96-647: Troodontidae / t r oʊ . ə ˈ d ɒ n t ɪ d iː / is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous . During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discoveries of complete and articulated specimens (including specimens which preserve feathers , eggs , embryos , and complete juveniles), have helped to increase understanding about this group. Anatomical studies, particularly studies of
192-471: A jigsaw puzzle . Rocks normally form relatively horizontal layers, with each layer younger than the one underneath it. If a fossil is found between two layers whose ages are known, the fossil's age must lie between the two known ages. Because rock sequences are not continuous, but may be broken up by faults or periods of erosion , it is very difficult to match up rock beds that are not directly next to one another. However, fossils of species that survived for
288-588: A population , or a species ( extinct or extant ). Clades are nested, one in another, as each branch in turn splits into smaller branches. These splits reflect evolutionary history as populations diverged and evolved independently. Clades are termed monophyletic (Greek: "one clan") groups. Over the last few decades, the cladistic approach has revolutionized biological classification and revealed surprising evolutionary relationships among organisms. Increasingly, taxonomists try to avoid naming taxa that are not clades; that is, taxa that are not monophyletic . Some of
384-479: A "ladder", with supposedly more "advanced" organisms at the top. Taxonomists have increasingly worked to make the taxonomic system reflect evolution. When it comes to naming , this principle is not always compatible with the traditional rank-based nomenclature (in which only taxa associated with a rank can be named) because not enough ranks exist to name a long series of nested clades. For these and other reasons, phylogenetic nomenclature has been developed; it
480-446: A branch of mammals that split off after the end of the period when the clade Dinosauria stopped being the dominant terrestrial vertebrates 66 million years ago. The original population and all its descendants are a clade. The rodent clade corresponds to the order Rodentia, and insects to the class Insecta. These clades include smaller clades, such as chipmunk or ant , each of which consists of even smaller clades. The clade "rodent"
576-623: A clade can be described based on two different reference points, crown age and stem age. The crown age of a clade refers to the age of the most recent common ancestor of all of the species in the clade. The stem age of a clade refers to the time that the ancestral lineage of the clade diverged from its sister clade. A clade's stem age is either the same as or older than its crown age. Ages of clades cannot be directly observed. They are inferred, either from stratigraphy of fossils , or from molecular clock estimates. Viruses , and particularly RNA viruses form clades. These are useful in tracking
672-552: A collision that formed the Moon about 40 million years later, may have cooled quickly enough to have oceans and an atmosphere about 4,440 million years ago . There is evidence on the Moon of a Late Heavy Bombardment by asteroids from 4,000 to 3,800 million years ago . If, as seems likely, such a bombardment struck Earth at the same time, the first atmosphere and oceans may have been stripped away. Paleontology traces
768-602: A common ancestor. Ideally the "family tree" has only two branches leading from each node ("junction"), but sometimes there is too little information to achieve this, and paleontologists have to make do with junctions that have several branches. The cladistic technique is sometimes fallible, as some features, such as wings or camera eyes , evolved more than once, convergently – this must be taken into account in analyses. Evolutionary developmental biology , commonly abbreviated to "Evo Devo", also helps paleontologists to produce "family trees", and understand fossils. For example,
864-456: A constant rate. These " molecular clocks ", however, are fallible, and provide only a very approximate timing: for example, they are not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first evolved, and estimates produced by different techniques may vary by a factor of two. Earth formed about 4,570 million years ago and, after
960-403: A data source that is not limited to animals with easily fossilised hard parts, and they reflect organisms' behaviours. Also many traces date from significantly earlier than the body fossils of animals that are thought to have been capable of making them. Whilst exact assignment of trace fossils to their makers is generally impossible, traces may for example provide the earliest physical evidence of
1056-573: A fortunate accident during other research. For example, the 1980 discovery by Luis and Walter Alvarez of iridium , a mainly extraterrestrial metal, in the Cretaceous – Paleogene boundary layer made asteroid impact the most favored explanation for the Cretaceous–Paleogene extinction event – although debate continues about the contribution of volcanism. A complementary approach to developing scientific knowledge, experimental science ,
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#17327919032261152-496: A fourth subfamily of troodontids, the Sinovenatorinae. A simplified version of their analysis is shown below. Dromaeosauridae Eosinopteryx Anchiornis Aurornis Xiaotingia IGM 100/44 Byronosaurus Xixiasaurus Clade In biological phylogenetics , a clade (from Ancient Greek κλάδος (kládos) 'branch'), also known as a monophyletic group or natural group ,
1248-407: A large, curved claw on their retractable second toes, similar to the "sickle-claw" of the dromaeosaurids . However, the sickle-claws of troodontids were not as large or recurved as in dromaeosaurids, and in some instances could not be held off the ground and "retracted" to the same degree. In at least one troodontid, Borogovia , the second toe could not be held far off the ground at all and the claw
1344-507: A long, low opening in the upper jaw (the maxillary fenestra ). Features of the pelvis also suggested they were less advanced than dromaeosaurids. New discoveries of primitive troodontids from China (such as Sinovenator and Mei ), however, display strong similarities between Troodontidae, Dromaeosauridae and the primitive bird Archaeopteryx , and most paleontologists, including Holtz, now consider troodontids to be much more closely related to birds than they are to ornithomimosaurs, causing
1440-821: A minor group until the first jawed fish appeared in the Late Ordovician . The spread of animals and plants from water to land required organisms to solve several problems, including protection against drying out and supporting themselves against gravity . The earliest evidence of land plants and land invertebrates date back to about 476 million years ago and 490 million years ago respectively. Those invertebrates, as indicated by their trace and body fossils, were shown to be arthropods known as euthycarcinoids . The lineage that produced land vertebrates evolved later but very rapidly between 370 million years ago and 360 million years ago ; recent discoveries have overturned earlier ideas about
1536-812: A modified version of the Tsuihiji et al. (2014) analysis. It was in turn based on data published by Gao et al. (2012), a slightly modified version of the Xu et al. (2011) analysis, focusing on advanced troodontids. A simplified version is shown below. Dromaeosauridae Sinovenator Eosinopteryx Liaoningvenator Anchiornis Xiaotingia Talos Mei Byronosaurus IGM 100/140 SPS 100/44 Sinornithoides Gobivenator Linhevenator Philovenator Troodon Saurornithoides Zanabazar In 2014, Brusatte, Lloyd, Wang and Norell published an analysis on Coelurosauria , based on data from Turner et al. (2012) who named
1632-555: A rapid increase in knowledge about the history of life on Earth and to progress in the definition of the geologic time scale , largely based on fossil evidence. Although she was rarely recognised by the scientific community, Mary Anning was a significant contributor to the field of palaeontology during this period; she uncovered multiple novel Mesozoic reptile fossils and deducted that what were then known as bezoar stones are in fact fossilised faeces . In 1822 Henri Marie Ducrotay de Blainville , editor of Journal de Physique , coined
1728-543: A relatively short time can be used to link up isolated rocks: this technique is called biostratigraphy . For instance, the conodont Eoplacognathus pseudoplanus has a short range in the Middle Ordovician period. If rocks of unknown age are found to have traces of E. pseudoplanus , they must have a mid-Ordovician age. Such index fossils must be distinctive, be globally distributed and have a short time range to be useful. However, misleading results are produced if
1824-422: A revised taxonomy based on a concept strongly resembling clades, although the term clade itself would not be coined until 1957 by his grandson, Julian Huxley . German biologist Emil Hans Willi Hennig (1913–1976) is considered to be the founder of cladistics . He proposed a classification system that represented repeated branchings of the family tree, as opposed to the previous systems, which put organisms on
1920-503: A single tooth, and small maniraptoran teeth from the Middle Jurassic of England were identified as those of indeterminate troodontids in 2023. Over the Cretaceous , troodontids radiated throughout western North America , Asia , and Europe , suggesting a mostly Laurasian distribution for the group. However, in 2013, a single diagnostic tooth from the latest Cretaceous ( Maastrichtian ) Kallamedu Formation of southern India
2016-608: A steady increase in brain size after about 3 million years ago . There is a long-running debate about whether modern humans are descendants of a single small population in Africa , which then migrated all over the world less than 200,000 years ago and replaced previous hominine species, or arose worldwide at the same time as a result of interbreeding . Life on earth has suffered occasional mass extinctions at least since 542 million years ago . Despite their disastrous effects, mass extinctions have sometimes accelerated
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#17327919032262112-429: A suffix added should be e.g. "dracohortian". A clade is by definition monophyletic , meaning that it contains one ancestor which can be an organism, a population, or a species and all its descendants. The ancestor can be known or unknown; any and all members of a clade can be extant or extinct. The science that tries to reconstruct phylogenetic trees and thus discover clades is called phylogenetics or cladistics ,
2208-416: A theropod family. Since 1969, Troodontidae has typically been allied with Dromaeosauridae , in a clade (natural group) known as Deinonychosauria , but this was by no means a consensus. Holtz (in 1994) erected the clade Bullatosauria, uniting Ornithomimosauria (the "ostrich-dinosaurs") and Troodontidae, on the basis of characteristics including, among others, an inflated braincase ( parabasisphenoid ) and
2304-499: A third subfamily of troodontids, Jinfengopteryginae. Their analysis included more basal troodontid species but failed to resolve many of their interrelationships, resulting in large " polytomies " (sets of species where the branching order in the family tree is uncertain). An updated version of the Brusatte et al. analysis was provided by Shen et at. (2017b), who included more taxa and recovered greater resolution. Shen et at. named
2400-546: Is a grouping of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree . In the taxonomical literature, sometimes the Latin form cladus (plural cladi ) is used rather than the English form. Clades are the fundamental unit of cladistics , a modern approach to taxonomy adopted by most biological fields. The common ancestor may be an individual,
2496-595: Is composed only of eukaryotic cells, and the earliest evidence for it is the Francevillian Group Fossils from 2,100 million years ago , although specialisation of cells for different functions first appears between 1,430 million years ago (a possible fungus) and 1,200 million years ago (a probable red alga ). Sexual reproduction may be a prerequisite for specialisation of cells, as an asexual multicellular organism might be at risk of being taken over by rogue cells that retain
2592-482: Is hard to decide at what level to place a new higher-level grouping, e.g. genus or family or order ; this is important since the Linnaean rules for naming groups are tied to their levels, and hence if a group is moved to a different level it must be renamed. Paleontologists generally use approaches based on cladistics , a technique for working out the evolutionary "family tree" of a set of organisms. It works by
2688-476: Is in turn included in the mammal, vertebrate and animal clades. The idea of a clade did not exist in pre- Darwinian Linnaean taxonomy , which was based by necessity only on internal or external morphological similarities between organisms. Many of the better known animal groups in Linnaeus's original Systema Naturae (mostly vertebrate groups) do represent clades. The phenomenon of convergent evolution
2784-443: Is often said to work by conducting experiments to disprove hypotheses about the workings and causes of natural phenomena. This approach cannot prove a hypothesis, since some later experiment may disprove it, but the accumulation of failures to disprove is often compelling evidence in favor. However, when confronted with totally unexpected phenomena, such as the first evidence for invisible radiation , experimental scientists often use
2880-594: Is one that contained an extinct "crocodile-like" marine reptile, which eventually came to be known as the mosasaurid Mosasaurus of the Cretaceous period. The first half of the 19th century saw geological and paleontological activity become increasingly well organised with the growth of geologic societies and museums and an increasing number of professional geologists and fossil specialists. Interest increased for reasons that were not purely scientific, as geology and paleontology helped industrialists to find and exploit natural resources such as coal. This contributed to
2976-401: Is our only means of giving rocks greater than about 50 million years old an absolute age, and can be accurate to within 0.5% or better. Although radiometric dating requires very careful laboratory work, its basic principle is simple: the rates at which various radioactive elements decay are known, and so the ratio of the radioactive element to the element into which it decays shows how long ago
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3072-515: Is responsible for many cases of misleading similarities in the morphology of groups that evolved from different lineages. With the increasing realization in the first half of the 19th century that species had changed and split through the ages, classification increasingly came to be seen as branches on the evolutionary tree of life . The publication of Darwin's theory of evolution in 1859 gave this view increasing weight. In 1876 Thomas Henry Huxley , an early advocate of evolutionary theory, proposed
3168-489: Is still controversial. As an example, see the full current classification of Anas platyrhynchos (the mallard duck) with 40 clades from Eukaryota down by following this Wikispecies link and clicking on "Expand". The name of a clade is conventionally a plural, where the singular refers to each member individually. A unique exception is the reptile clade Dracohors , which was made by haplology from Latin "draco" and "cohors", i.e. "the dragon cohort "; its form with
3264-456: Is the scientific study of life that existed prior to the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossils to classify organisms and study their interactions with each other and their environments (their paleoecology ). Paleontological observations have been documented as far back as the 5th century BC. The science became established in
3360-503: Is thought to have been propelled by coevolution with pollinating insects. Social insects appeared around the same time and, although they account for only small parts of the insect "family tree", now form over 50% of the total mass of all insects. Humans evolved from a lineage of upright-walking apes whose earliest fossils date from over 6 million years ago . Although early members of this lineage had chimp -sized brains, about 25% as big as modern humans', there are signs of
3456-582: The Middle Ages the Persian naturalist Ibn Sina , known as Avicenna in Europe, discussed fossils and proposed a theory of petrifying fluids on which Albert of Saxony elaborated in the 14th century. The Chinese naturalist Shen Kuo (1031–1095) proposed a theory of climate change based on the presence of petrified bamboo in regions that in his time were too dry for bamboo. In early modern Europe ,
3552-538: The Neogene - Quaternary . In deeper-level deposits in western Europe are early-aged mammals such as the palaeothere perissodactyl Palaeotherium and the anoplotheriid artiodactyl Anoplotherium , both of which were described earliest after the former two genera, which today are known to date to the Paleogene period. Cuvier figured out that even older than the two levels of deposits with extinct large mammals
3648-637: The Permian–Triassic extinction event . Amphibians Extinct Synapsids Mammals Extinct reptiles Lizards and snakes Extinct Archosaurs Crocodilians Extinct Dinosaurs Birds Naming groups of organisms in a way that is clear and widely agreed is important, as some disputes in paleontology have been based just on misunderstandings over names. Linnaean taxonomy is commonly used for classifying living organisms, but runs into difficulties when dealing with newly discovered organisms that are significantly different from known ones. For example: it
3744-516: The Permian–Triassic extinction event . A relatively recent discipline, molecular phylogenetics , compares the DNA and RNA of modern organisms to re-construct the "family trees" of their evolutionary ancestors. It has also been used to estimate the dates of important evolutionary developments, although this approach is controversial because of doubts about the reliability of the " molecular clock ". Techniques from engineering have been used to analyse how
3840-454: The embryological development of some modern brachiopods suggests that brachiopods may be descendants of the halkieriids , which became extinct in the Cambrian period. Paleontology seeks to map out how living things have changed through time. A substantial hurdle to this aim is the difficulty of working out how old fossils are. Beds that preserve fossils typically lack the radioactive elements needed for radiometric dating . This technique
3936-526: The " jigsaw puzzles " of biostratigraphy (arrangement of rock layers from youngest to oldest). Classifying ancient organisms is also difficult, as many do not fit well into the Linnaean taxonomy classifying living organisms, and paleontologists more often use cladistics to draw up evolutionary "family trees". The final quarter of the 20th century saw the development of molecular phylogenetics , which investigates how closely organisms are related by measuring
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4032-463: The 18th century as a result of Georges Cuvier 's work on comparative anatomy , and developed rapidly in the 19th century. The term has been used since 1822 formed from Greek παλαιός ( 'palaios' , "old, ancient"), ὄν ( 'on' , ( gen. 'ontos' ), "being, creature"), and λόγος ( 'logos' , "speech, thought, study"). Paleontology lies on the border between biology and geology , but it differs from archaeology in that it excludes
4128-555: The Early Cambrian , along with several "weird wonders" that bear little obvious resemblance to any modern animals. There is a long-running debate about whether this Cambrian explosion was truly a very rapid period of evolutionary experimentation; alternative views are that modern-looking animals began evolving earlier but fossils of their precursors have not yet been found, or that the "weird wonders" are evolutionary "aunts" and "cousins" of modern groups. Vertebrates remained
4224-463: The Earth's organic and inorganic past". William Whewell (1794–1866) classified paleontology as one of the historical sciences, along with archaeology , geology, astronomy , cosmology , philology and history itself: paleontology aims to describe phenomena of the past and to reconstruct their causes. Hence it has three main elements: description of past phenomena; developing a general theory about
4320-412: The ability to reproduce. The earliest known animals are cnidarians from about 580 million years ago , but these are so modern-looking that they must be descendants of earlier animals. Early fossils of animals are rare because they had not developed mineralised , easily fossilized hard parts until about 548 million years ago . The earliest modern-looking bilaterian animals appear in
4416-515: The appearance of moderately complex animals (comparable to earthworms ). Geochemical observations may help to deduce the global level of biological activity at a certain period, or the affinity of certain fossils. For example, geochemical features of rocks may reveal when life first arose on Earth, and may provide evidence of the presence of eukaryotic cells, the type from which all multicellular organisms are built. Analyses of carbon isotope ratios may help to explain major transitions such as
4512-625: The atmosphere increased their effectiveness as nurseries of evolution. While eukaryotes , cells with complex internal structures, may have been present earlier, their evolution speeded up when they acquired the ability to transform oxygen from a poison to a powerful source of metabolic energy. This innovation may have come from primitive eukaryotes capturing oxygen-powered bacteria as endosymbionts and transforming them into organelles called mitochondria . The earliest evidence of complex eukaryotes with organelles (such as mitochondria) dates from 1,850 million years ago . Multicellular life
4608-415: The bodies of ancient organisms might have worked, for example the running speed and bite strength of Tyrannosaurus , or the flight mechanics of Microraptor . It is relatively commonplace to study the internal details of fossils using X-ray microtomography . Paleontology, biology, archaeology, and paleoneurobiology combine to study endocranial casts (endocasts) of species related to humans to clarify
4704-401: The causes of various types of change; and applying those theories to specific facts. When trying to explain the past, paleontologists and other historical scientists often construct a set of one or more hypotheses about the causes and then look for a " smoking gun ", a piece of evidence that strongly accords with one hypothesis over any others. Sometimes researchers discover a "smoking gun" by
4800-763: The characteristics and evolution of humans as a species. When dealing with evidence about humans, archaeologists and paleontologists may work together – for example paleontologists might identify animal or plant fossils around an archaeological site , to discover the people who lived there, and what they ate; or they might analyze the climate at the time of habitation. In addition, paleontology often borrows techniques from other sciences, including biology, osteology , ecology, chemistry , physics and mathematics. For example, geochemical signatures from rocks may help to discover when life first arose on Earth, and analyses of carbon isotope ratios may help to identify climate changes and even to explain major transitions such as
4896-520: The chronological order in which rocks were formed, is useful to both paleontologists and geologists. Biogeography studies the spatial distribution of organisms, and is also linked to geology, which explains how Earth's geography has changed over time. Although paleontology became established around 1800, earlier thinkers had noticed aspects of the fossil record. The ancient Greek philosopher Xenophanes (570–480 BCE) concluded from fossil sea shells that some areas of land were once under water. During
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#17327919032264992-479: The clade Bullatosauria to be abandoned. One study of theropod systematics by members of the Theropod Working Group [ Reasonator search ] has uncovered striking similarities among the most basal dromaeosaurids, troodontids, and Archaeopteryx . This clade is together called Paraves by Novas and Pol. The extensive cladistic analysis conducted by Turner et al. , (2012) supported
5088-445: The date when lineages first appeared. For instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an ancestor of B and C, then A must have evolved more than X million years ago. It is also possible to estimate how long ago two living clades diverged – i.e. approximately how long ago their last common ancestor must have lived – by assuming that DNA mutations accumulate at
5184-594: The development of mammalian traits such as endothermy and hair. After the Cretaceous–Paleogene extinction event 66 million years ago killed off all the dinosaurs except the birds, mammals increased rapidly in size and diversity, and some took to the air and the sea. Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early Cretaceous between 130 million years ago and 90 million years ago . Their rapid rise to dominance of terrestrial ecosystems
5280-561: The development of the body plans of most animal phyla . The discovery of fossils of the Ediacaran biota and developments in paleobiology extended knowledge about the history of life back far before the Cambrian. Increasing awareness of Gregor Mendel 's pioneering work in genetics led first to the development of population genetics and then in the mid-20th century to the modern evolutionary synthesis , which explains evolution as
5376-482: The different levels of deposits represented different time periods in the early 19th century. The surface-level deposits in the Americas contained later mammals like the megatheriid ground sloth Megatherium and the mammutid proboscidean Mammut (later known informally as a "mastodon"), which were some of the earliest-named fossil mammal genera with official taxonomic authorities. They today are known to date to
5472-436: The eardrum may have aided in the detection of low-frequency sounds. In some troodontids, ears were also asymmetrical, with one ear placed higher on the skull than the other, a feature shared only with some owls . The specialization of the ears may indicate that troodontids hunted in a manner similar to owls, using their hearing to locate small prey. Although most paleontologists believe that they were predatory carnivores ,
5568-409: The end of the 20th century have been particularly important as they have provided new information about the earliest evolution of animals, early fish, dinosaurs and the evolution of birds. The last few decades of the 20th century saw a renewed interest in mass extinctions and their role in the evolution of life on Earth. There was also a renewed interest in the Cambrian explosion that apparently saw
5664-410: The evolution of the human brain. Paleontology even contributes to astrobiology , the investigation of possible life on other planets , by developing models of how life may have arisen and by providing techniques for detecting evidence of life. As knowledge has increased, paleontology has developed specialised subdivisions. Vertebrate paleontology concentrates on fossils from the earliest fish to
5760-466: The evolutionary history of life back to over 3,000 million years ago , possibly as far as 3,800 million years ago . The oldest clear evidence of life on Earth dates to 3,000 million years ago , although there have been reports, often disputed, of fossil bacteria from 3,400 million years ago and of geochemical evidence for the presence of life 3,800 million years ago . Some scientists have proposed that life on Earth
5856-555: The exceptional events that cause quick burial make it difficult to study the normal environments of the animals. The sparseness of the fossil record means that organisms are expected to exist long before and after they are found in the fossil record – this is known as the Signor–Lipps effect . Trace fossils consist mainly of tracks and burrows, but also include coprolites (fossil feces ) and marks left by feeding. Trace fossils are particularly significant because they represent
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#17327919032265952-515: The focus of paleontology shifted to understanding evolutionary paths, including human evolution , and evolutionary theory. The last half of the 19th century saw a tremendous expansion in paleontological activity, especially in North America. The trend continued in the 20th century with additional regions of the Earth being opened to systematic fossil collection. Fossils found in China near
6048-449: The following: At the end of the 18th century Georges Cuvier 's work established comparative anatomy as a scientific discipline and, by proving that some fossil animals resembled no living ones, demonstrated that animals could become extinct , leading to the emergence of paleontology. The expanding knowledge of the fossil record also played an increasing role in the development of geology, particularly stratigraphy . Cuvier proved that
6144-580: The fossil record: different environments are more favorable to the preservation of different types of organism or parts of organisms. Further, only the parts of organisms that were already mineralised are usually preserved, such as the shells of molluscs. Since most animal species are soft-bodied, they decay before they can become fossilised. As a result, although there are 30-plus phyla of living animals, two-thirds have never been found as fossils. Occasionally, unusual environments may preserve soft tissues. These lagerstätten allow paleontologists to examine
6240-687: The history and driving forces behind their evolution. Land plants were so successful that their detritus caused an ecological crisis in the Late Devonian , until the evolution of fungi that could digest dead wood. During the Permian period, synapsids , including the ancestors of mammals , may have dominated land environments, but this ended with the Permian–Triassic extinction event 251 million years ago , which came very close to wiping out all complex life. The extinctions were apparently fairly sudden, at least among vertebrates. During
6336-533: The history of Earth's climate and the mechanisms that have changed it – which have sometimes included evolutionary developments, for example the rapid expansion of land plants in the Devonian period removed more carbon dioxide from the atmosphere, reducing the greenhouse effect and thus helping to cause an ice age in the Carboniferous period. Biostratigraphy , the use of fossils to work out
6432-542: The immediate ancestors of modern mammals . Invertebrate paleontology deals with fossils such as molluscs , arthropods , annelid worms and echinoderms . Paleobotany studies fossil plants , algae , and fungi. Palynology , the study of pollen and spores produced by land plants and protists , straddles paleontology and botany , as it deals with both living and fossil organisms. Micropaleontology deals with microscopic fossil organisms of all kinds. Instead of focusing on individual organisms, paleoecology examines
6528-434: The index fossils turn out to have longer fossil ranges than first thought. Stratigraphy and biostratigraphy can in general provide only relative dating ( A was before B ), which is often sufficient for studying evolution. However, this is difficult for some time periods, because of the problems involved in matching up rocks of the same age across different continents . Family-tree relationships may also help to narrow down
6624-538: The interactions between different ancient organisms, such as their food chains , and the two-way interactions with their environments. For example, the development of oxygenic photosynthesis by bacteria caused the oxygenation of the atmosphere and hugely increased the productivity and diversity of ecosystems . Together, these led to the evolution of complex eukaryotic cells, from which all multicellular organisms are built. Paleoclimatology , although sometimes treated as part of paleoecology, focuses more on
6720-463: The internal anatomy of animals that in other sediments are represented only by shells, spines, claws, etc. – if they are preserved at all. However, even lagerstätten present an incomplete picture of life at the time. The majority of organisms living at the time are probably not represented because lagerstätten are restricted to a narrow range of environments, e.g. where soft-bodied organisms can be preserved very quickly by events such as mudslides; and
6816-456: The investigation of evolutionary "family trees" by techniques derived from biochemistry , began to make an impact, particularly when it was proposed that the human lineage had diverged from apes much more recently than was generally thought at the time. Although this early study compared proteins from apes and humans, most molecular phylogenetics research is now based on comparisons of RNA and DNA . Fossils of organisms' bodies are usually
6912-518: The latter term coined by Ernst Mayr (1965), derived from "clade". The results of phylogenetic/cladistic analyses are tree-shaped diagrams called cladograms ; they, and all their branches, are phylogenetic hypotheses. Three methods of defining clades are featured in phylogenetic nomenclature : node-, stem-, and apomorphy-based (see Phylogenetic nomenclature§Phylogenetic definitions of clade names for detailed definitions). The relationship between clades can be described in several ways: The age of
7008-409: The logic that, if groups B and C have more similarities to each other than either has to group A, then B and C are more closely related to each other than either is to A. Characters that are compared may be anatomical , such as the presence of a notochord , or molecular , by comparing sequences of DNA or proteins . The result of a successful analysis is a hierarchy of clades – groups that share
7104-522: The many small, coarsely serrated teeth, large denticle size, and U-shaped jaws of some species (particularly Troodon ) suggest that some species may have been omnivorous or herbivorous . Some suggest that the large denticle size is reminiscent of the teeth of extant iguanine lizards. In contrast, a few species, such as Byronosaurus , had large numbers of needle-like teeth, which seem best-suited for picking up small prey, such as birds, lizards and small mammals . Other morphological characteristics of
7200-769: The monophyly of Troodontidae. There are multiple possibilities of the genera included in Troodontidae as well as how they are related. Very primitive species, such as Anchiornis huxleyi , have alternately been found to be early troodontids, early members of the closely related group Avialae, or more primitive paravians by various studies. The cladogram below follows the results of a study by Lefèvre et al. 2017. Avialae Dromaeosauridae Jinfengopteryx Mei Sinovenator Sinusonasus Sinornithoides Byronosaurus Gobivenator Troodon Borogovia Saurornithoides Zanabazar Shen et al. (2017a) explored troodontid phylogeny using
7296-409: The most informative type of evidence. The most common types are wood, bones, and shells. Fossilisation is a rare event, and most fossils are destroyed by erosion or metamorphism before they can be observed. Hence the fossil record is very incomplete, increasingly so further back in time. Despite this, it is often adequate to illustrate the broader patterns of life's history. There are also biases in
7392-402: The most primitive troodontids, like Sinovenator , demonstrate striking anatomical similarities with Archaeopteryx and primitive dromaeosaurids , and demonstrate that they are relatives comprising a clade called Paraves . The oldest definitive troodontid known is Hesperornithoides from the Late Jurassic of Wyoming . The slightly older Koparion of Utah is only represented by
7488-414: The outcome of events such as mutations and horizontal gene transfer , which provide genetic variation , with genetic drift and natural selection driving changes in this variation over time. Within the next few years the role and operation of DNA in genetic inheritance were discovered, leading to what is now known as the "Central Dogma" of molecular biology . In the 1960s molecular phylogenetics ,
7584-537: The predatory behavior of troodontids, Fowler and colleagues theorize that the longer legs and smaller sickle claws (as compared to dromaeosaurids) indicate a more cursorial lifestyle, though the study indicates that troodontids were still likely to have used the unguals for prey manipulation. The proportions of the metatarsals, tarsals and unguals of troodontids appear indicative of their having nimbler, but weaker feet, perhaps better adapted for capturing and subduing smaller prey. This suggests an ecological separation from
7680-452: The principal types of evidence about ancient life, and geochemical evidence has helped to decipher the evolution of life before there were organisms large enough to leave body fossils. Estimating the dates of these remains is essential but difficult: sometimes adjacent rock layers allow radiometric dating , which provides absolute dates that are accurate to within 0.5%, but more often paleontologists have to rely on relative dating by solving
7776-432: The radioactive element was incorporated into the rock. Radioactive elements are common only in rocks with a volcanic origin, and so the only fossil-bearing rocks that can be dated radiometrically are a few volcanic ash layers. Consequently, paleontologists must usually rely on stratigraphy to date fossils. Stratigraphy is the science of deciphering the "layer-cake" that is the sedimentary record, and has been compared to
7872-556: The region, it has been suggested that the existence of Gondwanan troodontids should be regarded as provisional. Troodontids are a group of small, bird-like, gracile maniraptorans . All troodontids have unique features of the skull, such as large numbers of closely spaced teeth in the lower jaw. Troodontids have sickle-claws and raptorial hands , and some of the highest non- avian encephalization quotients , suggesting that they were behaviourally advanced and had keen senses. They had unusually long legs compared to other theropods, with
7968-482: The relationships between organisms that the molecular biology arm of cladistics has revealed include that fungi are closer relatives to animals than they are to plants, archaea are now considered different from bacteria , and multicellular organisms may have evolved from archaea. The term "clade" is also used with a similar meaning in other fields besides biology, such as historical linguistics ; see Cladistics § In disciplines other than biology . The term "clade"
8064-701: The same approach as historical scientists: construct a set of hypotheses about the causes and then look for a "smoking gun". Paleontology lies between biology and geology since it focuses on the record of past life, but its main source of evidence is fossils in rocks. For historical reasons, paleontology is part of the geology department at many universities: in the 19th and early 20th centuries, geology departments found fossil evidence important for dating rocks, while biology departments showed little interest. Paleontology also has some overlap with archaeology , which primarily works with objects made by humans and with human remains, while paleontologists are interested in
8160-478: The similarity of the DNA in their genomes . Molecular phylogenetics has also been used to estimate the dates when species diverged, but there is controversy about the reliability of the molecular clock on which such estimates depend. The simplest definition of "paleontology" is "the study of ancient life". The field seeks information about several aspects of past organisms: "their identity and origin, their environment and evolution, and what they can tell us about
8256-470: The slow recovery from this catastrophe a previously obscure group, archosaurs , became the most abundant and diverse terrestrial vertebrates. One archosaur group, the dinosaurs, were the dominant land vertebrates for the rest of the Mesozoic , and birds evolved from one group of dinosaurs. During this time mammals' ancestors survived only as small, mainly nocturnal insectivores , which may have accelerated
8352-445: The slower but more powerful Dromaeosauridae. Troodontid fossils were among the first dinosaur remains described. Initially, Leidy (1856) assumed they were lacertilian (lizards), but, by 1924, they were referred to Dinosauria by Gilmore , who suggested that they were ornithischians and allied them with the pachycephalosaurian Stegoceras in a Troodontidae. It was not until 1945 that C.M. Sternberg recognized Troodontidae as
8448-480: The spread of viral infections . HIV , for example, has clades called subtypes, which vary in geographical prevalence. HIV subtype (clade) B, for example is predominant in Europe, the Americas and Japan, whereas subtype A is more common in east Africa. Paleontology Paleontology ( / ˌ p eɪ l i ɒ n ˈ t ɒ l ə dʒ i , ˌ p æ l i -, - ən -/ PAY -lee-on- TOL -ə-jee, PAL -ee-, -ən- ), also spelled palaeontology or palæontology ,
8544-631: The study of anatomically modern humans . It now uses techniques drawn from a wide range of sciences, including biochemistry , mathematics , and engineering. Use of all these techniques has enabled paleontologists to discover much of the evolutionary history of life , almost back to when Earth became capable of supporting life, nearly 4 billion years ago. As knowledge has increased, paleontology has developed specialised sub-divisions, some of which focus on different types of fossil organisms while others study ecology and environmental history, such as ancient climates . Body fossils and trace fossils are
8640-629: The systematic study of fossils emerged as an integral part of the changes in natural philosophy that occurred during the Age of Reason . In the Italian Renaissance, Leonardo da Vinci made various significant contributions to the field as well as depicted numerous fossils. Leonardo's contributions are central to the history of paleontology because he established a line of continuity between the two main branches of paleontology – ichnology and body fossil paleontology. He identified
8736-407: The teeth, such as the detailed form of the denticles and the presence of blood grooves, also seem to indicate carnivory. Analyses of barium / calcium and strontium /calcium ratios, which are higher in carnivores due to bioaccumulation , found low ratios in teeth of Stenonychosaurus , suggesting that it had a diet ranging from mixed to plant-dominant omnivory. Though little is known directly about
8832-406: The word "palaeontology" to refer to the study of ancient living organisms through fossils. As knowledge of life's history continued to improve, it became increasingly obvious that there had been some kind of successive order to the development of life. This encouraged early evolutionary theories on the transmutation of species . After Charles Darwin published Origin of Species in 1859, much of
8928-454: Was "seeded" from elsewhere , but most research concentrates on various explanations of how life could have arisen independently on Earth. For about 2,000 million years microbial mats , multi-layered colonies of different bacteria, were the dominant life on Earth. The evolution of oxygenic photosynthesis enabled them to play the major role in the oxygenation of the atmosphere from about 2,400 million years ago . This change in
9024-423: Was coined in 1957 by the biologist Julian Huxley to refer to the result of cladogenesis , the evolutionary splitting of a parent species into two distinct species, a concept Huxley borrowed from Bernhard Rensch . Many commonly named groups – rodents and insects , for example – are clades because, in each case, the group consists of a common ancestor with all its descendant branches. Rodents, for example, are
9120-414: Was identified as a troodontid, suggesting that troodontids either also inhabited Gondwana or managed to disperse to India from elsewhere prior to its separation as an island continent . The potential Gondwanan occurrence of troodontids is supported by the existence of Middle Jurassic remains, which suggest that they originated prior to the breakup of Pangaea . However, due to the lack of other remains from
9216-409: Was straight, not curved or sickle-like. Troodontids had unusually large brains among dinosaurs, comparable to those of living flightless birds. Their eyes were also large, and pointed forward, indicating that they had good binocular vision . The ears of troodontids were also unusual among theropods, having enlarged middle ear cavities, indicating acute hearing ability. The placement of this cavity near
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