In phylogenetics , long branch attraction ( LBA ) is a form of systematic error whereby distantly related lineages are incorrectly inferred to be closely related. LBA arises when the amount of molecular or morphological change accumulated within a lineage is sufficient to cause that lineage to appear similar (thus closely related) to another long-branched lineage, solely because they have both undergone a large amount of change, rather than because they are related by descent. Such bias is more common when the overall divergence of some taxa results in long branches within a phylogeny . Long branches are often attracted to the base of a phylogenetic tree , because the lineage included to represent an outgroup is often also long-branched. The frequency of true LBA is unclear and often debated, and some authors view it as untestable and therefore irrelevant to empirical phylogenetic inference. Although often viewed as a failing of parsimony-based methodology, LBA could in principle result from a variety of scenarios and be inferred under multiple analytical paradigms.
42-615: The Sphaerolichida is a suborder of mites belonging to the order Trombidiformes . Mite Mites are small arachnids (eight-legged arthropods ). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes , which were historically grouped together in the subclass Acari . However, most recent genetic analyses do not recover the two as each other's closest relative within Arachnida, rendering
84-425: A cephalothorax (with no separate head) or prosoma, and an opisthosoma or abdomen. Segmentation has almost entirely been lost and the prosoma and opisthosoma are fused, only the positioning of the limbs indicating the location of the segments. At the front of the body is the gnathosoma or capitulum. This is not a head and does not contain the eyes or the brain, but is a retractable feeding apparatus consisting of
126-408: A prion disease of sheep. The mite Varroa destructor is a serious pest of honey bees , contributing to colony collapse disorder in commercial hives . This organism is an obligate external parasite, able to reproduce only in bee colonies. It directly weakens its host by sucking up the bee's fat, and can spread RNA viruses including deformed wing virus . Heavy infestation causes the death of
168-583: A colony, generally over the winter. Since 2006, more than 10 million beehives have been lost. Various mites prey on other invertebrates and can be used to control their populations. Phytoseiidae , especially members of Amblyseius , Metaseiulus , and Phytoseiulus , are used to control pests such as spider mites. Among the Laelapidae , Gaeolaelaps aculeifer and Stratiolaelaps scimitus are used to control fungus gnats , poultry red mites and various soil pests. Mites were first observed under
210-600: A few are associated with allergies or may transmit diseases. The scientific discipline devoted to the study of mites is called acarology . The mites are not a defined taxon, but is used for two distinct groups of arachnids, the Acariformes and the Parasitiformes . The phylogeny of the Acari has been relatively little studied, but molecular information from ribosomal DNA is being extensively used to understand relationships between groups. The 18 S rRNA gene provides information on relationships among phyla and superphyla, while
252-522: A few phylogenomic studies have found strong support for monophyly of Acari and a sister relationship between Acariformes and Parasitiformes, although this finding has been questioned, with other studies suggesting that this likely represents a long branch attraction artefact. Mites are tiny members of the class Arachnida ; most are in the size range 250 to 750 μm (0.01 to 0.03 in) but some are larger and some are no bigger than 100 μm (0.004 in) as adults. The body plan has two regions ,
294-439: A million or more species as yet undescribed. The tropical species Archegozetes longisetosus is one of the strongest animals in the world, relative to its mass (100 μg): It lifts up to 1,182 times its own weight, over five times more than would be expected of such a minute animal. A mite also holds a speed record: for its length, Paratarsotomus macropalpis is the fastest animal on Earth. The mites living in soil consist of
336-765: A network of sinuses and most mites lacks a heart, movement of fluid being driven by the contraction of body muscles. But ticks, and some of the larger species of mites, have a dorsal, longitudinal heart. Gas exchange is carried out across the body surface, but many species additionally have between one and four pairs of tracheae . The excretory system includes a nephridium and one or two pairs of Malpighian tubules . Several families of mites, such as Tetranychidae, Eriophyidae, Camerobiidae, Cunaxidae, Trombidiidae, Trombiculidae, Erythraeidae and Bdellidae have silk glands used to produce silk for various purposes. Additionally, water mites (Hydrachnidia) produce long thin threads that may be silk. The sexes are separate in mites; males have
378-418: A pair of testes in the mid-region of the body, each connected to the gonopore by a vas deferens , and in some species there is a chitinous penis ; females have a single ovary connected to the gonopore by an oviduct , as well as a seminal receptacle for the storage of sperm . In most mites, sperm is transferred to the female indirectly; the male either deposits a spermatophore on a surface from which it
420-448: A range of taxa. Oribatida and Prostigmata are more numerous in soil than Mesostigmata, and have more soil-dwelling species. When soil is affected by an ecological disturbance such as agriculture, most mites (Astigmata, Mesostigmata and Prostigmata) recolonise it within a few months, whereas Oribatida take multiple years. Many mites are parasitic on plants and animals. One family of mites, Pyroglyphidae , or nest mites, live primarily in
462-542: A small scale, walking is used to access other suitable locations in the immediate vicinity. Some species mount to a high point and adopt a dispersal posture and get carried away by the wind, while others waft a thread of silk aloft to balloon to a new position. Parasitic mites use their hosts to disperse, and spread from host to host by direct contact. Another strategy is phoresy ; the mite, often equipped with suitable claspers or suckers, grips onto an insect or other animal, and gets transported to another place. A phoretic mite
SECTION 10
#1732783874197504-478: Is a result of the way clustering algorithms work: terminals or taxa with many autapomorphies (character states unique to a single branch) may by chance exhibit the same states as those on another branch ( homoplasy ). A phylogenetic analysis will group these taxa together as a clade unless other synapomorphies outweigh the homoplastic features to group together true sister taxa. These problems may be minimized by using methods that correct for multiple substitutions at
546-525: Is affecting tree topology is the SAW method, named for Siddal and Whiting. If long branch attraction is suspected between a pair of taxa (A and B), simply remove taxon A ("saw" off the branch) and re-run the analysis. Then remove B and replace A, running the analysis again. If either of the taxa appears at a different branch point in the absence of the other, there is evidence of long branch attraction. Since long branches can't possibly attract one another when only one
588-498: Is demonstrated that for certain unknown "true" trees, some methods can show bias for grouping long branches, ultimately resulting in the inference of a false sister relationship. Often this is because convergent evolution of one or more characters included in the analysis has occurred in multiple taxa. Although they were derived independently, these shared traits can be misinterpreted in the analysis as being shared due to common ancestry. In phylogenetic and clustering analyses , LBA
630-409: Is different, and we cannot learn anything, as all trees have the same length. Similarly, A can be - and C can be +. The only remaining possibility is that A and C are both -. In this case, however, we view either A and C, or B and D, as a group with respect to the other (one character state is ancestral, the other is derived, and the ancestral state does not define a group). As a consequence, when we have
672-399: Is in the analysis, consistent taxon placement between treatments would indicate long branch attraction is not a problem. Assume for simplicity that we are considering a single binary character (it can either be + or -) distributed on the unrooted "true tree" with branch lengths proportional to amount of character state change, shown in the figure. Because the evolutionary distance from B to D
714-440: Is just a hitch-hiker and does not feed during the time it is carried by its temporary host. These travelling mites are mostly species that reproduce rapidly and are quick to colonise new habitats. Mites are tiny and apart from those that are of economic concern to humans, little studied. The majority are beneficial, living in the soil or aqueous environments and assisting in the decomposition of decaying organic material, as part of
756-491: Is located on the ventral surface between the fourth pair of legs. Some species have one to five median or lateral eyes but many species are blind, and slit and pit sense organs are common. Both body and limbs bear setae (bristles) which may be simple, flattened, club-shaped or sensory. Mites are usually some shade of brown, but some species are red, orange, black or green, or some combination of these colours. Many mites have stigmata (openings used in respiration). In some mites,
798-405: Is picked up by the female, or he uses his chelicerae or third pair of legs to insert it into the female's gonopore. In some of the Acariformes , insemination is direct using the male's penis. The spermatophora in all mites are aflagellate. The eggs are laid in the substrate , or wherever the mite happens to live. They take up to six weeks to hatch, according to species, with the next stage being
840-453: Is small, we assume that in the vast majority of all cases, B and D will exhibit the same character state. Here, we will assume that they are both + (+ and - are assigned arbitrarily and swapping them is only a matter of definition). If this is the case, there are four remaining possibilities. A and C can both be +, in which case all taxa are the same and all the trees have the same length. A can be + and C can be -, in which case only one character
882-538: Is some other evidence that suggests that the phylogeny is incorrect. For example, two different sources of data (i.e. molecular and morphological) or even different methods or partition schemes might support different placement for the long-branched groups. Hennig's Auxiliary Principle suggests that synapomorphies should be viewed as de facto evidence of grouping unless there is specific contrary evidence (Hennig, 1966; Schuh and Brower, 2009). A simple and effective method for determining whether or not long branch attraction
SECTION 20
#1732783874197924-539: The ITS2 , and the 18S ribosomal RNA and 28S ribosomal RNA genes, provide clues at deeper levels. The mite fossil record is sparse, due to their small size and low preservation potential. The oldest fossils of acariform mites are from the Rhynie Chert , Scotland, which dates to the early Devonian , around 410 million years ago while the earliest fossils of Parasitiformes are known from amber specimens dating to
966-417: The carbon cycle . Two species live on humans, namely Demodex folliculorum and Demodex brevis ; both are frequently referred to as eyelash mites. The majority of mite species are harmless to humans and domestic animals , but a few species can colonize mammals directly, acting as vectors for disease transmission, and causing or contributing to allergenic diseases. Mites which colonize human skin are
1008-638: The chelicerae , the pedipalps and the oral cavity. It is covered above by an extension of the body carapace and is connected to the body by a flexible section of cuticle . Two-segmented chelicerae is the ancestral condition in Acariformes, but in more derived groups they are single-segmented. And three-segmented chelicerae is the ancestral condition in Parasitiformes, but has been reduced to just two segments in more derived groups. The pedipalps differ between taxa depending on diet; in some species
1050-516: The microscope by the English polymath Robert Hooke . In his 1665 book Micrographia , he stated that far from being spontaneously generated from dirt, they were "very prettily shap'd Insects". In 1898, Arthur Conan Doyle wrote a satirical poem, "A Parable", with the conceit of some cheese mites disputing the origin of the round cheddar cheese in which they all lived. The world's first science documentary featured cheese mites , seen under
1092-552: The appendages resemble legs while in others they are modified into chelicerae-like structures. The oral cavity connects posteriorly to the mouth and pharynx . Most mites have four pairs of legs (two pairs in Eriophyoidea ), each with six segments, which may be modified for swimming or other purposes. The dorsal surface of the body is clad in hardened tergites and the ventral surface by hardened sclerites ; sometimes these form transverse ridges. The gonopore (genital opening)
1134-642: The body of honey bees , and Acarapis woodi (family Tarsonemidae ) lives in their tracheae . Hundreds of species are associated with other bees, mostly poorly described. They attach to bees in a variety of ways. For example, Trigona corvina workers have been found with mites attached to the outer face of their hind tibiae. Some are thought to be parasites, while others are beneficial symbionts . Mites also parasitize some ant species, such as Eciton burchellii . Most larvae of Parasitengona are ectoparasites of arthropods, while later life stages in this group tend to shift to being predators. Plant pests include
1176-407: The cause of several types of itchy skin rashes, such as gamasoidosis , rodent mite dermatitis , grain itch , grocer's itch , and scabies ; Sarcoptes scabiei is a parasitic mite responsible for scabies, which is one of the three most common skin disorders in children. Demodex mites, which are common cause of mange in dogs and other domesticated animals, have also been implicated in
1218-428: The disease rickettsialpox . House dust mites , found in warm and humid places such as beds, cause several forms of allergic diseases, including hay fever , asthma and eczema , and are known to aggravate atopic dermatitis . Among domestic animals , sheep are affected by the mite Psoroptes ovis which lives on the skin, causing hypersensitivity and inflammation. Hay mites are a suspected reservoir for scrapie ,
1260-528: The group non- monophyletic . Most mites are tiny, less than 1 mm (0.04 in) in length, and have a simple, unsegmented body plan. The small size of most species makes them easily overlooked; some species live in water, many live in soil as decomposers , others live on plants, sometimes creating galls , while others are predators or parasites . This last type includes the commercially destructive Varroa parasite of honey bees , as well as scabies mites of humans. Most species are harmless to humans, but
1302-417: The human skin disease rosacea , although the mechanism by which demodex contributes to the disease is unclear. Ticks are well known for carrying diseases, such as Lyme disease and Rocky Mountain spotted fever . Chiggers are known primarily for their itchy bite , but they can also spread disease in some limited circumstances, such as scrub typhus . The house-mouse mite is the only known vector of
Sphaerolichida - Misplaced Pages Continue
1344-561: The microscope; the short film was shown in London's Alhambra music hall in 1903, causing a boom in the sales of simple microscopes. Long branch attraction LBA was first recognized as problematic when analyzing discrete morphological character sets under parsimony criteria, however Maximum Likelihood analyses of DNA or protein sequences are also susceptible. A simple hypothetical example can be found in Felsenstein 1978 where it
1386-502: The mid- Cretaceous , around 100 million years ago. Most fossil acarids are no older than the Tertiary (up to 65 mya ). Members of the superorders Opilioacariformes and Acariformes (sometimes known as Actinotrichida) are mites, as well as some of the Parasitiformes (sometimes known as Anactinotrichida). Recent genetic research has suggested that Acari is polyphyletic (of multiple origins). A study using molecular data from
1428-752: The mitochondria and nucleus recovered Acariformes as sister to the Solifugae (camel spiders) and Parasitiformes as sister to the Pseudoscorpionida , with other arachnid orders separating these two groupings on the phylogenetic tree, as shown below. Palpigradi False scorpions [REDACTED] Ixodida (ticks) [REDACTED] Parasitic mites, inc. Varroa [REDACTED] Trombidiformes (chiggers, velvet mites, etc) [REDACTED] Sarcoptiformes (dust & fur mites, etc) [REDACTED] Camel spiders [REDACTED] other Arachnids including spiders and scorpions However,
1470-612: The most diverse and successful of all invertebrate groups. They have exploited a wide array of habitats , and because of their small size go largely unnoticed. They are found in freshwater (e.g. the water mites or Hydrachnidia ) and saltwater (most Halacaridae ), in the soil, in forests, pastures, agricultural crops, ornamental plants, thermal springs and caves. They inhabit organic debris of all kinds and are extremely numerous in leaf litter. They feed on animals, plants and fungi and some are parasites of plants and animals. Some 48,200 species of mites have been described, but there may be
1512-506: The nests of birds and other animals. These mites are largely parasitic and consume blood , skin and keratin . Dust mites , which feed mostly on dead skin and hair shed from humans instead of consuming them from the organism directly, evolved from these parasitic ancestors. Ticks are a prominent group of mites that are parasitic on vertebrates, mostly mammal and birds, feeding on blood with specialised mouthparts. Parasitic mites sometimes infest insects. Varroa destructor attaches to
1554-412: The problem arises when sequences from two (or more) lineages evolve rapidly. There are only four possible nucleotides and when DNA substitution rates are high, the probability that two lineages will evolve the same nucleotide at the same site increases. When this happens, a phylogenetic analysis may erroneously interpret this homoplasy as a synapomorphy (i.e., evolving once in the common ancestor of
1596-459: The same site, by adding taxa related to those with the long branches that add additional true synapomorphies to the data, or by using alternative slower evolving traits (e.g. more conservative gene regions). The result of LBA in evolutionary analyses is that rapidly evolving lineages may be inferred to be sister taxa, regardless of their true relationships. For example, in DNA sequence-based analyses,
1638-564: The six-legged larvae. After three moults, the larvae become nymphs, with eight legs, and after a further three moults, they become adults. Longevity varies between species, but the lifespan of mites is short compared to many other arachnids. Mites occupy a wide range of ecological niches . For example, Oribatida mites are important decomposers in many habitats. They eat a wide variety of material including living and dead plant and fungal material, lichens and carrion; some are predatory , though no oribatid mites are parasitic . Mites are among
1680-477: The so-called spider mites (family Tetranychidae ), thread-footed mites (family Tarsonemidae ), and the gall mites (family Eriophyidae ). Among the species that attack animals are members of the sarcoptic mange mites (family Sarcoptidae ), which burrow under the skin. Demodex mites (family Demodecidae ) are parasites that live in or near the hair follicles of mammals , including humans. Being unable to fly, mites need some other means of dispersal. On
1722-505: The stigmata are associated with peritremes: paired, tubular, elaborated extensions of the tracheal system. The higher taxa of mites are defined by these structures: Mite digestive systems have salivary glands that open into the preoral space rather than the foregut. Most species carry two to six pairs of salivary glands that empty at various points into the subcheliceral space. A few mite species lack an anus: they do not defecate during their short lives. The circulatory system consists of
Sphaerolichida - Misplaced Pages Continue
1764-454: The two lineages). The opposite effect may also be observed, in that if two (or more) branches exhibit particularly slow evolution among a wider, fast evolving group, those branches may be misinterpreted as closely related. As such, "long branch attraction" can in some ways be better expressed as "branch length attraction". However, it is typically long branches that exhibit attraction. The recognition of long-branch attraction implies that there
#196803