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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, is hard or even impossible to test.

Mayden recorded about 24 concepts, [7] and the philosopher of science John Wilkins counted 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.

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. Species named in this manner are called morphospecies. In the s, 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.

A mate-recognition species is a group of sexually reproducing organisms that recognise one another as potential mates. In microbiology , genes can move freely even between distantly related bacteria, possibly extending to the whole bacterial domain. See [20] This concept was narrowed in to a similarity of The average nucleotide identity method quantifies genetic distance between entire genomes, using regions of about 10, base pairs. With enough data from genomes of one genus, algorithms can be used to categorize species, as for Pseudomonas avellanae in , [22] and for all sequenced bacteria and archaea since DNA barcoding has been proposed as a way to distinguish species suitable even for non-specialists to use.

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 “. 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. An evolutionary species, suggested by George Gaylord Simpson in , 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”.

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. 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.

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. An evolutionarily significant unit ESU or “wildlife species” [47] is a population of organisms considered distinct for purposes of conservation. 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.

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. 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. 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.

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, 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. Books and articles sometimes intentionally do not identify species fully, using the abbreviation “sp.

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. 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.

When a species’s identity is not clear, a specialist may use “cf. The abbreviations “nr. With the rise of online databases, codes have been devised to provide identifiers for species that are already defined, including:. 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.

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. Most modern textbooks make use of Ernst Mayr ‘s definition, [75] [76] 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 [77]. It has been argued that this definition is a natural consequence of the effect of sexual reproduction on the dynamics of natural selection.

It is difficult to define a species in a way that applies to all organisms. 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. 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. The evolutionary biologist James Mallet concludes that. 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 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. Blackberries belong to any of hundreds of microspecies of the Rubus fruticosus species aggregate.

The butterfly genus Heliconius contains many similar species. The Hypsiboas calcaratus — fasciatus species complex contains at least six species of treefrog.

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 hybridise freely where their geographical ranges overlap.

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.

Ring species thus present a difficulty for any species concept that relies on reproductive isolation. 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 the result of misclassification leading to questions on whether there really are any ring species.

Seven “species” of Larus gulls interbreed in a ring around the Arctic. Opposite ends of the ring: a herring gull Larus argentatus front and a lesser black-backed gull Larus fuscus in Norway.

A greenish warbler , Phylloscopus trochiloides. Presumed evolution of five “species” of greenish warblers around the Himalayas. 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 evolve to become distinct or reproductively isolated as species is called speciation. 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.

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.

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. 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.

The resulting single species has been termed as a ” compilospecies “. 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. 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.

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. 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. 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 , 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.

In the 18th century, the Swedish scientist Carl Linnaeus classified organisms according to shared physical characteristics, and not simply based upon differences. 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. 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.

Jean-Baptiste Lamarck , in his Zoological Philosophy , described the transmutation of species , proposing that a species could change over time, in a radical departure from Aristotelian thinking. In , 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. 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. From Wikipedia, the free encyclopedia. Basic unit of taxonomic classification, below genus. For other uses, see Species disambiguation. Main article: Chronospecies. Main article: Viral quasispecies. Main article: Species description. Main article: Lumpers and splitters. Main article: Sensu. Main article: Species concept. The willow warbler and chiffchaff are almost identical in appearance but do not interbreed.

Main article: Species complex. Main article: Hybrid biology. Carrion crow. Hybrid with dark belly. Main article: Ring species. Main article: Speciation. Main article: Horizontal gene transfer. Further information: Extinction and Extinction event. Main article: Aristotle’s biology.

Main article: Great chain of being. The New York Times. ISSN Retrieved 25 January Associated Press. PLOS Biology.

PMC PMID Scientific American. Archived from the original on 14 March Retrieved 14 March Systematic Zoology. JSTOR Claridge, M. A hierarchy of species concepts: the denouement of the species problem. Systematics Association. Kingfisher Books. ISBN New York: W. Evolutionary Genetics. Oxford University Press. The British Journal for the Philosophy of Science.

S2CID Systematic Biology. Vrba, E. Monograph No. Species and Speciation. Pretoria: Transvaal Museum. In Calow, P. Encyclopaedia of Ecology and Environmental Management. Archived PDF from the original on 19 October Archived from the original on 7 February International Journal of Systematic Bacteriology. Microbiology Today.

Archived PDF from the original on 25 March This article contains quotations from this source, which is available under the Creative Commons Attribution 4. September Nature Biotechnology. Barcode of Life. Archived from the original on 1 July Retrieved 11 October Molecular Ecology Notes. Fungal Genetics and Biology. In Hodkinson, T.

Inside each sculpture there is a personal memento of the victim. Syracuse University holds a memorial week every year called “Remembrance Week” to commemorate its 35 lost students. Every 21 December, a service is held in the university’s chapel at UTC , marking the moment the bomb on board the aircraft was detonated.

In addition, the university annually awards 35 scholarships to seniors to honour each of the 35 students killed. At Cornell University funds from the Libyan payment were used to establish a memorial professorship in honor of student Kenneth J. This mother tragically lost her son in the bombing of the Pan Am Flight While in Lockerbie, 7 years after the flight, she meets the women who witnessed and were affected by the crash itself while she attempts to find closure.

The main UK memorial is at Dryfesdale Cemetery about one mile 1. There is a semicircular stone wall in the garden of remembrance with the names and nationalities of all the victims along with individual funeral stones and memorials. Inside the chapel at Dryfesdale there is a book of remembrance. There are memorials in Lockerbie and Moffat Roman Catholic churches, where plaques list the names of all victims.

In Lockerbie Town Hall Council Chambers, there is a stained-glass window depicting flags of the 21 countries whose citizens lost their lives in the disaster. There is also a book of remembrance at Lockerbie public library and another at Tundergarth Church.

The Air Accidents Investigation Branch reassembled a large part of the fuselage to aid with the investigation; this has been retained as evidence and stored in a hangar at Farnborough Airport since the bombing. It was announced in April that part of the wreckage was transferred to a secure location in Dumfries , Scotland, and that it remains evidence in the ongoing criminal investigation. From Wikipedia, the free encyclopedia.

Transatlantic flight terrorist bombed in For the hijacking in Karachi, see Pan Am Flight For other uses, see Flight disambiguation. Main article: Pan Am Flight bombing investigation. Main article: Pan Am Flight bombing trial. Further information: Release of Abdelbaset al-Megrahi. Main article: Pan Am Flight conspiracy theories. Archived from the original on 26 March Retrieved 8 June Aviation Safety Network.

Retrieved 27 February BBC News. The Independent. Archived from the original on 24 May Retrieved 13 June The Guardian. ISSN Retrieved 21 December Federal Aviation Administration. Archived from the original on 30 April San Francisco Chronicle. Associated Press. The jumbo jet that crashed The Pan Am The first ever delivered to an airline—also Pan Am—entered the fleet the previous month, said David Jimenez, spokesman for Boeing Commercial Airplanes, which builds s in Everett.

The Washington Post. Retrieved 21 May Times of London. Archived from the original on 21 December Lockerbie, the real story. Bloomsbury Publishing Ltd. Archived from the original on 14 May Retrieved 5 June Retrieved 8 September Retrieved 16 March Medico-Legal Journal. PMID S2CID Retrieved 15 October The New York Times.

Retrieved 5 April Retrieved 30 July Omnibus Press. ISBN Retrieved 21 August Retrieved 5 August Retrieved 2 September Retrieved 17 December Retrieved 13 December Remembering Lockerbie 20 years on.

The Scotsman. Executive Order US Government Printing Office. Retrieved 15 May The Times. Retrieved 31 October Archived from the original on 1 February Archived from the original on 15 October Retrieved 23 February Archived from the original on 26 February Archived from the original on 16 March Retrieved 10 June Channel 4 News.

Advocate Report of Steven E. Clark, Professor of Psychology at the University of California. Archived from the original on 23 September Archived from the original PDF on 17 December Retrieved 10 November Scottish Criminal Cases Review Commission. Retrieved 15 September Retrieved 9 May Retrieved 1 November Retrieved 3 January Archived from the original on 3 March Archived from the original on 3 May BBC interview of Dr.

Fresh doubts on Lockerbie conviction. Glasgow Herald. Archived from the original on 29 January Retrieved 15 January Scottish Courts. Retrieved 23 August Kuwait Times. Retrieved 8 February Archived from the original on 27 September Retrieved 25 August Retrieved 20 May Retrieved 10 September White House.

Gulf of Sidra incident. US Department of State Bulletin. Flashback: The Berlin disco bombing. Retrieved 4 June Revealed: Gaddafi’s air massacre plot. Archived from the original on 23 October Yorkshire Post. Retrieved 2 November The Firm magazine. Archived from the original on 24 October Times of Malta.

Daily Telegraph. November Archived from the original on 28 December Retrieved 12 January Retrieved 24 January Archived from the original on 13 August Archived from the original on 7 July Retrieved 16 September Archived from the original on 10 April Archived from the original on 3 December Archived from the original on 8 March Archived from the original on 27 January Retrieved 14 August Al Jazeera.

Retrieved 3 November Retrieved 29 August Retrieved 19 October New York Post. Archived from the original on 30 August Retrieved 7 March Los Angeles Times. Retrieved 9 August Victims of Pan Am Flight Archived from the original on 14 October ABC News. Orlando Sentinel. Adventures in Education.

University of Rochester. Retrieved 17 March Pan AM the bombing, the betrayals, and a bereaved family’s search for justice. New American Library.

Syracuse, NY: Syracuse University. Document No. Retrieved 31 May Retrieved 18 July Archived from the original on 17 April Retrieved 8 January Retrieved 25 April Washington, DC: U. Government Printing Office. Emerson, Steven ; Duffy, Brian Cox, Matthew; Foster, Tom Johnston, David Lockerbie: The True Story. Trail of the Octopus.

Ashton, John; Ferguson, Ian Ashton, John Megrahi: You Are My Jury. Bannon, Kevin How Abdelbaset al-Megrahi became convicted for the Lockerbie Bombing. Kerr, Morag Adequately Explained by Stupidity? Brown, David A. Aviation Week and Space Technology. Shifrin, Carole A. Cowell, Alan 29 June New York Times. The Scottish indictment against Megrahi and Fhimah , 13 November Retrieved 27 February The U. Retrieved 27 February Aviation Safety Network summary report.

Retrieved 27 February The cost of the trial. Retrieved 26 February The Lockerbie Trial. Retrieved 29 December Rowan, Roy 27 April Archived from the original on 13 February Retrieved 25 February Retrieved 26 February Website set up by supporters of Megrahi , not recently updated. Retrieved 27 February “Court told how jet’s radar blip broke up at 7. Retrieved 28 February Cassell, Andrew 21 December Rizzo, Patrick 29 May The Namibian.

Archived from the original on 4 January Kirkup, James 11 April Archived from the original on 6 July Duffy, Brian 18 November Retrieved 16 June Thatcher, Margaret The Downing Street Years.

Koechler, H. Documents Related to the I. Observer Mission. Studies in International Relations, Vol. Retrieved Cohen, Dan and Susan Dornstein, Ken Leppard, David On the Trail of Terror.

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Pan Am. Juan Trippe. List of Pan Am destinations. Pan Am Express.