera pardus ) of Africa and Asia. In contrast, the scientific names of species are chosen to be unique and universal (except for some inter-code homonyms ); they are in two parts used together : the genus as in Puma , and the specific epithet as in concolor . A species is given a taxonomic name when a type specimen
7686-526: The " jigsaw puzzles " of biostratigraphy (arrangement of rock layers from youngest to oldest). Classifying ancient organisms is also difficult, as many do not fit well into the Linnaean taxonomy classifying living organisms, and paleontologists more often use cladistics to draw up evolutionary "family trees". The final quarter of the 20th century saw the development of molecular phylogenetics , which investigates how closely organisms are related by measuring
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#17327827813247812-406: The "cynical species concept", and arguing that far from being cynical, it usefully leads to an empirical taxonomy for any given group, based on taxonomists' experience. Other biologists have gone further and argued that we should abandon species entirely, and refer to the "Least Inclusive Taxonomic Units" (LITUs), a view that would be coherent with current evolutionary theory. The species concept
7938-463: The 18th century as a result of Georges Cuvier 's work on comparative anatomy , and developed rapidly in the 19th century. The term has been used since 1822 formed from Greek παλαιός ( 'palaios' , "old, ancient"), ὄν ( 'on' , ( gen. 'ontos' ), "being, creature"), and λόγος ( 'logos' , "speech, thought, study"). Paleontology lies on the border between biology and geology , but it differs from archaeology in that it excludes
8064-555: The Early Cambrian , along with several "weird wonders" that bear little obvious resemblance to any modern animals. There is a long-running debate about whether this Cambrian explosion was truly a very rapid period of evolutionary experimentation; alternative views are that modern-looking animals began evolving earlier but fossils of their precursors have not yet been found, or that the "weird wonders" are evolutionary "aunts" and "cousins" of modern groups. Vertebrates remained
8190-463: The Earth's organic and inorganic past". William Whewell (1794–1866) classified paleontology as one of the historical sciences, along with archaeology , geology, astronomy , cosmology , philology and history itself: paleontology aims to describe phenomena of the past and to reconstruct their causes. Hence it has three main elements: description of past phenomena; developing a general theory about
8316-405: The abbreviation "sp." in the singular or "spp." (standing for species pluralis , Latin for "multiple species") in the plural in place of the specific name or epithet (e.g. Canis sp.). This commonly occurs when authors are confident that some individuals belong to a particular genus but are not sure to which exact species they belong, as is common in paleontology . Authors may also use "spp." as
8442-412: The ability to reproduce. The earliest known animals are cnidarians from about 580 million years ago , but these are so modern-looking that they must be descendants of earlier animals. Early fossils of animals are rare because they had not developed mineralised , easily fossilized hard parts until about 548 million years ago . The earliest modern-looking bilaterian animals appear in
8568-570: The amount of hybridisation is insufficient to completely mix their respective gene pools . A further development of the recognition concept is provided by the biosemiotic concept of species. In microbiology , genes can move freely even between distantly related bacteria, possibly extending to the whole bacterial domain. As a rule of thumb, microbiologists have assumed that members of Bacteria or Archaea with 16S ribosomal RNA gene sequences more similar than 97% to each other need to be checked by DNA–DNA hybridisation to decide if they belong to
8694-515: The appearance of moderately complex animals (comparable to earthworms ). Geochemical observations may help to deduce the global level of biological activity at a certain period, or the affinity of certain fossils. For example, geochemical features of rocks may reveal when life first arose on Earth, and may provide evidence of the presence of eukaryotic cells, the type from which all multicellular organisms are built. Analyses of carbon isotope ratios may help to explain major transitions such as
8820-625: The atmosphere increased their effectiveness as nurseries of evolution. While eukaryotes , cells with complex internal structures, may have been present earlier, their evolution speeded up when they acquired the ability to transform oxygen from a poison to a powerful source of metabolic energy. This innovation may have come from primitive eukaryotes capturing oxygen-powered bacteria as endosymbionts and transforming them into organelles called mitochondria . The earliest evidence of complex eukaryotes with organelles (such as mitochondria) dates from 1,850 million years ago . Multicellular life
8946-474: The biological species concept, "the several versions" of the phylogenetic species concept, and the idea that species are of the same kind as higher taxa are not suitable for biodiversity studies (with the intention of estimating the number of species accurately). They further suggested that the concept works for both asexual and sexually-reproducing species. A version of the concept is Kevin de Queiroz 's "General Lineage Concept of Species". An ecological species
9072-505: The biological species concept, a cladistic species does not rely on reproductive isolation – its criteria are independent of processes that are integral in other concepts. Therefore, it applies to asexual lineages. However, it does not always provide clear cut and intuitively satisfying boundaries between taxa, and may require multiple sources of evidence, such as more than one polymorphic locus, to give plausible results. An evolutionary species, suggested by George Gaylord Simpson in 1951,
9198-415: The bodies of ancient organisms might have worked, for example the running speed and bite strength of Tyrannosaurus , or the flight mechanics of Microraptor . It is relatively commonplace to study the internal details of fossils using X-ray microtomography . Paleontology, biology, archaeology, and paleoneurobiology combine to study endocranial casts (endocasts) of species related to humans to clarify
9324-401: The causes of various types of change; and applying those theories to specific facts. When trying to explain the past, paleontologists and other historical scientists often construct a set of one or more hypotheses about the causes and then look for a " smoking gun ", a piece of evidence that strongly accords with one hypothesis over any others. Sometimes researchers discover a "smoking gun" by
9450-763: The characteristics and evolution of humans as a species. When dealing with evidence about humans, archaeologists and paleontologists may work together – for example paleontologists might identify animal or plant fossils around an archaeological site , to discover the people who lived there, and what they ate; or they might analyze the climate at the time of habitation. In addition, paleontology often borrows techniques from other sciences, including biology, osteology , ecology, chemistry , physics and mathematics. For example, geochemical signatures from rocks may help to discover when life first arose on Earth, and analyses of carbon isotope ratios may help to identify climate changes and even to explain major transitions such as
9576-520: The chronological order in which rocks were formed, is useful to both paleontologists and geologists. Biogeography studies the spatial distribution of organisms, and is also linked to geology, which explains how Earth's geography has changed over time. Although paleontology became established around 1800, earlier thinkers had noticed aspects of the fossil record. The ancient Greek philosopher Xenophanes (570–480 BCE) concluded from fossil sea shells that some areas of land were once under water. During
9702-433: The concept of a chronospecies can be applied. During anagenesis (evolution, not necessarily involving branching), some palaeontologists seek to identify a sequence of species, each one derived from the phyletically extinct one before through continuous, slow and more or less uniform change. In such a time sequence, some palaeontologists assess how much change is required for a morphologically distinct form to be considered
9828-445: The date when lineages first appeared. For instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an ancestor of B and C, then A must have evolved more than X million years ago. It is also possible to estimate how long ago two living clades diverged – i.e. approximately how long ago their last common ancestor must have lived – by assuming that DNA mutations accumulate at
9954-594: The development of mammalian traits such as endothermy and hair. After the Cretaceous–Paleogene extinction event 66 million years ago killed off all the dinosaurs except the birds, mammals increased rapidly in size and diversity, and some took to the air and the sea. Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early Cretaceous between 130 million years ago and 90 million years ago . Their rapid rise to dominance of terrestrial ecosystems
10080-561: The development of the body plans of most animal phyla . The discovery of fossils of the Ediacaran biota and developments in paleobiology extended knowledge about the history of life back far before the Cambrian. Increasing awareness of Gregor Mendel 's pioneering work in genetics led first to the development of population genetics and then in the mid-20th century to the modern evolutionary synthesis , which explains evolution as
10206-482: The different levels of deposits represented different time periods in the early 19th century. The surface-level deposits in the Americas contained later mammals like the megatheriid ground sloth Megatherium and the mammutid proboscidean Mammut (later known informally as a "mastodon"), which were some of the earliest-named fossil mammal genera with official taxonomic authorities. They today are known to date to
10332-449: The dunes sagebrush lizard and its habitat, either singly or in combination, are of sufficient imminence, intensity, and magnitude to indicate that the species is in danger of extinction throughout all of its range." Species A species ( pl. : species) is a population of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring , typically by sexual reproduction . It
10458-718: The dunes sagebrush lizard as endangered under the Endangered Species Act throughout its range in New Mexico and Texas. The final determination was originally due in July 2011, but was delayed to allow the scientific community to continue research to see if the listing was necessary. In June 2012 the Fish and Wildlife Service decided not to list it, citing the "unprecedented commitments to voluntary conservation agreements now in place in New Mexico and Texas" that provide for
10584-409: The end of the 20th century have been particularly important as they have provided new information about the earliest evolution of animals, early fish, dinosaurs and the evolution of birds. The last few decades of the 20th century saw a renewed interest in mass extinctions and their role in the evolution of life on Earth. There was also a renewed interest in the Cambrian explosion that apparently saw
10710-435: The evidence to support hypotheses about evolutionarily divergent lineages that have maintained their hereditary integrity through time and space. Molecular markers may be used to determine diagnostic genetic differences in the nuclear or mitochondrial DNA of various species. For example, in a study done on fungi , studying the nucleotide characters using cladistic species produced the most accurate results in recognising
10836-410: The evolution of the human brain. Paleontology even contributes to astrobiology , the investigation of possible life on other planets , by developing models of how life may have arisen and by providing techniques for detecting evidence of life. As knowledge has increased, paleontology has developed specialised subdivisions. Vertebrate paleontology concentrates on fossils from the earliest fish to
10962-466: The evolutionary history of life back to over 3,000 million years ago , possibly as far as 3,800 million years ago . The oldest clear evidence of life on Earth dates to 3,000 million years ago , although there have been reports, often disputed, of fossil bacteria from 3,400 million years ago and of geochemical evidence for the presence of life 3,800 million years ago . Some scientists have proposed that life on Earth
11088-555: The exceptional events that cause quick burial make it difficult to study the normal environments of the animals. The sparseness of the fossil record means that organisms are expected to exist long before and after they are found in the fossil record – this is known as the Signor–Lipps effect . Trace fossils consist mainly of tracks and burrows, but also include coprolites (fossil feces ) and marks left by feeding. Trace fossils are particularly significant because they represent
11214-515: The focus of paleontology shifted to understanding evolutionary paths, including human evolution , and evolutionary theory. The last half of the 19th century saw a tremendous expansion in paleontological activity, especially in North America. The trend continued in the 20th century with additional regions of the Earth being opened to systematic fossil collection. Fossils found in China near
11340-449: The following: At the end of the 18th century Georges Cuvier 's work established comparative anatomy as a scientific discipline and, by proving that some fossil animals resembled no living ones, demonstrated that animals could become extinct , leading to the emergence of paleontology. The expanding knowledge of the fossil record also played an increasing role in the development of geology, particularly stratigraphy . Cuvier proved that
11466-580: The fossil record: different environments are more favorable to the preservation of different types of organism or parts of organisms. Further, only the parts of organisms that were already mineralised are usually preserved, such as the shells of molluscs. Since most animal species are soft-bodied, they decay before they can become fossilised. As a result, although there are 30-plus phyla of living animals, two-thirds have never been found as fossils. Occasionally, unusual environments may preserve soft tissues. These lagerstätten allow paleontologists to examine
11592-687: The history and driving forces behind their evolution. Land plants were so successful that their detritus caused an ecological crisis in the Late Devonian , until the evolution of fungi that could digest dead wood. During the Permian period, synapsids , including the ancestors of mammals , may have dominated land environments, but this ended with the Permian–Triassic extinction event 251 million years ago , which came very close to wiping out all complex life. The extinctions were apparently fairly sudden, at least among vertebrates. During
11718-533: The history of Earth's climate and the mechanisms that have changed it – which have sometimes included evolutionary developments, for example the rapid expansion of land plants in the Devonian period removed more carbon dioxide from the atmosphere, reducing the greenhouse effect and thus helping to cause an ice age in the Carboniferous period. Biostratigraphy , the use of fossils to work out
11844-542: The immediate ancestors of modern mammals . Invertebrate paleontology deals with fossils such as molluscs , arthropods , annelid worms and echinoderms . Paleobotany studies fossil plants , algae , and fungi. Palynology , the study of pollen and spores produced by land plants and protists , straddles paleontology and botany , as it deals with both living and fossil organisms. Micropaleontology deals with microscopic fossil organisms of all kinds. Instead of focusing on individual organisms, paleoecology examines
11970-434: The index fossils turn out to have longer fossil ranges than first thought. Stratigraphy and biostratigraphy can in general provide only relative dating ( A was before B ), which is often sufficient for studying evolution. However, this is difficult for some time periods, because of the problems involved in matching up rocks of the same age across different continents . Family-tree relationships may also help to narrow down
12096-538: The interactions between different ancient organisms, such as their food chains , and the two-way interactions with their environments. For example, the development of oxygenic photosynthesis by bacteria caused the oxygenation of the atmosphere and hugely increased the productivity and diversity of ecosystems . Together, these led to the evolution of complex eukaryotic cells, from which all multicellular organisms are built. Paleoclimatology , although sometimes treated as part of paleoecology, focuses more on
12222-463: The internal anatomy of animals that in other sediments are represented only by shells, spines, claws, etc. – if they are preserved at all. However, even lagerstätten present an incomplete picture of life at the time. The majority of organisms living at the time are probably not represented because lagerstätten are restricted to a narrow range of environments, e.g. where soft-bodied organisms can be preserved very quickly by events such as mudslides; and
12348-456: The investigation of evolutionary "family trees" by techniques derived from biochemistry , began to make an impact, particularly when it was proposed that the human lineage had diverged from apes much more recently than was generally thought at the time. Although this early study compared proteins from apes and humans, most molecular phylogenetics research is now based on comparisons of RNA and DNA . Fossils of organisms' bodies are usually
12474-445: The lizard's range was sprayed with herbicide to clear the land for cattle grazing, and the lizards are now extinct at these locations. The dune systems are also heavily interrupted by oil industry activities. These interruptions allow Mesquite to invade areas where shinnery oak (and dunes sagebrush lizards) were once dominant. While herbicide spraying has been outlawed in the dunes sagebrush lizard's New Mexico distribution, development for
12600-441: The lizards use "blowouts" as their primary microhabitat . Blowouts are sandy, bowl-shaped interruptions in the shinnery oak sand dune system which look like small meteor craters. The roots of the shinnery oak shrubs provide structure for the dunes sagebrush lizards' burrows, where the lizards retreat from the blowouts when the sand surface is too hot or cold. Habitat destruction is their primary threat. Shinnery oak through much of
12726-409: The logic that, if groups B and C have more similarities to each other than either has to group A, then B and C are more closely related to each other than either is to A. Characters that are compared may be anatomical , such as the presence of a notochord , or molecular , by comparing sequences of DNA or proteins . The result of a successful analysis is a hierarchy of clades – groups that share
12852-415: The long-term conservation of the species. In May 2024, the Fish and Wildlife Service published a final rule to list the lizard as an endangered species, citing future energy development, sand mining and climate change as the biggest threats to its survival. The Fish and Wildlife Service estimates that 35% of the lizard's approximately 520,161 acres (2,105.02 km) of shinnery oak habitat has been degraded to
12978-409: The most informative type of evidence. The most common types are wood, bones, and shells. Fossilisation is a rare event, and most fossils are destroyed by erosion or metamorphism before they can be observed. Hence the fossil record is very incomplete, increasingly so further back in time. Despite this, it is often adequate to illustrate the broader patterns of life's history. There are also biases in
13104-483: The numerous fungi species of all the concepts studied. Versions of the phylogenetic species concept that emphasise monophyly or diagnosability may lead to splitting of existing species, for example in Bovidae , by recognising old subspecies as species, despite the fact that there are no reproductive barriers, and populations may intergrade morphologically. Others have called this approach taxonomic inflation , diluting
13230-426: The oil industry has not ceased. Competition from other lizard species may be a threat, as well. Uta stansburiana , the side-blotched lizard, seems to be more of a habitat generalist than the dunes sagebrush lizard, and may be able to take advantage of recent habitat changes, introducing skewed resource competition that is not natural for that ecosystem. The United States Fish and Wildlife Service proposed listing
13356-414: The outcome of events such as mutations and horizontal gene transfer , which provide genetic variation , with genetic drift and natural selection driving changes in this variation over time. Within the next few years the role and operation of DNA in genetic inheritance were discovered, leading to what is now known as the "Central Dogma" of molecular biology . In the 1960s molecular phylogenetics ,
13482-593: The paper is accepted for publication. The type material is usually held in a permanent repository, often the research collection of a major museum or university, that allows independent verification and the means to compare specimens. Describers of new species are asked to choose names that, in the words of the International Code of Zoological Nomenclature , are "appropriate, compact, euphonious, memorable, and do not cause offence". Books and articles sometimes intentionally do not identify species fully, using
13608-674: The person who named the species, while the antonym sensu lato ("in the broad sense") denotes a wider usage, for instance including other subspecies. Other abbreviations such as "auct." ("author"), and qualifiers such as "non" ("not") may be used to further clarify the sense in which the specified authors delineated or described the species. Species are subject to change, whether by evolving into new species, exchanging genes with other species, merging with other species or by becoming extinct. The evolutionary process by which biological populations of sexually-reproducing organisms evolve to become distinct or reproductively isolated as species
13734-459: The point it is no longer likely to have viable populations of the species, and that 47% of the species' geographic range has experienced habitat loss to the point of causing significant population declines. Areas where habitat quality was high were disconnected to other sand dune habitats and relatively small. Rangewide population data was not available at the time of listing, but the Fish and Wildlife Service concluded that "the risk factors acting on
13860-452: The principal types of evidence about ancient life, and geochemical evidence has helped to decipher the evolution of life before there were organisms large enough to leave body fossils. Estimating the dates of these remains is essential but difficult: sometimes adjacent rock layers allow radiometric dating , which provides absolute dates that are accurate to within 0.5%, but more often paleontologists have to rely on relative dating by solving
13986-432: The radioactive element was incorporated into the rock. Radioactive elements are common only in rocks with a volcanic origin, and so the only fossil-bearing rocks that can be dated radiometrically are a few volcanic ash layers. Consequently, paleontologists must usually rely on stratigraphy to date fossils. Stratigraphy is the science of deciphering the "layer-cake" that is the sedimentary record, and has been compared to
14112-487: The result of misclassification leading to questions on whether there really are any ring species. The commonly used names for kinds of organisms are often ambiguous: "cat" could mean the domestic cat, Felis catus , or the cat family, Felidae . Another problem with common names is that they often vary from place to place, so that puma, cougar, catamount, panther, painter and mountain lion all mean Puma concolor in various parts of America, while "panther" may also mean
14238-586: The ring. Ring species thus present a difficulty for any species concept that relies on reproductive isolation. However, ring species are at best rare. Proposed examples include the herring gull – lesser black-backed gull complex around the North pole, the Ensatina eschscholtzii group of 19 populations of salamanders in America, and the greenish warbler in Asia, but many so-called ring species have turned out to be
14364-701: The same approach as historical scientists: construct a set of hypotheses about the causes and then look for a "smoking gun". Paleontology lies between biology and geology since it focuses on the record of past life, but its main source of evidence is fossils in rocks. For historical reasons, paleontology is part of the geology department at many universities: in the 19th and early 20th centuries, geology departments found fossil evidence important for dating rocks, while biology departments showed little interest. Paleontology also has some overlap with archaeology , which primarily works with objects made by humans and with human remains, while paleontologists are interested in
14490-508: The same species. This concept was narrowed in 2006 to a similarity of 98.7%. The average nucleotide identity (ANI) method quantifies genetic distance between entire genomes , using regions of about 10,000 base pairs . With enough data from genomes of one genus, algorithms can be used to categorize species, as for Pseudomonas avellanae in 2013, and for all sequenced bacteria and archaea since 2020. Observed ANI values among sequences appear to have an "ANI gap" at 85–95%, suggesting that
14616-529: The same species. When two species names are discovered to apply to the same species, the older species name is given priority and usually retained, and the newer name considered as a junior synonym, a process called synonymy . Dividing a taxon into multiple, often new, taxa is called splitting . Taxonomists are often referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to recognising differences or commonalities between organisms. The circumscription of taxa, considered
14742-478: The similarity of the DNA in their genomes . Molecular phylogenetics has also been used to estimate the dates when species diverged, but there is controversy about the reliability of the molecular clock on which such estimates depend. The simplest definition of "paleontology" is "the study of ancient life". The field seeks information about several aspects of past organisms: "their identity and origin, their environment and evolution, and what they can tell us about
14868-470: The slow recovery from this catastrophe a previously obscure group, archosaurs , became the most abundant and diverse terrestrial vertebrates. One archosaur group, the dinosaurs, were the dominant land vertebrates for the rest of the Mesozoic , and birds evolved from one group of dinosaurs. During this time mammals' ancestors survived only as small, mainly nocturnal insectivores , which may have accelerated
14994-506: The species concept and making taxonomy unstable. Yet others defend this approach, considering "taxonomic inflation" pejorative and labelling the opposing view as "taxonomic conservatism"; claiming it is politically expedient to split species and recognise smaller populations at the species level, because this means they can more easily be included as endangered in the IUCN red list and can attract conservation legislation and funding. Unlike
15120-631: The study of anatomically modern humans . It now uses techniques drawn from a wide range of sciences, including biochemistry , mathematics , and engineering. Use of all these techniques has enabled paleontologists to discover much of the evolutionary history of life , almost back to when Earth became capable of supporting life, nearly 4 billion years ago. As knowledge has increased, paleontology has developed specialised sub-divisions, some of which focus on different types of fossil organisms while others study ecology and environmental history, such as ancient climates . Body fossils and trace fossils are
15246-629: The systematic study of fossils emerged as an integral part of the changes in natural philosophy that occurred during the Age of Reason . In the Italian Renaissance, Leonardo da Vinci made various significant contributions to the field as well as depicted numerous fossils. Leonardo's contributions are central to the history of paleontology because he established a line of continuity between the two main branches of paleontology – ichnology and body fossil paleontology. He identified
15372-485: The theoretical difficulties. If species were fixed and clearly distinct from one another, there would be no problem, but evolutionary processes cause species to change. This obliges taxonomists to decide, for example, when enough change has occurred to declare that a lineage should be divided into multiple chronospecies , or when populations have diverged to have enough distinct character states to be described as cladistic species. Species and higher taxa were seen from
15498-877: The time of Aristotle until the 18th century as categories that could be arranged in a hierarchy, the great chain of being . In the 19th century, biologists grasped that species could evolve given sufficient time. Charles Darwin 's 1859 book On the Origin of Species explained how species could arise by natural selection . That understanding was greatly extended in the 20th century through genetics and population ecology . Genetic variability arises from mutations and recombination , while organisms themselves are mobile, leading to geographical isolation and genetic drift with varying selection pressures . Genes can sometimes be exchanged between species by horizontal gene transfer ; new species can arise rapidly through hybridisation and polyploidy ; and species may become extinct for
15624-540: The wild. It is difficult to define a species in a way that applies to all organisms. The debate about species concepts is called the species problem. The problem was recognised even in 1859, when Darwin wrote in On the Origin of Species : I was much struck how entirely vague and arbitrary is the distinction between species and varieties. He went on to write: No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of
15750-406: The word "palaeontology" to refer to the study of ancient living organisms through fossils. As knowledge of life's history continued to improve, it became increasingly obvious that there had been some kind of successive order to the development of life. This encouraged early evolutionary theories on the transmutation of species . After Charles Darwin published Origin of Species in 1859, much of
15876-454: Was "seeded" from elsewhere , but most research concentrates on various explanations of how life could have arisen independently on Earth. For about 2,000 million years microbial mats , multi-layered colonies of different bacteria, were the dominant life on Earth. The evolution of oxygenic photosynthesis enabled them to play the major role in the oxygenation of the atmosphere from about 2,400 million years ago . This change in
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