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89-540: Nothofagus , also known as the southern beeches , is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and east and southeast Australia, New Zealand, New Guinea , and New Caledonia . The species are ecological dominants in many temperate forests in these regions. Some species are reportedly naturalised in Germany and Great Britain. The genus has

178-423: A mutation–selection balance . It is predicted that a viral quasispecies at a low but evolutionarily neutral and highly connected (that is, flat) region in the fitness landscape will outcompete a quasispecies located at a higher but narrower fitness peak in which the surrounding mutants are unfit, "the quasispecies effect" or the "survival of the flattest". There is no suggestion that a viral quasispecies resembles

267-400: A "classical" method of determining species, such as with Linnaeus, early in evolutionary theory. However, different phenotypes are not necessarily different species (e.g. a four-winged Drosophila born to a two-winged mother is not a different species). Species named in this manner are called morphospecies . In the 1970s, Robert R. Sokal , Theodore J. Crovello and Peter Sneath proposed

356-424: A 'smallest clade' idea" (a phylogenetic species concept). Mishler and Wilkins and others concur with this approach, even though this would raise difficulties in biological nomenclature. Wilkins cited the ichthyologist Charles Tate Regan 's early 20th century remark that "a species is whatever a suitably qualified biologist chooses to call a species". Wilkins noted that the philosopher Philip Kitcher called this

445-428: A connected series of neighbouring populations, each of which can sexually interbreed with adjacent related populations, but for which there exist at least two "end" populations in the series, which are too distantly related to interbreed, though there is a potential gene flow between each "linked" population. Such non-breeding, though genetically connected, "end" populations may co-exist in the same region thus closing

534-464: A cooler time. Sexual reproduction has since been shown to be possible. The genus Nothofagus was first formally described in 1850 by Carl Ludwig Blume who published the description in his book Museum botanicum Lugduno-Batavum, sive, Stirpium exoticarum novarum vel minus cognitarum ex vivis aut siccis brevis expositio et descriptio . Nothofagus means "false beech", which Blume chose to indicate that Nothofagus species were different from beeches in

623-432: A different species from its ancestors. Viruses have enormous populations, are doubtfully living since they consist of little more than a string of DNA or RNA in a protein coat, and mutate rapidly. All of these factors make conventional species concepts largely inapplicable. A viral quasispecies is a group of genotypes related by similar mutations, competing within a highly mutagenic environment, and hence governed by

712-508: A genetic boundary suitable for defining a species concept is present. DNA barcoding has been proposed as a way to distinguish species suitable even for non-specialists to use. One of the barcodes is a region of mitochondrial DNA within the gene for cytochrome c oxidase . A database, Barcode of Life Data System , contains DNA barcode sequences from over 190,000 species. However, scientists such as Rob DeSalle have expressed concern that classical taxonomy and DNA barcoding, which they consider

801-505: A heavier crop of seeds and is known as the beech mast . In New Zealand, the beech mast causes an increase in the population of introduced mammals such as mice, rats, and stoats . When the rodent population collapses, the stoats begin to prey on native bird species, many of which are threatened with extinction. This phenomenon is covered in more detail in the article on stoats in New Zealand . Species A species ( pl. : species)

890-465: A lineage should be divided into multiple chronospecies , or when populations have diverged to have enough distinct character states to be described as cladistic species. Species and higher taxa were seen from the time of Aristotle until the 18th century as categories that could be arranged in a hierarchy, the great chain of being . In the 19th century, biologists grasped that species could evolve given sufficient time. Charles Darwin 's 1859 book On

979-492: A misnomer, need to be reconciled, as they delimit species differently. Genetic introgression mediated by endosymbionts and other vectors can further make barcodes ineffective in the identification of species. A phylogenetic or cladistic species is "the smallest aggregation of populations (sexual) or lineages (asexual) diagnosable by a unique combination of character states in comparable individuals (semaphoronts)". The empirical basis – observed character states – provides

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1068-449: A particular species, including which genus (and higher taxa) it is placed in, is a hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further information comes to hand, the hypothesis may be corroborated or refuted. Sometimes, especially in the past when communication was more difficult, taxonomists working in isolation have given two distinct names to individual organisms later identified as

1157-493: A peak abundance during the Miocene. Their distribution contracted westwards during the late Miocene due to the aridification of Patagonia. Although the genus now mostly occurs in cool, isolated, high-altitude environments at temperate and tropical latitudes , the fossil record shows that it survived in climates that appear to be much warmer than those that Nothofagus now occupies. Nothofagus species are used as food plants by

1246-400: A perfect model of life, it is still a 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

1335-641: A rich fossil record of leaves, cupules , and pollen, with fossils extending into the late Cretaceous period and occurring in Australia, New Zealand, Antarctica, and South America. The leaves are toothed or entire, evergreen or deciduous . The fruit is a small, flattened or triangular nut , borne in cupules containing one to seven nuts. Many individual trees are extremely old, and at one time, some populations were thought to be unable to reproduce in present-day conditions where they were growing, except by suckering ( clonal reproduction ), being remnant forest from

1424-551: A series of continental glaciations in the Quaternary Period that followed. In ICS terminology, from upper (later, more recent) to lower (earlier): The Pliocene Epoch is subdivided into two ages: The Miocene Epoch is subdivided into six ages: In different geophysical regions of the world, other regional names are also used for the same or overlapping ages and other timeline subdivisions. The terms Neogene System (formal) and Upper Tertiary System (informal) describe

1513-400: A short way of saying that something applies to many species within a genus, but not to all. If scientists mean that something applies to all species within a genus, they use the genus name without the specific name or epithet. The names of genera and species are usually printed in italics . However, abbreviations such as "sp." should not be italicised. When a species' identity is not clear,

1602-404: A specialist may use "cf." before the epithet to indicate that confirmation is required. The abbreviations "nr." (near) or "aff." (affine) may be used when the identity is unclear but when the species appears to be similar to the species mentioned after. With the rise of online databases, codes have been devised to provide identifiers for species that are already defined, including: The naming of

1691-523: A species as groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups. It has been argued that this definition is a natural consequence of the effect of sexual reproduction on the dynamics of natural selection. Mayr's use of the adjective "potentially" has been a point of debate; some interpretations exclude unusual or artificial matings that occur only in captivity, or that involve animals capable of mating but that do not normally do so in

1780-400: A species as determined by a taxonomist. A typological species is a group of organisms in which individuals conform to certain fixed properties (a type), so that even pre-literate people often recognise the same taxon as do modern taxonomists. The clusters of variations or phenotypes within specimens (such as longer or shorter tails) would differentiate the species. This method was used as

1869-488: A species. All species definitions assume that an organism acquires its genes from one or two parents very like the "daughter" organism, but that is not what happens in HGT. There is strong evidence of HGT between very dissimilar groups of prokaryotes , and at least occasionally between dissimilar groups of eukaryotes , including some crustaceans and echinoderms . The evolutionary biologist James Mallet concludes that there

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1958-685: A species. Generally the term includes the unknown element of a distinct act of creation. Many authors have argued that a simple textbook definition, following Mayr's concept, works well for most multi-celled organisms , but breaks down in several situations: Species identification is made difficult by discordance between molecular and morphological investigations; these can be categorised as two types: (i) one morphology, multiple lineages (e.g. morphological convergence , cryptic species ) and (ii) one lineage, multiple morphologies (e.g. phenotypic plasticity , multiple life-cycle stages). In addition, horizontal gene transfer (HGT) makes it difficult to define

2047-519: A taxonomic decision at the discretion of cognizant specialists, is not governed by the Codes of Zoological or Botanical Nomenclature, in contrast to the PhyloCode , and contrary to what is done in several other fields, in which the definitions of technical terms, like geochronological units and geopolitical entities, are explicitly delimited. The nomenclatural codes that guide the naming of species, including

2136-506: A traditional biological species. The International Committee on Taxonomy of Viruses has since 1962 developed a universal taxonomic scheme for viruses; this has stabilised viral taxonomy. Most modern textbooks make use of Ernst Mayr 's 1942 definition, known as the Biological Species Concept as a basis for further discussion on the definition of species. It is also called a reproductive or isolation concept. This defines

2225-447: A variation on the morphological species concept, a phenetic species, defined as a set of organisms with a similar phenotype to each other, but a different phenotype from other sets of organisms. It differs from the morphological species concept in including a numerical measure of distance or similarity to cluster entities based on multivariate comparisons of a reasonably large number of phenotypic traits. A mate-recognition species

2314-515: A variety of reasons. Viruses are a special case, driven by a balance of mutation and selection , and can be treated as quasispecies . Biologists and taxonomists have made many attempts to define species, beginning from morphology and moving towards genetics . Early taxonomists such as Linnaeus had no option but to describe what they saw: this was later formalised as the typological or morphological species concept. Ernst Mayr emphasised reproductive isolation, but this, like other species concepts,

2403-438: Is "an entity composed of organisms which maintains its identity from other such entities through time and over space, and which has its own independent evolutionary fate and historical tendencies". This differs from the biological species concept in embodying persistence over time. Wiley and Mayden stated that they see the evolutionary species concept as "identical" to Willi Hennig 's species-as-lineages concept, and asserted that

2492-400: Is a group of sexually reproducing organisms that recognise one another as potential mates. Expanding on this to allow for post-mating isolation, a cohesion species is the most inclusive population of individuals having the potential for phenotypic cohesion through intrinsic cohesion mechanisms; no matter whether populations can hybridise successfully, they are still distinct cohesion species if

2581-452: Is a population of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring , typically by sexual reproduction . It 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

2670-458: Is a set of organisms adapted to a particular set of resources, called a niche, in the environment. According to this concept, populations form the discrete phenetic clusters that we recognise as species because the ecological and evolutionary processes controlling how resources are divided up tend to produce those clusters. A genetic species as defined by Robert Baker and Robert Bradley is a set of genetically isolated interbreeding populations. This

2759-414: Is called speciation . Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book The Origin of Species . Speciation depends on a measure of reproductive isolation , a reduced gene flow. This occurs most easily in allopatric speciation, where populations are separated geographically and can diverge gradually as mutations accumulate. Reproductive isolation

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2848-412: 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,

2937-403: Is described formally, in a publication that assigns it a unique scientific name. The description typically provides means for identifying the new species, which may not be based solely on morphology (see cryptic species ), differentiating it from other previously described and related or confusable species and provides a validly published name (in botany) or an available name (in zoology) when

3026-475: Is due to the comparatively fine divisibility of time units as time approaches the present, and due to geological preservation that causes the youngest sedimentary geological record to be preserved over a much larger area and to reflect many more environments than the older geological record. By dividing the Cenozoic Era into three (arguably two) periods ( Paleogene , Neogene, Quaternary ) instead of seven epochs,

3115-671: Is further weakened by the existence of microspecies , groups of organisms, including many plants, with very little genetic variability, usually forming species aggregates . For example, the dandelion Taraxacum officinale and the blackberry Rubus fruticosus are aggregates with many microspecies—perhaps 400 in the case of the blackberry and over 200 in the dandelion, complicated by hybridisation , apomixis and polyploidy , making gene flow between populations difficult to determine, and their taxonomy debatable. Species complexes occur in insects such as Heliconius butterflies, vertebrates such as Hypsiboas treefrogs, and fungi such as

3204-726: Is hard or even impossible to test. Later biologists have tried to refine Mayr's definition with the recognition and cohesion concepts, among others. Many of the concepts are quite similar or overlap, so they are not easy to count: the biologist R. L. Mayden recorded about 24 concepts, and the philosopher of science John Wilkins counted 26. Wilkins further grouped the species concepts into seven basic kinds of concepts: (1) agamospecies for asexual organisms (2) biospecies for reproductively isolated sexual organisms (3) ecospecies based on ecological niches (4) evolutionary species based on lineage (5) genetic species based on gene pool (6) morphospecies based on form or phenotype and (7) taxonomic species,

3293-403: Is no easy way to tell whether related geographic or temporal forms belong to the same or different species. Species gaps can be verified only locally and at a point of time. One is forced to admit that Darwin's insight is correct: any local reality or integrity of species is greatly reduced over large geographic ranges and time periods. The botanist Brent Mishler argued that the species concept

3382-468: Is not valid, notably because gene flux decreases gradually rather than in discrete steps, which hampers objective delimitation of species. Indeed, complex and unstable patterns of gene flux have been observed in cichlid teleosts of the East African Great Lakes . Wilkins argued that "if we were being true to evolution and the consequent phylogenetic approach to taxa, we should replace it with

3471-675: Is often seen as an analogous climate to the projected climate of the near future as a result of anthropogenic global warming . Towards the end of the Pliocene, decreased heat transport towards the Antarctic resulting from a weakening of the Indonesian Throughflow (ITF) cooled the Earth, a process that exacerbated itself in a positive feedback as sea levels dropped and the ITF diminished and further limited

3560-586: Is similar to Mayr's Biological Species Concept, but stresses genetic rather than reproductive isolation. In the 21st century, a genetic species could be established by comparing DNA sequences. Earlier, other methods were available, such as comparing karyotypes (sets of chromosomes ) and allozymes ( enzyme variants). An evolutionarily significant unit (ESU) or "wildlife species" is a population of organisms considered distinct for purposes of conservation. In palaeontology , with only comparative anatomy (morphology) and histology from fossils as evidence,

3649-618: 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. Neogene During this period, mammals and birds continued to evolve into modern forms, while other groups of life remained relatively unchanged. The first humans ( Homo habilis ) appeared in Africa near

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3738-594: Is threatened by hybridisation, but this can be selected against once a pair of populations have incompatible alleles of the same gene, as described in the Bateson–Dobzhansky–Muller model . A different mechanism, phyletic speciation, involves one lineage gradually changing over time into a new and distinct form (a chronospecies ), without increasing the number of resultant species. Horizontal gene transfer between organisms of different species, either through hybridisation , antigenic shift , or reassortment ,

3827-523: 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

3916-704: The Late Cretaceous . During the Campanian Nothofagus diversified and became dominant within Antarctic ecosystems, with the appearance of all four modern subgenera by the end of the stage. Nothofagus shows a progressive decline in the Antarctic pollen record through the Maastrichtian , before substantially recovering after the Cretaceous-Paleogene boundary . Nothofagus persisted in Antarctica deep into

4005-640: The Northern Hemisphere . In the past, they were included in the family Fagaceae , but genetic tests revealed them to be genetically distinct, and they are now included in their own family, Nothofagaceae . The following is a list of species, hybrids and varieties accepted by the Plants of the World Online as of April 2023: Four subgenera are recognized, based on morphology and DNA analysis: In 2013, Peter Brian Heenan and Rob D. Smissen proposed splitting

4094-745: The Paleogene . The Early Miocene was relatively cool; Early Miocene mid-latitude seawater and continental thermal gradients were already very similar to those of the present. During the Middle Miocene , Earth entered a warm phase known as the Middle Miocene Climatic Optimum (MMCO), which was driven by the emplacement of the Columbia River Basalt Group . Around 11 Ma, the Middle Miocene Warm Interval gave way to

4183-399: The fly agaric . Natural hybridisation presents a challenge to the concept of a reproductively isolated species, as fertile hybrids permit gene flow between two populations. For example, the carrion crow Corvus corone and the hooded crow Corvus cornix appear and are classified as separate species, yet they can hybridise where their geographical ranges overlap. A ring species is

4272-498: The jaguar ( Panthera onca ) of Latin America or the leopard ( Panthera pardus ) of Africa and Asia. In contrast, the scientific names of species are chosen to be unique and universal (except for some inter-code homonyms ); they are in two parts used together : the genus as in Puma , and the specific epithet as in concolor . A species is given a taxonomic name when a type specimen

4361-497: The larvae of hepialid moths of the genus Aenetus , including A. eximia and A. virescens . Zelopsis nothofagi is a leaf hopper, endemic to New Zealand, which is found on Nothofagus . Cyttaria is genus of ascomycete fungi found on or associated with Nothofagus in Australia and South America. Misodendrum are specialist parasitic plants found on various species of Nothofagus in South America. Additionally,

4450-406: The "cynical species concept", and arguing that far from being cynical, it usefully leads to an empirical taxonomy for any given group, based on taxonomists' experience. Other biologists have gone further and argued that we should abandon species entirely, and refer to the "Least Inclusive Taxonomic Units" (LITUs), a view that would be coherent with current evolutionary theory. The species concept

4539-525: The Cenozoic, despite the increasingly inhospitable conditions, with the final records from the late Neogene , around 15-5 million years old, which were small tundra-adapted prostrate shrubs , similar to Salix arctica (Arctic willow). Nothofagus first appeared in southern South America during the late Campanian. During the Paleocene and Eocene they were mostly restricted to southern Patagonia, before reaching

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4628-497: The Miocene-Pliocene boundary. The first hominins , the ancestors of humans, may have appeared in southern Europe and migrated into Africa. The first humans (belonging to the species Homo habilis ) appeared in Africa near the end of the period. About 20 million years ago gymnosperms in the form of some conifer and cycad groups started to diversify and produce more species due to the changing conditions. In response to

4717-692: The Neogene and the Pliocene end at 2.58 Ma, that the Gelasian be transferred to the Pleistocene, and the Quaternary be recognized as the third period in the Cenozoic, citing key changes in Earth's climate, oceans, and biota that occurred 2.58 Ma and its correspondence to the Gauss-Matuyama magnetostratigraphic boundary . In 2006 ICS and INQUA reached a compromise that made Quaternary a sub-era, subdividing Cenozoic into

4806-622: 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

4895-546: The Pliocene Warm Interval (PWI), interrupting the longer-term cooling trend. The Pliocene Thermal Maximum (PTM) occurred between 3.3 and 3.0 Ma. During the Pliocene, Green Sahara phases of wet conditions in North Africa were frequent and occurred about every 21 kyr, being especially intense when Earth's orbit's eccentricity was high. The PWI had similar levels of atmospheric carbon dioxide to contemporary times and

4984-405: The abbreviation "sp." in the singular or "spp." (standing for species pluralis , Latin for "multiple species") in the plural in place of the specific name or epithet (e.g. Canis sp.). This commonly occurs when authors are confident that some individuals belong to a particular genus but are not sure to which exact species they belong, as is common in paleontology . Authors may also use "spp." as

5073-570: The amount of hybridisation is insufficient to completely mix their respective gene pools . A further development of the recognition concept is provided by the biosemiotic concept of species. In microbiology , genes can move freely even between distantly related bacteria, possibly extending to the whole bacterial domain. As a rule of thumb, microbiologists have assumed that members of Bacteria or Archaea with 16S ribosomal RNA gene sequences more similar than 97% to each other need to be checked by DNA–DNA hybridisation to decide if they belong to

5162-532: The atmosphere, C 4 plants expanded and reached ecological dominance in grasslands during the last 10 million years. Also Asteraceae (daisies) went through a significant adaptive radiation . Eucalyptus fossil leaves occur in the Miocene of New Zealand, where the genus is not native today, but have been introduced from Australia. The Neogene traditionally ended at the end of the Pliocene Epoch, just before

5251-726: The beetle, Brachysternus prasinus , has been known to live in Nothofagus in Chile and in parts of Argentina. The geographic range of B. prasinus is highly dependent on the availability and distribution of Nothofagus on which B. prasinus is believed to feed. B. prasinus have been observed in the Nothofagus forests near the cities of Coquimbo and Llanquihue in Chile as well as the areas of Neuquén and Chubut in Western Argentina. The species of subgenus Brassospora are evergreen, and distributed in

5340-474: The biological species concept, "the several versions" of the phylogenetic species concept, and the idea that species are of the same kind as higher taxa are not suitable for biodiversity studies (with the intention of estimating the number of species accurately). They further suggested that the concept works for both asexual and sexually-reproducing species. A version of the concept is Kevin de Queiroz 's "General Lineage Concept of Species". An ecological species

5429-505: The biological species concept, a cladistic species does not rely on reproductive isolation – its criteria are independent of processes that are integral in other concepts. Therefore, it applies to asexual lineages. However, it does not always provide clear cut and intuitively satisfying boundaries between taxa, and may require multiple sources of evidence, such as more than one polymorphic locus, to give plausible results. An evolutionary species, suggested by George Gaylord Simpson in 1951,

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5518-428: 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

5607-433: The concept of a chronospecies can be applied. During anagenesis (evolution, not necessarily involving branching), some palaeontologists seek to identify a sequence of species, each one derived from the phyletically extinct one before through continuous, slow and more or less uniform change. In such a time sequence, some palaeontologists assess how much change is required for a morphologically distinct form to be considered

5696-416: 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. About 14% of these had been described by 2011. All species (except viruses ) are given a two-part name , called a "binomial". The first part of a binomial is the genus to which the species belongs. The second part

5785-459: The cooler, seasonal climate, tropical plant species gave way to deciduous ones and grasslands replaced many forests. Grasses therefore greatly diversified, and herbivorous mammals evolved alongside it, creating the many grazing animals of today such as horses , antelope , and bison . Ice age mammals like the mammoths and woolly rhinoceros were common in Pliocene . With lower levels of CO 2 in

5874-482: The distribution of Nothofagus derives from the break-up of Gondwana (i.e. vicariance ), or if long-distance dispersal has occurred across oceans. In South America, the northern limit of the genus can be construed as La Campana National Park and the Vizcachas Mountains in the central part of Chile. Nothofagus first appeared in Antarctica during the early Campanian stage (83.6 to 72.1 million years ago) of

5963-680: The end of the period. Some continental movements took place, the most significant event being the connection of North and South America at the Isthmus of Panama , late in the Pliocene. This cut off the warm ocean currents from the Pacific to the Atlantic Ocean, leaving only the Gulf Stream to transfer heat to the Arctic Ocean . The global climate cooled considerably throughout the Neogene, culminating in

6052-435: The evidence to support hypotheses about evolutionarily divergent lineages that have maintained their hereditary integrity through time and space. Molecular markers may be used to determine diagnostic genetic differences in the nuclear or mitochondrial DNA of various species. For example, in a study done on fungi , studying the nucleotide characters using cladistic species produced the most accurate results in recognising

6141-464: The genus into four, turning the four recognized subgenera into the new genera Fuscospora , Lophozonia and Trisyngyne , with the five South American species of subgenus Nothofagus remaining in genus Nothofagus . The proposed new genera are not accepted at the World Checklist of Selected Plant Families . The following additional species are listed as extinct: The pattern of distribution around

6230-460: The heat transported southward by the Leeuwin Current . By the end of the period the first of a series of glaciations of the current Ice Age began. Marine and continental flora and fauna have a modern appearance. The reptile group Choristodera went extinct in the early part of the period, while the amphibians known as Allocaudata disappeared at the end of it. Neogene also marked the end of

6319-452: The lower montane forests, and below the conifer -dominated upper montane forests. Nothofagus grows in mixed stands with trees of other species or in pure stands, particularly on ridge crests and upper slopes. The Central Range has the greatest diversity of species, with fewer species distributed among the mountains of western and northern New Guinea, New Britain, and Goodenough and Normanby islands. The New Caledonian species are endemic to

6408-406: The main island (Grand Terre), most commonly on soils derived from ultramafic rocks between 150 and 1350 meters elevation. They occur in isolated stands, forming a low or stunted and irregular and fairly open canopy. The conifers Agathis and Araucaria are sometimes present as emergents, rising 10 to 20 meters above the Nothofagus canopy. Every four to six years or so, Nothofagus produces

6497-477: The much cooler Late Miocene. The ice caps on both poles began to grow and thicken, a process enhanced by positive feedbacks from increased formation of sea ice. Between 7 and 5.3 Ma, a decrease in global temperatures termed the Late Miocene Cooling (LMC) ensued, driven by decreases in carbon dioxide concentrations. During the Pliocene, from about 5.3 to 2.7 Ma, another warm interval occurred, being known as

6586-478: The numerous fungi species of all the concepts studied. Versions of the phylogenetic species concept that emphasise monophyly or diagnosability may lead to splitting of existing species, for example in Bovidae , by recognising old subspecies as species, despite the fact that there are no reproductive barriers, and populations may intergrade morphologically. Others have called this approach taxonomic inflation , diluting

6675-595: The old classical Tertiary and Quaternary, a compromise that was rejected by International Union of Geological Sciences because it split both Neogene and Pliocene in two. Following formal discussions at the 2008 International Geological Congress in Oslo, Norway, the ICS decided in May 2009 to make the Quaternary the youngest period of the Cenozoic Era with its base at 2.58 Mya and including

6764-503: The older definition of the beginning of the Quaternary Period; many time scales show this division. However, there was a movement amongst geologists (particularly marine geologists ) to also include ongoing geological time (Quaternary) in the Neogene, while others (particularly terrestrial geologists) insist the Quaternary to be a separate period of distinctly different record. The somewhat confusing terminology and disagreement amongst geologists on where to draw what hierarchical boundaries

6853-585: The paper is accepted for publication. The type material is usually held in a permanent repository, often the research collection of a major museum or university, that allows independent verification and the means to compare specimens. Describers of new species are asked to choose names that, in the words of the International Code of Zoological Nomenclature , are "appropriate, compact, euphonious, memorable, and do not cause offence". Books and articles sometimes intentionally do not identify species fully, using

6942-616: The periods are more closely comparable to the duration of periods in the Mesozoic and Paleozoic Eras. The International Commission on Stratigraphy (ICS) once proposed that the Quaternary be considered a sub-era (sub-erathem) of the Neogene, with a beginning date of 2.58 Ma, namely the start of the Gelasian Stage . In the 2004 proposal of the ICS, the Neogene would have consisted of the Miocene and Pliocene Epochs. The International Union for Quaternary Research (INQUA) counterproposed that

7031-674: The person who named the species, while the antonym sensu lato ("in the broad sense") denotes a wider usage, for instance including other subspecies. Other abbreviations such as "auct." ("author"), and qualifiers such as "non" ("not") may be used to further clarify the sense in which the specified authors delineated or described the species. Species are subject to change, whether by evolving into new species, exchanging genes with other species, merging with other species or by becoming extinct. The evolutionary process by which biological populations of sexually-reproducing organisms evolve to become distinct or reproductively isolated as species

7120-501: The reptilian genera Langstonia and Barinasuchus , terrestrial predators that were the last surviving members of Sebecosuchia , a group related to crocodiles. The oceans were dominated by large carnivores like megalodons and livyatans , and 19 million years ago about 70% of all pelagic shark species disappeared. Mammals and birds continued to be the dominant terrestrial vertebrates, and took many forms as they adapted to various habitats. An explosive radiation of ursids took place at

7209-487: The result of misclassification leading to questions on whether there really are any ring species. The commonly used names for kinds of organisms are often ambiguous: "cat" could mean the domestic cat, Felis catus , or the cat family, Felidae . Another problem with common names is that they often vary from place to place, so that puma, cougar, catamount, panther, painter and mountain lion all mean Puma concolor in various parts of America, while "panther" may also mean

7298-573: The ring. Ring species thus present a difficulty for any species concept that relies on reproductive isolation. However, ring species are at best rare. Proposed examples include the herring gull – lesser black-backed gull complex around the North pole, the Ensatina eschscholtzii group of 19 populations of salamanders in America, and the greenish warbler in Asia, but many so-called ring species have turned out to be

7387-610: The rocks deposited during the Neogene Period . The continents in the Neogene were very close to their current positions. The Isthmus of Panama formed, connecting North and South America . The Indian subcontinent continued to collide with Asia , forming the Himalayas . Sea levels fell, creating land bridges between Africa and Eurasia and between Eurasia and North America. The global climate became more seasonal and continued an overall drying and cooling trend which began during

7476-508: The same species. This concept was narrowed in 2006 to a similarity of 98.7%. The average nucleotide identity (ANI) method quantifies genetic distance between entire genomes , using regions of about 10,000 base pairs . With enough data from genomes of one genus, algorithms can be used to categorize species, as for Pseudomonas avellanae in 2013, and for all sequenced bacteria and archaea since 2020. Observed ANI values among sequences appear to have an "ANI gap" at 85–95%, suggesting that

7565-529: The same species. When two species names are discovered to apply to the same species, the older species name is given priority and usually retained, and the newer name considered as a junior synonym, a process called synonymy . Dividing a taxon into multiple, often new, taxa is called splitting . Taxonomists are often referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to recognising differences or commonalities between organisms. The circumscription of taxa, considered

7654-502: The southern Pacific Rim suggests the dissemination of the genus dates to the time when Antarctica, Australia, and South America were connected in a common land-mass or supercontinent referred to as Gondwana . However, genetic evidence using molecular dating methods has been used to argue that the species in New Zealand and New Caledonia evolved from species that arrived in these landmasses by dispersal across oceans. Uncertainty exists in molecular dates and controversy rages as to whether

7743-502: The species concept and making taxonomy unstable. Yet others defend this approach, considering "taxonomic inflation" pejorative and labelling the opposing view as "taxonomic conservatism"; claiming it is politically expedient to split species and recognise smaller populations at the species level, because this means they can more easily be included as endangered in the IUCN red list and can attract conservation legislation and funding. Unlike

7832-453: The tropics of New Guinea, New Britain, and New Caledonia. In New Guinea and New Britain Nothofagus is characteristic of lower montane rain forests between 1000 and 2500 meters elevation, occurring infrequently at elevations as low as 600 meters, and in upper montane forests between 2500 and 3150 meters elevation. Nothofagus is most commonly found above the Castanopsis - Lithocarpus zone in

7921-537: The wild. It is difficult to define a species in a way that applies to all organisms. The debate about species concepts is called the species problem. The problem was recognised even in 1859, when Darwin wrote in On the Origin of Species : I was much struck how entirely vague and arbitrary is the distinction between species and varieties. He went on to write: No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of

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