A cladogram (from Greek clados "branch" and gramma "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to descendants, nor does it show how much they have changed, so many differing evolutionary trees can be consistent with the same cladogram. A cladogram uses lines that branch off in different directions ending at a clade , a group of organisms with a last common ancestor . There are many shapes of cladograms but they all have lines that branch off from other lines. The lines can be traced back to where they branch off. These branching off points represent a hypothetical ancestor (not an actual entity) which can be inferred to exhibit the traits shared among the terminal taxa above it. This hypothetical ancestor might then provide clues about the order of evolution of various features, adaptation, and other evolutionary narratives about ancestors. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational phylogenetics are now very commonly used in the generation of cladograms, either on their own or in combination with morphology.
44-458: See text Statilia is a genus of praying mantis that resembles dead or living grass. Statilia species live in Australia, Africa, Asia and islands. The Mantodea Species File lists: Praying mantis See text Mantises are an order ( Mantodea ) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family
88-446: A t e s {\displaystyle n.states} , c i occupies a range from 1 to ( n . s t a t e s − 1 ) / ( n . t a x a − ⌈ n . t a x a / n . s t a t e s ⌉ ) {\displaystyle (n.states-1)/(n.taxa-\lceil n.taxa/n.states\rceil )} . The retention index (RI)
132-445: A metric to measure how consistent a candidate cladogram is with the data. Most cladogram algorithms use the mathematical techniques of optimization and minimization. In general, cladogram generation algorithms must be implemented as computer programs, although some algorithms can be performed manually when the data sets are modest (for example, just a few species and a couple of characteristics). Some algorithms are useful only when
176-406: A prothorax , a mesothorax , and a metathorax . In all species apart from the genus Mantoida , the prothorax, which bears the head and forelegs, is much longer than the other two thoracic segments. The prothorax is also flexibly articulated, allowing for a wide range of movements of the head and fore limbs while the remainder of the body remains more or less immobile. Mantises also are unique to
220-537: A character in a phylogenetic analysis as they do not contribute anything to our understanding of relationships. However, homoplasy is often not evident from inspection of the character itself (as in DNA sequence, for example), and is then detected by its incongruence (unparsimonious distribution) on a most-parsimonious cladogram. Note that characters that are homoplastic may still contain phylogenetic signal . A well-known example of homoplasy due to convergent evolution would be
264-412: A cladogram can be roughly categorized as either morphological (synapsid skull, warm blooded, notochord , unicellular, etc.) or molecular (DNA, RNA, or other genetic information). Prior to the advent of DNA sequencing, cladistic analysis primarily used morphological data. Behavioral data (for animals) may also be used. As DNA sequencing has become cheaper and easier, molecular systematics has become
308-422: A dataset, the degree to which each character carries phylogenetic information, and the fashion in which additive characters are coded, rendering it unfit for purpose. c i occupies a range from 1 to 1/[ n.taxa /2] in binary characters with an even state distribution; its minimum value is larger when states are not evenly spread. In general, for a binary or non-binary character with n . s t
352-515: A few ground-dwelling species are found actively pursuing their prey. They normally live for about a year. In cooler climates, the adults lay eggs in autumn, then die. The eggs are protected by their hard capsules and hatch in the spring. Females sometimes practice sexual cannibalism , eating their mates after copulation. Mantises were considered to have supernatural powers by early civilizations, including ancient Greece , ancient Egypt , and Assyria . A cultural trope popular in cartoons imagines
396-457: A larger clade. The incongruence length difference test (ILD) is a measurement of how the combination of different datasets (e.g. morphological and molecular, plastid and nuclear genes) contributes to a longer tree. It is measured by first calculating the total tree length of each partition and summing them. Then replicates are made by making randomly assembled partitions consisting of the original partitions. The lengths are summed. A p value of 0.01
440-411: A mantis has two sets of wings: the outer wings, or tegmina , are usually narrow and leathery. They function as camouflage and as a shield for the hindwings, which are clearer and more delicate. The abdomen of all mantises consists of 10 tergites , with a corresponding set of nine sternites visible in males and seven visible in females. The abdomen tends to be slimmer in males than females, but ends in
484-452: A more and more popular way to infer phylogenetic hypotheses. Using a parsimony criterion is only one of several methods to infer a phylogeny from molecular data. Approaches such as maximum likelihood , which incorporate explicit models of sequence evolution, are non-Hennigian ways to evaluate sequence data. Another powerful method of reconstructing phylogenies is the use of genomic retrotransposon markers , which are thought to be less prone to
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#1732858471156528-408: A pair of cerci in both sexes. Mantises have stereo vision . They locate their prey by sight; their compound eyes contain up to 10,000 ommatidia . A small area at the front called the fovea has greater visual acuity than the rest of the eye, and can produce the high resolution necessary to examine potential prey. The peripheral ommatidia are concerned with perceiving motion; when a moving object
572-462: A similar series of tubercles along the tibia and the apical claw near its tip, give the foreleg of the mantis its grasp on its prey. The foreleg ends in a delicate tarsus used as a walking appendage, made of four or five segments and ending in a two-toed claw with no arolium . Mantises can be loosely categorized as being macropterous (long-winged), brachypterous (short-winged), micropterous (vestigial-winged), or apterous (wingless). If not wingless,
616-497: A wide binocular field of vision and precise stereoscopic vision at close range. The dark spot on each eye that moves as it rotates its head is a pseudopupil . This occurs because the ommatidia that are viewed "head-on" absorb the incident light , while those to the side reflect it. As their hunting relies heavily on vision, mantises are primarily diurnal . Many species, however, fly at night, and then may be attracted to artificial lights . They have good night vision. Mantises in
660-410: Is a character state that is shared by two or more taxa due to some cause other than common ancestry. The two main types of homoplasy are convergence (evolution of the "same" character in at least two distinct lineages) and reversion (the return to an ancestral character state). Characters that are obviously homoplastic, such as white fur in different lineages of Arctic mammals, should not be included as
704-430: Is noticed, the head is rapidly rotated to bring the object into the visual field of the fovea. Further motions of the prey are then tracked by movements of the mantis's head so as to keep the image centered on the fovea. The use of stereoscopic vision differs from humans or primates because they specifically utilize this vision for capturing and spotting prey. The eyes are widely spaced and laterally situated, affording
748-411: Is obtained for 100 replicates if 99 replicates have longer combined tree lengths. Some measures attempt to measure the amount of homoplasy in a dataset with reference to a tree, though it is not necessarily clear precisely what property these measures aim to quantify The consistency index (CI) measures the consistency of a tree to a set of data – a measure of the minimum amount of homoplasy implied by
792-463: Is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They have triangular heads with bulging eyes supported on flexible necks. Their elongated bodies may or may not have wings, but all Mantodea have forelegs that are greatly enlarged and adapted for catching and gripping prey; their upright posture, while remaining stationary with forearms folded, has led to
836-526: Is usually done by comparison to the character states of one or more outgroups . States shared between the outgroup and some members of the in-group are symplesiomorphies; states that are present only in a subset of the in-group are synapomorphies. Note that character states unique to a single terminal (autapomorphies) do not provide evidence of grouping. The choice of an outgroup is a crucial step in cladistic analysis because different outgroups can produce trees with profoundly different topologies. A homoplasy
880-460: The Dictyoptera in that they have tympanate hearing, with two tympana in an auditory chamber in their metathorax. Most mantises can only hear ultrasound . Mantises have two spiked, grasping forelegs ("raptorial legs") in which prey items are caught and held securely. In most insect legs, including the posterior four legs of a mantis, the coxa and trochanter combine as an inconspicuous base of
924-691: The Early Cretaceous . Fossils of the group are rare: by 2022, 37 fossil species are known. Fossil mantises, including one from Japan with spines on the front legs as in modern mantises, have been found in Cretaceous amber. Most fossils in amber are nymphs; compression fossils (in rock) include adults. Fossil mantises from the Crato Formation in Brazil include the 10 mm (0.39 in) long Santanmantis axelrodi , described in 2003; as in modern mantises,
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#1732858471156968-863: The Mantidae and Thespidae especially were considered polyphyletic , so the Mantodea have been revised substantially as of 2019 and now includes 29 families. † Extinct Genera Chaeteessidae Mantoididae Metallyticidae Thespidae Angelidae Coptopterygidae Liturgusidae Photinaidae Acanthopidae Chroicopteridae Leptomantellidae Amorphoscelidae Nanomantidae Gonypetidae Epaphroditidae Majangidae Haaniidae Rivetinidae Amelidae Eremiaphilidae Toxoderidae Hoplocoryphidae Miomantidae Galinthiadidae Empusidae Hymenopodidae Dactylopterygidae Deroplatyidae Mantidae Mantises are thought to have evolved from cockroach-like ancestors. The earliest confidently identified mantis fossils date to
1012-431: The character, "presence of wings". Although the wings of birds, bats , and insects serve the same function, each evolved independently, as can be seen by their anatomy . If a bird, bat, and a winged insect were scored for the character, "presence of wings", a homoplasy would be introduced into the dataset, and this could potentially confound the analysis, possibly resulting in a false hypothesis of relationships. Of course,
1056-434: The characteristic data are molecular (DNA, RNA); other algorithms are useful only when the characteristic data are morphological. Other algorithms can be used when the characteristic data includes both molecular and morphological data. Algorithms for cladograms or other types of phylogenetic trees include least squares , neighbor-joining , parsimony , maximum likelihood , and Bayesian inference . Biologists sometimes use
1100-448: The common name praying mantis . The closest relatives of mantises are termites and cockroaches ( Blattodea ), which are all within the superorder Dictyoptera . Mantises are sometimes confused with stick insects ( Phasmatodea ), other elongated insects such as grasshoppers ( Orthoptera ), or other more distantly related insects with raptorial forelegs such as mantisflies ( Mantispidae ). Mantises are mostly ambush predators , but
1144-685: The dataset). The rescaled consistency index (RC) is obtained by multiplying the CI by the RI; in effect this stretches the range of the CI such that its minimum theoretically attainable value is rescaled to 0, with its maximum remaining at 1. The homoplasy index (HI) is simply 1 − CI. This measures the amount of homoplasy observed on a tree relative to the maximum amount of homoplasy that could theoretically be present – 1 − (observed homoplasy excess) / (maximum homoplasy excess). A value of 1 indicates no homoplasy; 0 represents as much homoplasy as there would be in
1188-1101: The family Liturgusidae collected at night have been shown to be predominately males; this is probably true for most mantises. Nocturnal flight is especially important to males in locating less-mobile females by detecting their pheromones . Flying at night exposes mantises to fewer bird predators than diurnal flight would. Many mantises also have an auditory thoracic organ that helps them avoid bats by detecting their echolocation calls and responding evasively. Mantises are generalist predators of arthropods . The majority of mantises are ambush predators that only feed upon live prey within their reach. They either camouflage themselves and remain stationary, waiting for prey to approach, or stalk their prey with slow, stealthy movements. Larger mantises sometimes eat smaller individuals of their own species, as well as small vertebrates such as lizards, frogs, fish, and particularly small birds. Most mantises stalk tempting prey if it strays close enough, and will go further when they are especially hungry. Once within reach, mantises strike rapidly to grasp
1232-534: The female mantis as a femme fatale . Mantises are among the insects most commonly kept as pets . The name mantodea is formed from the Ancient Greek words μάντις ( mantis ) meaning "prophet", and εἶδος ( eidos ) meaning "form" or "type". It was coined in 1838 by the German entomologist Hermann Burmeister . The name "mantid" properly refers only to members of the family Mantidae , which was, historically,
1276-494: The front legs were adapted for catching prey. Well-preserved specimens yield details as small as 5 μm through X-ray computed tomography . Extinct families and genera include: Because of the superficially similar raptorial forelegs , mantidflies may be confused with mantises, though they are unrelated. Their similarity is an example of convergent evolution ; mantidflies do not have tegmina (leathery forewings) like mantises, their antennae are shorter and less thread-like, and
1320-423: The leg; in the raptorial legs, however, the coxa and trochanter combine to form a segment about as long as the femur , which is a spiky part of the grasping apparatus (see illustration). Located at the base of the femur is a set of discoidal spines, usually four in number, but ranging from none to as many as five depending on the species. These spines are preceded by a number of tooth-like tubercles, which, along with
1364-554: The only family in the order. The other common name, praying mantis, applied to any species in the order (though in Europe mainly to Mantis religiosa ), comes from the typical " prayer -like" posture with folded forelimbs. The vernacular plural "mantises" (used in this article) was confined largely to the US, with "mantids" predominantly used as the plural in the UK and elsewhere, until the family Mantidae
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1408-705: The only reason a homoplasy is recognizable in the first place is because there are other characters that imply a pattern of relationships that reveal its homoplastic distribution. A cladogram is the diagrammatic result of an analysis, which groups taxa on the basis of synapomorphies alone. There are many other phylogenetic algorithms that treat data somewhat differently, and result in phylogenetic trees that look like cladograms but are not cladograms. For example, phenetic algorithms, such as UPGMA and Neighbor-Joining, group by overall similarity, and treat both synapomorphies and symplesiomorphies as evidence of grouping, The resulting diagrams are phenograms, not cladograms, Similarly,
1452-462: The order Orthoptera with the cockroaches (now Blattodea ) and ice crawlers (now Grylloblattodea ). Kristensen (1991) combined the Mantodea with the cockroaches and termites into the order Dictyoptera , suborder Mantodea. Evolutionary relationships based on Evangelista et al. 2019 are shown in the cladogram : (Mantises) (Cockroaches and termites) One of the earliest classifications splitting an all-inclusive Mantidae into multiple families
1496-559: The prey with their spiked raptorial forelegs. Some ground and bark species pursue their prey in a more active way. For example, members of a few genera such as the ground mantises Entella , Ligaria , and Ligariella run over dry ground seeking prey, much as tiger beetles do. Some mantis species such as Euantissa pulchra can discriminate between different types of prey, and approached spiders mimicking non-aggressive ant species much more than spiders that mimicked aggressive ant species. Cladogram The characteristics used to create
1540-499: The problem of reversion that plagues sequence data. They are also generally assumed to have a low incidence of homoplasies because it was once thought that their integration into the genome was entirely random; this seems at least sometimes not to be the case, however. Researchers must decide which character states are "ancestral" ( plesiomorphies ) and which are derived ( synapomorphies ), because only synapomorphic character states provide evidence of grouping. This determination
1584-436: The program settles on a local minimum rather than the desired global minimum. To help solve this problem, many cladogram algorithms use a simulated annealing approach to increase the likelihood that the selected cladogram is the optimal one. The basal position is the direction of the base (or root) of a rooted phylogenetic tree or cladogram. A basal clade is the earliest clade (of a given taxonomic rank[a]) to branch within
1628-464: The raptorial tibia is more muscular than that of a similar-sized mantis and bends back farther in preparation for shooting out to grasp prey. Mantises have large, triangular heads with a beak-like snout and mandibles . They have two bulbous compound eyes , three small simple eyes, and a pair of antennae . The articulation of the neck is also remarkably flexible; some species of mantis can rotate their heads nearly 180°. The mantis thorax consists of
1672-405: The results of model-based methods (Maximum Likelihood or Bayesian approaches) that take into account both branching order and "branch length," count both synapomorphies and autapomorphies as evidence for or against grouping, The diagrams resulting from those sorts of analysis are not cladograms, either. There are several algorithms available to identify the "best" cladogram. Most algorithms use
1716-460: The term parsimony for a specific kind of cladogram generation algorithm and sometimes as an umbrella term for all phylogenetic algorithms. Algorithms that perform optimization tasks (such as building cladograms) can be sensitive to the order in which the input data (the list of species and their characteristics) is presented. Inputting the data in various orders can cause the same algorithm to produce different "best" cladograms. In these situations,
1760-406: The tree. It is calculated by counting the minimum number of changes in a dataset and dividing it by the actual number of changes needed for the cladogram. A consistency index can also be calculated for an individual character i , denoted c i . Besides reflecting the amount of homoplasy, the metric also reflects the number of taxa in the dataset, (to a lesser extent) the number of characters in
1804-423: The user should input the data in various orders and compare the results. Using different algorithms on a single data set can sometimes yield different "best" cladograms, because each algorithm may have a unique definition of what is "best". Because of the astronomical number of possible cladograms, algorithms cannot guarantee that the solution is the overall best solution. A nonoptimal cladogram will be selected if
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1848-403: Was further split in 2002; at present, only some 80 out of 430 known genera are mantids, the rest are in other families. Over 2,400 species of mantis in about 430 genera are recognized. They are predominantly found in tropical regions, but some live in temperate areas. The systematics of mantises have long been disputed. Mantises, along with stick insects ( Phasmatodea ), were once placed in
1892-399: Was proposed as an improvement of the CI "for certain applications" This metric also purports to measure of the amount of homoplasy, but also measures how well synapomorphies explain the tree. It is calculated taking the (maximum number of changes on a tree minus the number of changes on the tree), and dividing by the (maximum number of changes on the tree minus the minimum number of changes in
1936-416: Was that proposed by Beier in 1968, recognizing eight families, though it was not until Ehrmann's reclassification into 15 families in 2002 that a multiple-family classification became universally adopted. Klass, in 1997, studied the external male genitalia and postulated that the families Chaeteessidae and Metallyticidae diverged from the other families at an early date. However, as previously configured,
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