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47-436: Caponiidae is a family of ecribellate haplogyne spiders that are unusual in a number of ways. They differ from other spiders in lacking book lungs and having the posterior median spinnerets anteriorly displaced to form a transverse row with the anterior lateral spinnerets. Most species have only two eyes, which is also unusual among spiders. A few species of Caponiidae variously have four, six or eight eyes. In some species

94-576: A body plan from a common ancestor, and that taxa were branches of a single tree of life . The word homology, coined in about 1656, is derived from the Greek ὁμόλογος homologos from ὁμός homos 'same' and λόγος logos 'relation'. Similar biological structures or sequences in different taxa are homologous if they are derived from a common ancestor . Homology thus implies divergent evolution . For example, many insects (such as dragonflies ) possess two pairs of flying wings . In beetles ,

141-404: A common ancestor after having been subjected to adaptive modifications for different purposes as the result of natural selection . The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin 's theory of evolution in 1859, but had been observed before this from Aristotle's biology onwards, and it

188-465: A gene in an organism is duplicated, the two copies are paralogous. They can shape the structure of whole genomes and thus explain genome evolution to a large extent. Examples include the Homeobox ( Hox ) genes in animals. These genes not only underwent gene duplications within chromosomes but also whole genome duplications . As a result, Hox genes in most vertebrates are spread across multiple chromosomes:

235-477: A pattern of gene expression in the growing zones ( meristems ) is described by the ABC model of flower development . Each of the four types of flower parts is serially repeated in concentric whorls, controlled by a small number of genes acting in various combinations. Thus, A genes working alone result in sepal formation; A and B together produce petals; B and C together create stamens; C alone produces carpels. When none of

282-636: A sister group of the Tetrablemmidae plus the four families inside the superfamily Dysderoidea . The subfamily Nopinae consists at least of the genera Nops , Nopsides , Orthonops and Tarsonops . The remaining genera are unlikely to form a monophyletic group. The family is found in Africa and the Americas from Argentina to the United States. Calponia is a contraction of Californian Caponia , because

329-487: Is a synapomorphy for fleas. Patterns such as these lead many cladists to consider the concept of homology and the concept of synapomorphy to be equivalent. Some cladists follow the pre-cladistic definition of homology of Haas and Simpson, and view both synapomorphies and symplesiomorphies as homologous character states. Homologies provide the fundamental basis for all biological classification, although some may be highly counter-intuitive. For example, deep homologies like

376-445: Is believed that the precursor of all Araneomorphae was cribellate ( symplesiomorphy ), and that this function was lost in some araneomorph spiders secondarily (Coddington & Levy, 1991). Many of these still retain a colulus , which is thought to be a reduced cribellum, and is of unknown function. However, some "ecribellate" spiders seem to have evolved independently, without cribellate precursors (Foelix, 1979). In Austrochilidae ,

423-549: Is defined in terms of shared ancestry. Two segments of DNA can have shared ancestry because of either a speciation event ( orthologs ) or a duplication event ( paralogs ). Homology among proteins or DNA is typically inferred from their sequence similarity. Significant similarity is strong evidence that two sequences are related by divergent evolution of a common ancestor. Alignments of multiple sequences are used to indicate which regions of each sequence are homologous. Homologous sequences are orthologous if they are descended from

470-461: Is named in honor of one of the collectors, P.A. Goloboff. As of April 2019, the World Spider Catalog accepts the following genera: Ecribellate In certain groups of diatoms it refers to microscopically punctured regions of the frustule , or outer layer. In certain groups of spider species, so-called cribellate spiders, the cribellum is a silk spinning organ. Unlike

517-443: Is shared due to common ancestry. Primary homology may be conceptually broken down further: we may consider all of the states of the same character as "homologous" parts of a single, unspecified, transformation series. This has been referred to as topographical correspondence. For example, in an aligned DNA sequence matrix, all of the A, G, C, T or implied gaps at a given nucleotide site are homologous in this way. Character state identity

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564-404: Is strong evidence that two sequences are related by divergent evolution from a common ancestor. Alignments of multiple sequences are used to discover the homologous regions. Homology remains controversial in animal behaviour , but there is suggestive evidence that, for example, dominance hierarchies are homologous across the primates . Homology was noticed by Aristotle (c. 350 BC), and

611-450: Is taken to be homologous. As implied in this definition, many cladists consider secondary homology to be synonymous with synapomorphy , a shared derived character or trait state that distinguishes a clade from other organisms. Shared ancestral character states, symplesiomorphies, represent either synapomorphies of a more inclusive group, or complementary states (often absences) that unite no natural group of organisms. For example,

658-404: Is the hypothesis that the particular condition in two or more taxa is "the same" as far as our character coding scheme is concerned. Thus, two Adenines at the same aligned nucleotide site are hypothesized to be homologous unless that hypothesis is subsequently contradicted by other evidence. Secondary homology is implied by parsimony analysis , where a character state that arises only once on a tree

705-594: The pax6 genes that control the development of the eyes of vertebrates and arthropods were unexpected, as the organs are anatomically dissimilar and appeared to have evolved entirely independently. The embryonic body segments ( somites ) of different arthropod taxa have diverged from a simple body plan with many similar appendages which are serially homologous, into a variety of body plans with fewer segments equipped with specialised appendages. The homologies between these have been discovered by comparing genes in evolutionary developmental biology . Among insects,

752-460: The cribellate and ecribellate (not cribellate) type. The distinction can be used to study evolutionary relationships. However, in 1967 it was discovered that there are many families with both cribellate and ecribellate members ( Lehtinen , 1967). Some species, such as Amaurobius ferox , are also capable of switching between cribellate and ecribellate silk, primarily using cribellate silk for webs and ecribellate silk for trophic eggs. Today, it

799-412: The embryo in the same manner and from similar origins, such as from matching primordia in successive segments of the same animal, are serially homologous . Examples include the legs of a centipede , the maxillary and labial palps of an insect , and the spinous processes of successive vertebrae in a vertebrate's backbone . Male and female reproductive organs are homologous if they develop from

846-488: The frustule , or outside layer of many forms of diatom also are called cribella. In such species of diatom the frustule consists of a thin siliceous plate with many small pores. Homology (biology) In biology , homology is similarity in anatomical structures or genes between organisms of different taxa due to shared ancestry , regardless of current functional differences. Evolutionary biology explains homologous structures as retained heredity from

893-465: The stinger of the female honey bee is a modified ovipositor , homologous with ovipositors in other insects such as the Orthoptera , Hemiptera , and those Hymenoptera without stingers. The three small bones in the middle ear of mammals including humans, the malleus , incus , and stapes , are today used to transmit sound from the eardrum to the inner ear . The malleus and incus develop in

940-423: The "same organ in different animals under every variety of form and function", and contrasting it with the matching term "analogy" which he used to describe different structures with the same function. Owen codified 3 main criteria for determining if features were homologous: position, development, and composition. In 1859, Charles Darwin explained homologous structures as meaning that the organisms concerned shared

987-496: The HoxA–D clusters are the best studied. Some sequences are homologous, but they have diverged so much that their sequence similarity is not sufficient to establish homology. However, many proteins have retained very similar structures, and structural alignment can be used to demonstrate their homology. It has been suggested that some behaviours might be homologous, based either on sharing across related taxa or on common origins of

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1034-458: The anterior middle of the carapace. Caponiidae are unusual in the degree to which the eye number varies. In this they surpass even the family Cybaeidae in which some species have two eyes, some six, and some eight. In some species of the Caponiidae paired eyes meet in the midline, giving the spider in effect, an odd number of eyes. The following genera have eyes as follows: Their habits are for

1081-538: The behaviour in an individual's development; however, the notion of homologous behavior remains controversial, largely because behavior is more prone to multiple realizability than other biological traits. For example, D. W. Rajecki and Randall C. Flanery, using data on humans and on nonhuman primates , argue that patterns of behaviour in dominance hierarchies are homologous across the primates. As with morphological features or DNA, shared similarity in behavior provides evidence for common ancestry. The hypothesis that

1128-477: The cribellum is developed only in the second nymphal stage, so the ecribellate and cribellate conditions change during the spider ontogenesis . Only about 180 genera in 23 families (1991) still contain cribellate members, although the diverse Australian cribellate fauna is still mostly undescribed. However, that fauna may be an example of high diversity in Australian animals that are only relicts in other regions of

1175-482: The embryo from structures that form jaw bones (the quadrate and the articular) in lizards, and in fossils of lizard-like ancestors of mammals. Both lines of evidence show that these bones are homologous, sharing a common ancestor. Among the many homologies in mammal reproductive systems , ovaries and testicles are homologous. Rudimentary organs such as the human tailbone , now much reduced from their functional state, are readily understood as signs of evolution ,

1222-510: The embryos develop. The implication that the ancestors of snakes had hind legs is confirmed by fossil evidence: the Cretaceous snake Pachyrhachis problematicus had hind legs complete with hip bones ( ilium , pubis , ischium ), thigh bone ( femur ), leg bones ( tibia , fibula ) and foot bones ( calcaneum , astragalus ) as in tetrapods with legs today. As with anatomical structures, sequence homology between protein or DNA sequences

1269-472: The explanation being that they were cut down by natural selection from functioning organs when their functions were no longer needed, but make no sense at all if species are considered to be fixed. The tailbone is homologous to the tails of other primates. In many plants, defensive or storage structures are made by modifications of the development of primary leaves , stems , and roots . Leaves are variously modified from photosynthetic structures to form

1316-415: The fibres have a surface that absorbs waxes from the epicuticle of insect prey on contact. This creates a powerful adhesion without any liquid glue that tends to dry out. The spider cribellum is a functional homolog of the anterior median spinnerets of Mesothelae and Mygalomorphae , which do not have a cribellum. The presence or absence of a cribellum is used to classify araneomorph spiders into

1363-455: The first pair of wings has evolved into a pair of hard wing covers , while in Dipteran flies the second pair of wings has evolved into small halteres used for balance. Similarly, the forelimbs of ancestral vertebrates have evolved into the front flippers of whales , the wings of birds , the running forelegs of dogs , deer , and horses , the short forelegs of frogs and lizards , and

1410-445: The genes are active, leaves are formed. Two more groups of genes, D to form ovules and E for the floral whorls, complete the model. The genes are evidently ancient, as old as the flowering plants themselves. Developmental biology can identify homologous structures that arose from the same tissue in embryogenesis . For example, adult snakes have no legs, but their early embryos have limb-buds for hind legs, which are soon lost as

1457-561: The grasping hands of primates including humans. The same major forearm bones ( humerus , radius , and ulna ) are found in fossils of lobe-finned fish such as Eusthenopteron . The opposite of homologous organs are analogous organs which do similar jobs in two taxa that were not present in their most recent common ancestor but rather evolved separately . For example, the wings of insects and birds evolved independently in widely separated groups , and converged functionally to support powered flight , so they are analogous. Similarly,

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1504-597: The insect-trapping pitchers of pitcher plants , the insect-trapping jaws of the Venus flytrap , and the spines of cactuses , all homologous. Certain compound leaves of flowering plants are partially homologous both to leaves and shoots, because their development has evolved from a genetic mosaic of leaf and shoot development. The four types of flower parts, namely carpels , stamens , petals , and sepals , are homologous with and derived from leaves, as Goethe correctly noted in 1790. The development of these parts through

1551-406: The most part unknown. At least some species are known to hunt other spiders. The fact that they are ecribellate and haplogyne suggests that they might be relatively primitive . Calponia harrisonfordi from California seems to be the most primitive member of the family. Their phylogenetic relationships have long been enigmatic, but in the early 1990s it was determined that they are probably

1598-422: The number of eyes will increase when the spiderling changes its skin as it grows towards adulthood. These spiders of about 2 to 5 millimetres (0.079 to 0.197 in) are rarely noticed, but generally look like somewhat faded woodlouse hunter spiders in the genus Dysdera . The carapace (cephalothorax or prosoma) is orange and the abdomen (opisthosoma) light gray. The two-eyed species have their two eyes in

1645-400: The presence of wings is a synapomorphy for pterygote insects, but a symplesiomorphy for holometabolous insects. Absence of wings in non-pterygote insects and other organisms is a complementary symplesiomorphy that unites no group (for example, absence of wings provides no evidence of common ancestry of silverfish, spiders and annelid worms). On the other hand, absence (or secondary loss) of wings

1692-435: The principle of connections, namely that what is important is the relative position of different structures and their connections to each other. Embryologist Karl Ernst von Baer stated what are now called von Baer's laws in 1828, noting that related animals begin their development as similar embryos and then diverge: thus, animals in the same family are more closely related and diverge later than animals which are only in

1739-414: The same order and have fewer homologies. Von Baer's theory recognises that each taxon (such as a family) has distinctive shared features, and that embryonic development parallels the taxonomic hierarchy: not the same as recapitulation theory . The term "homology" was first used in biology by the anatomist Richard Owen in 1843 when studying the similarities of vertebrate fins and limbs, defining it as

1786-423: The same ancestral sequence separated by a speciation event: when a species diverges into two separate species, the copies of a single gene in the two resulting species are said to be orthologous . The term "ortholog" was coined in 1970 by the molecular evolutionist Walter Fitch . Homologous sequences are paralogous if they were created by a duplication event within the genome. For gene duplication events, if

1833-444: The same embryonic tissue, as do the ovaries and testicles of mammals , including humans . Sequence homology between protein or DNA sequences is similarly defined in terms of shared ancestry. Two segments of DNA can have shared ancestry because of either a speciation event ( orthologs ) or a duplication event ( paralogs ). Homology among proteins or DNA is inferred from their sequence similarity. Significant similarity

1880-762: The single species Calponia harrisonfordi has, like the African genus Caponia eight eyes. The species name is in honor of Harrison Ford , recognizing his efforts on behalf of the American Museum of Natural History . The Chilean caponiid fauna differs from that of the rest of the Neotropics in lacking members of the Nopinae (named after the genus Nops ). Three genera newly described by Norman I. Platnick in 1994 were thus named Notnops , Taintnops and Tisentnops , emphasizing this fact. The only Taintnops species, T. goloboffi ,

1927-432: The three groups. Thus, in the pterosaurs, the "wing" involves both the forelimb and the hindlimb. Analogy is called homoplasy in cladistics , and convergent or parallel evolution in evolutionary biology. Specialised terms are used in taxonomic research. Primary homology is a researcher's initial hypothesis based on similar structure or anatomical connections, suggesting that a character state in two or more taxa share

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1974-522: The usual spinnerets of spiders, the cribellum consists of one or more plates covered in thousands of tiny spigots , tiny holes that hardly project from the surface, in contrast to the elongated spigots that project from spinnerets. These minute spigots produce extremely fine fibers, merely tens of nanometres thick, which are combed out by the spider's calamistrum , producing silk with a woolly texture. The fibers are so small in diameter that they are strongly subject to Van der Waals forces . In addition,

2021-436: The wings of a sycamore maple seed and the wings of a bird are analogous but not homologous, as they develop from quite different structures. A structure can be homologous at one level, but only analogous at another. Pterosaur , bird and bat wings are analogous as wings, but homologous as forelimbs because the organ served as a forearm (not a wing) in the last common ancestor of tetrapods , and evolved in different ways in

2068-824: The world, like the marsupials (Coddington & Levy, 1991). Cribellate taxa are not very speciose, and for nearly all cribellate-ecribellate sister clades the cribellate lineage is less diverse (Coddington & Levy, 1991), for example: 22 families of araneomorph spiders, namely Agelenidae , Amaurobiidae , Amphinectidae , Austrochilidae , Ctenidae , Deinopidae , Desidae , Dictynidae , Eresidae , Filistatidae , Gradungulidae , Hypochilidae , Miturgidae , Neolanidae , Nicodamidae , Oecobiidae , Psechridae , Stiphidiidae , Tengellidae , Titanoecidae , Uloboridae and Zoropsidae contain at least some cribellate spiders (Griswold et al. 1999). While some of these families are entirely cribellate, others contain both cribellate and ecribellate species. The perforated regions of

2115-438: Was described late in the 18th century. The French zoologist Etienne Geoffroy Saint-Hilaire showed in 1818 in his theorie d'analogue ("theory of homologues") that structures were shared between fishes, reptiles, birds, and mammals. When Geoffroy went further and sought homologies between Georges Cuvier 's embranchements , such as vertebrates and molluscs, his claims triggered the 1830 Cuvier-Geoffroy debate . Geoffroy stated

2162-426: Was explicitly analysed by Pierre Belon in 1555. A common example of homologous structures is the forelimbs of vertebrates , where the wings of bats and birds , the arms of primates , the front flippers of whales , and the forelegs of four-legged vertebrates like horses and crocodilians are all derived from the same ancestral tetrapod structure. In developmental biology , organs that developed in

2209-680: Was explicitly analysed by Pierre Belon in his 1555 Book of Birds , where he systematically compared the skeletons of birds and humans. The pattern of similarity was interpreted as part of the static great chain of being through the mediaeval and early modern periods: it was not then seen as implying evolutionary change. In the German Naturphilosophie tradition, homology was of special interest as demonstrating unity in nature. In 1790, Goethe stated his foliar theory in his essay "Metamorphosis of Plants", showing that flower parts are derived from leaves. The serial homology of limbs

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