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Species

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The hierarchy of biological classification's eight major taxonomic ranks. A genus contains one or more species. Intermediate minor rankings are not shown.LifeDomainKingdomPhylumClassOrderFamilyGenusSpecies
The hierarchy of biological classification's eight major taxonomic ranks. A genus contains one or more species. Intermediate minor rankings are not shown.

In biology, a species is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It 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 is between 8 and 8.7 million.[1][2][3] However, only about 14% of these had been described by 2011.[3]

All species (except viruses) are given a two-part name, 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 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 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 an incredibly useful tool to scientists and conservationists for studying life on Earth, regardless of 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 were seen from the time of Aristotle until the 18th century as fixed 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 a variety of reasons. Viruses are a special case, driven by a balance of mutation and selection, and can be treated as quasispecies.

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Biology

Biology

Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary information encoded in genes, which can be transmitted to future generations. Another major theme is evolution, which explains the unity and diversity of life. Energy processing is also important to life as it allows organisms to move, grow, and reproduce. Finally, all organisms are able to regulate their own internal environments.

Biodiversity

Biodiversity

Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic, species, and ecosystem level. Biodiversity is not distributed evenly on Earth; it is usually greater in the tropics as a result of the warm climate and high primary productivity in the region near the equator. Tropical forest ecosystems cover less than 10% of earth's surface and contain about 90% of the world's species. Marine biodiversity is usually higher along coasts in the Western Pacific, where sea surface temperature is highest, and in the mid-latitudinal band in all oceans. There are latitudinal gradients in species diversity. Biodiversity generally tends to cluster in hotspots, and has been increasing through time, but will be likely to slow in the future as a primary result of deforestation. It encompasses the evolutionary, ecological, and cultural processes that sustain life.

Chronospecies

Chronospecies

A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is physically, morphologically, and/or genetically distinct from the original ancestors. Throughout the change, there is only one species in the lineage at any point in time, as opposed to cases where divergent evolution produces contemporary species with a common ancestor. The related term paleospecies indicates an extinct species only identified with fossil material. That identification relies on distinct similarities between the earlier fossil specimens and some proposed descendant although the exact relationship to the later species is not always defined. In particular, the range of variation within all the early fossil specimens does not exceed the observed range that exists in the later species.

Binomial nomenclature

Binomial nomenclature

In taxonomy, binomial nomenclature, also called binominal nomenclature or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, although they can be based on words from other languages. Such a name is called a binomial name, a binomen, binominal name or a scientific name; more informally it is also historically called a Latin name.

Botanical nomenclature

Botanical nomenclature

Botanical nomenclature is the formal, scientific naming of plants. It is related to, but distinct from taxonomy. Plant taxonomy is concerned with grouping and classifying plants; botanical nomenclature then provides names for the results of this process. The starting point for modern botanical nomenclature is Linnaeus' Species Plantarum of 1753. Botanical nomenclature is governed by the International Code of Nomenclature for algae, fungi, and plants (ICN), which replaces the International Code of Botanical Nomenclature (ICBN). Fossil plants are also covered by the code of nomenclature.

Boa constrictor

Boa constrictor

The boa constrictor, also known as the red-tailed boa, is a species of large, non-venomous, heavy-bodied snake that is frequently kept and bred in captivity. The boa constrictor is a member of the family Boidae. The species is native to tropical South America. A staple of private collections and public displays, its color pattern is highly variable yet distinctive. Four subspecies are recognized. This article focuses on the species Boa constrictor as a whole, and on the nominate subspecies B. c. constrictor.

Boa (genus)

Boa (genus)

Boa is a genus of non-venomous boas found in Mexico, the Caribbean, and Central and South America. Five extant species, and one extinct, are currently recognized.

Asexual reproduction

Asexual reproduction

Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the full set of genes of their single parent and thus the newly created individual is genetically and physically similar to the parent or an exact clone of the parent. Asexual reproduction is the primary form of reproduction for single-celled organisms such as archaea and bacteria. Many eukaryotic organisms including plants, animals, and fungi can also reproduce asexually. In vertebrates, the most common form of asexual reproduction is parthenogenesis, which is typically used as an alternative to sexual reproduction in times when reproductive opportunities are limited. Komodo dragons and some monitor lizards can also reproduce asexually.

Cladistics

Cladistics

Cladistics is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived characteristics (synapomorphies) that are not present in more distant groups and ancestors. However, from an empirical perspective, common ancestors are inferences based on a cladistic hypothesis of relationships of taxa whose character states can be observed. Theoretically, a last common ancestor and all its descendants constitute a (minimal) clade. Importantly, all descendants stay in their overarching ancestral clade. For example, if the terms worms or fishes were used within a strict cladistic framework, these terms would include humans. Many of these terms are normally used paraphyletically, outside of cladistics, e.g. as a 'grade', which are fruitless to precisely delineate, especially when including extinct species. Radiation results in the generation of new subclades by bifurcation, but in practice sexual hybridization may blur very closely related groupings.

Aristotle

Aristotle

Aristotle was an Ancient Greek philosopher and polymath. His writings cover a broad range of subjects including physics, biology, zoology, metaphysics, logic, ethics, aesthetics, poetry, drama, music, rhetoric, psychology, linguistics, economics, politics, meteorology, geology, and government. As the founder of the Peripatetic school of philosophy in the Lyceum in Athens, he began the wider Aristotelian tradition that followed, which set the groundwork for the development of modern science.

Charles Darwin

Charles Darwin

Charles Robert Darwin was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended from a common ancestor is now generally accepted and considered a fundamental concept in science. In a joint publication with Alfred Russel Wallace, he introduced his scientific theory that this branching pattern of evolution resulted from a process he called natural selection, in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding. Darwin has been described as one of the most influential figures in human history and was honoured by burial in Westminster Abbey.

Allopatric speciation

Allopatric speciation

Allopatric speciation – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from each other to an extent that prevents or interferes with gene flow.

Definition

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, is hard or even impossible to test.[4][5] Later biologists have tried to refine Mayr's definition with the recognition and cohesion concepts, among others.[6] 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,[7] and the philosopher of science John Wilkins counted 26.[4] 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, a species as determined by a taxonomist.[8]

Typological or morphological species

All adult Eurasian blue tits share the same coloration, unmistakably identifying the morphospecies.[9]
All adult Eurasian blue tits share the same coloration, unmistakably identifying the morphospecies.[9]

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.[10][11] The clusters of variations or phenotypes within specimens (such as longer or shorter tails) would differentiate the species. This method was used as 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.[12][13]

In the 1970s, Robert R. Sokal, Theodore J. Crovello and Peter Sneath proposed 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.[14] 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.[15]

Recognition and cohesion species

A mate-recognition species is a group of sexually reproducing organisms that recognise one another as potential mates.[16][17] 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 the amount of hybridisation is insufficient to completely mix their respective gene pools.[18] A further development of the recognition concept is provided by the biosemiotic concept of species.[19]

Genetic similarity and barcode species

A region of the gene for the cytochrome c oxidase enzyme is used to distinguish species in the Barcode of Life Data Systems database.
A region of the gene for the cytochrome c oxidase enzyme is used to distinguish species in the Barcode of Life Data Systems database.

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 the same species.[20] This concept was narrowed in 2006 to a similarity of 98.7%.[21]

The average nucleotide identity 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,[22] and for all sequenced bacteria and archaea since 2020.[23]

DNA barcoding has been proposed as a way to distinguish species suitable even for non-specialists to use.[24] 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.[25][26] However, scientists such as Rob DeSalle have expressed concern that classical taxonomy and DNA barcoding, which they consider a misnomer, need to be reconciled, as they delimit species differently.[27] Genetic introgression mediated by endosymbionts and other vectors can further make barcodes ineffective in the identification of species.[28]

Phylogenetic or cladistic species

The cladistic or phylogenetic species concept is that a species is the smallest lineage which is distinguished by a unique set of either genetic or morphological traits. No claim is made about reproductive isolation, making the concept useful also in palaeontology where only fossil evidence is available.
The cladistic or phylogenetic species concept is that a species is the smallest lineage which is distinguished by a unique set of either genetic or morphological traits. No claim is made about reproductive isolation, making the concept useful also in palaeontology where only fossil evidence is available.

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)".[29] The empirical basis – observed character states – provides the evidence to support hypotheses about evolutionarily divergent lineages that have maintained their hereditary integrity through time and space.[30][31][32][33] Molecular markers may be used to determine diagnostic genetic differences in the nuclear or mitochondrial DNA of various species.[34][29][35] For example, in a study done on fungi, studying the nucleotide characters using cladistic species produced the most accurate results in recognising the numerous fungi species of all the concepts studied.[35][36] Versions of the phylogenetic species concept that emphasise monophyly or diagnosability[37] 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.[38] Others have called this approach taxonomic inflation, diluting the species concept and making taxonomy unstable.[39] 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.[40]

Unlike 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.[29] Therefore, it applies to asexual lineages.[34][35] 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.[35]

Evolutionary species

An evolutionary species, suggested by George Gaylord Simpson in 1951, 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".[7][41] 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 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.[42] A version of the concept is Kevin de Queiroz's "General Lineage Concept of Species".[43]

Ecological species

An ecological species 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.[44]

Genetic species

A genetic species as defined by Robert Baker and Robert Bradley is a set of genetically isolated interbreeding populations. This is similar to Mayr's Biological Species Concept, but stresses genetic rather than reproductive isolation.[45] In the 21st century, a genetic species can be established by comparing DNA sequences, but other methods were available earlier, such as comparing karyotypes (sets of chromosomes) and allozymes (enzyme variants).[46]

Evolutionarily significant unit

An evolutionarily significant unit (ESU) or "wildlife species"[47] is a population of organisms considered distinct for purposes of conservation.[48]

Chronospecies

A chronospecies is defined in a single lineage (solid line) whose morphology changes with time. At some point, palaeontologists judge that enough change has occurred that two species (A and B), separated in time and anatomy, once existed.
A chronospecies is defined in a single lineage (solid line) whose morphology changes with time. At some point, palaeontologists judge that enough change has occurred that two species (A and B), separated in time and anatomy, once existed.

In palaeontology, with only comparative anatomy (morphology) from fossils as evidence, the concept of a chronospecies can be applied. During anagenesis (evolution, not necessarily involving branching), 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, palaeontologists assess how much change is required for a morphologically distinct form to be considered a different species from its ancestors.[49][50][51][52]

Viral quasispecies

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.[53] A viral quasispecies is a group of genotypes related by similar mutations, competing within a highly mutagenic environment, and hence governed by 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 a traditional biological species.[54][55][56] The International Committee on Taxonomy of Viruses has since 1962 developed a universal taxonomic scheme for viruses; this has stabilised viral taxonomy.[57][58][59]

Mayr's biological species concept

Ernst Mayr proposed the widely used Biological Species Concept of reproductive isolation in 1942.
Ernst Mayr proposed the widely used Biological Species Concept of reproductive isolation in 1942.

Most modern textbooks make use of Ernst Mayr's 1942 definition,[60][61] 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 a species as[62]

groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups.[62]

It has been argued that this definition is a natural consequence of the effect of sexual reproduction on the dynamics of natural selection.[63][64][65][66] 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 the wild.[62]

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Morphology (biology)

Morphology (biology)

Morphology is a branch of biology dealing with the study of the form and structure of organisms and their specific structural features.

Genetics

Genetics

Genetics is the study of genes, genetic variation, and heredity in organisms. It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar working in the 19th century in Brno, was the first to study genetics scientifically. Mendel studied "trait inheritance", patterns in the way traits are handed down from parents to offspring over time. He observed that organisms inherit traits by way of discrete "units of inheritance". This term, still used today, is a somewhat ambiguous definition of what is referred to as a gene.

Ernst Mayr

Ernst Mayr

Ernst Walter Mayr was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, philosopher of biology, and historian of science. His work contributed to the conceptual revolution that led to the modern evolutionary synthesis of Mendelian genetics, systematics, and Darwinian evolution, and to the development of the biological species concept.

John Wilkins

John Wilkins

John Wilkins, was an Anglican clergyman, natural philosopher, and author, and was one of the founders of the Royal Society. He was Bishop of Chester from 1668 until his death.

Eurasian blue tit

Eurasian blue tit

The Eurasian blue tit is a small passerine bird in the tit family, Paridae. It is easily recognisable by its blue and yellow plumage and small size.

Drosophila

Drosophila

Drosophila is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit. They should not be confused with the Tephritidae, a related family, which are also called fruit flies ; tephritids feed primarily on unripe or ripe fruit, with many species being regarded as destructive agricultural pests, especially the Mediterranean fruit fly.

Gene pool

Gene pool

The gene pool is the set of all genes, or genetic information, in any population, usually of a particular species.

Gene

Gene

In biology, the word gene can have several different meanings. The Mendelian gene is a basic unit of heredity and the molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protein-coding genes and noncoding genes.

Cytochrome c oxidase

Cytochrome c oxidase

The enzyme cytochrome c oxidase or Complex IV, is a large transmembrane protein complex found in bacteria, archaea, and mitochondria of eukaryotes.

Enzyme

Enzyme

Enzymes are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. Almost all metabolic processes in the cell need enzyme catalysis in order to occur at rates fast enough to sustain life. Metabolic pathways depend upon enzymes to catalyze individual steps. The study of enzymes is called enzymology and the field of pseudoenzyme analysis recognizes that during evolution, some enzymes have lost the ability to carry out biological catalysis, which is often reflected in their amino acid sequences and unusual 'pseudocatalytic' properties.

Microbiology

Microbiology

Microbiology is the scientific study of microorganisms, those being unicellular, multicellular, or acellular. Microbiology encompasses numerous sub-disciplines including virology, bacteriology, protistology, mycology, immunology, and parasitology.

Bacteria

Bacteria

Bacteria are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among the first life forms to appear on Earth, and are present in most of its habitats. Bacteria inhabit soil, water, acidic hot springs, radioactive waste, and the deep biosphere of Earth's crust. Bacteria are vital in many stages of the nutrient cycle by recycling nutrients such as the fixation of nitrogen from the atmosphere. The nutrient cycle includes the decomposition of dead bodies; bacteria are responsible for the putrefaction stage in this process. In the biological communities surrounding hydrothermal vents and cold seeps, extremophile bacteria provide the nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and methane, to energy. Bacteria also live in symbiotic and parasitic relationships with plants and animals. Most bacteria have not been characterised and there are many species that cannot be grown in the laboratory. The study of bacteria is known as bacteriology, a branch of microbiology.

The species problem

It is difficult to define a species in a way that applies to all organisms.[67] The debate about species concepts is called the species problem.[62][68][69][70] The problem was recognised even in 1859, when Darwin wrote in On the Origin of Species:

No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species. Generally the term includes the unknown element of a distinct act of creation.[71]

When Mayr's concept breaks down

Palaeontologists are limited to morphological evidence when deciding whether fossil life-forms like these Inoceramus bivalves formed a separate species.
Palaeontologists are limited to morphological evidence when deciding whether fossil life-forms like these Inoceramus bivalves formed a separate species.

A simple textbook definition, following Mayr's concept, works well for most multi-celled organisms, but breaks down in several situations:

The willow warbler and chiffchaff are almost identical in appearance but do not interbreed.
The willow warbler and chiffchaff are almost identical in appearance but do not interbreed.
The willow warbler and chiffchaff are almost identical in appearance but do not interbreed.

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).[81] In addition, horizontal gene transfer (HGT) makes it difficult to define a species.[82] 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.[83] There is strong evidence of HGT between very dissimilar groups of prokaryotes, and at least occasionally between dissimilar groups of eukaryotes,[82] including some crustaceans and echinoderms.[84]

The evolutionary biologist James Mallet concludes that

there 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.[18]

Wilkins writes that biologists such as the botanist Brent Mishler[85] have argued that the species concept is not valid, and that "if we were being true to evolution and the consequent phylogenetic approach to taxa, we should replace it with a 'smallest clade' idea" (a phylogenetic species concept).[86] Wilkins states that he concurs[87] with this approach, while noting the difficulties it would cause to taxonomy. He cites the ichthyologist Charles Tate Regan's early 20th century remark that "a species is whatever a suitably qualified biologist chooses to call a species".[86] Wilkins notes that the philosopher Philip Kitcher called this the "cynical species concept",[88] and arguing that far from being cynical, it usefully leads to an empirical taxonomy for any given group, based on taxonomists' experience.[86]

Aggregates of microspecies

The species concept is further weakened by the existence of microspecies, groups of organisms, including many plants, with very little genetic variability, usually forming species aggregates.[89] 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,[90] complicated by hybridisation, apomixis and polyploidy, making gene flow between populations difficult to determine, and their taxonomy debatable.[91][92][93] Species complexes occur in insects such as Heliconius butterflies,[94] vertebrates such as Hypsiboas treefrogs,[95] and fungi such as the fly agaric.[96]

Hybridisation

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.[97]

Ring species

A ring species is 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.[98] Such non-breeding, though genetically connected, "end" populations may co-exist in the same region thus closing the ring. Ring species thus present a difficulty for any species concept that relies on reproductive isolation.[99] However, ring species are at best rare. Proposed examples include the herring gulllesser black-backed gull complex around the North pole, the Ensatina eschscholtzii group of 19 populations of salamanders in America,[100] and the greenish warbler in Asia,[101] but many so-called ring species have turned out to be the result of misclassification leading to questions on whether there really are any ring species.[102][103][104][105]

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On the Origin of Species

On the Origin of Species

On the Origin of Species is a work of scientific literature by Charles Darwin that is considered to be the foundation of evolutionary biology; it was published on 24 November 1859. Darwin's book introduced the scientific theory that populations evolve over the course of generations through a process of natural selection. The book presented a body of evidence that the diversity of life arose by common descent through a branching pattern of evolution. Darwin included evidence that he had collected on the Beagle expedition in the 1830s and his subsequent findings from research, correspondence, and experimentation.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Inoceramus

Inoceramus

Inoceramus is an extinct genus of fossil marine pteriomorphian bivalves that superficially resembled the related winged pearly oysters of the extant genus Pteria. They lived from the Early Jurassic to latest Cretaceous.

Asexual reproduction

Asexual reproduction

Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the full set of genes of their single parent and thus the newly created individual is genetically and physically similar to the parent or an exact clone of the parent. Asexual reproduction is the primary form of reproduction for single-celled organisms such as archaea and bacteria. Many eukaryotic organisms including plants, animals, and fungi can also reproduce asexually. In vertebrates, the most common form of asexual reproduction is parthenogenesis, which is typically used as an alternative to sexual reproduction in times when reproductive opportunities are limited. Komodo dragons and some monitor lizards can also reproduce asexually.

Bacteria

Bacteria

Bacteria are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among the first life forms to appear on Earth, and are present in most of its habitats. Bacteria inhabit soil, water, acidic hot springs, radioactive waste, and the deep biosphere of Earth's crust. Bacteria are vital in many stages of the nutrient cycle by recycling nutrients such as the fixation of nitrogen from the atmosphere. The nutrient cycle includes the decomposition of dead bodies; bacteria are responsible for the putrefaction stage in this process. In the biological communities surrounding hydrothermal vents and cold seeps, extremophile bacteria provide the nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and methane, to energy. Bacteria also live in symbiotic and parasitic relationships with plants and animals. Most bacteria have not been characterised and there are many species that cannot be grown in the laboratory. The study of bacteria is known as bacteriology, a branch of microbiology.

Prokaryote

Prokaryote

A prokaryote is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word prokaryote comes from the Greek πρό and κάρυον. In the two-empire system arising from the work of Édouard Chatton, prokaryotes were classified within the empire Prokaryota. But in the three-domain system, based upon molecular analysis, prokaryotes are divided into two domains: Bacteria and Archaea. Organisms with nuclei are placed in a third domain, Eukaryota. In biological evolution, prokaryotes are deemed to have arisen before eukaryotes.

Apomixis

Apomixis

In botany, apomixis is asexual reproduction without fertilization. Its etymology is Greek for "away from" + "mixing". This definition notably does not mention meiosis. Thus "normal asexual reproduction" of plants, such as propagation from cuttings or leaves, has never been considered to be apomixis, but replacement of the seed by a plantlet or replacement of the flower by bulbils were categorized as types of apomixis. Apomictically produced offspring are genetically identical to the parent plant.

DNA barcoding

DNA barcoding

DNA barcoding is a method of species identification using a short section of DNA from a specific gene or genes. The premise of DNA barcoding is that by comparison with a reference library of such DNA sections, an individual sequence can be used to uniquely identify an organism to species, just as a supermarket scanner uses the familiar black stripes of the UPC barcode to identify an item in its stock against its reference database. These "barcodes" are sometimes used in an effort to identify unknown species or parts of an organism, simply to catalog as many taxa as possible, or to compare with traditional taxonomy in an effort to determine species boundaries.

Phylogenetics

Phylogenetics

In biology, phylogenetics is the study of the evolutionary history and relationships among or within groups of organisms. These relationships are determined by phylogenetic inference methods that focus on observed heritable traits, such as DNA sequences, protein amino acid sequences, or morphology. The result of such an analysis is a phylogenetic tree—a diagram containing a hypothesis of relationships that reflects the evolutionary history of a group of organisms.

Hybrid (biology)

Hybrid (biology)

In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Generally, it means that each cell has genetic material from two different organisms, whereas an individual where some cells are derived from a different organism is called a chimera. Hybrids are not always intermediates between their parents, but can show hybrid vigor, sometimes growing larger or taller than either parent. The concept of a hybrid is interpreted differently in animal and plant breeding, where there is interest in the individual parentage. In genetics, attention is focused on the numbers of chromosomes. In taxonomy, a key question is how closely related the parent species are.

Ring species

Ring species

In biology, a ring species is a connected series of neighbouring populations, each of which interbreeds with closely sited 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 and the next. Such non-breeding, though genetically connected, "end" populations may co-exist in the same region (sympatry) thus closing a "ring". The German term Rassenkreis, meaning a circle of races, is also used.

Common chiffchaff

Common chiffchaff

The common chiffchaff, or simply the chiffchaff, is a common and widespread leaf warbler which breeds in open woodlands throughout northern and temperate Europe and the Palearctic.

Taxonomy and naming

A cougar, mountain lion, panther, or puma, among other common names: its scientific name is Puma concolor.
A cougar, mountain lion, panther, or puma, among other common names: its scientific name is Puma concolor.

Common and scientific names

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 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; they are in two parts used together: the genus as in Puma, and the specific epithet as in concolor.[106][107]

Species description

The type specimen (holotype) of Lacerta plica, described by Linnaeus in 1758
The type specimen (holotype) of Lacerta plica, described by Linnaeus in 1758

A species is given a taxonomic name when a type specimen 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[108] (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 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.[109][110][111] 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".[112]

Abbreviations

Books and articles sometimes intentionally do not identify species fully, using 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.[113]

Authors may also use "spp." as 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.[113]

When a species's identity is not clear, 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.[113]

Identification codes

With the rise of online databases, codes have been devised to provide identifiers for species that are already defined, including:

Lumping and splitting

The naming of 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 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.[118][119][113] The circumscription of taxa, considered a taxonomic decision at the discretion of cognizant specialists, is not governed by the Codes of Zoological or Botanical Nomenclature.

Broad and narrow senses

The nomenclatural codes that guide the naming of species, including 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 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.[113][120][121]

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Felidae

Felidae

Felidae is the family of mammals in the order Carnivora colloquially referred to as cats. A member of this family is also called a felid. The term "cat" refers both to felids in general and specifically to the domestic cat.

Jaguar

Jaguar

The jaguar is a large cat species and the only living member of the genus Panthera native to the Americas. With a body length of up to 1.85 m and a weight of up to 158 kg (348 lb), it is the largest cat species in the Americas and the third largest in the world. Its distinctively marked coat features pale yellow to tan colored fur covered by spots that transition to rosettes on the sides, although a melanistic black coat appears in some individuals. The jaguar's powerful bite allows it to pierce the carapaces of turtles and tortoises, and to employ an unusual killing method: it bites directly through the skull of mammalian prey between the ears to deliver a fatal blow to the brain.

Leopard

Leopard

The leopard is one of the five extant species in the genus Panthera, a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, and on the Indian subcontinent to Southeast and East Asia. It is listed as Vulnerable on the IUCN Red List because leopard populations are threatened by habitat loss and fragmentation, and are declining in large parts of the global range. The leopard is considered locally extinct in Hong Kong, Singapore, South Korea, Jordan, Morocco, Togo, the United Arab Emirates, Uzbekistan, Lebanon, Mauritania, Kuwait, Syria, Libya, Tunisia and most likely in North Korea, Gambia, Laos, Lesotho, Tajikistan, Vietnam and Israel. Contemporary records suggest that the leopard occurs in only 25% of its historical global range.

Binomial nomenclature

Binomial nomenclature

In taxonomy, binomial nomenclature, also called binominal nomenclature or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, although they can be based on words from other languages. Such a name is called a binomial name, a binomen, binominal name or a scientific name; more informally it is also historically called a Latin name.

Genus

Genus

Genus is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclature, the genus name forms the first part of the binomial species name for each species within the genus.E.g. Panthera leo (lion) and Panthera onca (jaguar) are two species within the genus Panthera. Panthera is a genus within the family Felidae.

Holotype

Holotype

A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several examples, but explicitly designated as the holotype. Under the International Code of Zoological Nomenclature (ICZN), a holotype is one of several kinds of name-bearing types. In the International Code of Nomenclature for algae, fungi, and plants (ICN) and ICZN, the definitions of types are similar in intent but not identical in terminology or underlying concept.

Plica plica

Plica plica

Plica plica is a species of lizard in the family Tropiduridae, the Neotropical ground lizards. Its common names include collared tree lizard, collared tree runner,tree runner, and harlequin racerunner. In Guyana it is known as wakanama.

Carl Linnaeus

Carl Linnaeus

Carl Linnaeus, also known after his ennoblement in 1761 as Carl von Linné, was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the modern system of naming organisms. He is known as the "father of modern taxonomy". Many of his writings were in Latin; his name is rendered in Latin as Carolus Linnæus and, after his 1761 ennoblement, as Carolus a Linné.

Species description

Species description

A species description is a formal scientific description of a newly encountered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species that have been described previously or are related. To be considered valid, a species description must follow guidelines established over time. Naming requires adherence to respective codes, for example: in zoology, the International Code of Zoological Nomenclature (ICZN); plants, the International Code of Nomenclature for algae, fungi, and plants (ICN); viruses, the International Committee on Taxonomy of Viruses (ICTV). The species description often contains photographs or other illustrations of type material along with a note on where they are deposited. The publication in which the species is described gives the new species a formal scientific name. Some 1.9 million species have been identified and described, out of some 8.7 million that may actually exist. Millions more have become extinct throughout the existence of life on Earth.

Species complex

Species complex

In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each other, further blurring any distinctions. Terms that are sometimes used synonymously but have more precise meanings are cryptic species for two or more species hidden under one species name, sibling species for two species that are each other's closest relative, and species flock for a group of closely related species that live in the same habitat. As informal taxonomic ranks, species group, species aggregate, macrospecies, and superspecies are also in use.

Available name

Available name

In zoological nomenclature, an available name is a scientific name for a taxon of animals that has been published conforming to all the mandatory provisions of the International Code of Zoological Nomenclature for the establishment of a zoological name.

International Code of Zoological Nomenclature

International Code of Zoological Nomenclature

The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the International Commission on Zoological Nomenclature. The rules principally regulate:How names are correctly established in the frame of binominal nomenclature Which name must be used in case of name conflicts How scientific literature must cite names

Change

Species are subject to change, whether by evolving into new species,[122] exchanging genes with other species,[123] merging with other species or by becoming extinct.[124]

Speciation

The evolutionary process by which biological populations evolve to become distinct or reproductively isolated as species is called speciation.[125][126] Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book The Origin of Species.[127] 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 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.[122] A different mechanism, phyletic speciation, involves one lineage gradually changing over time into a new and distinct form, without increasing the number of resultant species.[128]

Exchange of genes between species

Horizontal gene transfers between widely separated species complicate the phylogeny of bacteria.
Horizontal gene transfers between widely separated species complicate the phylogeny of bacteria.

Horizontal gene transfer between organisms of different species, either through hybridisation, antigenic shift, or reassortment, 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.[129][82][130][123]

Louis-Marie Bobay and Howard Ochman suggest, based on analysis of the genomes of many types of bacteria, that they can often be grouped "into communities that regularly swap genes", in much the same way that plants and animals can be grouped into reproductively isolated breeding populations. Bacteria may thus form species, analogous to Mayr's biological species concept, consisting of asexually reproducing populations that exchange genes by homologous recombination.[131][132]

Extinction

A species is extinct when the last individual of that species dies, but it may be functionally extinct well before that moment. It is estimated that over 99 percent of all species that ever lived on Earth, some five billion species, are now extinct. Some of these were in mass extinctions such as those at the ends of the Ordovician, Devonian, Permian, Triassic and Cretaceous periods. Mass extinctions had a variety of causes including volcanic activity, climate change, and changes in oceanic and atmospheric chemistry, and they in turn had major effects on Earth's ecology, atmosphere, land surface and waters.[133][134] Another form of extinction is through the assimilation of one species by another through hybridization. The resulting single species has been termed as a "compilospecies".[135]

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Evolution

Evolution

In biology, evolution is the change in heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation tends to exist within any given population as a result of genetic mutation and recombination. Evolution occurs when evolutionary processes such as natural selection and genetic drift act on this variation, resulting in certain characteristics becoming more common or more rare within a population. The evolutionary pressures that determine whether a characteristic is common or rare within a population constantly change, resulting in a change in heritable characteristics arising over successive generations. It is this process of evolution that has given rise to biodiversity at every level of biological organisation.

Charles Darwin

Charles Darwin

Charles Robert Darwin was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended from a common ancestor is now generally accepted and considered a fundamental concept in science. In a joint publication with Alfred Russel Wallace, he introduced his scientific theory that this branching pattern of evolution resulted from a process he called natural selection, in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding. Darwin has been described as one of the most influential figures in human history and was honoured by burial in Westminster Abbey.

Natural selection

Natural selection

Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which is intentional, whereas natural selection is not.

Allele

Allele

An allele is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution.The word "Allele" is a short form of "allelomorph". "The chromosomal or genomic location of a gene or any other genetic element is called a locus and alternative DNA sequences at a locus are called alleles."

Bateson–Dobzhansky–Muller model

Bateson–Dobzhansky–Muller model

The Bateson–Dobzhansky–Muller model, also known as Dobzhansky–Muller model, is a model of the evolution of genetic incompatibility, important in understanding the evolution of reproductive isolation during speciation and the role of natural selection in bringing it about. The theory was first described by William Bateson in 1909, then independently described by Theodosius Dobzhansky in 1934, and later elaborated in different forms by Herman Muller, H. Allen Orr and Sergey Gavrilets.

Horizontal gene transfer

Horizontal gene transfer

Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the evolution of many organisms. HGT is influencing scientific understanding of higher order evolution while more significantly shifting perspectives on bacterial evolution.

Bacteria

Bacteria

Bacteria are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among the first life forms to appear on Earth, and are present in most of its habitats. Bacteria inhabit soil, water, acidic hot springs, radioactive waste, and the deep biosphere of Earth's crust. Bacteria are vital in many stages of the nutrient cycle by recycling nutrients such as the fixation of nitrogen from the atmosphere. The nutrient cycle includes the decomposition of dead bodies; bacteria are responsible for the putrefaction stage in this process. In the biological communities surrounding hydrothermal vents and cold seeps, extremophile bacteria provide the nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and methane, to energy. Bacteria also live in symbiotic and parasitic relationships with plants and animals. Most bacteria have not been characterised and there are many species that cannot be grown in the laboratory. The study of bacteria is known as bacteriology, a branch of microbiology.

Hybrid (biology)

Hybrid (biology)

In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Generally, it means that each cell has genetic material from two different organisms, whereas an individual where some cells are derived from a different organism is called a chimera. Hybrids are not always intermediates between their parents, but can show hybrid vigor, sometimes growing larger or taller than either parent. The concept of a hybrid is interpreted differently in animal and plant breeding, where there is interest in the individual parentage. In genetics, attention is focused on the numbers of chromosomes. In taxonomy, a key question is how closely related the parent species are.

Antigenic shift

Antigenic shift

Antigenic shift is the process by which two or more different strains of a virus, or strains of two or more different viruses, combine to form a new subtype having a mixture of the surface antigens of the two or more original strains. The term is often applied specifically to influenza, as that is the best-known example, but the process is also known to occur with other viruses, such as visna virus in sheep. Antigenic shift is a specific case of reassortment or viral shift that confers a phenotypic change.

Extinction

Extinction

Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point. Because a species' potential range may be very large, determining this moment is difficult, and is usually done retrospectively. This difficulty leads to phenomena such as Lazarus taxa, where a species presumed extinct abruptly "reappears" after a period of apparent absence.

Extinction event

Extinction event

An extinction event is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. It occurs when the rate of extinction increases with respect to the background extinction rate and the rate of speciation. Estimates of the number of major mass extinctions in the last 540 million years range from as few as five to more than twenty. These differences stem from disagreement as to what constitutes a "major" extinction event, and the data chosen to measure past diversity.

Endling

Endling

An endling is the last known individual of a species or subspecies. Once the endling dies, the species becomes extinct. The word was coined in correspondence in the scientific journal Nature. Alternative names put forth for the last individual of its kind include ender and terminarch.

Practical implications

Biologists and conservationists need to categorise and identify organisms in the course of their work. Difficulty assigning organisms reliably to a species constitutes a threat to the validity of research results, for example making measurements of how abundant a species is in an ecosystem moot. Surveys using a phylogenetic species concept reported 48% more species and accordingly smaller populations and ranges than those using nonphylogenetic concepts; this was termed "taxonomic inflation",[136] which could cause a false appearance of change to the number of endangered species and consequent political and practical difficulties.[137][138] Some observers claim that there is an inherent conflict between the desire to understand the processes of speciation and the need to identify and to categorise.[138]

Conservation laws in many countries make special provisions to prevent species from going extinct. Hybridization zones between two species, one that is protected and one that is not, have sometimes led to conflicts between lawmakers, land owners and conservationists. One of the classic cases in North America is that of the protected northern spotted owl which hybridises with the unprotected California spotted owl and the barred owl; this has led to legal debates.[139] It has been argued that the species problem is created by the varied uses of the concept of species, and that the solution is to abandon it and all other taxonomic ranks, and use unranked monophyletic groups instead. It has been argued, too, that since species are not comparable, counting them is not a valid measure of biodiversity; alternative measures of phylogenetic biodiversity have been proposed.[140][141]

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Conservation biology

Conservation biology

Conservation biology is the study of the conservation of nature and of Earth's biodiversity with the aim of protecting species, their habitats, and ecosystems from excessive rates of extinction and the erosion of biotic interactions. It is an interdisciplinary subject drawing on natural and social sciences, and the practice of natural resource management.

External validity

External validity

External validity is the validity of applying the conclusions of a scientific study outside the context of that study. In other words, it is the extent to which the results of a study can be generalized to and across other situations, people, stimuli, and times. In contrast, internal validity is the validity of conclusions drawn within the context of a particular study. Because general conclusions are almost always a goal in research, external validity is an important property of any study. Mathematical analysis of external validity concerns a determination of whether generalization across heterogeneous populations is feasible, and devising statistical and computational methods that produce valid generalizations.

Ecosystem

Ecosystem

An ecosystem consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the system through photosynthesis and is incorporated into plant tissue. By feeding on plants and on one another, animals play an important role in the movement of matter and energy through the system. They also influence the quantity of plant and microbial biomass present. By breaking down dead organic matter, decomposers release carbon back to the atmosphere and facilitate nutrient cycling by converting nutrients stored in dead biomass back to a form that can be readily used by plants and microbes.

Northern spotted owl

Northern spotted owl

The northern spotted owl is one of three spotted owl subspecies. A western North American bird in the family Strigidae, genus Strix, it is a medium-sized dark brown owl native to the Pacific Northwest. An important indicator species, the northern spotted owl remains threatened due to continued population decline from human-caused habitat destruction and competition with invasive species, its main competitor being the barred owl.

Barred owl

Barred owl

The barred owl, also known as the northern barred owl, striped owl or, more informally, hoot owl or eight-hooter owl, is a North American large species of owl. A member of the true owl family, Strigidae, they belong to the genus Strix, which is also the origin of the family's name under Linnaean taxonomy. Barred owls are largely native to eastern North America, but have expanded their range to the west coast of North America where they are considered invasive. Mature forests are their preferred habitat, but they can also acclimate to various gradients of open woodlands. Their diet consists mainly of small mammals, but this species is an opportunistic predator and is known to prey upon other small vertebrates such as birds, reptiles, and amphibians, as well as a variety of invertebrates.

Biodiversity

Biodiversity

Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic, species, and ecosystem level. Biodiversity is not distributed evenly on Earth; it is usually greater in the tropics as a result of the warm climate and high primary productivity in the region near the equator. Tropical forest ecosystems cover less than 10% of earth's surface and contain about 90% of the world's species. Marine biodiversity is usually higher along coasts in the Western Pacific, where sea surface temperature is highest, and in the mid-latitudinal band in all oceans. There are latitudinal gradients in species diversity. Biodiversity generally tends to cluster in hotspots, and has been increasing through time, but will be likely to slow in the future as a primary result of deforestation. It encompasses the evolutionary, ecological, and cultural processes that sustain life.

History

Classical forms

In his biology, Aristotle used the term γένος (génos) to mean a kind, such as a bird or fish, and εἶδος (eidos) to mean a specific form within a kind, such as (within the birds) the crane, eagle, crow, or sparrow. These terms were translated into Latin as "genus" and "species", though they do not correspond to the Linnean terms thus named; today the birds are a class, the cranes are a family, and the crows a genus. A kind was distinguished by its attributes; for instance, a bird has feathers, a beak, wings, a hard-shelled egg, and warm blood. A form was distinguished by being shared by all its members, the young inheriting any variations they might have from their parents. Aristotle believed all kinds and forms to be distinct and unchanging. His approach remained influential until the Renaissance.[142]

Fixed species

John Ray believed that species breed true and do not change, even though variations exist.
John Ray believed that species breed true and do not change, even though variations exist.

When observers in the Early Modern period began to develop systems of organization for living things, they placed each kind of animal or plant into a context. Many of these early delineation schemes would now be considered whimsical: schemes included consanguinity based on colour (all plants with yellow flowers) or behaviour (snakes, scorpions and certain biting ants). John Ray, an English naturalist, was the first to attempt a biological definition of species in 1686, as follows:

No surer criterion for determining species has occurred to me than the distinguishing features that perpetuate themselves in propagation from seed. Thus, no matter what variations occur in the individuals or the species, if they spring from the seed of one and the same plant, they are accidental variations and not such as to distinguish a species ... Animals likewise that differ specifically preserve their distinct species permanently; one species never springs from the seed of another nor vice versa.[143]

Carl Linnaeus created the binomial system for naming species.
Carl Linnaeus created the binomial system for naming species.

In the 18th century, the Swedish scientist Carl Linnaeus classified organisms according to shared physical characteristics, and not simply based upon differences.[144] He established the idea of a taxonomic hierarchy of classification based upon observable characteristics and intended to reflect natural relationships.[145][146] At the time, however, it was still widely believed that there was no organic connection between species, no matter how similar they appeared. This view was influenced by European scholarly and religious education, which held that the categories of life are dictated by God, forming an Aristotelian hierarchy, the scala naturae or great chain of being. However, whether or not it was supposed to be fixed, the scala (a ladder) inherently implied the possibility of climbing.[147]

Mutability

In viewing evidence of hybridisation, Linnaeus recognised that species were not fixed and could change; he did not consider that new species could emerge and maintained a view of divinely fixed species that may alter through processes of hybridisation or acclimatisation.[148] By the 19th century, naturalists understood that species could change form over time, and that the history of the planet provided enough time for major changes. Jean-Baptiste Lamarck, in his 1809 Zoological Philosophy, described the transmutation of species, proposing that a species could change over time, in a radical departure from Aristotelian thinking.[149]

In 1859, Charles Darwin and Alfred Russel Wallace provided a compelling account of evolution and the formation of new species. Darwin argued that it was populations that evolved, not individuals, by natural selection from naturally occurring variation among individuals.[150] This required a new definition of species. Darwin concluded that species are what they appear to be: ideas, provisionally useful for naming groups of interacting individuals, writing:

I look at the term species as one arbitrarily given for the sake of convenience to a set of individuals closely resembling each other ... It does not essentially differ from the word variety, which is given to less distinct and more fluctuating forms. The term variety, again, in comparison with mere individual differences, is also applied arbitrarily, and for convenience sake.[151]

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Aristotle's biology

Aristotle's biology

Aristotle's biology is the theory of biology, grounded in systematic observation and collection of data, mainly zoological, embodied in Aristotle's books on the science. Many of his observations were made during his stay on the island of Lesbos, including especially his descriptions of the marine biology of the Pyrrha lagoon, now the Gulf of Kalloni. His theory is based on his concept of form, which derives from but is markedly unlike Plato's theory of Forms.

Aristotle

Aristotle

Aristotle was an Ancient Greek philosopher and polymath. His writings cover a broad range of subjects including physics, biology, zoology, metaphysics, logic, ethics, aesthetics, poetry, drama, music, rhetoric, psychology, linguistics, economics, politics, meteorology, geology, and government. As the founder of the Peripatetic school of philosophy in the Lyceum in Athens, he began the wider Aristotelian tradition that followed, which set the groundwork for the development of modern science.

Bird

Bird

Birds are a group of warm-blooded vertebrates constituting the class Aves, characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight skeleton. Birds live worldwide and range in size from the 5.5 cm (2.2 in) bee hummingbird to the 2.8 m common ostrich. There are about ten thousand living species, more than half of which are passerine, or "perching" birds. Birds have wings whose development varies according to species; the only known groups without wings are the extinct moa and elephant birds. Wings, which are modified forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in some birds, including ratites, penguins, and diverse endemic island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly seabirds and some waterbirds, have further evolved for swimming.

Fish

Fish

Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of living fish species are ray-finned fish, belonging to the class Actinopterygii, with around 99% of those being teleosts.

Hylomorphism

Hylomorphism

Hylomorphism is a philosophical doctrine developed by the Ancient Greek philosopher Aristotle, which conceives every physical entity or being (ousia) as a compound of matter (potency) and immaterial form (act), with the generic form as immanently real within the individual. The word is a 19th-century term formed from the Greek words ὕλη and μορφή.

Crane (bird)

Crane (bird)

Cranes are a family, the Gruidae, of large, long-legged, and long-necked birds in the group Gruiformes. The 15 species of cranes are placed in three genera, Antigone, Balearica, and Grus. Unlike the similar-looking but unrelated herons, cranes fly with necks outstretched, not pulled back. Cranes live on most continents, with the exception of Antarctica and South America.

Eagle

Eagle

Eagle is the common name for many large birds of prey of the family Accipitridae. Eagles belong to several groups of genera, some of which are closely related. Most of the 68 species of eagles are from Eurasia and Africa. Outside this area, just 14 species can be found—2 in North America, 9 in Central and South America, and 3 in Australia.

Crow

Crow

A crow is a bird of the genus Corvus, or more broadly a synonym for all of Corvus. Crows are generally black in colour. The word "crow" is used as part of the common name of many species. The related term "raven" is not pinned scientifically to any certain trait, but is rather a general grouping for larger Corvus spp.

Old World sparrow

Old World sparrow

Old World sparrows are a group of small passerine birds forming the family Passeridae. They are also known as true sparrows, a name also used for a particular genus of the family, Passer. They are distinct from both the New World sparrows, in the family Passerellidae, and from a few other birds sharing their name, such as the Java sparrow of the family Estrildidae. Many species nest on buildings and the house and Eurasian tree sparrows, in particular, inhabit cities in large numbers. They are primarily seed-eaters, though they also consume small insects. Some species scavenge for food around cities and, like gulls or pigeons, will eat small quantities of a diversity of items.

Genus

Genus

Genus is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclature, the genus name forms the first part of the binomial species name for each species within the genus.E.g. Panthera leo (lion) and Panthera onca (jaguar) are two species within the genus Panthera. Panthera is a genus within the family Felidae.

John Ray

John Ray

John Ray FRS was a Christian English naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after "having ascertained that such had been the practice of his family before him". He published important works on botany, zoology, and natural theology. His classification of plants in his Historia Plantarum, was an important step towards modern taxonomy. Ray rejected the system of dichotomous division by which species were classified according to a pre-conceived, either/or type system, and instead classified plants according to similarities and differences that emerged from observation. He was among the first to attempt a biological definition for the concept of species, as "a group of morphologically similar organisms arising from a common ancestor". Another significant contribution to taxonomy was his division of plants into those with two seedling leaves (dicotyledons) or only one (monocotyledons), a division used in taxonomy today.

Great chain of being

Great chain of being

The great chain of being is a hierarchical structure of all matter and life, thought by medieval Christianity to have been decreed by God. The chain begins with God and descends through angels, humans, animals and plants to minerals.

Source: "Species", Wikipedia, Wikimedia Foundation, (2023, March 13th), https://en.wikipedia.org/wiki/Species.

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See also
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