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Horse Evolution

Horse evolution

. It should be noted that the animals shown are only selected examples - the evolution of horses was by no means a linear process. See the article for details.]] The evolution in the structure of their teeth, odd-toed limbs, obvious mobility of the upper lip, and other aspects by which they join the evolutionary line of odd-toed, hoofed mammals, the Perissodactyls. The tapirs and rhinoceroses remained adapted to their original style of life, conserving forms suitable for life in tropical forests, but the evolutionary line of the horse adapted to life on dryer land in the much-harsher climatic conditions of the steppes. The first ancestors of the modern horse walked on several spread-out toes, an accommodation to life spent walking on the soft, moist grounds of primeval forests. As the mainland was drying out, the steppes began to appear, and with it came large numbers of dry land predators. This in turn required the horse’s predecessors to possess more speed in order to survive. The ability to run faster was accomplished by the lengthening of limbs and the lifting of some toes from the ground in such a way that the weight of the body was gradually placed on one of the longest toes, the third. On solid ground, pushing-off with a single toe and equipped at the last evolutionary link with a hoof, the horse was able to reach fast speeds. The modern horse is a single-toed, hoofed mammal, while his predecessors were multi-odd-toed animals. The evolutionary development of the odd-toed predecessors to the present single-toed Family Equidae can be observed in the Tertiary period of North America, where layers of many well-preserved fossil skeletons of horse predecessors have been discovered. Paleontologists have been able to piece together a more complete picture of the modern horse’s evolutionary line than of any other mammal species. The evolutionary line of horses began in the lower Eocene epoch in a form called Hyracotherium. This “horse” was approximately the size of a fox and had various characteristics reminding us of its older predecessors: a relatively short head, 44 teeth with uneven, dull and bumpy molars, a short neck, a “springy”, arched back, and “wrist” and hock joints that are still low to the ground. The limbs are relatively long, obviously showing the beginnings of adaptation to gaining more speed. The forelimbs had developed five toes out of which only four were equipped with a small hoof; the fifth large “toe–thumb” was off the ground. The hind limbs had three out of the five toes equipped with small hooves, while the first and fifth toes did not touch the ground. During the Eocene, the Hyracotheriums branched out into various types resembling the fox, mainly in size (from 250 mm to 450 mm in height). Thousands of complete, fossilized skeletons of these animals have been found in the Eocene layers of North America, mainly in the Wind River basin of Wyoming. Similar limbs of four-toed horse ancestors have also been discovered in the Eocene layers of Europe. This mammal was called Hyracotherium. Orohippus, the successor of Hyracotherium, appeared in the middle of the Eocene. It has also been called Protorohippus. It still very much resembled the Hyracotherium; same size, slimmer body, elongated head, slimmer forelimbs and longer hind legs, all of which are characteristics of a good jumper. The outer toes of Hyracotherium are no longer present in the Orohippus; hence on each forelimb there were four fingers (toes) and on each hind leg three toes, and the first of his premolar teeth were dwarfed. In the early stages of the Oligocene epoch, the North American environment was changing. During the still-warm and dry weather conditions, the forests were yielding to flatlands, which were home to grasses and various kinds of brush. In some places these plains were covered in sand; hence the type of environment resembling our present prairies. The younger Pliocene epoch revealed the Mesohippus, which was one of the more widespread types of mammals in North America at the time. Mesohippus walked on three toes (fingers) on both front and hind feet. The third toe was stronger than the outer ones and thus more weighted. Although the first and fifth toe remained, they were very small. Judging by its slim limbs, Mesohippus, which was about 500 mm tall, was a fast animal. At the end of Oligocene epoch and the beginning of the Miocene epoch, the Mesohippus evolved into a form known as Miohippus. Miohippus had some parts bigger than its predecessor. It had branched out into two branches from which one adjusted to the life in the primeval forests, while the other remained suited to life on the prairies. The forest form led to the birth of Kalobatippus (Miohippus intermedius), whose second and fourth finger again elongated for travel on the softer primeval forest grounds. The Kalobatippus managed to relocate to Asia via the Bering Strait land bridge, and from there moved into Europe, where its fossils were formerly described under the name Anchitherium. Kalobatippus is then believed to have evolved into a form known as Hyohippus, which became extinct near the beginning of Pliocene. From the Miohippus that remained on the steppes evolved the North American breed of Parahippus. This little “horse” was the size of a small pony, with a prolonged skull and the facial structure resembling horses of today. The third toe became stronger and larger, hence carrying the main weight of the body. His four premolars resembled the molar teeth and the first almost disappeared. The incisive teeth of the Parahippus, like its predecessors, had a crown as humans do; however, the top incisors had a trace of shallow crease marking the beginning of the core/cup. In the middle of the Miocene epoch, Merychippus appeared, following after the Parahippus. Meryhippus’ “crunching” on hard steppe, grassy plants caused it to grow relatively longer. It also had wider molars than its predecessors. The hind legs, which were relatively short, still had the side toes equipped with small hooves, though they probably only touched the ground when running. From the numerous varieties of Meryhippus evolved three new kinds of equids: Hipparion, Protohippus and Pliohippus. According to shapes, the most diverted of the three was the Hipparion. The main difference here was in the structure of tooth enamel. In comparison with other horses, the inside, or tongue side had a completely isolated parapet. On his slim legs, the Hipparion had three toes equipped with small hooves, but the side toes did not touch the ground. The American Hipparion, also known as Neohipparion, proliferated in several kinds of equids several of which managed to migrate to Asia and Europe during the Pliocene Epoch. (The European Hipperia differs from the American Hipparion in the smaller body size – the most known discovery of these fossils was near Athens.) The complete and well-preserved skeleton of the North American Hipparion shows an animal the size of a small pony. They were very slim, similar to antelopes, and equally timid. They easily adjusted to life on dry prairies. According to newer research, they’re being excluded from the direct ancestry of horses, as it’s believed Hipparion more likely led to the evolution of zebras and asses. Outside the diversities of head features, today’s horses, asses, and zebras differ only slightly in their teeth and bones. The branch of Protohippus that is believed to be the line leading to the modern horse died off in the Pliocene. The form of Meryhippus that lead to present horses was the Pliocene Pliohippus. It still had long extra toes on both sides of the hoof, but they were externally barely visible, callused stubs. The long and slim limbs of Pliohippus reveal a quick-footed steppe animal. The Pliohippus became the Plesippus, and further evolution into the form of genuine Equus occurred during the upper Pliocene. It still remains uncertain how these horses came from their “home land” of North America to Europe. At the end of the Pliocene, the climate in North America began to cool down significantly. The animals were forced to move south. One part of the Plesippus species escaped to South America, and the other moved across the land bridge around the Bering Strait into Asia and Europe. A portion also remained in the southern section of North America. The Ice Age spread five times over Europe and North America and five times again receded (the interglacial periods). This of course lasted many millennia and it is estimated that approximately one million years elapsed from the Ice Age (the Quaternary period) to our era. The oldest species of true horse, Equus stenonis, was discovered in Italy, and is believed to have evolved from the Plesippus at the end of the Tertiary and beginning of the Quaternary periods. Equus stenonis suddenly proliferated into two branches, one lighter in body mass and one heavier. The North American genuine form of the diluvium horse was named Equus scotti and did not differ in any way from the European form; however, some types exceeded the modern horse (Equus scotti var. giganteus) in size. In South America the Plesippus was evolving into a form named Hippidium. The Hippidium was relatively short-legged with an especially long nose that also formed the lower part of the skull. It continued to live on the South American pampas for a long time, but eventually died off. All the horses in North America became extinct as well, perhaps due to some mass contagion, but it is also theorized that humans hunted them to extinction, as the appearance of humans in the Americas occurred at about the same time as the extinction of most large mammals in the Americas.

Horses in Historical Times

At the end of the 15th century, when the first Europeans came to America, there were no horses; the cultural tribes of Indians (in today’s Mexico and Peru) did not even have a name for the animal. The Spanish imported predecessors of all the horses back to America. Runaway horses and cattle went wild on the pampas and proliferated into large herds, only to be caught again later and domesticated.

Emergence of the Genuine Equus Species

The species of genuine horse came to walk only on the end of the third toe and from both side toes, as did their predecessors. Skeletal remnants of show obvious wear on the back of both sides of metacarpal and metatarsal bones, commonly called the “splint bones”. They are the remnants of the second and the fourth toe. It is often believed that the splint bones on the modern horse are a useless attachment, but they indeed play an important role in supporting the carpal joints (front knee) and even the tarsal joints (hock). It is not unheard of that foals are occasionally born with three toes equipped with hooves. This is called a phylogenetic atavism caused by arrested development in a certain embryonic stage. Throughout the phylogenetic development, the teeth of the horse underwent significant changes. The type of the original omnivorous teeth with short, "bumpy" molars, with which the prime members of the evolutionary line distinguished themselves, gradually changed into the teeth common to herbivorous mammals. They became long (as much as 100 mm), roughly cubical molars equipped with a flat grinding surface. In conjunction with the teeth, during the horse’s evolution the elongation of the facial part of the skull is apparent, and can also be observed in the backward set eyeholes. In addition, the relatively short neck of the equine ancestors became longer with equal elongation of the legs. This is because they adapted to finding food by grazing on the steppes. Finally, the size of the body grew as well, not only due to plentiful food, but also to the increase in variety. Recent studies by a team of geneticists headed by C. Vila indicate that the horse line split from the zebra/donkey line in the window 4 mya to 2 mya. Equus ferus, ancestor species to Equus caballus, appeared 630,000 to 320,000 years bp. Equus caballus was formed from several subspecies of Equus ferus by selective breeding widely over Eurasia for an extended time. The details of this process are currently a target of research by archaeologists and geneticists. Category:Horses Category:Prehistoric horses Category:Cenozoic

External Links

- [http://www.sciencemag.org/cgi/content/abstract/291/5503/474?ijkey=ee6b658a7e39e6054958efcf7f95ef0c57496273&keytype2=tf_ipsecsha Widespread Origins of Domestic Horse Lineages]

Evolution

, based on rRNA gene data, showing the separation of the three domains, bacteria, archaea, and eukaryotes, as described initially by Carl Woese.]] In biology, evolution is the process by which populations of organisms acquire and pass on novel traits from generation to generation, affecting the overall makeup of the population and even leading to the emergence of new species. The terms organic evolution or biological evolution are often used to distinguish this meaning from other usages. The development of the modern theory of evolution began with the introduction of the concept of natural selection in a joint 1858 paper by Charles Darwin and Alfred Russel Wallace. This theory achieved a wider readership in Darwin's 1859 book, The Origin of Species. Darwin and Wallace proposed that evolution occurs because a heritable trait that increases an individual's chance of successfully reproducing will become more common, by inheritance, from one generation to the next, and likewise a heritable trait that decreases an individual's chance of reproducing will become rarer. This work was groundbreaking, and overturned other evolutionary theories, such as that advanced by Jean Baptiste Lamarck. Because of its potential implications for the origins of humankind, the theory has been at the center of many social and religious controversies since its first inception (see Creation-evolution controversy). In the 1930s, scientists combined Darwinian natural selection with the re-discovered theory of Mendelian heredity to create the modern synthesis, now one of the fundamental scientific theories of biology. In the modern synthesis, "evolution" is defined as a change in the frequency of alleles within a population from one generation to the next. The basic mechanisms that produce these changes are natural selection, genetic drift, and genetic variation. The primary sources of genetic variation are mutation, sex, and gene flow.

Overview of evolution

Evidence of evolution

The process of evolution has left behind numerous records which reveal the history of species. While the best-known of these are the fossils, fossils are only a small part of the overall physical record of evolution. Fossils, taken together with the comparative anatomy of present-day plants and animals, constitute the morphological record. By comparing the anatomies of both modern and extinct species, biologists can reconstruct the lineages of those species with some accuracy. Using fossil evidence, for instance, the connection between dinosaurs and birds has been established by way of so-called "transitional" species such as Archaeopteryx. The development of genetics has allowed biologists to study the genetic record of evolution as well. Although we cannot obtain the DNA sequences of most extinct species, the degree of similarity and difference among modern species allows geneticists to reconstruct lineages with greater accuracy. It is from genetic comparisons that claims such as the 98-99% similarity between humans and chimpanzees come from, for instance. Other evidence used to demonstrate evolutionary lineages includes the geographical distribution of species. For instance, monotremes and most marsupials are found only in Australia, showing that their common ancestor with placental mammals lived before the submerging of the ancient land bridge between Australia and Asia. Scientists correlate all of the above evidence – drawn from paleontology, anatomy, genetics, and geography – with other information about the history of the earth. For instance, paleoclimatology attests to periodic ice ages during which the climate was much cooler; and these are found to match up with the spread of species such as the woolly mammoth which are better-equipped to deal with cold.

Morphological evidence

Fossils are important for estimating when various lineages developed. As fossilization on an organism is an uncommon occurrence, usually requiring hard parts (like bone) and death near a site where sediments are being deposited, the fossil record only provides sparse and intermittent information about the evolution of life. Fossil evidence of organisms without hard body parts, such as shell, bone, and teeth, is sparse but exists in the form of ancient microfossils and the fossilization of ancient burrows and a few soft-bodied organisms. Fossil evidence of prehistoric organisms has been found all over the Earth. The age of fossils can often be deduced from the geologic context in which they are found; and their absolute age can be verified with radiometric dating. Some fossils bear a resemblance to organisms alive today, while others are radically different. Fossils have been used to determine at what time a lineage developed, and transitional fossils can be used to demonstrate continuity between two different lineages. Paleontologists investigate evolution largely through analysis of fossils. Phylogeny, the study of the ancestry of species, has revealed that structures with similar internal organization may perform divergent functions. Vertebrate limbs are a common example of such homologous structures. Bat wings, for example, are very similar to hands. A vestigial organ or structure may exist with little or no purpose in one organism, though they have a clear purpose in other species. The human wisdom teeth and appendix are common examples.

Genetic sequence evidence

Comparison of the genetic sequence of organisms reveals that phylogenetically close organisms have a higher degree of sequence similarity than organisms that are phylogenetically distant. For example, neutral human DNA sequences are approximately 1.2% divergent (based on substitutions) from those of their nearest genetic relative, the chimpanzee, 1.6% from gorillas, and 6.6% from baboons. Sequence comparison is considered a measure robust enough to be used to correct erroneous assumptions in the phylogenetic tree in instances where other evidence is scarce. Further evidence for common descent comes from genetic detritus such as pseudogenes, regions of DNA which are orthologous to a gene in a related organism, but are no longer active and appear to be undergoing a steady process of degeneration. Since metabolic processes do not leave fossils, research into the evolution of the basic cellular processes is done largely by comparison of existing organisms. Many lineages diverged when new metabolic processes appeared, and it is theoretically possible to determine when certain metabolic processes appeared by comparing the traits of the descendants of a common ancestor.

History of evolutionary thought

metabolic.]] The idea of biological evolution has existed since ancient times, notably among Hellenists such as Epicurus and Anaximander, but the modern theory was not established until the 18th and 19th centuries, by scientists such as Jean-Baptiste Lamarck and Charles Darwin. While transmutation of species was accepted by a sizeable number of scientists before 1859, it was the publication of Charles Darwin's The Origin of Species by Means of Natural Selection which provided the first cogent mechanism by which evolutionary change could occur: his theory of natural selection. Darwin was motivated to publish his work on evolution after receiving a letter from Alfred Russel Wallace, in which Wallace revealed his own discovery of natural selection. As such, Wallace is sometimes given shared credit for the theory of evolution. Darwin's theory, though it succeeded in profoundly shaking scientific opinion regarding the development of life, could not explain the source of variation in traits within a species, and Darwin's proposal of a hereditary mechanism (pangenesis) was not compelling to most biologists. It was not until the late 19th and early 20th centuries that these mechanisms were established. pangenesis, proposed the theory of punctuated equilibrium in 1972.]] When Gregor Mendel's work regarding the nature of inheritance in the late 19th century was "rediscovered" in 1900, it led to a storm of conflict between Mendelians (Charles Benedict Davenport) and biometricians (Walter Frank Raphael Weldon and Karl Pearson), who insisted that the great majority of traits important to evolution must show continuous variation that was not explainable by Mendelian analysis. Eventually, the two models were reconciled and merged, primarily through the work of the biologist and statistician R.A. Fisher. This combined approach, applying a rigorous statistical model to Mendel's theories of inheritance via genes, became known in the 1930s and 1940s as the modern evolutionary synthesis. In the 1940s, following up on Griffith's experiment, Avery, McCleod and McCarty definitively identified deoxyribonucleic acid (DNA) as the "transforming principle" responsible for transmitting genetic information. In 1953, Francis Crick and James Watson published their famous paper on the structure of DNA, based on the research of Rosalind Franklin and Maurice Wilkins. These developments ignited the era of molecular biology and transformed the understanding of evolution into a molecular process: the mutation of segments of DNA (see molecular evolution). George C. Williams' 1966 Adaptation and natural selection: A Critique of some Current Evolutionary Thought marked a departure from the idea of group selection towards the modern notion of the gene as the unit of selection. In the mid-1970s, Motoo Kimura formulated the neutral theory of molecular evolution, firmly establishing the importance of genetic drift as a major mechanism of evolution. Debates have continued within the field. One of the most prominent public debates was over the theory of punctuated equilibrium, proposed in 1972 by paleontologists Niles Eldredge and Stephen Jay Gould to explain the paucity of transitional forms between phyla in the fossil record.

Social and religious controversies

Stephen Jay Gould from 1871 reflects part of the social controversy over whether humans and apes share a common lineage.]] There has been constant controversy surrounding the ideas presented by The Origin of Species since it was first printed in 1859. Since the early twentieth century, however, the idea that biological evolution of some form occurred and is responsible for speciation has been almost completely uncontested within the scientific community. Most controversy over the theory has come because of its philosophical, cosmological, and religious implications, and supporters as well as detractors have interpreted it as generally indicating that human beings are, like all animals, evolved, and that this account of the origins of humankind is squarely at odds with many religious interpretations. The idea that humans are "merely" animals, and are genetically very closely related to primates, have been independently argued as repellent notions by generations of detractors. Others also intepreted the truth of the theory to imply varying types of social changes — one prominent example is the idea of eugenics, formulated by Darwin's cousin Francis Galton, which argues for the improvement of human heredity by means of political policies. Others have found different political interpretations which have been used as arguments both for and against the theory. The questions raised about the relation of evolution to the origins of humans has made it an especially tenacious issue with religious traditions. It has prominently been seen as opposing a "literal" interpretation of the account of the origins of humankind as described in Genesis, the first book of the Bible. In many countries — notably in the United States — this has led to what has been called the Creation-evolution controversy, which has focused primarily on struggles over teaching curriculum.

Science of evolution

Science: fact and theory

The word "evolution" has been used to refer both to a fact and a theory, and it is important to understand both these different meanings of evolution, and the relationship between fact and theory in science.

Evolution as fact and theory

When "evolution" is used to describe a fact, it refers to the observations that populations of one species of organism do, over time change into new, or several new, species. In this sense, evolution occurs whenever a new strain of bacterium evolves that is resistant to antibodies that had been lethal to prior strains. Another clear case of evolution as fact involves the hawthorn fly, Rhagoletis pomonella. Different populations of hawthorn fly feed on different fruits. A new population spontaneously emerged in North America in the 19th century some time after apples, a non-native species, were introduced. The apple feeding population normally feeds only on apples and not on the historically preferred fruit of hawthorns. Likewise the current hawthorn feeding population does not normally feed on apples. A current area of scientific research is the investigation of whether or not the apple feeding race may further evolve into a new species. Some evidence, such as the fact that six out of thirteen alozyme loci are different, that hawthorn flies mature later in the season, and take longer to mature, than apple flies, and that there is little evidence of interbreeding (researchers have documented a 4-6%hybridization rate) suggests that this is indeed ocurring. (see Berlocher and Bush 1982, Berlocher and Feder 2002, Bush 1969, McPheron et. al. 1988, Prokopy et. al. 1988, Smith 1988) When "evolution" is used to describe a theory, it refers to an explanation for why and how evolution (for example, in the sense of "speciation") occurs. An example of evolution as theory is the modern synthesis of Darwin and Wallace's theory of natural selection and Mendel's principles of genetics. This theory has three major aspects: # Common descent of all organisms from a single ancestor or ancestral gene pool. # Manifestation of novel traits in a lineage. # Mechanisms that cause some traits to persist while others perish. When people provide evidence for evolution, in some cases they are providing evidence that evolution occurs; in other cases they are providing evidence that a given theory is the best explanation yet as to why and how evolution occurs.

The meaning of, and relationship between, fact and theory in science

:Main article: Theory The modern synthesis, like its Mendelian and Darwinian antecedents, is a scientific theory. In plain English, people use the word "theory" to signify "conjecture", "speculation", or "opinion". In this popular sense, "theories" are opposed to "facts" — parts of the world, or claims about the world, that are real or true regardless of what people think. In scientific terminology however, a theory is a model of the world (or some portion of it) from which falsifiable hypotheses can be generated and tested through controlled experiments, or be verified through empirical observation. In this scientific sense, "facts" are parts of theories – they are things, or relationships between things, that theories must take for granted in order to make predictions, or that theories predict. In other words, for scientists "theory" and "fact" do not stand in opposition, but rather exist in a reciprocal relationship – for example, it is a "fact" that every apple ever dropped on earth (under normal, controlled conditions) has been observed to fall towards the center of the planet in a straight line, and the "theory" which explains these observations is the current theory of gravitation. In this same sense evolution is a fact and modern synthesis is currently the most powerful theory explaining evolution, variation and speciation. Within the science of biology, modern synthesis has completely replaced earlier accepted explanations for the origin of species, including Lamarckism and creationism.
Who studies evolution?
Scholars in a number of academic disciplines and subdisciplines document the fact of evolution, and contribute to the theory of evolution.
Physical anthropology
Physical anthropology emerged in the late 1800s as the study of human osteology, and the fossilized skeletal remains of other hominids. At that time anthropologists debated whether their evidence supported Darwin's claims, because skeletal remains revelaed temporal and spacial variation among hominids, but Darwin had not offered an explanation of the mechanisms that produce variation. With the recognition of Mendelian genetics and the rise of the modern synthesis, however, evolution became both the fundamental conceptual framework for, and object of study of, physical anthropologists. In addition to studying skeletal remains, they began to study genetic variation among human populations (i.e. population genetics; thus, some physical anthropologists began calling themselves biological anthropologists.
Evolutionary biology
Evolutionary biology is a subfield of biology concerned with the origin and descent of species, as well as their change over time. At first it was an interdisciplinarity field including scientists from many traditional taxonomically oriented disciplines. For example, it generally includes scientists who may have a specialist training in particular organisms such as mammalogy, ornithology, or herpetology but use those organisms as systems to answer general questions in evolution. Evolutionary biology as an academic discipline in its own right emerged as a result of the modern evolutionary synthesis in the 1930s and 1940s. It was not until the 1970s and 1980s, however, that a significant number of universities had departments that specifically included the term evolutionary biology in their titles.
Evolutionary developmental biology
Evolutionary developmental biology is an emergent subfield of evolutionary biology that looks at genes of related and unrelated organisms. By comparing the explicit nucleotide sequences of DNA/RNA, it is possible to experimentally determine and trace timelines of species development. For example, gene sequences support the conclusion that chimpanzees are the closest primate ancestor to humans, and that arthropods (e.g., insects) and vertebrates (e.g., humans) have a common biological ancestor.
Ancestry of organisms
vertebrates In biology, the theory of universal common descent proposes that all organisms on Earth are descended from a common ancestor or ancestral gene pool (which is called having "common descent"). Evidence for common descent may be found in traits shared between all living organisms. In Darwin's day, the evidence of shared traits was based solely on visible observation of morphologic similarities, such as the fact that all birds — even those which do not fly — have wings. Today, the theory of evolution has been strongly confirmed by genetics. For example, every living cell makes use of nucleic acids as its genetic material, and uses the same twenty amino acids as the building blocks for proteins. All organisms use the same genetic code (with some extremely rare and minor deviations) to translate nucleic acid sequences into proteins. The universality of these traits strongly suggests common ancestry, because the selection of these traits seems somewhat arbitrary, . The evolutionary process can be exceedingly slow. Fossil evidence indicates that the diversity and complexity of modern life has developed over much of the age of the earth. Geological evidence indicates that the Earth is approximately 4.6 billion years old. (See Timeline of evolution.) Studies on guppies by David Reznick at the University of California, Riverside, however, have shown that the rate of evolution through natural selection can proceed 10 thousand to 10 million times faster than what is indicated in the fossil record. Information about the early development of life includes input from the fields of geology and planetary science. These sciences provide information about the history of the Earth and the changes produced by life. A great deal of information about the early Earth has been destroyed by geological processes over the course of time.
History of life
planetary sciences in the Siyeh Formation, Glacier National Park. In 2002, William Schopf of UCLA published a controversial paper in the journal Nature arguing that formations such as this possess 3.5 billion year old fossilized algae microbes. If true, they would be the earliest known life on earth.]] The chemical evolution from self-catalytic chemicals to life (see Origin of life) is not a part of biological evolution. Not much is known about the earliest developments in life. However, all existing organisms share certain traits, including cellular structure, and genetic code. Most scientists interpret this to mean all existing organisms share a common ancestor, which had already developed the most fundamental cellular processes, but there is no scientific consensus on the relationship of the three domains of life (Archea, Bacteria, Eukaryota) or the origin of life. Attempts to shed light on the earliest history of life generally focus on the behavior of macromolecules, particularly RNA, and the behavior of complex systems. The emergence of oxygenic photosynthesis (around 3 billion years ago) and the subsequent emergence of an oxygen-rich, non-reducing atmosphere can be traced through the formation of banded iron deposits, and later red beds of iron oxides. This was a necessary prerequisite for the development of aerobic cellular respiration, believed to have emerged around 2 billion years ago. In the last billion years, simple multicellular plants and animals began to appear in the oceans. Soon after the emergence of the first animals, the Cambrian explosion (a period of unrivaled and remarkable, but brief, organismal diversity documented in the fossils found at the Burgess Shale) saw the creation of all the major body plans, or phyla, of modern animals. This event is now believed to have been triggered by the development of the Hox genes. About 500 million years ago, plants and fungi colonized the land, and were soon followed by arthropods and other animals, leading to the development of land ecosystems with which we are familiar.
The Modern Synthesis
The current understanding of the mechanistics of evolution differs considerably from the theory first outlined by Charles Darwin. Importantly, advances in genetics pioneered by Gregor Mendel led to a sophisticated understanding of the basis of variation and the mechanisms of inheritance. In addition natural selection has come to be seen as only one of a number of forces acting in evolution. A notable milestone in this regard was the formulation of the neutral theory of molecular evolution by Motoo Kimura.
Heredity
Gregor Mendel first proposed a gene-based theory of inheritance, discretizing the elements responsible for heritable traints into the fundamental units we now call genes, and laying out a mathematical framework for the segregation and inheritance of variants of a gene, which we now refer to as alleles. Later research identified the molecule DNA as the genetic material, through which traits are passed from parent to offspring, and identified genes as discrete elements within DNA. Though largely faithfully maintained within organisms, DNA is both variable across individuals and subject to a process of change or mutation. Non-DNA based forms of heritable variation exist, which may change the way in which genes are expressed or maintained. The processes that produce these variations leave the genetic information intact and are often reversible. This is called epigenetic inheritance and may include phenomena such as DNA methylation, prions, and structural inheritance. Investigations continue into whether these mechanisms allow for the production of specific beneficial heritable variation in response to environmental signals. If this were shown to be the case, then some instances of evolution would lie outside of the typical Darwinian framework, which avoids any connection between environmental signals and the production of heritable variation.
Sexual reproduction
In addition to passing genetic material from parent to offspring, nearly all organisms employ sex to exchange genetic material. This, combined with meiotic recombination, allows genetic variation to be propagated through an interbreeding population. These mechanisms allow individual variations to be propagated more or less independently, so that the population as a whole can retain beneficial variation and eliminate harmful variation (rather than both of these effects competing within a single asexual organism). However, these mechanisms are not perfect, and so some variation is co-propagated as a result of linkage, producing some odd effects (see Muller's ratchet).
Mechanisms of evolution
Evolution consists of two basic types of processes: those that introduce new genetic variation into a population, and those that affect the frequencies of existing variation. There are three known processes that affect the survival of a characteristic (or, more specifically, the frequency of an allele):
- Natural selection
- Changes in population structure
- Genetic drift These basic mechanisms of evolution have all been observed in the present and in evidence of their existence in the past. Their study is being used to guide the development of new medicines and other health aids such as the current effort to prevent a H5N1 (i.e. bird flu) pandemic
Variation
Without genetic variation, populations cannot evolve. The two principle sources of genetic variation are mutations and gene flow. Other forms of genetic variation due to gene transfer include horizontal gene transfer, antigenic shift, and reassortment. Viruses can transfer genes between species [http://66.102.7.104/search?q=cache:tpICVNWaTbgJ:non.fiction.org/lj/community/ref_courses/3484/enmicro.pdf+sex+evolution+%22Horizontal+gene+transfer%22+-human+Conjugation+RNA+DNA&hl=en]. Bacteria can incorporate genes from other dead bacteria, exchange genes with living bacteria, and can have plasmids "set up residence seperate from the host's genome" [http://www2.nau.edu/~bah/BIO471/Reader/Pennisi_2003.pdf]. "Genes that move between species play by rules that microbial experts are just beginning to discern" [http://66.102.7.104/search?q=cache:gto6eXfbGIEJ:www.niagara.edu/eli/Science%252016%2520July%25202004.GeneSwap.doc+sex+evolution+%22Horizontal+gene+transfer%22+-human+Conjugation+RNA+DNA&hl=en].
Mutation
The ultimate source of all genetic variation is mutations. They are permanent, transmissible changes to the genetic material (usually DNA or RNA) of a cell, and can be caused by "copying errors" in the genetic material during cell division and by exposure to radiation, chemicals, or viruses. In multicellular organisms, mutations can be subdivided into germline mutations that occur in the gametes and thus can be passed on to progeny, and somatic mutations that often lead to the malfunction or death of a cell and can cause cancer. Mutations that are not affected by natural selection are called neutral mutations. Their frequency in the population is governed entirely by genetic drift and gene flow. It is understood that a species' genome, in the absence of selection, undergoes a steady accumulation of neutral mutations. The probable mutation effect is the proposition that a gene that is not under selection will be destroyed by accumulated mutations. This is an aspect of genome degradation. Not all mutations are created equal; simple point mutations (substitutions), which comprise the vast majority of genetic variation, usually can only alter the function or level of expression of existing genes. Gene duplications, which may occur via a number of mechanisms, are believed to be the major mechanism for the introduction of new genes; most genes belong to larger "families" of genes derived from a common ancestral gene (two genes from a species that are in the same family are dubbed "paralogs"). Finally, large chromosomal rearrangements (like the fusion of two chromosomes in the chimp/human common ancestor that produced human chromosome 2) almost invariably result in a speciation event.
Gene flow
Gene flow (or gene admixture) is introduction of variation into a population from an outside population. It is the only mechanism whereby two populations can become closer genetically while increasing their variation. Migration of one population into an area occupied by a second population can result in gene flow. Gene flow operates when geography and culture are not obstacles. When gene flow is impeded by non-geographic obstacles, the situation is termed reproductive isolation and is considered to be the hallmark of speciation.
Drift
Genetic drift describes changes in allele frequency from one generation to the next due to sampling variance. The frequency of an allele in the offspring generation will vary according to a probability distribution of the frequency of the allele in the parent generation. Thus, over time, allele frequencies will tend to "drift" upward or downward, eventually becoming "fixed" - that is, going to 0% or 100% frequency. Fluctuations in allele frequency between successive generations may result in some alleles disappearing from the population. Two separate populations that begin with the same allele frequencies therefore might drift by random fluctuation into two divergent populations with different allele sets (for example, alleles that are present in one have been lost in the other). Many aspects of genetic drift depend on the size of the population (generally abbreviated as N). This is especially important in small mating populations, where chance fluctuations from generation to generation can be large. The relative importance of natural selection and genetic drift in determining the fate of new mutations also depends on the population size and the strength of selection: when N times s (population size times strength of selection) is small, genetic drift predominates. When N times s is large, selection predominates. Thus, natural selection is 'more efficient' in large populations, or equivalently, genetic drift is stronger in small populations. Finally, the time for an allele to become fixed in the population by genetic drift (that is, for all individuals in the population to carry that allele) depends on population size, with smaller populations requiring a shorter time to fixation.
Population structure
An important facet of evolution occurs through changes in population structure. The movement of populations and changes in their size can have profound impacts on evolution over and above those governed by selection and drift. Migration can result in admixture leading to the introduction of new genetic variation, or it may result in geographic isolation which may in turn lead to reproductive isolation or speciation. Populations may also shrink or grow over time, producing "bottlenecks" or "explosions" respectively. Since population size has a profound effect on the relative strengths of genetic drift and natural selection, changes in population size can alter the dynamics of these processes considerably. Such changes may also produce dramatic and dangerous crashes in the level of genetic variation in the population, or allow rapid increases in standing genetic variation. The free movement of alleles through a population may also be impeded by population structure. For example, most real-world populations are not actually fully interbreeding; geographic proximity has a strong influence on the movement of alleles within the population. Many models of evolution rely on simplifying assumptions of constant population size and fully interbreeding populations for mathematical convenience. An example of the effect of population structure is the so-called founder effect, resulting from a migration and population bottleneck. In this case, a single, rare allele may suddenly increase very rapidly in frequency if it happened to be prevalent in a small number of "founder" individuals. The frequency of the allele in the resulting population can be much higher than otherwise expected, especially for deleterious, disease-causing alleles.
Selection and adaptation

Natural selection
Natural selection comes from differences in survival and reproduction as a result of the environment. Differential mortality is the survival rate of individuals to their reproductive age. Differential fertility is the total genetic contribution to the next generation. Note that, whereas mutations and genetic drift are random, natural selection is not, as it preferentially selects for different mutations based on differential fitnesses. For example, rolling dice is random, but always picking the higher number on two rolled dice is not random. The central role of natural selection in evolutionary theory has given rise to a strong connection between that field and the study of ecology. Natural selection can be subdivided into two categories:
- Ecological selection occurs when organisms that survive and reproduce increase the frequency of their genes in the gene pool over those that do not survive.
- Sexual selection occurs when organisms which are more attractive to the opposite sex because of their features reproduce more and thus increase the frequency of those features in the gene pool. Natural selection also operates on mutations in several different ways:
- Purifying or background selection eliminates deleterious mutations from a population.
- Directional selection increases the frequency of a beneficial mutation.
- Balancing selection maintains variation within a population through a number of mechanisms, including:
- Heterozygote advantage or overdominance, where the heterozygote is more fit than either of the homozygous forms (exemplified by human sickle cell anemia conferring resistance to malaria)
- Frequency-dependent selection, where rare variants have a higher fitness, because of thier rarity.
- Stabilizing selection favors average characteristics in a population, thus reducing gene variation but retaining the mean.
- Disruptive selection favors both extremes, and results in a bimodal distribution of gene frequency. The mean may or may not shift.
Adaptation
Through the process of natural selection, species become better adapted to their environments. Adaptation is any evolutionary process that increases the fitness of the individual, or sometimes the trait that confers increased fitness, e.g. a stronger prehensile tail or greater visual acuity. Note that adaptation is context-sensitive; a trait that increases fitness in one environment may decrease it in another. Evolution does not act in a linear direction towards a pre-defined "goal" — it only responds to various types of adaptionary changes. The belief in a telelogical evolution of this sort is known as orthogenesis, and is not supported by the scientific theory of evolution. One example of this misconception is the erroneous belief humans will evolve more fingers in the future on account of their increased use of machines such as computers. In reality, this would only occur if more fingers offered a significantly higher rate of reproductive success than those not having them, which seems very unlikely at the current time. Most biologists believe that adaptation occurs through the accumulation of many mutations of small effect. However, macromutation is an alternative process for adaptation that involves a single, very large scale mutation.
Speciation and extinction
macromutation Speciation is the creation of two or more species from one. This may take place by various mechanisms. Allopatric speciation occurs in populations that become isolated geographically, such as by habitat fragmentation or migration. Sympatric speciation occurs when new species emerge in the same geographic area. Ernst Mayr's peripatric speciation is a type of speciation that exists in between the extremes of allopatry and sympatry. Peripatric speciation is a critical underpinning of the theory of punctuated equilibrium. Extinction is the disappearance of species (i.e. gene pools). The moment of extinction generally occurs at the death of the last individual of that species. Extinction is not an unusual event in geological time — species are created by speciation, and disappear through extinction. The Permian-Triassic extinction event was the Earth's most severe extinction event, rendering extinct 90% of all marine species and 70% of terrestrial vertebrate species. In the Cretaceous-Tertiary extinction event many forms of life perished (including approximately 50% of all genera), the most often mentioned among them being the extinction of the non-avian dinosaurs (See Image 5).
See also

Notes and references
# "Ancient microfossils from Western Australia are again the subject of heated scientific argument: are they the oldest sign of life on Earth, or just a flaw in the rock?" "[http://www.abc.net.au/science/news/space/SpaceRepublish_497964.htm]" # Understanding Evolution, from California's Berkeley University. "[http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_17] [http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_16] # Li WH, Saunders MA (2005) Initial sequence of the chimpanzee genome and comparison with the human genome. Nature 437: 69–87. Britten RJ (2002) Divergence between samples of chimpanzee and human DNA sequences is 5%, counting indels. Proc Natl Acad Sci U S A 99: 13633–13635. # Two sources: 'Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees'. and 'Quantitative Estimates of Sequence Divergence for Comparative Analyses of Mammalian Genomes' "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=11170892] [http://www.genome.org/cgi/content/full/13/5/813]" # Pseudogene evolution and natural selection for a compact genome. "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=10833048]" # Reference for emergence of new race of apple maggot flies http://www.nd.edu/~aforbes/ # Evaluation of the Rate of Evolution in Natural Populations of Guppies (Poecilia reticulata) "[http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=9072971&query_hl=2]" # The use of evolutionary principles to guide disease diagnosis and drug development with respect to bird flu (i.e. H5N1 virus) [http://www.cdc.gov/ncidod/EID/vol11no10/05-0644.htm] # Understanding Evolution, from California's Berkeley University: "Sex can introduce new gene combinations into a population. This genetic shuffling is another important source of genetic variation."[http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_17]
- Berlocher, S.H. and G.L. Bush. 1982. An electrophoretic analysis of Rhagoletis (Diptera: Tephritidae) phylogeny. Systematic Zoology 31:136-155.
- Berlocher, S.H. and J.L. Feder. 2002. Sympatric speciation in phytophagous insects: moving beyond controversy? Annual Review of Entomology 47:773-815.
- Bush, G.L. 1969. Sympatric host race formation and speciation in frugivorous flies of the genus Rhagoletis (Diptera: Tephritidae). Evolution 23:237-251.
- Darwin, Charles November 24 1859. On the Origin of Species by means of Natural Selection or the Preservation of Favoured Races in the Struggle for Life. London: John Murray, Albemarle Street. 502 pages. Reprinted: Gramercy (May 22, 1995). ISBN 0517123207
- Prokopy, R.J., S.R. Diehl and S.S. Cooley. 1988. Behavioral evidence for host races in Rhagoletis pomonella flies. Oecologia 76:138-147.
- Zimmer, Carl. Evolution: The Triumph of an Idea. Perennial (October 1, 2002). ISBN 0060958502
- Larson, Edward J. Evolution: The Remarkable History of a Scientific Theory (Modern Library Chronicles). Modern Library (May 4, 2004). ISBN 0679642889
- Mayr, Ernst. What Evolution Is. Basic Books (October, 2002). ISBN 0465044263
- McPheron, B. A., D. C. Smith and S. H. Berlocher. 1988. Genetic differentiation between host races of Rhagoletis pomonella. Nature. 336:64-66.
- Gigerenzer, Gerd, et al., The empire of chance: how probability changed science and everyday life (New York: Cambridge University Press, 1989).
- Smith, D. C. 1988. Heritable divergence of Rhagoletis pomonella host races by seasonal asynchrony. Nature. 336:66-67.
- Williams, G.C. (1966). Adaptation and Natural Selection: A Critique of some Current Evolutionary Thought . Princeton, N.J.: Princeton University Press.
- Sean B. Carroll, 2005, Endless Forms Most Beautiful: The New Science of Evo Devo and the Making of the Animal Kingdom, W. W. Norton & Company. ISBN 0393060160
- Bill Bryson, A Short History of Nearly Everything, Black Swan Books (2004), ISBN 0-552-99704-8
External links

- [http://www.talkorigins.org Talk.Origins Archive] — see also talk.origins
- [http://evolution.berkeley.edu/ Understanding Evolution] @ [http://berkeley.edu Berkeley]
- [http://nationalacademies.org/evolution/ National Academies Evolution Resources]
- [http://www.evowiki.org/index.php/Main_Page EvoWiki] — A wiki whose goal is to promote general evolution education, and provide mainstream scientific responses to the arguments of antievolutionists.
- [http://www.chains-of-reason.org/chains/evolution-by-natural-selection/introduction.htm Evolution by Natural Selection] — An introduction to the logic of the theory of evolution by natural selection
- [http://www.pbs.org/wgbh/evolution/index.html Evolution] — Provided by PBS.
- [http://www.newscientist.com/channel/life/evolution Everything you wanted to know about evolution] — Provided by New Scientist.
- [http://evol.allenpress.com/evolonline/?request=index-html International Journal of Organic Evolution]
- [http://science.howstuffworks.com/evolution.htm/printable Howstuffworks.com — How Evolution Works]
- [http://pages.britishlibrary.net/charles.darwin/ Charles Darwin's writings]
- [http://www.genomenewsnetwork.org/categories/index/genome/evolution.php Evolution News from Genome News Network (GNN)]
- [http://www.nap.edu/books/0309063647/html/ National Academy Press: Teaching About Evolution and the Nature of Science]
- [http://www.evolution.mbdojo.com/evolution-for-beginners.html Evolution for beginners]
- [http://www.rmcybernetics.com/science/cybernetics/ai.htm RMCybernetics - AI] Evolution can create emergent behavior in a computer program.
- [http://www.sciencefriday.com/pages/2005/Nov/hour2_111805.html NPR - Science Friday: links to museums, articles and books.]
Evolution Simulators

- [http://www.truthtree.com/evolve.shtml Isolated species evolves to interact more efficiently with its environment (java applet)]
- [http://obermuhlner.com/public/Projects/Applets/Blobs/index.html Evolution in a predator-prey relationship (java applet)] Category:Evolutionary biology Category:Theories ko:진화 ja:進化 th:วิวัฒนาการ

Climate

The climate (ancient Greek: κλίμα) is the weather averaged over a long period of time. The Intergovernmental Panel on Climate Change (IPCC) glossary definition is: : Climate in a narrow sense is usually defined as the “average weather”, or more rigorously, as the statistical description in terms of the mean and variability of relevant quantities over a period of time ranging from months to thousands or millions of years. The classical period is 30 years, as defined by the World Meteorological Organization (WMO). These quantities are most often surface variables such as temperature, precipitation, and wind. Climate in a wider sense is the state, including a statistical description, of the climate system.[http://www.grida.no/climate/ipcc_tar/wg1/518.htm]

Climate vs weather

In the most succinct words, weather is the combination of events in the atmosphere and climate is the overall accumulated weather in a certian location. The exact boundaries of what is climate and what is weather are not well defined and depend on the application. For example, in some senses an individual El Niño event could be considered climate; in others, as weather. When the original conception of climate as a long-term average came to be considered, perhaps towards the end of the 19th century, the idea of climate change was not current, and a 30 year average seemed reasonable (but see note 1). Given the current availability of long-term trends in the temperature record, it is harder to give a precise contradiction-free definition of climate: over a 30 year period, averages may shift; over a shorter period, the statistics are less stable.

Climate determinants

In a given geographical region, the climate generally does not vary over time on the scale of a human life span. However, over geological time, climate can vary considerably for a given place on the Earth. For example, Scandinavia has been through a number of ice ages over hundreds of thousands of years (the last one ending about 10,000 years ago). Paleoclimatology is the study of these past climates, their origin, and by extension, the origin of today's climate. Over historic time spans there are a number of static variables that determine climate including: altitude, proportion of land to water, and proximity to oceans and mountains. Other climate determinants are more dynamic: The Thermohaline circulation of the ocean distributes heat energy between the equatorial and polar regions; other ocean currents do the same between land and water on a more regional scale. Degree of vegetation coverage affects solar heat absorption, water retention, and rainfall on a regional level. Alterations in the quantity of atmospheric greenhouse gases determines the amount of solar energy retained by the planet, leading to global warming (or cooling). The variables which determine climate are numerous and the interactions complex but there is general agreement that the broad outlines are understood, at least in so far as the determinates of historical climate change are concerned.

Climate indices

Scientists use climate indices in their attempt to characterize and understand the various climate mechanisms that culminate in our daily weather. Much in the way the Dow Jones Industrial Average, which is based on the stock prices of 30 companies, is used to represent the fluctuations in the stock market as a whole, climate indices are used to represent the essential elements of climate. Climate indices are generally identified or devised with the twin objectives of simplicity and completeness, and each typically represents the status and timing of the climate factor they represent. By their very nature, indices are simple, and combine many details into an generalized, overall description of the atmosphere or ocean which can be used to characterize the factors which impact the global climate system. Because the climate indices are generally determined from measurements made in a localized area, they can have impacts in other areas around the globe, through processes sometimes called teleconnections. References:
- [http://www.arctic.noaa.gov/essay_bond.html Why and how do scientists study climate change in the Arctic? What are the Arctic climate indices?]
- [http://www.arctic.noaa.gov/climate.html Climate index and mode information]

Classifications

In the original sense, climate is a concept used to divide the world into regions sharing similar climatic parameters. Climate regions can be classified on the basis of temperature and precipitation alone. Examples of such climate schemes are the Köppen climate classification or the Thornthwaite climate classification schemes. For more details about specific climates, please see:
- Tropical climate
- Subtropical climate
- Arid climate
- Semiarid climate
- Mediterranean climate
- Temperate climate
- Oceanic climate
- Continental climate
- Alpine climate
- Subarctic climate
- Polar climate
- Climate of Antarctica To understand a climate of a specific place or area, please see the article on that place or area.

Historical climates

- Climate changes of 535-536
- Medieval climate optimum

National climates

- Climate of the Alps
- Climate of India
- Climate of the United Kingdom

External links

- [http://climateapps2.oucs.ox.ac.uk/cpdnboinc/ Climate Prediction Project]
- [http://www.worldclimate.com WorldClimate]
- [http://www.atmosphere.mpg.de/enid/1442 ESPERE Climate Encyclopaedia]
- [http://www.weatherbase.com Weatherbase]
- [http://www.climate-zone.com Global Climate Data]
- [http://www.limaperunet.com/climate/climateall.html The Climate of Peru]
- [http://www.arctic.noaa.gov/climate.html Climate index and mode information]
- [http://www.arctic.noaa.gov/essay_bond.html Why and how do scientists study climate change in the Arctic? What are the Arctic climate indices?]
- [http://www.arctic.noaa.gov/detect/ A near-realtime Arctic Change Indicator Website]
- [http://www.beringclimate.noaa.gov/ A current view of the Bering Sea Ecosystem and Climate]

Notes

# In "Climatology" by W G Kendrew (OUP; 3rd edition 1949; chapter 38; page 359) we find: "A well-known cycle is one with a mean period of about 35 years... which was worked out by Bruckner... the reality of this cycle seems to be well established, though it is of little use for actual forecasting; it is a basis of the choice of 35 years as the period estimated to give true mean values of climate elements." Category:Ecology ko:기후 ja:気候 simple:Climate

Forest

:This article is about forests as communities of trees. For other uses of the word, see Forest (disambiguation). Forest (disambiguation) A forest is an area with a high density of trees (or, historically, a wooded area set aside for hunting). These plant communities cover large areas of the globe and function as carbon dioxide sinks, animal habitats, hydrologic flow modulators, and soil conservers, constituting one of the most important aspects of the Earth's biosphere. Forests can be found in all regions capable of sustaining tree growth, at altitudes up to the tree-line, except where natural fire frequency is too high, or where the tree growing environment has been impaired by natural processes or humans. Forests sometimes contain many tree species in a small area (e.g. tropical rain and temperate deciduous forests), but other forest types have relatively few species over large areas (e.g. taiga and arid montane coniferous forests). As a general rule, forests dominated by angiosperms are species-rich, while those dominated by gymnosperms are not so rich, although exceptions do exist (e.g., species-poor aspen and birch stands in northern latitudes). Forests are often home to many animal and plant species, and biomass per unit area is high compared to other vegetation types. Much of this biomass occurs below-ground in the root systems and as partially decomposed plant detritus. The woody component of forests contains lignin, which is relatively slow to decompose compared with other organic materials such as cellulose or carbohydrate. Forests are differentiated from woodlands by the extent of canopy coverage: in a forest the branches and foliage of separate trees often meet or interlock, although there can be gaps of varying sizes within an area referred to as forest. A woodland has a more continuously open canopy, with trees spaced further apart, which allows more sunlight to penetrate to the ground between them (see also: savanna). savanna Among the major forested biomes are:
- rain forest (tropical and temperate)
- taiga
- temperate hardwood forest
- tropical dry forest

Classification

tropical dry forest Forests can be classified in different ways and to different degrees of specificity. One such way is in terms of the biome in which they exist combined with leaf longevity of the dominant species (whether they are evergreen or deciduous). Another distinction is whether the forests composed predominantly of broadleaf trees, coniferous (needle-leaved) trees, or mixed.
- Boreal forests occupy the subarctic zone and are generally evergreen and coniferous.
- Temperate zones support both broadleaf deciduous forests (e.g., temperate deciduous forest) and evergreen coniferous forests (e.g., Temperate coniferous forests and Temperate rainforests). Warm temperate zones support broadleaf evergreen forests, including laurel forests.
- Tropical and subtropical forests include tropical rainforests, tropical and subtropical moist forests, tropical and subtropical dry forests, and tropical and subtropical coniferous forests.
- Physiognomy classifies forests based on their overall physical structure or developmental stage (e.g. old growth vs. second growth).
- Forests can also be classified more specifically based on the dominant tree species present, resulting in numerous different forest types (e.g., ponderosa pine/Douglas-fir forest).

Forest management

The scientific study of forests is referred to as forest ecology, while the management of forests is often referred to as forestry, often with the goal of sustainable resource extraction. Forest ecologists concentrate on forest patterns and processes, usually with the aim of elucidating cause and effect relationships. Foresters often focus on wood extraction and silviculture, including tree regeneration and growth processes. Forests can be damaged by logging, forest fires, acid rain, herbivores, and diseases, among other things. In the United States, most forests have historically been affected by humans to some degree, though in recent years environmental protection has helped regulate or moderate large scale or severe impacts. For more comprehensive information on this sub-topic visit the [http://www.iifm.ac.in Indian Institute of Forest Management] in India.

Predator

, but the goose is too wary.]] A predator is an animal or other organism (such as a carnivorous plant) that hunts and kills other organisms for food in an act called predation. Predators are either carnivores or omnivores. The difference between a predator and a parasite is that for a predator killing the prey is necessary for consuming it, but for parasites it is not even desirable because a parasite lives on or in its host. Some might consider herbivores to be predators as well, but this is arguable as most herbivores only consume parts of their food species, leaving the remainder alive. However, where the "prey" consists of single-celled algae, the activities of the herbivorous grazer is generally of the same nature as that of a carnivore. As usual in ecology as most fields of study, there is seldom consensus on the distinctions; some ecologists prefer functional definitions like the one outlined above, others rather look at the ecological dynamics the relationships between the species create. There may be hierarchies of predators; for example, though small birds prey on insects, they may in turn be prey for snakes, which may in turn be prey for hawks. A predator at the top of its food chain (that is, one that is preyed upon by no organism) is called an apex predator; examples include the Great White Shark, Tiger and Crocodile. Sometimes a predator may have a profound influence on the balance of organisms in a particular ecosystem; introduction or removal of this predator, or changes in its population, can have drastic cascading effects on the equilibrium of many other populations in the ecosystem. In this instance the organism may be described as an apex predator or keystone predator. The Volterra-Lotka equations describe a simple mathematical model of the interaction between predators and their prey. Category:Ecology

North America

North America is a continent in the northern hemisphere bordered on the north by the Arctic Ocean, on the east by the North Atlantic Ocean, on the south by the Caribbean Sea, and on the west by the North Pacific Ocean. It covers an area of 24,497,994 km² (9,458,728 sq mi), or about 4.8% of the Earth's surface. As of July 2002, its population was estimated at more than 514,600,000. It is the third largest continent in area, after Asia and Africa, and is fourth in population after Asia, Africa, and Europe. Both North and South America are named after Amerigo Vespucci, who was the first European to suggest that the Americas were not the East Indies, but a previously undiscovered (by Europeans) New World. North America occupies the northern portion of the landmass generally referred to as the New World, the Western Hemisphere, the Americas, or simply America. North America's only land connection is to South America at the narrow Isthmus of Panama. (For geopolitical reasons, all of Panama – including the segment east of the Panama Canal in the isthmus – is often considered a part of North America alone.) According to some authorities, North America begins not at the Isthmus of Panama but at the Isthmus of Tehuantepec, with the intervening region called Central America and resting on the Caribbean Plate. Most, however, tend to see Central America as a region of North America, considering it too small to be a continent on its own. Greenland, although a part of North America geographically, is not considered to be part of the continent politically.

Physical features

Greenland, plutonic, metamorphic rock types of North America. ]] Plate tectonics recognizes the vast majority of North America as being the surface of the North American Plate. Parts of California and western Mexico are known for being the edge of the Pacific Plate, with the two plates meeting along the San Andreas fault. The continent can be divided into four great regions (each of which contains many sub-regions): the Great Plains stretching from the Gulf of Mexico to the Canadian Arctic; the geologically young, mountainous west, including the Rocky Mountains, the Great Basin, California and Alaska; the raised but relatively flat plateau of the Canadian Shield in the northeast; and the varied eastern region, which includes the Appalachian Mountains, the coastal plain along the Atlantic seaboard, and the Florida peninsula. Mexico, with its long plateaus and cordilleras, falls largely in the western region, although the eastern coastal plain does extend south along the Gulf. The western mountains are split in the middle, into the main range of the Rockies and the coast ranges in California, Oregon, Washington, and British Columbia with the Great Basin – a lower area containing smaller ranges and low-lying deserts – in between. The highest peak is Denali in Alaska. Since 1931, Rugby, North Dakota, has officially been recognized as being at the geographic center of North America. The location is marked by a 4.5 metre (15 foot) field stone obelisk. Image:North america terrain 2003 map.jpg|North America bedrock and terrain. Image:North america basement rocks.png|North American cratons and basement rocks. Image:North America Tectonic Elements.jpg|Tectonic elements of North America Image:North america craton nps.gif|North American craton.

Territories and regions

craton On the main continent landmass, there are three large and relatively populous countries:
- Canada - many large islands off the shore of North America belong to Canada, including Vancouver Island and the Queen Charlotte Islands on the west, Prince Edward Island, Newfoundland and Cape Breton Island on the east, and the Canadian Arctic islands (including Ellesmere Island, Baffin Island, and Victoria Island) in the north
- Mexico - the Revillagigedo archipelago and numerous smaller islands off its coast belong to Mexico
- The United States - the 48 contiguous states and Alaska are part of North America, while the state of Hawaii in the Pacific Ocean is not; the Aleutian Islands south of Alaska also belong to the U.S. At the southern end of the continent, in a relatively small area known as Central America, are the countries of:
- Belize
- Costa Rica
- El Salvador
- Guatemala
- Honduras
- Nicaragua
- Panama ¹ At the southeastern end of the continent lies a chain of islands territories called the Antilles, the Caribbean or the West Indies, which include the countries:
- Antigua and Barbuda
- Bahamas
- Barbados
- Cuba
- Dominica
- Dominican Republic
- Grenada
- Haiti
- Jamaica
- Saint Kitts and Nevis
- Saint Lucia
- Saint Vincent and the Grenadines
- Trinidad and Tobago ¹ And the dependencies:
- Anguilla (British overseas territory)
- Aruba ² (part of the Kingdom of the Netherlands)
- Cayman Islands (British overseas territory)
- Guadeloupe (French région d'outre-mer)
- Martinique (French région d'outre-mer)
- Montserrat (British overseas territory)
- Navassa Island (U.S. territory)
- Netherlands Antilles ¹ (part of the Kingdom of the Netherlands)
- Puerto Rico (U.S. commonwealth)
- Turks and Caicos Islands (British overseas territory)
- British Virgin Islands (British overseas territory)
- U.S. Virgin Islands (territory of the USA) Lying in the Atlantic Ocean but considered part of the continent are the dependencies:
- Bermuda, a British overseas territory found about 1,072 km (670 mi.) southeast of New York City
- Greenland, the largest island in the world and a self-governing dependency of Denmark, which is located in the far north of the continent to the east of Nunavut.
- Saint Pierre and Miquelon, a French collectivité d'outre-mer off the south coast of Newfoundland, is the last of France's once vast possessions in America north of the Caribbean. ¹ These states and dependencies have territory both in North and South America.
² These dependencies lie in South America, but are considered North American because of cultural and historical reasons.
See here for details.

Usage

The United States, Canada, and the other English-speaking nations of the Americas (Belize, Guyana, and the Anglophone Caribbean) are sometimes grouped under the term Anglo-America, while the remaining nations of North and South America are grouped under the term Latin America. Alternatively, Northern America is used to refer to Canada and the U.S. together (plus Greenland and Bermuda), while Central America is mainland North America south of the United States. The West Indies generally include all islands in the Caribbean Sea. In this respect, Latin America generally includes Central America and South America and, sometimes, the West Indies. The term Middle America is sometimes used to refer to Mexico, Central America, and the Caribbean collectively. The term "North America" may mean different things to different people. The term in common usage is often taken to mean "the United States and Canada, only" by some people of the United States and Canada, excluding Mexico and the countries of Central America, unless the context makes it clear that they are to be included (such as with specific reference to Mexico, when talking about NAFTA). For example, guides to wild flora and fauna published by the National Audubon Society for "North America" frequently include only species found in Canada and the U.S. This may be attributed to the fact that culturally and economically, the U.S. and Canada are more alike to each other than they are to the rest of North America. Mexicans, however, are acutely aware that Mexico is a part of North America and object to this usage. Central Americans, however, are generally content to be called Central Americans – largely because of their shared history, which includes several attempts at supranational integration in the region and in which Mexico, their much larger northern neighbor, was never involved.

Political divisions and regions

Notes:
¹ Continental regions as per UN categorisations/map.
² Depending on definitions, Aruba, Netherlands Antilles, Panama, and Trinidad and Tobago have territory in one or both of North and South America.
³ Due to ongoing activity of the Soufriere Hills volcano beginning 1995, much of Plymouth, Montserrat's de jure capital, was destroyed and government offices relocated to Brades.

External links

- http://www.america-norte.com/america-norte-mapa.htm Category:Continents Category:North America zh-min-nan:Pak Bí-chiu ko:북아메리카 ja:北アメリカ simple:North America th:ทวีปอเมริกาเหนือ

Skeleton

In biology, the skeleton or skeletal system is the biological system providing support in living organisms. (By extension, non-biological outline structures such as gantries or buildings may also acquire skeletons.) Skeletal systems are commonly divided into three types - external (an exoskeleton), internal (an endoskeleton), and fluid based (a hydrostatic skeleton), though hydrostatic skeletal systems may be classified separately from the other two since they lack hardened support structures. Large external skeletal systems support proportionally less weight than endoskeletons of the same size, and thus many larger animals, such as the vertebrates, have internal skeletal systems. Examples of exoskeletons are found in arthropods, shellfish, and some insects: the skeleton forms a hard shell-like covering protecting the internal organs. The phyla arthropoda and mollusca both have exoskeletons. Since exoskeletons necessarily limit growth, phyla with exoskeletons have come up with various solutions. Most molluscs have calcareous shells and as they grow, the diameter of the shell is enlarged without altering its shape. On the other hand, arthropods shed their exoskeletons to grow, a process known as molting (or ecdysis). During molting the arthropod breaks down their old exoskeleton and then regenerates a new one which they then harden through various processes (such as calcification or sclerotization). An internal skeletal system consists of rigid structures within the body, moved by the muscular system. If the structures are mineralized or ossified, as they are in humans and other mammals, they are referred to as bones. Cartilage is another common component of skeletal systems, supporting and supplementing the skeleton. The human ear and nose are shaped by cartilage. Some organisms have a skeleton consisting entirely of cartilage and without any calcified bones at all, for example sharks. The bones or other rigid structures are connected by ligaments and connected to the muscular system via tendons. Hydrostatic skeletons are similar to a water-filled balloon. Located internally in cnidarians (coral, jellyfish, etc.) and annelids (leeches), among others, these animals can move by contracting the muscles surrounding the fluid-filled pouch, creating pressure within the pouch that causes movement. Animals such as earthworms use their hydrostatic skeletons to change their body shape as they move forward from long and skinny to short and stumpy.

Species

In biology, a species is the basic unit of biodiversity. In scientific classification, a species is assigned a two-part name in Latin. The genus is listed first (and capitalized), followed by a specific epithet. For example, humans belong to the genus Homo, and are in the species Homo sapiens. The name of the species is the whole binomial not just the second term (the specific epithet). The binomial, and most other purely formal aspects of the biological codes of nomenclature, were formalized by Carolus Linnaeus in the 1700's and as a result are called the "Linnaean system". At that time, species were thought to represent independent acts of creation by God, and were therefore considered objectively real and immutable. Since the advent of the theory of evolution, the conception of species has undergone vast changes in biology, however no consensus on the definition of the word has yet been reached. The most commonly cited definition of "species" was first coined by Ernst Mayr. By this definition, called the biological species concept or isolation species concept, species are "groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups". However, many other species concepts are also used (see other definitions of species below). The scientific name of a species is properly typeset in italics. When an unknown species is being referred to this may be done by using the abbreviation "sp." in the singular or "spp." in the plural in the place of the second part of the scientific name. Note that the word "specie" is not the singular of "species". It refers to coined money.

Definitions of species

The definition of a species given above as taken from Mayr, is somewhat idealistic. Since it assumes sexual reproduction, it leaves the term undefined for a large class of organisms that reproduce asexually. Biologists frequently do not know whether two morphologically similar groups of organisms are "potentially" capable of interbreeding. Further, there is considerable variation in the degree to which hybridization ma

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